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THE NORTH AMERICAN SPECIES OF
THE BEE GENUS UTHURGE
(HYMENOPTERA: MEGACHILIDAE)

Roy R. Snelling

Number 343
5 July 1983

Natural History Museum of Los Angeles County « ftOO Exposition -Boulevard •* Los Angeles, California 9001]?

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THE NORTH AMERICAN SPECIES OF
THE BEE GENUS LITHURGE
(HYMENOPTERA: MEGACHILIDAE)

Roy R. Snelling1

ABSTRACT. The North American species of Lithurge are re-
viewed, and a key is given for the separation of both sexes of all
species. Supraclypeal structures of the females are illustrated, as are
the labra) structures of the males. Of the previously described native
forms, L. apicalis var. opuntiae Cockerell and L. arizonensis Cock-
erell are synonyms of L. apicalis Cresson (NEW SYNONYMY); L.
apicalis subsp. lilloralis Cockerell is a valid species and a senior
synonym of L. bruesi Mitchell (NEW SYNONYMY). One new
species, L. listrota. is described from southern California desert re-
gions. All native American species belong to the subgenus Lithur-
gopsis.

Two representatives of the nominate subgenus occur in the New
World. The Palaearctic species, L. chrysurus (Fonscolombe). is ad-
ventive in the eastern United States. A South American species, L.
huberi Ducke, belongs to the Indo-Australian species group of L.
atratus F. Smith and may have been introduced into Brazil in historic
times.

The South American genera Trichothurgus Moure and Lithur-
gomma Moure are merged into a single genus, Trichothurgus, and
the tribe Trichothurgini is dropped.

INTRODUCTION

The megachilid subfamily Lithurginae includes about 50
species in both Old and New World faunas. Old World species
all belong to the nominate subgenus of Lithurge Latreille and
are present in the southern Palaearctic, Ethiopian and Indo-
Australian regions. These species, as far as known, provision
their nests with pollen from plants of the families Malvaceae
and Compositae.

For the most part. New World species of Lithurge belong
to the subgenus Lithurgopsis Fox and are present in southern
North America and South America. One Palaearctic species,
L. chrysurus (Fonscolombe), is adventive in the eastern United
States (Roberts, 1978); a Brazilian species, L. huberi Ducke,
also belongs to the nominate subgenus and is probably in-
troduced from the Indo-Australian region. Although most
Lithurgopsis utilize pollen from Cactaceae in their larval cells,
a few Argentinian species use malvaceous pollens (J. L. Neff,
in litt.). Some Chilean and Argentinian species, formerly
placed in Lithurge, are now placed in the genera Tricho-
thurgus and Lithurgomma, both described by Moure (1949);

Contributions in Science, Number 343, pp. 1-11

Natural History Museum of Los Angeles County, 1983

at least one of these is an oligolege on Compositae (J. L. Nelf,
in litt.).

THE NEW WORLD LITHURGINAE

Most New World Lithurginae have been described in the
genus Lithurgus Berthold, 1827, an invalid emendation of
Lithurge Latreille, 1825. Fox (1902) noted that males of the
New World species possess tarsal arolia and proposed to
remove them to a new genus, Lithurgopsis. Subsequent work-
ers have consistently treated Lithurgopsis as a subgenus of
Lithurge (e.g., Michener, 1944; Hurd, 1979).

In addition to the presence of tarsal arolia in Lithurgopsis
males, there are other features by which members of the
subgenus may be separated from the nominotypical subge-
nus. In females of Lithurgopsis the facial prominence, when
present, occurs only on the supraclypeal area; in Lithurge,
sensu stricto, the basal portion of the clypeus is also involved.
The first flagellar segment is consistently short in both sexes
of Lithurgopsis ; it is little, if any, longer than the second and
often much shorter; the second segment is as long as broad
or longer. In Lithurge, sensu stricto, the first flagellar segment
is as long as broad, or longer, the second segment not more
than half as long as the first; the second segment is often
conspicuously broader than long.

Lithurge (L.) chrysurus (Fonscolombe, 1834) is a Palaearc-
tic species that is now introduced and established in the
eastern United States. Roberts (1978) reported this Medi-
terranean species from New Jersey, discussed its nesting bi-
ology and behavior, and described the larva and pupa.

Lithurge (L.) huberi (Ducke, 1907), described from Brazil,
is the only South American representative of the nomino-
typical subgenus. I have seen specimens of both sexes of this
species. Morphologically, L. huberi is a member of the group
of forms related to L. atratus (F. Smith). Nominate species
of this group range from India to Australia, Melanesia, and
Polynesia; according to Michener ( 1965) there may be but a

1. Entomology Section, Natural History Museum of Los Angeles
County.

ISSN 0459-8113

single species involved. There are no obvious morphological
features by which L. huberi may be separated from L. alratus.
It seems likely that L. huberi is adventive in Brazil, intro-
duced via infested wood within historic times.

Although most New World Lithurginae belong to the genus
Litharge, Moure ( 1 949) described two genera, Trichothurgus
and Lithurgomma, which he placed in the new tribe Tricho-
thurgini. Species of Trichothurgini occur in Peru and Argen-
tina but are primarily Chilean. Trichothurgus and Lithur-
gomma differ from Litharge in the elongate labrum and in
the shape of the first gastric tergum, which is not flattened
as in Litharge. The species of these two genera are large,
robust bees with abundant body hairs that conceal most in-
tegumentary surfaces.

In Trichothurgus, the facial prominence, if present, is on
the supraclypeal area and the frons lacks a shiny area in front
of the anterior ocellus. Species of Lithurgomma have a clyp-
eal prominence, in females only, and there is a large, flat-
tened, shiny area in front of the anterior ocellus. These dif-
ferences are trivial and do not, in my opinion, justify the
status of Trichothurgus and Lithurgomma as separate genera.
My view is that Lithurgomma is a junior synonym of Tricho-
thurgus (NEW SYNONYMY). The characters of Moure’s
Trichothurgini will differentiate the expanded genus Tricho-
thurgus from Litharge. Concurrently, the tribe Trichothur-
gini is superfluous; there is no need to place the two genera
of Lithurginae in separate tribes.

The known species of Trichothurgus are: T. a/biceps (Friese),
T. alpestris (Friese), T. aterrimus (Cockerell), T. colloncu-
rensis Ogloblin, T. herbsti (Friese), T. holomelan (Moure),
T. laticeps (Friese), T. neoqueensis (Friese), T. osmioides
(Friese), T. pseudocellalus (Moure), T. shajovskoyi Ogloblin,
and T. wagenknechti (Moure).

The few South American species of Lithurge have never
been critically examined. The North American species were
reviewed by Mitchell (1938), who gave a key for the species
then known. The biology of no American species has been
studied in detail; some notes on L. apicalis (Cresson) were
published by Parker and Potter (1973), and Brach (1979)
reported some observations on L. gibbosa (F. Smith). The
nesting biology and behavior of the introduced species, L.
chrysurus, were described by Roberts (1978).

SPECIMENS EXAMINED

During the course of this study, specimens from the following
institutional collections were examined: American Museum
of Natural Flistory (AMNH), Arizona State University
(ARSU), British Museum (Natural History) (BMNH), Cal-
ifornia Academy of Sciences (CAS), California Department
of Food and Agriculture (CDFA), National Museum of Nat-
ural History (USNM), Natural History Museum of Los An-
geles County (LACM), San Diego Natural History Museum
(SDM), Texas A and I University (TAI), Texas A and M
University (TAMU), University of Arizona (UNAR), Uni-
versity of California, Berkeley (UCB), University of Califor-
nia, Davis (UCD), University of California, Riverside (UCR),
University of Colorado (UCOL), University of Kansas

(UKAN), University of Nebraska (UNEB), and Utah State
University (UTSU). Specimens from the personal collection
of T. Griswold are indicated (GRIS).

SYSTEMATICS

In the review by Mitchell (1938), five species were treated
in the North American fauna, one with three subspecies. A
sixth species was mentioned but not treated as it had been
too recently described and was unknown to Mitchell. The
present study recognizes eight species, including one adven-
tive species and one that was previously undescribed. The
various subspecies attributed to L. apicalis by Mitchell ( 1 938)
are discussed below. The morphological characteristics of
each species are sufficiently distinctive that the previously
described species are not redescribed; the keys and figures
given here are adequate for their recognition.

Superficially, the various species tend to be quite similar.
Females are most notably different from one another in the
shape of the elevations or processes of the supraclypeal area.
There are differences, also, in the density of the punctures of
the clypeus and supraclypeal area, but there is sufficient infra-
specific variability to render these unreliable as diagnostic
characters.

Males are most readily separated by differences in labral
structure. When fresh, individuals of this sex should have
the mandibles spread so that the external surface of the la-
brum is clearly visible. Although the mandibles of a dry
specimen can be spread after relaxing the bee, there is a
greater chance that the specimen will be damaged.

KEY TO NORTH AMERICAN LITHURGE
Males

la. Tarsal arolia present; first flagellar segment shorter than

second (subgenus Lithurgopsis) 2

b. Tarsal arolia absent; first flagellar segment distinctly

longer than second (subgenus Lithurge)

chrysurus (Fonscolombe)

2a. Labrum with a single median tubercle, or none .... 3
b. Labrum with a pair of erect, slender tubercles (Fig. 4)

littoralis (Cockerell)

3a. Labrum without median tubercle, with a low, transverse

basal ridge or swelling (Figs. 6, 10) 4

b. Labrum with an erect, median tubercle, without trans-
verse basal ridge or swelling (Figs. 8, 12) 6

4a. Supraclypeal area gently convex, its upper portion slop-
ing toward antennal sockets, and with a definite median
area that is less closely punctate than area near suban-
tennal suture or entire supraclypeal area sparsely and

irregularly punctate 5

b. Supraclypeal area uniformly flat between clypeal base
and antennal sockets, continguously punctate, punctures

coarse and subcontiguous along midline

planifrons (Friese)

5a. First flagellar segment, on shortest side, distinctly broad-
er than long, shorter than pedicel; supraclypeal area about

2 Contributions in Science, Number 343

Snelling: North American Lithurge

1.4 times wider than long; legs usually reddish

echinocacti (Cockerell)

b. First flagellar segment, on shortest side, about as long as
wide, about as long as pedicel; supraclypeal area about

1.8 times wider than long; legs brown

apicalis (Cresson)

6a. Labrum with a low, often obsolescent, ridge extending
obliquely from basal corner to base of median tubercle

(Figs. 2, 12) 7

b. Labrum deeply concave on either side of tubercle and
with a short, curved ridge from base of tubercle to mid-
length of lateral margin (Fig. 8)

socorroensis (Mitchell)

7a. Supraclypeal area slightly protuberant, densely punctate;
ocelloccipital distance greater than interocellar distance

gibbosa (F. Smith)

b. Supraclypeal area flat, sparsely punctate, at least in mid-
dle; ocelloccipital distance less than interocellar distance
listrota new species

Females

la. First flagellar segment shorter than second; facial prom-
inence, when present, not as below (subgenus Lithur-

gopsis) 2

b. First flagellar segment distinctly longer than second; fa-
cial prominence high, triangular in lateral view, its lower
portion sloping toward base of clypeus (subgenus Li-
tharge) chrysurus (Fonscolombe)

2a. Supraclypeal protuberance nearly as broad as face, ex-
tending laterad of subantennal sutures (Figs. 3, 5); malar
area distinct and with deep posterior pit (Figs. 3, 5) 3

b. Supraclypeal protuberance, when present, much narrow-
er than width of face, not extending laterad of suban-
tennal sutures (Figs. 7, 9, 1 1 ); malar area linear, mandible
nearly contiguous with eye, posterior pit usually absent

(Figs. 1 , 7, 9) 4

3a. Labral tubercle with median and sublateral impressions;

clypeus without preapical ridge (Fig. 3)

/ itt oralis (Cockerell)

b. Labral tubercle divided in middle only (Fig. 5); clypeus
with preapical ridge which is interrupted in middle (Fig.

5) apicalis (Cresson)

4a. Supraclypeal area distinctly elevated on each side of mid-
dle or with a single, transverse, bowed elevation .5
b. Supraclypeal area flat, with no protuberance, shiny and
very sparsely punctate (Fig. 1) ... listrota new species
5a. Supraclypeal area with prominent rounded elevation on
either side of middle (Figs. 7, 9); malar area without

posterior pit 6

b. Supraclypeal elevation a high, transverse, bowed ridge
(Fig. 1 1 ); malar area with broad, shallow, posterior pit

gibbosa (F. Smith)

6a. Dorsal margin of pronotum with conspicuous fascia of
dense, plumose, white hairs; ocelloccipital distance less

than interocellar distance echinocacti (Cockerell)

b. Dorsal margin of pronotum without fascia of dense, plu-

mose, white hairs; ocelloccipital distance greater than
interocellar distance socorroensis (Mitchell)

Lithurge ( Lithurgopsis ) apicalis (Cresson)

Figures 5, 6

Lithurgus apicalis Cresson, 1875:724. 9.

Lithurgopsis apicalis var. opuntiae Cockerell, 1 902; 182.9.
NEW SYNONYMY.

Lithurgus arizonensis Cockerell, 1937:108. 9. NEW SYN-
ONYMY.

RANGE

South Dakota and Wyoming to western Texas (?), west to
Utah, Nevada, and southern California; northern Mexico.

Although this species has been recorded from Texas by
previous authors (Cockerell, 1911; Mitchell, 1938), I have
seen no material of L. apicalis from that State. All specimens
from Texas previously identified as L. apicalis, that I have
examined, proved to be L. littora/is.

BIONOMICS

Parker and Potter (1973) published some notes on the nesting
habits of L. apicalis. The larva has been described by Rozen
(1973).

Females provision the cells with pollen from species of
Opuntia, but both sexes have been taken at flowers of other
genera. Hurd ( 1979) recorded this species on flowers of Echi-
nocactus, Ence/ia californica, Eriogonum, and Sphaeralcea.
Previously unpublished floral records include: Agave neva-
densis, Argemone platyceros, Argemone sp., Asclepias erosa,
Baileya multiradiata, Cirsium californicum, Cirsium sp.,
Calochortus concolor, Chi/opsis linearis, Cleome serrulata,
Cnicus sp., Grindelia squarrosa, Prosopis juliflora, Robinia
neomexicana, Senecio longi/obus, Verbena stricta, Verbesina
encelioides, and “sweet pea.”

DISCUSSION

For many years, L. apicalis has been divided into two sub-
species: L. a. apicalis and L. a. opuntiae. The former, with
ferruginous hairs on the last tergum of the female, is a north-
ern form, found in South Dakota, Wyoming, Nebraska, Kan-
sas, Colorado, Utah, and Nevada. The southern population,
L. a. opuntiae, with dark brown hairs on the last tergum of
the female, ranges from New Mexico to southern California.
Both forms, however, occur in New Mexico, Arizona, and
California. In many areas the two occur together, along with
many specimens of intermediate character.

In view of the broad zone of mtergradalion, were mixing
of the forms occurs and which actually includes much of the
range of L. a. opuntiae, it hardly seems desirable to separate
the two forms nomenclatorially. Accordingly, L. a. opuntiae
its here treated as a synonym of L. apicalis.

Cockerell (1937) described L. arizonensis from two fe-
males collected in the Baboquivari Mountains of Arizona.
These were said to differ from L. a. opuntiae by their more

Contributions in Science, Number 343

Snelling: North American Lithurge 3

Figures 1-6. Lithurge spp. Figs. 1, 3, 5, lower portion of face of female of L. listrota (1), L. littoralis (3). and L. apicalis (5). Scale line = 1.0
mm. Figs. 2, 4, 6, labrum of male of L. listrota (2), L. littoralis (4), and L. apicalis (6). Scale line = 0.5 mm.

4 Contributions in Science, Number 343

Snelling: North American Lithurge

flattened and coarsely punctate clypeus and by the deeper,
sparser facial punctures. I have examined the type of L.
arizonensis (in the CAS) and find no significant differences
between this specimen and other individuals of L. apicalis.
All the differences cited by Cockerell fall well within the range
of character variation that 1 attribute to this species.

The male from Port Isabel, Texas, described by Cockerell
( 1 9 1 7) as L. apicalis subsp. littora/is, is a distinct species and
a senior synonym of L. bruesi (see below).

TYPE MATERIAL

Lithurgus apicalis Cresson: female, in Academy of Natural
Sciences, Philadelphia. Lithurgopsis apicalis var. opuntiae
Cockerell: no types designated, and none of the original ma-
terial can be identified with surety. Lithurgus arizonensis
Cockerell: female, in California Academy of Sciences, San
Francisco.

SPECIMENS EXAMINED (566 29, 423 33)

UNITED STATES. ARIZONA, Apache Co.: White Mts.
(CAS); Carrizo (UNAR). Cochise Co.: Chiricahua Mts. (Bar-
foot Park, 8600 ft. elev.; Cave Creek; Pinery Canyon; Portal
and vicinity; Southwest Research Station; Sunny Flat; AMNFI,
ARSU, LACM, UNAR, UCB, UCD, UKAN, UTSU); Hua-
chuca Mts. (Ramsey Canyon; UKAN, UNAR); Mustang Mts.
(UKAN); 6 mi. SE of Willcox (AMNH); Douglas (UCB).
Coconino Co.: 24 mi. W of Cameron (UKAN, UNEB); Fre-
donia (UTSU); Grand Canyon Natl. Park (LACM, UCD,
UNAR); Flagstaff and vicinity (UCB, UNAR, USNM); Oak
Creek Canyon (CAS, LACM, UKAN, UNEB, UTSU); Se-
dona (UKAN, UNAR). Gila Co.: Florence (ARSU); 18 mi.
SE of Globe (UKAN); 18 mi. S of Payson (UNAR); San
Carlos Lake (UNAR). Greenlee Co.: 23 mi. N of Clifton
(ARSU). Maricopa Co.: Granite Reef Dam (ARSU). Navajo
Co.: Fort Apache (UNAR); Cedar Creek, 15 mi. W of Fort
Apache (UNAR); 16 mi. SW of Kayenta (ARSU); Marsh
Pass (USNM, UTSU). Pima Co.: Baboquivari Mts. (Brown
Canyon; Elkhorn Ranch Canyon; AMNH, CAS, LACM,
UKAN, LINAR); Redington and Redington Pass (LINAR);
Rincon Mts. (Madrona Ranger Station; 1.5 mi. NE of X-9
Ranch; UNAR); Santa Rita Mts. (Gardner Canyon; Madera
Canyon; LACM, UKAN, UNAR); Santa Catalina Mts. (Sa-
bino Canyon; UNAR); Silver Bell bajada (LACM); Tucson
and vicinity (UCD, UNAR, UNEB, USNM, UTSU). Pinal
Co.: Oracle (USNM); Superior (B. Thompson Arboretum;
UNAR). Santa Cruz Co.: Atascosa Mts. (Sycamore Canyon;
UNAR); Nogales (USNM); Patagonia (UNAR); Sonoita
(LACM). Yavapai Co.: Ash Fork (USNM); Hell Canyon
(ARSU); Jerome and vicinity (UKAN, UNEB); Mingus Mt.
(UTSU); 5 mi. NE of Paulden (ARSU); Prescott and vicinity
(LACM, UCB); 24 mi. SE of Wikieup (ARSU). County un-
known: White River (UNAR; this locality name occurs in
three different counties). CALIFORNIA, Inyo Co.: Bristle-
cone Natl. Forest (Schulman Grove; UCD, UTSU); Argus
Mts. (LACM); Mountain Spring (UCB); Panamint Mts. (Ben-
nett Peak, 9980 ft. elev.; Frenchman’s Canyon, 6800-7200
ft. elev.; GRIS, UCR, UTSU); White Mts. (Wyman Canyon.
8500 ft. elev.; UCB). Los Angeles Co.: Eagle Rock (UKAN).

Mono Co.: Benton (UCD); Cottonwood Creek (9300 ft. elev.;
UCB); Mammoth Lake (UCD, LITSU). Riverside Co.: Anza
and vicinity (CDFA, LACM, UCB, UCD, UTSU); Banning
(CAS); Deep Canyon (UCB, UCR); Joshua Tree Natl. Mon-
ument (UCR); Mt. San Jacinto (UTSU); 4 mi. S of Palm
Desert (UCB); Palm Springs (UCD); Pinon Flat (LACM,
UCB, UKAN); Ribbonwood (UCB); Santa Rosa Peak (8000
ft. elev.; CAS). San Bernardino Co.: Caruthers Canyon (5650
ft. elev.; GRIS); Cedar Canyon (5100 ft. elev.; GRIS); 9.5
mi. NNE of Cima (4200-4265 ft. elev.; LACM); Cima Dome
(5050 ft. elev.; GRIS); Clark Mtn. (6000 ft. elev.; GRIS);
Deep Creek Public Camp (LACM); Dove Spring (4600 ft.
elev.; GRIS); Falls Public Camp (UCB); Forest Home (CAS,
UCR); Granite Mts. (4050 ft. elev., GRIS); Joshua Tree
(LACM); Lower Covington Flat (LACM); Mitchell’s Caverns
(GRIS); s. fork, Santa Ana River (UNEB); Wheaton Springs
(UCB). San Diego Co.: Anza-Borrego Desert State Park
(LACM); 5 mi. W of Borrego Springs (UCD); Jacumba
(LACM); 10-1 1 mi. E of Julian (LACM); Laguna (SDM);
Mt. Laguna (UCB); Oak Grove (UCR); San Felipe Wash
(LACM, UCR); 2 mi. N of Warner Springs (LACM, UCB,
UCD, UTSU). COLORADO, Alamosa Co.: Great Sand
Dunes Natl. Monument (LACM, UCOL, UKAN). Archuleta
Co.: Arboles (USNM); Yellow Jacket (UCOL). Boulder Co.:
Boulder and vicinity (AMNH, UCOL, UKAN); Gregory
Canyon (AMNH); Pinecliffe (UKAN). Chaffee Co.: Buena
Vista (7900-8000 ft. elev.; AMNH). Costilla Co.: Fort Gar-
land (UNEB). Crowley Co.: Fowler (UCOL). Delta Co.: 5
mi. N of Cedaredge (UCOL); Delta (UCOL); Paonia (CAS).
£7 Paso Co.: Manitou (UCB, UKAN, USNM); Ute Creek
(LJNEB). Fremont Co.: Canon City (UKAN); 10 mi. NE of
Cotopaxi (5900 ft. elev.; UKAN). Garfield Co.: Glenwood
Springs (AMNH). Huerfano Co.: Cuchara Dam (UCOL);
Lathrop State Park (UCOL); 2-3 mi. W of Walsenberg
(UCOL, UKAN). Mesa Co.: Colorado Natl. Monument
(UCOL). Moffat Co.: Dinosaur Natl. Monument (Castle Rock;
Red Rock Ranch; UCOL). Montrose Co.: Naturita (UKAN).
Ouray Co.: Ridgway (AMNH). KANSAS, Hamilton Co.:
(UKAN). Kingman Co.: Calista (UKAN). Pratt Co.: 8 mi.
NE of Isabel (UKAN). NEBRASKA, Arthur Co.: 4 mi. N of
Arthur (UKAN). Sioux Co.: Warbonnet Canyon (UNEB).
NEVADA, Clark Co.: (SDM); Charleston Mts. (Lee Canyon;
Willow Creek Camp. AMNH, UCD). Humboldt Co.: Win-
nemucca (UTSU). NEW MEXICO, Bernalillo Co.: Cedro
Canyon (UKAN); Cienega Canyon (UKAN); Sandia Mts.
(UKAN). Chavez Co.: Elk (UNAR). Dona Ana Co.: Las
Cruces (AMNH, BMNH, UCOL, UKAN, UTSU). Grant
Co.: 13 mi. N of Silver City (6900 ft. elev.; AMNH). Gua-
dalupe Co.: Vaughn (UNEB). Hidalgo Co.: Post Office Can-
yon (ARSU). Lincoln Co.: Alto (UKAN); Capitan (UNEB);
Lincoln (LJKAN); Nogal (UKAN, UNEB); Ruidoso and vi-
cinity (AMNH, UKAN). McKinley Co.: Pinedale (USNM).
Otero Co.: High Rolls (AMNH). Rio Arriba Co.: Ghost Ranch
(UNAR); Echo Canyon (UCR). Sandoval Co.: Jemez Spring
(AMNH); 10 mi. N of Jemez Spring (7500 ft. elev.; AMNH);
San Ysidro (AMNH). San Juan Co.: Bloomfield (UTSU).
Santa Fe Co.: Lamy (USNM); Nambe (UTSU); Santa Fe
and vicinity (CAS, UKAN, UNEB). Torrance Co.: Durran

Contributions in Science, Number 343

Snelling: North American Lithurge 5

(UCB). OKLAHOMA, Custer Co.: Weatherford (UKAN).
Cimarron Co.: Black Mesa (UTSU). SOUTH DAKOTA,
Custer Co.: Custer (UNEB). UTAH, Beaver Co.: Beaver
(UTSU); 16 mi. E of Beaver (Keat’s Lake, 8800 ft. elev.;
AMNH). Cache Co.: Logan (UCD). Duchesne Co.: Roosevelt
(UTSU). Garfield Co.: Escalante River (UTSU). Grand Co.:
Castle Valley (UTSU). Juab Co.: Eureka (UTSU). Kane Co.:
Kanab (UTSU); Navajo Mt. (UTSU). San Juan Co.: 5 mi.
W of Monticello (Dalton Springs, 850 ft. elev.; AMNH).
Tooele Co.: (UCD). Uintah Co.: Vernal (UTSU). Utah Co.:
Thistle (UTSU). WYOMING, Albany Co.: Laramie and vi-
cinity (UKAN, UTSU). Sweetwater Co.: Green River
(AMNH). Weston Co.: Newcastle (USNM). MEXICO. BAJA
CALIFORNIA: Canon del Tajo (LACM); 3 mi. S of Encinas
(UCB) SONORA: San Bernardo (LACM).

Lithurge ( Lithurgopsis ) echinocacti (Cockerell)

Figures 9, 10

Lithurgus echinocacti Cockerell, 1898:453. 9.

RANGE

New Mexico to southern California; northwestern Mexico,
south to Nayarit.

BIONOMICS

In Arizona, L. echinocacti has been observed nesting in rotted
wood (Parker and Potter, 1973). Females collected near Sa-
huarita, Pima County, Arizona, are noted to have been “bor-
ing in post attached to house.”

Known pollen sources for females are various species of
Echinocactus. Other lloral records for this bee include Ar-
gemone sp., Baccharis sp., Baileya pleniradiata. Chilopsis
linearis. Ferocactus wis/izeni. Heterotheca sp., Parkinsonia
aculeata. and Tamarix pentandra.

DISCUSSION

This appears to be an uncommon species. Females may be
easily recognized by the combination of two conical protu-
berances on the supraclypeal area, the presence of a pubescent
fascia on the dorsal margin of the pronotum, and the usually
red legs. Males are similar to those of L. apicalis in that both
lack labral tubercles, but differ in the narrower supraclypeal
area.

TYPE MATERIAL

Lithurgus echinocacti Cockerell: female, present location un-
known.

SPECIMENS EXAMINED (38 99, 97 <?<$)

UNITED STATES. ARIZONA, Cochise Co.: Apache Pass
(LACM); 12 mi. E of Benson (Texas Canyon, 4800 ft. elev.;
LACM); Portal and vicinity (LACM, UCB, UCR). Graham
Co.: Dripping Spring, Whitlock Mts. (LINAR). Pima Co.:
Florida Wash (SDM); Santa Rita Mts. (Box Canyon; Madera

Canyon; IBP site; AMNH, LACM, UCD, UNAR, UTSU);
Silver Bell bajada (LACM); Mountain View (LACM, UCR);
Santa Catalina Mts. (Sabino Canyon; Peppersauce Canyon;
AMNH, UCR, UNAR); Continental (UCR, USNM); Ba-
boquivari Mts. (Brown Canyon; Kitts Peak; AMNH, UKAN,
UNAR); Sahuarita (UNAR); Tucson and vicinity (AMNH,
CAS, UCB, UKAN, UNAR, UTSU). Pinal Co.: Oracle and
vicinity (CAS, LACM, UCR); Ray (ARSU); Superior (B.
Thompson Arboretum; UNAR). Santa Cruz Co.: 20 mi. N
of Nogales (Tumacacori; UNAR); Pena Blanca (UCD). CAL-
IFORNIA, Inyo Co.: 5 mi. E of Valley Wells (UCR). NEW
MEXICO, Hidalgo Co.: 19 mi. N of Rodeo (UTSU). MEXI-
CO. BAJA CALIFORNIA: 13 mi. SE of Millers Landing
(ARSU); 1 1 mi. E of Rosalillito (UNAR); 13 km. NW of
Rosarito (LACM); Valle de Santa Maria (LACM). BAJA
CALFORNIA SUR: 48 km. S of Loreto (LACM); Los Ba-
rriles (CDFA). NAYARIT: 43 mi. N of Tepic (UCB). SO-
NORA: Guaymas (UTSU); Rio Mayo (UTSU); San Bernar-
do (LACM).

Lithurge ( Lithurgopsis ) gihbosa (F. Smith)

Figures 11, 12, 13

Lithurgus gibbosus F. Smith, 1853:147. 9.

Lithurgus compressus F. Smith, 1853:147. <3.

RANGE

North Carolina to Florida, west to Kansas, Oklahoma, and
Texas.

BIONOMICS

Notes on the nesting biology of L. gibbosa were published
by Brach (1979). Females provision cells with pollen from
species of Opuntia. Mitchell (1962) has recorded this bee
from flowers of the genera Cirsium. Helianthus, Ilex. Pon-
taderia, and Rudbeckia. I have also seen a few males taken
at Centaurea americana. Ratibida sp., and Teucrium sp. in
Texas.

DISCUSSION

The female of L. gibbosa is easily recognized by the high,
bowed supraclypeal process, similar to that of members of
the Palaearctic species allied to L. cornuta (Fabricius).

Males are recognizable by the combination of densely
punctate supraclypeal area, the presence of a single labral
tubercle, and the long ocelloccipital distance. The labral tu-
bercle is a curved, transverse, subapical ridge; in the center,
this ridge is elevated. Males from Florida and Georgia have
this median elevation abrupt and quite prominent (Fig. 12).
The median elevation in males from Texas is less pronounced
(Fig. 13). This difference between specimens from the two
areas is consistent but does not correlate with other features.

TYPE MATERIAL

Lithurgus gibbosus F. Smith: females, British Museum (Nat-

6 Contributions in Science, Number 343

Snelling: North American Lithurge

Figures 7-13. Lithurge spp. Figs. 7, 9, 1 1, lower portion of face of female of L. socorroensis (7). L. echinocacti (9), and L. gibbosa (11). Scale
line = 1.0 mm. Figs. 8, 10, 12, 13, labrum of male of L. socorroensis (8), L. echinocacti (10), and L. gibbosa from Florida and Texas (12 and
13, respectively). Scale line = 0.5 mm.

Contributions in Science, Number 343

Snelling: North American Lithurge 7

ural History). Lithurgus compressus F. Smith: male, British
Museum (Natural History).

SPECIMENS EXAMINED (123 29, 98 33)

UNITED STATES (western records only). KANSAS, King-
man Co.: Calista (UKAN). OKLAHOMA, Caddo Co.: Red
Rock Canyon State Park (UKAN). TEXAS. Angelina Co.:
10 mi. N of Lufkin (AMNH). Bastrop Co.: Bastrop and vi-
cinity (UKAN, TAMU); McDade (UKAN). Bexar Co.:
(TAMU, UTSU); San Antonio (USNM). Brazos Co.: (TAMU,
UCB); College Station (TAMU). Crockett Co.: (TAMU). De
Witt Co.: Cuero (AMNH). Gillespie Co.: Fredricksburg
(USNM). Goliad Co.: (TAMU). Gonzales Co.: Palmetto State
Park (UCB). Howard Co.: Big Spring (USNM). Jackson Co.:
Edna (UKAN). Kerr Co.: Kerrville (LISNM). La Salle Co.:
Cotulla (USNM). Lee Co.: Fedor (AMNH, USNM); Gid-
dings (UKAN). Matagorda Co.: Matagorda (UKAN). Mav-
erick Co.: (AMNH). Robertson Co.: (TAMU). Shackleford
Co.: 3 mi. S of Throckmorton (UKAN). Torrant Co.: Fort
Worth (TAMU). Zava/la Co.: Nueces River (USNM).

Lithurge ( Lithurgopsis ) listrota, new species

Figures 1, 2

DIAGNOSIS

Both sexes are separable from all previously described Li-
thurgopsis by the flat supraclypeal area with a shiny im-
punctate median line that extends onto the clypeus; the male
is further distinguished from L. planifrons, the only other
flat-faced species, by possessing a labial tubercle.

DESCRIPTION

FEMALE. Measurements (mm). Head width 3.2, head
length 2.7, wing length 7.5, total length 12.

Head. About 1.2 times broader than long; eyes moderately
convergent below, upper interorbital distance about 1 .2 times
lower; malar space linear, without posterior pit. Transverse
basal ridge of labrum low, broadly interrupted in middle.
Apical margin of clypeus not thickened. Supraclypeal area
flat, about 1 .4 times broader than long. Clypeus polished
between subcontiguous to dense punctures; supraclypeal area
broadly impunctate and polished in middle, elsewhere sub-
contiguously punctate, interspaces polished. Interocellar dis-
tance slightly less than ocelloccipital distance or ocellocular
distance. First flagellar segment shorter than pedicel or sec-
ond flagellar segment; median flagellar segments distinctly
broader than long.

Thorax. Anterior half of mesoscutal dorsum sharply sca-
brous; posterior half of mesoscutum and all of scutellum
rugosopunctate.

Color. Blackish brown; legs medium brown; tegula and
underside of flagellum ferruginous.

Pilosity. Mostly whitish; reddish hairs along apical margin
of clypeus, lower margin of mandible, and inner side of basi-
tarsi; first five gastric terga with distinct fasciae of dense,
appressed white hairs; posterior margin of pronotum with

fascia of white, plumose hairs; discs of terga two through four
with sparse, very short, fuscous hairs; fifth segment with
conspicuously longer, denser dark hairs; apical segment
densely covered with long, dark brown hairs; scopa whitish;
all sterna, except last, with conspicuous apical fascia of dense,
appressed, white hairs, last segment with brown hairs.

MALE. Measurements (mm). Head width 2.5 to 3.2, head
length 2.0 to 2.6, wing length 5.7 to 7.5, total length 8.3 to
10.5.

Head. Head 1 .2 to 1.3 times broader than long; eyes mod-
erately convergent below, upper interorbital distance about
1.2 times lower interorbital distance. Labrum with subbasal,
median, erect tubercle and with an obsolete ridge extending
from base of tubercle to each basal corner. Supraclypeal area
flat, about 1 .4 times broader than long. Clypeus weakly shiny
between continguous to subcontiguous punctures. Supra-
clypeal area polished and nearly impunctate in middle, with
subcontiguous punctures laterad that are coarser than those
of clypeus. Interocellar distance a little greater than either
ocelloccipital distance or ocellocular distance. First flagellar
segment about as long as pedicel, a little shorter than second
flagellar segment.

Thorax. Mesoscutum and scutellum with distinct, contig-
uous punctures.

Color. As described for female.

Pilosity. Similar to that of female, but hairs at apex of
clypeus whitish, dark hairs of discs of gastric terga longer;
fasciae of second and third terga interrupted in middle; apical
fasciae of sterna longer than in female; apical sternite with
pale hairs.

TYPE MATERIAL

All type material is from San Bernardino County, California.
Holotype female: Cedar Canyon, 4650 ft. elev. (Sec. 36, T13N,
R14E), 18 June 1980 (T. Griswold), on Eriogonum fasci-
culatum ssp. polifolium. Allotype: same locality, 13 June
1980 (T. Griswold). Paratypes: 2 22, 5 33, same locality, 18
and 30 June 1980 (T. Griswold); 1 9, Fenner Valley, 3670
ft. elev. (Sec. 29, T11N, R15E). 17 June 1980 (T. Griswold),
on Chilopsis linearis ; 1 3, Colton Hills, 3500 ft. elev. (Sec.
12, T10N, R14E), 17June 1 980 (T. Griswold), on Ferocactus
acanthodes ; 1 2, 7 33, Wild Horse Canyon, 4300 ft. elev. (Sec.
18, T1 IN, R15E), 6 June 1980 (T. Griswold), on Opuntia
echinocarpa ; 1 3, same locality, 27 May 1980 (T. Griswold);

1 2, Van Winkle Mts., 10 June 1965 (G. E. Wallace); 1 1 33,
9.5 mi. NNEofCima, 4200-4265 ft. elev. (Morningstar Mine
Road), 1-4 June 1977 (R. R. Snelling and C. D. George), on
Encelia californica ; 1 3, 10 mi. NE of Earp, 31 March 1959
(A. S. Menke and L. A. Stange). Holotype and allotype in
LACM; paratypes in LACM, UCR. and GRIS.

ADDITIONAL SPECIMENS

CALIFORNIA, Riverside Co.: 1 2, 30 May 1963, on Opuntia
ramosissima\ 2 33, 3 May 1963, on Echinocactus sp.; 1 3, 26

8 Contributions in Science, Number 343

Snelling: North American Lithurge

April 1963, on Cereus sp.; all from Deep Canyon (E. I. Schlin-
ger; UCR).

ETYMOLOGY

From Greek, listrotos (leveled), in reference to the flat su-
praclypeal area of both sexes.

BIONOMICS

Little is known of this species. Floral records within the
Cactaceae include the genera Cereus, Echinocactus, Fero-
cactus, and Opuntia but are so few that no clear preference
can be established. The few females available all bear pollen
from Cactaceae. Noncactus floral visits are known for both
sexes and are presumed nectar sources.

In the Mojave Desert, from which most specimens were
collected, the elevational range is from 3500 to 4650 feet
(approximately 1067 to 1417 meters). Many specimens do
not bear elevation data, but most, if not all, seem to fall
within this range. Most specimens have been collected in
June, but the earliest record is 3 1 March. Within the Mojave
Desert, captures range between 27 May and 30 June. Earlier
records are all from the margins of the Colorado Desert (Deep
Canyon and 10 miles northeast of Earp).

DISCUSSION

Both sexes of L. listrota may be readily separated from all
known species of Lithurgopsis by the flattened supraclypeal
area with a shiny, impunctate median line. Since the females
of other species all possess some conspicuous modification
of the supraclypeal area, the lack of such in the case of the
L. listrota female is especially obvious.

Males of L. listrota differ less conspicuously from those of
the other species, since males of none have prominent ele-
vations on the supraclypeal area. But, in these other species,
the supraclypeal area is weakly depressed along the midline
and weakly elevated toward the subantennal suture on either
side. This contour is particularly evident in such species as
L. apicalis. Only in L. listrota and L. planifrons is the su-
praclypeal area completely flat, even though it is slightly
elevated above the adjacent areas of the face; males of L.
planifrons have the median line coarsely and contiguously
punctate and lack an erect labial tubercle.

Of the known North American species, only one, L. api-
calis, is sympatric with L. listrota. Males of L. apicalis tend
to be larger than those of L. listrota and possess a relatively
strongly elevated supraclypeal area. In addition, the labrum
does not bear a median tubercle but has instead a low sub-
basal ridge.

It is possible that the ranges of L. listrota and L. echinocacti
may overlap in southern California or in Baja California.
The male of L. echinocacti lacks a median tubercle on the
labrum, and the legs are often ferruginous.

Geographically, the nearest species with a labial tubercle
is L. socorroensis in southern Arizona. In this species, there
is a distinct ridge extending laterad from the median tubercle
to the margin of the labrum near midlength. In L. listrota

males, the ridge extends obliquely to the anterolateral corner
of the labrum. In addition, the clypeus of L. socorroensis
males is coarsely punctate, with the punctures discrete, es-
pecially in the middle. In contrast, the clypeus of L. listrota
is finely and contiguously punctate in most areas, with the
punctures often distorted.

Lithurge ( Lithurgopsis ) littoralis (Cockerell)

Figures 3, 4

Lithurgus apicalis littoralis Cockerell, 1917:191. 3.
Lithurgus bruesi Mitchell, 1927:104. <3. NEW SYNONY-
MY.

RANGE

Illinois to New Mexico, south to the State of Oaxaca, Mexico.

BIONOMICS

Parks ( 1930) observed the behavior of this bee (as L. bruesi)
in Texas. He noted that females visited Opuntia lindheimeri,
O. ellisiana, and Echinocactus setispinosus for pollen. Nectar
sources for both sexes included Monarda punctata, M. ci-
triodora, and Vitex negundo incisa. Previously unpublished
floral records include Acacia sp., Callirhoe involucrata, Col-
ubrina texensis, Opuntia macrorhiza, and “composite.”

DISCUSSION

The type of L. littoralis, from Port Isabel, Texas, is in the
British Museum (Natural History) and was made available
by G. R. Else. This type has been compared with specimens
of L. bruesi ; there is no doubt that the two are conspecific.
Mitchell ( 1 938) first suggested that this might be the case but
had not seen the type of L. littoralis.

TYPE MATERIAL

Lithurgus apicalis littoralis Cockerell: male, in British Mu-
seum (Natural History). Lithurgus bruesi Mitchell: male, in
Museum of Comparative Zoology.

SPECIMENS EXAMINED (86 2$, 87 33)

UNITED STATES. ILLINOIS, Jersey Co.: Principia Col-
lege, Elsah (UKAN). KANSAS, Bourbon Co.: Fort Scott
(UKAN). Johnson Co.: Olathe (UKAN). NEW MEXICO,
Lincoln Co.: Nogal (UKAN). OKLAHOMA, Cimarron Co.:
14-18 mi. N of Boise City (UTSU). TEXAS, Aransas Co.:

1 0 mi. N of Rockport (UKAN, UNEB). Bexar Co.: (AMNH.
LACM, TAMU, UKAN, UTSU); San Antonio (TAMU); Ft.
Sam Houston (UCB). Brewster Co.: Big Bend Natl. Park (The
Basin; Window Trail; AMNH, TAMU, UCB, UKAN); 12
mi. NW of Alpine (UKAN). Cameron Co.: 18 mi. N of
Harlingen (UKAN); McAllen (AMNH); Port Isabel (BMNH).
Duval Co.: 8 mi. W of Fremont (TAI). Goliad Co.: (TAMU).
Hidalgo Co.: Edinburg (AMNH). Howard Co.: Big Spring
(USNM). Jeff Davis Co.: Davis Mts. (AMNH, UKAN). Jim
Wells Co.: (TAMU). Kenedy Co.: 3 mi. S of Sarita (TAI).

Contributions in Science, Number 343

Snelling: North American Lithurge 9

Kerr Co.: Kerrville (UKAN). Kleberg Co.: Kingsville and
vicinity (TAI, UKAN); Bishop (UKAN). La Salle Co.: Co-
tulla (USNM). Maverick Co.: Quemado (UKAN). Presidio
Co.: 10 mi. SE of Presidio (TAMU). San Patricio Co.:
(UKAN). Sutton Co.: Sonora (UKAN); 45 mi. S of Sonora
(UKAN). Tom Green Co.: (UKAN). Travis Co.: Austin
(TAMU). Uvalde Co.: Garner State Park (TAMU); Sabinal
(USNM); Uvalde (UKAN, UNEB). Val Verde Co.: Del Rio
(AMNH); 1 9 mi. S of Langtry (LACM). Victoria Co.: Victoria
(USNM). Webb Co.: 22 mi. S of Laredo (UKAN). Zavalla
Co.: Nueces River (USNM). County unknown: “Lopeno”
(UKAN). MEXICO. COAHUILA: 15 mi. N of Saltillo
(AMNH). HIDALGO: Pachuca (UTSU). NUEVO LEON: 4
mi. W of El Cercado (UCB, UKAN). OAXACA: Matatlan
(UCB). PUEBLA: 10.5 mi. E of Azumbilla (TAMU).

Litluirge ( Lithurgopsis ) planifrons (Friese)

Litburgus planifrons Friese, 1908:62. <3.

RANGE
Central Mexico.

DISCUSSION

No authentic material of L. planifrons has been available for
study; the species is based on a unique male, and the original
description is broad enough that it can be applied to almost
any North American Lithurge male. Were it not for the de-
scriptive specific epithet chosen by Friese, this would remain
a species of questionable identity.

Three males from Chamela, Jalisco, Mexico, collected by
S. H. Bullock, are, I believe, representatives of this species.
More important than that they agree with the original de-
scription is that the face is flat. The supraclypeal area is not
at all elevated above the level of the clypeus and does not,
in its upper portion, slope toward the antennal sockets. The
only other species with a flat supraclypeal area is L. listrota ;
males of L. listrota possess a prominent erect tubercle on the
labrum, lacking in L. planifrons. In addition, males of L.
listrota have a conspicuous median impunctate line on the
supraclypeal area.

Since the male of L. planifrons lacks an erect labral tu-
bercle, it resembles males of both L. apica/is and L. ecbino-
cacti. From both of these species, L. planifrons is separable
by the flat supraclypeal area, which does not have a median
impunctate line. Instead, the supraclypeal area is conspicu-
ously more coarsely punctate in the middle than on either
side; these coarse punctures are subcontiguous, and a similar
central line of coarse punctures is present on the clypeus.
The supraclypeal area of L. apica/is and L. echinocacti is
distinctly convex between the clypeal base and the antennal
sockets (often depressed in the center in L. apica/is ), and the
middle is distinctly shiny and impunctate (sometimes the
entire supraclypeal area sparsely punctate in L. apicalis).

The type locality for L. planifrons was originally cited as

“Jacubaya, Mexico.” This is a misprint for Tacabaya, now
a suburb of Mexico City.

Lithurge ( Lithurgopsis ) socorroensis (Mitchell)

Figures 7, 8

Litburgus socorroensis Mitchell, 1938:152-154. 2 <3.
RANGE

Southern Arizona to Costa Rica; Revillagigedo Archipelago.

BIONOMICS

Little is known of this species, not previously recorded from
the mainland. Both sexes have been taken at flowers of Opun-
tia sp. on Socorro Island, the type locality. Males were taken
in Arizona on “thistle.” In Costa Rica, both sexes have been
collected on Opuntia sp. and males were on Cydista heter-
ophylla. Paullinia costaricensis, Pterocarpus rohrii, Securi-
daca sylvestris, and “pink bignon vine.”

DISCUSSION

Females of this species are similar to those of L. echinocacti
in the structure of the supraclypeal area. In L. socorroensis,
the legs are brown, rather than red, and there is no pubescent
fascia on the dorsal margin of the pronotum. The male can
be recognized by the structure of the labrum, as noted in the
key.

TYPE MATERIAL

Litburgus socorroensis Mitchell: female holotype and allo-
type in California Academy of Sciences.

SPECIMENS EXAMINED (18 22, 24 3<5)

UNITED STATES. ARIZONA, Pima Co.: Santa Catalina
Mts. (UNAR); Rincon Mts. (Madrona Ranger Station;
UNAR); Baboquivari Mts. (UCR); Continental (UTSU).
MEXICO. JALISCO: Guadalajara (UTSU). MORELOS:
Cuernavaca (UKAN). PUEBLA: Petlalcingo and vicinity
(UCD, UTSU). ISLAS REVILLAGIGEDO: Socorro Island
(CAS, LACM, UTSU). SINALOA: 50 mi. NW of Mazatlan
(UTSU). VERA CRUZ: 8 mi. SW of Vera Cruz (UTSU).
EL SALVADOR. LA LIBERTAD: 5 km. NW of Quezaltepec
(Hacienda Capolinas; UCR). COSTA RICA. GUANACAS-
TE: Hacienda Comelco (LACM, UCB).

Lithurge ( Lithurge ) chrysurus (Fonscolombe)

Litburgus chrysurus Fonscolombe, 1834:220. 2 <3.

RANGE

Countries bordering western Mediterranean Sea; adventive
in eastern United States.

BIONOMICS

According to Roberts (1978), females construct communal
nests in both soft and hard woods, causing serious damage.

10 Contributions in Science, Number 343

Snelling: North American Lithurge

Cells are usually separated by wood-dust partitions and are
packed with a pollen and nectar mass gathered from Cen-
taurea maculosa (Compositae), an introduced weedy plant.
When partitions are absent, two or three regularly spaced
larvae feed on the single large pollen mass. When larval
feeding is completed, the mature larva defecates and spins a
pupal cocoon. Immatures overwinter as prepupae, and pu-
pation occurs during the following spring.

DISCUSSION

Friese (1911) gives a complete synonymy for L. chrysurus.
Roberts (1978) reported this Palaearctic species from New
Jersey, where it had been established for at least three years;
nesting biology, behavior, and immature morphology were
all described by Roberts. In North America, this bee is pres-
ently known only in New Jersey, where it is oligolectic on
Centaurea maculosa, a plant also native to countries bor-
dering the western Mediterranean Sea.

ACKNOWLEDGMENTS

For the loan of specimens utilized in this study, I am indebted
to; P. H. Arnaud, Jr. (CAS), G. R. Else (BMNH), D. K.
Faulkner (SDM), M. Favreau (AMNH), S. Frommcr (UCR),
J. E. Gillaspy (TAI), T. Griswold and F. D. Parker (GRIS,
UTSU), F. F. Hasbrouck (ARSU), the late P. D. Hurd, Jr.
(USNM), L. S. Kimsey and R. O. Schuster (UCD), U. N.
Lanham (UCOL), E. G. Linsley (UCB), S. J. Merrit (TAMU),
C. D. Michener (UKAN), B. Ratcliffe (UNEB), M. S. Was-
bauer (CDFA), and F. G. Werner (UNAR).

I am especially grateful to G. R. Else for his assistance in
locating, and making available, the type of L. apica/is litto-
ralis.

LITERATURE CITED

Berthold, A. A. 1827. Latreille’s natiirliche Familien des
Theirreiche. Weimar, 610 pp.

Brach, V. 1979. Notes on the biology of Lithwgus gibbosus
Smith in Florida (Hymenoptera: Megachilidae). Bull. So.
Calif. Acad. Sci. 77:144-147.

Cockerell, T.D. A. 1898. New and little-known Hymenop-
tera taken by Prof. C.H.T. Townsend and Mr. C.M.
Barber in New Mexico in 1898. Ann. Mag. Nat. Hist.
(7) 2:448-457.

. 1902. The genus Lilhurgopsis. Entomol. News 13:

182.

. 1911. Bees in the collection of the United States

National Museum. 2. Proc. U.S. Natl. Mus. 40:24 1-264.
. 1917. In W.P. Cockerell, Collecting bees in south-
ern Texas. Jour. N.Y. Entomol. Soc. 25:187-193.

. 1937. Bees collected by Mr. and Mrs. J.L. Sperry

and Mr. R.H. Andrews in Arizona. Bull. So. Calif. Acad.
Sci. 36:107-1 10.

Cresson, E.T. 1875. Report upon the collections of Hy-
menoptera, In Wheeler, Rept. Geogr. Geol. Explor. Surv.
West of 100th Meridian, vol. 5, pp. 707-728.

Ducke, A. 1 907. Contribution a la connaissance de la faune
hymenopterologique du nord-est du Bresil. Rev. Ento-
mol. 26:73-96.

Fonscolombe, B. 1834. Notices sur les genres d’Hyme-
nopteres Lithwgus et Philoxera. Ann. Soc. Entomol.
France 3:219-224.

Fox, W.J. 1902. Lilhurgopsis, a new genus of bees. Ento-
mol. News 13:137-140.

Friese, H. 1908. Die Apidae (Blumenwespen) von Argen-
tina nach den Reisenergebnissen der Herren A.C. Jen-
sen-Haarup und P. Jorgensen in den Jahren 1 904-1907.
Deutsch. Entomol. Zeits. 1908:1-94.

. 1911. Apidae 1. Megachilinae. Das Tierreich 28:

1-440.

Hurd, P.D., Jr. 1979. In K.V. Krombein et al.. Catalog of
Hymenoptera of America north of Mexico. Washington,
D.C. 2:1 199-2209.

Latreille, P.A. 1825. Families naturelles du regne animal.
Paris, 570 pp.

Michener, C.D. 1944. Comparative external morphology,
phylogeny and a classification of the bees (Hymenop-
tera). Bull. Amer. Mus. Nat. Hist. 82:151-326.

. 1965. A classification of the bees of the Australian

and South Pacific regions. Bull. Amer. Mus. Nat. Hist.
130:1-362.

Mitchell, T.B. 1927. New megachilid bees. Psyche 34: 104-
121.

. 1938. The genus Lithwgus (Hymenoptera: Mega-
chilidae). Psyche 45:146-155.

. 1962. Bees of the eastern United States. N. Car.

Agric. Exp. Sta. Tech. Bull. 152:1-557.

Moure, J.S. 1949. Las especies Chilenas de la sub-famiiia
Lithurginae (Hym.-Apoidea). Arq. Mus. Paranaense 7:
265-286.

Parker, F.D., and H.W. Potter. 1973. Biological notes on
Lithurgus apica/is Cresson (Hymenoptera: Megachili-
dae). Pan-Pacific Entomol. 49:294-299.

Parks, H.B. 1 930. Notes on Texas bees. Bull. Brookl. Ento-
mol. Soc. 25:263-264.

Roberts, R.B. 1978. The nesting biology, behavior and im-
mature stages of Litharge chrysurus, an adventitious
wood-bonng bee in New Jersey (Hymenoptera: Mega-
gachilidae). Jour. Kans. Entomol. Soc. 51:735-745.

Rozen, J.G., Jr. 1973. Immature stages of lithurgine bees
with descriptions of the Megachilidae and Fideliidae
based on mature larvae (Hymenoptera, Apoidea). Amer.
Mus. Novitates 2527, 14 pp.

Smith, F. 1853. Catalogue of hymenopterous insects in the
collection of the British Museum. Part 1, 197 pp. British
Museum, London.

Submitted 20 May 1982; accepted 17 December 1982.

Contributions in Science, Number 343

Snelling: North American Lithurge 11

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Primed a? Ail Sea Pre

ii- kit.i.’ m i

IB

, e: .

. lac., Lawrence, Kansas

MHi-

as

Li ipft
lllkg

iMIF

THE TERTIARY SPONGES APHROCALLISTES
AND EURETE FROM

WESTERN WASHINGTON AND OREGON
J. Keith Rigby1 and David E. Jenkins2

ABSTRACT. Extensive collections of sponges from the Eocene to
Miocene of southwestern Washington and from the Oligocene and
Miocene of northwestern Oregon consist mainly of Aphrocallistes
polytretos n. sp. This new sponge has irregular undulating to tubular
walls that look like liny-celled honeycombs because of their coarse
canals. The upper cellular part of the skeleton is made of flattened
hexiradiates, but the basal part is irregularly dictyonine.

The less common Eurete goedertin. sp., a branching zigzag tubular
sponge, occurs in the Oligocene part of the Lincoln Creek Formation
in southwestern Washington. Root tufts of large monaxial spicules
occur in concretions with both Eurete and Aphrocallistes but may
not be part of either of these sponges.

Distributions of living species of both genera suggest by analogy
that the fossils lived at a depth of 300-350 meters. Most occur in
coarse clastic units or in calcareous concretions with a mixed diverse
fish-echinoid-sponge assemblage. This fauna may have been swept
into still deeper water by pulses of high-energy currents, for most
sponges and echinoids appear fragmented. Only the root tufts appear
to have grown in place.

INTRODUCTION

Numerous fossil sponges have been recovered from concre-
tions eroded out of relatively deep-water marine Eocene to
Oligocene sediments near Knappton (Localities 2, 5-7, Fig-
ure 1), from Eocene limestone in a quarry near Chinook
(Locality 1, Figure 1) in southwestern Washington, from
Miocene concretionary beds at Astoria, and from Oligocene
deposits near Yaquina (Locality 4, Figure 1) in northwestern
Oregon. Near Knappton, the major locality, the sponges are
apparently scattered through the formation but are most
common in a zone where sponge-bearing concretions are
particularly abundant. The major sponge fauna occurs pre-
sumably below beds that have produced the mollusks being
studied by Ellen James Moore, of the U.S. Geological Survey,
and above beds that have produced the silicified micro-
mollusks (bivalves and gastropods) being studied by Carole
Hickman, of the University of California Museum of Pa-
leontology. Fossil barnacles have been described by Victor
Zullo (1982) from localities both above and below the major
sponge zone.

The fossils come, in large part, from concretions weath-
ering out of a Holocene landslide block in the Lincoln Creek
Formation and occur as float along the Columbia River ter-
race. The slump block is in the sea cliff at the head of the
bay between Knappton and Grays Point. If, as Moore (per-
sonal communication, December, 1981) suggests, “there is
a certain amount of stratigraphy preserved in the float from
the landslide block,” the sponges then are stratigraphically
just below the main mollusk-bearing beds. “Because the
landslide moved generally parallel with the strike of the beds,
a proper stratigraphic sequence seems to have been main-
tained,” according to Moore (personal communication, Au-
gust, 1982).

The designation Lincoln Formation was initially proposed
by Weaver (1912:10-12) for Oligocene beds of western
Washington. Weaver! 1937) later suggested retaining the name
Lincoln Formation for the entire middle Oligocene of Wash-
ington. Weaver and others ( 1 944:592) noted the Lincoln beds
overlie the Keasey Formation and unconformablv underlie
the Miocene Astoria Formation. The Lincoln Creek For-
mation in the Knappton area occurs in an east-dipping homo-
cline, west of outcrops of the Astoria Formation, in the
Portuguese Point-Grays Point area west of Frankfurt. Wash-
ington (Wells, 1979).

The name Lincoln Creek Formation was proposed by Beik-
man and others (1967) to replace the Lincoln Formation,
which has been preempted several times. In the type area,
the formation consists of 700-3000 m (2000-9000 feet) of
clastic rocks of predominantly Oligocene age. Lincoln Creek
beds unconformably overlap Eocene and older units near
Oligocene paleotopographic high areas but rest conformably
on Eocene beds within the basins. The Lincoln Creek For-
mation is overlain by the Astoria (?) Formation in the type
region. Molluscan and foraminiferal correlations suggest that
the Lincoln Creek Formation ranges from the Late Eocene

1. Brigham Young University, Provo, Utah 84602.

2. Union Oil Company of California, 2323 Knoll Drive, Ventura,
California 93003.

Contributions in Science, Number 344, pp. 1-13
Natural History Museum of Los Angeles County, 1983

ISSN 0459-8113

Figure 1. Index maps to sponge localities in southwestern Washington and northwestern Oregon. Arrows indicate collecting localities. A,
Ona Beach locality (4, LACMIP 6148) south of Newport and north of Seal Rocks on the coast, Yaquina 15-minute quadrangle, Lincoln
County, Oregon. B, Bear River locality where calcified sponges and mollusks occur in “reefoidaP' limestone (1, LACMIP 5802). Chinook IVi-
minute quadrangle. Pacific County. Washington. C, Knappton localities in the Lincoln Creek Formation where it is exposed in the sea cliff,
(8, LACMIP 5787, Oligocene; 2, LACMIP 5842, Upper Oligocene; 5, LACMIP 5843, Lower Oligocene; 6, LACMIP 5844, Eocene; 7, LACMIP
5852, Oligocene) on the north shore of the mouth of the Columbia River, Knappton 71/2-minute quadrangle. Pacific County, Washington.

2 Contributions in Science, Number 344

Rigby and Jenkins: Tertiary Sponges

Foraminiferal

Molluscan

WASHINGTON

OREGON

\

Stages

Stages

Knappton

Area

Bear River Area

Seal Rocks Area

c

<D

O

o

Saucesian

Ne wportian

Astoria (?) Formation

Astoria (?) Formation

Astoria (?) Formation

Piilarian

7

?

Nye Mudstone

Unit 4

Juanian

c 5842

o

Yaquina Formation

Oligocene

ro Unit 3

5852

o

6148

Zemorrian

Matlockian

ll

(D

Q)

- Unit 2

U 5843

c

o

o

c

_l

00

in

Lincoln Creek Formation

Alsea Formation

o

© £

Unit 1

5844

a. o

Refugian

Galvinian

Q. o
=> LU

?

?

Siltstone of Cliff Point
5802

Figure 2. Stratigraphic distribution of localities in the Knappton, Bear River, and Yaquina (Seal Rocks) areas showing relationships of the
informal units of James and Gail Goedert. Natural History Museum of Los Angeles County localities, and biostratigraphic zones of the Tertiary
section (modified from Zullo, 1982; Wells, 1979; Snavely and others, 1975; Armentrout, 1981; Rau, 1981).

to latest Oligocene (Rau, 1958, 1964, 1981; Armentrout,
1975. 1977, 1981).

James L. Goedert (Zullo, 1982:2-3) has subdivided the
Lincoln Creek beds in the Knappton outcrops into four in-
formal units (Figure 2). Unit 1 is a barnacle-bearing concre-
tionary unit associated with the abundant Ll-shaped trace
fossil, Tisoa. Unit 2 is characterized by sponges, small aturiid
nautiloids, many decapod crustaceans, and marine vertebrate
remains associated with tisoid burrows. Unit 3 is character-
ized by a glass sponge fauna and Unit 4 by an abundance of
marine vertebrate and invertebrate fossils, but few sponges.
Zullo ( 1 982:2-3, fig. 2) concluded that lower Unit 1 is Upper
Eocene, Unit 2 is Lower Oligocene, and LJnits 3 and 4 are
Upper Oligocene (Figure 2).

The sponges noted here from Oregon are reportedly from
the Astoria Formation at the type locality in Astoria and
from the Yaquina Formation near Seal Rocks. Moore ( 1 963)
described outcrops and faunas of Astoria beds in the type
area and southward along the Oregon coast for approximately
70 miles. One specimen of Aphrocallistes has been figured
from the type area of the Astoria Formation by Moore ( 1 963,
pi. 32, fig. 16), but the precise locality from which the early
collection was made is uncertain. No additional sponges have
been reported from other exposures along the sea cliff be-
tween Astoria and Newport. However, a few sponges have

been recovered from the Yaquina Formation south of New-
port, in the Seal Rocks area (Figure 1).

Moore (1963: 1 2) described Astoria beds in the coastal area
as mainly marine silty shale and fine-grained sandstone, with
mterbedded siltstone, coarse-grained sandstone, conglom-
erate, and tuff. She reported that concretions up to a foot
and a half in diameter are common in some places. Such
concretions have produced nearly all the sponges from the
Astoria, Yaquina, and Lincoln Creek localities. Some con-
cretions contain a varied molluscan fauna, fragments of car-
bonized wood, many with teredinid borings, marine mam-
mal bones, fish bones and scales, a varied foraminiferal fauna,
decapod crustaceans, and barnacle fragments. Many concre-
tions are coarse-grained glauconitic sandstone that shows
evidence of vigorous transport. These assemblages may have
been mixed, that is may have been derived from a variety
of depositional sites, by downslope movement.

The sponges, in general, are delicate and silicified and can
be freed from the calcareous matrix. Some concretions were
partially etched in dilute acetic acid and screened for recovery
of all microorganisms. In general, the concretions are highly
argillaceous, often sandy and glauconitic. For complete re-
moval of matrix, the partially etched calcareous muds were
gently brushed and washed from the sponge material. Some
fine cleaning of specimens was done with a microsandblaster

Contributions in Science, Number 344

Rigby and Jenkins: Tertiary Sponges 3

air-abrasive unit. Some delicate etched specimens were hard-
ened in weak acetone-soluble plastic solutions.

LOCALITIES

The fossil sponges described here were all collected by James
L. and Gail H. Goedert, except for the sponge reported by
Ellen J. Moore (1963:89, pi. 32, fig. 16), and are deposited
in the Natural History Museum of Los Angeles County, In-
vertebrate Paleontology Section (abbreviated LACMIP).

LOCALITY 1. Limestone in the siltstone of Cliff Point of
Wells (1979) in a quarry in the bluff on the south side of
Bear River, 2.2 km (1.4 miles) northeast of Goulter Ranch,
on the section line between Sections 20 and 21, T. 10 N., R.

1 0 W., on the Chinook VA-minute quadrangle. Pacific Coun-
ty, Washington. LACMIP locality 5802, Upper Eocene.

LOCALITY 2. Lincoln Creek Formation in association
with other invertebrate fossils, mammal and fish bones, and
plant fragments in calcareous, argillaceous concretions that
range from fine-grained to coarsely clastic, from a landslide
area at the head of the bay between Knappton and Grays
Point in the center N 'A N ‘A of Sec. 9, T. 9 N., R. 9 W.. on
the Knappton 7‘A-minute quadrangle in Pacific County,
Washington. The locality is approximately 1.6 km (1 mile)
northeast of Knappton. LACMIP Locality 5842, Upper Oli-
gocene.

LOCALITY 3. Astoria Formation at Astoria, as cited by
Moore ( 1 963:90) at her locality la, from collections of James
D. Dana discussed by Dali (1909:140), Miocene.

LOCALITY 4. Sponges from the Yaquina Formation, in
fossiliferous concretions from rocks exposed at Ona Beach,
0.4 km (0.25 mile) south of the mouth of Beaver Creek, west
of U.S. Highway 101, 2 km (1.3 miles) north of the com-
munity of Seal Rocks, just west of the northwest corner of
Sec. 1 9, T. 12 S., R. 1 1 W., Lincoln County, Oregon, Yaquina
15-minute quadrangle. LACMIP Locality 6148, Miocene.

LOCALITY 5. Lower Oligocene part of the Lincoln Creek
Formation, from concretions weathering out of the sea cliff
in the northwestern part of the bay between Grays Point and
Knappton. approximately 305 m (1000 feet) south and 430
m (1400 feet) east of the northwest corner of Sec. 9, T. 9 N.,
R. 9 W., on the Knappton 7‘A-minute quadrangle, in Pacific
County. Washington. The locality is approximately 300 m
( 1 000 feet) west of Locality 2, and 300 m ( 1 000 feet) northeast
of Locality 7. LACMIP Locality 5843, Lower Oligocene.

LOCALITY 6. LIpper Eocene part of the Lincoln Creek

Formation, concretions weathering out of the sea cliff in the
western part of the bay between Grays Point and Knappton,
approximately 0.8 km (0.5 mile) northeast of Knappton; 122
m (400 feet) east and 520 m ( 1 700 feet) south of the northwest
corner of Sec. 9, T. 9 N.. R. 9 W., on the Knappton 7‘A-
minute quadrangle. Pacific County, Washington. LACMIP
Locality 5844. Upper Eocene.

LOCALITY 7. Oligocene part of the Lincoln Creek For-
mation, collected from the “sponge zone” in concretions
weathering out of the sea cliff, approximately 1.3 km (0.8
mile) northeast of Knappton, 305 m (1000 feet) south and
580 m (1900 feet) east of the northwest corner of Sec. 9, T.
9 N., R. 9 W.. Knappton 7‘A-minute quadrangle. Pacific
County, Washington. This is the most productive sponge
locality of those cited here. LACMIP Locality 5852, Oligo-
cene.

LOCALITY 8. Lincoln Creek Formation on the Columbia
River, approximately 122 m (400 feet) east of the boundary
between Sections 8 and 9, T. 9 N., R. 9 W., approximately
0.8 km (0.5 mile) northeast of Knappton. Knappton 7‘A-
minute quadrangle. Pacific County, Washington. A general
Knappton locality of older collections before units were es-
tablished. LACMIP Locality 5787, Oligocene.

SYSTEMATICS

Class Hexactinellida Schmidt, 1870
Order Hexactinosa Schrammen, 1903
Suborder Scopularia F. E. Schulze, 1885
Family Aphrocallistidae Gray, 1867
Genus Aphrocallistes Gray, 1858

Aphrocallistes polytretos n. sp.

Figures 3-10, 13-17, 21

DIAGNOSIS. Branching tubular to cuplike or irregular
undulating leaf- or ear-shaped sponges with upper walls per-
forated by honeycomblike, subprismatic to polygonal pri-
mary canals in full diarhysis and open on both sides of the
wall. Basal attachment and lower part of walls irregularly
dictyonme with fewer canals. Each canal of upper wall out-
lined by a complex net of siliceous skeletal strands produced
by hexiradiate beams whose rays have been distorted to be
more or less in the plane of the porous canal walls. No sculp-

Figures 3 through 10. Aphrocallistes polytretos n. sp. (3) Basal view of two specimens showing dense irregular attachment areas grading up
into distal radially canalled area. Paratype, LACMIP 6601, Locality 2, XI. (4) Natural vertical section through an irregularly dictyonine base
and associated lower canalled area. Definition of canals increases away from base, along with increase in modification of spicule structure
toward hexiradiates. Paratype, LACMIP 6602, Locality 8, X2. (5) Side view of well-preserved paratype with honeycomblike wall and tubular
to cuplike form but with broken upper end. Base of this same specimen is shown in upper right of Figure 3. LACMIP 6601, Locality 2, X2.
(6) Fragments of sponge showing coarse conglomeratic and glauconitic nature of common matrix. LACMIP 6603, Locality 7, XL (7) Side
view of broad flattened, moderately fine-textured paratype. LACMIP 6604, Locality 7, XL (8) Side view of holotype showing annular tubular
form and honeycombed walls. Reverse side has concave-up. dictyonine-spiculed diaphragmhke interruptions that probably marked pulses in
growth. LACMIP 6600, Locality 2, XL (9, 10) Stereoscopic photomicrographs of skeletal net of the holotype showing canal patterns, shape,
and hexiradiate skeleton typical of the middle and distal part of the sponge. LACMIP 6600. Locality 2, X5.

4 Contributions in Science, Number 344

Rigby and Jenkins: Tertiary Sponges

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Rigby and Jenkins: Tertiary Sponges 5

ture is evident, except for the undulating surfaces. Polygonal
canals are generally 1.0- 1.5 mm across in a wall 2-5 mm
thick. Upper part of skeleton of irregular flattened hexiradiate
dictyonal beams with largest, earliest formed beams essen-
tially horizontal but connected with irregularly diagonal and
vertical strands in a cross-bracing crudely hexiradiate pat-
tern, like an expanded metal screen, with roughly hexagonal
openings often subdivided into triangular sections. All beams
fused into a solid network; each is minutely spinose.

Canals much larger and less oblique, and skeleton in ma-
ture parts much more hexiradiate-dominated and much less
clearly dictyonine, than in Aphrocallistes vastus whiteavesi-
anus Lambe and A. vastus vastus Schulze. Canals much
larger but more closely spaced and more commonly prismatic
than in A. alveolites Roemerand A. cylindrodactylus Schram-
men.

DESCRIPTION. Numerous nearly complete siliceous
fragments to broken bits of the sponge occur, often as several
pieces with different growth forms in single concretions. The
holotype is a subtubular undulating sponge with a diameter
of 30-35 mm. It is approximately 80 mm tall and is sub-
divided into three subequal sections with saucer-shaped,
noncanalled partitions with a texture like that of bases on
other specimens.

The wall expands upward from a thickness of 2. 0-2. 5 mm
in the lower part of the fragment, to 2. 5-3. 5 mm in the middle
segment, and to a maximum of 4. 0-4. 5 mm at the upper end
of the fragment. The arcuate saucer-shaped noncellular par-
titions are generally 0. 5-1.0 mm thick in the center of the
tubular opening. The lower segment is approximately 1 5 mm
high, the middle segment is 20 mm high, and the uppermost
is 20 mm high, above the uppermost cellular separation.
Other fragments show the same range in wall thickness, al-
though most are only undulating irregular platelike bits or
tube sections. Some nearly complete cup-shaped sponges show
the massive base, a few millimeters across, and a tubular
form 8-30 mm in diameter and up to 90 mm long or high.

Individual canals pierce the walls, essentially normally, in
full diarhysis. They are 1.0-1. 8 mm across on the outside
but are only 0.8-1. 5 mm across on the inside, showing ex-
pansion of the canals related to their radial orientation. They
are closely packed, although not in predictable horizontal or
vertical series, and are so spaced that 7-8 occur in 10 mm.
both horizontally and vertically on the exterior. Many of the
canals have rectangular cross sections, but with rounded mar-

gins. Hexagonal and pentagonal openings also occur, com-
monly in the middle unmodified parts of the segments. All
canals have porous walls so that there could have been com-
plete interconnection with adjacent ones. Canal walls are
perforated at irregular intervals with distinct rounded open-
ings 0.10-0.22 mm across. These are less common than the
much smaller and more irregularly sized triangular openings
that occur between the diagonal and vertical beams in the
hexiradiate-based skeletal net.

The skeleton is composed of fused solid beams that are
difficult to separate into single spicular elements. Walls are
principally outlined by horizontal fibers that tend to bifurcate
horizontally at the wall intersections and vertically in the
middle part of the wall. These are the first beams secreted
at any level in the skeleton and are the most massive units.
They are spaced 5-7 horizontal beams per mm. measured
vertically along the canal wall. Most of these strands are 0.06-
0.08 mm across, but in some areas where the skeletal net is
dense, they have been thickened to 0.10-0.12 mm in di-
ameter. These dominantly horizontal units are crossbraced
by diagonal and vertical units in the basic flattened hexira-
diate skeleton. Diagonal beams are next to horizontal ones
in size and are generally 0.03-0.06 mm across. Vertical beams
are apparently the last added and are the smallest, ranging
from tiny hairlike spines 0.01 mm across up to rods 0.04-
0.05 mm in diameter, fused at both ends. Diagonal beams
are less continuous than horizontal ones, and vertical ones
are the least continuous. However, vertical beams can be
traced completely along the canal wall, in some areas, parallel
to the trend of the canals.

Most skeletal elements are shared between two adjacent
canals in the thin wall. Elsewhere, particularly in proximal
parts of the sponge, double walls occur, with each canal hav-
ing its own subparallel fused series of elements. These strands
bifurcate to produce triangular or polygonal radial openings
along the common juncture between three or four adjacent
canals.

Beams are apparently solid and appear to have been built
by enlargement, with layered secretions over initial small
needles or spines. These needles are common in young parts
of the skeleton, pointing upward and horizontally from ad-
jacent thicker strands. Some beams are so enlarged that they
combine to produce a nearly solid fused wall in the older
parts of the net. Exchange between canals is moderately lim-
ited in these areas, but in the outer, more open-textured parts.

Figures 11 through 17. Root tufts and Aphrocallistes polytretos n. sp. Figures 11, 12. Root tufts. (11) Large concretion with aligned oxeate
root tuft spicules in fine-grained calcareous siltstone, which has undergone some bioturbation (lower right). LACMIP 6611, Locality 7, Xl.
(12) Large monaxial root tuft spicules in fine-grained siltstone showing packing, shape, and general preservation that indicate they have not
been moved after formation of the tuft. LACMIP 6612. Locality 5, X2. Figures 13 through 17. Aphrocallistes polytretos n. sp. (13) Photo-
micrograph showing irregular spicules of basal attachment and dense, noncanalled pattern, in left center of Figure 17. LACMIP 6605, Locality
5, X10. (14) Part of a concretion containing Aphrocallistes with irregular branching growth form in variously colored fine-grained matrix,
which produces blotchy appearance. Paratype, LACMIP 6606, Locality 2, XL (15) Photomicrograph of lower part of sponge showing thick
canal walls and change from dictyonine-based skeleton in lower right to hexiradiate skeleton in upper left. Side of same specimen in Figure
4. Paratype. LACMIP 6602. Locality 8, X10. (16) Photomicrograph of dictyonine part of basal skeleton of paratype illustrated in Figure 4, in
left center, but rotated 90 degrees. Somewhat swollen spicule nodes and irregular fabric are characteristic. LACMIP 6602, Locality 2, X10. (17)
Irregular basal region and part of honeycomblike tubular wall of paratype. Base shown enlarged in Figure 13. LACMIP 6605, Locality 2, X2.

6 Contributions in Science, Number 344

Rigby and Jenkins: Tertiary Sponges

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Rigby and Jenkins: Tertiary Sponges 7

interchange could have been effected, even through double
walls where two skeletal layers separate adjacent canals.

The entire surface of each spicular strand, particularly the
thickened ones, is covered by minute conical spines 0.005-
0.01 mm high and across. Some tiny elements extend as fine
needles from small conical bases. These cones and spines are
generally 0.01-0.02 mm apart and appear to be scattered
irregularly over the entire surface of the major strands.

DISCUSSION. Generic and family placement of these
sponges is based on their peculiar flattened hexactine hexi-
radiate skeleton. The sponges superficially appear like lithis-
tids, composed of orchoclad dendroclones. The “clads” are
fused to “clads" ofother spicules to produce a flattened roughly
hexagonal series of openings with one spicule axis essentially
normal to the long dimensions or axes of the canals, but
individual lithistid spicules cannot be identified.

The large canals might be considered as peculiar porous
calicles, such as in the sclerosponge Merlia or the chaetaetids.
However, in sclerosponges, the calicles rise from an imper-
vious base rather than forming perforate walls around a sub-
cylindrical or platelike porous sponge.

Schrammen ( 19 12:358), in a summary table of stratigraph-
ic ranges of Cretaceous fossils of northwestern Germany,
noted that Aphrocallistes ranges from the Late Cretaceous
into Recent. He also listed the genus from Tertiary rocks of
Russia, Japan, and Oran, but cited no references for those
occurrences. Schrammen (1912:219-222) differentiated three
species of the genus on general body shape, wall thickness,
presence or absence of terminal sieve plates, or whether the
forms are many branched or relatively simple structures. In
general, the German species have canals considerably more
widely separated and distinctly smaller than our specimens.

Schrammen (1912:185) summarized the observed depth
ranges of living species of Aphrocallistes. In general, their
range is from approximately 100-1700 m. Reid (1968:549)
cited occurrences of A. vastus as shallow as 27.5 m from
Puget Sound. He also listed other occurrences off Oregon and
Washington in water 97 and 108 m deep. Living species
described by Schulze (1887) occur essentially in the North
Pacific, from localities in the Philippine Islands, Japan, the
Aleutian Islands, and along the coast of Oregon and Wash-
ington, California, and Vancouver Island. Thus, it is not
surprising to find Aphrocallistes in Eocene to Miocene beds
in Oregon and Washington.

Aphrocallistes was collected from Astoria beds at Astoria,
Oregon, on the south bank of the Columbia River (Dana,

1849). That specimen was noted by Dali (1909:140) and
figured by Moore (1963, pi. 32, fig. 16) as an unidentified
ophiuroid (?) as an incidental occurrence to her extensive
treatment of the Miocene mollusks from the Astoria For-
mation of Oregon.

Canal patterns in living Aphrocallistes vastus whiteavesi-
anus Lambe, illustrated by Reid (1964, text-fig. 47), and
Aphrocallistes vastus vastus Schulze show markedly oblique
canals and clearly defined dictyonine-based skeletons in a
moderately rectangular arrangement. Development of hex-
iradiate patterns is more clearly shown in Aphrocallistes bea-
trix Gray, as figured by Reid (1964, text-fig. 49b). Even in
that species, however, there is a moderately oblique orien-
tation to some canals in the gastral region, and wall thick-
nesses tend to be somewhat greater than in the Tertiary species
from Washington and Oregon. Canal dimensions in each of
these are considerably smaller than in the fossil A. polytretos.
There is striking similarity of skeletal arrangements in A.
beatrix to that of the Tertiary fossils. The walls are somewhat
thicker in the Recent species, and the canals tend to be more
circular than distinctly prismatic, such as in the fossil forms.
Such rounded canals are the general pattern, as well, in the
Cretaceous sponges described by Schrammen (1912:21 9-222).
A. polytretos is similar to Aphrocallistes beatrix in having the
dictyonal strands obscure, in contrast to Aphrocallistes vastus
varieties.

Aphrocallistes beatrix Gray (1858:1 14-1 1 5) was described
from the Azores in the Atlantic Ocean. Some of the limited
modern distribution is unquestionably related to sampling
difficulty and spacing of samples in the 100-1000 m range
in the modern oceans.

Associated benthonic foraminifersand mollusks, including
the nautiloid Aturia, suggest water depths between 300 and
600 m, which is also within the range of modern Aphrocal-
listes. The assemblage may have been transported from mod-
erately shallow waters of 300-400 m and accumulated ulti-
mately in depths as great as 2000-4000 m.

Specific characteristics are the size of individual canals and
skeletal elements and the general growth form of the sponges.

Our species is fairly variable and ranges at least from Late
Eocene to Miocene. These Tertiary sponges show a general
increase in canal size, a decrease in canal wall thickness, and
a tendency for a more prismatic canal pattern, when com-
pared to the Cretaceous species from northwestern Germany.

Schrammen (191 2:220) noted a terminal or oscular sieve-
like diaphragm on branches of A. alveolites Roemer and A.

Figures 18 through 22. Eurete goederti n. sp. and Aphrocallistes polytretos n. sp. Figures 18 through 20 and 22. Holotype of Eurete goederti
n. sp. LACMIP 6609, Locality 7. (18) Photomicrograph of gastral part of dictyonine skeletal net showing marked regularity of enlarged strands
and somewhat smaller cross-connecting beams. Spicule nodes are little swollen. Figure 23 is a drawing of part of the net in the upper left. X5.
(19) Holotype of Eurete goederti and associated fragments of Aphrocallistes in bioclastic matrix of a concretion from Locality 7. The regularly
spaced zigzag pattern of the branching sponge is clearly shown. Proximal is to upper right and distal to lower left, where circular cross section
shows near shadowed boundary (arrow). XL (20) Photomicrograph of thin walls of the holotype, with diverging walls that are 3 or 4 spicule
complexes thick. Gastral surface is to the right. X5. (22) Well-preserved part of the skeleton showing thickness of the walls in lower part and
regular nature of skeleton in middle and upper part. Parts of three dichotomous branches, producing zigzag structure of the sponge, are shown.
X3. Figure 21. Aphrocallistes polytretos n. sp., paratype illustrated in Figure 1 7, here enlarged to show smaller hexactine-based spicules partially
filling canals. LACMIP 6605, Locality 2, XI 5.

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Rigby and Jenkins: Tertiary Sponges

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Rigby and Jenkins: Tertiary Sponges 9

cylindrodactylus Schrammen. Diaphragms are present in the
Washington Oligocene specimens, but in general these are
concave-up divisions within tubular parts of the sponge and
must mark pulses in growth of the sponge rather than convex-
up terminations like those on the German Cretaceous sponges.
Orientation in the Tertiary sponges is well defined by de-
velopment of irregular basal areas, just as the terminations
are well defined in Schrammen’s A. cylindrodactylus by the
rounded fingerlike growth.

TYPE SPECIMENS AND AVAILABLE MATERIAL.
Holotype LACMIP 6600 from Locality 7. the type locality
of the species, and paratypes LACMIP 6601, 6603, 6607,
6608 from Locality 7, LACMIP 6605, 6606 from Locality
5, LACMIP 6604 from Locality 7, LACMIP 6602 from Lo-
cality 8. In addition 22 specimens or concretions were ob-
tained from Locality 1. 18 from 2, 1 from 4, 1 from 5, 2
from 6, and 12 from Locality 7. Some concretions contain
several specimens or fragments.

ETYMOLOGY. Poly, Gr., many; tretos, perforated, re-
ferring to the many coarse canals through the cellular-ap-
pearing wall.

Family Euretidae Zittel, 1877
(fide Schulze, 1887)

Subfamily Euretinae Reid. 1958
Genus Eurete Semper, 1868

Eurete goederti n. sp.

Figures 18-20, 22, 23

DIAGNOSIS. Dendroid or irregular zigzag heteroto-
mously branching axial tube with short lateral branches, both
approximately 8-10 mm in diameter; branches approxi-
mately 20 mm apart. Walls 0. 5-1.1 mm thick; skeletal net
euretoid, lacking prominent ostia and postica; moderately
open skeleton lacking well-defined gastral and dermal cortex;
strands originate on gastral surface and parallel that surface
before arching steeply through the wall. Gastral strands ap-
proximately 0.10-0.23 mm apart and 0.10 mm in diameter,
with nodes 0.16-0.18 mm in diameter spaced 0.4-0. 5 mm
apart along strands and cross connected by beams 0.06-0.09
mm across to form rectangular meshes. Nodes slightly to
distinctly swollen but not spherical.

Branches more closely and regularly spaced and slightly
larger than in E. lithodendron Reid, and much better devel-
oped than closely spaced, rudimentary branches in E. (?)
setosum Reid. Lacks cortex and ostia that are developed in
both E. lithodendron and E. (?) setosum. Growth form and
branches more regular than in E. schmidti treubi Ijima.
Spherical nodes absent at spicule centra, although common
in related species.

DESCRIPTION. Several fragments are in the collection.
The most complete, the holotype, is a fragment approxi-
mately 1 2 cm high. It is an elongated branched form in which
the axial tube has undergone heterotomous division to pro-
duce a dichotomous structure in which one of the two branch-
es ceases to grow and the other grows to divide again. This

produces a zigzag growth pattern with angles of approxi-
mately 120-130 degrees between segments of the axial tube.
The axial tubes and branches are 8-10.5 mm in diameter.
The short lateral branches are spaced approximately 20 mm
apart along the axial tube and are approximately 1 cm long.
Oscula occur at the ends of the lateral rudimentary branches
and range from openings to somewhat constricted.

Walls of tubes and branches are 0.5- 1.1 mm thick and
have a generally thin, but well-defined gastral layer of thick-
ened strands and beams. Most of the wall is composed of
relatively open-textured endosomal skeleton. A dermal layer
is not differentiated in the well-developed dictyonal frame-
work. The thickened gastral layer usually affects only strands
and beams at the immediate gastral surface.

Neither ostia nor postica are clearly defined, although some
irregular interruptions in the outer part of the skeleton may
represent ostia. These are subcircular openings, 0. 1 5-0.3 mm
across, and are irregularly developed in the visible exterior.
No similar interruptions are evident in the inner part of the
skeleton, and water must have passed through the wall via
the open skeletal meshes.

Strands of the dictyonal skeleton originate near or at the
gastral surface and parallel that surface before swinging
abruptly derntally. They terminate as unattached conical to
spinelike rays at the outer margin of the wall, where they are
often nearly at right angles to the dermal surface. In the gastral
layer, strands are parallel and spaced 0.15-0.4 mm apart.
They are closest immediately distal to insertion of a new
strand, often by “branching,” and are farthest apart imme-
diately proximal to the insertion. Spicular nodes are spaced
by beams 0.4-0. 5 mm long along the gastral strands and are
connected laterally by beams 0.10-0.25 mm long, measured
horizontally on the gastral surface. Beams of gastral strands
are 0.08-0. 1 1 mm in diameter midway between nodes, with
most 0.10 mm across. Lateral gastral beams are 0.06-0.09
mm across, with most in the upper end of that range. Nodes
are 0.10-0.22 mm in diameter but without much swelling
other than slight flaring where rays meet. Nodes throughout
the skeleton are not spherical but appear almost rectangular.
Most such nodes are 0. 1 6-0. 1 8 mm across in the endosomal
part of the wall.

Endosomal and dermal beams are slightly smaller than
gastral ones, with common diameters of 0.06-0. 10 mm and
lengths of 0.2-0. 3 mm, thus spacing nodes three-dimen-
sionally that distance within the wall. This part of the wall
is less linearly organized than the gastral part and is 2-4
spicules thick. Free beams form only low rounded knobs or
cones on the gastral surface but pointed spines on the dermal
surface.

Skeletal mesh openings are generally rectangular in the
inner part of the skeleton but become much more irregular
and commonly triangular in the middle and outer part of the

wall.

Adnate small free hexactines occur commonly throughout
the skeleton. They are generally attached to a beam by one
ray, and the other rays of the tiny spicules are free. They
range from small spicules with rays only 0. 1 5 mm long and
0.005 mm in diameter up to larger forms with rays 0.3 mm

10 Contributions in Science, Number 344

Rigby and Jenkins: Tertiary Sponges

Figure 23. Eurele goederti n. sp., drawing of part of the skeleton
of the holotype showing smaller attached hexactines and larger beams
of gastral dictyonine net. Vertical strands are dominant elements in
this part of the net. Distal is toward the top, and proximal is toward
the base. LACMIP 6609, Locality 7, approximately X40.

long and 0.008-0.0 10 mm across in basal ray diameter. They
are frequently oriented with rays parallel to the major dic-
tyonal elements, in the general plane of those elements, or
at approximately 45 degrees to those elements where they
project out into the “cubic” meshes (Figure 23). Beams in
the skeleton all show granular to distinctly spinose surfaces,
many of the tiny spines only 0.005 mm across and 0.10 mm
or less high.

DISCUSSION. Eurete goederti n. sp. is most similar to
Cretaceous E. lithodendron Reid (1961:34-38, pi. 7, figs. 4a,
b) in general growth form but is slightly larger. In addition,
branching is more distant in E. goederti n. sp., and it lacks
a dermal cortex and ostia, which are present in E. litho-
dendron. Modern E. schmidti treubi Ijima (1927:170; Reid,
1961:37, fig. 2a) also has a similar growth form but branches
considerably more irregularly.

E. (?) setosum Reid (1958:38-40), from the Cretaceous of
Great Britain, is also a tubular branching form, but its branches
are often reduced to rudimentary structures, in contrast to
the distinct branches in E. goederti. In addition E. (?) setosum
has a dermal cortex and ostia in places, and branches are
spaced relatively closely together. Branching in E. goederti
is considerably more regular than in either E. lithodendron
Reid. E. (?) setosum Reid, or in the living E. schmidti treubi.
The northwestern United States fossil species also lacks
spherical spicule nodes that are common in the other species.

Depth ranges of living species of Eurete were summarized

by Schrammen (1912:1 84-185), based upon work by Schulze
and Carter. Shallowest occurrence cited by Schrammen is
220 m for Eurete carteri Schulze, and deepest is 717 m for
E. erectum Schulze. Most species are listed from depths of
300 or 360 meters. Reid ( 1 968:549) listed Aphrocallistes vas-
tus Schulze as having been collected in depths of 27.5 m
(Puget Sound), 97 m (Oregon), and 108 m (Vancouver, Brit-
ish Columbia) but noted that the Puget Sound site is the
shallowest authenticated record of modern dictyonine hex-
actinellids. By analogy it seems likely that the Washington
Oligocene species had a depth range of 100-350 m. The
fragmental nature of our material, however, suggests that the
assemblage may have been transported. Downslope transport
is suggested by occurrence of these and associated fossils in
thin, coarse, sandy to glauconitic beds, intercalated as high-
energy pulses in the dominantly fine-grained sequence.

TYPE SPECIMENS AND AVAILABLE MATERIAL.
The holotype, LACMIP 6609, is the largest fragment and
occurs with Aphrocallistes fragments, pieces of wood, fish
fragments, and foraminifers in a coarse sandy siltstone from
Locality 7, the type locality of the sponge species. The para-
type (LACMIP 6610) and an additional, more fragmental
piece of Eurete are also from the same locality. One concre-
tion from Locality 8 also contains tiny scraps of dictyonal
skeleton that are probably from E. goederti ; these are too
small to determine growth habit and other relationships but
have the same proportions in the gastral net as the holotype
of E. goederti.

ETYMOLOGY. Goederti, named for James Goedert, on
whose collections this and parallel studies of other groups
are largely based.

Order, Family, Genus Uncertain

Hexactinellid Root Tufts

Figures 11,12

Several concretions contain moderately well organized root
tufts of hexactinellid sponges. These are composed of con-
centrically layered, aligned, large, doubly tapering oxeas (?),
with maximum diameters of 1. 1-1.4 mm at approximately
midlength. Neither entire length nor preserved tips were ob-
served on any single spicule, but spicule fragments at least 3
or 4 cm long occur in fine-grained calcareous siltstone where
sharp tips and double taper are well shown. They form clus-
ters 3 or 4 cm across and are commonly isolated from other
sponges. In one small concretion from Locality 2, isolated
large root tuft spicules occur embedded in irregular basal
spicular masses of Aphrocallistes. Bases of other specimens
of the species do not have such spicules and apparently were
cemented to bivalve or gastropod fragments or pebbles. This
implies that the tuft spicules are not an integral part oi Aphro-
callistes but that the sponge from Locality 2 overgrew ex-
posed root tuft spicules, which formed a solid substrate
“island” on the muddy bottom. Concretions, without Aphro-
callistes, from the same locality contain both root tuft clusters
and fragments of Eurete, but the sponge fragments and root
spicules are not mtergrown and may be unrelated biologi-
cally. The root tufts may represent sponges not otherwise

Contributions in Science, Number 344

Rigby and Jenkins: Tertiary Sponges 1 1

preserved in the collections. Taxonomic relationships of root
tufts are unknown.

FIGURED SPECIMENS AND AVAILABLE MATE-
RIAL. Figured specimen LACMIP 6612 is from Locality 5,
and 6611 is from Locality 7. Five other concretions con-
taining tuft fragments were collected from Locality 2, and
one was collected from Locality 5.

Similar root tufts of large bundled oxeas, which show con-
centric layering, occur at two other localities (Armentrout.
personal communication, 1982). One such tuft was collected
by Susan Bee, from Portland State University, from Late
Eocene siltstone of the upper member of the Keasey For-
mation. The fossil was float in a 5-m high railroad cut south
of and upslope from the abandoned railroad trestle across
Highway 47, between Buxton and Vernonia, approximately
20 m (80 feet) west and 210 m (700 feet) north of the south-
east corner of Sec. 8, T. 3 N., R. 4 W., on the Vernonia 7 V2-
minute quadrangle, Washington County, Oregon (Locality
Tok-Ml 12 of Warren and others, 1945).

The other tuft was collected from Late Eocene siltstone of
the Lincoln Creek Formation by J.M. Armentrout. It came
from the upper 3-m ( 10-foot) interval at the top of a cliff on
the south side of Canyon River, just upstream from a logging
road bridge, approximately 240 m (800 feet) west and 67 m
(220 feet) north of the southeast corner of Sec. 13, T. 21 N.,
R. 6 W., on the Grisdale 15-minute quadrangle. Grays Har-
bor County, Washington (Locality CR9 and 10 of Armen-
trout, 1973).

ACKNOWLEDGMENTS

The sponges and associated fossils were collected by James
L. and Gail H. Goedert over a several-year period. Edward
Wilson of the Natural History Museum of Los Angeles Coun-
ty suggested the project, arranged for loans of the collections,
and reviewed the manuscript. Ellen J. Moore of the U.S.
Geological Survey; Lein F. Hintze and James L. Baer of the
Department of Geology, Brigham Young University; Victor
Zullo, Department ofGeology, University of North Carolina;
and John M. Armentrout of Mobil Oil Corporation, Dallas,
Texas, critiqued the manuscript and added stratigraphic and
biostratigraphic information. Armentrout provided data on
additional occurrences of root tufts from the Keasey and
Lincoln Creek formations. Drafts of the manuscript were
typed by Ann Bracken and Camille Crezee. David Jenkins
did preliminary work on part of the collections while on a
graduate internship in the Department of Geology at Brig-
ham Young University. Some costs of manuscript and illus-
tration preparation were covered by National Science Foun-
dation grant DEB 78-25229.

LITERATURE CITED

Armentrout, J.M. 1973. Molluscan paleontology and bio-
stratigraphv of the Lincoln Creek Formation, late Eocene-
Oligocene, southwestern Washington. (Ph.D. disserta-
tion) University of Washington. Seattle, 478 pp.

. 1975. Molluscan biostratigraphy of the Lincoln

Creek Formation, southwest Washington. In Weaver,

D. (ed.). Future Energy Horizons of the Pacific Coast;
Paleogene Symposium and selected technical papers.
Annual Meeting of the Pacific Sections, American As-
sociation of Petroleum Geologists, Society of Economic
Paleontologists and Mineralogists, Society of Economic
Geologists, Long Beach, California, pp. 14-48.

. 1977. Cenozoic molluscan stages of Oregon and

Washington. Abstracts with Programs. Geological So-
ciety of America 9(7):882— 883.

. 1981. Correlation and ages of Cenozoic biostrati-
graphic units in Oregon and Washington. In Armen-
trout, J.M. (ed.). Pacific Northwest Cenozoic Biostratig-
raphy, Geological Society of America Special Paper 1 84,
pp. 137-148.

Beikman, H.N., W.W. Rau, and H.C. Wagner. 1967. The
Lincoln Creek Formation. Grays Harbor Basin, south-
western Washington. U.S. Geological Survey Bulletin
1244-1:11-11 14.

Dali, W.H. 1909. Contributions to the Tertiary paleontol-
ogy of the Pacific coast, I. The Miocene of Astoria and
Coos Bay, Oregon. U.S. Geological Survey Professional
Paper 59, 278 pp.

Dana, J.D. 1849. Geological observations of Oregon and
northern California, U.S. Exploration Expedition 1 838—
1842 under the command of Charles Wilkes, Geology,
v. 10, pp. 61 1-678; Appendix, pp. 722-723, 729-730;
Atlas.

Gray, J.E. 1858. On Aphrocallistes, a new genus of Spon-
giadae from Malacca. Proceedings of the Zoological So-
ciety of London 26: 114-115.

. 1867. Notes on the arrangement of sponges, with

a description of some new genera. Proceedings of the
Zoological Society of London. 1867, pp. 492-558.

Ijima. I. 1927. The Hexactmellida ofthe Siboga Expedition.
Siboga-Expeditie, Leiden, v. 6, 383 pp.

Moore, E. J. 1963. Miocene marine mollusks from the
Astoria Formation of Oregon. U.S. Geological Survey
Professional Paper 419. 109 pp.

Rau, W.W. 1958. Stratigraphy and foraminiferal zonation
in some of the Tertiary rocks of southwestern Washing-
ton. U.S. Geological Survey Oil and Gas Investigations
Chart OC-57, 2 sheets.

. 1964. Forantinifera from the northern Olympic

Peninsula, Washington. U.S. Geological Survey Profes-
sional Paper 375-G:G 1-G33.

. 1981. Pacific Northwest Tertiary benthic forami-
niferal biostratigraphic framework — An overview. In
Armentrout. J.M. (ed.). Pacific Northwest Cenozoic Bio-
stratigraphy, Geological Society of America Special Pa-
per 184, pp. 76-84.

Reid, R.E.H. 1958-64. A monograph of the Upper Creta-
ceous Hexactinellida of Great Britain and Northern Ire-
land. Palaeontographical Society, London. 1958, Part I.
pp. i-xlvi; 1958, Part II. pp. xlvii-xlviii, 1-26; 1961,
Part III, pp. 27-48; 1964, Part IV, pp. xlix-cliv.

. 1968. Bathymetric distributions of Calcarea and

Hexactinellida in the present and the past. Geological
Magazine 1 05( 6): 546— 5 59 .

12 Contributions in Science, Number 344

Rigby and Jenkins: Tertiary Sponges

Schrammen, A. 1912. Die Kieselspongien der oberen Kreide
von Nordwestdeutschland; II Teil, Triaxonia (Hexacti-
nellida). Palaeontographica, Supplement 5. pp. 177-385.

Schulze, F.E. 1887. Report on the Hexactinellida collected
by H.M.S. Challenger during the years 1873-76. Report
of Scientific Results of the Voyage of the H.M.S. Chal-
lenger. Zoology 21:1-513.

Semper, C. 1869. Einige neue Kieselschwamme der Phil-
ippen: Hyalonema Schultzei n. sp. und Eurete n. g.
Verhandlungen der Physikalisch-Medicinischen Gesell-
schaft in Wurzburg, new series, 1 868, 1 : 29— 30 (also pub-
lished in 1 868, Annals and Magazine of Natural History
2:372-373).

Snavely, P.D., Jr., N.S. MacLeod. W.W. Rau, W.O. Addi-
cott, and J.E. Pearl. 1975. Alsea Formation — An Oli-
gocene marine sedimentary sequence in the Oregon Coast
Range. U.S. Geological Survey Bulletin 1 395-F:Fl-F20.

Warren, W.C., R.M. Grivetti, and H. Norbisrath. 1945.
Geology of northwestern Oregon. U.S. Geological Sur-
vey Oil and Gas Investigations. Preliminary Map 42.

Weaver, C.E. 1912. A preliminary report on the Tertiary
paleontology of western Washington. Washington Geo-
logical Survey Bulletin 15:1-80.

. 1937. Tertiary stratigraphy of western Washington

and northwestern Oregon. Washington University (Se-
attle) Publications Geology 4:1-266.

Weaver, C.E. , and others. 1944. Correlation of the marine
Cenozoic formations of western North America (Chart
no. 1 1 ). Geological Society of America Bulletin 55:569-
598.

Wells, R.E. 1979. Geologic map of the Cape Disappoint-
ment-Naselle River area. Pacific County, Washington.
U.S. Geological Survey Open-File Report 79-389.

Zullo. V.A. 1982. Arcoscalpellum Hoek and Solidobalanus
Hoek (Cirripedia, Thoracia) from the Paleogene of Pa-
cific County, Washington, with a description of a new
species of Arcoscalpellum. Natural History Museum of
Los Angeles County, Contributions in Science 336:1-9.

Received 22 September 1982; accepted 7 February 1983.

Contributions in Science, Number 344

Rigby and Jenkins: Tertiary Sponges 13

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Number 345
7 October 1983

* a mm species of

AMERICAN SALAMANDERS, /WITH, A REVIEW.
DPICAL GENERA
IA, CAUDATA, PLETHGIKMIIJME)

!8wK

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NEW GENERA AND A NEW SPECIES OF
CENTRAL AMERICAN SALAMANDERS, WITH A REVIEW
OF THE TROPICAL GENERA
(AMPHIBIA, CAUDATA, PLETHODONTIDAE)

David B. Wake1 2 and
Paul Elias'

ABSTRACT. A new genus and species of plethodontid bolitoglos-
sine salamander is described from material collected in northwestern
Guatemala. Bradvtriton silus new genus, new species, is unique in a
combination of structural characteristics that includes a laterally
compressed tail, stocky body with no clearly defined neck, and short,
slender limbs bearing syndactylous hands and feet. To diagnose the
new genus, an analysis of the entire neotropical assemblage of pleth-
odontid salamanders was undertaken. Approximately 138 species
belong to the supergenus Bolitoglossa and 1 1 genera are recognized.
The genus Chiropterotriton is shown to be polyphyletic; thus, two
new genera are described. Notolriton new genus, includes the picadoi
group of Chiropterotriton beta. Dendrotriton new genus, includes the
bromeliacia group of Chiropterotriton beta. Those species formerly
called Chiropterotriton alpha remain as the sole representatives of
the genus. Eight of the eleven neotropical genera are shown to be
monophyletic. Of the three exceptional genera, both Dendrotriton
and Nototriton are nonparaphyletic relative to all genera except Oed-
ipina, but Dendrotriton is easily distinguished from Oedipina. No-
totriton may be paraphyletic relative to Oedipina. but the two genera
can be readily distinguished on the basis of major differences in
ecology and shape. Pseudoeurycea comprises morphologically gen-
eralized species that may not be far removed from the morphology
of the ancestral stock of the entire neotropical group. Only Nyctanolis
and Chiropterotriton are more plesiomorphic than Pseudoeurycea.
Lists of species assigned to the genera are provided. Potential rela-
tionships within the supergenus Bolitoglossa are discussed, but par-
allelism and convergence have been so extensive that no definitive
statement concerning generic affinities is possible.

RESUMEN. Un nuevo genero y especie de salamandra plethodon-
tida bolitoglossina se describe en base a material recolectado en el
noroeste de Guatemala. Bradyiriton silus. nuevo genero, nueva es-
pecie, es peculiar por su combinacion de caracteres que incluye una
cola comprimida lateralmente, un cuerpo macizo con cuello esca-
samente definido, y patas delgadas y cortas provistas de manos y
pies syndactilos. Para diagnosticar este nuevo genero se realizo un
analisis de! grupo completo de salamandras plethodontidas neotro-
picales. Las aproximadamente 138 especies pcrtenecen al superge-
nero Bolitoglossa, en el cua! se reconocen 1 1 generos. El genero

Chiropterotriton se sefiala como politiletico y por lo tanto dos nuevos
generos se describen. Nototriton. nuevo genero, incluye el grupo
picadoi Ac Chiropterotriton beta. Dendrotriton, nuevo genero, inclu-
ye el grupo bromeliacia de Chiropterotriton beta. Las especies pre-
viamente incluidas en Chiropterotriton alfa permanecen como las
unicas represenlantes de este genero. Ocho de los once generos neo-
tropicales se senalan como monofileticos. De los tres generos res-
tantes, tanto Dendrotriton como Nototriton son monofileticos en
relacion a todos los generos excepto Oedipina, aunque Dendrotriton
es facilmente distinguible de Oedipina. Nototriton podria ser para-
liletico en relacion a Oedipina. pero estos dos generos pueden ser
rapidamente reconocidos en base a sus marcadas diferencias en eco-
logia y forma corporal. Pseudoeurycea comprende especies morfo-
Iogicamente generalizadas que podrian estar no muy alejadas de la
morfologia del grupo ancestral del ensamble neotropical analizado.
Solo Nyctanolis y Chiropterotriton son mas plesiomorficos que Pseu-
doeurycea. Listas de las especies asignadas a cada genero se presen-
tan. Tambien se discuten las potenciales relaciones dentro del su-
pergenero Bolitoglossa. pero paralelismos y convergences han sido
tan prevalentes que ninguna opinion definitiva puede plantearse en
cuanto a alinidades genericas.

INTRODUCTION

In the summer of 1974, the junior author collected several
species of salamanders in a remote area in northwestern Gua-
temala. Included in this collection were three species that
obviously were undescribed. Subsequent morphological
analysis showed that two of the new species have combi-
nations of traits that require them to be placed into two new
genera. One of these ( Nyctanolis ) has been described else-
where (Elias and Wake, 1983). In this paper we describe the
second new genus. We also report the results of a detailed

1. Museum of Vertebrate Zoology and Department of Zoology,
University of California, Berkeley, California 94720.

2. Research Associate in Herpetology, Natural History Museum
of Los Angeles County, Los Angeles, California 90007.

Contributions in Science, Number 345, pp. 1-19
Natural History Museum of Los Angeles County, 1983

ISSN 0459-8113

analysis of all of the currently recognized genera of tropical
salamanders. As a result of this analysis, which was a nec-
essary background for the description of the new genera and
species, two additional new genera are erected. These are
described herein, and diagnostic characters are given for all
existing genera of tropical salamanders. In addition, all rec-
ognized species of tropical salamanders are referred to a ge-
nus, and relationships among the genera are examined. This
analysis reinforces previous views (Wake, 1966; Wake and
Lynch, 1976) that there has been very extensive parallelism
and convergence during the adaptive radiation of pletho-
dontid salamanders in the New World tropics.

BACKGROUND

Although several genera of New World tropical salamander
species were described in the nineteenth century, and some
knowledge of the diversity of the group was available at that
time, the most authoritative taxonomic work on these sal-
amanders during the early part of the twentieth century (Dunn,
1926) placed all tropical species in a single plethodontid ge-
nus, Oedipus. Dunn considered Oedipus to be “a large, mod-
em genus of some 30 species. The extremes are quite different
but there are many connecting links.” There was little in-
crease in knowledge of tropical salamanders until the mid-
I 930’s, when Schmidt, Taylor, and other workers began pub-
lishing their results (for historical summary, see Wake, 1 972,
and Smith and Smith, 1976). Taylor (1940) showed that
Oedipus was a preoccupied name, and substituted the old
name Bolitoglossa for the entire assemblage, except for a
group of diminutive species, which he placed in Cope’s ( 1 869)
old genus Thorius. Shortly thereafter, Taylor (1944) under-
took a radical revision of all the neotropical salamanders, in
which he described four new genera (Chiropterotriton. Par-
vimolge, Magnadigita, Pseudoeurycea), resurrected Oedi-
pina and Haptoglossa. and continued to recognize Bolito-
glossa and Thorius. Since that time, there has been relative
stability in the generic classification of the group. Lineatriton
was established by Tanner (1950), Magnadigita was placed
in the synonymy of Bolitoglossa by Wake and Brame ( 1 963),
and Haptoglossa was placed in the synonymy of Oedipina
by Brame (1968). The entire assemblage was characterized
and the genera defined by Wake ( 1966), who established the
supergenus Bolitoglossa for this group. This supergenus, and
the supergenera Hydromantes and Batrachoseps, were in-
cluded in the tribe Bolitoglossini, subfamily Plethodontinae,
of the family Plethodontidae. Many species have been dis-
covered since 1926; herein, we recognize 138.

The most recent comprehensive treatment of the super-
genus Bolitoglossa! Wake and Lynch, 1976) dealt mainly with
ecology, distribution, and biogeography and offered little new
morphological or taxonomic analysis. Since the publication
of that paper, major new discoveries have been made; here,
we attempt to give phylogenetic perspective to the available
information.

The present effort is not definitive, for continued new dis-
coveries indicate that our knowledge of the group, even at
the generic level, remains incomplete. We are aware of many

undescribed species, and a number of described species-
including some crucially important ones— are known from
only one or a very few individuals. Nevertheless, we believe
that the broad outlines of relationship are sufficiently clear
to justify the substantial revision we undertake here.

MATERIALS AND METHODS

We have not considered in detail the species of Bolitoglossa,
Oedipina, and Thorius, all large, monophyletic genera, which
are either under intense study presently ( Bolitoglossa by D.B.
Wake, P. Alberch, A. Larson, and colleagues, Thorius by J.
Hanken), or have been analyzed recently (Oedipina by Brame,
1968). Instead, we have emphasized apparently polyphyletic
and paraphyletic groups, as well as newly discovered species
that do not easily fit into any existing genus. In these critical
cases, we have analyzed doubly cleared and stained speci-
mens, histological sections, and dissections of fixed material.
As a point of departure for the choice and analysis of char-
acters, we have relied on the literature, which will be cited
where appropriate.

We have had access to adequate samples of most of the
species. Important species for which we have lacked sufficient
material but nevertheless can offer useful information include
(present generic designation used): Chiropterotriton barbouri,
C. richardi, and Parvimolge praecellens. These are discussed
in appropriate places in the following account.

As noted above, one of the new genera has recently been
described (Elias and Wake, 1983). We begin this work with
an account of the characters that are used in our generic
revision. We then describe a newly discovered genus and
species and erect two new generic names to encompass pre-
viously known species. Finally, we present a preliminary
phylogenetic analysis.

CHARACTERS USED FOR ANALYSIS

A necessary first step in the process of phylogenetic recon-
struction is the identification of monophyletic taxa (sensu
Eldredge and Cracraft, 1980). Our goal is to deduce mono-
phyletic groups from a matrix of morphological character
states. There are incomplete data for too many species to
justify an extensive analysis at the species level. Accordingly,
we have relied on the literature and our own previous work
to recognize groups of species. We have treated these groups
as hypotheses and have investigated whether or not each is
a monophyletic taxon. A group that is found to be mono-
phyletic is then treated as a genus. We believe that the generic
level of classification should combine species into monophy-
letic units that are separated from other such units by mor-
phological gaps. Ideally, the gaps will coincide with ecological
and behavioral differences as well. Thus, our objective is to
define generic-level units that are meaningful in both phy-
logenetic and ecological terms.

In the analysis that follows, each genus is scored for eigh-
teen characters (Table 1). In certain genera, there is some
variation in these characters among, or even within, species.
Only if a derived condition is characteristic of every known
population and species (i.e., only if it appears in most adults)

2 Contributions in Science, Number 345

Wake and Elias: Tropical Salamanders

is the genus as a whole scored as derived. Thus, our character
scoring should reflect the most primitive common denomi-
nator for each genus discussed and should represent the sit-
uation in the common ancestor of the included species. The
disadvantage of this conservative approach is that it masks
a certain amount of parallelism.

The characters used in our analysis are described below.
All of the characters are treated as two-state characters, with
primitive (plesiomorphic) and derived (apomorphic) states
identified on the basis of outgroup analysis (Eldredge and
Cracraft, 1980). Our outgroups are the genera Hydromantes
and Batrachoseps, the other members of the tribe Bolito-
glossini; and the members of the tribe Plethodontini. Most
of the osteological characters have already been discussed in
detail by Wake (1966) and Lynch and Wake (1978). By con-
vention, we code the characters (0) to indicate primitive and
(1) to indicate derived states.

1. Mesopodial mineralization. Mesopodials are cartilagi-
nous throughout life (0) or are mineralized in adults (1).

2. Tibial spur. A well-developed rodlike structure, free
from the shank of the bone, is present proximally (0) or is
absent or reduced to a low ridge ( 1 ).

3. Carpal fusion. The ulnare and intermedium are discrete
(0) or fused ( 1 ).

4. Tarsal fusion. Distal tarsals four and five are discrete
(0) or fused (1).

5. Mental glands. Clusters of individual glands form a
small patch located near the tip of the chin in male tropical
salamanders (Truffelli, 1954). The patch is either externally
visible, and ovoid, circular, or subtriangular, with rounded
or low columnar individual internal glands (0), or is exter-
nally obscure, with long, tubular individual internal glands
that are oriented posteriorly from their openings near the
chin (1). The tubular glands extend posteriorly as twisted
tubes covered by the skin of the gular area.

6. Stylus of operculum. The stylus, or columella, is not well
developed in any of the tropical species, but it may be present,
with a distinct rodlike shape (0) or reduced to a broad bulge
or be entirely absent (1).

7. Preorbital processes of vomers. These slender processes,
which may or may not bear teeth, are either present between
the internal nares and the orbit (0) or absent (1).

8. Prefrontal bones. These bones are either present in vary-
ing degrees of development (0) or absent (1).

9. Septomaxillary bones. These bones are not well devel-
oped in any tropical salamanders, but small ossicles may be
present (0) or absent (1).

10. Frontal processes of premaxillary bone. Where only a
single premaxillary bone is present, the frontal processes are
either fused together at the point of origin on the dorsal
surface of the pars dentalis and for some distance dorsopos-
teriorly (1) or separated for their entire length (0). We score
those few instances in which the processes arise separately
and subsequently fuse as 0.

1 1 . Premaxillary bones. The pars dentalis of plethodontid
salamanders is either divided (0) or fused so that only a single
bone is present ( 1 ).

12. Skull roof. The parietal bones may either meet or ap-

proach each other very closely on the midline, thus forming
a complete skull roof (0), or be widely separated and joined
by a connective tissue sheet, thus forming an incomplete skull
roof ( 1 ).

1 3. Sublingual fold. A small to large fold of glandular skin
underlying the tongue may be present (0) or absent (1).

14. Limb length. Limbs are moderate to short in length,
so that they overlap slightly or not at all when adpressed (0),
or are very long, so that they overlap by more than two costal
interspaces ( 1 ).

1 5. Tarsal arrangement. Distal tarsal four is discrete from
and larger than distal tarsal five and articulates with the
fibulare (0), or distal tarsal five is discrete from and larger
than four and articulates with the centrale (1). When distal
tarsals four and five are fused, the character is scored as 0.

16. Trunk vertebral shape. The ratio of centrum length to
posterior centrum diameter is low (2.5-4), and the vertebrae
are relatively wide and short (0), or the ratio is high (greater
than 4), and the vertebrae are relatively narrow and elongate
(1).

17. Number of trunk vertebrae. There are either 14 (0) or
18 or more (1).

18. Tail shape. The tail is either round or ovoid in cross
section (0) or strongly compressed laterally, with a dorsal,
glandular ridge (1).

The eighteen characters are listed in Table 1, along with
the states that characterize each of the 1 1 genera we recognize
in this paper. This table alone can be used to diagnose all of
the genera. In most instances the genera are diagnosed by
autapomorphies (derived characters unique to a given taxon),
combinations of synapomorphies (derived characters shared
by two or more taxa), or both. In some instances, determi-
nation that a given genus is monophyletic requires an analysis
of the distribution of character states over all taxa; such cases
are detailed in the Discussion. All 1 1 genera can be readily
diagnosed on the basis of unique combinations of traits, in-
cluding both apomorphic and plesiomorphic states. Bolito-
glossa probably should be divided into at least two taxa
(Wake and Lynch, 1 976), but it is a well-defined (if unwieldy),
monophyletic taxon as currently recognized. However, our
generic concept does require that we recognize four new ge-
neric taxa. One of these, Nyctanolis, is a recently discovered
monotypic genus that has been described elsewhere (Elias
and Wake, 1983). A second newly discovered genus, also
monotypic, is described immediately below. The other two
new genera result from subdivision of an existing genus and
are described below in the context of our generic diagnoses.

DESCRIPTION OF A NEW GENUS
AND SPECIES

Bradytriton new genus

TYPE SPECIES. Bradytriton si/us new species.

DIAGNOSIS. A plethodontid salamander belonging to
the subfamily Plethodontinae, tribe Bolitoglossini, superge-
nus Bolitoglossa. Bradytriton is a short-snouted, stout-bod-
ied, short-tailed taxon with small limbs and diminutive, syn-

Contributions in Science, Number 345

Wake and Elias: Tropical Salamanders 3

Table 1. Distribution of character states in genera of neotropical plethodontid salamanders. See text for listing of characters. 0 = primitive

state; 1 = derived state.

Genus

Character

1

2

3

4

5

6

7

8

9

10

1 1

12

13

14

15

16

17

18

Bolitoglossa

0

0

0

1

0

1

0

0

1

0

1

0

1

0

0

0

0

0

Bradytriton

0

0

1

1

1

1

0

0

1

0

1

0

0

0

0

0

0

1

Chiropterotriton

0

0

0

0

0

0

0

0

0

0

1

0

0

0

1

0

0

0

Dendrotriton

0

1

0

0

0

0

0

1

0

0

1

0

0

0

0

0

0

0

Lineatriton

1

0

0

1

0

0

0

0

1

1

1

0

0

0

0

1

0

0

Nototriton

0

0

1

1

0

1

0

0

1

1

I

0

0

0

0

0

0

0

Nyctanolis

0

0

0

0

0

0

0

0

1

0

0

0

0

1

0

0

0

0

Parvimolge

1

0

]

1

1

1

1

0

1

0

1

0

0

0

0

0

0

0

Pseudoeurycea

0

0

0

0

0

0

0

0

0

0

1

0

0

0

0

0

0

0

Oedipina

0

1

1

1

1

1

0

1

1

1

1

0

0

0

0

0

1

0

Thorius

1

0

1

1

0

1

1

0

1

0

1

1

0

0

0

0

0

0

dactylous hands and feet, readily distinguished from most
other members of the supergenus on the basis of those fea-
tures. It differs from all other members of the supergenus in
having a laterally compressed tail with a relatively massive
dorsal glandular ridge. Thorius and Parvimolge have short
limbs, but these genera are much smaller and differ in other
ways as well: Thorius has an incomplete brain case, and both
genera frequently have mineralized mesopodial and hyo-
branchial ossifications as adults. Nototriton and Dendrotriton
have long, slender tails. All Bolitoglossa lack sublingual folds.
Most Pseudoeurycea have long tails and long limbs, and all
lack the mesopodial fusions (ulnare-intermedium in manus;
fourth and fifth distal tarsal in pes) that characterize Bradytri-
ton.

ETYMOLOGY. From bradys, Gr., referring to the slow
and lethargic movements of the living animals, and triton,
Gr., a commonly used term for salamanders.

Bradytriton silus new species

Figures 1 through 6

HOLOTYPE. Museum of Vertebrate Zoology (MVZ)
131587, an adult female from Finca Chiblac, 15 km NE
Barillas, Depto. Fluehuetenango, Guatemala, elevation 1,310
m (4,300 ft), collected by P. Elias, 6 September 1974.

PARATYPES. MVZ 131586, LACM 134566, same data
as the holotype; MVZ 131589-131594 (6 specimens), same
locality and collector as holotype, collected 7 September 1974;
MVZ 1 34635-1 34637 (3 specimens), El Rayo, elevation 1 ,370
m (4,500 ft), 3 km S buildings of Finca Chiblac, 10 km NE
Barillas, Huehuetenango, Guatemala, collected by J. Jackson
and P. Elias on 1 September 1975; MVZ 134638, same lo-
cality and collectors as preceding series, collected on 31 Au-
gust 1975; MVZ 173063, Finca Chiblac, 10 km NE Barillas,
Depto. Huehuetenango, Guatemala, elevation 1,370 m (4,500
ft), collected by H.B. Shaffer and P. Elias on 2 July 1977;
MVZ 173064, same data as preceding number, collected be-
tween August 1975 and October 1977 by J. Jackson and P.
Elias.

DIAGNOSIS (measurements in millimeters). See Gener-
ic Diagnosis. A stout species of moderate size (standard length,
SL, in four adult males, 39.1-53.3, mean 45.3; seven adult
females 44.5-53.0, mean 49.0; Table 2) with a very stout,
short tail (SL/tail length in three adult males is 1 .3-1 .7, mean
1.4; in six adult females, 1.2-1. 7, mean 1.4), short limbs
(when adpressed, fore and hind limbs leave 2-3, mean 2.3,
costal interspaces uncovered in four adult males; 2.5-3, mean
2.9, in seven adult females), and small, syndactylous hands
and feet. The relatively broad head (SL/head width in four
adult males is 5.9-6. 1 , mean 6. 1 ; in seven adult females, 5.7-
6.6, mean 6.2) is essentially continuous with the body, and
there is no neck. Maxillary teeth (41-50, mean 44.0, in four
adult males; 40-59, mean 47.9, in seven adult females) and
vomerine teeth (16-22, mean 18.5, in four adult males; 17-
25, mean 20.6, in seven adult females) are moderate in num-
ber. The species is reddish brown in color, with obscure
streaking and flecking of lighter and darker pigments.

DESCRIPTION. This stout, short-limbed species has a
very short, truncate snout and diminutive hands and feet.

o I 2 3 4

Figure 1. Dorsal view of MVZ 131587, the holotype of Bradytriton
silus new genus and species.

4 Contributions in Science, Number 345

Wake and Elias: Tropical Salamanders

The nostrils are small. Labial protuberances are poorly de-
veloped in females, which have an especially short snout,
but are large and wide in adult males; the protuberances
produce a broad muzzle rather than the elongate snout typical
of other tropical salamanders. Mental glands are present in

males but are not externally demarcated; the glands consist
of a cluster of moderately long tubules that converge in the
region of the mandibular symphysis, where they open indi-
vidually to the ventral surface of the anterior mtermandibular
region. The tongue is adetoglossal, with a distinct small pad.

Figure 2. Parasagittal section through the head of a specimen (MVZ 134637) of Bradytriton silus new genus and species. The section is near
the midline and passes through the mandibular symphysis.

Left. The arrow points to an enlarged premaxillary tooth and indicates the part of the section that is magnified in the view on the right.
When the mouth is closed, the premaxillary teeth extend outside the mouth and lie near the ventral and anterior parts of the lower jaw, near
the openings of the mental gland. The long, tubular ducts of the mental gland are evident immediately posterior to the mandible. Note that
no external ventral swelling is present. Also well displayed in this section is the complex free tongue characteristic of the tribe Bolitoglossim.
The hyoglossal muscles are attached to the apparently flexible anterior tip of the basibranchial. Immediately anterior to the tongue is the large
sublingual fold, characteristic of all of the tropical salamanders except Bo/itoglossa.

Right. Enlargement of the premaxillary tooth at the end of the arrow in the left part of this figure. The pedicel of the tooth is attached to
the posteroventral border of the premaxillary bone (here oriented so that the border appears to be ventral). The crown is relatively large and
is unicuspid. The cusp is strongly hooked.

Table 2. Data for type series of Bradytriton silus new genus and species.*

Sex

Standard

length

Mead

length

Head

width

Foot

width

Hind

limb

length

Fore

limb

length

Tail

length

Maxil-

lary

tooth

number

Premax-

illary

tooth

number

Vomerine

tooth

number

I.imb

inter-

val

MVZ 131589

6

53.3

11.3

8.7

3.3

12.1

9.4

—

50

3

17

3.0

MVZ 134636

S

46.4

10.3

7.6

3.1

1 1.0

9.5

36.0

41

5

16

2.0

MVZ 134637

6

42.3

oo

bo

6.8

2.3

9.6

8.7

31.6

43

6

19

2.0

MVZ 134638

6

39.1

9.0

6.6

2.3

8.7

8.0

23.2

42

8

22

2.0

MVZ 131591

2

53.0

10.4

8.7

3.2

10.7

9.5

30.6

59

8

22

3.0

MVZ 131593

2

51.5

10.3

8.8

2.5

9.2

9.6

-

48

8

20

3.0

MVZ 131586

2

50.1

9.7

7.8

2.9

1 1.0

10.1

40.4

46

9

19

3.0

MVZ 134635

2

48.8

10.1

7.4

2.7

1 1.0

9.2

34.1

46

9

25

3.0

MVZ 131587**

2

48.5

9.5

7.5

2.5

9.4

9.5

36.6

52

6

17

3.0

MVZ 131592

2

46.6

9.7

8.2

2.9

1 1.0

9.5

34.5

44

10

23

2.5

LACM 134566

2

44.5

9.3

6.8

2.5

8.6

8.9

33.3

40

9

18

3.0

MVZ 131590

J

35.6

8.1

6.7

2.2

7.5

7.3

21.5

31

8

16

2.5

MVZ 131594

J

34.4

7.8

6.1

1.4

7.2

7.1

22.3

37

8

18

2.0

* All measurements are in millimeters.
** Holotype.

Contributions in Science, Number 345

Wake and Elias: Tropical Salamanders 5

10mm

Figure 3. Radiograph of the broken tail ofa specimen (MVZ 131586)
of Bradytriton silus new genus and species. Dorsal to the top. Note
the large glandular mass above the vertebral column.

The sublingual fold is well developed. The profile of the
rounded head is undifferentiated from the trunk, and there
is no apparent neck region. A deep unpigmented groove ex-
tends beneath the eye, following its curvature, but does not
extend to the lip. The eyes are moderate in size and protrude
slightly beyond the margin of the jaw. The postorbital groove
is an indistinct furrow that extends posteriorly from the eye
and intersects a deep vertical groove at right angles. The latter
groove passes behind the base of the mandible and becomes
a clearly defined nuchal groove. The gular fold is pronounced.
Vomerine teeth are few to moderate in number and are ar-
ranged in a single, curved row that extends laterally beyond
the lateral margin of the internal nares. Maxillary and vo-
merine teeth increase in number to some extent with in-
creasing body size. The maxillary tooth row extends back to

Figure 4. Dorsal views of the left foot (left) and of the right hand
(right) of an adult male (MVZ 1 73064) Bradytriton silus new genus
and species. Cartilage is stippled. The phalangeal elements are very
poorly ossified and are less distinct than illustrated here (see text).
Note the syndactylous nature of the digits.

Figure 5. Dorsal view of the skull of an adult male (MVZ 1 73063)
Bradytriton silus new genus and species. Bone is outlined, and car-
tilage is stippled. The external nares and the nasolacrimal foramina
are black.

a point nearly posterior to the eyeball. Premaxillary teeth are
3-8, mean 5.5, in four adult males, 6-10, mean 8.4, in seven
adult females; the teeth of males are very large and hooked
and protrude under or through the lip (Fig. 2). The trunk is
stout and cylindrical. The tail is of moderate length and is
strongly tapered near its tip. It has a marked basal constric-
tion and is strongly compressed laterally. The tail appears
stout when viewed laterally but seems narrow from a dorsal
perspective (Figs. I. 3). Postiliac glands are distinct. The
limbs are short and slender. Hands and feet are diminutive
and syndactylous; the greatest foot width is about ‘/15 SL and
the foot is barely wider than the lower limb. The digital tips
are free, but adjacent digits are fused for most of their length
(Fig. 4). There are no subdigital pads. The toes, in order of
decreasing length, are 3, 2, 4, 5, 1; the fingers, 3, 2, 4, 1.

MEASUREMENTS OF THE HOLOTYPE (in milli-
meters). Head width 7.5; snout to gular fold (head length)
9.2; head depth at posterior angle of jaw 4.6; eyelid width
1.9; anterior rim of orbit to snout 2.0; horizontal orbit di-
ameter 2.4; interorbital distance 2.3; vomerine teeth virtually
continuous with parasphenoid tooth patch; snout to fore limb
12.6; distance separating internal nares 1.8; distance sepa-
rating external nares 1.9; snout projection beyond mandible
0.3; snout to posterior angle of vent (standard length) 48.5;

6 Contributions in Science, Number 345

Wake and Elias: Tropical Salamanders

snout to anterior angle of vent 44.4; axilla to groin 28. 1 ; tail
length 36.6; tail width at base 3.4; tail depth at base 4.8; fore
limb length (to tip of longest toe) 9.5; hind limb length 9.4;
hand width 1.7; foot width 2.5.

COLORATION IN LIFE. The dorsal coloration of this
species is reddish brown. This color is broken by irregular
Hecks of black on the anterior trunk and dominated by black
on the head. Brown coloration extends halfway down the
lateral surfaces. The cheeks, sides of the tail, and lower flanks
are black, overlain with dense white flecks. The legs are brown
proximally but grade distally to black with white flecking.
The feet are black marked with white flecks dorsally. The
eye is chestnut brown.

COLORATION IN ALCOHOL. The dorsal surfaces of
the head and trunk are dark gray. This color gives way to
lighter gray and then to brown on the tail. Black flecks are
superimposed on the brown at the tail tip. The arms and legs
are dark gray with some light flecks. The lips, chin, throat,
and undersides of the legs are dark gray with light gray flecks.
The ventral surface of the trunk is unmarked dark gray. The
ventral surface of the tail lightens distally to brown inter-
rupted by darker flecks. The palmar surfaces and the tips of
the nasal cirri are pale gray.

HABITAT. This species lives in an area of very humid
cloud forest that received 5 to 6 meters of rainfall annually.
All specimens were taken within 2 km of the type locality
and were found in direct contact with the substratum under
cover objects. Sympatric with Bradytriton were three species
of Bolitoglossa, one species of Nyctano/is, various hylid and
leptodactylid frogs, lizards of the genera Anolis, Sce/oporus,
and Lepidophyma, and one member of the snake genus Lep-
todeira (Elias, in press).

OSTEOLOGY AND RELATED MORPHOLOGY. In-
formation has been derived from two cleared and stained
male specimens, from radiographs of the entire sample, and
from histological sections of the head and neck of one male.

The skull (Fig. 5) is short and very broad. The facial portion
of the skull is poorly developed and is little expanded in front
of the eyes. The anterior cranial elements are surprisingly
weak for a moderate-sized species, and the bones are highly
variable in shape, position, and number from one individual
to the next, and even from one side to the other in the same
individual. The premaxillary is relatively broad and has a
large pars dentalis. However, the bone virtually “floats” at
the anterior end of the skull; in most individuals, it does not
contact the maxillaries, and in some it is separated from those
bones by a sizeable gap. Even when a contact exists, the
articulation is tenuous. The palatal portions of the premax-
illary are greatly reduced or absent and do not contact the
anterior extensions of the vomers. The frontal processes arise
separately from the pars dentalis and are separated for their
whole length. They are very narrow and columnar basally
but become markedly compressed vertically to form the lat-
eral margins of the gland-filled internasal fontanelle. The
processes are generally divergent for their entire length and
become flattened and slightly expanded near their distal tips.
These tips overlap the expanded facial portions of the frontals
in relatively firm (for this species) articulations. The frontal

Figure 6. Dorsal view of the hyobranchial apparatus of an adult
male (MVZ 173063) Bradytriton silus new genus and species. The
entire structure is cartilaginous. The paired ceratohyals ordinarily
lie closer together but have been moved somewhat laterally for clarity
of illustration.

processes terminate distinctly posterior to the posterior tip
of the nasals, a little behind the osseous anterior margin of
the orbit. The mternasal fontanelle is relatively narrow, ex-
cept near its posterior terminus. In three adult males the
nasals are strongly protuberant, extending well anterior to
the jaw outline, but they are only slightly to moderately pro-
tuberant (Fig. 5) in the other males. The nasals protrude
slightly in two females, including the holotype. Nasals are
very irregular in size, shape, and degree of ossification but

Contributions in Science, Number 345

Wake and Elias: Tropical Salamanders 7

tend to be roughly triangular to quadrangular. The posterior
tip is poorly defined, and the anterior margin is very irregular.
Tiny fragments of disconnected bone frequently occur around
the anterior and lateral margins of the nasal. In one cleared
and stained individual, one of these fragments is very large
and is situated as if it were an entirely separate bone inter-
calated between the nasal and the prefrontal (Fig. 5). This
fragment is only slightly smaller than the prefrontal. The
nasals only slightly overlap the anterior parts of the frontals.
They have a variable degree of contact with the maxillary
facial processes but a very narrow contact with the prefron-
tals. The posterolateral margin of the nasals may be slightly
evacuated for the passage of the nasolacrimal duct, which
passes through a shallow but distinct channel in the lateral
part of the prefrontal and enters the nasal capsule through a
foramen in the anterior margin of the prefrontal. The fora-
men may form a half-circle in the prefrontal. The prefrontals
are about one-third the size of the nasals. Some small, bony
fragments are present in some individuals at the anteroven-
tral end of the prefrontals. The prefrontals are relatively long
and narrow and are extensively overlapped by the facial pro-
cesses of the maxillary. The prefrontals overlap the frontals
only slightly. The maxillaries are relatively short and straight
with narrow, pointed anterior and posterior projections on
the dental portion. The posterior tips extend nearly to the
posterior margin of each eyeball. Maxillary teeth are bicuspid
and moderate in size. They are borne along all but the tip of
the dental portion of the maxilla. The relatively well-devel-
oped palatal portion of the maxillaries is separated by a strong
articulation from the lateral-most part of the vomerine body.
There are no septomaxillary bones.

The large, morphologically specialized premaxillary teeth
of males are attached to the premaxillary in such a way that
their elongate crowns project directly anteriorly and pierce
the lip (Fig. 2). Once through the lip, the unicuspid (appar-
ently only the lingual cusp is retained) makes a sharp 90°
turn caudad before terminating in a point. When the mouth
is closed, the tooth tip is very close to the opening of the
mental gland. This gland, which has a restricted exit at the
mandibular symphysis, is a cluster of tubules converging to
form a small cluster of apertures (Fig. 2). The glands and
teeth constitute a functional complex that serves to transfer
the glandular secretion to females during courtship in a man-
ner analogous to vaccination (Arnold, 1977). Teeth of similar
shape occur in some species of Pseudoeurycea (Taylor, 1941).

The large, strong vomers are in limited contact only at
their extreme tips, posterior to the large intervomerine fon-
tanelle. A distinct, narrow process at the anterior end of the
vomer extends toward the premaxillary but does not touch
that bone; rather, it is appressed against the ventral surface
of the nasal capsule and follows that structure to curve up-
wards at its anterior tip. Immediately medial and anterior
to the internal naris, the body of the vomer is strengthened
by a dorsal bony ridge that is apparently unique among boli-
toglossine salamanders. The large, stout, but relatively short,
preorbital process falls far short of the lateral margin of the
vomerine body. The process is blunt tipped, and it has a
dorsal dimension that is very unusual in that it seems to

become continuous with the antorbital cartilage. The vo-
merine teeth are borne in a single curving row along the base
of the vomer body and nearly to the tip of the preorbital
process. The bicuspid teeth are about the size of the maxillary
teeth.

Frontals are well developed and articulate firmly with each
other middorsally, except at their anterior end. The posterior
part of each bone is a large, broad lobe that broadly overlaps
the parietal. A small, anterior lobe of the parietal abuts the
lateral margins of the posterior lobe of the frontal but does
not significantly overlap the frontal. The facial portion of the
frontal is rather poorly developed. Anteriorly, the frontal is
drawn into a narrow point lying ventral or lateral to the
frontal processes of the premaxillary.

The parietals are large bones, firmly articulated to each
other and to all surrounding bones. The stout, well-developed
lateral spurs extend anteroventrally, overlapping the cartilag-
inous insertion of the ascending process of the palatoquadrate
cartilage into the braincase. The semicircular canals produce
prominent bulges in the relatively large otic capsules. A small
but prominent spinous process is present at the extreme lat-
eral margin of each capsule. A fibrous mass of connective
tissue extends from this process to the quadrate. The squa-
mosal lies in a distinct depression in the lateral wall of the
capsule. The large parasphenoid is relatively broad and blunt-
tipped anteriorly; the orbitosphenoids are relatively widely
separated. Posterior vomerine teeth are in two bilateral patches
that diverge posteriorly. The patches in two males contain
36-39 and 44-48 small, bicuspid teeth, respectively. The
operculum has no stilus, although a slight protrusion is pres-
ent. The well-developed quadrate is attached to the otic cap-
sules by relatively large, strong squamosals. A small process
extends posteriorly from the squamosal and is connected
by a ligament to the proximal tip of the ceratohyal.

The lower jaw is rather weak. The prearticular is relatively
large, but low, and the dentary is very slender. The arc of
the lower jaw is relatively flat and broad.

The hyobranchial apparatus is typical of that of generalized
members of the supergenus Bolitoglossa in being cartilagi-
nous and lacking a urohyal (Fig. 6). The rather slender cer-
atohyals have a discrete and well-developed medial process.
The very narrow anterior process is drawn into a point that
extends upward into the sublingual fold. The long and slender
epibranchials are slightly more than twice the length of the
basibranchial. The basibranchial is slightly less than twice
the length of the second ceratobranchial. The second cera-
tobranchials are much stouter than the slender first cerato-
branchials. The first ceratobranchials are relatively well
developed in comparison with those of other tropical pleth-
odontids. The radii of the first basibranchial are continuous
with the main part of the element, and there is no sign of a
fibrocartilage joint. The broad-based and relatively short ra-
dii are joined by a distinct connective tissue strand. The most
distinctive feature of the entire hyobranchial apparatus is the
very well-developed anterior projection that is unique among
the tropical genera (Figs. 2, 6). This broad-based and rela-
tively stout projection is apparently somewhat flexible, for
the cartilage cells at its base are surrounded by less intercel-

8 Contributions in Science, Number 345

Wake and Elias: Tropical Salamanders

lular matrix than are the cells either proximal or distal to it.
This process apparently represents a primitive condition and
is reminiscent of the process seen in Ensatina (Lombard and
Wake, 1977), although the process in Ensatina is less con-
tinuous with the basibranchial. The anterior basibranchial
process is clearly a less-derived homologue of the lingual
cartilage that is present in many tropical plethodontids.

The vertebral column consists of relatively stout vertebrae,
with well-developed centra. There are one cervical, fourteen
trunk, one sacral, two caudosacral, and a varying number of
caudal vertebrae (from 20 to 26 in individuals having com-
plete tails). The atlas has no special features, but the neural
arch is barely completed in both cleared and stained adults.
The first trunk vertebra is distinctly shorter than the re-
maining vertebrae and has a high, well-developed neural
crest. Lower crests are present on the next few trunk verte-
brae. The centrum is large and may be largely filled with
mineralized cartilage, for only the terminal concavities are
clearly filled by unmineralized intervertebral cartilage. All
trunk vertebrae have well-developed, separated, pointed pro-
cesses (hyperapophyses) on the posterior border of the neural
arch. The nerve route pattern is typical of bolitoglossine sal-
amanders (Edwards, 1976): the first trunk vertebra has a
single foramen in front of the transverse processes, the second
has single foramina both in front of and behind the transverse
processes, and the third and succeeding vertebrae (to the end
of the tail) have single foramina behind the transverse pro-
cesses. The transverse processes of the trunk vertebrae are
well developed. The dorsal and ventral rib bearers are sep-
arated for their entire lengths. The long, relatively straight
processes extend beyond the lateral margins of the zygapoph-
yses. They are sharply angled posteriorly in the first few
vertebrae but are nearly perpendicular to the column over
most of its length. The dorsal bearer is immediately dorsal
to the ventral bearer, but it tends to be slightly more pos-
teriorly oriented. Moderately long ribs with distinctly sepa-
rated heads are present on all but the last trunk vertebra.
Transverse processes of the first caudosacral vertebra are
stout and blunt-ended and are swept somewhat posteriorly.
Transverse processes of the second caudosacral vertebra are
short, stout, and blunt-ended; they are nearly perpendicular
to the column or are slightly anteriorly oriented. The first
caudal vertebra is short and is associated with a distinctly
constricted tail base. The relatively short transverse processes
located at the extreme anterior end of the centrum are sharply
oriented in an anterior direction. The transverse processes
of succeeding caudal vertebrae are progressively shorter and
barely exceed the anterior zygapophyses in length; they are
borne on the extreme anterior end of each vertebra, at the
base of the zygapophyses. The sharply tapered tail is deep as
a result of a thick glandular layer lying dorsal to the vertebral
column but is relatively narrow. Caudal vertebrae generally
lack neural crests, although there may be low and irregularly
formed crests on the first one or two vertebrae; however, the
caudal vertebrae have very well-developed hyperapophyses.

The small hands and feet are highly distinctive (Fig. 4).
They are very reduced in size and have relatively little ossified
tissue. The digits are fused to one another for most of their

lengths but are free at their tips. Indeed, the digits are so
poorly developed that it is somewhat artificial to present a
phalangeal formula. Some phalanges are entirely cartilagi-
nous, others may have a tiny speck of ossified tissue, and
only the proximal phalanx of the longest finger and toe is
consistently well ossified. An unusual feature is the cartilag-
inous tip of many terminal phalanges. In other plethodontid
salamanders, these tips are well ossified and often specialized
in structure. The poor degree of development suggests that
Bradytriton does not use its limbs for specialized locomotory
activity. A few individuals can be scored as having a pha-
langeal formula of 1, 2, 3, 2, for the hand and 1, 2, 3, 3, 2,
for the foot, based on the presence of tiny specks of radio-
opaque material visible in radiographs. Even the metapodial
elements, which are cylindrical in shape, are small and poorly
ossified, and the first digit of the hand has an especially small
element. As a result of the fusion of the ulnare and inter-
medium, there are seven carpals; similarly, there are eight
tarsals as a result of the fusion of distal tarsals 4 and 5. A
distinct spur projects from the shank of the tibia for nearly
its entire length, and there is a low but distinct tibial crest.

RANGE. Bradytriton silus is known only from the im-
mediate vicinity of the type locality on the eastern slopes of
the Sierra de los Cuchumatanes in extreme northwestern
Guatemala.

ETYMOLOGY. From silus, L. for “pug-nosed” referring
to the characteristic truncated appearance of the snout.

GENERIC SYNOPSES

Bolitoglossa Dumeril, Bibron,
and Dumeril

Oedipus Tschudi
Eladinea Miranda-Ribeiro
Magnadigita Taylor
Palmatotnton Smith

TYPE SPECIES. Bolitoglossa mexicana Dumeril, Bi-
bron, and Dumeril.

DIAGNOSTIC CHARACTERS. Small to very large sal-
amanders with partially to fully webbed hands and feet, dis-
tinguished from members of all other tropical salamander
genera in lacking a sublingual fold.

REFERRED SPECIES. Bolitoglossa adspersa (Peters); B.
altamazonica (Cope); B. alvaradoi Taylor; B. arborescandens
Taylor; B. biseriata Tanner; B. borburata Trapido; B. capi-
tana Brame and Wake; B. cerroensis Taylor; B. chica Brame
and Wake; B. colonnea (Dunn); B. compacta Wake, Brame,
and Duellman; B. cuchumatana (Stuart); B. cuna Wake,
Brame, and Duellman; B. dofleini (Werner); B. dunni
(Schmidt); B. engelhardti (Schmidt); B. epimela Wake and
Brame; B. equatoriana Brame and Wake; B. flavimembris
(Schmidt); B. flaviventris (Schmidt); B. franklini (Schmidt);
B. hartwegi Wake and Brame; B. helmrichi (Schmidt); B.
hypacra (Brame and Wake); B. hgnicolor ( Peters); B. lincolni
McCoy and Walker; B. macrinii (Lafrentz); B. marmorea
(Tanner and Brame); B. medemi Brame and Wake; B. me-
liana Wake and Lynch; B. mexicana Dumeril, Bibron and

Contributions in Science, Number 345

Wake and Elias: Tropical Salamanders 9

Figure 7. Feet of Chiropterotriton and Pseudoeurycea.

A. Left foot of an adult Chiropterotriton multidentatus (39.6 nint SL) from Hidalgo, Mexico. Cartilage is stippled. Note that distal tarsal 5
is larger than distal tarsal 4 and articulates with the centrale. This arrangement is unique to Chiropterotriton within the supergenus Bolitoglossa.

B. Left foot of an adult Pseudoeurycea leprosa (53.9 mm SL) from Veracruz, Mexico. Cartilage is stippled. Note that distal tarsal 5 is smaller
than distal tarsal 4 and that it does not articulate with the centrale; this is the primitive pattern in plethodontid salamanders. The foot of this
species is relatively much smaller than that of C. multidentatus, for although the feet as illustrated are about the same size, the specimen of
Pseudoeurycea is much larger. Note that the fifth toe is smaller in Pseudoeurycea than in Chiropterotriton.

Dumeril; B. minutula Wake, Brame, and Duellman; B. mo-
no (Cope); B. mu/leri (Brocchi); B. nicefori Brame and Wake;
B. occidentalis Taylor; B. odonnelli (Stuart); B. orestes Brame
and Wake; B. palmata (Werner); B. pandi Brame and Wake;
B. peruviana (Boulenger); B. phalarosoma Wake and Brame;
B. platydacty/a (Gray); B. ramosi Brame and Wake; B. re-
splendent McCoy and Walker; B. riletti Holman; B. robusta
(Cope); B. rostrata (Brocchi); B. rufescens (Cope); B. salvinii
(Gray); B. savagei (Brame and Wake); B. schizodactyla Wake
and Brame; B. schmidti (Dunn); B. silverstonei Brame and
Wake; B. sima (Vaillant); B. sooyorum Vial; B. striatula (No-
ble); B. stuarti Wake and Brame; B. subpalmata (Boulenger);
B. taylori Wake, Brame, and Myers; B. vallecula Brame and
Wake; B. veracrucis Taylor; B. walkeri Brame and Wake; B.
yucatana (Peters).

RANGE. Bolitoglossa has by far the widest range of any
of the tropical salamander genera. It occurs almost contin-
uously from the lowlands of eastern San Luis Potosi, Mexico,

in the north, south to the Amazonian lowlands of southern
Peru, the mountains near Cochabamba, Bolivia, and south-
ern Minas Gerais, Brazil.

COMMENT. Many species have been added to this large
genus during the past two decades, but no revisionary study
has been undertaken. A number of undescribed species are
known to us. We here place Bolitoglossa omniumsanctorum
in the synonymy of B. morio on the basis of close similarity
of the type specimens of B. omniumsanctorum to specimens
of B. morio in size, form, and coloration.

Wake and Brame (1969) and Wake and Lynch (1976) dis-
cussed a possible subdivision of this large, cumbersome ge-
nus. Wake and Lynch (1976) referred to “alpha” and “beta”
sister groups and suggested that these might eventually be
recognized as separate genera. The beta group is defined on
the basis of an apomorphic character, a tail base specializa-
tion (Wake and Dresner, 1967). Members of the alpha group
frequently have osteological reductions and foot specializa-

10 Contributions in Science, Number 345

Wake and Elias: Tropical Salamanders

tions, but we know of no apomorphic character or combi-
nation of characters that would unambiguously define the
alpha group. These diagnostic problems, combined with the
absence of marked ecological or phenetic differentiation be-
tween the two groups, lead us to postpone formal division.

Bradytriton new genus

TYPE SPECIES. Bradytriton si/us new species.

DIAGNOSTIC CHARACTERS. This is a short-snouted,
stout-bodied salamander with small limbs and small, syn-
dactylous hands and feet. It differs from all other tropical
salamander genera in having a laterally compressed tail with
a well-developed dorsal glandular ridge.

REFERRED SPECIES. Bradytriton situs new species.

RANGE. The unique species is known only from the im-
mediate vicinity of the type locality on the northeastern slopes
of the Sierra de los Cuchumatanes in northwestern Guate-
mala.

Chiropterotriton Taylor

TYPE SPECIES. Oedipus multident atus Taylor.

DIAGNOSTIC CHARACTERS. Small to moderately
large salamanders with relatively broad, partially webbed
hands and feet and broad-tipped fingers and toes. Chirop-
terotriton differs from all other tropical salamander genera
in its arrangement of tarsal cartilages: in Chiropterotriton,
distal tarsal five is larger than distal tarsal four and articulates
with the centrale; in other genera, four is larger than five,
and five is excluded from articulation with the centrale (Fig.
7).

REFERRED SPECIES. Chiropterotriton arboreus (Tay-
lor); C. chiropterus (Cope); C. chondrostega (Taylor); C. di-
midiatus (Taylor); C. lavae (Taylor); C. magnipes Rabb; C.
mosaueri (Woodall); C. multidentatus (Taylor); C. prisons
Rabb.

RANGE. The genus is restricted to eastern Mexico, from
west-central Tamaulipas in the north to the mountains of
northern Oaxaca in the south.

COMMENT. Most species of this genus have been stud-
ied in some detail by Rabb (1955, 1958, 1965). The major
unstudied unit is that comprising C. chiropterus and C. lavae.

Dendrotriton new genus

TYPE SPECIES. Oedipus bromeliacia Schmidt.

DIAGNOSIS. Small, slender, long-tailed, arboreal sala-
manders with long legs, broad hands and feet, and long,
broad-tipped digits. Dendrotriton differs from Oedipina by
its short trunk (14 vs. 18 or more trunk vertebrae) and long
limbs; from Nyctanolis in having a single premaxillary; from
Chiropterotriton in having a larger fourth than fifth tarsal;
from Bolitog/ossa in possessing a sublingual fold; from Bra-
dytriton, Nototriton, Parvimolge, Oedipina and Thorius by
its lack of carpal and tarsal fusions; from Lineatriton by its
short vertebrae and long legs; and from Pseudoeurycea by its
lack of prefrontals and tibial spurs.

REFERRED SPECIES. Chiropterotriton bromeliacia

(Schmidt); C. cuchumatanus Lynch and Wake; C. mega-
rhinus Rabb; C. rabbi Lynch and Wake; C. xolocalcae (Tay-
lor).

ETYMOLOGY. From dendron, Gr. for tree, referring to
the arboreal habits of all known species, and triton, Gr., a
commonly used term for salamanders.

RANGE. Southwestern Chiapas, Mexico, through west-
ern Guatemala.

COMMENT. This group has recently been studied in de-
tail by Lynch and Wake (1975), who presented photographs
of all five species.

Lineatriton Tanner

TYPE SPECIES. Spelerpes lineolus Cope.

DIAGNOSTIC CHARACTERS. A very slender, elon-
gate, fossorial form with diminutive limbs, hands and feet,
and an extremely long tail. Lineatriton is distinguished from
all other tropical salamander genera in having very elongated,
narrow vertebrae. Oedipina has a similar body form and
ecology but differs from Lineatriton in having 18 or more,
rather than 14, trunk vertebrae.

REFERRED SPECIES. Lineatriton lineola (Cope).

RANGE. Lineatriton is restricted to two disjunct areas of
the Gulf slope of Mexico: along the lower slopes of the moun-
tains of west-central Veracruz and the area around Los Tux-
tlas in eastern Veracruz.

COMMENT. This is one of the most distinctive of the
tropical genera and is apparently monotypic. It has been little
studied since the work of Tanner (1950).

Nototriton new genus

TYPE SPECIES. Spelerpes picadoi Stejneger.

DIAGNOSIS. Small, long-tailed salamanders of arboreal,
terrestrial, or semifossorial habitats. The species have mod-
erately long to short legs and small hands and feet. The in-
termedium and ulnare of the manus and distal tarsals four
and five of the pes are fused. The genus is distinguished from
other genera with similar fusions as follows: from Oedipina
by having only 14 rather than 18 or more trunk vertebrae;
from Bradytriton by having a long, slender tail and slender
body; from Parvimolge by having the tail longer than the
body, by having frontal processes fused together at their base
(all species but richardi), and by lacking mesopodia! and
hyobranchial mineralizations; from Thorius by having a
complete skull roof over the brain case; from Bo/itoglossa
by having carpal fusions and a sublingual fold.

REFERRED SPECIES. Chiropterotriton barbouri
(Schmidt); Bo/itoglossa diminuta Robinson; Chiropterotriton
nasalis (Dunn); C. picadoi (Stejneger); C. richardi (Taylor);
C. veraepacis Lynch and Wake.

ETYMOLOGY. From notos, L., referring to the southerly
distribution of the genus, and triton, L., Gr., a commonly
used term for salamanders.

RANGE. Nototriton ranges from eastern Guatemala to the
Meseta Central of Costa Rica.

COMMENT. Most of Nototriton consists of what has been

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Wake and Elias: Tropical Salamanders 11

called the picadoi group (Wake and Lynch, 1976) or the
nasa/is group (Lynch and Wake, 1978) of Chiropterotriton.
The species richardi may not be referable to this genus; it
was originally described as a member of the genus Parvimolge
and has been considered problematic by Rabb (1955), Wake
(1966), Wake and Lynch (1976), and Lynch and Wake (1978).
It is more appropriately placed in Nototriton than elsewhere,
but it differs from other members of the genus in possessing
an unusual combination of ancestral and derived characters
(Lynch and Wake, 1978). Table 2 was constructed by ignoring

N. richardi, but we have nonetheless tentatively assigned the
species to this genus because it does not come as close to
fitting into any other genus. Further, we are reluctant to es-
tablish a new genus for such a poorly known form.

The species described as Bolitoglossa diminuta by Rob-
inson (1976) is also most appropriately placed in Nototriton.
Radiographs of the holotype reveal that it has frontal pro-
cesses of the premaxillary that are fused, and it has a very
long tail for its small body size (53% of total length). It was
reported to lack a sublingual fold, and we have been unable
to see one; however, the holotype, a mature female, is very
small (31.1 mm SL), and it may be impossible to determine
the status of this character without histological sections. No-
totriton richardi has a very reduced sublingual fold. It was
on the basis of the apparent absence of this fold in the ho-
lotype of B. diminuta that the senior author urged Robinson
to describe the species as a Bolitoglossa, even though Rob-
inson had originally intended to place it in the genus Chi-
ropterotriton.

Nyctano/is Elias and Wake

TYPE SPECIES. Nyctanolis pernix Elias and Wake.

DIAGNOSTIC CHARACTERS. This is a large, long-
legged, long-fingered, long-toed, long-tailed taxon that differs
from all other tropical salamander genera in having paired
premaxillary bones.

REFERRED SPECIES. Nyctanolis pernix Elias and
Wake.

RANGE. The unique species is known from the north-
eastern slopes of the Sierra de los Cuchumatanes in NW
Guatemala; near the Lagos de Montebello in nearby Chiapas,
Mexico; and from the Sierra de las Minas in eastern Gua-
temala.

Oedipina Keferstein

Oedipina Keferstein
Ophiobatrachus Gray
? Haptoglossa Cope
Oedopinola Hilton

TYPE SPECIES. Oedipina uniformis Keferstein.

DIAGNOSTIC CHARACTERS. Slender, elongate sala-
manders of moderate to large size with very small limbs,
hands, and feet, and a very long tail. Oedipina is distinguished
from all other tropical salamander genera, including Linea-
triton, the only genus that resembles it externally, by having
18 to 22 (rather than 14) trunk vertebrae.

REFERRED SPECIES. Oedipina alfaroi Dunn; O. al-
tura Brame; O. carab/anca Brame; O. col/aris (Stejneger); O.
complex (D\inn)\ O. cyclocauda Taylor; O. e/ongata (Schmidt);

O. grandis Brame and Duellman; O. ignea Stuart; O. par-
vipes (Peters); O. paucidentata Brame; O. poelzi Brame; O.
pseudouniformis Brame; O. stuarti Brame; O. taylori Stuart;

O. uniformis Keferstein.

RANGE. Oedipina ranges from south-central Chiapas,
Mexico, through all of Central America and western Colom-
bia to northwestern Ecuador.

COMMENT. This is an exceptionally well-defined genus.
Since the revision by Brame (1968), only one additional
species, O. grandis, has been described, and there have been
no other taxonomic changes.

Parvimolge Taylor

TYPE SPECIES. Oedipus townsendi Dunn.

DIAGNOSTIC CHARACTERS. Diminutive salaman-
ders with relatively stocky bodies, short legs with small hands
and feet, and short, stout tails. It is distinguished from other
genera with small species as follows: from Bolitoglossa in
having a sublingual fold; from Dendrotriton and Chiroptero-
triton in having distal tarsals four and five fused and the
intermedium and ulnare fused; from Nototriton in having a
short tail and lacking fused frontal processes of the premax-
illary; from Thorius in having a complete skull roof over the
braincase.

REFERRED SPECIES. Parvimolge townsendi (Dunn).

COMMENT. Two other species, praecellens and rich-
ardi, have been included in the genus. Neither seems to be
closely related to townsendi, and we here refer them to the
genera Pseudoeurycea and Nototriton, respectively.

Pseudoeurycea Taylor

TYPE SPECIES. Spelerpes leprosus Cope.

DIAGNOSTIC CHARACTERS. A diverse group of
moderate-sized to very large salamanders with moderate to
long limbs, well-developed hands and feet in which the mid-
dle digits are markedly longer than the outer ones (Fig. 7),
and a tail about as long as the body. The species are very
generalized in morphology. The genus differs from Nyctanolis
in having a single premaxillary and shorter limbs and tail;
from Dendrotriton in having prefrontals and tibial spurs;
from Bolitoglossa in having a sublingual fold; from Chirop-
terotriton in having a fourth distal tarsal that articulates with
the fibulare and is larger than the fifth (Fig. 7); from all other
genera in having separate distal tarsals four and five.

REFERRED SPECIES. Pseudoeurycea altamontana
(Taylor); P. anitae Bogert; P. bellii (Gray); P. brunnata
Bumzahem and Smith; P. cephalica (Cope); P. cochranae
(Taylor); P. conanti Bogert; P. exspectata Stuart; P.firscheini
Shannon and Werler; P. gadovii (Dunn); P. ga/eanae (Tay-
lor); P. goebeli (Schmidt); P.juarezi Regal; P. leprosa (Cope);

P. melanomolga (Taylor); P. mystax Bogert; P. nigroma-
culata (Taylor); P. praecellens (Rabb); P. rex (Dunn); P. rob-
ertsi (Taylor); P. scandens Walker; P. smithi (Taylor); P.
unguidentis Smith and Taylor; P. werleri Darling and Smith.

12 Contributions in Science, Number 345

Wake and Elias: Tropical Salamanders

RANGE. The genus ranges from eastern Sonora, Mexico,
in the northwest, and west-central Tamaulipas, Mexico, in
the northeast, through most of eastern and south-central
Mexico into western and southern Guatemala.

COMMENT. Few characters bind this assemblage ofgen-
eralized species together. The genus lacks any autapomor-
phies and is mainly defined by exclusion. A few species stand
out from the rest as distinct morphological entities (e.g., P.
bellii, P. praecel/ens, P. werleri). The gadovii, rex, and leprosa
groups are the core of the genus and appear to be closely
interrelated (Maxson and Wake, 1981). We here assign prae-
cel/ens to Pseudoeurycea. Rabb (1955) placed it in the genus
Parvimolge with some hesitation. It differs from P. townsendi
in that it is larger and stouter and has prefrontal bones. Rabb
(1955) reported that P. praecel/ens had some mineralization
of the hyobranchial apparatus, as in P. townsendi ; however,
we have been unable to see this feature in our radiographs
of the unique holotype. The species fits reasonably well in
Pseudoeurycea, and Rabb suggested that it might prove to
be a relative of P. cephalica.

Thor ius Cope

TYPE SPECIES. Thorius pennatulus Cope.

DIAGNOSTIC CHARACTERS. Diminutive, slender
salamanders with short limbs and small hands and feet. Tho-
rius is distinguished from all other tropical salamanders by
two characters: the suborbital groove intersects the lip, and
the skull is incompletely ossified, leaving the braincase un-
covered by bone (the parietal bones have an especially large
gap separating them).

REFERRED SPECIES. Thorius duhitus Taylor; T. mac-
dougalli Taylor; T. maxil/abrochus Gehlbach; T. minutissi-
mus Taylor; T. narisovalis Taylor; T. pennatulus Cope; T.
pulmonaris Taylor; T. schmidti Gehlbach; T. troglodytes
Taylor.

RANGE. The genus ranges from the eastern margins of
the Mexican Plateau into the mountains of northern and
central Oaxaca, and in the Sierra Madre del Sur of southern
Oaxaca and Guerrero, Mexico.

COMMENT. The genus has recently been studied in de-
tail by Freeman (1977) and Hanken (1980), but the taxo-
nomic results of these unpublished theses are not yet avail-
able. All of the described species (except perhaps T.
maxillabrochus) are valid, but there are numbers of addi-
tional undescribed species.

DISCUSSION

The supergenus Bolitoglossa was redefined by Elias and Wake
(1983) on the basis of hyobranchial and tail base character-
istics. All genera considered here have the features of the
supergenus. Our main departure from Wake (1966) is that
two formerly diagnostic features are deemphasized. First,
Nyctanolis has two premaxillary bones; thus, the supergenus
Bolitoglossa can no longer be characterized by the presence
of a single, fused bone. Second, Nyctanolis and some species
of Pseudoeurycea have a lateral parietal spur that is only

poorly developed: in Nyctanolis the spur may be little more
than a low ridge on the underside of the bone, and in some
Pseudoeurycea the “spur” is really only a small lateral lobe.
Still, at least some form of parietal “spur” is present in all
members of the supergenera Batrachoseps and Bolitoglossa.

With the new genera, the supergenus Bolitoglossa includes
1 1 genera. Nyctanolis, Bradytriton, Parvimolge, and Linea-
triton are monotypic; the other genera include from five (Den-
drotriton) to over sixty (Bolitoglossa) species. Additional un-
described species that are known to us will increase the number
of species in such large genera as Bolitoglossa, Pseudoeurycea,
and Thorius but will not affect the monotypic genera.

The genera Bolitoglossa, Lineatriton, Oedipina, Pseudoeu-
rycea, and Thorius are essentially unchanged by our analysis.
The content of Parvimolge is reduced to a single species by
transfer of the species praecel/ens to Pseudoeurycea. The ge-
nus most strongly affected is Chiropterotriton. As a result of
the shift of the old bromeliacia species group (Lynch and
Wake, 1975) to the new genus Dendrotriton and of the old
picadoi( Wake and Lynch, 1 976) or nasalis (Lynch and Wake,

1 978) species group to the new genus Nototriton, Chiroptero-
triton has been reduced to a group of nine species that inhabit
eastern Mexico. The species formerly known as Bolitoglossa
diminuta is included in Nototriton. The other two new genera,
Nyctanolis and Bradytriton, are based on newly discovered
species.

Hecht and Edwards (1976a, 1 976b) and Hecht (1976) dis-
cuss the relative information content of different types of
shared derived character states (synapomorphies) for the re-
construction of phylogeny. Ordered by increasing significance
(basically, the degree of confidence one has in their inter-
pretation) in phylogenetic reconstruction are five classes of
synapomorphies:

“(I) Shared and derived character states which are the result
of loss. Such loss characters are the zero state because
there is no developmental evidence.

(II) Shared and derived character states which are the results
of simplification or reduction of complex structures as
indicated by comparative or developmental anatomy.

(III) Shared and derived character states which are the result
of growth and developmental processes dependent on
size, age, and hormonal and other physiological rela-
tionships. Characters related to allometric functions are
the best examples.

(IV) Shared and derived character states which are highly
integrated functionally and are subject to directional se-
lection.

(V) Shared and derived character states which are unique
and innovative in structure (the most important type of
information).” (Lrom Hecht and Edwards, 1976a.)

Our analysis in this paper is based on the 18 characters out-
lined earlier. The apomorphic characters we have chosen
include seven that are autapomorphic (Table 1). These are
not useful in phylogenetic analysis but are included because
they aid in diagnosing genera. The 1 1 remaining characters
vary in degree of utility. Six involve morphological trans-
formation, and five involve losses of elements. We interpret

Contributions in Science, Number 345

Wake and Elias: Tropical Salamanders 13

our loss characters as falling into the third category of Hecht
and Edwards (1976) rather than the first, for we have some
ontogenetic information concerning the way in which four
of the five characters are lost. We believe that loss of septo-
maxillaries, prefrontals, preorbital processes of vomers, and
columellae (or stylar processes of the opercula) are all pae-
domorphic characters (Lynch and Wake, 1978; Alberch et
al., 1979; Alberch and Alberch, 1981; Elias, in press), but
whether they should all be counted as independent characters
or considered to be parts of a single “global” paedomorphic
process is unresolved.

The six remaining characters ( 1, 3, 4, 5, 10, 1 1 ) all involve
transformations. Characters 10 and 1 1 relate to ontogenetic
phenomena and fall into Hecht and Edwards's third category.
Transformation characters 1, 3, 4, and 5 are parts of inte-
grated, functionally significant systems; they fall in the fourth
category of Hecht and Edwards.

Other authors have used additional characters in generic
diagnoses of tropical salamanders. Some of these characters
(e.g., a posteriorly directed spur on the squamosal in Thorius)
are autapomorphies and of little use in phylogenetic analysis.
Others occur in all members of one genus but also appear
irregularly in one or a few species in some other genera (e.g.,
the complex tail base present in some Pseudoeurycea). Other
characters, such as the shape and degree of ossification of the
intervertebral articulation (used by Taylor, 1 944), are subject
to extreme ontogenetic variation (Wake, 1 970), and we have
not had sufficient material to perform a complete analysis.
Still other characters, such as external shape of hands and
feet, shape of terminal digits, length of ribs, and fusion or
nonfusion of transverse processes, have proven impossible
to codify consistently for all neotropical species, but these
may well have utility for liner analysis of intra- and inter-
generic relationships in the future. The hyobranchial appa-
ratus (features of the tip of the first basibranchial, including
length and shape of radii and detailed structure of the lingual
cartilage and associated musculature) holds much promise
as a source of additional characters (Tanner, 1952; Lombard
and Wake, 1977). However, this structure requires detailed
histological study, and we have examined too few species to
discern broad patterns. Future studies should involve com-
parative ontogenetic analyses, for heterochronic modes of
evolution have been strongly implicated in salamanders
(Wake, 1966; Alberch et al., 1979; Alberch and Alberch,
1981).

Biochemical, immunological, and cytological characters are
useful in analyzing phylogenetic relationships in the super-
genus Bo/itoglossa. Recently Maxson and Wake (1981) used
immunological approaches to study albumin evolution in
Pseudoeurycea and Chiropterotriton (sensu lato). Extensive
evolution of albumin has taken place in these groups, and
immunological methods will be most useful at the intrage-
neric level of comparison. In reference to the present generic
review, the principal result of interest from the immunolog-
ical work is the finding that Chiropterotriton (antisera from
C. multidentatus) and Dendrotriton (antisera from D. bro-
meliacia) are more similar to Pseudoeurycea (antisera from
five species) than they are to each other. This finding cor-

roborates our conclusion that the two genera are distinct. No
antisera were prepared for species of Nototriton, but one-way
tests were made to some species of the genus. Immunological
distances between species of Chiropterotriton and Dendro-
triton are about equivalent to those between Chiropterotriton
and Nototriton, but immunological distances between Den-
drotriton and Nototriton are considerably less. Distances
within Dendrotriton (maximum 21) are much less than be-
tween Dendrotriton and Nototriton (minimum 64).

Some data on chromosomes are available (Leon and Kezer,
1978; J. Kezer and S. Sessions, pers. comm.). All members
of the supergenus Bo/itoglossa so far examined have a haploid
chromosome number of 13, as do members of the genus
Batrachoseps. All other plethodontids examined have a hap-
loid number of 14. The species of Dendrotriton (D. brome-
liacia, D. cuchumatanus, D. rabbi ) and Nototriton ( N .
picadoi, N. veraepacis) so far examined have a distinct het-
eromorphism (presumably XY) in the thirteenth pair of chro-
mosomes. This pattern has been found in some species of
Thorius and Oedipina but not in Chiropterotriton, Pseu-
doeurycea, Lineatriton, Bo/itoglossa, or Parvimo/ge (chro-
mosomes of Nyctanolis and Bradytriton have not been ex-
amined). These data lend further support to our decision to
separate Dendrotriton and Nototriton from Chiropterotriton.

From the 1 8 characters analyzed we are unable to conclu-
sively demonstrate the monophyletic status of 3 of the 1 1
bolitoglossine genera: Dendrotriton, Nototriton, and Pseu-
doeurycea (Table 1 ). A genus appearing in Table 1 must have
at least one apomorphic character (a synapomorphy for the
species of the genus) that is not shared with any other given
genus in order for us to consider it to be monophyletic.

All of the apomorphies shown for Dendrotriton in Table
1 are shared with Oedipina ; nevertheless, in other characters,
which proved impossible to score for all neotropical genera,
Dendrotriton shows derived states not found in Oedipina.
Species of Dendrotriton, all of which are arboreal, show such
derived features as relatively long legs, with long, well-de-
veloped digits and expanded terminal phalanges. Oedipina,
all of which are semi- to fully fossorial, have very short legs,
with short, erratically developed digits and unspecialized
phalanges. The limb states of the two genera are derived in
opposite directions from the ancestral state. Thus, despite
the absence of defining synapomorphies in our formal tab-
ulation, we consider Dendrotriton to be a well-defined, mono-
phyletic assemblage.

The situation with Pseudoeurycea is far more complex.
This genus is something of a “waste-basket,” notwithstand-
ing arguments of Baird (1951) to the contrary. It has only a
single apomorphy (fused premaxillaries, character 1 1, Table
1 ), and this is shared with all genera of the supergenus except
Nyctanolis. All of the other genera are easily distinguished
from Pseudoeurycea on the basis of a variety of synapo-
morphies and autapomorphies. The species of Pseudoeurycea
are relatively diverse in external structure and behavior but
are basically similar in their generalized osteology, such that
there is no clear osteological basis for subdividing the genus
into monophyletic assemblages. The generotype is P. leprosa;
it and some allied species have a plesiomorphic character

14 Contributions in Science, Number 345

Wake and Elias: Tropical Salamanders

(septomaxillary bones) and an apomorphic character (a com-
plex tail base, not coded here, but rather similar to the tail
base seen in Bolitog/ossa beta. Wake and Lynch, 1976). It
may eventually be possible to formally recognize this group
taxonomically, but the remaining species in the present genus
Pseudoeurycea would still constitute a very disparate assem-
blage. Maxson and Wake (1981) have shown that species of
Pseudoeurycea are greatly divergent in albumin structure.
One group that is not well defined morphologically can be
defined reasonably well on a biochemical basis (the combined
gadovii and rex groups of Wake and Lynch, 1976, together
with some members of other groups). However, an electro-
phoretic analysis in progress (Yang and Wake, unpublished
data) discloses that most species are very distinct from one
another and that groups of species are not readily apparent.
Pseudoeurycea may be a paraphyletic taxon, the members of
which have retained relatively conservative morphologies.
In this respect, the genus presents a taxonomic problem sim-
ilar to that encountered in the genus Plethodon (Larson et
al., 1981), another taxon based largely on plesiomorphic
characteristics. We do not think that Pseudoeurycea, as pres-
ently constituted, represents the remnants of the ancestral
stock of all tropical salamanders, for in many respects mem-
bers of Nyctanolis and Chiropterotriton are more generalized
osteologically. For example, C. priscus has relatively large,
well -developed septomaxillary bones, primitive elements that
occur only occasionally in the few species of Pseudoeurycea
that have them. Thus, our character scorings must be used
cautiously when making phylogenetic interpretations.

Notototnton, although it is readily distinguished from most
genera, has no apomorphies that are not shared with Oedi-
pina. This situation is superficially similar to the problem of
Dendrotriton versus Oedipina, but the three synapomorphies
shared by Nototriton and Oedipina are a different set. Fur-
ther, unlike Dendrotriton, Nototriton may be a paraphyletic
group that includes lineages of a pr e-Oedipina stock (see
Lynch and Wake, 1978). For example, N. richardi, a poorly
known species of which fewer than ten specimens have been
collected, appears to be more primitive than other Nototriton
in some respects but more derived in others. It could be
placed in Oedipina, but it lacks an increased number of trunk
vertebrae (character 1 7), the synapomorphy that most strong-
ly distinguishes the species of Oedipina from all other tropical
plethodontids. The remaining species of Nototriton form a
well-defined assemblage (see Lynch and Wake, 1978). As
recognized here, Nototriton is a diverse group consisting of
several rare, secretive, and poorly known species. At this
time we are unable either to define autapomorphies for the
genus or to subdivide it into clearly monophyletic sublin-
eages. By joining Oedipina and Nototriton to form a single
genus, we could claim monophyly for the group as a whole,
but this would severely blur the limits of the present genus
Oedipina, one of the most tightly knit, well-diagnosed, and
highly apomorphic genera of salamanders. Species of Noto-
triton share a number of structural and ecological similarities,
and they are sharply distinguished from Oedipina by the
synapomorphies of the latter group. Accordingly, we choose
to recognize both Oedipina and Nototriton, although the lat-

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supergenus

Bolitoglossa

Figure 8. Maximum parsimony cladogram of genera in the tribe
Bolitoglossini. The numbered characters (Table 1 ) are present in
their derived condition in all taxa above the point in the dendrogram
where the lines occur. Reversals are circled. The tree treats character
9 as if it were a synapomorphy in its reversed form (i.e., resembling
the presumed primitive condition). Parallelisms are not illustrated.

ter genus may ultimately not prove to be a monophyletic
assemblage.

We used the data in Table 1 to compute Wagner trees,
using the method of cladistic inference by parsimony (Farris,
1970), but with one modification. We fixed the root of the
tree a priori by declaring Nyctanolis to be the first derivative
branch. We believe that the premaxillary character is con-
siderably more substantial than any other in the matrix (be-
cause it is conservative and relatively well understood; see
Wake, 1966), and we accordingly used the rooting method
to weight this character heavily. The resulting tree is pre-
sented in Figure 8.

There are reasons for questioning the validity of this max-
imum parsimony tree. Although there are six stems char-
acterized by synapomorphies, two of these are reversed later
in the dendrogram. Only three of the six synapomorphic

Contributions in Science, Number 345

Wake and Elias: Tropical Salamanders 15

characters (3, 4, 5) are “high-quality” characters, i.e., char-
acters that fall into Hecht and Edwards’s (1976a) category 4
and constitute parts of integrated systems. The significance
of character 5 is unclear; the long, tubular mental glands of
certain of the genera (e.g., Oedipina, Bradytriton ) are unusual
but may represent extreme cases of subtler variation that
would require histological documentation. Note that the den-
drogram requires a reversal of this character in the stem
leading to Thorius. It seems more likely that tubular glands
have evolved in parallel, a hypothesis that is supported by
the observation that some species of the unrelated North
American genus Eurycea have such glands. Characters 3 and
4 involve fusions of carpals and tarsals. Although these char-
acters may be more complex than those that involve loss of
elements, they nevertheless require little more than simple
developmental modifications. We know that character 3 has
evolved independently in extra-tropical plethodontids (e.g..
Amides hardii and some species of Batrachoseps\ Wake, 1 966).
However, among plethodontids character 4 (tarsal fusions)
appears to be unique to the genera reported here. It is possible
that distal tarsals 4 and 5 are fused in Balrachoseps, rather
than 5 being lost as Wake (1966) has suggested.

Because we have fixed the root of the tree with Nyctanolis,
character 1 1 will be a common feature of most possible den-
drograms. Characters 6 and 7 fall into Hecht and Edwards’s
third weighting category, for we have some ontogenetic in-
formation concerning them (unpublished and Elias, in press).
We think that both characters 6 and 7 reflect parallel evo-
lution, for both characters refer to features that appear rel-
atively late in the development of related species. Character
7 (loss of preorbital processes of the vomer) has evolved
independently in some species of Balrachoseps (Marlow et
al., 1979). Scoring for character 6 (loss of stylus of the oper-
culum) requires a certain degree of subjectivity, because the
process, where it appears at all, is invariably very small in
members of the supergenus Bolitoglossa.

The final significant character in Figure 8 is the presence
or absence of septomaxillary bones (character 9). Most mem-
bers of the supergenus Bolitoglossa lack septomaxillaries, and
a questionable feature of this tree is that it contains a stem
for which the synapomorphy is a postulated reversal (reap-
pearance of septomaxillaries). This reversal unites Pseu-
doeurvcea, Chiropterotriton. and Dendrotriton, three genera
that share no additional synapomorphies and have little else
in common. Because the first two of these genera are exten-
sively plesiomorphic, no system of analysis can deal with
them easily.

Septomaxillary bones are not well developed in any trop-
ical salamander. They appear regularly in some Chiroptero-
triton (Rabb, 1956; Wake, 1966) but are absent in others
(e.g., C. dimidiatus). Septomaxillary bones are extremely small
and variably present in four of the five species of Dendro-
triton (Lynch and Wake, 1975, 1978). Septomaxillaries also
occur in at least some individuals of several species of Pseu-
doeurycea (P. cephalica, P. leprosa, P. werleri, and P. nigro-
maculata) and occasionally in other bolitoglossine genera.
Wake (1966) reports their presence in individuals of Parvi-
molge townsendi, Oedipina uniformis, Thorius pennatulus,

and three species of Bolitoglossa. We hypothesize that trop-
ical salamanders as a group have undergone osteological re-
duction, and that such reduction may have characterized the
ancestral stock. Loss of the septomaxillaries can be inter-
preted as a manifestation of a general paedomorphic trend
in the group (Wake, 1966; Alberch and Alberch, 1981). Be-
cause these bones are so well developed in Chiropterotriton
relative to other members of the supergenus Bolitoglossa. we
consider it unlikely that a reversal of this character (i.e., re-
evolution of septomaxillaries following their loss in an ances-
tor) has occurred in the genus. Chiropterotriton is the only
tropical genus in which the presence of septomaxillaries is
universal in adults of most species (all but the paedomorphic
C. dimidiatus). The loss of septomaxillaries in certain Chi-
ropterotriton species parallels the loss of the bones in other
genera, and the same underlying mechanism could be in-
volved in all cases.

Paedomorphosis is not necessarily a unidirectional pro-
cess, and some reversals are to be expected. Thus, septo-
maxillaries may have reappeared in Dendrotriton and Pseu-
doeurycea. following their loss in the respective ancestral
stocks. The bones occur irregularly even in those species
where they are found at all. The rare appearance of septo-
maxillaries in other genera is likely to be atavistic, i.e., based
on a reversal of an original paedomorphic event.

Summarizing our arguments, linkage of Chiropterotriton.
Dendrotriton. and Pseudoeurycea on the basis of a postulated
shared reversal of a single character is not warranted. If the
septomaxillary character is omitted from the Wagner anal-
ysis, the resulting dendrogram includes a stem uniting these
same three genera. This time the stem is based on another
shared reversal, separation of tarsal elements (character 4).
We consider such a reversal to be unlikely but not impossible.
Instances are known in which species with fused tarsals show
polymorphism for the trait, so reversals could occur (Wake,
1966; Wake and Brame, 1969; Alberch and Alberch, 1981).
We conclude that parsimony methods must be used cau-
tiously where extensive parallelism pervades an evolutionary
radiation.

An alternative method of analyzing our data is illustrated
in Figure 9. This dendrogram is based on the zoogeographic
and phyletic arguments of Wake and Lynch (1976, fig. 32),
who recognized a "Mexican Group” of genera including
Pseudoeurycea. Chiropterotriton ( sensu stricto ), Parvimolge,
Lineatriton. and Thorius. Using the present suite of osteo-
logical characters, we can find no justification for recognition
of a clade including these five genera; so we have indicated
a five-way division separating them at the second level of
the dendrogram and grouping them to the left of the figure.
Although this dendrogram has relatively little internal struc-
ture, it does distinguish two groups of three and four genera,
respectively.

The dendrogram in Figure 9 differs from that in Figure 8
in several respects. The former dendrogram contains fewer
unique synapomorphies (three rather than six) and more par-
allelisms (ten rather than five), but it has no reversals. A five-
way division occurs above the first synapomorphy. The three
characters involved in unique synapomorphies (1,7, 11) fall

16 Contrikiutions in Science, Number 345

Wake and Elias: Tropical Salamanders

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supergenus

Bolitoglossa

Figure 9. An alternative tree of the genera in the tribe Bolitoglos-
sini. There are no reversals in this tree, but all parallelisms (asterisks)
are indicated.

into Hecht and Edwards’s (1976a) third and fourth cate-
gories, the kinds of characters that they consider to have
relatively high systematic value. The stem uniting Lineatri-
ton, Parvimolge, and Thorius has three synapomorphies. For
illustrative purposes, we have indicated all parallelisms in
Figure 9, and it can be seen that two of these three synapo-
morphies evolved in parallel with other stems. The stem
uniting Bradytriton, Nototriton, Oedipina, and Bolitoglossa
also has three synapomorphies (4, 6, 9), but none of them is
unique.

The tropical plethodontid genera constitute a highly de-
rived lineage. The species are so specialized, in fact, that little
further morphological innovation seems possible. Rather,
variation on similar themes has occurred repeatedly, with
the result that characters useful in the taxonomy of other
salamander groups have been subject to extensive parallel
evolution. We have long known that such features as elon-
gation of body and tail, diminution in body size, changes in

relative limb length and foot size, and increased digital web-
bing have evolved several times in the group (Wake, 1966;
Wake and Fynch, 1976; Alberch and Alberch, 1981). Ap-
parently the evolution of other morphological traits is also
complex, and close evolutionary interlinking of seemingly
unrelated traits is a possibility (see Alberch et al., 1979; Al-
berch and Alberch, 1981). The incorporation of biochemical
and immunological approaches to phyletic analysis holds
promise for unraveling the complex history of this group,
but the long evolutionary history of the neotropical sala-
manders poses serious problems for all known analytical
techniques. At one time, the tropical salamanders were con-
sidered to form a reasonably compact congeneric group ( Dunn,
1926). The intervening years have seen a steady erosion of
this concept as new salamanders have been discovered and
novel analytical techniques have been applied. Based pri-
marily on its possession of a common tongue projection
mechanism (Lombard and Wake, 1 977), the supergenus Boli-
toglossa does appear to be monophyletic, but the group is
more ancient and highly diverse than was thought previ-
ously. The present effort is but one more developmental stage
in our attempt to understand the complex history of the
tropical salamanders.

ACKNOWLEDGMENTS

Work reported in this paper has been under way for several
years and has involved examination of specimens housed in
a number of collections. We especially thank the curators of
the collections of the Museum of Natural History, University
of Kansas; National Museum of Natural History; American
Museum of Natural History; Field Museum of Natural His-
tory; Museum of Comparative Zoology, Harvard University;
Museum of Zoology, University of Michigan; and the Nat-
ural History Museum of Los Angeles County for loan of
specimens and for hospitality and access to their collections
during our visits. Monica Frelow assisted in several phases
of our study. Gene M. Christman of the Museum of Verte-
brate Zoology and James Hendel of the Scientific Photo-
graphic Laboratory, University of California, Berkeley, as-
sisted in preparation of the figures. We have benefitted from
discussions concerning several aspects of this study with Ar-
den H. Brame, James Kezer, James F. Lynch, Stanley Ses-
sions, Marvalee H. Wake, and Kay P. Yanev. We are in-
debted to several colleagues for extensive help in our field
work, especially Jeremy L. Jackson, James F. Lynch, Theo-
dore J. Papenfuss, Robert Seib, and H. Bradley Shaffer, but
also to others too numerous to mention. The work of this
laboratory on tropical salamanders has been supported by
the National Science Foundation (current grant DEB-78
03008). Finally, we thank the Guatemalan Forest Service for
making it possible for us to work in Guatemala, and the
Direccion General de la Fauna Silvestre for collecting permits
for Mexico.

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Contributions in Science, Number 345

Wake and Elias: Tropical Salamanders 17

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. 1969. Systematics and evolution of neotropical sal-
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Submitted 6 October 1981; accepted 15 December 1982.

Contributions in Science, Number 345

Wake and Elias: Tropical Salamanders 19

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mm

Number 346

7 October I9S3

, , .

THE CRANIAL MORPHOLOGY ilF THE FOSSIL BEAVER
DIPOIDES SMITH. I (ROBENITA: MAMMALIA)

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Printed at Allen Press, Inc., Lawrence. Kansas

THE CRANIAL MORPHOLOGY OF THE FOSSIL BEAVER
DIPOIDES SMITHI (RODENTIA: MAMMALIA)

Hugh M. Wagner1

ABSTRACT. The cranial morphology of the genus of fossil beaver,
Dipoides, has never been described or compared with that of other
fossil and extant beavers. An extremely well-preserved skull re-
covered from north-central Oregon in 1974 allows for identification
of nearly all features of the cranium. Variation in skull morphology
in rodents is generally recognizable at the generic level, and intra-
specific variation is usually reflected in differences in the dentition.
The skull of Dipoides closely resembles that of Eucastor and Cas-
toroides and differs noticeably from that of Castor. The regions of
the skull with similarities to Dipoides, Eucastor, and Castoroides are
( 1 ) the bones of the lacrimal region; (2) the arrangement of foramina
in the basicranium and interorbital region; and (3) the development
of the alisphenoid wing of the internal pterygoid fossa. Castor differs
in these three areas. Close examination of the crania of these four
genera confirms that Eucastor, Dipoides, and Castoroides represent
one group of beavers that has been phyletically distinct from that
leading to the extant beaver. Castor, since the early Miocene.

INTRODUCTION

Species in the genus Dipoides Jager, 1835, are small beavers
that lived during the Hemphillian Mammal Age in North
America. The genus appears to have evolved from Eucastor
Leidy, 1 858, during the late Miocene, approximately 8.5 mya
(Wagner, 1981).

The genus Dipoides was first described by G.F. Jager in
1835 from late Miocene deposits of Germany. Subsequently,
other species of Dipoides have been described from late Mio-
cene deposits of Eurasia and North America. R.A. Stirton
(1935) reviewed the Tertiary beavers, presenting a phylogeny
based on dental characters, with Dipoides in the lineage lead-
ing to Castoroides Foster, 1838. Later, Olson (1940) under-
took a comparative study of the cranial foramina of the Cas-
toridae. He described and compared the cranial foramina of
Paleocastor Leidy, 1869, Eucastor, Castoroides, and Castor
Linneaus, 1758, noting evolutionary trends in foramina with-
in the Castoridae. Dipoides was not included in that study.

J.A. Shotwell (1955) reviewed the genus Dipoides, basing
his taxonomic conclusions on the dental morphology, pri-

1. Department of Physics, University of California, Irvine, Cal-
ifornia 92717, and Research Associate, Vertebrate Paleontology,
Natural History Museum of Los Angeles County.

marily P4. He concluded with the description of a new species,
D. smithi Shotwell, 1955, from McKay Reservoir in north-
central Oregon. Stirton (1965) described the cranial mor-
phology of Castoroides ohioensis Foster, 1838, in detail, in-
creasing the knowledge of this giant castorid. The specimen
of D. smithi that I describe in the present paper was recovered
from the type locality of the species at McKay Reservoir (Fig.
1 ) in 1974 and provides new evidence concerning the cranial
morphology of the genus and helps clarify its relationship to
other genera of castorids.

METHODS AND MATERIALS

The abbreviations used in the text and tables are CNHM
(Chicago Natural History Museum), UO (University of Or-
egon), and UCMP (University of California Museum of Pa-
leontology). All measurements are in millimeters. The com-
parative specimens are:

Eucastor cf. E. tortus Leidy, 1858, CNHM P15787, Big
Springs Canyon local fauna, Bennett County, South Da-
kota, Ogallala Group, Clarendonian.

Castoroides ohioensis Foster, 1838, CNHM P3800, Gravel
Pit near Bellflower, McClean County, Illinois Pleisto-
cene.

Castor canadensis Kuhl, 1820, UCMP 126-158, South
Dakota, Recent.

Dipoides smithi Shotwell, 1955, UCMP 113422, McKay
Reservoir local fauna, Umatilla County, Oregon, ? Shu-
tler Formation, Hemphillian (Figs. 2, 3 and 4).

SYSTEMATICS

Family Castoridae Gray, 1 92 1
Subfamily Castoroidinae Trouessant, 1888
Genus Dipoides Jager, 1835

Dipoides smithi Shotwell, 1955

Figures 2, 3, and 4

REFERRED MATERIAL. UCMP 113422, nearly com-
plete skull lacking only the projection of the premaxillaries

Contributions in Science, Number 346, pp. 1-6
Natural History Museum of Los Angeles County, 1983

ISSN 0459-8113

Figure 1. The skull of Dipoides smithi was recovered from UCMP
locality V74163 at McKay Reservoir in Umatilla County in north-
central Oregon, designated by an X.

and incisors (Figs. 2 and 3); UCMP 113433, left dentary
fragment with P4, M,_2 (Fig. 4), both from locality UCMP
V74 1 63 = U02222, McKay Reservoir, Umatilla County,
Oregon, ? Shutler Formation, Hemphillian.

DESCRIPTION AND COMPARISON. The dental
morphology of UCMP 1 13422 agrees with the diagnosis of
D. smithi (Shotwell, 1955). Upper molars and premolars lack
parastria and have S-shaped occlusal patterns characteristic
of this species. In the sample of P4’s of D. smithi from McKay
Reservoir, 16 percent of 24 P4’s had complete or vestigial
parastriids and paraflexids.

The incisors and most of the ventral region of the pre-
maxillary bone are absent in this specimen. The dorsal sur-
face of the skull is long and relatively flat. Over the orbits
the frontal bone forms a broad, Hat surface (Fig. 2). The nasal
region is straight, with neither the lateral nor dorsal inflation
of Castor canadensis. A small protuberance exists on the
dorsal lip of the fossa for the masseter profundis anterior
muscle at the premaxillary-maxillary suture. This protuber-
ance is similar to that in Castoroides ohioensis ; Eucastor has
only a slight one and Castor lacks one. Frontoparietal crests
arise above the orbits and extend posteriorly along the fron-
tal-parietal suture, where they join to form a strong median
sagittal crest between the parietal bones. The lambdoidal
crests are well developed, extending laterally nearly to the
tip of the mastoid processes.

The anterior and posterior temporal foramina are both
present on the parietal-squamosal suture and in the same
position as in Eucastor and Castoroides. Approximately ninety
percent of the surfaces of the parietal bones are covered with
rugosities. Eucastor and Castoroides also have similar ru-
gosities although not as extensive anteriorly as in this spec-
imen of Dipoides. In Castor, such rugosities are not as well
developed. The dorsal surface of the cranium in Dipoides is
more inflated than in Eucastor but not as inflated as in Cas-
tor. The occipital plate intersects a plane extended posterior

Figure 2. Dipoides smithi : Shotwell, 1955, UCMP 1 1 3422, McKay
Reservoir local fauna, UCMP locality V74 1 63, nearly complete skull
lacking the incisor region; A, dorsal view; B, right lateral view; C,
ventral view; scale indicated.

from the palate at approximately seventy degrees. By com-
parison, in Eucastor, Castoroides, and Castor, the occipital
plate intersects the same plane at nearly ninety degrees. A
small mastoid foramen pierces the mastoid bone slightly lat-
eral to the mastoid-supraoccipital suture. In size and position
the mastoid foramen is very similar to that in Castoroides.
In Castor, this foramen is much larger and situated on the
mastoid-supraoccipital suture. The occipital condyles appear
very similar to those in Castor and do not show the transverse
flattening of those of Castoroides.

2 Contributions in Science, Number 346

Wagner: Cranial Morphology of Dipoides smithi

Anterior and ventral to the infraorbital foramen there is a
very prominent masseter superficialis process (Fig. 3, B). A
prominent flange or sheath of bone extends dorsally from the
lateral edge of this process, closing off the ventral region of
the infraorbital foramen from the masseter lateralis profun-
dus anterior fossa (Fig. 3, B). This flange bends posteriorly
toward its dorsal limit and does not extend above the in-
fraorbital foramen. The morphology of the masseteric su-
perficialis process and infraorbital complex in Dipoides is
nearly identical to that of Castoroides. In Eucastor the mas-
seteric superficialis process is not well developed, and the
structure of the infraorbital region resembles that of Castor.

The zygomatic arch is wide, but not to the extreme seen
in Castor or Castoroides. The orbit of Dipoides is propor-
tionally larger than in Castor and Castoroides and appears
to be directed more laterally than in Castor. The structure
of the zygomatic arch and lacrimal region is similar to that
in Castoroides and Eucastor and differs from that in Castor.
The contribution that the jugal bone makes to the zygomatic
arch of Dipoides, Eucastor, and Castoroidesd s less than in
Castor. In the former three genera, the jugal-maxilla suture
is on the lateral surface of the zygomatic arch as contrasted
to Castor where this suture is in the lacrimal area. The an-
terior margin of the orbit is composed of the maxilla, because
the jugal does not extend into the lacrimal region. In Castor,
however, the jugal-maxilla suture extends anteriorly on the
dorsal surface of the zygomatic arch, and the jugal meets the
lacrimal in the anterior region of the orbit.

Dipoides has a simple lacrimal region in which only the
frontal and maxilla meet on the dorsal surface of the skull.
The lacrimal bone does not have a dorsal component in this
specimen. The structure of the lacrimal region in Dipoides
and Castoroides is similar by having only the two dorsal
elements. Eucastor differs slightly from Dipoides and Cas-
toroides, because a small portion of the lacrimal bone is
visible on the dorsal surface at the maxillary-frontal suture.
Castor has a much more complicated dorsal lacrimal area in
which the jugal, maxillary, lacrimal, and frontal bones all
meet.

The pattern of foramina within the inner orbit of Dipoides
is very similar to that in Eucastor and distinctly different
from the pattern in either Castoroides or Castor. However,
the resemblance to Castoroides is greater than to Castor. The
sphenoid foramen is the anterior foramen, piercing the max-
illary in an anterior direction. Directly behind the sphenoid
foramen is a slightly smaller one that enters the cranium
medially. This foramen completely penetrates the cranium
as does the presphenoid canal of Olson (1940) (alisphenoid-
rotundum foramen of Stirton (1965)). This foramen appears
to be an anterior presphenoid canal. Directly above the an-
terior presphenoid canal on the orbitosphenoid-frontal su-
ture is the ethmoid foramen. Posterior and ventral to the
ethmoid foramen is a well-developed optic foramen entirely
enclosed in the orbitosphenoid bone. A large posterior pre-
sphenoid canal is situated posterior to the optic foramen in
a deep sulcus at the anterior end of the sphenoidal fissure.
Near the anteroposterior margin of the alisphenoid wing of
the internal pterygoid fossa is the masticatory-buccinator

Figure 3. Dipoides smithi, Shotwell, 1955, skull with structures
labeled, UCMP 1 13422; A, dorsal view; B, right lateral view; C,
ventral view; scale indicated roman type represents bones and struc-
ture, italics are foramina. Abbreviations: af, alisphenoid foramen;
als, alisphenoid; apf, anterior palatine foramen; atf anterior tem-
poral foramen; awipf, alisphenoid wing of internal pterygoid process;
bo, basioccipital; earn, external auditory meatus; ef. ethmoid fora-
men; eo, exoccipital; fr, frontal; hf, hypoglossal foramen; iof, in-
fraorbital foramen; ip, interparietal; j, jugal; m, mastoid; mf, mastoid
foramen; mlf-fo, median lacerate foramen-foramen ovale; mp, mas-
toid process; ms-bcf masticatory-buccinator foramen; msp, mas-
seteric superficialis process; mx, maxillary; n, nasal; of, optic fora-
men; p, parietal; pgf, postglenoid foramen; pi, palatine; pp, paroccipital
process; prsp. can., presphenoid canal; ps, premaxillary; pt, ptery-
goid; ptf, posterior temporal foramen; sf, stylomastoid foramen; so,
supraoccipital; sphf sphenoidal foramen; sq, squamosal; ty, tym-
panic; vc, ventral choana.

Contributions in Science, Number 346

Wagner: Cranial Morphology of Dipoides smithi 3

I cm.

Figure 4. Dipoides smithi, UCMP 1 13433, UCMP V74163, col-
lected near the skull figured in this paper. Left dentary fragment with
P4,M,_2 showing normal occlusal wear. This specimen has the more
typical P4 of D. smithi, where no paraflexid is present.

foramen. Below the masticatory-buccinator foramen, a por-
tion of the alisphenoid canal is visible, as it passes through
the alisphenoid bone. Anteroventral to the alisphenoid canal
is a large foramen that appears to enter the pulp cavity of
M3. The pattern of foramina anterior to and including both
presphenoid canals in Dipoides is very similar to that in
Eucasior and Castor. Eucastor resembles Dipoides by having

the multiple presphenoid canals. By contrast, in Castoroides,
the presphenoid canal is single and proportionally much
smaller and more posterior in its position.

Posterior to the presphenoid canal, the morphology of the
alisphenoid in Dipoides does not resemble that in Castor but
is very similar to that in Castoroides. In Dipoides and Cas-
toroides, the alisphenoid forms a smooth lateral surface of
the sphenoid fissure from the anterior sulcus of the presphe-
noid canal posterior to the masticatory-buccinator foramen.
The masticatory-buccinator foramen is directed posteriorly
through the margin of the alisphenoid. In Castor, part of the
alisphenoid forms a strap-like structure, forming the lateral
surface of the alisphenoid canal. The masticatory-buccinator
foramen and alisphenoid foramen share a common orifice
that is at the posterior end of the straplike structure of the
alisphenoid. This common foramen is directed anteriorly,
entering directly into the sphenoid fissure. The alisphenoid
foramen pierces the broad alisphenoid wing of Dipoides and
Castoroides, and a branch of the trigeminal nerve (V2) enters
the skull through a bone canal enclosed in the alisphenoid
wing of the internal pterygoid. Castor differs in that the ali-
sphenoid foramen does not pass through the alisphenoid wing,
but the trigeminal nerve enters via the alisphenoid foramen.
This region of the skull of the specimen of Eucastor was not
well enough preserved to allow comparison.

The postglenoid foramen is large and nearly equal in size

Table 1. A comparison of cranial characters in Eucastor cf. E. tortus, CNHM PI 5787; Dipoides smithi, UCMP 113422; Castoroides ohioensis,
CNHM P3800; and Castor canadensis, UCMP 126-158.

Characters

E. cf. E. tortus

D. smithi

C. ohioensis

C. canadensis

Protuberance on dorsal lip of anterior

slight

slight

well-developed

absent

masseteric fossa

Temporal foramina present

yes

yes

yes

yes

Rugosities present on parietal bone

slight

well-developed

well-developed

slight

Inflation of cranium

slight

slight

slight

well-developed

Mastoid foramen

small

small

small

large

Masseteric superficialis process well developed

no

yes

yes

yes

Masseteric superficialis process anterior to

no

yes

yes

no

anterior masseteric fossa

Lateral vertical flange on infraorbital foramen

yes

no

no

yes

Parallel tooth row on maxillary

yes

no

no

yes

Dorsal choana or invagination on pterygoid or

no

no

yes

yes

basioccipital bones

Posterior carotid foramen

absent

absent

present

present

Alisphenoid foramen pierces medial

not known

yes

yes

no

alisphenoid wing of internal pterygoid fossa

Jugal extends into lacrimal region

no

no

no

yes

Multiple presphenoid canals

yes

yes

no

yes

Bulla highly inflated

yes

yes

no

yes

Mastoid process extends beyond external

no

no

no

yes

auditory meatus

4 Contributions in Science, Number 346

Wagner: Cranial Morphology of Dipoides smithi

to the external auditory meatus. The external auditory me-
atus is posterior and ventral to the postglenoid foramen and
is appressed posteriorly against the mastoid process and di-
rected laterally. Its anterior lip has the greatest lateral exten-
sion. The condition of the postglenoid foramen and external
auditory meatus is very similar to that of Castoroides. Castor
and Eucastor differ, having a much smaller postglenoid fo-
ramen; the meatus has a more dorsolateral orientation, and
its ventral lip has the greatest lateral extension. In Dipoides
and Castor, the meatus projects laterally beyond the mastoid
process. In Castoroides, it does not.

The internal pterygoid fossa is broad, bounded laterally by
the alisphenoid wing and medially by the palatal wing of the
pterygoid, which is a septum supporting the hamular process.
The alisphenoid foramen is present just anterior to the con-
joined median lacerate foramen and foramen ovale on the
alisphenoid wing of the internal pterygoid fossa (Fig. 3, C).
The surface of the pterygoid fossa has depressions. The ven-
tral choana is triangular in outline, bounded laterally by the
palatal wings of the pterygoid. In Castoroides the outline of
the ventral choana is similar to that of Dipoides, but in Castor
it is nearly circular.

There is a rugosity on the anterior median border of the
bulla, resembling that of Castor, where the hamuiar process
meets the tympanic. In Castor these rugosities extend farther
posteriorly, so it seems probable that the hamular processes
of Dipoides were not as long as in Castor. The overall struc-
ture of the internal pterygoid fossa in Dipoides resembles that
in Castoroides, in which the alisphenoid wing of the internal
pterygoid fossa is expanded, and the alisphenoid foramen
enters the skull through the medial surface of the alisphenoid.
This region of the skull of Eucastor was not preserved well
enough to allow comparison.

Dipoides does not possess the dorsal choana (mesoptery-
goid fossa) of Castoroides (see Stirton, 1965). This fossa is
unique to Castoroides. Castor does have a shallow invagi-
nation in the basioccipital, but this invagination is posterior
to the mesopterygoid fossa of Castoroides.

A small anterior carotid foramen in Dipoides pierces the
lateral margin of the basisphenoid anterior to the medial
extension of the bulla. It is similar in position to that of
Castor. In Dipoides and Eucastor, the posterior carotid fo-
ramen is not visible along the medial margin of the bulla
anterior to the posterior lacerate foramen as it is in Castor
and Castoroides. A large posterior lacerate foramen is present
along the posteromedial border of the bulla. Directly pos-
terior to the posterior lacerate is a large hypoglossal foramen.
A small stylomastoid foramen is present, wedged between
the base of the mastoid process and the boney tube of the
tympanic leading to the externa! auditory meatus. The par-
occipital process extends posteriorly almost as far as do the
occipital condyles (see Table 1). The very tip of the paroc-
cipital process is hooked ventromedially.

The bullae are very inflated. Their ventral extension does
not reach as far ventrally as the greatest depth of the skull
at the tooth row (see Fig. 2, B; Table 2). In Castor, the bulla,
though exhibiting no greater inflation, extends ventrally to a

Table 2. Measurements of cranial and dental characters of Dipoides

smithi, UCMP 113422 (in millimeters).

Posterior margin lambdoidal crest to anterior
tip of nasals 90.5*

Dorsal width of rostrum anterior to
zygomatic arch I 7.0

Least width between orbital fossae 14.7

Greatest width between external auditory meatus 45.7

Greatest width between mastoid processes 23.3

Greatest width of zygomatic arches 64.9

Greatest width between masseteric

superficialis processes 23.3

Length from dorsal margin of foramen magnum
to greatest height of occiput 15.2

Depth of skull between dorsal surface of skull
and occlusal surface of M2 36.8

Depth of skull between sagittal crest and bulla 30.6

Least width of palate between P4’s 5.2

Least width of palate between M3’s 14.4

Length from anterior border of PJ to posterior
border of M3 21.9

Length of occlusal surface of P4 5.8

Width of occlusal surface of P4 5.5

Length of occlusal surface of M1 4.9

Width of occlusal surface of M1 4.5

Length of occlusal surface of M2 5.0

Width of occlusal surface of M2 4.4

Length of occlusal surface of M3 5.1

Width of occlusal surface of M3 3.8

* Approximate.

level in line with the tooth row, which is considerably more
ventral than in Dipoides, Eucastor, or Castoroides.

In ventral view, the skull of Dipoides closely resembles
those of Eucastor and Castor in general outline. The incisors
form a crescentic arc that terminates above P4. The dilator
naris fossa is deep in Dipoides, with a well-developed, grooved
median ridge present on the maxilla. In Eucastor, the dilator
naris fossa is deep, as in Dipoides ; however, in the former,
the maxilla slopes steeply toward the maxillary-premaxillary
suture, and in the latter a relatively flat platform is present.
The masseteric superficialis process of Dipoides and Casto-
roides is posterior to the anterior flaring of the maxillary.
The cheek tooth row diverges posteriorly from the midline
of the skull. Castoroides has a similar appearance with closely
placed P4’s and widely separated M3’s; whereas in Eucastor
and Castor, the cheek tooth rows are nearly parallel. The
tooth wear in Dipoides forms a flat transverse plane differing
from that in Castor wherein the upper tooth surface is con-
cave with the labial margin having greater elevation than the
lingual edge. There are two nearly parallel medial grooves
that run from the premaxillary-maxillary suture posterior to
ML These grooves resemble those in Eucastor and Castor,

Contributions in Science, Number 346

Wagner: Cranial Morphology of Dipoides smithi 5

though in Dipoides they are deeper. The anterior palatine
foramina are present on the suture between the M2’s. The
posterior palatine foramen is situated directly posterior to
M3. A ridge is present posterior to the posterior palatine
foramen forming the posterolateral margin of the palate. The
posterior margin of the palate forming the roof of the mouth
of Dipoides describes a smooth concave margin. Eucastor
differs markedly from this condition in having an invagi-
nation in the palatine bone that extends anteriorly to the
anterior margin of M3. Castor and Castoroides resemble Di-
poides in this feature.

DISCUSSION

A detailed examination of the cranial morphology of Di-
poides smithi reveals close similarities to Eucastor cf. E.
tortus and Castoroides ohioensis. The similarities in the po-
sitions of the foramina in the orbital region and basicranium
and in the bone development of the skull in these three
species indicate a close phyletic relationship supporting Stir-
ton’s (1935) conclusions based on dental morphology. The
similarities of the skull in Dipoides, Eucastor, and Casto-
roides are ( 1 ) the relationship of bones of the lacrimal region;
(2) the arrangement of foramina in the basicranium and in-
terorbital region; and (3) the development of the alisphenoid
wing of the internal pterygoid fossa. Castor differs markedly
from other genera in all three characteristics.

In Dipoides and Castoroides, the dorsal surface of the lac-
rimal region is composed of the maxilla, premaxilla, and
frontal bones. Eucastor has a lacrimal component. The most
anterior extension of the jugal does not reach the lacrimal
region. The jugal does contribute to this region of the skull
in Castor, a definite difference from the other mentioned taxa.

In the Eucastor, Dipoides, Castoroides group, Eucastor and
Dipoides differ in the morphology of the infraorbital foramen
from Eucastor to Dipoides. The structure of the infraorbital
foramen in Dipoides and Castoroides is derivable from the
condition found in Eucastor. The multiple presphenoid ca-
nals in Eucastor, Dipoides, and some specimens of Castor
appear to be a primitive trait evolved prior to Eucastor and
retained in Eucastor, Dipoides, and Castor but lost in Cas-
troides.

The nonparallel upper cheek tooth rows of Dipoides and
Castoroides are derived compared with Eucastor and Castor.
Similar nonparallel tooth rows are seen in some South Amer-
ican histricomorph rodents. It is probable that this represents
a modification related to a different masticatory adaptation.
For example, the South American rodents that exhibit this

type of modification are primarily aquatic forms, with teeth
composed of lophs for grinding, very similar to the castorid
genera.

ACKNOWLEDGMENTS

Appreciation is expressed to the Oregon Museum of Science
and Industry (OMSI) for providing me the opportunity to
prospect and collect fossil specimens from eastern Oregon
during the summer of 1974, in conjunction with the Paleon-
tological Research Team, during which time the specimen
of Dipoides smithi described in this paper was collected. All
members of the Paleontology Research Team, sponsored by
OMSI, deserve credit for their participation, sharp eyes, and
enthusiasm during the field season. It was a pleasure having
Bruce Batten, Steve Damiani, Kathy Harvey, Roy West, and
Kathy Wolfram as students and companions during the six-
week program. The near perfectly preserved skull of Dio-
poides smithi recovered from McKay Reservoir was found
by Bruce Batten. The illustrations of D. smithi were drawn
by Jaime P. Lufkin, an artist from the Museum of Paleon-
tology, University of California, Berkeley.

I thank the curators of the Museum of Paleontology, Uni-
versity of California, Berkeley, The University of California
Museum of Vertebrate Zoology, and the Natural History
Museum of Los Angeles County for the loan and use of
comparative collections. I also thank Lawrence G. Barnes
for the helpful and constructive comments offered in his
review of this manuscript.

LITERATURE CITED

Jager, G.F. 1835. Ueber die fossilen Saugethiere, welche
in Wtirttemberg gefunded worden sind. Le Abth; 17-18.
Olson, E.C. 1940. Cranial foramina of North American
beavers. Jour. Paleontology 14:495-501.

Shotwell, J.A. 1955. Review of the Pliocene beaver Di-
poides. Jour. Paleontology 29:129-144.

Stirton, R.A. 1935. A review of the Tertiary beavers. Univ.
Ca. Pubs., Geol. Sci., Bull. 23:391-458.

. 1965. Cranial morphology of Castoroides. Dr. D.N.

Wadia Comm. Vol. Nim. & Metal. Inst, of India, pp.
273-285.

Wagner, H.M. 1981. Geochronology of the Mehrten For-
mation in Stanislaus County, California. Ph.D. disser-
tation, Dept. Earth Sci., Univ. of California, Riverside,
347 pp.

Submitted 22 June 1982; accepted 3 May 1983.

6 Contributions in Science, Number 346

Wagner: Cranial Morphology of Dipoides smithi

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SCIENTIFIC PUBLIC. ATHENS COMMITTEE

'Craig C. Black Museum Director
Donald Output

Daniel M. Cohen, Committee Chairman
John M. Harris
Charles L. Hogue

Robin A. Simpson, Managing Editor
Gary D. Wallace
Edward C. Wilson

IAR 2 8 1984
Lib rabies

STUDIES ON THE TAXONOMY AND DISTRIBUTION OF
AMERICAN CENTRIDINE BEES
(HYMENOPTERA: ANTHOPHORIDAE)

Roy R. Snelling1

ABSTRACT. This study focuses primarily on the Centridini of
North America south of the United States and includes a key to the
three genera of Centridini: Centris, Ptilotopus, and Epicharis. Within
the genus Centris keys are provided for species in the subgenera
Paracentris, Centris, Xanthemisia, Acritocentris, Melanocentris,
Trachina, Hemisiella, and Heterocentris ; one new monotypic sub-
genus is described: Ptilocentris (type-species: Centris festiva F. Smith,
1854).

The following new species are described: C. ( Xerocentris ) griseola
(Mexico); C. (C.) aethiocesta (El Salvador to Panama); C. (Acrito-
centris) satana (Mexico, United States); C. (Melanocentris) agi/oides
(Mexico to Costa Rica); C. (M.) gelida (Mexico, Guatemala); C.
(Trachina) eurypatana (Mexico); C. (T.) xochipillii (Mexico). Centris
(C.) meaculpa is proposed as a new name for C. (C.) erubescens
Snelling, 1974, not C. costaricensis var. erubescens Friese, 1925.
Centris atripes var. ferrisi Cockerell is a valid species in Paracentris ;
the types are males, not females as stated by Cockerell.

The following new synonymy is proposed in Centris: C. cockerelli
resoluta Cockerell = C. (Paracentris) cockerelli W. Fox; C. clypeata
Friese, C. anthracina Snelling = C. (P.) nigrocaerulea F. Smith; C.
limbata Friese = C. ( P .) atripes Mocsary; C. strawi Snelling = C.
(Acritocentris) albiceps Friese; C. robusta Cockerell, C. segregata
Crawford = C. (C.) inermis Friese; C. citrotaeniata Gribodo, C.Jla-
vifrons var. rufescens Friese, C. flavifrons var. nigritula Friese = C.
(C.) flavifrons (Fabricius); C. chlorura Cockerell = C. (Ptilocentris)
festiva F. Smith; C. ignita F. Smith, C. bakerella Friese, Epicharis
cisnerosi Cockerell = C. (Melanocentris) agilis F. Smith; C. fusci-
ventris var. scutellata Mocsary = C. (M.) fusciventris Mocsary; C.
melanochlaena F. Smith, Epicharis zamoranensis Cockerell = C. (A/.)
obsoleta Lepeletier; C. schwarzi Cockerell = C. (Trachina) labiata
Friese; C. confinis Perez = C. (Hemisiella) nitida F. Smith; C. ruae
Cockerell = C. (H.) transversa Perez; C. dentipes F. Smith, C. ru-
fomaculata Cockerell, C. lanipes subtarsata Cockerell = C. (H.) tri-
gonoides Lepeletier; C. costaricensis var. erubescens Friese = C. (H.)
vittata Lepeletier; C. triangulifera Cockerell = C. (Heterocentris)
labrosa Friese.

Ptilotopus is elevated to generic rank from its previous status as
a subgenus of Centris ; P. zonalis Mocsary of Panama is the only
species in North America.

A key is given for all the subgenera of Epicharis and keys are given
for the species of each subgenus known to occur in North America.

Two new species are described in the subgenus Epicharana: E. an-
gulosa (Costa Rica) and E. bova (Costa Rica, Panama). The following
are new synonyms: E. salazari Cockerell = E. (Epicharana) elegans
F. Smith; E. rustica var . /lava Friese = E. (Epicharana) rustica (Oliv-
ier); E. phenacura Cockerell, E. conura Cockerell = E. (Parepicharis)
metatarsalis Friese. North American distribution data are cited for
all species.

Included is a synonymic list of the recognized North and Central
American Centridini.

RESUMEN. Este estudio se refiere principalmente a los Centridini
de Norte America al sur de los Estados Unidos e incluye una clave
para los tres generos de Centridini: Centris, Ptilotopus y Epicharis.
Dentro del genero Centris se incluyen claves para ias especies de los
subgeneros Paracentris, Centris, Xanthemisia, Acritocentris. Mela-
nocentris, Trachina, Hemisiella y Heterocentris ; un subgenero mo-
notipico nuevo se describe, Ptilocentris (especie-tipo: Centris festiva
F. Smith, 1854).

Ptilotopus se eleva al rango generico desde su estatus previo de
subgenero de Centris ; P zonalis Mocsary de Panama es la unica
especie en Norte America.

Se da una clave para todos los subgeneros de Epicharis y para las
especies de cada subgeneros conocidos en Norte America.

Se describen ocho nuevas especies para el genero Centris y dos
para el genero Epicharis. Estas al igual que los nuevos sinommos se
incluyen en el “Abstract.”

Se citan datos de distribucion de Norte America para todas las
especies. Se incluye una lista de los Centridini reconocidos de Norte
America y de las sinonimias conocidas.

INTRODUCTION

The present paper is a continuation of my earlier work on
the Centris of North and Central America (Snelling, 1956,
1 966, 1974). New data on the distribution of previously treat-
ed species, as well as investigations into the systematics of
previously unstudied groups are presented below. Although
my previous investigations involved only the genus Centris,

1. Section of Entomology, Natural History Museum of Los An-
geles County, 900 Exposition Blvd., Los Angeles, California 90007.

Contributions in Science, Number 347, pp. 1-69
Natural History Museum of Los Angeles County, 1984

ISSN 0459-8113

the study has now expanded to include the genera Epicharis
and Ptilotopus, the latter newly elevated to generic status
from its previous position as a subgenus of Centris.

SPECIMENS EXAMINED

Material utilized in this study is from the following institu-
tional and private collections: American Museum of Natural
' History (AMNH), Robert W. Brooks, personal collection
’ (RWB), California Academy of Sciences (CAS), Cornell Uni-
versity (CORN), Florida State Department of Plant Industry
(DPIF), Museum of Comparative Zoology (MCZ), Museum
National d’Histoire Naturelle, Paris (MNHN), Natural His-
tory Museum of Los Angeles County (LACM), John L. Neff,
personal collection (NEFF), Oregon State University (ORSU),
D. Roubik, personal collection (ROUB), United States Na-
tional Museum of Natural History (USNM), University of
California, Berkeley (UCB), University of California, Davis
(UCD), University of Kansas (UKAN), and Thomas J. Za-
vortink, personal collection (TJZ).

SPECIMEN DATA

New data on distribution, capture dates, and plant associa-
tion are fully cited for many species since there are few pub-
lished records. Localities are organized by country and by
state, province or department within that country. The des-
ignation “state,” “department,” or “province” is omitted to
conserve space. Thus, a locality cited under “COSTA RICA,
SAN JOSE” is from San Jose Province in Costa Rica.

Specimen data are cited for most species, but some species
are common, widely distributed and abundant. For such
species, only general range is given.

TERMINOLOGY

In general, the morphological terminology employed below
is that which is traditional is apoid systematics, following
Michener (1944, 1954, 1 965). A few terms, however, should
be explained, in order that they may be correctly employed
(in the sense of this paper) by others.

Antennal socket diameter. Since the antennal socket is not
circular it follows that two different measurements are pos-
sible. In this study, the antennal socket diameter is measured
perpendicular to the long axis of the head (i.e., the transverse
diameter); measurement is made from the summit of the rim
on the opposite side. Associated measurements include the
interantenna/ distance and the antennocular distance.

Interantenna/ distance is measured as the least distance
between the summits of the two antennal socket rims.

Antennocular distance is the least distance from the inner
eye margin to the summit of the rim of the adjacent antennal
socket.

Several features of the clypeus should be defined for con-
sistent usage. The clypeal disc is the central portion of the
clypeus, from base to apical margin and roughly bounded on
either side by an imaginary line extended distad from the
junction of the subantennal sutures with the base of the clyp-

eus. Clypeal length, when compared to clypeal width, is mea-
sured along the midline, from the base to the apical margin.
Clypeal width is measured as the greatest distance between
the apicolateral extremities of the clypeus (i.e., that portion
nearest the inner eye margin). The distance between the clyp-
eus and the eye is measured as the least distance between the
apicolateral extremity and the nearest point on the inner eye
margin.

Elaiospathe is a new term: It is a combination of the Greek
words elaion (oil) and spathe (a paddle for stirring or mixing;
a broad blade). This term refers to the modified oil-gathering
structures present in female centridine and exomalopsine
bees. The elaiospathe most commonly consists of a row of
long, erect, apically spatulate and curved setae, usually on
the pro- and mesobasitarsi; in Centridini they are situated
anteriorly on the “inner” surface of the segment, in some
Exomalopsini they are external and may be present in males
as well as females, and are limited to the probasitarsi. How-
ever, oil-collecting structures may also be present on the
abdominal venter, at least in some species of Tapinotaspis
( Tapinotaspoides). For an excellent survey of the morphology
and function of the elaiospathe, see Neff and Simpson (198 1).

The lower frontal width is measured as the distance be-
tween the inner eye margins at the level of the apicolateral
angle of the clypeus. It is compared with the upper frontal
width to determine the degree of convergence of the inner
eye margins. The upper frontal width is the minimum dis-
tance between the inner eye margins at about the level of the
ocelli.

The diameter of the anterior ocellus is the greatest trans-
verse (perpendicular to long axis of head) diameter of the
anterior ocellus. Interocellar distance is the minimum dis-
tance between the posterior, or lateral, ocelli. Ocellocular
distance is the minimum distance between one of the pos-
terior ocelli and the nearest point on the eye margin. Ocel-
loccipital distance is measured in dorsal view and is the min-
imum distance from the posterior margin of one of the
posterior ocelli to the occipital declivity. Transocellar dis-
tance is measured as the maximum distance between the
outer margins to the two posterior ocelli. In all cases, the
ocellus is, for purposes of measurement, considered to be
only the transparent amber-colored lens.

Pilosity. In general, centridines, like most other anthopho-
rids, are densely hairy bees, with much of the head, thorax,
and first abdominal tergum clothed with long, plumose, de-
cumbent to fully erect hairs. Similarly, the legs are hairy, but
hairs tend to be decumbent to subappressed on the outer
faces of the tibiae and basitarsi and are often stout and simple.
Long, plumose hairs are usually present, but sparse, on the
femora. On the abdomen, the hairs on the discs of the seg-
ments generally become longer and more erect on succeeding
segments, plumose hairs as a rule being absent from the discs
of the second and third terga.

The abdominal sterna normally have long, plumose hairs
which tend to be longest and most abundant along the mid-
line. As a rule, males are more densely pubescent on the
sterna than are the females.

Punctation. The description of characteristics related to

2 Contributions in Science, Number 347

Snelling: American Centridini

the punctation of the integument of bees and other Hyme-
noptera have long been the bane of taxonomists since the
terminology has always been undefined and subject to con-
siderable variation in interpretation. Punctures which one
author considers to be fine, another might describe as mod-
erate in size. The distances between punctures, whether dense,
close, or sparse, have likewise been troublesome. A few years
ago I (Snelling, 1980) introduced a system of terminology to
solve this difficulty in the genus Hylaeus. After considering
the objections by Harris (1979), I can find no valid reason
not to apply the same terminology of absolute measurements
to the Centridini. I do not agree that punctation need be
described relative to the size of the entity. Admittedly, I was
initially uncomfortable at describing as “coarse” those punc-
tures on a centridine bee that I was accustomed to considering
to be moderate-sized. This is, however, strictly a perceptual
problem, one which has ceased to disturb my sense of “right-
ness.”

Therefore, I propose to expand the usage of my system to
include the Centridini. Such a system must be considered,
at this point, to be experimental; some adjustment or mod-
ification may prove to be necessary as this method is tried
on other groups. A micrometer disk in one ocular of a mi-
croscope is, obviously, necessary and I use a minimum mag-
nification of 64 x; several punctures of about the same ap-
parent size are measured and averaged. The appropriate
terminology is as follows:

minute— puncture diameter 0.010-0.019 mm
fine— puncture diameter 0.020-0.035 mm
moderate— puncture diameter 0.036-0.055 mm
coarse— puncture diameter 0.056-0.070 mm
very coarse — puncture diameter over 0.070 mm

Since punctures are often not uniform in size on a given
segment or area, it may be necessary to combine terms into
phrases such as “fine to moderate” (puncture diameter vary-
ing between 0.020 and 0.055 mm), though usually a more
limited size range, such as moderate, may prevail.

The relative density of punctures may also be standardized
and I use the following terminology below:

Contiguous— punctures so close that they are often de-
formed; interspaces are compressed and sharp-edged.

Subcontiguous— punctures separated by more or less flat
interspaces up to about 0.30 times a puncture diameter; some
punctures may be deformed.

Dense — punctures separated by more or less flat inter-
spaces between 0.30 and 0.70 times a puncture diameter;
punctures usually round but may be elongate.

Close — punctures separated by more or less flat interspaces
0.70 to 1.50 times a puncture diameter.

Sparse — punctures separated by more or less flat inter-
spaces 1.50 to 3.00 times a puncture diameter.

Scattered — puncture interspaces are very irregular and range
from about 3.00 to 6.00 or more times a puncture diameter.

Variations in puncture density may be expressed by com-
bining terms: “sparse to scattered.”

Clypeal punctation is usually described from the middle
one-third of the disc.

Mesoscutal punctation is described from the area between
the parapsidal line and the median line at the level of the
tegula; punctures are commonly less close posteromesally on
the segment and are closer laterad of the parapsidal line.
Scutellar and metanotal sculpture is described from the mesal
one-third of the segment. The middle of the mesopleural disc
is the standard for that segment.

The first abdominal segment has a nearly vertical anterior
and a horizontal posterior face; the latter is referred to as the
disc of the segment. Tergal punctation is described from the
mesal one-third of the postgradular area of the segments
beyond the first, and anterior to the apical zone.

The distal portion of the abdominal terga, especially of the
second and following segments, except the last, is commonly
depressed in bees; this area is usually called the apical depres-
sion and it is often sculptured differently from the disc. In
centridines this area is usually not obviously depressed, es-
pecially across the middle two-thirds of the segment, but is
usually more sparsely and less coarsely sculptured than the
disc; herein I refer to this as the apical zone.

In the description of new taxa, measurements and pro-
portions are based on the primary type specimens. The cor-
responding range of measurements and proportions of para-
types only (if any) are indicated parenthetically after those
of the primary type.

NESTING BIOLOGY

The little that is known of centridine nesting biology was
reviewed by Coville, Frankie, and Vinson (1983). They con-
cluded that the choice of nesting substrate by the female bees
is roughly correlated with taxonomic groupings. Thus, those
species in the subgenera Wagenknechtia, Paracentris, Xero-
centris, Centris s. str., Exallocentris, and Melanocentris nor-
mally excavate original tunnels in soil. About one-half of the
species cited utilize flat soil and about one-half excavate in
more or less vertical banks or mud walls; two species evi-
dently are capable of choosing either flat or vertical surfaces.

Species belonging to the subgenera Hemisiella and Het-
erocentris generally utilize preexisting cavities in a variety of
substrates. Some species, at least, seem to be highly oppor-
tunistic in choice of substrate and may choose old bee cells
and burrows in soil, abandoned Sceliphron nests, or holes in
wood. Of two species of Trachina cited, one nested in the
walls of a living arboreal termitarium and the other chose
flat soil.

Species of Ptilotopus, herein treated as a separate genus,
were uniform in excavating their nests in the walls of living
arboreal termitaria. There is some suggestion that the ar-
rangement of cells within the nest may similarly substantiate
the present groupings of subgenera, but the data are very
fragmentary.

SYSTEM A I ICS

The general interpretation of the Centridini has been that
there are two genera, Centris and Epicharis (e.g., Michener,
1954). One result of my recent work in this tribe is that I
now believe that Ptilotopus , formerly treated as a subgenus

Contributions in Science, Number 347

Snelling: American Centridini 3

of Centris, must be elevated to generic status; justification
for this decision will be provided below. The three genera of
Centridini which 1 recognize may be separated as follows.

KEY TO GENERA OF CENTRIDINI

la. Marginal cell of forewing shorter than distance from its
apex to tip of wing; occipital llagelliform setae usually
absent, but if present, not reaching as far back as anterior

margin of tegula 2

b. Marginal cell of forewing longer than distance from its
apex to tip of wing; occipital llagelliform setae present
and usually extending to, or beyond, level of anterior

margin of tegula Epicharis

2a. Occipital llagelliform setae absent; mesosternum of fe-
male without tubercles; hypoepimeron flat or slightly
convex; pronotal lobe low and moderately convex, never

conspicuously tuberculate Centris

b. Occipital llagelliform setae present and extending to an-
terior margin of mesoscutum, or slightly beyond; meso-
sternum of female with a pair of stout tubercles hidden
in dense cluster of hairs; hypoepimeron usually promi-
nently coniform, but if not, its lower margin is sharply
elevated above adjacent portions of mesepisternum; pro-
notal lobe often tuberculate Ptilotopus

Genus Centris Fabricius

Centris is a primarily tropical genus, with 1 1 subgenera; these
subgenera are separated in keys by Michener (1951) and
Snelling (1974). The separation of Ptilotopus from Centris
(see below) removes the only group with flagelliform occipital
setae. The North American species of the subgenera Xero-
centris and Paracentris were treated by Snelling (1974). The
North American species of the remaining subgenera are treat-
ed below.

Subgenus Xerocentris Snelling

Centris subg. Xerocentris Snelling, 1974:3-4. Type-species;

Centris californica Timberlake, 1 940; original designation.

This subgenus was proposed for a number of species found
in arid regions of North and South America. Although Xero-
centris is related to Paracentris, females lack an elaiospathe
on the pro- and mesobasitarsi, present in Paracentris and
Centris s.s., and the pygidial plate is convex along the median
line. Males are much like those of Paracentris. but the ocel-
locular distance is less than the diameter of a lateral ocellus,
and the abdominal terga are covered with abundant fully
erect, often pale, hairs.

One new species of Xerocentris is described below.

Centris ( Xerocentris ) griseoia, new species

DIAGNOSIS

Female runs to C. vanduzeei Cockerell in my key (1974), but
mandible tndentate (quadridentate in C. vanduzeei). meso-
scutum and scutellum with mixed black and pale erect hairs
(wholly pale in C. vanduzeei ), prepygidial brush blackish

brown (bright fulvous in C. vanduzeei) and distal portion of
median ridge of pygidium low and rounded (cariniform in
C. vanduzeei). Male unknown.

DESCRIPTION

HOEOTYPE FEMALE. Measurements (mm). Head width
5.18 (5.03-5.28); head length 3.79 (3.23-3.33) wing length
10.1 (10.2-10.3); total length 13.0 (13.0-15.5).

Head. 1.58 (1 .5 1-1 .62) times broader than long; occipital
margin nearly Hat in frontal view; inner orbits strongly con-
vergent above, upper frontal width 0.84 (0.82-0.84) times
lower frontal width. Maxillary palp five-segmented. Man-
dible slender, tndentate, second preapical tooth absent; inner
tooth prominent, triangular, and nearly perpendicular to long
axis of mandible. Labrum unmodified; shiny between sub-
contiguous, moderate punctures. Clypeus flattened, weakly
protuberant; disc slightly shiny, surface roughened, irregu-
larly, obliquely to transversely rugose. Frons and vertex mod-
erately shiny to shiny between fine subcontiguous to dense
punctures, but with extensive smooth areas in front of, and
laterad to, ocelli; gena less than half as wide as eye, mod-
erately shiny between fine, dense punctures. Interantennal
distance 2.92 (2.32-3.67) times antennal socket diameter;
antennocular distance 1.62(1 .65-2.00) times antennal socket
diameter; scape short, robust, scape length 0.73 (0.67-0.70)
times length of first flagellar segment; first flagellar segment
4.83 (5.06-5.50) times length of second. Ocelli, in frontal
view, well below occipital margin; interocellar distance 2.23
(2.20-2.24) times ocellar diameter; ocellocular distance 1 .50
(1.52-1.58) times ocellar diameter; ocelloccipital distance
2.50 (2.52-2.65) times ocellar diameter.

Thorax. Mesoscutum shiny between subcontiguous to dense
moderate punctures; scutellum narrowly shiny and impunc-
tate across anterior margin but mostly weakly shiny, inter-
spaces conspicuously roughened between dense, moderate
punctures; metanotum vertical, shiny between sparse, fine
punctures; mesopleuron moderately shiny, moderately tes-
sellate between subcontiguous to dense, fine punctures; meta-
pleuron similar but a little shinier. Propodeal disc smooth
and shiny between scattered minute to fine punctures; side
of propodeum less shiny, punctures dense to sparse, minute.
Metabasitibial plate about twice longer than broad, strongly
narrowed apicad, secondary plate poorly defined and merging
into primary plate, anterior depression small (all specimens
worn, details uncertain); scopal hairs largely plumose along
margins, mostly simple elsewhere.

Abdomen. Discs of all terga moderately shiny and weakly
tessellate between dense to sparse, fine punctures; pygidium
elongate-triangular, apex narrowly rounded; median eleva-
tion rounded and not attaining apex.

Color. Generally blackish brown, abdominal terga with
weak blue-green reflections; mandibles dark ferruginous; fla-
gellum and legs brown; tergal margins yellowish; tegula pi-
ceous; wings slightly brownish, veins and stigma brown.

Pilosity. Of head, thorax and abdomen fully erect, whitish,
with that of vertex largely brownish; pale and blackish hairs
about equally abundant on mesoscutum, scutellum and pro-

4 Contributions in Science, Number 347

Snelling: American Centridini

notal lobes; tergal hairs shorter and sparser, plumose; fourth
tergite with a few, and fifth with many, long, curled blackish
bristles; prepygidial fimbria dark; sixth tergite with dark bris-
tles and hairs; sternites with dark discal hairs and pale distal
hair bands, those of fourth and fifth segments broadly inter-
rupted; sixth segment dark pubescent; scopal hairs white;
some brownish bristles on protibia; bristles of protarsus and
thoracic venter brown; bristles on inner side of meso- and
metabasitarsi brownish ferruginous.

TYPE MATERIAL

Holotype female and three paratypes. MEXICO, GUER-
RERO: Iguala, 2000 ft. elev., 4 Feb. 1954 (R.R. Dreisbach).
Holotype and two paratypes in UKAN; one paratype in
LACM.

ETYMOLOGY

Modern Latin griseolus (gray) because of the grayish ap-
pearance of the thoracic dorsum.

DISCUSSION

Although C. griseola superficially resembles C. vanduzeei
Cockerell of Lower California, it is easily separated from that
species by the characters given above in the Diagnosis. Es-
pecially distinctive is the tridentate mandible. The only other
species of Xerocentris with tridentate mandible is C. pallida
W. Fox, of the arid western deserts. In that species, the max-
illary palp is four-segmented, the pygidial apex is truncate
and with a broad median ridge which extends to the apex,
the thoracic integument is dull, and the scopal hairs are all
plumose.

The unknown male of C. griseola probably resembles the
female in that the clypeus will be black and the thoracic hairs
will be a mixture of dark and light hairs.

Pollen samples from two specimens were examined by J.L.
Neff. His conclusion (personal communication) was that in
both cases, the samples were “. . . a mixture of Cercidium
and Hoffmanseggia (or possibly Caesalpinia) with the former
predominating.”

Subgenus Paracentris Cameron

Paracentris Cameron, 1903:235-236. Type-species: Para-
centris fulvohirta Cameron, 1903; original designation.
Penthemisia Moure, 1950:390. Type-species: Centris chilen-
sis Spinola, 1851; original designation.

Hemisia, subg. Penthemisia: Michener, 1951:3-4.

Centris, subg. Trichocentris Snelling, 1956:3. Type-species:
Centris rhodoleuca Cockerell, 1923; original designation.
Centris, subg. Paracentris: Snelling, 1974:5-7.

The North American species of Paracentris were treated in
1974; 14 species were recognized, four of which were known
from only one sex. Since then, I have seen opposite sexes for
three of these species. One previously described form, C.
atripes ferrisi Cockerell, was not considered in 1974 because
it was known only from the deficient original description. I

have now seen the type specimens. It seems appropriate,
therefore, to present a new key to the North American species
of Paracentris to accommodate these new data.

KEY TO NORTH AMERICAN PARACENTRIS

la. Female, antenna 12-segmented and basitibial plate

present on metatibia 2

b. Male, antenna 1 3-segmented and basitibial plate absent

from metatibia 15

2a. Pubescence of head, thorax, and legs entirely or pre-
dominantly black, that of mesepistemum wholly black

3

b. Pubescence of head, thorax, and legs entirely or pre-
dominantly pale, that of mesepistemum pale, at least

in part 7

3a. Pubescence of thoracic dorsum entirely black 4

b. Pubescence of thoracic dorsum largely pale 6

4a. Integument of abdominal tergites black; punctures of
second tergite uniformly distributed across disc, not
notably sparser along midline, their hairs simple or

plumose, interspaces polished or roughened 5

b. Integument of abdominal tergites dark blue; punctures
of second tergite dense at sides, distinctly more sepa-
rated (sometimes sparse) in middle, their hairs decum-
bent and simple, interspaces polished

laevibullata Snelling

5a. Punctures of disc of second tergite fine, little greater in
diameter than hairs arising from them, hairs fine, plu-
mose, suberect; interspaces of second tergite roughened

and dull nigrocaeru/ea F. Smith

b. Punctures of second tergite conspicuously greater in
diameter than coarse, simple, decumbent hairs arising
from them; interspaces of second tergite polished and

shiny aterrima F. Smith

6a. Large species, 14.5-18.5 mm long; pubescence of ver-
tex and pronotal lobes black; clypeal punctures sparse,
obscure apicad, median impunctate line evanescent

mexicana F. Smith

b. Smaller species, 12.5-14.5 mm long; pubescence of ver-
tex and pronotal lobe whitish; clypeus with sharply
defined median impunctate line, punctures sharply de-
fined zacateca Snelling

7a. Clypeal integument entirely blackish 8

b. Clypeal integument at least partially yellowish, orange

or red 1 1

8a. Discs of second and third terga with evenly spaced,

distinct punctures throughout 9

b. Discs of second and third terga polished and shiny,
with scattered fine punctures which become coarse and
distinct laterad, most of disc virtually impunctate . . .

fisheri Snelling

9a. First flagellar segment a little longer than following three
segments combined; abdominal terga black; punctures
of scutellum separated by much less than a puncture
diameter 10

b. First flagellar segment a little shorter than following
three segments combined; abdominal terga with defi-

Contributions in Science, Number 347

Snelling: American Centridini 5

nite metallic bluish reflections; punctures of scutellum
mostly more separated, with many interspaces of more

than a puncture diameter lanosa Cresson

10a. Median line of clypeus broad, smooth, shiny, and ini-
punctate; punctures of disc mostly well separated; hairs
at side of dorsal face of first tergum short, light brown
anteriorly, becoming short and fuscous toward distal

margin angustifrons Snelling

b. Median line of clypeus narrow, roughened and dull;
punctures of disc mostly subcontiguous; some hairs of
dorsal face of first tergum conspicuously long, erect,

plumose, and pale whitish harbisoni Snelling

1 la. Small species, 8.5-1 3.5 mm long; clypeus with distinct
impunctate median line, punctures elsewhere separated
by less than twice a puncture diameter (if clypeal punc-
tures somewhat sparse, scape, femora, and tibiae fer-
ruginous) 12

b. Larger species, 15.5-18.5 mm long; clypeus polished
between scattered punctures, usually separated by more
than twice a puncture diameter, and median impunc-
tate line undefined (scape, femora, and tibiae dark

brownish) caesa/piniae Cockerell

12a. Color of mandible, labrum, and clypeus usually not as
below; clypeus polished and shiny, median impunctate
line present and usually well defined; tergal discs mod-
erately to strongly shiny, with or without bluish reflec-
tions; scape, femora, and tibiae brown or red ... 13
b. Mandible (except apex), labrum, and clypeus dull or-
ange-ferruginous; clypeus usually contiguously punc-
tate, dull and without median impunctate line, but may
be moderately shiny and with narrow impunctate line;
tergal discs dull and closely tessellate, with dull bluish

reflections; scape, femora, and tibiae brown

ferrisi Cockerell

1 3a. Basal face of first tergum with abundant long, plumose,
white hairs, some of which extend onto dorsal face in

middle; terga with definite bluish reflections 14

b. Basal face of first tergum with sparse plumose, white
hairs, those in middle shorter and not extending to
dorsal face in middle; terga black, without bluish re-
flections cockerel/i W. Fox

14a. Mandible, labrum, clypeus, scape, femora, and tibiae
mostly or entirely bright ferruginous; apical margins of
second and third terga with short bands of appressed
pale hairs; first flagellar segment shorter than following

three combined rhodopus Cockerell

b. Mandible, scape, and legs brown, labrum and most of
clypeus yellowish; second and third terga with all hairs
blackish; first flagellar segment longer than following

three combined atripes Mocsary

1 5a. Clypeus wholly black; pubescence of mesepisternum

wholly blackish 16

b. Clypeus largely or entirely whitish, yellowish or red-
dish; pubescence of mesepisternum pale or dark . . 18

1 6a. Pubescence of thoracic dorsum pale 17

b. Pubescence of thoracic dorsum blackish

aterrima F. Smith

17a. Pubescence of pronotal lobe, and usually of entire lat-

eral pronotal area, pale; first flagellar segment 2.6-3. 1

times longer than second zacateca Snelling

b. Pubescence of entire lateral pronotal area, including
lobe, blackish; first flagellar segment 3. 8-3. 9 times longer

than second mexicana F. Smith

1 8a. Pubescence of head and thorax at least partially pale

19

b. Pubescence of head and thorax blackish

nigrocaeru/ea F. Smith

1 9a. Metabasitarsus much broadened toward apex, about
twice longer than greatest width; metafemur swollen

and about twice longer than wide 20

b. Metabasitarsus slender and parallel-sided, at least 3.5
times longer than wide; metafemur usually about three

times longer than wide, but may be swollen 22

20a. Punctures equally dense on mesoscutum and scutellum;
mesepisternum finely, closely punctate; clypeus pale

yellow 21

b. Punctures of mesoscutum much sparser than those of
scutellum; mesepisternum with scattered, obscure, fine

punctures; clypeus orange-ferruginous

ectypha Snelling

2 1 a. First flagellar segment longer than following three com-
bined; punctures of clypeal disc mostly separated by a
puncture diameter or more .... angustifrons Snelling
b. First flagellar segment a little shorter than following
three combined; punctures of clypeal disc mostly sep-
arated by less than 0.75 times a puncture diameter

harbisoni Snelling

22a. Discs of second to fourth tergites closely and sharply
punctate, punctures separated by no more than twice
a puncture diameter and clearly greater in diameter

than hairs arising from them 23

b. Discs of second to fourth tergites polished and shiny
between scattered, obscure punctures little greater in

diameter than hairs arising from them

fisheri Snelling

23a. Clypeus polished or not, punctures mostly separated
by two puncture diameters or less; ocellocular distance
no more than 1.25 times diameter of anterior ocellus;
smaller species, head width 3. 8-5. 3 mm, almost always

less than 5.0 mm 24

b. Clypeus polished, nearly impunctate; ocellocular dis-
tance at least 1.65 times diameter of anterior ocellus;
larger species, head width 5.0-6. 5 mm, almost always

more than 5.3 mm caesalpiniae Cockerell

24a. Clypeus slightly shiny to shiny, punctures ofdisc mostly
subcontiguous, usually with distinct impunctate me-
dian line, sometimes with median line roughened and
dull; abdominal terga without pale hairs beyond first
segment; legs medium to dark brown and metatibia

mostly dark pubescent 25

b. Clypeus polished between sparse punctures and median
impunctate line very broad and poorly defined; discs
(or at least apical margins) of second to sixth terga with
subappressed (suberect in metander), mostly simple,
pale hairs; legs almost always ferruginous and metatibia
at least largely whitish pubescent . . rhodopus Cockerell

6 Contributions in Science, Number 347

Snelling: American Centridini

25a. Face narrow, eye length at least 1 .56 (and usually more
than 1.60) times interocular distance at level of anten-
nal sockets; ocellocular distance no more than, and
usually less than, diameter of anterior ocellus ... 26
b. Face broad, eye length no more than 1.51 (and usually
less than 1.45) times interocular distance at level of
antennal sockets; ocellocular distance a little greater

than diameter of anterior ocellus 27

26a. Paraocular area, mandible (mostly) and underside of
scape (usually) yellow; clypeus closely punctate and me-
dian impunctate line narrow; abdominal terga with

bluish reflections lanosa Cresson

b. Paraocular area and underside of scape dark, mandible
ferruginous; clypeus with median impunctate line broad
and punctures mostly separated by about a puncture
diameter; abdominal terga black, without bluish re-
flections cockerelli W. Fox

27a. Median area of clypeus sharply roughened and dull,
contrasting to shiny, closely punctate areas on either
side; discs of abdominal terga two to five sharply tes-

sellate and slightly shiny ferrisi Cockerell

b. Clypeal disc shiny, with distinct smooth, impunctate
median line; discs of abdominal terga two to five shiny,
not obviously tessellate atripes Mocsary

Centris ( Paracentris ) angustifrons Snelling

Centris ( Paracentris ) angustifrons Snelling, 1966:13-14. 9.

The type locality for C. angustifrons is Huachuca Mountains,
Arizona and this species was based on a single female col-
lected nearly 80 years ago. A few additional specimens are
now available, including the previously unknown males. The
males are very similar to those of C. harbisoni, especially in
the shape of the metabasitarsus, but are easily separated by
the characteristics cited in the key above.

The females of C. angustifrons and C. harbisoni are even
more similar, a fact I had not fully appreciated when I pre-
pared my 1974 key. The present key brings the two species
to the same couplet; the differences between them are noted
there and should present few difficulties.

NEW RECORDS

MEXICO, SONORA: 13, Aduana, 1 5 Mar. 1 962 (L.A. Stange,
UCD); 699, 233, Rio Cuchuhaqui, 8 mi. S Alamos, 1-13 Apr.
1975 (A. Brewster; LACM), on Parkinsonia sp. (299, 13),
Fourquieria sp. (13), Cercidium sp. (299), and Prosopis sp.
(299).

Centris ( Paracentris ) aterrima F. Smith

Centris aterrima F. Smith, 1854:378. 3.

Centris (Paracentris) aterrima: Snelling, 1974:7, 8 (key).

NEW RECORDS

UNITED STATES, ARIZONA, Cochise Co.: 19, Rustler Park,
Chiricahua Mountains, 4 Aug. 1971 (LACM); 699, 1 mi. E
Douglas, 14 Aug. 1969 (J.G. and K.C. Rozen; AMNH); 19,
Yaqui Canyon area, 5370-5700 ft. elev., Huachuca Moun-

tains, 29 Aug. 1972 (R.R. Snelling; LACM), on Acacia an-
gustissima. PimaCo.: 299, Arivaca, 1 8 July and 24 Aug. 1974
(J.L. Neff; LACM). Santa Cruz Co.: 19, 13, Sycamore Can-
yon, near Ruby, 16-17 Aug. 1961 (J.C. Bequaert; LACM);
13, Nogales, 24 Aug. 1939 (R.H. Crandall; LACM); 13, Pat-
agonia, 24 Aug. 1955 (F.G. Werner and G.D. Butler; LACM);
19, White Rock Campground, 0.5 mi. S Pena Blanca, 9 Aug.
1972 (D.C. Frack; LACM); 19, Pena Blanca, 27 July 1972
(D.C. Frack; LACM). MEXICO, CHIAPAS: 19, 333, 12 mi.
W Ocozocoautla, 26 July 1953 (E.E. Gilbert and C.D.
MacNeill; UCB). OAXACA: 13, 4 mi. N Pochutla, 150 m
elev., 1 1 Oct. 1975 (J.L. Neff; LACM), on “Mint 71 15”; 399,
4 mi. W Zanatepec, 200 m elev., 16 Sept. 1975 (J.L. Neff;
LACM. NEFF), on Krameria revoluta. SINALOA: 19, 30 mi.
E Villa Union, 570 m elev., 20 Mar. 1980 (J.L. Neff; NEFF).

Centris ( Paracentris ) atripes Mocsary

Centris atripes Mocsary, 1899:254: 3.

Centris limbata Friese, 1899:44. 9. NEW SYNONYMY.
Centris atriventris W. Fox, 1899:68. 9 3. Preoccupied.
Centris Foxi Friese, 1900b:350. New name for C. atriventris
W. Fox.

Centris (Paracentris) atripes: Snelling, 1974:8, 9-10 (key,
distr., var.).

Friese described C. limbata from a single female collected
by G. Birkmann at Fedor, Lee County, Texas. The most
distinctive feature of this species was the presence of distinct
bands of pale hairs on the metasomal terga. Since no spec-
imens resembling the description have been subsequently
collected. C. limbata remained an enigma.

At my request, R.W. Brooks examined the type of C. lim-
bata in the Berlin Museum and he provided several photo-
graphs and sketches. A photograph of the lateral view of the
type specimen clearly shows that the abdomen is glued to
the thorax. The abdomen is distinctly fasciate in dorsal view
and a prominent, apically truncate pygidial plate, without a
secondary plate, is present. In fact, both the abdominal bands
and the pygidial plate are typical, not of Centris, but of a
Me/issodes male! The head and thorax of the type are just
as clearly those of a C. atripes female. Since the species was
described as a Centris, 1 consider these parts to be the true
type; the abdomen is extraneous. This restriction negates any
possible nomenclatural confusion within Me/issodes.

NEW RECORDS

COSTA RICA, GUANACASTE: 19, 8 km NW Liberia, 9
Feb. 1975 (G.R. Frankie; TAMU), on Cassia biflora, 0900.
GUATEMALA: 19, 8 mi. NE El Progreso, 8 July 1965 (A.
Raske and C. Slobodchikoff; UCB); 13, Jicara, 8 May 1931
(J. Bequaert; AMNH). UNITED STATES, OKLAHOMA:
19, Norman, Cleveland Co., “8/4 1949” (W.T. Nailon;
UKAN).

Centris ( Paracentris ) cockerelli W. Fox

Centris lanosa: W. Fox, 1899:69; Cockerell, 1906:97; Lutz
and Cockerell, 1 920:556; Timberlake, 1940: 1 38; Snelling,
1956:7 (in part, misidentifications).

Contributions in Science, Number 347

Snelling: American Centridini 7

Centris cockerelli W. Fox, 1899:68. 9.

Centris cockerelli resoluta Cockerell, 1923:76-77. 9 <3. NEW
SYNONYMY.

Centris lanosa lanosa: Snelling, 1966:6 (misidentification).
Centris lanosa resoluta: Snelling, 1966:6.

Centris (Paracentris) cockerelli cockerelli: Snelling, 1974: 1 0-
1 1.

Centris (Paracentris) cockerelli resoluta: Snelling, 1974: 1 1 .

For complete literature citations of this common species, see
Snelling (1974) under C. cockerelli and C. c. resoluta. This
is the species long improperly called C. lanosa. Although I
have attempted to maintain recognition of C. c. resoluta. 1
no longer believe this to be correct. In the females there are
two principal phenotypes: an eastern form with whitish to
yellowish clypeus and a western form with a reddish yellow
clypeus. There are no apparent differences to be seen in the
males associated with these females. The two female forms
intergrade continuously from western Texas and Tamaulipas
through New Mexico and Chihuahua. In my opinion these
variants are the end-points of a cline of variation and are
not worthy of formal separation.

Centris ( Paracentris ) ferrisi Cockerell

Centris atripes subsp. ferrisi Cockerell, 1924:49. “9” = <3!

This was described from two specimens from La Paz. Baja
California Sur, Mexico; although Cockerell stated the spec-
imens to be females, both are males. The types are in the
CAS. In my key (1974), the males will run to C. atripes.
Males differ from those of C. atripes. and other species of
similar appearance, in that the integument of the abdominal
tergites is dull and sharply roughened between minute, ob-
scure punctures; the clypeus, especially basad. is distinctly
roughened between well-separated punctures. The females,
too, will key to C. atripes and, as in the male, have the tergal
sculpture distinct, dulling the surface; the facial marks are
dull ferruginous rather than yellow as in C. atripes', in C.
ferrisi the first flagellar segment is as long as the following
three combined, shorter in C. atripes. The male clypeus is
usually ferruginous, but is yellow in some specimens.

In addition to the types of C. ferrisi I have seen the fol-
lowing, all from MEXICO, BAJA CALIFORNIA SUR: 1 <3,
Cabo San Lucas, 8-14 Sept. 1978 (J.P. and K.E. Donahue;
LACM); 19, 1<3, Canon de la Zorra, 260 m elev., 1 1 km W
Santiago, 4-5 Sept. 1977 (R.R. Snelling; LACM), on Par-
kinsonia aculeata\ 299, 4 km N Los Barriles, 10 m elev., 4
Sept. 1 977 (R.R. Snelling; LACM); 19, 3.7 mi. W La Burrera,
1400 ft. elev., 7-8 Oct. 1975 (R.R. Snelling; LACM), on
Antigonon leptopus ; 1 <3, 2.5 mi. SE La Huerta, 2200 ft. elev.,
8-9 Oct. 1968 (E.L. Sleeper and F.J. Moore; LACM); 19, 2
mi. S La Paz, 6 Aug. 1966 (J.A. Chemsak; UCB), on P.
aculeata ; 13, 7 mi. SW La Paz (J.A. Chemsak; UCB), on
Wislizenia refract a var. mamillata\ 19, 23 km W La Paz,
24-27 Apr. 1975 (E.M. and J.L. Fisher; LACM); 19, 68 km
S Loreto, Km 76 on Hwy 1, 29 July 1977 (D. Weismann
and C. Mullinex; CAS); 13, La Paz, 15 Sept. 1983 (R.R.
Snelling; LACM), on A. leptopus', 19, 25 mi. W La Paz, 30

Aug. 1959 (E.W. Radford and F.G. Werner; CAS); 19, 3 mi.
E San Pedro, 15 Sept. 1983 (R.R. Snelling; LACM), on A.
leptopus\ 13, 52 mi. NW La Paz, 15 Mar. 1980 (J.L. Neff;
NEFF), on Cercidium penmsulare: 19, La Laguna (Sierra de
la Laguna), 1829 m elev., 23 Oct. 1977 (D.E. and W.R.
Breedlove; CAS); 13, 11 mi. NE Todos Santos, 16 Sept. 1983
(R.R. Snelling; LACM), on A. leptopus ; 19, 25 km E Todos
Santos, near La Burrera, 1829 m elev., 21 Oct. 1977 (D.E.
and W.R. Breedlove; CAS); 13, Miraflores, 17 Sept. 1983
(R.R. Snelling; LACM), on A. leptopus ; 19, 1 mi. S Agua
Caliente, 17 Sept. 1983 (R.R. Snelling; LACM), on A. lep-
topus', 299, 5 mi. W San Ignacio, 13 Sept. 1983 (R.R. Snelling;
LACM), on Tephrosta tenella ; 299, same, except 19 Sept.
1983.

Centris ( Paracentris ) fisheri Snelling

Centris ( Paracentris ) fisheri Snelling, 1974:12. 3.

This was described from two males collected near San Ig-
nacio, Baja California Sur, Mexico. The female will go to,
and closely resembles, C. harbisoni Snelling in my key (1974)
to species of Paracentris. but differs immediately in that the
abdominal terga are polished and very sparsely punctate; in
C. fisheri the first flagellar segment is shorter than the scape
and shorter than the combined lengths of the second to fourth
flagellar segments; the first flagellar segment is longer in C.
harbisoni.

NEW RECORDS

MEXICO, BAJA CALIFORNIA SUR: 1 199, 533, 5 mi. W
San Ignacio, 13 Sept. 1983 (R.R. Snelling; LACM), on Te-
phrosia tenella', 599, 1033, same, except 19 Sept. 1983; 19,
19 km NW Mulege, 8 Sept. 1977 (R.R. Snelling; LACM), on
Hojfmanseggia sp.; 13, 35 mi. N Loreto, 5 Oct. 1975 (R.R.
Snelling; LACM), on Wislizenia refract a\ 533, 9.6 mi. N Lore-
to, 14 Sept. 1983 (R.R. Snelling; LACM), on Antigonon lep-
topus', 19, Estacion Microondas “Ligui,” 48 km S Loreto, 425
melev., 14 Sept. 1 983 (R.R. Snelling; LACM), on A. leptopus:
233, 68 km S Loreto, Km 76 on Hwy 1, 29 July 1977 (D.
Weismann and C. Mullinex; CAS).

Centris ( Paracentris ) harbisoni Snelling

Centris (Paracentris) harbisoni Snelling, 1974: 14-16. 9 3.

NEW RECORDS

MEXICO, BAJA CALIFORNIA SUR: 19, San Jose del Cabo,
11-16 Sept. 1967 (J. Chemsak, A. and M. Michelbacher;
UCB); 13, 52 mi. NW La Paz, 1 5 Mar. 1980 (J.L. Neff; NEFF)
on Cercidium peninsulare', 499, 106 km N La Paz, 18 Mar.
1980 (J.L. Neff; NEFF), on Krameria parvifolia.

Centris (Paracentris) laevibullata Snelling

Centris (Paracentris) laevibullata Snelling, 1966:17-18. 9;
Snelling, 1974:7 (key).

This species was described from a female from Orizaba, Vera

8 Contributions in Science, Number 347

Snelling: American Centridini

Cruz, Mexico (type locality) and another collected 14 mi.
NW Zitacuaro, Michoacan, Mexico. The male is unknown.

NEW RECORDS

MEXICO, DUR.ANGO: 19, Durango, 13 Aug. 1962 (A.E.
Michelbacher; UCB). JALISCO: 19, 3.5 mi. E [Rio] Mag-
dalena, 5 Sept. 1965 (A.R. Gillogly; LACM); 19, 25 mi. W
Guadalajara, 4700 ft. elev., 29 Sept. 1957 (H.A. Scullen;
ORSU); 19, 13 mi. NW Lagos de Moreno, 3 Sept. 1975 (J.L.
Neff; LACM). MICHOACAN: 19, 7.7 km NE Patzcuaro,
2088 m elev., 23 Sept. 1 976 (C.D. George and R.R. Snelling;
LACM). NUEVO LEON: 19, 18 mi. W Linares, 2700 ft.
elev., 26 Sept. 1975 (J.A. Powell, J. Chemsak, and T. Fried-
lander; UCB). SAN LUIS POTOSI: 19, 52 mi. S. Tamazun-
chali, 5700 ft. elev., 7 Oct. 1957 (H.A. Scullen; ORSU).

Centris ( Paracentris ) lanosa Cresson

Centris lanosa Cresson, 1872:284. 3.

Centris subhyalina W. Fox, 1899:69. 9.

Centris birkmanii Friese, 1899:44. <5 9.

Centris (Paracentris) lanosa: Snelling, 1974:8, 16-17 (key,
syn.).

NEW RECORDS

UNITED STATES, FLORIDA: 19, Austin Carey, Alachua
Co., 22-24 May 1975 (G.B. Fairchild; DPIF); 19, Gainesville,
Alachua Co. (DPIF); 19, Trenton, Gilchrist Co., 14 Apr. 1925
(D.M. Bates; DPIF). KANSAS: 19, 3 mi. S Sawyer, Pratt Co.,
16 June 1962 (C.D. Michener and party; UKAN), on Amor-
pha canescens. OKLAHOMA: 19, near Ardmore, Carter Co.,
3 June 1961 (Umv. Kans. Mex. Exped.; UKAN); 19, Cleve-
land Co., 31 May 1951 (H.L. Parker; UKAN); 19, Ft. Sill,
Comanche Co., 24 June 1974(T.E. Rogers; LACM). TEXAS:
599, Bastrop, Bastrop Co., 2 May 1969 (Brothers, Krueger,
Michener; UKAN).

Centris ( Paracentris ) mexicana F. Smith

Centris mexicana F. Smith, 1854:378. “9” = <5!

Centris ( Paracentris ) mexicana: Snelling, 1974:7, 8, 17-18
(key, tax., distr.).

NEW RECORDS

MEXICO, OAXACA: 499, 233, Tamazulapam, 6200 ft. elev.,
2 Sept. 1965 (S.J. Arnold; UCB), on Salvia sp.; 13, Monte
Alban ruins, 3 Aug. 1964 (H.V. Daly; UCB). VERA CRUZ:
19, 2 66, Cotaxtla Exp. Sta., Cotaxtla, 9 Aug. 1962 (D.H.
Janzen; UCB).

Centris ( Paracentris ) nigrocaerulea F. Smith

Centris nigro-caerulea F. Smith, 1874:369. 9 6.

Centris clypeata Friese, 1899:41. 9 <3. Preoccupied. NEW
SYNONYMY.

Centris (Paracentris) anthracina Snelling, 1966:14-17. 9 6.
NEW SYNONYMY.

DISCUSSION

I have examined the female and male type specimens of C.
nigro-caerulea and they are conspecific with C. clypeata and
C. anthracina. Smith’s types are from an unspecified locality
in Mexico; the female is herewith designated lectotype and
the male as paralectotype. Both are in the collections of the
British Museum (Natural History).

NEW RECORDS

MEXICO, CHIAPAS: 333, 733, 3 mi. W Navenchuac, 1-2
Apr. 1953 (R.C. Bechtel and E. I. Schlinger; UCB); 19, be-
tween Tuxtla [Gutierrez] and Chilapa, 23 Jan. 1 974 (S. Buch-
mann; LACM), on Cassia sp. JALISCO: 333, “env. de Gua-
dalajara,” 1903 and May 1913 (L. Diguet; MNHN).
MORELOS: 19, Cuernavaca, 6000 ft. elev., 7 Feb. 1954
(R.R. Dreisbach; LIKAN). OAXACA: 13, Oaxaca, 12 Oct.
1963 (A.E. and M.M. Michelbacher; UCB); 499, 20 mi. S
Taxco, 6 Feb. 1954 (R.R. Dreisbach; UKAN). SONOR. 4:
13, Aduana (near Alamos), 15 Mar. 1962 (F.D. Parker; UCD);
13, Rio Cuchuhaqui, 8 mi. S Alamos, 1-13 Apr. 1975 (A.
Brewster; LACM), on Parkinsonia sp. GUATEMALA: 399,
San Miguel Duenas, 5200 ft. elev., 1-7 Dec. 1975 (S.W.T.
Batra; LACM); 19, Ciudad de Guatemala, Dec. 1911 (W.M.
Wheeler; MCZ); 19, Universidad del Valle, Ciudad de Gua-
temala, 25 Nov. 1975 (S.W.T. Batra; LACM). PANAMA.
CHIRIQUI: 2 33, Potrero Grande, 28 Jan. 1981 (D.W. In-
ouye; ROUB).

Subgenus Acritocentris Snelling, 1974

Centris subg. Acritocentris Snelling, 1974:36. Type-species:
Centris (Melanocentris) ruthannae Snelling, 1966; original
designation.

In the years subsequent to the description of this subgenus
and my treatment of its component species, I have been able
to examine many more specimens than were then available.
As a result, I can now provide a more adequate account of
these species. The key given then (Snelling, 1974) will not
always work for the females and should be replaced by the
new one given below.

KEY TO SPECIES OF ACRITOCENTRIS

1 a. Female, antenna 1 2-segmented and basitibial plate pres-
ent 2

b. Male, antenna 13-segmented and basitibial plate absent

4

2a. Pubescence of thoracic dorsum whitish 3

b. All pubescence blackish ruthannae Snelling

3a. First abdominal tergite nearly im punctate toward apical
margin, especially in middle; median area of clypeus dull
and roughened but not rugose, laterad with rounded,
separated punctures; basal shiny area of clypeus broad-
ened in middle a/biceps Friese

Contributions in Science, Number 347

Snelling: American Centridini 9

b. First abdominal tergite closely, though minutely, punc-
tate to apical margin; median area of clypeus rugose and
dull, laterad with coarse, close, elongate punctures; basal
shiny area narrow across entire clypeal base

agameta Snelling

4a. Thoracic dorsum dark pubescent 5

b. Thoracic dorsum pale pubescent 6

5a. Labrum, clypeus, paraocular area, and supraclypeal area

all yellow-maculate ruthannae Snelling

b. Entire face black satana, new species

6a. Face marks whitish, absent from paraocular and supra-
clypeal areas, clypeus broadly blackish along lateral and
basal margin, disc smooth and shiny . . . a/biceps Friese
b. Face marks yellowish, usually present on paraocular and
supraclypeal areas, lateral and basal margins of clypeus
narrowly, or not at all. blackish, disc dull and roughened
agameta Snelling

Centris ( Acritocentris ) albiceps Friese

Centris mexicana var. albiceps Friese, 1899:289. 3.

Centris (Melanocentris) strawi Snelling, 1 966:27-28. <5. NEW
SYNONYMY.

Centris ( Acritocentris ) strawi Snelling, 1974:37, 38 (in part)
(key, distr.).

1 suggested (1974) that C. mexicana var. albiceps might be
an older name for C. strawi. Friese based this name on a
specimen from an unknown locality in Mexico. The type is
in the Berlin Museum and was examined at my request by
R.W. Brooks who confirmed the above synonymy. The type
is not so marked, and bears two labels: “116 59/Mexico”
and “ Centrist mexicana/ v . albiceps/Friese 1898.”

The females which in 1974 1 believed to belong to C.
albiceps (as C. strawi) are now known to be those of C.
agameta. The true females of this species are very similar to
those of C. agameta. but the disc of the clypeus, although
dull and roughened, lacks the characteristic irregular rugulae
present in C. agameta. The first two abdominal terga are
much more weakly punctate toward the margins in C. al-
biceps. and in C. albiceps females the labrum and posterior
pronotal lobes are without pale hairs.

NEW RECORDS

MEXICO, CHIAPAS: 329, 9<5<5, Municipio Chiapo de Corzo,
El Chorreadero, 753 m elev., 1 Nov. 1976 (D.E. and J.A.
Breedlove; CAS); 12, Municipio Motozintla, betw. Motozin-
tla and Mazapa, 1219 m elev., 5 Oct. 1976 (D.E. and J.A.
Breedlove; CAS). HIDALGO: 229, Zimapan, 6400 ft. elev.,
8 Oct. 1957 (H.A. Scullen; ORSU). MICHOACAN: 229, Lake
Patzcuaro, 6800 ft. elev., 2 1 Sept. 1 957 (H.A. Scullen; ORSU);
13, 10 mi. N Morelia, 5900 ft. elev., 28 July 1962 (Univ.
Kans. Mex. Exped.; UKAN), on Leguminoseae; 12, 233, Ca-
rapan, 1 Sept. 1962 (D.H. Janzen; UCB). NUEVO LEON:
12, 30 mi. N Linares, 1500 ft. elev., 11 Oct. 1957 (H.A.
Scullen; ORSU). OAXACA: 13, 22 mi. SE Oaxaca, 5700 ft.
elev., 2 Sept. 1957 (H.A. Scullen; ORSU); 12, 5 mi. NW
Totolapan, 3800 ft. elev., 6 July 1953 (Univ. Kans. Mex.

Exped.; UKAN) on Lonchocarpus\ 522, 4 mi. N Totolapan,
1849 m elev., 15 Sept. 1975 (J.L. Neff; LACM, NEFF), on
“ Cassia 7064”; 12, 2 mi. S Totolapan, 15 Sept. 1975 (J.L.
Neff; LACM), on “ Cassia 7064”; 13, near Las Margaritas,
1400 m elev., 15 Sept. 1975 (J.L. Neff; LACM). PUEBLA:
13, 3 mi. NW Petlalcingo, 4600 ft. elev., 5 Sept. 1972 (Byers
and Thornhill; UKAN); 12, 22 km NW Izucar de Matamoros,
1158m elev., 2 1 Sept. 1 976 (C.D. George and R.R. Snelling;
LACM), on Cassia laevigata. QUERETARO: 12, 10 mi. S
Jet. Hwy 55 and 45, 30 Aug. 1963 (Scullen and Bollinger;
ORSU). SAN LUIS POTOSI: 12, 9 mi. E Ciudad [del] Maiz,
3975 ft. elev., 23 July 1962 (Univ. Kans. Mex. Exped.;
UKAN); 12, 5 mi. E Ciudad [del] Maiz, 4700 ft. elev., 22
Aug. 1954 (Univ. Kans. Mex. Exped.; UKAN); 12, 1 133, 6
mi. W Guadalcazar, 2 Sept. 1975 (J.L. Neff; LACM, NEFF),
on “mint 7031.” TAMAULIPAS: 3622, 38 mi. N El Mante,
1050 ft. elev., 1 1 Oct. 1957 (H.A. Scullen; ORSU); 1622, 9,
12, and 24 mi. S [Ciudad] Victoria, 11 Oct. 1957 (H.A.
Scullen; ORSU); 422, 15 mi. SW Ciudad Victoria, 5000 ft.
elev., 19 Sept. 1976 (J.A. Chemsak et al.; UCB); 2022, 35
mi. S Villagran. 15 Sept. 1977 (Chemsak and Michelbachers;
UCB), on Solanum\ 12, 7 mi. S Villagran, 26 Sept. 1975 (J.
Powell et al.; UCB); 12, 1 5 mi. NE Juamava, 2500 ft. elev.,
19 Sept. 1976 (J. Chemsak et al.; UCB). VEIL 4 CRUZ: 13,
3 mi. E Acultzingo, 1 600 m elev., 2 1 Aug. 1 977 (E.I. Schlin-
ger; UCB); 12, 233, E Citlaltepetl, 6000 ft. elev., 25 June 1964
(L.W. Swan; CAS).

Centris ( Acritocentris ) agameta Snelling

Centris ( Acritocentris ) agameta Snelling. 1974:37-38. 3.

NEW RECORDS

MEXICO, GUERRERO: 12, Acapulco, 6 Aug. 1954 (Univ.
Kans. Mex. Exped.; UKAN). JALISCO: 322, 13, Hwy 15,
0.2 mi. from Nayarit State line, 11 Aug. 1963 (D. Byers;
UKAN); 322, 6 mi. NE El Rincon, 3 Aug. 1971 (E.M. Fisher;
LACM); 12, Puente Grande, 5000 ft. elev., 20 Aug. 1954
(Univ. Kans. Mex. Exped.; UKAN); 12, Barra de Navidad,
6 Sept. 1966 (E.M. Fisher; UKAN); 13, 3 mi. NW Tequila,
4000 ft. elev., 15 July 1953 (Univ. Kans. Mex. Exped.;
LACM), on Vitex pyramidata: 13, 8 km W Tequila, 18 July
1951 (P.D. Hurd; UCB); 13, 7 km N Tequila, 6 Sept. 1975
(J.L. Neff; LACM); 13, 14 mi. NW [Rio] Magdalena, 3500
ft. elev., 19 July 1953 (Univ. Kans. Mex. Exped.; UKAN),
on Vitex pyramidata. MORELOS: 12, 14 mi. S Cuernavaca,
3 Aug. 1954 (Univ. Kans. Mex. Exped.; UKAN); 12, 12 mi.
E Cuernavaca, 4300 ft. elev., 14 Aug. 1954 (Univ. Kans.
Mex. Exped.; UKAN); 12, 8.7 mi. NE Yautepec, 4400 ft.
elev., 15 Aug. 1963 (Ordway and Roberts; UKAN); on Cu-
phea sp.; 13, 6.7 mi. S Yautepec, 29 July 1963 (Naumann
and Willis; UKAN). NAYARIT: 322, Ixtlan del Rio, 5 Aug.
1963 (P. Fonda-Bonardi; LACM). OAXACA: 322, 13, Salina
Cruz, 7 Sept. 1965 (D.H. Janzen; UKAN); 433, 20 mi. E El
Camaron, 7 Aug. 1956 (J.W. MacSwain; UCB). PUEBLA:
12, 13, 7 mi. N Izucar de Matamoros, 4450 ft. elev., 19 Aug.
1962 (Univ. Kans. Mex. Exped.; UKAN), on Cuphea sp.
SAN LUIS POTOSI: 13, 9 mi. E Ciudad [del] Maiz, 3975

10 Contributions in Science, Number 347

Snelling: American Centridini

ft. elev., 23 July 1962 (Univ. Kans. Mex. Exped.; UKAN).
TAMAULIPAS: 433, 40 km N Soto la Marina, 5 Sept. 1975
(E.M. and J.L. Fisher; LACM). SINALOA: 12, 733, 31 km
N Mazatlan, 76 m elev., 29 Sept. 1976 (C.D. George and
R.R. Snelling; LACM), on Antigonon leptopus.

Centris ( Acritocentris ) ruthannae Snelling

Centris ( Melanocentris ) ruthannae Snelling, 1 966:28-30. <3 2.
Centris ( Acritocentris ) ruthannae: Snelling, 1 974:37, 40 (distr.).

This species has been previously known only from southern
Arizona.

NEW RECORDS

MEXICO, CHIHUAHUA: 12, Temoris, 13 Sept. 1970 (T.A.
Sears et al.; UCD). SINALOA: 12, 2.5 mi. W Concordia, 25
Sept. 1977 (J.A. Chemsak, A. and M. Michelbacher; UCB).
SONOR.A: 12, Bahia San Carlos, 20 July 1965 (D.S. Verity;
LACM); 12, 20 km N Guaymas (Km 141), 16 Aug. 1979
(E.M. Fisher; LACM); 12, 13, Navajoa, 12 Sept. 1964 (A.E.
and M.M. Michelbacher; UCB), on morning glory, 0750-
0800; 222, 233, Cerro Masiaco, ca. 1 1 road mi. SSE Baca-
bachi, ca. 750 ft. elev., 29 Aug. 1 976 (J.P. and K.E. Donahue;
LACM); 13, 37 km SE Bacabachi (Km 90), 14 Aug. 1976
(E.M. Fisher; LACM), on Caesa/pinia sp.

Centris ( Acritocentris ) satana, new species

Figures 1-4

DIAGNOSIS

Male unique within Acritocentris by the entirely blackish
integument and pubescence; female unknown.

DESCRIPTION

HOLOTYPE MALE. Measurements (mm). Head width
6.15 (5.33-5.74); head length 4.56 (4.00-4.10); wing length
13.5 (13.0-13.5); total length 1 8.0 ( 1 5.0-1 7.0).

Head. 1.35 (1.33-1.40) times broader than long; occipital
margin, in frontal view, very weakly convex and barely raised
above level of tops of eyes; ocelli well below occipital margin
in frontal view; inner orbits weakly convergent above, upper
frontal width 0.91 (0.85-0.87) times lower frontal width.
Mandible slender, tridentate. Labrum about two-thirds as
long as broad, apical margin narrowly rounded; disc shiny,
moderately rugosopunctate to contiguously punctate. Disc of
clypeus obtusely raised along midline over basal one-half,
integument weakly to moderately shiny, surface with dense,
moderate punctures or with very irregular, fine, anastomos-
ing rugules; punctures on side of clypeus subcontiguous, elon-
gate. Remainder of head shiny between fine to moderate,
dense to subcontiguous punctures (sparse mesad in ocellocu-
lar area), but sparse, minute and obscure over most of gena.
Interantennal distance 2.35 (2. 1 3-2.33) times antennal sock-
et diameter; antennocular distance 1.00 (0.87-0.97) times
antennal socket diameter; scape moderately stout, 2.36 (2.29-
2.42) times longer than wide, scape length 1 .06 ( 1 .08-1 . 1 4)

times length of first flagellar segment; first flagellar segment
shorter than length of following three segments combined,
4.00 (3.22-4.06) times length of second; interocellar distance
1 .79 ( 1 .6 1-1 .90) times diameter of anterior ocellus; ocellocu-
lar distance 1 .29 (0.94-1 . 1 6) times diameter of anterior ocel-
lus; ocelloccipital distance 2.21 (1.97-2.09) times diameter
of anterior ocellus.

Thorax. Mesoscutum shiny between subcontiguous to
dense, moderate punctures; scutellum similar, but punctures
contiguous to subcontiguous and slightly elongate. Mesepi-
sternum moderately shiny, punctures dense, fine and oblique.
Metepisternum shiny, punctures fine and sparse anteriorly,
becoming dense to subcontiguous along posterior margin.
Metafemur stout, more than twice longer than thick; meta-
basitarsus about three times longer than broad.

Abdomen. Tergal discs shiny between dense to sparse, fine
to moderate punctures which become sparser mesad and on
succeeding segments; punctures in apical zone minute. Py-
gidial plate narrowly truncate to shallowly notched at apex.

Terminalia. Seventh sternite (Fig. 1) with lateral margins
of distal process convex, apical margin concave, with definite
mesoapical row of short hairs. Eighth sternite (Fig. 2) with
distal process slender, compressed; hairs numerous, long,
coarsely plumose. Dorsal process of gonocoxite (Fig. 4) long,
slender, nearly straight, slightly hooked at apex; gonostylus
slender, setae long and coarse along basal margin, shorter
and finer distad.

Pilosity. Hairs uniformly dark brown to blackish.

Color. Blackish brown, abdominal terga with strong bluish
reflections. Mandible, antenna and legs reddish brown, fem-
ora more conspicuously reddish. Wings dark brown, veins
and stigma black.

TYPE MATERIAL

Holotype male: Tepoxtlan, Morelos, MEXICO, 28 Aug. 1964
(E. Fisher and D. Verity), in Natural History Museum of Los
Angeles County. Paratypes, MEXICO: 13, Carapan, Mi-
choacan, 1 Sept. 1962 (D.H. Janzen; UCB); 13, 3.4 km NW
Tequila, 1295 m elev., Jalisco, 6 Sept. 1976 (C.D. George
and R.R. Snelling; LACM); 13, 5 mi. W Durango, Durango,
21 July 1964 (J. Powell; UCB); 233, 25 mi. W Hidalgo del
Parral, 6800 ft. elev., Chihuahua, 15 July 1964 (J.A. Chem-
sak, J. Powell; UCB); 13, “Guanajuato, Mexique” (Duges;
MNHN). UNITED STATES: 13, Atascosa Mts, 4800 ft. elev.,
5.6 mi. W Pena Blanca Lake, Santa Cruz Co., Arizona, 15
Aug. 1974 (T.J. Zavortink; TJZ), on Acacia angustissima,
1145-1200.

ETYMOLOGY

From Hebrew, literally an enemy or an evil spirit; in modern
zoological nomenclature an epithet for a species of black or
dark color or threatening appearance.

DISCUSSION

The wholly black pubescence will immediately separate C.
satana from C. agameta and C. albiceps, since both have

Contributions in Science, Number 347

Snelling: American Centridini 1 1

Figures 1-8. Male seventh and eighth sternites and genitalia (ventral and dorsal views), respectively, of: 1-4, Centris ( Acritocentris ) satana,
scale line = 1.00 mm; 5-8, C. (C.) aethiocesta. scale line = 0.50 mm.

12 Contributions in Science, Number 347

Snelling: American Centridini

pale dorsal thoracic pubescence. The black, rather than yel-
low, clypeus will distinguish C. satana from C. ruthannae.
Additionally, males of C. ruthannae have metallic blue re-
flections on the abdominal terga and the hairs of the second
and third segments are conspicuously plumose.

The female of C. satana is unknown, but very likely will
be black-haired like the male and will have a similarly black
abdomen, without metallic blue reflections.

Subgenus Exallocentris Snelling

Centris subg. Exallocentris Snelling, 1974:35. Type-species:
Centris ( Melanocentris ) anomala Snelling, 1966; mono-
basic and original designation.

Although I had originally placed this monotypic subgenus
near Melanocentris, Neff and Simpson (1981) have rightly
pointed out its affinities to Paracentris. In Exallocentris the
elaiospathe of the female pro- and mesobasitarsi is replaced
by dense pads of fine-branched setae and the secondary basi-
tibial plate is sharply marginate and projects over the primary
plate. Males differ from those of Paracentris in lacking
branched setae on the gonocoxite at the base of the gono-
stylus; a conspicuous pygidial plate is present, the innermost
mandibular tooth is truncate, and the lateral ocellus is sep-
arated from the inner eye margin by about its own diameter.
Males resemble those of Melanocentris, but the scutellum is
not bilobed on its dorsal surface and the upper inner man-
dibular carina ends near the base of the innermost tooth.

Centris ( Exallocentris ) anomala Snelling

Centris ( Melanocentris ) anomala Snelling, 1966:31-32. 6 2.
Centris (Exallocentris) anomala: Snelling, 1974:35-36 (tax.).

This species has previously been known only from the vi-
cinity of Guadalajara, Jalisco, Mexico. I have recently seen
a few specimens of the cleptoparasitic bee, Mesoplia dugesi
(Cockerell), which were collected with the type series of C.
anomala. One of the specimens bears the following note:
“Ericrocis??/Very evasive, flying low/Parasite ?? of the/big
Hemisia? not numerous.”

NEW RECORDS

MEXICO, HIDALGO: 18, 22 mi. SW Actopan, 6800 ft. elev.,
27 Aug. 1962 (Ordway and Marston; UKAN). JALISCO:
12, 103 mi. NE Guadalajara, 6200 ft. elev., 1 Oct. 1957 (H. A.
Scullen; ORSU); 1033, 8 mi. NE Chapala, 5150 ft. elev., 30
Sept. 1957 (H.A. Scullen; ORSU); 12, “env. de Guadalajara”
(L. Diguet; MNHN). MICHOACAN: 222, 33 km NE Arteaga
(Hwy 37, km 242), 980 m elev., 10 Nov. 1976 (E. Fisher
and P. Sullivan; LACM). OAXACA: 12, 9 mi. SE Nochixtlan,
7 Nov. 1963 (R.F. Smith; UCB); 333, Monte Alban, 12 Oct.
1963 (A.E. and M.M. Michelbacher; UCB). PUEBLA: 9 33,
“env. de Tehuacan” (L. Diguet; MNHN). ZACATECAS: 12,
10 mi. S Jalpa, 18 Sept. 1970 (R.M. Bohart; UCD).

Subgenus Xanthemisia Moure

Xanthemisia Moure, 1945:401. Type-species: Centris bicolor

Lepeletier, 1841; monobasic and original designation.

Hemisia subg. Xanthemisia: Michener, 1951:2, 3, 5-6.
Centris subg. Xanthemisia: Michener, 1954:140. Snelling,
1974:2, 3.

This subgenus was described for a small group of species
characterized, in the females, by possessing a low, blunt tooth
on the inner surface of the mandible, near the base of the
apical tooth. In this sex, too, the pygidial plate is abruptly
narrowed a short distance beyond the secondary plate so that
the distal portion is narrow and parallel-sided. Males of Xan-
themisia possess giant branched setae near the base of the
gonostylus, which is much broadened on a vertical plane and
the maxillary palp is four-segmented.

KEY TO NORTH AMERICAN XANTHEMISIA

la. Male, antenna 1 3-segmented, basitibial plate absent . .

2

b. Female, antenna 1 2-segmented, basitibial plate present

4

2a. Labrum and clypeus wholly black; pubescence of dorsum
of scutellum yellow; erect hairs of mesoscutum longer

than interocellar distance 3

b. Labrum, entirely, and clypeus mostly, yellow; pubes-
cence of dorsum of scutellum black; erect hairs of meso-
scutum shorter than interocellar distance

rubella F. Smith

3a. Pubescence of mesoscutum wholly yellow; abdomen
black and wholly blackish pubescent; ocellocular dis-
tance less than diameter of anterior ocellus

lutea Friese

b. Mesoscutum with interalar band of blackish pubescence;
abdomen reddish, with golden brown pubescence; ocel-
locular distance greater than diameter of anterior ocellus

carolae Snelling

4a. Clypeus immaculate and pubescence of dorsum of scu-
tellum yellow; abdomen blackish or dark brown; scopal

hairs dark 5

b. Clypeus usually with a pair of preapical spots; pubes-
cence of dorsum of scutellum dark brown; abdomen dull

ferruginous; scopal hairs pale rubella F. Smith

5a. Mesoscutum wholly yellow pubescent; abdomen black-
ish, with blackish pubescence; first flagellar segment
longer than following three combined .... lutea Friese
b. Mesoscutum wholly blackish pubescent; abdomen dark
brown with reddened apical zone and yellowish to yel-
lowish red pubescence; first flagellar segment no longer
than following three combined carolae Snelling

Centris ( Xanthemisia ) carolae Snelling

Centris ( Xanthemisia ) carolae Snelling, 1966:24-25. 3.

This species has been previously known only from the unique
male type from Tuxtla Chico, Chiapas, Mexico. A few fe-
males are now available. They may be separated from our
other species by the largely blackish pubescence on the tho-
racic dorsum, yellowish only on the scutellum and meta-

Contributions in Science, Number 347

Snelling: American Centridini 13

notum. The pubescence of the abdomen is dull reddish over
the brownish integument.

NEW RECORDS

EL SALVADOR: 29$, Cerro Verde, 6800 ft. elev., 29 June
1963 (M.E. Irwin and D.Q. Cavagnaro; UCB); 1$, same data
(CAS). COSTA RICA: 1$, “El Fuente,” 8 July 1937 (A. Al-
faro No. 177; AMNH).

Centris ( Xanthemisia ) lutea Friese

Centris lutea Friese, 1899:43. <3 9.

Centris (Xanthemisia) lutea : Michener, 1954:140 (distr.).

Among the North American Centris, this species is easily
known by the wholly blackish pubescence, except that of the
thoracic dorsum which is bright lemon-yellow. Michener
(1954) recorded this primarily South American species from
Panama.

NEW RECORDS

MEXICO, CHIAPAS: 2<3<3, 46 km N Chilpacingo, 580 m
elev., 4-6 Aug. 1977 (E.M. Fisher and P. Sullivan; LACM).
JALISCO: 1<3, 48 mi. N Guadalajara, 8 Sept. 1966 (R.J.
Hamton; LACM). OAXACA: 1<3, 20 mi. E El Camaron, 21
July 1956 (J.W. MacSwain; UCB). COSTA RICA, GUA-
NACASTE: 19, [Hacienda] Comelco, 8 km NW Bagaces, 21
Jan. 1 972 (P.A. Opler; UCB), on Bvrsonima sp.; 19, Hacienda
Comelco, 24 km NW Canas, 7 Feb. 1972 (E.R. Heithaus;
LACM), on Bvrsonima crassifolia, 0730-0830. SAN JOSE:
19, San Jose, no further data (USNM).

Centris ( Xanthemisia ) rubella F. Smith

Centris rubella F. Smith, 1854:372. 9.

Hemisia ( Xanthemisia ) rubella: Michener, 1951:6 (tax.).

The ferruginous abdomen and maculate female clypeus will
readily separate C. rubella from all other species of Xanthe-
mesia. This species has previously been known only from
South America.

NEW RECORDS

PANAMA, CANAL ZONE: 19, Barro Colorado Island, 19
Aug. 1968 (L.S. Kimsey; LACM), “wood nester”; 399, same
locality, 27 Apr., 3 May, 5 May 1980 (K.E. Steiner; LACM,
UCD), on Byrsonima crassifolia ; 19, Gatun, 20 May 1980
(K.E. Steiner; UCD), on B. crassifolia ; 399, same locality, 30
Oct. and 3 Nov. 1977 (K.E. Steiner; UCD), on Stigmaphyllon
hypargyreum.

Subgenus Centris Fabricius

Centris Fabricius, 1804:354. Type-species: Apis haemor-
rhoidalis Fabricius, 1775; designated by Internatl. Comm.
Zool. Nomencl., op. 567, 1959.

Hemisia Klug, 1807:227. Type-species: Apis haemorrhoi-
da/is Fabricius, 1 775; designated by Cockerell, 1906.
Centris subg. Cyanocentris Friese, 1900b:243. Type-species:
Apis versicolor Fabricius, 1775; designated by Sandhouse,
1943.

Centris subg. Poecilocentris Friese, 1900b:244. Type-species:
(Centris [Poecilocentris] fasciateIla Friese, 1900) = Centris
fasciata F. Smith, 1854; designated by Sandhouse, 1943.

Species in this subgenus normally have the abdomen metallic
blue or blue-green in both sexes and with pale tergal maculae
in the male and often in the female as well. In a few species,
such as C. inermis Friese, the abdomen is largely ferruginous
and in others, such as C. eisenii W. Fox, both sexes have
much of the terga taken up by yellow bands. In both sexes,
too, the mandibles, labrum, and clypeus are conspicuously
maculate, the clypeal maculation of the female usually in an
inverted T-shape or, more rarely, an inverted Y-shape. Both
sexes have five-segmented maxillary palps, the female with
slender, usually quadridentate, mandible, and a distinct sec-
ondary basitibial plate with an overhanging margin. Males
possess giant branched setae along the inner margin of the
styliform apical process of gonocoxite.

Some of the most exasperating taxonomic problems in
Centris are encountered in the nominate subgenus. Many of
the species are widely distributed and may be locally abun-
dant. The more widely distributed species tend to exhibit a
variety of phenotypes throughout their ranges, phenotypes
that are often strikingly different from one another. Many of
these variant populations have been given formal names. As
a rule, these divergent populations represent selected points
along a dine of variation and it is impossible to fix to any
of these a discrete distribution and set of identifying char-
acteristics.

The situation becomes more complex when a species is
distributed through the islands of the Caribbean. Since the
ranges are disjunct, the various insular populations tend to
breed true. Even here, however, there usually are discrete
clinal trends, proceeding from one end of the island distri-
bution to the other. But, a great deal of collecting must be
done in the Caribees before any understanding can be gained
of these forms.

Further complicating the taxonomy of this subgenus is the
fact that some species perhaps most, possess metanders, i.e.,
males which are unusually large and robust, with very stout
legs, and much more extensively maculate than normal males.
Metanders may be easily assumed to represent a different
species. In general, all large, robust, richly marked males
should be assumed to be probable metanders.

Yet another difficulty is that at least one species (C. iner-
mis) is dichromatic in both sexes. There is a color phase with
red abdomen and one with metallic blue-green abdomen.
There is some evidence that one phase tends to fly earlier
than the other, but there is a broad temporal overlap. Not
surprisingly, the two color phases have been thought to be
separate species. There are very likely more such cases in
this subgenus.

The following key is for those species which occur in Cen-

14 Contributions in Science, Number 347

Snelling: American Centridini

tral America; it does not include C. errans W. Fox, a species
found only in Florida.

KEY TO CENTRAL AMERICAN CENTRIS, S. S I R.

1 a. Male, antenna 1 3-segmented and basitibial plate absent

2

b. Female, antenna 12-segmented and basitibial plate

present 9

2a. Disc of fourth tergum with simple, usually blackish,
hairs only (plumose hairs may be present on apical

zone) 3

b. Disc of fourth tergum with abundant, usually whitish,
suberect plumose hairs which extend onto apical zone
(may be only a band across segment basad of apical

zone, often extended cephalad in middle) 7

3a. Second to sixth abdominal terga with broad preapical
yellow bands which cover most of each segment, nar-
row basal area bluish to rufescent, apical margin trans-
parent; sterna largely rufescent 4

b. Second to sixth abdominal terga largely bluish or fer-
ruginous, second usually with lateral spots only, fol-
lowing segments with lateral spots, narrow transverse
bands (often interrupted and usually hidden under mar-
gin of preceding segment) or immaculate; if bands are
present, they are dull reddish, usually interrupted in
middle, and fourth tergum with broad area along distal

margin with abundant plumose hairs 5

4a. Facial marks whitish; first flagellar segment at least
1.25 x scape length; hairs along apical band of fourth

tergum dark, mostly simple eisenii W. Fox

b. Facial marks distinctly yellowish; first flagellar segment
about as long as scape, rarely as much as 1.1 x scape
length; apical band of fourth tergum with abundant

plumose pale hairs aethyctera Snelling

5a. Ground color of abdominal segments ferruginous, sec-
ond segment maculate at side, rarely with a complete
or narrowly interrupted subbasal band; transverse bands
of remaining segments, if present, hidden under margin

of preceding segment 6

b. Ground color of first terga, and of sterna, bluish, sub-
lateral spots or narrowly interrupted bands on one or
more segments; rarely, second to sixth segments with
broad, transverse, median yellowish or rufescent bands

inermis Friese (part)

6a. Flairs of thoracic dorsum black-tipped

obscurior Michener

b. Hairs of thoracic dorsum uniformly ochreous

inermis Friese (part)

7a. Pubescence of thoracic dorsum uniformly ochreous to
ferruginous, no blackish hairs present; smaller species,
head width less than 6. 1 mm and usually less than 6.0

mm; (metander unknown) 8

b. Pubescence of thoracic dorsum pale anteriorly and pos-
teriorly, a broad interalar brown to black band present,
often replacing much pale pubescence, especially on
scutellum; larger species, head width at least 5.5 mm
and usually over 6.3 mm (metander larger, robust, legs

and abdomen abundantly yellow maculate or both areas
largely ferruginous with limited yellow maculae) . .

flavifrons group (see Text)

8a. Apical two or three abdominal terga reddish; mandible
mostly yellowish on outer face; pubescence of thoracic

dorsum usually ochreous adani Cockerell

b. Fourth and fifth, usually sixth, abdominal tergites blue,
except colorless margins; mandible with small pale spot
near base; pubescence of thoracic dorsum usually dark

ferruginous decolorata Lepeletier

9a. Terga 2-5 each with entire transverse yellow fascia

10

b. Terga 2-5 without yellow fasciae 11

10a. Hairs of thoracic dorsum black-tipped; scape yellow
beneath; fourth tergum with erect, plumose, white hairs;

HW less than 6.0 mm aethyctera Snelling

b. Hairs of thoracic dorsum not black-tipped; scape im-
maculate beneath; fourth tergum with simple hairs only;

HW at least 6.3 mm eisenii Fox

1 la. Thoracic dorsum with conspicuous interalar band of
black pubescence which may cover much of mesoscu-
tum (sometimes reduced to median patch), sharply

contrasting with pale scutellar hairs 12

b. Thoracic dorsum without interalar band of black pu-
bescence; scutellar hairs not contrastingly colored

14

12a. Larger species, head width greater than 6.5 mm, but if
as little as 6.25 mm, scopa is principally black; disc of
clypeus, in profile, distinctly convex basad and flat-
tened distad (Fig. 16) 13

b. Smaller species, head width 5.95-6.25 mm; scopa pale,
with golden brown hairs apicad on metabitarsus; disc
of clypeus, in profile, very weakly convex basad (Fig.

15) aethiocesta, new species

13a. Larger species, head width 7.17-7.7 mm; scopa with
pale hairs at least anterobasally on metatibia, often
largely pale; pygidial plate and basitibial plate as in

Figs. 38 and 29 flavifrons (Fabricius)

b. Smaller species, head width 6.27-7.2 mm; scopa black,
a few posteroapical hairs on metabasitarsus pale at tips;
pygidial and basitibial plates as in Figs. 39 and 30

flavofasciata Friese

14a. Abdomen mostly ferruginous, one or more terga often
partially bluish or greenish toward apical margin ..15
b. At least first three terga wholly dark blue or blue-green

17

15a. Hairs of thoracic dorsum not dark-tipped; if slightly
tipped, terga 2-4 with conspicuous apical hair bands;

erect plumose hairs of tergum 4 pale 16

b. Hairs of thoracic dorsum conspicuously dark-tipped
and terga 2-4 without apical hair bands; erect plumose

hairs of tergum 4 dark obscurior Michener

16a. Terga 2-4 with conspicuous apical hair bands; basal
margin of labrum convex across entire width, convex
ridge with fine, widely spaced longitudinal ridges; ex-
ternal stripe of protibia not reaching apical three-fourths

of segment meaculpa, new name

b. Terga 2-4 without apical hair bands; basal margin of

Contributions in Science, Number 347

Snelling: American Centridini 15

labrum smooth and flat between punctures; external
stripe of protibia reaching three-fourths, or more, of

tibial length inermis Friese (part)

17a. Apical margin of tergum 4 and all of tergum 5 ferru-
ginous; scopa entirely pale or entirely dark; terga 2-3

with or without apical hair bands 18

b. Terga 4 and 5 blue-green; scopa pale, with brownish
setae posteriorly and apically on basitarsus; terga 2-3

without apical hair bands decolorata Lepeletier

18a. Scopa black; terga 2-3 without apical hair bands; ex-
ternal stripe of protibia extending half, or more, length

of segment inermis Friese (part)

b. Scopa pale; terga 2-3 with apical hair bands, that of 2
interrupted in middle; external stripe of protibia less

than half length of segment, often absent

adani Cockerell

Centris ( Centris ) aethyctera Snelling

Centris ( Centris ) aethyctera Snelling, 1974:23-26. $ 2.

Centris aethyctera is a common species ranging from Mexico
to Panama. It can only be confused with C. eisenii in this
region, a larger species with the hairs of the thoracic dorsum
without black tips. In previous literature it has been confused
with the Antillean species, C.fasciata F. Smith (see Snelling,
1974), and all records of C. fasciata (or its synonym C.fas-
ciatella Friese) in Central America seem to be based on C.
aethyctera.

In Panamanian populations of C. aethyctera the bands of
the abdominal terga are conspicuously shorter. The abdomen
thus appears to be principally ferruginous, with short, trans-
verse yellow fasciae across the disc of each segment.

Centris ( Centris ) decolorata Lepeletier

Centris decolorata Lepeletier, 1 84 1 : 1 60. <3.

Centris ( Cyanocentris ) decolorata: Friese, 1900b:243, 325. 3
(not 2?).

Centris versicolor: Cheesman, 1929:142. Misidentification.
Centris (Centris) obscuriventris: Michener, 1954:138. Mis-
identification.

Centris ( Centris ) decolorata: Snelling, 1966:23 (distr. ).

Friese (1900b) associated females with C. decolorata. but
noted that the association might be incorrect. In the event
that this should prove to be true, he proposed that the females
be called C. obscuriventris. Whether or not his female spec-
imens actually are those of C. decolorata can only be deter-
mined after his specimens are examined. In the meantime,
C. obscuriventris must be considered a name of dubious va-
lidity and identity.

There is no doubt, however, that Central American spec-
imens, previously identified as C. obscuriventris. are conspe-
cific with C. decolorata. a common species throughout the
Caribbean islands, coastal northern South America, and east-
ern Central America. The range extends north to southern

Texas, but C. decolorata appears to be absent from Florida,
even though present on Cuba. The record from Cristobal,
Canal Zone, Panama, of C. versicolor by Cheesman (1929)
is based on misidentified specimens of C. decolorata.

Centris ( Centris ) errans W. Fox

Centris errans W. Fox, 1899:65. 2.

Centris versicolor: Lutz and Cockerell, 1920:561 (in part);

Mitchell, 1962:335-336 (misidentification).

Centris ( Centris ) versicolor: Hurd, 1979:2175 (in part, mis-
identification).

This species occurs in Florida and has often been cited as C.
versicolor (Fabricius, 1775). However, Moure (1960b) ex-
amined the type of C. versicolor and redescribed the type
specimen; the true C. versicolor is “probably confined to
lesser Antilles,” according to Moure. Until all the represen-
tatives of this very difficult complex can be reexamined, it
seems best to follow Moure’s restricted interpretation of C.
versicolor and to regard C. errans as a distinct species.

Centris ( Centris ) inermis Friese

Centris inermis Friese, 1899:46. 3 2.

Centris poecila var. segregata Crawford, 1906:159. 2. NEW
SYNONYMY.

Centris inermis var. gualanensis Cockerell, 1912:568. 3. 2.
Centris (Rhodocentris) robusta Cockerell, 1949:478-479. 3.
NEW SYNONYMY.

Centris inermis subsp. pallidifrons Cockerell, 1949:479. 3.
Centris ( Centris ) inermis: Snelling, 1974:30 (syn., var.).
Centris ( Centris ) segregata: Snelling, 1974:34-35 (tax., distr.).

This is a common Central American species, ranging from
Mexico to Panama and into northern South America. It is
also a very unusual species, one which suggests that the tax-
onomy of the nominate subgenus may be fraught with more
complexities than was previously supposed. Males of C. iner-
mis are morphologically bimodal; there is a metandric form,
very different in appearance from the normal males. Addi-
tionally, however, there are two very different color pheno-
types in both sexes.

The nominate phenotype, in both sexes, has the legs and
abdomen ferruginous; in females there is usually a conspic-
uous patch of metallic blue across the base of the fourth
tergite. The second phenotype was described as C. poecila
var. segregata and later considered to be a separate species.
In this form the legs are blackish brown and the first four
abdominal segments are dark metallic blue.

The recognition of C. segregata as a synonym of C. inermis
is possible thanks to the observations of R. Coville and G.
Frankie. They have found that emergent bees from nests
provisioned by C. inermis females were, sometimes, the blue-
abdomen form, C. segregata. This fact, coupled with the lack
of morphological differences between the two forms, is the
basis for the current synonymy. Additional support lies in

16 Contributions in Science, Number 347

Snelling: American Centridini

the complete allopatry of the two forms. Frankie and Coville
will publish their observations separately.

Centris {Cent r is) meaculpa, new name

Figure 33

Centris ( Cyanocentris ) nitens: Friese, 1900b:330. 2 (in part;
misidentification).

Centris (Centris) emhescens Snelling, 1974:27-28. 2. Preoc-
cupied.

My C. erubescens is a junior homonym of C. costaricensis
var. erubescens Friese, 1925, a synonym of C. vittata Le-
peletier. The new specific epithet is, of course, the Latin
phrase for “my fault” and seems apropos in this case.

In some of the specimens recorded below, the abdominal
tergites, especially the second to fourth segments, are exten-
sively bluish, rather than wholly ferruginous. They thus
somewhat resemble females of C. adani. However, in C.
adani the erect hairs of the the fourth tergite are longer and
are plumose at their tips, as noted in the key. In most spec-
imens of C. meaculpa the clypeus is more or less distinctly
obliquely rugulose, the rugulae directed toward the apical
middle portion of the segment. The clypeus of C. adani tends
to be smooth between well spaced punctures.

The female which Friese (1900b) recorded from Mexico
as C. nitens Lepeletier, a Brazilian species, is misidentified.
The specimen is in the Paris Museum and has been made
available to me; it is a female of C. meaculpa with the first
three abdominal terga primarily metallic bluish and the spec-
imen agrees well with the characteristics of this species, es-
pecially in the structure of the labrum and of the basitibial
plate.

When I originally described this species, I was not fully
aware of its apparent affinities with a small group of primarily
Brazilian species. Included in this group are C. aenea Le-
peletier, 1841, C. caixensis Ducke, 1907, and C. nitens Le-
peletier, 1841. Females of this group are similiar in that the
pubescence of the thorax is ochreous (paler on the sides), the
abdominal terga, except the fifth and sixth, are metallic green-
ish to bluish, the discs of the second and third segments
possess abundant appressed simple hairs, and the margins
of the second and third segments have a definite fascia of
appressed pale hairs which may be plumose. In these Bra-
zilian species the basal margin of the labrum is flat and is
smooth and shiny between sparse punctures. The labrum of
C. meaculpa has a definite convexity across the basal margin
and the convex area is crossed by widely spaced longitudinal
ridges.

Of the Brazilian species, C. caixensis appears to be the
most distinct. I have seen a single female, marked as “Type,”
from the collections of the Paris Museum: Maranhao, Caixas,
30 June 1907, collected by A. Ducke. The second to fifth
terga each bear a small, lateral, pale yellow mark; the basi-
tibial plate (Fig. 32) is short, and the secondary plate is short,
broad, and convexly transverse. In all of the other species,
including C. meaculpa (Fig. 33), the secondary plate is elon-
gate, with its anterior margin approximately parallel with.

and well removed from, the anterior margin of the primary
plate. The fascial hairs of the second and third terga of C.
caixensis are yellow and simple.

Of the two remaining species, C. aenea may be recognized
by the golden or somewhat coppery color of the discal and
fascial hairs of the second and third terga. In C. nitens the
discal hairs are fuscous, sharply contrasting with the golden
to coppery color of the fascial hairs. The secondary pygidial
plate is more abruptly narrowed in C. nitens , which is the
smaller of the two species.

This species was described from two females 67 km E
Escarcega, Campeche, Mexico. A few additional specimens
are now available.

NEW RECORDS

MEXICO, TAMA ULIPAS: 12, 8 mi. NW Nuevo Morelos,
22 July 1962 (Univ. Kans. Mex. Exped.; UKAN), on Kall-
stroemia sp. SAN LUIS POTOSP. 12, El Salto, 1800 ft. elev.,
8 June 1961 (Univ. Kans. Mex. Exped.; UKAN); 12, El Salto
(above falls), 21 July 1962 (Ordway and Roberts; UKAN),
on Leguminoseae. VENA CRUZ: 12, 22 mi. SE Jalapa, 1 100
ft. elev., 29 June 1953 (Univ. Kans. Mex. Exped.; UKAN);
222, 3 mi. SW Paso del Toro, 50 ft. elev., 23 June 1961
(Univ. Kans. Mex. Exped.; UKAN). YUCATAN: 12, Piste,
12 June 1967 (E.C. Welling; LACM). STATE UNKNOWN:
12, “Mexique, Cote Occid.,” no date except 1864 (L. Biart;
MNHN).

Centris ( Centris ) obscurior Michener
Centris (Centris) obscurior Michener, 1954:138-140. 2 6.
This species has previously been reported only from Panama.

NEW RECORDS

COSTA RICA, HEREDIA: 12, Puerto Viejo, Sarapiqui, 30
July 1965 (D.H. Janzen; UKAN); 822, 37<$<5, Finca La Selva,
near Puerto Viejo, 6 May to 27 July (D.R. Perry; LACM),
on Dipteryx panamensis, Dussia sp., Hymenolobium sp.,
Byrsonima sp., and Vochysia sp. PUNTARENAS: 12, 6 km
S San Vito, 19-21 Mar. 1967 (UKAN). SAN JOSE: 222, San
Jose, 1160 m elev., 17 July 1964 (M. Naumann; UKAN),
on Solatium sp.; 222, San Jose, 1160 m elev., 8 June 1963
(C.D. and D.R. Michener; UKAN), on Solatium wendlandy
222, Ciudad Universitaria, San Jose, 28 July 1965 (S.J. Ar-
nold; UCB), on Duranta repens\ 1622, 1 mi. ESE San Isidro
de General, 21 July 1965 (S.J. Arnold; UCB), on Rhyn-
ch a nth era mexicana, 0815-0950. MEXICO, CHIAPAS: 12,
Simojovel, 1-16 Aug. 1958 (J.A. Chemsak; UCB); 12, Santo
Domingo, 15 mi. S Simojovel, 8 July 1958 (J.A. Chemsak;
UCB); 3<5<5, 3 mi. SE San Juan del Bosque, 16 Aug. 1958
(J.A. Chemsak; UCB); 12, Yaxoquintela, 560 m elev., 30
Aug. 1978 (J.E. Rawlons; CORN).

Centris (Centris) flavifrons Group

To this group are assigned a number of medium-sized to
large species in which the integument of the head, thorax.

Contributions in Science, Number 347

Snelling: American Centridini 17

and legs is blackish with limited whitish or yellowish marks
on the face and legs. The abdomen is mainly metallic blue
to blue green, in the males with yellowish maculations, at
least in the second tergite; the apical segments may be red-
dish. In both sexes the thorax is either largely cinereous pu-
bescent and with a broad interalar band of blackish hairs, or
principally dark pubescent but with the scutellum and meta-
notum pale pubescent. The wings are light brownish to black-
ish.

Morphologically, the species tend to be very similar and
species limits are, at present, very poorly understood. The
following key will serve to separate those species which I am
presently able to recognize.

KEY TO SPECIES, C. FLA VIFRONS GROUP

la. Male, antenna 13-segmented and basitibial plate absent

2

b. Female, antenna 1 2-segmented and basitibial plate pres-
ent 6

2a. Ocellocular distance no more than 0.90 times diameter
of anterior ocellus and usually less; lower inner man-
dibular carina ending in a small, obscure tooth-like pro-
cess (Fig. 13); smaller species, head width less than 5.8

mm 3

b. Ocellocular distance at least 0.95 times diameter of an-
terior ocellus and often greater; lower inner mandibular
carina often terminating in a prominent tooth-like pro-
cess (Fig. 14); usually larger species, head width usually
more than 6.0 mm, but may be a little as 4.8 mm . . 4
3a. Clypeus moderately shiny, punctures fine and close, in-
terspaces very finely roughened; apical margin of third
tergite with transverse band of pale yellowish hairs; pu-
bescence of side of propodeum pale (Ecuador, Peru)

buchwaldi Friese

b. Clypeus shiny and subpolished between fine, close punc-
tures, except in broad, impunctate median line; third
tergum blackish pubescent along apical margin; side of
propodeum brown pubescent (El Salvador, Costa Rica,

Panama) aethiocesta, new species

4a. Larger species, head width always over 5.2 mm and
usually greater than 6.0 mm; clypeal disc, at least in large
part, shiny and weakly, or not at all, tessellate; second
tergum, at least, with transverse yellow fascia across base,
sometimes narrowly interrupted in middle (widespread)

5

b. Smaller species, head width less than 5.0 mm; clypeal
disc moderately shiny, conspicuously tessellate and
roughened, more weakly so along midline; second ter-
gum with a pair of widely separated spots (Eucador)

nigrofasciata Friese

5a. Lower inner mandibular carina ending in a long, acute
tooth-like process. Normal male with broad, basal, yel-
low fasciae across base of second to fourth terga which
are complete or narrowly interrupted, that of second
nearly as long as apical area beyond it; punctures of
clypeal disc numerous and close, interspaces more or
less distinctly tessellate, especially basad. Metander with

clypeus slightly roughened, punctures close to sparse;
clypeal disc entirely yellow (S Arizona to Panama) . . .

flavofasciata Friese

b. Lower inner mandibular carina ending in low, obtusely
triangular process (Fig. 14). Normal male with narrow
basal fasciae on second to fourth terga, that of second
often broadly interrupted in middle, or, when complete,
distinctly shorter in middle than apical area beyond it;
clypeal disc smooth and polished, punctures at side sparse.
Metander with clypeus smooth and polished, punctures
sparse to scattered; laterobasal black mark of clypeus
extending onto disc (widespread) . .flavifrons (Fabricius)
6a. Apical margin of third tergite with a narrow band of
white or yellowish red hairs which sharply contrast with
the black discal hairs, the pale hairs closer than those of

disc and projecting well beyond margin 7

b. Apical margin of third tergite, across middle one-half
with hairs short, black, no closer than short, black hairs
of disc and hardly projecting beyond margin of segment

' 8

7a. Scopa pale; marginal hairs of third tergite whitish, straight;
scape pale beneath; fifth tergite wholly ferruginous (Ec-
uador) nigrofasciata Friese

b. Scopa dark; marginal hairs of third tergite yellowish red,
their tips curved laterad; scape dark; fifth tergite mostly
metallic blue-green (Ecuador, Peru) . . buchwaldi Friese
8a. Head width usually exceeding 6.7 mm, rarely as little as
6.2 mm; scopa usually mostly dark; clypeal punctures
moderate in size and on side of disc separated by a
puncture diameter or more; hairs of side of propodeum

pale, but if dark, pleural hairs also dark 9

b. Head width less than 6.3 mm; scopa pale; clypeal punc-
tures fine and close; pleura with pale hairs and side of
propodeum with brown hairs (El Salvador, Costa Rica,

Panama) aethiocesta, new species

9a. Larger species, head width 7.0-7. 7 mm; scopa with pale
hairs, at least anterobasally on metatibia, often largely
pale; pygidial and basitibial plates as in Figs. 38 and 29

(widespread) flavifrons (Fabricius)

b. Smaller species, head width 6. 2-7. 2 mm; scopa black, a
few pale-tipped hairs posteroapically on metabasitarsus;
pygidial and basitibial plates as in Figs. 39 and 30 (S
Arizona to Panama) flavofasciata Friese

Centris ( Centris ) aethiocesta, new species

Figures 5-8, 13, 15, 31, 40

Centris ( Centris ) nigrofasciata: Michener, 1954:138. Mis-
identification.

DIAGNOSIS

Medium-sized species with thoracic pubescence whitish, but
with black interalar band and brown pubescence on propo-
deal side; abdomen metallic blue-green. Female with pale
scopa, broad basitibial plate, and clypeal disc flattened in
profile. Male with clypeal disc flattened in profile; abdominal
terga 4-7 metallic green, immaculate; punctures of clypeal

18 Contributions in Science, Number 347

Snelling: American Centridini

disc numerous and mostly separated by about a puncture
diameter.

DESCRIPTION

HOLOTYPE FEMALE. Measurements (mm). Head width
5.54 (5.95-6.26); head length 4.26 (3.95-4.21); wing length
1 1.0 (1 1.0-12.0); total length 15.5 (16.6-18.0).

Head. 1 .43 ( 1 .49-1 .53) times broader than long; occipital
margin nearly flat in frontal view and slightly below level of
tops of eyes; ocelli anterior to occipital margin; inner orbits
moderately convergent above, upper frontal width 0.90 (0.89-
0.92) times lower frontal width. Mandible slender, tridentate,
apical margin oblique. Labrum about twice broader than
long, apical margin narrowly rounded. Disc of clypeus weakly
depressed, nearly flat in profile (Fig. 15); broad median line
impunctate, punctures fine on either side and mostly sepa-
rated by about 1.5 times a puncture diameter, interspaces
shiny, a little less so basad. Frons slightly to moderately shiny
between fine, subcontiguous punctures; ocellocular area
moderately shiny, impunctate adjacent to ocellus, densely to
subcontiguously punctate near eye, with fine and minute
punctures; preoccipital area slightly shiny between dense to
subcontiguous fine punctures; gena shiny between irregularly
close to subcontiguous punctures which are minute near eye,
becoming fine to moderate ventrad. Interantennal distance
3.41 (3.13-3.48) times antennal socket diameter; antennoc-
ular distance 1.63 (1.57-1.63) times antennal socket diam-
eter; scape slender, 2.29 (2.23-2.36) times longer than wide,
scape length 0.72 (0.70-0.75) times length of first flagellar
segment; first flagellar segment 5.28 (5.28-5.50) times length
of second and distinctly longer than combined length of fol-
lowing three segments. Interocellar distance 2.09 (2.00-2. 1 2)
times diameter of anterior ocellus; ocellocular distance 1.31
( 1 .32-1 .45) times diameter of anterior ocellus; ocelloccipital
distance 1 .88 ( 1 .88-2.03) times diameter of anterior ocellus.

Thorax. Mesoscutum shiny between fine, dense punctures;
scutellum similar, but punctures dense to subcontiguous and
narrow median line impunctate; metanotum moderately shiny
between scattered minute punctures; meso- and metepisterna
shiny between dense, fine punctures. Basal area of propo-
deum moderately shiny, sharply tessellate and with sparse
fine punctures except along anterior margin; propodeal side
shiny between dense, fine punctures. Basitibial plate (Fig. 3 1 )
about twice longer than wide, lower margin broadly rounded;
lower margin of secondary plate rounded.

Abdomen. First three terga shiny between dense, minute
punctures; fourth and fifth terga shiny between dense, fine
punctures which are a little more separated than on basal
segments; pygidial plate (Fig. 40) narrow, apex narrowly trun-
cate; secondary plate with margins slightly concave toward
gradually narrowed apical ridge.

Color. Head, thorax, and legs blackish brown; scutellum
dull reddish; first to fifth abdominal terga metallic blue-green,
fourth and fifth more greenish; abdominal sterna brownish,
with obscure metallic greenish tints, especially laterad. The
following whitish: basal spot on mandible; labrum, except
apex and along basal margin (sometimes reduced to lateral

spots); inverted T-shaped clypeal mark (may be reduced to
narrow median stripe and sublateral, preapical spots); narrow
supraclypeal mark (present only in holotype); elongate spot
on malar area; broad mark on paraocular area, ending nar-
rowly on eye margin at, or slightly above, level of lower
margin of antennal socket; basal spot on pro- and mesotibia
(preapical protibial spot sometimes present). Tegula reddish.
Wings dark brown; veins and stigma blackish.

Pilosity. Hairs whitish on head, but with broad preoccipital
band of long, dark brown hairs; ocellar area pale-haired, but
with brownish hairs on frons anterior to ocelli. Hairs of tho-
rax whitish, but with broad interalar band of dark brown
hairs and brown hairs on metepisternum and side of pro-
podeum. Hairs on front face of first tergum pale; hairs on
dorsal face of first, and on second and third terga appressed,
simple, blackish, very short; hairs of fourth and fifth terga
long, erect, mostly plumose, whitish; prepygidial fimbria
golden reddish to brown. Hairs of legs mostly brownish black,
but with glistening whitish hairs on coxae, trochanters, and
posterior margins of profemora and protibia; scopal hairs
whitish, becoming brownish distad on metabasitarsus.

MALE. Measurements (mm). Head width 5.79(5.33-5.85);
head length 3.79 (3.44-3.79); wing length 12.0; total length
14.0

Head. 1.53 (1.51-1.56) times broader than long; occipital
margin nearly flat in frontal view and slightly below tops of
eyes, ocelli anterior to occipital margin; inner orbits very
strongly convergent above, upper frontal width 0.80 (0.72-
0.82) times lower frontal width. Mandible slender, tridentate,
inner tooth broad and with its apical margin concave. La-
brum about 1.6 times broader than long, moderately shiny
between sparse fine punctures and with interspersed minute
punctures. Disc of clypeus nearly flat in profile, about as in
female, moderately shiny, with broad median impunctate
line, punctures on either side fine, mostly separated by about
a puncture diameter. Punctation of remainder of head about
as in female. Interantennal distance 2.72 (2.47-2.81) times
diameter of antennal socket; antennocular distance 0.79 (0.77-
0.93) times diameter of antennal socket; scape stout, 2.18
(2.07-2.18) times longer than wide, 0.71 (0.70-0.79) times
length of first flagellar segment; first flagellar segment 5.60
(5.33-5.60) times length of second and longer than following
three segments combined. Interocellar distance 1.82 (1.84-
1 .94) times diameter of anterior ocellus; ocellocular distance
0.88 (0.84-0.90) times diameter of anterior ocellus; ocelloc-
cipital distance 1.76 (2.00-2.06) times diameter of anterior
ocellus.

Thorax. As described for female, but mesoscutal punctures
subcontiguous and scutellum slightly depressed along mid-
line. Metafemur about twice longer than thick (dorsoven-
trally); metabasitarsus about three times longer than broad,
posterior margin weakly curved.

Abdomen. As described for female; apex of last tergite
broadly bilobate.

Terminalia. Sides of distal process of seventh sternite (Fig.
5) slightly divergent distad, apical margin weakly incised;
setae short, sparse, simple. Distal process of eighth sternite
(Fig. 6) weakly narrowed subbasally; setae long, numerous.

Contributions in Science, Number 347

Sneiling: American Centridini 19

Gonostylus (Figs. 7, 8) a little broadened distad; ventral pro-
cess broadly rounded.

Pilosity. As described for female, but hairs of apical ab-
dominal sternite pale brown; hairs of metatibia and meta-
basitarsus mostly dark brownish, but with some along pos-
teroapical margins of metatibia golden brown and some along
posterior margin of metabasitarsus light brown, with pale
tips.

Color. Body color as described for female, except whitish
marks as follows: spot at base of mandible; entire labrum;
clypeus, except along laterobasal margin; broad supraclypeal
mark; spot on malar area; paraocular area, ending narrowly
on inner eye margin slightly above level of lower margin of
antennal socket; underside of scape; basal spot on all tibiae;
outer stripe on protibia; sublateral mark at base of second
tergite. Tarsi reddish brown to brown. Wings light brown.

TYPE MATERIAL

Holotype female and allotype: airstrip, Isla El Rey, Islas Per-
las, PANAMA, 22 Feb. 1981 (D.W. Roubik), flying around
Dioclea megacarpa, in Natural History Museum of Los An-
geles County. Paratypes: 12, same data as holotype; 5<3<3, Coi-
ba Island, Veraguas, PANAMA, 21-24 Oct. 1979 (D. Rou-
bik); 12, Fort Kobbe, Canal Zone, PANAMA, 9 Sept. 1958
(W.J. Hanson); 1 <3, Patilla Point, Canal Zone, PANAMA, 15
Jan. 1929 (C.H. Curran); 1<3, Bruja Point, Canal Zone, PAN-
AMA, 25 Jan. 1929 (C.H. Curran); 12, LaChorrera, Panama,
PANAMA, 22 May 1912 (A. Busck); 1 <5, 5 km E Comarca
de San Bias, Puerto Obaldia, PANAMA, 1 1 Oct. 1979 (D.
Roubik); 322, Tamarindo Beach, Guanacaste, COSTA RICA,
28 Feb. 1980 (G.W. Frankie); 222, same locality and collec-
tor, 30 Mar. 1 980, on Haematoxylon brasiletto\ 266, La Union,
Playa El Icacal, EL SALVADOR, 8 July 1975 (E.M. and J.L.
Fisher). Paratypes in AMNH, LACM, ROUB, UCB, UKAN,
and USNM.

ETYMOLOGY

Combines the Greek words aithiops (swart or dark) and kes-
tos (girdle or band).

DISCUSSION

Although this species superficially looks much like C. flavi-
frons and C. flavofasciata, it is smaller than either. The disc
of the clypeus, in profile, is distinctly flatter in C. aethiocesta
than in the other two species (compare Fig. 1 5 with Fig. 16)
and, in both sexes, the disc is more closely and abundantly
punctate. Small females of C. flavofasciata may be no larger
than females of C. aethiocesta but have the tibial scopa black
and the hairs of the side of the propodeum are whitish. The
much larger size (head width greater than 7.0 mm) will sep-
arate C. flavifrons females from C. aethiocesta, as will the
sparser clypeal punctation and the differently shaped basi-
tibial plate. Males of C. aethiocesta have the apical abdom-
inal segments dark and immaculate; in C. flavifrons and C.
flavofasciata males the apical segments are usually reddish
and commonly are yellowish maculate; in these two species,
the hairs of the propodeum are usually whitish.

Two South American species also resemble C. aethiocesta
and are of about the same size. Females of C. buchwaldi
Friese, 1 900, have a dark scopa, pale propodeal hairs, a band
of prostrate, simple, golden brown hairs across the apical
margin of the third tergite, and the erect hairs of the fourth
tergite are simple. In females of C. nigrofasciata Friese, 1 899,
there is a band of pale hairs across the apical margin of the
third tergite, the apical margin of the fourth tergite is fer-
ruginous, the fifth tergite is ferruginous and with simple erect
hairs, and the basitibial plate is more elongate, with a more
acute apex.

In the males of both C. buchwaldi and C. nigrofasciata the
clypeus is only slightly shiny, with the integument conspic-
uously roughened. Neither of these has brown hairs on the
side of the propodeum and in both species the fifth and sixth
tergites are ferruginous. Males of C. nigrofasciata also differ
in that the ocellocular distance is greater than the diameter
of the anterior ocellus.

Centris ( Centris ) flavifrons (Fabricius)

Figures 14, 29, 38

Apis flavifrons Fabricius, 1775:383. <5.

Apis flavifrons brasiliana Christ, 1791:1 40.

Centris flavifrons: Lepeletier, 1841:152. F. Smith, 1874:361.
Centris citrotaeniata Gribodo, 1894:267. 2. NEW SYN-
ONYMY.

Centris flavifrons var. nigritula Friese, 1899:46. 2. NEW
SYNONYMY.

Centris flavifrons var. rufescens Friese, 1899:46. <3. NEW
SYNONYMY.

Centris (Cyanocentris) flavifrons: Friese, 1 900b:3 1 7—3 1 8 (in
part) (tax., distr.).

Centris ( Centris ) flavifrons: Michener, 1954:137 (distr.).

Moure, 1 960b: 1 25-126 (tax.).

Centris (Centris) rufescens: Michener, 1 954: 1 37 (tax., status).
“Centris fulvifrons" Moure, 1960a: 10 (lapsus for C. flavi-
frons).

Centris ( Centris ) citrotaeniata: Moure, 1960a: 10-1 1 (tax.).
Centris (Centris) nigritula: Snelling, 1974:30-31 (tax. status).

Moure (1960b) examined and redescribed the type of C.
flavifrons, from “Brasilia.” He noted that, according to cur-
rent concepts, C. flavifrons ranges from southern Brazil to
Mexico. Throughout this range this is often a common bee
and one subject to considerable variation. It is now apparent
that my effort to separate C. f. nigritula as a distinct species
(Snelling, 1974) was futile. Since that time I have examined
several hundred specimens and seen so broad a range of
variation that it is now apparent that C. nigritula is not
worthy of recognition at any level.

Friese (1899) described C. flavifrons var. rufescens from a
male from Chiriqui, Panama. It was said to differ from the
nominate form in possessing wide, yellow abdominal bands
and reddish pubescence on the legs. Although Michener (1954)
thought that this seemed to represent a distinct species, I do
not agree. Neither Michener nor I has seen the type of var.
rufescens but I suspect that it is nothing more than a metan-
der. Metanders of both C. flavifrons and C. flavofasciata are

20 Contributions in Science, Number 347

Snelling: American Centridini

characterized by possessing broad abdominal fasciae and fer-
ruginous pubescence on the legs. Since nothing in Friese’s
brief description would indicate to which of these species this
form should be assigned, I have arbitrarily elected to follow
Friese and include it under C. flavifrons. as a synonym. In
Panama, C. flavifrons is more common than is C. flavofas-
ciata and it seems more likely than the var. rufescens belongs
with the former species.

Centris citrotaeniata was also described from Chiriqui,
Panama, based on a single female specimen. The type is in
the collections of the Museu Civico de Storia Naturale, Gen-
oa, Italy, and was examined by Moure ( 1 960a). Moure noted
that this species was close to “C. fulvifrons evidently a
lapsus for C. flavifrons. From the original description, as well
as the supplemental notes provided by Moure, it is clear that
C. citrotaeniata is equivalent to dark Panamanian individ-
uals of C. flavifrons. and so Gribodo’s species is here placed
in synonymy.

Variation in females of C. flavifrons chiefly involves the
extent of pale versus blackish pubescence on the thorax and
on the scopa. In females from Brazil the thoracic hairs are
mostly whitish, with a conspicuous broad interalar band of
blackish pubescence; a few dark brown hairs are present be-
low the wing bases and the scopa is uniformly pale. In ma-
teria! from Guyana and French Guiana, there is considerable
replacement of the pale pleural hairs by blackish hairs and
the scopa is largely dark, but with conspicuous white hairs
anteriorly on both the tibia and basitarsus. This dark variant
also has many dark hairs on the fourth and fifth terga. A
similar variant also appears sporadically in samples from
Panama and Costa Rica. The darkest phenotype is from
Trinidad, the var. nigritula. In these specimens, the thorax
is entirely blackish pubescent, with only the axilla, scutellum,
and thoracic venter whitish pubescent. Curiously, the scopa
is predominantly pale, with only a few brownish hairs distally
on the tibia and along the posterior margin of the basitarsus.
Some of the Panamanian females have the thoracic hairs
wholly pale, except for a small median patch of dark hairs
on the mesoscutum; in these individuals the scopa is mainly
dark.

The Central American populations vary on a smaller scale.
Specimens from Mexico typically have a large inverted
T-shaped clypeal mark, the labrum is mostly dark and the
lateral face mark fills the paraocular area below the level of
the tentorial pit. The transverse mesoscutal fascia of dark
hair is preceded by a narrow band of white pubescence, the
mesopleuron is largely dark pubescent, and the tibial scopa
is only about one-half pale. Coloration becomes progressively
darker toward the south and specimens from Costa Rica may
be much darker: the labrum wholly black, the clypeal mark
broken into two or three narrow segments, and the meso-
scutum with only a small cluster of pale hairs on the antero-
median portion; the scopa is almost entirely dark, with pale
hairs anteriorly on the metatibia and anterobasally on the
metabasitarsus. Another characteristic of the Costa Rican
specimens is that the hairs of the dorsal face of the first tergite
are blackish. In the Mexican specimens the hairs are pale at
the sides and across the basal portion of the dorsal face.

The tendency toward increasing melanism is partially re-
versed in Panama. A long series from Panama Province,
collected by R.W. Brooks, exhibits considerable variation.
In some specimens the interalar fascia is nearly absent and
in others it may cover up to two-thirds of the mesoscutum.
In most specimens the mesepisternum has only a few dark
hairs below the tegula and in others there may be a blackish
hair patch over the upper one-half or more of the segment.
The scopa varies from nearly wholly pale to about one-half
black. Face marks may be greatly reduced: mandible and
labrum entirely dark; clypeus with narrow median line and
widely separated apicolateral spots and scape wholly dark.
Reduction of face marks is not correlated with increased
melanism in pubescence. Many of these specimens will match
the color pattern of the type of C. citrotaeniata.

The darkest Costa Rican specimens superficially resemble
the form from Guyana, French Guiana, and Trinidad de-
scribed by Friese as the var. nigritula. They differ from that
form, however, in that the plumose hairs of the fourth tergite
are uniformly pale. In the var. nigritula there are many dark
hairs on the fourth tergite and, in some specimens, pale hairs
may be entirely lacking.

In general, then, this species becomes progressively more
melanic, proceeding from central Brazil to northeastern South
America. Westward across northern South America there is
a reversal, toward decreased melanism, in Venezuela, Co-
lombia, and Panama. The melanic trend is again evident in
Costa Rica and Nicaragua, but decreases once more at the
northern (Mexican) extremities of the range. There are nu-
merous localized phenotypes which may depart, in one way
or another, from the general dine.

The discussion thus far has focused primarily on the fe-
males, though the pubescent patterns of the males of C. flavi-
frons generally follow those of the females. There are, how-
ever, problems that are peculiar to the males. Both this species
and the similar C. flavofasciata possess metanders, males
which are unusually large and robust and much more exten-
sively marked with bright yellow, especially on the legs and
abdomen. While it is possible to separate normal males of
the two species, the metanders are much less convincingly
handled.

In part, this is due to a scarcity of fully developed metan-
ders which can be assigned with surety to one or the other
of these two species. And, in part, the metanders that are
available tend not to exhibit the differences which separate
the normal males. I do not believe that this in any way
invalidates the distinctions between the two species, but it
is merely another manifestation of a common problem within
some subgenera: within a given complex males may be much
less readily separated than their respective females. A final
difficulty is that, unlike the situation in such subgenera as
Xerocentris and Paracentris, the metandrous and normal
males form a more or less continuous series, with many
individuals of intermediate stature.

A series of about 30 males collected by J.A. Chemsak near
Mazatlan, Sinaloa, Mexico, 12 Dec. 1980 (UCB) consists
largely of metanders of C. flavofasciata. but with a few nor-
mal males and intermediates. While I have seen many males

Contributions in Science, Number 347

Snelling: American Centridini 21

of C. flavifrons that seem to be intermediates, I have only
three that are clearly fully developed metanders, two from
Brazil and one from Mexico.

Normal males of C. flavifrons possess the following char-
acteristics, each subject to moderate variation: the lower in-
ner mandibular carina terminates in a low, obtusely trian-
gular process (Fig. 14); the clypeal disc is smooth and polished
throughout between fine, scattered punctures and is not whol-
ly yellow; the laterobasal black mark of the clypeus extends
onto the disc and sometimes the clypeus bears only an in-
verted (though broad) T-mark. The second to fourth abdom-
inal terga are maculate; the maculation of the second segment
often consists merely of two somewhat elongate sublateral
spots, but may extend across the base of the segment as a
continuous or narrowly interrupted band; when it is a band,
it is almost always much shorter in the middle than the dark
area distad to it.

In normal males of C. flavofasciata the lower inner man-
dibular carina ends in an elongate, acute, tooth-like process.
The disc of the clypeus is wholly yellow and the areas on
either side of the shiny, impunctate median line are less shiny
and distinctly roughened between fine punctures that are
mostly separated by 1.0 to 1.5 times a puncture diameter.
The abdominal terga are almost always continuously fasciate
and the yellow band of the second segment is as long as, or
longer than, the dark area that follows it.

Most of these distinctions disappear in the metanders, or
are at least less certain. Some metanders of C. flavofasciata
have the clypeal disc nearly polished and the punctures about
as sparsely distributed as in C. flavifrons. While very nearly
all metanders of C. flavofasciata have the clypeal disc wholly
yellow, there are a few in which the laterobasal black mark
does extend slightly onto the disc. The abdominal banding
breaks down completely, since metanders of both species
have broad, continuous yellow fasciae on the second to sixth
segments and an interrupted band across the dorsum of the
first segment.

The only feature that seems to hold up is the shape of the
termination of the lower inner mandibular carina. In me-
tanders of C . flavifrons the terminus is low and obtuse, much
as in the normal males. It is an acutely tooth-like process in
C. flavofasciata. How consistent this difference is, I cannot
now state, only that it does hold true for the specimens ex-
amined thus far.

There is obviously much to be done before the phenom-
enon of metandry can be adequately dealt with taxonomi-
cally. Presumably, the metanders of such species as C. flavi-
frons and C. flavofasciata are behaviorally different from
their normal counterparts, perhaps in a manner similar to
the situation in such species as C. (Xerocentris) pallida W.
Fox, the only species studied thus far (Alcock et al., 1976,
1977). In that species metanders seek newly emerging females
at nest sites of the previous year. The normal males establish
territories at nectar/pollen sources and attempt to mate with
females there.

A large nesting aggregation of C. flavifrons was observed
in Panama. Normal males and the rufescent variety were
present. Males were observed to “. . . purposely attack Me-

soplia in the nest area— with a frequency far higher than the
pursuit of other males or females” (D. Roubik and D. Yane-
ga, in prep.).

Centris (Centris) flavofasciata Friese

Figures 16, 30, 39

Centris flavifrons var. flavofasciata Friese, 1899:46. <5.
Centris ( Cyanocentris ) flavifrons var. flavofasciata: Friese,

1900b:318. $ (distr. , tax.).

Centris (Centris) flavofasciata: Michener, 1954:137 (distr.).

Snelling, 1966:23 (distr.).

Centris flavofasciata is a common Mexican species which
ranges from southern Arizona to Panama. It is possible that
the range extends into northern South America, but I have
seen no specimens to substantiate that assumption. Most of
the features by which this species may be separated from
other members of the C. flavifrons groups are summarized
in the keys.

Females of C. flavofasciata consistently have a dark scopa,
with a few of the posteroapical hairs of the metabasitarsus
with pale tips. Most females have a small patch of dark hairs
below the base of the forewing; none has the side of the thorax
extensively dark pubescent, as is usually true of Central
American material of C. flavifrons. Those specimens of C.
flavifrons with little dark pleural pubescence are from Pan-
ama.

Facial maculations of C. flavofasciata tend to be somewhat
whitish in females, rather than definitely yellow as in C.
flavifrons and, in both sexes, are more extensive; in the male
the entire clypeal disc is yellow, not encroached upon by the
black laterobasal mark. In females of C. flavofasciata the
labrum is largely pale, rather than extensively dark and the
vertical and lateral arms of the inverted T-mark of the clyp-
eus are broad and never broken into segments.

Ptilocentris, new subgenus

DIAGNOSIS

Differs from other subgenera of Centris by the following com-
bination of characters. Maxillary palp five-segmented; tergal
integument metallic blue-green, first four terga with pubes-
cence long, dense, erect, plumose. Female: secondary basi-
tibial plate with distinct overhanging margin, pro- and me-
sobasitarsi with elaiospathe; clypeal disc evenly rounded into
lateral and basal faces, gently convex from side to side. Male:
metabasitarsus without posterior carina; genitalia without
branched setae; upper inner mandibular carina ending at base
of inner tooth; eighth sternite expanded at apex.

DESCRIPTION

Mandible narrow, neither subapically broadened nor bent;
tri- (male) or quadridentate (female); upper inner carina end-
ing near base of innermost tooth; lower inner carina not
subangularly produced, continuous to innermost tooth along
upper margin. Labrum of female about 1.3 times broader
than long, apex subacute; about as broad as long in male.

22 Contributions in Science, Number 347

Snelling: American Centridini

Clypeus broader than long and separated from inner eye
margin by less than diameter of anterior ocellus; disc not at
all flattened, low-convex from side to side and evenly round-
ed onto lateral faces and gently sloping toward base. First
flagellar segment about as long as following three combined
(female) or shorter (male). Ocellocular distance greater than
anterior ocellus diameter. Pro- and mesobasitarsi with an-
terior elaiospathe on inner surface; basitibial plate of female
with distinct secondary plate, its lower margin overhanging
primary plate; metafemur of male robust, without ventral
ridge or tubercle; metabasitarsus simple, without posterior
ridge or carina. Female pygidial plate V-shapcd, apex nar-
rowly rounded; secondary plate distinct, apex acute. Male
without distinct pygidial plate; seventh sternite (Fig. 9) sub-
apically broadened and apical margin deeply emarginate; with
abundant long, plumose hairs at apex, with short, simple
hairs at base of apical lobe; eighth sternite (Fig. 10) with
distal process long, its apex angularly spatulate, apical one-
half of shaft and broadened distal portion with long, plumose
hairs; genital capsule (Figs. 11, 12) robust, gonocoxite mas-
sive; gonostylus short, broad, with a few fine, simple setae.
Abdominal terga metallic blue-green and four basal segments
with abundant, erect, plumose hairs on discs.

TYPE-SPECIES

Centris festiva F. Smith, 1854, by present designation.

ETYMOLOGY

The subgeneric name combines Greek ptilon (feather) with
the generic name Centris and refers to the presence of abun-
dant long, erect, plumose hairs on the abdominal terga.

The subgeneric name proposed here was originally con-
ceived by J.S. Moure and appears on specimens of C. festiva
in various collections which he identified 20 years ago. It is
an appropriate name and I am pleased to adopt it here.

DISCUSSION

The type-species is the only known representative of this
subgenus.

Ptilocentris seems to be another segregate of the Paracen-
tris-Centris s. str. complex of subgenera. In my key to the
subgenera of Centris (Snelling, 1974) the female will go to
Centris s. str.

Females differ from those of the nominate subgenus in
lacking a stipital comb; in Centris s. str. the comb consists
of a long row of stout, close-set, coarse, acute bristles. The
labrum in Centris s. str. is much broader, with the apical
margin broadly rounded. The broad, distinctly flattened, or
slightly depressed, clypeal disc which abruptly slopes basad
and toward the lateral margins is also distinctive of Centris
s. str. In this regard, Ptilocentris is more like some species
of Paracentris from which it immediately differs in possessing
a sharply defined secondary basitibial plate with a distinct
margin which extends slightly over the disc of the primary
plate. Also, Paracentris is like Centris s. str. in the form of
the comb of the stipes.

The male of Ptilocentris is unique among the Paracentris -
Centris s. str. complex in the form of the gemlalic structures.
The shapes of the seventh and eighth stermtes are distinctive
and the lack of coarse, plumose setae on the genitalic capsule
is also an unusual feature in this complex. So, too, is the
short, broad gonostylus.

Centris (Ptilocentris) festiva F. Smith

Figures 9-12

Centris festiva F. Smith, 1854:375. 2.

Centris chlorura Cockerell, 1919:188-189. 2. NEW SYN-
ONYMY.

This attractive and unusual species is easily recognized by
the characteristic pattern of erect hairs on the discs of the
abdominal terga. In most specimens the second and third
terga have a basal zone of blackish hairs, but in some spec-
imens the hairs may be wholly pale; one such pale haired
specimen was the basis for C. chlorura, the type of which
has been examined.

There are no previous records of this species from Central
America. In South America, C. festiva is known to be present
in Colombia, Ecuador, and Peru.

NEW RECORDS

MEXICO, DISTRITO FEDERAL: 13, Los Venados, 6 Nov.
1938 (G. Vivas-Berthier; USNM). COSTA RICA, ALAJUE-
LA: 1<3, 5 km S Vara Blanca, 1 1 Nov. 1973 (P.A. Opler;
LACM). SAN JOSE: 12, San Jose, “1.6.19” (M. Valerio;
USNM). PANAMA, CHIRIQUI: 13, Osra Clara, 26 Jan.
1981 (D.W. Inouye; ROUB). According to Roubik (pers.
comm.): “Sandy Knapp also collected C. festiva in Chinqui
on 22 April 82 ... at Cerro Colorado, about 1000 m ele-
vation.”

Subgenus Melanocentris Friese

Centris subg. Melanocentris Friese, 1900b:241, 244. Type-
species: Centris atra Friese, 1900b; designated by Sand-
house, 1943.

Species belonging to the subgenus Melanocentris are medi-
um-sized to large bees, usually black (though with pale face
marks in the males) or with the abdomen more or less dusky
ferruginous. Less commonly the abdomen may have metallic
blue or green reflections. The pubescence is mostly dark but
there may be limited amounts of pale pubescence on the
thorax, the hind legs, and on the abdomen. In both sexes the
maxillary palp is five-segmented and the mandible is stout,
quadridentate in the female and tndentate in the male; in
the female the mandible is broadened and abruptly bent near
the apex.

Additional characteristics of the female include the pres-
ence of an elaiospathe on the pro- and mesobasitarsi, the
weakly bilobate scutellum, the upper inner carina of the man-
dible ends near the base of the subbasal tooth, and the margin
of the secondary basitibial plate overhangs the primary plate.
Males lack giant branched setae on the genitalia, the meta-

Contributions in Science, Number 347

Snelling: American Centridini 23

11 12

Figures 9-12. Centris (Plilocentris) festiva, male seventh and eighth sternites and genitalia (ventral and dorsal views). Scale line = 1.00 mm.

24 Contributions in Science, Number 347

Snelling: American Centridini

Figures 13-14, apex of left mandible of male; 13, Centris (C.) aethiocesta\ 14, C. (C.) flavifrons. Figs. 15-16, clypeal profile of female: 15, C.
(C.) aethiocesta\ 16, C. (C.) Jlavofasciata. Fig. 17, apex of left mandible of male C. (Melanocentris) fusciventris. Figs. 18-19, melabasitarsus
of male: 18, C. ( Trachina ) labiata\ 19, C. (T.) longimana.

Contributions in Science, Number 347

Snelling: American Centridini 25

basitarsus is without an elevated carina on the posterior mar-
gin, the scutellum is weakly swollen on either side of the
middle, and the upper inner mandibular carina ends near the
base of the middle tooth.

This is primarily a South American group, and much in
need of revisionary study. The few North American species
may be separated by the following key.

KEY TO NORTH AMERICAN MELANOCENTRIS

1 a. Male, antenna 1 3-segmented and basiti bial plate absent

2

b. Female, antenna 12-segmented and basitibial plate

present 8

2a. Integument of abdominal terga shiny, reddish or black-
ish, never strongly metallic; if dull, punctures subcon-
tiguous to dense, sharply defined at least on second
tergunr, metafemur swollen, but without ventral ridge

3

b. Integument of abdominal terga dull to slightly shiny,
dark blue, punctures on discs of second and third ter-
gites very fine, obscured by dense tessellation and well
separated; metafemur swollen, with low ventral ridge

which terminates slightly beyond middle

plumipes F. Smith

3a. Pilosity of thorax buff-colored to ferruginous (hairs of

mesoscutum may be dark-tipped) 4

b. Pilosity of thorax dark brown to blackish (one species
with light hairs on scutellum, metanotum, and pro-

podeum) 5

4a. Inner mandibular tooth broad, cutting margin incised
so that mandible approaches quadridentate condition
(Fig. 1 7); pilosity of third and following tergites blackish

fusciventris Mocsary

b. Inner mandibular tooth narrow, triangular, mandible
definitely tridentate; pilosity of third and following ter-
gites ferruginous flavilabris Mocsary

5a. Clypeus narrow, about 1 . 10 to 1 .25 times broader than
long; lower facial width no more than 1 .5 times broader
than interocular distance, latter always greater than

transocellar distance 6

b. Clypeus broader, at least 1.40 times broader than long;
lower facial width at least 1.6 (and usually more than
1 .8) times minimum interocular distance, latter usually

no more than transocellar distance, often less 7

6a. Abdomen dusky ferruginous; posterior part of thorax,
metatibia and metabasitarsus pale pilose; disc of second
tergum dull, punctures mostly separated by less than a

puncture diameter agiloides, new species

b. Abdomen brownish to blackish; pilosity of thorax and
leggs blackish; disc of second tergum moderately shiny
between punctures mostly separated by a puncture di-
ameter or more sericea Friese

7a. Dorsal face of scutellum depressed along midline and
slightly raised on either side, apex of eminence im-
punctate or nearly so; basal area of propodeum slightly
shiny and conspicuously tessellate; pubescence of legs
dark obsoleta Lepeletier

b. Dorsal face of scutellum neither depressed along mid-
line nor raised on either side, uniformly subcontig-
uously punctate; basal area of propodeum shiny, with
very weak tessellation; hairs pale on metatibia and me-
tabasitarsus agilis F. Smith

8a. Apex of pygidial plate broad, deeply and angularly in-
cised (Fig. 42); scopa pale; yellowish marks often pres-
ent on some part of lower face 9

b. Apex of pygidial plate narrowly truncate or acute, never
incised (Figs. 41,43); scopa often dark; face rarely mac-
ulate 10

9a. First four abdominal terga black, with strong metallic

blue reflections species A

b. Entire abdomen dusky ferruginous

flavilabris Mocsary

10a. Lower half of clypeal disc slightly shiny to shiny, but
conspicuously roughened or ridged between punctures;
labrum and side of clypeus black; paraocular area usu-
ally black; integument of frons and mesopleuron black,
without metallic bluish or greenish-bronze reflections

11

b. Lower half of clypeal disc smooth and polished between
punctures; most of labrum, at least side of clypeus, and
paraocular area yellowish maculate; frons and meso-
pleuron with metallic bluish or greenish-bronze reflec-
tions plumipes F. Smith

1 la. Discs of second and third terga moderately to strongly
shiny, punctures various but usually not subcontiguous
and much coarser than hairs arising from them (if sub-
contiguous, scopa dark); scopa often dark; abdomen

often black 12

b. Discs of second and third terga dull, contiguously punc-
tate, punctures much coarser than hairs arising from

them; scopa pale; abdomen ferruginous

agiloides, new species

12a. Punctures of disc of second tergum, when visible, little
larger than hairs arising from them and finer than those
on disc of following segment; pubescence of thorax and/
or scopa often partly pale; basitibial plate and pygidium

various 13

b. Punctures of disc of second tergum subcontiguous, much
coarser than hairs arising from them and conspicuously
coarser than punctures of apical zone and of following
segment; thoracic and scopal hairs black; secondary
basitibial plate acute (Fig. 24); pygidial plate narrowly

V-shaped, secondary plate short and broad

sericea Friese

13a. Dorsal face of first tergum, across middle one-third,
polished, without evident punctures and with only a
few widely scattered, appressed simple hairs; pygidial
plate broadly V-shaped and broadly truncate at apex,
lateral margins somewhat convex at about midlength,
secondary plate elongate and with median raised ridge
extending from its apex to apical truncation (not always

visible in worn specimens) (Fig. 43); scopa pale

agilis F. Smith

b. Dorsal face of first tergum pubescent and/or punctate
across middle half at least at summit of declivity; py-

26 Contributions in Science, Number 347

Snelling: American Centridini

gidium not as above (Fig. 41); scopa pale or dark .14
14a. Scopa entirely pale; abdomen largely reddish . ... 15
b. Scopa entirely black or pale on tibia and brownish on

basitarsus; abdomen dark 16

15a. Mesepistemal pubescence dark brownish; disc of sec-
ond tergite with distinct, minute, close punctures; apex

of pygidial plate narrowly truncate

gelida, new species (part)

b. Mesepistemal pubescence pale, at least in part; disc of
second tergite without obvious minute punctures, but
with a few scattered, moderate, shallow punctures; apex

of pygidial plate acute fusciventris Mocsary

1 6a. Pubescence of first and second terga and of scopa black-
ish; clypeal punctures, except along impunctate mid-
line, uniformly subcontiguous on disc

obsoleta (Lepeletier)

b. Pubescence of adbominal terga pale (brownish golden
on second segment); tibial scopa mostly pale; punctures

of clypeal disc very irregularly spaced

gelida, new species (part)

Centris ( Me/anocentris ) agilis F. Smith

Figure 43

Centris agilis F. Smith, 1874:361. 3.

Centris ignita F. Smith, 1874:362. 2. NEW SYNONYMY.
Centris bakeri Friese, 1912:199. <3. Preoccupied.

Centris bakerel/a Friese, 1913:89. New name for C. bakeri
Friese, 1912, not C. bakeri Cockerell, 1912. NEW SYN-
ONYMY.

Epicharis cisnerosi Cockerell, 1949:180. 2. NEW SYNON-
YMY.

F. Smith (1874) described the two sexes as C. agilis and C.
ignita. This is a common species in Mexico and one that is
variable in the color of the abdominal integument. In both
sexes, the abdominal terga, beyond the blackish first segment,
may be wholly ferruginous. In these specimens the dense,
short, simple discal hairs appear to be yellowish red. In other
specimens, one or more of the following three segments may
be blackish and when this is the case, the discal hairs appear
to be yellowish or somewhat whitish, imparting a distinctly
“frosted” appearance.

Friese’s C. bakeri and Cockerell’s Epicharis cisnerosi are
based on specimens with dark tergites. Although I have seen
no type material of C. bakeri, the description matches well
the characteristics of dark males of C. agilis. I have examined
the type of Epicharis cisnerosi ; it is a normal, dark female
of C. agilis.

NEW RECORDS

MEXICO, CHIAPAS: 12, 10 mi. NW Comitan, 9 Aug. 1963
(F.D. Parker and L.A. Stange; UCD); 12, Municipio Ocozo-
cautla, El Aguacero de Derna, 762 m elev., 1 Sept. 1976
(D.E. and J.A. Breedlove; CAS); 12, 1833, Municipio Angel
Albino Corzo, Rio Custepec, below Finca Gadow, 853 m
elev., 12 Sept. 1976 (D.E. and J.A. Breedlove; CAS); 1 <3,
Sumidero, Tuxtla Gutierrez, 17 Aug. 1964 (E. Fisher and D.

Verity; LACM) 13, 82 mi. W Tuxtla Gutierrez, 2100 ft. elev.,
30 Aug. 1957 (H.A. Scullen; ORSU); 222, NW slope Cerro
Baul, 1768 m elev., W of Rizo de Oro, 12 Oct. 1979 (D.E.
and J.A. Breedlove; CAS). COLIMA: 13, 10 mi. W Colima,
1 Aug. 1954 (M. Cazier, W. Gertsch, Bradts; AMNH).
GUERRERO: 13, 4 mi. W Chilpancingo, 1530 m elev., 27
Aug. 1977 (E.I. Schlinger; UCB). JALISCO: 12, 25 mi W
Guadalajara, 4700 ft. elev., 29 Sept. 1957 (H.A. Scullen;
ORSU); 222, 5.6 km E Plan de Barranca, 914 nr elev., 25
Sept. 1976 (C.D. George and R.R. Snelling; LACM), on Cas-
sia sp.; 12, 3.4 km E Plan de Barranca, 960 m elev., 25 Sept.
1976 (C.D. George and R.R. Snelling; LACM), on Cosmos
sulphureus. MICHOACAN: 222, Tuxpan, 6550 ft. elev., 19
Sept. 1957 (H.A. Scullen; ORSU). MORELOS: 13, no lo-
cality or date (Crawford; LACM); 13, 7 mi. NE Yautepec,
4000 ft. elev., 18 Aug. 1962 (Univ. Kans. Mex. Exped.;
UKAN) on Leguminosae; 12, 0.6 km S Teacalco, 1021 m
elev., 1 6 Sept. 1976 (C.D. George and R.R. Snelling; LACM),
on Crota/aria incana; 13, Lake Tequesquitengo, 5000 ft. elev.,
13 Sept. 1957 (H.A. Scullen; ORSU). NAYARIT: 222, 8 mi.
N Tepic, 1 Sept. 1962 (D.H. Janzen; LACM). OAXACA: 2 33,
El Camaron, 20 Aug. 1959 (L.A. Stange and A.S. Menke;
UCB); 233, 48 mi. E La Ventosa, 21 July 1963 (J. Doyen;
UCB); 12, near Pinotepa Nacional, 200 m elev., 12 Oct. 1975
(J.L. Neff; LACM), on Legume 7 1 23; 222, 4 mi. W Zanatepec,
7 Oct. 1975 (J.L. Neff; LACM), on Cassia 7070; 12, Mixtla,
5600 ft. elev., 22 Aug. 1963 (Scullen and Bolinger; ORSU).
PUEBLA: 12, 16.1 km NW Izucar de Matamoros, 1250 m
elev., 1 7 Sept. 1 976 (C.D. George and R.R. Snelling; LACM),
on Caesalpinia cacalaco: 322, 22 km NW Izucar de Mata-
moros, 1 158 m elev., 21 Sept. 1976 (C.D. George and R.R.
Snelling; LACM), on Cassia laevigata. SINALOA: 622, 14
mi. SE Espinal, 14 Sept. 1964 (A.E. Michelbacher; UCB);
222, 33 km SE Esquinapa, 1 1 Sept. 1 974 (E.M. Fisher; LACM).
VER.4 CRUZ: 13, Catemaco, 6 Oct. 1976 (E. Barrera; LACM).
HONDURAS: 12, Zamorano, 2 Nov. 1946 (G. Cisneros;
USNM) (type of Epicharis cisnerosi). COSTA RICA, SAN
JOSE. 12, San Jose, no date (M. Valerio, USNM).

Centris ( Melanocentris ) agiloides, new species

Figures 20-23

DIAGNOSIS

Abdomen ferruginous in both sexes, second tergite dull, sub-
contiguously punctate; male with clypeus narrow, pubescence
of thoracic dorsum blackish except on metanotum and meta-
femur without ventral ridge; female with narrowly truncate
pygidial plate and pale scopa.

DESCRIPTION

HOLOTYPE MALE. Measurements (mm). Head width
6.41 (6.00-6.62); head length 4.97 (4.56-5.03); wing length
15.0 (14.0-16.5); total length 19.0 (17.0-21.0).

Head. 1.29 (1.29-1.37) times broader than long; occipital
margin slightly concave between tops of eyes; ocelli well
anterior to occipital margin; inner orbits moderately con-
vergent above, upper frontal width 0.65 (0.62-0.71) times

Contributions in Science, Number 347

Snelling: American Centridini 27

Figures 20-23. Centris (Melanocentris) agiloides, male seventh and eighth sternites and genitalia (ventral and dorsal views). Scale line = 1.00
mm.

lower frontal width. Mandible stout, tridentate, inner tooth
acute and nearly as large as second tooth. Labrum about 1.5
times wider than long, apex subacute; disc shiny between
dense to subcontiguous moderate punctures. Clypeus narrow,
1.1-1. 2 times wider than long; median impunctate line nar-
row and poorly defined, basal area of disc moderately shiny,
roughened and with sparse fine punctures, distal one-half,

more or less, shiny and weakly or not roughened and with
close to dense, fine punctures. Frons and preoccipital area
moderately shiny between dense to subcontiguous fine punc-
tures, except usual nearly impunctate areas adjacent to ocelli;
gena shiny between sparse to close fine, punctures. Interan-
tennal distance 1.62 (1.50-1.77) times antennal socket di-
ameter; antennocular distance 0.46 (0.32-0.47) times anten-

28 Contributions in Science, Number 347

Snelling: American Centridini

nal socket diameter; scape stout, 2.05 ( 1 .90-2.2 1 ) times longer
than wide; scape length 0.75 (0.69-0.76) times length of first
flagellar segment; first flagellar segment longer than following
three segments combined, 5.94 (5.00-5.53) times length of
second. Interocellar distance 1.35 (1.26- 1.47) times diameter
of anterior ocellus; ocellocular distance 0.38 (0.30-0.50) times
diameter of anterior ocellus; ocelloccipital distance 2.22 (2.16-
2.50) times diameter of anterior ocellus.

Thorax. Mesoscutum moderately shiny between dense to
subcontiguous, fine punctures which are a little more sepa-
rated posteromedially; scutellum with a low, rounded em-
inence on either side, moderately shiny between dense to
subcontiguous, fine punctures, shiny along midline; meta-
notum dull, conspicuously tessellate and with scattered fine
punctures; mesepisternum and metepistemum moderately
shiny, punctures subcontiguous to contiguous, fine, and shal-
low. Basal area of propodeum moderately shiny, tessellate
between sparse, fine punctures; side and posterior face sim-
ilar, but punctures dense to subcontiguous. Metafemur stout,
about 1.9 times longer than thick; thickest at basal one-third;
without ventral ridge; metabasitarsus weakly curved, about
four times longer than wide.

Abdomen. Dorsum of first tergite moderately shiny, weakly
tessellate between sparse, fine punctures; remaining tergites
dull and sharply tessellate between subcontiguous, moderate
punctures, sparser on fourth and following segments; seventh
tergum weakly bilobate at apex.

Terminalia. Distal process of seventh sternite (Fig. 20)
short, not well differentiated from disc; apical emargination
broad and shallow; hairs sparse, mostly short and weakly
plumose or simple. Median expansion of eighth sternite (Fig.
21) evenly rounded; apex slightly convex; hairs abundant,
long, conspicuously plumose. Dorsal process of gonocoxite
(Figs. 22, 23) nearly triangular; gonostylus nearly reaching
level of apex of penis valve; ventral lobe of penis valve only
slightly extended laterad of dorsal lobe.

Pilosity. Blackish brown on front and top of head, on tho-
racic dorsum, upper mesepisternum, anterior and middle
legs, and first tergite; hairs on discs of second and third ter-
gites very short, simple, decumbent, longer and more erect
on following segments, with some very long, plumose, red-
dish brown hairs, especially laterad; hairs on underside of
head, posterior margin of scutellum, metanotum, and re-
mainder of thorax light brown. Metafemur and metabasi-
tarsus with scopa-like yellowish hairs.

Color. Head and thorax blackish brown, legs and antenna
more rufescent. Large labral mark, large discal spot on clyp-
eus and narrow paraocular mark all very pale yellowish. Ab-
domen dull ferruginous. Wings dark brown, veins and stigma
blackish.

FEMALE. Measurements (mm). Head width 6.56-7.08;
head length 4.82-5.13; wing length 14.5-16.0; total length
20.0-24.5

Head. 1.26-1.38 times broader than long; ocelli well an-
terior to nearly flat occipital margin in frontal view; inner
orbits moderately convergent above, upper frontal width 0.84-
0.90 times lower frontal width. Mandible quadridentate, in-
ner tooth acute and slightly larger than subbasal tooth. La-

brum about twice broader than long, apex broadly rounded;
disc shiny between subcontiguous to contiguous, moderate
punctures. Clypeus mostly moderately shiny, but disc with
variable area which is conspicuously shiny (up to distal two-
thirds of median area); median impunctate line poorly de-
fined; disc with sparse moderate punctures and a few low,
irregular rugulae which are usually convergent distad or curved
mesad at their lower ends. Punctation of frons and occipital
area as described for male, but interspaces (including ocellar
area) moderately shiny and conspicuously tessellate; gena as
described for male. Interantennal distance 2.00-2.31 times
antennal socket diameter; scape robust, 1 .98-2. 1 3 times longer
than wide; scape length 0.71-0.77 times length of first fla-
gellar segment; first flagellar segment slightly longer than fol-
lowing three segments combined, 4.52-5.40 times length of
second. Interocellar distance 1.55-1.72 times diameter of
anterior ocellus; ocellocular distance 1.08-1.21 times di-
ameter of anterior ocellus; ocelloccipital distance 1.71-2.00
times diameter of anterior ocellus.

Thorax. As described for male, but scutellum tessellate
and slightly shiny, except in basal middle where it is shiny.
Basitibial plate slender, apex narrowly rounded.

Abdomen. Dorsal face of first tergite moderately shiny and
weakly tessellate between close to dense, moderate punctures;
second and following terga dull, contiguously and finely punc-
tate to tergal margins; apex of pygidial plate narrowly trun-
cate.

Pilosity. On head and thorax, as described for male, but
hairs of metanotum and propodeum often very pale brown-
ish; scopa yellowish white. Hairs on dorsal face of first tergite
erect, dark, plumose; dark, very short, appressed, and simple
on disc of second segment, becoming progressively longer
and less appressed on following segments; fourth and fifth
terga with a few suberect to erect dark bristles on either side;
prepygidial fimbria dark reddish brown.

Color. As in male, but without pale face marks.

TYPE MATERIAL

Holotype male: Finca La Selva, 500 ft. elev., near Puerto
Viejo, Heredia Prov., COSTA RICA, 3 June 1975 (D.R.
Perry), on Dipteryx panamensis, 0703-1 100, in Natural His-
tory Museum of Los Angeles County. Allotype: same locality
and collector, 6 May 1979, on Dussia sp., 1045-1 1 1 5 (LACM).
Paratypes (all COSTA RICA): 1 1 <33, 69$, same data as allo-
type (LACM); 13, same locality and collector, 4 June 1975,
on Dipteryx panamensis, 1 100-1400 (LACM); 13, same lo-
cality and collector, 14 June 1975, on Dipteryx panamensis,
0830-1030 (LACM); 13, same locality and collector, 24 July
1978 (LACM), on Hymenolobium sp.; 13, 8 km S Puerto
Viejo, Heredia Prov., 28-29 May 1971 (P.A. Opler; UCB),
on Ipomaea sp., pink fl.; 13, Zapote de Upala (vie. Bijagua),
Alajuela Prov., 19 May 1972 (F. Cordero; UCB); 1$, Dulce
Nombre, Cartago Prov., 25 Aug. 1967 (R.W. McDiarmid;
LACM).

ADDITIONAL SPECIMENS (not paratypes)

MEXICO, CHIAPAS: 13, Mahosik’, Tenejapa, 4800 ft. elev.,
9-12 July 1966 (D.E. Breedlove and J. Emmel; CAS). SAN

Contributions in Science, Number 347

Snelling: American Centridini 29

LUIS POTOSI: 13, 26 mi. SW Tamazunchale, 4-5 July 1964
(E. Fisher and D. Verity; LACM); 12, 15 mi. Xililla, 1350
m elev., 15 Aug 1977 (E.I. Schlinger; UCB). VER.4 CRUZ:
12, Pehuela, 13 Sept. 1974 (M. Sousa; LACM); 422, same
locality, 2 Aug. 1974 (A. Delgado S.; LACM), on “C. Doylei
0800-1002; 12, “Rig. de Cordoba (A. Genin; MNHN). COS-
TA RICA, SAN JOSE: 2SS, San Jose, no date (M. Valerio;
USNM). PANAMA, PANAMA: 13, Cerro Azul, N of To-
cumen, 28 Apr. 1958 (W.J. Hanson; UKAN); 13, Curundu,
19 May 1981 (R.W. Brooks; RWB), on Genipa americana.

ETYMOLOGY

Combines the Latin suffix -oides (resembling) to agilis, be-
cause of the superficial resemblance to C. agilis.

DISCUSSION

Variation in non-meristic characters is negligible. In some
specimens, particularly those of Mexico, the pubescence of
the metanotum and propodeum is not conspicuously paler
than that of the thoracic dorsum. In some males, the posterior
femur, tibia, and basitarsus may be extensively reddish, but
mostly dark in other specimens. The supraclypeal mark may
be present or absent, and when it is present, it is transverse
and very narrow. Although the scape is usually black in the
males available, there is a distinct ventral maculation in the
males from Mexico and Panama. One of the Mexican males
has the clypeus largely pale.

In the females there is considerable variation in the extent
of the shiny area on the clypeal disc. In most specimens the
shiny portion of the disc occupies about one-half the length
of the disc; in a few this is exceeded (up to nearly the entire
length) and in others reduced to the apicomedian one-fourth
of the segment. Similarly, the development of the oblique
rugulae of the clypeal disc is variable, and the rugulae, while
never becoming a dominant feature, are always present.

The uniformly subcontiguously punctate, dull, red abdo-
men is diagnostic for C. agiloides. The only species with a
similarly dull abdomen is C. sericea but in both sexes of that
species the abdomen is black, the posterior legs are black-
haired, the abdominal punctures become much finer toward
the margins of the segments, and the punctures of the third
segment are much finer than those of the second.

Centris {Melanocentris) flavilabris Mocsary

Centris flavilabris Mocsary, 1899:253. 2.

Centris flavilabris var. boliviensis Mocsary, 1899: 253. 2.

This is a primarily South American species, not previously
recorded from Central America.

NEW RECORDS

COSTA RICA, GUANACASTE: 13, Volcan Miravalles, 2
km W Rio Navinjo, 15 Mar. 1973 (P.A. Opler; UCB). PUN-
TARENAS: 12. Rincon, 25 Apr. 1975 (C.L. Hogue; LACM).

Centris ( Melanocentris ) fusciventris Mocsary

Figure 15

Centris fusciventris Mocsary, 1899:252. 2.

Centris fusciventris var. scutellata Friese, 1 900b:273. 3 2. NEW
SYNONYMY.

Centris (Melanocentris) fusciventris: Moure, 1950:388 (tax.).
Centris ( Melanocentris ) fusciventris scutellata: Michener,

1 954: 144 (distr.).

The var. C. f scutellata was described from both sexes from
Chiriqui, Panama. The nominate form occurs widely in South
America (Brazil, Colombia, Bolivia, Venezuela). Differences
between the two forms are slight and fall within the range of
variation seen in South American material.

NEW RECORDS

COSTA RICA, HEREDIA: 322, 13, Finca La Selva, near
Puerto Viejo, 6 May 1979 (D.R. Perry; LACM), on Dussia
sp. PANAMA, COLON: 233, Puerto Pilon, 22 km NE Santa
Rita, 23 May 1982 (D. Roubik; ROUB). DARIEN: 3 33, Bay-
ano Bridge, 184 km SE Canglon, 16 May 1980 (D. Roubik;
ROUB). PANAMA: 2 33, Panama, Curundu, 17 May 1981
(R.W. Brooks, RWB), on Genipa americana.

Centris ( Melanocentris ) gelida, new species

DIAGNOSIS

Female only: scopa yellowish on metatibia, brown on meta-
basitarsus; abdomen mostly ferruginous, dorsum shiny (ob-
scured by pubescence) between minute punctures; dorsal face
of first tergite pubescent and minutely punctate in middle;
pubescence of first and second tergites pale. Male unknown.

DESCRIPTION

HOLOTYPE FEMALE. Measurements (mm). Head width
7.49 (7.38-7.64); head length 5.13 (5.03-5.13); wing length
18.0 (17.5-19.0); total length 22.0 (22.0-25.0).

Head. 1.46 (1.46-1.49) times broader than long; occipital
margin nearly flat in frontal view and slightly below tops of
eyes; ocelli anterior to occipital margin; inner orbits strongly
convergent above, upper frontal width 0.85 (0.8 1-0.84) times
lower frontal width. Mandible stout, quadndentate. Labrum
about twice broader than long. Disc of clypeus slightly de-
pressed, moderately shiny (duller, obviously tessellate latero-
basad), with broad, poorly defined median impunctate line,
punctures fine and irregularly spaced a puncture diameter or
more apart. Frons roughened and slightly shiny between fine,
dense punctures; vertex slightly shiny and with punctures
minute and subcontiguous in ocellocular area, shiny between
dense to sparse fine punctures in preoccipital area; gena shiny
between dense to close, minute punctures. Interantennal dis-
tance 2.04 (2.00-2.24) times antennal socket diameter; an-
tennocular distance subequal to antennal socket diameter;
scape stout, about twice longer than wide, scape length 0.72
(0.63-0.72) times length of first flagellar segment; first fia-

30 Contributions in Science, Number 347

Snelling: American Centridini

gellar segment slightly longer than following three combined,
4.33 (4.35-4.65) times longer than second. Interocellar dis-
tance 1.60 (1.42-1.59) times diameter of anterior ocellus;
ocellocular distance 1.07 (1.00-1.07) times diameter of an-
terior ocellus; ocelloccipital distance 1.72 (1.68-1.82) times
diameter of anterior ocellus.

Thorax. Mesoscutum and dorsum of scutellum shiny be-
tween fine, dense to close punctures, posterior face of scu-
tellum with subcontiguous, fine punctures; scutellum weakly
depressed along ntidline; metanotum slightly shiny, sharply
tessellate between sparse, minute punctures; mesepisternum
moderately shiny between subcontiguous to dense, fine punc-
tures; basal area of propodeum slightly depressed on either
side, moderately shiny and tessellate between sparse, fine
punctures; disc shinier, closely and more finely punctate; side
shiny between fine, close punctures. Basitibial plate about
twice longer than wide, secondary plate narrowly rounded at
apex, primary plate more broadly rounded. Scopal hairs plu-
mose nearly to tips.

Abdomen. Summit of first tergite shiny between sparse
micropunctures in middle and close to sparse, minute punc-
tures laterad; discs of second and third tergites shiny between
dense to close, minute punctures; fourth tergite less shiny,
densely, finely punctate; fifth tergite slightly shiny, subcon-
tiguously, finely punctate. Pygidium narrowly truncate at apex,
sides straight, secondary plate acute at apex.

Color. Color generally blackish brown, distal half of sec-
ond, all of following tergites and all sternites ferruginous;
tarsi and underside of flagellum dark ferruginous. Wings
strongly brownish, veins and stigma blackish brown.

Pilosity. Dark brown to blackish on head, thorax, and legs,
except most of metatibial scopa yellowish and metabasitarsal
scopa light brown; hairs of mesoscutum very dense, con-
cealing surface; hairs of first tergite moderately long, erect,
plumose, dirty white; hairs of second to fifth tergites short,
simple, subappressed, whitish to yellowish brown, especially
on discs of second and third segments; prepygidial and py-
gidial fimbriae ferruginous; sternites with sparse erect whitish
hairs, longer along midline and forming weak distal fimbriae,
that of fifth segment ferruginous.

TYPE MATERIAL

Holotype female: Santa Rita, 10 mi. E Nahuala, 7100 ft.
elev.. Dept. Solola, GUATEMALA, 3 Sept. 1965 (S.J. Ar-
nold), on Canavalia villosa, in California Academy of Sci-
ences. Paratypes: 22$, same data as holotype (UCB); 12, 25
mi. S El Bosque, 5500 ft. elev., Chiapas, MEXICO, 5 Sept.
1965 (S.J. Arnold; UCB); one paratype in LACM, two in
UCB.

ETYMOLOGY

The specific name is a Latin word for "frosted” and refers
to the characteristic appearance of the abdominal dorsum.

DISCUSSION

An additional female, not a paratype, is from Municipio
Motozintla, ridge between Cerro Boqueron and Niguivil,

2438-2743 m elev., Chiapas, MEXICO, 1 5 Dec. 1 976 (D.E.
and J.A. Breedlove; CAS). This specimen is assumed to be
conspecific with the type series, agreeing closely in most fea-
tures. It does differ, however, in having the abdomen uni-
formly blackish, rather than mostly dull reddish. Abdominal
color is somewhat variable in some members of this sub-
genus.

The only other species in Central America with a similarly
“frosted” abdomen is C. agi/is. In C. agilis the disc of the
first tergite is polished and nearly devoid of punctures and
pubescence across most of its breadth and the secondary plate
of the pygidium is extended distad as a cariniform ridge.

Centris ( Melanocentris ) obsoleta Lepeletier

Figure 39

Centris obsoleta Lepeletier, 1841:153. 2.

Centris melanochlaena F. Smith, 1874:360. 3. NEW SYN-
ONYMY.

Epicharis zamoranensis Cockerell, 1949:480. 2. NEW SYN-
ONYMY.

DISCUSSION

I have examined the type of C. melanochlaena, from Ori-
zaba, Mexico. It corresponds to the concept of C. obsoleta,
in the sense of Friese (1900b) and subsequent workers, and
it is typical of Central American males since the clypeus is
almost wholly yellowish.

I have also seen the type and one cotype of Epicharis
zamoranensis. Cockerell described the thoracic dorsum as
being devoid of hairs; this is true of the type, but not of the
cotype. The hairs are worn off in the type specimen. There
is no doubt that E. zamoranensis is conspecific with C. ob-
soleta, a common species in Central America and northern
South America.

NEW RECORDS

MEXICO, NAYARIT: 12, 1 6 mi. S Acaponeta, 12 Aug. 1963
(M.G. Naumann; UKAN). OAXACA: 13, 522, Salina Cruz,
7 Sept. 1965 (D.H. Janzen; UKAN); 12, 12 mi. S Chivela,
18 Aug. 1959 (A.S. Menke and L.A. Stange; UCD); 13, 8 km
W Tehuantepec, 9-10 Aug. 1974 (E.M. and J.L. Fisher;
LACM). PUEBLA: 12, 16.1 km S Izucarde Matamoros, 1280
m elev., 17 Sept. 1976 (C.D. George and R.R. Snelling;
LACM), on Caesalpinia cacalaco. SINALOA: 12, 35 km SE
Esquinapa, 1 1 Sept. 1974 (E.M. Fisher; LACM). HONDU-
RAS: 222, Zantorano, 2600 ft. elev., 5 Nov. 1946 (G. Cis-
neros; USNM, LACM) (type and cotype E. zamoranensis).
COSTA RICA, GUANACASTE: 13, Hacienda Comelco, 8
km NW Bagaces, 1 9 Nov. 1 97 1 (P. A. Opler; UCB), on Stach-
ytarpheta frantzii.

Centris ( Melanocentris ) sericea Friese

Figures 24-28

Centris sericea Friese, 1899:41. 2.

Dr. R.W. Brooks has examined the type of C. sericea, a
female from an unspecified Mexican locality, and found it

Contributions in Science, Number 347

Snelling: American Centridini 31

to be conspecific with those recorded below. I have rede-
scribed C. sericea to facilitate its comparison with similar
species.

DIAGNOSIS

Abdomen black and hind legs black pubescent in both sexes.
Male metafemur swollen, without ventral ridge; inner man-
dibular tooth acute; clypeus about 1.2 times broader than
long; punctures of second tergum separated by a puncture
diameter or more. Female with punctures of disc of second
tergum coarser than those of apical zone; pygidial plate with
margins strongly convergent apicad, apex subacute; thoracic
pubescence wholly dark.

DESCRIPTION

MALE. Measurements (mm). Head width 5.90-6.3 1 ; head
length 4.46-4.72; wing length 15.0-16.0; total length 1 8.5—
20.0.

Head. 1 .32-1 .34 times broader than long, occipital margin,
in frontal view, slightly concave between tops of eyes, ocelli
well anterior to occipital margin; inner orbits moderately
convergent above, upper frontal width 0.68-0.70 times lower
frontal width. Mandible robust, tridentate, inner tooth acute
and subequal to middle tooth. Labrum about twice broader
than long; apical margin narrowly rounded; disc shiny be-
tween subcontiguous. moderate punctures. Clypeus narrow,
about 1 .2 times broader than long; disc moderately shiny to
shiny (apical area), minutely roughened between close to
sparse, moderate punctures, without distinct median im-
punctate area. Frons slightly shiny between dense, fine punc-
tures, area in front of anterior ocellus and laterad of posterior
ocelli impunctate, tessellate and moderately shiny; vertex
moderately shiny between dense, fine punctures; preocciput
similar but punctures sparse; gena moderately shiny, punc-
tures sparse to close, minute near eye grading to fine toward
gular margin. Interantennal distance 1.75-1.89 times anten-
nal socket diameter; antennocular distance 0.48-0.61 times
antennal socket diameter; scape stout, 1 .97-2.08 times longer
than broad; scape length 0.69-0.77 times length of first fla-
gellar segment; first flagellar segment longer than following
three combined, 4.41-5.44 times length of second. Inter-
ocellar distance 1.54-1.67 times diameter of anterior ocellus;
ocellocular distance 0.51-0.69 times diameter of anterior
ocellus ocelloccipital distance 2.03—2 . 1 5 times diameter of
anterior ocellus.

Thorax. Mesoscutum shiny between dense to subcontig-
uous, moderate punctures, interspaces tessellate and mod-
erately shiny in posterior middle; scutellum slightly shiny,
tessellate between dense to subcontiguous, moderate punc-
tures, a very low convexity on either side of middle; meta-
notum moderately shiny and sharply tessellate between sparse,
weak, fine punctures; mes- and metepistema moderately shiny
between dense to subcontiguous, moderate punctures. Basal
area of propodeum moderately shiny and sharply tessellate
between sparse, fine punctures; side moderately chiny and
roughened between close to dense, fine punctures. Metafemur
robust, about 1.8 times longer than thick, without ventral

ridge; metabasitarsus slender, about 3.6 times longer than
broad.

Abdomen. Elevated basal area of dorsal face of first tergite
much shorter, in middle, than apical zone, slightly shiny and
tessellate between sparse, fine punctures; apical zone of first
tergite long, shiny between scattered fine punctures; disc of
second tergum moderately shiny between dense, moderate
punctures which are conspicuously coarser and closer than
those of apical zone; disc of third tergite moderately shiny
and tessellate between close, fine punctures which are re-
placed with fine setigerous papillae on apical zone; fourth to
sixth tergites moderately shiny and tessellate between close,
slightly coarser setigerous papillae.

Terminalia. Seventh stemite (Fig. 25) with apical incision
deeper and more angular than in C. agiloides (Fig. 18) and
hairs more numerous and more conspicuously plumose.
Eighth stemite (Fig. 26) more angularly expanded in middle
than in C. agiloides ( Fig. 19). Gonostylus(Figs. 27, 28) ending
well short of level of end of penis valve; ventral lobe of penis
valve extended well beyond margin of dorsal lobe.

Pilosity. Blackish brown on head, thorax, legs, and first
tergite; second and following tergites with whitish hairs, very
short and appressed on second segment, becoming progres-
sively longer and more erect on succeeding segments; golden-
brown hairs laterad on fourth and following terga, on pygidial
plate and on sternites.

Color. Integument blackish brown on most areas; antenna,
tegula, and legs dark reddish brown; apical abdominal seg-
ments light brown. The following pale yellow: large spot on
labrum; clypeus, except small sublateral brown spots near
base; transverse supraclypeal mark; paraocular area, not ex-
tending above lower margin of antennal socket; preapical
spot on underside of scape. Wings dark brown, veins and
stigma blackish.

FEMALE. Measurements (mm). Head width 6.67-7.08;
head length 4.92-5.33; wing length 16.0-17.0; total length
20.5-23.0.

Head. 1.31-1.37 times broader than long; occipital margin,
in frontal view, nearly flat and, in center, slightly above level
of tops of eyes; ocelli well below occipital margin; inner orbits
weakly convergent above, upper frontal width 0.85-0.93 times
lower frontal width. Mandible stout, quadridentate, inner
tooth acute and subequal to adjacent tooth. Labrum about
twice broader than long, apex narrowly rounded, shiny be-
tween subcontiguous, moderate punctures. Clypeus as de-
scribed for male, but about 1.3 times broader than long and
median impunctate line a little more evident, but still poorly
defined. Punctation of frons, occipital area, and gena as in
male. Interantennal distance 2.37-2.53 times antennal socket
diameter; scape stout, 2.00-2.27 times longer than broad,
scape length 0.61-0.73 times length of first flagellar segment;
first flagellar segment longer than following three segments
combined, 4.64-5.41 times longer than second. Interocellar
distance 1.66-1.95 times diameter of anterior ocellus; ocel-
locular distance 1 .24-1 .45 times diameter of anterior ocellus;
ocelloccipital distance 1.87-1.97 times diameter of anterior
ocellus.

Thorax. Punctation and form as described for male. Basi-

32 Contributions in Science, Number 347

Snelling: American Centridini

Figures 24-28. Centris ( Melanocentris ) sericea, female basitibial plate, male seventh and eighth stermtes and genitalia (ventral and dorsal
views). Scale line (25-28) = 1.00 mm.

tibial plate (Fig. 24) elongate, apices of primary and second-
ary plates narrowly rounded.

Abdomen. First tergum as described for male, but tessellate

area limited to lateral areas of dorsal face; disc of second
tergum slightly shiny and tessellate between fine, subcontig-
uous punctures which are conspicuously coarser than punc-

Contributions in Science, Number 347

Snelling: American Centridini 33

turesofapical zone; disc of third tergite slightly shiny between
dense, minute and scattered, fine punctures; fourth tergite
slightly shiny, more sharply tessellate between dense, fine
punctures; fifth tergite duller between subcontiguous, fine
punctures and scattered setigerous papillae. Margins of py-
gidial plate strongly convergent apicad, apex narrowly trun-
cate or subacute; secondary plate short and broad.

Pilosity. As described for male; prepygidial fimbria golden
brown.

Color. As described for male, but pale facial marks absent
and fifth tergite dull reddish.

SPECIMENS EXAMINED

MEXICO, MEXICO: 12, Canon de Lobos, 1200 m elev.,
12-13 July 1969 (M. Sousa; LACM). MORELOS: 222, Yau-
tepec, 3 1 July 1963 (F.D. Parker and L. A. Stange; UCD); 12,
4 mi. SW Yautepec, 3800 ft. elev., 2 July 1961 (C.D. Mich-
ener; UKAN), on Cassia sp.; 13, 4.3 mi. W Yautepec. 4000
ft. elev., 17 Aug. 1962 (Ordway and Marston; UKAN). NA-
YARIT: 233, Km 190, 1200 nr elev., NW of Guadalajara, 6
Sept. 1975 (J.L. Neff; LACM), on “legume 7046”; 922, 8 mi.
N Tepic, 1 Sept. 1962 (D.H. Janzen; UCB), on Crotalaria
sp.

DISCUSSION

Although the male of C. sericea closely resembles that of C.
obsoleta, the clypeus is broader in the latter species, at least
1.4 times broader than long. The inner margins of the eyes
are more strongly convergent above in C. obsoleta, so that
the distance between the eyes at their upper ends is less than
the distance between the lateral margins of the two posterior
ocelli. In C. sericea males the transocellar distance is less
than the distance between the eyes at their upper ends. These
same features will readily distinguish C. sericea from C. agi-
lis, as will the less shiny, uniformly dark abdomen and the
lack of the pale hairs on the hind legs. From males of C.
agiloides, this species may be separated by the blackish ab-
domen, lack of pale pubescence on the hind legs, the mostly
pale clypeus, and the generally more uniformly dark thoracic
pubescence, as well as differences in abdominal punctation.

Females of C. sericea, by virtue of their uniformly dark
color and narrowly truncate pygidial plate can only be con-
fused with those of C. obsoleta. They differ from that species,
however, in that the disc of the second tergite is densely
punctate, the punctures conspicuously coarser than those of
the apical zone and those on the disc of the following segment.
Additionally, the clypeus is less closely punctate and the
pygidial plate is more strongly narrowed distad and less
broadly truncate.

Centris (. Melanocentris ) p/umipes F. Smith

Centris plumipes F. Smith, 1854:373. 2 3.

This species, originally described from Santarem, Brazil, is
easily recognized by the features cited in the key.

NEW RECORDS

COSTA RICA, HEREDIA: 3 33, Finca La Selva, near Puerto
Viejo, 6 May 1979 (D.R. Perry; LACM), on Dussia sp.; 533,

same locality and collector, 24 July 1979 (LACM), on Hy-
menolobium sp.

Centris (Melanocentris) species A

Figure 42

This is known only from four females. In this species, pu-
bescence is dark, except light brownish on the posterior part
of the scutellum and yellowish on the scopa, the apex of the
pygidial plate is angularly emarginate, and the clypeus is
conspicuously maculate. Although the integument is gener-
ally blackish, the abdominal terga have weak metallic bluish
reflections.

SPECIMENS EXAMINED

COSTA RICA, HEREDIA: 12, Finca La Selva, near Puerto
Viejo, 6 May 1979 (D.R. Perry; LACM), on Dussia sp. PAN-
AMA, CANAL ZONE: 12, Barro Colorado Island. 9 July
1979 (H. Wolda; ROUB); 12, Frijoles, 19 May 1980 (K.E.
Steiner; UCD), on Byrsomma crassifolia\ 12, Pipeline Road,
3.7 mi. NW Gamboa, 9 May 1980 (K.E. Steiner; LACM),
on B. crassifo/ia.

Subgenus Trachina Klug

Trachina Klug, 1807:226. Type-species: Centris longimana
Fabricius, 1804; monobasic.

Paremisia Moure, 1945b:406. Type-species: “ Paremisia li-
neolata (Lepeletier, 1841)” = Bombus similis Fabricius,
1804.

Trachina is a member of a small group of subgenera, the
other two being Heterocentris and Hemisiella, characterized
by the three-segmented maxillary palp in both sexes, the
swollen hind legs of the male and the metabasitarsus of the
male with a sharp posterior carina or ridge which terminates
in a raised tooth (Fig. 18).

Michener (1951) suggested that these three subgenera might
well be united into a single genus. While there are some
similarities, it seems to me that Trachina, at least, is distinct
from both Heterocentris and Hemisiella. The latter two sub-
genera share a procoxal modification which appears to be
unique in Centris. There is a sharply depressed groove along
the inner, ventral margin of the procoxa; such a groove is
not present in Trachina or the other subgenera. Females of
Heterocentris and Hemisiella have some scopal hairs, es-
pecially basad on the metatibia, simple; in Trachina and
other subgenera, the scopal hairs are conspicuously plumose.
Trachina females possess a distinct secondary basitibial plate;
in Heterocentris and Hemisiella, the basitibial plate has a
basal convexity, but no secondary plate.

Both sexes of Trachina differ from species of Hemisiella
and Heterocentris in possessing a longitudinal, median yellow
line on the clypeus. Sometimes, this line may not be evident
when the black of the clypeus is reduced to a pair of subbasal,
sublateral spots. Clypeal maculae in Heterocentris and Hemi-
siella are usually present as a pair of subapical spots or a
subapical band (females, some males) or else the clypeus is
yellow except across the base (some males). Males of Tra-

34 Contributions in Science, Number 347

Snelling: American Centridini

Figures 29-37, left basitibial plate of female: 29, Centris (C. ) flavifrons ; 30, C. (C.) flavofasciata\ 31, C. (C.) aethiocesta ; 32, C. (C.) caixensis ;
33, C. (C.) meaculpa ; 34, C. ( Trachina ) eurypalana: 35, C. (T.) fuscata\ 36, C. (7’.) dentala\ 37, C. ( /'. ) xochipillii.

china further differ in that the apical lobe of the seventh
sternite is greatly reduced, so that the segment is essentially
a transverse band.

KEY TO NORTH AMERICAN TRACHINA

la. Male, antenna 1 3-segmented, basitibial plate absent

Contributions in Science, Number 347

Snelling: American Centridini 35

Figures 38-43, pygidial plate of female: 38, Centris (C.) flavifrons\ 39, C. (C.) flavofasciata: 40, C. ( C .) aelhiocesta\ 41, C. (Melanocentris)
obsolete ;; 42, C. (M.) species A; 43, C. (A/.) agilis.

36 Contributions in Science, Number 347

Snelling: American Centridini

b. Female, antenna 1 2-segmented, basitibial plate present

10

2a. Ocellocular distance greater than diameter of anterior
ocellus; in full frontal view, distance from anterior ocel-
lus to clypeal base a little shorter than distance from

anterior ocellus to occipital margin 3

b. Ocellocular distance less than diameter of anterior ocel-
lus; in full frontal view, distance from anterior ocellus
to clypeal base much greater than distance from an-
terior ocellus to occipital margin 4

3a. Thoracic pubescence uniformly blackish, except white
on scutellum and metanotum; metabasitarsus with sharp
ridge along posterior margin, terminating in sharp spi-

niform process beyond midlength (Fig. 18)

labiata Friese

b. Flairs of mesoscutum dark brownish distally, base and
branches pale, so that pubescence appears “clouded”;

metabasitarsus without posterior ridge (Fig. 19)

longimana (Fabricius)

4a. Abdomen largely or wholly ferruginous; if brownish or
blackish, hairs of mesoscutum red or pale ochreous .

5

b. First three, and most of fourth, terga blackish, apical
band of fourth and all of fifth to seventh terga pale
reddish; hairs of mesoscutum black except for band of

whitish hairs across anterior one-fourth

vidua Mocsary

5a. At least first four terga dark brown to blackish (fourth

may be ferruginous on apical margin 6

b. Abdomen largely or entirely ferruginous 7

6a. Terga 5-7 dark, with glistening whitish hairs; meso-
scutal hairs pale to dark ferruginous; in frontal view,

occiput arcuately raised above tops of eyes

si mi lis (Fabricius)

b. Terga 5-7 ferruginous, with yellowish to ferruginous
hairs; mesoscutal hairs pale ochreous; in frontal view,
occiput flat or slightly concave between tops of eyes

xochipillii, new species

7a. Procoxa without distal spine; wings uniformly light yel-
lowish brown 8

b. Procoxa with slender distal spine (hidden in dense pu-
bescence); forewing, basad of basal vein, clear and col-
orless and distinctly brownish beyond basal vein ....

eurypatana, new species

8a. Hairs of mesoscutum pale ochreous or ferruginous;
metafemur usually stout, subbasal ventral process usu-
ally tooth-like (Fig. 57, 58); second and third terga mod-
erately shiny, disc finely tessellate; middle and hind
legs with conspicuous brownish to blackish pubescence

9

b. Hairs of mesoscutum (and of most of thorax) whitish,
though often slightly dusky anteriorly; metafemur less
stout, subbasal ventral process low and not tooth-like
(Fig. 56); second and third terga shiny, discs weakly or
not at all tessellate; pubescence of legs whitish to pale
ferruginous, ferruginous on inner surface of metabasi-
tarsus heithausi Snelling

9a. Disc of seventh sternite triangular, apex acute (Fig. 46);
apical swelling of eighth sternite short and broad (Fig.

47); basal lobes of penis valve short and broad

dentata F. Smith

b. Disc of seventh sternite quadrate, apical margin straight
or concave (Fig. 44); apical swelling of eighth sternite
long and narrow (Fig. 45); basal lobes of penis valve

relatively slender fuscata Lepeletier

10a. Hairs of mesoscutum dark brownish to blackish and
either (a) with a transverse band of white hairs across
front of mesoscutum or (b) hairs of scutellum whitish

11

b. Hairs of mesoscutum whitish, yellowish, red or with
dark apices, but without sharply contrasting pale hairs

anteriorly or on scutellum 12

11a. Mesoscutum with a band of whitish hairs across an-
terior margin and hairs of scutellum and metanotum
dark; first four terga blackish (fourth pale reddish api-

cally), last two segments pale reddish

vidua Mocsary

b. Mesoscutum without pale hairs anteriorly, but whitish
hairs on posterior margin of scutellum and all of meta-
notum; terga wholly ferruginous labiata Friese

1 2a. First three terga dark brownish to blackish 13

b. First three terga ferruginous 14

13a. Terga 4 and 5 dark, margins colorless, with long, glis-
tening whitish hairs; scopa black; mesoscutal hairs usu-
ally dark reddish; tergum 4, across middle, with fine

dense piligerous punctures similis (Fabricius)

b. Terga 4 and 5 ferruginous, with yellowish hairs, scopa
pale yellowish; mesoscutal hairs ochreous; tergum 4,
across middle, with sparse fine piligerous punctures

xochipillii, new species

14a. Head width less than 5.75 mm; in frontal view, occiput
weakly convex between tops of eyes; ocellocullar dis-
tance less than 1.5 x diameter of anterior ocellus; hairs

of thoracic dorsum whitish to ferruginous 15

b. Head width more than 7.0 mm; in frontal view, occiput
arcuately raised above tops of eyes; occellocular dis-
tance about twice diameter of anterior ocellus; hairs of
thoracic dorsum with blackish shafts and pale branches

near base longimana (Fabricius)

15a. Thoracic pubescence ochreous to light brownish fer-
ruginous; scape distinctly maculate beneath; second and
third terga finely and closely punctate, surface mod-
erately shiny 16

b. Thoracic pubescence whitish, tinged with pale brown-
ish on mesoscutum; scape obscurely, if at all, maculate;
second and third terga shiny between fine punctures
mostly separated by twice a puncture diameter or more

heithausi Snelling

16a. Wings light to medium yellowish brown; lower margin
of secondary basitibial plate strongly oblique (Fig. 36)
or, if somewhat transverse, posterior margin is contin-
uous with that of primary plate (Fig. 35) 17

b. Wings blackish brown; lower margin of secondary basi-
tibial plate transverse, anterior margin curved basad

Contributions in Science, Number 347

Snelling: American Centridini 37

well before anterior margin of primary plate (Fig. 34)

eurypatana, new species

1 7a. Basitibial plate broad, lower margin of secondary plate
usually transverse across middle portion, posterior
margin continuous with that of primary plate (Fig. 35)

fuscata Lepeletier

b. Basitibial plate narrow, lower margin of secondary plate
strongly oblique throughout, posterior margin extended
over that of primary plate (Fig. 36) . dentata F. Smith

Centris ( Trachina ) dentata F. Smith

Figures 36, 46, 47, 58

Centris dentata F. Smith. 1854:374. <3.

Centris proximo Friese, 1899:45. <3 2.

Michener (1954) recorded C. dentata from Panama. In ad-
dition to material from Panama, I have seen several females
from Mixtepec, Oaxaca, Mexico, 28 Feb. 1974 (M. Sousa;
LACM). This appears to be an uncommon species in Central
America, although widely distributed in South America; the
types of both C. dentata and C. proximo were from Brazil.

Centris ( Trachina ) eurypatana, new species

Figures 34, 48-51, 57

DIAGNOSIS

Male distinguished from all other Trachina by the presence
of a slender spine at the apex of the procoxa. Female separable
from other Trachina by the following combination: pubes-
cence pale ochreous, abdomen ferruginous, wings dark brown,
posterior margin of secondary basitibial plate extended be-
yond that of primary plate.

DESCRIPTION

HOLOTYPE MATE. Measurements (mm). Head width
5.90; head length 4.10; wing length 13.0; total length 15.0.

Head. 1 .44 times broader than long; occipital margin weakly
convex in frontal view, ocelli well anterior to margin; inner
orbits strongly convergent above, upper frontal width 0.68
times lower frontal width. Mandible slender, tridentate, inner
tooth large and its lower edge slightly sinuate. Labrum about
1 .4 times broader than long, apical margin broadly rounded;
disc shiny between subcontiguous, fine punctures. Clypeus
about 1.4 times broader than long; disc shiny across apical
margin, otherwise slightly shiny and distinctly tessellate be-
tween dense to subcontiguous, fine punctures, impunctate
median line narrow and slightly elevated. Frons moderately
shiny between dense to subcontiguous, fine to moderate
punctures, sparsely punctate areas adjacent to ocelli mod-
erately shiny and distinctly tessellate; preoccipital area shiny
between dense subcontiguous, fine to minute punctures; gena
shiny between close to dense, fine punctures. Interantennal
distance 0.82 times antennal socket diameter, scape robust,
1.97 times longer than wide; scape length 0.70 times length
of first flagellar segment; first flagellar segment 5.63 times
longer than second, distinctly longer than combined lengths
of second to fourth segments. Interocellar distance 1 .90 times

diameter of anterior ocellus; ocellocular distance 0.55 times
diameter of anterior ocellus; ocelloccipital distance 2.84 times
diameter of anterior ocellus.

Thorax. Mesoscutum shiny between subcontiguous, fine
punctures which become well separated posteromedially;
scutellum shiny between scattered, minute to fine punctures;
metanotum dull, sharply tessellate and with scattered, minute
punctures; mesepisternum and metepisternum shiny be-
tween dense to subcontiguous, fine to moderate punctures.
Basal area of propodeum shiny and weakly tessellate between
sparse to close, moderate punctures; side and disc shiny be-
tween close to dense, fine to moderate punctures. Procoxa
with narrow, distal, spiniform process; metafemur stout, about
twice longer than thick, ventral process stout (Fig. 57); pro-
cess on anterior margin of metatibia slender, spiniform;
metabasitarsus about 3.2 times longer than broad, posterior
ridge ending at about midlength.

Abdomen. Dorsal face of first tergite moderately shiny and
tessellate between sparse, minute punctures; disc of second
tergite shiny and weakly tessellate between sparse to dense,
fine punctures, punctures of apical zone minute; third tergite
similar, but punctures dense; fourth tergite similar to third,
but punctures variably spaced from sparse to subcontiguous;
fifth tergite similar to fourth, but punctures sparse; apex of
seventh tergite distinctly bilobed, hidden under dense hairs.

Terminalia. Process of seventh sternite (Fig. 48) short,
broad, apical margin convex; margin with long, plumose
hairs, disc with a few short, simple hairs. Eighth sternite (Fig.
49) moderately flared preapically, apex acute; hairs fine, plu-
mose. Dorsal process of gonocoxite short, broad, apex round-
ed; distal tubercles of gonostylus not visible in dorsal view
(Figs. 50, 5 1 ).

Pilosity. Generally pale ochreous, slightly brownish across
vertex, on thoracic dorsum, and on legs, darker on meso-
and metatibiae and tarsi; some brown hairs on inner surface
of meso- and metatibiae and basitarsi. Second tergite with
long, suberect, plumose, blackish brown hairs across base,
discs of second to fourth terga with sparse, simple, dark hairs
which become progressively longer and more erect on suc-
ceeding segments; hairs on fifth to seventh terga long, sub-
erect to erect, slightly yellowish to ferruginous. Ventral ab-
dominal pubescence dense, light yellowish brown.

Color. Head and thorax blackish brown, abdomen ferru-
ginous; antenna and legs variably light to dark reddish brown.
The following yellow: mandible, except apical teeth; labrum;
clypeus, except a pair of brown submedian spots near base;
transverse supraclypeal stripe; paraocular area, upper end
constricted and terminating on eye margin at about midlevel
of antennal socket; broad ventral stripe on scape; dorsal,
apical spot on profemur; basal spot on pro- and mesotibia.
Tegula clear yellowish-brown. Wings clear and very light
brown basad of vein M, darker brown distad; veins and
stigma blackish brown.

FEMALE. Measurements (mm). Head width 5.87-6.10;
head length 4.05-4.27; wing length 11.5-12.5; total length
15.0-17.0.

Head. 1.43-1.44 times broader than long; in frontal view,
occipital margin gently convex, ocelli well anterior to margin;

38 Contributions in Science, Number 347

Snelling: American Centridini

46

Figures 44-47, seventh and eighth sternites of male: 44-45, Centris ( Trachina) fiuscata: 46-47, C. (T.) dentata. Scale line = 0.50 mm.

inner orbits moderately convergent above, upper frontal width
0.87-0.89 times lower frontal width. Mandible tridentate,
inner tooth large, blunt. Labrum about 1.5 times broader
than long, margin broadly rounded; disc shiny between sub-
contiguous, fine to moderate punctures. Clypeus about 1 .6
times broader than long, otherwise as described for male.
Frons and preoccipital area as described for male; gena shiny,
punctures close and minute near eye, becoming dense and
fine ventrad. Interantennal distance 2.79-3.00 times anten-
nal socket diameter; antennocular distance 1.61—1.71 times
antennal socket diameter; scape stout, 2.03-2. 1 7 times longer
than wide, scape length 0.67-0.72 times length of first fla-
gellar segment; first flagellar segment 5.63-5.88 times longer

than second, longer than following three segments combined.
Interocellar distance 2. 1 4—2.30 times diameter of anterior
ocellus; ocellocular distance 1.33-1.48 times diameter of an-
terior ocellus; ocelloccipital distance 2.80-3.00 times di-
ameter of anterior ocellus.

Thorax. As described for male. Procoxa without ventral
spine; basitibial plate (Fig. 34) narrowly rounded at apex,
posterior margin of secondary plate beyond that of primary
plate.

Abdomen. First two terga as described for male, third with
punctures of disc minute and sparse, becoming more minute
and scattered in apical zone; fourth tergum less shiny than
third, punctures very irregularly spaced, moderate and some-

Contributions in Science, Number 347

Snelling: American Centridini 39

what elongate, close to dense. Pygidial plate with margins
strongly convergent distad, apex narrowly truncate; second-
ary plate obsolete.

Pilosity. About as described for male, but some hairs on
disc of fourth tergite at least weakly plumose; scopa pale
yellowish; prepygidial fimbria light golden brown.

Color. As described for male, except mandibular apex more
extensively dark, dark clypeal spots reaching base, supra-
clypeal spot greatly reduced or absent and legs approximately
concolorous with abdomen. Wings darker brownish.

TYPE MATERIAL

Holotype male: Estacion Biologia Chamela, Jalisco, MEX-
ICO, 17 May 1980 (S.H. Bullock, #373), in Natural History
Museum of Los Angeles County. Allotype: same locality and
collector, 7 Apr. 1982 (#922; LACM). Paratypes (all MEX-
ICO): 12, same locality and collector, 12 May 1980 (#372);
222, 30 mi. E Villa Union, 570 m elev., Sinaloa, 10 Mar.
1980 (J.L. Neff; NEFF). One paratype each in collections of
the Estacion Biologia de Chamela and in the personal col-
lection of J.L. Neff, the remaining specimens in the Natural
History Museum of Los Angeles County.

ETYMOLOGY

The specific epithet combines the Greek eurys (broad) and
patanas (plate), in allusion to the broad secondary basitibial
plate of the female.

DISCUSSION

The male is easily distinguished from all previously described
species of Trachina by the presence of a distinct spine-like
process at the apex of the procoxa. It is otherwise very similar
to the males of C. dentata, C. fuscata, C. heithausi, and C.
xochipillii. All of these differ from males of C. eurypatana
in having lighter colored wings, particularly that of C. heit-
hausi, which further differs in the paler pubescence of the
head and thorax. Males of C. dentata have the ventral tooth
of the metafemur higher and more slender and the tooth on
the anterior margin of the metatibia is stouter than in C.
eurypatana. The brown, rather than ferruginous, first three
abdominal terga will differentiate C. xochipillii from C. eu-
rypatana.

The female of C. eurypatana is best recognized from other
members of this complex by the broad secondary basitibial
plate, ochreous pubescence, dark wings, and wholly red ab-
domen. In C. xochipillii the first three abdominal segments
are brown; in C. heithausi the thoracic pubescence is whitish
and the wings only faintly brown; in C. dentata the median
line of the clypeus is shiny and the punctures on either side
are moderate rather than fine, and the second tergite is duller,
more sharply tessellate and closely punctate. Since the sec-

ondary basitibial plate of C. fuscata does not overhang the
first, this species is easily separated from C. eurypatana.

Centris ( Trachina ) fuscata Lepeletier

Figures 34, 44, 45

Centris fuscata Lepeletier, 1841:167. <3.

Centris bimaculata Lepeletier, 1841:168. 2.

This species, also described from Brazil, is much more com-
mon in Central America than the similar C. dentata. Females
of C. fuscata, however, have a narrow secondary basitibial
plate (Fig. 35), a feature which will separate this species from
superficially similar species. Males most closely resemble
those of C. dentata, but have the ventral tooth of the meta-
femur short and stout; in C. dentata this tooth is long, slender,
and somewhat curved.

I have seen Central American material from Mexico, Gua-
temala, Costa Rica, and Panama. Michener (1954) has also
recorded C. fuscata from Panama and Lutz and Cockerell
(1920) cite it from Guatemala.

Centris ( Trachina ) heithausi Snelling

Figure 56

Centris ( Trachina ) heithausi Snelling, 1974:20-23. <3 2.

This species was described from many specimens from
Guanacaste Province, Costa Rica. I have seen numerous ad-
ditional specimens from the same general area. One female,
however, was collected 20 km SSE of Chiquimula, Depto.
Chiquimula, Guatemala, 25 Feb. 1966 (D.P. Gregory; UCB).

Centris ( Trachina ) labiata Friese

Figure 18

Centris labiata Friese, 1904:91. <3.

Centris schwarzi Cockerell, 1919:192. 2. NEW SYNONY-
MY.

This is evidently not a common species. Friese’s male type
is from San Carlos, Costa Rica. The type of C. schwarzi is
from Alta Vera Paz, Guatemala. I have seen the type of C.
schwarzi, in the USNM, and it is the same as females I had
already associated with C. labiata males. Both sexes are char-
acterized by the dark thoracic pubescence (but pale on the
scutellum and metanotum) and the red abdomen.

NEW RECORDS

MEXICO, OAXACA: 2<3<3, 20 mi. E El Cameron, 21 July
1956 (J.W. MacSwain; UCB); 2<3<3, 19 mi. W Tequisistlan,
29 Aug. 1970 (E.M. and J.L. Fisher; LACM). VERA CRUZ:
1<5, El Palmar, 28 Mar. 1954 (D.H. Janzen; LACM). YU-
CATAN: 12, Piste, 29 June 1967 (E.C. Welling; LACM).
BELIZE: 12, no further data (LACM).

Centris ( Trachina ) longimana Fabricius

Figure 19

Centris longimana Fabricius, 1804:356. 2 <3.

40 Contributions in Science, Number 347

Snelling: American Centridini

Figures 48-51. Centris (Track ina) eurypatana, male seventh and eighth sternites and genitalia (ventral and dorsal views). Scale line = 1.00
mm.

Centris personata F. Smith, 1874:362. <3.

Michener (1954) reported C. longimana from several Pan-
amanian localities. In addition to specimens from Panama,
I have seen material from Nicaragua and Costa Rica.

Centris ( Trachina ) similis (Fabricius)

Bombus similis Fabricius, 1 804:35 1 . 9.

Centris lineolata Lepeletier, 1841:158. 9.

Centris lineolata castaneiventris Mocsary, In Friese, 1899:
288.

Centris (Paremisia) similis: Moure, 1 960b: 1 30-13 1 .

This is a common species in northern South America (Trin-
idad, Guyana, French Guiana) south to Brazil and Peru.
There are no previous reports of its presence in Central
America. I have seen the following Central American spec-
imens.

NEW RECORDS

COSTA RICA, PUNTARENAS: 19, 1.8 mi. W Rincon, 4
Mar. 1971 (J.P. Donahue and C.L. Hogue; LACM). SAN
JOSE: 19. Pozo Azul, Junction Rios Parrita and Candelaria,
85 m elev., 9 Dec. 1961 (A. Wille; UKAN). PANAMA.
PANAMA: 2i3<3, 15 km E Chepo, Llano Carti Rd., 18 Jan.

Contributions in Science, Number 347

Snelling: American Centridini 41

56 57 58

Figures 52-55. Centris (Trachina) xochipillii, male seventh and eighth sternites and genitalia (ventral and dorsal views). Scale line = 1.00 mm.
Figs. 56-58, outline of ventral metafemoral process of male: 56, C. (T.) heithausr, 57, C. (T.) eurypatana\ 58, C. (T.) dentata.

42 Contributions in Science, Number 347

Snelling: American Centridini

1980 (D. Roubik, #6; ROUB); 292, 15, 15 km NE Chepo, 19
Dec. 1980 (D. Roubik, #48; ROUB).

Centris ( Trachina ) vidua Mocsary

Centris vidua Mocsary, 1899:252. 5.

This species was described from Honduras (San Pedro Sula)
and has been reported by Friese (1900b) from Orizaba, Vera
Cruz, Mexico. I have seen the following specimens of this
uncommon bee.

NEW RECORDS

BELIZE: 1 2, Belize (no further data; LACM). COSTA RICA,
CARTAGO: 15, Turrialba, 18 Oct. 1947 (A. Svihla; LACM).
PANAMA, BOCAS DEL TORO: 12, Almirante, Sept. 1963
(LACM). COLON: 222, 15, 5 km SW Colon, 30 Jan. 1980
(D. Roubik, #12; ROUB).

Centris ( Trachina ) xochipillii, new species

Figures 37, 52-55

DIAGNOSIS

At least first three terga brown, apical segments ferruginous;
male ocellocular distance less than ocellar diameter, occipital
margin at most weakly convex in frontal view, thoracic pu-
bescence ochreous; female with ochreous thoracic pubes-
cence, posterior margin of secondary plate of basitibial plate
overhanging that of primary plate, scopa yellowish.

DESCRIPTION

HOLOTYPE MALE. Measurements (mm). Head width
5.23 (5.13-5.74); head length 3.79 (3.59-4.05); wing length
13.5 (12.0-14.0); total length 15.0 (13.0-17.0).

Head. 1 .38 ( 1 .38— 1.51) times broader than long; in frontal
view, occipital margin flat or slightly convex, ocelli well be-
low margin; inner eye margins strongly convergent above,
upper frontal width 0.74 (0.69-0.75) times lower frontal width.
Mandible tridentate, inner tooth large, its lower margin slightly
concave in outline. Labrum about 1 .7 times wider than long,
apical margin broadly rounded; disc shiny between subcon-
tiguous, fine to moderate punctures. Clypeus about 1.6 times
broader than long; disc dull and densely tessellate at base,
shiny and weakly tessellate on about distal one-fourth, me-
dian line raised and impunctate, disc otherwise densely to
subcontiguously punctate, punctures fine to moderate. Frons
and ocellar area closely punctate, except usual nearly im-
punctate areas near ocelli; postocellar area varying from
moderately shiny between minute, dense punctures imme-
diately behind ocelli, to shiny between sparse to close, mod-
erate punctures at posterior margin; gena shiny between sparse
to close punctures, minute adjacent to eye and grading to
fine over most of area. Interantennal distance 2.30 (2.00-
2.56) times antennal socket diameter; antennocular distance
0.85 (0.63-0.88) times antennal socket diameter; scape stout,
1.90 (1.7 6—2.03) times longer than wide, scape length 0.63
(0.63-0.72) times length of first flagellar segment; first fla-

gellar segment longer than following three segments com-
bined, 5.63 (5.24-6.13) times length of second segment. In-
terocellar distance 2. 1 5 ( 1 .89-2. 1 1 ) times diameter of anterior
ocellus; ocellocular distance 0.70 (0.57-0.78) times diameter
of anterior ocellus; ocelloccipital distance 2.85 (2.50-2.96)
times diameter of anterior ocellus.

Thorax. Mesoscutum shiny between dense, fine punctures;
dorsal face of scutellum shiny, punctures sparse and minute
in center, becoming close and line laterad, dense and fine on
posterior face; metanotum slightly shiny, sharply tessellate
between scattered, minute punctures; mesepisternum shiny
between dense, fine punctures; metespisternum similar but
punctures more crowded toward posterior margin and lower
one-third with punctures minute and scattered. Basal area
shiny between sparse, fine punctures in middle, becoming
dense laterad; side of propodeum similar but punctures scat-
tered to close. Procoxa aspinose; metafemur robust, about
1.8 times longer than thick, ventral process stout; anterior
tooth of metatibia acute, moderately stout; metabasitarsus
about 3.5 times longer than wide, posterior carina ending at
about midlength in slender tooth.

Abdomen. Dorsal face of first tergum moderately shiny and
tessellate between sparse, minute punctures; disc of second
tergum shiny between sparse, minute punctures which extend
across apical zone nearly to margin; disc of third tergum
similar, but punctures a little larger and impunctate margin
broader; fourth tergum shiny and weakly tessellate between
sparse, moderate punctures; fifth and sixth terga similar, but
a little more distinctly tessellate.

Terminalia. Seventh sternite (Fig. 52) with distal process
longer and more narrowly rounded than in C. eurypatana
(Fig. 48) and fewer long marginal hairs. Apical swelling of
eighth sternite (Fig. 51) shorter than in C. eurypatana (Fig.
49), apex less acute. Dorsal process of gonocoxite (Fig. 55)
narrower and more acute than in C. eurypatana ; tubercles
at apex of gonostylus visible in dorsal view.

Pilosity. Pubescence generally ochreous, whitish on head,
especially on gena; brownish across vertex, anteriorly on me-
sobasitarsus, externally on metatibia and metabasitarsus;
reddish brown on remainder of hind legs; discs of second to
fourth terga with hairs simple, blackish and appressed on
second, longer and more erect on third, some suberect on
fourth; fifth with erect hairs, some plumose, mixed ferrugi-
nous and black; sixth and seventh terga with hairs mostly
plumose, pale ferruginous, brownish laterad.

Color. Blackish; first three terga, and most of fourth, dark
reddish brown; distal margin of fourth, and all of following
terga, ferruginous; sternites reddish; legs medium to light
reddish brown. The following dull yellow: most of mandible;
labrum; narrow median stripe and transverse distal band
(broader laterad) on clypeus; paraocular area, constricted
above and ending on eye margin at about midlength of an-
tennal socket; linear mark on underside of scape; small basal
spot on pro- and mesotibiae. Wings light brown, a little dark-
er apicad; veins and stigma dark brown.

FEMALE. Measurements (mm). Head width 5.79-6.10;
head length 4.21-4.36; wing length 1 1.0-13.0; total length
13.0-17.0.

Contributions in Science, Number 347

Snelling: American Centridini 43

Head. 1.36-1.42 times broader than long; in frontal view
occiput nearly flat and ocelli well anterior to margin; eyes
weakly convergent above, upper frontal width 0.88-0.9 1 times
lower frontal width. Mandible tridentate, inner tooth large,
subtruncate. Labrum about 1 .6 times broader than long, apex
broadly rounded; shiny between subcontiguous, moderate
punctures, but with distinct, slightly convex basal zone vir-
tually free of punctures. Clypeus about 1.7 times broader
than long, otherwise about as described for male, but im-
punctate median line moderately shiny and moderate-sized
punctures clearly dominant. Frons, occipital area, and gena
about as described for male. Interantennal distance 2.43-
2.77 times antennal socket diameter, antennocular distance
1 .47-1 .68 times antennal socket diameter; scape robust, scape
1.83-2.06 times longer than broad; scape length 0.70-0.76
times length of first flagellar segment; first flagellar segment
longer than following three segments combined, 5.00-5.41
times length of second segment. Interocellar distance 2.06-
2.24 times diameter of anterior ocellus; ocellocular distance
1.39-1.48 times diameter of anterior ocellus; ocelloccipital
distance 2.50-2.83 times diameter of anterior ocellus.

Thorax. Punctation as described for male. Basitibial plate
(Fig. 37) long, apex subacute; posterior margin of secondary
plate extending beyond that of primary plate.

Abdomen. First tergum similar to that of male, but with a
few moderate punctures across base of dorsal face; second
to fifth terga about as described for male. Pygidial plate
V-shaped, apex narrowly rounded, secondary plate indis-
tinct.

Pilosity. Generally as described for male, but only first
three terga dark reddish brown, last three ferruginous; yel-
lowish marks of face as in male, but scape entirely dark, or
with a small, obscure, yellowish blotch.

TYPE MATERIAL (all Oaxaca, MEXICO)

Holotype male, allotype, 233, 792 paratypes; Tehuantepec.
18 Feb. 1954 (R.R. Dreisbach), in University of Kansas,
Snow Entomological Museum. Additional paratypes: 12, 20
mi. W Tehuantepec, 18 Feb. 1954 (R.R. Dreisbach; UKAN);
13, 5 mi. W Tehuantepec, 7 Apr. 1953 (E.I. Schlinger; UCB);
833, 35 mi. N Tehuantepec, 2600 ft. elev., 5 Feb. 1966 (D.
Bolinger; ORSU); 13, 20 mi. E Juchitan jet., 500 ft. elev., 28
Jan. 1965 (D. Bolinger; ORSU). Two male and two female
paratypes in LACM, remainder returned to their respective
collections.

ETYMOLOGY

This species is named for the Aztec (Nahuatl) god of Spring
and of flowers, Xochipillt in pronoucing the name, the x has
an “sh” sound.

DISCUSSION

Aside from the variations in measurements and ratios noted
above, the females of this species are all very similar to one
another. Even the yellowish face marks appear to be quite
stable, but this may be due to the fact that all are from the

same locality. Much the same is true of the males. The prin-
cipal variation involves the extent of the shiny area of the
clypeus. In a few males, up to one-half of the discal area is
shiny, but generally the shiny portion is less extensive, and
may be limited to a very narrow band along the apical mar-
gin.

Most males possess a narrow, transverse supraclypeal mark,
but in one it is merely a small median spot. Other males,
including the holotype, lack a supraclypeal mark. All males
seen possess a broad stripe on the underside of the scape
which almost attains the apex of the segment. The black
laterobasal marks on the clypeus are consistently large.

The combination of pale ochreous pubescence and bicol-
ored abdomen will separate both sexes of C. xochipHlii from
all other species. Males of C. eurypatana, C. heithausi, and
C. xochipillii all have the basal margin of the clypeus more
strongly arched upward in the center than at either side; in
C.fuscata the margin is evenly, very slightly convex between
the subantennal sutures; C. dentata is intermediate between
these two types.

From C. eurypatana, males of C. xochipillii may be sep-
arated by the lack of procoxal spines, the broader clypeus
and the reduced yellow areas on the clypeus. In C. xochipillii
the pubescence is slightly yellowish, the disc of the second
tergum is only moderately shiny, and the punctures of the
apical zone of the second tergum are not conspicuously finer
than those of the disc; these features will separate C. xo-
chipillii from C. heithausi. Males of C. dentata have the
punctures of the clypeal disc distinct, rather than obscured
by dense tessellation, the disc of the second tergite is densely
punctate, and the ventral tooth of the metafemur slender and
curved. All of the species differ from C. xochipillii in having
the abdomen wholly ferruginous.

The female shares with those of C. dentata, C. heithausi,
and C. eurypatana the broad secondary plate on the basitibial
plate, thus differing from that of C. fuscata. It is separable
from all by the bicolored abdomen, from C. heithausi by the
ochreous rather than whitish pubescence, from C. dentata
by the immaculate scape and shiny, sparsely punctate disc
of the second tergite, and from C. eurypatana by the shiny
median clypeal line, immaculate scape, and much paler wings.

Subgenus Hemisiella Moure

Hemisiella Moure, 1945b:407-408. Type-species: “ Hemi-
siella lanipes (Fabricius, 1775)” = Apis lanipes Fabricius,
1775; original designation.

Centris subg. Hemisiella: Michener, 1951:7-8.

This is a large group, with most of the species occurring in
South America, but with one entering the southwestern United
States. The distinctions between this subgenus and Hetero-
centris are not great and possibly the two should be merged.
A detailed study of the extensive South American fauna might
resolve this question, but is beyond the scope of this paper.

In addition to the characters noted in the keys by Michener
(1951) and Snelling (1974), Hemisiella females may be sep-
arated from those of Heterocentris by the lack of compressed.

44 Contributions in Science, Number 347

Snelling: American Centridini

blade-like setae on the fourth and fifth sterna and by the
acute apex of the secondary pygidial plate.

KEY TO NORTH AMERICAN HEM I SI ELLA

la. Male, antenna 1 3-segmented; abdomen with seven vis-
ible terga 2

b. Female, antenna 1 2-segmented; abdomen with six vis-
ible terga 6

2a. Hairs of thoracic dorsum with conspicuous black tips

3

b. Hairs of thoracic dorsum without conspicuous black

tips 4

3a. Large species, head width over 5.2 mm; face broad,

frontal width greater than clypeocellar distance

vittata Lepeletier

b. Smaller species, head width less than 4.7 mm; face
narrower, frontal width less than clypeocellar distance

'• dichrootricha Moure

4a. Midline of clypeus not cariniform; paraocular area with
yellow mark; segments 10 and 11 of flagellum dark
beneath, contrasting with underside of segments 2-9

5

b. Midline of clypeus low-cariniform; paraocular area
without yellow mark; segments 10 and 1 1 of flagellum
beneath not contrasting with color of segments 2-9 . .

transversa Perez

5a. At least two terga, often entire abdomen, ferruginous,
apical hairs ferruginous; first flagellar segment less than

3.7 times second trigonoides Lepeletier

b. Abdominal terga blackish, apical segments with white
hairs; first flagellar segment at least 4.0 times second

nitida F. Smith

6a. Hairs of thoracic dorsum conspicuously black-tipped

7

b. Hairs of thoracic dorsum not black-tipped 8

7a. Clypeus broadly and deeply depressed across lower part
of disc; labrum at least twice wider than long; larger

species, head width at least 6.0 mm

vittata Lepeletier

b. Clypeus protuberant; disc flat or slightly convex; la-
brum no more than 1.5 times wider than long; smaller

species, head width less than 5.5 mm

dichrootricha Moure

8a. Abdomen and legs blackish; scopa black 9

b. Abdomen and middle and hind legs largely ferruginous;

scopa ferruginous trigonoides Lepeletier

9a. Fifth tergum usually with long, glistening, white hairs;
lower margin of clypeal marks parallel to apical margin
of clypeus; smaller species, head width 3. 9-4. 5 mm

transversa Perez

b. Fifth tergum with brown to black hairs only; lower
margin of clypeal marks oblique to apical margin of

clypeus; larger species, head width 4. 8-5. 3 mm

nitida F. Smith

Centris (Hemisiella) dichrootricha (Moure)

Hemisiella dichrootricha Moure, 1945b:408-409. 9.

Centris (Hemisiella) dichrootricha: Michener, 1954:143. 9 <3
(distr., tax.).

This species was described from the State of Guapore, Brazil;
Michener (1954) recorded it from Panama and noted char-
acteristics of the previously undescribed male. I have seen
many specimens from Panama and a few from Costa Rica.
The species is easily recognized by the characteristics cited
in the key.

Centris ( Hemisiella ) nitida F. Smith

Centris nitida F. Smith, 1874:368. 9.

Centris confinis Perez, 1905:40. 9. NEW SYNONYMY.

Perez (1905) described C. confinis from “Mexique?” The
type specimen, a female, is in the Museum National d'His-
toire Naturelle, Paris, and has been examined by me. Al-
though in poor condition, it is unquestionably a specimen of
C. nitida. The records from southern Arizona cited by Hurd
( 1 979) as C. confinis are based on specimens of C. transversa.

This is a common species through Central America, ex-
tending into northern South America; the type locality is
Honduras, without more precise locality. I have seen Central
American material of C. nitida from Mexico, Belize, Gua-
temala, Honduras, El Salvador, and Costa Rica. Although I
have seen no records from Panama, C. nitida must be present
there, as it is present in South America (Colombia, Ecuador).

Centris ( Hemisiella ) transversa Perez

Centris transversa Perez, 1905:39. 9 6.

Hemisiella transversa: Moure, 1945b:408.

Centris (Melanocentris) ruae Cockerell, 1949:474-475. 9.
NEW SYNONYMY.

Centris (Hemiesiella) transversa: Snell mg, 1966:26-27 (distr.).
Centris (Hemisiella) confinis: Hurd, 1979:2175 (misidenti-
fication).

This primarily Mexican species is found also in Guatemala
and Honduras. Although it has been suggested that this is a
synonym of C. nitida (see Lutz and Cockerell, 1920:560), the
two are distinct from one another. In addition to differences
noted in the key, females of C. transversa have the median
impunctate line of the clypeus distinctly raised, the flagellum
is uniformly dark, the dorsal thoracic hairs are brownish and
the hairs at the sides of the fourth and fifth sternites are
whitish. In C. nitida, the impunctate median line is not raised
or, if a little elevated, the raised area is broadly rounded
rather than narrow and sharply defined; the underside of the
flagellum is lighter in color than the upper side; the dorsal
thoracic hairs are yellowish; the hairs at the sides of the fourth
and fifth sternites are uniformly dark.

Males of C. transversa possess a somewhat coniform ven-
tral process on the metatrochanter, a prominent ventral pro-
cess near the base of the metafemur, and the pubescence of
the fifth and sixth sternites is mostly pale. The ventral process
of the metatrochanter of C. nitida is a depressed, narrow
apical spine and, in that species, the ventral surface of the
metafemur lacks a process and the pubescence of the fourth
to sixth sterna is dark.

Contributions in Science, Number 347

Snelling: American Centridini 45

The type of C. ( Melanocentris ) ruae is in the USNM (No.
58880) and is from Zamorano, Honduras. I have examined
the specimen and it is a normal specimen of C. transversa,
differing from Mexican specimens only in the reduction in
the number of white hairs on the apical terga. In spite of
Cockerell’s statement to the contrary, the size is normal and
the clypeal and labral markings are well within the range for
this species. The type agrees with other material from Hon-
duras.

Hurd (1979) reported C. (Hemisiella) confinis from Ari-
zona (Patagonia and Tumacacori) on flowers of Parkinsonia
aculeata. The specimens on which this report was based are
in UCB and I have examined them. They are C. transversa,
which I had previously (1966) recorded from Arizona. At
that time I conjectured that C. transversa was possibly ad-
ventive in Arizona, a view I no longer hold. Additional ma-
terial now shows the distribution of C. transversa to extend
north along the western flanks of the Sierra Madre Occidental
through Sonora to southern Arizona.

Centris ( Hemisiella ) trigonoides Lepeletier

Centris trigonoides Lepeletier, 1841:167. <5.

Centris dentipes F. Smith, 1874:366. <5. NEW SYNONYMY.
Centris hoplopoda Moure, 1943:160. <5.

Centris rufomaculata Cockerell, 1949:476. <5. NEW SYN-
ONYMY.

Centris ( Rhodocentris ) lanipes subtarsata Cockerell, 1949:
476-477. 6 2. NEW SYNONYMY.

Centris ( Hemisiella ) hoplopoda: Michener, 1954:142-143
(distr., tax.).

Centris (Hemisiella) trigonoides subtarsata: Snelling, 1966:
25-26 (distr., tax.).

Centris trigonoides is a common species, ranging from Mex-
ico to Argentina. Throughout this range it is subject to much
variation and some localized phenotypes have been named;
presumably there are additional synonyms to be recognized
among the many names applied to South American forms
of Hemisiella. The entire complex of forms in this difficult
group will have to be examined.

In an earlier paper (Snelling, 1966), I attempted to justify
recognition of C. lanipes subtarsata as a Central American
subspecies of C. trigonoides. Subsequent study of several
hundred additional specimens convinced me of the futility
of that effort.

The type male of C. dentipes (BMNH 17B.919) has been
examined and agrees with the current concept of C. trigo-
noides. The type male of C. rufomaculata (USNM 58883) is
merely an individual with abdomen mostly dark brownish.

Females of C. trigonoides consistently have the entire ab-
domen ferruginous in Central American samples, and cannot
be confused with any other species in our area. The abdomen
in males varies from wholly ferruginous to dark reddish brown
on the basal three or four tergites. The metatrochanter has
a prominent, thick, ventral spine, the ventral ridge of the

metafemur is high and cariniform, and the ventral pubes-
cence of the abdomen is yellowish to reddish.

Centris ( Hemisiella ) vittata Lepeletier

Centris vittata Lepeletier, 1841:168. <3 2.

Centris montezuma Cresson, 1879:213. 2 3.

Centris breviceps Friese, 1899:44. $ 2.

Centris Friesei Crawford, 1906:158. 2. Preoccupied.

Centris Costaricensis Crawford, 1907:21. New name for C.

friesei Crawford, 1906, not C. friesei Ducke, 1902.
Centris costaricensis var. erubescens Friese, 1 925:30. 2. NEW
SYNONYMY.

Both sexes of C. vittata are easily recognized by the large size
(length over 20 mm), black-tipped thoracic hairs, and the
transversely depressed clypeal disc. The posteroventral mar-
gin of the male metafemur is sharply angled, but is not a
cariniform ridge and the ventral spine of the metatrochanter
is reduced to an inconspicuous, obtuse tubercle.

This is a widespread species through South America, and
in Central America. 1 have seen specimens from Mexico,
Costa Rica, Panama, and Honduras. Friese's C. costaricensis
var. erubescens, described from Costa Rica, is a minor vari-
ant, not worthy of recognition.

Subgenus Heterocentris Cockerell

Gundlachia Cresson, 1865:195. Type-species: Centris ? cor-
nuta Cresson, 1865; type by monotypy. Preoccupied.
Heterocentris Cockerell, 1899:14. Type-species: Centris ?
cornuta Cresson, 1865; autobasic. New name for Gund-
lachia Cresson, 1865, not Gundlachia Pfeiffer, 1850 (Mol-
lusca), not Gundlachia Herrich-Schaeffer, 1866 (Insecta,
Lepidoptera).

Centris subg. Rhodocentris Friese, 1900b: 244. Type-species:
C. dijformis F. Smith, 1854; designated by Sandhouse,
1943.

Since Cresson’s generic name Gundlachia was preoccupied,
Cockerell (1899) proposed Heterocentris as a replacement
name; the type-species for Heterocentris automatically is C.
? cornuta which Cockerell (1906) later considered to be the
same as C. dijformis. There is no evidence that he actually
saw the type of C. cornuta, now in the Gundlach collection
of the Academia de Ciencias in Havana, Cuba. Indeed, there
is reason to suppose that he did not, for the original descrip-
tions of C. cornuta and C. dijformis do not agree in several
important peculiarities of head structure. Thus, Cresson de-
scribed the mandible of C. cornuta as “very long, narrow
and cleft at tip, shining black . . .” as opposed to Smith’s
“. . . mandibles large, very broad at their base, and armed
above with a stout tooth, their apex bidentate, having a lon-
gitudinal pale testaceus stripe” (italics mine). Of the clypeus,
Cresson stated: “clypeus short, very transverse, emarginate
on each side, with a large, very prominent, incurved, subacute
tooth on the middle, pale yellowish white, the anterior and
posterior margin and the tooth except its lateral base, black”
(italics mine); the labrum is said to possess a “long slender.

46 Contributions in Science, Number 347

Snelling: American Centridini

porrect, subacute spine.” According to Smith, the clypeus of
C. dijformis is “short, transverse, elevated, its anterior por-
tion vertical.” There is no mention of processes of any sort
on either clypeus or labrum. I think it as unlikely that Cresson
would have overlooked the very prominent mandibular pro-
cess as that Smith would have failed to mention such a con-
spicuous anomaly (in this genus) as a spinose clypeus. It is
my opinion that C. cornuta and C. difformis are very different
entities and that C. cornuta must be properly considered to
be the type-species of Heterocentris.

This possibly has unfortunate ramifications since the iden-
tity of C. cornuta is problematic. However, since the only
forms of Centris with unusual modifications of mandible,
labrum, and clypeus all belong to the accepted interpretation
of Heterocentris, the question is probably moot.

Just as the type-species of Heterocentris should be clearly
restricted to C. cornuta, so, too, should the type-species of
Rhodocentris be restricted to C. dijformis. Rhodocentris was
described as a new subgenus of Centris, not as a replacement
name for Gundlachia. It is clear that Sandhouse considered
C. dijformis to be the proper name for the species which she
selected as type. It is unclear why she chose to cite the type-
species as “( Centris cornuta Cresson, 1 865) = Centris diffor-
mis F. Smith, 1854,” unless it was to assure that Rhodocentris
was an automatic junior synonym of Heterocentris through
isogenotypy. Both specific names were available, since both
were originally included in Rhodocentris by Friese. Since it
is clear that Sandhouse was of the opinion that C. difformis
was the correct name for the taxon chosen as type-species I
think it best to consider that name to be the type; this would
eliminate the ambiguity of having two names involved as
possible type-species.

Heterocentris, together with Hemisiella and Trachina, is
part of a complex recognized by possessing three-segmented
maxillary palps in both sexes and the male with a carina
along the posterior margin of the metabasitarsus; this carina
usually terminates in a prominent tooth-like process. Males
of Heterocentris differ from those of both Trachina and
Hemisiella in the form of the dorsal face of the first abdom-
inal tergite: at the extreme side, the dorsal face is extended
caudad (most strongly so in C. labrosa ) and there is a con-
spicuous patch of erect, plumose, dark setae at the side of
the segment, their apices abruptly bent and llattened. This
is a feature unique in the genus. Additionally, the middle
mandibular tooth is smaller and nearer to the inner tooth
than to the apical tooth.

In females of Heterocentris the upper inner mandibular
carina is elevated near the base and the labrum is large, with
the disc depressed and the apical margin more or less flange-
like and with a pair of dentiform submedian processes. In
both sexes the apicolateral angle of the clypeus is contiguous
with the eyes, or nearly so. Females are additionally char-
acterized by the distinct, abruptly truncate secondary pygidial
plate and by the presence of large, flattened, spiniform setae
near the apical margins of the fourth and fifth abdominal
sterna.

The few Central American Heterocentris may be separated

by the following key. Since no males of C. difformis are
known to me, I am unable to include this sex in the key.

KEY TO CENTRAL AMERICAN HETEROCENTRIS

la. Male, antenna 13-segmented and basitibial plate absent

2

b. Female, antenna 12-segmented and basitibial plate pres-
ent 5

2a. Postgradular area of second tergum about four times as

long as pregradular area 3

b. Postgradular area of second tergum shorter than pre-
gradular area labrosa Friese

3a. Mandible tridentate 4

b. Mandible bidentate bicornuta Mocsary

4a. Clypeal disc smooth and shiny between punctures; apical
margin of labrum transverse or very weakly concave;
lower corner of pronotum with a few long, simple, red-
dish or yellowish setae among plumose hairs

analis (Fabricius)

b. Clypeal disc roughened and slightly shiny between punc-
tures; apical margin of labrum with distinct median
emargination; lower corner of pronotum with plumose

hairs only C. (Heterocentris) species

5a. Clypeus without lateral cornuti; labrum broader than

long; mandible tridentate 5

b. Clypeus with pair of long, slender cornuti; labrum longer
than broad; mandible bidentate bicornuta Mocsary
6a. Clypeus no more than 1 .8 times wider than long in mid-
dle, with disc flattened; mandible without long subbasal

process on outer face 7

b. Clypeus very short, more than twice wider than long,
transversely elevated; mandible with large, tooth-like
subbasal process on outer face .... difformis F. Smith
7a. Lower, lateral angle of pronotum with a cluster of long,
yellowish or reddish, simple setae as well as plumose
hairs; juncture of anterior and lateral faces of mesepi-
sternum without carina, rounded; hairs of thoracic dor-
sum rarely dark-tipped analis (Fabricius)

b. Lower, lateral angle of pronotum with long, plumose
hairs only; juncture of anterior and lateral faces of mes-
episternum with a short, lamelliform carina; hairs of
thoracic dorsum always dark-tipped . . . labrosa Friese

Centris ( Heterocentris ) analis (Fabricius)

Anthophora analis Fabricius, 1804:375. 2.

Centris totonaca Cresson, 1879:213. 2.

Centris otomita Cresson, 1879:214. <3.

Centris minuta Mocsary, 1899:254. <3.

Centris labrosa var. simplex Friese, 1 899:44. 2.

Centris (Melanocentris) durantae Cockerell, 1949:474. <3.
Centris (Melanocentris) petreae Cockerell, 1949:475. <3.
Centris (Melanocentris) petreae var. rufopicta Cockerell, 1 949:
475. <3.

Centris (Heterocentris) totonaca: Michener. 1954:140 (syn.).
Centris (Heterocentris) analis: Moure, 1 960b: 1 32-133 (syn.,
notes on type).

Contributions in Science, Number 347

Snelling: American Centridini 47

Moure (1960b) gives very complete literature citations of
this common species which ranges from Mexico to Brazil.
The presence of long, simple, ferruginous setae on the lower
corner of the pronotum is diagnostic for the female. The male
has similar setae, but they are less numerous than in the
female and are sometimes difficult to see among the more
numerous plumose hairs.

Centris (Heterocentris) bicornuta Mocsary

Centris bicornuta Mocsary, 1899:254. 9.

Heterocentris bicornuta: Moure, 1945b:502.

Centris (Heterocentris) bicornuta: Michener, 1951:6, 7.

Although widely distributed, C. bicornuta does not appear
to be a common species, though males are sometimes locally
abundant. The bicornute clypeus of the female is diagnostic
for that sex. Males may be easily separated from those of C.
analis and C. labrosa by the bidentate mandible. Since I have
seen no males of C. difformis, which presumably has biden-
tate mandibles, I am uncertain how to distinguish that species
from C. bicornuta. Presumably, however, the males of C.
difformis will be more than 1 5 mm long and the hairs of the
thoracic dorsum will be black-tipped. Males of C. bicornuta
are not more than about 10 mm long and the hairs of the
thoracic dorsum are not black-tipped.

Specimens of C. bicornuta have been seen from Mexico,
Guatemala, Costa Rica, and Panama, as well as from South
America (Brazil and Guyana). The species was described
from Brazil.

Centris ( Heterocentris ) difformis F. Smith

Centris difformis F. Smith, 1854:374. 9.

Centris difformis: Crawford, 1906:158. 9.

Heterocentris difformis: Moure, 1945b: 402, 403.

Centris ( Heterocentris ) difformis: Michener, 1951: 6, 7.

As discussed above, 1 do not believe that C. cornuta, de-
scribed from Cuba, is a synonym of C. difformis. The latter
species was originally described from Brazil and seems to be
rare in collections. According to Cresson ( 1879) C. difformis
occurs in Mexico, but I have seen no specimens from there.
Crawford (1906) recorded a female from Pozo Azul, Costa
Rica, as C. difformis. I have examined the specimen and
agree with Crawford's identification.

No males have been seen. Presumably the mandibles are
bidentate, as in C. bicornuta (Moure, 1945b. noted that C.
minuta and C. labrosa differed from his characterization of
Heterocentris mandibles as bidentate with the statement that
the mandible is falsely tridentate in these two species). Based
on the females, males of C. difformis should be conspicuously
larger than those of C. bicornuta and the hairs of the thoracic
dorsum should have blackish apices.

NEW RECORDS

PANAMA, CANAL ZONE: 399, Barro Colorado Island. 27
Apr. 1980 (K.E. Steiner; UCD), on Byrsonima crassifolia.

Centris ( Heterocentris ) labrosa Friese

Centris labrosa Friese, 1899:44. 9 (not 3).

Centris tarsata: Schwarz, 1934:13. Misidentification.
Heterocentris labrosa: Moure, 1945b:402.

Centris (Rhodocentris) triangulifera Cockerell, 1949: 477. 9.
NEW SYNONYMY.

Centris (Heterocentris) labrosa: Michener, 1954:104 (var.,
distr.).

This is a moderately common species which ranges from
Mexico to Brazil. I have seen specimens from throughout
Central America. It should be noted that the specimen re-
corded by Schwarz (1934) from Barro Colorado Island, Pan-
ama, as C. tarsata F. Smith, is actually C. labrosa. The type
of C. triangulifera. in the USNM (No. 58885), has been
examined and is a normal specimen of C. labrosa.

The unusually long pregradular area of the males is im-
mediately diagnostic for this sex of C. labrosa. Females are
likewise immediately recognizable by the presence of a dis-
tinct carina separating the anterior and lateral faces on the
lower half of the mesepisternum; in all other species, the
juncture of the two surfaces is rounded.

Centris ( Heterocentris ) species

A few males from Panama (Canal Zone and Panama Prov-
ince) cannot be assigned to any of the above species. It seems
unlikely that they are males of C. difformis for they are much
smaller (less than 15 mm long) than the one female of C.
difformis (about 20 mm long) I have seen; the two sexes tend
to be approximately equal in size in this genus.

They are very similar to males of C. analis. but lack the
simple setae on the lower lateral angle of the pronotum, the
clypeus is distinctly dull and roughened between the punc-
tures, and the mandible and scape are immaculate or with
traces of yellowish markings. Possibly these represent an un-
described species, but the available material is too limited
for any decision as to their identity.

Genus Ptilotopus Klug

Ptilotopus Klug, 1810:31. Type-species: Ptilotopus ameri-
canus Klug, 1810; monobasic.

Centris subg. Ptilotopus: Michener, 1951:10. Snelling, 1974:
2, 3.

In recent years Ptilotopus has usually been treated as a sub-
genus of Centris characterized by the bilobate scutellum with
defined bare areas, the prominent hypoepimeral tubercle, the
lack of a secondary basitibial plate in the female and the male
genitalia without giant branched setae (Michener, 1951;
Snelling. 1974). The largest and most spectacular centridines
are included in Ptilotopus.

Although in general habitus the species of Ptilotopus are
similar to those of Centris and have Centris- like wing ve-
nation, I am now of the opinion that Ptilotopus should be
removed from Centris. There are a number of features which
are not shared with any of the groups presently assigned to
Centris. Both sexes of Ptilotopus possess long, black flagel-
liform setae on the occipital margin; these setae project well
beyond the occipital hair fringe, although they are not as
spectacularly developed as in many Epicharis.

48 Contributions in Science, Number 347

Snelling: American Centridini

Contributions in Science, Number 347

Sneiling: American Centridini 49

The metatibia of Ptilotopus females has a well-defined ba-
sitibial plate. There is, however, no secondary plate; instead,
the disc of the basitibial plate is moderately depressed, the
depression filled with a dense mat of short, fine, subappressed
hairs. In Centris and Epicharis females, a secondary plate is
present and usually well defined. When it is not distinct the
plate has a discal convexity and is glabrous.

Females of Ptilotopus have a distinct tubercle on either
side of the midline of the mesosternum, anterior to the me-
socoxa. As a rule the tubercles are hidden in an exceptionally
dense tuft of short, stiffhairs. Although mesosternal tubercles
are known for some Epicharis males, they are not known to
be present in the females of either Centris or Epicharis.

A particularly unusual feature is to be seen in the structure
of the basal areas of the second and third abdominal tergites
of the females. In both Centris and Epicharis the gradulus
of each segment marks off a very narrow basal area across
most of the breadth of the segment; laterad. it is either eva-
nescent or deflected distad. In Ptilotopus females the gradulus
of the second and third tergites is strong and, in the middle,
is directed distad as a more or less triangular incursion onto
the disc of the segment. The area on either side of this tri-
angular extension is depressed and filled with a compact mat
of very short, erect, plumose hairs. This unusual feature is
not known to occur in Centris and in Epicharis is known
only in the subgenus Epicharitides.

Thus, although Ptilotopus is Centris-Yxkt in many features,
particularly in general body form and wing venation, there
are features, such as the presence of the flagelliform occipital
setae, presence of mesosternal tubercles in the female and
the modified structure of the female second and third ab-
dominal terga, which will readily separate Ptilotopus from
Centris. The short marginal cell of the forewing will readily
separate Ptilotopus from Epicharis, as will the mesosternal
tubercles and lack of a secondary basitibial plate in the fe-
male. Additionally, in Ptilotopus, the hypoepimeron has a
prominent tubercle and the scutellum has a conspicuous,
flattened, bare lobe on either side of the middle. The genitalic
structures of Ptilotopus are very distinctly Centra-like, with-
out the unusual modifications seen in Epicharis.

Ptilotopus zouata (Mocsary)

Figures 59-62

Centris zonata Mocsary, 1899:251. 9.

Centris pandora Friese, 1900b:241, 269 (new name for C.
zonata Mocsary, not C. zonata F. Smith, 1 854, now placed
in Epicharis).

This species, the only known North American Ptilotopus,
was described from Chiriqui Province, Panama; no addi-
tional material has been recorded since the original descrip-
tion.

The females have the pubescence of the head, thorax, and
legs bright ferruginous. Except for a broad band of short,
erect yellow pubescence across the second tergite, the ab-

domen is black pubescent. The male pubescence is all black,
except for a yellow band across the second tergite.

NEW RECORDS

PANAMA, PANAMA: 1499, Arraijan, 16 Oct. 1980(D. Rou-
bik; LACM, ROUB). VER.4GUAS: 13, El Maria, Coibu Is-
land, 22 Oct. 1979 (D. Roubik; LACM). CANAL ZONE: 12,
Gatun, 3 Nov. 1977 (K.E. Steiner; UCD), on Stigmaphyllon
hypargyreum.

Genus Epicharis Klug

Epicharis is an exclusively Neotropical genus of medium- to
large-sized bees, often quite colorful, allied to Centris. Friese
( 1 900b) monographed these bees, as a subgenus of Centris,
but most subsequent authors have treated Epicharis as a
separate genus. Since Friese’s monograph most work in this
genus has consisted ofdescriptions of new species and variant
forms. Moure ( 1945a), however, divided Epicharis into nine
genera, for which he proposed the subtribe Epicharitina with-
in within the Centridini. Michener ( 1954) recognized Moure’s
genera as subgenera within the single genus Epicharis and
subsequent authors have been in accord with this interpre-
tation.

Epicharis was described by Klug ( 1 807), but a type-species
was not designated until Latreille (1810) fixed Apis rustica
Olivier, 1789, as the type-species; A. rustica was not one of
the originally included species and, hence, is not available.
Lutz and Cockerell (1920) selected Centris umbraculata Fa-
bricius, 1804, one of the originally included species, as the
type-species. Moure ( 1 945a, 1 960b) held that the designation
made by Latreille is valid since Apis hirtipes Fabricius, 1 793,
one of the originally included names, is a synonym of A.
rustica.

Sandhouse (1943) accepted the designation of Lutz and
Cockerell, as did Michener (1954) when he named Epicha-
rana to replace Epicharis (s.s.) of Moure ( 1945a) and placed
Xanthepicharis Moure in synonymy with true Epicharis (s.s.).
It is my understanding of the ICZN Code that even though
A. hirtipes ultimately proved to be a synonym of A. rustica,
this does not validate the designation made by Latreille, since
Latreille was not aware that the two names applied to the
same species. There is, additionally, the fact that A. hirtipes
can never be anything other than a subjective synonym of A.
rustica ; the synonym is a generally accepted opinion which
has the potential for being incorrect. Stability would be best
served by accepting the security of the designation of Lutz
and Cockerell.

Of the nine subgenera recognized by Moure (1945a), five
are known to be present in North America; Hoplepicharis
Moure, 1945a; Epicharana Michener, 1945 (=“ Epicharis”
of Moure, 1945a); Epicharoides Radoszkowski, 1884; Pa-
repicharis Moure, 1945a; and Epicharitides Moure, 1945a.
The remaining four subgenera, including Epicharis proper
(=Xanthepicharis Moure, 1 945a), are limited to South Amer-
ica: Anepicharis, Triepicharis, and Cyphepicharis, all Moure,
1945a.

50 Contributions in Science, Number 347

Snelling: American Centridini

The taxonomy of Epicharis is difficult. These bees are
generally black or blackish and both sexes usually have con-
spicuous patterns of white or yellow and/or ferruginous
markings on various parts of the body. The species within a
subgenus tend to be morphologically very similar and dif-
ferences often are subtle. Previous descriptive work has tend-
ed to emphasize differences in maculations. This, coupled
with a scarcity of specimens, has resulted in a confused sit-
uation. I expect a reduction in the number of species of
Epicharis as more material becomes available and the species
better known. I have proposed some new synonymy below
and pointed out cases where I suspect further synonymy will
be in order. Two new species are described, based on mor-
phological characteristics.

The following key to the subgenera of Epicharis is modified
from that of Moure (1945a).

KEY TO SUBGENERA OF EPICHARIS

la. Female with secondary basitibial plate; male with py-
gidial plate obsolete, its margins more or less coextensive
with margins of seventh tergite, apex thin and bilobate

or bidentate 2

b. Female without secondary basitibial plate; male with
distinct, sharply marginate pygidial plate, no more than
one-half as wide as seventh tergite, its apex narrowly

truncate or rounded 4

2a. Lateral margins of female pygidial plate distinctly con-
cave in dorsal view, apex broadly truncate (Fig. 63);
female metatibia no longer than metabasitarsus; male
without scopa-like hairs on hind legs and metabasitarsus
with longitudinal keel on anterior margin which termi-
nates in spiniform process at apex, or without keel. . 3
b. Lateral margins of female pygidial plate nearly straight,
apex narrowly truncate (Fig. 64); female metatibia longer
than metabasitarsus; hind legs of male with long, scopa-
like hairs and metabasitarsus with prominent tooth at

basal one-third Hoplepicharis

3a. Female: frontal carina ending more than diameter of
anterior ocellus in front of that ocellus; pygidial trun-
cation narrower than diameter of anterior ocellus; disc
of fifth tergite with very short hairs mostly simple or
barbulate. Male: mesosternal tubercles absent; meta-
basitarsus without carinate ridge on anterior margin;
metatrochanter and metafemur without ventral patch of

short, dark plumose setae Epicharis, s.s.

b. Female: distance between frontal carina and anterior
ocellus less than diameter of anterior ocellus; pygidial
truncation broader than diameter of anterior ocellus;
disc of fifth tergite with very short hairs distinctly plu-
mose. Male: mesosternal tubercles present; metabasi-
tarsus with carinate ridge on anterior margin, terminat-
ing in tooth-like process (Fig. 65); metatrochanter and
metafemur with ventral patch of short, dark, plumose

setae Epicharana

4a. First flagellar segment short, in female usually no longer
than combined second and third but always shorter than
following three combined, and in male shorter than scape;

posterior margin of dorsal face of scutell um not, or weak-
ly, impressed in middle (if deeply impressed, maxillary

palp three-segmented) 5

b. First flagellar segment of female equal to length of next
three segments combined, of male longer than scape and
longer than following two segments combined; posterior
margin of dorsal face of scutellum deeply impressed in
middle; maxillary palp two-segmented . Parepicharis
5a. Lateral ridges of clypeal disc absent or weakly evident
only on basal one-half or less; male metabasitarsus with
carinate ridge on anterior margin, terminating in large

tooth at about midlength 6

b. Lateral ridges of clypeal disc strong and sharply defined
to near apical margin; male metabasitarsus without car-
inate ridge or tooth on anterior margin 7

6a. Maxillary palp two-segmented; fiagelliform occipital se-
tae distinct and extending beyond anterior margin of
mesoscutum; ocellocular distance of male less than di-
ameter of lateral ocellus in dorsal view . . Anepicharis
b. Maxillary palp three-segmented; fiagelliform occipital
setae short, not projecting beyond occipital hairs and not
reaching anterior margin of mesoscutum; ocellocular
distance of male equal to diameter of lateral ocellus in

dorsal view Triepicharis

7a. Dorsal face of scutellum flat; prepygidial fimbria of fe-
male preceded by a shiny, nearly glabrous area ... 8
b. Dorsal face of scutellum bigibbous; female without shiny,

glabrous area anterior to prepygidial fimbria

Cyphepicharis

8a. Jugal lobe of posterior wing about half as long as vannal
lobe and nearly attaining apex of cubital cell; female
metabasitarsus about twice longer than broad, posterior
margin nearly straight; male pygidial plate broad, cov-
ering most of seventh tergum Epicharoides

b. Jugal lobe of posterior wing about one-third as long as
vannal lobe, its apex well short of that of cubital cell;
female metabasitarsus about one-third longer than broad,
posterior margin strongly curved; male pygidial plate
narrow, covering less than one-half of seventh tergum
Epicharitides

Subgenus Epicharana Michener

Epicharis: Moure, 1 945a:294-295. Type-species: " Epicharis
rustica Olivier, 1 789” = Apis rustica Olivier, 1789. Not
Epicharis Klug, 1 807.

Epicharis subg. Epicharana Michener, 1954:144. Type-
species: Apis rustica Olivier, 1789; original designation.

DESCRIPTION

Maxillary palp three-segmented, third segment a little short-
er, and much narrower, than second; lateral ridge of clypeal
disc strong; malar area about as long as minimum thickness
of first flagellar segment; occipital margin rounded; occipital
fiagelliform setae long, extending beyond anterior margin of
tegula; metanotum bifaced, dorsal face about as long as pos-
terior face, juncture angular or crested; jugal lobe of posterior
wing shorter than cubital cell and less than half as long as
vannal lobe.

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Snelling: American Centridini 51

Female. Labrum with low, median longitudinal ridge; out-
er face of mesobasitarsus, on anterior one-third with mixed
long, coarse, simple setae and shorter, fine, plumose hairs,
posterior two-thirds with sparse longer, coarse, simple setae
and sparse, short, fine, plumose hairs; basitibial plate with
secondary plate; metatibia no longer than metabasitarsus;
third and fourth terga without basal specialized areas; in
dorsal view, margins of pygidial plate concave, apex broadly
truncate.

Male. Labrum without median ridge; first flagellar segment
shorter than scape, longer than second segment, much shorter
then second and third combined; ocellocular distance greater
than ocellar diameter; procoxa with flattened distal process;
mesosternum with prominent process on each side of mid-
line, anterior to mesocoxa; metatrochanter and base of meta-
femur with ventral mat of short blackish setae; metatibia
with carinate posteroventral ridge; metabasitarsus carinate
along anterior margin, ending in prominent tooth-like pro-
cess at apex of segment; pygidial plate broad, its margins
confluent with those of segment, apex bidentate.

This is the group that Moure (1945a) incorrectly inter-
preted as Epicharis s.s., as discussed above. Five species
occur in Central America, with a few more in South America.

KEY TO CENTRAL AMERICAN EPICHARANA

la. Abdominal terga blackish and immaculate (except male
of one species with dorsal fascia on first and lateral spot
on second) or ferruginous and immaculate; male clypeus

black 2

b. Abdominal terga reddish brown to ferruginous and dor-
sal face of first segment with narrowly interrupted trans-
verse yellow fascia (sometimes obscure); male clypeus

yellow elegans F. Smith

2a. Dorsal segments of abdomen black, first tergite with or
without transverse yellow fascia on dorsum; male meta-
tibia moderately convex in middle of anterior margin
(Fig. 66); disc of female clypeus, in basal one-third or
more, with more or less distinct, weakly depressed, me-
dian impunctate line 3

b. Dorsal segments of abdomen bright ferruginous and im-
maculate; male metatibia strongly, obtusely convex in
middle of anterior margin (Fig. 65); disc of female clyp-
eus uniformly, subcontiguously punctate in basal one-

third bova, new species

3a. Female: scutellum shinier than mesoscutum, punctures
of disc both less distinct and more separated than those
of mesoscutum; larger punctures of parapsis sparse, sep-
arated by more than twice a puncture diameter. Male:
first tergum without dorsal transverse yellow fascia, me-
sosternal tubercles nearly prostrate, their inner margins

rounded (Fig. 67) rustica (Olivier)

b. Female: scutellum no shinier than mesoscutum, micro-
punctures of disc as sharp and dense as those of meso-
scutum; larger punctures of parapsis dense, separated by
about a puncture diameter or less. Male: first tergite with
narrowly interrupted yellow fascia on dorsal face; me-

sostemal tubercles erect, obtuse, inner margin sharply
carinate (Fig. 68) angulosa, new species

Epicharis ( Epicharana ) angulosa, new species

Figures 66, 68, 69-72

DIAGNOSIS

Separable from other species of Epicharana by the following
unique combination of features: pubescence, except of hind
legs, black; abdomen black, except dorsally on first tergum
and laterally on second tergum of male. Male: mesosternal
tubercles angulate along inner margin, high, long and obtuse
in profile; metatibia moderately swollen (Fig. 66), posterior
ventral carina low, abruptly reduced beyond midlength. Fe-
male: shiny area at base of clypeal disc no greater in area
than triangular supraclypeal area and basal one-half of disc
densely and coarsely punctate, without definite impunctate
median line.

DESCRIPTION

HOLOTYPE MALE. Measurements (mm). Head width
5.95 (5.64-5.90); head length 4.00 (3.90-4.00); wing length
17.0 (15.0-16.5); total length 21.0 (20.0-22.0).

Head. 1 .48 ( 1 .45-1.49) times broader than long; in frontal
view, occipital margin nearly straight (except ocellar eleva-
tion) and slightly below level of tops of eyes; inner eye mar-
gins moderately convergent above, upper frontal width 0.77
(0.76-0.79) times lower frontal width. Mandible slender be-
yond middle, inner tooth large, blunt. Labrum quadrate,
slightly broader than long, apical margin transverse, disc shiny
and weakly tessellate between sparse to close punctures vary-
ing from fine to coarse. Clypeus about 1 .7 times broader than
long, apicolateral angle removed from eye by slightly less
than diameter of antennal socket; disc moderately shiny be-
tween dense, coarse punctures except near apical margin where
punctures are sparse and fine and integument is more or less
“wrinkled,” median impunctate line absent; discal carinae
moderately convergent above, distance between them at low-
er end about 1.7 times that at upper end; side slightly shiny
and distinctly roughened between variably sparse to subcon-
tiguous, fine to coarse punctures. Frons moderately shiny
and distinctly tessellate between dense, mixed fine and mod-
erate punctures; preocellar area slightly bulging on either side,
smooth, shiny, and impunctate; ocellocular area dull and
densely tessellate, subcontiguously micropunctate and with
scattered minute punctures; preocciput moderately shiny be-
tween dense, fine punctures; gena moderately shiny to shiny
between close to dense punctures, minute near eye, becoming
fine near margin. Interantennal distance 1.86 (1.88-2.05)
times antennal socket diameter; antennocular distance 0.59
(0.63-0.71) times antennal socket diameter; scape robust,
1.50 (1.50-1.64) times longer than broad, scape length 1.57
(1.53-1.61) times length of first flagellar segment; first fla-
gellar segment 1.31 (1 .22-1 .32) times longer than broad, much
shorter than following three segments combined, 1.15(1 .08-
1.13) times longer than second. Interocellar distance 1.70
(1.44-1.64) times diameter of anterior ocellus; ocellocular

52 Contributions in Science, Number 347

Snelling: American Centridini

Figures 63-64, pygidial plate of female: 63, Epicharis ( Epicharana ) rustica\ 64, E. ( Hoplepicharis ) lunulata. Figs. 65-66, metatibia of male:
65, E. ( Epicharana ) bova\ 66, E. (Epicharana) angulosa. Figs. 67-68, profile of right mesosternal process of male: 67, E. ( Epicharana ) rustica\
68, E. (Epicharana) angulosa.

distance 1 .96 ( 1 .89-1 .93) times diameter of anterior ocellus;
ocelloccipital distance 1.70 (1.50-1.67) times diameter of
anterior ocellus.

Thorax. Mesoscutum slightly shiny, uniformly densely,
finely punctate and with sparse moderate punctures which
become scattered distad; dorsal face of scutellum broadly
depressed in middle, punctation as scutum, but fine punc-
tures scattered; metanotum moderately shiny in median area,
dull elsewhere, roughened and tessellate, with minute obscure
punctures. Mesepisternum moderately shiny and tessellate
between sparse, shallow, fine punctures; metepisternum
moderately shiny between subcontiguous, minute punctures

(appearing finely reticulopunctate at certain angles). Meso-
sternal process, in profile, suberect and obtusely triangular
(Fig. 68); in ventral view, inner margin carinate. Propodeum
moderately shiny, minutely reticulopunctate and with sparse
to scattered fine punctures. Procoxal process broadly ellip-
tical, with acute apex; mesofemur stout, twice longer than
thick, greatest thickness a little basad of midlength; antero-
basal ventral depression of metafemur subcarinate along its
posterior margin; anterior edge of metatibia evenly curved
(Fig. 66), posterior carina low throughout, abruptly reduced
distad of middle.

Abdomen. Tergal discs moderately shiny between minute

Contributions in Science, Number 347

Snelling: American Centridini 53

sSSwrosj

Figures 69-72. Epicharis ( Epicharana ) angulosa, male seventh and eight stermtes and genitalia (ventral and dorsal views). Scale line = 1.00
m m .

54 Contributions in Science, Number 347

Snelling: American Centridini

punctures which are dense on basal segments, becoming pro-
gressively coarser and less close on succeeding segments; discs
also with scattered fine punctures on basal segments, becom-
ing progressively coarser (but still fine) on succeeding seg-
ments. Seventh tergite weakly raised in middle (broader ba-
sad) to form a weakly differentiated median plate; apex
bidentate, teeth long, slender, acute, emargination longer than
a semicircle.

Terminalia. Apical margin of distal lobe of seventh sternite
(Fig. 69) broadly, shallowly concave; setae long, some con-
spicuously plumose. Shoulders of disc of eighth sternite (Fig.
70) angular and sides of apical lobe of disc strongly conver-
gent. Genitalia as in Figs. 71 and 72.

Pilosity. Generally blackish brown; sides of pronotal collar,
pronotal lobe, mesoscutum (especially anteriorly and at side)
with admixed very pale brown hairs; hairs on side of tergites
and on abdominal venter mixed medium and light brown.
Flairs mostly long, erect, plumose and dense, but labrum
nearly bare. Metatibia and metabasitarsus with long yellow-
ish hairs. Abdominal terga, beyond first, with sparse discal
hairs which are short, simple and suberect on second seg-
ment, becoming progressively longer and more abundant
caudad; terga also with abundant minute, appressed, scale-
like hairs (arising from minute punctures) which become
progressively longer on succeeding segments.

Color. Blackish brown. The following pale yellow: oblique,
linear mark near base of mandible; labrum; oblique, linear
mark from clypeal margin, at tentorial pit, to malar area;
triangular supraclypeal mark; underside of scape; spot on
procoxal process; anterodistal spot on pro- and mesofemora;
large distal spot on metafemur; stripe on anterior margin of
protibia nearly to apex; broad stripe on anterior margin of
mesotibia nearly to apex, constricted in middle; anterior and
outer face of metabasitarsus; large, anterior spots on dorsal
face of first tergum, narrowly separated in middle; small lat-
erobasal spot on second tergum. Underside of flagellum light
brown; apical tarsal segments reddish. Wings uniformly dark
brown, veins and stigma blackish.

FEMALE. Measurements (mm). Head width 6.31; head
length 4.15; wing length 16.5; total length 23.0.

Head. 1.52 times broader than long; in frontal view, oc-
cipital margin straight, except ocellar elevation; inner eye
margins moderately convergent above, upper frontal width
0.84 times lower frontal width. Mandible stout, tridentate,
inner teeth obtuse and margin between them broadly con-
cave. Labrum about 1 . 1 times longer than broad, apex sub-
angularly rounded in middle; disc moderately shiny, rough-
ened and tessellate between dense to subcontiguous, mixed
fine and coarse punctures, median line slightly raised. Clyp-
eus about 1.6 times broader than long, apicolateral angle
separated from eye by 0.5 times diameter of antennal socket;
discal carinae moderately convergent basad, intercarinal dis-
tance at base about 0.66 times that at their distal end; sculp-
ture as described for male, but with shiny basal area which
is no greater than supraclypeal area. Punctation of frons,
preoccipital area, and gena as in male. Interantennal distance
2.20 times antennal socket diameter; antennocular distance
0.88 times antennal socket diameter; scape robust, 1 .9 1 times

longer than wide, scape length 1.05 times length of first fla-
gellar segment; first flagellar segment shorter than following
three combined, 3.81 times longer than second. Interocellar
distance 1 .44 times diameter of anterior ocellus; ocellocular
distance 1.78 times diameter of anterior ocellus; ocelloccip-
ital distance 1.56 times diameter of anterior ocellus.

Thorax. Punctation as in male. Metanotum with horizon-
tal basal face separated from vertical posterior face by a
cariniform ridge on either side of middle. Apex of basitibial
plate acute; secondary plate about twice longer than wide.

Abdomen. Punctation as described for male. Disc of py-
gidial plate not visible.

Pilosity. About as described for male, but pronotal collar
and lobe without pale hairs; bristles of thoracic venter with
pale tips; scale-like hairs of fifth tergum becoming longer,
more erect and plumose toward prepygidial fringe; hairs of
prepygidial fringe reddish brown and pale tips; scopa! hairs
pale yellowish.

Color. Generally blackish brown; mandible (except golden
apical blotch), underside of flagellum, apical segments, red-
dish brown; paraocular area with small, obscure yellowish
blotch near clypeal margin, below level of tentorial pit (larger
and more distinct on left side). Wings as in male.

TYPE MATERIAL

Holotype male, allotype, and one male paratype: Monte-
verde, 1500 m elev., Puntarenas Prov., COSTA RICA, 12
Sept. 1978 (G.W. Frankie), in LACM. Paratypes: 13, same
locality, 19 Aug. 1974 (D. Janzen; UKAN); 233, 4 km E San
Ignacio de Acosta, 4000 ft. elev., San Jose Prov., COSTA
RICA, 8 July 1963 (C.D. Michener et al.; UKAN).

ETYMOLOGY

The specific epithet is a Latin word meaning with corners,
referring to the angulate mesosternal processes of the male.

DISCUSSION

Aside from the variations noted above in the measurements
and proportions, the males available are very similar to one
another. The laterobasal spot on the second tergite may ex-
tend mesad as a narrow line and/or a short distance distad.
The procoxal process is immaculate in three paratypes and
in these specimens the mesotibial stripe is divided into a
basal spot and two narrowly separated stripes along the seg-
ment. One male has a narrow yellow stripe on the anterior
face of the metadistitarsus and yellow blotches on the outer
face of the metamediotarsi.

Males of this species are easily recognized by the combi-
nation of black abdomen with contrasting pale marks, sub-
erect and internally carinate mesosternal tubercles and black
labrum. Females are considerably less distinctive, but differ
from such species as E. elegans, E. rustica, and E. bova in
possessing an immaculate abdomen and largely black tho-
racic pubescence. The lack of a median impunctate line on

Contributions in Science, Number 347

Snelling: American Centridini 55

Figures 73-76. Epicharis (Epicharana) bova, male seventh and eighth sternites and genitalia (ventral and dorsal views). Scale line = 1.00 mm.

56 Contributions in Science, Number 347

Snelling: American Centridini

the clypeal disc will also separate E. angulosa from E. elegans
and E. rustica.

Epicharis ( Epicharana ) bova, new species

Figures 65, 73-76

DIAGNOSIS

Separable from all other species of Epicharana by the fol-
lowing unique combination of characters. Abdomen ferru-
ginous, immaculate; pubescence of head and thorax medium
brown to blackish brown, mesoscutum without pale hairs;
clypeus black. Male: mesostemal processes, in profile, sub-
erect but low and rounded, not carinate on inner margin;
anterior margin of metatibia strongly produced (Fig. 65).
Female: clypeal disc subcontiguously to densely punctate,
without median impunctate line.

DESCRIPTION

1 IOLOTYPE MALE. Measurements (mm). Head width
5.95; head length 4.15; wing length 16.0; total length 25.0.

Head. 1.43 times broader than long; in frontal view, oc-
cipital margin very weakly concave, except for ocellar ele-
vation; ocelli on occipital margin; upper frontal width 0.71
times lower frontal width. Mandible slender, inner tooth small,
obtuse. Labrum about 1.2 times broader than long, apical
margin broadly rounded; disc moderately shiny, weakly tes-
sellate between irregularly spaced (mostly sparse), fine and
coarse punctures. Clypeus about 1 . 5 times broader than long,
apicolateral angle separated from inner eye margin by about
0.8 times diameter of antennal socket; clypeus shiny and very
weakly tessellate between subcontiguous to dense, coarse
punctures which become moderate distad, median impunc-
tate line absent; discal carinae about twice as far apart distad
as at base; side shiny between dense, moderate to coarse,
elongate (especially distad) punctures. Frons moderately shiny
and distinctly tessellate between dense, mixed fine and mod-
erate punctures; preocellar area slightly protuberant on either
side, shiny and nearly impunctate; ocellocular area dull and
densely tessellate, subcontiguously, shallowly micropunctate
and with scattered minute punctures; preocciput moderately
shiny to shiny between close to dense punctures, minute near
eye, becoming fine near margin. Interantennal distance 2.00
times antennal socket diameter; antennocular distance 0.63
times antennal socket diameter; scape robust, 1.74 times
longer than wide, scape length 1.52 times length of first fla-
gellar segment; first flagellar segment shorter than following
three segments combined, 1.37 times longer than second
segment. Interocellar distance 1.61 times diameter of anterior
ocellus; ocellocular distance 1.96 times diameter of anterior
ocellus; ocelloccipital distance 1.61 times diameter of ante-
rior ocellus.

Thorax. Mesoscutum slightly shiny, uniformly densely,
finely punctate, and with sparse, moderate punctures which
become scattered distad; dorsal face of scuteilum broadly
depressed in middle, punctures as on mesoscutum, but fine
punctures scattered; dorsal face of metanotum moderately
shiny and distinctly tessellate between dense, fine punctures.

posterior face dull, densely tessellate and with scattered fine
punctures. Mesepisternum moderately shiny and tessellate
between sparse, fine punctures; metepisternum moderately
shiny between subcontiguous, minute punctures. Mesoster-
nal process suberect in profile, its apex narrowly rounded; in
ventral view, inner margin rounded. Propodeum moderately
shiny, subcontiguously micropunctate between sparse to
scattered, fine punctures. Procoxal process subcircular, apex
not produced; mesofemur stout, about twice longer than thick,
greatest thickness a little basad of midlength; anterobasal
ventral depression of metafemur subcarinate along its pos-
terior margin; anterior margin of metatibia abruptly convex
at about midlength and anterior carina plainly visible (Fig.
65), posterior carina low throughout, highest distad of middle
and abruptly reduced beyond highest point.

Abdomen. Tergal discs moderately shiny between minute
punctures which are dense on basal segments, becoming pro-
gressively coarser and less close on succeeding segments; discs
also with scattered fine punctures on basal segments, becom-
ing progressively coarser (but still fine) on succeeding seg-
ments. Seventh tergite with very weakly differentiated py-
gidial plate which is depressed along midline; apex bidentate,
teeth stout and acute, emargination between them semicir-
cular.

Terminalia. Apical margin of distal lobe of seventh sternite
(Fig. 73) deeply, angularly incised; setae long, widely spaced,
mostly simple. Shoulders of disc of eighth sternite (Fig. 74)
obtuse and sides of apical lobe of disc weakly convergent
distad. Genitalia as in Figs. 75 and 76.

Pilosity. Generally blackish brown on head, thorax, and
legs, but yellowish on metatibia and metabasitarsus; suberect
hairs and appressed scaliform hairs of terga two to six golden
brown, but fringes of fifth and sixth segments brown and a
few brown, simple, suberect hairs scattered on discs. Scale-
like hairs very short basad, becoming longer and more erect
on succeeding segments. Hairs of abdominal sterna golden
brown to blackish brown.

Color. Head, thorax, and legs blackish brown; abdominal
terga ferruginous, sterna light brown. The following yellow-
ish: large triangular mark near base of mandible; labrum;
narrow, oblique stripe on side of face, from tentorial pit to
malar area; small, triangular supraclypeal spot; small spot on
underside of scape; basal spot on protibia; dorsal, apical spot
on metafemur; outer face of metatibia and metabasitarsus
(metatibia black at base). Inner leg surfaces dark ferruginous;
tarsi ferruginous. Wings blackish brown, veins and stigma
black.

FEMALE. Measurements (mm). Head width 6.41; head
length 4.26-4.31; wing length 16.5; total length 24.0-27.0.

Head. 1.49-1.51 times broader than long; in frontal view,
occipital margin nearly straight, ocellar elevation anterior to
margin; upper frontal width 0.79-0.81 times lower frontal
width. Mandible stout, tridentate, inner teeth obtuse, margin
between them broadly concave. Labrum about 1.1 times
longer than broad, apex subangularly rounded in middle; disc
moderately shiny, roughened, and tessellate between dense
to subcontiguous, mixed fine and coarse punctures, median
line slightly raised. Clypeus about 1.4 times broader than

Contributions in Science, Number 347

Snelling: American Centridini 57

long, apicolateral angle separated from inner eye margin by
about 0.75 times antennal socket diameter; intercarinal dis-
tance at base about 0.36 times that at their distal end; sculp-
ture as described for male. Remaining cephalic sculpture as
described for male. Interantennal distance 2.28-2.37 times
antennal socket diameter; antennocular distance 0.92-0.95
times antennal socket diameter; scape robust, 1 .86-1.9 1 times
longer than broad, scape length 1.04-1.08 times length of
first flagellar segment; first flagellar segment shorter than fol-
lowing three segments combined, 3.45-3.85 times length of
second segment.

Thorax. Punctation as in male. Dorsal and posterior faces
of metanotum separated on either side of middle by short,
convex, cariniform ridge. Basitibial plate subacute at apex,
secondary plate about 2.8 times longer than wide.

Abdomen. Punctation as in male. Pygidial plate broadly
truncate at apex, secondary plate very weak.

Pilosity. About as described for male, with following dif-
ferences: discs of third, fourth, and fifth terga with some
suberect to erect dark brown, bristle-like hairs, longer and
more abundant on succeeding segments; prepygidial fringe
golden brown. Scopa of metatibia and metabasitarsus yel-
lowish.

Color. As described for male, but wholly without yellowish
markings and pronotum and propodeum reddish brown.

TYPE MATERIAL

Holotype male: Cerro Campana, Panama Prov., PANAMA,
4 May 1960 (W.J. Hanson), in Snow Entomological Museum,
University of Kansas. Allotype: 4 mi. S San Vito de Java,
Puntarenas Prov., COSTA RICA, 15 Aug. 1967 (R.W.
McDiarmid; LACM). Paratypes: IS, N of El Valle de Anton,
Code Prov., PANAMA, 12-13 Sept. 1964 (R.L. Dressier;
UKAN); IS, O.T.S. Field Station, Finca La Selva, Heredia
Prov., COSTA RICA, mid Aug. 1980 (J.M. MacDougal;
LACM), on Passiflora lobata, 0900.

ETYMOLOGY

The specific epithet is a Latin word for a swelling of the legs
and alludes to the swollen metatibia of the male.

DISCUSSION

Both sexes are easily separated from others in the subgenus
Epicharana by the features listed above in the Diagnosis.
The male is especially distinctive in the shape of the meta-
tibia, the anterior margin of which is conspicuously and
abruptly convex at about midlength. The low posterior carina
of the metatibia is like that of E. angulosa, as is the poste-
riorly subcarinate anterobasal depression on the underside
of the metafemur. The abdomen of E. angulosa is black and
the mesosternal tubercles are acute at their apices and sharply
carinate on their inner margins.

The only other species with a reddish abdomen is E. ele-
gans, in both sexes of which the abdomen is distinctly yel-
lowish red and the first tergum is yellowish maculate on the
disc. In males of E. elegans, and those of E. flava and E.

rustica, the clypeus is pale, the metatibia is regularly convex
along its anterior margin, there is no inner, anterior meta-
tibial carina, and the inner posterior metatibial carina is ex-
panded beyond midlength and folded mesad. In both sexes
of these species, E. elegans, E. Jlava, and E. rustica, there is
considerable pale pubescence intermixed with blackish on
the thoracic dorsum. Females of these three species have a
definite median impunctate line on the clypeus and the mi-
nute punctures of the scutellum are usually coarser and less
distinct than those of the mesoscutum.

Epicharis ( Epicharana ) elegans F. Smith

Epicharis elegans F. Smith, 1 86 1 : 1 52. 9 <5.

Epicharis elegans: Moure, 1945a:296 (tax.).

Epicharis salazari Cockerell, 1949:480-481. 9. NEW SYN-
ONYMY.

This is a common black and red species in Mexico and ranges
south to Costa Rica. Specimens from southern Mexico (Chia-
pas and Yucatan), Guatemala, El Salvador, and Costa Rica
have the abdomen more brownish and correspond to E.
salazari, described from El Salvador. These are minor vari-
ants and not worthy of separation from E. elegans.

NEW RECORDS

MEXICO, CHIAPAS: 499, La Revancha, 20 Aug. 1972 (T.W.
Taylor; LACM). GUERRERO: 699, 12.7 mi. N Iguala, 5200
ft. elev., 1 Aug. 1969 (Univ. Kans. Mex. Exped.; UKAN);
19, 23 mi. N Taxco, 1 700 ft. elev., 8 Aug. 1 954 (Univ. Kans.
Mex. Exped.; UKAN); 19, 3.7 mi. W Rio Balsas, 5 Aug. 1 965
(G.W. Byers and party; UKAN); 2 33, Iguala, 2400 ft. elev.,
8 Aug. 1954 (Univ. Kans. Mex. Exped.; UKAN). JALISCO:

18, Guadalajara, no date (Crawford; LACM); 399, Cocula,
4450 ft. elev., 27 Sept. 1957 (H.A. Scullen; ORSU); 299,
Puente Grande, 5000 ft. elev., 20 Aug. 1954 (Univ. Kans.
Mex. Exped.; UKAN). MORELOS: 599, 888, Lake Tequis-
quetengo, 5000 ft. elev., 13 Sept. 1957 (H.A. Scullen; ORSU);
13, Rancho Tetela, Cuernavaca, 24 June 1974 (K.E. Donahue
and S. Adams; LACM); 19, 13, Cuernavaca, no date (Craw-
ford; LACM); 233, Alpuyeca, 27 June and 3 July 1951 (P.D.
Hurd; UCB); 19, 7.3 mi. S Yautepec, 3000 ft. elev., 16 Aug.
1962 (Ordway and Naumann; UKAN); 399, 4 mi. SW Yau-
tepec, 3800 ft. elev., 2 July 1961 (C.D. Michener; UKAN),
on Cassia sp.; 299, 4.3 mi. W Yautepec, 4000 ft. elev., 17
Aug. 1962 (Ordway and Marston; UKAN); 13, 7 mi. NE
Yautepec, 4000 ft. elev., 18 Aug. 1962 (Univ. Kans. Mex.
Exped.; UKAN); 13, 14 mi. S Yautepec, 16 Aug. 1962 (Mar-
ston and Roberts; UKAN). OAXACA: 19, 25 mi. SE Oaxaca,
5600 ft. elev., 27 June 1963 (Scullen and Bolinger; ORSU);

19, 13, 47 mi. SE Oaxaca, 13 July 1952 (E.E. Gilbert and
C.D. MacNeil; UCB); 2 99, 5 mi. NW Totolapan, 4000 ft.
elev., 29 July 1970 (E.M. Fisher and P. Sullivan; LACM);
4799, 5 mi. NW Totolapan, 3800 ft. elev., 6 July 1 953 (Univ.
Kans. Mex. Exped.; UKAN), on Malpighia mexicana ; 19,
Salina Cruz, no date (F.K. Knab; USNM); 299, Mixtla, 5600
ft. elev., 22 Aug. 1963 (Scullen and Bolonger; ORSU); 433,
Monte Alban, 6000 ft. elev., 27 June 196 1 (Univ. Kans. Mex.

58 Contributions in Science, Number 347

Snelling: American Centridini

Exped.; UKAN); 1 <3, same, except 15 July 1955; 19, Tama-
zulapan, 6000 ft. elev., 28 June 1961 (Univ. Kans. Mex.
Exped.; UKAN); 19, 2 mi. NW Tamazulapan, 6000 ft. elev.,
28 June 1961 (Univ. Kans. Mex. Exped.; UKAN). PUEBLA:
999, 22 km NW Izucar de Matamoros, 1158m elev., 2 1 Sept.
1976 (C.D. George and R.R. Snelling; LACM), on Cassia
laevigata ; 1699, 16.1 km NW Izucar de Metamoros, 1280 m
elev., 1 7 Sept. 1976 (C.D. George and R.R. Snelling; LACM),
on Caesalpinia cacalaco\ 999, 788, 6.9 km S Izucar de Mat-
amoros, 1 250 m elev., 1 7 Sept. 1 976 (C.D. George and R.R.
Snelling; LACM), on Solatium rostratum (99) and Martynia
annua (88); 399, 388, Atlixco, 9 July 1970 (R.E. Beer and
party; L1KAN); 299, 3 mi. NW Petlalcingo, 4600 ft. elev., 29
Aug. and 5 Sept. 1 972 (Byers and Thornhill; UKAN); 19, 12
mi. NW Tehuitzingo, 4050 ft. elev., 29 June 1961 (Univ.
Kans. Mex. Exped.; UKAN); on Cassia sp.; 19, 10 mi. SE
Tehuitzingo, 3900 ft. elev., 3 July 1953 (Univ. Kans. Mex.
Exped.; UKAN). SAN LUIS POTOSI: 688, El Salto, 1600
ft. elev., 24 Aug. 1954 (Univ. Kans. Mex. Exped.; UKAN).
TAMAULIPAS: 19, 38 mi. N El Mante, 1050 ft. elev., 1 1
Oct. 1957 (H.A. Scullen; ORSU). YUCATAN: 599, Piste, 24
June 1967 (E.C. Welling; LACM). GUATEMALA: 299, “1923
F.4696” (UKAN); 19, “env. de Guatemala” (R. Guerin;
MNHN). EL SALVADOR: 19, Dept. Santa Tecla, Feb. 1947
(M. Salazar; USNM; type of E. salazari)] 499, Santa Tecla,
900 m elev., 25 Apr. -6 May 1972 (S. and L. Steinhaus;
DPIF). COSTA RICA, GUANACASTE: 19, Comelco, 8 km
NW Bagaces, 3 1 March 1971 (P.S. Opler; UCB), on Tabebuia
rosea\ 19, same, except 25 Nov. 1972; 18, same, except 5
Mar. 1971; 18, La Pacifica, 4 km NW Canas, 14 Mar. 1972
(P.A. Opler; UCB), on Inga vera\ 18, Hacienda Comelco, 24
km NW Canas, 21 Mar. 1971 (E.R. Heithaus; LACM), on
Stachytarpheta jamaicense, 0815; 18, same locality, 13 Mar.
1971 (E.R. Heithaus; LACM), on Centrosoma pubescens,
0750.

Epicharis ( Epicharana ) rustic a (Olivier)

Figures 63, 67

Apis rustica Olivier, 1789:64.

Apis hirtipes Fabricius, 1793:325. 9.

Epicharis rustica: F. Smith, 1854:368.

Centris ( Epicharis ) rustica: Friese, 1900b:253. 9 8.

Centris (Epicharis) rustica var. /lava Friese, 1900b:254. 8.
NEW SYNONYMY.

Epicharis rustica: Moure, 1945a:295-296. 8 9 (syn.).
Epicharis (Epicharana) rustica: Michener, 1954:144 (distr.).
Epicharis ( Epicharis ) rustica: Moure, 1 960b: 1 19-120 (syn.).
Epicharis (Epicharis) flava: Moure, 1960b: 120 (status).

Moure (1960b) reexamined the type female of Apis hirtipes
and reaffirmed its traditional place in the synonymy of E.
rustica. He concluded “. . . that with this discovery, the true
meaning of Epicharis is to be restored as in my revision of
Epicharis (1945) and Epicharana Michener, 1954, with the
same type species is to be considered a synonym.” I have
already alluded to be unavailability of Apis rustica to be the
type-species of Epicharis.

In point of fact, Moure’s determination is inconclusive.
Moure synonymized A. hirtipes under “. . . Epicharis rustica
as interpreted by older authors and Friese in his Monographic
der Bienengattung Centris (s. lat.).” This is by no means the
same as placing A. hirtipes in synonymy with Olivier’s species.
Moure admitted that he had been unable to find Olivier’s
type. In truth the identity of E. rustica is unknown and the
current interpretation of this species rests upon the very in-
secure foundation of assumptions made over a century ago
by workers whose concepts of species were different from
those of the present and who may not actually have seen the
relevant specimens.

For the present, it would seem best to continue to accept
the traditional interpretation of E. rustica and its appended
synonymy, as published by Moure (1960b). The alternative
would be to regard Olivier’s species as unidentifiable and to
use the next available name (A. hirtipes ), suffering the re-
sultant nomenclatural confusion. I adhere to the traditional
understanding, which includes the unavailability of E. rustica
as the type species for this genus.

This is a common bee, ranging from Costa Rica and Pan-
ama to Brazil and Peru. Friese’s var. flava, with whitish
cinereous pubescence on the thorax, was thought by Moure
( 1 960b) to be probably a good species. In general, specimens
from northeastern South America have almost entirely black
thoracic pubescence and would correspond to the “typical”
form of E. rustica. Similar dark specimens are also found in
Costa Rica, but most populations have considerable pale
pubescence on the dorsum and sides of the thorax. There
are, however, many variants, the most common being a form
with dark mesepisternal hairs, those of the mesoscutum and
scutellum pale, or largely so. These specimens of an inter-
mediate character occupy geographical areas between the two
extremes, as would be expected if E. rustica and E. flava
were conspecific, which I believe to be the case.

NEW RECORDS

COSTA RICA, GUANACASTE: 19 [Hacienda] Comelco, 5
Mar. 1971 (P.A. Opler; UCB), on Tabebuia rosea. HERE-
DIA: 19, Finca La Selva, 500 m elev., 23 June 1979 (D.R.
Perry; LACM). PUNTARENAS: 19, Monteverde, 1400 m
elev., 17 Sept. 1982 (C.D. Nagano and M. Hayes; LACM).
PANAMA, CANAL ZONE: 19, Barro Colorado Island, 15
July 1980 (H. Wolda; ROUB); 19, same, except 1 Aug. 1980.
CHIRIQUI: 399; 18, Dolega, 15 Mar. 1980 (D.W. Inouye;
ROUB). PANAMA: 399, Arraijan. 16 Oct. 1980 (D. Roubik;
ROUB); 499, Chihbre Cave, 26 July 1966 (R.D. Sage; UCB);
18, San Miguelito, 16 May 1974 (M. Gonzalez; LACM).

Subgenus Par epicharis Moure

Parepicharis Moure, 1945a:307-308. Type-species: “ Pare -
picharis zonata (Smith, 1854 )” = Epicharis zonata F.
Smith, 1854; monobasic and original designation.

DESCRIPTION

Maxillary palp two-segmented, second segment much longer
than first, more than five times longer than wide, gradually

Contributions in Science, Number 347

Snelling: American Centridini 59

narrower distad; lateral ridges of clypeal disc weak; malar
space linear, eye margin nearly contiguous with mandible
base; frontal carina ending before attaining anterior ocellus;
occipital margin abruptly rounded; flagelliform occipital se-
tae reaching about to level of anterior margin of tegula; pos-
terior margin of dorsal face of scutellum impressed; meta-
notum vertical; jugal lobe of posterior wing about as long as
cubital cell and one-half as long as vannal lobe.

Female. Labrum with median ridge weak or absent; outer
face of mesobasitarsus with mixed long, coarse plumose setae
and long, fine plumose hairs on anterior half, posterior half
with long, coarse plumose setae only; basitibial plate without
secondary plate; metatibia no longer than metabasitarsus;
third and fourth terga without specialized basal areas; mar-
gins of pygidial plate, in dorsal view, nearly straight, apex
broadly truncate.

Male. Labrum without median ridge; first flagellar segment
longer than scape and longer than following two segments
combined; ocellocular distance less than ocellar diameter;
procoxa without apicoventral process; mesosternal protu-
berances absent; metatrochanter and metafemur without
ventral seta patch; metatibia without carinate posteroventral
ridge; metabasitarsus with or without anterior carina-like
ridge, when present terminating in tooth a little beyond mid-
length; pygidial plate narrower than seventh tergite, margins
cariniform, apex narrow, rounded or truncate.

Parepicharis was proposed as a monotypic genus for Epi-
charis zonata. Subsequently, Moure and Seabra (1959) added
E. metatarsalis to Parepicharis. Of the two species, only E.
metatarsalis is found in Central America. I have seen ma-
terial of E. zonata from Guyana, Brazil, and Bolivia and
there are records of the species from Peru and Trinidad.

KEY TO SPECIES OF PAREPICHARIS

la. Male, antenna 13-segmented and ocellocular distance

less than diameter of anterior ocellus 2

b. Female, antenna 1 2-segmented and ocellocular distance

greater than diameter of anterior ocellus 3

2a. Metabasitarsus about twice as long as broad and with
distinct tooth on anterior margin . . metatarsalis Friese
b. Metabasitarsus about six times longer than broad and
without tooth on anterior margin .... zonata F. Smith
3a. First abdominal segment blackish, remaining segments
dull ferruginous, immaculate; prepygidial fimbria con-
sisting of long, closely ranked, plumose hairs

metatarsalis Friese

b. Abdomen brown, with conspicuous yellow maculations
on at least first three terga; prepygidial fimbria weak,
consisting of short, discretely separated, short-plumose

hairs which do not conceal underlying surface

zonata F. Smith

Epicharis ( Parepicharis ) metatarsalis Friese

Epicharis metatarsalis Friese, 1899:40. <5.

Epicharis phenacura Cockerell, 1917:200. 6. NEW SYN-
ONYMY.

Epicharis conura Cockerell, 1917:200. 9. NEW SYNONY-
MY.

Epicharis ( Parepicharis ) metatarsalis: Moure and Seabra,
1959:126 (distr., tax.).

The male of E. metatarsalis differs from that of E. zonata
most obviously by the metabasitarsus, which is only twice
as long as broad and with a distinct tooth on the anterior
margin beyond the middle. Males of E. zonata lack a tooth
on the metabasitarsus and the segment is about six times
longer than broad. Females of E. metatarsalis have the first
tergite blackish and the remaining segments dull yellowish
red; in E. zonata there are yellow maculae, of variable extent,
on the first three tergites which usually are dark brown to
blackish. Both sexes of E. metatarsalis are larger (25-27 mm
versus 20-23 mm).

Friese (1900b) recorded males of this species from St. Pa-
rime, Venezuela (type locality), and “Darien (=Colombia),”
now in Panama. Another male was reported by Moure and
Seabra (1959) from San Jose, Costa Rica. San Carlos, Costa
Rica is the type locality for both E. phenacura and E. conura.
The material now available indicates that E. phenacura and
E. conura are the opposite sexes of one species and that the
males are inseparable from E. metatarsalis.

NEW RECORDS

COSTA RICA, HEREDIA: 2399, 633, Finca La Selva, near
Puerto Viejo, 5 June-24 July (D.R. Perry; LACM), on Hy-
menolobium sp. (899, 288), Dipteryx panamensis (1 199, 233),
Tabebuia sp. (1<S), Dussia sp. (13), and Byrsonima sp. (499).

Subgenus Hoplepicharis Moure

Hoplepicharis Moure, 1945a:300-301 . Type-species: ‘'Hop-
lepicharis fasciata (Lepeletier & Serville, 1 828)” = Epi-
charis fasciata Lepeletier and Serville, 1828; original des-
ignation.

Epicharis subg. Hoplepicharis: Michener, 1954:145.

DESCRIPTION

Maxillary palp two-segmented, second segment shorter than
first, flattened; lateral ridges of clypeal disc strong; malar
space distinct, slightly longer than minimum thickness of
first flagellar segment; frontal carina sharp but ending well
in front of anterior ocellus; occipital margin abruptly round-
ed; flagelliform occipital setae extending nearly to level of
posterior tegular margin; posterior margin of dorsal face of
scutellum not impressed; metanotum bifaced, juncture sharply
angulate to crested; jugal lobe of posterior wing about as long
as cubital cell and about one-half as long as vannal lobe.

Female. Labrum with weak median ridge; outer face of
mesobasitarsus, on anterior half, with long, coarse, simple
setae and shorter, fine, plumose hairs, posterior half with
sparse long, coarse, simple setae and scattered short, fine,
plumose hairs; secondary basitibial plate present; metatibia
a little longer than metabasitarsus; third and fourth terga
without specialized basal areas; margins of pygidial plate

60 Contributions in Science, Number 347

Snelling: American Centridini

nearly straight and strongly convergent to narrowly truncate
apex.

Male. Labrum without median ridge; first flagellar segment
shorter than either scape or second flagellar segment; ocel-
locular distance greater than diameter of anterior ocellus;
procoxa with short, inconspicuous apicoventral process;
mesosternum without tubercles; metatrochanter and meta-
femur without ventral seta patch; metatibia without pos-
teroventral ridge; metabasitarsus with anterior carinate ridge
terminating in sharp tooth basad of midlength; pygidial plate
broad and indistinct, weakly bilobate at apex.

This is a small group, with two species known from South
America and two in Central America.

KEY TO CENTRAL AMERICAN HOPLEPICHARIS

a. Second abdominal tergite with basal yellow band, nar-

rowed across middle, followed by narrow blackish brown
band; most of remainder of this segment, and all of fol-
lowing segments, ferruginous (except short, yellow lon-
gitudinal stripe at side of third segment)

lunulata Mocsary

b. Similar, but second tergite, beyond yellow basal band,

and all of following segments, blackish (except short, yel-
low longitudinal stripe at side of third segment)

monozona Mocsary

Epicharis ( Hoplepicharis ) lunulata Mocsary

Figure 64

Epicharis lunulata Mocsary, 1898:499. <5 9.

Hoplepicharis lunulata: Moure, 1945a:302 (distr., syn.).
Epicharis ( Hoplepicharis ) lunulata: Michener, 1954:145
(distr.).

This attractive Central American species appears to be less
common than the superficially similar E. e/egans. Its range
extends from Mexico to Panama.

NEW RECORDS

MEXICO, CHIAPAS: 19, Simojovel, 18-31 July 1958 (J.A.
Chemsak; UCB). JALISCO: 19, Estacion Biologia UNAM,
Chamela, 1 1 Sept. 1981 (S.H. Bullock; LACM), on Psidium
sartorianum. NAYARIT: 299, 16 mi. NW Tepic, 19 July 1953
(Univ. Kans. Mex. Exped.; UKAN). OAXACA: 19, 6 mi. W
Zanatepec, 150 ft. elev., 9 July 1953 (Univ. Kans. Mex.
Exped.; UKAN), on Malpighia mexicana\ 299, 5 mi. NW
Totolapan, 3800 ft. elev., 6 July 1953 (Univ. Kans. Mex.
Exped.; UKAN), on Malpighia mexicana. VERA CRUZ: 19,
13, Cordoba, 1-10 Sept. (9), 1 Oct. 1964 (<3) (LACM). GUA-
TEMALA: 19, Cayuga, Nov. (Schaus and Barnes; USNM);
19, “F.4694” (UKAN). HONDURAS: 13, Tegucigalpa, 12
May 1981 (F.J. Dyer; USNM). COSTA RICA, GUANA-
CASTE: 499, 13, Hacienda Comelco, 8 km NW Bagaces and
24 km NW Canas, dates between 7 Mar. and 14 Oct. (E.R.
Heithaus [19], P.A. Opler [399, 13]; LACM, UCB), on Byr-
sonima sp. (19), Stachytarphe frantzii (19), Petastoma patel-
liferum (3) and Bignonaceae, 0700 (19); 19, near Turin
(10°20'N, 84°50'W), 1 Feb. 1960 (C.W. Palmer; UKAN).

PUNTARENAS: 19, 1 mi. ESE jet. Rio Canas and Hwy 2,
1000 ft. elev., 23 July 1965 (R.D. Sage and S.J. Arnold; UCB),
on Bixa orel/ana, 1000-1 100. SAN JOSE: 13, 4 km E San
Ignacio de Acosta, 4000 ft. elev., 8 July 1 963 (C.D. Michener
et ah; UKAN); 19, Playon, 8 km N Parrita, 30 ft. elev., 14-
19 Aug. 1962 (C.D. Michener and A. Wille; UKAN); 13,
San Jose, 25 July 1913 (UKAN).

Epicharis ( Hoplepicharis ) monozona Mocsary

Epicharis monozona Mocsary, 1898:498. 9.

Hoplepicharis monozona: Moure, 1945a:392 (distr., tax.).
Epicharis ( Hoplepicharis ) monozona: Michener, 1954:145
(distr.).

This species was based on a female from an unspecified Pan-
amanian locality. Moure (1945a) recorded another female
from Muzo, Rio Cantinero, Colombia, and Michener ( 1954)
reported two additional Panamanian specimens, both from
the Canal Zone: Las Cruces trail, near Corozal, and Fort
Clayton.

The male of E. monozona has not been previously re-
ported. It is separable from that of E. lunulata by the black,
rather than ferruginous, abdomen. Aside from the differences
in color, E. monozona and E. lunulata appear to be identical
and I suspect they will ultimately be found to be conspecific.

NEW RECORDS

PANAMA, PANAMA: 399, Arraijan, 16 Oct. 1980 (D. Rou-
bik; LACM, ROUB); 19, Chepo, 15 km E Carti, 8 June (D.
Roubik; ROUB); 13, Cerro Cantpana, 13 Apr. 1960 (W.J.
Hanson; UKAN); 19, Cerro Azul, N of Tocumen, 15 June
1958 (W.J. Hanson; UKAN). CANAL ZONE: 19, Pipeline
Road, Gamboa, 12 Jan. 1980 (D. Roubik; ROUB); 19, same,
except 10 June 1980; 19, 7 33, Pipeline Road, 20 Sept., 6 Oct.
1979 (K. Steiner; UCD), on Drymonia serrulata: 599, 13,
Gamboa, 28 Sept. 1979, same collector and host; 13, Barro
Colorado Island, 15 July 1958 (W.J. Hanson; UKAN); 333,
same locality, 15 Sept., 12 Oct., 28 Oct. 1979 (K. Steiner;
UCD), on D. serrulata. COLON: 19, Portobello, 30 Sept.
1979 (K. Steiner; UCD), on D. serrulata.

Subgenus Epicharoides Radoszkowski

Epicharoides Radoszkowski, 1884:20. Type-species: (Epi-
charoides bipunctatus Radoszkowski, 1884 ) = Epicharis
maculata F. Smith, 1874; monobasic and original desig-
nation.

Epicharoides: Moure, 1945a:309-310.

Epicharis subg. Epicharoides: Michener, 1954:144.

DESCRIPTION

Maxillary palp three-segmented, second segment longer than
first, third segment narrower and shorter than second; lateral
ridges of clypeal disc strong; malar space distinct, about as
long as minimum thickness of first flagellar segment; frontal
carina sharp, nearly reaching anterior ocellus; occipital mar-
gin slightly compressed but not ridge-like; flagelliform oc-

Contributions in Science, Number 347

Snelling: American Centridini 61

Figures 77-80. Epicharis (Epicharoides) albofasciata, male seventh and eighth stemites and genitalia (ventral and dorsal views). Scale line =
1.00 mm.

cipital setae short and curved laterad, not extending beyond
anterior margin of mesoscutum; posterior margin of dorsal
face of scutellum not impressed; metanotum wholly vertical;
jugal lobe of posterior wing about as long as cubital cell and
about one-half as long as vannal lobe.

Female. Labrum without median ridge; mesobasitarsis ex-
ternally with evenly distributed long coarse setae (some plu-
mose) and short, fine, long-plumose hairs; secondary basi-
tibial plate absent; metatibia no longer than metabasitarsus;
third and fourth terga without specialized basal areas; mar-

62 Contributions in Science, Number 347

Snelling: American Centridini

gins of pygidial plate, in dorsal view, slightly concave, apex
narrowly truncate, disc depressed.

Male. Labrum without median ridge; scape unusually ro-
bust, slightly longer than wide, first flagellar segment shorter
than either scape or second flagellar segment; ocellocular
distance greater than diameter of anterior ocellus; procoxa
without apicoventral process; mesosternal tubercles absent;
metatrochanter and metafemur without ventral seta patch;
metatibia without posteroventral ridge; metabasitarsus with-
out carinate anterior ridge; pygidial plate about one-half as
broad as seventh tergite, sharply margined, apex narrowly
rounded.

Moure (1945a) recognized three species in this subgenus,
and a fourth was added by Moure and Seabra (1959). Two
of these species are present in Central America and two are
confined to South America. The two Central American species
are black, with abundant, very variable yellow markings.

KEY TO CENTRAL AMERICAN EPICHAROIDES

a. Male clypeus with irregular black blotch on disc; female

with basal yellow band of second tergite broadly inter-
rupted, but if complete, it is much shorter in middle than
black band following it and basal bands of following terga
are broadly interrupted albofasciata F. Smith

b. Male clypeus without black blotch on disc; female with
basal yellow band of second tergum as long in middle as
black band following it or, if a little shorter, third and
fourth terga with complete transverse yellow fasciae . . .

maculata F. Smith

Epicharis {Epicharoides) albofasciata F. Smith

Figures 77-80

Epicharis albofasciata F. Smith, 1874:321. <3.

Epicharis maculata var. nigroclypeata Friese, 1899:40.
Epicharoides albofasciata: Moure, 1945a:310 (syn., distr.).
Epicharis (Epicharoides) maculata: Michener, 1954:145 (in
part).

This species and E. maculata are so similar as to be virtually
inseparable and records from the literature under either name
must be treated with caution. There are, however, consistent
differences in the male terminalia (compare Figs. 77-80 with
Figs. 81-84).

Females of the two species are especially difficult to sep-
arate. Those of E. albofasciata have either lateral spots on
the second tergite or a subbasal fascia which is shorter than
the black area following the band; the third and fourth tergites
apparently never possess entire transverse subbasal bands,
though the margins of the segments may be broadly ferru-
ginous. On the other hand, females of E. maculata possess
a subbasal band on the second segment which is usually,
though not always, at least as long as the dark band following
it; a shorter transverse band, often attenuated in the middle,
is present on the third tergite; a transverse band is also present
on the fourth segment, usually longer than that of the third.

Aside from the differences in genitalic structures, males of
E. albofasciata are recognizable by the presence of a black

blotch on the the clypeal disc, the very short fasciae of the
second to fourth tergites (evanescent or absent on third and
often on fourth as well) and the pronotum black, except two
widely separated spots on the collar.

From Central America, I have seen material of E. albo-
fasciata only from Costa Rica and Panama. At least some of
the specimens from Panama recorded by Michener (1954)
as E. maculata are this species; his figures 141-143 are based
on E. albofasciata.

NEW RECORDS

COSTA RICA, GUANACASTE: 299, Hacienda Comelco, 24
km NW Canas, 6-13 Mar. 1972 (E.R. Hcithaus; LACM), 1
on Securidaca tenuifolia ; 19, Hacienda Comelco, 8 km NW
Bagaces, 5 Mar. 1971 (P.A. Opler; UCB), on Delbergia re-
teusa\ 1<3, same locality, 28-31 Jan. 1972 (P.A. Opler; UCB),
on Securidaca sylvestris\ 13, Finca La Pacifica, 4 mi. NW
Canas, 10 July 1971 (P.A. Opler; UCB), on Petastoma pa-
telliferum ; 19, Liberia, 15-16 Feb. 1972 (P.A. Opler; UCB),
on Andira inermis. HEREDIA: 599, 1633, Finca La Selva,
near Puerto Viejo, 6 May to 18 June (D.R. Perry; LACM),
on Dipteryx panamensis (399, 833), Vochysia sp. (13), Byr-
sonima sp. (19), Dussia sp. (633), and Hymenolobium sp.
(13). PANAMA, PANAMA: 19, 24 Mar. 1980 (D. Roubik;
ROUB); 499, same, except 2 Apr. 1980; 299, same except 13
Apr. 1980; 19, Curundu, 3 1 Mar. 1981 (D. Roubik; ROUB).
CANAL ZONE: 19, Barro Colorado Island, 2 June 1981 (H.
Wolda; ROUB).

Epicharis ( Epicharoides ) maculata F. Smith

Figures 8 1-84

Epicharis maculata F. Smith, 1874:320. 9.

Epicharoides bipunctatus Radoszkowski, 1884:20.

Centris (Epicharis) variabilis Friese, 1 900b: 3 5 1 . (New name
for C. maculata F. Smith, not C. maculata (Lepeletier.)
Epicharoides maculata: Moure, 1945a:310 (syn., distr.).
Epicharis (Epicharoides) maculata: Michener, 1954: 145 (syn.,
distr.) (in part).

This species was described from Oaxaca, Mexico, and ranges
south to Venezuela and Colombia. In Central America it is
much more common than E. albofasciata. a primarily South
American species.

NEW RECORDS

GUATEMALA: 19, Secanquim, June 1984 (USNM); 19,
Quirigua, Aug. (Schaus and Barnes; USNM). MEXICO, OA-
XACA: 299, Salina Cruz, 10 Aug. 1964 (E. Fisher and D.
Verity; LACM). QUINTANA ROO: 19, 3 33, 8-14 May 1963
(E.C. Welling; LACM). SINALOA: 19, 5 mi. N Mazatlan, 29
July 1 973 (J.A. Chemsak, E.G. Linsley, A.E. and M.M. Mich-
elbacher; UCB), on Turnera diffusa. VERA CRUZ: 599, Te-
colulla, 19 June 1951 (P.D. Hurd; UCB). YUCATAN: 19,
Piste, July 1967 (E.C. Welling; LACM). COSTA RICA,
GUANACASTE: 19, Liberia, 15-16 Feb. 1972 (P.A. Opler;
UCB), on Andira inermis\ 299, Finca La Pacifica, 4 km NW
Canas, 17-20 Nov. 1972 (P.A. Opler; UCB), on “pink malp.

Contributions in Science, Number 347

Snelling: American Centridini 63

Figures 81-84. Epicharis (Epicharoides) maculata, male seventh and eighth sternites and genitalia (ventral and dorsal views). Scale line =
1 .00 mm.

vine”; 19, same except 30 Nov. 1 972, on Bixa orellana\ 1 133,
Hacienda Comelco, 24 km NW Canas, 13-14 Mar. 1971
(E.R. Heithaus; LACM), on Secundaca tenuifo!ia\ 333, Ha-
cienda Comelco, 8 km NW Bagaces, 28-3 1 Jan., 6 Mar. 1971

(P. A. Opler; UCB), on Secridaca sylvestns. HEREDIA: 1 099,
7133, Ftnca La Selva, near Puerto Viejo, 25 Apr. to 19 June
(D.R. Perry; LACM), on Vochysia sp. (533), Byrsonima sp.
(599), Diptervx panamensis (599, 6033), Dussia sp. (333), and

64 Contributions in Science, Number 347

Snelling: American Centridini

Hymenolobium sp. (1<5). PANAMA, PANAMA: 2<3<5, Capira,
Cerro Campana, 12 Aug. 1980 (D. Roubik; ROUB); 1<3, 15
km E Chepo, Llano Carti Rd., 22 Feb. 1980 (D. Roubik;
ROUB).

Subgenus Epicharitides Moure

Epicharitides Moure, 1945a: 311. Type-species: “Epichari-
tides cockerelli (Friese, 1900)” = Epicharis cockerelli Fnese,

1900a; original designation.

DESCRIPTION

Maxillary palp three-segmented, second segment broad, longer
than first, third segment narrower and a little shorter than
second; lateral ridges of clypeal disc strong; malar space dis-
tinct, but shorter than minimum thickness of first flagellar
segment; frontal carina sharp but short, ending in front of
anterior ocellus by much more than diameter of anterior
ocellus; occipital margin compressed and ridge-like or crest-
ed; flagelliform occipital setae short and curved laterad before
reaching anterior margin of mesoscutum; posterior margin
of dorsal face of scutellum not impressed; metanotum wholly
vertical, jugal lobe of posterior wing shorter than cubital cell
and less than one-half as long as vannal lobe.

Female. Labrum without median ridge; outer face of me-
sobasitarus with mixed long, coarse, simple setae and short,
fine, long-plumose hairs on anterior half, posterior half with
sparse long, coarse, simple setae only; secondary basitibial
plate absen; metatibia no longer than metabasitarsus; grad-
ulus of third and fourth terga, in middle, broadly deflected
apicad and area on either side depressed and covered with
dense mat of very short, plumose hairs; margins of pygidial
plate, in dorsal view, nearly straight, apex narrowly truncate,
disc depressed.

Male. Labrum without median ridge; first flagellar segment
shorter than either scape or second flagellar segment; ocel-
locular distance greater than diameter of anterior ocellus;
procoxa without apicoventral process; mesosternal tubercles
absent; metatrochanter and metafemur without ventral seta
patch; metatibia without posteroventral ridge; metabasitar-
sus without carinate ridge on anterior margin; pygidial plate
less than one-half as wide as seventh tergite, margins sharp,
apex narrowly truncate.

Moure (1945b) recognized five species in this subgenus
and gave a key for their separation. Two additional species
were described from Brazil by Moure and Seabra ( 1959). All
the included species were known, at that time, only from
South America, especially Brazil. One species is now known
to occur in Central America.

Epicharis ( Epicharitides ) species

Two females from Cerro Campana, Panama Province, Pan-
ama, 9 June 1960 (W.J. Hanson; UKAN) possibly represent
an undescribed species. In the key to species of Epicharitides
by Moure (1945b) they fail at couplet 7, failing to agree with
the characteristics cited for either E. obscura Friese or E.
duckei Friese. The abdomen is black ventrally as well as

dorsally, there is a narrow transverse lateral stripe on each
side of the dorsal face of the first tergum and the second
tergum has a broad basal yellow band, slightly expanded on
either side. The distal margin of this band is broadly curved
inward, but in the middle of the segment, the band is one-
half as long as the segment. The face is wholly black except
for the yellow basal portion of the labrum and a minute
median spot on the lower paraocular area.

These specimens may represent a previously undescribed
species, but in the absence of associated males, no nomen-
clatural action is possible at this time.

These specimens differ from E. duckei in the color pattern
of the abdominal terga. The first tergite bears a short, sub-
lateral yellow stripe and the second is yellow across the entire
base, with the distal one-third blackish, the yellow band a
little shorter in the middle one-half. The remainder of the
abdomen is blackish. Pubescence is wholly dark, except the
pale scopa and some pale hairs around the pronotal lobe.
Females of E. obscura agree with the Panamanian specimens
in having the pubescence largely dark, but the first tergum
is immaculate and the second to fourth or fifth terga are
maculate only at the sides.

Superficially, these females resemble unusually dark in-
dividuals of E. maculata, but may be readily separated by
the shorter jugal lobe of the hind wing, by the characteristic
shape of the metabasitarsus and by the presence of the mod-
ified areas on the abdominal terga. These differences will also
distinguish this bee from E. albofasciata.

SYNONYMIC LIST OF NORTH AND CENTRAL
AMERICAN CENTRIDINI

Centris Fabricius

Subgenus Xerocentris Snelling, 1974. Type-species: C. cali-
fornica Timberlake, 1940.

01. californica Timberlake, 1940. Calif., Nev.

02. griseola Snelling, n. sp. Mexico (Guerrero).

03. hoffmanseggiae Cockerell, 1897. N. Mex., Ariz., Calif.

= davidsoni Cockerell, 1904.

04. pallida W. Fox, 1899. Southwestern U.S., northwestern

Mexico.

= callognatha Cockerell, 1923.

= trichosoma Cockerell, 1923.

05. rhodomelas Timberlake, 1940. Calif.

06. tiburonensis Cockerell, 1923. Calif., Ariz., Son., B. Calif.

07. vanduzeei Cockerell, 1923. Mexico (B. Calif., B. Calif.

Sur).

Subgenus Paracentris Cameron, 1902. Type-species: C. ful-
vohirta (Cameron, 1902).

= Penthemisia Moure, 1950. Type-species: C. chilensis
Spinola, 1851.

= Trichocentris Snelling, 1956. Type-species: C. rho-
doleuca Cockerell, 1923.

08. angustifrons Snelling, 1966. Ariz., Son.

09. aterrima F. Smith, 1854. Ariz. to central Mexico.

1 0. atripes Mocsary, 1 899. Southwestern U.S. to Costa Rica.
= atriventris W. Fox, 1899.

Contributions in Science, Number 347

Snelling: American Centridini 65

= limbata Friese, 1899. N. SYN.

= Foxi Friese, 1900b.

11. caesalpiniae Cockerell, 1897. Tex. to Ariz., northern

Mexico.

= morsei Cockerell, 1897.

= marginata W. Fox, 1899.

12. cockerelli W. Fox, 1899. Southwestern U.S., northern

Mexico.

= resoluta Cockerell, 1923. N. SYN.

13. ectypha Snelling, 1974. B. Calif.

14. ferrisi Cockerell, 1924. B. Calif. Sur.

15. fisheri Snelling, 1974. B. Calif, B. Calif. Sur.

16. harbisoni Snelling, 1974. B. Calif

17. laevibullata Snelling, 1966. Central Mexico.

18. lanosa Cresson, 1872. Fla. to Tex., Okla., Kans.

= subhyalina W. Fox, 1899.

= birkmanii Friese, 1900a.

19. mexicana F. Smith, 1854. Texas to Ariz., south to Oa-

xaca, Mexico.

20. nigrocaerulea F. Smith, 1874. Central Mexico to Pan-

ama.

= clypeata Friese, 1899. Preoccupied. N. SYN.

= anthracina Snelling, 1966. N. SYN.

21. rhodopus Cockerell, 1 897. Southwestern U.S., northern

Mexico.

= pulchrior Cockerell, 1900.

= rhodoleuca Cockerell. 1923.

22. zacateca Snelling, 1966. Ariz., N. Mex., central Mexico.

Subgenus Xanthemisia Moure, 1945b. Type-species: C. bi-
color Lepeletier, 1841.

23. caro/ae Snelling, 1966. Mexico to Costa Rica.

24. lutea Friese, 1899. Central Mexico to Panama; South

America.

25. rubella F. Smith, 1854. Panama; South America.

Subgenus Exa/locentris Snelling, 1974. Type-species: C. an-
omala Snelling, 1966.

26. anomala Snelling, 1966. Central Mexico.

Subgenus Acritocentris Snelling, 1974. Type-species: C. ru-
thannae Snelling, 1966.

27. agameta Snelling, 1974. Northeastern and central Mex-

ico.

28. a/biceps Friese, 1899. Northeastern and central Mexico.
= strawi Snelling, 1966. N. SYN.

29. ruthannae Snelling, 1966. Ariz., Son.

30. satana Snelling, n. sp. Southern Arizona to central Mex-

ico.

Subgenus Centris Fabncius, 1804. Type-species: C. hae-
morrhoidalis (Fabricius, 1775).

= Hemisia Klug, 1807. Type-species: C. haemorrhoi-
dalis (Fabricius, 1775).

= Cyanocentris Friese, 1900b. Type-species: C. versi-
color (Fabricius, 1775).

= Poecilocentris Friese, 1900b. Type-species: C.fascia-
tella Friese, 1 900b.

31. adanae Cockerell. 1949. Southern Mexico to Panama.

32. aethiocesta, Snelling, n. sp. El Salvador to Panama.

33. aethyctera Snelling, 1974. Southern Mexico to Panama.

34. decolorata Lepeletier, 1841. Texas to Panama; Carib-

bean; northern South America.

= obscuriventris Friese, 1899?

35. eisenii W. Fox, 1899. Arizona to Panama.

36. errans W. Fox, 1899. Florida.

37. flavifrons (Fabricius, 1775) Central America; South

America.

= brasiliana Christ, 1791.

= citrotaemata Gribodo, 1894. N. SYN.

= nigritula Friese, 1899. N. SYN.

= rufescens Friese, 1899. N. SYN.

38. jlavo/asciata Friese, 1900a. Central America, northern

South America.

39. inermis Friese, 1899. Central America, northern South

America.

= segregata Crawford, 1906. N. SYN.

= gualanensis Cockerell, 1912.

= robusta Cockerell, 1949. N. SYN.

= pallidifrons Cockerell, 1949.

40. meaculpa Snelling, n. name. Eastern Mexico.

= erubescens Snelling, 1974. Preoccupied.

41. obscurior Michener, 1954. Southern Mexico to Panama.

Subgenus Ptilocentris Snelling, n. subg. Type-species: C. /es-
tiva F. Smith, 1854.

42. /estiva F. Smith, 1854. Costa Rica to Venezuela and

Peru.

= chlorura Cockerell, 1919. N. SYN.

Subgenus Melanocentris Friese, 1 900b. Type-species: C. atra
Friese, 1899.

43. agilis F. Smith, 1874. Mexico to Honduras.

= ignita F. Smith, 1874. N. SYN.

= bakeri Friese, 1912. N. SYN.

= bakerella Friese, 1913. N. SYN.

= Epicharis cisnerosi Cockerell. 1949. N. SYN.

44. agiloides Snelling, n. sp. Mexico to Costa Rica.

45. fl avilabris Mocsary, 1899. Costa Rica to South America.
= boliviensis Mocsary, 1899.

46. fusciventris Mocsary, 1899. Costa Rica to South Amer-

ica.

= scutellata Friese, 1900b. N. SYN.

47. gelida Snelling, n. sp. Mexico, Guatemala.

48. obso/eta Lepeletier, 1841. Mexico to South America.

= melanochlaena F. Smith, 1874. N. SYN.

= Epicharis zamoranensis Cockerell, 1949. N. SYN.

49. plumipes F. Smith, 1854. Costa Rica to South America.

50. sericea Friese, 1899. Central Mexico.

Subgenus Trachina Klug, 1810. Type-species: C. longimana
Fabricius, 1804.

= Paremisia Moure, 1945b. Type-species: C. lineo/ata
Lepeletier, 1841.

51. dentata F. Smith, 1854. Southern Mexico to South

America.

= proxima Friese, 1900a.

52. eurypatana Snelling, n. sp. Mexico (Jalisco).

53 . fuse at a Lepeletier, 1841. Southern Mexico to South

America.

66 Contributions in Science, Number 347

Snelling: American Centridini

54. heithausi Snelling, 1974. Guatemala to Costa Rica.

55. labiata Friese, 1904. Mexico to Costa Rica.

= schwarzi Cockerell, 1919. N. SYN.

56. longimana Fabricius, 1804. Nicaragua to South Amer-

ica.

= personata F. Smith, 1874.

57. similis (Fabricius, 1804). Costa Rica to northern South

America.

= lineolata Lepeletier, 1841.

58. vidua Mocsary, 1899. Belize to Panama.

59. xochipillii Snelling, n. sp. Mexico (Oaxaca).

Subgenus Hemisiella Moure, 1945b. Type-species: C. lanipes
(Fabricius, 1775).

60. dichrootricha Moure, 1945b. Panama; South America.

6 1 . nitida F. Smith, 1 874. Mexico to Panama; northern South

America.

= confinis Perez, 1905. N. SYN.

62. transversa Perez, 1905.

= ruae Cockerell, 1949. N. SYN.

63. trigonoides Lepeletier, 1841. Southern Mexico to South

America.

= dentipes F. Smith, 1874. N. SYN.

= hoplopoda Moure, 1943.

= rufomaculata Cockerell, 1949. N. SYN.

= subtarsata Cockerell, 1949. N. SYN.

64. vittata Lepeletier, 1841. Mexico to South America.

= montezuma Cresson, 1879.

= breviceps Friese, 1899.

= friesei Crawford, 1906. Preoccupied.

= costaricensis Crawford, 1907.

= erubescens Friese, 1925. N. SYN.

Subgenus Heterocentris Cockerell, 1899. Type-species: C.
cornuta Cresson, 1865.

65. analis (Fabricius, 1804). Mexico to South America.

= totonaca Cresson, 1879.

= otomita Cresson, 1879.

= minuta Mocsary, 1899.

= simplex Friese, 1899.

= durantae Cockerell, 1949.

= petreae Cockerell, 1949.

= petreae var. rufopicta Cockerell, 1949.

66. bicornuta Mocsary, 1899. Mexico to South America.

67. difformis F. Smith, 1854. Costa Rica; South America.

68. labrosa Friese, 1899. Costa Rica to South America.

= triangulifera Cockerell, 1949. N. SYN.

Pti lot opus Klug

Ptilotopus Klug, 1810. Type-species: P. americanus Klug,
1810.

69. zonatus Mocsary, 1899. Panama.

= pandora Friese, 1900b.

Epicharis Klug

Subgenus Epicharana Michener, 1954. Type-species: E. rus-
tica (Olivier, 1789).

70. angulosa Snelling, n. sp. Costa Rica.

7 1 . bova Snelling, n. sp. Costa Rica, Panama.

72. elegans F. Smith, 1861. Mexico to Costa Rica.

= salazari Cockerell, 1949. N. SYN.

73. rustica (Olivier, 1789). Costa Rica to South America.

= flava Friese, 1900b. N. SYN.

Subgenus Hoplepicharis Moure, 1945a. Type-species: E.fas-
ciata Lepeletier and Serville, 1828.

74. lunulata Mocsary, 1899. Mexico to Panama.

75. monozona Mocsary, 1899. Panama.

Subgenus Parepicharis Moure, 1945a. Type-species: E. zo-
nata F. Smith, 1854.

76. metatarsalis Friese, 1899. Costa Rica to Venezuela.

= phenacura Cockerell, 1917. N. SYN.

= conura Cockerell, 1917. N. SYN.

Subgenus Epicharoides Radoszkowski, 1884. Type-species:
E. maculata F. Smith, 1874.

77. albofasciata F. Smith, 1874. Costa Rica to South Amer-

ica.

= nigroclypeata Friese, 1899.

78. maculata F. Smith, 1874. Mexico to northern South

America.

= bipunctatus Radoszkowski, 1884.

= variabilis Friese, 1900b.

Subgenus Epicharitides Moure, 1945a. Type-species: E.
cockerelli Friese, 1900a.

79. undetermined species near obscura Friese. Panama.

ACKNOWLEDGMENTS

For the use of material cited in this study 1 am indebted to:
P.H. Arnaud, Jr., and W.J. Pulawski (CAS), R.W. Brooks
(RWB), R.E. Coville, G.R. Frankie, E.G. Linsley, and J.A.
Powell (UCB), G.C. Eickwort (CORN), H.E. Evans (MCZ),
M. Favreau and J.G. Rozen, Jr. (AMNH), S.S. Gingras and
the late P.D. Flurd, Jr. (USNM), C.D. Michener (UKAN),
A.R. Moldenke (ORSU), J.L. Neff (NEFF), D. Roubik
(ROUB), R.O. Schuster, L.S. Kimsey, and K.E. Steiner (UCD),
L.A. Stange (DPIF), and T.J. Zavortink (TJZ).

In particular, I wish to express my gratitude to G.R. Else,
S.S. Gingras, and S. Kelner-Pillault for making available crit-
ical type material from the British Museum (Natural His-
tory), U.S. National Museum of Natural History, and the
Museum d’Histoire Naturelle, respectively. Special thanks
also, to R.W. Brooks for important notes on Friese types at
the Berlin Museum.

For critically reading the manuscript and offering helpful
suggestions I am indebted to C.L. Hogue and to the various
reviewers.

Finally, I wish to express my thanks to Beatriz Larrain for
providing the Spanish Resumen for this paper.

LITERATURE CITED

Alcock, J., C.E. Jones, and S.L. Buchmann. 1976. Location
before emergence of the female bee, Centris pallida, by
its male (Hymenoptera: Anthophoridae). Journal of Zo-
ology 179:189-199.

Contributions in Science, Number 347

Snelling: American Centridini 67

. 1977. Male mating strategies in the bee Centris

pallida Fox (Anthophoridae: Hymenoptera). The Amer-
ican Naturalist 1 1 1:145-155.

Cameron, P. 1903. Descriptions of new species of Hyme-
noptera taken by Mr. Edward Whymper on the “Higher
Andes of the Equator.” Transactions, American Ento-
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68 Contributions in Science, Number 347

Snelling: American Centridini

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Submitted 18 April 1983; accepted 31 August 1983.

Contributions in Science, Number 347

Snelling: American Centridini 69

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SALAMANDERS OF THE NORTHWESTERN
HIGHLANDS OF GUATEMALA

Paul Elias'

ABSTRACT. The salamander faunas of the Cordillera de los Cu-
chumatanes and the Montanas de Cuilco of western Guatemala are
reviewed in light of extensive new collections. Thirteen species are
reported from the Cuchumatanes including one new member of the
Bolitoglossa mexicana species group, herein described as Bolito-
glossa jacksoni sp. nov., and four recognized species never before
collected in the massif ( Bolitoglossa hartwegi, B. mulleri, B. rufescens
and Dendrotriton rabbi). Four species are reported from the Cuilco
including two hitherto unknown from that mountain range ( Boli-
toglossa mono and Pseudoeurycea rex).

Bolitoglossa resplendens (sensu lato) lacks the features purported
to differentiate it from B. lincolni and is considered a junior syn-
onym. Based on gross morphological similarity, the Bolitoglossa
helmrichi and B. rostrata species groups are divided and reconsti-
tuted into four species groups (dunni, franklini. morio and veracrucis
groups).

The Cuchumatanes support fewer salamander species but a larger
number of species groups and genera than do the Pacific Uplands.
The diversity differences are correlated with greater age and con-
nectedness among high elevation salamander habitats in the Cu-
chumatanes relative to the Pacific Uplands.

Salamander species on the Cuchumatanes and Cuilco occur in
narrow elevational belts as they do on the Pacific Uplands. Con-
specific populations or related species tend to occur at comparable
elevations wherever found. Within-group variation in elevational
range does exist, however, and much of it is correlated with geo-
graphic variation in the elevation of the cloud line. The Bolitoglossa
veracrucis species group is exceptional among Nuclear Central Amer-
ican salamander groups in the extreme breadth of the combined
elevational ranges of its three member species (2,800 m; the next
broadest is 1,200 m). The great ecological scope of the Bolitoglossa
veracrucis group may have been a crucial factor in the invasion of
the lowlands by Bolitoglossa beta.

RESUMEN. Las faunas de salamandras de la Cordillera de los Cu-
chumatanes y de las Montanas de Cuilco en el oeste de Guatemala
se revisan en base a las extensas nuevas colecciones realizadas. Trece
especies se encuentran en los Cuchumatanes, incluyendo una nueva
especie, Bolitoglossa jacksoni, miembro del grupo espedfico de Bo-
litoglossa mexicana, y cuatro especies ya descritas pero nunca en-
contradas en esta sierra previamente (Bolitoglossa hartweig, B.

1. Museum of Vertebrate Zoology, University of California,
Berkeley, California 94720.

mulleri, B rufescens y Dendrotriton rabbi). Cuatro especies se
encuentran en el Cuilco incluyendo dos anteriormente desconocidas
en esta region (Bolitoglossa morio y Pseudoeurycea rex).

Bolitoglossa resplendens (sensu lato) no tiene las caracteristicas
que supuestamente la diferencian de la B. lincolni, y consecuente-
mente B. resplendens es un sinonimo. En base de similaridades en
morfologia externa, los grupos especificos de Bolitoglossa helmrichi
y B. rostrata descritos por Wake y Lynch (1976) son divididos y
redefinidos en cuatro grupos (dunni, franklini, morio y veracrucis).

En los Cuchumatanes habitan menos especies de salamandras pero
un numero mayor de grupos especificos y generos, en comparacion
con las montafias del lado Pacifico. Estas diferencias de diversidad
estan correlacionadas con una edad geologica mayor y con una mayor
conexion entre regiones de alta elevacion en comparacion con las
montafias del lado Pacifico.

Las especies de urodelos en los Cuchumatanes y el Cuilco se lo-
calizan en estrechas bandas elevacionales al igual que en las mon-
tafias del Pacifico. Poblaciones de la misma especie, o especies re-
lacionadas, generalmente se encuentran en la misma zona elevacional.
Pero, si se encuentra variacion al nivel de intra-grupo, la mayor
parte de esta variacion esta correlacionada con la variacion geografica
a la altura de la zona nublada. El grupo espedfico de Bolitoglossa
veracrucis es una exception entre todos los grupos de Centro America
Nuclear debido a la extrema distribucion elevacional de las tres
especies miembros (2,800 m); la segunda distribucion mas amplia
es 1,200 m. El gran alcance geologico de este grupo puede tener un
papel importante en la historia de la invasion de las tierras bajas
por Bolitoglossa beta.

INTRODUCTION

Nuclear Central America (NCA), the landmass bounded to
the northwest by the Isthmus of Tehuantepec and to the
southeast by the Nicaraguan depression, contains six major
highland masses isolated from one another by deep dry val-
leys (Fig. 1). The Cordillera de los Cuchumatanes and the
Montanas de Cuilco are two limestone capped ranges, re-
spectively about 3,000 and 400 square kilometers in extent
and 4,000 and 3,400 meters at greatest elevation (Fig. 2).

On the Caribbean slope of the Cuchumatanes, tropical rain
forest (following the ecological terminology of Schmidt, 1 936)
extends from near sea level to a 1,300 m "cloud line” (that
characteristic elevation at which the water carried by rising

Contributions in Science, Number 348, pp. 1-20
Natural History Museum of Los Angeles County, 1984

ISSN 0459-8113

humid air condenses in the form of fog). Above the cloud
line is a zone of extremely wet broadleaf and treefern forest,
the subtropical zone cloud forest, which grades upward into
drier but still humid subtropical forest that is frequently com-
posed of oak. At about 2,700 m, varying locally, the pine
and cypress temperate zone forests commence, continuing
to the 3,400-m treeline, above which bunchgrass predomi-
nates.

The above vegetational series is found on the unobstructed
face of the Caribbean slope; but, in the protected interior
valleys, variably drier systems occur, typically dominated by
pine-oak forest even at low elevation. Because of their pro-
tected position, the Montanas de Cuilco have an abbreviated
form of the ecological pattern found on the open escarpment:
as in the Cuchumatanes, pine and cypress dominate down
to about 2,700 m, where a wet broadleaf forest begins; how-
ever, a high cloud line occurs at about 2,000 m, and dry
scrubby pine-oak growth predominates below this level.

The salamander faunas of the Cordillera de los Cuchu-
matanes and the Montanas de Cuilco of western Guatemala
have been little sampled or reported upon. In the Cuchu-
matanes Stuart ( 1 943a, 1 943b) made the first collections and
reported four species referred to Oedipus : lincolni, cuchu-
matanus, rostratus, and rex. From later collections Stuart
(1952) described Magnadigita omniumsanctorum. The next
figure 1. Nuclear Cential America. The 2,100-m contour interval discovery in the Cuchumatanes was recorded by Lynch and
is suppled.

Figure 2. Montanas de Cuilco and the Cordillera de los Cuchumatanes with the 2,100-m contour interval and regional landmarks.

2 Contributions in Science, Number 348

Elias: Guatemalan Salamanders

Figure 3. Bolitoglossa rostrata, B. cuchumatana, B. mono, and B. lincolni, life size. UPPER LEFT: Boliloglossa rostrata, body, legs, and
flanks dark brown, bilateral shoulder stripes lighter brown with middorsal brown mottling. A unicolor brown morph also is found. UPPER
LEFT INSET: Feet of B. cuchumatana. Overall coloration like that of B. rostrata. UPPER RIGHT: B. morio, dark slate grey all over except
for light speckling on flanks and anterior surfaces of limbs, and light lateral spotting on tail (Cuilco specimen figured; Cuchumatan specimens
have light spotting on flanks as well as tail). BOTTOM CENTER: B lincolni, black ground color with a coral red dorsal swath and spotting
on limbs and (occasionally) venter. In some specimens, the dorsal swath is reduced to a series of red spots.

Wake (1975) with the naming of Chiropterotriton cuchu-
matanus. Finally, Elias and Wake (1983) and Wake and Elias
(1983) named two new monotypic genera, Nyctanolis pernix
and Bradytriton silus, both from the cordillera.

Prior to the present study the only salamanders that had

been collected in the Montanas de Cuilco were Bolitoglossa
resplendens ( sensu lato) and Chiropterotriton rabbi, both first
reported by Lynch and Wake (1975).

Bolitoglossa stuarti was described from the valley sepa-
rating the Cuilco from the Cuchumatanes (Wake and Brame,

Contributions in Science, Number 348

Elias: Guatemalan Salamanders 3

□

Figure 4. Relationship between maxillary tooth number (both sides
summed) and standard length in the Bolitog/ossa dunni complex.
Solid triangle = B. cuchumatana (Eastern Cuchumatanes); open tri-
angle = B. cuchumatana (Western Cuchumatanes); half-filled cir-
cle = B. helmrichi (Chuacus); solid circle = B helmrichi (Western
Minas); open circle = B. helmrichi (Xucaneb); solid square = B. en-
gelhardti (Pacific Uplands); open square = B. sp. (Eastern Cuchu-
matanes); half-filled square = B. dunni (Elonduras).

1969) and is the only salamander species known front the
valley. As a low elevation form this species will be mentioned
only in the discussion.

In the course of the present study twelve salamander species
were documented for the Cuchumatanes, bringing the total
number of known forms from six to thirteen for the mountain
range. Among the seven species previously unknown from
the massif were the two new monotypic genera described
elsewhere (Elias and Wake, 1983; Wake and Elias, 1983) and
the one new member of the Bo/itoglossa mexicana species
group described below. Four species were taken in the Mon-
tanas de Cuilco during this work. All of these were recognized
forms, but two were previously unknown from those moun-
tains.

This paper provides a species account for each of the sal-
amander species now known from the Cuchumatanes and
Cuilco, including taxonomic synonymies, type locality, geo-
graphic range, habits and habitat preferences and a discussion
of the geographic variation and possible evolutionary rela-
tionships of the taxon. The discussion section is divided into
three parts. In the first part, the species groups within Boli-
toglossa beta ( sensu Wake and Lynch, 1976) are redefined.
In the second part, contemporary patterns of distribution
and differentiation are examined to provide insight into the
history of the salamander fauna. In the third part, patterns
of elevational distribution are presented for ihe Cuchuma-

tanes and Cuilco and are discussed relative to the results of
Wake and Lynch (1976) for the Pacific Uplands.

SPECIES ACCOUNTS

Bolitoglossa cuchumatana (Stuart, 1943)

Figure 3

Oedipus cuchumatanus Stuart, 1943a: 14.

Magnadigita cuchumatana: Taylor, 1944:218.

Bolitoglossa cuchumatana: Wake and Brame, 1963:386.

TYPE LOCALITY. Oak woods 2 km N Nebaj, Depto.
Quiche, Guatemala, 1,900 m elevation.

RANGE. Stuart collected this form only at the type lo-
cality. Wake, Houck, and Lynch next found the species 25
years later about 15 km to the south and 600 m higher in
elevation. I found B. cuchumatana at two additional sites in
the western Cuchumatanes near Chiapas and at the eastern-
most end of the range, near San Miguel Uspantan.

HABITAT AND HABITS. B. cuchumatana have been
taken on the ground, under bark, and in arboreal bromeliads
where they have been found active on surface debris on wet
evenings from the 1 ,200-m cloud line through the subtropical
cloud forest to 2,500 m.

SYSTEMATICS. A series of species, including Bolito-
glossa engelhardti in the Pacific Uplands, B. helmrichi in the
Xucaneb and Western Minas, a newly discovered population
(possibly representing a new species) in the Eastern Minas,
B. cuchumatana in the Cuchumatanes, and B. dunni in Hon-

Table 1. Geographic variation in subcaudal coloration and degree
of interdigital webbing among the species of the Boltioglossa dunni
complex.

Species and
population

Subcaudal

coloration

Degree of
interdigital
webbing

B. helmrichi
(Chuacus)

no data

high

B. helmrichi
(Xucaneb)

orange

high

B. helmrichi
(Western Minas)

orange

high

B. sp.

(Eastern Minas)

tan

low

B. cuchumatana
(Western Cuchumatanes)

tan

medium

B. cuchumatana
(Eastern Cuchumatanes)

tan

medium

B. engelhardti
(Pacific Uplands)

tan

medium

B. dunni
(Honduras)

no data

medium

4 Contributions in Science, Number 348

Elias: Guatemalan Salamanders

Figure 5. BolHoglossa jacksoni, B. rufescens, B. mulleri, Dendrotriton rabbi, D. cuchumatanus, and Boliloglossa hartwegi. life size. UPPER
LEFT: Boliloglossa jacksoni (holotype figured), background yolky yellow, middorsal swath dark brown with narrow white border. TOP
CENTER: B. rufescens. tan overall with a slightly darker brown snout and flanks. Tone varies front light to dark brown both between individuals
and for one individual at different times of day. UPPER RIGHT: B. mulleri, black overall except for a narrow broken vertebral stripe of tan.
In some Cuchumatan specimens, the stripe is reduced or absent. LOWER LEFT: Dendrotriton rabbi, pattern composed of various shades of
brown. Note light interorbital bar. Animals may have a reddish or even greenish cast. Pattern and tone variable between individuals. BOTTOM
CENTER: D. cuchumatanus, similar to D. rabbi above. LOWER RIGHT: Bo/itoglossa hartwegi, dark brown overall with lighter paired
shoulder and pelvic stripes. Animals may be without markings altogether or may have small light streaks on dorsum at random.

Contributions in Science, Number 348

Elias: Guatemalan Salamanders 5

Figure 6. Bolitoglossa jacksoni sp. nov., an adult female (escaped) from the type locality.

duras, appears to form a relatively homogeneous assemblage
of related congeners. The group, which I shall call the dunni
complex, was first outlined by Stuart (1943a) and is char-
acterized by a moderate degree of foot webbing; by well-
developed subdigital pads and the associated hypertrophy of
the terminal phalanges; by a generally brown dorsal color
frequently differentiated into light shoulder stripes and dark-
er flanks; and by a common microhabitat (see Wake and
Lynch, 1976, for information on the ecology of B. engel-
hardti).

The species in this series are most similar to Bolitoglossa
rostrata, but differences in foot morphology (B. rostrata has
little mterdigital webbing) usually permit separation (see B.
rostrata and compare inset B. cuchumatana feet. Fig. 3). B.
rostrata occurs in limited sympatry with B. cuchumatana
with no hybridization.

Despite the general similarities, variation between the
species of the B. dunni complex is pronounced in the col-
oration of the subcaudal surface, the number of maxillary
teeth, and the morphology of the foot. Data were collected
for eight populations from the five B. dunni complex species
(Fig. 4, Table 1). B. helmrichi can be defined on the basis of
its high degree of interdigital webbing and orange subcaudal
coloration to include populations in the Xucaneb, the West-
ern Minas, and the Chuacus. B. cuchumatana , including all
Cuchumatan specimens, and B. engelhardti are similar in
their intermediate degree of interdigital webbing, tan sub-
caudal surface, and moderate tooth number, and are distin-
guishable from one another both on the basis of adult size

(B. engelhardti is smaller than any other member of the com-
plex) and in frequency of shoulder striping ( B . engelhardti is
usually unmarked, while B. cuchumatana almost always has
bilateral light striping). Bolitoglossa dunni occurs in both a
pink and dark morph, the latter with light spotting (Schmidt,
1933), and it achieves larger size than any other member of
the complex and has little webbing and low tooth numbers.
The population from the Eastern Minas differs sharply from
all other members of the B. dunni complex (and from all
known B. rostrata populations as well) in having extremely
high numbers of maxillary teeth. It has almost as little web-
bing as B. rostrata. That this population belongs in the B.
dunni complex rather than to B. rostrata is suggested only
by its occurrence at relatively low elevation.

Bolitoglossa hartwegi Wake and Brame, 1969

Figure 5

Bolitoglossa hartwegi Wake and Brame, 1969:10.

TYPE LOCALITY. 4.5 mi. W San Cristobal de Las Casas,
Chiapas, Mexico. 7,000 ft elevation.

RANGE. B. hartwegi has been known only from the gen-
eral vicinity of its type locality on the Mesa Central de Chia-
pas. It is here reported from Guatemala, both in the western
Cuchumatanes at Finca Chiblac, and in the easternmost
reaches of the range, near San Miguel Uspantan.

HABITAT AND HABITS. Specimens have been collect-
ed under bark on logs and stumps, under cover objects on
the ground, in rock crevices, and in arboreal bromeliads; the

6 Contributions in Science, Number 348

Elias: Guatemalan Salamanders

species is active on rainy nights. B. hartwegi have a broad
elevational range, occurring from the 1,200-m cloud line to
the lower temperate forest at over 2,700 m.

SYSTEMATICS. When these populations were discov-
ered in the Cuchumatanes during this study they were as-
signed to the Bolitoglossa veracrucis species group (see Dis-
cussion) based upon their well-ossified skull with a broad
pars dentalis of the premaxillary, and a derived foot mor-
phology. They were assigned to the species B. hartwegi based
on a mainly unicolor dorsum ( B . veracrucis is streaked and
mottled), a small size (B. stuarti is larger), and a mottled
venter (unicolor in B. stuarti).

Bolitoglossa jacksoni new species

Figures 5 and 6

HOLOTYPE. MVZ 134634, a young adult female from
the Las Nubes sector of Finca Chiblac, approximately 1 2 km
NNE of Santa Cruz Barillas, Depto. Fluehuetenango, Gua-
temala, at about 1,400 m elevation, collected by Jeremy L.
Jackson, 1 September 1975.

DIAGNOSIS. The new taxon is a large species of Boli-
toglossa (the young adult holotype is 49.1 mm and another
specimen was approximately 65 mm in standard length) with
low numbers of maxillary teeth (31 in the holotype) and
moderate numbers of vomerine teeth (21 in the holotype).
The species is a member of the Bolitoglossa alpha group on
the basis of its simple vertebral tail autotomy mechanism
(Wake and Dresner, 1967; Wake and Lynch, 1976). It is
distinguished from B. salvinii, B. mexicana. B. mulleri, B.
odonne/li. B. platydactyla. and B. flaviventris in having dark
coloration restricted to the middorsal region; separated from
all other Nuclear Central American Bolitoglossa both by col-
or pattern and its tail autotomy mechanism.

DESCRIPTION (OF HOLOTYPE). Nostril small, labial
protuberances of nasolabial grooves small and poorly de-
veloped; canthus rostralis moderate in length, rounded and
gently arched. Standard length 6.6 times head width; stan-
dard length 4.5 times snout-gular fold length (head length).
Deep, unpigmented groove below eye extends almost full
length of opening, following curvature of eye, but does not
extend to lip. Eye moderate in size, slightly protuberant in
life. Lightly indicated postorbital groove extends posteriad
and slightly ventrad as a shallow depression for 2.7 mm,
thence proceeds directly ventrad and passes behind posterior
end of mandible to produce a shallow but definite nuchal
groove parallel to and 4.0 mm anterior to the dearly defined
gular fold. The 21 vomerine teeth are arranged in single
arched rows on either side of the midline and extend laterally
to the level of the midpoint of the internal nares. The two
rows of 3 1 maxillary teeth extend posteriad to a level one-
third of the way through the eye. Five small undifferentiated
premaxillary teeth protrude from well behind the lip. The
tail is round in cross section and is 0.8 times the standard
length from base to tip; the postiliac gland is not evident.
Limbs are of moderate length, three costal grooves are ex-
posed between the toetips of the adpressed fore and hind
limbs; standard length is 4.4 times right hind limb length

and 4.9 times right fore limb length. Standard length is 10.0
times the hind foot width and both fore and hind feet are
fully webbed and lack subdigital pads; the toetips are rounded
in dorsal aspect. The fingers in order of decreasing length: 3,
2, 4, 1; toes in order of decreasing length: 3, 4, 2, 5, 1.

MEASUREMENTS (OF HOLOTYPE). Head width
(maximum), 7.4 mm; head length (snout to gular fold), 1 1.0
mm; head depth at posterior angle of jaw, 4.1 mm; eyelid
length, 3.0 mm; eyelid width, 1.9 mm; anterior rim of orbit
to snout, 2.6 mm; horizontal orbital diameter, 2.2 mm; m-
terorbital distance, 3.0 mm; distance between vomerine teeth
and parasphenoid tooth patch, 0.3 mm; snout to fore limb,

13.6 mm; distance separating internal nares, 2.2 mm; dis-
tance separating external nares, 2.1 mm; snout projection
beyond mandible, 0.6 mm; snout to posterior angle of vent
(standard length), 49.1 mm; snout to anterior angle of vent,

45.6 mm; axilla to groin, 27.4 mm; tail length, 43.4 mm; tail
width at base, 3.7 mm; tail depth at base, 3.3 mm; fore limb
length, 10.0 mm; hind limb length, 1 1.2 mm; width of hand,
3.7 mm; width of foot, 4.9 mm.

COLORATION IN ALCOHOL. The ground color is pale
yellow with a broad middorsal swath of dark brown. The
brown marking originates on the head at the level of the
eyelids and extends in breadth from the center of one eyelid
to the center of the other in a straight edge across the head.
Posteriorly the brown stripe diminishes gradually until it
disappears near the tail tip. Aside from the brown swath, the
animal is unmarked (Fig. 5).

COLORATION IN LIFE. This animal was a brilliant yolky
yellow. The dorsal swath was chocolate brown and was en-
tirely bordered by an edging of immaculate white less than
a millimeter in width. The eye was metallic gold (Fig. 6).

OSTEOLOGY. As the animal was preserved flat in stan-
dard pose, accurate measurements from a radiograph could
be taken. Skull, from the anterior border of the premaxilla
to the medial edge of the occipital condyles, 8.0 mm; width
of pars dentalis of the premaxilla, 1.3 mm; maximum width
of braincase (anterior to the otic capsules) 3.0 mm; maximum
width of skull base (across otic capsules) 4.9 mm; longest
axis of otic capsule, 2.6 mm; and straight line length of the
maxilla from anterior to posterior tip, 4.5 mm.

The preorbital process of the vomer extends well lateral
to the internal narial opening as a slender sliver that tapers
to a point. The frontal processes of the premaxilla arise and
continue well separated from one another, spreading grad-
ually as they rise dorsad and posteriad. Septomaxillae are
absent. Trunk vertebrae number 14, caudosacral 2, and cau-
dal 30. No tibial spur is evident; phalangeal formulae are 1,
2, 3, 3, 2 for the foot and 1, 2, 3, 2 for the hand. Phalanges
are broadened and slightly irregular in outline, and the ter-
minals of the three outer fingers and four outer toes are
laterally expanded such that each is broader than it is long.
The innermost toe and finger have terminals that are pointed
and curved along the rim of the pad toward the other digits.

VARIATION. Two specimens of this form were collected,
but one, a large adult female, escaped before it could be
preserved. Knowledge of variation is thus restricted to as-
pects of color and pattern that were preserved in photographs

Contributions in Science, Number 348

Elias: Guatemalan Salamanders 7

98° 96° 94° 92° 90° 88°

Figure 7. Distribution of the Boliloglossa mexicana species group (modified front Wake and Lynch, 1976).

of the escaped animal (Fig. 6, and Hanken et al., 1980). All
the hues described for the holotype were similar in the other
individual, but the latter had a more irregular dorsal pattern.
The brown dorsal swath was interrupted by a large yellow
nape spot and a break on the tail, and the white border was
incomplete. In addition a spot of brown was present on the
dorsal surface of the right hind foot.

RANGE. Bolitoglossa jacksoni has been taken only within
1 km of the type locality on the Caribbean escarpment of
the western Cuchumatanes (Fig. 7).

HABITAT AND HABITS. Both individuals were taken
under bark of felled hardwood logs in clearings in a sub-
tropical forest, which receives upward of 6 m of rain annually.

SYSTEMATICS. Bolitoglossa jacksoni is a member of the
B. mexicana species group ( sensu Wake and Lynch, 1976)
which also includes B. Jlaviventris, B. mexicana, B. mulleri,
B. odonnelli, B. platydactyla, and B. salvinii. The members
of the group share a variety of structural and ecological traits;
all have the primitive (alpha type) vertebral tail autotomy
mechanism (Wake and Dresner, 1 967); all have fully webbed
feet that lack subdigital pads but have expanded terminal
phalanges; all are large species, each represented by speci-
mens exceeding 65 mm standard length; and they occur only
in the tropical and low subtropical forests.

The species of the B. mexicana group are similar in general
structure and have been defined mainly on the basis of col-
oration. The color patterns of the six species are described
below; their geographic ranges are indicated in Figure 7.

Bolitoglossa Jlaviventris typically has paired dorsolateral
light stripes that begin on each eyelid and join to produce a
light snout. In the extreme condition, these stripes are paired
all the way onto the tail, but often they are irregularly de-
veloped and may form a light reticulation, especially pos-
teriorly. This species differs from others with dark flanks in
having a light colored and unmarked ventral surface. The
light ventral coloration sometimes invades the dorsal sur-
faces of the legs as well, but the flanks are always dark and
sharply set off from the light belly (see Wake and Lynch,
1976, fig. 24). The eye is golden.

Bolitoglossa mexicana has a broad middorsal light swath
that is typically broken into three longitudinal stripes by the
invasion of dark ground color onto the dorsal surface. The
dorsal pattern may range from virtually a single swath (as in
B. platydactyla ) to triple stripes, and then, through loss of
the vertebral line, to a bilinear pattern much as seen in B.
odonnelli (described below). The snout is heavily flecked,
and light flecking extends over all of the dark regions of the
body (see Wake and Lynch, 1976, frontispiece; Taylor and

8 Contributions in Science, Number 348

Elias: Guatemalan Salamanders

Smith, 1945, fig. 58; Stuart, 1943a, plate II, fig. 2). The eye
is dark.

Bolitoglossa mulleri typically has a narrow vertebral light
stripe. When maximally developed, the stripe extends from
the tail tip to the nape of the neck where it bifurcates before
terminating on the eyelids. The light pigmentation is fre-
quently reduced, and may be absent altogether leaving a
uniformly black animal (see Fig. 5 and Stuart, 1943a, plate
II, fig. 4). The eye is dark.

Bolitoglossa odonnelh has narrow paired light stripes. At
their maximum development, these stripes extend continu-
ously from the eyelids to the tip of the tail, but frequently
they are broken into rows of dots or may not extend as far
anteriorly or posteriorly. The body, including the snout, is
otherwise unmarked black (see Stuart, 1943a, plate II, fig.
3). The eye is dark.

Bolitoglossa platvdactyla has a single broad middorsal light
swath on a generally dark ground. The snout is dark and
speckled with light flecks; additional flecking may occur on
the fore limbs. A light swath begins on the eyelids and extends
almost to the tip of the tail (see Wake and Lynch, 1976, fig.
43; Stuart, 1943a, plate II, fig. 1); the eye is dark.

Finally, B. salvinii has the same distribution of light dorsal
markings as B. Jlaviventris'. paired dorsolateral lines that join
anteriorly to produce a light-colored snout and that frequent-
ly meet across the dorsal midline or break into reticulation.
In this species, the ventral surfaces are dark and undiffer-
entiated from the dark flanks, although a light ventral blotch
occasionally occurs (see Wake and Lynch, 1976, fig. 24;
Schmidt, 1936, fig. 17). The eye may be either light or dark.

Relationships among the B. mexicana group species re-
main obscure. The animals are scarce, and more samples
would be needed for a comprehensive analysis of the entire
assemblage. While primitive-derived polarities are not
understood for the color characters separating the species,
the seven taxa cluster in two basic color and pattern groups.
Stuart (1943a) considered B. platvdactyla, B. mexicana, B.
mulleri, and B. odonelH to form a group united by aspects
of their coloration, and the remaining three taxa, B. jacksoni,

B. Jlaviventris, and B. salvinii, themselves form a cluster. All
three of the latter forms have ( 1 ) some white edging between
dark and light areas; (2) at least some individuals with golden
eyes; (3) the same basic hues, yellow (to tan) and chocolate
brown; and (4) a dorsal swath of brown bordered bilaterally
and on the snout by yellow. These three species are the only
members of the mexicana group that ever have light ventral
coloration (apparently universal in B. jacksoni and B. jla-
viventris and occasionally, as small blotches, in B. salvinii).
An additional unique feature of the jacksoni-jlaviventris-sal-
vinii assemblage is that each is marked only with solid colors,
the light regions a single uniform yellow and the dark regions
an equally pure brown; speckling and mottling are never
observed.

ETYMOLOGY. I name this rare and beautiful new species
for Jeremy L. Jackson, the friend who aided me in long moldy
months of wet season collecting, and the captor of the first,
and now sole, representative of this species.

Bolitoglossa lincolni (Stuart, 1943a)

Figure 3

Oedipus lincolni Stuart, 1943a:9.

Magnadigita lincolni: Taylor, 1944:218.

Bolitoglossa lincolni: Wake and Brame, 1963:386.
Bolitoglossa resplendens McCoy and Walker, 1966:1.

TYPE LOCALITY. Monte at Salquil Grande, Depto. El
Quiche, Guatemala, 2,450 m elevation.

RANGE. Including the range of B. resplendens (herein
synonymized with B. lincolni ), this is one of the widest rang-
ing of Central American salamanders. It occurs on the Pacific
Uplands, in the Cuilco, in the Cuchumatanes, and on the
Mesa Central de Chiapas. B. lincolni is herein reported from
one new locality in the Cuchumatanes and one in the Cuilco.

HABITAT AND HABITS. Bolitoglossa lincolni were tak-
en in bromeliads and under logs and bark (see Wake and
Lynch, 1976, fig. 30). In addition they were found to be active
on wet nights. The species occurs in the upper subtropical

Table 2. Summary of geographic variation in Bolitoglossa lincolni.

Population

Dorsal

pattern

Ventral

spotting

Foot

webbing

Max. standard
length

(millimeters)*

Pacific Uplands

Swath to spots

>95% of pop.

Reduced

m

76

f

74

Cuilco

Swath to spots

<60% of pop.

Intermediate

m

58

f

79

Cuchumatanes

Swath

>95% of pop.

Extensive

m

66

f

78

Mesa Central

Swath

>95% of pop.

Reduced

m

62.3**

r

78.4**

* Males (m), females (0.

** Data from McCoy and Walker (1966).

Contributions in Science, Number 348

Elias: Guatemalan Salamanders 9

forest from well above the cloud line up to the lower tem-
perate zone.

SYSTEMATICS. The diagnostic characters differentiat-
ing Bolitoglossa resp/endens from B. lincolni are its greater
standard length and proportionately shorter limbs. In the
description of B. resplendens it was reported that a higher
incidence of ventral spotting and less extensive interdigital
webbing also distinguish B. resplendens from B. lincolni
(McCoy and Walker, 1966).

Examination of specimens from the four known popula-
tions indicates that the two supposedly diagnostic features
of B. resplendens reflect comparison to a limited sample of
unusually small B. lincolni. Nominate B. lincolni from the
Cuchumatanes are fully as large as the specimens in the B.
resplendens type series (McCoy and Walker, 1966) from the
Mesa Central de Chiapas (Table 2), and both populations
have proportionately longer limbs at smaller standard length
(Fig. 8). Specimens from the Cuilco and Pacific Uplands have
the same maximum size (Table 2) and limb/body length
allometry (Fig. 8) as the other two populations.

Color pattern, both dorsal and ventral, and degree of in-
terdigital webbing vary between the four populations (see
Table 2 and Fig. 3 for dorsum of Cuchumatan specimen) but
not in concordant patterns that suggest species level differ-
entiation. Thus, I refer the name Bolitoglossa resplendens to
the synonymy of Bolitoglossa lincolni.

Wake and Lynch (1982) reported the results of morpho-
metric and electrophoretic analysis of the Bolitoglossa frank-
lini species group. While there is general agreement with my
conclusions, they choose to continue to recognize Bolito-
glossa resplendens as a distinct species based upon the large
genetic distance between the Cuilco population and those
from the Cuchumatanes and Pacific Uplands. The genetic
affinities of the nominate populations of Bolitoglossa re-
splendens on the Mesa Central de Chiapas remain unknown.

Bolitoglossa morio (Cope, 1869)

Figure 3

Geotriton carbonarius (part) Cope, 1868:313.

Oedipus morio Cope, 1869:103.

Geotriton morio: Smith, 1877:64.

Spelerpes morio: Boulenger, 1882:70.

Spelerpes bocourti Brocchi, 1883:1 1 1.

Magnadigita morio: Taylor, 1944:218.

Magnadigita omniumsanctorum Stuart, 1952:4.

Bolitoglossa omniumsanctorum: Wake and Brame, 1963:386.
Bolitoglossa morio: Wake and Brame, 1963:386.

TYPE LOCALITY. Mountains of Guatemala.

RANGE. This species occurs throughout the Pacific Up-
lands, in the Chuacus, and in the Cuchumatanes. It is here
reported for the first time from the Cuilco.

HABITAT AND HABITS. B. morio were taken under
logs, in logs, and in bromeliads and were found to be active
on rainy nights. They occur in the upper subtropical and
lower temperate zone forests between roughly 1,800 and
3,000 m.

A

18-

•

• A

OA OO «Of

°

• o

• 2

oO A A o

| 14-

it A •

-C

°o°»

O' _

c

A*

A A •

Bolitoglossa lincolni

1 l0~

•

o

A Mesa Central de Chiapas

a Cuchumatanes

c —

A •

X

A

• Cuilco

6-

•

1 1 1 1

O Pacific uplands

i ~T — r~ — ~n

10 20 30 40 50

Axilla -groin length (mm)

Figure 8. Relationship between hind limb length and axilla-groin
length in four populations of Bolitoglossa lincolni.

SYSTEMATICS. Relationships of this species are ob-
scure, but its closest relative appears to be B. flavimembris
(see species group section, below). The characters that unite
the populations of this species are the presence of large light
(whitish, cream, or pink) blotches on the flanks and/or sides
of the tail; the slate grey to brownish ground color of the
dorsum; salt and pepper belly mottling; reduced webbing
such that all toes of the hind foot are free at their tips; and
the robust body form.

The newly discovered Cuilco population has all of the
specific characters, except that the lateral markings are re-
duced to small flecks (diameters less than that of the eye),
which are restricted to the sides of the tail rather than ex-
tending onto the trunk.

Bolitoglossa mulleri (Brocchi, 1883)

Figure 5

Spelerpes mulleri Brocchi, 1883:1 16.

Oedipus p/atydactylus (part) Dunn, 1926:400.

Oedipus mulleri: Schmidt, 1936:150.

Bolitoglossa mulleri: Taylor, 1944:219.

Oedipus mexicanus mulleri: Stuart, 1948:19.

Bolitoglossa moreleti mulleri: Duellman, 1963:220.
Bolitoglossa mexicana mulleri: Wake and Brame, 1963:386.

TYPE LOCALITY. Several localities in Alta Verapaz, re-
stricted by Stuart (1943a) to "les montagnes qui dominent
Coban.”

RANGE. This form had been taken only in the Xucaneb
complex until discovered in the Cuchumatanes during the
study described here. It is now known from Santa Cruz Bari-
llas and from a finca 3 km to the NE at 1,500 m (Fig. 7).

HABITAT AND HABITS. This species occurs in the low
pine-broadleaf cloud forest border in the Xucaneb and was
found in the Cuchumatanes in the cutover Barillas valley,
which was once a mix of the two forest types. This area is
at the elevation of the lower subtropical zone but is drier

10 Contributions in Science, Number 348

Elias: Guatemalan Salamanders

because of its protected position. Of the ten or so specimens
taken from the Cuchumatanes, all but one were found under
stones in a well-watered garden in the town of Barillas. The
other specimen was found under a piece of wood in a stand
of coffee. In the Xucaneb the species has been taken between
1,000 and 1,500 m.

SYSTEMATICS. The new population has a somewhat
reduced middorsal light stripe relative to animals from the
type locality: markings never extend anterior to the nape,
and some individuals are unmarked black.

Bolitoglossa mulleri is a member of the Bolitoglossa mex-
icana species group (sensu Wake and Lynch, 1 976). Variation
and relationships among the mexicana group species are dis-
cussed in the B. jacksoni account above.

Bolitoglossa rostrata (Brocchi, 1883)

Figure 3

Spelerpes rostratum Brocchi, 1883:1 12.

Oedipus rostratum: Dunn, 1924:99.

Oedipus rostratus: Dunn, 1926:384.

Magnadigita rostrata: Taylor, 1944:218.

Bolitoglossa rostrata: Wake and Brame, 1963:386.

TYPE LOCALITY. The mountains of Totonicapan, Gua-
temala.

RANGE. This species is one of the most broadly ranging
forms in Nuclear Central America, from the Mesa Central
de Chiapas through the Cuchumatanes and into the Pacific
Uplands. A newly discovered population at the eastern ex-
tremity of the Cuchumatanes indicates that the species may
be distributed throughout that cordillera.

HABITAT AND HABITS. B. rostrata were taken in and
under logs, under bark, in bunchgrass, and under rocks, and
were found to be active on rainy nights. The species occurs
only above 2,700 m and up to about 3,000 m, in temperate
forests and above the treeline in bunchgrass associations.

SYSTEMATICS. The species is distinguished by the fol-
lowing characters: it has almost unwebbed toes (as unwebbed
as any member of the genus); it is brown, typically with a
middorsal swath of light color or light shoulder stripes; and
it has a uniform light beige to dull yellow ventral and sub-
caudal color.

B. rostrata appears to be most closely related to the B.
dunni series (see B. cuchumatana account above) and where
it occurs in sympatry with B. cuchumatana the two species
are separated with difficulty. Bolitoglossa rostrata is distin-
guishable from B. cuchumatana on a number of subtle char-
acters; it occurs in an unstriped morph, is slightly less robust,
has a longer tail, and shows reduced interdigital webbing (Fig.

3).

Bolitoglossa rufescens (Cope, 1869)

Figure 5

Oedipus rufescens Cope, 1869: 104.

Geotriton rufescens: Smith, 1877:76.

Spelerpes rufescens: Boulenger, 1882:71.

Bolitoglossa rufescens: Taylor, 1 94 1 : 1 45.

Palmatotriton rufescens: Smith, 1945:4.

TYPE LOCALITY. Orizaba, Veracruz, Mexico.

RANGE. The range of this species is broad, extending
along the Atlantic versant throughout western Honduras,
Guatemala, and Chiapas, Mexico, and continuing up the
Atlantic coastal plain through the Mexican state of Veracruz
to San Luis Potosi. The species is well known from Chiapas
and the low Xucaneb on both sides of the Cuchumatanes,
but this is the first report of its occurrence in the Cuchu-
matanes proper. It occurs just below the cloud line at Finca
Chiblac in the northwestern Cuchumatanes.

HABITAT AND HABITS. This species almost always
occurs in the leaf bases of banana plants. All but one of the
four B. rufescens taken in the Cuchumatanes were found in
this situation. The exception was found crossing a trail on a
sultry, overcast afternoon at about 1 400 h. The species occurs
throughout the tropical zone forest from near sea level (Stuart,
1943a) up to the 1,200-m cloud line.

SYSTEMATICS. Specimens from the Cuchumatanes were
assigned to B. rufescens based upon derived skull and foot
structure and the absence of maxillary teeth.

Bradytriton silus Wake and Elias, 1983

Figure 9

Bradytriton silus Wake and Elias, 1983:3.

TYPE LOCALITY. Finca Chiblac, 15 km NE Barillas,
Huehuetenango, Guatemala, elevation 4,300 ft. (1,310 m).

RANGE. This species was discovered in the course of this
investigation. It is known only from the type locality.

HABITAT AND HABITS. Eleven of the twelve known
specimens were taken beneath pieces of wood in a clearing
that was being prepared for the cultivation of quinine. The
other individual was found under a log in the undisturbed
subtropical zone cloud forest surrounding the clearing. The
native forest at the type locality is a mixed hardwood growth
that receives over 6 m of rain yearly.

SYSTEMATICS. This species has unique specializations
and has been placed in a monotypic genus. Its possible re-
lationships are discussed by Wake and Elias (1983).

Dendrotriton cuchamatanus
(Lynch and Wake, 1975)

Figure 5

Chiropterotriton cuchumatanus Lynch and Wake, 1975:6.
Dendrotriton cuchumatanus: Wake and Elias, 1983.

TYPE LOCALITY. On highway 9N, 8.5 km (by road) SW
San Juan Ixcoy, Depto. Huehuetenango, Guatemala, eleva-
tion 2,860 m.

RANGE. This species is known only from the type locality.
Appropriate habitat is present elsewhere in the area, and this
species may range more widely than is presently known.

HABITAT AND HABITS. This species has been taken
under moss and bark on large fallen trees. The type locality
is a patch of great buttressed rain forest oaks on the Sub-

Contributions in Science, Number 348

Elias: Guatemalan Salamanders 1 1

Figure 9. Bradytriton silus, Pseudoewycea rex. and Nyctanolis pernix, life size. UPPER LEFT: Bradytriton situs, head, legs, and feet slate
grey, trunk and tail brick red-brown. LOWER LEFT: Pseudoewycea rex (Cuchumatan specimen), dark brown overall (in some Cuchumatan
specimens and in most Cuileo animals, there is light speckling, especially laterally). RIGHT: Nyctanolis pernix. background color black, spots
on eyelids and nape crimson, those on elbows and knees orange, trunk and tail spots yellow.

tropical-temperate zone border. This area receives a great
but undetermined amount of precipitation both in the form
of rain and as dense, cold fog.

SYSTEMATICS. The genus Dendrotriton has been re-
viewed by Lynch and Wake (1975). Their results suggest dose
relationship between D. cuchumatanus, D. bromeliacea, and

12 Contributions in Science, Number 348

Elias: Guatemalan Salamanders

20-

cn

c

0)

if)

c r>
CD

u

o

lo-

ci 3

A D cuchumatanus (WC)
3 D. rabbi (EC)

° " (WC)

;cu)

• %

• o

<D

C

a>

E

o

>

o

O OO AA A
A

O O A

O

rorViAAYx n A_

[— — r

50 60

Braincase width

T

~ I

70

Figure 10. Relationship between length of the preorbital process
of the vomer and braincase width in Dendrotriton rabbi and D.
cuchumatanus (in micrometer units). WC = Western Cuchuma-
tanes; EC = Eastern Cuchumatanes; CU = Cuilco.

D. megarhinus (the latter two are inhabitants of the Pacific
Uplands), based upon shared configurations of the vomer
and the nostril.

Dendrotriton rabbi (Lynch and Wake, 1975)

Figure 5

Chiropterotriton rabbi Lynch and Wake, 1975:2.
Dendrotriton rabbi: Wake and Elias, 1983.

TYPE LOCALITY. 9.5 km W, 8.5 km S (by air) La De-
mocracia, Depto. Fluehuetenango, Guatemala, elevation
2,100 to 2,500 m.

RANGE. This species, described and hitherto known only
from a spur of the Montanas de Cuilco, is here reported both
from the main massif of the Cuilco and from two localities
in the Cordillera de los Cuchumatanes.

HABITAT AND HABITS. Dendrotriton rabbi occur at
the subtropical-temperate border. In the Cuilco, they have
most frequently been found in bromeliads but only under
bark on logs and stumps in the Cuchumatanes, perhaps be-
cause bromeliads are absent from the appropriate elevations.
The species occurs from 2,100 to 2,700 m.

SYSTEMATICS. The newly discovered populations have
been referred to D. rabbi on the basis of a small nostril di-
ameter and the presence of a preorbital process on the vomer
in adults (as distinct from D. cuchumatanus, D. brome/iacea,
and D. megarhinus) and because some specimens from each
population have ossified septomaxillary bones (absent in D.
xolocalcae) (for discussion, see Lynch and Wake, 1975, 1978).

Lynch and Wake (1975) examined the ontogeny of the
nostril in all of the species of Dendrotriton and discovered
that all juveniles have similar, relatively large nostrils. They
found that patterns of relative growth determined the adult

condition; species that as adults have small nostrils experi-
ence either no growth or actual shrinkage in the absolute size
of the nostril through ontogeny, while large nostriled forms
experience nostril growth.

I have examined ontogenetic variation in the presence or
absence of the preorbital process of the vomer. The process
is large and tooth-bearing in small nostriled species but, if
present, is a nub in species with large nostrils. As with nostril
size, vomerine process length is initially equal in the large
nostriled D. cuchumatanus and small nostriled D. rabbi. Dif-
ferential growth produces the difference in the adult condition
in the two species (Fig. 10). In this case, interspecific differ-
ences are compounded by a differential in maximum adult
size. These data further indicate that the newly discovered
Cuchumatan populations clearly fall with the nominate D.
rabbi. With the new localities reported here, D. rabbi is known
to occur within 250 vertical m and about 25 km of D. cu-
chumatanus in the Cuchumatanes. The two species have been
taken in similar habitats without intervening ecological bar-
riers, and the proximity of the two suggests that the first case
of sympatry in the genus may yet be found.

Nyctanolis pernix Elias and Wake, 1983

Figure 9

Nyctanolis pernix Elias and Wake, 1983:2.

TYPE LOCALITY. Finca Chiblac, about 10 km NE Santa
Cruz Barillas, Depto. Huehuetenango, 1,370 m (4,500 ft.)
elevation.

RANGE. This species has been taken at the type locality,
just across the Mexican border in Chiapas and in the western
end of the Sierra de las Minas near Purulha, Baja Verapaz
(J. Campbell, pers. comm.).

HABITAT AND HABITS. The type series was found
under moss and bark during the day and on stumps and logs
on wet nights. The type locality is a region of quinine cul-
tivation in the subtropical zone, just above the cloud line.
The forest receives over 6 m of rain annually. The Mexican
specimen was taken in the mouth of a cave in the cloud
forest.

SYSTEMATICS. This species was discovered during the
course of this investigation and appears to have no close
relatives. The relationships of this monotypic genus were
discussed by Elias and Wake (1983).

Pseudoeurycea rex (Dunn, 1921)

Figure 9

Oedipus rex Dunn, 1921:143.

Pseudoeurycea rex: Taylor. 1944:209.

TYPE LOCALITY. Sierra Santa Elena (near Tecpam),
Guatemala, elevation 9,500 ft. (2,900 m).

RANGE. P. rex is known from the Cuchumatanes and
from several regions on the Pacific Uplands. The species is
here recorded for the first time from the Montanas de Cuilco.

HABITAT AND HABITS. This form occurs from 2,750
m elevation to over 4,000 m at the top of the highest moun-

Contributions in Science, Number 348

Elias: Guatemalan Salamanders 13

tains in its range and occurs in the temperate zone forest and
above the treeline in open bunchgrass communities. They
live under and within logs, under bark, moss, grass clumps,
and stones and were found active in the open on wet nights.
P. rex is often abundant, reaching densities higher than those
of any other Guatemalan salamander.

SYSTEMATICS. The Guatemalan members of the genus
Pseudoeurycea are four or five similar brown to black-colored
species. Species-level differences in foot webbing and color
pattern have proven difficult to document and tend to be
exceeded by intrapopulational variation. No careful quan-
titative work on variation in this assemblage has been pub-
lished, and my allocation of the Cuilco and Cuchumatan
populations is based upon their elevational distribution and
their general morphological similarities to known popula-
tions of P. rex.

DISCUSSION

SPECIES GROUPS IN BOLITOGLOSSA BETA

Since Wake and Lynch (1976) last outlined species groups
in the genus Bo/itog/ossa, examination of new material from
the Cuchumatanes, the Cuilco, the Pacific Uplands, and the
Sierra de las Minas has blurred certain distinctions and clar-
ified others among the species found in Nuclear Central
America (abbreviated NCA and including the entire land-
mass between the Isthmus of Tehuantepec and the Nicara-
guan Depression). Implicit in some of the comparisons made
in the preceding species accounts, and crucial to the discus-
sion sections to come, is a revision of the species groups for
Guatemalan Bolitoglossa.

Wake and Dresner (1967) found that a group of derived
structural characters involved in tail autotomy was shared
by certain species of Bolitoglossa. The derived group, referred
to by Wake and Lynch ( 1 976) as "Bolitoglossa beta,” includes
all Mexican and NCA species except the B. mexicana group,
as well as two forms from the Talamancan highlands further
south. Included are the following species: B. alvaradoi, B.
arboroscandens. B. cuchumatana, B. dofleini, B. dunni, B.
engelhardti, B. flavimembris, B. franklini, B. hartwegi, B.
helmrichi, B. lincolni, B. macrinii, B. melania, B. morio, B.
occidentalis, B. riletti. B. rostrata. B. rufescens, B. schmidti,
B. stuarti, B. veracrucis, B. yucatana. and at least one un-
described species (Wake and Lynch, 1982).

Species groups within Bolitoglossa beta were first recog-
nized informally. Taylor (1941), in his description of B. oc-
cidentalis. noted the affinity between his new species and B.
rufescens. Stuart (1943a) noted that B. dofleini and B. yu-
catana were close relatives. He also suggested that B. cuchu-
matana, B. dunni, B. engelhardti and B. helmrichi formed a
close group in terms of both morphology and ecology, and
in a later paper (1952) he named them the B. dunni species
group. In the description of B. lincolni Stuart (1943a) re-
ported the new species to be most closely affiliated with B.
franklini on morphological grounds, and later (Stuart, 1952)
included them with the then newly described B. nigroflaves-
cens in a B. franklini species group. Recently Wake and Lynch
( 1 982) reduced B. nigroflavescens to the status of a subspecies

within B. franklini. Stuart (1952) erected the B. morio group
to contain B. morio and B. omniumsanctorum, but Wake
and Elias (1983) synonymized the two.

By 1952, therefore, five species assemblages had been rec-
ognized within Bolitoglossa beta, leaving unassigned only the
following six of the nineteen species then known: B. flavi-
membris, B. macrinii, B. riletti, B. rostrata, B. schmidti and
B. veracrucis (see Table 3). Wake and Brame (1969) named
two new species, B. hartwegi and B. stuarti, and demonstrat-
ed their close morphological similarities to one another and
to B. veracrucis. In addition they reorganized Stuart’s ( 1 943a,

1 952) three species groups into two new ones, the B. rostrata
and B. helmrichi groups (Table 3). These two groups were
large and heterogeneous as compared to those described
above. The B. rostrata group included Stuart’s (1952) B.
morio and B. franklini groups, B. dunni and B. engelhardti
from his B. dunni group, the previously unassigned B. ma-
crinii, B. riletti and B. rostrata, and the recently described
species B. resplendens and B. brevipes. Their B. helmrichi
group included B. cuchumatana and B. helmrichi from
Stuart’s (1952) B. dunni group, the previously unassigned B.
flavimembris and B. veracrucis, and the new species B. hart-
wegi and B. stuarti.

Wake and Lynch (1976) assembled the first complete list-
ing of Bolitoglossa species groups. They tentatively assigned
every species to some group and recognized a total of six
species groups within Bolitoglossa beta. The B. dofleini group
was erected to include B. dofleini, B. yucatana and the poorly
known B. schmidti. The B. rufescens group was recognized
as containing B. occidentalis and B. rufescens. The two Tala-
mancan species from southern Central America, B. alvaradoi
and B. arboroscandens, were placed together in a B. alvaradoi
group, and the two species from north of the Isthmus of
Tehuantepec, B. macrinii and B. riletti, were put in a B.
macrinii group. The two large groups, B. rostrata and B.
helmrichi, of Wake and Brame (1969), were retained with
their original membership but for the switching of B. mono
from one to the other and the inadvertent omission of B.
omniumsanctorum altogether (see Table 3).

The recognition of polyphyletic groups (groups united by
characters not shared by their most recent common ancestor)
is a severe obstacle to phylogenetic reconstruction. Ensuring
that only nonpolyphyletic (= paraphyletic and monophyletic)
species groups are recognized is a first step towards an un-
derstanding of the phylogenetic history of Bolitoglossa beta.
To eliminate possibly polyphyletic groups I have recognized
only tight clusters of phenotypically similar species. Mor-
phological similarity between species group members is dem-
onstration of nonpolyphyletic origin of the group insofar as
such similarities are homologous (acquired by descent from
a similarly endowed common ancestor).

The B. rostrata and B. helmrichi groups of Wake and Lynch
(1976) are large, heterogeneous and united by no specific
characters. To compose tighter units more comparable to
other species groups in Bolitoglossa beta I have subdivided
and reorganized these two groups.

The B. rostrata group contains a series of species that has
been recognized as a phenetic cluster by many authors (Stuart,

14 Contributions in Science, Number 348

Elias: Guatemalan Salamanders

Table 3. The species groups of Bolitoglossa beta as defined by different authors. Species not assigned to a group until after 1952 (both newly described and described but previously
unaffiliated species) are indicated by an asterisk where they first appear.

Early classifications Wake and Brame, 1969 Wake and Lynch, 1976 Elias, 1983

Contributions in Science, Number 348

Elias: Guatemalan Salamanders 15

group I arbor oscande ns

1943a, 1952, as his B. frank/ini group; McCoy and Walker,
1966, Wake and Brame, 1969, and Wake and Lynch, 1976,
as the B. lincolni subgroup of their B. rostrata group; Wake,
Yang, and Papenfuss, 1980, and Wake and Lynch, 1982, as
the B. franklini species group). This assemblage includes B.
franklini, B. lincolni. B. melania, and one undescribed Chia-
pan form (Wake and Lynch, 1982, who also include B. re-
sp/endens). These species are characterized by large adult size,
stout bodies and tails, and a black ground color broken dor-
sally only by a sharply bounded swath, spotting, or reticu-
lation in some lighter color; red in some species to yellow or
grey-green in others (see B. lincolni. Fig. 3). B. melania is
solid black. These species all have toetips free of webbing
and subdigital pads. They are distributed allopatrically ex-
cept for an area of elevational parapatry with occasional
hybridization between the high elevation B. lincolni and low-
er elevation B. franklini on the Pacific Uplands (Wake, Yang,
and Papenfuss, 1980).

The remainder of the B. rostrata group of Wake and Lynch
(1976), including B. dunni, B. engelhardti and B. rostrata.
fits with B. cuchumatana and B. helmrichi of their B. helm-
richi group to form an assemblage characterized by blunt
rounded toetips, fully developed subdigital pads, and a dark
brown ground color, frequently marked with a lighter brown
dorsal swath or paired shoulder stripes (see B. rostrata in
Fig. 3 for typical color pattern and compare B. rostrata and
B. cuchumatana [inset] feet as examples of variation in foot
morphology within the group). Stuart (1943a, 1952) recog-
nized this assemblage as his B. dunni group but hesitated to
include B. rostrata because of its different ecology. Relying
on morphological similarities, I recognize this assemblage,
including B. rostrata, as a new B. dunni species group. The
members of the B. dunni group are all either geographically
or elevationally (in the case of B. rostrata and B. engelhardti
in the Pacific Uplands) allopatric, except that narrow sym-
patry is seen between the high elevation B. rostrata and the
lower B. cuchumatana in the Cuchumatanes.

Bolitoglossa hartwegi. B. stuarti, and B. veracrucis were
recognized by Wake and Brame (1969) to form a compact
subgroup within their B. helmrichi group. I choose to rec-
ognize the similarities of these species by isolating them in
a new B. veracrucis species group. The three species are united
by the combination of fully webbed feet that lack subdigital
pads and are differentiated from members of the B. rufescens
group by posession of a strongly articulated skull with a broad
pars dentalis of the premaxilla. The B. veracrucis group has
the broadest elevational range of any group of NCA sala-
manders (see elevational distribution section below), and no
two species occur sympatrically.

The two remaining species left over from Wake and Lynch’s
(1976) B. helmrichi group are B. flavimembris and B. morio.
B. morio is slightly larger than B. flavimembris, but the two
share a stout body and tail and a unicolor dorsum (see B.
morio in Fig. 3). That B. flavimembris is not affiliated with
the B. dunni group is indicated both by its coloration and
build and by its rather pointed toetips and poorly developed
subdigital pads (see Wake and Lynch, 1976, fig. 22). New
evidence that B. morio and B. flavimembris are relatives

comes from the discovery of populations of intermediate
morphology in the Pacific Uplands of Guatemala near the
Mexican border (D.B. Wake, pers. comm.). Given these in-
dications of relationship, and the absence of other affilia-
tions, I group B. flavimembris and B. morio together in a
new B. morio species group.

Aside from the changes proposed above I follow the species
group assignments of Wake and Lynch (1976); the full history
of species groups among the species of Bolitoglossa beta
including my revisions is presented in tabular form in
Table 3.

PATTERNS OF DISTRIBUTION AND
DIFFERENTIATION

Faunal Connections Between the Mexican Plateau,

NCA, and the Talamancas

Wake and Lynch (1976) found that the three most moun-
tainous areas of Middle America are especially high in sal-
amander species diversity; these are the Caribbean escarp-
ment of the Mexican Plateau, the mountains of NCA, and
the Talamancan highlands of Costa Rica and Panama. That
there has been faunal contact between these centers of di-
versity is demonstrated by the distribution of some genera
on more than one of the three upland areas. For example,
the NCA fauna includes some genera that range to the Mex-
ican Plateau ( Pseudoeurycea ), others that range to the Tala-
mancas ( Oedipina and Nototriton ), still others that occur on
all three upland areas (Bolitoglossa), as well as genera en-
demic to NCA ( Bradytriton , Dendrotriton, and Nyctanolis).
Each genus has a clear center of intrageneric diversity on one
of the three upland regions (except Bolitoglossa, in which the
alpha and beta subgeneric groups center on two different
uplands (Wake and Lynch, 1976)). This pattern suggests that
most intrageneric evolution proceeded on single upland areas
at a time when the three major uplands were isolated from
one another.

If we go one step further back, however, and consider the
origins of the genera themselves, we find that genera with
distributions centered on the same upland area (i.e., Bradytri-
ton, Dendrotriton, Nyctanolis, and the beta complex of Bo-
litoglossa in NCA) do not usually form a clearcut monophy-
letic group (Wake and Elias, 1983). This implies a complex
history of ancient faunal contact between the three great up-
land regions of Middle America.

Faunal Connections Among the Mountain Ranges of NCA

Within NCA, the uplands form discrete islands of habitat
for high elevation salamander species. Between these uplands
lie warmer, drier, rain-shadowed valleys that are generally
uninhabited by salamanders. Around the coastward periph-
ery of these mountains are warm, humid lowlands that sup-
port a restricted plethodontid fauna. Most of the salamanders
occurring in NCA are montane (including Bradytriton, Den-
drotriton, Nototriton, Nyctanolis, and Pseudoeurycea). A sin-
gle genus, Oedipina, is restricted to the lowlands. Only Bolito-
glossa occurs at all elevations, and even this genus is split

16 Contributions in Science, Number 348

Elias: Guatemalan Salamanders

1000

CUILCO

CUCHUMATANES

Figure 1 1 . Elevational distribution of salamander species on the Montanas de Cuilco and the Cordillera de los Cuchumatanes (all collections
combined).

into upland ( B . helmrichi, B. morio, B. franklini) and low-
land ( B . mexicana, B. dofleini, B. rufescens) species assem-
blages (the B. veracrucis group is exceptional in its broad
elevational range and will be discussed separately below).
The upland and lowland faunas intermingle narrowly in the
low cloud forest at about 1 ,000 m elevation, but the upland
groups are subject to special biogeographic limitations, which
give their current patterns of differentiation particular his-
torical interest.

Comparison of patterns of upland diversity obtaining in
the Cuchumatanes relative to the Pacific Uplands raises in-
teresting contrasts. There are fifteen upland salamander species
known from the Pacific Uplands (Wake and Lynch, 1976,
1982) and only nine (excluding B. hartwegi of the B. vera-
crucis group) from the Cuchumatanes, but the supraspecific
diversity is greater in the Cuchumatanes. The nine Cuchu-
matan species belong to five genera, and the four Bolitoglossa
fall into three species groups. In the Pacific Uplands, how-
ever, the fifteen species are in three genera, and all seven
Bolitoglossa are in the same three species groups that occur
in the Cuchumatanes. No genus or species group in the Pacific
Uplands is not also represented in the Cuchumatanes, but
two genera present in the Cuchumatanes are absent in the
Pacific Uplands. The two areas have four species in common:

Bolitoglossa lincolni (here including B. resplendens), B. mo-
rio, B. rostrata, and Pseudoeurycea rex.

The fact that salamander diversity is so deep in the Cu-
chumatanes may reflect that region's ancient history as a
land-positive area (Rosen, 1978).

The shallowness of salamander diversity on the Pacific
Uplands may reflect the topography of that area: the high-
lands are composed of many abutting cinder cones. Small
climatic fluctuations should have caused alternate fragmen-
tation and reunion of the geographic ranges of highland sal-
amander species. When a species’ range is subdivided and
then reunified after a period in this way, a complex of in-
trogressions, parapatries and sympatries might be expected
to result as partially differentiated sister populations are
thrown back into contact after a period of allopatry on neigh-
boring cinder cones. Just these sorts of interactions are found
among the Pacific Uplands salamanders (Wake and Lynch,
1976; Wake, Yang, and Papenfuss, 1980; Wake and Lynch,
1982).

ELEVATIONAL DISTRIBUTION

Schmidt (1936) documented the elevational ranges of sala-
mander species along a transect down the coastal escarpment

Contributions in Science, Number 348

Elias: Guatemalan Salamanders 17

of the Pacific Uplands. He found that species there inhabited
narrow, sharply bounded elevational ranges and that a given
species typically inhabited similar elevations at different
transect sites along the Pacific escarpment.

Wake and Lynch (1976) performed a detailed analysis of
elevational distribution along a transect near the volcano first
studied by Schmidt. They corroborated Schmidt’s results and
filled them out with detailed studies of elevational boundary
zones and possible interspecific competition.

Composite transects (grouping all collections) for the Cu-
chumatanes and Cuilco (Fig. 1 1) compared to the transect
results of Wake and Lynch (1976) for Volcan Tajumulco in
the Pacific Uplands indicate that related populations (either
conspecifics or members of related species) show similar el-
evational ranges wherever they occur. Many of the related
populations that have significant variation in elevational dis-
tribution between transects appear to vary in response to
geographic variation in the elevation of the cloud line.

The results for the Cuchumatanes and Cuilco reinforce the
generalization that salamander populations in NCA inhabit
narrow elevational belts, and recent information for all up-
lands in NCA indicates that with one exception no single
species or group of related species (meaning species groups
in Bolitoglossa and entire genera in other cases) occurs over
an elevational range as broad as 1,300 m. The exception is
Bolitoglossa hartwegi (with a 1 ,500-m elevational range) and
the Bolitoglossa veracrucis species group with a combined
elevational range of over 2,800 m. As Wake and Brame
(1969) have pointed out, and as these results reiterate, this
exceptional group may have played a key role in the invasion
of the lowlands by Bolitoglossa beta and must have unique
physiological attributes, which, if studied, might expand our
understanding of the history of salamanders throughout Mid-
dle America.

SPECIMENS EXAMINED

Salamanders are known from the following localities in the
Cordillera de Los Cuchumatanes, Depto. Huehuetenango,
and the Sierra de Cuilco, Depto. Quiche, Guatemala.

Bolitoglossa cuchumatana. CUCHUMATANES. Finca
Chiblac.ca. 10 km (airline) NE Santa Cruz Barillas, el. 1,300-

l, 500 m, 3 km S of buildings, MVZ 134524-72, 134574-
79, 134582-88, LACM 135482-85; El Porvenir, 134603-
lb; Finca Chiblac, 149306-14; 1.5 km S of buildings, 134591-
602; San Isidro; 134617-22; El Rayo, 134623-25. On ridge
ca. 4 km (airline) NW Santa Cruz Barillas, el. 2,000 m, MVZ
149315.

CUILCO. 6.7 mi. (rd.) S Nebaj on Rta. Depto. 3, el. 2,500

m, MVZ 160993-1005. On ridge 2.5 km W, 2 km N (airline),
San Miguel Uspantan, el. 2,200-2,500 m, MVZ 149301-05.
Oak forest about 2 km (airline) N Nebaj, el. 1,900 m, UMMZ
89110 (holotype), 8911 1-13.

Bolitoglossa hartwegi. CUCHUMATANES. Finca Chi-
blac, ca. 10 km (airline) NE Santa Cruz Barillas, el. 1,300-

1.500 m, MVZ 134629, 134631-33, LACM 135481; 3 km
S of buildings, MVZ 1 49324. 3 km (airline) NNW San Mateo
Ixtatan, el. 2,750 m, MVZ 160355-59.

CUILCO. On ridge 2.5 km W, 2 km (airline) N San Miguel
Uspantan, el. 2,200-2,500 m, MVZ 149325.

Bolitoglossa jacksoni. CUCHUMATANES. Finca Chi-
blac, ca. 12 km (airline) NE Santa Cruz Barillas, el. 1,400
m, MVZ 134634 (holotype).

Bolitoglossa lincolni. CUCHUMATANES. Just N San Juan
Ixcoy, UMMZ 123275-76. Ca. 9 mi. (rd.) SW El Reposo,
summit between El Reposo and Oja Blanca, el. 2,200-2,300
m, MVZ 161791, 119011. Planes de Pena Blanca, 1.5 km
(airline) NE Pena Blanca peak, el. 2,700 m, MVZ 149359-
64; 9.5 km W, 8.5 km (airline) S, La Democracia, el. 2,100-

2.500 m, MVZ 103834-38; 10 km W, 143681-87. Paraiso
Hoja Blanca Cumbre, UMMZ 129149-50.

CUILCO. Monte at Salquil Grande, el. 2,450 m, UMMZ
89107 (holotype), 89108-09. On ridge NW San Miguel Us-
pantan, el. 2,200-2,500 m, 2.5 km N Uspantan, MVZ
149365-69; 3.5 km (air) NNW, 160833-39.

Bolitoglossa morio. CUCHUMATANES. Planes de Pena
Blanca, 1.5 km (airline) NE Pena Blanca peak, el. 2,700 m,
MVZ 149335-43. Todos Santos, el. 2,500 m, UMMZ
102285-86 (holotype and paratype of Oedipus omnium-
sanctorum).

Bolitoglossa mulleri. CUCHUMATANES. Town of Santa
Cruz Barillas, el. 1,500 m, MVZ 131691-93, 149327-31,
161 102. Finca la Florida ca. 2 mi. (rd.) NE Santa Cruz Ba-
rillas, el. 1,500 m, MVZ 149326.

Bolitoglossa rostrata. CUCHUMATANES. Ca. 37-40 km
(rd.) N Huehuetenango, el. 2,860 m, MVZ 11341 8-540. Vi-
cinity Capzin, Rta. Nac. 9N, 50-60 km (rd.) N Huehuete-
nango, el. 2,800-3,200 m. 52.7 km N, MVZ 114722- 65; ca.
50 km N, km post 139, 115494-524, 117039. 2-8 km N,
W, and S (airline) San Mateo Ixtatan, el. 2,700-3,200 m; 2
km WSW, MVZ 150940-160299; 6 km WSW, 160300-333;
5 km SSW, 160334-49; 7 km SSW, 160350-51; 3 km NNW.
160353, 160360; 4 km NW, 163928-31, 163973; 2 km W,
171198-202; 4 km SW, 171220-41; 4 km N, 171242; 4 km
W, 171173-97; 4 km NW, 171112-43; 5 km SW, 171215;
5 km S, 171203-14; 6 km W, 171163-72; 8 km W, 171144-
62. Paquix-Todos Santos road, 4-7 km SE Todos Santos,
MVZ 149198-206, 108616-30, 108631-72. 13.6 mi. (rd.)N
Santa Eulalia along Rta. Nac. 9N, MVZ 1 49228. 5. 1 mi. (rd.)
N Santa Eulalia along Rta. Nac. 9N, MVZ 108673-95,
109296. Cumbre between Ixtiapoc and Villa Linda, el. 2,800-
3,400 m, UMMZ 120007-08. Cumbre between Yaiquich
and San Mateo Ixtatan, el. 3,000 m, UMMZ 120009. Todos
Santos, el. 9,000-10,000 ft., UMMZ 120465-66, 120487.
Mina Villa Linda, UMMZ 130073. 9 km (rd.) SW San Juan
Ixcoy, el. 2,900 m, UMMZ 120011. Rta. Nac. 9N, 7.4 mi.
(rd.) SW San Juan Ixcoy, MVZ 108696-728. Rta. Nac. 9N,
10.6 mi. (rd.) SW San Juan Ixcoy, MVZ 108729-32. 8 km
W San Mateo Ixtatan, el. 3,100 m, MVZ 171 148-62. 3 km
NNW San Mateo Ixtatan, el. 8,700 ft., MVZ 150940-1000,
160201-91, 160353, 160360; 4 km NW, 2,750 m, 171112-
43. 2 km WSW San Mateo Ixtatan, el. 8,850 ft., MVZ 160292-

18 Contributions in Science, Number 348

Elias: Guatemalan Salamanders

99; 2 km W, 2,750 m, 1 7 1 198-202. 6 km WSW San Mateo
Ixtatan, el. 9,150-9,450 ft., MVZ 160330-33, 160352,
160354; 6 km W, 171 163-72. 5 km SSW San Mateo Ixtatan,
el. 8,650-9, 1 50 ft., MVZ 1 60343-49; 4 km SW, 2,835-2,965
m, 171220-41; 4 km W, 2,900 m, 171 173-97; 5 km SW,
2,965-3,045 m, 171215-19; 5 km S, 3,150 m, 171203-14.
7 km SSW San Mateo Ixtatan, el. 9,650 ft., MVZ 160350-
51.4 km NW San Mateo Ixtatan, el. 2,750 m, MVZ 1 63928—
31, 163973; 4 km N, 2,650 m, 171242.

CUILCO. Above Salquil Grande, el. 3,000 m, UMMZ
891 16-18. 6.7 mi. (rd.) S Nebaj by Rta. Depto. 3, el. 2,500
m, MVZ 160992. On ridge 2.5 km W, 2 km (airline) N San
Miguel Uspantan, el. 2,260-2,640 m, MVZ 149229-300.

Bolitoglossa rufescens. CUCHUMATANES. Vicinity Fin-
ca Chiblac, ca. 10 km (airline) NE Santa Cruz Barillas, el.
950-1,100 m, MVZ 134626-28.

Bolitoglossa rufescens. CUCHUMATANES. Vicinity Fin-
ca Chiblac, ca. 10 km (airline) NE Santa Cruz Barillas, el.
950-1,100 m, MVZ 134626-28.

Bolitoglossa stuarti (tentative identification). CUCHU-
MATANES. 15 km (airline) W La Democracia, el. 1,850 m,
UMMZ 126781.

Bradytriton silus. CUCHUMATANES. Finca Chiblac, 15
km NE Barillas, el. 4,300 ft. (1,3 10 m), MVZ 131586, 131587
(holotype), 131589-94, 134635-37, 134638, 1 73063-64;
LACM 134566.

Dendrotriton cuchumatanus. CUCHUMATANES. Along
Rta. Nac. 9N, 8.5 km (rd.) SW San Juan Ixcoy, el. 2,860 m,
MVZ 113002 (holotype), 1 13003-05, 113007-19, 113021-
22; LACM 105296-97.

Dendrotriton rabbi. CUCHUMATANES. Planes de Pena
Blanca, 1.5 km (airline) NE Pena Blanca peak, el. 2,700 m,
MVZ 105298-300, 149702-05, 149139. 2 km (airline) WSW
San Mateo Ixtatan, el. 8,850 ft., MVZ 160437. 3 km (airline)
NNW San Mateo Ixtatan, el. 2,750 m, 160422-36. Cloud
forest above summit of road between El Reposo and Oja
Blanca ca. 9 mi. by rd., SW El Reposo, el. 2,200-2,300 m,
MVZ 109297-301.

CUILCO. 9.5 km W, 8.5 km S (airline), La Democracia,
Montanas de Cuilco, el. 2,100-2,500 m, MVZ 1 14766-96,
143920-26, 103839 (holotype), 103840-78, LACM 105298-
300. On ridge 2.5 km W, 2 km (airline) N, San Miguel Us-
pantan, el. 2,400 m, MVZ 160895-96.

Nyctanolis pernix. CUCHUMATANES. Finca Chiblac, 10
km (air) NE Barillas, el. 1,370 m (4,500 ft.), MVZ 131583-
85, 134639-40, 134641 (holotype), 134642-44, 149370-73;
MCZ 100154.

Pseudoeurycea rex. CUCHUMATANES. Vicinity El Re-
tiro and Pena Blanca peak, el. 2,800-3,300 m, MVZ 149404-
33, 149464-520, 149522-58, 149693-700. Vicinity Capzin,
Rta. Nac. 9N, 50-60 km (rd.) N Huehuetenango, el. 2,900-
3,300 m, MVZ 160973-82, 134211-401.2-8 km N, W, and
S (airline) San Mateo Ixtatan, el. 2,750-3,200 m; 5 km, MVZ
171243-52; 7 km SSW, 160410-19; 6 km WSW, 160409; 4
km NW, 167796-803. Between Cumbre Yaiquich and San
Mateo Ixtatan, 3,000 m, UMMZ 120067. 10 km (rd.) SW
San Juan Ixcoy, el. 3,300 m, UMMZ 120068.

CUILCO. Above Salquil Grande, el. 3,000 m, UMMZ
89114-15.

ACKNOWLEDGMENTS

I dedicate this work to the memory of Lie. Mario Dary, whose
assistance during the field work was invaluable, and whose
killing was tragic and senseless.

Dr. L.C. Stuart was generous with his hospitality and un-
surpassed knowledge of Guatemala’s herpetofauna. The res-
idents of the departments of Huehuetenango and El Quiche
were universally tolerant of behavior that to them appeared
absurd and were friendly and hospitable at all times. The
staffs of various fincas, especially R. Klein-Holkenborg and
Antonio Diaz of Finca Chiblac, were very cooperative.

A variety of other collectors assisted me, either directly in
the field or indirectly through the sharing of experience and
specimens. Among these are J.L. Jackson, J. Johnson, E.J.
Koford, J.F. Lynch, T.J. Papenfuss. R. Seib, H.B. Shaffer,
L.C. Stuart, R.T. Tesucun, and D.B. Wake.

The students and staff of the Museum of Vertebrate Zo-
ology provided an atmosphere for discussion and research,
from which I profited immensely. Dr. David B. Wake ex-
pedited every aspect of this research. He guided the inves-
tigation through its entire development, provided space and
resources in Berkeley, and was the source of most field sup-
port (current grant NSF DEB 78-03008).

I should finally like to thank Gene M. Christman and
James Hendel of the University of California for their expert
figure preparation and K.F. Liem and F.A. Jenkins, Jr., of
Harvard University, under whose auspices I received support
and travel funds (on NIH Musculo-Skeletal Training Grant
#5 T32 GMO 71 17-04 and -05) during the preparation of
this manuscript.

The Spanish summary was prepared by Fabian Jaksic.

LITERATURE CITED

Boulenger, G.A. 1882. Cat. Batrach. Grad. British Mu-
seum, London. 127 pp., 9 plates.

Brocchi, P. 1 883. Etude des batraciens de I’Amerique Cen-
trale. Mission Scientifique au Mexique et dans l’Ame-
rique Centrale 3(2): 1-1 22, 21 plates.

Cope, E.D. 1868. Sixth contribution to the herpetology of
tropical America. Proc. Acad. Natur. Sci. Philadelphia
1868:305-313.

. 1869. A review of the species of the Plethodontidae

and Desmognathidae. Proc. Acad. Natur. Sci. Philadel-
phia 21:93-118.

Duellman, W.E. 1963. Amphibians and reptiles of the rain-
forests of southern El Peten, Guatemala. Univ. Kansas
Publ. Mus. Natur. Hist. 1 5(5):205-249.

Dunn, E.R. 1921. Two new Central American salamanders.
Proc. Biol. Soc. Washington 34:143-145.

. 1924. New salamanders of the genus Oedipus with

a synoptical key. Field Mus. Natur. Hist. Zool. Ser. 12:
95-100.

Contributions in Science, Number 348

Elias: Guatemalan Salamanders 19

. 1926. The salamanders of the family Plethodon-

tidae. Smith College, Northampton, Mass., viii + 441

pp.

Elias, P., and D.B. Wake. 1983. Nyctanolis pernix, a new
genus and species of plethodontid salamander from
northwestern Guatemala and Chiapas, Mexico. Pages 1-
12 in A. G. J. Rhodin and K. Miyata (eds.). Advances
in herpetology and evolutionary biology: Essays in honor
of Ernest E. Williams. Mus. Comp. Zool., Cambridge,
Mass.

Hanken, J., J.F. Lynch, and D.B. Wake. 1980. Salamander
invasion of the tropics. Natur. Hist. 89(1 2):46— 53.

Lynch, J.F., and D.B. Wake. 1975. Systematics of the Chi-
ropterotriton bromeliacea group (Amphibia: Caudata),
with description of two new species from Guatemala.
Los Angeles Co. Natur. Hist. Mus., Contrib. Sci. 265:
1-45.

. 1978. A new species of Chiropterotriton (Am-
phibia: Caudata) from Baja Verapaz, Guatemala, with
comments on relationships among Central American
members of the genus. Los Angeles Co. Natur. Hist.
Mus., Contrib. Sci. 294:1-22.

McCoy, C.J., and C.F. Walker. 1966. A new salamander
of the genus Bolitoglossa from Chiapas. Occ. Papers Mus.
Zool. Univ. Mich. 649:1-1 1.

Rosen, D.E. 1978. Vicariant patterns and historical expla-
nation in biogeography. Syst. Zool. 27:159-188.

Schmidt, K.P. 1933. New reptiles and amphibians from
Honduras. Zool. Ser. Field Mus. Nat. Hist. 322:15-22.

. 1936. Guatemalan salamanders of the genus Oe-
dipus. Zool. Ser. Field Mus. 20:135-166.

Smith, H.M. 1945. Herpetological collecting in banana fields
of Mexico. Ward’s Natur. Sci. Bull. 1:1-7.

Smith, W.H. 1877. The tailed amphibians. Thesis, Mich-
igan Univ.

Stuart, L.C. 1943a. Taxonomic and geographic comments
on Guatemalan salamanders of the genus Oedipus. Misc.
Publ. Mus. Zool. Univ. Mich. 56:1-33.

. 1 943b. Comments on the herpetofauna of the Sier-
ra de los Cuchumatanes of Guatemala. Occ. Papers Mus.
Zool. Univ. Mich. 471:1-29.

. 1948. The amphibians and reptiles of Alta Verapaz,

Guatemala. Misc. Publ. Mus. Zool. Univ. Mich. 69: 1 —
109.

. 1950. A geographic study of the herpetofauna of

Alta Verapaz, Guatemala. Contrib. Lab. Vert. Biol. Univ.
Mich. 45:1-77.

. 1952. Some new amphibians from Guatemala. Proc.

Biol. Soc. Washington 65:1-12.

Taylor, E.H. 1941. New amphibians from the Hobart M.
Smith Mexican Collections. Univ. Kansas Sci. Bull. 28:
141-167.

. 1944. The genera of plethodont salamanders in

Mexico, Pt. 1. Univ. Kans. Sci. Bull. 30:189-232.

Taylor, E.H. , and H.M. Smith. 1945. Summary of the col-
lections of amphibians made in Mexico under the Walter
Rathbone Bacon Traveling Scholarship. Proc. U.S. Natl.
Mus. 95:521-613.

Wake, D.B., and A.H. Brame, Jr. 1963. The status of the
plethodontid salamander genera Bolitoglossa and Mag-
nadigita. Copeia 1963:382-387.

. 1 969. Systematics and evolution of neotropical sal-
amanders of the Bolitoglossa helmrichi group. Natur.
Hist. Mus. Los Angeles County, Contrib. Sci. 1 75: 1-40.

Wake, D.B., and I.G. Dresner. 1967. Functional mor-
phology and evolution of tail autotomy in salamanders.
J. Morph. 122:265-306.

Wake, D.B., and P. Elias. 1983. New genera and a new
species of Central American salamanders, with a review
of the tropical genera (Amphibia, Caudata, Plethodon-
tidae). Natur. Hist. Mus. Los Angeles County, Contrib.
Sci. 345:1-19.

Wake. D.B., and J.F. Lynch. 1976. The distribution, ecol-
ogy, and evolutionary history of plethodontid salaman-
ders in tropical America. Bull. Natur. Hist. Mus. Los
Angeles Co. 25:1-65.

. 1982. Evolutionary relationships among Central

American salamanders of the Bolitoglossa franklini
group, with a description of a new species from Gua-
temala. Herpetologica 38:257-272.

Wake, D.B.. S.Y. Yang, and T.J. Papenfuss. 1980. Natural
hybridization and its evolutionary implications in Gua-
temalan plethodontid salamanders of the genus Bolito-
glossa. Herpetologica 36:335-345.

Submitted 6 October 1981; accepted 23 August 1983.

20 Contributions in Science, Number 348

Elias: Guatemalan Salamanders

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SYSTEMATICS OF XANTUSIID LIZARDS
OF THE GENUS LEPIDOPH YMA
IN NORTHEASTERN MEXICO

Robert L. Bezy

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Printed at Alien Press. Inc.. Lawrence. Kansas

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.

SYSTEMA TICS OF XANTUSIID LIZARDS
OF THE GENUS LEPIDOPHYMA
IN NORTHEASTERN MEXICO

Robert L. Bezy1

ABSTRACT. Discriminant analyses of variation among 30 scale
characters indicate that the 21 populations of Lepidophyma from
northeastern Mexico form six morphological groups. Two of 27
population samples from southern Mexico approach the northern
groups in discriminant space. When viewed in terms of univariate
differences and geographic distribution, the northern population
groups constitute four unique morphological entities that are con-
sidered to represent species units. The most distinctive is Lepido-
phyma gaigeae occurring in limestone habitats in the Sierra Madre
Oriental of Hidalgo and Queretaro. Lepidophyma occulor is known
from four localities in the semi-arid Jalpan region of Queretaro and
San Luis Potosi, and the cavemicolous L. micropholis is confined
to the Sierra del Abra of Tamaulipas and San Luis Potosi. The wide-
ranging L. sylvaticum includes four moderately divergent population
groups: northern Madrean (Tamaulipas to San Luis Potosi), southern
Madrean (San Luis Potosi to Hidalgo), Veracruzan, and western
(Mesa Central of San Luis Potosi to Nuevo Leon).

The karyotypes of L. gaigeae and L. occulor are unique within
the genus, while most L. sylvaticum are chromosomally identical to
L. micropholis. A heteromorphism in microchromosomes was ob-
served in six females of one population of L. sylvaticum. and could
represent either ZW sex chromosomes or allodiploidy. This same
population plus one in Queretaro have statistically significantly skewed
sex ratios that may be associated with hybridization.

RESUMEN. Los analisis discriminatorios de variacion entre 30
caracteres de las escamas, indican que las 2 1 poblaciones de Lepi-
dophyma del noreste de Mexico forman seis grupos morfologicos.
Dos de los muestreos de la poblacion del sur de Mexico se aproximan
a los grupos del norte en espacio discriminatorio. Cuando se ex-
aminan en terminos de diferencias univariadas y de distribucion
geografica, los grupos de la poblacion del norte constituyen cuatro
entidades morfologicas unicas que se considera representan unidades
de especie. La mas distintiva es Lepidophyma gaigeae que vive en
habitats de piedra caliza en la Sierra Madre Oriental de Hidalgo y
Queretaro. Lepidophyma occulor se conoce de cuatro localidades de
la region semiarida de Jalpan de Queretaro y San Luis Potosi, y el
cavemicola L. micropholis se encuentra confinado a la Sierra del
Abra de Tamaulipas y San Luis Potosi. L. sylvaticum de amplia
distribucion en y cerca de la Sierra Madre Oriental incluye cuatro
grupos poblacionales moderadamente divergentes: norte (Tamau-
lipas a San Luis Potosi), sur (San Luis Potosi a Hidalgo), Veracruz,
y occidental (Mesa Central de San Luis Potosi a Nuevo Leon).

Los cariotipos de L. gaigeae y L. occulor son unicos dentro del
genero mientras que la mayoria de L. sylvaticum son cromosomi-
camente identicos a L. micropholis. Se observo heteromorfismo de
microcromosomas en seis hembras de una poblacion de L. sylva-
ticum y podria representar ya sea cromosomas sexuales ZW o alo-
diploidia. Esta misma poblacion, mas una en Queretaro han torcido
estadisticaniente en forma significativa las proporciones en los sexos
que pudieran ser asociadas con hibridizacion.

INTRODUCTION

Lizards of the xantusiid genus Lepidophyma range from Pan-
ama to Nuevo Leon, Mexico, living principally in wet trop-

Contributions in Science, Number 349, pp. 1-16
Natural History Museum of Los Angeles County, 1984

ical lowland forests in the south, but becoming increasingly
restricted to montane and/or rimose habitats in the semi-
arid regions to the north. In the rugged ranges of the Sierra
Madre Oriental, and in the canyons and valleys along both
of its flanks, are a morphologically diverse array of Lepi-
dophyma populations. While most of these populations re-
main taxonomically unallocated, four have been named, and
two of these names have been alternatively associated with
species occurring to the south (e.g.. Smith, 1942; Walker,
1955).

In this paper, the problems of discordant morphological
variation, geographic isolation, and small sample sizes of the
populations of Lepidophyma in northeastern Mexico are han-
dled by treating each locality as a separate sample, and em-
ploying multivariate analyses of variation to identify groups
of morphologically similar populations. Additional multi-
variate comparisons with populations to the south, and anal-
yses of univariate differences among all population groups
are used to diagnose morphological species. Names are then
assigned to the units on the basis of included type or topotypic
material, the species of Lepidophyma recognized in north-
eastern Mexico are summarized in brief accounts, including
comments on chromosomal variation and skewed sex ratios
in certain populations, and a key is presented.

MATERIALS AND METHODS

A total of 1 52 specimens of Lepidophyma from Mexico north
of 19°N latitude were used in the analyses. This includes all
material studied from the area, except that referable to L.
gaigeae. One sample (N = 20) of the over 500 known spec-
imens of the species was used as a reference population. In
addition, 31 population samples (N = 188) from southern
and western Mexico were utilized in the comparative anal-
yses. The selection of 19°N latitude as the southern limit of
the study area is based on a distributional hiatus for the genus
in the transvolcanic region (ca. 1 9-20°N), and on preliminary
observations suggesting that the populations occurring to the
north of this distributional gap share a number of unique
morphological similarities.

The localities of the specimens were determined on avail-
able maps, and geographic samples were constituted with all
specimens from a given locality (or in a few instances by
pooling adjacent localities separated by less than 20 km) to
form a total of 2 1 population samples of Lepidophyma from
northern Mexico (Fig. 1 ). The specimens and localities are
listed in specimens examined, below.

1 . Section of Herpetology, Natural History Museum of Los An-
geles County, 900 Exposition Boulevard, Los Angeles, California
90007.

ISSN 0459-8113

The analyses use a total of 30 scale characters, 20 meristics,
and 1 0 ratios of the relative size or proportions of individual
scales. No significant sexual dimorphism, ontogenetic vari-
ation, or correlation was detected among the 30 characters.
The characters were selected largely on the basis of their
purported diagnostic strength in the genus (Bezy, 1973; Bezy
et al., 1982; Mosauer, 1936; Smith, 1942, 1973; Smith and
Alvarez del Toro, 1977; Taylor, 1939; Walker, 1955; Werler,
1957; Werler and Shannon, 1957).

Scale terminology follows Savage (1963). The characters
are defined below.

1.

2.

3.

4.

5.

6.

7.

8.

9.

10.

11.

12.

13.

14.

15.

16.

17.

18.

19.

20.

21.

22.

23.

24.

25.

26.

27.

28.

FPT

LTR

DBPVR

1WD1

IWV1

PTMP

DBPVT

GC1IL

GUL

PVTL

VL

4TL

4TLD

DOR

DAPVR

PVR

PVS

PVT1

PVT2

PVT3

RPOL

RPAW

RPFML

RPFL1

RMW

RNL

RML

RAPPSL

Femoral pores (total both sides).

Lateral rows of tubercles (axilla to groin).
Dorsals between paravertebral rows of tu-
bercles.

Dorsal interwhorls in first caudal segment.
Ventral interwhorls in first caudal segment.
Pretympanics (total both sides) separating
postocular from second postorbital suprala-
bial.

Distance between large paravertebral tuber-
cles within-row, expressed in number of mid-
dorsal scales.

Gulars contacting first pair of mfralabials.
Gulars (fold to second infralabials).

Large tubercles in paravertebral row (axilla
to groin).

Ventrals (gular to vent; includes preanals).
Fourth toe lamellae (ventral).

Fourth toe lamellae divided (i.e., with ca. mid-
ventral sutures).

Dorsals occiput to rump (above vent).
Dorsals in row immediately above paraver-
tebral row (axilla to groin).

Total scales in paravertebral row (axilla to
groin).

Scales in paravertebral row (a-g) smaller than
1.5 dorsals.

Scales in paravertebral row (a-g) larger than
1.5 dorsals.

Scales in paravertebral row (a-g) larger than

2.0 dorsals.

Scales in paravertebral row (a-g) larger than

3.0 dorsals.

Length of postocular/length of orbit.

Width of posterolateral preanal/width of pos-
teromedial preanal.

Prefrontal; length along midline/length along
lateral border.

Prefrontal: length of mid-line suture/length
along lateral border.

Width of median (prefrontal)/anterior width
of interparietal.

Length of nasal/length of postparietal.
Length of median (prefrontal)/length of fron-
tal.

Length (total both sides) of all anomalous su-
tures on postparietals/length of postparietals.

Figure 1. Location of the 21 population samples of Lepidophyma
in Mexico north of 19°N. Stippled area indicates approximate dis-
tribution of pine-oak woodland (after Leopold, 1959). Population
numbers are those used throughout the paper (see Specimens Ex-
amined for localities).

29. RPNH Height of postnasal/height of anterior loreal.

30. RSLH Height of second postorbital supralabial/

height of first postorbital supralabial.

Variation in the 30 characters was analyzed univariately
with BMDP1D for simple data description, and multivari-
ately with BMDP7M for stepwise discriminant analysis (Dix-
on, 1981). In all discriminant analyses the a priori groups
were individual population samples rather than population
groups or species.

RESULTS AND DISCUSSION

In the following sections the results of discriminant analyses
of populations of Lepidophyma from northern Mexico are
used to identify northern population groups, which in turn
are compared with populations from southern Mexico. The
northern population groups are then viewed relative to their
univariate differences and geographic relationships to arrive
at the definition of unique morphological units. Finally, names
are allocated to these units (morphospecies) on the basis of
included topotypic and/or type material, and each species is
briefly summarized.

NORTHERN MEXICO POPULATIONS

The initial discriminant analysis utilized 30 characters and
18 of the 21 populations from northeastern Mexico (Fig. 1,

2 Contributions in Science, Number 349

Bezy: Systematics of Lepidophyma in Mexico

_J I I I I —

-14 -7 0 7 14

Figure 2. Nineteen population samples of Lepidophyma from
northeastern Mexico plotted on the first two canonical variables for
25 characters. Population centroids are indicated by dots, and the
number of the sample is placed along the line enclosing all included
specimens (lower case letters). Upper case letters indicate population
groups identified by the analysis.

Table 1). In samples 3, 5, and 19 there are no individuals
on which all of the characters could be scored. The analysis
resulted in high resolution of the populations in that 99 per-
cent (127/128) of the individuals were “correctly” assigned
by the posterior classification to the locality sample of which
they were a member (one specimen of sample 7 was mis-
assigned to 6). A second analysis was performed excluding
five characters (IWD1, IWV1, RPFML, RPFL1, RML) in
order to allow inclusion of sample 5 (Fig. 2). Samples 3 and
19 were not included in any of the multivariate analyses due
to the limitations of the data available from them. While
both analyses produced similar results, the reduction in char-
acters of the second lowered the accuracy of the posterior
classification (96%; 126/131).

From the second analysis, eight population groups were
identified on the basis of overlap or juxtaposition of the
included samples and the distance between groups in dis-
criminant space (Fig. 2). The first canonical variable accounts
for 57 percent of the variation, is most heavily loaded with
LTR. PVS, PVR, DOR, and GUL (in order of decreasing
weight), and places group A at one end, and B, C, and D at
the other, with E, F, G, and H occupying intermediate po-
sitions. The second coordinate has heavy loadings for PVS,
PVR, PVT1, LTR, and FPT, accounts for 21 percent of the
variation, and effectively separates groups E, G, and H from
one another.

The sample comprising group A (21) is highly isolated in
discriminant space from all other populations, suggesting it
is not a member of the same morphological complex. Group
D is a discrete cluster of four overlapping populations (13-
lb) that is approximately equidistant from B (17, 18) and C
(20). The three populations of group E (9-1 1) form a mod-
erately tight cluster that is only weakly separated from the
loosely associated populations of G (5-8) and the one spec-
imen (12) comprising the intermediate group F. The three
populations of group H ( 1 , 2, 4) are well separated from their
nearest discriminant neighbor, group G.

(J I I I l_

-14 -7 0 7 14

Figure 3. Twelve population groups of Lepidophyma from Mexico
plotted on the first two canonical variables for 30 characters. Lines
enclose all individuals comprising each of the eight northern (A-G)
and four southern (I-L) population groups.

COMPARISONS WITH SOUTHERN GROUPS

Twenty-seven samples from southern Mexico were com-
pared with the 18 northern populations to identify those that
might be closest morphologically to northern groups. The
initial discriminant anlaysis utilized 30 characters and a total
of 288 specimens arrayed in 45 populations (Fig. 3), and
produced high resolution of the populations in that the ac-
curacy of the posterior classification was 98 percent. The first
canonical variable is most heavily loaded with PVS, DBPVT,
LTR, GUL, and PVR, and expresses 43 percent of the total
dispersion; the second expresses 2 1 percent and is dominated
by LTR, PVS, PVTL, FPT, and RPAW. The graph (Fig. 3)
was used primarily to identify those southern population
groups that are multivariately most similar to the northern
ones and which are further resolved in subsequent analyses
containing fewer populations.

All but one (Fj of the eight northern groups identified
in the previous analysis remain separated from each other,
although they are approached or overlapped by three south-
ern groups (J, K, L) (Fig. 3). Northern groups B, C, and D
were well separated from both southern and northern pop-
ulations and thus are not included in the subsequent analyses.
Southern group I is also strongly separated from all popu-
lations, and its nearest discriminant neighbor is another
southern group (J). Consequently, it was also excluded from
further analysis. In the following analyses, northern groups
E through H are compared in greater detail first with K and
L, and then with J.

The 10 populations of northern groups E, F, G, and H
were analyzed together with the four populations of southern
groups K and L (Fig. 4). The posterior classification was 98
percent ( 1 2 1/123) accurate, one specimen of sample 11 being
misassigned to 10 (both group E) and one of sample 43 to
44 (both group K). The first canonical variable accounts for
60 percent of the total dispersion, is heavily loaded with

Contributions in Science, Number 349

Bezy: Systematics of Lepidophyma in Mexico 3

Table 1. Variation among 30 scale characters for 21 population samples of Lepidophyma from northern Mexico. Sample size is (in parentheses)
under each of the population numbers (POP). In each cell the upper number is the mean; the middle, the standard error; and the lower, the
range. See text for character abbreviations and locality data.

POP

FPT

LTR

DBPVR

1 WD1

IWV1

PTMP

DBPVT

GC1IL

GUL

PVTL

VL

4TL

4TLD

DOR

DAPVR

PVR

1

31.3

18.1

3.17

3.0

2.1

2.1

2.83

.3

45.0

15.9

36.1

25.3

8.1

161.2

83.2

44.8

(15)

.37

.59

.093

0

.09

.07

.080

.13

.59

.21

.15

.33

.56

1.88

.99

1.13

29-34

15-22

2. 5-4.0

3

2-3

2-3

2. 5-3. 5

0-1

42-49

15-17

35-37

24-29

4-12

150-174

77-91

39-53

2

29.0

19.0

2.50

3.0

2.0

3.0

2.50

1.0

46.0

17.0

35.0

26.0

7.0

164.0

83.0

43.0

u)

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

29

19

2.5

3

2

3

2.5

1

46

17

35

26

7

164

83

43

3

29.0

-

3.50

2.0

2.0

2.0

3.00

1.0

46.0

-

-

20.0

5.0

-

-

-

U)

0

-

0

0

0

0

0

0

0

—

-

0

0

-

-

-

29

-

3.5

2

2

2

3.0

1

46

-

-

20

5

-

-

-

4

27.0

17.0

3.38

2.5

1.5

2.3

3.00

.3

45.0

16.0

35.3

23.0

6.3

172.5

91.0

47.0

(4)

.41

0

.125

.58

.29

.25

0

.25

.71

.41

.25

.41

.25

3.43

1.47

1.08

26-28

17

3.0-3. 5

2-3

1-2

2-3

3.0

0-1

43-46

15-17

35-36

22-24

6-7

163-178

87-94

44-49

5

28.0

31.0

5.00

-

-

1.0

4.50

3.0

56.0

15.0

38.0

31.0

15.0

207.0

98.0

63.0

U)

0

0

0

-

-

0

0

0

0

0

0

0

0

0

0

0

28

31

5.0

-

-

1

4.5

3

56

15

38

31

15

207

98

63

6

27.8

29.3

3.89

3.2

1.8

2.0

3.08

.5

48.3

16.8

35.8

26.3

14.6

182.1

93.5

51.0

(.32)

.45

.40

.122

.52

.11

.15

.106

.11

.49

.35

.23

.33

.53

1.96

1.26

.87

24-35

24-34

3. 0-5.0

2-4

0-3

0-4

1. 0-4.0

0-2

43-55

15-23

34-39

23-30

9-23

166-217

83-1 12

42-62

7

28.0

29.0

4.83

3.3

2.0

2.0

3.50

.3

51.0

16.7

35.3

25.3

13.3

206.3

107.0

57.0

(4)

1.53

1.00

.167

.58

0

0

.289

.25

1.78

.33

.88

.67

.88

5.36

5.51

5.57

26-31

27-30

4. 5-5.0

3-4

2

2

3. 0-4.0

0-1

48-55

17-18

34-37

24-26

12-15

196-214

96-113

46-64

8

26.2

27.8

4.08

3.5

2.5

2.2

3.08

1.0

45.5

18.0

36.0

27.7

14.2

191.3

97.8

53.8

(6)

.40

1.14

.201

.55

.22

.31

.201

0

1.09

1.83

.37

.56

.98

2.91

2.12

3.28

25-27

25-31

3. 5-5.0

3-4

2-3

1-3

2. 5-4.0

1

43-50

15-27

35-37

26-30

1 1-18

180-199

91-104

42-64

9

26.0

36.0

4.00

3.0

2.0

2.0

2.50

0

42.0

27.0

37.0

24.0

14.0

191.0

89.0

56.0

(!)

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

26

36

4.0

3

2

2

2.5

0

42

27

37

24

14

191

89

56

10

26.1

33.7

4.64

3.1

1.9

1.6

1.71

.6

44.0

24.9

34.8

25.5

17.7

180.1

91.0

52.2

(22)

.31

.44

.082

.47

.06

.15

.146

.11

.42

1.47

.24

.14

.52

1.33

1.25

1.18

24-30

31-38

4. 0-5.0

2-4

1-2

0-2

1. 0-3.0

0-1

41-49

15-39

33-37

24-27

13-23

165-188

78-99

40-66

1 1

25.2

34.2

4.25

4.0

2.2

2.2

3.25

1.1

47.8

19.0

36.6

25.7

18.8

184.6

95.5

61.4

(10)

.55

.51

112

.67

.20

.13

.186

.18

.83

1.09

.16

.45

.57

2.57

1.86

1.63

23-28

32-36

3. 5-4.5

3-5

1-3

2-3

2. 5-4.0

0-2

45-54

15-25

36-37

23-28

16-21

167-192

88-104

56-73

12

31.0

35.0

5.00

4.0

2.0

3.0

4.00

0

47.0

17.0

36.0

28.0

18.0

205.0

103.0

53.0

u)

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

31

35

5.0

4

2

3

4.0

0

47

17

36

28

18

205

103

53

13

32.7

29.3

5.67

5.0

3.0

.7

4.67

.3

62.0

16.0

36.3

25.3

12.3

243.3

126.7

70.3

(3)

1.45

1.20

.167

0

0

.67

.333

.33

3.21

.58

.33

.88

1.20

4.41

7.26

4.81

30-35

27-31

5. 5-6.0

5

3

0-2

4. 0-5.0

0-1

57-68

15-17

36-37

24-27

10-14

235-250

115-140

61-77

14

32.6

30.6

6.27

4.7

2.5

1.2

4.68

.4

62.4

16.2

35.9

26.5

13.9

244.2

122.5

73.4

(//)

.62

.59

.124

.47

.16

.25

.122

.16

.88

.23

.21

.43

.64

1.50

2.14

2.00

29-36

28-35

6. 0-7.0

4-5

2-3

0-2

4. 0-5.0

0-1

58-68

15-17

35-37

24-29

1 1-17

235-251

112-134

65-85

15

29.5

29.2

4.92

3.7

1.8

1.8

5.00

.5

56.8

15.8

36.3

26.0

13.2

237.5

128.0

62.5

(6)

.56

.40

.239

.52

.17

.17

0

.34

.70

.31

.21

.45

.65

2.43

2.36

2.14

28-3 1

28-31

4.0-5. 5

3-4

1-2

1-2

5.0

0-2

55-60

15-17

36-37

25-28

1 1-15

231-245

1 19-135

52-66

16

29.8

30.2

4.80

4.0

2.2

2.0

4.90

.8

58.6

15.6

36.0

27.0

16.8

239.4

120.4

60.8

(5)

.80

.49

.123

0

.20

0

.100

.20

.81

.40

.32

.71

.66

1.40

2.42

2.43

28-32

29-31

4. 5-5.0

4

2-3

2

4. 5-5.0

0-1

56-61

15-17

35-37

25-29

15-19

236-244

112-127

56-70

17

19.0

22.5

5.00

3.5

2.0

2.0

4.00

0

67.0

17.0

37.5

25.0

6.0

228.5

113.0

54.0

(3)

1.00

1.50

0

.71

0

0

0

0

4.00

0

.50

0

0

4.50

1.00

1.00

18-20

21-24

5.0

3-4

2

2

4.0

0

63-71

17

37-38

25

6

224-233

112-114

53-55

18

19.0

21.3

4.67

3.0

1.7

2.0

4.00

0

61.0

15.7

35.3

24.7

7.3

239.3

1 18.3

60.7

(3)

1.15

.67

.441

0

.33

0

0

0

.58

.67

.33

.33

.88

1.33

1.86

4.63

17-21

20-22

4.0-5. 5

3

1-2

2

4.0

0

60-62

15-17

35-36

24-25

6-9

238-242

1 16-122

53-69

19

19.5

24.0

5.00

4.0

2.0

2.0

4.50

—

66.0

17.0

36.5

24.0

8.5

-

-

-

(2)

.50

0

0

0

0

0

.500

-

1.00

1.00

.50

0

.50

-

-

-

19-20

24

5.0

4

2

2

4. 0-5.0

-

65-67

16-18

36-37

24

8-9

-

-

-

20

19.5

21.5

3.00

3.0

2.0

2.5

4.00

0

60.0

15.5

35.5

22.0

4.5

216.5

101.0

53.5

(2)

1.50

1.50

0

0

0

.50

0

0

1.00

.50

.50

0

1.50

3.50

1.00

4.50

18-21

20-23

3.0

3

2

2-3

4.0

0

59-61

15-16

35-36

22

3-6

213-220

100-102

49-58

21

33.5

46.5

4.02

2.2

2.0

3.8

2.54

.5

36.4

11.3

34.3

26.3

11.1

133.5

64.2

51.2

(20)

.28

.54

.057

.38

0

.24

.098

.11

.41

.73

.19

.32

.46

1.09

.69

1.08

32-37

43-50

3. 5-5.0

2-3

2

2-6

2. 0-3.0

0-1

33-39

6-18

33-36

25-30

7-16

126-142

59-68

44-58

4 Contributions in Science, Number 349

Bezy: Systematics of Lepidophyma in Mexico

Table 1. Continued.

POP

PVS

PVT 1

PVT2

PVT3

RPOL

RPAW

RPFML

RPFL1

RMW

RNL

RML

RAPPSL

RPNH

RSLH

1

13.9

30.0

20.2

15.7

.229

.603

.459

.382

.117

.136

.100

0

.704

1.079

(15)

.98

.55

.81

.21

.0249

.0311

.0409

.0475

.0514

.0086

.0491

0

.0118

.0476

7-20

26-34

15-28

15-17

. 1 5-56

.39-82

.1 1-.70

.1 1-.65

0-.56

o

O'

o

0-.63

0

.65-8 1

.91-1.61

2

15.0

28.0

17.0

36.0

.247

.788

.456

.456

0

.121

0

0

.754

.890

(/)

0

0

0

0

0

0

0

0

0

0

0

0

0

0

15

28

17

16

.25

.79

.46

.46

0

.12

0

0

.75

.89

3

—

—

-

-

.124

-

.401

.351

0

.309

0

0

.723

1.245

(/>

-

-

-

—

0

—

0

0

0

0

0

0

0

0

-

-

-

-

.12

-

.40

.35

0

.31

0

0

.72

1.25

4

15.0

32.0

26.5

16.3

.181

.706

.485

.485

0

.238

0

.221

.907

1.099

(4)

1.47

1.08

.87

.48

.0184

.0289

.0321

.0321

0

.0151

0

.2214

.1605

.0560

12-19

29-34

24-28

15-17

. 1 4-23

,64-.77

,42-,54

.42-. 54

0

,22-.28

0

0-.89

.73-1.39

.98-1.21

5

43.0

20.0

15.0

14.0

.241

.400

-

-

0

.072

-

0

.742

1.156

(1)

0

0

0

0

0

0

-

-

0

0

-

0

0

0

43

20

15

34

.24

.40

-

-

0

.07

-

0

.74

1.16

6

22.1

28.9

21.4

12.0

.181

.539

.564

.357

.272

.171

.375

.007

.781

1.172

(32)

1.25

.72

.84

.59

.0075

.0132

.0222

.0362

.0612

.0082

.0865

.0042

.0236

.0218

12-37

20-35

15-34

4-18

.1 1-.27

41-.69

.38-.S4

0-.71

0-.84

. 1 1— .29

0-1.26

0-.1 1

.66-1.40

.86-1.41

7

26.3

30.7

21.3

15.3

.158

.593

.623

.522

0

.133

0

0

.783

1.279

(4)

6.23

.67

2.85

.33

.0006

.0382

.0177

.0203

0

.0318

0

0

.0121

.0246

14-34

30-32

18-27

15-16

.16

.53-.66

.60-.66

.48-. 54

0

.08-. 19

0

0

o

00

f

SO

r-

1.23-1.32

8

27.5

26.3

19.3

14.0

.181

.545

.513

.344

.213

.164

.319

.46

.727

1.113

(6)

4.43

1.65

2.01

1.03

.0145

.0354

.0504

.0800

.1265

.0130

.2134

.3021

.0282

.0359

12-38

19-30

16-29

9-16

.13-.23

.40-.66

.36-68

0-.52

0— .77

. 1 3-22

0-1.32

0-1.65

.64-.81

1.01-1.23

9

24.0

32.0

27.0

0

.154

.300

.754

.422

.623

.132

.682

0

.787

1.253

(/)

0

0

0

0

0

0

0

0

0

0

0

0

0

0

24

32

27

0

.15

.30

.75

.42

.62

.13

.68

0

.79

1.25

10

21.0

31.2

24.0

2.5

.188

.497

.540

.455

.141

.167

.109

.017

.856

1.180

(22)

1.67

.94

1.02

.73

.0140

.0190

.0218

.0259

.0585

.01 10

.0483

.0173

.0283

.0338

7-42

22-37

17-33

0-13

.07-32

.31-68

.35-.82

. 1 4—65

0-.95

.09-27

0-.85

0-.38

.71-1.37

.95-1.55

1 1

31.5

29.9

19.1

2.9

.176

.589

.475

.480

.045

.172

.022

.140

.803

1.141

(10)

1.89

1.04

.99

.75

.01 19

.0364

.0322

.0433

.0284

.0160

.0137

.1102

.0209

.0369

21-40

26-35

16-25

0-6

. 1 0— .22

.39-. 75

.27-.69

.27-.81

0-.28

.12-28

0-.13

0-1.12

.71-9 1

.84-1.25

12

20.0

33.0

22.0

9.0

.200

.466

1.071

.283

.729

.106

.739

0

.750

1.141

(1)

0

0

0

0

0

0

0

0

0

0

0

0

0

0

20

33

22

9

.20

.47

1.07

.28

.73

.11

.74

0

.75

1.14

13

42.0

28.3

23.3

15.0

.146

.498

.562

.611

0

.188

0

0

.744

1.168

(3)

5.13

2.85

3.84

1.15

.0042

.1159

.0405

.0654

0

.0051

0

0

.0342

.1171

35-52

25-34

19-31

13-17

.14-. 15

.27-63

.49-63

.49-. 71

0

. 1 8-. 1 9

0

0

.70-8 1

.94-1.32

14

42.6

30.7

23.5

15.6

.168

.570

.579

.579

0

.129

0

0

.745

1.176

an

2.38

.76

1.26

.34

.0062

.0313

.0321

.0321

0

.0027

0

0

.0276

.0701

32-56

26-34

17-29

14-18

. 14— .20

,35-,74

.46-. 76

.46-. 76

0

.11-. 15

0

0

.66-.95

.95-1.76

15

26.7

35.8

30.3

17.7

.140

.728

.528

.528

0

.181

0

0

.754

1.11 1

(6)

2.20

1.22

1.23

.67

.0085

.0584

.0198

.0198

0

.0129

0

0

.0184

.0427

17-32

33-40

26-35

16-20

12-.18

46-.85

.48-. 62

.48-. 62

0

.14-22

0

0

.70-.83

.96-1.21

16

29.8

31.0

28.2

16.2

.189

.500

.544

.544

0

.129

0

0

.743

1.114

(5)

2.63

.45

.80

.58

.0177

.0185

.0468

.0468

0

.0123

0

0

.0236

.0259

25-40

30-32

26-30

15-18

. 14-.24

,44-.54

.42-.68

.42-.68

0

.10-15

0

0

.69-8 1

1.02-1.17

17

26.5

27.5

31.5

19.5

.151

.537

.670

.670

0

.136

0

0

.677

.954

(2)

5.50

4.50

1.50

1.50

.0217

.0020

.0049

.0049

0

.0232

0

0

.0636

.0087

21-32

23-32

30-33

18-21

. 1 3-. 1 7

.54

.67

.67

0

.1 1-.16

0

0

.61-74

,95-,96

18

30.3

30.3

30.3

20.7

.181

.566

.747

.747

0

.156

0

.037

.775

.819

(3)

4.91

.33

.33

1.45

.0206

.0496

.0247

.0247

0

.0102

0

.0370

.0039

.0343

19
(2)

20

22-39

30-31

30-31

18-23

. 1 5-,22

.178

.0018

.18

.145

.55-.66

.70-.77

.70-.78

0

.14-. 18

0

0-.1 1

.77-78

.76-.88

22.0

31.5

30.0

16.0

.629

.709

.709

0

.090

0

0

.776

.805

(2)

4.00

.50

0

0

.0251

.0592

.0443

.0443

0

.0059

0

0

.0281

.0216

18-26

31-32

30

16

. 1 2-. 1 7

,57-.69

,67-,75

,67-,75

0

.08-10

0

0

.75— .8 1

,78-,83

21

36.5

14.6

3.9

0

.208

.407

.491

.405

.094

.199

.137

.052

.744

.745

(20)

1.97

1.05

.74

0

.0056

.0168

.0264

.0367

.0400

.0076

.0587

.0323

.0087

.0155

22-50

8-22

0-1 1

0

. 1 5-26

.27-.54

.28-. 71

0-.63

0-.62

.15-. 26

0-.84

0-.59

O'

00

1

bo

O

.59-. 86

Contributions in Science, Number 349

Bezy: Systematics of Lepidophyma in Mexico 5

Figure 4. Fourteen samples of Lepidophyma of two southern (K.
and L) and four northern (E-H) population groups plotted on the
first two canonical variables for 30 characters. Dots are centroids;
sample numbers are along lines enclosing all included specimens.

4TLD, LTR, PVS, PVR, and RMW; the second explains 16
percent of the variation and is most weighted with PVS, PVR,
PVT 1 , FPT, and PVT2. Southern groups K and L are strongly
separated from H, their nearest discriminant neighbor among
northern groups. However, one population of group H (4) is
separated from the other two populations of the group in the
direction of group L. The relationships of population 4 are
discussed further on p. 7.

The 1 7 populations of group J were analyzed together with
northern groups E through H (Fig. 5). The accuracy of the
posterior classification was 98 percent (128/131), two spec-
imens of sample 1 I being misclassified as 10 (both group E).
The first variable accounts for 42 percent of the total dis-
persion and is influenced most by PVS, PVR, PVT1, FPT,
and LTR; the second expresses 26 percent and has heavy
loadings for PVS, PVT1, PVR, LTR, and DAPVR. The five
groups are separated from one another, although one pop-
ulation of group J (27) is separated from the remainder of
the group and is placed intermediate between J and E and
F, and two individuals of sample 37 (group J) approach
group G.

GROUP ANALYSES

The multivariate relationships of the 46 populations de-
scribed above are here considered in respect to univariate
similarities or differences between population groups (Table
2) and to geographic distributions (Fig. 6) in order to arrive
at the definition of morphologically diagnosable units of Lep-
idophyma occurring in northern Mexico. It is anticipated that
the resultant units should consist of groups of populations
that are overlapping orjuxtaposed in discriminant space, that
can be diagnosed by one or more univariate characters, that
are not linked to other groups by univariately, multivariately,
and geographically intermediate populations, and that thus
represent morphospecies. For a genus such as Lepidophyma,
in which sympatry is rare, discordant variation common,
and populations often disjunct and represented by small sam-
ple sizes, such morphologically defined units are initial species
hypotheses to be tested by securing additional samples and
information (e.g., allozyme data).

Groups E, F, and G are positioned nearest each other in
the four discriminant analyses (Figs. 2-5) and they overlap

Figure 5. Twenty-seven samples of Lepidophyma of one southern
(.1) and four northern (E-H) population groups plotted on the first
two canonical variables for 30 characters. Presentation as in Fig. 4.

in all individual characters (Table 2). The three appear to
represent a single species unit EFG distributed along the
Sierra Madre Oriental from southern Tamaulipas to Vera-
cruz (Fig. 6).

The nearest geographic and discriminant neighbor of group
H among northern populations is group G (Figs. 2 and 6),
from which it differs (=no overlap in range of variation) in
LTR (Table 2). The decision as to whether H should be
considered specifically distinct from EFG is complicated by

Figure 6. Distribution of eight population groups of Lepidophyma
in northeastern Mexico. Lines enclose the samples (numbers) in-
cluded in the groups (letters).

6 Contributions in Science, Number 349

Bezy: Systematics of Lepidophyma in Mexico

Figure 7. Lateral body surface of specimens of Lepidophyma of
groups A (upper, sample 21, AMNH 13879) and E (middle, sample
1 1, LACM 106742; and lower, sample 10, LACM 109771).

Figure 8. Lateral body surface of specimens of Lepidophyma of
groups G (upper, sample 6, UMMZ 1 02980; middle, sample 8, LACM
131 145) and H (lower, sample 2, EAL 4644).

the presence of intermediate states, observed in the lateral
tubercle rows, that are not expressed in the LTR counts. In
some specimens, the low number of lateral tubercle rows that
characterizes group H results from a slight reduction of some
of the rows in terms of the distance they extend above the
ventrals and the relative size of the tubercles which compose
them (Figs. 7-8). While uniform criteria were employed
throughout the study to determine which rows to include in
the counts, for some of the specimens in groups G and H
the decision was difficult and repeatability of the counts was
low. The difference in LTR between H and EFG is thus less
discrete than suggested by the counts and is bridged by in-
termediate morphological states. While additional specimens
and information (e.g., allozyme data) are needed to fully
evaluate this situation, it seems best not to place emphasis
on the differences in LTR number, and to recognize a single
species unit composed of groups E, F, G, and H.

In one of the analyses, population 4 is slightly separated
from the other two populations of group H (1, 2) in the
direction of southern group L from coastal Michoacan (Fig.
4). For a number of characters (e.g., LTR, IWD 1 , GUL, 4TL,
and 4TLD), the mean for population 4 is intermediate be-
tween L and the Nuevo Leon populations (1,2) (Tables 1

and 2). Nevertheless, population 4 differs from L in four
characters (FPT, PTMP, 4TLD, DOR), and from the Nuevo
Leon populations (1, 2) in one (FPT). Evaluation of these
differences is hampered by the small sample sizes of popu-
lation 4 (N = 4) and group L (N = 3). To estimate the range
of variation of 4 and L that would be expected with larger
sample sizes, three standard deviations of population 1 (N =

1 5) were added to and subtracted from the means of each of
the characters to encompass 99.7 percent of the population
(Simpson, Roe, and Lewontin, 1 960; 1 39). The estimated range
of population 4 overlaps the observed range of population 1
in all characters, but is separated from the estimated range
of group L in FPT (23-31 vs. 13-21) and PTMP (1. 5-3.0
vs. 3. 2-4. 8). While additional material is necessary to fully
evaluate the relationships of population 4, the information
at hand suggests that it should be considered a member of
group EFGH. Further collecting along the western flank of
the Sierra Madre Oriental seems likely to produce material
linking the Sierra Alvarez population (4) geographically and
morphologically with the Nuevo Leon populations (1,2) (Fig.
6).

Two populations of southern group J (27, 37) approach
EFGH in discriminant space (Fig. 5). The two groups differ

Contributions in Science, Number 349

Bezy: Systematics of Lepidophyma in Mexico 7

Table 2. Variation among 30 scale characters for eight population groups of Lepidophyma in northern Mexico. Presentation as in Table 1.

Group

FPT

LTR

DBPVR

IWD1

IWV1

PTMP

DBPVT

GC1IL

GUL

PVTL

VL

4TL

4TLD

DOR

DAPVR

PVR

A

33.5

46.5

4.03

2.2

2.0

3.8

2.54

.5

36.4

1 1.3

34.3

26.3

1 1.1

133.5

64.2

51.2

(20)

.28

.54

.057

.09

0

.24

.098

.11

.41

.73

.19

.32

.46

1.09

.69

1.08

32-37

43-50

3. 5-5.0

2-3

2

2-6

2. 0-3.0

0-1

33-39

6-18

33-36

25-30

7-16

126-142

59-68

44-58

B

19.0

21.8

4.80

3.2

1.8

2.0

4.00

0

63.4

16.2

36.2

24.8

7.0

235.0

1 16.2

58.0

(5)

.71

.66

.255

.20

.20

0

0

0

1.96

.49

.58

.25

.71

3.10

1.69

3.03

17-21

20-24

4.0-5. 5

3-4

1-2

2

4.0

0

60-71

15-17

35-38

24-25

6-9

224-242

1 12-122

53-69

C

19.5

21.5

3.00

3.0

2.0

2.5

4.00

0

60.0

15.5

35.5

22.0

4.5

216.5

101.0

53.5

(2)

1.50

1.50

0

0

0

.50

0

0

1.00

.50

.50

0

1.50

3.50

1.00

4.50

18-21

20-23

3.0

3

2

2-3

4.0

0

59-61

15-16

35-36

22

3-6

213-220

100-102

49-58

D

31.3

30.0

5.58

4.4

2.3

1.5

4.80

.5

60.2

16.0

36.1

26.4

14.1

241.5

123.9

67.9

(25)

.47

.33

.157

.13

.11

.16

.071

.12

.73

.16

.13

.28

.46

1.15

1.48

1.64

28-36

27-35

4. 0-7.0

3-5

1-3

0-2

4. 0-5.0

0-2

55-68

15-17

35-37

24-29

10-19

231-251

112-140

52-85

E

25.8

33.9

4.50

3.4

2.0

1.8

2.20

.7

45.1

23.1

35.4

25.5

17.9

181.8

92.4

55.1

(33)

.27

.33

.072

.11

.07

.12

.167

.10

.49

1.13

.23

.17

.41

1.24

1.06

1.18

23-30

31-38

3. 5-5.0

2-5

1-3

0-3

1 .0-4.0

0-2

41-54

15-39

33-37

23-28

13-23

165-192

78-104

40-73

F

31.0

35.0

5.00

4.0

2.0

3.0

4.00

0

47.0

17.0

36.0

28.0

18.0

205.0

103.0

53.0

(D

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

0

31

35

5.0

4

2

3

4.0

0

47

17

36

28

18

205

103

53

G

27.6

29.1

4.01

3.0

1.9

2.0

3.15

.6

48.3

17.0

35.9

26.5

14.5

185.7

95.2

52.1

(43)

.37

.36

.107

.08

0

.12

.093

.11

.50

.37

.19

.30

.43

1.95

1.19

.94

24-35

24-34

3. 0-5.0

2-4

0-3

0-4

1.0-4. 5

0-3

43-56

15-27

34-39

23-31

9-23

166-217

83-113

42-64

H

30.4

17.9

3.18

2.9

2.0

2.2

2.85

.3

45.1

16.0

35.9

24.9

7.7

163.6

85.5

45.2

(20)

.49

.45

.083

.06

.10

.08

.064

.11

.46

.18

.15

.33

.45

1.84

1.00

.89

26-34

15-22

2. 5-4.0

2-3

1-3

2-3

2. 5-3. 5

0-1

42-49

15-17

35-38

22-29

4-12

150-178

77-94

39-53

Table 3. Variation among 30 scale characters for nine species of Lepidophyma from Mexico. Presentation as in Table 1.

Species

FPT

LTR

DBPVR

IWD1

1WV1

PTMP

DBPVT

GC1IL

GUL

PVTL

VL

4TL

4TLD

DOR

DAPVR

PVR

gaigeae

33.5

46.5

4.03

2.2

2.0

3.8

2.54

.5

36.4

11.3

34.3

26.3

11.1

133.5

64.2

51.2

(20)

.28

.54

.057

.09

0

.24

.098

.11

.41

.73

.19

.32

.46

1.09

.69

1.08

32-37

43-50

3. 5-4.0

2-3

2

2-6

2. 0-3.0

0-1

33-39

6-18

33-36

25-30

7-16

126-142

59-68

44-58

occulor

19.2

22.2

4.44

3.3

1.9

2.1

4.1 1

0

63.2

16.2

36.1

23.9

6.8

229.7

1 1 1.9

56.7

(9)

.46

.55

.306

.17

.11

.11

.11 1

0

1.28

.36

.35

.44

.70

4.09

3.04

2.46

17-21

20-24

3. 0-5. 5

3-4

1-2

2-3

4. 0-5.0

0

59-71

15-18

35-38

22-25

3-9

213-242

100-122

49-69

micropholis

31.3

30.0

5.58

4.4

2.3

1.5

4.80

.5

60.2

16.0

36.1

26.4

14.1

241.5

123.9

67.9

(25)

.47

.33

.157

.13

.11

.16

.071

.12

.73

.16

.13

.28

.46

1.15

1.48

1.64

28-36

27-35

4. 0-7.0

3-5

1-3

0-2

4. 0-5.0

0-2

55-68

15-17

35-37

24-29

10-19

231-251

112-140

52-85

sylvaticum

27.6

28.5

4.01

3.2

2.0

2.0

2.78

.6

46.6

18.9

35.7

25.8

14.2

180.0

92.3

51.7

(98)

.27

.64

.074

.06

.05

.07

.083

.06

.33

.52

.12

.18

.46

1.36

.77

.70

23-35

15-38

2.5-5. 0

2-5

0-3

0-4

1. 0-4.5

0-3

41-56

15-39

33-39

20-31

4-23

1 50-2 1 7

77-113

39-73

tuxtlae

24.6

34.1

4.09

3.7

2.3

5.5

.29

1.6

42.6

39.3

39.6

25.8

16.3

172.0

79.6

45.0

(81)

.22

.20

.053

.06

.06

.11

.032

.09

.28

.43

.14

.16

.22

.97

.70

.43

20-29

30-40

3. 0-5.0

3-5

0-3

4-9

0-.9

0-4

37-49

30-47

37-42

23-28

13-20

150-190

69-98

37-55

pajapanensis

33.3

40.5

4.21

3.4

1.8

7.5

.14

.1

38.9

43.8

36.6

28.3

13.8

164.3

69.6

44.0

(19)

.35

.40

.088

.12

.10

.31

.037

.07

.53

1.02

.17

.24

.50

1.59

1.22

.84

30-36

37-43

4. 0-5.0

3-4

1-2

6-10

0-.5

0-1

35-43

39-51

35-38

26-30

10-17

152-175

59-82

37-49

flavimaculatum

33.7

28.5

4.45

3.8

2.1

6.8

2.99

.9

47.3

18.6

36.6

27.3

14.7

197.7

90.1

54.3

(49)

.56

.26

.084

.08

.05

.20

.119

.09

.41

.41

.13

.25

.34

1.64

1.17

1.00

27-41

25-32

3.0-5. 5

3-5

1-3

4-1 1

.5-4.0

0-2

40-55

15-27

34-38

23-31

10-22

173-221

75-118

43-69

smithii

23.1

17.5

4.21

3.0

2.1

2.7

3.18

.7

52.4

16.8

35.8

25.2

5.3

194.0

94.2

48.7

(36)

.56

.16

.102

.05

.08

.14

.067

.13

.39

.36

.18

.24

.33

1.43

1.15

.73

18-29

15-19

3.0-5. 5

2-4

1-3

2-4

2. 0-4.0

0-3

46-57

14-25

33-38

22-28

2-9

178-214

81-1 10

37-61

tarascae

17.0

16.7

2.67

2.7

2.0

4.0

2.50

.7

42.3

16.0

34.3

22.7

1.7

146.7

75.0

44.3

(3)

.58

.33

.333

.33

0

0

0

.33

.67

0

.33

.33

.33

1.67

1.00

1.20

16-18

16-17

2. 0-3.0

2-3

2

4

2.5

0-1

41-43

16

34-35

22-23

1-2

145-150

74-77

42-46

8 Contributions in Science, Number 349

Bezy: Systematics of Lepidophyma in Mexico

Table 2. Continued.

Group

PVS

PVT1

PVT2

PVT3

RPOL

RPAW

KPFML

RPFL1

RMW

RNL

RML

RAPPSL

RPNH

RSLH

A

36.5

14.6

3.9

0

.208

.407

.491

.405

.094

.199

.127

.052

.744

.745

(20)

1.97

1.05

.74

0

.0056

.0168

.0264

.0367

.0400

.0076

.0587

.0323

.0087

.0155

22-50

8-22

0-1 1

0

.15-. 26

.27-54

.28-. 71

0-.63

0-.62

. 1 5-. 2 6

0-.84

0-.59

.68-80

.59-86

B

28.8

29.2

30.8

20.2

.169

.555

.716

.716

0

.148

0

.022

.736

.873

(5)

3.34

1.59

.58

.97

.0151

.0281

.0233

.0233

0

.0104

0

.0222

.0314

.0381

21-39

23-32

30-33

18-23

. 13-.22

.50-.66

.67-.78

.67-78

0

.1 1-.18

0

0-.11

.61 — .78

.76-96

C

22.0

31.5

30.0

16.0

.145

.629

.709

.709

0

.090

0

0

.776

.805

(2)

4.00

.50

0

0

.0251

.0838

.0443

.0443

0

.0059

0

0

.0281

.0216

18-26

31-32

30

16

.12-. 17

,57-,69

.67-75

.67-75

0

.08-. 10

0

0

.75-81

.78-.83

D

36.2

31.7

26.0

16.2

.163

.585

.557

.563

0

.147

0

0

.747

1.146

(25)

1.99

.72

.96

.32

.0060

.0592

.0175

.0187

0

.0067

0

0

.0133

.0334

17-56

25-40

17-35

13-20

. 1 2-.24

,27-,85

.42-.76

.42-76

0

. 10-.22

0

0

.66-. 95

.94-1.76

E

24.3

30.8

22.6

2.5

.183

.519

.527

.462

.127

.167

.100

.054

.838

1.170

(33)

1.50

.70

.84

.53

.0100

.0283

.0193

.0214

.0432

.0088

.0376

.0356

.0202

.0252

7-42

22-37

16-33

0-13

.07-.32

.30-.75

.27-.82

. 19-.8 1

0-.94

.09-.28

0-.85

0-1.12

.71-1.37

.84-1.55

F

20.0

33.0

22.0

9

.200

.466

1.071

.283

.729

.106

.739

0

.750

1.141

< i )

0

0

0

0

0

.0193

0

0

0

0

0

0

0

0

20

33

22

9.0

.20

.46

1.07

.28

.73

.11

.74

0

.75

1.14

G

23.7

28.4

2.09

12.6

.180

.540

.560

.368

.237

.165

.338

.072

.772

1.171

(43)

1.30

.65

.73

.50

.0062

0

.0190

.0309

.0508

.0073

.0746

.0473

.0185

.0181

12-43

19-35

15-34

4-18

.1 1-.27

.4Q-.69

.36-. 81

0-.71

0-.84

.07-.29

0-1.32

0-1.65

.64-1.40

.86-1.41

H

14.2

30.3

21.3

15.9

.220

.633

.464

.406

.088

.156

.074

.044

.749

1.073

(20)

.78

.50

.87

.18

.0194

.0120

.0311

.0370

.0399

.0121

.0373

.0443

.0368

.0377

7-20

26-34

15-28

15-17

. 1 4-56

.39-.82

.1 1-.70

.1 1-.65

0-.56

.09-28

0-.63

0-.89

.65-1.39

.89-1.61

Table 3. Continued.

Species

PVS

PVT 1

PVT2

PVT3

RPOL

RPAW

RPFML

RPFL1

RMW

RNL

RML

RAPPSL

RPNH

RSLH

gaigeae

36.5

14.6

3.9

0

.208

.407

.491

.405

.094

.199

.127

.052

.744

.745

(20)

1.97

1.05

.74

0

.0056

.0168

.0264

.0367

.0400

.0076

.0587

.0323

.0087

.0155

22-50

8-22

0-1 1

0

. 1 5-. 26

.27-.54

.28—. 7 1

0-.63

0-1.49

. 1 5-26

0-.84

0-.59

.68-. 80

.59-. 86

occulor

26.9

29.9

30.6

19.0

.166

.576

.714

.714

0

.131

0

.016

.748

.854

(9)

2.76

1.18

.43

1.02

.0099

.0271

.0188

.0188

0

.0129

0

.0159

.0237

.0295

18-39

23-32

30-33

16-23

. 1 2-,22

.50-.69

.66-. 78

.67-.77

0

oo

OO

o

0

0-.1 1

.6 1 —.8 1

.76-. 96

micropholis

36.2

31.7

26.0

16.2

.163

.585

.557

.563

0

.147

0

0

.747

1.146

(25)

1.99

.72

.95

.32

.0060

.0283

.0175

.0187

0

.0067

0

0

.0133

.0334

17-56

25-40

17-35

13-20

.12-.24

.27-85

.42-.76

,42-,76

0

.09-.22

0

0

,66-.95

.94-1.76

sylvaticum

21.9

29.7

21.0

9.8

.190

.551

.532

.408

.171

.165

.200

.059

.790

1.152

(98)

.88

.40

.47

.62

.0062

.0109

.0141

.0172

.0287

.0052

.0373

.0253

.0133

.0144

7-43

19-37

15-34

0-18

.07-. 56

.30-. 82

.1 1-1.07

0 — .8 1

0-.95

.07-.31

0-1.32

0-1.65

.64-1.40

.84-1.61

tuxtlae

5.1

39.9

32.2

1.7

.224

.830

.734

.042

1.024

.108

1.216

.027

.734

.547

(81)

.47

.37

.63

.33

.0047

.0128

.0092

.0126

.0243

.0049

.0539

.0159

.0054

.0106

0-18

31-49

1 1-42

0-15

.1 1-.33

.59-1.12

O'

OO

f

OO

T

0-.57

0-1.31

.04-.29

0-3.15

0-.99

.63-.83

.32-77

pajapanensis

2.8

41.2

26.6

.1

.222

.540

.803

.157

1.087

.163

1.656

.002

.764

.485

(19)

.56

1.02

1.33

.06

.0081

.0189

.0131

.0246

.0264

.0113

.1192

.0015

.0116

.0349

0-8

34-49

15-40

0-1

. 1 3-28

.38-.69

.70-.90

0-.31

.85-1.29

. 1 0— .28

1.00-2.65

0-.03

.69-86

,32-.88

Jlavimaculatum

20.1

34.2

25.3

13.7

.204

.411

.699

.175

.834

.145

.978

.061

.736

.557

(49)

1.17

.65

.71

.76

.0089

.0199

.0126

.0128

.0447

.0075

.0687

.0356

.0063

.0154

6-44

24-43

17-35

1-24

.11-39

.19-69

,52-,88

0-.73

0-1.31

.07-.28

0-2.43

0-1.66

.58-83

.36-89

smithii

17.0

31.1

26.9

16.3

.181

.723

.713

.349

.608

.137

.783

.099

.794

.904

(36)

.75

.45

.60

.4!

.0065

.0459

.0141

.0414

.0681

.0052

.1050

.0398

.0093

.0106

8-29

25-39

18-32

13-27

.1 1-.28

.43-1.35

.54— .89

0-.79

0-1.07

.08-24

0-2.62

0-1.04

.66-.92

.61-1.22

tarascae

13.7

30.7

25.0

16.0

.299

.591

.361

.361

0

.154

0

0

.982

.756

(3)

1.33

.33

1.53

0

.0717

.0923

.0664

.0664

0

.0130

0

0

.0227

.0417

1 1-15

30-31

23-28

16

.19-.44

.41-.71

.29-49

.29-49

0

.14-18

0

0

.94-1.02

.69-. 83

Contributions in Science, Number 349

Bezy: Systematics of Lepidophyma in Mexico 9

in PTMP (0-3 in 95/97 EFGH and 4-11 in all J) and RSLH
(0.84-1 .60 in all EFGH and 0.36-0.79 in 49/50 J) (Table 2).
The two specimens of EFGH with a PTMP of 4 are from a
population (6) at the northern end of the range (Fig. 1), and
the specimen of J with RSLH of 0.89 is from a population
in Tabasco (37), that is separated from the southernmost
sample (12) of EFGH by 350 km and by intervening pop-
ulations that are clearly assigned to J. Thus the populations
of EFGH and J that are most similar in morphology are not
geographically intermediate, and the multivariate and uni-
variate ditferences between the two population groups are
sufficiently constant that they are judged to represent units
that are likely reproductively isolated.

Group D is multivariately closest to G (Fig. 2). The four
populations of D are from the Sierra del Abra ofTamaulipas
and San Luis Potosi (Fig. 6), and differ in DOR from all
populations of group EFGH including those in the Sierra
Madre Oriental to the west and Sierra Tamaulipas to the east
(Table 2). While two of the three specimens from the Sierra
Tamaulipas (sample 7, group G) approach group D in dis-
criminant space, they are not geographically intermediate
(Fig. 6) and do not bridge the gap between the two in DOR
(Table 2). It is concluded that group D should be considered
specifically distinct from EFGH.

Groups B and C are closest to D in discriminant space,
but differ in FPT (18-21 vs. 28-36) and LTR (20-24 vs. 27-
35). Evaluation of these differences is hampered by small
sample sizes of B (N = 4) and C (N = 2). As an estimate of
the range of variation that would be expected with larger
samples, three standard deviations of D were added to and
subtracted from the means of B and C. The estimated ranges
of B and C overlap each other for all characters but differ
from the observed range ofD for FPT (12-27 vs. 28-36) and
LTR (17-26 vs. 27-35). The populations of B and C are
located in the Jalpan Valley of Queretaro and San Luis Potosi
(Fig. 6), and their combined ranges of variation differ in six
characters from the populations of E occurring in the Sierra
Madre Oriental, 22 km to the east. While larger sample sizes
are necessary to fully evaluate the differences between groups
B and C, their multivariate juxtaposition and the small uni-
variate differences between them indicate they are probably
members of the same species. On the other hand the number
and magnitude of the univariate differences between BC and
its nearest geographic (E) and discriminant (D) neighbors are
such that they are not likely to be bridged by larger samples.
Group BC is considered specifically distinct from D and
EFGH.

Group A is strongly separated in discriminant space from
the populations of all other groups (Figs. 2-3). It is morpho-
logically and geographically closest to group E (Fig. 6), but
differs in 6 of the 30 characters (Table 2). The univariate and
multivariate differences between A and other population
groups clearly qualify it as a distinct morphospecies.

While the combined problems of small sample sizes, dis-
junct distributions, and discordant variation confound some
of the decisions, four unique morphological units of Lepi-
dophyma are recognizable in northeastern Mexico: A, BC,
D, and EFGH.

ALLOCATION OF NAMES

Several of the groups identified in the discriminant analyses
include lizards that are either types or are from or near the
type locality of named taxa: Group A: Population 21: L.
gaigeae Mosauer, 1936; B: 20: L. smithu occulor Smith, 1942;
D: 14: L. micropholis Walker, 1955; E: 11: L. sylvaticum
Taylor, 1939; G: 6: L. flavimaculatum tenebrarum Walker,
1955; I: 51: L. tuxtlae Werler and Shannon, 1957, 41: L.
pajapanensis Werler, 1957, 48: L. sawini Smith, 1973, 47:
L. alvarezi Smith, 1 973; J: 28: L. flavimaculatum A. Dumeril
in Dumeril and Dumeril, 1851; K: 44: L. smithii Bocourt,
1876; L: 46: L. tarascae Bezy, Webb, and Alvarez, 1982.

The oldest available names for the species units recognized
in northern Mexico (Fig. 9) from the foregoing discussions
are: A, L. gaigeae ; BC, L. occulor, D, L. micropholis-, and
EFGH, L. sylvaticum. These are summarized below.

The systematic relationships among populations of Lepi-
dophyma in southern Mexico currently are under study (Bezy,
in prep.); the southern population groups used in this paper
are considered to represent the following species: I — L. pa-
japanensis (Veracruz) and L. tuxtlae (Veracruz, Oaxaca,
Chiapas); J = L. flavimaculatum (Atlantic versant east of the
Isthmus ofTehuantepec in Oaxaca, Veracruz, Tabasco, Chia-
pas, Quintana Roo); K = L. smithii (Pacific versant of Guer-
rero, Oaxaca, Chiapas); and L = L. tarascae (coastal Mi-
choacan).

SPECIES ACCOUNTS
Lepidophyma gaigeae Mosauer
Group A; Figures 7, 10

Lepidophyma gaigeae Mosauer, 1936:3. Holotype: MCZ

42145: Durango, State of Hidalgo, Mexico.

Gaigeia gaigeae: Smith, 1939:24.

DIAGNOSTIC CHARACTERS. Differs from other
members of the genus except L. radula in having 43-50
subequal scales (rather than 1 5-42 discrete rows of enlarged
tubercles) along the side of the body (axilla to groin) (Figs.
7-8) and fewer dorsal scales (126-142 vs. 145-251) (Table
3). It differs from L. radula and L. dontomasi in having two
(rather than one) caudal interwhorls complete ventrally.

DISTRIBUTION. The species is known from Hidalgo (near
the type locality) and Queretaro (between El Lobo and Jalpan;
Dixon et al., 1972), where it occurs in limestone crevices
primarily in pine-oak woodland (Fig. 9).

REMARKS. The high degree of separation of L. gaigeae
from other populations in the discriminant analyses is con-
sistent with its proposed separate generic (Smith, 1942) or
subgeneric (Smith, 1973) status. Geographic variation and
relationships of this form to L. dontomasi and L. radula are
currently under study (Bezy, in prep.).

KARYOTYPE. Lepidophyma gaigeae has a diploid chro-
mosome number of 38 with nine pairs of macrochromo-
somes and 10 pairs of microchromosomes. The karyotype is
unique in the genus, but closest to those of L. flavimaculatum,
L. pajapanensis. and L. tuxtlae (Bezy, 1972).

10 Contributions in Science, Number 349

Bezy: Systematics of Lepidophyma in Mexico

Figure 9. Distribution of the four species of Lepidophyma recog-
nized in northeastern Mexico (north of 19°N). Lines enclose the
samples included in each species.

Lepidophyma occulor Smith

Group BC; Figure 10

Lepidophyma smithii occulor Smith, 1942:378. Holotype:

USNM 47133: Jalpan, Queretaro.

Lepidophyma flavimaculatum occulor: Walker, 1955:5.
Lepidophyma occulor: Bezy, 1972:15.

DIAGNOSTIC CHARACTERS. Differs from all other
species of Lepidophyma except L. micropholis in having more
gulars (59-71 vs. 33-57), and from L. micropholis in having
fewer femoral pores (17-21 vs. 28-36), fewer lateral tubercle
rows (20-24 vs. 27-35), and fewer divided fourth toe lamellae
(3-9 vs. 10-19) (Table 3).

DISTRIBUTION. Lepidophyma occulor is known from
four localities in the Jalpan Valley of Queretaro and San Luis
Potosi (Fig. 9), where it has been found beneath stones in
arid tropical scrub (Dixon et al., 1972).

REMARKS. The species alternatively has been considered
a subspecies of either L. smithii (Smith, 1942) or L. Jlavi-
maculatum (Walker, 1955) from both of which it is well
separated multivariately (Fig. 3), differing from the former
in lateral tubercle rows and gulars and from the latter in
femoral pores, lateral tubercle rows, pretympanics, gulars,
and divided fourth toe lamellae. It is multivariately closest
to L. micropholis from which it differs in femoral pores.

Figure 10. Living individuals of Lepidophyma gaigeac (upper,
LACM 127170), L. occulor (middle, sample 18, TCWC 35605), and
L micropholis (lower, sample 16. TCWC 60767).

lateral tubercle rows, and divided fourth toe lamellae (Table
3).

KARYOTYPE. Lepidophyma occulor has a diploid chro-
mosome number of 36, with the lowest number of micro-
chromosomes (18) known in the family Xantusiidae (Bezy,
1972).

Lepidophyma micropholis Walker

Group D; Figure 10

Lepidophyma micropholis Walker, 1955:6. Holotype: UMMZ
101298: cave at El Pachon, about 5 miles NNE of Antigua
Morelos, Tamaulipas.

DIAGNOSTIC CHARACTERS. Differs from all other
species in the genus (except L. occulor) in having more dorsal

Contributions in Science, Number 349

Bezy: Systematics of Lepidophyma in Mexico 1

!

Figure 1 1. Living individuals of southern Madrean (upper, sample
1 1 . LACM 106752) and northern Madrean (lower, sample 8, LACM
131 146) population groups of Lepidophyma sylvalicum.

scales (231-251 vs. 126-222) and from L. occulor in having
more lateral tubercle rows (27-35 vs. 20-24) (Table 3).

DISTRIBUTION. Lepidophyma micropholis occurs in
southern Tamaulipas and northern San Luis Potosi at four
localities situated along the Sierra del Abra (Fig. 9). This
cavemiferous, low-lying range constitutes the easternmost
front of the Sierra Madre Oriental in the highly dissected
region between the Rio Guayalejo and the Rio Tamuin
(Mitchell et al., 1 977). The lizards have been found primarily
in limestone caves (El Pachon and Quintero) and fissures.

REMARKS. This extensively cavernicolous species is
closest in scalation to L. occulor and L. sylvaticum, and future
work may demonstrate the existence of morphologically and
biochemically intermediate populations, similar to those oc-
curring between the epigean and troglodytic Astyanax mex-
icanus of the Sierra del Abra (Avise and Selander, 1972;
Mitchell et al., 1977).

KARYOTYPE. Lepidophyma micropholis has a diploid
chromosome number of 36 with 1 6 macrochromosomes and
20 microchromosomes (Bezy, 1972).

Lepidophyma sylvaticum Taylor

Group EFGH; Figures 7-8, 11-12

Lepidophyma sylvatica Taylor, 1 939: 131. Holotype; FMNH

100102: 7 mi. north of Zacaultipan, Hidalgo.

Gaigeia sylvatica: Smith, 1942:380.

Figure 12. Living individuals of northern Madrean (upper, sample
6. LACM 106752) and western (lower, sample 1, LACM 106781)
population groups of Lepidophyma sylvaticum.

Lepidophyma sylvaticum: Walker, 1955:9.

Lepidophyma flavimaculatum tenebrarum Walker, 1955:1.
NEW SYNONYMY. Holotype: UMMZ 101374: ±5 miles
NW (by road) of Gomez Farias in the Sierra Madre Ori-
ental at “Rancho del Cielo.”

DIAGNOSTIC CHARACTERS. Differs from L. gaigeae
and L. occulor in numbers of gulars (41-56 vs. 33-39 and
59-7 1 , respectively), from L. micropholis in numbers of dor-
sals (150-217 vs. 231-251), from L. tarascae in numbers of
femoral pores (23-35 vs. 16-18), from L. smithii in having
a parietal foramen, from L. flavimaculatum in numbers of
pretympanics (0-3 vs. 4-11, 99%) and ratio of supralabial
height (0.84-1.61 vs. 0.36-0.79, 99%), from L. tuxtlae in
ratio of supralabial height (0.84-1.61 vs. 0.32-0.77), and
from L. pajapanensis in numbers of pretympanics (0-4 vs.
6-10) (Table 3).

DISTRIBUTION. The 12 populations occur from Vera-
cruz to Nuevo Leon along the Sierra Madre Oriental and
adjacent Mesa Central and Sierra Tamaulipas (Fig. 9).

REMARKS. The four population groups included in L.
sylvaticum are moderately divergent from one another and
further work may indicate that one or more of them should
be given separate taxonomic recognition (i.e., subspecies).

The northern Madrean group (G) includes four populations
in southern Tamaulipas and northern San Luis Potosi, three
from along the main axis of the Sierra Madre Oriental and

12 Contributions in Science, Number 349

Bezy: Systematics of Lepidophyma in Mexico

AX

6 + 8

AK NA

4 5

a* • • V •

KK

7 9

XX /#/S DM Xa *K /lt*»

Figure 13. Karyotypes of Lepidophyma sylvaticum (sample 6; upper, LACM 106758, <5; lower, LACM 106763, 9).

one in the Sierra Tamaulipas. They differ from the three
populations (group E) of southern San Luis Potosi, Quere-
taro, and Hidalgo in lateral tubercle rows (34/39 northern
Madrean with 31 or less, 30/33 southern Madrean with 32
or more) (Table 2). The hiatus between the ranges of the two
corresponds roughly to the Rio Panuco gap in the Sierra
Madre Oriental. Should further work demonstrate a need to
accord them separate nomenclatural status, L. sylvaticum
Taylor, 1939 is applicable to the southern group, and L. f
tenebrarum Walker, 1 955 is available for the northern group.

The southernmost specimen (group F) assigned to L. syl-
vaticum is from ca. 170 km SE of the type locality and occurs
at the northern base of the Cordillera Volcanica in central
Veracruz (Fig. 6). It has similarities to both southern and
northern Madrean L. sylvaticum, being closest to the former
in lateral tubercle rows and femoral pores, and to the latter
in dorsals. No other Lepidophyma are known from the Cor-
dillera Volcanica. The nearest populations to the south are
L. tuxtlae and L. pajapanensis of the Tuxtlas region (which
differ from L. sylvaticum in numbers of large paravertebrals,
pretympanics, and dorsals, and in ratio of supralabial height),
and L. flavimaculatum of the northern Isthmus of Tehuan-
tepec (which differ in pretympanics, lateral tubercle rows,
and ratio of supralabial height) (Tables 2-3).

The western group of L. sylvaticum (H, Figs. 6, 8, 12)
includes one population in the Sierra Alvarez on the Mesa
Central of southeastern San Luis Potosi and two in canyons
around the northern base of the Sierra Madre Oriental below
the Cumbres de Monterey of Nuevo Leon. A fourth locality
(sample 3) is represented by a fragmentary specimen, but the
limited data obtainable from it suggests it is a member of
the western group. The group differs from all other L. syl-
vaticum in lateral tubercle rows, and from southern Madrean

samples of L. sylvaticum in numbers of fourth toe lamellae
and numbers of paravertebrals larger than three dorsal scales
(Table 2).

KARYOTYPE. Chromosomal information was obtained
from 16 specimens (66, 92, 1 juv.) of L. sylvaticum'. three
from sample 1 1, one from 10, eight from 6, and four from
1. A total of 271 metaphase spreads were studied.

In all four populations the karyotype was found to consist
of a diploid number of 36 with 16 macrochromosomes and
20 microchromosomes (Fig. 13). There are five metacentric
to submetacentric (Nos. I, 2, 2A, 5, 7), two subtelocentric
(3, 4), and one acrocentric (9) pairs of macrochromosomes
(pair numbering after Bezy, 1972). No secondary constric-
tions were observed. The cells of six specimens (52, 1 juv.)
of sample 6 were found consistently to have a pair of het-
eromorphic chromosomes involving the largest pair of mi-
crochromosomes, with a metacentric member, ca. 1.5 times
the size of the next largest micro (Fig. 1 3).

The karyotype of L. sylvaticum appears identical in all
respects (except the heteromorphism) to that of L. micro-
pholis (Bezy, 1972). It differs from that of L. flavimaculatum,
L. tuxtlae, and L. pajapanensis in having one less pair of
macrochromosomes (the large metacentric 2A presumably
was formed from centric fusions involving pairs 6 and 7), a
pair 3 that lacks terminal satellites, and a submetacentric
rather than subtelocentric pair 7; from that of L. smithii in
that pair 2A is more metacentric (rather than submetacen-
tric), pair 3 lacks satellites, and the smallest macro pair is
acrocentric rather than subtelocentric; and from that of L.
occulor in having one more pair of macrochromosomes, one
less pair of microchromosomes, and an acrocentric (rather
than submetacentric) pair 9.

Lepidophyma sylvaticum thus differs karyotypically from

Contributions in Science, Number 349

Bezy: Systematics of Lepidophyma in Mexico 13

the species that are closest to it in scalation (L. flavimacu-
latum, L. smithii, and L. occulor) except L. micropholis. Such
chromosomal differences in themselves would not be ex-
pected to present reproductive barriers, and some cases of
extraordinary geographic variation in karyotypes have been
documented in species of other lizard families (e.g.. Hall and
Selander, 1973; Sites, 1983). However, among lizards chro-
mosomal divergence most often is associated with differen-
tiation at or above the species level. The karyotypic identity
of L. micropholis and L. sylvaticum serves to underscore the
morphologic and biogeographic relationships which suggest
that the former may be a troglodytic derivative of the latter,
and that future work might demonstrate a morphologic and
genetic continuum between the two.

The microchromosomal heteromorphism found in the fe-
males of sample 6 was not detected in samples 1, 10, and
1 1. It occurs in all females (five; plus one juvenile) of sample
6, but is absent in the two females of sample 1 and in all
males studied (two each from samples 1, 6, and 1 1, and one
from 1 0). It may constitute a sex chromosomal heteromorph-
ism (ZW) present in population 6, absent in population 1,
and of unknown occurrence in populations 10 and 11 (no
preparations from females available). On the other hand, it
may represent a heterozygous condition where unsampled
homozygous individuals (for the large macrochromosome)
occur in the population, or in adjacent populations. Until
additional material is obtained, all that can be said is that it
is a heteromorphic condition which has been found only in
females of population 6, and not detected in any other pop-
ulation in the family.

SEX RATIO. Two samples of L. sylvaticum have sex ra-
tios (2/<3 + 2) that differ significantly (0.05 level, Fisher exact
test, Yates correction) from 0.50: sample 6 with 0.89 (25/
28) and sample 10 with 0.85 (1 1/13). The skewed sex ratios
of these samples have been discussed earlier in relationship
to the unisexual populations of L. flavimaculatum occurring
in Panama and Costa Rica (Bezy, 1 972). Although the sample
sizes of populations 6 and 10 of L. sylvaticum remain smaller
than desirable, both are now sufficiently large to conclude
that the sex ratios differ significantly (0.05 level) from those
of other Lepidophyma populations in eastern Mexico: L.
tuxtlae (30/59) and L. gaigeae ( 1 50/260).

In addition to previously discussed factors that may be
responsible for the observed skewed sex ratios (Bezy, 1972),
temperature-dependent sex determination has now been doc-
umented for lizards (Bull, 1980), and comparable effects could
be operative in Lepidophyma. Theoretical considerations
would predict that this may not be the case in that temper-
ature-dependent sex determination is thought to interfere
with the evolution of both viviparity (ovoviviparity) and sex
chromosomes (Bull, 1980). Viviparity appears to be univer-
sal among xantusiids (Blackburn, 1982), and the population
(6) of L. sylvaticum with the most aberrant sex ratio is the
only one in the family with heteromorphism, possibly in-
dicating the presence of sex chromosomes. Experimental data
are needed to evaluate the role of environmental factors,
particularly temperature, in determining sex ratio in xantu-
siids.

The aberrant sex ratios and the heteromorphic chromo-

14 Contributions in Science, Number 349

somes of populations of L. sylvaticum could be a conse-
quence of hybridization. Both conditions frequently are found
in hybrid populations, the skewed sex ratios perhaps resulting
from a disruption in the balance of the sex-determining
mechanism or an increased expression of lethals in one of
the sexes (Darevsky et al., 1978; White, 1973). The identi-
fication of populations homozygous for the large microchro-
mosome and additional data (e.g., allozymes) would be re-
quired to establish the existence of and possible participants
in such a hybridization.

KEY TO THE SPECIES OF LEPIDOPHYMA
OF NORTHEASTERN MEXICO

la. Side of body lacking vertical rows of enlarged keeled
tubercles (Fig. 7), but with 43-50 subequal scales (axilla
to groin); less than 145 dorsal scales (occiput to rump);
tail with two interwhorls complete dorsally and ventrally

/.. gaigeae

lb. Side of body with enlarged, keeled tubercles arranged in
1 5-42 vertical rows (A-G) separated by smaller granular
scales (Figs. 7-8); 145 or more dorsal scales (O-R); tail
usually with more than 2 mterwhorls complete dorsally

2

2a. Total femoral pores 2 1 or less (gulars 59 or more; divided
4th toe lamellae 9 or less; lateral tubercle rows 24 or

less) L. occulor

2b. Total femoral pores 23 or more 3

3a. Dorsal scales (O-R) 231 or more (gulars 55 or more;
divided 4th toe lamellae 1 0 or more; lateral tubercle rows
27 or more) L. micropholis

3b. Dorsal scales (O-R) 2 1 7 or less (gulars 56 or less; divided
4th toe lamellae 4-23; lateral tubercle rows 15-38) . . .
L. sylvaticum

SPECIMENS EXAMINED

The 351 specimens and 52 population samples studied from
Mexico are listed below. Sample numbers are in parentheses
preceding localities.

L. gaigeae

HI DALGO: (2 1 ): La Placita, 8 km S Jacala (UIMNH 26 1 80-
86, 26191-99, 26204, 26207-09).

L. micropholis

SAN LUIS POTOSI: (15): 6 mi. E Valles (BCB 13837-42);
(16): 5.5 mi. S, 1.4 mi. E Valles (TCWC 60621, 60766-67).
TAMAULIPAS: (13): Gruta de Quintero, 1.5 mi. S Quintero
(AMNH 93409, LACM 66662, SAM 885); (14): cave at El
Pachon, ca. 5 mi. (by rd) NNE Antigua Morelos (LACM
106767-68. UAZ 28762, 28767-69, UMMZ 101299,
102886-88); 11.3 mi. S Ciudad Mante, Hwy 85 (TCWC
57256).

L. occulor

QUERETARO: (18): 2.5 mi. S Conca, Hda. Conca (TCWC
35605-06, 48499); (19): Jalpan (LJSNM 47134-35); (20): 1.2
mi. E Landa de Matamoros ( TCWC 2969 1 ); 1.5 mi. E Landa

Bezy: Systematics of Lepidophyma in Mexico

]J

(TCWC 33063). SAN LUIS POTOSI: (17): Boa Capulin
(LSUMZ 2379-80).

L. sylvaticum

HIDALGO: (11): 5.8 mi. (by Hwy 105) S Tianquistengo
(LACM 106741-48); 4.0 mi. (by Hwy 105) S Tianquistengo
(LACM 106721); 3 mi. S Tianquistengo (UIMNH 26230).
NUEVO LEON: (1): La Boca (KU 92612-13); ca. 7 km NE
Santiago, Presa La Boca (LACM 106781-792); (2): 5 mi. N
Las Ajuntas (EAL 4644). QUERETARO: ( 1 0): El Madrono,
3.5 mi. W (rd) El Lobo [and vie] (LACM 109771, SAM 1 104,
TCWC 29692-29707, 32291, 33064, 35607, UMMZ
129749). SAN LUIS POTOSI: (3): Buenavista (ca. 20 mi.
NE Cerritos) (AMNH 64025); (4): Alvarez (58 kilo) (MCZ
24507-08); between San Francisco and Alvarez (MCZ
157826); Valle de los Fantasmos (SDNHM 60482); (8): 27
km (by Mex 80) W El Naranjo (LACM 1 3 1 145-48); 3.8 mi.
(by Hwy 80) NNE Ciudad del Maiz (LACM 131144); 5 mi.
NE Ciudad del Maiz (TCWC 35582); (9): Huichihuagan
(FMNH 39631). TAMAULIPAS: (5): 8 mi. S, 6 mi. W Vic-
torio. Sierra Madre Oriental (KU 33992); (6): Rancho del
Cielo [and vie] (AMNH 107273, LACM 106751-60,
106762-65, LSUMZ 10989, UMMZ 101301, 101375,
102977-81, 109763-67); (7): Sierra de Tamaulipas, Santa
Maria (UMMZ 102889-90); 10 mi. W, 2 mi. S Piedra (KU
33993-94). VERACRUZ: (12): 4 km W Tlapacoyan (KU
26909).

L. flavimaculatum

CHIAPAS: (22): Palenque (LACM 65117-19); Ruinas de
Palenque (EAL 3030-31, FSM 32915-16, KU 94104-05);
San Juanito, Palenque (USNM 1 1 1486-87); (23): 4.5 km S
Pichucalco (KU 94106); (24): El Estoracan, ca. 50 km N
Cintalapa (AMNH 73468); (25): Chiapa, 1 mi. W (TNHC
27517-18); (26): Ocozocoautla Selvas El Ocote (MCZ 5432 1—
22); 16.1 mi. NW Ocozocoautla (LACM 61259); 26 km N
Ocozocoautla (UTEP 5367-68); 32 km NW Ocozocoautla,
Selva del Ocote (JFC); 25 mi. (by rd to Malpaso) NW Oco-
zocoautla (UAZ 28764, 28805-07); 12 km N Berriozabal
(UTEP 5365-66); (27): ca. 5 km S Solusuchiapa (UAZ 31635);
(28): Lago Miramar, near San Quintin (JFC); (29): Lacanja
(LACM 114244). OAXACA: (30): 2.8 mi. N Rio Sarabia
(UMMZ 1 15096); (31): 2 km S Tolosita (KU 39676); (32):
Rio Mono Blanco, Juchitan (UIMNH 36832); (33): La Gloria
(UIMNH 35515); (34): Mogone (UIMNH 40811); (35): 50.5
mi. S Acayucan, Hwy 185 (TNHC 25182). QUINTANA
ROO: (36): 4. 1 km NE Felipe Carrillo Puerto (UMRC 79-
252). TABASCO: (37): Teapa (LACM 61260-61, LSUMZ
6878-79, UIMNH 47883, UMMZ 1 13777); (38): Soledad
(UIMNH 47884). VERACRUZ: (39): 20 km E Jesus Car-
ranza (KU 24453); 25 km SE Jesus Carranza (KU 26920-
21); 35 km SW Jesus Carranza (KU 26919); (40): Rio de las
Playas (USNM 118638).

L. pajapanensis

VERACRUZ; (41): Sontecomapan, Los Tuxtlas [and vie]
(TCWC 21365, UAZ 28765, 28808-1 1, UTAR 3107, 31 10,

3116, TCWC 21365); Coyame, 9 mi. (by rd) SE Catemaco
(UAZ 28804); Univ. Mex. Biol. Exp. Sta., ca. 33 km ENE
Catemaco (TCWC 53351); Coloma de Bastonal, above Que-
zalapam (TCWC 19133); Laguna Catemaco, nr Cuezalapan
(UMMZ 126363-64); 4 mi. SETebanca, Los Tuxtlas (TCWC
21364); S slope Volcan San Martin (KU 97290, UMMZ
1 18220, 126362); (42): 35 km SE Jesus Carranza (KU 269 1 3).

L. smithii

CHIAPAS: (43): La Esperanza (UIMNH 10952-56, 10958-
59, 10963, 10965, 10968-69, 10970-71, 10975-79, 10997-
98); (44); Tonala (UIMNH 26227-29). GUERRERO: (45):
2 km W Puerto Marquez (CU 9676-79, 9692-93, 9772,
LACM 128590, 130027-29).

L. tarascae

MICHOACAN: (46): near Mexiquillo, Aquila District (ENCB
9221-22, LACM 134226).

L. tuxtlae

CHIAPAS: (47): 25 mi. (by rd to Malpaso) NW Ocozocoautla
(UAZ 28780-82). OAXACA: (48): Vista Hermosa (KU
87396-98); 30 mi. (by rd) NE Llano de las Flores (UMMZ
125870); (49): Mts nr La Gloria (UIMNH 37236); (50): Finca
San Carlos, Matias Romero Oaxaca (FSM 32918). VERA-
CRUZ: (51): Volcan San Martin (TCWC 22102-03, TNHC
29792-93, UIMNH 80695-99, UMMZ 1 18219, 121165,
122112, 126360-61); S slope Volcan San Martin Tuxtla (KU
59560); Rancho El Tular, 1 5 mi. N San Andres Tuxtla (USNM
139731); Rio Tecolapan, 2.4 mi. NNW Tapalapan (UMMZ
115098-99); Salto de Eyipantla (TCWC 19134); Montepio
(FSM 32917); Sontecomapan [and vie] (CM 41470, FSM
32914, TCWC 19135, 26717, UAZ 28770-79, UTAR 3101,
3103-04, 3108-09, 31 1 1-13, 3115, 3127); 7.7 mi. NW Son-
tecomapan (UTAR 3728-30, 3733-34); E of Lago Catemaco,
1 2.7 mi. from Catemaco by rd (LACM 106795); 1 8 mi. NNE
Catemaco (JCL 67); Univ. Mex. Bio. Exp. Sta., ca. 33 km
ENE Catemaco (TCWC 53352—53); between Laguna Cate-
maco and Volcan Martin (UMMZ 121 166); Coyame (UAZ
28763); midway between Coyame and Tebanca (UMMZ
121164); 4 mi. SE Tebanca (TCWC 21366); 5.6 mi. ESE
Tebanca (UTAR 3156); Rio Quetzalapan [and vie] (TCWC
19136, 21367-69, UTAR 3133, 3139-40); Coloma de Bas-
tonal (TCWC 19137); Dos Arroyos, 5 mi. E Zapoapan (TCWC
21370-71); (52): 25 km SE Jesus Carranza (KU 26912).

ACKNOWLEDGMENTS

I thank the following individuals for assistance with aspects
of the study: Kathryn Bolles, Salvador Contreras B., Robb
Dean, James R. Dixon, Jerry D. Johnson, Carl S. Lieb, Rob-
ert I. Lonard. James F. Lynch, Barbara T. Warburton, and
John W. Wright for held assistance and/or collecting live
material; Ticul Alvarez and officials of the Fauna del Silvestre
for granting the necessary collecting permits; John P. Karges
and John W. Wright for discussions, advice, and/or review
of the manuscript; John DeLeon for photography; Kathryn

Contributions in Science, Number 349

Bezy: Systematics of Lepidophyma in Mexico 15

Bolles for illustration; Beatriz Larrain for translation; Terri
Togiai for typing the manuscript; and C.J. Cole (AMNH),
B.C. Brown (BCB), A.E. Leviton (CAS), C.J. McCoy (CM),
J.M. Savage (CRE), F.H. Pough (CU), E.A. Liner (EAL), T.
Alvarez (ENCB), R.F. Inger, H. Marx, ELK. Voris (FMNH),
W. Auffenberg (FSM), W.E. Duellman (KU), J.F. Copp (JFC),
D.A. Rossman (LSUMZ), P. Alberch, E.E. Williams (MCZ),
FEW. Greene, D.B. Wake (MVZ), T.M. Uzzell (PAS), S.A.
Minton (SAM), G.K. Pregill (SDNHM), J.R. Dixon (TCWC),
R.F. Martin (TNHC), C.H. Lowe (UAZ), D.F. Hoffmeister
(UIMNH), A.G. Kluge (UMMZ), J.C. Lee (UMRC), G.R.
Zug (USNM), W.F. Pyburn (UTA), and R.G. Webb (UTEP)
for loan of specimens under their charge (museum abbre-
viations follow Leviton et al., 1980).

LITERATURE CITED

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. 1973. A new species of the genus Lepidophyma

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Bezy, R.L., R.G. Webb, and T. Alvarez. 1982. A new species
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Werler, J.E. 1957. A new lizard of the genus Lepidophyma
from Volcan San Martin Pajapan, Veracruz. Herpeto-
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Edition. Cambridge University Press, London.

Submitted 16 May 1983; accepted 13 September 1983.

16 Contributions in Science, Number 349

Bezy: Systematics of Lepidophyma in Mexico

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PilMli

MEGAPALEONTOLOGY OF THE EOCENE LLAJAS FORMATION,
SIMI VALLEY, CALIFORNIA

Richard L. Squires

Contributions in Science, Number 350
Natural History Museum of Los Angeles County
13 July 1984

ISSN 0459-8113

Natural History Museum of Los Angeles County
900 Exposition Boulevard
Los Angeles, California 90007

CONTENTS

ABSTRACT 1

INTRODUCTION 1

HISTORICAL REVIEW 2

MEGAFOSSILS 2

DEPOSITIONAL ENVIRONMENTS AND STRATIGRAPHIC DISTRIBUTION

OF MEGA FOSSILS 4

PAI.EOCL.IMATE 7

AGE 9

CORRELATION 1 1

BIOGEOGRAPHY 1 1

SYSTEMATIC MATERIALS AND METHODS 13

SYSTEMATICS 13

Phylum Protista 13

Order Foraminiferida 13

Family Discocyclinidae 13

Pseudophragmina ( Proporocyclina ) clarki (Cushman, 1920) 14

Phylum Coelenterata 14

Order Scleractinia 14

Family Caryophylliidae 14

ITrochocyathus striatus (Gabb, 1864) 14

Turbino/ia dickersoni Nomland, 1916 15

Phylum Brachiopoda 15

Order Terebratulida 1 5

Eogryphus tolmani Hertlein and Grant, 1944 15

Phylum Annelida 1 5

Order Sedentaria 1 5

Family Serpulidae 15

Rotularia tejonense ( Arnold, 1910) 15

Phylum Mollusca 15

Class Scaphopoda 15

Family Dentaliidae 15

Dentalium stentor Anderson and Hanna, 1925 16

Dentalium (Laevidentalium) calafium Vokes, 1939 16

Class Gastropoda 16

Order Archaeogastropoda 16

Family Neritidae 16

Nerita cf. N. ( Amphinerita ) eorex Vokes, 1939 16

Velates perversus ( Gmelin, 1791) 16

Order Mesogastropoda 17

Family Turritellidae 17

Turrit ella meganosensis protumescens Merriam and Turner, 1937 17

Turrite/la andersoni Dickerson, 1916 18

Turritella andersoni lawsoni Dickerson, 1916 18

Turritella buwa/dana Dickerson, 1916 18

Turritella uvasana infera Merriam, 1941 19

Turritella uvasana applinae Hanna, 1927 19

Family Architectonicidae 19

Architectonica ( Architectonica ) l/ajasensis Sutherland, 1966 19

Architectonica ( Stellaxis ) cognata Gabb, 1864 19

Family Cerithiidae 20

Benoistia umpquaensis Turner, 1938 20

Family Cerithiopsidae 20

Cerithiopsis llajasensis n. sp 20

Family Epitoniidae 21

Cirsotrema sp 21

Family Calyptraeidae 21

Calyptraea diegoana (C onrad, 1855) 21

Family Xenophoridae 21

Xenophora stocki Dickerson, 1916 21

Family Strombidae 21

Ectinochilus (Macilentos) macilentus (White, 1889) 21

Family Seraphsidae 23

Paraseraphs erraticus (C ooper, 1894) 23

Family Cypraeidae 23

Cypraea castacensis Stewart, 1927 23

Family Naticidae 24

Eocernina hannibali Dickerson, 1914 24

Tejonia moragai (Stewart, 1927) 24

Pachycrommium clarki (Stewart, 1927) 24

Polinices (Euspira) nuciformis (Gabb, 1864) 25

Neverita (Neverita) globosa Gabb, 1 869 25

Sinum obliquum (Gabb, 1864) 26

Natica ( Naticarius ) aff. N. (TV.) uvasana Gabb, 1864 26

Family Cassididae 26

Galeodea ( Cahagaleodea) californica Clark, 1942 26

Galeodea (Gomphopages) susanae Schenck, 1926 27

Phalium (Semicassis) tuberculiformis ( Hanna, 1924) 27

Family Cymatiidae 27

Cymatium (Septa) janetae Squires, 1983a 27

Ranella katherineae Squires, 1983a 29

Family Bursidae 29

Olequahia domenginica (Vokes, 1939) 29

Rane/lina pilshryi Stewart. 1927 29

Family Ficidae 29

Ficopsis cooperiana Stewart, 1927 29

Ficopsis remondii crescentensis Weaver and Palmer, 1922 30

Order Neogastropoda 30

Family Muricidae 30

Laevityphis (Laevityphis) antiquus (Gabb, 1864) 30

Family Buccinidae 30

A ncistrolepisl carolineae n. sp 30

Family Nassariidae 31

Molopophorus cretaceus (Gabb, 1864) 31

Family Fasciolariidae 31

Clavilithes tabulatus (Dickerson, 1913) 31

Clavilithes n. sp. A Clark and Vokes, 1936 31

Clavilithes n. sp. B Squires, 1983a 31

Fusinus teglandae Hanna, 1927 33

Fusinus aff. F. ucalius X okes, 1939 33

Family Olividae 33

Pseudoliva lineata Gabb, 1864 33

Strepsidura ficus (Gabb, 1864) 33

Ancilla (Spirancilla) gabbi Cossmann, 1899 34

Olive/ 1 a mathewsonii Gabb, 1864 34

Family Mitridae 34

Proximitra ? cretacea (Gabb, 1864) 34

Family Tudiclidae 34

PseudoperissolaxblakeipraeblakeiXoV.es, , 1939 34

Family Harpidae 35

Eocithara mutica californiensis (Vokes, 1937) 35

Family Voiutidae 35

Cryptochorda (Cryptochorda) californica (Cooper, 1894) 35

Lyria andersoni Waring, 1917 35

Lyrischapa lajollaensis ( Hanna, 1927) 36

Family Cancellariidae 36

Bonellitia (Admetula) paucivaricata (Gabb, 1864) 36

Family Turridae 36

Pleurofusia fresnoensis (Arnold, 1910) 36

Fusiturricula ( Crenaturricula ) crenatospira (Cooper, 1894) 36

Fusiturricula ( Crenaturricula ) crenatospira domenginica Vokes, 1939 . . 37

Surculites mathewsonii (Gabb, 1864) 37

Domenginelia claytonensis ( Gabb, 1864) 37

Fxilia ilajasensis Bentson, 1940 39

Family Conidae 39

Conus caleocius \ okes, 1939 39

Conus hornii umpquaensis Turner, 1938 39

Family Terebridae 39

Terebra calij arnica Gabb, 1869 39

Order Cephalaspidea 39

Family Cylichnidae 39

Cylichnina tantilla (Anderson and Hanna, 1925) 40

Scaphander (Mirascapha) costatus (Gabb, 1864) 40

Family Philinidae 40

Megistostoma gabbianum ( Stoliczka, 1868) 40

Class Bivalvia 40

Order Nuculoida 40

Family Nuculidae 40

Acila (Truncaci/a) decisa (Conrad. 1855) 41

Family Nuculanidae 41

Nuculana (Saccella) gabbii (Gabb, 1864) 41

Order Arcoida 41

Family Glycymerididae 41

Glycymeris (Glycymeris) roseca nyonensis Hanna, 1927 41

Glycymeris (Glycymerita) sagittata ( Gabb, 1864) 42

Order Mytiloida 42

Family Mytilidae 42

Brachidontes (Brachidontes) cowlitzensis (Weaver and Palmer, 1922) .. 42

Family Pinnidae 42

Pinna lewisi Waring, 1917 42

Pinna Ilajasensis Squires, 1983a 43

Order Pterioida 43

Family Malleidae 43

Nayadina (Exputens) Ilajasensis (Clark, 1934) 43

Family Spondylidae 43

Spondylus carlosensis Anderson, 1905 43

Family Anomiidae 45

Anomia mcgoniglensis Hanna, 1927 45

Family Ostreidae 45

Ostrea idriaensis Gabb, 1869 45

Order Veneroida 45

Family Lucinidae 45

Claibornites diegoensis { Dickerson, 1916) 45

Family Carditidae 46

Venericardia (Pacifcor) hornu calafia Stewart, 1930 46

Venericardia ( Pacificor ) aragonia joaquinensis (Vokes, 1939) 46

Glyptoactis (G/yptoactis) domenginica (V okes, 1939) 46

Family Crassatellidae 47

Crassatella uvasana Conrad, 1855 47

Family Cardiidae 49

Acanthocardia (Schedocardia) breweru (Gabb, 1864) 49

Nemocardium linteum (Conrad, 1855) 49

Family Solenidae 50

Solena (Eosolen) novacularis (Anderson and Hanna, 1928) 50

Family Tellinidae 50

Macoma rosa Hanna, 1927 50

Family Psammobiidae 50

Gari cf. G. eoundulata Yokes. 1939 50

Family Veneridae 50

Callista ( Costacallista ) cf. C. (C.) hornii (Gabb, 1864) 51

Callocardia (Nitidavenus) tejonensis (Waring, 1914) 51

Pitar (Calpitaria) uvasanus ( Conrad, 1855) 51

Pitar (Lamelliconcha) joaquinensis Vokes, 1939 51

Marcia {Mercimonia) bunkeri (Hanna, 1927) 53

Order Myoida 53

Family Corbulidae 53

Corbula (Caryocorbula) dickersoni ( Weaver and Palmer, 1 922) 53

Family Teredinidae 53

Teredo ? sp 53

Order Pholadomyoida 54

Family C’uspidariidae 54

Cardiomya aff C. russelli (Hanna, 1927) 54

Class Cephalopoda 54

Order Nautilida 54

Family Aturiidae 54

Aturia myrlae Hanna, 1927 54

Order Sepiida 55

Family Indeterminate 55

Spirulimorph sepiid 55

Phylum Arthropoda 55

Infraorder Brachyura 55

Family Goneplacidae 55

Glyphithyreus weaveri (Rathbun, 1926) 55

Phylum Kehinodermata 55

Order Spatangoida 56

Family Schizasteridae 56

Schizaster diabloensis Kew, 1920 56

Phylum Chordata 56

Order Lamniformes 56

Family Odontaspididae 56

Odontaspis sp 56

Family Lamnidae 57

Isurus cf. /. praecursor (Leriche, 1906) 57

LOCALITIES 57

ACKNOWLEDGMENTS 66

LITERATURE CITED 67

MEGAPALEONTOLOGY OF THE EOCENE LLAJAS FORMATION
SIMI VALLEY, CALIFORNIA

Richard L. Squires1

ABSTRACT. The paleontology and stratigraphic distribution of the
megafossils in the late early through early middle Eocene age Llajas
Formation, Simi Valley, Los Angeles and Ventura Counties, south-
ern California, are described in detail. One hundred and seven species
and subspecies, representing 88 identifiable genera, are recorded for
132 localities. Twenty-six of these species are previously unreported
from the Llajas Formation.

Illustrations, synonymies, primary type material information. West
Coast molluscan stage ranges, geographic distributions, local occur-
rences, and remarks are provided for the taxa, which include one
large foraminifer, two corals, one brachiopod, one annelid, two
scaphopods, 65 gastropods, 29 bivalves, one nautiloid, one sepiid,
one brachyuran, one spatangoid, and two sharks. The sepiid can be
identified only to the family level. A new species of Cerithiopsis and
a new species of Ancistrolepis! are named. Crassatella uvasana and
C. u. semidentata are shown to be identical.

Most of the megafossils occur in channel-fill deposits in shallow-
marine strata but have undergone minimal postmortem transport.
The Llajas megafauna lived in warm waters less than 80 m depth.

Mollusks indicative of the lower of the two faunal zones of the
West Coast provincial molluscan “Capay Stage” (lower Eocene) are
present in the lowermost fossil-bearing beds of the Llajas Formation.
The remaining fossil-bearing part of the Llajas contains mollusks
indicative of the “Domengine Stage” (upper lower through lower
middle Eocene). Age refinement of the Llajas extends the molluscan
stage ranges of several molluscan species. The Llajas megafauna is
similar to those in late early-early middle Eocene age formations
from San Diego, California through southwestern Oregon. These
megafaunas were in the same faunal province that extended, at least,
from southern California into Washington. Most of the genera were
already present on the West Coast by Llajas time and several species
were carry-overs from earlier times. A few genera were early Eocene
immigrants from the Caribbean and Old World Tethyan regions,
and some were early middle Eocene immigrants from the same
regions. Several genera originated on the West Coast during the early
middle Eocene.

INTRODUCTION

Since the early 1900’s, paleontologists have collected mega-
fossils from the late early through early middle age Eocene
Llajas Formation in the Simi Valley area, Los Angeles and

Contributions in Science, Number 350, pp. 1-76
Natural History Museum of Los Angeles County, 1984

Ventura Counties, southern California. The fossils are pre-
dominantly mollusks and nearly every paleontological mu-
seum on the West Coast has a representative collection. To
date, however, there has been no comprehensive taxonomic
and stratigraphic analysis of the Llajas megafauna. Research-
ers, therefore, have been hindered in performing detailed
comparative studies on the taxonomy, biogeography, and
time-correlation of the Llajas megafossils. The objectives of
this paper are 1) to tabulate the species and show their dis-
tribution within the formation, 2) to provide synonymies
and illustrations of the species, 3) to provide taxonomic re-
finements of certain known taxa and describe two new gas-
tropods, 4) to interpret the paleoenvironment and biogeog-
raphy of the megafauna, 5) to refine the age assignment and
correlation of the Llajas Formation, and 6) to refine the West
Coast molluscan stage ranges of the taxa.

The Llajas Formation crops out in the southwestern Santa
Susana Mountains and along the south side of Simi Valley
(Fig. 1 ). It disconformably overlies the late Paleocene through
earliest Eocene age marine Santa Susana Formation and,
except where local faults occur, is unconformably overlain
by the early late Eocene through Oligocene age nonmarine
Sespe Formation (Figs. 2 and 3).

Outcrops of the Llajas Formation are typically extensively
covered. Best exposures are at the 545-m-thick type section
in the southwestern Santa Susana Mountains (Fig. 1 ). In 1981
and 1 983b, I reported that the Llajas Formation in the south-
western Santa Susana Mountains consists mostly of a trans-
gressive (retrogradational) sequence of facies that grades ver-
tically from coastal alluvial fan, to shallow marine, to outer
shelf and slope with incised channels. The outer shelf and
slope facies was subsequently covered by regressive (progra-
dational) shallow-marine facies. The regressive sequence is

1. Department of Geological Sciences, California State Univer-
sity, Northridge, California 91330, and Research Associate, Inver-
tebrate Paleontology, Natural History Museum of Los Angeles Coun-
ty.

ISSN 0459-8113

118° 40'

incomplete due to erosional truncation by the Sespe For-
mation (Fig. 2).

HISTORICAL REVIEW

In the last 70 years, starting with Waring (1914), there have
been many investigations dealing with the megafossils of the
Llajas Formation. These previous studies have included par-
tial faunal lists (Waring, 1917; Clark, 1926; Hanna, 1927;
McMasters, 1932; Cushman and McMasters, 1936; Turner,
1938; Vokes, 1939; Lewis, 1940; Seiden, 1972), descriptions
of new taxa (Waring, 1914, 1917; Schenck, 1926; Stewart,
1927, 1930; Clark, 1934, 1942; Merriam and Turner, 1937;
Vokes, 1937, 1939; Bentson, 1940; Merriam, 1941; Hertlein
and Grant, 1944; Sutherland, 1966; Squires, 1983a), and
taxonomic refinements or miscellaneous comments on known
species. In this last category, the most notable papers are
those of Schenck (1931), Gardner and Bowles (1934), Clark
and Vokes (1936), Turner (1938), Vokes (1935, 1939), Mer-
riam (1941), Hanna and Hertlein (1943), Verastegui (1953),
Marincovich (1977), Givens (1979), Squires (1979), and Saul
(1983).

As the name “Llajas Formation” was not formalized until
1936 by Cushman and McMasters, earlier workers referred
to such strata by various names (Fig. 4). Clark and Vokes
(1936), Turner (1938), and Vokes (1939) used the terms
“Lower Llajas” and “Upper Llajas” but did not define them.

Various aspects of the microfossils of the Llajas Formation
have been reported on by McMasters (1932), Cushman and
McMasters (1936), Laiming ( 1 940a, 1940b, 1943), Mallory
(1959), Schymiczek and Squires (1981), Schymiczek (1983a,
1983b), and Filewicz and Hill (1983).

MEGAFOSSILS

One hundred and seven taxa, more than 90 percent of which
are mollusks, were identified from the Llajas Formation.
Taxa identified to species and subspecies include 64 gastro-
pods, 28 bivalves, two scaphopods, two solitary corals, and
one species each of large foraminifer, brachiopod, annelid,
nautiloid, brachyuran, and spatangoid. One gastropod, one
bivalve, and two sharks are identified only to genus. A spi-
rulimorph sepiid could only be identified to the family level.
All of these taxa are illustrated in Figures 5 through 1 3. Other

2 Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks

NONMARINE

Figure 2. Stratigraphic column of the Llajas Formation, showing
depositional environments based on Squires (1981, 1983b).

taxa, too poorly preserved for even generic determination,
include a bryozoan, two brachiopods, calcareous worm tubes,
two gastropods, a nautiloid, a myliobatoid, and scattered
Teredo! -bored wood fragments.

The identifications of Llajas species and subspecies studied
in this report are based on published figures and descriptions
and selected comparisons with type specimens and non-type
specimens on deposit at 1) Department of Earth and Space
Sciences, University of California at Los Angeles, 2) Uni-
versity of California, Museum of Paleontology, Berkeley, 3)
Natural History Museum of Los Angeles County, and 4)
California State University, Northridge.

Megafossils were collected at 121 localities in the south-
western Santa Susana Mountains and 1 1 localities along the
south side of the Simi Valley. All of the localities are de-
scribed in the “Localities” section, and the relative strati-

Figure 3. Correlation of the Llajas Formation with Eocene West
Coast molluscan stages (after Saul, 1983); millions of years before
present (M.Y.B.P.) scale, epochs, subepochs, standard ages, planktic
foraminifera zones, and calcareous nannoplankton zones (all after
Berggren, Kent, and Flynn, in press); and West Coast benthic fo-
raminifera stages (after Poore, 1980).

graphic position of each one is shown in Figures 14 through
19.

Species and subspecies with holotype localities in the Llajas
Formation are listed in Table 1. Species not previously re-
ported from the Llajas Formation are listed in Table 2.

Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks 3

Vyvvv^v/ Unconformity Conformable contact

Nature of contact not discussed

* Present only on south side of Simi Valley

Figure 4. Comparative concepts of Paleogene stratigraphy in the
Simi Valley area (after Squires, 1983b).

Some megafossil species previously reported (Waring, 1914,
191 7; Clark, 1921:158-159, 1926:1 14-1 16; Clark, 1942;Kew,
1924:25, 29; Hanna, 1927:260; Vokes, 1935, 1939:23-26,
30-31; Clark and Vokes, 1936; Turner, 1938:33-37; Bent-
son, 1940; Ingram, 1940) from the Llajas Formation (or
generally equivalent strata) were not found during the present
study. There are serious problems in trying to substantiate
the records of these species because 1) locality information
is either lacking or vague, and/or 2) previously used for-
mation names (i.e., Meganos and Domengine) are not the
exact equivalents of the Llajas Formation (Fig. 3). In addi-
tion, because these species names usually have only been
listed, without illustration or reference to catalogued museum
specimens, there is no way to verify the identifications. Some
of these so-called “Llajas” species can be shown to be from
the underlying late Paleocene through earliest Eocene age
Santa Susana Formation. Based on analysis of Zinsmeister’s
(1974, 1983a) work on the megafauna of the Santa Susana
Formation, the following species are from this unit: Cucul-
laria morani [=Cucullaea], mentioned by Waring (1914,
1917); Polinices hornii, Mesalia martinezensis [ = Turritella ],
Turritella infragranulata, and Septifer elegans, mentioned
by Waring (1917). Clark (1942) reported that the type locality
of Chedevi/lea stewart i and Tibia (Eotibia) Uajasensis is in
the lower portion of the Llajas Formation. According to Keen
and Bentson (1944:143), this locality (University of Califor-
nia, Museum of Paleontology, Berkeley, locality 7015) plots
within Paleocene strata as mapped by Nelson (1925).

The marine strata of the Llajas Formation have been thor-
oughly bioturbated, but the only identifiable trace fossils are
Ophiomorpha, Thalassinoides, and Chronditesl. The first two
ichnogenera are fairly common in the shallow-marine facies,
especially in certain beds where this facies interfingers with
the coastal alluvial-fan facies. An Ophiomorpha burrow in
one of these beds is figured by Squires (1981:930, fig. 6C;

Table 1. Megafossils with holotype localities in the Llajas For-
mation.

Brachiopoda

Eogryphus tolmani Hertlein and Grant, 1944
Gastropoda

A ncistrolepisl carolineae n. sp.

Architectonica ( Architectonica ) Uajasensis Sutherland, 1966
Cerithiopsis Uajasensis n. sp.

Conus caleocius Vokes, 1939
Cymatium (Septa) janetae Squires, 1983
Eocithara mutica californiensis (Vokes, 1937)

Exilia Uajasensis Bentson, 1940

Fusiturricula (Crenaturricula) crenatospira domenginica
Vokes, 1939

Galeodea (Caliagaleodea) californica Clark, 1942
Galeodea ( Gomphopages ) susanae Schenck, 1926
Lyria andersoni Waring, 1917
Pachycrommium clarki (Stewart, 1927)

Ranella katherineae Squires, 1983

Tectarius ligniticus Vokes, 1939 (see Benoistia umpquaensis)
Turritella andersoni lawsoni secondaria Merriam, 1941
Turritella andersoni susanae Merriam, 1941
Turritella meganosensis protumescens Merriam and
Turner, 1937

Turritella uvasana etheringtoni Merriam, 1941
(see T. u. applinae)

Turritella uvasana infera Merriam, 1941
Bivalvia

Nayadina (Exputens) Uajasensis (Clark, 1 934)

Callocardia (Nitidavenus) tejonensis (Waring, 1914)

Pinna lewisi Waring, 1917

Pinna Uajasensis Squires, 1983

Venericardia (Paciftcor) hornii calafia Stewart, 1930

1983b, fig. 6E). Chronditesl is common in the outer shelf
and slope facies.

DEPOSITIONAL ENVIRONMENTS AND
STRATIGRAPHIC DISTRIBUTION
OF MEGAFOSSILS

Based on sedimentologic and lithologic features of the Llajas
Formation, on comparative studies of modem and ancient
sedimentary sequences, and ecology of representative genera
of mollusks and benthic foraminifers, the following facies
have been recognized in the Llajas Formation: coastal allu-
vial fan, shallow marine, and outer shelf and slope with in-
cised channels. In a vertical sense, most of the formation is
transgressive (retrogradational) in that shallower facies are
overlain by deeper facies. The uppermost part of the for-
mation, however, represents a regressive (progradational)
event in which the outer shelf and slope facies and associated

4 Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks

Table 2. Megafossils previously unreported from the Llajas For-
mation.

Anthozoa

ITrochocyathus striatus (Gabb, 1 864)

Turbinolia dickersoni Nomland, 1916

Annelida

Rotularia tejonense (Arnold, 1909)

Scaphopoda

Dentalium ( Laevidentalium ) calaftum Vokes, 1939
Dentalium stentor Anderson and Hanna, 1925

Gastropoda

Ancistrolepis ? carolineae n. sp.

Cerithiopsis llajasensis n. sp.

Cirsotrema sp.

Laevityphis ( Laevityphis ) antiquus (Gabb, 1864)

Molopophorus cretaceus (Gabb, 1864)

Nerita cf. N. (Amp hi writ a) eorex Vokes, 1939
Neverita ( Neverita ) globosa Gabb, 1 869
Olivella mathewsonii Gabb, 1 864
Terebra California? Gabb, 1869

Bivalvia

Anomia mcgoniglensis Hanna, 1927
Cardiomya aff. C. russet 1 1 (Hanna, 1927)

Corbula ( Caryocorbula) dickersoni Weaver and Palmer, 1922

Gari aff. G. eoundulata Vokes, 1939

Glycymeris (Glycymeris) rosecanyonensis Hanna, 1927

Macoma rosa Hanna, 1927

Marcia (Mercimonia) bunkeri (Hanna, 1927)

Pilar (Calpitaria) uvasanus (Conrad, 1855)

Spondylus carlosensis Anderson, 1905
Teredo? sp.

Vertebrata

Isurus cf. I. praecursor (Leriche, 1906)

Odontaspis sp.

turbidite-filled incised channels were covered by shallow-
marine facies (Squires, 1981, 1983b) (Fig. 2).

The coastal alluvial-fan facies, equivalent to the basal con-
glomerate depicted in Figure 1, is unfossiliferous. The zone
of interfingering between this facies and the shallow-marine
facies contains fairly abundant mollusks. Many of these mol-
lusks are the same species that occur in the stratigraphically
higher shallow-marine facies. Some of the mollusks, how-
ever, are restricted in their occurrence in the Llajas For-
mation to this zone of interfingering. These mollusks are
Nerita cf. N. ( Amphi nerita ) eorex, V elates perversus, Turri-
tella meganosensis protumescens, Turritella andersoni, and
Venericardia ( Pacificor ) aragonia joaquinensis. Unidentifi-
able ostreid remains also characterize this zone of interfin-
gering. At CSUN locality 542, such remains form an ostreid
coquinite. Turritella meganosensis protumescens typically
occurs as robust, thick-shelled fragments up to 5.5 cm in
length. This taxon belongs to the T. reversa stock of Merriam
(1941). According to Saul (1983), the presence of large mem-
bers of this stock, as well as the presence of Venericardia
(Pacificor) aragonia joaquinensis, in the Llajas Formation

I’able 3. Explanation of reference abbreviations for Tables 4 and
5.

A1

Abbott, 1968a

A2

Abbott, 1974

A&D

Abbott and Dance, 1982

C

Cossmann, 1901

D

Dance, 1976

D&D

Dell and Dance, 1963

E

Eisenberg, 1981

G

Grasse, 1968

Ha

Habe, 1968

H&G

Hertlein and Grant, 1972

K

Keen, 1971

K&C

Keen and Coan, 1974

Ki

Kira, 1965

L

Lindner, 1978

Ma

Marincovich, 1977

Me

McMillan, 1968

Me

Merriam, 1941

Ml

Morris, 1966

M2

Morris, 1975

N 1

Nordsieck, 1968

N2

Nordsieck, 1969

O

Olsson, 1961

R

Rios, 1970

S

Schenck, 1936

T

Tebble, 1976

W&D

Weaver and duPont, 1970

indicate nearshore, shallow-water conditions. The presence
of ostreid coquinites indicates similar conditions. Nerita oc-
curs along shorelines today (Tables 3 and 4), and Velates, a
closely allied but extinct genus, probably occurred in the same
type of environment.

Most of the fossils in the formation occur in the shallow-
marine facies proper, which can be divided into a transgres-
sive phase and a regressive phase. The transgressive phase
makes up most of the lower part of the formation. It consists
of alternating laminated and bioturbated sandstone with scat-
tered fossiliferous beds mostly confined to channels. The
laminated sandstone represents storm-influenced stratifica-
tion and the biogenic reworking is inter-storm activity
(Squires, 1981). These deposits probably accumulated where
the coastal alluvial-fan rivers emptied into the ocean. The
resulting sediment buildup would have been subjected to
reworking by waves (Squires, 1 983b). The fossils have under-
gone a small distance of postmortem transport and constitute
indigenous death assemblages (Squires, 1981). Many fossils
show preservation of delicate features, such as protoconchs,
outer lips, and ribs. There is much variability in the taxo-
nomic composition of the fossils in these channel-fill depos-
its. At many localities, only one or two species of megafossils

Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks 5

Table 4. Recent bathymetry of extant molluscan genera of the Llajas Formation.

Genus (subgenus)

Depth range
(m)

*MFR depth
(m)

**References

Scaphopoda

Dentalium

4-2320

7-100

A2,K,K&C,M2,R

Gastropoda

Ancilla

6-87

21-70

A2,R

Ancistrolepis

40-200

100

K

Architectonica

0-134

2-37

A2,K

Calyptraea

0-70

0-37

A1,M1,M2,K

Cerithiopsis

0-823

24-60

A2.M1

Cirsotrema

33-274

73-137

A2,Ha,Ki

Conus

0-550

9-55

K,M 1 ,R

Cymatium (Septa)

0-146

24-60

A2.M1.M2

Cypraea

0-140

9-24

A1,M1,M2,R

Fusinus

0-3616

18-60

A 1 ,A2,K,K&C,Ki,M 1 ,M2

Fusiturricula

56-65

60

R

Galeodea

7-330

8

Mc.Nl

Lyria

0-133

2-24

Ha,K,Ki,W&D

Natica (Naticarius)

0-280

32-60

Ma,R

Nerita

0-9

0-9

K,M1,M2

Never it a (Never it a)

1 1-2860

658-1281

M

Olivella

0-90

0-27

A1,A2,K,K&C,R

Phalium ( Semicassis )

18-100

37-55

Ha.Ki

Polinices (Euspira)

0-4794

15-200+

Ma,M 1

Pseudoliva

0-3

0-3

C,L

Ranella

55-915

110

D&D

Scaphander

20-5200

50-160

A1.A2.K

Sinum

0-170

24-48

K,K&C,Ma,R

Terebra

0-280

4-18

K,M 1

Turritella

4-185

26-56

K,K&C,Ki,Ml

Xenophora

0-200

20-144

A1 ,A2,Ha,K,Ki,M2,R

Bivalvia

Acanthocardia

0-360

5

N2,T

Acila (Truncacila)

7-1256

20-100

A1,A2

Anomia

0-550

0-9

A2,K,M1,M2

Brachidontes

0-31

0-24

K,M1,M2,R

Callista (Costacallista)

30-214

45-100

A2,R

Cardiomya

4-2400

60-190

K,Ml,M2,Ha,R

Corbula (Caryocorbula)

0-823

10-27

A2,K

Gari

0-150

10-50

A2,Ha,Ml

Glycymeris

0-110

9-60

A1,K,M1,M2

Macoma

0-1545

0-45

A1,K&C,M1,R

Nemocardium

9-640

25-90

A2,Ha,H&G,K,Ki,M 1 ,K.&C

Nuculana (Saccella)

2-1097

10-80

A2,K

Ostrea

0-91

0-24

H&G,K,M1,M2

Pinna

0-200

0-24

A2,Ha,K,Ml,M2

Pitar (Lamelliconcha)

0-110

30-80

K,0

Solena

Nearshore

K

Spondylus

3-140

15-50

A1,K,M1,M2,R

Teredo

Bores into wood

A2,M2

* Most frequently reported; ** see Table 3 for abbreviations.

6 Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks

are present. Turritella andersoni lawsoni is usually one of
these species. At other localities, such as CSUN locality 371,
as many as 45 species of megafossils have been found.

The regressive phase of the shallow-marine facies makes
up the uppermost part of the formation. It consists of bio-
turbated silty sandstone with minor occurrences of laminated
sandstone and scattered fossiliferous units. These deposits
were in slightly deeper waters than those of the transgressive
phase and were less affected by storm reworking (Squires,
1983b). Fossils are unabraded, and at CSUN locality 475
articulated Pinna lewisi specimens were found. The fossils
in the regressive phase are also interpreted to be indigenous
death assemblages.

Where the sandstone of the transgressive and regressive
phases of the shallow-marine facies grades into muddy silt-
stone of the outer shelf and slope facies, the sandstone is
more silty and more bioturbated, and most of the fossils are
confined to a few beds with indistinct contacts due to bio-
turbation (Squires, 1981).

One such bed, informally known as the “Stewart bed”
(Squires, 1979, 1981, 1983b), is the most distinctive fossil-
iferous unit in the shallow-marine facies. This 1-m-thick
layer occurs in the uppermost part of the transgressive phase
of the shallow-marine facies in the northern part of the study
area and can be traced laterally for about 10 km eastward
from the type section of the formation. Fossils in the lower
part of the “Stewart bed” show less evidence of postmortem
transport than those elsewhere in the shallow-marine facies.
Many of the taxa are represented by nearly complete growth
series (with only the early juvenile individuals lacking). Ar-
ticulation of valves is also common. There is a nearly 1:1
ratio of opposite valves of the bivalves Crassatella uvasana
and Venericardia (Pacificor) hornii calafia. Some specimens
of these bivalves, especially Crassatella, are articulated but
are not in growth position. Specimens of all taxa show pres-
ervation of delicate morphologic features. There is random
orientation of the faunal remains. Most of the specimens
occur in the lower 50 cm of the bed and seem to be somewhat
concentrated in indistinct pods. Based on the above features,
as well as the lack of sedimentary structures indicative of
high energy, it is interpreted that a residual (winnowed) pa-
leocommunity, as defined by Fagerstrom (1964), occurs at
many exposures of the lower part of the “Stewart bed”
(Squires, 1981).

The uniformity of the lithology throughout the “Stewart
bed” suggests uniform depositional and environmental con-
ditions. The megafauna is also fairly uniform throughout the
bed. The most abundant faunal elements are carnivorous
naticid gastropods ( Eocernina and Pachycrommium), her-
bivorous gastropods (Turritella and Ectinochilus), infaunal
suspension feeder bivalves ( Crassatella and Venericardia ),
and carnivorous solitary ahermatypic scleractinian corals
( Trochocyathusl ). This Eocernina-Turrite/la-Crassatella-
Trochocyathus ? paleocommunity is best developed at and in
the vicinity of CSUN locality 374, with 50 species of mega-
fossils present. The shallow-marine “Stewart bed” was de-
posited near the shelf/slope break based on calcareous nan-
nofossil studies (Filewicz and Hill, 1 983) and on the presence

of outer shelf and slope deposits directly above it. The pres-
ence of abundant large specimens of Trochocyathusl in the
“Stewart bed,” as well as in a few thin beds a few meters
above the “Stewart bed” at the type section, is consistent
with a shelf/slope break environment for this part of the
Llajas. Solitary ahermatypic scleractinian corals tolerate a
wide range of temperatures and live at all depths, attaining
their best development along margins of the continental
shelves (Wells, 1957; Heckel, 1972).

Megafossils which characterize the transgressive phase of
the shallow-marine facies are Pseudophragmina ( Proporo -
cyclina) c/arki, Cylichnina tantilla, Ectinochilus (Macilentos)
macilentus, Eocernina hannibali, Pachycrommium clarki,
Turritella andersoni lawsoni, Turritella buwa/dana, Brachio-
dontes ( Brachidontes ) cowlitzensis, and Dent ahum ( Laevi -
dentalium) calafium. The only taxon that characterizes the
regressive phase of the shallow-marine facies is Macoma
rosa.

Megafossils which commonly occur in both the transgres-
sive and regressive phases of the shallow-marine facies are
Ca/yptrea diegoana, Phalium ( Semicassis ) tuberculiformis,
Turritella uvasana app/inae, Corbula (Caryocorbula) dick-
ersoni, and Glycvmeris (Glycymeris) rosecanyonensis.

Many of the molluscan genera of the shallow-marine facies
of the Llajas are extant and most commonly occur today in
seas less than 80 m depth (Tables 3 and 4). The discocyclinid
foraminifers lived in very shallow water (below tide level to
perhaps 100 m) (Vaughan, 1945). Small calcareous benthic
foraminifers present in the transgressive and regressive phas-
es of the shallow-marine facies of the formation are indicative
of inner to middle neritic conditions (Schymiczek, 1983a,
1983b).

Megafossils are scarce in the outer shelf and slope facies,
but foraminifers indicative of bathyal depths are common
(Squires, 1981; Schymiczek, 1983a, 1983b). The mollusks
that occur in this facies also occur in the shallow-marine
facies. Apparently, many of the outer shelf and slope facies
mollusks are shallow-marine species that were transported
into the deeper water facies. This is especially true for the
mollusks at CSUN locality 54 1 . This locality is from the base
of a turbidite-filled channel in the outer shelf and slope facies.
Some of the fossils occur as fragments.

PALEOCLIMATE

West Coast Eocene megafauna have long been assigned to
tropical or subtropical environments (Arnold, 1909; Dick-
erson, 1917; Smith, 1919; Clark and Vokes, 1936; Ber-
thiaume, 1938; Vokes, 1940; Durham, 1950). Among the
Eocene molluscan genera that Durham (1950) listed as par-
ticularly characteristic of tropical (20°C or warmer), the fol-
lowing also occur in the Llajas Formation: the gastropods
Anci/la, Architectonica, Conus, Cypraea, Ectinochilus [=Ri-
mella ], Eocernina, Eocithara [=Harpa], Ficopsis, Paraser-
aphs [=Terebellum\, Pseudoliva, Terebra, Turritella, Ve/ates,
Xenophora, the bivalves Corbula, Pinna, Pilar, Spondylus,
large Venericardia, Crassatella [=Crassatellites\, and the
cephalopod Aturia.

Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks 7

Table 5. Recent, marine faunal regions (Ross, 1974:4) of extant molluscan genera of the Llajas Formation.

Genus (subgenus)

Boreal

Warm

temperate

Tropical

*References

Scaphopoda

Dentalium

X

X

M

D,K,R

Gastropoda

A ncilla

M

A2,D,R

A ncistrolepis

M

A&D,E,Ha,K

Architectonica

X

M

A2,K,M2

Calyptraea

X

X

M

A1,D,K,M1,M2

Cerithiopsis

X

X

M

A2,K,K&C,M 1

Cirsotrema

X

M

A2,Ha,Ki,K

Conus

X

X

M

K,M 1 ,R

Cymatium (Septa)

X

M

A2,D,E,K,M 1 ,M2

Cypraea

X

M

A1,K,L,M1,R

Fusinus

X

X

M

Al,A2,D,K,Ki,M 1 ,M2

Fusiturricula

M

A&D.R

Galeodea

M

X

D,E,Ha,Mc,N 1

Lyria

X

M

A2,Ha,K,Ki,W&D

Natica ( Naticarius )

X

M

Ma.R

Nerita

M

K,L,M1,M2

Neverita (Neverita)

M

X

M

Olivella

X

X

M

A2,D,K,K&C,R

Phalium (Semicassis)

X

X

M

E,Ha,Ki,L

Polmices (Euspira)

X

X

M

Ma

Pseudoliva

M

C,G,L

Ranella

X

M

D,E

Scaphander

X

M

X

A 1 ,A2,D,K,M2

Sinum

X

M

K,K&C,Ma,R

Terebra

X

M

D,K,M 1

Turrit ella

X

M

K,K&C,Ki,M 1 ,Me

Xenophora

X

M

A 1 ,A2,Ha,K,Ki,M2,R

Bivalvia

Acanthocardia

X

M

D,E,L,N2,T

Acila (Truncacila)

M

M

X

A1,S

Anomia

X

M

X

A2,D,K,L,M1,M2

Brachidontes

X

X

M

D,K,M 1 ,M2,R

Callista (Costacallista)

X

M

A2,L,R

Cardiornya

X

X

M

Ha,K,Ml,M2,R

Corbula (Caryocorbula)

X

M

A2,K

Gari

X

X

M

A2,D,Ha,L,Ml

Glycymeris

X

X

M

A1,K,M1,M2

Macoma

M

M

X

A1,D,M1,R

Nemocardium

X

X

M

A2,D,Ha,H&G,K,Ki,Ml

Nuculana (Saccella)

X

X

M

A2,K

Ostrea

X

X

M

D,H&G,K,M 1 ,M2

Pinna

X

X

M

A2,D,Ha,K,Ml,M2

Pitar (Lamelliconcha)

M

K.O

Solena

M

K

Spondylus

X

M

D,K,M 1 ,M2

Teredo

X

M

X

A2,M2

x = Present, M = Most frequently reported occurrence. * See Table 3 for abbreviations.

8 Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks

Berthiaume (1938) and Durham (1950) also mentioned
that discocyclinid foraminifers are indicative of tropical or
subtropical environments. All the recorded species of Amer-
ican Discocyclinidae occur in tropical, subtropical, and south
temperate latitudes (Vaughan, 1945).

More recent and independent evidence supports a tropical
to subtropical environment interpretation for West Coast
Eocene faunas. Mineralogical analyses of kaolinite and quartz-
rich, buried-soil profiles in northwest Baja California and
southwest California indicate that humid tropical climatic
conditions prevailed during Paleocene to mid-Eocene time
(Peterson and Abbott, 1979).

Based on oxygen isotope compositions of Tertiary planktic
foraminifer tests throughout the world. Savin, Douglas, and
Stehli (1975) concluded that ocean temperatures remained
warm and relatively constant from Paleocene through middle
Eocene time. Kennett (1982) has similarly stated that during
the Paleocene and Eocene, world climates were relatively
warm and equable with low pole-to-equator temperature gra-
dients. He further stated that a late Paleocene warming trend
culminated in a period of peak warming during the early
Eocene to early middle Eocene. This interval of time coin-
cides with the age of the Llajas Formation. Such equable
conditions ceased in earliest Oligocene time with the ap-
pearance of a worldwide cooling trend. This cooling was
associated with the development of circum-Antarctic cir-
culation (Kennett et ah, 1975; Kennett, 1982; Zinsmeister,

1982) .

Although 56 percent of the molluscan genera collected from
the shallow-marine facies of the Llajas Formation are extant
(Tables 3 and 5), the percentage is not high enough to permit
accurate generic comparison studies with modern molluscan
faunas like those done by Addicott (1970) for Oligocene,
Miocene, and Pliocene molluscan faunas of the West Coast.
Generic comparison studies with modem molluscan faunas
are also difficult to do with early Tertiary faunas because of
the likelihood of significant changes in environmental tol-
erance in progressively older faunas (Addicott, 1 970). Never-
theless, a general comparison with modern molluscan faunas
can be made. Most of the extant molluscan genera of the
shallow-marine facies of the Llajas most commonly occur
today in tropical seas (Tables 3 and 5). It should be noted
also that most of these genera can occur in warm-temperate
seas as well.

The taxa Ancilla, Pseudoliva, Nerita, Pitar ( Lamel/icon -
cha), and Solena are particularly significant as they are con-
fined to tropical waters today. In addition, several of the
extinct generic and subgeneric taxa in the Llajas fauna, in-
cluding Ectinochilus, Eocernina, Eocithara [=Harpa\, Fi-
copsis, Gomphopages, Laevityphis, Lyriscapha [= Volutocris-
tata ], and Exputens, have modern analogues that are also
tropical or subtropical in distribution (Clark and Vokes, 1936;
Palmer, 1967; Givens, 1974).

The presence of large Turritella ( i.e., T. meganosensis pro-
tumescens) in the Llajas is indicative of shallow waters that
were warmer than are presently found at this latitude (Saul,

1983) .

A small percentage of the Llajas megafauna does have a

temperate water aspect to it. Acanthocardia and Ranella are
most commonly found in temperate seas today. Ancistrole-
pis ? and Neverita ( Neverita ) are strictly cold-water gastropods
(Tables 3 and 5). Acanthocardia is most abundant in the
“Stewart bed,” and Ranella and Ancistrolepisl were found
only in this bed. Neverita (Neverita) was found at about the
same horizon as the “Stewart bed.” Most likely, their pres-
ence in or near the “Stewart bed,” which was deposited near
the shelf/slope break, was due to “tropical submergence.”
This phenomenon has been observed in modern mollusks
(Ekman, 1953). Many cold-water mollusks that occur in shal-
low water at higher latitudes have been found to inhabit only
deeper water of similar temperature in the tropics.

AGE

The Llajas Formation is late early Eocene through early mid-
dle Eocene in age, based on mollusks, benthic foraminifers,
and calcareous nannofossils. Assignment to European Stan-
dard Ages, various standard plankton zones, and West Coast
provincial benthic foraminifer and molluscan stages in shown
in Figure 3.

Prior to 1936, early workers using mollusks reported the
age of the Llajas Formation as Eocene (Waring, 1914, 1917;
Kew, 1919, 1924) or middle Eocene (Clark, 1921, 1926;
Cushman and McMasters, 1936).

Clark and Vokes (1936) informally proposed five mollus-
can provincial Eocene “Stages”; namely “Meganos,” “Ca-
pay,” “Domengine,” “Transition,” and “Tejon.” They rec-
ognized two faunal zones in their “Capay Stage,” and they
assigned the Llajas Formation to the upper zone of the “Ca-
pay Stage” through the “Transition Stage.” Givens (1974)
showed that their upper faunal zone of the “Capay” should
be considered part of the “Domengine Stage,” and he re-
stricted the use of “Capay Stage” to their lower faunal zone
of the “Capay Stage.” It is in this restricted sense that “Capay
Stage” is used herein.

Saul (1983) regarded the “Meganos Stage” as late Paleo-
cene-early Eocene, the restricted “Capay Stage” of Givens
( 1 974) as early Eocene, the “Domengine” as late early through
early middle Eocene, and the “Transition” as middle Eocene.
These ages are used for this present report (Fig. 3). Weaver
et al. ( 1 944) regarded the “Tejon” as late Eocene, and Givens
and Kennedy (1979) regarded it as late middle Eocene and/
or late Eocene. Such ages are used for this present report (Fig.
3).

Merriam and Turner (1937) reported that in the Llajas
Formation the “Capay Stage” is represented by the “basal
conglomerate” because of the presence of Turritella mega-
nosensis protumescens, a subspecies regarded by them as an
index fossil for the “Capay Stage.” Based on this present
study, this subspecies occurs only where the basal conglom-
erate (coastal alluvial-fan facies) interfingers with the shal-
low-marine facies.

Vokes (1939) assigned the lower part of the Llajas For-
mation to the lower “Capay Stage” based, in part, on the
presence of Galeodea sutterensis. I was unable to find any G.
sutterensis in the Llajas. Vokes did not figure his Galeodea

Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks 9

species from the Llajas Formation, nor did he give any lo-
cality information or catalog numbers of the specimen(s).
Vokes (1939) assigned the middle part of the Llajas For-
mation to the upper “Capay Stage” based on the presence
of Galeodea susanae, which can be documented as a Llajas
species (i.e., CSUN locality 371). The “G. susanae Zone” of
Clark and Vokes (1936) was originally assigned to the upper
part of the “Capay Stage,” but it was shown to be faunally
indistinguishable from the “Domengine Stage” by Givens
(1974) and was reassigned to the “Domengine Stage.”

Vokes (1939) also used the presence of Fusiturricula (Cre-
naturricula) crenatospira in the “Lower Llajas” as an indi-
cator of the “Capay Stage.” He noted the presence of F. (C.)
crenatospira domenginica in the “Upper Llajas” and consid-
ered this subspecies confined to the “Domengine Stage.” In
this present study, F. (C.) crenatospira was found only in the
“Stewart bed” (just above the middle of the formation) where
it is associated with F. (C.) crenatospira domenginica. F. (C.)
crenatospira domenginica. furthermore, occurs in strata be-
low the “Stewart bed.”

Merriam (1941) reported the presence of Turritella an-
dersoni susanae and T. uvasana infera from the “basal con-
glomerate” of the Llajas Formation. In this present study,
T. andersoni susanae is considered equivalent to T. ander-
soni. As with T. meganosensis protumescens, these taxa occur
in the Llajas Formation only where the coastal alluvial-fan
facies interfingers with the shallow-marine facies. T. ander-
soni is a “Capay Stage” index fossil. Although it has not been
previously reported from the Llajas Formation, Weaver et
al. (1944) assigned the lower part of the Llajas Formation to
their “Capay” Turritella andersoni Zone. The rest of the
Llajas Formation was assigned to the “Domengine” and low-
er “Transition” “Stages” by Weaver et al. (1944).

Squires (1981, 1983a, 1983b) and Saul ( 1 983) reported the
lowermost mollusk-bearing beds of the Llajas Formation to
be late early Eocene in age and the rest of the fossil-bearing
beds as early middle Eocene in age.

Based on the above discussions, it is concluded that the
zone of interfingering between the coastal alluvial-fan facies
and the shallow-marine facies is assignable to the Turritella
uvasana infera fauna of the restricted “Capay Stage” of Giv-
ens (1974) which is equivalent to the lower of the two faunal
zones of the “Capay Stage” of Clark and Vokes ( 1 936), Vokes
(1939), and Weaver et al. (1944). Due to the presence of
many “Domengine” molluscan species associated with the
“Capay” species where the coastal alluvial-fan facies inter-
fingers with the shallow-marine facies, it is probable that
most of the “Capay Stage” is not represented in the Llajas
Formation. The part that is present is probably near the
“Capay-Domengine” boundary. Saul (1983) put this bound-
ary near the P8-P9 boundary of the “Standard” planktic
foraminiferal zonation scheme as used by Berggren, Kent,
and Flynn (in press) (Fig. 3). Saul (1983) also noted that most
of the “Capay Stage” is probably missing in the Simi Valley
area. The uppermost “Capay Stage” part of the Llajas For-
mation, therefore, probably corresponds to latest early Eocene
time.

Fossil-bearing rocks of the Llajas Formation, therefore.

cannot be assigned to any molluscan stage older than the
Turritella uvasana infera fauna of the “Capay,” as used in
the restricted sense of Givens (1974). As noted by Saul (1983),
no mollusks restricted to Clark and Vokes’ ( 1936) older “Me-
ganos Stage” have been recovered from the Llajas Formation.
Although T. uvasana infera and T. andersoni susanae have
been reported from the upper 100 m of the Meganos-age
Santa Susana Formation, specimens are variants slightly dif-
ferent from those at the type localities low in the Llajas
(Merriam, 1941; Saul, 1983). Filewicz and Hill (1983) re-
ported a hiatus of approximately 4 million years between the
Llajas Formation and the underlying Santa Susana Forma-
tion. In this present study, this hiatus is considered to be
more on the order of 3 million years (Fig. 3).

The remaining part of the Llajas Formation is associated
with megafossils that have been reported (Vokes, 1939; Mer-
riam, 1941; Verastegui, 1953; Rehder, 1973; Givens, 1974,
1979; Givens and Kennedy, 1979; Saul, 1983) elsewhere on
the West Coast only from the “Domengine Stage.” These
taxa are Conus caleocius, Eocithara mutica californiensis,
Fusinus teglandae, Fusiturricula ( Crenaturricula ) crenato-
spira domenginica. Lyriscapha lajollaensis , Molopophorus
cretaceus, Olequahia domenginica, Proximitral cretacea,
Pseudoliva lineata, Turritella andersoni lawsoni, Turritella
uvasana applinae, Xenophora stocki, Claibornites diegoensis,
Pitar ( Lamelliconcha ) joaquinensis, and Venericardia (Pa-
cificor) hornii calafia.

Additional evidence for a “Domengine” age for the Llajas
Formation is the overlap or joint occurrence of taxa that
have been reported (Givens, 1974; Givens and Kennedy,
1979) elsewhere on the West Coast as having their lowest
stratigraphic occurrence in the “Domengine Stage,” includ-
ing Ficopsis cooperiana, Laevityphis (Laevityphis) antiquus,
Ranellina pilsbryi, Terebra californica, Glycymeris ( Glycy -
meris) rosecanyonensis, and Glyptoactis (Glyptoactis) do-
menginica, with taxa that have been reported (Givens, 1 974;
Givens and Kennedy, 1979) elsewhere as having their highest
stratigraphic occurrence in the “Domengine Stage,” includ-
ing Architectonica (S tel /axis) cognata. Ectinochilus ( Maci -
lentos) macilentus, Pseudoperissolax blakei praeblakei, Ve-
lates perversus, and Nayadina ( Exputens ) llajasensis.

Based on calcareous nannofossils and mollusks in the Lla-
jas Formation, Saul (1983) reported the “Domengine Stage”
to be of late early through early middle Eocene age.

Strata assignable to the middle Eocene “Transition Stage”
were not recognized in the Llajas Formation in this present
study. The unique association of molluscan species that is
utilized in recognizing this “stage” (Givens, 1974; Givens
and Kennedy, 1979) was not found in the Llajas Formation.

Workers using benthic foraminifers have reported the age
of the Llajas Formation as middle Eocene (Cushman and
McMasters, 1 936) or early to middle Eocene (Laiming, 1 940a,
1940b, 1943; Mallory, 1959; Schymiczek, 1983a, 1983b).

Laiming (1940a, 1940b, 1943) assigned the Llajas For-
mation to the B-3 (upper part), B-2, B-l, and B-1A “Zones”
of his tentative benthic foraminiferal zonation scheme. He
correlated these “zones” to the upper “Capay” through “Do-
mengine” molluscan “Stages” of Clark and Vokes (1936).

10 Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks

Mallory (1959) reported that the lower Llajas corresponds
to his Penutian Stage (lower Eocene, “Capay”) and that the
bulk of the formation corresponds to his Ulatisian Stage
(middle Eocene, “Domengine to Transition”). An updated
version of these benthic foraminiferal stages, now known to
be time-transgressive, is given in Poore (1980). Schymiczek
(1983a, 1983b) assigned the Llajas Formation to the Ulati-
sian Stage. Givens and Kennedy (1979) have concluded that
the “Domengine Stage” is correlative with the Ulatisian (and
probably in part Penutian) Stage.

Using the calcareous nannofossil biostratigraphic zones of
Okada and Bukry (1980) and the absolute dates assigned to
these zones by Berggren, Kent, and Flynn (in press), Filewicz
and Hill (1983) reported that most of the shallow-marine
(transgressive) facies of the Llajas Formation is of latest early
Eocene or earliest middle Eocene age ( Discoaster lodoensis-
D. sublodoensis Zones, CPI 1-CP12). The uppermost 30 m
or so of this facies (Discoasteroides kuepperi Subzone, CP 1 2a)
and the outer shelf and slope facies are of middle Eocene age
(Rhabosphaera inflata Subzone, CP12b-CP13, to Nannotet-
rina quadrata Zone, CPI 3). Rhabdosphaera inflata first oc-
curs 4 m above the “Stewart bed.” These results are sum-
marized in Figure 3.

According to Filewicz and Hill (1983), the R. inflata Sub-
zone plots within the TE2. 1 global, eustatic sea-level cycle
of Vail, Mitchum, and Thompson (1977), and the sudden
shift to upper bathyal waters just above the “Stewart bed”
may be in response to both a steady increase in sea level and
tectonism.

CORRELATION

Correlations between California and Oregon/Washington can
be easily accomplished with molluscan assemblages of mid-
dle and early late Eocene age because these mollusks occupied
a single faunal province (Armentrout, 1975).

Twenty-two (or 42 percent) of the species listed by Givens
and Kennedy (1979) for the “Domengine Stage” Mount So-
ledad, Ardath Shale, and Scripps Formations of the La Jolla
Group near San Diego, California also occur in the Llajas
Formation. All three formations yield Turritella andersoni
lawsoni. In addition, the Ardath Shale yields T. uvasana
applinae and has planktic foraminifers correlative with P10/
1 1 Zones and calcareous nannofossils correlative with the
CPI 2 Zone (Givens and Kennedy, 1979).

Most of the megafauna of the late early through middle
age Eocene Maniobra Formation, Orocopia Mountains,
southeastern California is conspecific with that of the Llajas
Formation. The lower part of the formation has calcareous
nannofossils indicative of CP9 through CPI 1 . Turritella an-
dersoni lawsoni is present in the upper part of the formation
(Crowell and Susuki, 1959; Advocate, 1982).

As mentioned earlier, the zone of interfingering between
the coastal alluvial-fan facies and the shallow-marine facies
of the Llajas Formation can be correlated to the Turritella
uvasana infera fauna described by Givens ( 1 974). This fauna
is from the lowermost part of the Juncal Formation, Pine
Mountain area, California. Fifteen (or 40 percent) of the

species restricted to the Turritella uvasana applinae fauna
described by Givens (1974) from the upper half of the Juncal
Formation in the same area occur also in the Llajas For-
mation above the zone of interfingering between the coastal
alluvial-fan facies and the shallow-marine facies. This Juncal
Formation fauna corresponds to the upper of the two faunal
zones of the “Capay Stage” of Clark and Vokes ( 1 936) through
the “Domengine Stage” (Givens, 1974, fig. 6).

Twenty-seven (or 69 percent) of the megafossil species
listed by Squires (1977) from the “Domengine-Transition”
“Stages” unnamed stratigraphic unit E-,, lower Piru Creek
area, Transverse Ranges, California, also occur in the Llajas
Formation.

In central California, the megafaunas of both the Avenal
Sandstone (Stewart, 1946) and the Domengine Formation
(Vokes, 1939) are similar to that of the Llajas.

In southwestern Oregon, the megafaunas in the Umpqua
and Tyee Formations, as used by Turner (1938), are similar
to that in the Llajas Formation. Turner (1938:32) assigned
most of the Umpqua Formation megafauna to the “Capay
Stage.” He locally subdivided the Umpqua into lower and
upper portions, and he considered the upper portion of his
Glide section as possibly intermediate between the “Capay”
and “Domengine” “Stages.” He assigned the Tyee mega-
fauna to the “Domengine Stage.”

Turner’s “lower Umpqua” now corresponds to the Rose-
burg Formation, and his “upper Umpqua” includes both the
Lookingglassand Flournoy Formations (Baldwin, 1974). The
base of the Lookingglass Formation is considered by Baldwin
to coincide with the division between Turner’s “lower” and
“upper Umpqua.” Based on a study of planktic foraminifers.
Miles (1981) assigned the Roseburg Formation to Zone P7-
8 of the “Standard” zonation, the Lookingglass Formation
to Zone P7-8, and the Flournoy Formation to Zone P10. He
assigned the Tyee Formation to an early middle Eocene age.

Miles (1981), in his study of the southwestern Oregon
Eocene strata, placed the “Capay-Domengine” boundary near
the P9-10 boundary, but Saul (1983) has pointed out that
this molluscan stage boundary actually should lie near the
P8-9 boundary. Using Miles’ (1981) figures 3 and 7 and
Saul’s ( 1 983) refinement of the “Capay-Domengine” bound-
ary, the Roseburg and Lookingglass Formations can be shown
to be equivalent to the “Capay Stage” and the Flournoy
Formation to be equivalent to the “Domengine Stage.”

Miles (1981, fig. 7) assigned most of the Llajas Formation
to early Eocene time, but such an age assignment does not
conform to the findings of this present report or to the findings
of Filewicz and Hill (1983), Schymiczek (1983a, 1983b), or
Saul (1983).

BIOGEOGRAPHY

The widespread aspect of West Coast warm-water Eocene
faunas can be demonstrated by the recognition of some of
Clark and Vokes’ (1936) Eocene “stages” from southern Cal-
ifornia (latitude 33°N) to as far north as the Gulf of Alaska
(latitude 60°N) (Addicott, 1970). About 30 percent of the
Llajas molluscan species, for example, ranged from San Die-

Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks 11

go into Oregon/Washington. Two other species, Acila (Trun-
cacila) decisa and Nuculana ( Saccella ) gabbii, ranged from
southern California to Kamchatka.

Clark and Vokes (1936) used the presence of Velates per-
versus, which occurs in the lower part of the Llajas, as evi-
dence of a seaway connection in the tropical Central Amer-
ican region between the Atlantic Ocean and the waters of the
West Coast of North America. Velates is predominantly
Tethyan in its distribution and has been found in Western
Europe, Africa, Asia, Burma, West Pacific, Jamaica, Florida,
Panama, and southern and central California (Vokes, 1935;
Clark and Vokes, 1936; Palmer, 1967). It was a shallow-
water mollusk indicative of tropical and subtropical climate,
as are species of the subgenus Pacificor of the bivalve Ve-
nericardia (Palmer, 1967) which also occur in the Llajas.

The discocyclinid foraminifer Pseudophragmina (Propo-
rocyclina ) clarki has been found in Florida, Mexico, and Peru,
as well as along the West Coast (Cole and Applin, 1964;
Blondeau and Brabb, 1983). The presence of this species in
the Llajas Formation is additional evidence of the seaway
connection mentioned above.

Clark and Vokes (1936) also cited the presence of Clavi-
lithes n. sp. A and Cryptochorda ca/ifornica from the Llajas
as further evidence of a Central American seaway connection
because these species are closely related, respectively, to
Clavilithes scalaris and Cryptochorda stromboides from the
Paris Basin Eocene. Other Llajas megafauna species used as
evidence for the seaway are Clavilithes tabulatus, Lyria an-
dersoni, and Eocithara mutica californiensis, which are closely
related, respectively, to Clavilithes parisiensis, Lyria maga,
and Eocithara mutica of the Paris Basin Eocene (Clark and
Vokes, 1936; Vokes, 1937). Some other Llajas species which
have been noted as similar to ones from the Paris Basin
Eocene are Callocardia (Nitidavenus) tejonensis [=Isocardia]
which resembles Cytherea nitida from France (Vokes, 1939)
and Benoistia umpquaensis which closely resembles B. brevi-
cula from France (Givens and Kennedy, 1976).

Gardner and Bowles (1934) commented upon middle
Eocene gastropod species from the Chiapas locality, Isthmus
of Tehuantepec, southern Mexico (Atlantic side), that are
closely related to ones from the Llajas Formation. These
species from Chiapas are Cernina ( Eocernina ) chiapasensis,
Amaurellina malinchae, Amaurellina cortezi, and Voluto-
cristata chiapasensis, which are closely related, respectively,
to Eocernina hannibali, Tejonia moragai, Pachycrommium
clarki, and Lyriscapha lajollaensis from the Llajas. Gardner
and Bowles (1934) inferred that the two areas were once
connected by a seaway.

Givens (1979), in a detailed study of Lyrischapa, conclud-
ed that this genus, which is present in the shallow-marine
(transgressive) facies of the Llajas, probably evolved in the
Carribbean region during Paleocene or early Eocene time. It
subsequently spread northward during the middle Eocene or
it immigrated from the Old World Tethyan biogeographic
province. Entrance into the Pacific Coast region of North
America was by way of a narrow seaway connection across
southern Central America (Costa Rica, Panama) or north-
western South America (Givens, 1979).

According to Zinsmeister (1983a), the first indication of
the influx of large numbers of mollusks from the Gulf Coast
and the Caribbean into western North America was in the
late Paleocene, and this influx of immigrants continued in
the early Eocene. “It is not known whether these immigrants
came in distinct pulses associated with major tectonic events
in Central America or represent a continuous long-term west-
ward migration of Caribbean taxa into the Pacific as a result
of a general climatic warming during the Eocene” (Zins-
meister, 1983a:66).

Most of the megafauna genera of the Llajas Formation
were already present on the West Coast by Llajas time. In
addition, several species of the Llajas megafauna were carry-
overs from earlier times. The Llajas taxa Calyptraea die-
goana, Pseudoperissolax blakei praeblakei, Olivella mathew-
sonii. Scaphander ( Mirascapha ) costatus, Surculites mathew-
sonii, Acila (Truncacila) decisa, Nemocardium linteum, and
Nuculana ( Saccella ) gabbii have been reported from late Pa-
leocene “Martinez Stage” strata on the West Coast (Nelson,
1925; Weaver, 1953; Smith, 1 975; Zinsmeister, 1974, 1983a).
The Llajas taxa Turbinolia dickersoni, Polinices ( Euspira )
nuciformis, and Schizaster diabloensis have been reported
from late Paleocene strata on the West Coast (Clark and
Woodford, 1927; Quayle, 1932; Marincovich, 1977). The
Llajas taxa Neverita ( Neverita ) globosa, Turritella uvasana
infera, and Brachidontes ( Brachidontes ) cowlitzensis have been
reported from late Paleocene-early Eocene “Meganos Stage”
strata on the West Coast (Givens, 1974; Saul, 1983).

Several genera immigrated into the West Coast region dur-
ing Llajas time. Equatorial paleocirculation of surface waters
was still largely unrestricted worldwide during Llajas time
and until at least 45 million years ago (middle Eocene), even
though the Tethyan Seaway was closing up due to compres-
sive tectonic forces (Kennett, 1982). As mentioned earlier,
migration from points east of the West Coast was accom-
plished by means of a narrow seaway across southern Central
America or northwestern South America (Woodring, 1966;
Givens, 1977, 1979; Zinsmeister, 1983a). Molluscan genera/
subgenera present in the Llajas Formation and coeval strata
but not in older Tertiary strata of the West Coast will be
discussed below. Unless otherwise noted, biogeographic data
for these genera were obtained from Wenz (1938-1944),
Palmer and Brann (1965-1966), Cox et al. (1969), Palmer
(1974), Marincovich (1977), and Moore (1983), as well as
from the “Systematics” portion of this present report.

Architectonica ( Stellaxis ), Paraseraphs, and Strepsidura
possibly arrived during the early Eocene (uppermost “Ca-
pay” part), most likely from the Caribbean province. Do-
menginella apparently originated on the West Coast at this
time, possibly as a northern variant of Scobinella, as sug-
gested by Vokes (1939).

Benoistia and Megistostoma arrived during the late early-
early middle Eocene (“Domengine”) on the West Coast from
the Tethyan province. Eocithara (see Rehder, 1973), Natica
( Naticarius ), Ranella (see Squires, 1983a), and possibly Co-
nus, Phalium ( Semicassis ), and Marcia (Mercimonia) ar-
rived during this time, most likely from the Caribbean region.
Lyriscapha (see Givens, 1979) may have arrived from either

12 Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks

the Caribbean or Tethyan regions. Cymatium (Septa) (see
Squires, 1983a), Galeodea ( Ca/iagaleodea ), Olequahia, Te-
jonia , Glyptoactis (Glyptoactis), and possibly Proxi mitral
originated on the West Coast at this time. Ancistrolepisl prob-
ably arrived during the early middle Eocene from the Pacific
Northwest.

In summary, the influx of mollusks into western North
America that Zinsmeister ( 1 983a) commented on continued,
at least, into the early middle Eocene. As noted by Smith
(1975), the megafauna of the Llajas Formation is of mixed
origins, having cosmopolitan, Tethyan, Caribbean, and North
American elements.

SYSTEMATIC MATERIALS AND METHODS

From 1978 to 1983, an intensive and meticulous search of
the Llajas Formation was undertaken by the author in order
to amass megafossil specimens that would represent the com-
position and the stratigraphic distribution of the fauna more
accurately than the cursory collections on which previous
workers based their reports. About 5000 specimens were
obtained from 132 localities, permitting the addition to the
known fauna of many rare species that otherwise would not
have been found. Numerous return visits were made to sev-
eral of the richest localities. Several hundred man-hours were
spent quarrying CSUN locality 374 (“Stewart bed”) and
CSUN locality 371. Because specimens with preservation
superior to those previously figured were obtained and be-
cause many of the holotypes of species erected for Llajas
Formation specimens are from localities that are either un-
known or only vaguely known, most of the holotypes are not
refigured.

Systematic arrangement of the generic and higher taxo-
nomic categories follows that of Cole (1964) for the large
foraminifer; Wells (1956) for the scleractinians; Muir-Wood,
Elliott, and Hatari (1965) for the brachiopod; Howell (1962)
for the annelid tube; Ludbrook (1960) for the scaphopod;
Cox et al. (1969) and Vokes (1980) for the bivalves; Miller
(1947) for the nautiloid; Jeletzky (1966) for the sepiid;
Glaessner (1969) for the brachyuran; Fischer (1966) for the
echinoid; and Compagno (1973) for the sharks. The system-
atic arrangement of Wenz (1938-1944) is followed generally
for the gastropods.

Most of the figured specimens are on deposit in the Natural
History Museum of Los Angeles County, Invertebrate Pa-
leontology section. Figured specimens borrowed from the
paleontology collections of the University of California, Los
Angeles, Department of Earth and Space Sciences are on
deposit there. Additional unfigured specimens are on deposit
in the Department of Geological Sciences Paleontology col-
lection, California State University, Northridge.

The synonymies are selective. Works that include original
figures and/or descriptions are listed. References that add
pertinent and documentable biostratigraphic information are
also included. More complete synonymies of many of the
gastropods and bivalves can be found in Stewart (1927, 1 930).

Primary type material, molluscan stage range, geographic
distribution, local occurrence, and remarks are listed for all

the species. Unless otherwise noted, such data are derived
from references listed in the synonymies or from new data
obtained in the course of this present study. “Primary type
material” refers to the holotype, paratype(s), syntypes, lec-
totype, paralectotype(s), or neotype of the senior subjective
synonym of each taxon. In the case of homonyms, the junior
homonym “primary type material” is listed, and if the new
name “primary type material” is different it is listed also.
The molluscan stages are for the West Coast, and they are
from Clark and Vokes ( 1 936) and Weaver et al. ( 1 944), with
refinements made by Givens (1974) and Saul (1983). The
stages are provisional, hence the names are placed in quo-
tation marks. The relative age of each stage and correlation
with various biostratigraphic zones are shown in Figure 4.
Any taxa stage range extensions that are the result of this
present study of the Llajas material are so mentioned under
the “Remarks” for each species. Locality information for all
the localities mentioned in this report is given in the “Lo-
calities” section.

Letter abbreviations used for catalog and/or locality num-
bers are:

ANSP = Academy of Natural Sciences of Philadelphia
CAS = California Academy of Sciences
CIT = California Institute of Technology
CSUN = California State University, Northridge
MCZ = Museum of Comparative Zoology, Harvard
HS = Herman Schymiczek (CSUN master’s thesis field sta-
tion)

LACMIP = Los Angels County Museum, Invertebrate Pa-
leontology Section

SU = Stanford University (collections now housed at the
California Academy of Sciences)

UCMP = University of California Museum of Paleontology
(Berkeley)

UCLA = University of California, Los Angeles
UCR = University of California, Riverside
UO = University of Oregon

USGS = United States Geological Survey (Washington, D.C.
register)

USNM = United States National Museum of Natural His-
tory

UW = University of Washington

SYSTEMATICS
Phylum Protista
Subphylum Sarcodina
Class Rhizopodea
Subclass Granuloreticulosia
Order Foraminiferida
Suborder Rotaliina
Superfamily Orbitoidacea
Family Discocyclinidae Galloway, 1928

Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks 13

Genus Pseudophragmina Douville, 1923

Type Species. By original designation, Orthophragmina
flondana Cushman, 1917.

Pseudophragmina ( Proporocyclina ) clarki
(Cushman, 1920)

Figures 5a-b

Orbitolites sp. A Arnold, 1 9 1 0:pl. 3, fig. 6.

Orthophragmina clarki Cushman, 1920:41-42, pi. 7, figs.
4-5.

Discocyclina clarki (Cushman). Schenck, 1929:221, figs. 1-
2, 5, text fig. 7. Keenan, 1932:pl. 4, figs. 1-2. Vaughan,
1936:255-256, pi. 43, figs. 1-2. Berthiaume, 1938:496, pi.
61, fig. 12. Turner, 1938:7, 12, 21. Stewart, 1946:table 1.
Shepard, Lankford, and Milow, 1957:columnar section.
Discocyclina cloptoni Vaughan. Cushman and McMasters,
1936:516, pi. 77, figs. 6-9. [Misidentification fide Schy-
miczek, 1983b:50.] Vaughan, 1945:98-99, pi. 42, fig. 2.
Pseudophragmina ( Proporocyclina ) clarki (Cushman).
Vaughan, 1945:104, 106. Cole, 1958:419-420, pi. 52, figs.
3-11. Crowell and Susuki, 1959:589. Mallory, 1959:259.
Cole and Applin, 1964:47, pi. 9, figs. 1-4. Blondeau and
Brabb, 1983:47, pi. 6, fig. 17, and table 5.
Pseudophragmina clarki (Cushman). Schymiczek and Squires,
1981:989. Squires, 1983b:fig. 9i.

Primary Type Material. Holotype and paratype, USNM
collections, upper Lodo Formation, northeast side of Do-
mengine Creek near the comer of the SW Vi of section 29,
T 18 S, R 15 E, Domengine Ranch quadrangle, California.
Molluscan Stage Range. “Capay” through “Domengine.”
Geographic Distribution. Florida, Mexico, Peru through
southwestern Oregon.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 374, 457, 4691, 470b, 472, 473, 477, 479,
487, 491, 504, 51 1, 522, 527a, 527b, 548.

Remarks. Specimens are essentially confined to the shal-
low-marine (transgressive) facies. Most occur in several 50-
cm-thick beds approximately 100 m above the base of the
formation (e.g., localities 457, 4691, 487, 511, 527b). In such
beds, the tests are concentrated in small pods. Turrit ella an-
dersoni lawsoni may or may not occur with the tests, but if
present it is usually the only associated megafossil or the
most abundant megafossil. The tests of P. (P.) clarki are
mostly complete, unabraded, and 1.5 to 7 mm in diameter.

The stratigraphic highest occurrence of the specimens in
the Llajas Formation is at locality 473, about 14 m above
the “Stewart bed” at the type section. The bed that locality
473 occurs in probably represents a shallow-marine (trans-
gressive) facies bed that interfingers with the outer shelf and
slope facies.

Phylum Coelenterata
Subphylum Cnidaria
Class Anthozoa

Subclass Zoantharia
Order Scleractinia
Suborder Caryophylliina
Superfamily Caryophylliicae
Family Caryophylliidae Gray, 1847
Subfamily Caryophylliinae Gray, 1847

Genus Trochocyathus
Milne-Edwards and Haime, 1848

Type Species. By subsequent designation (Milne-Edwards
and Haime, 1850b), Turbinolia mitrata Goldfuss, 1827.

ITrochocyathus striatus (Gabb, 1864)

Figure 5c

Trochosmilia striata Gabb, 1864:207-208, pi. 26, fig. 195.
Trochocyathus striatus (Gabb). Vaughan, 1900:101-102, pi.

7, fig. 24; pi. 8, figs. 1-3. Dickerson, 1913:265. Hanna,

1927:269.

Primary Type Material. UCMP holotype 12256, Division
B, near Mt. Diablo, California.

Molluscan Stage Range. “Capay”?; Domengine.

Geographic Distribution. Simi Valley through central Cal-
ifornia.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 374, 438, 444, 445, 447, 451, 458, 459,
471c, 472, 473, 477, 479, 480, 493, 494, 495, 496, 500, 510,
525.

Remarks. ITrochocyathus striatus occurs primarily in the
“Stewart bed.” At a few localities (e.g., locality 374) there is
a partial growth series, with the early juveniles missing. Most
specimens in the Llajas Formation are complete, unattached,
and unabraded.

Specimens at locality 473 occur in a bed that probably
represents a shallow-marine facies bed that interfingers with
the outer shelf and slope facies.

The genus and species of the most abundant coral in the
Llajas Formation are questioned because of a lack of modem
comparative taxonomic work on middle Eocene solitary
scleractinian ahermatypic corals. The type of this taxon is
largely imbedded in matrix and probably cannot be well
enough prepared to make a valid comparison (Durham, 1981,
pers. commun.).

Dickerson (1913) reported T. striatus from “Capay”-age
strata in central California, but he gave no specific locality
information nor did he figure the species. For these reasons,
the “Capay” age occurrence is uncertain.

Subfamily Turbinoliinae
Milne-Edwards and Haime, 1848

Genus Turbinolia Lamarck, 1816

Type Species. By subsequent designation (Milne-Edwards
and Haime, 1850b), Turbinolia sulcata Lamarck, 1816.

14 Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks

Turbinolia dickersoni Nomland, 1916

Figures 5d-e

Turbinolia dickersoni Nomland, 1916:61, pi. 3, figs. 5-8.

Palmer, 1923:306, pi. 56, figs. 2, 4. Quayle, 1932:98-99,

pi. 6, figs. 1-5.

Primary Type Material. UCMP holotype 12020 (missing
since 1932), Cerros Shale Member of the Lodo Formation,
UCMP locality 1817.

Molluscan Stage Range. Upper Paleocene through “Do-
mengine.”

Geographic Distribution. Simi Valley through central Cal-
ifornia.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN locality 467.

Remarks. Only four specimens were found in the Llajas
Formation. This small coral is distinguished from the much
larger ITrochocyathus striatus by the presence of a stellate
columella. This columella does not project noticeably above
the corallum wall.

The type locality of this species is in the lower part of the
Cerros Shale Member (late Paleocene age) or the Lodo For-
mation (Marincovich, 1977:252; Moore, 1983:A88).

Clark (1926:114) reported this species from “Domengine”-
age strata in central California, but such a report is uncertain
because of his lack of locality data. The presence of this
species at CSUN locality 467 does extend, with certainty, its
molluscan stage range into the “Domengine Stage.”

Phylum Brachiopoda
Class Articulata
Order Terebratulida

Suborder, Superfamily, and Family UNKNOWN
Genus Eogryphus Hertlein and Grant, 1 944

Type Species. By original designation, Eogryphus tolmani
Hertlein and Grant, 1944.

Eogryphus tolmani Hertlein and Grant, 1944

Figure 5f

Eogryphus tolmani Hertlein and Grant, 1944:89-90, pi. 5,

figs. 1-3, 7; pi. 18, figs. 1, 9-11; text figure 22.

Primary Type Material. UCLA holotype 6203, CAS para-
types 7279-7282; all from upper Llajas Formation, upper
Las Llajas Canyon, 3000 feet north and 1 500 feet east of the
southwest corner of section 23, T 3 N, R 1 7 W (Hertlein and
Grant, 1944:89).

Molluscan Stage Range. “Domengine.”

Geographic Distribution. Simi Valley, California.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 438, 468a, 515, 525.

Remarks. Many specimens were found at and near the
vicinity of locality 438. Many of the specimens are articu-
lated. Plotted on the topographic map available in 1944, the
locality of the type specimens probably plots about 1 500 feet

(457 m) east of CSUN locality 5 1 5 in an area in which slope
wash has covered the exposures. The southwest corner of
section 23 was unmarked and had to be projected. The lo-
cality of the type specimens, however, is in the general vi-
cinity of all the CSUN localities in which specimens of Eogry-
phus tolmani were found.

Smith (1975:470) reported Eogryphus cf. E. tolmani from
the “Martinez Stage” (Paleocene) part of the Lodo Forma-
tion, central California.

Phylum Annelida
Class Polychaetia
Order Sedentaria

Family Serpulidae Lamarck, 1818
Genus Rotularia Defrance, 1827

Type Species. By original designation, Serpula spirulaea
Lamarck, 1818.

Rotularia tejonense (Arnold, 1910)

Figure 5g

Spiroglyphus ? tejonensis Arnold, 1 9 1 0:5 1 , pi. 4, fig. 18. Dick-
erson, 191 6:pl. 37, figs. 5a-b. Vokes, 1939:162-163, pi.

20, figs. 20-22. Stewart, 1946:pl. 11, fig. 21.

ITubulostium tejonense (Arnold). Keen and Bentson, 1944:

195.

Rotularia tejonense { Arnold). Nilsen, 1973:table 1. Squires,

1977:table 1 .

Primary Type Material. USNM holotype 165658, Avenal
Formation, USGS locality 4617.

Molluscan Stage Range. “Capay” through “Transition.”

Geographic Distribution. Simi Valley through central Cal-
ifornia.

Local Occurrence. Interfingering coastal alluvial-fan facies
and shallow-marine (transgressive) facies: CSUN locality 703.
Shallow-marine (transgressive) facies: CSUN localities 374,
444, 445, 494, 521, 546.

Remarks. Most specimens were found in the “Stewart bed”
encrusting other shells, most notably Ostrea.

According to Clark (1921:table 1), R. tejonense [ =Spiro -
glyphus (?) tejonensis] occurs in “Meganos” age strata. Such
a report, however, cannot be substantiated at this time be-
cause of his lack of locality information and because his rock
unit names are not clearly defined.

Phylum Mollusca
Class Scaphopoda
Family Dentaliidae Gray, 1834
Genus Dentalium Linne, 1758

Type Species. By subsequent designation (Montfort, 1810),
Dentalium elephantinum Linne, 1758.

Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks 15

Dentalium stentor Anderson and Hanna, 1925

Figure 5h

Dentalium stentor Anderson and Hanna, 1925:145, pi. 13,

fig. 17.

Dentalium stentorl Anderson and Hanna. Squires, 1977 :ta-

ble 1.

Primary Type Material. CAS holotype 819, Tejon For-
mation, CAS locality 792.

Molluscan Stage Range. “Domengine” through “Tejon.”

Geographic Distribution. Simi Valley through southern San
Joaquin Valley, California.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 371, 373, 374, 444, 445, 447, 449, 451, 493,
494, 500, 510.

Remarks. Abundant large, thick-shelled specimens were
found locally in the “Stewart bed” (locality 374 and vicinity).
Although some specimens are nearly complete (up to 75 mm
in height), no apices were found. Due to the absence of apices,
the specimens could not be assigned with certainty to any
subgenus of Dentalium.

Subgenus Laevidenta/ium Cossman, 1888

Type Species. By original designation, Dentalium incertum
Deshayes, 1825.

Dentalium ( Laevidenta/ium ) calafium
Vokes, 1939

Figure 5i

Dentalium (Laevidentalium) calafium Vokes, 1939:105, pi.

16, figs. 30-31. Squires, 1983b:fig. 9h.

Primary Type Material. UCMP lectotype 15750, Domen-
gine Formation, UCMP locality A- 1027; UCMP paralecto-
type 15751, Domengine Formation?, UCMP locality A-1003.

Molluscan Stage Range. "Domengine.”

Geographic Distribution. Simi Valley through central Cal-
ifornia.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 37 1,373, 455, 466, 467, 484, 49 1 , 499, 500,
507, 508, 510, 529b, 538, 539, 548.

Remarks. Specimens occur in channel-fill deposits in the
Llajas Formation. They are small, smooth, and fragmented
specimens. The presence of the apical notch on the convex
side of the shell serves to distinguish this taxon.

Vokes (1939) designated two syntypes (UCMP 15750 and
15751) but no holotype for his species. In accordance with
the International Code of Zoological Nomenclature (1964)
article 74, therefore, UCMP 15750 is herein designated as
the holotype of D. (L.) calafium.

Class Gastropoda
Subclass Prosobranchia
Order Archaeogastropoda
Superfamily Neritacea

Family Neritidae Rafinesque, 1815
Subfamily Neritinae Rafinesque, 1815
Genus Nerita Linne, 1758

Type Species. By subsequent designation (Montfort, 1810),
Nerita peloronta Linne, 1758.

Nerita cf. N. ( Amphinerita ) ear ex
Vokes, 1939

Figure 6 a

Local Occurrence. Interfingering coastal alluvial-fan facies
and shallow-marine (transgressive) facies: CSUN localities
700, 701.

Remarks. Specimens were found only in the “Capay Stage”
portion of the Llajas Formation. A total of 22 specimens was
found, and all but two specimens were from locality 700.
Due to the presence of well-cemented matrix in the aperture
area of each specimen, the labial areas could not be studied.
The small specimens have a very low spire, large body whorl,
and a smooth shell surface. Apparently, there is no umbilicus,
and the outer lip is not dentate.

The specimens most closely resemble N. (A.) eorex Vokes
(1939:180-181, pi. 22, figs. 24, 26, 29) from the Domengine
Formation, Reef Ridge area, California. Vokes (1939) re-
garded this species as the only smooth-shelled Nerita from
the West American Eocene.

Genus Velates de Montfort, 1810

Type Species. By original designation, Velates conoideus
Montfort, 1810 [=Nerita perversa Gmelin, 1791].

Velates perversus (Gmelin, 1791)

Figures 6b-c

Nerita perversa Gmelin, 1791:vol. 1, pt. 6, p. 3686.

Velates conoideus Montfort, 1810:355.

Velates perversus (Gmelin). Vokes, 1935:382-383, pi. 25,

figs. 1-5; pi. 26, figs. 1-2. Clark and Vokes, 1936:875, pi.

1, figs. 7-8. Givens, 1974:61, pi. 5, figs. 5-6, 13. Givens

and Kennedy, 1979:83.

Primary Type Material. No holotype was designated by
Gmelin (1791). Gmelin did not have a collection but worked
mainly from the literature, according to Smith (1970:459).
Gmelin (1791) lists the Chemnitz Cabinet as the source of
his Nerita perversa, and if the Cabinet is still extant, the
lectotype could be designated.

Molluscan Stage Range. “Capay” through “Domengine.”

Geographic Distribution. Western Europe, Africa, Asia,
Burma, West Pacific, Jamaica, southern and central Califor-
nia.

Local Occurrence. Interfingering coastal alluvial-fan facies
and shallow-marine (transgressive) facies: CSUN localities
700, 701, 702.

Remarks. Specimens were found only in the “Capay Stage”
portion of the Llajas Formation. Except at locality 701, only
fragments were found. A very large specimen (diameter 70

16 Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks

Figure 5. Llajas Formation large benthic forammifer, solitary corals, brachiopod, calcareous worm tube, and scaphopods. a and b. Pseudo-
phragmina ( Proporocyclina ) clarki (Cushman, 1920). a, test exterior, LACMIP hypotype 6500, CSUN loc. 511, x 8. b, equatorial section,
LACMIP hypotype 6501, HS loc. HS-A-1 1(4), x 25. c. ITrochocyathus striatus (Gabb, 1864), lateral view, LACMIP hypotype 6502, CSUN
loc. 374, x 1. d and e. Turbinolia dickersoni Nomland, 1916, LACMIP hypotype 6503, CSUN loc. 467. d, dorsal view, x 7. e, lateral view,
x 5. f. Eogryphus tolmani Hertlein and Grant, 1944, dorsal view, LACMIP hypotype 6504, CSUN loc. 438, x 0.75. g. Rotularia tejonense
(Arnold, 1910), LACMIP hypotype 6505, CSUN loc. 444, x 1.5. h. Dentalium stentor Anderson and Hanna, 1925, partial specimen, side
view, LACMIP hypotype 6506, CSUN loc. 444, x 1 . i. Dentalium ( Laevidentalium) calafium Vokes, 1939, partial specimen, apical notch
view, LACMIP hypotype 6507, CSUN loc. 373, x 4.5.

cm) was found at locality 701. The best specimens from the
Llajas are in the UCMP collection and are from a locality
equivalent to CSUN locality 702 (i.e., UCMP locality 7193).
UCMP specimen 37435 (Figs. 6b-c) is from this locality.
Vokes (1935) figured two hypotypes from this same locality.

The most characteristic feature of this species is the pres-
ence of seven or eight small teeth on the inner lip.

Order Mesogastropoda

Superfamily Cerithiacea

Family Turritellidae Woodward, 1851

Genus Turritella Lamarck, 1799

Type Species. By monotypy, Turbo terebra Linne, 1758.

Turritella meganosensis protumescens

Merriam and Turner, 1937

Figure 6d

Turritella meganosensis n. subsp. Clark, 1929:pl. 10, figs.
1-2.

Turritella meganosensis protumescens Merriam and Turner,

1937:104, pi. 6, figs. 8-10. Turner, 1938:85, pi. 22, fig. 15.

Merriam, 1941:75-76, pi. 8, figs. 1-2, 5, 6, 8. Weaver,

1943:369-370, pi. 74, figs. 14, 18. Saul, 1983:pl. 2, fig. 1.

Primary Type Material. UCMP holotype 15353, “basal
conglomerate” of the Llajas Formation, UCMP locality 7195.

Molitiscam Stage Range. “Capay.”

Geographic Distribution. Simi Valley, California through
southwestern Oregon.

Local Occurrence. Interfingering coastal alluvial-fan facies
and shallow-marine (transgressive) facies: CSUN localities
452, 454, 469a, 469b, 469c, 501, 542, 545.

Remarks. According to Merriam and Turner (1937:104),
this taxon occurs in the basal conglomerate of the Llajas
Formation. This is misleading, as specimens do not occur in
the basal conglomerate proper (equivalent to the coastal al-
luvial-fan facies). They actually are confined to beds that
represent shallow-marine facies interfingering with the coast-
al alluvial-fan facies. At a few of the localities, the thick-
shelled, robust T. meganosensis protumescens exhibits growth
series.

Contributions in Science, Number 350

Squires: Simi Vaiiey Eocene Mollusks 17

Turritella andersoni Dickerson, 1916

Figure 6e

Turritella andersoni Dickerson, 1916:501-502, pi. 42, figs.
9a-b. Turner, 1938:83, pi. 22, figs. 4-6. Merriam, 1941:
76-77, pi. 9, figs. 1-2; pi. 10, figs. 1, 3-5, 8; pi. 12, figs.
1-3. Givens, 1974:62, pi. 5, figs. 7-10.

Turritella andersoni susanae Merriam, 1941:79, pi. 11, fig.
6. Saul, 1983:pl. 2, fig. 5.

Primary Type Material. UCMP holotype 12131, Domen-
gine Formation, UCMP locality 1817.

Molluscan Stage Range. Upper “Meganos”?, “Capay.”
Geographic Distribution. Simi Valley, California through
southwestern Oregon.

Local Occurrence. Upper part of Santa Susana Formation?;
interfingering coastal alluvial-fan facies and shallow-marine
(transgressive) facies of the Llajas Formation: CSUN local-
ities 469d, 700.

Remarks. Only a single specimen was found at locality
469d. This specimen (Fig. 6e) consists of only 2Vi whorls,
but the characteristic noded three primary spiral ribs, the
median one situated closer to the anterior than the posterior
primary, are present.

Although the holotype (UCMP 15295, UCMP locality
A-993) of T. a. susanae is from strata equivalent to the
interfingering coastal alluvial-fan facies and shallow-marine
(transgressive) facies, specimens have been reported by Mer-
riam (1941) from the upper part of the underlying Santa
Susana Formation. Saul (1983) identified these specimens
with T. andersoni n. sp. because none has the sculpture of
the Llajas specimens. Saul (1983), however, did note that
some of the Santa Susana specimens are very close to T.
andersoni Dickerson, especially from localities near the Lla-
jas-Santa Susana contact.

Turritella andersoni lawsoni Dickerson, 1916

Figures 6f-g

Turritella lawsoni Dickerson, 1916:502, pi. 42, figs. lOa-b.
Hanna, 1927:308, pi. 49, fig. 5. Hanna and Hertlein, 1943:
fig. 62-15. Stewart, 1946:pl. 11, figs. 27-29.

Turritella andersoni Dickerson, Waring, 1917 :pl. 15, fig. 18.
Turritella andersoni lawsoni Dickerson. Vokes, 1939:161.
Merriam, 1941:77-78, pi. 9, figs. 3-8; pi. 12, fig. 4. Crowell
and Susuki, 1959:pl. 2, fig. 12. Givens, 1974:62-63, pi. 5,
figs. 11-12, 14. Givens and Kennedy, 1979:82-83, table
1. Saul, 1983:pl. 2, figs. 10-1 1. Squires, 1983b:fig. 9e.
Turritella andersoni lawsoni secondaria Merriam, 1941:78-
79, pi. 9, fig. 9. Crowell and Susuki, 1959:pl. 2, fig. 11.
Turritella andersoni lawsoni forma secondaria Merriam. Saul,
1983:pl. 2, fig. 12.

Primary Type Material. UCMP holotype 12128, Domen-
gine Formation, UCMP locality 2295.

Molluscan Stage Range. “Domengine.”

Geographic Distribution. San Diego through central Cal-
ifornia.

Local Occurrence. Shallow-marine (transgressive) facies:

CSUN localities 374, 439, 440, 441, 442, 443, 444, 445, 447,
449, 453, 455, 457, 468b, 469j, 470c, 471a, 471b, 471e, 481,
485, 487, 490, 493, 494, 510, 516, 522, 523, 527a, 529a,
529b, 537, 539, 548, 704.

Remarks. Turritella andersoni lawsoni is one of the most
abundant megafaunal components in the Llajas Formation.
At many localities (e.g., 442 and 471a), it is the only mega-
fossil present. T. andersoni lawsoni is restricted in the Llajas
Formation to the shallow-marine (transgressive) facies. Its
highest stratigraphic occurrence is the “Stewart bed,” where
it is abundant. Some specimens show faint spiral ribs, where-
as others do not. Some specimens, especially at locality 374,
have the spiral ribs on one side but not on the other (weath-
ered) side.

Some specimens of T. a. lawsoni from localities 374 and
548 are characterized by the presence of a well-developed
anterior primary rib and represent the form described by
Merriam (1941:78-79, pi. 9, fig. 9) as variety secondaria.
These Llajas specimens of T. a. lawsoni which show the well-
developed anterior primary rib are probably just ecotypes.

The primary type material of secondaria is UCMP 33998
from UCMP locality 7004 = CSUN locality 374.

Turritella buwaldana Dickerson, 1916

Figure 6h

Turritella buwaldana Dickerson, 1916:500-501, pi. 42, figs.

7a-b. Hanna, 1927:307, pi. 49, figs. 7-8, 12. Merriam,

1941:86-87, pi. 21, figs. 3-9; pi. 22, figs. 1-14. Stewart,

1946:pl. 11, fig. 24. Givens, 1974:63, pi. 5, fig. 15. Saul,

1983:pl. 2, figs. 13-15. Squires, 1983b:fig. 9f.

Turritella kewi Dickerson, 1916:501, pi. 42, fig. 8.

Primary Type Material. UCMP holotype 12130, Domen-
gine Formation, UCMP locality 672.

Molluscan Stage Range. Upper “Meganos”?, “Capay”?,
“Domengine” through “Tejon.”

Geographic Distribution. San Diego, California through
southwestern Oregon.

Local Occurrence. Upper part of the Santa Susana For-
mation?; shallow-marine (transgressive) facies of the Llajas
Formation: CSUN localities 371, 372, 373, 455, 467, 469e,
469h, 469j, 484, 486, 499, 500, 505, 514, 538, 539, 540,
548, 705.

Remarks. T. buwaldana in the Llajas Formation is char-
acterized by its small size and presence of three primary spiral
ribs and two posterior secondaries.

T. buwaldana has been reported from the upper part of
the Santa Susana Formation by Merriam (1941), but he noted
that these forms are probably subspecifically distinct from
those in the Llajas Formation.

Saul (1983) reported T. b. crooki Merriam and Turner
from the “Capay” portion of the Llajas Formation. Although
none was found in the present study, it is possible that T. b.
crooki and T. buwaldana are the same. More taxonomic data
are needed to resolve the question (Saul, 1983, pers. com-
mun.).

18 Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks

Turritella uvasana infera Merriam, 1941

Figure 6i

Turritella uvasana infera Merriam, 1941:90, pi. 40, figs. 2-
4. Givens, 1974:65-66, pi. 6, figs. 5-7. Saul, 1983:pl. 2,
fig. 4.

Primary Type Material. UCMP holotype 33993, “basal
conglomerate” of the Llajas Formation, UCMP locality
A-994 = CSUN locality 452; UCMP paratypes 1 5439 and
1 5443, upper part of the Santa Susana Formation, UCMP
locality 7000.

Molluscan Stage Range. Upper “Meganos” through “Ca-
pay.”

Geographic Distribution. Simi Valley and Pine Mountain
area, southern California.

Local Occurrence. Upper part of the Santa Susana For-
mation; interfingering coastal alluvial-fan facies and shallow-
marine (transgressive) facies of the Llajas Formation: CSUN
localities 452, 705.

Remarks. According to Merriam (1941:90), this taxon oc-
curs in the basal conglomerate of the Llajas Formation. This
is misleading, as specimens do not occur in the basal con-
glomerate proper (equivalent to the coastal alluvial-fan fa-
cies). They actually are confined to beds that represent shal-
low-marine facies interfingering with the coastal alluvial-fan
facies.

T. u. infera has been reported from the upper part of the
Santa Susana Formation, but the specimens are slightly dif-
ferent variants that have more rounded whorl profiles and
heavier ribbing than those from the type locality in the Llajas
Formation (Merriam, 1941; Saul, 1983).

Adult whorls of the Llajas specimens have six to seven
primary spiral ribs with the posteriormost two more closely
spaced and somewhat weaker than the other spiral ribs. Sec-
ondary ribs are generally absent.

The illustrated specimen (Fig. 6i) is the same one used by
Saul (1983:pl. 2, fig. 4).

Turritella uvasana applinae Hanna, 1927

Figure 6j

Turritella applini Hanna, 1927:307, pi. 49, figs. 1, 4. Clark,
1929:pl. 10, figs. 8, 18.

Turritella uvasana applini Hanna. Merriam, 1941:93-94, pi.
16, figs. 5-6; pi. 18, fig. 2.

Turritella uvasana etheringtoni Merriam, 1941:94, pi. 15,
figs. 12-15. Squires, 1977:table 1.

Turritella uvasana applinae Hanna. Givens, 1974:66, pi. 6,
figs. 3-4; pi. 7, fig. 19. Givens and Kennedy, 1979:82-83,
table 1. Saul, 1983:pl. 2, fig. 18. Squires, 1983b:fig. 9g.

Primary Type Material. UCMP holotype 30971, La Jolla
Formation, UCMP locality 3993; UCMP paratype 33894,
La Jolla Formation, UCMP locality 3990.

Molluscan Stage Range. “Domengine.”

Geographic Distribution. San Diego through Pine Moun-
tain area, southern California.

Local Occurrence. Shallow-marine (transgressive) facies:

CSUN localities 371, 374, 440, 444, 449, 469e, 469i, 47 Id,
471e, 481, 484, 486, 491, 500, 507, 529a, 539, 548, 704.
Outer shelf and slope facies: CSUN locality 483. Shallow-
marine (regressive) facies: CSUN localities 475, 489, 512b,
51 2d.

Remarks. Llajas specimens of T. u. infera superficially re-
semble T. u. applinae, but the latter has a more convex whorl
profile, many more primary spiral ribs, and the presence of
numerous well-developed secondary ribs.

The primary type material for T. u. etheringtoni is from
the Llajas Formation: UCMP holotype 33875, UCMP lo-
cality 7003; UCMP paratypes 33876 and 33877, UCMP lo-
cality 7004 = CSUN locality 374. Specimens of T. u. appli-
nae are especially abundant at locality 374.

Family Architectonicidae Gray, 1850
Genus Architectonica Roding, 1798

Type Species. By subsequent designation (Gray, 1847),
Trochus perspectivus Linne, 1758.

Subgenus Architectonica s.s.

Architectonica {Architectonica) llajasensis
Sutherland, 1966

Figure 6k

Architectonica llajasensis Sutherland, 1966:1-4, figs. 1-2.

Primary Type Material. LACMIP holotype 1140, Llajas
Formation, LACMIP locality 461-B.

Molluscan Stage Range. “Domengine.”

Geographic Distribution. Simi Valley, California.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN locality 371.

Remarks. Only three specimens were found, and these are
from the locality in Devil Canyon. Judging from the mega-
fauna associated with Sutherland’s (1966) holotype speci-
men, the type locality of A. llajasensis is probably near CSUN
locality 444 in the “Stewart bed.”

This species is placed in Architectonica s.s. because the
umbilicus is bordered by two serrated ridges (see Sutherland,

1 966), a characteristic of typical Architectonica (Stewart, 1 927:
343).

Subgenus Stellaxis Dali, 1892

Type Species. By original designation, Solarium alveatum
Conrad, 1833.

Architectonica ( Stellaxis ) cognat a

Gabb, 1864

Figure 61

Architectonica cognata Gabb, 1864:1 17, pi. 20, figs. 72, 72a,
72c, not d and e as stated [not 72b = A. alveata (Conrad)
fide Stewart, 1927:344],

Stellaxis cognata (Conrad). Waring, 1917:98.

Architectonica cognata Gabb. Givens and Kennedy, 1979:
83, table 1 .

Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks 19

Architectonica (Stellaxis) cognata Gabb. Stewart, 1927:343-
344, pi. 28, figs. 7-8. Turner, 1938:90, pi. 18, fig. 17.Vokes,
1939:163-164. Stewart, 1946:pl. 11, fig. 4. Weaver, 1943:
363-364, pi. 73, fig. 20; pi. 103, fig. 19. Givens, 1974:68,
pi. 7, figs. 1-3.

Primary Type Material. ANSP lectotype 4223, Tejon For-
mation s.l., 1 1 km south of Martinez, California.

Molluscan Stage Range. Uppermost “Capay” through
“Domengine.”

Geographic Distribution. San Diego, California through
southwestern Oregon.

Local Occurrence. Interfingering coastal alluvial-fan facies
and shallow-marine (transgressive) facies: CSUN localities
452, 702, 703. Shallow-marine (transgressive) facies: CSUN
localities 371, 372, 373, 374, 444, 447, 455, 486, 490, 491,
538, 704.

Remarks. This species is characterized mostly by a single
keel on the periphery of the whorls (Givens, 1974:68). The
Llajas specimens have this prominent keel, as well as a less
prominent sutural cord, thereby producing an apparent dou-
ble-keel look. The sutural cord is exposed due to the removal
of the shell material in the suture region.

A previously assigned “Capay” age (Turner, 1938; Givens,
1974) for the lower range limit of this species was based on
occurrence in the “upper Umpqua” Formation of south-
western Oregon. The “upper Umpqua,” however, is actually
equivalent to both the “Capay” and “Domengine” “Stages”
(Baldwin, 1974; Miles, 1981). A lower range limit of upper-
most “Capay,” nevertheless, can be substantiated by the oc-
currence of this species in the zone of interfingering between
the coastal alluvial-fan facies and the shallow-marine (trans-
gressive) facies of the Llajas Formation.

Family Cerithiidae Fleming, 1828
Subfamily Cerithiinae Fleming, 1828
Genus Benoistia Cossmann, 1899

Type Species. By original designation, Cerithium muri-
coides.

Benoistia umpquaensis Turner, 1938

Figure 6m

Benoistia umpquaensis Turner, 1938:82, pi. 21, figs. 8, 10.

Weaver, 1943:387, pi. 76, figs. 1 1-12.

Tectanus ligniticus Vokes, 1939:164-165, pi. 21, figs. 1,
3, 4.

Benoistea umpquaensis Turner. Givens and Kennedy, 1976:
964-965, pi. 1, figs. 14-21.

Primary Type Material. UCMP holotype 33192, “upper
Umpqua” Formation, UCMP locality A-661.

Molluscan Stage Range. “Domengine.”

Geographic Distribution. San Diego, California through
southwestern Oregon.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN locality 374.

Remarks. Only a single specimen was found, and it is from

the “Stewart bed.” See Givens and Kennedy (1976) for a
detailed morphologic description of this species.

The holotype of Tectarius ligniticus is UCMP 1 5838, Lla-
jas Formation, UCMP locality 7005.

Family Cerithiopsidae H. and A. Adams, 1854
Genus Cerithiopsis Forbes and Hanley, 1853

Type Species. By monotypy, Murex tubercularis Montagu,
1803.

Cerithiopsis llajasensis n. sp.

Figures 6n-o

Diagnosis. A species of Cerithiopsis characterized by te-
leoconch sculpture of four to five primary spiral ribs crossed
by equal-strength collabral costae, with nodes at the inter-
sections. The varices are irregularly spaced.

Cerithiopsis llajasensis n. sp. is similar to C. orovillensis
Dickerson (1916:489-490, pi. 39, fig. 7) from Eocene strata
in northern California. C. llajasensis differs from the holo-
type of C. orovillensis in the following features: larger, a more
elongate-slender shape, equal strength spiral and collabral
sculpture, four to five rather than only four primary spiral
ribs on the teleoconch whorls, equal-strength primary spiral
ribs, less nodose sculpture, more varices, and an absence of
two strong spiral keels above the base of the body whorl.

Cerithiopsis llajasensis n. sp. is similar also to C. excelsus
Dali ( 1 909:75, pi. 3, fig. 9) from Eocene strata in southwestern
Oregon. Givens (1974) regarded C. excelsus as C. excelsum
(Dali). C. llajasensis differs from the holotype of C. excelsum
in the following features: suture more adpressed, absence of
a shoulder on the whorls, equally spaced collabral sculpture,
fewer primary spiral ribs, equal-strength primary spiral ribs,
many more varices, and more swollen varices. The cancellate
sculpture of C. llajasensis is equal in strength over the shell
whereas in C. excelsum this is not the case. C. excelsum has
two prominent spiral ribs and unequally spaced collabral
sculpture, thereby producing an uneven cancellate sculpture.

Description. Shell small, turriform and elongate, with many
convex whorls. Suture moderately impressed, undulating.
Protoconch mammillated, about 3 Vi whorls, smooth. Teleo-
conch sculpture cancellate, nodes at intersections of primary
spiral ribs and collabral costae. Spiral and collabral sculpture
equal strength. Four to five primary spiral ribs per whorl;
seven on body whorl proper, five more on base of body whorl.
Ribs equally spaced, usually separated by a secondary riblet.
No secondary riblets on base of body whorl. Fourteen to 16
collabral costae per whorl. Costae equally spaced, extending
from suture to suture. No collabral ornament on base of body
whorl. Swollen varices irregularly spaced, usually two per
whorl.

Columella smooth, covered by a thin callus. Aperture ovate.
Anterior canal short, shallowly notched, bent backwards. Di-
mensions of largest specimen, height (incomplete) 22.5 mm,
body whorl width 6 mm.

Primary Type Material. LACMIP holotype 6515, LAC-
MIP paratype 6516, Llajas Formation, CSUN locality 371.

20 Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks

Molluscan Stage Range. “Domengine.”

Geographic Distribution. Simi Valley, California.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 371, 373, 489, 491, 498, 517.

Remarks. Specimens of this new species are abundant at
locality 371, but preservation is only fair in most of the
specimens. The type specimens have the best overall pres-
ervation. Fifteen specimens were found at locality 373, one
specimen each from localities 489 and 491, and three spec-
imens each from localities 498 and 5 1 7. Preservation is not
as good at these other localities as it is at locality 37 1 .

The bent-backwards anterior canal of this species is well
developed enough to suggest assignment to Cerithiopsis. Bit-
tium is similar to Cerithiopsis in having a short and shallow
anterior canal, but according to Houbrick (1977) the canal
is not bent backwards in Bittium.

Etymology. The species is named for the Llajas Formation.

Superfamily Epitoniacea
Family Epitoniidae Lamarck, 1822
Genus Cirsotrema Morch, 1852

Type Species. By monotypy, Seal ana varicosum Lamarck,
1822.

Cirsotrema sp.

Figure 6p

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN locality 374.

Remarks. Only a single specimen was found, and it is from
the “Stewart bed.” This genus is characterized by thick axial
varices more or less reflexed posteriorly and usually not con-
tinuous from whorl to whorl; coarse primary spiral ribs; and
basal disk bounded by a prominent enlargement of the var-
ices. A species identification could not be made as parts of
the aperture and basal keel area are missing.

Superfamily Calyptraeacea
Family Calyptraeidae Blainville, 1824
Genus Calyptraea Lamarck, 1799

Type Species. By monotypy. Patella chinensis Linne, 1758.

Calyptraea diegoana (Conrad, 1855)

Figure 6q

Trochita diegoana Conrad, 1855:7, 17; 1857:327, pi. 5, fig.
42.

Galerus excentricus Gabb, 1864:136, pi. 20, fig. 95; pi. 29,
fig. 232a. Dickerson, 1913:264.

Calyptraea calabasaensis Nelson, 1925:419, pi. 54, figs.
8a-b.

Calyptraea (Galerus) calabasaensis Nelson. Clark and Wood-
ford, 1927:120, pi. 21, figs. 10-13.

Calyptraea diegoana (Conrad). Stewart, 1927:340-341, pi.
27, fig. 15. Turner, 1938:89-90, pi. 20, figs. 1-2. Weaver,
1943:351-352, pi. 71, figs. 16, 20; pi. 103, fig. 3; 1953:29.

Stewart, 1946:pl. 11, fig. 5. Kleinpell and Weaver, 1963:
186, pi. 24, fig. 7. Hickman, 1969:79, 82, pi. 11, figs. 7-
8; 1980:33-34, pi. 2, figs. 18-21. Smith, 1975:469, table
1. Givens and Kennedy, 1979:table 2.

Primary Type Material. USNM holotype 1856, Tejon?
horizon, San Diego, California.

Molluscan Stage Range. “Martinez” through lower Oli-
gocene.

Geographic Distribution. San Diego, California through
Washington.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 371, 455, 491, 548. Shallow-marine (re-
gressive) facies: CSUN localities 462, 475, 51 2d.

Remarks. Specimens were most abundant at locality 37 1 .
At all localities, the specimens are unattached.

Family Xenophoridae Philippi, 1853

Genus Xenophora Fischer von Waldheim, 1807

Type Species. By subsequent designation (Gray, 1847),
Trochus conchyliophorus Bom, 1780.

Xenophora stocki Dickerson, 1916

Figure 6r

Xenophora stocki Dickerson, 1916:502-503, pi. 37, figs. 4a-
b. Givens, 1974:71, pi. 7, fig. 8. Givens and Kennedy,
1979:tables 1, 3.

Primary Type Material. UCMP holotype 11838, Rose
Canyon Shale, UCMP locality 2226.

Molluscan Stage Range. “Domengine.”

Geographic Distribution. San Diego through Pine Moun-
tain area, southern California.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 374, 444, 458, 47 Id, 472, 477, 479.

Remarks. Xenophora stocki is most abundant in the “Stew-
art bed,” especially at locality 374. Specimens at all localities
occur as internal molds with only remnants of shell material.

Superfamily Strombacea
Family Strombidae Rafinesque, 1815
Genus Ectinochilus Cossmann, 1889

Type Species. By original designation, Strombus canalis.

Subgenus Macilentos Clark and Palmer, 1923

Type Species. By original designation, Rimella macilenta
White, 1889.

Ectinochilus ( Macilentos ) macilentus
(White, 1889)

Figure 6s

Rimella macilenta White, 1889:19, pi. 3, figs. 10-12.
Ectinochilus (Macilentos) macilentus (White). Clark and
Palmer, 1923:280, pi. 51, figs. 9-10. Givens, 1974:72, pi.
7, figs. 13, 16. Squires, 1977:table 1; 1983b:fig. 9b.

Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks 21

Figure 6. Llajas Formation gastropods. Unless otherwise indicated, views are apertural. a. Nerita cf. N. (Amphinerita) eorex Vokes, 1939,
LACMIP hypotype 6508, CSUN loc. 700, x 6. b and c. Velates perversus (Gmelin, 1791), UCMP hypotype 37435, UCMP loc. 7193 = CSUN
loc. 702, xl.3. b, abapertural view. d. Turritella meganosensis protumescens Merriam and Turner, 1937, LACMIP 6509, CSUN loc. 454,
x 1. e. Turritella andersoni Dickerson, 1916, LACMIP hypotype 6510, CSUN loc. 469d, x2. f and g. Turritella andersoni lawsoni Dickerson,
1916. All parts from CSUN loc. 374 and x 1.25. f, LACMIP hypotype 6511. g, LACMIP hypotype 6512. h. Turritella buwaldana Dickerson,

22 Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks

Rimella (Macilentos) macilenta White. Vokes, 1939:15 5—
156, pi. 20, figs. 1, 2, 4, 5.

Ectinochilus maci/entus (White). Stewart, 1946:93, pi. 11,
figs. 12-15. Givens and Kennedy, 1979:83, table 1.

Primary Type Material. USNM holotype 201 14, Domen-
gine Formation, about 3 km north of New Idria, section 16,
T 17 S, R 12 E, Priest Valley quadrangle, Fresno County,
California; CAS paratype 769, Llajas Formation, CAS lo-
cality 393.

Molluscan Stage Range. “Capay” through “Domengine.”
Geographic Distribution. San Diego through central Cal-
ifornia.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 371, 372, 373, 374, 440, 444, 445, 447, 449,
455, 458, 460, 471d, 472, 481, 484, 491, 493, 494, 498, 499,
508, 518, 538, 539. Outer shelf and slope facies: CSUN lo-
cality 482.

Remarks. This species is common in the shallow-marine
(transgressive) facies. The two localities at which it is most
abundant are 371 and 374. At 371 specimens are very well
preserved and some have the protoconch or the fragile outer
lip intact.

The diagnosis of the subgenus Macilentos by Clark and
Palmer (1923) was based on the paratype and other speci-
mens from the Llajas Formation, rather than on White’s
holotype from central California.

Family Seraphsidae Jung, 1974
Genus Paraseraphs Jung, 1974

Type Species. By original designation, Paraseraphs tetanus
Jung, 1974 [=Terebellum fusiforme of authors, not of La-
marck].

Paraseraphs erraticus (Cooper, 1894)

Figure 7a

Tornatina erratica Cooper, 1894:47, pi. 2, fig. 35. Waring,
1 9 1 7:pl. 15, fig. 11.

Terebellum californicum Vokes, 1939:157, pi. 20, figs. 7-8,
1 1.

Terebellum ( Terebellum ) erraticum (Cooper). KJeinpell and
Weaver, 1963:189, pi. 25, figs. 8-9.

Paraseraphs erraticus (Cooper). Jung, 1974:41, pi. 12, figs.
8-14; pi. 13, figs. 1-3. Givens and Kennedy, 1979:87,
tables 1, 3.

Primary Type Material. CAS holotype 608, Eocene strata.
Rose Canyon, San Diego, California.

Molluscan Stage Range. Uppermost “Capay” through
“Transition.”

Geographic Distribution. San Diego through central Cal-
ifornia.

Local Occurrence. Interfingering coastal alluvial-fan facies
and shallow-marine (transgressive) facies: CSUN localities
469d, 702. Shallow-marine (transgressive) facies: CSUN lo-
calities 371, 540.

Remarks. Only a few specimens were found, and most are
from locality 371. The largest specimen (Fig. 7a) is 35 mm
in height and is nearly twice the size of the holotype.

The presence of P. erraticus in the zone of interfingering
between the coastal alluvial-fan facies and the shallow-ma-
rine (transgressive) facies of the Llajas Formation extends
the molluscan stage range of this taxon into the uppermost
“Capay.” Previously, the lower range limit had been reported
as the “Domengine Stage” (Vokes, 1939; Jung, 1974).

Superfamily Cypraeacea
Family Cypraeidae Rafinesque, 1815
Subfamily Cypraeinae Rafinesque, 1815
Genus Cypraea Linne, 1758

Type Species. By subsequent designation (Montfort, 1810),
Cypraea tigris Linne, 1758.

Cypraea castacensis Stewart, 1927

Figure 7b

Cypraea castacensis Stewart, 1927:370, pi. 28, fig. 10. In-
gram, 1942:13, pi. 1, figs. 5-6.

Eocypraea castacensis (Stewart). Vokes, 1939:154, pi. 20,

figs. 9, 14.

Primary Type Material. UCMP holotype 1 1690, Tejon?
Formation, UCMP locality 452.

Molluscan Stage Range. “Domengine.”

Geographic Distribution. Simi Valley through central Cal-
ifornia.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 374, 444, 491.

Remarks. Specimens are relatively rare in the Llajas For-

1916, LACMIP hypotype 6513, CSUN loc. 373, x3. i. Turritella uvasana infera Merriam, 1941, abapertural view, UCLA hypotype 59359,
UCLA loc. 6616, xl.25. j. Turritella uvasana applinae Hanna, 1927, LACMIP hypotype 6514, CSUN loc. 374, xl. k. Architectonica
(Architectonica) llajasensis Sutherland, 1966, dorsal view, LACMIP 6515, CSUN loc. 371, x3. 1. Architectonica ( Stellaxis ) cognata Gabb,
1864, dorsal view, LACMIP hypotype 6516, CSUN loc. 374, x 1, m. Benoistia umpquaensis Turner, 1938, abapertural view, UCLA hypotype
59274, CSUN loc. 374, xl. n and o. Cerithiopsis llajasensis n. sp. All parts from CSUN loc. 371. n, LACMIP holotype 6517, x3.5. o,
abapertural view, LACMIP paratype 6518, x3. p. Cirsotrema sp., UCLA hypotype 59275, CSUN loc. 374, x 1.5. q. Calyptraea diegoana
(Conrad, 1855), side view, LACMIP hypotype 6519, CSUN loc. 371, x 1.5. r. Xenophora stocki Dickerson, 1916, internal mold, side view,
LACMIP hypotype 6520, CSUN loc. 374, xl. s. Ectinochilus ( Macilentos ) macilentus (White, 1889), LACMIP hypotype 6521, CSUN loc.
371, x 1.5.

Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks 23

mation. One of Vokes’ (1939:154, pi. 20, fig. 9) hypotypes
is from Devil Canyon, just north of locality 373.

Superfamily Naticacea

Family Naticidae Forbes, 1838

Subfamily Ampullospirinae Cox, 1930

Genus Eocernina Gardner and Bowles, 1934

Type Species. By original designation, Natica hannibali
Dickerson, 1914.

Eocernina hannibali (Dickerson, 1914)

Figure 7c

Natica hannibali Dickerson, 1 9 1 4: 1 1 9, pi. 12, figs. 5a-b; 1916:
508, pi. 38, figs. 9a-b.

Natica (Cryptonatica) hannibali Dickerson. Waring, 1917:
pi. 15, figs. 21-23.

Ampullina hannibali (Dickerson). Hanna, 1927:306, pi. 48,
figs. 1-3, 10.

Ampullina (Globularia) hannibali (Dickerson). Clark, 1929:
pi. 11, fig. 12.

Cernina ( Eocernina ) hannibali (Dickerson). Turner, 1938:
87-88, pi. 19, fig. 3. Vokes, 1939:172, pi. 22, figs. 1, 3.
Weaver, 1943:348-349, pi. 71, figs. 8-9, 21, 23.
Eocernina hannibali ( Dickerson). Hanna and Hertlein, 1943:
fig. 62-17. Marincovich, 1977:229-231, pi. 18, fig. 14; pi.
19, figs. 1-4. Givens and Kennedy, 1979:87, tables 1. 3.
Squires, 1983b:fig. 9c.

Globularia ( Eocernina ) hannibali (Dickerson). Stewart, 1946:
pi. 11, fig. 18. Givens, 1974:75, pi. 9, figs. 1, 3. Squires,

1 977:table 1 .

Primary Type Material. CAS holotype 243, Umpqua For-
mation, CAS locality 25.

Molluscan Stage Range. “Capay” through “Transition.”
Geographic Distribution. San Diego, California through
southwestern Oregon.

Local Occurrence. Interfingering coastal alluvial-fan facies
and shallow-marine (transgressive) facies: CSUN localities
469a, 542, 545, 702, 703. Shallow-marine (transgressive) fa-
cies: CSUN localities 37 1 , 372, 373, 374, 439, 440, 444, 445,
447, 449, 450, 451, 455, 458, 460, 465, 469e, 469g, 470b,
471b, 47 le, 477, 479, 480, 484, 486, 488, 493, 494, 497,
498, 499, 500, 508, 509, 5 10, 5 14, 5 1 7, 523, 538, 704. Outer
shelf and slope facies: CSUN locality 482. Shallow-marine
(regressive) facies: CSUN localities 512c, 51 2d.

Remarks. E. hannibali is one of the most common species
of the Llajas Formation. It occurs in all the marine facies of
the formation and is particularly characteristic of the shallow-
marine (transgressive) facies. According to Marincovich
( 1 977:23 1 ), this species may occur in prolific numbers. This
is especially true for the “Stewart bed” in the vicinity of
localities 374 and 444. There, E. hannibali is the most com-

mon megafossil with nearly complete growth series present
also.

Genus Tejonia Hanna and Hertlein, 1943

Type Species. By monotypy, Natica alveata Conrad, 1855,
not Troschel, 1852.

Tejonia moragai (Stewart, 1927)

Figure 7d

Natica alveata Conrad, 1855:10; 1857:321, pi. 2, figs. 8,8a.
Not Natica alveata Troschel, 1852:159, pi. 5, fig. 3.
Amauropsis alveata (Conrad). Gabb, 1864:110, pi. 19, fig.
59; pi. 2 1 , fig. 111. Dickerson, 1 9 1 6:pl. 38, fig. 7. Anderson
and Hanna, 1925: 1 19-120, pi. 6, fig. 2; pi. 7, fig. 1; pi. 15,
fig. 17.

Amauropsis alveatus (Conrad). Arnold, 1 907 :pl. 39, fig. 8.
Amaurellina moragai Stewart, 1927:334-336, pi. 28, fig. 3
[new name for Natica alveata Conrad, 1855, preoccupied].
Clark, 1929:pl. 14, figs. 3, 9. Kleinpell and Weaver, 1963:
188, pi. 25, figs. 1-2.

Amaurellina moragai lajollaensis Stewart, 1927:336, pi. 28,
fig. 2.

Amaurellina hendoni Turner, 1938:86-87, pi. 20, figs. 7, 8,
11. Vokes, 1939:173, pi. 22, figs. 7, 10. Weaver, 1943:345,
pi. 70, figs. 12-13, 16-17.

Tejonia lajollaensis (Stewart). Hanna and Hertlein, 1 943:
172, fig. 62-30. Givens, 1974:74, pi. 8, fig. 5.

Tejonia moragai (Stewart). Givens, 1974:74, pi. 8, fig. 8.
Marincovich. 1977:232-236, pi. 19, figs. 6-12. Squires,
1977:table 1. Givens and Kennedy, 1979:tables 1, 3-4.

Primary Type Material. Holotype unknown for Natica
alveata Conrad (USNM collection?), Tejon Formation,
Grapevine Canyon, Tejon quadrangle, Kern County, Cali-
fornia. UCMP “holotype” 31387 of Amaurellina moragai
Stewart, Tejon Formation, UCMP locality 7200. See Marin-
covich, 1977:235 for a discussion of Stewart’s (1927) “ho-
lotype.”

Molluscan Stage Range. “Domengine” through “Tejon.”
Geographic Distribution. San Diego, California through
southwestern Oregon.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 371, 373, 467. Shallow-marine (regressive)
facies: CSUN locality 489.

Remarks. At locality 373, T. moragai is fairly common
and occurs as a growth series. Some specimens have their
protoconchs present. Elsewhere, only single specimens were
found. This species is characterized by its strongly tabulate
adult whorls.

Genus Pachycrommium Woodring, 1928

Type Species. By original designation, Amaura guppyi
Gabb, 1873.

Pachycrommium clarki (Stewart, 1927)

Figure 7e

Amauropsis alveata (Conrad). Arnold, 1910:pl. 4, fig. 21.
Waring, 191 7:pl. 15, fig. 25. Dickerson, 191 5:pl. 5, fig. 9

24 Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks

[not Natica alveata Conrad, 1855 = Tejoma moragai
(Stewart, 1927), treated herein].

Amaurellina ( Euspirocrommium ) clarki Stewart, 1927:336-
339, pi. 26, figs. 8-9 [new name, in part, for Amauropsis
alveata (Conrad, 1855), preoccupied]. Clark, 1929:pl. 1 1,
fig. 10. Turner, 1938:86, pi. 20, fig. 3. Weaver, 1943:345,
pi. 70, figs. 10, 18. Kleinpell and Weaver, 1963:188, pi.
27, fig. 15.

Amaurellina clarki Stewart. Gardner and Bowles, 1934:246,
figs. 6, 8.

Amaurellina ? multiangulata Vokes, 1939: 1 74, pi. 22, figs. 2,
8, 13.

Pachycrommiuml clarki (Stewart). Vokes, 1939:175, pi. 22,
figs. 11, 30. Givens, 1974:73, pi. 8, figs. 6, 10. Squires,
1977:table 1.

Amaurellina ( Euspirocrommium ?) clarki Stewart. Stewart,
1946:pl. 1 1, fig. 3.

Pachycrommium clarki (Stewart). Marincovich, 1977:238-
241, pi. 20, figs. 4-10. Squires, 1983b:fig. 9d.

Primary Type Material. UCMP holotype 31385, UCMP
paratype 31386 of Amaurellina ( Euspirocrommium ) clarki
Stewart, Llajas Formation, UCMP locality 7004 = CSUN
locality 374.

Molluscan Stage Range. “Capay” through “Tejon. ”
Geographic Distribution. San Diego, California through
northern Washington.

Local Occurrence. Interfingering coastal alluvial-fan facies
and shallow-marine (transgressive) facies: CSUN localities
452, 469a, 702, 703. Shallow-marine (transgressive) facies:
CSUN localities 37 1, 372, 373, 374, 444, 447, 455, 457, 466,
469e, 47 le, 479, 484, 486, 488, 491, 492, 493, 494, 498,
500, 505, 506, 507, 508, 5 1 3, 5 14, 5 1 8, 528, 537, 538. Outer
shelf and slope facies: CSUN locality 482. Outer shelf and
slope channel facies: CSUN locality 541.

Remarks. This species is especially abundant at localities
371 and 374, and at both localities it occurs as a growth
series.

Subfamily Polinicinae
Finlay and Marwick, 1937

Genus Polinices Montfort, 1810

Type Species. By original designation, Polinices albus
Montfort, 1810.

Subgenus Euspira Agassiz in
J. Sowerby, 1838b

Type Species. By subsequent designation (Harris, 1897),
Ampullaria sigaretina Lamarck, 1804.

Polinices ( Euspira ) nuciformis (Gabb, 1864)

Figure 7f

Lunatia nuciformis Gabb, 1864:107, pi. 28, fig. 218. Dick-
erson, 19 1 6: pi. 39, fig. 4.

Lunatia cowlitzensis Dickerson, 1915:57, pi. 4, figs. 12a-b.
Natica nuciformis (Gabb). Anderson and Hanna, 1925:1 16,
pi. 10, fig. 8.

Polinices (Euspira) nuciformis (Gabb). Clark and Woodford,
1927: 121, pi. 21, figs. 16-17. Turner, 1938:88, pi. 20, figs.
4-5. Clark, 1 938:703-704, pi. 4, figs. 26,31. Vokes, 1 939:

168, pi. 21, figs. 12-14. Weaver, 1943:342-343, pi. 70,
figs. 1-2; pi. 103, fig. 2.

Euspira nuciformis ( Gabb). Stewart, 1927:323-324, pi. 30,
fig. 1 6; 1 946:pl. 1 1, fig. 16. Weaver, 1 953:29. Givens, 1974:
77, pi. 7, fig. 14. Squires, !977:table 1.

Polinices (Euspira) nuciformis var. cowlitzensis (Dickerson).

Weaver, 1943:343, pi. 69, figs. 10-11, 13-19.

Polinices (Euspira) nuciformis (Gabb). Marincovich, 1977:
281-285, pi. 26, figs. 6-9.

Primary Type Material. ANSP lectotype 4213, ANSP
paralectotypes 42 1 3 (two specimens), Tejon Formation, Live
Oak Canyon, Kern County, California.

Molluscan Stage Range. Upper Paleocene through “Te-
jon.”

Geographic Distribution. San Diego, California through
southwestern Oregon.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN locality 374.

Remarks. Only three specimens were found in the Llajas
Formation.

Genus Neverita Risso, 1826

Type Species. By monotypy, Neverita josephina Risso,
1826.

Subgenus Neverita s.s.

Neverita ( Neverita ) globosa Gabb, 1869

Figure 7g

Neverita globosa Gabb, 1869:161, pi. 27, fig. 39. Dickerson,
1916:510, pi. 39, figs. 5a-b. Stewart, 1927:326-327, pi. 28,
fig. 6. Clark and Woodford, 1927:121-122, pi. 22, figs. 5-
10. Turner, 1938:89, pi. 19, figs. 6-7, 13-15. Vokes, 1939:

169, pi. 21, figs. 9, 15-19. Givens and Kennedy, 1979:
tables 1-3.

Neverita weaver i Dickerson, 1915:57, pi. 4, figs. lOa-b.
Neverita nomlandi Dickerson, 1917:173-174, pi. 30, figs.
2a-b.

Polinices weaven (Dickerson). Turner, 1938:86, pi. 20, figs.
14, 16.

Neverita globosa reefensis Vokes, 1 939: 1 69, pi. 21, figs. 24-
25.

Polinices (Neverita) weaveri (Dickerson). Weaver, 1943:340,
pi. 68, figs. 16-17; pi. 69, fig. 3.

Polinices (Neverita) globosa (Gabb). Weaver, 1943:339, pi.

68, figs. 21, 24; pi. 69, figs. 5-6; pi. 100, fig. 29.

Polinices (Neverita) nomlandi (Dickerson). Weaver, 1943:
340, pi. 69, figs. 8, 9, 12.

Neverita (Neverita) globosa Gabb. Givens, 1974:76. Marm-
covich, 1977:312-316, pi. 28, figs. 10-15; pi. 29, figs.
1-3.

Neverita (Glossaulaxl) globosa Gabb. Givens and Kennedy,
1976:965-966, pi. 2, figs. 5-14, 16, 18-19.

Neverita globosa Gabb. Givens and Kennedy, 1979:tables
1-3.

Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks 25

Primary Type Material. MCZ holotype 27859, Domen-
gine? Formation, 1 6 km west of Griswold’s, on the road from
San Juan to New Idria, and southeast of the “Sheep Well,”
T 15 S, R 9 E, Priest Valley quadrangle, San Benito County,
California.

Molluscan Stage Range. “Meganos” through upper Eocene.
Geographic Distribution. San Diego, California through
western Washington.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN locality 704.

Remarks. Only three specimens were found in the Llajas
Formation.

Subfamily Sininae Wenz, 1941
Genus Sinum Roding, 1798

Type Species. By subsequent designation (Dali, 1915), He-
lix haliotoidea Linne, 1758.

Simon obliquum (Gabb, 1 864)

Figure 7h

Naticina obliqua Gabb, 1 864: 1 09, pi. 21, fig. 11 2. Dickerson,
1 9 1 5 :pl. 5, figs. 5a-b.

Simon occidentis Weaver and Palmer, 1922:32-33, pi. 11,
hgs. 8, 26. Weaver, 1943:351, pi. 71, fig. 15.

Sinum coryliforme Anderson and Hanna, 1925:120, pi. 9,
fig. 10; pi. 10, fig. 15; pi. 15, fig. 8.

Sinum obliquum (Gabb). Stewart, 1927:327, pi. 30, fig. 7a.
Clark, 1938:704, pi. 3, figs. 32, 37. Weaver, 1 943:350-
351, pi. 71, fig. 13; pi. 103, fig. 6. Hickman, 1969:85-88,
pi. 11, figs. 9-10. Marincovich, 1977:347-350, pi. 33, figs.
1-12. Squires, 1977:table 1. Givens and Kennedy, 1979:
table 4.

Primary Type Material. ANSP lectotype 4215, ANSP
paralectotypes 4215 (seven specimens), Tejon Formation,
Fort Tejon area, Kern County, California.

Molluscan Stage Range. “Capay” through lower Oligo-
cene.

Geographic Distribution. San Diego, California through
southwestern Washington.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 371, 539.

Remarks. S', obliquum is most common at locality 371
where it is well preserved and occurs as a growth series. At
locality 539, only a single specimen was found.

Subfamily Naticinae Forbes, 1838

Genus Natica Scopoli, 1777

Type Species. By subsequent designation (Anton, 1839),
Nerita vitellus Linne, 1758.

Subgenus Naticarius Dumeril, 1806

Type Species. By monotypy, Nerita canrena Linne, 1758.

Natica ( Naticarius ) aff.

N. ( N .) uvasana Gabb, 1864

Figure 7i

Local Occurrence. Interfingering coastal alluvial-fan facies
and shallow-marine (transgressive) facies: CSUN locality 545.
Shallow-marine (transgressive) facies: CSUN localities 371,
373, 486.

Remarks. This taxon is most common at locality 37 1 where
32 specimens make up a growth series. The largest specimen
is 18 mm in height. Elsewhere, only a few specimens were
found.

The Llajas specimens agree with the description of N. ( N .)
uvasana given by Marincovich (1977:390) in possessing ir-
regularly spaced collabral wrinkles immediately below the
suture, an open umbilicus, and a robust umbilical callus and
funicle. The Llajas specimens differ in that they lack a parietal
callus.

Superfamily Tonnacea
Family Cassididae Latreille, 1825
Genus Galeodea Link, 1 807

Type Species. By monotypy, Buccinum echinophorum
Linne, 1758.

Subgenus Caliagaleodea Clark, 1 942

Type Species. By original designation, Caliagaleodea cal-
ifornica C lark, 1942.

Galeodea ( Caliagaleodea ) californica
Clark, 1942
Figure 7j

Galeodea ( Caliagaleodea ) californica Clark, 1942:1 18-1 19,

pi. 19, figs. 15-19.

Galeodea californica Clark. Givens and Kennedy, 1979:

table 1.

Primary Type Material. UCMP holotype 34376, UCMP
paratype 34377, Llajas Formation, UCMP locality 7004 =
CSUN locality 374.

Molluscan Stage Range. “Domengine.”

Geographic Distribution. San Diego through Simi Valley,
California.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 374, 444, 445, 447, 458.

Remarks. Nearly all the specimens of this species were
found at the type locality. The 14 specimens at locality 374
make up a nearly complete growth series, but most occur as
internal molds. A few other specimens were found elsewhere
in the “Stewart bed” in the vicinity of locality 374.

Subgenus Gomphopages Gardner, 1939

Type Species. By original designation, Galeodea turneri
Gardner, 1939.

26 Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks

Galeodea ( Gomphopages ) susanae Schenck, 1926

Figure 7k

Galeodea susanae Schenck, 1926:85, pi. 15, figs. 3-7. Turner,
1938:92, pi. 18, fig. 18. Weaver, 1943:402-403, pi. 78,
figs. 2-3.

Galeodea ( Gomphopages ) susanae Schenck. Durham, 1942:
184. Givens, 1974:78.

Primary Type Material. CAS holotype 1753, CAS para-
type 1754, Llajas Formation, CAS locality 372; CAS para-
type 1755, Llajas Formation, CAS locality 364; CAS para-
type 1756, near Roseburg, Oregon.

Molluscan Stage Range. “Domengine,” “Transition”?.
Geographic Distribution. Simi Valley, California through
southwestern Oregon.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN locality 371.

Remarks. The 16 specimens from locality 371 make up a
growth series, and they are extremely well preserved. There
is excellent preservation of spines and, in a few cases, the
long anterior canal. The largest specimen is 53 mm in height.

Genus Phalium Link, 1807

Type Species. By subsequent designation (Dali, 1909),
Buccinum glaucum Linne, 1758.

Subgenus Semicassis Morch, 1852

Type Species. By subsequent designation (Flarris, 1897),
Cassis japonica.

Phalium ( Semicassis ) tuberculiformis
(Hanna, 1924)

Figure 71

Mono (Sconsia) tuberculatus Gabb, 1864:104, pi. 19, fig. 57.

Arnold, 1907:pl. 39, fig. 9.

Not Cassidaria tuberculata Risso, 1826:186.

Morio tuberculatus Gabb. Dickerson, 1913:264.

Galeodea tuberculata (Gabb). Dickerson, 191 6:pl. 42, fig. 2.
Galeodea (Morio) tuberculata (Gabb). Waring, 1 9 1 7:pl. 15,
fig. 17.

Galeodea tuberculiformis Hanna, 1924:167 [new name for
Morio ( Sconsia ) tuberculatus Gabb, 1864, preoccupied].
Schenck, 1926:83-84, pi. 14, figs. 12-16. Stewart, 1927:
380-38 1 , pi. 28, fig. 11. Vokes, 1 939: 1 49-1 50, pi. 1 9, figs.
19, 21, 23-27.

Coalingodea tuberculiformis (Hanna). Durham, 1942:186, pi.
29, figs. 5, 9. Givens, 1974:78-79, pi. 8, fig. 7. Squires,
1977:table 1.

Cassis (Coalingodea) tuberculata (Gabb). Abbott, 1968b: 59-
60, pi. 34.

Phalium (Semicassis) tuberculiformis (Hanna). Givens and
Kennedy, 1979:82, 95, tables 1, 3.

Primary Type Material. ANSP lectotype 4343 of Morio
(Sconsia) tuberculatus Gabb and Galeodea tuberculiformis
Hanna, Tejon? Formation, Martinez, California.

Molluscan Stage Range. “Capay”?, “Domengine” through
“Transition.”

Geographic Distribution. San Diego, California through
northwestern Washington.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 371, 372, 373, 374, 440, 444, 445, 455, 458,
471b, 480, 488, 494, 514, 537. Outer shelf and slope channel
facies: CSUN locality 541. Shallow-marine (regressive) fa-
cies: CSUN localities 462, 51 2d.

Remarks. This taxon is most common and best preserved
at localities 371 and 374, where it occurs in growth series.
The beaded appearance of the whorls and the toothed outer
lip are well preserved.

Dickerson (1913) reported this species from “Capay”-age
strata in central California, but he gave no specific locality
information nor did he figure the species. For these reasons,
the “Capay”-age occurrence is uncertain.

The presence of P. (S.) tuberculiformis in the Llajas For-
mation is one of the earliest occurrences of Phalium on the
West Coast. It is also known from similar age strata in San
Diego County (Givens and Kennedy, 1979).

Family Cymatiidae Iredale, 1913
Genus Cymatium Roding, 1798

Type Species. By subsequent designation (Dali, 1 904), Mu-
rex femorale Linne, 1758.

Subgenus Septa Perry, 1810

Type Species. By monotypy, Septa scarlatina Perry, 1810
(=Murex rubecula Linne, 1758).

Cymatium {Septa) janetae Squires, 1983a

Figure 7 m

Cymatium (Septa) janetae Squires, 1983a:355-357, figs.

2a-d.

Primary Type Material. UCLA holotype 59191, Llajas
Formation, CSUN locality 444; UCLA paratype 59192, Lla-
jas Formation, CSUN locality 445; UCLA paratype 59193,
Llajas Formation, CSUN locality 371.

Molluscan Stage Range. “Domengine.”

Geographic Distribution. Simi Valley, California.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 371, 374, 444, 498.

Remarks. Only juvenile specimens occur at localities 371
and 498. Only adult specimens occur in the “Stewart bed”
at and near localities 374 and 444. The figured specimen
(Fig. 7m) is the holotype. C. (S’.) janetae is, at present, the
earliest species worldwide referable to Septa (Squires, 1 983a).

Genus Ranella Lamarck, 1816

Type Species. By subsequent designation (Children, 1 823),
Ranella gigantea Lamarck, 1816 [=Murex o/earium Linne,
1758],

Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks 27

Figure 7. Llajas Formation gastropods (continued). Unless otherwise indicated, views are apertural. a. Paraseraphs erraticus (Cooper, 1894),
abapertural view, LACM1P hypotype 6522, CSUN loc. 371, x 1. b. Cypraea castacensis Stewart, 1927, LACM1P hypotype 6523, CSUN loc.
374, x 1.25. c. Eocermna hannibali (Dickerson, 1914), hypotype LACMIP 6524, CSUN loc. 374, x 1. d. Tejonia moragai (Stewart, 1927),
LACMIP hypotype 6525, CSUN loc. 467, x l. e. Pachycrommium clarki (Stewart, 1927), LACMIP hypotype 6526, CSUN loc. 371, x 1. f.
Polinices ( Euspira ) nuciformis (Gabb, 1864), UCLA hypotype 59276, CSUN loc. 374, x 1 . g. Neverita (N event a) globosa Gabb, 1 869, LACMIP
hypotype 6527, CSUN loc. 704, x 1.5. h. Sinum obliquum (Gabb, 1864), side view, LACMIP hypotype 6528, CSUN loc. 371, x2. i. Natica

28 Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks

Ranella katherineae Squires, 1983a

Figure 7n

“ Ranella ” sp. Smith, 1970:523.

Ranella katherineae Squires, 1983a:357-359, figs. 2e-g.

Primary Type Material. UCLA holotype 45969, Llajas
Formation, CSUN locality 374.

Molluscan Stage Range. “Domengine.”

Geographic Distribution. Simi Valley, California.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN locality 374.

Remarks. Only a single, nearly complete adult specimen
was found in the Llajas Formation. This specimen is the
holotype and the figured specimen (Fig. 7n). R. katherineae
is one of the earliest West Coast species of Ranella s.s. (Squires,
1983a).

Family Bursidae Thiele, 1925
Genus Olequahia Stewart, 1927

Type Species. By original designation, Cassidaria wash-
ingtoniana Weaver, 1912.

Olequahia domenginica (Vokes, 1939)

Figure 7o

Ranella domenginica Vokes, 1939:147-148, pi. 19, figs. 6,
20.

Olequahia hornii domenginica (Vokes). Stewart, 1946:
table 1.

Olequahia domenginica (Vokes). Givens, 1974:80, pi. 9, figs.
4-5. Givens and Kennedy, 1979:table 1.

Primary Type Material. UCMP syntypes 1 5803 and 1 5804,
Domengine Formation, UCMP locality 672.

Molluscan Stage Range. “Domengine Stage.”

Geographic Distribution. San Diego through central Cal-
ifornia.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 371, 374, 444, 445, 47 Id, 494.

Remarks. Specimens are most common in the “Stewart
bed” at locality 374 where they form a growth series.

Vokes (1939:147-148) used non-type, supplementary
specimens from the vicinity of locality 374 to augment his
description of this taxon.

Genus Ranellina Conrad, 1865

Type Species. By original designation, Ranellina maclurii
Conrad, 1865.

Ranellina pilsbryi Stewart, 1927

Figure 7p

Not Clavatulal californica Conrad, 1855:11; 1857:322, pi.
2, fig. 11.

Fusus californicus (Conrad). Gabb, 1864:85-86, pi. 28, figs.
205-205a.

Nyctilochus hornii (Gabb). Dickerson, 1915 :pl. 7, fig. 8. Dick-
erson, 1916:pl. 42, fig. 3.

Clavilithes californicus (Conrad). Anderson and Hanna, 1925:
63-64, pi. 13, figs. 1-2.

Ranellina pilsbryi Stewart, 1927:384-385, pi. 30, figs. 8-9
[new name for Fusus californicus (Conrad) Gabb, 1864,
misidentified]. Turner, 1938:91, pi. 16, fig. 3. Vokes, 1939:
148, pi. 19, figs. 10, 17. Weaver, 1943:417, pi. 82, fig. 7.
Givens, 1974:80, pi. 9, fig. 12. Givens and Kennedy, 1979:
tables 1,4.

Primary Type Material. UCMP holotype 31382, UCMP
paratype 31383 of Ranellina pilsbryi Stewart, Tejon For-
mation, UCMP locality 7182.

Molluscan Stage Range. “Domengine” through “Tejon.”
Geographic Distribution. San Diego, California to south-
western Oregon.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 371, 373, 486, 498.

Remarks. This species is most common at locality 371
where seven specimens were found.

Smith (1975:table 1) reported “ Ranellina ” sp. afif. R. pils-
bryi from “Martinez”-age strata of central California.

Family Ficidae Conrad, 1867

Genus Ficopsis Conrad, 1866

Type Species. By subsequent designation (Stewart, 1927),
Hemifusus remondii Gabb, 1864.

Ficopsis cooperiana Stewart, 1927

Figure 8a

Fusus (Hemifusus) cooperii Gabb, 1864:86, pi. 28, fig. 207.
Not Fusus cooperi Conrad, 1834:148.

Ficopsis cooperii (Gabb). Dickerson, 1915:61-62, pi. 6, fig.

11; 1916:492-493, pi. 37, fig. 7. Clark, 1929:pl. 9, fig. 13.
Ficopsis cooperiana Stewart, 1927:378-379 [new name for
Fusus (Hemifusus) cooperii Gabb, 1864, preoccupied].
Clark, 1929:pl. 9, fig. 13. Givens, 1974:81, pi. 9, figs. 7,
9. Givens and Kennedy, 1979:87, table 3.

Primary Type Material. UCMP holotype 11691 of Fusus
(Hemifusus) cooperii Gabb and Ficopsis cooperiana Stewart,
Eocene strata, UCMP locality 2226.

(Naticarius) aff. N. (N.) uvasana Gabb, 1864, LACMIP hypotype 6529, CSUN loc. 371, x 2. j. Galeodea (Caliagaleodea) californica Clark,
1942, internal mold, abapertural view, LACMIP topotype and hypotype 6530, CSUN loc. 374, xl. k. Galeodea (Gomphopages) susanae
Schenck, 1926, LACMIP hypotype 6531, CSUN loc. 371, x 1.5. 1. Phalium (Semicassis) tuberculiformis (Hanna, 1924), LACMIP hypotype
6532, CSUN loc. 371, xl. m. Cymatium (Septa) janetae Squires, 1983a, UCLA holotype 59191, CSUN loc. 444, xl. n. Ranella katherineae
Squires, 1983a, UCLA holotype 45969, CSUN loc. 374, x0.5. o. Olequahia domenginica (Vokes, 1939), LACMIP hypotype 6533, CSUN
loc. 374, x l. p. Ranellina pilsbryi Stewart, 1927, LACMIP hypotype 6534, CSUN loc. 371, x2.5.

Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks 29

Molluscan Stage Range. “Domengine” through “Transi-
tion.”

Geographic Distribution. San Diego through central Cal-
ifornia.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN locality 374.

Remarks. Only a few specimens were found in the Llajas
Formation.

Ficopsis remondii crescentensis
Weaver and Palmer, 1922

Figure 8 b

Ficopsis angulatus Weaver, 1905:1 19, pi. 13, fig. 5.

Not Pyrula angiilata Edwards, 1866:pl. 4.

Ficopsis remondii (Gabb) var. crescentensis Weaver and
Palmer, 1922:39-40, pi. 11, fig. 14 [new name for Ficopsis
angulatus Weaver, 1 905, preoccupied], Stewart, 1930:40-
41. Turner, 1938:93, pi. 15, fig. 19. Weaver, 1943:399, pi.
77, fig. 10.

Ficopsis remondii crescentensis Weaver and Palmer. Vokes,
1939:152-153. Givens, 1974:82, pi. 9, fig. 11. Squires,
1977:table 1. Givens and Kennedy, 1979:87, tables 1, 3.
Ficopsis crescentensis Weaver and Palmer. Stewart, 1946:pl.
1 1, fig. 17.

Primary Type Material. UCMP holotype 1 1887 of Ficop-
sis angulatus Weaver, Eocene strata, UCMP locality 337.
UW holotype 205 (CAS 7616) of Ficopsis remondii crescen-
tensis Weaver and Palmer, Crescent? Formation, UW local-
ity 358.

Molluscan Stage Range. “Capay” through “Transition.”
Geographic Distribution. San Diego, California through
northwestern Washington.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 371, 373, 374, 477, 479, 486.

Remarks. Specimens are most common at locality 371
although the largest specimens occur at locality 374. This
taxon is characterized by the tri-carination of the body whorl
and the cancellate sculpture.

Order Neogastropoda

Superfamily Muricacea

Family Muricidae da Costa, 1776

Subfamily Muricinae da Costa, 1776

Genus Laevityphis Cossmann, 1903

Type Species. By original designation, Typhis coronarius
Deshayes, 1865.

Subgenus Laevityphis s.s.

Laevityphis ( Laevityphis ) antiquus
(Gabb, 1864)

Figure 8c

Typhis antiquus Gabb, 1864:82, pi. 18, fig. 31. Stewart, 1927:
387-388, pi. 27, figs. 7-8.

Laevityphis ( Laevityphis ) antiquus (Gabb). Keen, 1944:58,

63. Givens, 1974:82.

Laevityphis antiquus (Gabb). Givens and Kennedy, 1979:

table 3.

Primary Type Material. ANSP lectotype 4335, Tejon?
Formation, Martinez, California.

Molluscan Stage Range. “Domengine” through “Tejon.”

Geographic Distribution. San Diego through central Cal-
ifornia.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN locality 371.

Remarks. Only a single specimen was found in the Llajas
Formation.

Family Buccinidae Rafinesque, 1815
Genus Ancistrolepis Dali, 1895

Type Species. By original designation, Chrysodomus eu-
cosmius Dali, 1891.

Ancistrolepis ? carolineae n. sp.

Figures 8d-e

Diagnosis. Bucciniform shell with two spiral ribs on spire
and about six spiral ribs on body whorl, interspaces char-
acteristically concave with numerous fine lirae. Due to the
fact that the canal area has been broken off on each of the
Llajas specimens, positive generic assignment cannot be made.

Ancistrolepis' ? carolineae n. sp. is very similar to A. clarki
Tegland (1933: 1 3 1-132, pi. 12, fig. 14) and A. clarki variety?
Tegland (1933:132, pi. 12, figs. 15-17) from the Oligocene
Blakeley Formation, northwestern Washington. A.1 caroli-
neae differs from A. clarki and A. clarki var.? in the following
features: less convex whorls, narrower and shorter spire,
equally spaced primary spiral ribs on the spire, less inflated
spiral ribs, and smaller shell size. Unfortunately, the anterior
canal area is missing in the Llajas specimens and comparison
to the ornamented canal areas of A. clarki and A. clarki var.?
cannot be made. A. clarki and A. clarki var.? differ from each
other only in the nature of the canal area ornamentation.

Description. Medium shell, bucciniform, with five or six
angulate whorls. Suture moderately impressed. Protoconch
smooth with low, rounded whorls and shallow sutures.

Spire whorls with two very prominent, primary spiral ribs
that divide the whorls into three equal concave areas covered
with numerous fine lirae. Body whorl with five to six primary
spiral ribs, interspaces concave and covered with numerous
fine lirae.

Columella smooth and twisted. Aperture straight. Thick-
ened outer lip. Anterior canal area missing. Dimensions of
largest specimen (Figs. 8d-e): height (incomplete) 32 mm,
body whorl width 20 mm.

Primary Type Material. UCLA holotype 59401, UCLA
paratypes 59402 (three specimens), Llajas Formation, CSUN
locality 374.

Molluscan Stage Range. “Domengine.”

Geographic Distribution. Simi Valley, California.

30 Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN locality 374.

Remarks. Specimens of this new species were found only
at CSUN locality 374 (“Stewart bed”). A total of 12 speci-
mens was found, and preservation is mostly as internal molds.
A few specimens (i.e., the primary type material) have por-
tions of the original shell material.

Ancistrolepis Dali is differentiated from Chrvsodomus in
three respects; namely, a shorter canal, a degenerate radula,
and in the nature of the operculum. Although only the canal
feature can be used in fossil material, Tegland (1933) con-
sidered that the outline of the shell and detail of sculpturing
of A. clarki was so close to the Recent species, Ancistrolepis
eucosimus Dali (1895:709, pi. 29, fig. 7), the type species of
the genus, that reference to that genus rather than Chryso-
domus was warranted. The Llajas specimens of A.? carolineae
n. sp. are very similar to those of A. clarki and A. clarki var.?,
and they are not similar to West Coast Paleogene species of
Chrysodomus in terms of sculpture.

If A.1 carolineae does belong to Ancistrolepis, it would
extend the earliest occurrence of this genus to the early middle
Eocene, and A. carolineae would be the oldest known species
referable to this genus. It would be the first occurrence of
this genus in California.

Etymology. The species is named for Caroline Squires.

Superfamily Buccinacea
Family Nassariidae Iredale, 1916
Genus Mo/opophorus Gabb, 1869

Type Species. By monotypy, Bullia (Molopophorus) striata

Gabb, 1869.

Molopophorus cretaceus (Gabb, 1 864)

Figure 8f

Nassa cretacea Gabb, 1864:97, pi. 18, fig. 49.
Molopophorus cretaceus ( Gabb). Stewart, 1927:391-392, pi.

28, fig. 9. Vokes, 1939:141-142, pi. 19, fig. 4.

Primary Type Material. ANSP lectotype 4197, Domen-
gine Formation, Bulls Head Point, near Martinez, California.

Mollusean Stage Range. “Domengine.”

Geographic Distribution. Simi Valley through central Cal-
ifornia.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN locality 467.

Remarks. Only two specimens were found in the Llajas
Formation, and the apertures are covered or smashed.

Family Fasciolariidae Gray, 1853
Genus Clavilithes Swainson, 1840

Type Species. By subsequent designation (Grabau, 1904),
Fusus parisiensis Mayer-Eymar, 1877 [=Fusus longaevus La-
marck, 1803, not Solander, 1766],

Clavilithes tabulatus (Dickerson, 1913)

Figure 8g

Clavella tabulata Dickerson, 1913:283, pi. 12, fig. 7.
Clavilithes tabulatus (Dickerson). Clark and Vokes, 1936:
874, pi. 1, fig. 3 (holotype refigured). Givens, 1974:85, pi.
10, figs. 4-5.

Clavilithes cf. C. tabulatus (Dickerson). Crowell and Susuki,
1959:588-589, pi. 2, figs. 6-7.

Primary Type Material. UCMP holotype 1 1753, Capay
Formation, UCMP locality 1853.

Mollusean Stage Range. “Capay” through "Domengine.”
Geographic Distribution. Orocopia Mountains, southern
California through central California.

Local Occurrence. Interfingering coastal alluvial-fan facies
and shallow-marine (transgressive) facies: CSUN locality 703.
Shallow-marine (transgressive) facies: CSUN localities 371,
374, 704.

Remarks. Only two specimens were found. The presence
of C. tabulatus in the middle part of the Llajas extends the
mollusean stage range of this taxon into the “Domengine
Stage.” Previously it had been reported as confined to the
“Capay Stage” (Clark and Vokes, 1936; Crowell and Susuki,
1959; Givens, 1974).

Clavilithes n. sp. A Clark and Yokes, 1936

Figure 8h

Clavilithes n. sp. Clark and Vokes, 1936:874, pi. 1, fig. 1.
Givens and Kennedy, 1976:973, pi. 4, figs. 9, 12.

Type Material. UCMP hypotype 15468, Llajas Forma-
tion, UCMP locality 3296 (Clark and Vokes, 1936). UCR
hypotype 4865/13, middle Eocene strata, UCR locality 4865
(Givens and Kennedy, 1976). LACMIP hypotype 6540, Lla-
jas Formation, CSUN locality 704.

Mollusean Stage Range. “Domengine.”

Geographic Distribution. San Diego through Simi Valley,
California.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN locality 704.

Remarks. Two specimens of this taxon were found with
C. tabulatus at CSUN locality 704. C. n. sp. A is character-
ized by the presence of nodes on the spire. It is not named
at this time pending further taxonomic research by Jack
Mount, Rutgers University.

Clavilithes n. sp. B Squires, 1983a

Figure 8i

Clavilithes n. sp. Squires, 1983a:359, figs. 2h-i.

Type Material. UCLA hypotype 59194, Llajas Formation,
CSUN locality 445; UCLA hypotype 59195, Llajas For-
mation, CSUN locality 444.

Mollusean Stage Range. “Domengine.”

Geographic Distribution. Simi Valley, California.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 374, 444, 445.

Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks 31

Figure 8. Llajas Formation gastropods (continued). Unless otherwise indicated, views are apertural. a. Ficopsis cooperiana Stewart, 1927,
side view, UCLA hypotype 59277, CSUN loc. 374, x 2. b. Ficopsis remondii crescentensis Weaver and Palmer, 1922, LACMIP hypotype
6535, CSUN loc. 374, xl. c. Laevityphis ( Laevityphis ) antiquus (Gabb, 1864), LACMIP hypotype 6536, CSUN loc. 371, x2. d and e.
Ancistrolepisl carolineae n. sp. All parts UCLA holotype 5940 1 , CSUN loc. 374, x 1 .25. e, abapertural view. f. Molopophorus cretaceus (Gabb,
1864), abapertural view, LACMIP hypotype 6537, CSUN loc. 467, x7. g. Clavilithes tabulatus (Dickerson, 1913), LACMIP hypotype 6538,
CSUN loc. 371, xQ.5. h. Clavilithes n. sp. A Clark and Yokes, 1936, LACMIP hypotype 6539, CSUN loc. 704, x 1.25. i. Clavilithes n. sp. B

32 Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks

Remarks. This taxon occurs with C. tabulatus at CSUN
locality 374. C. n. sp. B differs from C. n. sp. A in the presence
of an overhanging carina and in the lack of nodes on the
spire. C. n. sp. B is not named at this time pending further
taxonomic research by Jack Mount, Rutgers University.

Subfamily Fusininae Swainson, 1 840
Genus Fusinus Rafinesque, 1815

Type Species. By monotypy, Murex co/us Linne, 1758.

Fusinus teglandae Hanna, 1927

Figure 8j

Fusinus teglandi Hanna, 1927:315, pi. 51, fig. 9.

Fusinus teglandae Hanna. Givens, 1974:85, pi. 10, figs.

6-7.

Primary Type Material. UCMP holotype 31 124, Ardath
Shale, UCMP locality 5062.

Molluscan Stage Range. “Domengine.”

Geographic Distribution. San Diego through Pine Moun-
tain, southern California.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN locality 374.

Remarks. Only three poorly preserved specimens were
found in the Llajas Formation. The upper spire is missing
in all three specimens.

Fusinus aff. F. ucalius Vokes, 1939

Figure 8 k

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN locality 371.

Remarks. Only a single specimen was found in the Llajas
Formation. It is unabraded and nearly complete, lacking only
the protoconch. This specimen agrees with the description
of Fusinus ucalius Vokes (1939:137) in having swollen col-
labral costae, strong spiral cords, and a long straight anterior
canal. It differs in that the collabral costae are not obsolete
or even subobsolete on the posterior parts of the spire whorls.
These collabral costae on the Llajas specimen extend from
suture to suture. According to Vokes ( 1 939), F. voetus Hanna
(1927:315, pi. 51, figs. 1, 2, 4) and F. meganosensis Clark
and Woodford (1927: 1 1 1, pi. 19, fig. 1 3) differ from F. ucali-
us in the same respect. The Llajas specimen differs from F.
voetus by having a less elongate aperture, more swollen col-
labral costae, more angulate spire whorls, and only eight
(rather than 15) collabral costae on the body whorl. The
Llajas specimen differs from F. meganosensis by having much
stronger collabral costae and spiral cords. In addition, on the

body whorl of F. meganosensis, the collabral costae tend to
become obsolete posteriorly.

Superfamily Volutacea
Family Olividae Latreille, 1825
Genus Pseudoliva Swainson, 1 840

Type Species. By original designation, Buccinum plumbea
Chemnitz, 1780? [=Buccinum crassa Gmelin, 1788?].

Pseudoliva lineata Gabb, 1864

Figure 81

Pseudoliva lineata Gabb, 1864:99, pi. 18, fig. 52. Stewart,
1927:400, pi. 28, fig. 14a. Vokes, 1939:139, pi. 18, fig. 23.

Primary Type Material. ANSP lectotype 4200, Tejon For-
mation s.l., northeast of Martinez, California.

Molluscan Stage Range. “Domengine.”

Geographic Distribution. Simi Valley through Martinez,
California.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 371, 373, 484.

Remarks. Only three specimens were found in the Llajas
Formation. All show the typical Pseudoliva medial spiral
groove. P. lineata is distinguished by a complete lack of nodes
on the shoulder of the whorl, as noted by Turner (1938:78).
As mentioned by Vokes (1939:139), Llajas Formation spec-
imens show a spire that is somewhat higher than that of the
lectotype figured by Stewart (1927, pi. 28, fig. 14a).

Genus Strepsidura Swainson, 1 840

Type Species. By original designation, Strepsidura costata
Swainson, 1840 [=Fusus ficu/nea Lamarck, 1822, =Murex
turgida Solander, 1766],

Strepsidura ficus (Gabb, 1864)

Figure 8m

Whitneya ficus Gabb, 1864:104, pi. 28, fig. 216. Dickerson,
1915:69, pi. 9, figs. 5a-d.

Strepsidura ficus (Gabb). Stewart, 1927:404-405, pi. 29, fig.
1 1 . Kleinpell and Weaver, 1 963: 1 93, pi. 27, figs. 1 -3. Giv-
ens, 1974:87, pi. 10, fig. 10.

Primary Type Material. ANSP lectotype 433 1 , Tejon For-
mation s.l.. Fort Tejon area, California.

Molluscan Stage Range. Uppermost “Capay” through
“Tejon.”

Geographic Distribution. Simi Valley through Fort Tejon
area, southern California.

Squires, 1983a, spire only, UCLA hypotype 59195, CSUN loc. 444, x0.5. j. Fusinus teglandae Hanna, 1927, abapertural view, LACMIP
hypotype 6540, CSUN loc. 374, xO.75. k. Fusinus aff. F. ucalius Vokes, 1939, LACMIP hypotype 6541, CSUN loc. 371, x 1.5. I. Pseudoliva
lineata Gabb, 1864, LACMIP hypotype 6542, CSUN loc. 371, x 1.5. m. Strepsidura ficus (Gabb, 1864), abapertural view, LACMIP hypotype
6543, CSUN loc. 452, xl.25. n. Ancilla ( Spirancilla ) gabbi Cossmann, 1899, LACMIP hypotype 6544, CSUN loc. 489, x4. o. Olivella
mathewsonii Gabb, 1864, LACMIP hypotype 6545, CSUN loc. 467, x2.5. p. Proximitral cretacea (Gabb, 1864), LACMIP hypotype 6546,
CSUN loc. 371, x 2.5.

Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks 33

Local Occurrence. Interfingering coastal alluvial-fan facies
and shallow-marine (transgressive) facies: CSUN localities
452, 545, 703.

Remarks. In some of the specimens, collabral costae are
fairly well developed on the earlier whorls.

The presence of S. ficus in the zone of interfingering be-
tween the coastal alluvial-fan facies and the shallow-marine
(transgressive) facies of the Llajas Formation extends the
molluscan stage range of this species into the uppermost
“Capay.” Previously, the lower range limit had been reported
as the “Dontengine Stage” (Givens, 1974).

Subfamily Olivinae Swainson, 1840

Genus Ancilla Lamarck, 1799

Type Species. By monotypy, Ancilla cinnamonea La-
marck, 1801.

Ancilla ( Spirancilla ) gabbi Cossmann, 1899
Figure 8n

Ancillaria elongata Gabb, 1864:100, pi. 18, fig. 54. Hanna,
1927:323, pi. 53, figs. 9-13. Stewart, 1927:411.

Not Ancillaria elongata Gray, 1847:357, pi. 1, fig. 5.

Ancilla gabbi Cossmann, 1899:60 [new name for Ancillaria
elongata Gabb, 1864, preoccupied]. Turner, 1938:72, pi.
18, fig. 6. Weaver, 1943:500, pi. 95, fig. 18.

Ancilla (Spirancilla) gabbi Cossmann. Vokes, 1939:131, pi.
18, figs. 6, 10.

Primary Type Material. Holotype lost, UCMP syntypes
12521 (two specimens) of Ancillaria elongata Gabb and An-
cilla gabbi Cossmann, Ardath Shale, near San Diego, perhaps
300 m east of the summit of Soledad Mountain on the east
side of the canyon in the bottom of Rose Creek.

Molluscan Stage Range. “Domengine.”

Geographic Distribution. San Diego, California through
northwestern Oregon.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN locality 371. Shallow-marine (regressive) facies: CSUN
locality 489.

Remarks. Only two specimens were found in the Llajas
Formation. Both agree with Gabb’s (1864) observations of
callused sutures.

Genus Olivella Swainson, 1831

Type Species. By subsequent designation (Dali, 1 909), Oli-
vella purpurata Swainson, 1 83 1 [=Oliva dama Mawe, 1 823],

Olivella mathewsonii Gabb, 1864

Figure 8o

Olivella mathewsonii Gabb, 1864:100, pi. 18, fig. 53. An-
derson and Hanna, 1925:80, pi. 8, fig. 19. Stewart, 1927:
410-411, pi. 29, fig. 13. Weaver, 1943:500-501, pi. 103,
fig. 7. Givens, 1974:87. Smith, 1975:469, table 1. Squires,
1977:table 1.

Primary Type Material. ANSP lectotype 4202, Tejon For-
mation s.L, Martinez, California.

Molluscan Stage Range. Lower “Martinez” through “Te-
jon.”

Geographic Distribution. Simi Valley, California through
northwestern Washington.

Local Occurrence. Interfingering coastal alluvial-fan facies
and shallow-marine (transgressive) facies: CSUN locality
469a. Shallow-marine (transgressive) facies: CSUN localities
467, 484, 498.

Remarks. Two specimens were found at locality 469a. At
each of the other localities, only one specimen was found.
The specimens agree with Gabb’s (1864) observations of
three columellar plicae at the anterior end of the shell. This
taxon may be the same as O. m. umpquaensis Turner, 1938:
72, pi. 18, figs. 13-14.

Family Mitridae Swainson, 1831
Subfamily Vexillinae Thiele, 1929
Genus Proximitra Finlay, 1927

Type Species. By original designation, Vexillum rutido-
lomum Suter, 1913.

Proximitra ? cretacea (Gabb, 1 864)

Figure 8p

Mitra cretacea Gabb, 1864:103, pi. 28, fig. 215. Stewart,
1927:406, pi. 27, figs. 9-10.

Uromitra (?) cretacea (Gabb). Vokes, 1939:134-135, pi. 18,
fig. 19.

Proximitra ? cretacea (Gabb). Givens, 1974:87. Squires, 1977:
table 1.

Primary Type Material. ANSP holotype 4302, Tejon For-
mation s.L, Martinez, California.

Molluscan Stage Range. “Domengine.”

Geographic Distribution. Simi Valley through central Cal-
ifornia.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN locality 371.

Remarks. This taxon was found at only one locality in the
Llajas Formation, but 46 specimens were collected.

Family Tudiclidae Finlay and Marwick, 1937
Genus Pseudoperissolax Clark, 1918

Type Species. By original designation. Busy coni blakei
Conrad, 1855.

Pseudoperissolax blakei praeblakei Vokes, 1939

Figure 9a

Not Busycoril blakei Conrad, 1855:1 1; 1857:332, pi. 2, fig.
13.

Perissolax blakei (Conrad). Gabb, 1864:92 (in part), pi. 21,
fig. 110.

Pseudoperissolax blakei (Conrad) (subsp.?). Stewart, 1927:
429-430, pi. 28, fig. 1.

Pseudoperissolax blakei praeblakei Vokes, 1939:145-146, pi.

34 Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks

19, figs. 14, 22. Givens, 1974:88, pi. 10, figs. 15-16. Smith,
197 5:pl. 2, fig. 16.

Primary Type Material. UCMP holotype 15799, UCMP
paratype 15800, Arroyo Hondo Formation, UCMP locality
1817.

Molluscan Stage Range. Lower “Martinez” through “Do-
mengine.”

Geographic Distribution. San Diego?, Simi Valley through
central California.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN locality 374.

Remarks. Only two specimens were found in the Llajas
Formation. They are mostly internal molds with shell present
only in the spire area. The long anterior canal is missing in
the figured specimen (Fig. 9a).

A discussion of the taxonomic reasons for the use of the
family Tudiclidae in place of the normally used family Vasi-
dae is given in Zinsmeister (1983b).

Family Harpidae Bronn, 1 849

Genus Eocithara P. Fischer, 1883

Type Species. By monotypy, Harpa mutica Lamarck, 1 803.

Eocithara mutica californiensis (Vokes, 1937)

Figure 9b

Harpa ( Eocithara ) mutica n. sp. Clark and Vokes, 1936:pl.
2, fig. 5.

Harpa (Eocithara) mutica californiensis Vokes, 1937:1 1, pi.
2, figs. 2, 4, 6, 8.

Eocithara mutica californiensis (Vokes). Rehder, 1973:225,
pi. 193.

Primary Type Material. UCMP holotype 30438, UCMP
paratypes 30439-30449, Llajas Formation, UCMP locality
3296.

Molluscan Stage Range. “Domengine.”

Geographic Distribution. Simi Valley, California.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 373, 374.

Remarks. Four specimens were found in the Llajas For-
mation. Preservation is generally excellent with the thin col-
labral ridges present.

Family Volutidae Rafinesque, 1815
Subfamily Volutinae Rafinesque, 1815
Genus Cryptochorda Morch, 1858

Type Species. By monotypy, Buccinum stromboides
Herrmannsen.

Subgenus Cryptochorda s.s.

Cryptochorda (Cryptochorda) calif ornica
(Cooper, 1894)

Figure 9c

Ancilla ( Oliverato ) californica Cooper, 1894:43, pi. 1, figs.
6-11. Dickerson, 1913:264; 1914:115, pi. 12, figs. 4a-b.

Oliverato californica Cooper. Dickerson, 1913:286-287, pi.
1 3, figs. 4a-b.

Caricella stormsiana Dickerson, 1913:287, pi. 13, figs.
3a-b.

Cryptochorda californica (Cooper). Clark, 1929:pl. 4, figs. 6,
16; pi. 9, figs. 5-6. Clark and Vokes, 1936:874, pi. 1, fig.
5. Turner, 1938:72, pi. 18, figs. 11, 15. Vokes, 1939:139-
140. Weaver, 1943:499, pi. 95, figs. 19, 23.

Primary Type Material. CAS syntypes 8a-d (four speci-
mens), Capay Formation, Marysville Buttes, Sutter County,
California.

Molluscan Stage Range. “Capay” through “Domengine.”
Geographic Distribution. Simi Valley, California through
southwestern Oregon.

Local Occurrence. Interfingering coastal alluvial-fan facies
and shallow-marine (transgressive) facies: CSUN locality 703.

Remarks. Only two specimens were found in the Llajas
Formation.

According to Clark (1921 :table 1 ; 1 929), C. (C.) californica
[=Ancilla ( Oliverato ) californica ] occurs in “Meganos”-age
strata. Such a report, however, cannot be substantiated at
this time because of his lack of locality information and
because his rock unit names are not clearly defined.

Genus Lyria Gray, 1847

Type Species. By original designation, Voluta nucleus La-
marck, 1811.

Lyria andersoni Waring, 1917

Figure 9d

Cance/laria irelaniana Cooper, 1 894:42, pi. 1, fig. 5. Arnold,
1910:52, pi. 4, fig. 22. [Misidentification.]

Lyria andersoni Waring, 1917:97, pi. 15, fig. 12.

Lyria andersoni Waring. Clark, 1929:pl. 9, figs. 7-8. Clark
and Vokes, 1936:876, pi. 1, fig. 17. Turner, 1938:73, pi.
1 8, fig. 5. Vokes, 1 939: 1 36, pi. 1 8, figs. 22, 24. Hanna and
Hertlein, 1943:170, fig. 62-21.

Primary Type Material. SU holotype 195, SU paratype
196, Llajas Formation, SU locality 2696.

Molluscan Stage Range. “Domengine.”

Geographic Distribution. Simi Valley, California through
southwestern Oregon.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 371, 372, 439, 455, 466, 484, 498, 499, 507.

Remarks. This taxon helps to characterize the lower and
middle parts of the shallow-marine (transgressive) facies. It
is most common in the middle part. Preservation is generally
good to excellent.

Subfamily Fulgorariinae
Pilsbry and Olsson, 1954

Genus Lyrischapa Aldrich, 1911

Type Species. By monotypy, Lyrischapa harrisi Aldrich,
1911.

Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks 35

Lyrischapa lajollaensis (Hanna, 1927)

Figure 9e

Pejonia lajollaensis Hanna, 1927:320, pi. 52, figs. 1-2.
Volutospira ( Pejonia ) lajollaensis (Hanna). Clark, 1929, pi.

9, figs. 11-12.

Volutocristata lajollaensis (Hanna). Gardner and Bowles,
1934:246, fig. 13. Givens, 1974:88.

Lyrischapa lajollaensis (Hanna). Givens, 1979:124-126, pi.
3, figs. 1-2; pi. 4, figs. 1-3. Givens and Kennedy, 1979:
table 1.

Primary Type Material. Holotype lost, UCMP neotype
14634, Ardath Shale, UCMP locality 5062.

Molluscan Stage Range. “Domengine.”

Geographic Distribution. San Diego through Pine Moun-
tain area, southern California.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 373, 374, 444, 445, 510, 543, 704.

Remarks. This taxon is most common in the “Stewart bed”
at locality 374, where some rather large specimens (47 mm
width) occur.

Hypotype specimens figured by Givens (1979) are from
the Llajas Formation.

Family Cancellariidae
Forbes and Hanley, 1853

Genus Bonellitia Jousseaume, 1887

Type Species. By original designation, Cance/laria bonellii
Bellardi, 1872?

Subgenus Admetula Cossmann, 1889

Type Species. By original designation, Cancellaria evulsa
Solander, 1776.

Bonellitia ( Admetula ) paucivaricata
(Gabb, 1864)

Figure 9f

Tritonium paucivaricatum Gabb, 1864:95, pi. 28, figs. 209,
209a.

Cancellaria stantoni Dickerson, 1913:282, pi. 12, figs. 2a-b.
Ad mete (Bonellitia) stantoni (Dickerson). Weaver and Palm-
er, 1922:40-42, pi. 11, figs. 1, 5.

Cancellaria paucivaricata (Gabb). Anderson and Hanna,
1925:81, pi. 8, figs. 3-4.

Bonellitia (Admetula) paucivaricata (Gabb). Stewart, 1927:
413, pi. 29, fig. 5. Turner, 1938:71, pi. 15, figs. 12-13.
Weaver, 1943:508, pi. 96, figs. 10, 12, 13; pi. 103, figs. 9,

10, 17. Givens, 1974:89. Squires, 1977:table 1.

Primary Type Material. ANSP lectotype 4 1 94, Tejon For-
mation, Tejon Pass area, California.

Molluscan Stage Range. “Meganos”?, “Capay” through
“Tejon.”

Geographic Distribution. Simi Valley, California through
southwestern Washington.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN locality 371.

Remarks. Only seven specimens were found in the Llajas
Formation. According to Clark (1921:table 1; 1926:1 15), B.
(A.) paucivaricata [= Cancellaria stantoni and Admeta stan-
toni] occurs in “Meganos”-age strata. Such a report, however,
cannot be substantiated at this time because of his lack of
locality information and because his rock unit names are not
clearly defined. Clark and Woodford (1927), however, re-
ported a questionable occurrence of B. (A.) paucivaricata
[=Admete (Bonellitia) cf. stantoni ] from localities of late Pa-
leocene/early Eocene age strata.

Superfamily Conacea
Family Turridae Swainson, 1840
Subfamily Turriculinae Powell, 1942
Genus Pleurofusia de Gregorio, 1890

Type Species. By original designation, Pleurotoma (Pleu-
rofusia) longirostropis de Gregorio, 1890.

Pleurofusia fresnoensis (Arnold, 1910)

Figure 9g

Pleurotoma fresnoensis Arnold, 1910:53, pi. 4, fig. 23.
Surcu/a clarki Dickerson, 1913:278, pi. 1 1, fig. 3.
Pleurofusia fresnoensis (Arnold). Vokes, 1939:1 17-1 18, pi.

17, figs. 15-16. Givens, 1974:90, pi. 11, fig. 9. Givens and

Kennedy, 1979:95, tables 1, 3.

Primary Type Material. USNM holotype 165631, Do-
mengine Formation, USGS locality 4619.

Molluscan Stage Range. “Capay” through “Transition.”

Geographic Distribution. San Diego through central Cal-
ifornia.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 371, 538, 539.

Remarks. Specimens are most common at locality 371.

According to Clark (1921 :table 1 ; 1 926: 1 1 6), P. fresnoensis
[=Turris (Surcu/a) clarki and Surcula clarki] occurs in “Me-
ganos”-age strata. Such a report, however, cannot be sub-
stantiated at this time because of his lack of locality infor-
mation and because his rock unit names are not clearly defined.

Smith (1975:table 1) reported Pleurofusia sp. aff. P. fres-
noensis from “Martinez”-age strata of central California.

Genus Fusiturricula Woodring, 1928

Type Species. By original designation, Turris (Surcula)
fusinella.

Subgenus Crenaturricula Vokes, 1939

Type Species. By original designation, Surcula crenato-
spira Cooper, 1894.

Fusiturricula ( Crenaturricula ) crenatospira
(Cooper, 1894)

Figure 9h

Surcula crenatospira Cooper, 1894:39, pi. 1, figs. 2-4. Dick-
erson, 1913:278, pi. 11, fig. 4.

36 Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks

Fusiturricula ( Crenaturricula ) crenatospira (Cooper). Vokes,
1939:1 14-115, pi. 17, figs. 4-5.

Primary Type Material. CAS syntypes 9a-b (two speci-
mens), Capay Formation, UCMP locality 1853.

Molluscan Stage Range. “Capay” through “Domengine.”
Geographic Distribution. Simi Valley through Marysville
Buttes, California.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN locality 374.

Remarks. Only two specimens were found, and these are
from the “Stewart bed.” According to Vokes (1939:15) this
taxon is confined to and is diagnostic of the “Capay Stage.”
The presence of it in the “Stewart bed” extends its provincial
range into the “Domengine Stage.”

According to Clark (1921:table 1; 1929), F. (C.) crenato-
spira [=Turris ( Surcula ) crenatospira and Turricula crenato-
spira] occurs in “Meganos”-age strata. Such a report, how-
ever, cannot be substantiated at this time because of his lack
of locality information and because his rock unit names are
not clearly defined.

Fusiturricula ( Crenaturricula ) crenatospira
domenginica Vokes, 1939

Figure 9i

Fusiturricula { Crenaturricula ) crenatospira domenginica
Vokes, 1939:1 15-1 16, pi. 17, figs. 6-7.

Primary Type Material. UCMP holotype 15768, Llajas
Formation, UCMP locality 7002; UCMP paratype 15769,
Llajas Formation, UCMP locality 3304.

Molluscan Stage Range. “Domengine.”

Geographic Distribution. Simi Valley through central Cal-
ifornia.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 371, 374, 444, 445, 498, 548.

Remarks. At locality 374, this taxon occurs along with F.
(C.) crenatospira, which is characterized by the presence of
collabral sculpturing below the nodes on the spire and body
whorls. On F. ( C .) crenatospira domenginica, the collabral
sculpturing is obsolete. Contrary to Vokes’ (1939:1 15-116)
comments, the two taxa have approximately the same num-
ber of nodes on the body whorl.

Genus Surculites Conrad, 1865

Type Species. By monotypy, Surcula (Surculites) annosus
Conrad, 1865.

Surculites mathewsonii (Gabb, 1864)

Figure 9j

Fusus mathewsonii Gabb, 1 864:83, pi. 1 8, fig. 33. Dickerson,
1 9 1 4:pl. 16, fig. 2.

Bela clathra Gabb, 1869:152, pi. 26, fig. 31.

IPieurotoma decipiens Cooper, 1894:40, pi. 2, fig. 32.
Potamidesl davisiana Cooper, 1894:44, pi. 1, fig. 13.
Surcula davisiana (Cooper). Dickerson, 1 9 1 3:279, pi. 12, figs.
6a-b.

Surcula (?) sp. Waring, 1917 :pl. 15, fig. 16.

Surcula decipiens (Cooper). Hanna, 1927:324, pi. 54, figs.

6, 8.

Surculites mathewsonii (Gabb). Stewart, 1927:420-421, pi.

26, figs. 12-14. Clark, 1929:pl. 9, figs. 3-4. Turner, 1938:
69-70, pi. 17, figs. 6, 10. Vokes, 1939:123, pi. 17, figs. 8,
19. Weaver, 1943:526, pi. 97, figs. 24, 29; pi. 98, figs. 1,
5; 1953:29. Givens, 1974:90, pi. 11, figs. 5, 7. Zinsmeister,
1974:164, pi. 17, fig. 6; 1983a:table 1. Squires, 1977:table
1. Givens and Kennedy, 1979:87, tables 1, 3.

“ Surculites ” mathewsonii (Gabb). Smith, 1975:pl. 2, fig. 15.

Primary Type Material. ANSP lectotype 4180, Tejon For-
mation s.l., near Martinez, California.

Molluscan Stage Range. Lower “Martinez” through
“Transition.”

Geographic Distribution, San Diego, California through
southwestern Oregon.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 371, 372, 374, 444, 445, 447, 479, 498.

Remarks. Specimens are fairly common at locality 37 1 but
are more common and larger at locality 374 of the “Stewart
bed.” At locality 374, the specimens are internal molds. Spec-
imens at locality 371 are strongly angulated, which is in
keeping with one of the main characteristics of this species
as discussed by Vokes (1939:123).

Genus Domenginella Vokes, 1939

Type Species. By original designation, Tunis claytonensis
Gabb, 1864.

Domenginella claytonensis (Gabb, 1864)

Figure 9k

Turris claytonensis Gabb, 1864:92, pi. 18, fig. 46.

Surcula gesteri Dickerson, 1916:499, pi. 42, fig. 4.
Scobinella claytonensis ( Gabb). Stewart, 1927:417-418, pi.

27, figs. 11-12.

Domenginella claytonensis (Gabb). Vokes, 1939:122-123, pi.
17, figs. 18, 20.

Primary Type Material. ANSP lectotype 4 1 90, Tejon For-
mation s.l., near Martinez, California.

Molluscan Stage Range. Uppermost “Capay” through
“Domengine.”

Geographic Distribution. Simi Valley through central Cal-
ifornia.

Local Occurrence. Interfingering coastal alluvial-fan facies
and shallow-marine (transgressive) facies: CSUN locality
469a. Shallow-marine (transgressive) facies: CSUN localities
371, 372, 455, 466, 484, 486, 498, 517, 538, 539, 548.

Remarks. This taxon is most common at locality 371,
where most of the specimens are well-preserved juveniles.

The presence of D. claytonensis in the zone of interfingering
between the coastal alluvial-fan facies and the shallow-ma-
rine (transgressive) facies of the Llajas Formation extends
the molluscan stage range of this taxon into the uppermost

Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks 37

Figure 9. Llajas Formation gastropods (continued). Unless otherwise indicated, views are apertural. a. Pseudoperissolax blakei praeblakei
Vokes, 1939, internal mold, side view, LACMIP hypotype 6547, CSUN loc. 374, xl.5. b. Eocithara mutica californiensis (Vokes, 1937),
abapertural view, LACMIP hypotype 6548, CSUN loc. 373, xl.5. c. Cryptochorda (Cryptochorda) californica (Cooper, 1894), abapertural
view, LACMIP hypotype 6549, CSUN loc. 703, x 1.25. d. Lyria andersoni Waring, 1917, LACMIP hypotype 6550, CSUN loc. 498, x2. e.
Lyrischapa lajollaensis (Hanna, 1927), LACMIP hypotype 6551, CSUN loc. 373, xl. f. Bonellita (Admetula) paucivariata (Gabb, 1864),
LACMIP hypotype 6552, CSUN loc. 374, x4.5. g. Pleurofusia fresnoensis (Arnold, 1910), LACMIP hypotype 6553, CSUN loc. 371, x3. h.
Fusiturricula (Crenaturricula) crenatospira (Cooper, 1894), abapertural view, UCLA hypotype 59278, CSUN loc. 374, x 1.5. i. Fusiturricula
( Crenaturricula ) crenatospira domenginica Vokes, 1939, abapertural view, LACMIP hypotype 6554, CSUN loc. 371, xl. j. Surculites ma-

38 Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks

“Capay.” Previously, the lower range limit had been reported
as the “Domengine Stage” (Vokes, 1939).

Genus Exilia Conrad, 1860

Type Species. By monotypy, Exilia pergracilis Conrad,
1860.

Exilia llajasensis Bentson, 1940

Figure 91

Exilia llajasensis Bentson, 1940:212, pi. 1, fig. 28; pi. 2, fig.
16.

Primary Type Material. UCMP holotype 12144, Llajas
Formation, UCMP locality A-3042; UCMP paratype 12140,
Llajas Formation, UCMP locality 3296.

Molluscan Stage Range. “Domengine.”

Geographic Distribution. Simi Valley, California.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 486, 539.

Remarks. Only two specimens were found in the Llajas
Formation. In the primary type specimens, the anterior canal
is missing. As can be seen in Figure 91, the long anterior canal
is covered by 10 spiral ribs.

Family Conidae Rafinesque, 1815
Genus Conus Linne, 1758

Type Species. By subsequent designation (Children, 1 823),
Conus marmoreus Linne, 1758.

Conus caleocius Yokes, 1939

Figure 9m

Conus caleocius Vokes, 1939:127-129, pi. 18, figs. 1, 7.

Primary Type Material. UCMP holotype 15785, Llajas
Formation, UCMP locality 3310.

Molluscan Stage Range. “Domengine.”

Geographic Distribution. Simi Valley through central Cal-
ifornia.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 371, 498.

Remarks. Four specimens were found. Three of these are
from locality 371.

Conus hornii umpquaensis Turner, 1938

Figure 9n

Conus hornii umpquaensis Turner, 1938:69, pi. 15, figs. 1-
2. Vokes, 1939:127, pi. 18, figs. 2-3. Stewart, 1946:pl. 1 1,

fig. 6. Weaver, 1943:510-51 1, pi. 96, fig. 18. Givens and

Kennedy, 1979:87, tables 1, 3.

Primary Type Material. UCMP holotype 33656, “upper
Umpqua” Formation, UO locality 144.

Molluscan Stage Range. “Domengine” through “Transi-
tion.”

Geographic Distribution. San Diego, California through
southwestern Oregon.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 37 1 , 373, 374, 449, 455, 484, 486, 49 1 , 498,
538, 539.

Remarks. This taxon characterizes the lower half of the
shallow-marine (transgressive) facies. It is especially abun-
dant at locality 371, where it forms a growth series.

Family Terebridae H. and A. Adams, 1854
Genus Terebra Bruguiere, 1789

Type Species. By subsequent designation (Lamarck, 1 799),
Buccinum subulata Linne, 1758.

Terebra californica Gabb, 1869

Figure 9o

Terebra californica Gabb, 1869:162, pi. 27, fig. 41.

Terebra californica Gabb. Anderson and Hanna, 1925:82,

pi. 8, fig. 18. Stewart, 1927:424, pi. 26, fig. 5. Vokes, 1939:

113, pi. 16, fig. 38. Givens, 1974:92.

Primary Type Material. ANSP holotype 4209, Tejon For-
mation s.l., Martinez, California.

Molluscan Stage Range. “Domengine” through “Tejon.”

Geographic Distribution. Simi Valley through central Cal-
ifornia.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN locality 467.

Remarks. Only two specimens were found. They are very
small, with heights of 3 mm.

Subclass Euthyneura
Order Cephalaspidea

Superfamily Cylichnacea A. Adams, 1850
Family Cylichnidae A. Adams, 1850
Genus Cy/ichnina Monterosato, 1884

Type Species. By original designation. Bulla umbilicata
Montagu, 1803.

thewsonii (Gabb, 1864), abapertural view, LACMIP hypotype 6555, CSUN loc. 371, x2.5. k. Domenginella claytonensis (Gabb, 1864), side
view, LACMIP hypotype 6556, CSUN loc. 371, x2.5. 1- Exilia llajasensis Bentson, 1940, LACMIP hypotype 6557, CSUN loc. 539, x2.5.
m. Conus caleocius V okes, 1939, LACMIP hypotype 6558, CSUN loc. 498, x 2. n. Conus hornii umpquaensis Turner, 1938, LACMIP hypotype
6559, CSUN loc. 371, x2.5. o. Terebra californica Gabb, 1869, LACMIP hypotype 6560, CSUN loc. 467, x 8. p. Cylichnina tantilla (Anderson
and Hanna, 1925), LACMIP hypotype 6561, CSUN loc. 371, x2.5. q. Scaphander (Mirascapha) costatus (Gabb, 1864), abapertural view,
LACMIP hypotype 6562, CSUN loc. 493, x 1.5. r. Megistostoma gabbianum (Stoliczka, 1868), internal mold, abapertural view, LACMIP
hypotype 6553, CSUN loc. 458, x 1.5.

Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks 39

Cylichnina tantilla
(Anderson and Hanna, 1925)

Figure 9p

Cylichnella tantilla Anderson and Hanna, 1925:140, pi. 7,
figs. 4, 8-9.

Cylichnina tantilla (Anderson and Hanna). Stewart, 1927:
439-441, pi. 27, figs. 2-4. Turner, 1938:67-68, pi. 20, figs.
9-10. Vokes, 1939:110, pi. 16, figs. 28, 33, 39. Weaver,
1943:548-549, pi. 100, figs. 10-12, 1 4-1 5. Stewart, 1946:
pi. 11, fig. 11. Givens, 1974:93. Squires, 1977:table 1;
1983b, fig. 9a.

Primary Type Material. CAS holotype 958, CAS paratypes
959 and 960, Tejon Formation, CAS locality 711.

Molluscan Stage Range. “Domengine” through “Tejon.”
Geographic Distribution. Simi Valley, California through
western Washington.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 371, 372, 373, 374, 455, 460, 466, 467,
469e, 469f, 484, 486, 491, 492, 498, 499, 507, 508, 517,
537, 538, 540, 543. Shallow-marine (regressive) facies: CSUN
locality 475.

Remarks. This easily recognized taxon is especially abun-
dant at localities 371, 373, and 455. At these localities it
occurs as growth series.

Genus Scaphander Montfort, 1810

Type Species. By original designation. Bulla lignaria Linne,
1767.

Subgenus Mirascapha Stewart, 1927

Type Species. By original designation, Cylichna costata
Gabb, 1864.

Scaphander ( Mirascapha ) costatus
(Gabb, 1864)

Figure 9q

Cylichna costata Gabb, 1864:143-144, pi. 21, fig. 107. Ar-
nold, 1 907 :pl. 39, fig. 10. Zinsmeister, 1974:170-171, pi.
12, figs. 20-22.

Scaphander costata (Gabb). Hanna, 1927:329, pi. 57, figs. 2,
3, 5.

Scaphander (Mirascapha) costatus (Gabb). Stewart, 1927:
437-438, pi. 27, fig. 5. Turner, 1938:67, pi. 17, fig. 16.
Vokes, 1939:109, pi. 16, figs. 29, 35. Givens, 1974:93-94.
Scaphander costatus (Gabb). Weaver, 1943:545, pi. 100, fig.
2; pi. 103, fig. 21. Weaver, 1953:29. Givens and Kennedy,
1979:88, table 3.

Primary Type Material. ANSP lectotype 4338, Tejon For-
mation s.l., near Martinez, California.

Molluscan Stage Range. “Martinez” through “Transi-
tion.”

Geographic Distribution. San Diego, California through
western Washington.

Local Occurrence. Shallow-marine (transgressive) facies:

CSUN locality 493. Shallow-marine (regressive) facies: CSUN
locality 475.

Remarks. Only a few specimens were found in the Llajas
Formation.

Family Philinidae Gray, 1850

Genus Megistostoma Gabb, 1864

Type Species. By monotypy, Megistostoma striata Gabb,
1864.

Megistostoma gabbianum (Stoliczka, 1868)

Figure 9r

Megistostoma striata Gabb, 1864:144, pi. 21, figs. 108a-b.
Not Bullaea striata Deshayes, 1824:37, pi. 5, figs. 1-3.
Bullaea gabbiana Stoliczka, 1868:434 [new name for Meg-
istostoma striata Gabb, 1864, preoccupied].

Philine (Megistostoma) gabbiC ossmann, 1895:127 [new name
for Megistostoma striata Gabb, 1864, preoccupied],
Megistostoma gabbianum (Stoliczka). Stewart, 1 927:44 1-442,
pi. 26, figs. 1-2. Vokes, 1939:112, pi. 17, figs. 1-3. Givens
and Kennedy, 1979:88, table 3.

Megistostoma caminoensis Hanna, 1927:330, pi. 57, figs. 9-
10. Turner, 1938:68, pi. 20, fig. 15. Weaver, 1943:541, pi.
99, fig. 24.

Primary Type Material. ANSP holotype 4216 of Megis-
tostoma striata Gabb, Bullaea gabbiana Stoliczka, and Phi-
line (Megistostoma) gabbi Cossmann, Tejon Formation s.l.,
near Martinez, California.

Molluscan Stage Range. “Domengine” through “Transi-
tion.”

Geographic Distribution. San Diego, California through
southwestern Oregon.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 374, 458. Shallow-marine (regressive) facies:
CSUN locality 462.

Remarks. Only a few specimens were found in the Llajas
Formation, and they occur as internal molds.

Class Bivalvia
Subclass Palaeotaxodonta
Order Nuculoida
Superfamily Nuculacea
Family Nuculidae Gray, 1824
Genus Acila H. & A. Adams, 1858

Type Species. By subsequent designation (Stoliczka, 1871),
Nucula divaricata Hinds, 1843.

Subgenus Trunc acila Schenck, in
Grant and Gale, 1931

Type Species. By original designation Nucula castrensis
Hinds, 1843.

40 Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks

Acila ( Truncacila ) decisa (Conrad, 1855)

Figure 10a

Nucula decisa Conrad, 1855:1 1-12; 1 8 5 7:pl. 3, fig. 19.

Acila gabbiana Dickerson, 1916:481, pi. 36, fig. 1. Anderson
and Hanna, 1925:176, pi. 9, fig. 12.

Nucula {Acila) stil/waterensis Weaver and Palmer, 1922:6,
pi. 8, fig. 8.

Acila lajollaensis Hanna, 1927:270, pi. 25, figs. 1, 3, 5, 7-8,
12, 15.

Acila {Truncacila) decisa (Conrad). Schenck, 1 936:53-56, pi.
3, figs. 1-9, 11-15; pi. 4, figs. 1-2; text figure 7 (22, 23,
25). Turner, 1938:41-42, pi. 5, figs. 2-3. Vokes, 1939:41,
pi. 1, figs. 7-8. Weaver, 1943:22-23, pi. 6, figs. 1, 4, 8; pi.
7, figs. 8-9. Givens, 1974:38, pi. 1, fig. 1. Zinsmeister,
1974:67-68, pi. 6, fig. 3. Squires, 1977:table 1. Moore,
1983:A10, pi. 1, fig. 14. Zinsmeister, 1983a:table 1.

Primary Type Material. UCMP neotype 31132, Ardath
Shale, UCMP locality 5062.

Molluscan Stage Range. “Martinez” through upper Eocene
{Turritella schencki delaguerrae Zone of Kleinpell and Weav-
er, 1963).

Geographic Distribution. San Diego, California through
Kamchatka, Alaska.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN locality 467.

Remarks. Specimens are abundant at locality 467. Pres-
ervation is excellent. The specimens are disarticulated, but
the valves constitute a growth series.

Superfamily Nuculanacea
Family Nuculanidae H. and A. Adams, 1858
Genus Nuculana Link, 1 807

Type Species. By original designation. Area rostrata
Chemnitz, 1784 [=Arca pernula Muller, 1776],

Subgenus Saccella Woodring, 1925

Type Species. By original designation. Area fragilis Chem-
nitz, 1784.

Nuculana ( Saccella ) gabbii (Gabb, 1864)

Figure 10b

Not Leda protextal Gabb, 1860:303, pi. 48, fig. 23.

Leda protextal Gabb, 1864:199 (in part), pi. 26, fig. 185.
Nuculana gabbii Conrad, 1866:3 [ nomen nudum],

Leda gabbii Conrad. Gabb, 1869:197 [new name for Leda
protextal Gabb, 1864, misidentified]. Stanton, 1896:1041,
pi. 64, fig. 8. Waring, 1917:76, pi. 13, fig. 6. Dickerson,
191 5:pl. 1, fig. 1; 1916, pi. 36, fig. 3. Clark and Woodford,
1927:85-86, pi. 14, fig. 2. Clark, 1929:pl. 3, fig. 12.

Leda vaderensis Dickerson, 1915:52, pi. 1, fig. 3.

Leda vogdesi Anderson and Hanna, 1925: 1 77-1 79, pi. 2, figs.
8-9.

Saccella gabbii (Gabb). Stewart, 1930:55-58, pi. 7, fig. 3 [not
pi. 10, fig. 4. =N. (S'.) alaeformis fide Zinsmeister, 1974],
Weaver, 1953:28.

Nuculana {Saccella) gabbii (Gabb). Vokes, 1939:41-42.
Kleinpell and Weaver, 1963:195, pi. 28, fig. 1. Givens,
1974:39, pi. 1, fig. 3. Moore, 1983:A16, pi. 2, figs. 7-8.
Nuculana (Calorhadia) gabbii { Gabb). Zinsmeister, 1974:69-
70, pi. 6, fig. 7; 1983a, pi. 1, fig. 3.

Primary Type Material. ANSP lectotype 4476 of Leda
gabbii Gabb, 1869, Tejon Formation s.l., Martinez, Califor-
nia.

Molluscan Stage Range. “Martinez” through upper Eocene
( T urritella schencki delaguerrae Zone of Kleinpell and Weav-
er, 1963).

Geographic Distribution. Simi Valley, California through
Kamchatka, Alaska.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN locality 467.

Remarks. Only five disarticulated valves were found in the
Llajas Formation. There is considerable variation in the
amount of elongation of the rostrum in these specimens, with
the illustrated specimen (Fig. 10b) showing the most.

According to Zinsmeister (1974), Stewart ( 1930) somehow
confused N. {S.) gabbii with N. {S.) alaeformis and designated
N. (S.) alaeformis (pi. 10, fig. 4) as the lectotype (ANSP 4476)
of N. {S.) gabbii.

Subclass Pteriomorpha
Order Arcoida
Superfamily Limnopsacea
Family Glycymerididae Newton, 1922
Subfamily Glycymeridinae Newton, 1922
Genus Glycymeris da Costa, 1778

Type Species. By tautonymy. Area orbicularis da Costa,
1778 [=Arca glycymeris Linne, 1758],

Subgenus Glycymeris s.s.

Glycymeris ( Glycymeris ) rosecanyonensis
Hanna, 1927
Figure 10c

Glycymeris rosecanyonensis Hanna, 1927:273-274, pi. 27,
figs. 4-5, 9, 1 1. Clark, 1929:pl. 6, fig. 8. Givens and Ken-
nedy, 1979:tables 1, 3.

Glycymeris { Glycymeris ) rosecanyonensis Hanna. Givens,
1974:42. Moore, 1983:A49-A50, pi. 10, figs. 12-13.
Glycymeris { Glycymerisa ) rosecanyonensis Hanna. Squires,
1977:table 1.

Primary Type Material. UCMP holotype 30989, Ardath
Shale, UCMP locality 3990.

Molluscan Stage Range. “Domengine” through “Transi-
tion.”

Geographic Distribution. San Diego through lower Piru
Creek, southern California.

Local Occurrence. Shallow-marine (transgressive) facies:

Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks 41

CSUN localities 373, 467, 47 Id, 520. Shallow-marine (re-
gressive) facies: CSUN localities 476, 489, 512a, 512b.

Remarks. Only a few specimens of this small-sized bivalve
were found in the Llajas Formation.

Subgenus Glycymerita
Finlay and Marwick, 1937

Type Species. By original designation, Glycymeris concava
Marshall, 1917.

Glycymeris ( Glycymerita ) sagittata
(Gabb,’ 1864)

Figure lOd

Axinaea ( Limopsis ?) sagittata Gabb, 1864:197-198, pi. 31,
figs. 267, 267a.

Glycimeris hannibali Dickerson, 1916:483, pi. 36, figs.
8a-b.

Glycymeris sagittatus (Gabb). Dickerson, 1 9 1 6:pl. 36, figs.
5a-b.

Glycymeris sagittata (Gabb). Anderson and Hanna, 1925:
181-182, pi. 1, fig. 6. Stewart, 1930:71-73, pi. 12, fig. 10;
1946:pl. 12, fig. 3. Vokes, 1939:45-46, pi. 1, figs. 18-20.
Weaver, 1943:54-55, pi. 9, figs. 17-18; pi. 11, fig. 15.
Kleinpell and Weaver, 1963:196-197, pi. 28, fig. 10; pi.
29, figs. 1-2.

Glycimeris sagittatus (Gabb). Turner, 1938:43-44, pi. 6, figs.

1-3.

Glycymeris ( Glycymerita ) sagittata (Gabb). Givens, 1974:
42-43. Squires, 1977:table 1. Moore, 1 983:A54-A55, pi.
12, fig. 17.

Primary Type Material. ANSP lectotype 4422, Tejon For-
mation, near Fort Tejon (N Vi of section 29, T 10 N, R 19
W, Kern County, California).

Molluscan Stage Range. “Capay” through “Tejon,” Oli-
gocene?.

Geographic Distribution. Simi Valley, California through
southwestern Washington.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 371, 373, 374, 444, 500, 704.

Remarks. Specimens are uncommon in the Llajas For-
mation, and leaching has obscured some of the exterior sculp-
ture.

Order Mytiloida
Superfamily Mytilacea
Family Mytilidae Rafinesque, 1815
Genus Brachidontes Swainson, 1840

Type Species. By monotypy, Modiola sulcata Lamarck,
1819 (not 1 805) [=Mytilus citrinus Roding, 1 798, =Arca mo-
diolus Linne, 1767],

Subgenus Brachidontes s.s.

Brachidontes ( Brachidontes ) cowlitzensis
(Weaver and Palmer, 1922)

Figure lOe

Modiola ornata Gabb, 1864:184-185, pi. 24, fig. 166.

Not Mytilus ornatus Orbigny, 1843:283, pi. 342, figs. 10-12.
Modiolus ornatus Gabb. Arnold, 1907:pl. 38, fig. 4.
Brachydontes ornatus (Gabb). Anderson and Hanna, 1925:
188, pi. 3, fig. 4.

Modiolus ( Brachydontes ) ornatus Gabb. Clark and Wood-
ford, 1927:89, pi. 14, fig. 10. Clark, 1929:pl. 3, fig. 6.
Modiolus ( Brachydontes ) cowlitzensis Weaver and Palmer,
1922:16-17, pi. 9, fig. 19 [new name for Modiola ornata
Gabb, 1864, preoccupied],

Brachidontes cowlitzensis ? (Weaver and Palmer). Stewart,
1930:100-103, pi. 8, fig. 12.

Brachidontes cowlitzensis I'M caver and Palmer). Turner, 1938:
45-46, pi. 6, figs. 7-8. Kleinpell and Weaver, 1963:197,
pi. 29, fig. 3. Wolfe, 1977:3. Givens and Kennedy, 1979:
table 2.

Volsella (Brachidontes) cowlitzensis (Weaver and Palmer).

Weaver, 1943:1 13-1 14, pi. 26, fig. 4.

Brachidontes ( Brachidontes ) cowlitzensis (Weaver and Palm-
er). Givens, 1974:43. Squires, 1977, table 1. Moore, 1983:
A66-A67, pi. 17, fig. 1.

Primary Type Material. ANSP lectotype 4450 of Modiola
ornata Gabb, Domengine? Formation, Martinez, California.
CAS holotype 7406 of Modiolus (Brachydontes) ornatus
Weaver and Palmer, Cowlitz Formation, UW locality 329.

Molluscan Stage Range. “Meganos” through lower Oli-
gocene (Turritella variata lorenzana Zone of Kleinpell and
Weaver, 1963).

Geographic Distribution. San Diego, California through
Gulf of Alaska.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 371, 374, 455, 466, 471d, 492, 514, 528.
Shallow-marine (regressive) facies: CSUN locality 51 2d.

Remarks. Specimens are most common at locality 371,
where they are fragile, well-preserved single valves.

Superfamily Pinnacea
Family Pinnidae Leach, 1819
Genus Pinna Linne, 1758

Type Species. By subsequent designation (Children, 1823),
Pinna rudis Linne, 1758.

Pinna lewisi Waring, 1917

Figure lOf

Pinna lewisi Waring, 1914:785; 1917:94, pi. 15, fig. 24.
Pinna (Pinna) lewisi Waring. Moore, 1983:A79, pi. 21, fig.
10.

Primary Type Material. CAS/SU holotype 5194, Llajas
Formation, SU locality 2696.

42 Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks

Molluscan Stage Range. “Domengine.”

Geographic Distribution. Simi Valley through south end
of San Joaquin Valley, California.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN locality 548. Shallow-marine (regressive) facies: CSUN
locality 475.

Remarks. Only six specimens were found, and five are from
locality 475. Preservation is mostly as internal molds, and
four specimens are articulated. The following supplementary
comments are based on an examination of the holotype. It
is mostly an internal mold with 95 percent of the shell miss-
ing. There are 16 to 18, very closely spaced radial ribs and
about nine comarginal ribs on each valve. The holotype and
the Llajas specimens have a median sulcus.

Pinna llajasensis Squires, 1983a

Figure lOg

Pinna n. sp. Vokes, 1939:50, pi. 2, fig. 14. Moore, 1983:A79,

pi. 22, fig. 1.

Pinna llajasensis Squires, 1983a:359-360, fig. 21.

Primary Type Material. UCLA holotype 59196, Llajas
Formation, CSUN locality 458.

Molluscan Stage Range. “Domengine.”

Geographic Distribution. Simi Valley through central Cal-
ifornia.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 374, 458.

Remarks. Only two specimens were found and these are
from the “Stewart bed.” Both are articulated, incomplete
specimens (Squires, 1983). The holotype is the figured spec-
imen (Fig. lOg).

P. llajasensis differs from P. lewisi in having fewer radial
ribs (only 1 2) which are more widely spaced. P. llajasensis
also has no comarginal sculpture.

In the original description by Vokes (1939) of his Pinna
n. sp., he mentioned that his specimen (UCMP 32595, Do-
mengine Formation, UCMP locality 2287) had 23 ribs. The
following supplementary comments are based on an exam-
ination of the holotype. There are 12 ribs per single valve
and about 23 for the entire surface including both valves.

Order Pterioida
Suborder Pteriina
Superfamily Pteriacea
Family Malleidae Lamarck, 1819
Genus Nayadina Munier-Chalmas, 1864

Type Species. By monotypy, Nayadina herberti Munier-
Chalmas, 1864.

Subgenus Exputens Clark, 1934

Type Species. By subsequent designation (Vokes, 1939),
Exputens llajasensis Clark, 1934.

Nayadina ( Exputens ) llajasensis (Clark, 1934)

Figures lOh-i

Exputens llajasensis Clark, 1934:270-271, pi. 37, figs. 1 1-
18. Vokes, 1939:51.

Nayadina ( Exputens ) llajasensis (Clark). Givens, 1974:44,
pi. 1, fig. 9. Moore, 1983:A86-A87, pi. 26, figs. 10, 13.

Primary Type Material. UCMP holotype 32391, UCMP
paratypes 32390, 32393, Llajas Formation, UCMP locality
7004 = CSUN locality 374.

Molluscan Stage Range. “Capay” through “Domengine.”
Geographic Distribution. Simi Valley through Pine Moun-
tain area, southern California.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 374, 445, 447, 471b, 473, 479, 480.

Remarks. Specimens occur mainly in the “Stewart bed”
at and in the vicinity of locality 374. In addition, a few
specimens were found in glauconitic siltstone at locality 473,
about 14 m above the “Stewart bed” at the type section. The
bed that locality 473 occurs in probably represents a shallow-
marine (transgressive) facies bed that interfingers with the
outer shelf and slope facies. At localities 374 and 473, a few
of the specimens are articulated.

Superficially, this genus may resemble Hiatella, but Elia-
tella is a heterodont with two weak teeth whereas Nayadina
lacks hinge teeth. A view of the hinge of N. ( E .) llajasensis
is given in Figure lOi.

Superfamily Pectinacea
Family Spondylidae Gray, 1826
Genus Spo ndylus Linne, 1758

Type Species. By subsequent designation (Schmidt, 1818),
Spondylus gaederopus Linne, 1758.

Spondylus carlosensis Anderson, 1905

Figure lOj

Spondylus carlosensis Anderson, 1905:194, pi. 13, fig. 1. Ar-
nold, 1 9 1 0:pl. 2, figs. 6-7. Dickerson, 191 5:pl. 1, fig. 7.
Anderson and Hanna, 1925: 189-190, text figure 10. Vokes,
1939:57, pi. 3, figs. 10, 13. Kleinpell and Weaver, 1963:
199, pi. 31, fig. 6.

Primary Type Material. CAS holotype 56, Domengine
Formation, west and north of Coalinga, NW Va of section 35,
T 20 S, R 14 E, Fresno County, California.

Molluscan Stage Range. “Domengine.”

Geographic Distribution. Simi Valley through central Cal-
ifornia.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 374, 438, 444, 525.

Remarks. Specimens are relatively uncommon in the Lla-
jas Formation, and they consist of fragments of single valves.
The most complete and best preserved specimens are from
locality 374.

Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks 43

Figure 10. Llajas Formation bivalves, a. Acila ( Truncacila ) decisa (Conrad, 1855), right valve, LACMIP hypotype 6554, CSUN loc. 467,
x3.5. b. Nuculana ( Saccella ) gabbi (Gabb, 1864), right valve, UCLA hypotype 59279, CSUN loc. 467, x3. c. Glycymeris ( Glycymeris )
rosecanyonensis Hanna, 1927, right? valve, LACMIP hypotype 6565, CSUN loc. 489, x4.5. d. Glycymeris (Glycymerita) sagittata (Gabb,
1864), right? valve, LACMIP hypotype 6566, CSUN loc. 371, x2. e. Brachidontes ( Brachidontes ) cowlitzensis (Weaver and Palmer, 1922),
right valve, LACMIP hypotype 6567, CSUN loc. 371, x 1.5. f. Pinna lewisi (Waring, 1917), internal mold of right? valve, LACMIP hypotype

44 Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks

Superfamily Anomiacea
Family Anomiidae Rafinesque, 1815
Genus Anomia Linne, 1758

Type Species. By subsequent designation (Schmidt, 1818),

Anomia ephippium Linne, 1758.

Anomia mcgoniglensis Hanna, 1927

Figure 10k

Anomia mcgoniglensis Hanna, 1927:278, pi. 31, figs. 1, 2, 5,
7. Turner, 1938:46, pi. 6, figs. 4-6. Weaver, 1943:100, pi.
22, figs. 4-5. Givens and Kennedy, 1976:974, pi. 4, figs.
13-16. Givens and Kennedy, 1979:table 2.

Primary Type Material. UCMP syntypes 31009-31010,
Delmar Formation, UCMP locality 3981.

Molluscan Stage Range. “Domengine.”

Geographic Distribution. San Diego, California through
southwestern Oregon.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN locality 438.

Remarks. Only three specimens were found in the Llajas
Formation. They occur as single valves unattached to any
hard substrate.

Suborder Ostreina
Superfamily Ostreacea
Family Ostreidae Rafinesque, 1815
Subfamily Ostreinae Rafinesque, 1815
Genus Ostrea Linne, 1758

Type Species. By subsequent designation (ICZN opin. 94
and 356), Ostrea edulis Linne, 1758.

Ostrea idriaensis Gabb, 1869

Figure 101

Ostrea idriaensis Gabb, 1869:203, pi. 33, figs. 103b-d; pi.
34, figs. 103, 103a. Waring, 1917:78-79, pi. 13, fig. 10.
Hanna, 1927:276, pi. 30, figs. 1-2; pi. 31, figs. 3-4. Stewart,
1930:126-127, pi. 8, fig. 3; pi. 17, fig. 1. Vokes, 1935:291-
304, pi. 22-24. Merriam and Turner, 1937:table 2. Turner,
1938:46, pi. 6, fig. 9. Weaver, 1943:78-79, pi. 15, fig. 5.
Givens, 1974:44. Givens and Kennedy, 1 979:tables 2, 4.
Not Ostrea haleyi Hertlein, 1933:277-282, pi. 18, figs. 5-6.

Primary Type Material. MCZ lectotype 15048, Domen-
gine Formation, about 3 km east of the Hacienda at New
Idria, N V2 of section 15, T 17 S, R 12 E, Priest Valley
quadrangle, San Benito County, California.

Molluscan Stage Range. “Capay” through "Tejon.”

Geographic Distribution. San Diego, California through
western Washington.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 374, 444, 458, 468a, 478, 485, 548.

Remarks. Ostreid fragments occur at localities 502, 503,
and 542 in the interfingering coastal alluvial-fan facies and
shallow-marine (transgressive) facies, but generic identifi-
cations could not be made. Similarly, ostreid fragments occur
at localities 463 and 526 in the lower part of the shallow-
marine (transgressive) facies, but generic identifications could
not be made. Ostreid fragments and single valves of O. id-
riaensis are fairly common in the uppermost part of the
“Stewart bed.” The figured specimen (Fig. 101) is one of the
largest specimens.

Subclass Heterodonta
Order Veneroida
Superfamily Lucinacea
Family Lucinidae Fleming, 1828
Subfamily Milthinae Chavan, 1969
Genus Claibornites Stewart, 1930

Type Species. By original designation, Lucina rotunda Lea,
1833.

Claibornites diegoensis (Dickerson, 1916)

Figure 10m

Lucina diegoensis Dickerson, 1916:484, pi. 37, figs. la-b.
Claibornites diegoensis ( Dickerson). Givens, 1974:45-46, pi.

1, fig. 15.

Primary Type Material. UCMP holotype 1 1788, Ardath
Shale, UCMP locality 2226.

Molluscan Stage Range. “Domengine.”

Geographic Distribution. San Diego through Pine Moun-
tain area, southern California.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 374, 445.

Remarks. At locality 374, only two single valves were found.
An articulated specimen was found at locality 445.

6568, CSUN loc. 475, x 1. g. Pinna liajasensis Squires, 1983a, internal mold of right valve, UCLA holotype 59196, CSUN loc. 458, x0.5.
h and i. Navadina (Exputens) liajasensis (Clark, 1934). All parts from CSUN loc. 374. h, right valve, LACMIP topotype and hypotype 6569,
x 1. i, hinge line, LACMIP topotype and hypotype 6570, x 1.4. j. Spondylus carlosensis Anderson, 1905, left valve, LACMIP hypotype 6571,
CSUN loc. 374, x l. k. Anomia mcgoniglensis Hanna, 1927, right valve, UCLA hypotype 59280, CSUN loc. 438, x 1.5. 1. Ostrea idriaensis
Gabb, 1869, left valve, LACMIP hypotype 6572, CSUN loc. 458, x0.5. m. Claibornites diegoensis (Dickerson, 1916), right valve, UCLA
hypotype 59281, CSUN loc. 374, x 1. n and o. Venericardia ( Pacifcor ) hornii calafia Stewart, 1930. All parts from CSUN loc. 374 and x0.5.
n, left valve, LACMIP topotype and hypotype 6573. o, right valve hinge line, LACMIP topotype and hypotype 6574. p. Venericardia (Pacificor)
aragonia joaquinensis (Vokes, 1939), UCMP hypotype 37433, UCMP loc. 7193 = CSUN loc. 702, x 1. q and r. Glyptoactis domenginica
(Vokes, 1939), LACMIP hypotype 6575, CSUN loc. 371, x2.5. q, left valve exterior, r, left valve interior.

Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks 45

Superfamily Carditacea

Family Carditidae Fleming, 1 820

Subfamily Venericardiinae Chavan, 1969

Genus Venericardia Lamarck, 1801

Type Species. By subsequent designation (Schmidt, 1818),
Venericardia imbricata Lamarck, 1801.

Subgenus Pacificor Verastegui, 1953

Type Species. By original designation, Venericardia ( Pa-
cificor) mulleri Verastegui, 1953.

Venericardia ( Pacificor ) hornii calafia
Stewart, 1930

Figures lOn-o

Venericardia hornii calafia Stewart, 1930:168-170, pi. 11,
fig. 2. Turner, 1938:50, pi. 14, fig. 4. Weaver, 1943:1 34—
135, pi. 28, figs. 6-7; pi. 31, figs. 4-5.

Venericardia ( Pacificor ) calafia Stewart. Verastegui, 1953:
28-30, pi. 15, figs. 3-5, 7; pi. 16, figs. 1-3; pi. 17, figs.
1-2.

Venericardia ( Pacificor ) hornii calafia Stewart. Givens, 1 974:
47, pi. 4, fig. 1. Saul, 1983:74, 76, pi. 2, figs. 9, 16-17.

Primary Type Material. UCMP holotype 31450, Llajas
Formation, UCMP locality 7004 = CSUN locality 374.
Molluscan Stage Range. “Domengine.”

Geographic Distribution. Simi Valley, California through
southwestern Oregon.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 374, 445, 451, 458, 494, 548, 704.

Remarks. A growth series of this taxon was found at lo-
cality 374. At this locality, most of the specimens are single
valves, but there are several well-preserved, large-sized ar-
ticulated specimens. At locality 548, only a single articulated
juvenile was found.

Saul (1983) considered V. (P.) oregonensis Verastequi from
the Lookingglass Formation of southwestern Oregon to be
an immature V. (P.) hornii calafia. V. (P.) hertleini Veras-
tequi is also probably V. (P.) hornii calafia (Saul, 1984, pers.
commun.).

Venericardia ( Pacificor ) aragonia
joaquinensis {\ okes, 1939)

Figure lOp

Venericardia aragonia var. Turner, 1938:49, pi. 13, figs.
6-9.

Megacardita ( Venericor ) hornii joaquinensis Vokes, 1939:
69-70, pi. 8, figs. 1-2; pi. 9, figs. 1-2.

Venericardia ( Leuroactis ) schencki Verastegui, 1953:50-51,
pi. 4, figs. 6-8.

Venericardia ( Leuroactis ) alisoensis Verastegui, 1 953:52-53,
pi. 10, figs. 1-3.

Venericardia ( Leuroactis ) joaquinensis (Vokes). Verastegui,
1953:60-61, pi. 11, figs. 1-4; pi. 12, figs. 4-6.

Venericardia (Leuroactis) vokesi Verastegui, 1953:61-62, pi.

14, figs. 1-3.

Venericardia (Pacificor) aragonia joaquinensis (V okes). Saul,

1983:pl. 2, figs. 7-8.

Primary Type Material. UCMP holotype 15616, Avenal
Formation, UCMP locality 4170; UCMP paratype 15617,
Avenal Formation, UCMP locality 4169; UCMP paratype
15618, Avenal Formation, UCMP locality A-819.

Molluscan Stage Range. Uppermost “Capay” through
“Domengine.”

Geographic Distribution. Simi Valley through central Cal-
ifornia.

Local Occurrence. Interfingering coastal alluvial-fan facies
and shallow-marine (transgressive) facies: CSUN localities
452, 700, 702, 706.

Remarks. Most specimens in the Llajas Formation are
fragments of single valves. Extraction of identifiable speci-
mens from the rock is usually difficult. At locality 700, only
a single juvenile was found.

The synonymy of this taxon is based on work by Saul
(1984, pers. commun.). V. (P.) a. smileyi (Vokes) may be
conspecific with V. (P. ) a. joaquinensis, but more taxonomic
work is needed to resolve the matter (Saul, 1 984, pers. com-
mun.).

The figured specimen (Fig. lOp) is UCMP hypotype 37433
from UCMP locality 7193 = CSUN locality 702. It is the
same specimen as used by Saul (1983, pi. 2, fig. 8).

Verastegui (1953) reported V. (L.) schencki from the Santa
Susana Formation, but his vague locality information makes
it impossible to know exactly where he collected his speci-
mens.

The presence of V. (P.) a. joaquinensis in the zone of in-
terfingering between the coastal alluvial-fan facies and the
shallow-marine (transgressive) facies of the Llajas Formation
refines the lower limit of the molluscan stage range of this
taxon as uppermost “Capay.”

Subfamily Carditesinae Chavan, 1969
Genus Glyptoactis Stewart, 1930

Type Species. By original designation, Venericardia hadra
Dali, 1903.

Subgenus Glyptoactis s.s.

Glyptoactis ( Glyptoactis ) domenginica
(Vokes, 1939)

Figures lOq-r

Venericardia ( Glyptoactis ?) domenginica Vokes, 1939:66, pi.

5, figs. 7-9.

Venericardia (Glyptoactis) domenginica Vokes. Verastegui,

1953:43-44, pi. 13, fig. 1.

Glyptoactis domenginica (Vokes). Givens, 1974:47. Squires,

1977:table 1. Givens and Kennedy, 1979:tables 1, 3.

Primary Type Material. UCMP holotype 15611, Domen-
gine Formation, UCMP locality A-1219; UCMP paratypes
15612-15613, Tejon Formation, UCMP locality A-1003.

46 Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks

Molluscan Stage Range. “Domengine” through “Transi-
tion.”

Geographic Distribution. San Diego through Mt. Diablo,
California.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 371, 507, 518, 529a.

Remarks. This taxon is confined to the shallow-marine
(transgressive) facies of the Llajas Formation. Specimens are
most abundant at locality 37 1 where there is a growth series.
Preservation is excellent, showing the angular nodes on the
ribs. No articulated specimens were found.

This species is placed in the subgenus Glyptoactis s.s. be-
cause the radial ribs are not tripartite. It is the oldest species
of this subgenus.

Superfamily Crassatellacea
Family Crassatellidae Ferussac, 1822
Subfamily Crassatellinae Ferussac, 1822
Genus Crassatella Lamarck, 1799

Type Species. By subsequent designation (Schmidt, 1818),
Mactra cygnaea Lamarck, 1799 (not Chemnitz, 1782) [=C.
gibba Lamarck, 1801, = Venus ponderosa Gmelin, 1791],

Crassatella uvasana Conrad, 1855

Figures lla-g

Crassatella uvasana Conrad, 1855:9; 1857, pi. 2, fig. 5. Gabb,
1864:214-215, pi. 32, fig. 284. Stewart, 1930:141-143, pi.
12, fig. 9. Turner, 1938:47-48. Givens, 1974:48.
Crassatella alta Conrad, 1855:9; 1857:321 [not Conrad, 1832:
21, pi. 7],

Crassatella grandis Gabb, 1864:181, pi. 24, fig. 163; 1869:
189.

Astarte semidentata Cooper, 1894:48, pi. 3, figs. 44-45.
Crassatellites grandis (Gabb). Arnold, 1910:13, pi. 2, figs.
10, 10a, pi. 3, fig. 14. Dickerson, 1915:80, pi. 1, fig. 8; pi.
2, figs, la-b [not Waring, 1917:74, pi. 12, fig. 16 = Cras-
satella branneri fide Nelson, 1925:410],

Crassatellites uvasana (Conrad). Arnold and Hannibal, 1913:
569. Dickerson, 1915:80, pi. 2, fig. 2. Waring, 1917:59, pi.
8, fig. 10.

Crassatellites mathewsonii (Gabb). Dickerson, 191 6:pl. 36,
figs. 9a-b (probably C. semidentata (Cooper) fide Turner,
1938:47-48).

Crassatellites uvasanus (Conrad). Anderson and Hanna, 1 925:
172-174, pi. 4, figs. 2-3, text figure 7.

Crassatellites semidentata (Cooper). Hanna, 1927:282, pi.
35, figs. 1-2.

Crassatella semidentata (Cooper). Turner, 1938:47-48.
Crassatella uvasana semidentata (Cooper). Vokes, 1939:64-
65, pi. 4, figs. 4, 6, 8, 10, 12. Givens, 1974:48. Squires,
1977:table 1. Givens and Kennedy, 1979:tables 1, 3.
Crassatella uvasana uvasana (Conrad). Givens and Kenne-
dy, 1979:table 4.

Primary Type Material. Holotype undetected, USNM col-
lection, Tejon Formation, Grapevine Canyon, Tejon quad-
rangle, Kern County, California.

Molluscan Stage Range. “Domengine” through “Tejon.”

Geographic Distribution. San Diego through central Cal-
ifornia.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 374, 444, 445, 447, 458, 471c, 481, 488,
494. Outer shelf and slope channel: CSUN locality 541.

Remarks. Ninety-six specimens were found, and most oc-
cur as single valves. Twelve specimens are articulated. A
single, abraded fragment was found at the base of a turbidite-
filled channel at locality 54 1 . Numerous well-preserved spec-
imens of C. uvasana occur in the “Stewart bed” at localities
374 and 444. Heights range from 20 to 90 mm at locality
374 and from 20 to 65 mm at locality 444. There is a growth
series at both localities. Many of the juvenile specimens and
a few of the adult specimens are articulated at both localities.
For the disarticulated specimens, the ratio of right valves to
left valves is approximately 1:1. Based on criteria listed by
Fagerstrom ( 1 964), the C. uvasana specimens at these “Stew-
art bed” localities are clearly in situ.

There is a gradual change in the external appearance of
the shells from juvenile to adult individuals at these localities
(Figs. 1 la-c). The juvenile specimens have higher and more
prominent beaks, a less deep lunule, a less steep posterior
dorsal slope, a more concave anterior dorsal margin, and a
less obvious escutcheon on the left valve. The adult speci-
mens have a more triangular shape and lower, more rounded
beaks. The escutcheon on the right valve is much larger than
that on the left. Intermediate-sized specimens (with a height
of about 45 mm) are transitional between the two.

The comarginal ribbing is identical in both the juvenile
and adult specimens. There are numerous very fine, closely
spaced comarginal ribs. At regular intervals there are comar-
ginal rugae. Upon decortication, especially in the beak areas,
shell between the rugae has a tendency to chip off, leaving
depressed areas outlined by the rugae. Only the adult spec-
imens show cancellate sculpture along the ventral margins
of the valves.

Interior shell features of juvenile and adult specimens of
C. uvasana are identical (Figs. 1 1 d— g). The resilifer extends
halfway to the lower margin of the hinge plate and the pit is
large. The anterior muscle scar is reniform, and the pallial
line is deeply impressed. In both juvenile and adult speci-
mens the right valve has the following features: anterior ven-
tral margin of the hinge plate tends to be swollen and fits
into a socket in the left valve; anterior cardinal is large,
wedge-shaped, vertically below the beak, its upper end mar-
gining part of the resilifer; the floor of the lunule tends to
wrap around the anterior cardinal in some specimens (Fig.

1 If); posterior cardinal is weak; edge of the escutcheon is
swollen and projects as a long ridge that fits into a socket in
the left valve.

In both juvenile and adult specimens of C. uvasana. the
left valve has the following features: posterior ventral margin
of the hinge plate is slightly swollen; anterior and posterior
cardinals are about equal in size; floor of lunule tends to wrap

Contributions in Science, Number 350

Squires: Simni Valley Eocene Mollusks 47

Figure 11. Llajas Formation bivalves (continued), a through g. Crassatella uvasana Conrad, 1855. All parts from CSUN loc. 374. a, left
valve, LACMIP hypotype 6576, x 1. b, left valve, LACMIP hypotype 6577, x 1. c, left valve, LACMIP hypotype 6578, x 1. d, right valve
interior of juvenile, LACMIP hypotype 6579, x 1. e, left valve hinge line of juvenile, LACMIP hypotype 6580, x 1.5. f, right valve hinge line
of adult, LACMIP hypotype 6581, xl. g, left valve interior of adult, LACMIP hypotype 6582, x 1.

48 Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks

around the anterior cardinal in some specimens, ventral mar-
gin of lunule tends to be swollen. In the left valve, as well as
in the right valve, the posterior part of the hinge plate is
largely occupied by a smooth surface without hinge struc-
tures.

If it had not been for the presence of a growth series, the
juvenile specimens of C. uvasana would have been identified
as C. uvasana semidentata (Cooper), and the adult specimens
would have been identified as C. uvasana. Based on the
transition of shell characters from the juvenile and adult, as
well as the identical shell characters mentioned above, it is
concluded that C. uvasana semidentata is an unjustified sub-
species.

According to Hanna ( 1 927) and Givens ( 1 974), the nature
of the beaks is the main distinguishing character between C.
uvasana and C. uvasana semidentata. In actuality, this char-
acter can be used to distinguish juvenile from adult speci-
mens of C. uvasana as the beaks become more incurved with
age. It is interesting to note that if a juvenile specimen is
placed on a flat surface, viewed laterally, and elevated along
the venter (simulating additional shell material), the beak
area then looks identical to that of an adult specimen.

The molluscan stage range of C. u. semidentata had been
reported as “Domengine” through “Transition” and that of
C. uvasana as “Tejon” (Givens, 1 974). Because the two taxa
are identical, the molluscan stage range of C. uvasana can
be extended to “Domengine” through “Tejon.”

Superfamily Cardiacea

Family Cardiidae Lamarck, 1 809

Subfamily Cardiinae Lamarck, 1809

Genus Acanthocardia Gray, 1851

Type Species. By subsequent designation (Stoliczka, 1 870),
Cardium aculeatum Linne, 1758.

Subgenus Schedocardia Stewart, 1930

Type Species. By original designation, Cardium hatche-
tigbeense Aldrich, 1886.

Acanthocardia ( Schedocardia ) brewerii
(Gabb, 1864)

Figures 12a-b

Cardium brewerii Gabb, 1864:173, pi. 24, fig. 155. Arnold,

1 907 :pl. 39, fig. 5. McLaughlin and Waring, 191 5:fig. 14.
Waring, 1 9 1 7:pl. 14, fig. 9. Anderson and Hanna, 1925:
165-166, pi. 1, fig. 3. Clark, 1929:pl. 12, fig. 7.
Plagiocardium ( Schedocardia ) brewerii (Gabb). Stewart, 1 930:
256-258, pi. 12, fig. 6. Turner, 1938:52-53, pi. 9, figs. 6-
7. Vokes, 1939:75, pi. 1 1, figs. 1-4. Stewart, 1946:pl. I 1,
fig. 20.

Plagiocardium brewerii (Gabb). Merriam and Turner, 1937:
table 2.

Loxocardium (Schedocardia) brewerii) Gabb). Weaver, 1943:
153-154, pi. 35, figs. 15, 16, 18; pi. 38, figs. 1, 9; pi. 104,
fig. 12.

Cardium ( Trachycardium) brewerii brewerii (Gabb). Klein-
pell and Weaver, 1963:201-202, pi. 34, figs. 1-2.
Acanthocardia (Schedocardia) brewerii (Gabb). Givens, 1974:
48-49, pi. 1, fig. 17.

Acanthocardia brewerii (Gabb). Givens and Kennedy, 1979:
table 4.

Primary Type Material. ANSP lectotype 4560, Tejon For-
mation, east of north end of Grapevine Canyon, Kern Coun-
ty, California.

Molluscan Stage Range. “Capay” through “Tejon.”
Geographic Distribution. San Diego, California through
southwestern Washington.

Local Occurrence. Interfingering coastal alluvial-fan facies
and shallow-marine (transgressive) facies: CSUN localities
545, 702. Shallow-marine (transgressive) facies: CSUN lo-
calities 371, 374, 548, 704.

Remarks. This taxon is somewhat uncommon in the Llajas
Formation. It is most common at locality 374 where it occurs
as single valves. The largest specimens are also from this
locality.

Subfamily Protocardiinae Keen, 1951

Genus Nemocardium Meek, 1876

Type Species. By subsequent designation (Sacco, 1899),
Cardium semiasperum Deshayes, 1858.

Nemocardium linteum (Conrad, 1855)

Figure 1 2c

Cardium linteum Conrad, 1855:3, 9; 1857 :pl. 2, fig. 1. An-
derson and Hanna, 1925:166-167, pi. 3, fig. 3.

Cardium cooperii Gabb, 1864:172, pi. 24, figs. 154, 154a.
Arnold, 1 907 :pl 38, figs. 2-2a. Waring, 191 7:pl. 13, fig.
3. Hanna, 1927:285, pi. 41, figs. 6-7.

Cardium dalh Dickerson, 1913:289, pi. 14, figs. 4a-c.
Cardium marysvillensis Dickerson, 1916:482 [new name for
Cardium dalli Dickerson, 1913, preoccupied].

Cardium (Protocardium) marysvillensis Dickerson. Clark and
Woodford, 1927:94, pi. 15, fig. 12.

Nemocardium linteum (Conrad). Stewart, 1930:275-277, pi.
8, fig. 6. Turner, 1938:52, pi. 10, fig. 10. Vokes, 1939:7 6—
77, pi. 11, figs. 6, 9. Weaver, 1943:159-160, pi. 38, fig. 3;
1953:28. Stewart, 1946:pl. 11, fig. 19. Zinsmeister, 1974:
97-98, pi. 9, figs. 7-9; 1 983a:pl. 2, fig. 7. Givens and Ken-
nedy, 1 979:table 4.

Cardium (Nemocardium) linteum Conrad. Kleinpell and
Weaver, 1963:202, pi. 34, fig. 4.

Nemocardium (Nemocardium) linteum (Conrad). Givens,
1974:49. Squires, 1977:table 1.

Primary Type Material. USNM holotype 1834, Domen-
gine Formation, near Martinez, California.

Molluscan Stage Range. “Martinez” through “Tejon.”
Geographic Distribution. San Diego, California through
southwestern Oregon.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 373, 374, 438, 444, 445, 458, 479, 488, 493,
513, 525, 548.

Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks 49

Remarks. This taxon is most common in the upper part
of the shallow-marine (transgressive) facies. At locality 438,
six specimens were found, one of which was articulated. In
the “Stewart bed” at locality 374, 21 specimens were found
and one was articulated.

Superfamily Solenacea
Family Solenidae Lamarck, 1809
Genus Solena Morch, 1853

T ype Species. By subsequent designation (Stoliczka, 1871),
Solen obliquus Spengler, 1794.

Subgenus Eosolen Stewart, 1930

Type Species. By original designation, Solen plagiaulax
Cossmann, 1906.

Solena ( Eosolen ) novacularis
(Anderson and Hanna, 1928)

Figure 12d

Solen novacula Anderson and Hanna, 1925:147, pi. 6, fig. 9.

Hanna, 1927:294, pi. 43, fig. 1.

Not Solen novacula Montagu, 1803:47.

Solen novacularis Anderson and Hanna, 1928:65-66 [new
name for Solen novacula Anderson and Hanna, 1925,
preoccupied],

Solena ( Eosolen ) coosensis Turner, 1938:62-63, pi. 9, figs.
1-2. Vokes, 1939:96, pi. 15, fig. 5. Givens, 1974:49-50,
pi- 2, fig. 1.

Solena coosensis Turner. Weaver, 1943:229, pi. 52, fig. 16;
pi. 53, fig. 13.

Solena novacularis (Anderson and Hanna). Givens and Ken-
nedy, 1979:table 4.

Primary Type Material. CAS holotype 882 of Solen no-
vacula Anderson and Hanna and of Solen novacularis An-
derson and Hanna, Tejon Formation, CAS locality 792.

Molluscan Stage Range. Uppermost “Capay”?, “Domen-
gine” through “Tejon.”

Geographic Distribution. San Diego, California through
southwestern Oregon.

Local Occurrence. Interfingering coastal alluvial-fan facies
and shallow-marine (transgressive) facies: CSUN locality 701?
Shallow-marine (transgressive) facies: CSUN localities 374,
444, 445, 449, 458, 477, 494, 500.

Remarks. This species helps to characterize the “Stewart
bed.” At many localities of the “Stewart bed,” and especially
at locality 374, specimens are fairly common and many are
articulated. A solenid at locality 701 may be this species.

A previously assigned “Capay” age (Turner, 1 938; Givens,
1974) for the lower range limit of this species was based on
occurrence in the “upper Umpqua” Formation of south-
western Oregon. The “upper Umpqua,” however, is actually
equivalent to both the “Capay” and “Domengine” “Stages”
(Baldwin, 1974; Miles, 1981). A lower range limit of upper-
most “Capay,” nevertheless, is apparent because of the pos-
sible occurrence of this species in the zone of interfingering

between the coastal alluvial-fan facies and the shallow-ma-
rine (transgressive) facies of the Llajas Formation.

Superfamily Tellinacea
Family Tellinidae Blainville, 1814
Subfamily Macominae Olsson, 1961
Genus Macoma Leach, 1819

Type Species. By monotypy, Macoma tenera Leach, 1819
[=Tellina calcarea Gmelin, 1791].

Macoma rosa Hanna, 1927

Figure 12e

Macoma rosa Hanna, 1927:292, pi. 41, figs. 2-5, 8. Clark,

1929:pl. 6, fig. 15. Givens and Kennedy, 1979:table 1.

Primary Type Material. UCMP holotype 3101)4, Ardath
Shale, UCMP locality 3993; UCMP paratype 31095, Ardath
Shale, UCMP locality 5089; UCMP paratypes 3 1 096-3 1 097,
Ardath Shale, UCMP locality 5085.

Molluscan Stage Range. “Domengine.”

Geographic Distribution. San Diego through Simi Valley,
California.

Local Occurrence. Shallow-marine (regressive) facies:
CSUN localities 462, 489, 512a, 51 2d, 544.

Remarks. This taxon helps to characterize the shallow-
marine (regressive) facies. Specimens are usually few in num-
ber and preserved as external and internal molds of articu-
lated individuals.

Family Psammobiidae Fleming, 1828
Subfamily Psammobiinae Fleming, 1828
Genus Gari Schumacher, 1817

Type Species. Pending decision by the ICZN, Gari vulgaris
Schumacher, 1817 ( =Solen amethystus Wood, 1818).

Gari cf. G. eoundulata Vokes, 1939

Figure 12f

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 374, 47 Id.

Remarks. Specimens are relatively rare in the Llajas For-
mation. They are preserved as internal molds of single valves,
making species identification impossible.

Superfamily Veneracea
Family Veneridae Rafinesque, 1815
Subfamily Pitarinae Stewart, 1930
Genus Callista Poli, 1791

Type Species. By subsequent designation (Meek, 1876),
Venus chione Linne, 1758.

50 Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks

Subgenus Costacallista Palmer, 1927

Type Species. By original designation, Venus erycina Linne,
1758.

Callista ( Costacallista ) cf.

C. (C.) hornii (Gabb, 1864)

Figure 12g

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 373, 465, 466, 467, 704.

Remarks. A few specimens were found as single valves.
Hinge-line details were imperfectly preserved, hence a certain
species identification could not be made.

Genus Callocardia A. Adams, 1864

Type Species. By monotypy, Callocardia guttata A. Adams,
1864.

Subgenus Nitidavenus Vokes, 1939

Type Species. By original designation, Cytherea nitida
Deshayes, 1858.

Callocardia ( Nitidavenus ) tejonensis
(Waring, 1914)

Figure 12h

Isocardia tejonensis Waring, 1914:784-785; 1917:93, pi. 15,
fig. 14.

cf. “ Isocardia tejonensis" Waring. Turner, 1938:58, pi. 11,
figs. 1-4.

Nitidavenus tejonensis (Waring). Vokes, 1 939:83-84, pi. 12,
figs. 11, 13-16.

Primary Type Material. SU holotype 189, SU paratypes
5188-5190, Llajas Formation, SU locality 2696.

Molluscan Stage Range. “Capay” through “Domengine.”
Geographic Distribution. Simi Valley, California through
southwestern Oregon.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 371, 372.

Remarks. Only four single valves were found in the Llajas
Formation.

Genus Pi tar Romer, 1857
Type Species. By monotypy, Venus lumens Gmelin, 1791.
Subgenus Calpitaria Jukes-Browne, 1908

Type Species. By original designation, Cytherea sulcataria
Deshayes, 1824.

Pitar (Calpitaria) uvasanus (Conrad, 1855).

Figures 12i-j

Meretrix uvasana Conrad, 1855:9; 1857:pl. 2, fig. 3. Gabb,
1864:163-164, pi. 30, fig. 248.

Meretrix tejonensis Dickerson, 1915:53-54, pi. 3, figs. 3a-b,
not 2a-b. [Unjustified new name for Meretrix uvasana
Conrad of Gabb, 1864, fide Anderson and Hanna, 1925:
160-161 and Stewart, 1930:236.]

Not Pitaria tejonensis (Dickerson). Anderson and Hanna,
1925:160-161, pi. 3, fig. 5.

Pitaria uvasana (Conrad). Anderson and Hanna, 1925:161,
pi. 5, figs. 3-4.

Pitar (Calpitaria) uvasanus (Conrad). Stewart, 1930:235-236,
pi. 12, fig. 7. Weaver, 1943:178-179, pi. 47, fig. 13. Givens,
1974:53.

Pitar uvasanus uvasanus (Conrad). Givens and Kennedy,
1979:tables 3-4.

Primary Type Material. ANSP neotype 4457, Tejon For-
mation, Grapevine Canyon, Kern County, California.
Molluscan Stage Range. “Domengine” through “Tejon.”
Geographic Distribution. San Diego, California through
western Oregon.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 371, 374.

Remarks. Specimens are fairly uncommon and generally
consist of fragments of single valves. The occurrence of this
taxon in the Llajas Formation extends its range into the
“Domengine Stage.” Previously, it had been reported as
ranging from the “Transition” through the “Tejon” (Givens,
1974).

Meretrix tejonensis Dickerson, 1915:53, pi. 3, figs. 2a-b =
Pitar ( Lamelliconcha ) dickersoni Givens, fide Givens, 1974:
53-54.

Subgenus Lamelliconcha Dali, 1902

Type Species. By original designation, Cytherea concinna
J. Sowerby, 1835a.

Pitar ( Lamelliconcha ) joaquinensis
Vokes, 1939

Figure 12k

Meretrix hornii Gabb. Arnold, 1 9 10:pl. 3, fig. 9. [Misiden-
tification.]

Pitar (Lamelliconcha) joaquinensis Vokes, 1939:85-86, pi.

13, figs. 9-12. Givens, 1974:54, pi. 3, fig. 7.

Pitar ? joaquinensis Vokes. Stewart, 1946:pl. 12, fig. 12.
Pitar joaquinensis Vokes. Givens and Kennedy, 1979:
table 1.

Primary Type Material. UCMP holotype 15674, Domen-
gine Formation, UCMP locality A- 1027; UCMP paratype
1 5675, Domengine? Formation, UCMP locality 4175; UCMP
paratype 15676, Domengine Formation, UCMP locality
A- 1027; UCMP paratype 1 5677, Avenal Formation, UCMP
locality A- 1280.

Molluscan Stage Range. “Domengine.”

Geographic Distribution. Simi Valley through central Cal-
ifornia.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN locality 371.

Remarks. Only a few single valves were found in the Llajas
Formation.

Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks 51

Figure 12. Llajas Formation bivalves (continued), a and b. Acanthocardia (Schedocardia ) breweri (Gabb, 1864), right valve, LACMIP hypotype

6583, CSUN loc. 371, x2. a, exterior, b, interior, c. Nemocardium linteum (Conrad, 1855), internal mold of right? valve, LACMIP hypotype

6584, CSUN loc. 374, x 1. d. Solena ( Eosolen ) novacularis (Anderson and Hanna, 1928), left valve, LACMIP hypotype 6585, CSUN loc.
374, x l. e. Macoma rosa Hanna, 1927, internal mold of right valve, LACMIP hypotype 6586, CSUN loc. 489, x 1.5. f. Gari cf. G. eoundulata
Vokes, 1939, internal mold of right valve, LACMIP hypotype 6587, CSUN loc. 47 Id, x 1. g. Callista ( Costacallista ) cf. C. (C.) hornii (Gabb,
1864), right valve, LACMIP hypotype 6588, CSUN loc. 465, x2. h. Callocardia (Nitidavenus) tejonensis (Waring, 1914), left valve, LACMIP

52 Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks

Subfamily Tapetinae H. and A. Adams, 1857

Genus Marcia H. and A. Adams, 1857

Type Species. By subsequent designation (Kobelt, 1883),
“ Venus pinguis Chemnitz” [= V. opima Gmelin, 1791],

Subgenus Mercimonia Dali, 1902

Type Species. By original designation, Venus bernayi Coss-
mann, 1888.

Marcia ( Mercimonia ) bunkeri (Hanna, 1927)

Figure 121

Dosinia bunkeri Hanna, 1927:287, pi. 42, figs. 4, 6.
Mercimonia bunkeri (Hanna). Turner, 1938:60, pi. 10, figs.
5-9. Vokes, 1939:77, pi. 11, fig. 12. Givens and Kennedy,
1979:87, table 3.

Marcia ( Mercimonia ) bunkeri (Hanna). Weaver, 1943:192,
pi. 45, fig. 5; pi. 47, figs. 8, 11.

Primary Type Material. UCMP holotype 30950, Ardath
Shale, UCMP locality 5069; UCMP paratype 30951, Ardath
Shale, UCMP locality 3976.

Molluscan Stage Range. “Domengine” through “Transi-
tion.”

Geographic Distribution. San Diego, California through
southwestern Oregon.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN locality 374.

Remarks. Only three specimens were found in the Llajas
Formation, and of these, two are articulated.

Order Myoida
Suborder Myina
Superfamily Myacea
Family Corbulidae Lamarck, 1818
Subfamily Corbulinae Lamarck, 1818
Genus Corbula Bruguiere, 1797

Type Species. By subsequent designation (Schmidt, 1818),
Corbula sulcata Lamarck, 1801.

Subgenus C laryocorbula Gardner, 1926

Type Species. By original designation, Corbula alaba-
miensis Lea, 1833.

Corbula ( Caryocorbula ) dickersoni
(Weaver and Palmer, 1922)

Figure 12m

Corbula dickersoni Weaver and Palmer, 1922:24-25, pi. 9,
figs. 9-10. Clark, 1938:700, pi. 1, fig. 17. Weaver, 1943:
257-258, pi. 61, figs. 13, 16-17, 20.

Corbula ( Caryocorbula ) dickersoni Weaver and Palmer.
Vokes, 1939:98, pi. 16, figs. 1, 5, 9. Givens, 1974:57, pi.
4, fig. 7.

Primary Type Material. CAS holotype 7452, CAS syn-
types 7452A-B, Cowlitz Formation, UW locality 329.

Molluscan Stage Range. Uppermost “Capay” through
“Tejon.”

Geographic Distribution. Simi Valley, California through
southwestern Washington.

Local Occurrence. Interfingering coastal alluvial-fan facies
and shallow-marine (transgressive) facies: CSUN locality 545.
Shallow-marine (transgressive) facies: CSUN localities 371,

373, 455, 484, 498, 501, 507, 540. Shallow-marine (regres-
sive) facies: CSUN locality 475.

Remarks. This taxon is most common at locality 371,
where the specimens are similar-sized single valves.

The presence of C. (C.) dickersoni in the zone of interfin-
gering between the coastal alluvial-fan facies and the shallow-
marine (transgressive) facies of the Llajas Formation extends
the molluscan stage range of this taxon into the uppermost
“Capay.” Previously, the lower range limit had been reported
as the “Domengine” (Vokes, 1939; Givens, 1974).

Suborder Phoiadina

Superfamily Pholadacea

Family Teredinidae Rafinesque, 1815

Subfamily Teredininae Rafinesque, 1815

Genus Teredo Linne, 1758

Type Species. By subsequent designation (ICZN, 1926,
opin. 94), Teredo navalis Linne, 1758.

Teredol sp.

Figure 12n

Local Occurrence. Interfingering coastal alluvial-fan facies
and shallow-marine (transgressive) facies: CSUN locality
469c. Shallow-marine (transgressive) facies: CSUN localities

374, 438, 444, 469, 470a, 488, 534. Outer shelf and slope

hypotype 6589, CSUN loc. 371, x 1. i and j. Pitar ( Calpitaria) uvasanus (Conrad, 1855), left valve, LACMIP hypotype 6590, CSUN loc. 371,
x 1.5. i, exterior, j, interior, k. Pitar ( La melliconcha) joaquinensis Vokes, 1939, partial specimen, right valve, LACMIP hypotype 6591, CSUN
loc. 371, x l . l. Marcia ( Mercimonia ) bunkeri ( Hanna, 1927), left? valve, UCLA hypotype 59282, CSUN loc. 374, x 1 . m. Corbula (Caryocorbula)
dickersoni (Weaver and Palmer, 1922), right valve, LACMIP hypotype 6592, CSUN loc. 371, x 2.75. n. Teredo ? sp., in petrified wood,
LACMIP hypotype 6593, CSUN locality 524, x0.5. o. Cardiomya aff. C. russelli (Hanna, 1927), internal mold of right valve, LACMIP
hypotype 6594, CSUN loc. 51 2d, x4.

Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks 53

Figure 13. Llajas Formation nautiloid, crab, spatangoid, and shark teeth, a. Aturia myrlae Hanna, 1927, lateral view, LACMIP hypotype
6595, CSUN loc. 374, xl.75. b. Unidentifiable spirulimorph sepiid, ventral view of partial phragmocone, UCLA hypotype 59197, CSUN
loc. 493, x 2.6. c. Glyphithyreus weaveri (Rathbun, 1926), carapace and portion of right cheliped, dorsal view, UCLA hypotype 59273, CSUN
loc. 548, x l . d. Schizaster diabloensis Kew, 1920, internal mold, dorsal view, LACMIP hypotype 6596, CSUN loc. 482, x 1. e. Odontaspis
sp., LACMIP hypotype 6597, CSUN loc. 374, x 1. f. Isurus cf. I. praecursor (Leriche, 1906), LACMIP hypotype 6598, CSUN loc. 371, x3.

facies: CSUN locality 446. Outer shelf and slope channel
facies: CSUN locality 541.

Remarks. Teredo ? sp. occurs in pieces of petrified wood.
In most cases. Teredo ? sp. is represented by calcareous-lined
burrows, but unlined burrows can be present in the same
piece of wood. Generic determination is uncertain, and it is
very possible that future workers will assign these fossils to
another genus. Tertiary teredinids are in need of much taxo-
nomic work.

Subclass Anomalodesmata
Order Pholadomyoida
Superfamily Poromyacea
Family Cuspidariidae Dali, 1886
Genus Cardiomya A. Adams, 1864

Type Species. By monotypy, Neaera gouldiana Hinds,
1843.

Cardiomya aff. C. russelli (Hanna, 1927)

Figure 12o

Local Occurrence. Shallow-marine (regressive) facies:
CSUN locality 5 1 2d.

Remarks. Only a single specimen was found, and it is an
internal mold of a right valve. The specimen agrees closely
with the description of C. russelli given by Hanna (1927:
280-28 1 ) in possessing radial ribs and a long, straight smooth
rostrum. The Llajas specimen differs in that it has at least
19 radial ribs rather than 16.

Class Cephalopoda
Subclass Nautiloidea
Order Nautilida
Superfamily Nautilacea
Family Aturiidae Hyatt, 1894
Genus Aturia Bronn, 1838

Type Species. By subsequent designation (Herrmannsen,
1846), Nautilus aturi Basterot, 1825, and virtual tautonymy.

Aturia myrlae Hanna, 1927

Figure 13a

Aturia myrli Hanna, 1927:331, pi. 57, figs. 1, 6.

Aturia myrlae Hanna. Schenck, 1931:454-456, pis. 67-68;
text figures 4-3, 4-19. Yokes, 1939:107, pi. 16, fig. 36.

54 Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks

Miller, 1947:100-101, pi. 76, figs. 2-3; pi. 77, figs. 1-2;

pi. 92, figs. 6-7.

Primary Type Material. UCMP holotype 31089, Ardath
Shale, UCMP locality 3989.

Molluscan Stage Range. “Capay” through “Domengine.”

Geographic Distribution. San Diego through central Cal-
ifornia.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 374, 458.

Remarks. Specimens were found only in the “Stewart bed.”
The best preserved specimens are from locality 374 where
one juvenile (Fig. 13a) and an adult specimen were found.
As reported by Schenck (1931), a hypotype of A. myrlae is
from UCMP 7004 = CSUN locality 374.

Subclass Coleoidea
Order Sepiida
Family Indeterminate

Spirulimorph Sepiid

Figure 13b

Spirulimorph sepiid Squires, 1983a:360, figs. 2j-k.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN locality 493.

Remarks. Only a single partial phragmocone and associ-
ated external mold were found in the Llajas Formation. In
the absence of the rostrum and the apical part of the phrag-
mocone, the specimen and mold are unidentifiable. This is
the first sepiid in the Eocene of western North America
(Squires, 1983a).

Phylum Arthropoda

Class Malacostraca

Subclass Eumalacostraca

Order Decapoda

Suborder Pleocyemata

Infraorder Brachyura

Section Brachyrhyncha

Superfamily Xanthoidea

Family Goneplacidae MacLeay, 1838

Subfamily Carcinoplacinae
Milne-Edwards, 1852

Genus Glyphithyreus Reuss, 1859

Type Species. By original designation, Glyphithyreus for-
mosus Reuss, 1859 ( = Plagioloph us wetherelli Bell, 1858).

28

Location of measured section

Contact

Fault

Generalized attitude of beds
Section number
Paved road

Improved secondary road
Unimproved secondary road

|Tsp| Sespe Format
[ T1 | Llajas Format
P US Shallow-mar

□

ne (regressive) facies

Outer shelf & slope facies with
localized turbidite-filled channels

Shallow-marine (transgressive) facies

Coastal alluvial-fan facies

Santa Susana Formation

Figure 14. Index map to the Llajas Formation showing locations
of areas used as fossil-collecting locality maps in Figures 15-19. An
explanation of symbols used on the locality maps is also given.

Glyphithyreus weaveri (Rathbun, 1926)

Figure 13c

Cancer ? sp. Weaver, 1905:123, pi. 13, fig. 11.

Cancer (?) sp. A Dickerson, 1 9 1 6:pl. 42, fig. 11.
Plagiolophus weaveri Rathbun, 1926:35-37, pi. 9, figs. 5-6.
Orr and Kooser, 1971:157, text figure 3a, figs. 4a-c, figs.
5a-i.

Primary Type Material. USNM holotype 353351, Eocene
strata. Salt Creek, 5.2 km north-northeast of Joaquin Rocks,
south part of NW Vi of section 15, T 18 S, R 14 E, Fresno
County, California.

Molluscan Stage Range. “Capay” through “Domengine.”
Geographic Distribution. San Diego, California through
southwestern Oregon.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 471b, 548.

Remarks. Although fragments of brachyuran chelipeds were
found at widely scattered localities in the Llajas Formation,
carapaces were found only at localities 471b and 548. At
locality 548, most of the specimens have also one or both
chelipeds intact.

Phylum Echinodermata
Subphylum Echinozoa
Class Echinoidea
Subclass Euechinoidea

Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks 55

• 462

. 476

• 475

• 489

. 541
. 473

. 472,458, 459,

n477, 478 ,
479, 480

.481
•440
. 439
* 537
•453

• 442

• 441

I ®

. 452,705

• 454,706

• 542

Figure 15. Geologic map showing CSUN fossil-collecting localities, Llajas Formation, type section area. Accompanying columnar section
shows stratigraphic position of the fossil-collecting localities. See Figure 14 for explanation of symbols.

Superorder Atelostomata

Order Spatangoida

Suborder Hemiasterina

Family Schizasteridae Lambert, 1902

Genus Schizaster L. Agassiz, 1836

Type Species. By subsequent designation (ICZN, 1954,
opin. 209), Schizaster studeri L. Agassiz, 1836.

Schizaster diabloensis Kew, 1920

Figure 1 3d

Schizaster diabloensis Kew, 1920:150-151, pi. 41, figs. 5a-
c. Clark and Woodford, 1927:123, pi. 22, fig. 14. Clark,
1929:pl. 4, fig. 13. Grant and Hertlein, 1938:120.

Primary Type Material. UCMP holotype 1 1387, Eocene
strata, UCMP locality 1427.

Molluscan Stage Range. Upper Paleocene through “Do-
mengine.”

Geographic Distribution. Simi Valley through Marysville
Buttes, California.

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 374, 458, 478. Outer shelf and slope facies:
CSUN locality 482. Shallow-marine (regressive) facies: CSUN
locality 547.

Remarks. Preservation is as internal molds and most spec-
imens are somewhat crushed. At locality 547, 5. diabloensis
is the only megafossil present.

Clark and Woodford (1927) reported this species from the
type section area of the Meganos Formation. Marincovich
(1977:251) considered the formation to be late Paleocene in
age.

Phylum Chordata

Class Chondrichthyes

Subclass Elasmobranchii

Order Lamniformes

Family Odontaspididae
Muller and Henle, 1837

Genus Odontaspis L. Agassiz, 1838

Type Species. By monotypy, Odontaspis ferox Risso, 1810.

Odontaspis sp.

Figure 13e

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN localities 374, 445.

56 Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks

Figure 16. Geologic map showing CSUN fossil-collecting localities, Llajas Formation, lower Las Llajas Canyon area. Accompanying columnar
section shows stratigraphic position of the fossil-collecting localities. See Figure 14 for explanation of symbols.

Remarks. Only a few specimens were found in the Llajas
Formation.

Family Lamnidae Muller and Henle, 1838
Genus Isurus Rafinesque, 1 809

Type Species. By original designation, Isurus oxyrhynchus
Rafinesque, 1809.

Isurus cf. I. praecursor (Leriche, 1906)

Figure 13f

Local Occurrence. Shallow-marine (transgressive) facies:
CSUN locality 371.

Remarks. Only one specimen was found in the Llajas For-
mation.

LOCALITIES

CSUN fossil-collecting localities made by the author in the
course of this study are listed first. Localities of other insti-
tutions mentioned in this report follow in alphabetical order.

All CSUN localities are in the Llajas Formation, Simi
Valley area, California. Unless otherwise noted, they are in
the United States Geological Survey 7.5-minute topographic

quadrangle of Santa Susana, California (195 1), photorevised
1969. In some cases, a locality is in the Santa Susana quad-
rangle, but the section corner used for referencing is in the
United States Geological Survey 7.5-minute topographic
quadrangle of Oat Mountain, California (1952), photorevised
1969. These localities are so noted. Abbreviations used are
United States Geological Survey (USGS), feet (ft.), meters
(m), township (T), range (R), north (N), south (S), east (E),
and west (W). Distances are given in both English and metric
units, but map contour elevations are given in English units
only.

CSUN localities equivalent to localities of University of
California Museum of Paleontology (Berkeley) (UCMP);
University of California, Los Angeles (UCLA); and Califor-
nia Institute of Technology (CIT) are so indicated. CSUN
localities are denoted also as to which marine facies (Squires,
1981) they belong. The general location and the relative
stratigraphic position of each CSUN locality are shown in
Figures 14 through 19.

CSUN LOCALITIES

371. At elevation of 2000 ft. on south side of a side canyon
to Devil Canyon, 1275 ft. (389 m) south and 1150 ft. (350

Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks 57

1 km

N

BASE MAP FROM SANTA SUSANA SUSANA (1951) AND OAT MOUNTAIN (1952). CA.,
7.5-MINUTE QUADRANGLES, BOTH PHOTOREVISED 1969.

Figure 17. Geologic map showing CSUN fossil-collecting localities, Llajas Formation, Devil Canyon area. Accompanying columnar section
shows stratigraphic position of the fossil-collecting localities. See Figure 14 for explanation of symbols.

m) west of NE comer of section 26, T 3 N, R 1 7 W. Locality
is in the Santa Susana quadrangle, but the section comer is
in the Oat Mountain quadrangle. [Shallow-marine (trans-
gressive) facies.]

372. At elevation of 2035 ft. on west side of stream bank
of Devil Canyon, 800 ft. (244 m) south and 1200 ft. (366 m)
west of NE comer of section 26, T 3 N, R 17 W. Locality is
in the Santa Susana quadrangle, but the section comer is in
the Oat Mountain quadrangle. [Shallow-marine (transgres-
sive) facies.]

373. At elevation of 1900 ft. on east side of stream bank
of Devil Canyon, 450 ft. (137 m) south and 850 ft. (259 m)
west of NE comer of section 26, T 3 N, R 17 W. Locality is
in the Santa Susana quadrangle, but the section corner is in
the Oat Mountain quadrangle. [Shallow-marine (transgres-
sive) facies.]

374. At elevation of 1700 ft. on a small cliff on south side
of a side canyon to Las Llajas Canyon, 1950 ft. (594 m) north
and 1825 ft. (556 m) east of SE comer of section 29, T 3 N,
R 17 W. Locality is in the “Stewart bed” and is equivalent
to UCMP locality 7004, UCLA locality 2312, and CIT lo-
cality 206. [Shallow-marine (transgressive) facies.]

438. At elevation of 1800 ft. on east side of stream bank

of Las Llajas Canyon, 7000 ft. (2 1 34 m) N86°E of NE comer
of section 29, T 3 N, R 1 7 W. Locality is equivalent to UCLA
locality 2775. [Shallow-marine (transgressive) facies.]

439. At elevation of 1425 ft. on a ridge, 800 ft. (244 m)
south and 1900 ft. (579 m) east of NW corner of section 31,
T 3 N, R 1 7 W. [Shallow-marine (transgressive) facies.]

440. At elevation of 1475 ft. on a ridge, 600 ft. (183 m)
south and 1875 ft. (571 m) east of NW comer of section 31,
T 3 N, R 17 W. [Shallow-marine (transgressive) facies.]

441 . At elevation of 1 1 60 ft. on west side of Chivo Canyon
near its mouth, 1890 ft. (576 m) south and 400 ft. (122 m)
west of NE comer of section 31, T 3 N, R 17 W. [Shallow-
marine (transgressive) facies.]

442. At elevation of 1 1 85 ft. on west side of Chivo Canyon
near its mouth, 1600 ft. (488 m) south and 510 ft. (155 m)
west of NE comer of section 31, T 3 N, R 17 W. [Shallow-
marine (transgressive) facies.]

443. At elevation of 1 230 ft. on east side of Chivo Canyon
near its mouth, 1370 ft. (418 m) south and 200 ft. (61 m)
west of NE comer of section 31, T 3 N, R 17 W. [Shallow-
marine (transgressive) facies.]

444. At elevation of 1 585 ft. on a small cliff on north side
of a side canyon to Las Llajas Canyon, 2500 ft. (762 m) north

58 Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks

468b, 516

• 438,468a, 515, 525

514 . 513

• 467 ’ 524

• 523
’ 521

. 518

• 520

. 460
• 517

• 466

• 465

• 463

Figure 18. Geologic map showing CSUN fossil-collecting localities, Llajas Formation, upper Las Llajas Canyon area. Accompanying columnar
section shows stratigraphic position of the fossil-collecting localities. See Figure 14 for explanation of symbols.

and 1375 ft. (419 m) east of SE comer of section 29, T 3 N,
R 17 W. Locality is in the “Stewart bed.” [Shallow-marine
(transgressive) facies.]

445. At elevation of 1485 ft. on a small cliff on west side
of Las Llajas Canyon, 2820 ft. (859 m) north and 850 ft. (259
m) east of SE comer of section 29, T 3 N, R 1 7 W. Locality
is in the “Stewart bed.” [Shallow-marine (transgressive) fa-
cies.]

446. At elevation of 1 760 ft. in roadcut along ridge on west
side of Las Llajas Canyon, 3460 ft. (1055 m) north and 200
ft. (61 m) east of SE corner of section 29, T 3 N, R 17 W.
[Outer shelf and slope facies.]

447. At elevation of 1550 ft. on a small cliff on west side
of Las Llajas Canyon, 2780 ft. (847 m) north and 550 ft. (168
m) east of SE comer of section 29, T 3 N, R 17 W. Locality
is in the “Stewart bed.” [Shallow-marine (transgressive) fa-
cies.]

449. At elevation of 1700 ft. on a small cliff on west side
of Las Llajas Canyon, 1825 ft. (556 m) north and 200 ft. (61
m) west of SE comer of section 29, T 3 N, R 17 W. Locality
is in the “Stewart bed.” [Shallow-marine (transgressive) fa-
cies.]

450. At elevation of 1600 ft. on a small cliff on west side
of Las Laljas Canyon, 2450 ft. (747 m) north and 125 ft. (38
m) east of SE comer of section 29, T 3 N, R 17 W. Locality

is in the “Stewart bed.” [Shallow-marine (transgressive) fa-
cies.]

451. At elevation of 1715 ft. on a small cliff on a hillside,
2100 ft. (640 m) north and 2100 ft. (640 m) west of SE corner
of section 29, T 3 N, R 17 W. Locality is in the “Stewart
bed.” [Shallow-marine (transgressive) facies ]

452. At elevation of 1 285 ft. on west side of a small canyon
branching north near mouth of Las Llajas Canyon, 1650 ft.
(503 m) south and 550 ft. (168 m) east of NW corner of
section 32, T 3 N, R 1 7 W. [Interfingering coastal alluvial-
fan facies and shallow-marine (transgressive) facies.]

453. At elevation of 1 275 ft. on west side of Chivo Canyon,
800 ft. (243 m) south and 700 ft. (213 m) west of NE comer
of section 3 1 , T 3 N, R 1 7 W. [Shallow-marine (transgressive)
facies.]

454. At elevation of 1 3 1 0 ft. on west side of a small canyon
branching north near mouth of Las Llajas Canyon, 1 100 ft.
(335 m) south and 810 ft. (247 m) east of NW comer of
section 32, T 3 N, R 17 W. [Interfingermg coastal alluvial-
fan facies and shallow-marine (transgressive) facies.]

455. At elevation of 1475 ft. on east side of stream bank
of Las Llajas Canyon, 2500 ft. (762 m) north and 625 ft. ( 191
m) east of SE comer of section 29, T 3 N, R 17W. [Shallow-
marine (transgressive) facies.]

457. At elevation of 1 300 ft. on east side of Chivo Canyon,

Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks 59

700 m

TsP _/

■ L X

-V

\

1

1

•

\

544

Tl

19

cv'L-

//

0>/‘

o

CO

A

-\ Tss

Tsp

• 544

• 547

d

■ - v

eSEEeje*-,

548

543

704

. 546
,703 . .

! 702
'700, 701

BASE MAP FROM SANTA SUSANA (1951, PHOTOREVISED 1969) AND THOUSAND OAKS (1950, PHOTOREVISED 1967), CA.. 7.5-MINUTE OUADRANGLES.

Figure 19. Geologic maps showing CSUN fossil-collecting localities, Llajas Formation, at various areas along the south side of Simi Valley.
Accompanying columnar sections shows stratigraphic position of the fossil-collecting localities. See Figure 14 for explanation of symbols, a.
Long Canyon area. b. Lower Bus Canyon area. c. Simi Arroyo-Peppertree Lane area.

900 ft. (274 m) north and 300 ft. (91 m) west of SE comer
of section 30, T 3 N, R 17 W. Bed is the same one exposed
at CSUN localities 4691 and 511. [Shallow-marine (trans-
gressive) facies.]

458. At elevation of 1300 ft. along south side of a ranch
road that leads up south side of Chivo Canyon, 1550 ft. (472
m) north and 400 ft. (122 m) east of SW comer of section
29, T 3 N, R 1 7 W. Locality is in the “Stewart bed." [Shallow-
marine (transgressive) facies.]

459. At elevation of 1400 ft. on north side of Chivo Can-
yon, 1500 ft. (457 m) north and 400 ft. (122 m) west of SE
comer of section 30, T 3 N, R 17 W. Locality is in the
“Stewart bed.” [Shallow-marine (transgressive) facies.]

460. At elevation of 2150 ft. on a hillside just north of a
ranch road, 6700 ft. (2042 m) N67°E of NE corner of section
29, T 3 N, R 17 W. [Shallow-marine (transgressive) facies.]

462. At elevation of 1225 ft. on a hillside just north of an
oilfield road, 2250 ft. (686 m) south and 2675 ft. (815 m)
west of NE comer of section 36, T 3 N, R 18 W. Locality is
near the Llajas-Sespe contact. [Shallow-marine (regressive)
facies.]

463. At elevation of 1650 ft. in streambed. Las Llajas
Canyon, 3725 ft. (1 1 35 m) N89°E of SE comer of section 29,
T 3 N, R 17 W. [Shallow-marine (transgressive) facies.]

465. At elevation of 2100 ft. on south side of a hill, 5480
ft. (1670 m) N70°E of NE comer of section 29, T 3 N, R 1 7
W. [Shallow-marine (transgressive) facies.]

466. At elevation of 2 1 50 ft. along east side of a hill, 5700
ft. (1738 m) N69°E of NE comer of section 29, T 3 N, R 17
W. [Shallow-marine (transgressive) facies.]

467. At elevation of 1 900 ft. on west side of stream bank
of El Toro Canyon, 7900 ft. (2408 m) N69°W of SE comer
of section 23, T 3 N, R 1 7 W. Locality is in the Santa Susana
quadrangle, but the section comer is in the Oat Mountain
quadrangle. [Shallow-marine (transgressive) facies.]

468a. At elevation of 1975 ft. on north side of stream bank
of Las Llajas Canyon, 6675 ft. (2034 m) N69°W of SE comer
of section 23, T 3 N, R 1 7 W. Locality is in the Santa Susana
quadrangle, but the section comer is in the Oat Mountain
quadrangle. Bed is the same one exposed at CSUN locality
515. [Shallow-marine (transgressive) facies.]

468b. Bed 1 0 ft. (3 m) stratigraphically above locality 468a.

60 Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks

Bed is the same one exposed at CSUN locality 516. [Shallow-
marine (transgressive) facies.]

469a. At elevation of 1400 ft. on north side of a small
canyon branching west from Chivo Canyon, 100 ft. (31 m)
north and 1650 ft. (503 m) west of SE comer of section 30,
T 3 N, R 17 W. Locality is at the type section of the Llajas
Formation, 144 ft. (44 m) above the base of the formation.
[Interfingering coastal alluvial-fan facies and shallow-marine
(transgressive) facies.]

469b. Bed 26 ft. (8 m) stratigraphically above locality 469a,
on a traverse bearing N10°W from locality 469a. [Interfin-
gering coastal alluvial-fan facies and shallow-marine (trans-
gressive) facies.]

469c. Bed 1 3 ft. (4 m) stratigraphically above locality 469b,
on a traverse bearing N10°W from locality 469b. [Interfin-
gering coastal alluvial-fan facies and shallow-marine (trans-
gressive) facies.]

469d. Bed 1 3 ft. (4 m) stratigraphically above locality 469c,
on a traverse bearing N10°W from locality 469c. [Interfin-
gering coastal alluvial-fan facies and shallow-marine (trans-
gressive) facies.]

469e. Bed 20 ft. (6 m) stratigraphically above locality 469d,
on a traverse bearing N10°W from locality 469d. [Shallow-
marine (transgressive) facies.]

469f. Bed 6.5 ft. (2 m) stratigraphically above locality 469e,
on a traverse bearing N10°W from locality 469e. [Shallow-
marine (transgressive) facies.]

469g. Bed 6.5 ft. (2 m) stratigraphically above locality 469f,
on a traverse bearing N10°W from locality 469f. [Shallow-
marine (transgressive) facies.]

469h. Bed 1 6 ft. (5 m) stratigraphically above locality 469g,
on a traverse bearing N10°W from locality 469g. [Shallow-
marine (transgressive) facies.]

4691. Bed 9 ft. (3 m) stratigraphically above locality 469h,
on a traverse bearing N10°W from locality 469h. [Shallow-
marine (transgressive) facies.]

469j. Bed 75 ft. (19 m) stratigraphically above locality
469g, on a traverse bearing N10°W from locality 469g. [Shal-
low-marine (transgressive) facies.]

469k. Bed 20 ft. (6 m) stratigraphically above locality 469g,
on a traverse bearing N10°W from locality 469j. [Shallow-
marine (transgressive) facies.]

4691. Bed 13 ft. (4 m) stratigraphically above locality 469k,
on a traverse bearing N55°W from locality 469k. Bed is the
same one exposed at CSUN localities 457 and 511. [Shallow-
marine (transgressive) facies.]

470a. At elevation of 1550 ft. on south side of an amphi-
theater on west side of Chivo Canyon, 500 ft. (152 m) north
and 2100 ft. (640 m) west of SE comer of section 30, T 3 N,
R 17 W. Locality is at the type section of the Llajas For-
mation, 266 ft. (81 m) stratigraphically above locality 4691.
[Shallow-marine (transgressive) facies.]

470b. Bed 62 ft. (19 m) stratigraphically above locality
470a, on a traverse bearing N60°W from locality 470a. [Shal-
low-marine (transgressive) facies.]

470c. Bed 33 ft. (10 m) stratigraphically above locality
470b, on a traverse bearing N60°W from locality 470b. [Shal-
low-marine (transgressive) facies.]

471a. At elevation of 1450 ft. on west side of an amphi-
theater on west side of Chivo Canyon, 600 ft. (183 m) north
and 2125 ft. (655 m) west of SE comer of section 30, T 3 N,
R 17 W. Locality is at the type section of the Llajas For-
mation, 125 ft. (38 m) stratigraphically above locality 470c.
[Shallow-marine (transgressive) facies.]

471b. Bed 144 ft. (44 m) stratigraphically above locality
471a, on a traverse bearing N27°W from locality 471a. [Shal-
low-marine (transgressive) facies.]

471c. Bed 20 ft. (6 m) stratigraphically above locality 471b,
on a traverse bearing N27°W from locality 471b. [Shallow-
marine (transgressive) facies.]

47!d. Bed 39 ft. (12 m) stratigraphically above locality
47 lc, on a traverse bearing N27°W from locality 471c. [Shal-
low-marine (transgressive) facies.]

471e. Bed 79 ft. (24 m) stratigraphically above locality
47 Id, on a traverse bearing N27°W from locality 47 Id. [Shal-
low-marine (transgressive) facies.]

472. At elevation of 1600 ft. on a ridge, 1000 ft. (305 m)
north and 2300 ft. (701 m) west of SE corner of section 30,
T 3 N, R 1 7 W. Locality is at the type section of the Llajas
Formation and in the “Stewart bed,” 46 ft. (14 m) strati-
graphically above locality 47 le. [Shallow-marine (transgres-
sive) facies.]

473. At elevation of 1650 ft. on a ridge, 1050 ft. (320 m)
north and 2300 ft. (701 m) west of SE corner of section 30,
T 3 N, R 1 7 W. Locality is at the type section of the Llajas
Formation, 46 ft. (14 m) stratigraphically above locality 472.
The bed that this locality occurs in probably represents a
shallow-marine (transgressive) facies bed that interfingers with
the outer shelf and slope facies.

475. At elevation of 1625 ft. on a ridge, 1300 ft. (396 m)
north and 3100 ft. (945 m) west of SE corner of section 30,
T 3 N, R 1 7 W. Locality is at the type section of the Llajas
Formation, approximately 446 ft. (136 m) stratigraphically
above locality 473. [Shallow-marine (regressive) facies.]

476. Bed 13 ft. (4 m) stratigraphically above locality 475,
on a traverse bearing N15°W from locality 475. [Shallow-
marine (regressive) facies.]

477. At elevation of 1625 ft. on a ridge, 1200 ft. (366 m)
north and 1700 ft. (518 m) west of SE comer of section 30,
T 3 N, R 17 W. Locality is in the “Stewart bed.” [Shallow-
marine (transgressive) facies.]

478. At elevation of 1735 ft. just below a saddle along a
ridge, 680 ft. (270 m) north and 2670 ft. (814 m) west of SE
comer of section 30, T 3 N, R 1 7 W. Locality is in the
“Stewart bed.” [Shallow-marine (transgressive) facies.]

479. At elevation of 1775 ft. just below a saddle along a
ridge, 650 ft. (198 m) north and 2700 ft. (823 m) west of SE
comer of section 30, T 3 N, R 17 W. Locality is in the
“Stewart bed.” [Shallow-marine (transgressive) facies.]

480. At elevation of 1400 ft. near a canyon bottom, 200
ft. (61 m) north and 3400 ft. (1036 m) west of SE comer of
section 30, T 3 N, R 17 W. Locality is in the “Stewart bed.”
[Shallow-marine (transgressive) facies.]

481. At elevation of 1500 ft. on a ridge, 325 ft. (99 m)
south and 1800 ft. (549 m) east of NE comer of section 31,
T 3 N, R 1 7 W. [Shallow-marine (transgressive) facies.]

Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks 61

482. At elevation of 1435 ft. in a roadcut, 1 100 ft. (335
m) south and 300 ft. (91 m) west of NE corner of section 36,
T 3 N, R 18 W. [Outer shelf and slope facies.]

483. At elevation of 1525 ft. on west side of a ridge, 925
ft. (282 m) south and 175 ft. (53 m) west of NE comer of
section 36, T 3 N, R 18 W. [Outer shelf and slope facies.]

484. At elevation of 2340 ft. on a ridge, 6000 ft. (1829 m)
N68°W of SE comer of section 26, T 3 N, R 17 W. Locality
is in the Santa Susana quadrangle, but the section corner is
in the Oat Mountain quadrangle. [Shallow-marine (trans-
gressive) facies.]

485. Bed 1 35 ft. (4 1 m) stratigraphically below locality 484,
on a traverse bearing S23°E from locality 484. [Shallow-
marine (transgressive) facies.]

486. Bed 69 ft. (21 m) stratigraphically below locality 485,
on a traverse bearing S23°E from locality 485. [Shallow-
marine (transgressive) facies.]

487. Bed 1 74 ft. (53 m) stratigraphically below locality 486,
on a traverse bearing S23°E from locality 486. [Shallow-
marine (transgressive) facies.]

488. At elevation of 2175 ft. on a ridge, 5650 ft. (1722 m)
N64°W of SE comer of section 26, T 3 N, R 17 W. Locality
is in the Santa Susana quadrangle, but the section comer is
in the Oat Mountain quadrangle. Locality is in the “Stewart
bed.” [Shallow-marine (transgressive) facies.]

489. At elevation of 1250 ft. on east side of a canyon, 125
ft. (38 m) south of NW corner of section 31, T 3 N, R 17
W. [Shallow-marine (regressive) facies.]

490. At elevation of 1900 ft. in a saddle along a ridge, 600
ft. (183 m) north and 800 ft. (244 m) east of SE corner of
section 29, T 3 N, R 17 W. [Shallow-marine (transgressive)
facies.]

491. At elevation of 1 840 ft. on east side of a small canyon,
500 ft. (145 m) north and 1 100 ft. (320 m) east of SE comer
of section 29, T 3 N, R 1 7 W. [Shallow-marine (transgressive)
facies.]

492. At elevation of 2230 ft. in a saddle along a ridge, 1 700
ft. (518 m) north and 3225 ft. (983 m) east of SE comer of
section 29, T 3 N, R 1 7 W. [Shallow-marine (transgressive)
facies.]

493. At elevation of 2225 ft. in saddle along a ridge, 1825
ft. (556 m) north and 3450 ft. (1052 m) east of SE comer of
section 29, T 3 N, R 17 W. Locality is in the “Stewart bed.”
[Shallow-marine (transgressive) facies.]

494. At elevation of 2050 ft. on a small cliff, 5350 ft. (1630
m) N62°W of SE comer of section 26, T 3 N, R 1 7 W. Locality
is in the Santa Susana quadrangle, but the section corner is
in the Oat Mountain quadrangle. Locality is in the “Stewart
bed.” [Shallow-marine (transgressive) facies.]

495. At elevation of 2050 ft. on a small cliff, 4725 ft. (1440
m) N60°W of SE comer of section 26, T 3 N, R 1 7 W. Locality
is in the Santa Susana quadrangle, but the section comer is
in the Oat Mountain quadrangle. Locality is in the “Stewart
bed.” [Shallow-marine (transgressive) facies.]

496. At elevation of 2050 ft. on a hillside, 4100 ft. (1250
m)N59°W ofSE comer of section 26, T 3 N, R 17 W. Locality
is in the Santa Susana quadrangle, but the section comer is

in the Oat Mountain quadrangle. Locality is in the “Stewart
bed.” [Shallow-marine (transgressive) facies.]

497. At elevation of 2185 ft. on south side of ridge, 3685
ft. (1 123 m) N58°W of SE comer of section 26, T 3 N, R 17
W. Locality is in the Santa Susana quadrangle, but the section
comer is in the Oat Mountain quadrangle. Locality is in the
“Stewart bed.” [Shallow-marine (transgressive) facies.]

498. At elevation of 1850 ft. on west side of stream bank
of Devil Canyon, 1550 ft. (472 m) south and 600 ft. (183 m)
west of NE comer of section 26, T 3 N, R 1 7 W of the USGS

7.5-minute topographic quadrangle of Oat Mountain, Cali-
fornia (1952), photorevised 1969. [Shallow-marine (trans-
gressive) facies.]

499. At elevation of 1850 ft. on west side of stream bank
of Devil Canyon, 1400 ft. (427 m) south and 630 ft. (192 m)
west of NE comer of section 26, T 3 N, R 17 W of the USGS

7.5-minute topographic quadrangle of Oat Mountain, Cali-
fornia (1952), photorevised 1969. [Shallow-marine (trans-
gressive) facies.]

500. At elevation of 1875 ft. on east side of stream bank
of Devil Canyon, 800 ft. (244 m) south and 675 ft. (206 m)
west of NE comer of section 26, T 3 N, R 1 7 W of the USGS

7.5-minute topographic quadrangle of Oat Mountain, Cali-
fornia (1952), photorevised 1969. [Shallow-marine (trans-
gressive) facies.]

501. At elevation of 1500 ft. on east side of a small canyon
branching northwest of Las Llajas Canyon, 1400 ft. (427 m)
north and 1400 ft. (427 m) west of SE comer of section 29,
T 3 N, R 1 7 W. [Interfingering coastal alluvial-fan facies and
shallow-marine (transgressive) facies.]

502. At elevation of 2150 ft. on a ridge, 5700 ft. (1737 m)
N70°W of SE comer of section 26, T 3 N, R 17 W. Locality
is in the Santa Susana quadrangle, but the section comer is
in the Oat Mountain quadrangle. [Interfingering coastal al-
luvial-fan facies and shallow-marine (transgressive) facies.]

503. At elevation of 2160 ft. on a ridge, 5725 ft. (1745 m)
N69°W of SE comer of section 26, T 3 N, R 17 W. Locality
is in the Santa Susana quadrangle, but the section comer is
in the Oat Mountain quadrangle. [Interfingering coastal al-
luvial-fan facies and shallow-marine (transgressive) facies.]

504. At elevation of 2225 ft. on a ridge, 5750 ft. (1753 m)
N71°W of SE comer of section 26, T 3 N, R 17 W. Locality
is in the Santa Susana quadrangle, but the section comer is
in the Oat Mountain quadrangle. [Shallow-marine (trans-
gressive) facies.]

505. Bed 98 ft. (30 m) stratigraphically above locality 504,
on a traverse bearing N23°W from locality 504. [Shallow-
marine (transgressive) facies.]

506. At elevation of 2280 ft. on a ridge, 5600 ft. (1706 m)
N65°W of SE comer of section 26, T 3 N, R 17 W. Locality
is in the Santa Susana quadrangle, but the section comer is
in the Oat Mountain quadrangle. [Shallow-marine (trans-
gressive) facies.]

507. Bed 33 ft. (10 m) stratigraphically above locality 506,
on a traverse bearing N6°E from locality 506. [Shallow-ma-
rine (transgressive) facies.]

508. Bed 6.5 ft. (2 m) stratigraphically above locality 507,

62 Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks

on a traverse bearing N6°E from locality 507. [Shallow-ma-
rine (transgressive) facies.]

509. Bed 49 ft. (1 5 m) stratigraphically above locality 508,
on a traverse bearing N6°E from locality 508. [Shallow-ma-
rine (transgressive) facies.]

510. Bed 108 ft. (33 m) stratigraphically above locality
509, on a traverse bearing N6°E from locality 509. [Shallow-
marine (transgressive) facies.]

511. At elevation of 1225 ft. on north sideofChivo Canyon
near a major fork in Chivo Canyon, 800 ft. (244 m) north
and 800 ft. (244 m) west of SE corner of section 30, T 3 N,
R 17 W. Bed is the same one exposed at CSUN localities
457 and 4691. [Shallow-marine (transgressive) facies.]

512a. At elevation of 1 275 ft. on east side of a small canyon
branching north of Chivo Canyon, 2275 ft. (693 m) south
and 525 ft. (160 m) east of NW corner of section 29, T 3 N,
R 17 W. [Shallow-marine (regressive) facies.]

512b. Bed 20 ft. (6 m) stratigraphically above locality 5 1 2a,
on a traverse bearing N35°W from locality 512a. [Shallow-
marine (regressive) facies.]

512c. Bed 16 ft. (5 m) stratigraphically above locality 512b,
on a traverse bearing N35°W from locality 512b. [Shallow-
marine (regressive) facies.]

51 2d. Bed 30 ft. (9 m) stratigraphically above locality 512b,
on a traverse bearing N35°W from locality 512b. [Shallow-
marine (regressive) facies.]

513. At elevation of 1870 ft. (570 m) in the streambed of
Las Llajas Canyon, 6825 ft. (2080 m) N71°W of SE corner
of section 23, T 3 N, R 1 7 W. Locality is in the Santa Susana
quadrangle, but the section comer is in the Oat Mountain
quadrangle. [Shallow-marine (transgressive) facies.]

514. At elevation of 1875 ft. on north side of stream bank
of Las Llajas Canyon, 6800 ft. (2073 m) N71°W ofSE corner
of section 23, T 3 N, R 1 7 W. Locality is in the Santa Susana
quadrangle, but the section comer is in the Oat Mountain
quadrangle. [Shallow-marine (transgressive) facies.]

515. At elevation of 1925 ft. (587 m) on the north side of
stream bank of Las Llajas Canyon, 6750 ft. (2057 m) N69°W
of SE comer of section 23, T 3 N, R 17 W. Locality is in the
Santa Susana quadrangle, but the section corner is in the Oat
Mountain quadrangle. Bed is the same one exposed at CSUN
locality 468a. [Shallow-marine (transgressive) facies.]

516. At elevation of 1975 ft. (602 m) on north side of
stream bank of Las Llajas Canyon, 6500 ft. (1981 m) N67°W
of SE comer of section 23, T 3 N, R 1 7 W. Locality is in the
Santa Susana quadrangle, but the section corner is in the Oat
Mountain quadrangle. Bed is the same one exposed at CSUN
locality 468b. [Shallow-marine (transgressive) facies.]

517. At elevation of 1950 ft. in a small canyon bottom,
between the letters “n” and “y” in the word “Canyon,” 5100
ft. (1555 m) N47°E of NE comer of section 29, T 3 N, R 17
W. [Shallow-marine (transgressive) facies.]

518. At elevation of 2125 ft. on a small ridge, 5475 ft.
(1669 m) N40°E of NE comer of section 29, T 3 N, R 17 W.
[Shallow-marine (transgressive) facies.]

520. At elevation of 2070 ft., 6500 ft. (1981 m) N78°E of
NE comer of section 29, T 3 N, R 17 W. [Shallow-marine
(transgressive) facies.]

521. At elevation of 2 1 1 5 ft. on a ridge, 6360 ft. ( 1 938 m)
N88°E of NE comer of section 29, T 3 N, R 1 7 W. [Shallow-
marine (transgressive) facies.]

522. Bed 131 ft. (40 m) stratigraphically above CSUN
locality 37 1 , on a traverse bearing S 1 4°W from locality 371.
[Shallow-marine (transgressive) facies.]

523. At elevation of 2100 ft. on west side of a canyon
branching south of Las Llajas Canyon, 5275 ft. (1608 m)
N70°W of SE comer of section 23, T 3 N, R 17 W. Locality
is in the Santa Susana quadrangle, but the section comer is
in the Oat Mountain quadrangle. [Shallow-marine (trans-
gressive) facies.]

524. At elevation of 2035 ft. on ridge along south side of
Las Llajas Canyon, 6375 ft. (1943 m) N72°W of SE corner
of section 23, T 3 N, R 1 7 W. Locality is in the Santa Susana
quadrangle, but the section corner is in the Oat Mountain
quadrangle. [Shallow-marine (transgressive) facies.]

525. At elevation of 2050 ft. on a hillside, 6325 ft. (1928
m) N74°W of SE comer of section 23, T 3 N, R 1 7 W. Locality
is in the Santa Susana quadrangle, but the section comer is
in the Oat Mountain quadrangle. [Shallow-marine (trans-
gressive) facies.]

526. At elevation of 2055 ft. on a ridge, 2290 ft. (698 m)
N67°W of SE comer of section 26, T 3 N, R 17 W. Locality
is in the Santa Susana quadrangle, but the section corner is
in the Oat Mountain quadrangle. [Shallow-marine (trans-
gressive) facies.]

527a. At elevation of 2075 ft. on a ridge, 2325 ft. (709 m)
N65°W of SE comer of section 26, T 3 N, R 17 W. Locality
is in the Santa Susana quadrangle, but the section corner is
in the Oat Mountain quadrangle. [Shallow-marine (trans-
gressive) facies.]

527b. Bed 1 3 ft. (4 m) stratigraphically above locality 527a,
on a traverse bearing due north of locality 527a. [Shallow-
marine (transgressive) facies.]

528. At elevation of 2185 ft. on a ridge, 1400 ft. (427 m)
north and 2250 ft. (686 m) west of SE comer of section 26,
T 3 N, R 17 W. Locality is in the Santa Susana quadrangle,
but the section corner is in the Oat Mountain quadrangle.
[Shallow-marine (transgressive) facies.]

529a. At elevation of 1400 ft. on east side of stream bank
of Las Llajas Canyon, 1 300 ft. (396 m) north and 350 ft. (107
m) east of SE comer of section 29, T 3 N, R 1 7 W. [Shallow-
marine (transgressive) facies.]

529b. Bed 26.5 ft. (7 m) stratigraphically above locality
529a, on a traverse bearing N27°E of locality 529a. [Shallow-
marine (transgressive) facies.]

537. At elevation of 1 500 ft. on west side of an amphi-
theater on west side of Chivo Canyon, 790 ft. (241 m) north
and 2250 ft. (686 m) west of SE comer of section 30, T 3 N,
R 17 W. Locality is at the type section of the Llajas Lor-
mation. [Shallow-marine (transgressive) facies.]

538. At elevation of 1 500 ft. on a ridge on west side of Las
Llajas Canyon, 1500 ft. (457 m) north and 350 ft. (107 m)
west of SE comer of section 29, T 3 N, R 17 W. [Shallow-
marine (transgressive) facies.]

539. Bed 6 ft. (1.8 m) stratigraphically above locality 538,

Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks 63

on a traverse bearing N5°E of locality 538. [Shallow-marine
(transgressive) facies.]

540. Bed 38 ft. (11.5 m) stratigraphically below CSUN
locality 486, on a traverse bearing S28°E from locality 486.
[Shallow-marine (transgressive) facies.]

541. At elevation of 1 325 ft. on north bank of stream bank
of Chivo Canyon, 2800 ft. (853 m) north and 1 175 ft. (358
m) east of SE corner of section 29, T 3 N, R 1 7 W. [Outer
shelf and slope channel facies.]

542. At elevation of 1315 ft. on ridge near mouth of Las
Llajas Canyon, 2200 ft. (671 m) north and 800 ft. (244 m)
east of SW comer of section 32, T 3 N, R 1 7 W. [Interfingering
coastal alluvial-fan facies and shallow-marine (transgressive)
facies.]

543. At elevation of 1460 ft. on divide between Bus and
Montgomery Canyons, 600 ft. (183 m) south and 1500 ft.
(457 m) east of NW corner of section 28, T 2 N, R 18 W of
the USGS 7.5-minute topographic quadrangle of Thousand
Oaks, California (1950), photorevised 1967. [Shallow-ma-
rine (transgressive) facies.]

544. At elevation of 1 230 ft. on a small ridge, 1 025 ft. (3 1 2
m) north and 415 ft. (127 m) west of SE corner of section
19, T 2 N, R 18 W of the USGS 7.5-minute topographic
quadrangle of Thousand Oaks, California (1950), photore-
vised 1967. [Shallow-marine (regressive) facies.]

545. At elevation of 1 120 ft. on east side of Bus Canyon,
580 ft. (1 77 m) south and 14 10 ft. (430 m) west of NE comer
of section 28, T 2 N, R 18 W of the USGS 7.5-minute
topographic quadrangle of Thousand Oaks, California (1950),
photorevised 1967. [Interfingering coastal alluvial-fan facies
and shallow-marine (transgressive) facies.]

546. At elevation of 1 500 ft. on east side of divide between
Bus and Trough Canyons, 700 ft. (213 m) south and 200 ft.
(61 m) east of NW corner of section 27, T 2 N, R 18 W of
the USGS 7.5-minute topographic quadrangle of Thousand
Oaks, California (1950), photorevised 1967. [Shallow-ma-
rine (transgressive) facies.]

547. At elevation of 1 0 1 0 ft. on east side of a roadcut, 650
ft. (198 m) south of NE corner of section 21, T 2 N, R 18
W of the USGS 7.5-minute topographic quadrangle of Thou-
sand Oaks, California (1950), photorevised 1967. [Shallow-
marine (regressive) facies.]

548. At elevation of 947 ft. on south side of Simi Arroyo
just above the streambed, 1 100 ft. (335 m) north and 1750
ft. (553 m) west of SE corner of section 12, T 2 N, R 18 W.
Locality is equivalent to UCLA locality 5837. [Shallow-ma-
rine (transgressive) facies.]

700. At elevation of 1 100 ft. along east side of road in Bus
Canyon, 750 ft. (229 m) south and 1500 ft. (457 m) west of
NE comer of section 28, T 2 N, R 18 W of the USGS 7.5-
minute topographic quadrangle of Thousand Oaks, Califor-
nia (1950), photorevised 1967. [Interfingering coastal allu-
vial-fan facies and shallow-marine (transgressive) facies.]

701. At elevation of 1 160 ft. along east side of Bus Canyon,
800 ft. (244 m) south and 1400 ft. (427 m) west of NE corner
of section 28, T 2 N, R 18 W of the USGS 7.5-minute
topographic quadrangle of Thousand Oaks, California (1950),
photorevised 1967. [Interfingering coastal alluvial-fan facies
and shallow-marine (transgressive) facies.]

702. At elevation of 1420 ft. along a ridge, 900 ft. (274 m)
south and 680 ft. (207 m) west of NE corner of section 28,
T 2 N, R 18 W of the USGS 7.5-minute topographic quad-
rangle of Thousand Oaks, California (1950), photorevised
1967. Locality is equivalent to UCMP locality 7193 and
UCLA locality 7070. [Interfingering coastal alluvial-fan fa-
cies and shallow-marine (transgressive) facies.]

703. At elevation of 1430 ft. along a ridge, 780 ft. (238 m)
south and 580 ft. (177 m) west of NE corner of section 28,
T 2 N, R 18 W of the USGS 7.5-minute topographic quad-
rangle of Thousand Oaks, California (1950), photorevised
1 967. Locality is a few meters stratigraphically above locality
702. [Lowermost part of shallow-marine (transgressive) fa-
cies.]

704. At elevation of 1 320 ft. along divide between Bus and
Trough Canyons, 180 ft. (55 m) south and 90 ft. (27 m) west
of NE comer of section 28 of the USGS 7.5-minute topo-
graphic quadrangle of Thousand Oaks, California (1950),
photorevised 1967. [Shallow-marine (transgressive) facies.]

705. At elevation of 1240 ft. along a dip-slope just south
of the mouth of Las Llajas Canyon, 2450 ft. (747 m) north
and 450 ft. (137 m) east of SW corner of section 32, T 3 N,
R 17 W. [Interfingering coastal alluvial-fan facies and shal-
low-marine (transgressive) facies.]

706. At elevation of 1250 ft. along a small ridge on east
side of Chivo Canyon where it bends sharply to the northeast,
700 ft. (213 m) north and 590 ft. (180 m) west of SE corner
of section 30, T 3 N, R 1 7 W. [Interfingering coastal alluvial-
fan facies and shallow-marine (transgressive) facies.]

CAS LOCALITIES

25. On east bank of Little River at its confluence with
Umpqua River, near center of section 19, T 26 S, R 3 W.
Roseburg quadrangle, Douglas County, Oregon.

364. Exact location unknown. Aliso Creek (now known as
Devil Canyon), section 25, T 3 N, R 17 W, near Chatsworth,
California, Los Angeles County.

372. Exact location unknown. Aliso Canyon (now known
as Devil Canyon), section 23, T 3 N, R 17 W, near Chats-
worth, California, Los Angeles County.

393. Devil Canyon, SE 'A of the NW 'A of section 26, T 3
N, R 17 W, Santa Susana quadrangle, California.

711. East side of Grapevine Creek near point where it
enters valley floor, Tejon quadrangle, Kern County, Califor-
nia.

792. West side of Tecuya Creek, about 1 mile south of
where stream flows out on valley floor, Tejon quadrangle,
Kem County, California.

HERMAN SCHYMICZEK LOCALITY

HS-A-1 1(4). Bed 92 ft. (28 m) stratigraphically above CSUN
locality 4691, on a traverse bearing due north from locality
4691.

LACMIP LOCALITY

461-B. On the northern slope of a small canyon intersecting
Las Llajas Canyon from the east. It is 200 ft. (61 m) from

64 Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks

the top of the Llajas Formation, Las Llajas Canyon, Santa
Susana Mountains, Ventura County, California.

SU LOCALITY

2696. Chivo Canyon, 5 km N20°E of Bench Mark 961 at
Santa Susana, Santa Susana quadrangle, Ventura County,
California.

UCMP LOCALITIES

337. About 5 km south of Martinez, on the east side of
the road to Walnut Creek, Concord quadrangle. Contra Costa
County, California.

452. On road in Grapevine Creek, SW ‘A of the SE ‘A of
section 20, T 10 N, R 19 W, Tejon quadrangle, Kern County,
California.

672. South portion of crest of Parson’s Peak, SE ‘A of the
NW 'A of section 24, T 18 S, R 14 E, Coalinga quadrangle,
Fresno County, California.

1427. SW ‘A of section 1 1, NE of Wall Point, south side
of Mount Diablo, California.

1817. Opposite the place where Urruttia Canyon enters
Salt Creek, 100 ft. (30 m) up fourth small draw from west
end of ridge, SW ‘A of the NW ‘A of section 15, T 18 S, R
14 E, Coalinga quadrangle, Fresno County, California.

1853. Marysville Buttes, N Vi of section 28, T 16 N, R 1
E, Marysville Buttes quadrangle, Sutter County, California.

2226. Longitude 1 17°14'W, latitude 33°50'N, SE of Sole-
dad Mountain, north of Ladrillo Station, Southern Pacific
Railroad, Rose Canyon, La Jolla quadrangle, San Diego
County, California.

2287. West side of Domengine Canyon, SW comer of the
SW 'A of the SE ‘A of section 29, T 18 S, R 15 E, Fresno
County, California.

2295. Longitude 120. 1°W, latitude 36.1°N, near Domen-
gine Ranch, Fresno County, California.

3296. Exact location unknown. Aliso Canyon (now known
as Devil Canyon), SE ‘A of the SE ‘A of section 23, T 3 N, R
1 7 W, Santa Susana quadrangle, Ventura County, California.

3304. Exact location unknown. Aliso Canyon (now known
as Devil Canyon, section 36, T 3 N, R 17 W, Santa Susana
quadrangle, Los Angeles County, California.

3310. Exact location unknown. Probably Simi Hills, Santa
Susana quadrangle, California.

3976. Seven-eighths of a mile (1.4 km) north of Ladrillo
Station, Southern Pacific Railroad, in first canyon east of
Rose Canyon, elevation 259 ft.. La Jolla quadrangle, San
Diego County, California.

3981. At 15 m above high-tide level in small gully 0.4 km
south of mouth of Soledad Valley, La Jolla quadrangle, San
Diego County, California.

3989. 0.2 mi. (0.3 km) north of the top of the “y” of
“Tecolote Valley,” on the east side of the valley, elevation
100 ft., La Jolla quadrangle, San Diego County, California.

3990. On the east side of canyon in bottom of Rose Creek,
0.3 mi. (0.5 km) east of “t” of “Soledad Mountain,” La Jolla
quadrangle, San Diego County, California.

3993. In bottom of Rose Creek where creek makes a strong
bend to west, 0.2 mi. (0.3 km) south of Bench Mark 176, 2

mi. (3.2 km) east of La Jolla, La Jolla quadrangle, San Diego
County, California.

4169. About 500 ft. (152 m) east of ranch house in Big Tar
Canyon on east line of section 18, near point where road
crosses creek, Garza Peak quadrangle. Kings County, Cali-
fornia.

4170. On west side of Big Tar Canyon, where it crosses
the Eocene section, Garza Peak quadrangle, Kings County,
California.

4175. No locality data available. Probably Domengine
Ranch area, north of Coalinga, Domengine Ranch quadran-
gle, Fresno County, California.

5062. In sea cliff south of mouth of Soledad Valley, due
west of midpoint between “P” and “u” of Pueblo,” La Jolla
quadrangle, San Diego County, California.

5069. 0.4 inches SSE of the “a” in “Morena” in a branch
canyon of Tecolote Valley to the north, elevation 1 50 ft., La
Jolla quadrangle, San Diego County, California.

5080. North side of Soledad Valley, at McGonigle Canyon,
NE ‘A of section 36, T 14 S, R 4 W, La Jolla quadrangle, San
Diego County, California.

5085. 2.62 inches north of the top of the “S” of “Soledad
Mountain,” on the north side of the creek, on a small ridge
formed by the creek and sea cliff, elevation 75 ft.. La Jolla
quadrangle, San Diego County, California.

5089. 300 ft. (91 m) north of the Scripps Institution pier,
in the conglomerate above the mudstone. In the sea cliff,
elevation 10 ft., La Jolla quadrangle, San Diego County,
California.

7000. Exact location unknown. Las Llajas Canyon, in first
canyon on north side of road, Santa Susana quadrangle, Cal-
ifornia.

7002. At elevation of 1750 ft., in road near top of last
western spur which extends northward out of Las Llajas Can-
yon. In shales striking north just south of east-west fault
which is made prominent by the Meganos Conglomerate
which parallels it on the north side, Santa Susana quadrangle,
Ventura County.

7003. At elevation of 1 760 ft. in sandstone exposed in deep
washout on south side of second large canyon which enters
Llajas Canyon from the east. Washout is about 100 yards
east of mouth of canyon. Fossils are found in ledge on west
side of washout and near top. Sandstone here dips with slope
of canyon side and from west wall of east-west fault which
crosses Llajas Canyon south of this locality, Santa Susana
quadrangle, Ventura County.

7004. About 300 ft. (91 m) east of locality 7003 in next
small canyon which enters Llajas Canyon from the east just
south of the most northerly extension of the 1 500-ft. contour,
Santa Susana quadrangle, Ventura County. Locality is equiv-
alent to CSUN locality 374.

7005. No locality data available. Probably Las Llajas Can-
yon area, Santa Susana quadrangle, California.

7015. About 5500 ft. (1676 m) S69°E of Bench Mark at
Santa Susana, Santa Susana quadrangle, Ventura County.

7182. Boulder from second draw on east side of Live Oak
Canyon, Tejon Ranch, Kern County, California.

7193. About 600 ft. (183 m) east of and apparently slightly
stratigraphically beneath locality 7194, which is along the

Contributions in Science, Number 350

Squires: Sinii Valley Eocene Mollusks 65

top of a 1400-ft. ridge extending NW from third main ridge
west of Runkle’s Ranch on the ridge on which there is an
abandoned well and road leading up to it, Simi Valley, Cal-
ifornia. Locality is equivalent to CSUN locality 702 and
UCLA locality 7070.

7195. In the creek bed about 60 ft. (18 m) north of the
second falls or 300 ft. (91 m) north of the mouth of the first
small draw which enters Las Llajas Canyon west of the point
where the Meganos Conglomerate crosses the road, Santa
Susana quadrangle, Ventura County, California.

7200. No locality data available. Probably Live Oak Can-
yon area, Tejon Ranch, Kern County, California.

A-661. On east bank of Little River between the highway
bridge and the first bend of the stream east of the junction
with the North Umpqua River, center of section 19, T 26 S,
R 3 W, Douglas County, Oregon.

A-819. Lowest reef bed on side of hill just east of and
above first saddle south of Big Tar Canyon, Garza Peak
quadrangle. Kings County, California.

A-993. Second draw past Marrland Canyon (now known
as Las Llajas Canyon) at second small falls up draw approx-
imately 600 ft. (183 m), Santa Susana quadrangle, Ventura
County, California.

A-994. About 225 to 300 ft. (69 to 9 1 m) down the canyon
from locality A-993 on west side of canyon about 60 ft. (18
m) from streambed, Santa Susana quadrangle, Ventura
County, California.

A-1003. Exact location unknown. Pine Canyon, Mount
Diablo, Contra Costa County, California.

A-1027. Valdes Ranch, on branch of Silver Creek, Valle-
citos, center of east part of SW *A of section 4, T 16 S, R 12
E. Approximately where 1 20°40' parallel crosses most north-
erly intermittent stream indicated on section 4, Panoche
quadrangle, Fresno County, California.

A-1219. Base of Domengine Formation on west side, near
top, of long ridge extending NW of 2126-ft. hill on line be-
tween sections 9 and 16, T 19 S, R 1 5 E, Domengine Ranch
quadrangle, California.

A-1280. Near center of north edge of section 20, on hill
immediately south of point where the Big Tar-McLure Val-
ley road crosses saddle at head of steam running into McLure
Valley, 45 ft. (14 m) below uppermost fossiliferous layer,
Garza Peak quadrangle. Kings County, California.

A-3042. On ridge in blue clay shale member above con-
glomerate member at base of Llajas Formation, center of SE
■A of the SE 'A of section 10, T 3 N, R 17 W, Santa Susana
quadrangle, Los Angeles County, California.

UCLA LOCALITY

6616. West end of ridge south of Las Llajas Canyon, 2525
ft. (770 m) north and 575 ft. (175 m) east of SW corner of
section 32, T 3 N, R 1 7 W, Santa Susana quadrangle, Ventura
County, California.

UCR LOCALITY

4865. In a roadcut on the southwest side of California State
Highway 78, 5.2 km SE of Vista and 4.8 km NW of San

Marcos, at longitude 117°12'39"W, latitude 33°09'47"N, in
northern San Diego County, California.

UO LOCALITY

144. About 2.5 mi. (4 km) up Fall Creek from Little River
on Mathews farm in NE 'A of section 6, T 27 S, R 3 W,
Douglas County, Oregon.

USGS LOCALITIES

4617. On SW flank of Reef Ridge, north of McLure Valley,
2‘A mi. (3.6 km) SSE of El Cerrito oil well, in section 27, T
23 S, R 1 7 E, Cholame quadrangle. Kings County, California.

4619. North of Coalinga, 15 mi. (24 km) SW of Domen-
gine’s Ranch, T 18 S, R 15 E, Coalinga quadrangle, Fresno
County, California.

UW LOCALITIES

329. On north bank of the Cowlitz River at bend 1.5 to
2.5 km east ofVader, section 28, T 1 N, R 2 W, Lewis County,
Washington.

358. Joice Station, ‘A mi. (0.4 km) east of Tongue Point
Railroad, Port Crescent, section 22, T 3 1 N, R 8 W, Clallam
County, Washington.

ACKNOWLEDGMENTS

Acknowledgment is made to the donors of The Petroleum
Research Fund, administered by the American Chemical So-
ciety, for support of this research under grant number PRF
1 1472-B2.

The following ranchers kindly permitted access to the field:

G. Haigh, W. Haigh, D. Poe, G. Boyle, and J. Runkle. Sincere
thanks are extended to the following California State Uni-
versity, Northridge, geology students for their assistance in
the field: D. Advocate, D. Carpenter, G. Davis, S. Drewry,
T. Santochi, M. Stettner, and G. Wood. G. Davis helped in
photographing the fossils.

The following people kindly provided the loan of speci-
mens: F. J. Collier, United States National Museum of Nat-
ural History; F. A. Jenkins, Loyola Marymount University;

H. A. Lowenstam, California Institute of Technology; E. Nes-
bitt and J. Peck, University of California, Berkeley; P. U.
Rodda, California Academy of Sciences; and L. R. Saul,
University of California, Los Angeles.

I am grateful to A. G. Beu (New Zealand Geological So-
ciety), C. R. Givens (Nicholls State University), and L. R.
Saul for their valuable comments on molluscan taxonomy
and identification. L. R. Saul also helped greatly in the doc-
umentation of the early Eocene molluscan fauna of the Llajas
Formation. B. J. Welton (Chevron Research, California)
kindly identified the shark teeth. M. Filewicz and M. Hill,
III (Union Oil of California) shared their calcareous nan-
nofossil data. D. R. Lindberg (University of California,
Berkeley) and T. Susuki (University of California, Los An-
geles) provided information as to the location of some fossil
localities. R. Simpson and E. C. Wilson (Los Angeles County
Museum) gave most helpful editorial comments. C. R. Giv-

66 Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks

ens and W. J. Zinsmeister (Purdue University) critically re-
viewed the manuscript, and their suggestions greatly im-
proved it.

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Accepted for publication 7 February 1984.

76 Contributions in Science, Number 350

Squires: Simi Valley Eocene Mollusks

.

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MOLLUSCAN PALEONTOLOGY AND BIOSTRATIGRAPHY OF THE
LOWER MIOCENE UPPER PART OF THE LINCOLN CREEK
FORMATION IN SOUTHW ESTERN W ASHINGTON

Ellen J. Moore

Contributions in Science, Number 351
Natural History Museum of Los Angeles County
13 July 1984

Vessels loading lumber at Knappton, Washington, in 1908. When it was founded in 1867, Knappton was called Cementville,
and concretionary rocks served as the raw material for the cement. (Photograph courtesy of Dale Estoos and David Lee Myers.)

ISSN 0459-8113

Natural History Museum of Los Angeles County
900 Exposition Boulevard
Los Angeles, California 90007

PREFACE

Knappton, Washington, on the Columbia River north of Astoria, Oregon, was named for
Jabez Burrell Knapp, who built a cement plant there in 1867, calling the town Cementville.
The plant included a large kiln and a cement-barrel factory, but was unable to continue
production for more than a few years because of lack of sufficient raw material: fossiliferous
concretions and concretionary layers. Knapp then built a sawmill, which supported the
company town for 70 years. In 1941, the planing mill caught fire and the blaze quickly
spread to other buildings. The wooden water main burned through, leaving the fire fighters
helpless, and the town was destroyed. Only a monument and pilings remain at the site to
reflect its past history.

A few fossil mollusks were collected from the Knappton area in the early 1900’s and
deposited in the National Museum of Natural History at Washington, D.C., and the Cali-
fornia Academy of Sciences at San Francisco. The small collections contain usually poorly
preserved material from both the upper part of the Lincoln Creek Formation and the over-
lying Astoria Formation, which were not differentiated at that time.

Controversy over the age of the Astoria Formation, whose type locality is just across the
river, began in 1865 and revolved especially around the presence of the fossil cephalopod
Aturia. This cephalopod also was known to occur in the upper part of the Lincoln Creek
Formation on the Washington side of the Columbia River, and casual references to Knappton
occasionally appeared in the scientific literature. It was not until 1979, however, that a
detailed geologic map of the Knappton area was prepared by Ray E. Wells and issued by
the U.S. Geological Survey.

The first major collections of fossils from Knappton were made by James L. Goedert and
his wife, Gail. The presence of vertebrate fossils, chiefly whales, at Knappton led Goedert
to contact the Los Angeles County Museum of Natural History in 1979. His offer to donate
the collections to the Los Angeles Museum, and the recognition of the importance of the
invertebrate fossils in those collections by Edward C. Wilson, Curator of Invertebrate Pa-
leontology, has led to publications on the barnacles by Victor A. Zullo, the sponges by J.
Keith Rigby and D.E. Jenkins, and the mollusks described herein.

This paper is dedicated to James L. Goedert whose generosity has made it possible for
specialists to study the remarkable Knappton fossils.

CONTENTS

ABSTRACT 1

INTRODUCTION 1

PREVIOUS WORK 2

CHRONOSTRATIGRAPHY 3

PALEOECOLOGY 6

MOLLUSCAN PALEONTOLOGY 8

Abbreviations 9

Gastropods 9

Trochidae 9

Epitoniidae 10

Naticidae 10

Cassididae 1 0

Buccinidae? 12

Neptuneidae 12

Fusinidae 16

Volutidae 16

Turridae 24

Acteonidae 25

Pelecypods 25

Nuculidae 25

Nuculanidae 25

Solemyidae 25

Limopsidae 25

Mytilidae 26

I.imidae 26

Ostreidae 28

Lucinidae 28

Thyasiridae 28

Cardiidae 30

Tellinidae 30

Periplomatidae 30

Teredinidae 30

Scaphopod 34

Dentaliidae 34

Cephalopods 34

Aturidae 34

Sepiidae? 34

FOSSILS OTHER THAN MOLLUSKS 34

Sponges 34

Corals 36

Brachiopod 36

Echinoids 36

LOCALITIES 36

ACKNOWLEDGMENTS 39

LITERATURE CITED 39

MOLLUSCAN PALEONTOLOGY AND BIOSTRATIGRAPHY OF THE
LOWER MIOCENE UPPER PART OF THE LINCOLN CREEK
FORMATION IN SOUTHWESTERN WASHINGTON

Ellen J. Moore1

ABSTRACT. The uppermost part of the Lincoln Creek Formation
in the Knappton area of southwestern Washington is assigned to the
lower Miocene upper part of the Juanian Molluscan Stage (=Sauce-
sian Foraminiferal Stage) and the Liracassis apta Molluscan Zone.
This part of the Lincoln Creek Formation is correlated with the
upper part of the Pysht Formation of the Twin River Group in
northwestern Washington.

Fossils, invertebrates and vertebrates, are preserved in concretions
that erode out of landslides and accumulate as float along the Co-
lumbia River. The molluscan fauna of the upper part of the Lincoln
Creek Formation consists of 33 species in 30 genera, including five
newly described species in the genera “ Bathybembix ,” Ancistrolepis,
Musashia, Modiolus, and Acesta. Eastern Pacific species in the genus
Musashia are reviewed. The fauna lived at depths between 100 and
350 m, a bathymetric range substantiated by the inferred ranges of
16 associated phyla also preserved as fossils at Knappton. The nau-
tiloid cephalopod Aturia, which is common in the collections, in-
dicates that the water temperature may have been as high as 1 6°C
at a depth of 100 m. The abundance of preserved organic material
suggests that free oxygen was depleted in the sediment below the
level of bioturbation.

INTRODUCTION

Fossiliferous concretions that have eroded out of landslide
blocks from the upper part of the Lincoln Creek Formation
occur as float along the Columbia River near the site of
Knappton in southwestern Washington (Figs. 1, 2). The Lin-
coln Creek Formation in this area is part of a homocline,
and the sequence dips eastward. The upper part consists of
poorly bedded, locally laminated, bioturbated, concretionary
dark-gray siltstone. The formation is unconformable upon
the upper Eocene siltstones of Cliff Point unit (Wells, 1979).

The invertebrate fauna described in this report is from the
uppermost part of the Lincoln Creek Formation (LAM Loc.
5842) and is early Miocene in age. The fauna is assigned to
the upper part of the Juanian Molluscan Stage (=Saucesian
Foraminiferal Stage) and the Liracassis apta [Echinophoria
apta] Molluscan Zone. The Lincoln Creek Formation is over-
lain by the lower Miocene part of the Astoria Formation,

Contributions in Science, Number 351, pp. 1-42
Natural History Museum of Los Angeles County, 1984

assigned to the Pillarian Molluscan Stage and the Vertipecten
fucanus Molluscan Zone (Fig. 3).

Although the fossiliferous concretions are collected as float
on the bank of the Columbia River, an approximate stratig-
raphy is preserved in the landslides because wedge-shaped
blocks move south toward the river bank parallel with the
strike of the rocks. One locality, informally called the “glass
sponge bed” (LAM Loc. 5852), and others below it, called
the “decapod crustacean bed” (LAM Loc. 5843) and the
“gooseneck barnacle bed” (LAM Loc. 5844), all lie strati-
graphically below the major mollusk-bearing unit (the Aturia
bed) described here (LAM Loc. 5842). Faunas in these four
beds are segregated in the float in proper stratigraphic po-
sition. The lack of mixing is also characteristic of faunas
typical of the overlying Astoria Formation (LAM Loc. 5863).

The geology in the area is complicated by landslides, faults,
by few road cuts, and by vegetative cover (Figs. 4, 5). Gen-
erally, only sections 100 m or less can be measured in tidal
exposures, and the relationship between exposures is often
difficult to discern (Figs. 6, 7). Mapping by Wells (1979)
portrays the complexities in the area.

Despite the poor exposures, some interpretations can be
made. The concretions are continuously being reworked from
the landslides and new accumulations appear with sufficient
regularity to be collected every two weeks at low tide. The
concretions therefore are randomly distributed throughout
the unit.

Most of the concretions are spherical and composed of
fine-grained siltstone with calcareous cement. A few are ce-
mented by quartz in the central part and calcite at the rim.
I believe that the concretions formed early in diagenesis,
because the mollusks preserved in the concretions are almost
always complete specimens that are neither broken nor

1 . U.S. Geological Survey, 345 Middlefield Road MS-9 1 5, Menlo
Park, California 94025, and Research Associate, Invertebrate Pa-
leontology Section, Natural History Museum of Los Angeles County.

ISSN 0459-8113

124° 123°

Figure 1. Index map of the Pacific coast showing localities men-
tioned in the text.

crushed. Delicate features of snails such as the long, narrow
siphonal canal of Priscofusus and the T-shaped axial ribs of
Ancistrolepis are preserved intact. Some of the concretions

do not have any obvious organic center, but most contain
fossils, many are bioturbated, and most contain fecal pellets.

Of particular interest as an aid to diagenetic interpretation
is the preservation of complex sequences of laminated calcite,
sparry calcite, barite, and quartz, precipitated in that order,
within the phragmocone chambers of the cephalopod Aturia.
This sequential mineralization is currently being studied, but
it is obvious now that the laminated calcite was precipitated
first followed by sparry calcite, barite, and quartz.

PREVIOUS WORK

James L. Goedert, in collaboration with Gail H. Goedert,
began collecting fossils in the Knappton area in the late 1970’s
and donated the collections to the Natural History Museum
of Los Angeles County. The first invertebrate collections were
made over a relatively large area and assigned the locality
number LAM 5787. Later observations led Goedert to realize
that the concretions and other fossiliferous material on the
terrace represented separate faunal zones. Thus he separated
subsequent collections into three localities: the lowest (LAM
Loc. 5844) contains abundant gooseneck barnacles ( Arco -
scalpellum) and the trace fossil Tisoa; the middle (LAM Loc.
5843) contains many decapod crustaceans; and the upper-
most (LAM Loc. 5842) contains siliceous sponges, mollusks
including large specimens of the cephalopod Aturia, and
abundant marine vertebrates. Still later, he was able to sep-
arate the siliceous sponge-bearing locality (LAM Loc. 5852)
from the other three localities. At that time, he divided the
collecting area into four informal faunal units. Beginning at
the base of the section these are; Unit I (LAM Loc. 5844),
Unit II (LAM Loc. 5843), Unit III (LAM Loc. 5852), and
Unit IV (LAM Loc. 5842) (Fig. 3).

Victor A. Zullo (1982) described the barnacles from Units
I and IV. From Unit I, he described two species of gooseneck
barnacles, Arcoscalpellum knapptonensis and A. raricosta-
tum, and assigned the unit to the upper Eocene. From Unit
IV, Zullo described the archaeobalinid Solidoba/anus (Hes-
peribalanus) sp. alf. S. (//.) sookensis (Cornwall) and assigned
the unit to the upper Oligocene.

J. Keith Rigby and David E. Jenkins (1983) described
sponges from Units II, III, and IV. Eurete goederti was de-
scribed from Unit III and Aphrocallistes polytretos was de-
scribed from Units II, III, and IV. Rigby and Jenkins also
identified A. polytretos in three other places: a limestone
quarry in the Bear River area northwest of Knappton (LAM
Loc. 5802) that is in the upper Eocene siltstones of CliffPoint
unit (Wells, 1979); in the type area of the lower and middle
Miocene Astoria Formation at Astoria, Oregon; and in the
upper Oligocene and lower Miocene Yaquina Formation,
south of Newport, Oregon. Thus E. goederti is early Miocene
in age and the range of A. polytretos is late Eocene to middle
Miocene.

J. Dale Nations, Northern Arizona University, is studying
the decapod crustaceans from Units II and IV, Carole S.
Hickman, University of California, is studying micromol-
lusks from the section at Knappton, and Bruce J. Welton,
Chevron Oil Field Research Company, the fish. Birds and
marine mammals, collected throughout the section but most

2 Contributions in Science, Number 351

Moore: Lincoln Creek Formation Mollusks

123°48' 123°46’

Formation, Ta = Astoria Formation, Tb = Tertiary basaltic sill, and Qls = Quartemary landslide material.

commonly from Unit IV, are in the vertebrate collections of
the Natural History Museum of Los Angeles County.

CHRONOSTRATIGRAPHY

Molluscan stages were proposed for the Pacific northwest
Tertiary section by Addicott (1976c) and by Armentrout
(1975, 1977), and molluscan zones by Durham (1944), with
revisions and additions by both Addicott (1976c) and Ar-
mentrout (1977). Subsequent work (Allison, 1978; Marin-
covich, 1979; Moore, 1984) has demonstrated the usefulness
of these stages and zones (Fig. 3).

Stratigraphic sections representing the major Oligocene part
of the Juanian Molluscan Stage have been extensively studied
and their molluscan faunas described and illustrated (Teg-
land, 1933; Weaver, 1942; Durham, 1944; Armentrout, 1973;
Addicott, 1976a, 1976b).

The upper part of the Lincoln Creek Formation is assigned

to the upper part of the Juanian Molluscan Stage, equivalent
to the upper part of the Liracassis apta Molluscan Zone (Fig.
3). The upper part of the formation is of early Miocene age
and equivalent in age to the earliest part of the Saucesian
Foraminiferal Stage.

The upper part of the Lincoln Creek Formation in south-
western Washington, assigned to the part of the Juanian that
is of late Oligocene age, contains a molluscan fauna that
remained essentially the same in species composition
throughout the late Oligocene. The strata exposed near
Knappton, which represent the highest exposed part of the
Lincoln Creek Formation and the part of the Juanian Mol-
luscan Stage that is of earliest Miocene age, contains five new
species, representing about 1 5% of the molluscan fauna in
the unit. I interpret this as indicating that this part of the
section is rarely preserved and that the fauna is transitional
between the well-known part (upper Oligocene) of the Jua-
nian and the Pillarian (lower Miocene). The fauna has more

Contributions in Science, Number 351

Moore: Lincoln Creek Formation Mollusks 3

Figure 3. Stratigraphic position of fossil localities in the Lincoln Creek Formation and adjacent formations in southwestern Washington.

species in common with the Juanian than with the Pillarian,
which may indicate that it is closer in age to the Juanian or
lived at depths more commonly represented in the Juanian.
The mollusks from the upper part of the Lincoln Creek

Formation are similar enough to the molluscan fauna from
the upper part of the Pysht Formation of the Twin River
Group, exposed along the Strait of Juan de Fuca, to suggest
a partial correlation of those formations. Addicott (1976b:

4 Contributions in Science, Number 351

Moore: Lincoln Creek Formation Mollusks

Figure 4. View looking southwest toward Knappton monument at
upper left and past fossil localities along the coast at the right, [m
monument, f = fossil localities.]

442) considered the molluscan fauna in the upper part of the
Pysht Formation to belong to the Liracassis apta Molluscan
Zone. Although L. apta was not collected from the Pysht
Formation, Addicott believed that this was due to that unit’s
relatively shallow-water depositional environment rather than
to the organism’s extinction. On the basis of foraminiferal
evidence, Addicott assigned the upper part of the Pysht For-
mation to the Saucesian. “ Bathybembix” hickmanae n. sp.,
Bruclarkia yaquinana, and Megasurcula sp. cf. M. wyno-
ocheensis suggest correlation of the upper part of the Lincoln
Creek Formation with the upper part of the Pysht Formation.

Musashia ( Nipponomelon ) shikamai n. sp. and Bruclarkia
yaquinana, from the upper part of the Lincoln Creek For-
mation at Knappton, were previously reported from strata

Figure 5. Landslide exposure of upper part of the Lincoln Creek
Formation along the Columbia River.

Figure 6. Tidal exposure of upper part of the Lincoln Creek For-
mation on the Columbia River terrace.

no older than the Clallam Formation (Addicott, 1976c: 14-
15) of early Miocene age, equivalent to the Pillarian Mol-
luscan Stage. The occurrence of these species in the upper
part of the Lincoln Creek Formation extends their range

Figure 7. Concretionary layer in upper part of the Lincoln Creek
Formation exposed in a landslide block along the Columbia River.

Contributions in Science, Number 351

Moore: Lincoln Creek Formation Mollusks 5

Depth in meters

o 1000 2000 3000 4000

Gastropods

Bathybembix

Sthenorytis

Ancistrolepis

Musashia (Musashia) • • • •

Musashia (Nipponomelon) •

Aforia

Pelecypods

Acila (Acila)

Portlandia (Portlandia) * •

Acharax

Limopsis

Modiolus

Acesta ( Plicacesta )

Lucmoma

Thyasira (Conchocele) • • •

Cochlodesma

Cephalopod

Aturia, based on Nautilus • •

Figure 8. Depth range of living species of genera of fossils found in concretions in the Aturia bed near the top of the Lincoln Creek Formation,
at Knappton, Washington. The overlapping ranges indicate a depth for the assemblage of about 100 to 350 meters, equivalent to the outer
continental shelf or the upper continental slope.

downward into the upper part of the Juanian Molluscan Stage.
This further refines the Juanian Molluscan Stage and indi-
cates an interval in the earliest part of the Saucesian when a
single molluscan province may have extended from south-
western to northwestern Washington.

The generic composition of the Knappton fauna (“ Bath -
ybembix Liracassis, Ancistrolepis, and Musashia ) is similar
to other assemblages of mollusks collected from the Lincoln
Creek Formation and its correlative units, and the conditions
of deposition seem also to be similar.

Musashia ( Nipponomelon ) weaveri survived throughout the
entire Oligocene. Musashia (N.) shikamai n. sp., also present
in the Clallam Formation (Addicott, 1976c, pi. 3, fig. 27)
and in the upper part of the Poul Creek Formation, replaced
M. weaveri in the upper part of the Lincoln Creek Formation.

Liracassis durhami Kanno, which occurs with Liracassis
apta at Knappton (Moore, 1984), was present during the
upper part of the Juanian Stage, then became locally extinct,
but survived through the Pillarian Stage in the upper part of
the Poul Creek Formation in the Gulf of Alaska. Although
common in the collections from Knappton, L. durhami is
rare in other exposures of the Lincoln Creek Formation.

Ancistrolepis clarki teglandae occurs in the Oligocene part
of the Juanian Stage and was preceded by A. clarki clarki in
the Matlockian Stage. Ancistrolepis jimgoederti n. sp. occurs

in the upper part of the Lincoln Creek at Knappton, in the
lower Miocene part of the Juanian Stage.

The uppermost part of the Lincoln Creek Formation at
Knappton is assigned to the earliest Miocene on the basis of
its molluscan fauna. Two fossil localities of Goedert, units 3
and 4 are of early Miocene age; units 1 and 2 may be of late
Oligocene age. The upper part of the Lincoln Creek For-
mation at Knappton is correlative with the upper part (lower
Miocene) of the Pysht Formation. The placement of the up-
per part of the Lincoln Creek Formation in the lower Miocene
corroborates the assignment of the upper part of the Juanian
Stage to the lower Miocene (Addicott, 1976b:442).

PALEOECOLOGY

The Lincoln Creek Formation in southwestern Washington
represents the accumulation of sediment in a relatively stable
basin environment that shallowed to the southeast. The mol-
luscan fauna of the lowermost Miocene part of the Lincoln
Creek Formation at Knappton lived at dephs between 100
and 350 m (Fig. 8).

The mollusk-bearing concretions are highly bioturbated
with numerous burrows. Preserved fecal pellets occur as small
spherical ooids within the body chambers of gastropods,
elongate pelloids within the septal chambers of the cepha-

6 Contributions in Science, Number 351

Moore: Lincoln Creek Formation Mollusks

lopod Aturia (Figs. 156, 157), and both types scattered
throughout the concretions. Seventeen phyla are represented
in the fauna, and they indicate a healthy community of or-
ganisms living in water of normal salinity. The abundance
of Aturia suggests that the water temperature may have been
as high as 16°C with shallow embayments nearby with tem-
peratures as high as 24°C for egg laying (Cochram, Rye, and
Landman, 1981:477). The abundance of preserved organic
material suggests that the sediment below the level of bio-
turbation was dysaerobic, reduced in oxygen (0. 1-0.5 ml/
liter), and that regular sedimentation gradually buried the
organic material without destroying it or the organisms that
lived there.

Zullo (1982 and written commun., 1982) described Soli-
dobalanus (Hesperibalanus) aff. S. (H.) sookensis (Cornwall)
from the upper part of the Lincoln Creek Formation at
Knappton. Although this archeobalanid barnacle is repre-
sentative of a sessile-benthic group usually found at subtidal
to inner-shelf depths, the basal plates of the Knappton spec-
imens suggest attachment to shells or wood, which could
have been transported to the site of deposition.

Rigby and Jenkins (1983) described and assigned sponges
from the upper part of the Lincoln Creek Formation to the
genera Aphrocallistes and Eurete; the distribution of living
species of both genera suggests that they lived at a depth of
approximately 300 to 350 m.

The foraminifers in the upper part of the Lincoln Creek
Formation in the Grays River quadrangle, just east of
Knappton, indicate water depths of 300 to 900 m (Rau in
Wolfe and McKee, 1972:42).

James C. Ingle, Jr. (written commun., 1982) examined thin
sections made from the centers of mollusk-bearing concre-
tions. He concluded: “All of the evidence in your thin sec-
tions suggests deposition occurred on a continental slope or
marginal basin associated with impingement of the oxygen
minimum layer producing anaerobic or dysaerobic condi-
tions .... The core of this oxygen depleted water mass com-
monly occurs at a depth between 200 and 600 m off Cali-
fornia today and the foraminifera identified in your thin
sections support this depth range ( Globulimina , Epistomi-
nella, Bolivina, Uvigerina). There is evidence of redeposition
of some of the material from shallower environments with
neritic-littoral echinoderm spines and thick walled porcela-
neous foraminifera present in several samples. In addition,
the rare glauconite fragments were likely redeposited from
the adjacent shelf-edge or outer neritic area.”

A sample of sediment from the head of the active landslide
at the west end of the major fossil-bearing concretion locality
(LAM 5842) and samples from a measured section strati-
graphically below have yielded Zemmorian age foraminiferal
assemblages and suggest, as a conservative estimate, that the
water depth was midslope, 1000 m or possibly deeper (Kris-
tin McDougall, written commun., 1982; James C. Ingle, Jr.,
written commun., 1983).

Bruce J. Welton, studying the fish remains from Knappton,
has found bones, scales, and teeth to be abundant at all the
localities and bony-fish otoliths to be common in almost all
samples. Preliminary identifications show that the fish in-

clude seven genera of sharks representing six families, and
one family of bony fish, in addition to numerous unidentified
otoliths and isolated bones. According to Welton (written
commun., 1982),

Ecologically, the sharks are represented by two epipe-
lagic genera ( Cetorhinus and Eugomphodus) and five
genera ( Centrophorus , Chlamydoselachus, Notorynchus,
Scymnodon, and Pristiophorus) with closely related liv-
ing species which are predominantly benthic and deep
water forms. Several taxa are broad ranging bathy met-
rically (Notorhvnchus and Pristiophorus) but collectively
the assemblage is taxonomically right for deep water. A
precise depth would be difficult to substantiate but all
forms would be expected to occur together at a depth of
600 to 1500 ft [180 to 460 m].

Silicified otoliths are usually poorly preserved, how-
ever, the majority (98%) are of mesopelagic lantemfishes
(Family Myctophidae). Many compare favorably to the
genus Diaphus. The extant D. theta occurs today in the
N. Pacific from N. Baja California to the Gulf of Alaska
and Japan, at depths from surface (over deep water) to
2600 ft [790 m].

If one considers only the present day bathymetric dis-
tribution of the genera of sharks and bony fishes known
to occur at Knappton, the assemblage from all four lo-
calities would have to be characterized as a mix of epi-
pelagic and deep water benthic sharks and mesopelagic
teleosts. A bottom depth of 600 to 1500 ft [180 to 460
m] would not be unreasonable.

Although the sample is small, the absence of other
selachian taxa (e.g. Heterodontus, Squatina, Squalus,
Triakis, Mustelus, Galeorhinus, and assorted skates and
rays) strongly dictates against both a shallow water (shelf)
origin of the fauna or resedimentation of a shallow water
assemblage into deeper water by turbidites or related
processes.

From LAM Vertebrate Locality 4510 (=LAM Invertebrate
Locality 5842) Welton has identified Scymnodon sp., cf. Dia-
phus sp., and Myctophidae with a combined modem bathy-
metric distribution of 300 to 800 m.

Because Aturia is 20 times as abundant as any other mol-
lusk in the fauna, particular attention should be paid to its
inferred ecologic requirements. Nautilus, the structurally
similar closest living relative of Aturia, implodes at a water
depth of 785 m (Kanie et al., 1980), which presumably sets
a maximum living depth for Aturia and the associated mega-
fauna and microfauna. Nautilus eggs are probably laid in
shallow water, 100 m or less. After hatching, the young de-
scend to 250 to 350 m (Hamada, Obata, and Okutani, 1980:
47). The first seven septa in Nautilus have shown low 5I80
values, indicating Nautilus hatched in warm, shallow water,
about 24°C (Cochram, Rye, and Landman, 1981:477). The
eighth and later septa have higher <5lftO values, suggesting
that the juvenile Nautilus subsequently migrates to deeper,
colder water, about 16°C. Indirect evidence for the need of
shallow warm water for egg laying and hatching of Aturia
lies in the present distribution of Nautilus in the south Pacific

Contributions in Science, Number 351

Moore: Lincoln Creek Formation Mollusks 7

Other authors:

Musashia ( Musashia ) n. sp. Armentrout in MS (1973)

Musashia ( Musashia ) sp. of Allison and Marincovich (1981, pi. 3,
figs. 8, 14)

Musashia (Musashia) sp. of Allison and Marincovich (1981, pi. 3,
figs. 12, 13)

Musashia (Musashia) sp. of Allison and Marincovich (1981, pi. 3,
figs. 16, 17)

Miopleiona sp. A Durham (1944:178; UCMP 35421, 35422)
Miopleiona sp. B Durham (1944:178; UCMP 35423)

Miopleiona weaveri Tegland (1933:127-128, pi. 11, figs. 1-5)
Miopleiona scowensis Durham (1944:177-178, pi. 17, fig. 15)
Psephaea (Miopleiona) cf. P. (M.) weaveri (Tegland) of Addicott
(1970, pi. 13, figs. 15, 19)

Psephaea (Miopleiona) indurata (Conrad) of Moore (1963:43-44, pi.

7, figs. 1, 2, 3-9, 11; pi. 8, figs. 1-4, 5)

Miopleiona oregonensis Dali (1909:35-36, pi 18, figs. 3, 7)
Miopleiona sp. Clark (1918, pi. 23, fig. 13; UCMP 11244
Psephaea (Miopleiona) weaveri (Tegland) of Addicott (1970, pi. 13,
fig. 17)

Miopleiona indurata (Conrad) of Clark (1918:185; UCMP 12030)
Psephaea corrugata Clark (1932:831, pi. 21, figs. 5, 11)

Miopleiona sp. Loel and Corey (1932:241 ; UCMP 12136)
Psephaea (Miopleiona) cf. P. (M.) indurata (Conrad) of Addicott
(1970, pi. 13, figs. 6, 8)

Musashia indurata (Conrad) of Addicott (1976c, pi. 3, fig. 27)
Musashia n. sp. of Addicott (1976a, pi. 4, fig. 18)

Postellaria indurata Conrad (1849:727-728, pi. 19, fig. 12)
Miopleiona indurata (Conrad) of Weaver (1942:491, pi. 94, figs. 5,

8, 13)

This report:

Musashia (Musashia) n. sp. a
Musashia (Musashia?) sp. b

Musashia (Musashia?) n. sp. c

Musashia (Nipponomelon?) n. sp.?

Musashia (Musashia) n. sp. a
Musashia sp.

Musashia (Nipponomelon) weaveri (Tegland)

Musashia (Nipponomelon) weaveri (Tegland)

Musashia (Nipponomelon) weaveri (Tegland)

Musashia ( Miopleiona) indurata (Conrad)

Musashia (Nipponomelon) oregonensis (Dali)

Musashia (Nipponomelon?) sp.

Musashia (Nipponomelon) weaveri (Tegland)

Musashia (Nipponomelon?) sp. cf. M. (N.) weaveri (Tegland)
Musashia (Neopsephaea) corrugata (Clark)

Musashia (Nipponomelon) shik amain, sp.

Musashia (Nipponomelon) shikamai n. sp.

Musashia (Nipponomelon) shikamai n. sp.

Musashia (Nipponomelon) n. sp.?

Musashia (Miopleiona) indurata (Conrad)

Musashia (Miopleiona) indurata (Conrad)

Figure 9. Allocation of Eastern Pacific Tertiary volutids assigned to the genus Musashia.

and in the fact that Aturia became extinct in the eastern
Pacific at the close of the early Miocene. A combination of
cooling and marine regression may have eliminated suitable
sites for reproduction. Aturia also probably inhabited a shelf
or slope environment where it could have come into shallow
warm water for nocturnal feeding and reproduction and eas-
ily returned to deeper water for resting and escape from pred-
ators.

A depth between 100 and 350 m for the organisms pre-
served in the concretions is indicated on the basis of all the
mollusks including the most abundant element of the fauna,
Aturia. Foraminifers from nearby sediment at the modem
landslide at Knappton indicate a greater depth (about 1000
m). If the difference is real, a possible explanation is that an
early Miocene submarine landslide transported the Aturia-
bearing sediment into deeper water before lithification.

Other fossil remains present in the biota but not yet studied
in detail include radiolarians, coelenterates, echinoderms,
bryozoans, brachiopods, decapods, polychaetes, trace fossils,
marine mammals (cetaceans), birds, seeds, and wood.

The pelecypods are mostly infaunal at shallow subbottom
depths; the spantagoid echinoids and marine worms also are
infaunal, perhaps to depths as much as 6 cm. The gastropods
are mostly epifaunal, although the cassids may have plowed
through the sediment in search of their echinoid prey. The
sponges, coral, and crabs were also mostly epifaunal.

Temperature data obtained from living or closely related

molluscan species are somewhat ambiguous. Whereas mol-
lusks such as Ancistrolepis, Aforia, Portlandia, Acharax,
Acesta ( Plicacesta ), Acesta (Acesta), and Lucinoma suggest
temperatures between 5 and 8°C, Aturia may have required
a temperature of at least 1 6°C.

MOLLUSCAN PALEONTOLOGY

The molluscan fauna consists of 33 taxa and many of these
have been treated by Tegland (1931, 1933), Durham (1944),
and Addicott (1970, 1976b, 1976c). Taxonomic notes rather
than formal systematic descriptions are used for all but new
species to avoid redundancy. Newly described species are
treated more formally.

The following taxa are included, and, unless otherwise in-
dicated, all are from LAM Locality 5842 (Unit IV) and are
illustrated at natural size.

Gastropods:

“ Bathybembix" hickmanae n. sp.

Epitonium ( Nitidiscala ?) sp.

Sthenorytis sp.

Unidentified naticids
Liracassis durhami Kanno
Liracassis apt a (Tegland)

Buccinid?

Bruclarkia yaquinana (Anderson and Martin)
Ancistrolepis jimgoederti n. sp.

8 Contributions in Science, Number 351

Moore: Lincoln Creek Formation Mollusks

Priscofususl sp. cf. P. geniculus (Conrad)

Musashia (Musashia) n. sp.

Musashia ( Nipponomelon ) shikamai n. sp.

Musashia (Miopleiona) n. sp.

Aforia wardi (Tegland)

Turriculal sp.

Megasurculal sp. cf. M. wynoocheensis (Weaver)
Microglyphus n. sp.?

Pelecypods:

Acila ( Acila ) gettysburgensis (Reagan)

Portlandia ( Portlandia ) chehalisensis (Arnold)

Acharax dalli (Clark)

Limopsis nitens (Conrad)

Modiolus addicotii n. sp.

Acesta ( Acesta ) twinensis (Durham)

Acesta ( Plicacesta ) wilsoni n. sp.

CrassostreaP. sp.

Lucinoma hannibali (Clark)

Thyasira (Conchocele) disjuncta (Gabb)

Nemocardiuml sp. cf. N. lorenzanum (Arnold)

Macoma sp. cf. M. twinensis Clark
Cochlodesma bainbridgensis Clark
Teredinid
Scaphopod:

Dentalium ( Fissidentaliuml ) sp. cf D. porterensis Weaver
Cephalopods:

Aturia angustatata (Conrad)

Sepiid?

ABBREVIATIONS

CAS: California Academy of Sciences, San Francisco.
LACMIP: Natural History Museum of Los Angeles County,
Invertebrate Paleontology Section, California.

LACMP: Natural History Museum of Los Angeles County,
Invertebrate Paleontology Section, California.

LAM: Natural History Museum of Los Angeles County, Cal-
ifornia.

CAS/SU: Stanford University, Stanford, California. (The
Stanford University collections are now housed at the Cal-
ifornia Academy of Sciences.)

SU: Stanford University, Stanford, California.

UC: University of California, Berkeley.

UCMP: University of California, Museum of Paleontology,
Berkeley.

USGS: U.S. Geological Survey, Washington, D.C., Cenozoic
locality register.

USGS M: U.S. Geological Survey, Menlo Park, California,
Cenozoic locality register.

USNM: National Museum of Natural History, Washington,
DC.

UW: University of Washington, Seattle, Washington.

GASTROPODS

Trochidae

The genus Bathybembix is used here in a broad sense fol-
lowing Hickman (1980:16) who is currently undertaking a

detailed revision of the large tuberculate trochid gastropods
allied to Bathybembix. The Pacific Northwest fossil species,
Turcicula Columbiana Dali (1909:99-100, pi. 3, figs. 2, 1 1)
and T. washingtoniana Dali (1909:99-100, pi. 17, figs. 1, 2;
pi. 18, fig. 4) were considered by Rehder (1955:225) to “be-
long to Bathybembix, or are more closely related to that genus
than to any other.” Noda ( 1 975:60) believed that “ Turcicula ”
Columbiana and “7’.” washingtoniana differ sufficiently to
warrant a new subgeneric name. Certainly “77” washingtoni-
ana needs more careful scrutiny in terms of generic or subge-
neric allocation. Other Pacific coast fossil species that have
been assigned to Turcicula or to Bathybembix are: Turcicula
arnoldi Durham (1944:153-154, pi. 15, fig. 10), Turcicula
sanctacruzana Arnold (1908:373, pi. 33, fig. 4), Turcicula
turbonata Clark (1932:826, pi. 20, fig. 1 1), and Bathybembix
nitor Hickman (1980:17-18, pi. 2, figs. 1, 2). In addition,
Armentrout (1973), in his study of the Lincoln Creek For-
mation in Washington, recognized three new species which
he assigned to Bathybembix.

“ Bathybembix ” hickmanae n. sp.

Figures 10-12, 18

Bathybembix aff. B. arnoldi (Durham). Addicott, 1976b, figs.

6u, w.

“ Bathybembix ” hickmanae is a thin-shelled, moderately large
trochid with five whorls. The body whorl is characterized by
two spiral cords separated by an almost vertical angulation.
The whorls of the spire also have two spiral cords, one at
the suture separated from the cord above by the same type
of vertical angulation. The outer shell layer is preserved only
in small patches, but the spirals may have been keeled and
tuberculate on the shoulder of the body whorl and the spire
whorls; the rest of the shell may have been smooth. The
available specimens are poorly preserved and no nacreous
shell material is apparent on any of shell patches preserved,
whereas nacreous shell material is commonly preserved on
both “ Bathybembix ” columbiana and “ Bathybembix ” wash-
ingtoniana.

HOLOTYPE. LACMIP 6623, height 42 mm, width 35
mm; paratypes LACMIP 6621, height 41 mm, width 35 mm;
LAM 6622, height 28 mm.

TYPE LOCALITY. LAM 5842.

“ Bathybembix ” hickmanae somewhat resembles “R.” ar-
noldi (Durham) and “B.” sanctacruzana (Arnold) in outline
but differs in having both a steeper and longer slope between
the spirals and the suture on the body whorl. In addition,
“R.” hickmanae has the anterior spiral on the spire whorls
at the suture, whereas “2?.” arnoldi and “B.” sanctacruzana
have a space between the anterior spiral and the suture. The
vertical angulation between spirals on the body whorl and
the higher spire in proportion to width separate “2?” hick-
manae from “5.” washingtoniana (Dali). In addition, “2?”
washingtoniana has a strong keel on the periphery of the
body whorl. The vertical angulation between spirals on the
body whorl distinguishes “ B .” hickmanae from “2?” tur-
bonata (Clark). “ Bathybembix ” columbiana (Dali, 1 909: 1 00,
pi. 3, figs. 2, 11) has a higher spire and larger nodes than

Contributions in Science, Number 351

Moore: Lincoln Creek Formation Mollusks 9

“5.” hickmanae. “ Bathybembix” hickmanae differs from
“ Bathybembix” nitor (Hickman) in having a quadrate rather
than an ovate aperture.

Traditionally, the species assigned to Bathybembix have
been assumed to indicate deep, often bathyal depths. The
bathymetric distribution of living species in Japan, assigned
to Turcicula, Bathybembix, Ginebis, and Convexia, is 100 to
1000 m (Noda, 1975:58, fig. 3).

This species is named in honor of Carole S. Hickman.

Epitoniidae

Epitonium ( Nitidiscala ?) sp.

Figures 13, 14, 16

A latex impression of the mold of a specimen preserved in
a concretion is illustrated along with the original external and
internal molds. It is a thick-shelled epitoniid with seven whorls
preserved each of which probably bore 14 or 15 thick axial
ribs (seven are exposed). No spiral sculpture is preserved.

The rounded whorls, deep sutures, slim high spire, and
lack of spiral sculpture between the axial ribs suggest Niti-
discala.

This species somewhat resembles Epitonium ( Cirsotrema )
saundersiEe^and (1933:133, pi. 13, figs. 7-9; Durham, 1937:
491-492, pi. 57, fig. 21) which occurs in the Liracassis rex
Molluscan Zone (Durham, 1944: 1 58). Epitonium saundersi,
however, has spiral sculpture and less rounded but wider
whorls than E. (TV.?) sp.

Sthenorytis sp.

Figures 15, 19, 20

Sthenorytis sp. may have had five, or possibly more, rapidly
enlarging well-rounded whorls, including a very large body
whorl set off from the axis at an angle of about 40°, as is
typical of Sthenorytis. The suture presumably is deep, and
the body whorl may have had 16 varices. The varices are
rather evenly spaced, triangular in cross section, sharp edged,
and project about 4 mm beyond the shell.

The only described species resembling S. sp. is Sthenorytis
ventricosum (Clark, 1918:164, pi. 23, fig. 14) from the San
Ramon Sandstone, California. It has a smaller body whorl
(31 mm wide) than S. sp. (36 mm wide) and is 45 mm high
compared to about 55 mm for S’, sp. The varices number

about 1 2 and are rounded on S. ventricosum; S. sp. has about
1 6 varices that are triangular and sharp-edged.

Three Pacific coast Tertiary species are assigned to Sthe-
norytis: S.? crescentense (Durham, 1937), S. ventricosum
(Clark, 1918), and S. stearnsi (Dali, 1 892). The geologic range
of these species is Eocene to Pliocene and the geographic
range is northwestern Washington to southern California.

Sthenorytis lives today no farther north than the Gulf of
California and Cape San Lucas; it lives in warm water in the
Pacific and the Atlantic and is usually found on sandy bot-
toms (Durham, 1937:499). Woodring (1959: 184) reported S.
pernobilis (Fischer and Bemardi) from Cape Hatteras to the
Lesser Antilles at depths of 134 to 220 m, Keen (1971:434,
436) recorded 5”. dianae (Hinds) from Baja California Sur in
82 to 145 m and S. turbinum (Dali) from the Gulf of Cali-
fornia to the Galapagos Islands in 82 to 550 m, and Clench
and Turner (1950:225-226) recorded S. pernobilis (Fischer
and Bemardi) from North Carolina to the Lesser Antilles at
134 to 284 m.

Naticidae

Unidentified naticid

Figure 17

Naticids that may represent Polinices ( Euspira ) are repre-
sented by three specimens, none with the umbilical area well
enough preserved for positive identifications.

Cassididae

Two species of Liracassis, L. durhami and L. apt a, were
found in the upper part of the Lincoln Creek Formation.

Liracassis durhami Kanno

Figures 21-23, 25, 26

Liracassis durhami Kanno, 1971:112-113, pi. 13, figs.
14a-b.

Liracassis durhami Kanno has nodes on the shoulder that
usually form oblique ridges to the suture (Figs. 21-23, 25),
but may be separated from the suture by a narrow unsculp-
tured band or confined to two spiral straps. Secondary spirals
are absent on the body whorl except for one specimen which

Figures 10-23. “ Bathybembix ” hickmanae n. sp., Epitonium ( Nitidiscala ?) sp., Sthenorytis sp., unidentified naticid, and Liracassis durhami
Kanno.

Figures 10-12, 18. “ Bathybembix ” hickmanae n. sp. 10. Paratype LACMIP 6621, height 41 mm, width 35 mm. Showing cross section

of spiral cords. 11. Paratype LACMIP 6622, height 28 mm. Showing nodes on shoulders of spire and body whorl, x 1.5. 12, 18. Holotype
LACMIP 6623, height 42 mm, width 35 mm. Showing patches of preserved shell.

Figures 13, 14, 16. Epitonium ( Nitidiscala ?) sp. 13. Latex impression of specimen shown in Figure 16, xl.5. 14. Internal mold, height
25 mm, width 10 mm. LACMIP 6624a. 16. External mold from which latex impression was taken shown in Figure 13, x2.0; height 26 mm,
width 10 mm. LACMIP 6624.

Figures 15, 19, 20. Sthenory’tis sp. 15. View looking down upon spire, x 1.5; height 55 mm. LACMIP 6625a. 19. View of base of same
specimen in Figure 15 showing triangular cross section of varices, x 1.5. LACMIP 6625b. 20. Portion of body whorl of specimen shown in
Figure 19, width 37 mm.

Figure 17. Unidentified naticid, x 1.5; height 23 mm, width 20 mm. LACMIP 6626.

Figures 21-23. Liracassis durhami Kanno. 21. Showing one secondary rib in interspace, x 1.5; width 46 mm. LACMIP 6627. 22. Showing
ridges on shoulder of body whorl; height 57 mm, width 46 mm. LACMIP 6628. 23. Showing spire; height 63 mm, width 48 mm. LACMIP
6493.

10 Contributions in Science, Number 351

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Contributions in Science, Number 351

Moore: Lincoln Creek Formation Mollusks 1 1

has a single intercalary in one interspace (Fig. 26). Liracassis
durhami always has a rounded body whorl without nodes
below the shoulder; this character distinguishes it from Lira-
cassis rex (Tegland, 1931:413-415, pi. 60, fig. 12; pi. 61, figs.
1-4; pi. 62, figs. 1-6), depicted in Figure 32 and the tabulate
form of L. apta (Figs. 27, 31, 33, 39, 42). From the round
form of L. apta (Figs. 24, 28, 30, 47), L. durhami is distin-
guished by its more concave body-whorl shoulder, ridgelike
nodes on the shoulder, and the absence of intercalaries on
the body whorl. Liracassis durhami always has nodes on the
shoulder of the body whorl, whereas the round form of L.
apta may not. Liracassis petrosa (Moore, 1963, pi. 10, figs,
7,17) differs from L. durhami in having intercalaries between
primary spirals on the body whorl and nodes that are more
fluted on the periphery of the body-whorl shoulder, forming
less oblique and generally smaller ridges on the shoulder.

Liracassis durhami ranges from the northeastern Gulf of
Alaska to southwestern Washington; it occurs in the upper
part of the Poul Creek Formation and the lower part of the
Yakataga Formation, Alaska, as well as in the upper part of
the Lincoln Creek Formation, Washington. In the lower part
of the Yakataga Formation in Alaska, L. durhami is asso-
ciated with Pillarian mollusks (Scott McCoy, pers. commun.,
1980). In Washington, L. durhami ranges from the upper
Galvinian Stage through the Juanian Stage. The infrequent
occurrence of L. durhami compared to L. apta in the upper
part of the Lincoln Creek Formation may indicate that south-
western Washington was at the southern limit of its range.
Liracassis apta is restricted to the Juanian, but has a greater
geographic range than L. durhami, having been found as far
south as California (Ham, 1952:8).

Liracassis apta (Tegland)

Figures 24, 27-31, 33, 39, 42, 46, 47

Galeodea apta Tegland, 1931:415-417, pi. 63, figs. 1-10.

Liracassis apta is common in the upper part of the Lincoln
Creek Formation. The largest specimens of L. apta usually
have round shoulders (Figs. 28, 30), but it is not uncommon
to find large specimens with tabulate shoulders nor is it un-
usual to find small specimens with round shoulders. The
largest specimens of the round-shouldered form almost never
have nodes on either the shoulder or the rest of the body
whorl. The tabulate form always has nodes on the shoulder
of the body whorl and may have them on spiral cords anterior
to the shoulder. Of the Liracassis species studied (Moore,
1984), L. apta is the only one showing the wide range of
variation first noted by Tegland (1931:401, 406), who dis-

tinguished three varieties. Tegland’s varieties 1 and 2 fit into
the tabulate form and variety 3 into the round-shouldered
form. Liracassis apta has intercalaries in the interspaces be-
tween primary spiral cords (Fig. 24) as do L. rex, from the
type section of the Blakeley Formation of Weaver (1912),
Washington, and L. petrosa, from the Astoria Formation,
Oregon, but L. durhami is distinguished by having smooth
spaces between the primary spirals. Liracassis rex always has
nodes on the concave shoulder of its body whorl; these nodes
are not confined to the shoulder but continue adapically as
ridges to the suture, whereas in L. apta the nodes, if present,
are confined to the shoulder periphery. Liracassis rex always
has at least one, and commonly more than one, row of nodes
below the shoulder; L. apta never has more than one row
and commonly has none at all. The nodes on the periphery
of the shoulder of L. petrosa are fluted, whereas they are
rounded in L. apta. Liracassis apta is restricted to the Jua-
nian Stage, and its geographic range is from California to the
western Gulf of Alaska.

Studies of living cassids have shown that these carnivores
eat only echinoids, and that different species within each
cassid genus prefer certain distantly related genera of echi-
noids. That different echinoids can serve as food for closely
related cassids makes clear how two different species of Lira-
cassis, such as L. durhami and L. apta, could live in the same
biologic community, but in different niches.

Buccinidae?

Unidentified buccinid

Figure 36

A single specimen of a poorly preserved gastropod may be
a buccinid. Although many gastropod families can be elim-
inated from consideration on the basis of outline, sculpture,
suture, aperture, or siphonal canal, the specimen is not iden-
tified with complete confidence as a buccinid. The evenly
rounded whorls, slightly impressed suture, elongate-oval ap-
erture, seeming lack of columellar plaits, a probably short,
straight siphonal canal, and cancellate sculpture suggest buc-
cinid genera such as Cymatophos or Antillophos.

Neptuneidae

Bruclarkia yaquinana (Anderson and Martin)

Figures 41, 44

Agasoma yaquinana Anderson and Martin, 1914:75, pi. 4,
figs. 5a-b.

Figures 24-33. Liracassis apta (Tegland), Liracassis durhami Kanno, and Liracassis rex (Tegland).

Figures 24, 27-31, 33. Liracassis apta (Tegland). 24. Round form without body-whorl nodes; height 44 mm, width 37 mm. LACMIP

6495. 27. Tabulate form; height 55 mm. LACMIP 6629. 28. Round form; height 85 mm, width 62 mm. LACMIP 6630. 29. Showing round
body whorl and tabulate spire; height 63 mm, width 49 mm. LACMIP 6631. 30. Round form; height 63 mm. LACMIP 6632. 31. Tabulate
form with nodes on body whorl; height 47 mm, width 42 mm. LACMIP 6633. 33. Tabulate form with beads on body whorl; height 40 mm,
width 33 mm. LACMIP 6634.

Figures 25, 26. Liracassis durhami Kanno. 25. Showing smooth interspaces and ridges on shoulder; height 43 mm, width 39 mm. LACMIP

6496. 26. Showing straplike spiral cords; height 70 mm, width 66 mm. LACMIP 6494.

Figure 32. Liracassis rex (Tegland). Showing nodes on body whorl of holotype UCMP 32067; height 75 mm, width 64 mm. Blakeley
Formation, Restoration Point, Seattle, Washington.

12 Contributions in Science, Number 351

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32

Contributions in Science, Number 351

Moore: Lincoln Creek Formation Mollusks 13

Bruclarkia yaquinana is represented by a single incom-
pletely preserved specimen (Figs. 41. 44). As noted by Ad-
dicott ( 1 970:90-9 1 ; 1 976c:23), B. yaquinana is characterized
by four or five coarsely noded spiral cords on the body whorl.
Bruclarkia oregonensis (Conrad, 1848:433, fig. 13; Moore,
1963: pi. 3, figs. 2, 3, 8. 11, 13) has finer spiral sculpture and
is a larger, more rounded species. In the San Joaquin Valley,
California, the stratigraphic occurrence of B. yaquinana (bas-
al part of the Jewett Sand) and B. oregonensis (Olcese Sand)
is mutually exclusive (Addicott, 1970:91). In the Clallam
Formation of northwestern Washington, B. oregonensis oc-
curs almost exclusively at the top of the formation, and B.
yaquinana at and near the base (Addicott, 1 976c:23-24).

Ancistrolepis jimgoederti n. sp.

Figures 34, 35, 37, 38, 40, 43, 45, 56, 58, 59
The shell of Ancistrolepis jimgoederti is large and thin with
eight or nine subtabulate whorls that bear T-shaped spiral
cords that are strongly undercut and, on the holotype (Figs.
35, 37, 38), preserve three secondary spiral cords. The in-
terspaces on the body whorl bear secondary spiral cords and
one specimen has four moderately prominent and three very
fine secondary cords preserved in one interspace. On this
same specimen, fine vertical striations are preserved in one
interspace, perhaps reflecting the periostracum. The speci-
men with the most whorls preserved (eight) is 68 mm in
height (incomplete), and the specimen with the largest body
whorl has a maximum width of 36 mm. The largest T-shaped
spiral cord preserved projects 3.2 mm beyond the shell body.
The T-shaped spirals are preserved only in concretionary
matrix (Fig. 58) from which they were subsequently exposed
by preparation (Figs. 34, 35, 37, 38, 40). The body of the
shell is so thin that the spiral cords exfoliate easily from the
shell, leaving preserved a completely erroneous shell outline
(Figs. 43, 45). The T-shaped spirals are now replaced by
sparry calcite, but do not seem to have been hollow folds of
the shell, as in the genus Ecphora from the Miocene of the
eastern United States (Vokes, 1957, pi. 25, fig. 2).

HOLOTYPE. LACMIP 6636, height 55 mm, width 35
mm; paratypes LACMIP 6635, height 40 mm, width 29 mm;
LACMIP 6637, height 67 mm, width 35 mm; LACMIP 6638,
height 65 mm, width 35 mm; LACMIP 6646, width 35 mm;
and LACMIP 6647, height 70 mm.

TYPE LOCALITY. LAM 5842.

Species, such as Ancistrolepis clarki (Tegland, 1933:131 —

132, pi. 12, fig. 14), A. landesi (Tegland, 1933:132-133, pi.

13, figs. 1-4), and A. clarki teg/andae (Durham, 1944:177,
pi. 17, fig. 2), may also have had T-shaped spirals that were
removed by exfoliation. The concretionary fragments re-
maining with the holotype of A. clarki teglandae, and the
specimen itself (Figs. 52, 54), show no indication of T-shaped
spirals, but some specimens of A. jimgoederti also show no
indication of these spirals (Fig. 43).

Grant and Gale (1931:657) noted the similarity in spiral
sculpture between Ancistrolepis and Beringius and suggested
that perhaps Ancistrolepis should be considered a section or
synonym of Beringius (Dali, 1887:304; type species Chry-
sodomus crebicostatus Dali). Clifford M. Nelson, U.S. Geo-
logical Survey, kindly called my attention to the fact that the
whorl proportions, aperture, and fasciole of A. jimgoederti
do not fit Neptunea in the strict sense, and that the species
is more closely related to Ancistrolepis. Species from the
eastern Pacific Tertiary that have been assigned to Ancistro-
lepis are: Ancistrolepis rearensis (Clark, 1932), Ancistrolepis
macneili Kanno (1971), Ancistrolepis clarki clarki Tegland
(1933), Ancistrolepis clarki teglandae Durham (1944), An-
cistrolepis landesi Tegland ( 1 933), and Ancistrolepis packardi
Durham (1944).

The spiral ribs on the penultimate whorl of Ancistrolepis
rearensis (C lark, 1932:831, pi. 20, figs. 14, 1 5) are described
as having a fairly prominent collar and thus would resemble
those on A. jimgoederti. Ancistrolepis rearensis has convex
whorls rather than subtabulate ones and lacks the concavity
just below the suture, present on A. jimgoederti. Kanno (1971:
1 18) placed A. clarki teglandae into synonymy with A. rea-
rensis without discussion, and Addicott (1976c:23) cited it
as a junior synonym. Ancistrolepis clarki teglandae (Figs. 52,
54) has a round body whorl and a shorter, wider spire than
A. jimgoederti. Ancistrolepis macneili Kanno (1971:1 19, pi.

14, fig. 7) is much more inflated and has a shorter spire in
proportion to the body whorl than does A. jimgoederti. The
body whorl of A. clarki clarki is evenly rounded to the suture;
the body whorl of A. jimgoederti is deeply concave between
the suture and the first spiral cord and the whorls of the spire
are more tabulate. Ancistrolepis clarki clarki is also wider
and has fewer whorls (six or seven) than A. jimgoederti. An-
cistrolepis landesi has a high spire, similar to A. jimgoederti,
but the body whorl is convex to the suture rather than con-
cave just below the suture. In addition, the primary spiral
cords on the body whorl of A. landesi are grouped together

Figures 34 47. Ancistrolepis jimgoederti n. sp., unidentified buccinid, Liracassis apta (Tegland), and Bruclarkia yaquinana (Anderson and
Martin).

Figures 34, 35, 37, 38, 40, 43, 45. Ancistrolepis jimgoederti n. sp. 34. Paratype LACMIP 6635. Showing secondary spirals, x 1.5; height
40 mm, width 29 mm. 35, 37, 38. Holotype LACMIP 6636; height 55 mm, width 35 mm. Figures 35 and 37 show primary spirals sculptured
by secondaries. Figure 38 is the same specimen tipped to display undercut spiral cords. 40. Same specimen as Figure 34. Showing angular
spiral cord, x 1.5. 43. Paratype LACMIP 6637. Showing cast from which shell has exfoliated; height 67 mm, width 35 mm. 45. Showing spiral
cords in concretion; height 65 mm, width 35 mm. Paratype LACMIP 6638.

Figure 36. Unidentified buccinid, x2.0; height 28 mm, width 13 mm. USGS Loc. M 7891, USNM 363986.

Figures 39, 42, 46, 47. Liracassis apta (Tegland). 39. Tabulate form with body whorl nodes; height 45 mm, width 35 mm. LACMIP 6497.
42. Tabulate form showing nodes and intercalaries; height 49 mm. LACMIP 6639. 46, 47. Rear and apertural view of round form; height 68
mm, width 45 mm. LACMIP 6640.

Figures 41, 44. Bruclarkia yaquinana (Anderson and Martin). Views of aperture and side showing configuration and sculpture, xl.5;
height 30 mm, width 22 mm. LACMIP 6641.

14 Contributions in Science, Number 351

Moore: Lincoln Creek Formation Mollusks

on the middle of the body whorl whereas they are equally
spaced from the suture on A. jimgoederti. The entire shell of
A. landesi bears fine, evenly spaced secondary spiral sculp-
ture. The one specimen of A. jimgoederti that has the sec-
ondary spiral sculpture preserved (Figs. 34, 37, 40) shows
the spirals to be of unequal strength and spacing. The living
species Ancistrolepis grammatus (Dali, 1907:158; 1925:3, pi.
30, fig. 8) has T-shaped spiral cords on a thin shell with
subtabulate whorls. The body whorl, however, has nine
T-shaped spirals of almost equal width compared to five on
A. jimgoederti and the spire whorls have five rather than
three or four spirals. The siphonal canal of A. jimgoederti is
probably slightly longer and more strongly recurved than on
A. grammatus and also bears finer spiral sculpture.

Weaver (1942:427) assigned Ancistrolepis clarki clarki to
Neptunea, and renamed it N. teglandae, as N. clarki was a
homonym of Neptunea clarki (Meek). The reassignment of
A. clarki clarki to Ancistrolepis makes this action by Weaver
no longer necessary.

Ancistrolepis landesi and A. clarki clarki occur in the type
Blakeley Formation in the Liracassis rex Molluscan Zone.
Ancistrolepis clarki teglandae occurs in the upper part of the
Pysht Formation of the Twin River Group in the Liracassis
apta Molluscan Zone. Ancistrolepis rearensis was originally
collected from the upper part of the Poul Creek Formation
in the Liracassis apta Molluscan Zone and the geographic
and stratigraphic ranges were subsequently extended by Ad-
dicott (1976c:23) to the Clallam Formation, Washington, in
the Vertipecten fucanus Molluscan Zone. Ancistrolepis jim-
goederti was collected from the upper part of the Lincoln
Creek Formation, in the upper (Saucesian) part of the Lira-
cassis apta Molluscan Zone.

Chrysodomus eucosimium Dali (1891:187-188), the ge-
notype of Ancistrolepis, was collected off the coast of Un-
alaska in the Bering Sea. Ancistrolepis grammatus was col-
lected from Tsugaru Strait, Japan, at a depth of 550 m where
the surface temperature averages about 1 8°C.

This species is named in honor of James L. Goedert.

Fusinidae

Priscofususl sp. cf. P. geniculus (Conrad)

Figures 50, 53, 55, 60, 61

Priscofususl sp. cf. P. geniculus (Conrad) is represented by
three specimens; two free of matrix (Figs. 50, 53, 55, 61) and

the third preserved in a concretion with the shell missing on
most of the specimen, but replaced by sparry calcite where
still embedded (Fig. 60). Priscofusus geniculus (Conrad, 1849:
728, pi. 20, fig. 3) has been described and illustrated by Moore
(1963:40-41, pi. 6, figs. 13, 15-18) and by Addicott (1970:
101-102, pi. 12, figs. 21, 22, 26, 28-30) and occurs in the
Astoria Formation, Oregon, and the Jewett Sand, California.
The Priscofusus reported from the Clallam Formation (Ad-
dicott, 1 976c:24, pi. 2, fig. 1 2) and the Nye Mudstone (Moore,
1963:41, pi. 6, figs. 12, 19) may represent a new species.

Volutidae

Neogene volutids of the eastern Pacific Tertiary have in re-
cent years commonly been assigned to the genus Musashia
(Hayashi, 1960) and the subgenus Musashia or Miopleiona
(Dali, 1907). A new species of Miopleiona from the Eugene
Formation in Oregon (Howe, 1 922) extends the geologic range
of that subgenus into the late Eocene or early Oligocene.

Nipponomelon (Shikama, 1967), a subgenus previously re-
ported from the Miocene to Holocene in Japan, is used here
for most of the northeastern Pacific volutids, thus extending
the geographic range of the subgenus across the Pacific and
the geologic range into the Oligocene.

Musashia ( Musashia ) has a smooth shell or only thin axial
ribs; axial ribs, if present, may be only on the posterior por-
tion of the whorls (Fig. 70); the suture is slightly impressed.
Musashia (Nipponomelon) has thin axial ribs, a slightly im-
pressed suture, and only rarely a sutural collar (Figs. 66, 68).
Musashia ( Miopleiona ) has very thick keel-like ribs markedly
curved near the suture, which is deeply impressed and chan-
neled (Figs. 51, 57, 64, 67). Shikama (1967) thought that
Miopleiona was intermediate between the subgenera Mu-
sashia and Nipponomelon and was uncertain as to its proper
assignment, although he placed it in the genus Musashia. On
the basis of the suture and the thick axial ribs, Miopleiona
could perhaps be elevated to generic rank. The type species
of Miopleiona is Musashia ( Miopleiona ) indurata (Conrad,
1849). The markedly curved axial ribs and deeply channeled
suture of M. (M.) indurata set it apart from all other described
species of volutids, but the undescribed new species from the
Eugene Formation, Oregon (Howe, 1922), also has a deeply
channeled suture (Fig. 48) and is here assigned to Miopleiona.
The suture is so deeply channeled on Miopleiona that even
internal molds can be identified as belonging in the subgenus.

Figures 48-61. Musashia (Miopleiona) n. sp., Musashia (Nipponomelon) shikamai n. sp., Priscofususl sp. cf. P. geniculus (Conrad), Musashia
(Nipponomelon) indurata (Conrad), Ancistrolepis clarki teglandae Durham, and Ancistrolepis jimgoederti n. sp.

Figure 48. Musashia ( Miopleiona ) n. sp. Plaster cast of a specimen from the Eugene Formation, Oregon; height 95 mm. CAS/SU 2358.

Figure 49. Musashia (Nipponomelon) shikamai n. sp. Paratype. Latex impression showing fine spiral sculpture, height 82 mm. LACMIP
6642.

Figures 50, 53, 55, 60, 61. Priscofususl sp. cf. P. geniculus (Conrad). 50, 53. Internal mold showing traces of axial ribs, x 1.5; height 21
mm, width 18 mm. LACMIP 6643. 55, 61. Almost complete internal mold showing spire outline and axial ribs; height 26 mm, width 15
mm. LACMIP 6644. 60. Showing siphonal canal; height 51 mm, width 18 mm. LACMP 6645.

Figures 51, 57. Musashia (Nipponomelon) indurata (Conrad). Showing twisted axial ribs and deeply impressed suture, x0.8; height 125
mm, width 59 mm. From the Astoria Formation, Lincoln County, Oregon. CAS 037058.

Figures 52, 54. Ancistrolepis clarki teglandae Durham. Latex impression of holotype external molds, UCMP 35417, showing spiral
sculpture; height 48 mm.

Figures 56, 58, 59. Ancistrolepis jimgoederti n. sp. 56, 59. Paratype LACMIP 6646. Showing siphonal canal, xl.5; width 35 mm. 58.
Paratype LACMP 6647. Showing T-shaped cross section of spiral ribs preserved in concretion, x2.0; height 70 mm.

16 Contributions in Science, Number 351

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58

59

Armentrout (1973) reported an undescribed species of vo-
lutid from the Lincoln Creek Formation and assigned it to
Musashia (Musashia), an assignment with which I concur.
Although several species of Musashia ( Musashia ) have been
described from the Cenozoic of Japan, Armentrout’s material
documents the occurrence of the subgenus in western North
America. It has a smooth shell, a slightly impressed suture
that is markedly inclined, and a spire that is short in relation
to the length of the body whorl (Figs. 62, 65, 76, 80, 87).

Subgeneric characters used by Shikama (1967) to differ-
entiate volutids, such as the character of the protoconch and
the number of initial and last columellar plaits, are useful
for the allocation of living species, but are difficult to use
with incompletely preserved fossils. The character of the su-
ture and axial ribs serve best to distinguish fossil forms both
subgenerically and specifically. On the basis of thin axial ribs
and a slightly impressed suture, I assign most of the eastern
Pacific Tertiary volutids to the subgenus Nipponomelon (Fig.
68). My allocation of eastern Pacific Tertiary species is shown
in Figure 9.

Hayashi (1960:2) in his description of the genus Musashia
noted that sexual dimorphism is very pronounced and that
the large convex shells may be female. It is well to bear this
in mind when looking at closely related eastern Pacific Ter-
tiary species. In the Knappton fauna, however, slim forms
are more common that convex forms, which would lead to
the presumably false conclusion that males were more abun-
dant than females.

The oldest recorded occurrence I have found for Musashia
is Musashia ( Nipponomelon ?) caucasica (Korobkov, 1949:
694-695, text figs. 1,2; 1 955:205-206, pi. 4, figs. 6, 6a) from
the middle Eocene in the Caucasus of the U.S.S.R. The oldest
record of Miopleiona is Musashia (Miop/eiona) n. sp. from
the Eugene Formation, Oregon, of late Eocene to middle
Oligocene age. The oldest record of Nipponomelon in the
eastern Pacific is in the lower Oligocene part of the Lincoln
Creek Formation, Washington. Musashia and Nipponome-
lon may have originated in the western Pacific; Miopleiona
is indigenous to Alaska, Washington, Oregon, and California,
and did not invade the western Pacific.

Musashia ( Musashia ) n. sp.

Figures 62, 65, 76, 80, 87

Miopleiona sp. A Durham, 1944:178.

Musashia ( Musashia ) evelynae Armentrout, 1973, in MS:

338-339, pi. 5, figs. 25, 27.

Musashia ( Musashia ) n. sp. has a slim shell with a very low
spire compared to the body-whorl height. The suture is very
slightly impressed and markedly inclined. The shell is smooth,
without axial ribs, and only growth lines are preserved. The
aperture is elongate oval, the siphonal fasciole probably
straight and with a rather thick posterior callus and with two
columellar plaits, the anteriormost one bladelike.

Musashia ( Musashia ) n. sp. is the only species assigned to
this subenus in the eastern Pacific. The type species of Mu-
sashia is M. ( M .) hirasei (Sowerby) (Figs. 69, 70). The only
other described species at all similar to M. n. sp. is Musashia
{Musashia?) nagaoi Shikama (1967:1 1 1-1 12, pi. 13, figs. 9-
12) from the late Oligocene and early Miocene in Japan.
Shikama (1967:1 12) considered M. nagaoi to be unique
among Japanese fulgorids because it lacks axial ribs and ra-
dial striations; this is equally true for M. n. sp. in the eastern
Pacific. Musashia nagaoi has a much more inflated body
whorl than M. n. sp. and the suture of M. nagaoi is not as
steeply inclined. Musashia {Musashia) n. sp. is being de-
scribed by J. M. Armentrout.

Musashia ( Nipponomelon ) shikamai n. sp.

Figures 49, 63, 72, 74, 75, 77, 78, 82, 83, 88, 89

Psephaea {Miopleiona) cf. P. (M.) indurata (Conrad). Ad-
dicott, 1970:105, pi. 13, fig. 8; not pi. 13, fig. 6 {=M.
indurata).

Musashia indurata (Conrad, 1849). Addicott, 1976c:25, pi.
3, fig. 27. Not Rostellaria indurata Conrad, 1849.

Musashia {Nipponomelon) shikamai is large, slender, and
high spired with about nine whorls. The shell bears narrow
axial ribs that are closely spaced and usually twisted near the
suture. On large specimens the axial ribs disappear toward
the anterior end. Narrow axial folds between the ribs pre-
sumably represent growth lines; the entire shell is sculptured
by closely spaced subrounded spiral cords. The suture is
slightly impressed and no subsutural band is preserved. The
number of axial ribs ranges from 16 to 19, with 18 or 19
being the most common. The protoconch is not preserved.
The aperture is assumed to be elongate oval. The siphonal
fasciole is not preserved but may have been straight and long.

HOLOTYPE. LACMIP 6652, height 135 mm, width 52
mm; paratypes LACMIP 6642, height 87 mm; LACMIP
6648, height 73 mm, width 32 mm; LACMIP 6649, height
41 mm, width 20 mm; LACMIP 6650, height 90 mm, width
34 mm; LACMIP 6654, height 67 mm, width 25 mm; LAC-

Figures 62-70. Musashia {Musashia) n. sp., Musashia (Nipponomelon) shikamai n. sp., Musashia (Miopleiona) indurata (Conrad), Musashia
(Nipponomelon) prevostiana magna (Kuroda and Habe), and Musashia (Musashia) hirasei (Sowerby).

Figures 62, 65. Musashia (Musashia) n. sp. U W 16444a; height 57 mm, width 25 mm. 62. Showing smooth shell. 65. Showing columnar
plaits.

Figure 63. Musashia (Nipponomelon) shikamai n. sp. Paratype LACMIP 6648. Showing narrow axial ribs; height 73 mm, width 32 mm.

Figures 64, 67. Musashia (Miopleiona) indurata (Conrad). Showing wide axial ribs and deeply impressed suture; height 66 mm, width 35
mm. USNM 363987.

Figures 66, 68. Musashia (Nipponomelon) prevostiana magna (Kuroda and Habe). CAS 028423; height 1 70 mm, width 60 mm. 66. Looking
down on apex to show suture. 68. View of aperture.

Figures 69, 70. Musashia (Musashia) hirasei (Sowerby). CAS 028422; height 165 mm, width 56 mm. 69. Looking down on apex to show
suture. 70. View of aperture.

18 Contributions in Science, Number 351

Moore: Lincoln Creek Formation Mollusks

68

69

70

MIP 6655, height 104 mm, width 36 mm; and LACMIP
6651, height 67 mm, width 26 mm.

TYPE LOCALITY. LAM 5842.

The species closely similar to M. (N.) shikamai are M. ( N .)
weaveri (Tegland, 1933) and M. (TV.) miensis (Araki, 1960).
Musashia weaveri (Figs. 71, 81, 94) has a slimmer shell with
fewer (11 to 14) and wider axial ribs and a slightly angulated
shoulder rather than a smoothly rounded one as in M. shi-
kamai. Musashia miensis is slimmer and more elongate in
outline, the axial ribs are not twisted near the suture, and
the preserved radial sculpture is not as well developed as in
M. shikamai.

Musashia n. sp. of Addicott (1976a: 108, pi. 4, fig. 18) is
similar to M. shikamai but has a few more axial ribs and a
concave area below the suture. The latter character, however,
could reflect sexual dimorphism.

Musashia ( Miopleiona ) indurata (Conrad, 1849) is distin-
guished from M. shikamai by its deeply impressed, chan-
neled suture and by its strongly twisted keel-like axial ribs.

Musashia indurata (Conrad) of Addicott ( 1 976c: 25, pi. 3,

fig. 27) does not have the deeply impressed, channeled suture
and wide, keel-like axial ribs of M. ( Mioleiona ) indurata and
is here assigned to M. shikamai.

On the basis of the incomplete holotype of Miopleiona
scowensis Durham (1944:177-178, pi. 17, fig. 15), I am un-
able to find any characters to distinguish it from M. weaveri
and believe it should be synonymized with that species.

Musashia ( Neopsephaea ) corrugata (Clark, 1 932:831-832,
pi. 21, figs. 4, 5, 11; Addicott et al., 1971, figs. 2y, aa-bb),
from the Poul Creek Formation, Alaska (Figs. 73, 90, 92,
93, 95) is slimmer than M. shikamai and has a more sharply
inclined suture and fewer axial ribs that are thicker and more
widely spaced than in M. shikamai.

Musashia ( Musashia ) sp. of Allison and Marincovich (1981,
pi. 3, figs. 12, 13; not pi. 3, figs. 8, 14, 16, 17) has a much
wider body whorl in proportion to spire height and fewer
(about 14) axial ribs than M. shikamai. The specimen figured
by Allison and Marincovich ( 1 98 1 , pi. 3, figs. 8, 14), although
poorly preserved, probably belongs in the subgenus Musa-
shia.

Figures 71-77. Musashia (Nipponomelon) weaveri (Tegland), Musashia (Nipponomelon) shikamai n. sp., Musashia (Neopsephaea) corrugata
(Clark), and Musashia (Musashia) n. sp.

Figures 71. Musashia (Nipponomelon) weaveri (Tegland). Showing spacing of axial ribs and spiral sculpture; height 125 mm, width 40
mm. UC locality A 1806, Blakeley Formation of Weaver (1912), Bainbridge Island, Washington. UCMP 35420.

Figures 72, 74, 75, 77. Musashia (Nipponomelon) shikamai n. sp. 72. Showing inflation of body whorl x0.8; height 135 mm, width 52
mm. Holotype LACMIP 6652. 74. Showing spiral sculpture x2.0; height 41 mm, width 20 mm. Paratype LACMIP 6649. 75. Showing
deflection of ribs near suture; height 90 mm, width 34 mm. Paratype LACMP 6650. 77. Showing spacing of ribs and spiral sculpture. Paratype
LACMIP 6651.

Figure 73. Musashia (Neopsephaea) corrugata (Clark). Showing inclined suture and sculpture; height 120 mm, width 33 mm. Upper part
of the Poul Creek Formation, Yakataga Reef, Alaska. USNM 363988.

Figure 76. Musashia (Musashia) n. sp. Showing outline of shell and inclined suture x 1.5; height 52 mm, width 20 mm. LAM Loc. 5843;
LACMIP 6653.

Figures 78-86. Musashia (Nipponomelon) shikamai n. sp., Musashia ? sp., Musashia (Musashia) n. sp., Musashia (Nipponomelon) weaveri
(Tegland), and Musashia (Nipponomelon) prevostiana magna (Kuroda and Habe).

Figures 78, 82, 83. Musashia (Nipponomelon) shikamai n. sp. 78. Paratype LACMIP 6654. Showing inclined suture x 1.5; height 67 mm,
width 25 mm. 82. Showing narrow, closely spaced axial ribs; height 73 mm, width 32 mm. Paratype LACMIP 6648, shown in Figure 63. 83.
Paratype LACMIP 6655. Showing siphonal canal and spiral sculpture; height 104 mm, width 36 mm.

Figure 79. Musashia ? sp. Immature? specimen x 1.5; height 24 mm, width 10 mm. LACMIP 6656.

Figure 80. Musashia (Musashia) n. sp. Showing smooth shell and inclined suture; height 57 mm, width 22 mm. UW 16444.

Figures 81, 84, 86. Musashia (Nipponomelon) weaveri (Tegland). 81. Showing widely spaced axial ribs and spiral sculpture x 1.5; height
40 mm, width 23 mm. USNM 363989. 84, 86. From the basal part of the Jewett Sand, California. USNM 650185; height 73 mm, width 30
mm.

Figure 85. Musashia (Nipponomelon) prevostiana magna (Kuroda and Habe). Rear view showing narrow, closely spaced axial ribs and
suture; height 170 mm, width 60 mm. CAS 028423.

Figures 87-95. Musashia (Musashia) n. sp., Musashia (Nipponomelon) shikamai n. sp., Musashia (Neopsephaea) corrugata (Clark), and
Musashia (Nipponomelon) weaveri (Tegland).

Figure 87. Musashia (Musashia) n. sp. Showing smooth shell and inclined suture x 1.5; height 62 mm, width 19 mm. USNM 363992.
USGS Loc. 25764, Lincoln Creek Formation, Grisdale Quadrangle, Washington.

Figure 88, 89. Musashia (Nipponomelon) shikamai n. sp. 88. USGS Loc. M4050, Clallam Formation, Washington. Hypotype (Addicott,
1976c, pi. 3, fig. 27) USNM 216000; height 74 mm, width 34 mm. 89. UCMP Loc. 3229, Monterey Group of Wagner and Schilling (1923).
Hypotype (Addicott, 1970, pi. 13, fig. 6) UCMP 12136, a latex impression xl.5; height 52 mm, width 21 mm.

Figures 90, 92, 93, 95. Musashia (Neopsephaea) corrugata (Clark). 90. Paratype UCMP 12399; height 55 mm, width 30 mm. Poul Creek
Formation, Alaska. 92, 95. Holotype UCMP 12399; height 85 mm, width 29 mm. Poul Creek Formation, Alaska. 93. Showing spire and
aperture of specimen from Yakataga Reef, Alaska; height 120 mm, width 33 mm. USNM 363988. Same specimen shown in Figure 73.

Figures 91, 94. Musashia (Nipponomelon) weaveri (Tegland). 91. View looking down on apex showing suture, x 1.5; height 40 mm, width
23 mm. Same specimen shown in Figure 81. USGS Loc. 4093, USNM 363989. 94. Showing outline, axial ribs, and spiral sculpture; height
125 mm, width 40 mm. Same specimen shown in Figure 7 1 . UCMP Locality A 1806, Blakeley Formation of Weaver (1912), Bainbridge Island,
Washington. UCMP 35420.

20 Contributions in Science, Number 351

Moore: Lincoln Creek Formation Mollusks

77

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Contributions in Science, Number 351

Moore: Lincoln Creek Formation Mollusks 23

Musashia (Musashia) n. sp., described above, differs from
M. shikamai by having a smooth shell without axial ribs or
radial threads, a much larger body whorl in proportion to
height, and a strongly inclined suture.

In outline and axial sculpture, M. shikamai resembles Mu-
sashia {Nipponomelon) elegantula Shikama (1967) from the
early Pliocene, Japan, but differs from that species by not
having a subsutural band.

The early whorls of a few specimens of M. shikamai from
the upper part of the Lincoln Creek Formation at Knappton,
Washington, are filled with barite.

OCCURRENCE ELSEWHERE. Upper part of the Poul
Creek Formation, Alaska, upper part of the Pysht Formation
of the Twin River Group and the Clallam Formation, north-
western Washington, the Nye Mudstone, Oregon (Howe,
1922:138, pi. 10, fig. 3 as Miop/eiona clatsopensis Howe, n.
sp.), and the Freeman-Jewett Silt of Matthews (1955) and
the Vaqueros Formation, southern California.

Musashia ( Miopleiona ) n. sp.

Figure 48

Miopleiona n. sp. Howe, 1922:137.

Musashia ( Miopleiona ) n. sp., collected from the Eugene For-
mation in Oregon, is a large volutid with a thick shell that
bears perhaps as many as 30 narrow, keel-like axial ribs. The
suture is deeply impressed, channeled, and inclined. The
species is represented by a single plaster cast of a specimen
consisting of half of two whorls. The preserved portion is
identical in all characters to Musashia ( Miopleiona ) indurata
except that the axial ribs are half as wide and twice as closely
spaced as on M. (M.) n. sp.

There is no doubt in my mind that M. (M.) n. sp. was
collected from the Eugene Formation on the University of
Oregon campus. The record is based on a specimen, now
missing and presumed lost, collected by Professor Earl L.
Packard, a paleontologist of note, and I have no reason to
suspect the locality data. The specimen was first mentioned
by Howe ( 1 922: 1 37) as “ Miopleiona n. sp., (very large), lower
Oligocene, Eugene,” and later by Schenck (1928: 1 1) as Mio-
pleiona n. sp. from the Eugene Formation, Oregon. A plaster
cast was made of the specimen for Schenck and deposited in
the Stanford University collection, now housed at the Cali-
fornia Academy of Sciences. Hickman (1969) did not de-
scribe any volutids from the Eugene Formation. Presumably
the plaster cast was not seen at that time, or the locality
description was considered suspect.

Musashia ( Miopleiona ) n. sp. is the oldest known species
of Miopleiona, as the subgenus is used here. Its occurrence
in the Eugene Formation extends its geologic range from the
Miocene into the early Oligocene or late Eocene.

The Eugene Formation has been extensively collected, in
part because it is exposed in almost all excavations made for
buildings on the University of Oregon campus. The single,
incomplete specimen of M. (M.) n. sp. indicates its rareness.
Hickman ( 1 969:22) suggested that the Eugene molluscan fau-
na lived at a depth of 55 m. The molluscan fauna of the

upper part of the Lincoln Creek Formation at Knappton
probably lived at a depth no shallower than 100 m and the
related Musashia ( Nipponomelon ) is common in that part of
the unit. Musashia ( Miopleiona ) indurata is usually found
in the finer-grained, deeper-water facies of the Astoria For-
mation. I suggest that the molluscan fauna in the Eugene
Formation lived in shallower water than was common for
Miopleiona.

Turridae

Aforia wardi (Tegland)

Figure 99

Leucosyrinx clallamensis wardi Tegland, 1933:124, pi. 10,
figs. 5-8.

Aforia wardi is of medium size and pagodaform, with nine
strongly angulated whorls and a U-shaped sinus on the shoul-
der (Fig. 99). The shell is smooth above the angulations but
sculptured by fine spiral threads below, and these spirals
extend down the siphonal canal on the body whorl. Aforia
campbelli (Durham, 1944:183, pi. 14, fig. 4) differs in having
the whorl angulation closer to the suture, and the angulation
is rounded rather than bladelike. Javidpour (1973) discussed
the phylogeny of eastern Pacific Tertiary species of Aforia.
The correlation diagram (Javidpour, 1973:198, fig. 17) is
misleading in that Aforia campbelli is shown in the upper
Oligocene part of the Lincoln Creek Formation, whereas it
should have been placed in the middle Oligocene part of the
unit as stated in the text (Javidpour, 1973:196, 199-200).
Aforia was placed in the subfamily Turriculinae by McLean
(1971:119), following Powell ( 1 942).

Living species of Aforia in the eastern Pacific have been
recorded from depths of 6 to 2870 m (Abbott, 1 974:265) and
in the western Pacific from depths of 55 to 90 m (Kira, 1962:
102). Powell (1969:411-414) said that Aforia prefers cold
water ranging from —0.6° to +5.4°C and is bipolar, going
deeper under equatorial waters.

Based on the illustrations by Powell (1969:411, pi. 322,
figs. 1-4; 414, pi. 323, figs. 1-3) of the type species of Aforia,
Pleurotoma circinata Dali, characters such as apical angle
and position of whorl angulation are not useful in distin-
guishing species. If larger collections of well-preserved spec-
imens of Aforia become available, future workers may see
fit to synonymize some of the species proposed for eastern
Pacific Tertiary Aforia.

OCCURRENCE ELSEWHERE. Lower part of the Blak-
eley Formation in the Liracassis rex Molluscan Zone, Wash-
ington.

Turriculal sp.

Figure 97

Turriculal sp. is represented by one incompletely preserved
specimen on which the siphonal canal is not exposed. The
sinus is U-shaped and confined to the shoulder slope. The
shell is sculptured by moderately strong spiral cords that are
not noded. Turricula washingtonensis { Weaver, 1912:78, pi.

24 Contributions in Science, Number 351

Moore: Lincoln Creek Formation Mollusks

3, fig. 31; 1942:533, pi. 98, figs. 16, 17, 22) differs from
Turriculal sp. in having nodes.

Megasurculal sp.
cf. M. wynoocheensis (Weaver)

Figure 96

Megasurculal sp. cf. M. wynoocheensis (Weaver) is repre-
sented by a single poorly preserved and somewhat crushed
specimen. Megasurcula wynoocheensis (Weaver, 1912:70-
71, pi. 11, figs. 87-89, 94) is a middle Miocene and possibly
early Miocene species (Addicott, 1976c:27,pl. 3, figs. 16, 17).

Actenoidae

Microglyphus ? n. sp.?

Figure 98

Microglyphus ? n. sp.? has three or possibly four whorls and
a body whorl that is very inflated with the maximum width
at the middle of the whorl. The spiral cords, bounded by
incised grooves, are not equidimensional. The single speci-
men is very small (2. 1 mm high, 1 .9 mm wide) and, although
it may be an immature individual, the number of whorls
indicates that it probably is a very small species. In addition
to its small size, Microglyphus ? n. sp.? differs from other
described Tertiary actenoids in having a more globose body
whorl with the maximum inflation at the middle of the whorl.

PELECYPODS

Nuculidae

Acila ( Acila ) gettysburgensis (Reagan)

Figures 100-102, 105

Nucula {Acila) gettysburgensis Reagan, 1909:171, 175, 177,
pi. 1, fig. 3.

Acila (Acila) gettysburgensis is represented by five specimens.
Acila (A.) gettysburgensis ranges from the Matlockian through
the Pillarian Molluscan Stages.

Nuculanidae

Portlandia ( Portlandia ) chehalisensis (Arnold)

Figures 103, 104, 106, 107

Malletia chehalisensis Arnold, 1908:365, pi. 33, fig. 9.

Portlandia ( Portlandia ) chehalisensis is represented by seven
specimens, some well preserved (Fig. 107) and one double-
valved (Figs. 103, 104). One incomplete specimen has con-
centric Saccella- like ridges on the midportion of the shell
near the ventral margin (Fig. 106). One single valve is 31.8
mm long, 18.5 mm high, and 7.0 mm thick; perhaps the
largest specimen of the species collected. The largest speci-
men noted by Hickman (1969:3 1) measured 26 mm in length.

Yoldia reagani Dali (1922:306) was considered a synonym
of Portlandia chehalisensis (Hickman, 1969:30).

Living eastern Pacific species of Portlandia occur no farther
south than latitude 54°N and are found at depths of 10 to
2560 m and temperatures from —2° to +6°C (Bernard, 1983:
13).

Solemyidae

Acharax dal/i (Clark)

Figures 108-1 11, 114

Solemya dalli Clark, 1925:73, pi. 9, fig. 3.

Acharax dalli is represented by six specimens, all but one
double-valved. Fingerlike projections of the periostracum are
partially preserved on some specimens (Fig. 108). Acharax
ventricosa (Conrad, 1849:723, pi. 17, figs. 7, 8), a species
found in the Astoria Formation in Oregon and Washington,
is higher in proportion to length than A. dalli.

The eastern Pacific Holocene species Acharax johnsoni
(Dali, 1891) lives at a depth between 800 and 3000 m at
temperatures of 1° to 9°C (Bernard, 1983:9). Vokes (1955:
536-537) said that living species of Acharax are found at
depths of 5 to 3 1 80 m and that the controlling factor in their
distribution may be water temperature.

Limopsidae

Limopsis nitens (Conrad)

Figures 112, 113, 115, 116

Pectunculus nitens Conrad, 1849:726, pi. 18, figs. 9a-b.

Limopsis nitens occurs as numerous single valves (Figs. 1 12,
1 13) and occasional paired valves (Figs. 115, 116). Radial
lines of sculptural punctures are preserved on some speci-
mens.

The lithology of the concretions from Knappton and the
clustering together of many specimens is similar to the con-
cretion presumably from the Astoria Formation, at Astoria,
Oregon, that contains the lectotype of L. nitens (Moore, 1963:
61-62, pi. 15, figs. 2, 5). Weaver (1942:76) suggested that
the lectotype was collected at Knappton, rather than at As-
toria, because he had found nodules containing large numbers
of L. nitens at Knappton and had not found any specimens
at Astoria. Howe (1922:70) did not find any specimens of L.
nitens at Astoria and I found none in the Astoria Formation
farther south (Moore, 1963:62). The rock containing the lec-
totype of L. nitens may have come from Knappton, or the
upper part of the Lincoln Creek Formation may have been
exposed on the Columbia River terrace at Astoria when Dana
made his collection in 1841, yet no other mollusks typical
of the Lincoln Creek Formation were collected by him.

Most species of Limopsis live in deep water (Keen, 1971:
54); Limopsis diegensis Dali has been collected at depths of
120 to 1500 m and at temperatures between 3° and 27°C
(Bernard, 1983: 1 7). The fact that Limopsis nitens most com-

Contributions in Science, Number 351

Moore: Lincoln Creek Formation Mollusks 25

monly occurs in clumps suggests that the species was gre-
garious.

Mytilidae

Modiolus addicotti n. sp.

Figures 117, 125

Modiolus n. sp.? aff. M. restorationensis Van Winkle. Ad-
dicott, 1976c:28, pi. 5, fig. 5.

Modiolus addicotti is a rather small Modiolus with weakly
inflated valves, a markedly thin shell, and a convex dorsal
margin. The posterior end is only moderately enlarged, and
slightly longer near the ventral margin. The anterior end is
small and evenly curved. The umbones are close to the an-
terior margin. Patches of preserved shell are light brown and
iridescent with mostly evenly spaced growth lines but with
a few bunched together forming low ridges.

HOLOTYPE. L ACM IP 6672, length 47 mm, height 25
mm.

TYPE LOCALITY. LAM 5842.

Modiolus addicotti differs from Modiolus restorationensis
Van Winkle ( 1 9 1 8:82, pi. 4, fig. 5) in having a convex dorsal
margin and a narrower posterior end.

Modiolus lives intertidally to 360 m in the eastern Pacific,
but most species are found at depths no greater than 50 m
(Bernard, 1983:19).

OCCURRENCE ELSEWHERE. Lowermost part of the
Clallam Formation, northwestern Washington.

This species is named in honor of Warren O. Addicott.

Limidae

Acesta ( Acesta ) twinensis (Durham)

Figures 1 19, 123, 124

Lima twinensis Durham, 1944:139, pi. 13, fig. 11.

Acesta ( Acesta ) twinensis is represented by one incomplete
double-valved specimen and four incomplete single valves

all of which are preserved intact with their original inflation.
The anterior ears are small and well defined by a deep con-
cave groove along the anterior margin (Fig. 1 24). The anterior
margin is straight, not concave, and joins the ventral margin
without an abrupt break in alignment. The posterior ears are
large and indistinctly delineated (Figs. 1 19, 124). The shells
are large (maximum estimated height 140 mm), thin, and
smooth in the center but with rounded ribs of varying widths
at the shell margins (Fig. 124). The shells are inflated, and
the largest specimen suggests a thickness of 25 mm (one
valve). Portions of the brown translucent outer shell layer
are preserved on most specimens, but this shell layer tends
to stay attached to the enclosing rock when the specimens
are broken away.

Acesta twinensis is distinguished by its sharply truncated
anterior margin, which differentiates it from Acesta robertsae
(Durham, 1944), an early Oligocene species that has a more
rounded anterior margin.

Acesta ( Acesta ) oregonensis Clark (1925:84, pi. 14, figs. 3,
4), a species from the upper Eocene and lower Oligocene
Keasey Formation, Oregon, has an arcuate anterior margin.

Acesta twinensis ranges from the Matlockian through the
Juanian Molluscan Stages. Living species of Acesta ( Acesta )
are found in the eastern Pacific at depths between 600 and
2200 m and at temperatures of 1° to 8°C (Bernard, 1983:22).
One species has been collected in the western Pacific near
Japan at a depth of 185 m.

OCCURRENCE ELSEWHERE. Blakeley Formation,
Washington.

Acesta ( Plicacesta ) wilsoni n. sp.

Figures 118, 132, 134

Acesta ( Plicacesta ) wilsoni is of moderate size and subovate
in outline, with a thicker shell than Acesta ( Acesta ) and radial
ribs of varying widths and spacing that are rounded and most
prominent on the middle portion of the shell but that persist
to the shell margins. Beaks small; anterior auricle presumed
small and delineated; posterior auricle large and not delin-

Figures 96-117. MegasurculaP sp. cf. wynoocheensis (Weaver), Turricula ? n. sp.?, Microglyphus ? n. sp.?, Aforia wardi (Tegland), Acila ( Acila )
gettysburgensis (Reagan), Portlandia (Portlandia) chehalisensis (Arnold), Acharax dalli (Clark), Limopsis nitens (Conrad), and Modiolus
addicotti n. sp.

Figure 96. MegasurculaP sp. cf. wynoocheensis (Weaver). Rear view, x 1.5; height 32 mm, width 20 mm. USGS Loc. 7891, USNM 363990.

Figure 97. Turricula ? n. sp.? Showing spiral sculpture, x2.0; height 15 mm, width 10 mm. LACMIP 6657.

Figure 98. Microglyphus ? n. sp.? Showing outline and spiral sculpture, x5.0; height 2.7 mm, width 2.4 mm. LACMIP 6658.

Figure 99. Aforia wardi (Tegland). Showing pagodaform outline, x 1.5; height 28 mm, width 14 mm. LACMIP 6659.

Figures 100-102, 105. Acila (Acila) gettysburgensis (Reagan). 100. Showing primary bifurcation, x 2; length 13 mm, height 1 1 mm. LACMIP
6660. 101. Showing sulcus xl.5; height 17 mm. LACMIP 6661. 102. Showing secondary bifurcation, x2. LACMIP 6662. 105. Showing
outline and bifurcation, x 1.5; length 25 mm, height 20 mm. LACMIP 6663.

Figures 103, 104, 106, 107. Portlandia (Portlandia) chehalisensis (Arnold). 103, 104. Double-valved specimen showing sculpture and
lunule, x 1.5; length 21 mm, height 12 mm, width 8 mm (both valves). LACMIP 6664. 106. Showing Saccella- like sculpture, x2; length 22
mm, height 13 mm. LACMIP 6665. 107. Showing outline and sculpture, x 1.5; length 32 mm, height 18 mm. LACMIP 6666.

Figures 108-111, 114. Acharax dalli (Clark). 108, 114. Showing extensions of fingerlike periostracum and dorsal side; length 28 mm, height
24 mm, width 16 mm (both valves). LACMIP 6667. 109, 110. Showing sculpture of double-valved specimen, x 1.5; length 45 mm, height 20
mm, width 1 1 mm (both valves). LACMIP 6668. 111. Showing sculpture x 1.5; length 35 mm, height 13 mm. LACMIP 6669.

Figures 112, 113, 115, 116. Limopsis nitens (Conrad). 112. Showing outline, x3; 10 mm long, 7 mm high. LACMIP 6670. 113. Showing
radial punctae, x3; length 9 mm, height 7 mm. LACMIP 6671. 115, 116. Double-valved specimen showing thickness of valves, x5; length
8 mm, height 6 mm, width 5 mm (both valves). USGS Loc. 7891, USNM 363991.

Figure 117. Modiolus addicotti n. sp. Holotype LACMIP 6672. Showing outline of valve; length 47 mm, height 25 mm.

26 Contributions in Science, Number 351

Moore: Lincoln Creek Formation Mollusks

115

110

Contributions in Science, Number 351

Moore: Lincoln Creek Formation Moilusks 27

eated. No concentric sculpture is preserved. The holotype
(incomplete) is 60 mm high and 50 mm wide and the thick-
ness of one valve is 13 mm (Figs. 1 18, 132). The paratype
(incomplete) is 63 mm high and 40 mm wide and the thick-
ness of one valve is 1 1 mm (Fig. 134).

HOLOTYPE. LACMIP 6673, length 51 mm, height 61
mm, width 10 mm (one valve); paratype LAM 6686, length
41 mm, height 61 mm.

TYPE LOCALITY. LAM 5842.

Acesta wilsoni is smaller than Acesta (Plicacesta) oakvil-
lensis (Clark, 1925:84, pi. 15, figs. 1, 3) and has fewer ribs
and a more rounded anterior margin. A topotype specimen,
earlier illustrated by Weaver (1942, pi. 21, fig. 1), is figured
for comparison (Fig. 126). Acesta cf. A. oakvi/lensis Clark of
Addicott (1976b, figs. 6a, c) has narrower ribs that are more
widely and evenly spaced than on A. wilsoni and may be a
new species.

Acesta wilsoni and A. oakvi/lensis are the only Tertiary
species of Plicacesta known in North America. Acesta n. sp.
of Addicott (1976b, figs. 6x, z, ab) may also be a Plicacesta.

The Holocene species Acesta ( Plicacesta ) sphoni (Hertlein,
1963) was collected at latitude 33°N at a depth between 455
and 550 m and a temperature between 4° and 9°C. Acesta
(Plicacesta) smithi (Sowerby) occurs off Honshu, Japan, at
depths between 90 and 185 m (Kira, 1962:145).

This species is named in honor of Edward C. Wilson.

Ostreidae

Crassostrea! sp.

Figure 127

Crassostrea ? sp. is represented by a single double-valved
specimen preserved in a concretion and broken upon re-
moval from the matrix. Indigenous Holocene species of Cras-
sostrea live intertidally to a depth of 7 m in the eastern Pacific
(Bernard, 1983:23).

Lucinidae

Lucinoma hannibali (Clark)

Figures 120-122

Phacoides ( Lucinoma ) hannibali Clark, 1925:89, pi. 22, figs.

2, 4.

Lucinoma hannibali is represented by six double-valved
specimens (Figs. 1 20-122) from the upper part of the Lincoln

Creek Formation, one with the shell replaced by barite. The
specimens range in height from 27 mm to 52 mm. Lucinoma
acutilineata (Conrad, 1849:725, pi. 18, figs. 2, 2a, 2b) has a
shorter more concave dorsal margin than L. hannibali. Vari-
ation has been noted (Moore, 1963:70) in the spacing of
concentric lamellae within single lots of the Holocene species
Lucinoma annulata (Reeve, 1850) and by Addicott (1976c:
30) in the Oligocene to Miocene species L. acutilineata, yet
specimens of L. hannibali from the upper part of the Lincoln
Creek Formation have concentric lamellae rather consis-
tently less densely spaced (Figs. 120-122) than the lamellae
on L. acutilineata from the lower part of the Astoria For-
mation. Lucinoma acutilineata has been found in the Eugene
Formation (upper Eocene to middle Oligocene) in Oregon
(Hickman, 1969:38, 42) and in the lower and middle Mio-
cene Astoria Formation (Moore, 1963:70-71, pi. 15, figs. 7-
10, 12) in Oregon and Washington. If L. hannibali and L.
acutilineata are distinct species, and I believe that they are,
L. acutilineata may have preferred somewhat shallower water
(50 m or less) than L. hannibali, and the two species coexisted
at different depths. Lucinoma annulata lives today from lat-
itude 3 3° to 47°N at depths of 2 5 to 750 m and L. aequizonata
(Steams, 1891) lives from latitude 34° to 37°N at depths of
400 to 650 m (Bernard, 1983:29). Lucinoma hannibali has
a wider escutcheon and less concave dorsal margin than L.
columbiana (Clark and Arnold, 1923:144-145, pi. 25, figs.
2a-b) from the Sooke Formation, Vancouver Island, and the
Blakeley Formation of Weaver (1912). Lucinoma hannibali
ranges from the Matlockian through the Juanian Molluscan
Stages.

Thyasiridae

Thyasira ( Conchocele ) disjuncta (Gabb)

Figures 136, 138, 142

Conchocele disjuncta Gabb, 1866:28; 1869:99, pi. 7, figs.

48a-b.

Thyasira (Conchocele) disjuncta is larger, more quadrate, and
has a more truncated anterior end than Thyasira bisecta
(Conrad, 1849:724, pi. 17, figs. 10, 10a) from the Astoria
Formation in Oregon (Moore, 1963:72, pi. 23, figs. 8, 14,
1 5). Thyasira disjuncta occurs in the Clallam Formation (Ad-
dicott, 1976c:30, pi. 6, fig. 7) and is living today (Bernard,
1983:29). The presence of two internal casts (Figs. 136, 138)
and one specimen with the outer shell preserved (Fig. 142)

Figures 118-125. Acesta (Plicacesta) wilsoni n. sp., Acesta ( Acesta ) twinensis (Durham), Lucinoma hannibali (Clark), and Modiolus addicotti
n. sp.

Figure 118. Acesta (Plicacesta) wilsoni n. sp. showing outline and radial ribs, x 1.5; length 51 mm, height 61 mm, width 10 mm (one
valve). Holotype LACMP 6673.

Figures 119, 123, 124. Acesta (Acesta) twinensis (Durham). 119, 123. Showing posterior ears. 119. Length 74 mm, height 93 mm, width
22 mm (one valve). LACMP 6675. 123. Length 92 mm, height 75 mm. LACMP 6674. 124. Showing anterior ear and axial ribs, x 1.5; length
60 mm, height 70 mm. LACMP 6676.

Figures 120-122. Lucinoma hannibali (Clark). 120. Showing profile of lunule; length 29 mm, height 27 mm, width 1 1 mm (both valves).
LACMP 6677. 121. Showing sulcus; length 39 mm, height 34 mm, width 16 mm (both valves). LACMP 6678. 122. Showing concentric
sculpture; length 40 mm, height 37 mm, width 27 mm (both valves). LACMP 6679.

Figure 125. Modiolus addicotti n. sp. Holotype LACMP 6672. Tipped to show configuration of double-valved specimen, x 1.5; length 47
mm, height 25 mm. Same specimen shown in Figure 1 17.

28 Contributions in Science, Number 351

Moore: Lincoln Creek Formation Mollusks

in the upper part of the Lincoln Creek Formation extends
the range of the species downward into the upper part of the
Juanian Molluscan Stage. Thyasira disjuncta lives today from
latitude 48° to 54°N at depths of 100 to 750 m and at tem-
peratures between 0° and 7°C. Thyasira bisecta lives from
latitude 43° to 57°N at depths between 50 and 300 m and
temperatures between —1° and +11°C (Bernard, 1983:29).
The occurrence of T. disjuncta in the Lincoln Creek For-
mation and of T. bisecta in the Astoria Formation is related
to depth of water; the Astoria Formation represents a shal-
lower-water facies than the Lincoln Creek Formation.

Cardiidae

Nemocardium ? sp. cf. N. lorenzanwn (Arnold)

Figure 130

Nemocardium ? sp. cf. N. lorenzanum (Arnold, 1908:366, pi.
33, fig. 6) is represented by two poorly preserved single valves;
one is illustrated (Fig. 1 30).

Tellinidae

Macoma sp. cf. M. twinensis Clark

Figure 128

Macoma sp. cf. M. twinensis Clark is represented by a single
valve that does not have the hinge exposed (Fig. 128). In

size, outline, and position of umbo, the specimen resembles
M. twinensis Clark (1925:96, pi. 12, fig. 7), which ranges
from late Oligocene to early Miocene.

Periplomatidae

Cochlodesma bainbridgensis Clark

Figures 139, 141

Cochlodesma bainbridgensis Clark, 1925:86, pi. 13, figs.
3, 4.

Cochlodesma bainbridgensis Clark has a thin, internally
nacreous, fragile shell, that is sculptured with concentric un-
dulations (Figs. 139, 141). Five specimens and one possible
juvenile (Fig. 131) were collected from the upper part of the
Lincoln Creek Formation. As noted by Moore (1976:53, pi.
16, figs. 4, 6-11), the variation in outline is great and not
useful in the discrimination of fossil species. Holocene species
of the closely related genus Periploma live intertidally to a
depth of 380 m (Bernard, 1983:64).

Teredinidae

Figures 172, 177, 179

Teredinid burrows are preserved in wood and the tubes are
filled with quartz (Fig. 1 72) or with sediment (Figs. 177, 179).
No pallets are preserved, enabling generic differentiation, but

Figures 126-1 34. Acesta ( Plicacesta ) oakvillensis (Clark), Crassostreal sp. , Macoma sp. cf. M. twinensis Clark, Flabellum sp. , Nemocardium ?
sp. cf. N. lorenzanum (Arnold), Cochlodesmal sp., Lima ( Plicacesta ) wilsoni n. sp., and Aturia angustata (Conrad).

Figure 126. Acesta ( Plicacesta ) oakvillensis (Clark). Showing closely spaced axial ribs; length 79 mm, height 95 mm. UC Loc. A368, lower
part of the Lincoln Creek Formation, Grays Harbor County, Washington. UCMP 32405.

Figure 127. Crassostreal sp. Showing configuration; length 70 mm, height 102 mm. LACMP 6680.

Figure 128. Macoma sp. cf. M. twinensis Clark. Showing outline and concentric lines, x 1.5; length 15 mm, height 10 mm. LACMP 6681.
Figure 129. Flabellum sp. x 1.5; height 22 mm. LACMP 6682.

Figure 130. Nemocardium ? sp. cf. N. lorenzanum (Arnold). Showing configuration and radial ribs, x 1.5; height 20 mm. LACMP 6683.
Figure 131. Cochlodesmal sp. Showing outline, x2; length 1 1 mm, height 8 mm. LACMP 6684.

Figures 132, 134. Lima ( Plicacesta ) wilsoni n. sp. 132. Showing outline and radial ribs, x 1.5; length 51 mm, height 61 mm, width 10 mm
(one valve). Holotype LACMP 6673. 134. Showing radial ribs x 1.5; length 41 mm, height 61 mm. Paratype LACMP 6686.

Figure 133. Aturia angustata (Conrad). Immature specimen, x 1.5; height 24 mm. LACMP 6687.

Figures 135-145. Aturia angustata (Conrad), Thyasira ( Conchochele ) disjuncta (Gabb), and Cochlodesma bainbridgensis Clark.

Figures 135, 137, 140, 143-145. Aturia angustata (Conrad). 135. Showing outer shell and growth lines, x0.7; height 150 mm. LACMP
6688. 137. Cross section showing funnel-shaped septal structures; height 29 mm. LACMP 6689. 140. Cross section showing septal structures;
height 37 mm. LACMP 6690. 143. Apertural view of broken specimen showing siphuncular orifices; height 90 mm. LACMP 6691. 144. Cross
section showing septal structures; height 60 mm. LAM Loc. 5843, LACMP 6692. 145. Side view showing sutures; height 65 mm, width 27
mm (maximum diameter). Same specimen shown in Figure 150. LACMP 6693.

Figures 136, 138, 142. Thyasira ( Conchochele ) disjuncta (Gabb). 136. Showing sulcus and configuration; length 70 mm, height 67 mm.
LACMP 6694. 138. Showing sulcus and configuration; length 52 mm, height 43 mm. LACMP 6695. 142. Showing outer shell and concentric
lines, x 1.5; length 37 mm, height 32 mm. LACMP 6696.

Figures 139, 141. Cochlodesma bainbridgensis Clark. 139. Showing concentric undulations, x 1.5; length 34 mm, height 27 mm. LACMP
6697. 141. Showing outline, x 1.5; length 30 mm, height 25 mm. LACMP 6698.

Figures 146-154. Aturia angustata (Conrad) and a crab claw.

Figures 146, 147, 149, 150, 152-154. Aturia angustata (Conrad). 146. Thin section showing radiating calcite within phragmocone chamber
in crossed-polarized light, x70. LACMP 6699. 147. Showing suture, x0.8; height 95 mm. LAM Loc. 5287, LACMP 6700. 149. Showing
silicified siphuncular neck and orifice; height 43 mm. LACMP 670 1 . 150. Showing sutures; height 65 mm, width 27 mm (maximum diameter).
Same specimen shown in Figure 145. LACMP 6693. 152. Broken specimen showing siphuncular orifice and neck; height 1 14 mm. LACMP
6702. 153. Cross section showing funnel-shaped siphuncular necks, x2.0; length 45 mm. LACMP 6703. 154. Broken specimen showing
siphuncular orifice; height 75 mm. LACMP 6704.

Figures 148, 151. Crab claw; length 40 mm. LACMP 6705.

30 Contributions in Science, Number 351

Moore: Lincoln Creek Formation Mollusks

i \ V-

, r;?! a

132

137

143

144

145

Contributions in Science, Number 351

Moore: Lincoln Creek Formation Mollusks 33

on the basis of the size and configuration of the burrows, two
different genera may be present.

SCAPHOPOD

Dentaliidae

Denta/ium ( Fiss ide ntalium 7 ) sp.
cf. D. porter ensis (Weaver)

Dentalium ( Fiss identalium?) sp. cf. D. porterensis (Weaver,
1912:79, pi. 13, fig. 1 13) is circular in cross section, slightly
tapered, and has perhaps 32 (16 exposed on one half) fine
radial riblets crossed by strong concentric threads that pro-
duce a basket-weave sculpture. The preservation does not
permit comparison with other ribbed Tertiary dentaliids.

CEPHALOPODS

Aturidae

Aturia august at a (Conrad)

Figures 133, 135, 137, 140, 143-145, 146,

147, 149, 150, 152-154, 155-159

Nautilus angustatus Conrad, 1848:728, pi. 20, figs. 5, 6.

Aturia angustata has been described in detail by Schenck
(1931:457-462) and Miller (1947:85-88), and the type spec-
imen figured and discussed by Moore (1963:85-86, pi. 31,
figs. 1, 5).

A total of 1 80 specimens of A. angustata is in the Knappton
collections, making this cephalopod by far the most abundant
mollusk collected. Taking into account a possible bias in
favor of collecting Aturia, this is still a large number. Kummel
( 1 956:330-33 1 ) called attention to the rarity of post-Triassic
nautiloids saying that no large collection representing a pop-
ulation had ever been assembled from a single horizon and
locality. Stenzel (in Ladd, 1957:893) noted that there are in
excess of 1000 mollusks representing other classes for every
nautiloid shell and that the proportion may actually surpass
10,000 to 1.

The shell of the preserved portion of the living chamber
is commonly slightly broken but more frequently is intact.
The outer shell layer is dark brown and the entire shell or
venter is thin, thinner than that of the living Nautilus. Faint,
closely spaced growth lines can be seen on some specimens
(Fig. 135). The lateral lobes are tongue-shaped and ascending
on young specimens (Figs. 133, 145, 150), but not on more
mature specimens (Fig. 147).

The specimens are believed to range in size from 30 to
180 mm in greatest diameter. The smallest specimens (25
mm) are not complete and so were probably 5 to 10 mm
larger, and the largest specimen measures 170 mm but is

incomplete and has an estimated size of 1 80 mm. The largest
number of specimens sufficiently complete to make size mea-
surement meaningful (22%) are 90 mm in greatest diameter.
Presumably, this means that many of the specimens had not
reached maturity before death. About 24% of the specimens
are 100 to 180 mm in maximum diameter and are assumed
to have been mature. The specimens are not crushed, and
none shows any indication that it imploded as a result of
having been transported to great depths.

The suture is simple with a broad flattened ventral saddle,
a narrow pointed lateral lobe on the umbilical slope and
dorsal area, and a broad saddle on the dorsal area divided
by a deep, narrow lobe (Figs. 145, 147, 150). The siphuncle
is moderate in size, subdorsal and marginal in position (Figs.
137, 140, 143, 144, 149, 152, 154, 155, 157), and located
near the apex in the adapical flexure of the septa. The si-
phuncular tube consists of a series of cone-in-cone necks, or
long funnel-shaped connecting rings (Fig. 157) without the
long gaps between the necks that are present in Nautilus.

The phragmocone chambers may be filled with sediment
(Fig. 1 54) but are more commonly partially filled with calcite
or completely filled with calcite, barite, quartz, or combi-
nations of these minerals (Fig. 1 53). A phragmocone chamber
of one specimen is filled with glauconite. Some of the spec-
imens have empty phragmocone chambers except for a cal-
cite buttress, and these chambers may be followed or pre-
ceded by sediment-filled chambers, indicating that the
sediment did not enter through the siphuncular tube but
entered through a puncture in the shell. The body chamber,
of course, is always filled with sediment.

Most of the shells of Aturia angustata are preserved in
concretions as almost complete specimens, but some are frag-
ments that may have weathered out of concretions or not
have been so preserved. The specimens that show a sequence
of mineralization, which is currently being studied in detail,
begin with a buttressing of the shell walls with as many as
nine layers of radial calcite (Fig. 146), followed by the dis-
solution of the aragonitic shell, and then the filling of the
shell cavities and the remaining chamber voids with calcite,
barite, and/or quartz in that sequence.

Sepiidae?

A trace fossil that may represent the cuttlebone of a sepiid
is illustrated (Fig. 178).

FOSSILS OTHER THAN MOLLUSKS
SPONGES

Two sponges have been described by J.K. Rigby and D.E.
Jenkins (1983) from the upper part of the Lincoln Creek
Formation: Aphrocallistes polytretos (Fig. 180) and Eurete

Figures 155-159. Aturia angustata (Conrad). 155. Broken specimen showing siphuncular orifices, x3; height 80 mm. LACMP 6706. 156.
Showing phragmocone chambers filled with wood fragments and sediment. LACMP 6707. 157. Silicified specimen etched in dilute hydrochloric
acid to show cone-m-cone, funnel-shaped septal necks, x 1.5; 47 mm greatest diameter. LACMP 6708. 158. Fecal pellets probably formed by
a manne worm in phragmocone chamber, x3. LACMP 6709. 159. Fecal pellets in phragmocone chamber, x5. LACMP 6710.

34 Contributions in Science, Number 351

Moore: Lincoln Creek Formation Mollusks

Contributions in Science, Number 351

Moore: Lincoln Creek Formation Mollusks 35

goederti. In addition, hexactinellid root tuffs were identified
that may represent a third sponge. Although the sponges are
most common stratigraphically just below the major mollusk
locality, they also occur within it. Aphrocallistes lives at depths
of 100 to 1700 m (Schulze, 1887) and Eurete between 220
and 715 m, with the majority of species living at depths
between 300 and 360 m (Rigby and Jenkins, 1983).

CORALS

Dendrophyllia hannibali Nomland (1916:67, pi. 6, figs. 1-3)
was found in one concretion that also contains abundant fish
debris and a small patch of the siliceous sponge, Eurete goe-
derti Rigby and Jenkins (1983). The specimens are poorly
exposed and recrystallized (Fig. 175), but some show septa
(Fig. 176).

Dendrophyllia hannibali was described by Nomland (1916:
67) as colonial, branching, and forming several vertical series
that unite when coming in contact. It has deep nearly round
calices with about 42 to 48 septa. The maximum number of
preserved septa counted on the specimens described by Nom-
land is about 20, but recrystallization is believed to have
destroyed many of the septa.

Dendrophyllia is a scleractinian, ahermatypic (nonreef-
building; capable of living in cold deep water) coral. Ac-
cording to Wells (1956:F362, F435), “the greatest develop-
ment of ahermatypic corals occurs near and down the edges
of continental slopes and the equivalent bathymetric zone
around oceanic islands in depths from 175 to about 800 m
. . . in temperatures of 4° to 21°C.” Dendrophyllia is cos-
mopolitan in its modern distribution and is known from the
Eocene through the Holocene at depths ranging from 0 to
1370 m.

Flabellum sp. (Fig. 129) has also been collected.

BRACHIOPOD

Laqueusl sp. cf. L. vancouverensis Davidson is poorly pre-
served and only three (or possibly four) specimens have been

collected. The outer shell is smooth (Figs. 166, 168, 169), or
may possibly on some specimens be finely ribbed (Fig. 167),
and the inner fibrous layer is punctate. The specimens re-
semble L. vancouverensis (Davidson, 1887:1 13, pi. 18, figs.
10-1 3b) more than any other described species. Gradational
variation in sculpture between subspecies of the terebratellids
(Hertlein and Grant, 1 944: 1 32) seems to be sufficient to per-
haps allow for both smooth-shelled and finely ribbed forms
in one species.

ECHINOIDS

Most of the echinoids (Figs. 160-162, 164, 170) are tests of
a spatangoid (heart urchin) that was probably buried in living
position since so many of its spines are attached (Porter M.
Kier, written commun., 1980). The species probably lived
in a burrow at a depth of one to several centimeters within
the sediment. All the tests are broken, perhaps by the weight
of the overburden as the attached spines suggest the speci-
mens were not transported. The echinoids have not been
found in the center of spherical concretions typical of the rest
of the fauna.

A single specimen thought to be a madreporite, a sievelike
structure that provides access to the water-vascular system
(Fig. 163), was also collected.

LOCALITIES

NATURAL HISTORY MUSEUM OF LOS ANGELES
COUNTY

5787. From landslide block in upper part of the Lincoln
Creek Formation between Knappton and Grays Point, NW
'/» sec. 9, T. 9 N., R. 9 W., Knappton 71/2-minute quadrangle
(1973 edition), on the Columbia River, Washington. (Gen-
eral locality that includes 5842, 5843, 5844, and 5852.)

5802. From a limestone quarry in the siltstones of Cliff
Point unit (Wells, 1979) in the bluff on the south side of Bear
River, 2.3 km northeast of Goulter Ranch, on the section
line between secs. 20 and 21, T. 10 N., R. 10 W., Chinook

Figures 160-170. Spatangoid echinoids and a brachiopod.

Figures 160-162, 164, 170. Spatangoid echinoids. 160. Showing test outline and spines, *6.0; 8 mm greatest diameter of test. LACMP
6491. 161. Showing ambulacral area and spines, x3.0. LACMP 6710. 162. Showing test outline and attached spines, x3.0; 20 mm greatest
diameter of test. LACMP 6492. 164. Showing broken test with preserved ambulacral areas, x2.0; 52 mm greatest diameter of test. LACMP
6711. 170. Showing outline of several tests with associated wood fragments. LACMP 6712.

Figure 163. Madreporite? of spatangoid echinoid, x 12.0; 2.4 mm greatest diameter. LACMP 6713.

Figures 165-169. Laqueus ? sp. cf. L. vancouverensis Davidson. 165. View of apex, xl.5. LACMP 6714. 166. Showing configuration and
narrow axial ribs; length 23 mm, width 23 mm. LACMP 6715. 167. Showing radial ribs, x 1.5; width 23 mm. LACMP 6716. 168. Showing
configuration; height 17 mm. LACMP 6717. 169. Showing configuration and smooth shell; length 35 mm, width 34 mm. Same specimen
shown in Figure 165. LACMP 6714.

Figures 171-180. Teredinid bores, crab claw, Dendrophyllia hannibali Nomland, a trace fossil, and Aphrocallistes polytretos Rigby and
Jenkins.

Figures 171, 172, 177, 179. Teredinid-bored wood. 171. View of bored wood within Aturia. Same specimen shown in Figure 156, x3.0.
LACMP 6707. 177, 179. Teredinid tubes in wood. LACMP 6716. 177. Cross-sectional view. 179. Longitudinal view, xQ.8.

Figures 173, 174. Crab claw showing nodes, x 1.5; length 35 mm. USNM 363992.

Figures 175, 176. Dendrophyllia hannibali Nomland. 175. Showing configuration, xl.5. LACMP 6719. 176. Showing septa, x 3.0. LACMP
6720.

Figure 178. Trace fossil, possibly cuttlebone of sepiid, x 1.5; length 45 mm. LACMP 6721.

Figure 180. Aphrocallistes polytretos Rigby and Jenkins. Showing irregular branching growth. LACMP 6722.

36 Contributions in Science, Number 351

Moore: Lincoln Creek Formation Mollusks

Contributions in Science, Number 351

Moore: Lincoln Creek Formation Mollusks 37

38 Contributions in Science, Number 351

Moore: Lincoln Creek Formation Mollusks

7'/2-minute quadrangle, about 1 5 km northwest of Knappton,
Pacific County, Washington.

5842. From landslide block in upper part of the Lincoln
Creek Formation between Knappton and Grays Point, in the
center of the N 'A, N V2 sec, 9, T. 9 N., R. 9 W„ Knappton
7'/2-minute quadrangle, on the Columbia River, Washington.

5843. From landslide block in upper part of the Lincoln
Creek Formation between Knappton and Grays Point, 305
m south and 430 m east of NW cor. sec. 9, T. 9 N., R. 9 W.,
Knappton 7 '/2-minute quadrangle, on the Columbia River,
Washington.

5844. From landslide block in upper part of the Lincoln
Creek Formation between Knappton and Grays Point, 122
m east and 520 m south of NW cor. sec. 9, T. 9 N., R. 9 W.,
Knappton 7 ‘/2-minute quadrangle, on the Columbia River,
Washington.

5852. From landslide block in upper part of the Lincoln
Creek Formation between Knappton and Grays Point, NE
'/», NW 'A sec. 9, T. 9 N., R. 9 W., Knappton 7 '/2-minute
quadrangle, on the Columbia River, Washington.

5863. From the Astoria Formation between Knappton and
Grays Point, SE 'A, NW ‘A sec. 10, T. 9 N„ R. 9 W„ Knappton
7'/2-minute quadrangle, on the Columbia River, Washington.

U.S. GEOLOGICAL SURVEY, MENLO PARK,
CALIFORNIA

M7891. The same locality as LAM 5842, but collected by
E.J. Moore.

ACKNOWLEDGMENTS

I profited from discussions regarding this study with Kristin
A. McDougall and George W. Moore, U.S. Geological Sur-
vey, James C. Ingle, Jr., Stanford University, and Barry Roth,
California Academy of Sciences. The constructive comments
of Warren O. Addicott and George L. Kennedy, U.S. Geo-
logical Survey, are greatly appreciated. The fossil photo-
graphs were taken by Kenji Sakamoto and the manuscript
typed by Marion Anderson, U.S. Geological Survey. I am
indebted to Carole S. Hickman, University of California at
Berkeley, and to Peter U. Rodda, Barry Roth, Robert Van
Syoc, and Tony Summers, California Academy of Sciences,
for the loan of specimens. I am indebted to Edward C. Wil-
son, Los Angeles Natural History Museum, for arranging the
loans of the Knappton collections and for his support and
encouragement throughout the study.

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Contributions in Science, Number 351

Moore: Lincoln Creek Formation Mollusks 41

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Submitted 28 September 1983; accepted 17 February 1984.

42 Contributions in Science, Number 351

Moore: Lincoln Creek Formation Mollusks

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FISHES OF THE GENUS NANSENIA (MICROSTOMATIDAE)
WITH DESCRIPTIONS OF SEVEN NEW SPECIES

Kouichi Kawaguchi1 and John L. Butler2

ABSTRACT. The microstomatid fishes of the genus Nansenia are
reviewed on a worldwide basis and 1 3 valid species are recognized.
Of the 10 nominal taxa six species are recognized: N. Candida (with
N. sanrikuensis as a possible synonym), N. groenlandica, N. atlan-
tica, N. oblita, N. crassa, and N. ardesiaca (with N. tanakai, N.
schmitti, N. macrolepis, and N. robusta as synonyms). Seven new
species are described based on specimens collected from the Atlantic,
Pacific, and Antarctic oceans. Original descriptions are presented for
N. ahlstromi, N. antarctica, N. pelagica, N. megalopa, N. longicau-
da, N. tenera, and N. tenuicauda. The species of Nansenia are sep-
arated into two groups based on the number of branchiostegal rays,
either three or four; species of each group are distinguished on the
basis of the differences in vertebral and gill raker counts, predorsal
length in percent of standard length, and proportional size of the
caudal peduncle. A key is presented.

Five species, N. atlantica, N. oblita, N. megalopa, N. tenera, and
N. tenuicauda occur only in the Atlantic; three, N. ahlstromi, N.
Candida, and N. crassa, are limited to the Pacific. Nansenia ardesiaca
is widespread in the slope waters of southeast Asian seas, extending
northward to off southern Japan and westward to off South Africa.
Nansenia groenlandica occurs in the subarctic Atlantic and subant-
arctic eastern Pacific. Antarctic waters are inhabited by a circumpolar
species, N. antarctica. Nansenia pelagica and N. longicauda are found
in both the Atlantic and Pacific oceans; the latter species restricted
to the Northern Hemisphere. Distributional patterns of Nansenia
are discussed in relation to those of other midwater fishes.

Six species exhibit allometric growth of head length, predorsal
length, and eye diameter. Allometry may be an adaptation to me-
sopelagic and benthopelagic habitats.

INTRODUCTION

Fishes of the microstomatid genus Nansenia are distributed
widely in the world ocean. They are found in oceanic and
coastal waters from the subarctic to the subantarctic. Col-
lections from both midwater and bottom trawls suggest that
these fishes inhabit the epi- and mesopelagic zones and also
the benthopelagic zone of continental or insular slopes. In
areas of high productivity examples of Nansenia often exceed

1. Ocean Research Institute, University of Tokyo.

2. Southwest Fisheries Center, National Marine Fisheries Service,
National Oceanic and Atmospheric Administration, La Jolla.

200-300 mm in standard length and are sometimes captured
in large numbers.

Notwithstanding the considerable numbers of adults and
juveniles taken with trawls and larvae collected in plankton
nets, the species are poorly known. Cohen (1958) examined
specimens of six of seven nominal species that had been
referred to Nansenia and concluded that the paucity of ma-
terial precluded a critical revision. Since 1958 no compre-
hensive taxonomic work has been done on the genus, al-
though four additional species have been described (Blache
and Rossignol, 1962; Lavenberg, 1965; Abe, 1976; Kana-
yama and Amaoka, 1983). In general, specimens of Nansenia
are soft-bodied, poorly ossified and hence often collected in
poor condition. As a result the nominal species are inade-
quately known and a considerable number of species remain
undescribed. Some species are distributed worldwide, but in
previous works geographical variation and dines were not
considered, and specimens from new localities were named
as new species. The degree of ossification is different, both
among species and growth stages, which, combined with the
lack of a complete size series for all species, makes compar-
ative osteology difficult. For all these reasons it is not possible
at this time to present a comprehensive definition of the
genus.

The present study is a preliminary one that aims to di-
agnose each species, both previously described and ones de-
scribed in this paper with the objective of contributing in-
formation that will lead to a clearer definition of Nansenia
and to the osteology, phytogeny, zoogeography, early life his-
tory, and ecology of each species.

METHODS AND MATERIALS

Measurements were taken with dividers and recorded to the
nearest 0.1 mm. Unless otherwise indicated, definitions fol-
low Hubbs and Lagler ( 1 964) and include the following: stan-
dard length (SL); length of head (HL); depth of body (BD)—
vertical through origin of base of dorsal fin; least depth of
caudal peduncle (CPD); caudal peduncle length (CPL); di-
ameter of eye (ED)— horizontal distance between opposite

Contributions in Science, Number 352, pp. 1-22
Natural History Museum of Los Angeles County, 1984

margins of socket; snout length (Sn); interorbital length (In-
tor); width of body (BW)— width just behind the head; pre-
dorsal length (Pre D); preanal length (Pre A); preventral length
(PreV)— distance from tip of snout to structural base of out-
ermost ventral fin ray. Size of specimen is recorded in stan-
dard length, with standard deviation and range in parenthe-
ses. Center of body is the midpoint of standard length. If
allometric growth is indicated for a species, data are pre-
sented for different size groups.

Numbers of vertebrae and median fin rays were counted
from radiographs. The ultimate double rays of the anal and
dorsal fins were counted as one. The short spine sometimes
found at the base of the outermost pelvic fin ray was not
counted. In other counts, all elements were included. The
upturned ural centra were counted as one. Teeth were count-
ed on both sides of the upper and lower jaws. Frequency
distributions of gill raker and vertebrae counts are shown
with the number of specimens in parentheses following each
count.

In the materials section of each species, catalogue numbers
are followed by the number of specimens and their size range
in parentheses. The station number is preceded by the cruise
number unless other wise indicated. The sampling depth is
indicated by m and the length of wire out by mwo. The
following institutional abbreviations are used.

UBC Institute of Fisheries, University of British Colom-
bia, Vancouver, Canada.

CAS California Academy of Sciences, San Francisco, USA.

FMNH Field Museum of Natural History, Chicago, USA.
ISH Institute fur Seefischerei, Hamburg, West Germany.
KU Kochi University, Japan.

KYO Department of Fisheries, Kyoto University, Japan.

LACM Natural History Museum of Los Angeles County,
USA.

OSU School of Oceanography, Oregon State University,
Corvallis, USA.

SIO Scripps Institution of Oceanography, University of
California, La Jolla, USA.

SWFC Southwest Fisheries Center, National Marine Fish-
eries Service, La Jolla, USA.

UF Florida State Museum, University of Florida,
Gainesville, USA.

USNM National Museum of Natural History, Smithsonian
Institution, Washington, D.C., USA.

ZMUC Zoologiske Museum, University of Copenhagen,
Denmark.

Genus Nansenia Jordan and Evermann, 1896

Nansenia Jordan and Evermann, 1896:528 (type species by
monotypy, Microstomus gron/andicus Reinhardt, 1839; cf.
Follet and Cohen, 1958).

Bathymacrops Gilchrist, 1922:531 (type species by mono-
typy, Bathymacrops macrolepis Gilchrist, 1922).
Euproserpa (subgenus) Fowler, 1934:256 (type species by
original designation. Microstoma schmitti Fowler, 1934).

The definition of the genus Nansenia adopted here basically
follows Chapman (1942, 1943, 1948) and Cohen (1964). The
placement of the genus Nansenia in the family Microstom-
idae follows Ahlstrom et al., 1984. Body slender, subcylin-
drical or slightly compressed. Dorsal adipose fin present on
the last quarter of body. Predorsal length less than 6 1 percent
of SL. Snout shorter than half of eye, which is directed lat-
erally and is not tubular. Branchiostegals three or four. Anal
fin rays 8-10 (rarely 11). Pectoral fins inserted on sides of
the body. Parietals large and meeting on the midline. No
teeth on premaxillary or maxillary. Teeth present on pala-
tines, head of vomer and dentaries.

REMARKS. The difference between Nansenia and Bathy-
lagus (presently placed in a separate family, Bathylagidae) is
not clear, despite efforts to discover adequate diagnostic char-
acters. Characters concerned with the swimbladder, the oto-
liths, and morphology of the larvae, which may be diagnostic,
are not always known for all members of both genera at
different growth stages. Their value for distinguishing the two
genera remains to be confirmed.

The arrangement of the parietals, which have been re-
ported to meet broadly on the midline in Nansenia, but not
meeting in Bathylagus (Chapman, 1942, 1943, 1948; Cohen,
1964), is probably diagnostic if it is partly modified to pa-

Table 1. Counts and measurements used in the key to the species with three branchiostegal rays and distributional information.

Species

Vertebrae

Gill

rakers

Predorsal
distance
(% SL)

Ratio

CPL/CPD1

Distribution

N. ahlstromi

35-36

40

50.5-55.8

1.0-1. 3

Subtropical Eastern Pacific, 20°N-35°N

N. Candida

44-47

26-31

49.4-57.6

1. 8-2.2

Subarctic Pacific, 30°N-55°N

N. groenlandica

42-45

37-45

43.7-45.6

2. 8-4.0

Subarctic Atlantic, 40°N-70°N;

Eastern South Atlantic

N. antarctica

49-50

35-41

45.9-47.7

2.0-2. 5*

Southern oceans, 40°S-60°S

2. 5-3.0**

1 Caudal peduncle length to caudal peduncle depth.
* <1 10 mm SL.

**>110 mm SL.

2 Contributions in Science, Number 352

Kawaguchi and Butler: Fishes of the Genus Nansenia

Table 2. Counts and measurements used in the key to the species with four branchiostegal rays and distributional information.

Species

Vertebrae

Gill

rakers

Predorsal

length

(% SL)

Caudal
peduncle
length
(% SL)

Caudal

peduncle

length

(% SL)

Ratio

CPL/CPD1

Distribution

N. pelagica

38-39

36-41

50.4-57.8

1 1.5-13.3

8.1-1 1.3

1.1-1. 5

Tropical and Subtropical
Atlantic and Pacific

N. atlantica

41-42

30-36

51.9-56.9

1 1.3-12.2

8. 2-8. 6

1.4-1. 5

Tropical Atlantic

N. oblita

42-45

30-36

55.0-56.1

9.3-12.2

8. 2-9. 6

1. 1-1.2

Mediterranean and Eastern
North Atlantic

N. crassa

43-46

35-37

49.0-60.2

13.0-15.2

6. 3-7. 7

1. 7-2.2

Eastern Tropical and
Subtropical Pacific

N. megalopa

44-45

21-23

42.1-44.2

15.6-17.6

5.2-5. 7

2.8-3. 2

Tropical Atlantic

N. longicauda

47-50

23-27

41.8-46.0

15.4-18.6

4.3-5. 6

3.0-4. 1

North Atlantic and North
Pacific, 20°N-40°N

N. tenera

42-43

43-46

43.4-45.8

12.6-15.0

6. 2-6.5

1. 9-2.4

Subpolar and Temperate
Atlantic

N. ardesiaca

46-48

27-35

45.7-48.5

13.0-15.0

6. 1-7.5

1. 7-2.3

Off Japan, Southeast
Asia and East Africa

N. tenuicauda

46

38-42

44.0-45.8

13.5-13.9

4. 4-5.0

2.7-3. 1

South Atlantic

1 Caudal peduncle length to caudal peduncle depth.

rietals of Bathylagus not meeting or meeting at a point in
the anteriormost part on midline. Branchiostegal counts are
three or four in Nansenia, but mostly two in Bathylagus.
However at least one undescribed bathylagid examined dur-
ing the present study had exceptionally three branchiostegals.
Anal fin ray counts are also useful in separating the two
genera: 8-10 (rarely 1 1) in Nansenia and 12-28 (rarely 1 1)
in Bathylagus. Better diagnostic characters for the genus Nan-
senia must await a revision of the family Bathylagidae.

KEY TO SPECIES

The species of Nansenia are separated into two groups based
on branchiostegal counts of three or four. Four species have
three branchiostegals and nine have four.

Counts and measurements used in the keys and distribu-
tional data are presented for all species in Tables 1 and 2
and Figures 21 and 22. After identification is made with the
key, specimens should be checked against the information
in the tables and also against the detailed descriptions of each
species. The number of branchiostegal rays of Nansenia is
stable within species, but careful examination under a dis-
secting microscope is necessary to recognize the innermost
small ray (Fig. 1).

KEY TO SPECIES OF NANSENIA WITH THREE
BRANCHIOSTEGAL RAYS

1 a. Ratio of caudal peduncle length to caudal peduncle depth

less than 1.5; vertebrae 35-36 ahlstromi

1 b. Ratio of caudal peduncle length to caudal peduncle depth
greater than 1.5; vertebrae 40-42 2

2a. Gill rakers on first arch 25-31; predorsal length more

than 49 percent of SL Candida

2b. Gill rakers on first arch 35-45, predorsal length less than
49 percent of SL in specimens larger than 50 mm SL .

3

3a. Vertebrae 42 (eastern South Pacific), 43-45 (Atlantic)

groenlandica

3b. Vertebrae 49-50 antarctica

KEY TO SPECIES OF NANSENIA WITH FOUR
BRANCHIOSTEGAL RAYS

la. Vertebrae 38-39 pelagica

lb. Vertebrae 41-50 2

2a. Predorsal length 49 or more percent of SL 3

2b. Predorsal length less than 49 percent of SL 5

3a. Length of caudal peduncle more than 12.5 percent of SL

and depth of caudal peduncle less than 8 percent of SL

with CPL/CPD ratio more than 1.6 crassa

3b. Length of caudal peduncle less than 12.5 percent of SL
and depth of caudal peduncle more than 8 percent of SL

with CPL/CPD ratio less than 1.6 4

4a. Proximal part of adipose fin densely pigmented; verte-
brae 41-42; gill raker count 30-36 atlantica

4b. Proximal part of adipose fin not pigmented; vertebrae

42-45; gill raker count 28-30 oblita

5a. Length of caudal peduncle more than 1 5.2 percent of SL
and depth of caudal peduncle less than 6 percent of SL;

gill rakers 21-27 6

5b. Length of caudal peduncle less than 15.2 percent of SL
and depth of caudal peduncle more than 6 percent of SL
except in N. tenuicauda 7

Contributions in Science, Number 352

Kawaguchi and Butler: Fishes of the Genus Nansenia 3

a

b

Figure 1. Typical patterns of branchiostegal rays of Nansenia. a.
N. ardesiaca, N. ahlstromi, N. crassa, and N. megalopa ; b. N. pe-
lagica and N. atlantica; c. N. tenera and N. longicauda; d. N. ant-
arctica; e. N. groenlandica.

6a. Vertebrae 44-45; gill rakers 2 1-23 megalopa

6b. Vertebrae 47-50; gill rakers 23-27 longicauda

7a. Vertebrae 42-43; gill rakers 43-46 tenera

7b. Vertebrae 46-48; gill rakers 27-42 8

8a. Gill rakers 27-35; CPL/CPD ratio less than 2.5

ardesiaca

8b. Gill rakers 38-42; CPL/CPD ratio more than 2.5 ....
tenuicauda

SPECIES WITH THREE BRANCHIOSTEGAL RAYS

Nansenia ahlstromi new species

Figures 2, 21

HOLOTYPE. SIO 57-87 (1 , ca. 95 mm, slightly damaged
mature female), R/V Spencer F. Baird, 29°15'N, 126°07'W,
2240-0730, 14-15 May 1955, 10 ft.-IKMT, 0-754 m.

PARATYPES. SIO 63-425 (1, 38.7), R/V Horizon,
27°46'N, 1 29°14.9'W, 0534-1006, 2 Apr. 1962, 10 ft.-
IKMT, 4500 mwo; LACM 43546-1 (1, 32.0), R/V David
Starr Jordan, 24°00'N, 145°00'W, 2348-0021, 21-22 May
1972, 6 ft.-IKMT 600 mwo; SWFC 7210-24.139 (1, 58.5
decomposed), R/V David Starr Jordan, 24°00'N, 139°00'W,
0040-0159, 29 Oct. 1972, 50 ft. universal MWT, 0-494 m.

DIAGNOSIS. Differs from other Nansenia in having three
branchiostegal rays and 35-36 vertebrae, the least of any
known species.

DESCRIPTION. Counts and measurements are based on
three juvenile specimens, 32.0-58.5 mm SL. Only the ver-
tebrae were counted for the damaged adult specimen, ca. 95
mm SL. D 8-9; A 7-8; P 12; V 9-1 1; gill rakers 12 + 28 in
two specimens and 11 + 29 in one, total 40; branchiostegal
rays three; vertebrae 35 in one specimen, 36 in three speci-
mens. Pyloric caeca not counted due to paucity of specimens.
About 30 conical teeth irregularly arranged on head of vomer;
about 100 compressed teeth closely set on dentaries in ho-
lotype.

Head relatively large compared with body, reflecting the
reduced number of vertebrae. Eye large, diameter 2. 3-2. 9
times in head length in three specimens, 38.7-ca. 95 mm SL.

Head length and eye diameter indicate remarkable allometric
growth (Fig. 4). Supraorbital bone damaged and lost in the
holotype. Pectoral fin base low, its upper end above ventral
margin of body about one-third of the distance between ven-
tral margin and lateral line. Origin of dorsal fin base behind
center of body. Ventral fin base below or just behind posterior
end of dorsal fin base in the three juveniles, 32.0-58.5 mm
SL, but well behind dorsal fin base in the holotype. Adipose
fin base over posterior half of the anal fin base.

Tip of lower jaw, anterior part of gular area, and base of
adipose fin darkly pigmented in the 38.7-mm SL specimen
(SIO 63-425). The same pigment pattern is present in the
holotype.

SIZE. The holotype of ca. 95 mm is an adult bearing eggs.

DISTRIBUTION. Restricted to the subtropical eastern
North Pacific.

ETYMOLOGY. It is with great pleasure that we name this
species Nansenia ahlstromi in recognition of the contribu-
tions of the late Elbert Halvor Ahlstrom to our knowledge
of pelagic fishes.

Nansenia Candida Cohen, 1958

Figures 3, 4, 22

Nansenia Candida Cohen, 1958:52-54, fig. 1 (orig. descr.)

Eastern North Pacific.

INansenia sanrikuensis Kanayama and Amaoka, 1 983:77—

79 (orig. descr.) Western North Pacific.

MATERIALS. CAS 51023 (1, alizarin specimen dissect-
ed), one of the paratypes, R/V Hugh M. Smith, 41°39'N,
139°02'W, 30 May 1956, from the stomach of Alepisaurus
sp.; OSU 8, 9 (2, 46.4, 80.7), 49°16'N, 132°42'W, 0-60 m;
OSU 10, 12 (2, 44.0, 136.9), 41°59.5'N, 126°30.6'W, 0-200
m; OSU 11 (1, 150.5), 44°37.3'N, 125°17.3'W, 1500-500 m;
OSU 960 (1, 150.5), 44°37.3'N, 125°38.0'W, 0-200 m; OSU
1132,1133 (2, 79.0, 72.2), 44°25'N, 1 29°35'W, 0-75 m; OSU
1 166 (1, 32.1), MT-648, 44°16'N, 125°15'W, surface tow, 40
mwo; OSU 1 170 (1,23.5), MT-652, 44°3 l'N, 125°17'W, 0-
1400 m; OSU 1191 (2, 95.8, 99.6), MT-757, 44°38'N,
1 28°35'W, 0-210 m; OSU 1212 (1, 140.0), 44°39.8'N,
125°00'W, 0-200 m; OSU 1216 (1, 108.2), MT-749, 44°39'N,
1 25°36'W, 0-200 m; OSU 1 2 1 7 ( 1 , 60.3), MT-727, 44°39'N,
1 28°00'W, 0-1000 m; OSU 2003 (1, 179.2), haul 2054,
44°49.7'N, 1 25°34.0'W, 0-600 m; OSU 2388 ( 1 , 1 29. 1 ), MT-
2398, 44°39'N, 125°29'W, 0-500 m; USNM 197380 (1, 60.5),
20 miles southeast of San Clemente Is., California, spit up
by albacore; USNM 195874 (1, 122), sta. BB-76, North Pa-
cific, 0-30 m; USNM 195875 (1, 75.5), sta. BB- 1 76, 48°03'N,
1 34°20'W; USNM 195876 (1, 156), sta. BB-199, 49°15.6'N,
145°56.7'W, 100 mwo; SIO 66-5 1-9F (1, 24), 40°35'N,
125°51.5'W; SIO 55-73-9A (1, ca. 64, damaged), 39°01'N,
165°10'W, from Alepisaurus stomach; UBC 65-607 (2, 75.5,

81.0) and UBC 65-609 (2, 76.5, 87.5), off Queen Charlotte
Sound, B.C., Canada; UBC 65-6 10 ( 1 6, 76.0-88.0) and UBC
65-615 (1, 77.5), off Queen Charlotte Sound; UBC 65-623
(16, 118.0-187.0), 52°11'N, 133°11'W; UBC 65-524 (1,

121.0) , 52°1 3'N, 1 33°1 2'W; LACM 34258-1 (2, 63-71),

4 Contributions in Science, Number 352

Kawaguchi and Butler: Fishes of the Genus Nansenia

h

■* 1 0 mm

Figure 2. Nansenia ahlstromi. Paratype, SIO 63-425, 38.7 mm SL.

R/V Alaska, off San Clemente Is., California; LACM 9006-
23 (1, 35), R/V Velero, Catalina Basin off California.

DIAGNOSIS. Differs from other Nansenia in the follow-
ing combination of characters, three branchiostegal rays and
26-31 gill rakers on the first arch. Vertebral counts higher
(44-47) in N. Candida than in N. ahlstromi (35-36).

DESCRIPTION. D 9-10; A 8-9; P 10-1 1 (rarely 9); V 9-
10 (rarely 1 1); gill rakers 8-1 1 + 1 + 17-20, total 26-3 1 with
26 in one specimen, 27 in five, 28 and 29 in three respectively,

30 in one and 31 in three; branchiostegal rays 3; vertebrae
44-47 with 44 in two specimens, 45 in five, 46 in eight and
47 in two. The above counts are based on 18 specimens,
46.2-179.2 mm SL. Pyloric caeca seven in two specimens,
nine in one. About 12-21 conical teeth on head of vomer,
about 80-100 teeth on dentaries, resembling a closely spaced
picket fence, in three specimens more than 130 mm SL.

Body slender, moderately compressed, with body depth to
width ratio 1.5-1. 7. Eye diameter 3.0 to 3.4 times in head

Figure 3. Nansenia Candida. OSU 2003, mature female, 179.2 mm SL.

Contributions in Science, Number 352

Kawaguchi and Butler: Fishes of the Genus Nansenia 5

%

70

PreD

(D

O

0)

0.

40-

30-

HL

20- *

o

0 ' ’ ' >-

0 50 100 150 200

S L (mm)

Figure 4. Nansenia Candida. Predorsal, head lengths, and eye di-
ameter in relation to standard length. Triangles indicate the values
for postlarvae and young juveniles smaller than 23 mm SL.

length, viewed laterally, its dorsal margin (supraorbital bone)
touching or slightly higher than dorsal margin of head, yel-
lowish iridescent pigment, roughly crescent-shaped, present
in the posterior half of iris. Aphakic space developed, its
horizontal space nearly equal to width of anteriormost part
of iris, and about half the width of its posterior part. Aphakic
space less well developed in juveniles smaller than 100 mm
SL. Allometric growth in head length, eye diameter and pre-
dorsal length is shown in Figure 4.

Upper end of pectoral fin base on midpoint between lateral

line and ventral margin of body, or slightly lower; its lower
end separated from ventral margin of body by a distance
more than length of pectoral fin base. Dorsal fin originating
behind center of body. Ventral fin base below posterior part
of dorsal fin. Origin of anal fin slightly in advance of adipose
fin. Skin around the anterior end of adipose and caudal fin
bases densely pigmented. Body color in formalin preserved
specimens brown. However, Cohen (1958) noted that in life
the entire fish is probably a bright silver.

SIZE. The largest specimen examined, 179.2 mm SL, had
mature ovaries.

DISTRIBUTION. Nansenia Candida has been collected
in the subarctic eastern Pacific east of 170°W between 35°N
and 55°N and in the California Current region north of 30°N
(Fig. 22).

REMARKS. Kanayama and Amaoka (1983) described
Nansenia sanrikuensis based on two large adult specimens,
206.0 and 238.8 mm SL. This species is closely related to
N. Candida, and both species occur in the subarctic North
Pacific. But N. Candida has been reported only from the
eastern part of the Pacific between the latitude of southern
California and Canada, and N. sanrikuensis was collected in
the western Pacific off northern Honshu, Japan. Kanayama
and Amaoka (1983) reported that N. sanrikuensis differs in
having a smaller eye (4. 8-5. 8 vs. 8. 3-9. 6 percent of SL),
shorter snout (3. 5-3.6 vs. 5. 1-5.8), lower gill raker counts
(25-27 vs. 30) and higher vertebral counts (48-49 vs. 44-
47). Their proportional data and gill raker counts for N.
Candida are based on the original description by Cohen ( 1 958)
which was taken from three specimens, 54.4-72.4 mm SL,

Figure 5. Nansenia groenlandica. ISH 194-59, 1 12.5 mm SL.

6 Contributions in Science, Number 352

Kawaguchi and Butler: Fishes of the Genus Nansenia

for measurements and five for meristic counts. However,
their vertebral counts are based on the present study. Spec-
imens studied by Kanayama and Amaoka ( 1 983) and Cohen
(1958) differ greatly in size. In the present study we found
considerable allometric growth in eye diameter, snout length,
and head length of N. Candida. We conclude that proportions
of snout, eye, and head are invalid for distinguishing the two
species. The ranges of gill raker counts are also overlapping
between the two (25-27 vs. 26-31) as N. Candida has a
wider range than Cohen (1958) described. At present only
vertebral counts (48-49 vs. 44-47) separate these species.
Considering the adjacent ranges, the paucity of specimens of
N. sanrikuensis and the possibility of a geographical cline,
the status of N. sanrikuensis is questionable.

%

50

40

30

20 -

10

Pre D

~o °~o

HL

~o O 7T°~

ED

Nansenia groenlandica (Reinhardt, 1840)

Figures 5, 6, 21

0 I—

40

60

80

100

S L |mm|

— i -i —

120 140

Microstomas gronlandicus Reinhardt, 1840:8 (orig. descr.

Fiskenesset, Greenland).

Nansenia groenlandica: Jordan and Evermann, 1896:528;

Schmidt, 1918:12, figs. 1-12; Cohen, 1964:24, fig. 8.

MATERIALS. Atlantic: ISH 21/75 (2, 57.0-67.0), FFS
Walther Herwig sta. 10-1/75, 41°18'N, 27°49'W, 0-117 m;
ISH 29/75 (1, 32.5), FFS Walther Herwig sta. 14-1/75,
41°26'N, 27°09'W, 0-183 m; ISH 194/59 (1, 112.5), FFS
Anton Dorn sta. 3257/59, 6 1°48'N, 1 3°08'W, 0-1400 m; ISH
314/73 (2, 132.0, damaged), FFS Walther Herwig sta. 678/
73, 65°09'N, 32°50'W, 0-1 100 m; ISH 448/73 (8, 60.5-103.5),
FFS Walther Herwig sta. 693/73, 57°55'N, 28°23'W, 0-415
m; ISH 473/73 (1, 89.0), FFS Walther Herwig sta. 693/73,
56°28'N, 26°44'W, 0-2500 m; ISH 540/73 (3, 80.0-108.0),
FFS Walther Herwig sta. 696/73, 55°39'N, 25°47'W, 0-2500
m; ISH 540/73 (3, 80.0-108.0), FFS Walther Herwig 696/
73, 55°39'N, 25°47'W, 0-410 m; ISH 748/73 (1, 57.0), FFS
Walther Herwig 707/73, 50°03'N, 19°39'W, 0-255 m; USNM
186071 (damaged, head only), M/V Delaware. 40°34'N,
64°07'W, from stomach of Makaira\ USNM 221368 (3, dam-
aged, ca. 63-84), FFS Walther Herwig sta. 695/73, 55°43'N,
25°53'W, 0-2600 m. Pacific: LACM 1 1292 (1, 63.7), R/V
Eltanin sta. 1972, 39°34'S, 127°18'W, IKMT, 0-4493 m;
LACM 10288 (2, damaged, ca. 30), R/V Eltanin sta. 190,
34°5 l'S, 74°04'W, 0-1580 m.

DIAGNOSIS. Differs from other Nansenia in having three
branchiostegal rays, 42-45 vertebrae (43-45 in the Atlantic,
42 in Pacific specimens), and 37-45 gill rakers on the first
arch.

DESCRIPTION. D 9-10; A 8-10; P 1 1-13; V 1 l-12(rare-
ly 10); gill rakers 12-15 + 1 + 23-29, total 37-45 with 37(2),
38(5), 40(1), 41(2), 42(1), 44(2), and 45(1); branchiostegal
rays three; vertebrae 42(1) in the South Pacific, 43-45 with
43(8), 44(9), and 45(1) in the North Atlantic. Counts are
based on 19 specimens, 48.0 to 132.0 mm SL. Pyloric caeca
seven and eight in two specimens; 20-22 conical teeth irreg-
ularly arranged on vomer, 50-65 teeth closely set on dentaries
in four specimens.

Body slender, body depth-width ratio ranges from 1.0-
1.2. Eye large, its diameter 2.5 to 2.8 times in head length.

Figure 6. Nansenia groenlandica. Predorsal, head lengths, and eye
diameter in relation to standard length.

Viewed laterally, about one-third of eye circumference pro-
truded above the dorsal margin of the head. White or yel-
lowish tissue developed over posterior half of iris, sometimes
covering whole iris. Aphakic space slightly developed ante-
riorly, its horizontal space nearly equal to half to two-th(rds
width of the anterior part of iris. Aphakic space less well
developed in juveniles smaller than 100 mm SL.

Upper end of pectoral fin base separated from ventral mar-
gin of body by one-third of distance between lateral line and
ventral margin or slightly higher; its lower end above ventral
margin by a space more than the length of pectoral fin base.
Dorsal fin originating in front of center of body. Ventral fin
base below posterior end of dorsal fin base or just behind it.
Adipose fin base above last two anal fin rays. No remarkable
inflection in allometric growth for specimens between 48 and
132 mm SL (Fig. 6).

Skin around adipose fin base uniformly pigmented com-
pared to the lateral body surface. Caudal fin base and pos-
terior part of caudal peduncle densely pigmented, sparser in
specimens larger than 100 mm SL. Body of preserved spec-
imens usually brown or light brown, some with remains of
guanine. The entire body surface of the ISH 1 94-59 specimen,
although all scales had come off, is covered with guanine.
This specimen may have been preserved in alchohol.

SIZE. The badly damaged specimen (caudal segment miss-
ing) taken from a marlin stomach, USNM 1 8607 1 , identified
as N. groenlandica only by its branchiostegal ray count and
locality, is a mature male and its standard length can be
estimated as about 180 mm based on its head length. The
largest four intact specimens, 103.0 to 132.0 mm, collected
in September have undeveloped gonads.

DISTRIBLTTION. This species occurs widely in the arctic
and subarctic waters of the North Atlantic Ocean. The south-
ern limit of its distribution is near 40°N. Present results agree
well with the work of Schmidt (1918) in the eastern North
Atlantic. Three specimens collected in the subantarctic or

Contributions in Science, Number 352

Kawaguchi and Butler: Fishes of the Genus Nansenia 7

CM I
0

23456789 10

I I I I CM
12 13 14 15

Figure 7. Nansenia antarctica. LACM 10875 (one of the three catalogued), 198.9 mm SL, holotype.

transitional waters of the eastern South Pacific show no mor-
phological difference from the Atlantic specimens, but have
slightly fewer vertebrae. Mukhacheva (1972) reported a sim-
ilar distribution pattern for the midwater fish, Gonostoma
bathyphilum, which is widely distributed in the North At-
lantic between 30°N and 60°N, the South Atlantic south of
30°S, and also in the eastern South Pacific south of 30°S.

Nansenia antarctica new species

Figures 7, 8, 21

HOLOTYPE. LACM 10875 (one of three), 198.9 mm SL,
R/V Eltanin sta. 1204, 55°57'S, 159°23'W, 0560-1035, 10
Aug. 1964, I KMT, 0-4145 m.

PARATYPES. LACM 10875 (two of three) (2, 197.8-
207.5), collection data as for holotype. LACM 10658-15 (2,
79.2-225.0), R/V Eltanin sta. 858, 64°42'S, 78°34'W, 1905-

%

o * > 1 • —

0 50 100 150 200

S L (mm)

Figure 8. Nansenia antarctica. Predorsal, head lengths, and eye
diameter in relation to standard length.

0030, 17-18 Nov. 1963, I KMT; USNM 247255 (1, 158.6),
R/V Eltanin cr. 35, sta. 2300, 52°00'S, 124°02'E, 20 Sep.
1968, I KMT, 0-750 m (bottom depth 3914-4362 m).

OTHER MATERIALS. LACM 11194 (2, 98.9-108.9),
R/V Eltanin sta. 1661, 61°30'S, 108°26'W, 0134-0430, 26
Apr. 1966, I KMT, 0-5036 m; LACM 11212(1, 49.9), R/V
Eltanin sta. 1686, 57°39'S, 115°12'W, 1255-1540, 6 May
1966, I KMT, 0-4286 m; USNM 247254 (7, 31.0-35.0),
R/V Eltanin cr. 21, sta. 20G, 0-850 m; ISH 688/71 (1, 112.5),
FFS Walther Herwig sta. 363-III/71, 40°18'S, 39°04'W, 8
Mar. 1971, 0-800 m; ISH 542/71 (1, 118.2), FFS Walther
Herwig sta. 354-11/71, 39°19'S, 48°09'W, 6 Mar. 1971, 0-ca.
2000 m.

DIAGNOSIS. Differs from other Nansenia in having three
branchiostegal rays, 47-50 vertebrae, and 35-41 gill rakers.

DESCRIPTION. D 9-10; A 9-10; P 12-14; V 1 1-12; gill
rakers 10-13 + 1 + 23-27, total 35-41 with 35(2), 37(2),
38(2), 39(2), and 41(1); branchiostegals three; vertebrae 49-
50 with 49(3) and 50(5). Pyloric caeca seven in one specimen;
27 irregularly arranged conical teeth on head of vomer and
about 75 compressed teeth on dentaries on one specimen of
207.5 mm SL.

Body slender, anterior part slightly compressed in speci-
mens larger than 190 mm, subcylindrical in smaller speci-
mens. Caudal peduncle more compressed than anterior part
of body. Body depth-width ratio 1. 1-1.7. Eye large, diameter
2. 4-2. 9 times in head length, with its dorsal margin pro-
truding slightly above margin of head. Golden pigment,
roughly crescent-shaped, on posterior half of iris. The ratio
of width of the posterior part of iris to that of the anterior
part is between two and four, whereas the ratio is less than
two in other species having three branchiostegal rays. Apha-
kic space extremely well developed in front of lens (some-

8 Contributions in Science, Number 352

Kawaguchi and Butler: Fishes of the Genus Nansenia

CM

CM

0 1 23456789 10

12 13 14 15

Figure 9. Nansenia megalopa. ISH 2013-71 (one of the two catalogued), 133.0 mm SL, holotype.

times lens placed anteriorly), its horizontal space usually three
to four times as wide as the width of the iris at the ante-
riormost part, in specimens larger than 100 mm SL. This
space is less well developed in smaller juveniles.

Upper end of pectoral fin base just below midpoint between
the lateral line and ventral margin of body, its lower end
separated from the ventral margin by a distance 1 .8-3.0 times
as long as its base. Origin of dorsal fin base in front of the
center of the body. Ventral fin base just below the posterior
end of the dorsal fin base. Adipose fin base above the pos-
terior half of anal fin base.

Skin badly abraded in most specimens, light brown in
alcohol. Dark colored scale pockets, which are extremely
fragile, a few sometimes remaining along lateral line and the
dorsal and ventral sides of body. Caudal and adipose fin bases
usually slightly pigmented.

SIZE. The largest specimens studied, 207.5 mm, had ma-
ture ovaries with developing eggs.

DISTRIBUTION. This species occurs in the southern part
of the subantarctic region between the antarctic and sub-
tropical convergences of the Pacific (Fig. 21). Its distribution
probably extends northward to the subtropical convergence
in the Atlantic sector.

ETYMOLOGY. Named for its region of occurrence.

SPECIES WITH FOUR BRANCHIOSTEGAL RAYS

Nansenia megalopa new species

Figures 9, 2 1

Nansenia sp. 2 (?), Parin and Golovan, 1976:251-252, fig.

2, 04°10'N, off west Africa, 0-510 m.

HOLOTYPE. ISH 2013 a/71, 133.0 mm, FFS Walther

Herwig sta. 463-III/71, 08°11'S, 14°12'W, 8 Apr. 1971, 0-
640 m.

PARATYPES. ISH 1730/71 (1, 126.5), FFS Walther Her-
wig sta. 447-III/71, 18°36'S, 04°18'W, 4 Apr. 1971; ISH
20 1 3b/7 1 (1, 108.5), FFS Walther Herwig sta. 463-III/71,
08°1 l'S, 14°12'W, 8 Apr. 1971, 0-640 m; ISH 2491/71 (1,
139.5), FFS Walther Herwig sta. 486-III/71, 07°32'N,
20°54'W, 14 Apr. 1971, 0-ca. 1300 m; UF 29912 (1, 126.0),
R/V Geronimo cr. 2, sta. 82, 03°28'S, 00°14'W, Gulf of
Guinea, 6 Aug. 1963, 0-710 m.

DIAGNOSIS. Differs from other Nansenia in having four
branchiostegal rays, 44-45 vertebrae and 21-23 gill rakers
on the first arch. This species is also separable from all other
Nansenia by having a very large eye, eye diameter to snout
length ratio >2.8 to 1.

DESCRIPTION. D 9-10; A 8-10; P 12-13; V 9-10; gill
rakers 6-7 + 1 + 14-15, total 21-23 with 21(1), 22(2), and
23(2); branchiostegal rays four; vertebrae 44-45 with 44(3)
and 45(2).

Pyloric caeca not counted; teeth on head of vomer, usually
embedded in the tooth ridge and often indiscernible; 35-50
teeth on both dentaries.

Body slender and subcylindrical, body depth to width ratio
1.2-1. 4 in specimens of 108.5-139.5 mm SL. Eye large, its
diameter 2. 1 to 2.4 in head length. A roughly crescent-shaped,
silvery pigment patch on the posterior half of the iris; pupil
elliptical with a large aphakic space in front of the lens. Upper
end of pectoral fin base around midpoint between lateral line
and ventral margin of body. Dorsal fin base originating in
front of center of body. Ventral fin base below posterior end
of dorsal fin base. Adipose fin origin above posterior third
of anal fin base. Bases of ventral and adipose fins densely
pigmented.

Contributions in Science, Number 352

Kawaguchi and Butler: Fishes of the Genus Nansenia 9

H 10 mm

Figure 10. Nansenia longicauda. a. SIO 76-7-9, 133 mm SL, holotype; b. ISH 60-66, 78.0 mm SL, paratype.

SIZE. A specimen of 126.5 mm SL has ripe ovarian eggs.

DISTRIBUTION. All specimens were collected in the
oceanic area of the tropical Atlantic between 10°N and 20°S.

ETYMOLOGY. Nansenia megalopa from Greek megale,
great, and from ops, eye, referring to the extraordinarily large
eye.

Nansenia longicauda new species

Figures 10, 1 1, 22

HOLOTYPE. SIO 76-9, 133.0 mm, 30°37.0'N,

147°24.0'W, 2027-2350, 10 Nov. 1971,IKMWT, 1800mwo.

PARATYPES. Pacific: USNM 207530 (1, 104.6), Univ.
of Hawaii, samp. no. 71-2-11, 21°20'N, 158°20'W, 28 Feb.
1971, 610-650 m; USNM 215703 (1, 97.2), R/V Townsend
Cromwell, CR-52, sta. 16, 21°32.7'N, 158°21.8'W, 0355-
1100, 12 Feb. 1971. Atlantic: ISH 60/66(1,78.0), FFS Wal-
ther Herwig sta. 177-66, 33°45'N, 16°00'W, 10 May 1966,
MT1600, 0-600 m.

OTHER SPECIMENS EXAMINED. Pacific specimens:
SIO 71-300 (1, 65.0), R/V Thomas Washington sta. Aries

9-H7, 27°24.5'N, 155°25.5'W, 0720-1100, 30 Sep. 1971,
lOft.-IKMT, 3000 mwo; USNM 207531 (1, 46.5), Univ.
of Hawaii, samp. no. 71-6-10, 21°00'N, 158°20'W, 10 Jun.
1971, 560-600 m; USNM 215470 (1, 67.0), R/V Townsend
Cromwell, samp. no. 73-8-29, 21°20'N, 158°20'W, 0745-

%

50 -

40 -

-o-

o

-o-®

O O

-®- Pre D

30

= 20
“ 10

HL

t 0

30 40 50 60 70 80 90 100 110 120 130 140

S L ( mm )

Figure 11. Nansenia longicauda. Predorsal, head lengths, and eye
diameter in relation to standard length.

10 Contributions in Science, Number 352

Kawaguchi and Butler: Fishes of the Genus Nansenia

Figure 12. Nansenia tenera. a. ZMUC P 1 962 1-23 (one of the three cataloged), 129.0 mm SL, holotype; b. ISH 540-73, 117.0 mm SL,
paratype.

1054, 29 Aug. 1973, 400-1 100 m; SIO 71-302 (1, 67.0), R/V
Thomas Washington Aries 9-H9, 27°24.5'N, 155°25.5'E, 30
Sep. 1971. Atlantic specimens: USNM 246799 (1 , 41.5), Acre
6-22; USNM 246800 (1, 39.1), Acre 9-24. Both specimens
were collected around 38°18'N and 64°12'W.

DIAGNOSIS. Differs from other Nansenia in having four
branchiostegal rays, 23-27 gill rakers on the first arch, 47-
50 vertebrae and a slender caudal peduncle, the least depth
of which is 4. 3-5. 6 percent of SL. The body shape of this
species resembles that of N. megalopa. Higher gill raker counts
(23-27 vs. 21-23), higher vertebral counts (47-50 vs. 44-
45) and longer snout (2.3-4. 1 vs. 2. 3-2.9 percent of SL)
separate N. longicauda from N. megalopa.

DESCRIPTION. D 10 (rarely 11); A 10-11 (rarely 9); P
13-14 (rarely 1 1); V 10-11 (rarely 9); gill rakers 7-8 + 1 +
15-18, total 23-27 with 23(1), 24(4), 25(2), and 26(2); bran-
chiostegal rays four; vertebrae 47-50 with 47(1), 48( 1 ), 49(5),
and 50(4). These counts are based mainly on 1 1 specimens,
39.1-133.0 mm SL. Pyloric caeca not counted. Irregularly
sized and arranged conical teeth on head of vomer, which
are embedded deeply in tooth ridge, about 24; compressed
teeth closely set on dentaries, about 50 in a 104.6-mm spec-
imen.

Body slender and subcylindrical with body depth to width
ratio 1.3-1. 6 in specimens larger than 45 mm SL. The ratio
is larger in smaller specimens. Eye very large, indicating con-
siderable allometric growth (Fig. 11); its diameter varying
linearly from 6.4 percent of SL in a 39.1 -mm specimen to
8.8 percent in a 133.0-mm specimen; eye diameter 2. 2-2. 6

times in head length in specimens larger than 50 mm, and
2. 7-3.0 times in a specimen smaller than 50 mm.

Dorsal margin of eye protuding above dorsal margin of
head in a specimen larger than 100 mm SL. Pupil oval, with
aphakic space developed anteriorly. Base of uppermost pec-
toral fin at or slightly higher than midpoint between lateral
line and ventral margin of the body. Origin of dorsal fin base
well behind center of body. Predorsal length less than 47
percent of standard length with smaller values in larger spec-
imen due to allometric growth (Fig. 1 1). Base of outermost
ventral fin below or just behind posterior end of dorsal fin
base. Adipose fin base just in front of vertical through center
of anal fin base.

Tip of snout, upper jaw, anterior part of gular area, bases
of ventral and caudal fins densely pigmented.

SIZE. The largest specimen is the 133.0-mm SL holotype.

DISTRIBUTION. Nansenia longicauda occurs in the sub-
tropical and temperate waters of the Pacific and Atlantic
oceans. Based on collection data, this species appears to be
an oceanic mesopelagic species (Fig. 22).

ETYMOLOGY. The specific name is derived from Latin,
longus meaning long and cauda meaning tail referring to the
long and slender caudal peduncle.

Nansenia tenera new species

Figures 12, 21

HOLOTYPE. ZMC PI 962 1-23 (1, 129.0), Dana sta.
3975-1, 35°42'S, 18°37'E, 31 Jan. 1930, 3000 mwo.

Contributions in Science, Number 352

Kawaguchi and Butler: Fishes of the Genus Nansenia 1 1

-I 10 mm

-i 10 mm

C

i 1 10 mm

Figure 13. Nansenia ardesiaca. a. FMNH 57086, 164.5 mm SL, holotype from off Japan; b. USNM 93354, 136.0 mm SL, one of the seven
paratypes of N. schmitti from off the Philippines; c. ZMUC P19624-25, 183.5 mm SL, from off the northeast coast of South Africa.

PARATYPES. ZMC PI 962 1-23 (2, 1 12.0-1 17.4), collec-
tion data as for the holotype; ZMC not catalogued (1, 105.6),
Dana sta. 3975-11, 35°42'S, 18°37'E, 31 Jan. 1930, 2500 mwo;
ISH 1009/73 (1, 117.0), FFS Wa/ther Herwig 696-73,
55°39'N, 25°42'W, 22 Sep. 1973, 400-410 m; ISH 829/71
(1, 101.0), FFS Walther Herwig 376/71, 39°55'S, 26°02'W,
11 Mar. 1971, 0-2000 m.

DIAGNOSIS. Differs from other species of Nansenia in
having four branchiostegal rays, 42-43 vertebrae, and 43-
46 gill rakers.

DESCRIPTION. D 1 1-12; A 8-9; P 1 1-12; V 1 1-12; gill
rakers 13-15 + 1 + 28-32, total 43-46 with 43(1), 44(3),
45(1), and 46(1); branchiostegal rays four; vertebrae 42 in
one North Atlantic specimen, 43 in the five South Atlantic
specimens. Pyloric caeca 7-9 in three specimens; about 16
conical teeth irregularly arranged on head of vomer; about
60 compressed teeth on dentaries in a 1 12-mm specimen.

Eye large, its diameter 2. 4-2. 7 in head length; nearly one-
third to one-quarter of eye protruding above the dorsal mar-
gin of head. A golden colored, crescent-shaped pigment on

12 Contributions in Science, Number 352

Kawaguchi and Butler: Fishes of the Genus Nansenia

1 0 mm

Figure 14. Nansenia tenuicauda. ISH 1010-71, 69.0 mm SL, paratype.

posterior half of iris. Aphakic space developed; pupil acorn-
shaped with tip anteriorly directed. This type of pupil was
not observed in any other species.

Upper end of pectoral fin base below midpoint between
lateral line and ventral margin of body; its lower end above
ventral margin of body by a space greater than length of
pectoral fin base. Dorsal fin originating in front of center of
body. Ventral fin base above posterior half of anal fin base
in the South Atlantic specimens, but above the midpoint in
one specimen from the North Atlantic.

Body color of preserved specimens light brown. The entire
body surface of three specimens, especially the ventral side,
is sparsely covered with guanine, which resembles silver pow-
der. Caudal fin base densely pigmented in the specimens from
the South Atlantic, but the pigmentation extends forward
becoming sparser anteriorly in the North Atlantic specimen.
SIZE. A specimen of 1 12.0 mm SL has maturing gonads.
DISTRIBUTION. Five specimens were collected in the
South Atlantic between 35°S and 39°S, and one specimen
from the subarctic region of the North Atlantic (55°39'N).

ETYMOLOGY. The specific name tenera is from Latin,
soft, referring to the consistency of the body.

Nansenia ardesiaca Jordan and Thompson, 1914

Figures 13, 22

Nansenia groenlandica: Tanaka, 1911:15-16, pi. Ill, fig. 13
(non Reinhardt, 1840) (Sagami Bay, Japan).

Nansenia ardesiaca Jordan and Thompson, 1914:210-21 1,
pi. XXIV, fig. 2 (orig. descr., off Japan).

Nansenia tanakai Schmidt, 1918:15 (orig. descr. Japan).
Bathymacrops microlepis Gilchrist, 1922:53-54, pi. IX, fig.

2 (orig. descr., east coast of South Africa).

Microstoma (Euproserpa) schmitti Fowler, 1934:256-257, fig.
18 (orig. descr. Philippines).

Nansenia macrolepis: Cohen, 1958:56 (east coast of South
Africa).

Nansenia robusta Abe, 1976:27-31, figs. 1-6 (orig. descr.

South China Sea).

MATERIALS. Off Japan: FMNH 57086, 164.5 mm SL,
Okinose, Sagami Bay, central Japan (holotype of N. ardesia-
ca)', KYO 4434 (2, 157.5, 144.0), southern Japan, KU 7013
(1,1 20.0), Mimase fish market, Shikoku Is., Japan; KU 13012
(1, 141.5), Tosa Bay, Shikoku Is., 430-460 m; KU 131 25—
13130, 13243, 13244, 13279 (9, 124.0-155.0), 15 km east
of Ashizuri Misaki Pt., Shikoku Is., bottom trawl 420-
555 m.

Southeast Asian Seas: USNM 92327 (holotype of N.
schmitti) (1, 194.5), R/V Albatross, sta. D5445, Atalaya Pt.,
Batag Is., off Samar, the Philippines, 3 Jun. 1909; USNM
93354 (1, 136.0), R/V Albatross, sta. 5589, 04°12.10'N,
118°38.08'E, Mabul Is., off the Philippines (one of seven
paratypes of N. schmitti)', KU 15792 (1, 88.5, damaged),
R/V Hakuho Maru, Cr. KH72-1, sta. 20, 05°40.9'N,
1 19°46.3'E, bottom trawl 460-500 m; KU 16644 (1, 207.0),
R/V Hakuho Maru, KH72-1, sta. 26, 09°27.0'S, 127°58.6'E,
Timor Sea, bottom trawl 690-850 m.

Off South Africa: USNM 2034391-10 (10, 174.0-192.0),
R/V Anton Bruun, 22°25'S, 35°54'E, shrimp trawl, 0-740 m;
ZMC P19624-25 (2, 183.5, 185.0), Galathea exped. 1950-
1952, sta. 203, 25°36'S, 35°21'E off Natal, 2015 m; USNM
not catalogued, SOSC ref. no. 1701-2 (2, 150.0, 165.0), IIO
Exped. R/V Anton Bruun, Cr. no. 8, sta. 397C, 26°07'S,
34°1 l'E, bottom trawl 600-665 m; CAS-SU-31501 (1, 127.5),
off South Africa.

DIAGNOSIS. Differs in having four branchiostegal rays,
predorsal length less than 49 percent of SL, length of caudal
peduncle not more than 1 5 percent of SL, number of gill
rakers on first arch 27-35.

DESCRIPTION. D 9-10; A 9-10; P 12-14 (rarely 1 1); V
11 (rarely 10, 12); gill rakers 9-11 + 1 + 19-23, total 29-
35 with 29(1), 30(1), 31(1), 32(5), 34(2), and 35(2) in 12
specimens from off Japan; 9 + 1 + 18-21, total 27-31 with

Contributions in Science, Number 352

Kawaguchi and Butler: Fishes of the Genus Nansenia 13

Figure 15. Nansenia pelagica. ISH 649-74, 102.0 mm SL, holotype.

27(1), 30(1), and 31(1) in three specimens from off the Phil-
ippines; 8-10 + 1 + 19-21, total 28-31 with 28(1), 29(2),
30(7), and 3 1(4) in 14 specimens from off South Africa; bran-
chiostegal rays four; vertebrae 46-48 with 46(6), 47(12), and
48(2).

Gill raker counts were rather variable and higher on the
average in the specimens from off Japan. Ranges of these
counts, however, overlap among specimens from the three
different regions. No significant geographical variation was
found in other counts and measurements.

Pyloric caeca 8, 8, and 9 in three specimens from off South
Africa, and 7, 9, and 9 in two Japanese specimens; conical
teeth on vomerine series 25-35 and teeth on dentaries 65-
75.

Eye large, diameter 2. 3-2. 5 times in head length, dorsal
margin protruding above the margin of head. A roughly cres-
cent-shaped, yellowish pigment patch present in the posterior
half of iris. Aphakic space well developed. Anterior hori-
zontal space nearly equal to width of iris, about twice width
of posterior space. Aphakic space probably less well devel-
oped in smaller specimens, as is the case in other species.

Upper end of pectoral fin base above midpoint between
lateral line and ventral margin of body, its lower end well
above ventral margin of body by a distance more than (usu-
ally 1.5 to twice) length of pectoral fin base. Dorsal fin origin
in front of center of body. Ventral fin base below or just
behind posterior end of dorsal fin base. Adipose fin base
above posterior half of anal fin base. Tip of mouth, snout,
adipose, caudal and ventral fin bases pigmented.

SIZE. This species reaches about 200 mm SL. A specimen
of 196 mm SL is reported to have ripe eggs (Abe, 1976).

DISTRIBUTION. Based on previous records, N. ardesia-
ca occurs in slope waters of Southeast Asian Seas, as far north

as off southern Japan. It also occurs in slope waters off the
east coast of South Africa (Fig. 22).

Nansenia tenuicauda new species

Figures 14, 22

HOLOTYPE. ISH 658/71, 84.0 mm, FFS Walther Herwig
sta. 363-11/71, 40°18'S, 39°12'W, 8 Mar. 1971, MT1600, 0-
328 m.

PARATYPES. ISH 1010-71 (2, 70.5, 69.0), FFS Walther
Herwig sta. 399-11/71, 40°34'S, 09°50'W, 18 Mar. 1971,
MT1600, 0-310 m.

DIAGNOSIS. Differs in having four branchiostegal rays,
predorsal length less than 49 percent of SL and 38-42 gill
rakers on the first arch. Slender caudal peduncle the least
depth of which is less than 5.5 percent of SL also distinguish
this species from other species of Nansenia.

DESCRIPTION. D 10-11; A 8-10; P 11-13; V 11-12;
gill rakers 1 1-1 5 + 1 + 25-26, total 38-42 with 38(1), 39(1),
and 42(1); branchiostegal rays four; vertebrae 46. Counts are
based on three specimens.

Pyloric caeca eight in one specimen. Conical teeth ca. eight,
irregularly arranged on head of vomer and ca. 26 compressed
teeth on dentaries. A pair of teeth on the symphysis longest
of all. Other teeth of lower jaw, as usually observed in other
species of Nansenia. are progressively smaller from the back
of each dentary to the front.

Body subcylindrical, with a depth to width ratio of 1.3-
1 .4. Eye large, its diameter 2. 8-3.0 times in head length; pupil
ovoid in shape with aphakic space developed posteriorly.

Upper end of pectoral fin base just below the midpoint
between lateral line and ventral margin of the body. Dorsal
fin origin well in front of center of the body. Ventral fin base

14 Contributions in Science, Number 352

Kawaguchi and Butler: Fishes of the Genus Nansenia

below the posterior end of dorsal fin base. Adipose fin base
above base of the last two anal fin rays.

Base of caudal fin densely pigmented, but pigment becom-
ing progressively sparser anteriorly. Body color of preserved
specimens light brown.

SIZE. The specimens examined, 69.0-84.0 mm SL, are
probably juveniles.

DISTRIBUTION. Restricted to the area near the sub-
tropical convergence in the South Atlantic, around 40°S.

ETYMOLOGY. From Latin, tenuis, slender, and from
cauda, tail, in reference to the slender caudal peduncle.

Nansenia pelagica new species

Figures 15, 16, 22

HOLOTYPE. ISH 649/74 (1, 102.0), sta. A.D. 61-11/74,
02°27'N, 34°52'W, 24 Jan. 1974, MT1600, 0-350 m.

PARA TYPES. Atlantic: ISH 1865/66 (1,75.5), FFS Wal-
ther Herwig sta. 182-66, 10°46'N, 23°54'W, 16 May 1966,
0-300 m; ISH 442/66 (1, 93.0), FFS Walther Herwig sta.
1 84-66, 06°25'N,24°34'W, 17May 1 966, 0-320 m; ISH 57 1/
66 (1, 101.0), FFS Walther Herwig sta. 186-66, 01°24'S,
25°58'W, 19 May 1966, 0-330 m; ISH 619/66 (1, 64.8), FFS
Walther Herwig sta. 187-66, 05°34'S, 26°58'W, 20 May 1966,
0-320 m; ISH 2066/71 (2, 101.5, 107.5), FFS Walther Her-
wig 467-71, 05°30'S, 16°28'W, 9 Apr. 1971, 0-1900 m.
Pacific: USNM 207532 (1, 83.7), 21°20'N, 158°20'W (off
Hawaii), 16 Sep. 1970, 0-725 m.

OTHER MATERIALS. Atlantic: ISH 928/68 (1, 58.5),
FFS Walther Herwig sta. 17-68, 04°43'S, 26°39'W, MT1600,
0-ca. 2000 m. Pacific: USNM 201704 (1, 27.0), R/V Swan,
28°48'N, 1 4 1 °59'W, 6 ft. - 1 KMT, 200-247 m; USNM 212102
(2, ca. 60.5, ca. 54, damaged), from the stomach of a Thunnus
albacares collected off the New Hebrides Is., 20°00'S,
170°03'E.

DIAGNOSIS. Differs from other Nansenia in having four
branchiostegal rays and 38-39 vertebrae. Although it is sim-
ilar to N. atlantica, and their distributions overlap in the
tropical Atlantic, N. pelagica differs from N. atlantica in
having lower vertebral counts (38-39 vs. 41-42), a longer
head (27-30 percent of SL vs. 20-26 percent), and a dark
pigment spot on the gular area that becomes obscure with
growth in specimens larger than 90 mm SL.

DESCRIPTION. D 9-10; A 8-9; P 9-10 (rarely 1 1); V
10-1 1; gill rakers 10-13 + 1 + 24-27, total 36-4 1 with 36(2),
37(1), 38(3), 39(2), and 41(1); branchiostegal rays four, ver-
tebrae 38-39 with 38(6) and 39(4). Counts are based on 10
specimens, 58.5-107.5 mm. Gill raker count of the one Pa-
cific specimen was higher than counts from the Atlantic (41
vs. 36-38). Other counts showed no remarkable difference
between the Pacific and Atlantic specimens.

Pyloric caeca seven and eight in the two Atlantic speci-
mens. About 35 conical teeth on vomerine series and about
60 teeth, resembling a closely spaced picket fence, on the
lower jaw of the specimens of 68.4 mm SL.

Body slender and slightly compressed, with a body depth-
width ratio of 1.3-1. 6 in specimens larger than 58 mm SL.
Eye large, diameter 2. 7-2. 8 in head length in specimens 58.5-

%

60

O

o

50

o

o

— o o__

Pre D

CO

40-

o

30

o

H L

=

o °

o

^ ° ~

20

10

E D

o

o

0 . * • * • * • * • —

10 20 30 40 50 60 70 80 90 100 110

S L ( mm )

Figure 16. Nansenia pelagica. Predorsal, head lengths, and eye
diameter in relationship to standard length.

93.0 mm SL, and 2.4 to 2.5 times in head length in specimens
99.5-107.5 mm SL. Yellowish pigment, in a roughly cres-
cent-shaped patch, develops on posterior half of iris with
growth. Aphakic space developed in front of lens. This space
is inconspicuous in juveniles.

Upper end of pectoral fin base just below midpoint between
lateral line and ventral margin of body, its lower end above
ventral margin of body by a distance about 1.5 times as long
as length of pectoral fin base. Origin of dorsal fin base just
behind center of body. Adipose fin base over anterior half
of anal fin base.

Body skinned in most specimens, but lateral line pockets
remaining in some. Bases of caudal, dorsal, ventral and adi-
pose fins, and dorsal margin darker than lateral side of body.
Inside of opercle lined with dark membrane. Dark pigment
spot on gular area clearly recognizable, especially in juveniles
less than 70 mm SL.

SIZE. The largest specimen examined, probably an adult,
was 107.5 mm SL.

DISTRIBLITION. Nansenia pelagica occurs in the trop-
ical Atlantic between 1 5°N and 1 0°S. In the Pacific it is known
only from the subtropical eastern North Pacific between Ha-
waii and North America and the tropical western South Pa-
cific off the New Hebrides Islands.

ETYMOLOGY. In reference to the pelagic life of this
species.

Nansenia atlantica Blache and Rossignol, 1962

Figures 17, 21

Nansenia atlantica Blache and Rossignol, 1 962: 105-106, fig.

1 (orig. descr. 16 mm SL, 01°55'S, 8°30'E; 15 mm SL,

03°36'S, 09°10'E).

MATERIALS. ISH 335/66 (2, 88.5, one damaged), FFS
Walther Herwig sta. 182-66, 10°46'N, 23°54'W, 0-300 m;
ISH 457/68 (4, 90.5-1 12.8, one damaged), FFS Walther Her-
wig sta. 11-1-68, 16°14'N,22°24'E, 0-160 m; ISH 686/68 (1,
ca. 161, damaged), FFS Walther Herwig sta. 13-11-68,
08°2 1 'N, 24°10'W, 0-520 m; ISH 13-11/68 (1, 167.5), FFS
Walther Herwig sta. 1 1-III-68, 16°08'N, 22°22'W, 0-580 m.

Contributions in Science, Number 352

Kawaguchi and Butler: Fishes of the Genus Nansenia 15

-i 1 0 mm

H

Figure 17. Nansenia atlantica. ISH 475-68, 104.5 mm SL, showing proximal part of adipose fin densely pigmented.

DIAGNOSIS. Differs from other species of Nansenia in
having four branchiostegal rays, origin of dorsal fin base in
front of center of the body, 41-42 vertebrae and dark pigment
spot at the base of adipose fin. This species is closely related
to N. oblita as all counts overlap. The densely pigmented
proximal part of the adipose fin base in N. atlantica distin-
guishes the two.

DESCRIPTION. D 9-10; A 8-9; P 12-13; V 10-11; gill
raker on the first arch 9-13 + 1 + 19-23, total 30-36 with
30(1), 31(2), 33(2), 35(1), and 36(1); branchiostegal rays four;

vertebrae 41-42 with 41(3) and 42(5). Counts are based on
five to eight specimens.

Pyloric caeca seven in three specimens and nine in one
specimen. Conical teeth on head of vomer 35, about 140
teeth on dentaries in one specimen.

Body slender and compressed laterally with body depth-
width ratio, 1.4-1. 6. Eye diameter 2. 7-3.0 times in head
length, viewed laterally its dorsal margin touching or pro-
truding slightly higher than dorsal margin of head. Yellowish
iridescent pigment pattern, roughly crescent-shaped, recog-

i 1 10 mm

Figure 18. Nansenia oblita. USNM 92241, 39.1 mm SL, juvenile.

16 Contributions in Science, Number 352

Kawaguchi and Butler: Fishes of the Genus Nansenia

10 mm

Figure 19. Nansenia crassa. LACM 9808-32, 165.2 mm SL.

nizable in the posterior half of iris. Pupil elliptical with aphakic
space in front of lens.

Upper end of pectoral hn base well below midpoint be-
tween lateral line and ventral margin of body. Dorsal hn base
originating just below center of body. Ventral hn base just
in front of posterior end of dorsal hn base. Adipose hn base
above center of anal hn base.

Body skin brown and easily rubbed off. The bases of adi-
pose and caudal hns densely pigmented.

REMARKS. The original description of N. atlantica was
based on two juvenile specimens of 16 and 1 5 mm SL. There
is a considerable size gap between the type material and the
smallest specimen available. Therefore, the present speci-
mens are assigned to N. atlantica based on the coincidence
of: (1) counts of branchiostegal and anal hn rays; (2) locality;
(3) a wholly pigmented dark body surface. In the tropical
Atlantic, two species of the Br-4 group occur, namely N.
atlantica and N. pelagica. Juveniles of the latter differ from
the former in the absence of complete pigmentation on the
body.

SIZE. The largest specimens of 167.5 and 161 mm SL,
collected in January, were mature males. An immature fe-
male of 1 12.0 mm SL was collected at the same time.

DISTRIBUTION. All specimens, including the types, were
collected in the eastern tropical Atlantic between 20°N and
10°S (Fig. 21).

Nansenia oblita (Facciola, 1887)

Figures 18, 21

Microstoma argent eum oblitum Facciola, 1887:193 (orig.

descr., Messina, Mediterranean).

Microstoma rissoanum Sarato, 1890 (orig. descr., Nice).
Microstoma oblitum: Belloti, 1888:224, hgs. 3a, 3aA.
Nansenia oblita: Schmidt, 1918:19-22, hgs. 13-14; Cohen,

1958:55.

Doubtful references:

Leuroglossus oblitus: Kuroda, 1947:25 (listed, no descrip-
tion, material unavailable); Kuroda, 1951:318.

Nansenia oblita: Matsubara, 1955:216 (cited from Kuroda,
1947).

MATERIALS. USNM 40075 (2, 40.5-48.5), from Mes-
sina, originally deposited in Royal Zool. Mus. Florence;
USNM 92241 (3, 34.2-46.0), Mediterranean, originally de-
posited in Milano Mus.

DIAGNOSIS. Differs in having four branchiostegal rays,
predorsal length longer than 50 percent of standard length,
gill raker count 28-30 and proximal part of adipose hn un-
pigmented.

DESCRIPTION. D 10-11; A 9-10; P 10-11; V 10-11;
gill rakers on the hrst arch 7-8 + 1 + 20-21, total 28-30
with 28(1) and 30(2); branchiostegal rays four on three spec-

°/o

70

0 1 1 1 t t—

0 50 100 150 200 230

S L ( mm )

Figure 20. Nansenia crassa. Predorsal, head lengths, and eye di-
ameter in relation to standard length.

Contributions in Science, Number 352

Kawaguchi and Butler: Fishes of the Genus Nansenia 17

Figure 21. Distributions of specimens examined of seven species of Nansenia. A single symbol may indicate more than one record.

imens; vertebrae 42-45 with 42(1), 43(1), 44(2), and 45(1).
Fin ray counts are based on four specimens.

Pyloric caeca eight in one specimen (seven reported by
Schmidt, 1918); about 18 conical teeth irregularly arranged
on head of vomer and about 30 compressed teeth on den-
taries in a specimen of 48.5 mm SL. Measurements are based
on four juveniles, 34.2-48.5 mm. The present counts and
measurements are not very different from those presented
for Mediterranean specimens by Schmidt (1918), except for
the slightly lower pectoral and ventral fin ray counts in our
material.

Body laterally compressed and stubby rather than slender
with body depth-width ratio, 1.3-1. 6. Eye diameter 3.0 to
3.4 in head length in juveniles (this ratio may be higher in
adults due to negative allometry as observed in the closely
related N. crassa)\ pupil nearly round in a specimen of 34.2
mm SL, horizontally elliptical in specimens larger than 39. 1
mm.

Upper end of pectoral fin base above or on midpoint be-
tween lateral line and ventral margin of body, its lower end
separated from ventral margin of body by a distance more

than length of pectoral fin base. Dorsal fin origin behind
center of body. Ventral fin base below posterior end of dorsal
fin base. Adipose fin base above the middle of anal fin base
or slightly in front of it.

Entire body covered with guanine; base of caudal and pro-
current caudal fin rays pigmented.

SIZE. Largest specimen reported by Belloti (1888) about
1 8 cm. Therefore the present material is probably all juvenile.

DISTRIBUTION. Common in the western Mediterra-
nean (Schmidt, 1918). Although Schmidt (1918) reported
two juveniles, 21 and 31.5 mm, from the temperate eastern
Atlantic south of the British Isles (48°43'N, 12°05'W), we
have seen only Mediterranean specimens, in spite of consid-
erable fishing effort by FFS Walther Herwig in the eastern
North Atlantic.

Nansenia crassa Lavenberg, 1965

Figures 19, 20, 21

Nansenia crassa Lavenberg, 1965:282-285, fig. 1, table 1
(orig. descr.).

18 Contributions in Science, Number 352

Kawaguchi and Butler: Fishes of the Genus Nansenia

MATERIALS. LACM 4425, holotype, 212.0 mm, R/V
Ve/ero, sta. IV8296, 33°1 7'N, 118°40'W, 1 336-1642, 9 Nov.
1962, 0-580 m; LACM 4426, paratype, 202 mm, R/V Ve-
lero, sta. IV7374, 33°28'N, 1 18°18'W, 0200-0550, 29 Jim.,
1961, 0-865 m; LACM 4427, paratype, 266 mm (275 mm
in original), R/V Velero, sta. IV8934, 33°14'N, 118°33'W,
1559-1858, 18 Sep. 1963, 0-485 m; LACM 9031-6(1,26.3),
R/V Velero IV, 33°20.0'N, 1 1 8°38.22' W, 0704-1022, 10ft.-
IKMT, 0-504 m; LACM 9068-1 (1, 44.5), R/V Velero IV,
33°36'48"N, 118°26'26"W, 1241-1445, lOft.-IKMT, 0-ca.
500 m; LACM 9681 (2, 33.5, 39.0), R/V Velero, sta. 11616,
Valero Basin, Mexico; LACM 9808-32 (1, 165.2), R/V Ve-
lero, San Clemente Basin, Calif.; LACM 9809-1 (2, 34.0,
35.0), R/V Velero, sta. 10682, 29°17'00"N, 118°10'30"W, 10
ft.-IKMT; LACM 9965-9 (4, 19.0-22.0), R/V Velero, sta.
9891, San Juan Seamount; LACM 35549 (1, 203.6), R/V
Velero, San Clemente, Calif.; SIO H51-190 (1, 238.0),
32°49'N, 1 17°41'W, 0-548 m; SIO 57-87 (1, 93.0), 29°I5'N,
! 26°07 W. off Baja Calif.

DIAGNOSIS. Differs from other species of the genus in
the following characters: ( 1 ) four branchiostegal rays; (2) pre-
dorsal length more than 49 percent of standard length; (3)
43-46 (usually 44-45) vertebrae; (4) length of caudal pe-
duncle more than 13 percent of SL and depth of caudal
peduncle less than 8.0 percent of SL (CPL/CPD ratio 1.7-
2.2) in the specimens more than 30 mm SL. Gill raker count
(35-37) also distinguishes this species from the closely related
N. oblita (28-30).

DESCRIPTION. D 9-10; A 9 (rarely 8); P 11-13; V 10-
1 1; gill rakers on the first arch 12-14 + 1 + 22-23, total 35-
37 with 35(2), 36(2), and 37(2); branchiostegal rays four;
vertebrae 43-46 with 43(1), 44(3), 45(5), and 46(2).

Pyloric caeca seven in two specimens; conical teeth in
vomerine series 20-35, teeth on dentaries 150-180 in five
specimens larger than 160 mm SL.

Morphometries based on five juveniles and seven adults
show remarkable allometric growth in head and predorsal
lengths (Fig. 20).

Eye diameter 3.3 to 4.0 in head length with larger values
in smaller specimens due to allometry; viewed laterally dorsal
margin (supraorbital bone) protruding slightly above dorsal
margin of head (less in juveniles); yellowish, crescent-shaped,
pigment patch in posterior part of iris, sometimes obscure
due to condition of preservation. Aphakic space well devel-
oped with its horizontal space about 1.5 times as large as
width of anterior part of iris.

Upper end of pectoral fin base well below midpoint be-
tween lateral line and ventral margin of body; its lower end
above ventral margin of body by a distance more than length
of its base. Dorsal fin origin behind center of body. Ventral
fin base below or slightly in advance of posterior end of dorsal
fin base. Adipose fin base above center of anal fin base or
slightly in front of it.

Skin around adipose fin base and caudal fin base pig-
mented. Ventral fin base slightly pigmented. Body color in
preserved specimen brown.

SIZE. The largest specimen examined was 266 mm SL,
with mature ovaries in September.

DISTRIBUTION. Nansenia crassa is found in the Cali-
fornia Current region between 25°N and 35°N, where its
distribution overlaps that of N. Candida off mid- and south-
ern California. It also occurs in the eastern tropical Pacific
along the equator between 8°N and 7°S west to at least 1 26°W
(Ahlstrom, 1971, 1972). The pattern of occurrence of adult
specimens is associated with slope waters off California, sug-
gesting that the adults there are benthopelagic life on the
continental or insular slope. Larvae and juveniles are pelagic
in the upper 200-m layer and are more broadly distributed
than are adults. The larvae of N. crassa can be distinguished
from those of N. Candida. The geographical distributions of
the larvae of the two species correspond largely to the dis-
tributions of the adults, although there is a larger area of
overlap in the larvae. Specimens between 60 mm and 160
mm SL are completely lacking in our material. The size at
which the specimens begin to disappear from our material
corresponds to an abrupt change in allometric growth (Fig.
20). At this size, 50-60 mm SL, N. crassa may change its
vertical distributional pattern from pelagic to benthopelagic
life and become unavailable to either pelagic micronekton
nets or bottom trawls with coarse mesh.

DISCUSSION

DISTRIBUTION PATTERN AND
ZOOGEOGRAPHY

There are two types of distribution patterns, coastal and
oceanic, among the species of Nansenia.

Nansenia ardesiaca is restricted to slope waters associated
with insular or continental slopes, indicating a pseudo-ocean-
ic distribution (Krefft, 1974). Sampling data suggest that this
species is pelagic during larval and juvenile stages, but be-
comes benthopelagic as an adult. A distribution pattern sim-
ilar to that of N. ardesiaca is not uncommon among slope
water species of myctophids such as Diaphus watasei, D.
garmani, D. suborbitalis, Benthosema fibulatum, and B.
pterotum (Nafpaktitis, 1978; Gjosaeter, 1981). These myc-
tophids are distributed in Southeast Asian Seas and extend
to off southern Japan and westward in the Indian Ocean to
the Mozambique Channel, although few samples exist from
the slope waters off India. Of the remaining 1 1 species, three
(N. crassa, N. oblita, and N. Candida) show an intermediate
pattern of distribution between coastal and oceanic (Figs. 2 1 ,
22). Off southern California N. crassa has a coastal distri-
bution, however the larvae are oceanic in the eastern tropical
Pacific (Ahlstrom, 1971, 1972). The other nine species are
all oceanic.

The 1 3 species may also be grouped into cold-water forms,
four species; warm-water forms, seven species (sensu John-
son, 1982); and transitional forms, two species.

Of the four cold-water species, N. Candida occurs only in
the subarctic eastern Pacific. Kanayama and Amaoka (1983)
described Nansenia sanrikuensis, the validity of which re-
quires further confirmation, from the subarctic waters of the
western Pacific off Japan. Species assemblages have been
recognized in the myctophids of both the eastern and western

",

Contributions in Science, Number 352

Kawaguchi and Butler: Fishes of the Genus Nansenia 19

Figure 22. Distributions of specimens examined of six species of Nansenia. A single symbol may indicate more than one record.

parts of the subarctic Pacific, which includes Diaphus theta,
Lampanyctus ritteri, L. jordani, Stenobrachius leucopsarus,
S. nannochir, Symbo/ophorus calif orniensis, and Tarleton-
beania crenularis (Wisner, 1974). Clinal variation occurs in
the eye diameter of D. theta (Nafpaktitis, 1978; Kawaguchi
and Shimizu, 1978) and in the shape of the caudal luminous
scales of S. californiensis (Wisner, 1974 and pers. commun.)
A similar species-subspecies problem exists in Tarleton-
beania (Wisner, 1959; Becker, 1966). Definitive comparative
examination of these species assemblages based on materials
not only from both sides of the subarctic Pacific, but from
the central area is needed. At present, it is not possible to
establish either a clear break in the distribution of the eastern
and western populations (or species) or clinal variations in
characters.

Other cold-water species, N. tenera and N. groenlandica,
belong to a subpolar-temperate group based on the system
of distribution patterns recognized for Atlantic mid water fishes
by Backus et al. (1977). These two species occur in both
hemispheres, although N. groenlandica in the southern hemi-
sphere is restricted to the subpolar-temperate region of the
eastern South Pacific. However, some bipolar species of the
subpolar-temperate group in the Atlantic such as Lampa-
nyctus macdonaldi (Wisner, 1974; Backus et al., 1977) and
Gonostoma bathyphilum (Mukhacheva, 1972) are found
westward to the eastern South Pacific and occur north to
Chile. Therefore additional sampling might establish the
presence of N. groenlandica in the subantarctic Atlantic. Of
the remaining cold-water species, N. antarctica is probably

circumpolar along the Antarctic convergence, although col-
lections are needed in the Indian Ocean sector.

Two of the seven warm -water species, N. atlantica and N.
megalopa, occur only in the eastern tropical Atlantic. Ac-
cording to the Atlantic faunal regions recognized by Backus
et al. (1977), both species inhabit the Guinean Province of
Atlantic Tropical Region (Fig. 21). The distribution of N.
atlantica appears to be restricted to the more productive area
near the Mauritanian upwelling and to just off the Guinean
coast. The other two warm-water species, N. longicauda and
N. pelagica, are widely distributed in the Atlantic and Pacific
oceans with the former restricted to the subtropical region
between 20°N and 40°N and the latter to the tropical region
between 20°N and 20°S in the Atlantic and between 30°N
and 20°S in the Pacific. Nansenia ahlstromi is restricted to
the central Eastern North Pacific. There is nothing to add to
the previous discussion of the distribution of the remaining
two warm-water species, N. oblita and N. ardesiaca.

BODY COLOR

There have been many descriptions of the body color of
Nansenia, which suggest two patterns, silvery and colored.
During the course of this study, however, it has been observed
that the silvery color pattern is variable, depending upon
preservative; it is easily lost in formalin. For example, the
type of N. ardesiaca (Fig. 1 3a), which is now dark colored,
was originally described as “color brilliant silvery . . . scales
of a bright, silvery appearance” (Jordan and Evermann, 1 896).

20 Contributions in Science, Number 352

Kawaguchi and Butler: Fishes of the Genus Nansenia

Some specimens examined (Fig. 1 3b) still bear complete gua-
nine after more than 50 years of preservation in ethanol in
a dark place. But these specimens are usually so soft that
careful handling is necessary, indicating that they were not
fixed well in formalin.

INTERRELATIONSHIPS

Various character states studied here are shared indepen-
dently among 13 species and it is difficult to discuss the
relationships within the species of Nansenia. Although the
key separates species on the number of branchiostegal rays,
loss of a branchiostegal ray could have occurred more than
once. It is also possible to separate the species of this genus
into three groups based on morphology, mainly of the caudal
peduncle. With the exception of N. antarctica, the species of
Nansenia are separable into the following three distinctive
groups based on the ratio of caudal peduncle length to caudal
peduncle depth (CPL/CPD, Tables 1 and 2):

Stubby-tailed group CPL/CPD = 1.0-1. 5

Intermediate group CPL/CPD = 1.7-2. 3

Slender-tailed group CPL/CPD = 2.7-4. 1

The ratios are not overlapping. The stubby-tailed group in-
cludes four species: N. ahlstromi (Br 3), N. oblita (Br 4), N.
atlantica (Br 4), and N. pe/agica (Br 4). The slender-tailed
group includes: N. groenlandica (Br 3), TV. tenuicauda (Br 4),
N. megalopa (Br 4), and N. longicauda (Br 4). Of the re-
maining five, four species, N. Candida (Br 3), N. crassa (Br
4), N. ardesiaca (Br 4), and N. tenera (Br 4), are included in
the intermediate group. Nansenia antarctica (Br 3) shows
remarkable allometric growth in the caudal peduncle; the
range of the CPL/CPD ratio for specimens less than 1 10 mm
SL was 2. 5-3.0; but the ratio for specimens larger than 1 10
mm SL was 2.0-2. 5. This overlaps the range between inter-
mediate and slender-tailed groups.

No correspondence is recognized between groupings based
on the number of branchiostegal rays or morphology. This
suggests that natural groups cannot be defined by branchio-
stegal ray counts alone.

ACKNOWLEDGMENTS

We thank the late E.H. Ahlstrom whose interest in and en-
thusiasm for fish systematics were largely responsible for this
study. We are greatly indebted to H.G. Moser and his col-
leagues of the Southwest Fisheries Center for their continuous
help and kindness during the course of this study. The fol-
lowing people and their institutions allowed us to examine
the materials in their care: E. Bertelsen (ZMUC); G. Krefft
and A. Post (ISH); R.J. Lavenberg (LACM); R.H. Rosenblatt
(SIO); W.G. Pearcy (OSU); S.H. Weitzman (USNM); R.K.
Johnson (FMNH); W.N. Eschmeyer (CAS); G.H. Burgess
(UF); O. Okamura (KU). We would also like to thank D.M.
Cohen (LACM), R.H. Rosenblatt (SIO), and D.L. Stein (OSU)
for reading the manuscript and for their many helpful com-
ments. The authors thank I. Barrett, director of the Southwest
Fisheries Center and Professor Emeritus R. Marumo of Uni-
versity of Tokyo for giving them a chance to complete this
study.

Partial financial support from the Japanese Ministry of
Education, special project research “Studies on biological
processes in the sea and technical development of marine
resources” (No. 57108013) is also acknowledged.

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pp.

Accepted 28 March 1984.

22 Contributions in Science, Number 352

Kawaguchi and Butler: Fishes of the Genus Nansenia

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KENTRIOBON O&SCUkUS (KEI.I,OG€? 1931), A FOSSIL DOLPHIN
(MAMMALIA: KENll?i.ODONlIiT]f4!E;): FROM THE MIOCENE
SHARKTOOTH HILL BONEBED IN CAIJFORNIA

Lawrence G. Barnes and Edward MSfclseil

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issued its ureeulair intervals m three major series: the articles in each series are numbered
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sit Contributions in 'Science, a misce

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Science liuUetln, a miscellaneous series of monographs describing original research
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KENTRIODON OBSCURUS (KELLOGG, 1931), A FOSSIL DOLPHIN
(MAMMALIA: KENTRIODONTIDAE) FROM THE MIOCENE
SHARKTOOTH HILL BONEBED IN CALIFORNIA

Lawrence G. Barnes' and Edward Mitchell1 2

ABSTRACT. The fossil odontocete genus Kentriodon Kellogg, 1927,
is the type genus of the delphinoid family Kentriodontidae, a family
which has no living representatives. The genus Kentriodon has pre-
viously included only one named species, a small dolphin called
Kentriodon pernix Kellogg, 1927, from the Middle Miocene Calvert
Formation in Maryland and Virginia on the east coast of North
America. Some bones from Japan and California have previously
been tentatively referred to the genus. We have concluded that the
species Grypolithax obscura Kellogg, 1931, heretofore known only
by isolated periotics from the Middle Miocene Sharktooth Hill Bone-
bed in California, belongs in Kentriodon. The genus Grypolithax
Kellogg, 1931, is therefore a junior synonym of Kentriodon Kellogg,
1927. A second species named on isolated periotics from the Shark-
tooth Hill Bonebed, Grypolithax pavida Kellogg, 1 93 1 , is a synonym
of G. obscura. The appropriate binomen for the species is Kentriodon
obscurus (Kellogg, 1931). We assign to this species a partial skull
and additional isolated periotics from the same bonebed.

Other isolated periotics from rocks stratigraphically below the
Sharktooth Hill Bonebed in California resemble those of Kentriodon
spp. and a closely related kentriodontid, Delphinodon dividum True,
1912, which was also originally described from the Calvert For-
mation. Some of these have been referred to in previous literature,
but are illustrated and described here for the first time.

These newly reported specimens from California reinforce some
previous correlations between the upper part of the Round Mountain
Silt in California and the Calvert Formation in Maryland and Vir-
ginia. They also provide additional indications of a general pattern
of generic cosmopolitanism and specific endemism among the Mio-
cene odontocetes of the North Atlantic and North Pacific Oceans.

INTRODUCTION

Fossil odontocetes of the genus Kentriodon Kellogg, 1927,
are small Miocene dolphins in the extinct family Kentri-
odontidae. Kellogg (1928:33, 68) assigned the genus to the
family Delphinidae. Slijper later (1936:556) named a new
subfamily Kentriodontinae, which he placed within the Del-
phinidae, to include Kentriodon as well as Delphinodon div-
idum True, 1912, another Miocene dolphin. Barnes (1978)
altered the context and rank of Slijper’s family group name
by recognizing the family Kentriodontidae within the super-
family Delphinoidea. He recognized three subfamilies (Kam-
pholophinae, Kentriodontinae, and Lophocetinae) within the
Kentriodontidae, and the species included in this family are
now known by described fossils from Europe, Japan, New
Zealand, and both the east and west coasts of North America.
The importance of this family and its relationships to some
other odontocete families was commented on by Barnes.
He stated that kentriodontids might comprise the group from
which other living families of delphinoids, including modem
dolphins in the Delphinidae, have evolved, and they might
also be expected to occur widely in rocks of appropriate age

Contributions in Science, Number 353, pp. 1-23
Natural History Museum of Los Angeles County, 1984

around the world. Our interests in delphinoid evolution and
in the Middle Miocene cetaceans in the Sharktooth Hill Local
Fauna in California have led us to the present study.

The type species of Kentriodon, K. pernix Kellogg, 1927,
was named on the basis of two skulls, one of which was
collected with earbones and an articulated partial postcranial
skeleton (the holotype), of Middle Miocene age from the
Calvert Formation in Maryland. Until the present study, this
genus has had assigned to it only this one named species,
however, some references in the paleontologic literature have
suggested the former presence of the genus in the North Pa-
cific Ocean. The genus Kentriodon was questionably iden-
tified from Miocene rocks in Japan by Shikama, Hasegawa,
and Otsuka (1973; also cited in Okazaki, 1976:25), but spec-
imens documenting this identification have not been de-
scribed. Bames (1976:326) identified as cf. Kentriodon and
as a related kentriodontid, cf. Delphinodon dividum True,
1912, some isolated periotics of early Middle Miocene age
from the lower part of the Round Mountain Silt in Kern
County, California. The source of these is stratigraphically
lower than the Middle Miocene Sharktooth Hill Bonebed.
Other kentriodontid genera have been recognized based on
skulls from Miocene rocks in Europe and California (Kellogg,
1925:4—6; Bames, 1976, 1978), and from late Oligocene rocks
in New Zealand (Fordyce, 1980:328).

Kellogg (1931) prepared a preliminary description of the
mammals from the Sharktooth Hill Bonebed that comprise
a part of what we now call the Sharktooth Hill Local Fauna
(Mitchell, 1965:33; Mitchell and Tedford, 1973: fig. 3; Bames,
1 976:326-327). In this study, Kellogg named ten new species
of small odontocetes based solely on isolated periotics from
the bonebed. For many years the true identities and rela-
tionships of these species have remained unknown. The
problem of disparate skeletal parts has in some instances
precluded objective morphologic and taxonomic compari-
sons between the odontocetes from the Sharktooth Hill Bone-
bed and other taxa (even some studied by Kellogg himself)
known by skulls and skeletons from elsewhere in the world.
For example, Bames (1978) showed that Liolithax kernensis
Kellogg, 1931, from the Sharktooth Hill Bonebed is conge-
neric with Lophocetus pappus Kellogg, 1955, from the Cal-
vert Formation in Maryland.

In the present study, we show that Grypolithax obscura
Kellogg, 1931, from the Sharktooth Hill Bonebed is conge -

1. Section of Vertebrate Paleontology, Natural History Museum
of Los Angeles County, Los Angeles, California 90007.

2. Arctic Biological Station, Fisheries and Oceans, Ste. Anne de
Bellevue, Quebec, Canada H9X 3R4; and Research Associate, Nat-
ural History Museum of Los Angeles County.

ISSN 0459-8113

neric with Kentriodon pernix. Another species Kellogg named
from the bonebed, G. pavida Kellogg, 1931, is synonymous
with G. obscura and the properly emended binomen for the
species from California is Kentriodon obscurus (Kellogg, 1931).
We refer a partial skull and several more isolated periotics
from the Sharktooth Hill Bonebed to K. obscurus, and with
these we have been able to more fully characterize the species
and compare it with K. pernix. We also describe and illustrate
the specimens of kentriodontids that Barnes (1976) men-
tioned from stratigraphically lower levels of the Round
Mountain Silt in California.

MATERIALS AND METHODS
INSTITUTIONAL ACRONYMS

The specimens examined in the course of this study are in
the collections of the following scientific institutions: Cali-
fornia Academy of Sciences, San Francisco, California (CAS);
Natural History Museum of Los Angeles County, Los An-
geles, California (LACM); University of California Museum
of Paleontology, Berkeley, California (UCMP); National Mu-
seum of Natural History, Smithsonian Institution, Washing-
ton, D.C. (USNM).

Institutional locality numbers are given for specimens where
appropriate. Qualified investigators may obtain precise lo-
cality information by contacting the appropriate institutions.

COMPARATIVE MATERIAL

We have made extensive reference to and comparisons with
the type species of Kentriodon, K. pernix from the Middle
Miocene Calvert Formation in Maryland. We believe that
the holotype (USNM 8060) of K. pernix, which was collected
from Zone 5 relatively low in the Calvert Formation, and
Kellogg’s original (1927a) published referred skull (USNM
1 0670) from Zone 3 slightly lower in the formation, represent
the same species. These are the only skulls of K. pernix that
have been described in the literature.

We have identified one other small skull (USNM 21027)
as K. pernix. It was found at Plum Point, Maryland, where
Zone 6 of the Calvert Formation (also relatively low in the
rock unit) is exposed at sea level (see Clark, Shattuck, and
Dali, 1 904: pi. 5). These three specimens, therefore, comprise
our current concept of the species, but provide only limited
information on individual and ontogenetic variation within
the species.

There are several additional fossils of Kentriodon from the
Calvert Formation in Virginia and Maryland in the USNM
collections that are undescribed in the scientific literature.
Some of these are significantly different from both the ho-
lotype and referred specimens of K. pernix. Most are from
zones higher in the Calvert Formation, and are therefore
geologically younger than the holotype and referred speci-
mens of K. pernix. We conclude that one or more species in
addition to K. pernix are represented by these additional
specimens, but a detailed variability and taxonomic study of
Kentriodon from the Calvert Formation is beyond the scope
of the present study. In the absence of such a study of these

other Kentriodon specimens, we made some observations on
them that have influenced our diagnoses and comparisons.
Additionally, we have illustrated some of the periotics (Figs.
8-9) to demonstrate variable characters within the genus.

TERMINOLOGY

The morphology of a kentriodontid periotic is shown in Fig.

1 . The terminology used is derived or adapted from Denker
(1902), Boenninghaus (1904), and Kellogg (1928). Cranial
terminology is derived from Kellogg (1927a) and Fraser and
Purves (1960). Where we employ family group names with
different rank than originally proposed, we cite the author of
the emended rank following the original author.

Statistical analysis follows Simpson, Roe, and Lewontin
(1960).

SYSTEMATICS

Class Mammalia Linnaeus, 1758

Order Cetacea Brisson, 1762

Suborder Odontoceti Flower, 1867

Superfamily Delphinoidea (Gray, 1821)
Rower, 1864

Family Kentriodontidae (Slijper, 1936)
Barnes, 1978

Kentriodontinae Slijper, 1 936:556; as a subfamily of the fam-
ily Delphinidae.

Kentriodontidae. Slijper, 1958: label in fig. 36, emended rank
without explanation in text.

Kentriodontidae. Barnes, 1978:3; emended rank, as a family
of the superfamily Delphinoidea.

DISCUSSION. Barnes (1978) stated that a lack of fossae
in bones of the basicranium, reflecting a presumed lack of
extensive development of air sinuses of the middle ear air
sinus system, was a characteristic feature of the family Ken-
triodontidae. Aside from the usual, primitive odontocete
combination of peribullary and pterygoid air sinuses, we find
in Kentriodon spp. osteological evidence only for a middle
sinus adjacent to the glenoid fossa and for a postorbital lobe
of the pterygoid sinus. We find no fossae that would provide
evidence for a posterior sinus in the exoccipital, for any si-
nuses in the basioccipital or above the optic nerve (as in
phocoenids), for a large orbital lobe, or for an anterior sinus
extending onto the posterior part of the palate. These are all
locations where various modem odontocetes have been shown
to have air sinuses in their skulls (Fraser and Purves, 1960).
The fossa for the lobe of the pterygoid sinus within the pter-
ygoid hamulus is characteristically small in Kentriodontidae
compared with Delphinidae, Phocoenidae, Ziphiidae or Phy-
seteridae. No fossil kentriodontid skull has yet been de-
scribed with an ossified pterygoid hamulus flooring the ven-
tral surface of this sinus as in modem delphinids, phocoenids,
and monodontids, and it cannot be determined from the

2 Contributions in Science, Number 353

Barnes and Mitchell: Kentriodon from California

specimens presently available whether or not these animals
had an incomplete or non-ossified hamulus as in modem
platanistoid dolphins (see Fraser and Purves, 1960).

Subfamily Kentriodontinae Slijper, 1936

Kentriodontinae Slijper, 1936:556; as a subfamily of the fam-
ily Delphinidae.

Kentriodontinae. Bames, 1978:24; emended context, as a
subfamily of the family Kentriodontidae.

Kentriodon Kellogg, 1927

Kentriodon Kellogg, 1927a:4.

Grypolithax Kellogg, 1931:393.

EMENDED DIAGNOSIS OF GENUS. A genus of the
subfamily Kentriodontinae differing from Delphinodon divi-
dum by having a skull with a longer rostrum, the mesorostral
gutter not roofed over by the premaxillae at the anterior end,
a more concave lateral margin of the supraorbital process,
palatal surfaces of maxillae more transversely convex, and a
fossa for the postorbital lobe of the pterygoid air sinus on
the ventral surface of the frontal; differing from Leptodelphis,
Microphocaena, Pithanodelphis, and Sarmatodelphis by hav-
ing a more convex lateral margin of the supraorbital process,
flat rather than bulbous or convex nasal bones, and more
widely separated posterior ends of the maxillae at the cranial
vertex with a concomitantly wider exposure of the frontals,
differing from Leptodelphis and Pithanodelphis by having a
less elevated cranial vertex; and differing from Liolithax,
Delphinodon, Lophocetus, and perhaps other described gen-
era of Kentriodontidae by having the anterior-most pre-
maxillary tooth on each side elongated into a small tusk and
pointing anteriorly from the tip of the rostrum.

TYPE SPECIES. Kentriodon pernix Kellogg, 1927; type
by original monotypy.

INCLUDED SPECIES. Kentriodon pernix Kellogg, 1 927;
and Kentriodon obscurus (Kellogg, 1931), new combination.

Kentriodon obscurus (Kellogg, 1931),
new combination

Figures 2-7, 8d-t, 9d-t, 13b, 14b

Grypolithax obscura Kellogg, 1931:394.

Grypolithax pavida Kellogg, 1931:396.

EMENDED DIAGNOSIS OF SPECIES. A species of
Kentriodon characterized by and differing from K. pernix by
having skull with vertically short postorbital process of fron-
tal, posterior part of alveolar row curved medially on palatal
surface instead of extending along lateral margin of rostrum,
posterior maxillary alveoli directed more laterally than ven-
trally, lateral margin of maxilla adjacent to posterior part of
alveolar row thickened and squared off instead of being thin
and rounded, alveoli for maxillary teeth averaging 1.5 mm
in diameter instead of 3 mm, antorbital notch narrow and
directed anteriorly instead of wide and directed anterolat-
erally, supraorbital process of frontal thicker and more arched,

2 16 13

6 7

Figure 1. Right periotic of a kentriodontid, cf. Kentriodon sp.,
UCMP 83605, with anatomical structures labeled; a, cerebral or
dorsal view; b, tympanic or ventral view, natural size. 1— anterior
process, 2 — aquaeductus cochleae, 1— aquaeductus Fallopii. 4 —
aquaeductus vestibuli, 5— canal for facial nerve, 6— cochlear portion,
1 —fenestra ovalis, 8— fenestra rotunda, 9— foramen cent rale, 10 —
foramen singulare, 11— fossa for head of the malleus, 12 —fossa
incudis, 1 3 — internal acoustic meatus, 14 — posterior articular facet
for tympanic bulla, 1 5 — posterior process, 16 — tractus spiralis fora-
minosus.

medial margins of dorsal premaxillary surfaces on proximal
part of rostrum between antorbital notches flat instead of
inclined medially toward mesorostral gutter, palatine bone
and pterygoid sinus fossa extended farther anteriorly onto
palate instead of ending at level of antorbital processes, post-
orbital lobe of pterygoid sinus of middle ear air sinus system
occupying large fossa on ventral surface of supraorbital pro-
cess of frontal, internal acoustic meatus of periotic more
elliptical and less circular in shape.

HOLOTYPE. CAS 4349, right periotic, collected by
Charles Morrice in 1924.

TYPE LOCALITY. CAS locality 905, Sharktooth Hill
Bonebed, Kern County, California.

PARATYPE. CAS 4347, left periotic collected by Charles
Morrice in 1924 from CAS locality 905.

REFERRED SPECIMENS FROM THE SHARK-
TOOTH HILL BONEBED. LACM 21256, an incomplete

Contributions in Science, Number 353

Barnes and Mitchell: Kentriodon from California 3

skull lacking the extremity of the rostrum, the occipital shield,
and the basicranium, from LACM locality 1625, and 31
isolated periotics as follows: CAS 4348, left (holotype of
Grypolithax pavida ), CAS 4350, left (paratype of G. pavida),
both from CAS locality 905; LACM 21134, right, LACM
449 1 6, left, LACM 63800, left, all from LACM locality 1 625;
LACM 2 1 238, right, and LACM 48925, left, both from LACM
locality 1557; LACM 39696, left, from LACM locality 3232;
LACM 41473, left, from LACM locality 1 655; LACM 58893,
left, from LACM locality 6688; LACM 75371, left, from
LACM locality 3208; LACM 96150, right, LACM 121998,
LACM 121999, LACM 123477-123481, eight right, LACM
1 23482 and 1 23483 left, all from LACM locality 3 1 62; LACM
98712, left, from LACM locality 3160; LACM 104094, left,
from LACM locality 1622; LACM 123471-123473, three
right, from LACM locality 4672; LACM 123474, left, from
LACM locality 4874; LACM 123475, right, and LACM
123476, left, from LACM locality 4956.

FORMATION AND AGE. The holotype, paratype, and
all specimens here referred to Kentriodon obscurus were col-
lected from several localities scattered over several square
kilometers in the Sharktooth Hill Bonebed in the upper part
of the Round Mountain Silt. This bonebed is a single thin
stratum and is the source of the Sharktooth Hill Local Fauna
(Woodetal., 1941; Mitchell, 1 966:28-29; Mitchell and Ted-
ford, 1973; Barnes, 1976). It is correlated with the Barstovian
North American land mammal age, the “Temblor” provi-
sional provincial molluscan stage, the Relizian or Luisian
foraminiferal stage, and is approximately between 1 3 and 1 5
million years old (Wood et al., 1941:31, pi. 1 ; Weaver et al.,
1944:582, pi. 1; Evemden et al., 1964; Addicott, 1972; Sav-
age and Barnes, 1972:133, 1 40; Berggren and Van Couvering,
1974: hg. 11; Barnes, 1976:326-327; 1978:5-6; Repennmg
and Tedford, 1977: table 1). Based on these correlations, the
upper part of the Round Mountain Silt, including the bone-
bed, is approximately contemporaneous with the Calvert
Formation in Virginia and Maryland (Gazin and Collins,
1950:3; Kellogg and Whitmore, 1957:1022; Ray, 1976: fig.
1).

DESCRIPTION AND COMPARISONS. Skull. The re-
ferred skull (LACM 2 1 256, Figs. 2-7) of Kentriodon obscurus
from the Sharktooth Hill Bonebed is incomplete and has
suffered pre-depositional breakage and abrasion. The basi-
cranium, occipital area, rostral extremity, vomer and ptery-
goid sinus fossae have been broken off. The sutures between
the remaining bones were unfused, and the two halves were
separated along the midline when the skull was found in the
field. We have re-assembled it in its presumed original con-
figuration based on comparisons with specimens of Kentrio-
don pernix from the Calvert Formation. Due to some dis-
tortion, however, the left supraorbital process fits incorrectly
and attaches too low on the skull (Fig. 4a). On the right side
(Fig. 4b) the supraorbital process is in the correct position.
The measurements of the skull are as follows: total length as
preserved, 1 93.5 mm; breadth of rostrum at antorbital notch-
es, 67.7 mm; breadth of cranium at antorbital processes,
122.3 mm; interorbital width, 1 15.2 mm.

The skull is nearly the same size as both the holotype and

Kellogg’s ( 1 927a) referred skull of K. pernix. All three appear
to represent young adult animals because none has the ad-
vanced suture fusion and/or extreme development of rugos-
ities and processes seen on skulls of extremely old individuals
of living delphinoids. Additionally, in the holotype skeleton
of K. pernix most of the vertebral epiphyses, including those
on cervicals and caudals, are tightly appressed to the vertebral
centra but not fused, indicating that the individual had not
yet achieved physical maturity. Overall skull shape and pro-
portions are similar in the two species, as far as is known. It
is in cranial details that the two species differ.

The open mesorostral gutter on the distal part of the ros-
trum is apparently characteristic of Kentriodon pernix (Kel-
logg, 1927a: pis. 2, 6), but is uncharacteristic of species in
the family Kentriodontidae as a whole (Barnes, 1978: figs.
14-17). The premaxillae on the referred skull of K. obscurus
are partly broken away distally, but in addition to having the
usual wide open mesorostral gutter at the proximal end, they
begin to diverge anteriorly as though they would have also
left the gutter exposed distally as in K. pernix. There is no
indication that the medial margins of the premaxillae were
elevated adjacent to the posterior part of the mesorostral
gutter as they are in K. pernix. The anterior (= rostral) parts
of the premaxillae are comprised of bone which is denser
and of smoother surface than the adjacent maxillae.

On the skull of K. obscurus, the premaxillary foramina are
located on a transverse line between the antorbital notches
(Figs. 2-3). These foramina are located more posteriorly on
both the holotype and the referred skull of K. pernix. The
premaxillary sulci associated with these foramina are some-
what damaged from breakage and abrasion, but between the
right and left sides, the typical odontocete condition of three
sulci (anteromedial, posteromedial, and posterolateral, see
Barnes, 1 978: 1 3; Fordyce, 198 1 : 1034, text-fig. 2) can be seen.
Medial to the anteromedial sulcus and the premaxillary fo-
ramen, the premaxillary surface is rough and indicates the
area where the nasal plug muscle was attached (Lawrence
and Schevill, 1956: fig. 23; Mead, 1975). The part of the
premaxilla that in life underlay the premaxillary sac (the
premaxillary sac fossa of Fordyce, 1981:1035, text-fig. 2;
Mead, 1975) lies posterior to the posteromedial and pos-
terolateral sulci and lateral to the nares. This area is also
roughened, but this is the result of postmortem abrasion
because one small remnant of surficial bone on the left pre-
maxilla indicates the previous existence of a thin, smooth
bone surface on the premaxillary sac fossa as is typical of
delphinoids. The more anterior location in K. obscurus, com-
pared with K. pernix, of the premaxillary foramina, the three
associated sulci, and concomitantly the anterior margin of
the premaxillary sac fossae, is primitive, because these struc-
tures have moved progressively posteriorly during the odon-
tocete cranial telescoping process (Miller, 1923). The area
that was occupied by the premaxillary sacs is more elongate
anteroposteriorly in K. obscurus, and we consider this to be
primitive as well. These sacs in Kentriodon might have been
symmetrical because the premaxillary sac fossae are not
asymmetrical as they are in modem delphinids (cf. Mead,
1975).

4 Contributions in Science, Number 353

Barnes and Mitchell: Kentriodon from California

Both species of Kentriodon have relatively small bony nares
that are narrowly pointed anteriorly. This is also typical of
many other species of Kentriodontidae (see Barnes, 1978:
figs. 14-17), and contrasts with the large, round nares of
species of Delphinidae.

The lateral margin of the maxilla dorsal to the posterior
end of the tooth row and medial to the antorbital notch
typically is formed into a relatively thick, horizontally pro-
jecting shelf in most kentriodontids. In K. obscurus, this part
of the maxilla is proportionally wider and has a more squared
off and vertical lateral margin than in K. pernix, in which
the maxilla has a thinner, more rounded margin. The ant-
orbital notches of K. obscurus are relatively narrower than
those of K. pernix, because in K. obscurus, both the lacrimal
and the maxilla project in a more anteromedial direction,
thereby giving the antorbital process a different shape (Fig.

13).

On the supraorbital process, both the frontal and maxilla
are significantly thicker and thus more convex or domed in
K. obscurus than in K. pernix. The lateral margins of the
maxillae have been both chipped and abraded over the orbit
on the skull of K. obscurus. The frontal is therefore probably
exposed more in dorsal view than it was in life.

The postorbital process of the frontal of K. obscurus is
narrower, shorter, and more tapered distally than in K. per-
nix. These differences do not appear to be related to ontogeny
because the juvenile skull that we refer to K. pernix (USNM
21027) from the Calvert Formation is smaller than the skull
of K. obscurus, has characters indicating physical immatu-
rity, but has substantially longer postorbital processes. Ad-
ditionally, the postorbital processes on the three skulls of
Kentriodon pernix from the Calvert Formation have a hook-
like shape, in contrast to the straight process of K. obscurus.

Among modem Delphinidae, there is a fossa located at the
posterolateral comer of the palate lateral to the pterygoid
sinus fossa and which marks the position of an air sinus called
the anterior sinus (see Fraser and Purves, 1960). This area
of the palate of K. obscurus is convex and shows no devel-
opment of such a fossa. On skulls of K. pernix, this area of
the palate is distinctly less convex so that a transverse section
through the proximal part of the rostrum is nearly V-shaped.
Neither species has the type of fossa or concave area that in
some living odontocetes marks the location of an anterior
sinus, and there probably was no such sinus present in either
species of Kentriodon.

The fossa for the lobe of the pterygoid air sinus that filled
the pterygoid hamulus is larger and extends farther anteriorly
in K. obscurus than in K. pernix. The roof of this pterygoid
sinus fossa is marked by a tapered cavity on the ventral
surface of the left palatine bone. The palatines concomitantly
extend farther anteriorly and are more pointed in K. obscurus.
The anterior-most extent of the palatines in K. obscurus is
35 mm anterior to the antorbital notch, whereas the same
parameter on the holotype of K. pernix is only 10 mm and
on Kellogg’s referred skull (USNM 10670) it is 14.4 mm.

As on the referred skull (USNM 10670) of K. pernix, the
ventral surface of the vomer was only exposed in a very
narrow opening between the maxillae at about the middle of

the rostrum of K. obscurus (Figs. 14a, b). In both species,
posterior palatine foramina flank this area, and slightly far-
ther anteriorly a single anterior palatine foramen occurs in
each maxilla adjacent to the posterior-most palatal exposure
of the premaxillae.

The teeth of K. obscurus were significantly smaller in di-
ameter than those of K. pernix (Figs. 14a, b). In K. pernix,
throughout the tooth row, the alveoli have uniform diameter
(3 mm) and nearly equal spacing. In K. obscurus, however,
the alveoli that are still intact on the right side located just
posterior to the mid-length of the rostrum are only 1.5 mm
in diameter; one-half the size of those in K. pernix. The
alveolar rows on both sides of the specimen of K. obscurus
are incomplete because of breakage and abrasion. A row of
eight consecutive alveoli in a distance of 32 mm in the right
maxilla indicates that the species had more teeth than K.
pernix, which Kellogg (1927a:32) estimated at about 40 on
each side of each upper jaw. The alveoli of K. obscurus are
oriented in the maxilla so that they face more laterally than
in K. pernix. The alveolar rows do not extend as far poste-
riorly on the palate, and the posterior end of each tooth row
curves medially toward the midline.

Posterior to the orbit, K. obscurus has a large, oval fossa
in the ventral side of the supraorbital process of the frontal
that measures approximately 15 by 20 mm. In modem del-
phinoids, a fossa in this location holds the postorbital lobe
of the pterygoid air sinus (Fraser and Purves, 1960). We
conclude that K. obscurus had a relatively large sinus here.
On both the holotype and Kellogg’s referred skull (USNM
10670) of K. pernix, there is only a slight depression at this
place. This apparent difference in air sinus size is not clearly
diagnostic because the juvenile referred skull, USNM 21027
of K. pernix, has a deep recess for the sinus at this place.
Without a larger sample of specimens we cannot determine
whether the size or the extent of invasion of bone by the
sinus is variable in K. pernix, whether it is larger in geolog-
ically more recent individuals of K. pernix that occur higher
in the Calvert Formation, or whether it is a significant taxo-
nomic character that separates species.

Periotic. When Kellogg (1931) described Grypolithax ob-
scura and G. pavida from the Sharktooth Hill Bonebed, he
noted only a few differences between their holotypes. He
noted that the periotics of G. pavida had a flatter cerebral
surface. We now benefit from having a larger sample to study
and find that, within the anticipated range of morphology of
what we interpret as one species, the presence of a flat cerebral
surface is variable.

Among the 31 isolated periotics from the Sharktooth Hill
Bonebed that we refer to Kentriodon obscurus, there are at
least six (LACM63800, 48925, 21238, 4 1473, 7537 1,1 23476)
that are nearly identical in morphology to the periotic of the
holotype of K. pernix (USNM 8060). Each of the others has
some minor degree of difference. There was no periotic found
with the skull we have referred to K. obscurus, and except
for the holotype of K. pernix, there has been found only one
other skull of Kentriodon sp. in the Calvert Formation that
has an associated periotic. This specimen, USNM 187313,
which we believe is a different and probably undescribed

Contributions in Science, Number 353

Barnes and Mitchell: Kentriodon from California 5

6 Contributions in Science, Number 353

Barnes and Mitchell: Kentriodon from California

Figure 2. Kentriodon obscurus (Kellogg, 1931), referred skull, LACM 21256 from LACM locality 1625, dorsal view, natural size.

fmx

/

Contributions in Science, Number 353

Barnes and Mitchell: Kentriodon from California 7

Figure 3. Kentriodon obscurus (Kellogg, 1931), referred skull, LACM 21256 from LACM locality 1625. dorsal view, natural size. Abbreviations: aon — antorbital notch, fmx
maxillary foramen, fpmx— premaxillary foramen, Fr— frontal. La— lacrimal, mrg— mesorostral gutter, Mx— maxilla, n — nanal opening, Pmx — premaxilla.

8 Contributions in Science, Number 353

Barnes and Mitchell: Kentriodon from California

Kentriodon obscurus (Kellogg, 1931), referred skull, LACM 21256 from LACM locality 1625, a, left lateral view; b, right lateral view, natural size.

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Contributions in Science, Number 353

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Table 1. Some characters differentiating Kentriodon pernix and K.
obscurus. An asterisk denotes the more derived character states, as
deduced from the morphology of Agorophius and species of Squal-
odontidae.

Kentriodon obscurus Kentriodon pernix

1. Premaxillary foramina be-
tween level of antorbital
notches.

2. Lateral maxillary margin
anterior to antorbital
notches thick and squared
off.*

3. Antorbital notch narrow.

4. Premaxillae flat in area of
attachment of nasal plug
muscles.*

5. Postorbital process of
frontal short and thick.*

6. Alveoli for teeth small
(circa 1.5 mm in diame-
ter).

7. Posterior end of alveolar
row bends medially on
palate.

8. Teeth implanted in mid-
dle and posterior part of
alveolar row so as to face
ventrolaterally.*

9. Palatines and pterygoid
air sinus fossae on palate
extend farther anteriorly
beyond location of antor-
bital processes.*

10. Posterior part of palate
lateral to fossae for ptery-
goid air sinus convex.

1 1. Large fossa in ventral sur-
face of frontal for postor-
bital lobe of pterygoid air
sinus.*

12. Internal acoustic meatus
of periotic elliptical in
shape.

1 . Foramina posterior to
level of notches.*

2. Lateral margin thin and
rounded.

3. Antorbital notch wider.*

4. Premaxillae elevated me-
dially.

5. Process long and curved.

6. Alveoli larger (circa 3 mm
in diameter).*

7. Posterior end of alveolar
row follows lateral margin
of palate.*

8. Teeth face more ventrally.

9. Palatines and fossae do
not extend beyond level
of antorbital processes.

10. Palatal surface less con-

1 1. Fossa small or absent.

12. Meatus circular in shape.*

species of Kentriodon was collected from Zone 14, near the
top of the Calvert Formation, and the skull, periotic, and
postcranial skeleton show several significant differences from
the holotype and referred specimen of K. pernix. The periotic
of this specimen, USNM 1 8731 3 (Figs. 8a, 9a), has a prom-
inent, elevated, flat area on its cerebral surface that is more
pronounced than on any of the other periotics referred to

Barnes and Mitchell: Kentriodon from California 9

10 Contributions in Science, Number 353

Barnes and Mitchell: Kentriodon from California

I

Figure 6. Kentriodon obscurus (Kellogg, 1931), referred skull, LACM 21256 from LACM 1625, ventral view, natural size.

sod

Contributions in Science, Number 353

Barnes and Mitchell: Kentriodon from California 1 1

Figure 7. Kentriodon obscurus (Kellogg, 1931), referred skull, LACM 21256 from LACM locality 1625, ventral view, natural size. Abbreviations: aon-antorbital notch, fap—
anterior palatine foramen, fio— orbital apertures of the infraorbital foramen. Fr— frontal. La— lacrimal, Mx — maxilla, n — nanal opening, pos— fossa for postorbital lobe of the
pterygoid sinus of the middle ear air sinus system, Pmx— premaxilla, pts — fossa for pterygoid sinus of the middle ear air sinus system, Vo— cleft that originally contained the
palatal exposure of the vomer.

Figure 8. Periotics of Kentriodon Kellogg, 1 927: a, Kentriodon sp., Calvert Fm., Virginia, USNM 187313, right; b, Kentriodon pernix Kellogg,
1927, holotype, USNM 8060, left; c, Kentriodon sp., Calvert Fm., Virginia, USNM 214754, right; Kentriodon obscurus (Kellogg, 1931) from
the Sharktooth Hill Bonebed; d, LACM 41473, left; e, CAS 4349, holotype, right; f, CAS 4348, left (holotype of Grypolithax pavida Kellogg,
1931); g, CAS 4350, left (paratype of Grypolithax pavida)-, h, LACM 75371, right; i, LACM 63800, left; j, LACM 48925, left; k, CAS 4347, i

paratype, left; 1, LACM 21238, right; m, LACM 121998, right; n, LACM 58893, left; o, LACM 104094, left; p, LACM 44916, left; q, LACM
98712, left; r, LACM 21 134, right; s, LACM 39696, left; t, LACM 96150, right; all figures are stereophotographs of the cerebral (or dorsal)
surface, natural size.

i

l

(

Kentriodon. Such a flat surface is present, however, in varying
lesser degrees on eight of the periotics from the Sharktooth
Hill Bonebed referred to K. obscurus (including the holotype
of Grypolithax pavida ), as well as on the holotype periotic
of K. pernix. Another isolated periotic (USNM 214754, Figs.
8c, 9c) from the Calvert Formation in Virginia closely match-
es the holotype periotic of K. pernix, and although it has a
more rounded cerebral surface, we refer it to that species.

All of the periotics from the Sharktooth Hill Bonebed that
we refer to Kentriodon obscurus resemble the holotype peri-

otic of K. pernix by having the following characters: (1) sim-
ilar size; (2) relatively small anterior and posterior processes;

(3) an overall sinuosity in either cerebral or ventral view
owing to the fact that the anterior process is bent medially
and the posterior process is bent laterally; (4) cochlear portion
relatively small and broadly joined to the body of the periotic,
not narrowly joined to the body and extended medially as
in many primitive odontocetes; (5) posterior process bent |
ventrally at a sharp angle from the body of the periotic there-
by forming a sharp peak or angle on the posterior part of the

12 Contributions in Science, Number 353

Barnes and Mitchell: Kentriodon from California

Contributions in Science, Number 353

Barnes and Mitcheil: Kentriodon from California 13

Figure 9. Periotics of Kentriodon Kellogg, 1 927: a, Kentriodon sp., Calvert Fm., Virginia, USNM 187313, right; b, Kentriodon pernix Kellogg,
1927, holotype, USNM 8060, left; c, Kentriodon sp., Calvert Fm., Virginia, USNM 214754, right; Kentriodon obscurus (Kellogg, 1931) from
the Sharktooth Hill Bonebed; d, LACM 41473, left; e, CAS 4349, holotype, right; f, CAS 4348, left (holotype of Grypolithax pavida Kellogg,
1931); g, CAS 4350, left (paratype of Grypolithax pavida); h, LACM 75371, right; i, LACM 63800, left; j, LACM 48925, left; k, CAS 4347,
paratype, left; 1, LACM 21238, right; m, LACM 121998, right; n, LACM 58893, left; o, LACM 104094, left; p, LACM 44916, left; q, LACM
98712, left; r, LACM 21134, right; s, LACM 39696, left; t, LACM 96150, right; all figures are stereophotographs of the tympanic (or ventral)
surface, natural size.

cerebral surface; (6) anterior process bent anteroventrally
and having a groove or pit on its medial side; (7) the cleft
between the anterior process and the cochlear portion bearing
a small crease between the groove for the tensor tympani
muscle and the cochlear portion; (8) articular facet for the
auditory bulla on the posterior process large, concave and
striated; and (9) a raised rugosity lateral to the fossa for the
head of the malleus.

Statistical analysis of the Sharktooth Hill Bonebed sample

of K. obscurus periotics (Table 2) proved useful. The param-
eters chosen were width and length measurements. The mea-
surement of the cochlear portion is probably less variable
individually or allometrically than the other two, because the
sizes of the anterior and posterior processes apparently change
during ontogeny. Measurements of the holotype of K. ob-
scurus fall within the range of measurements for the sample,
and they differ from the mean less than the standard devia-
tion in all three parameters. The same situation exists when

14 Contributions in Science, Number 353

Barnes and Mitchell: Kentriodon from California

Contributions in Science, Number 353

Barnes and Mitchell: Kentriodon from California 15

Table 2. Statistical comparisons of the holotype periotic of Kentriodon pernix and the holotype and referred periotics of K. obscurus from the
Sharktooth Hill Bonebed.

Bonebed sample of K. obscurus*

K. obscurus holotype

K. pernix holotype

Measurement

Number

Observed

range

Mean

Standard

deviation

Coefficient
of variation

Measure-

ment

Difference
from mean

Measure-

ment

Difference
from mean

Total length

27

25.2-29.4

27.2

1.2

4.41

26.9

-0.3

28.3

1.1

Width at pars
cochlearis

29

16.5-19.5

18.0

0.8

4.44

18.2

0.2

17.4

-0.6

Antero-posterior
dimension of
pars cochlearis

28

13.2-15.9

14.7

0.7

4.76

14.1

-0.6

14.2

-0.5

* Sample includes paratype of K. obscurus and holotype and paratype of Grypolithax pavida.

the holotype of K. pernix is compared to the sample of K.
obscurus, except that it differs somewhat more from the mean
than does the holotype of K. obscurus, but is still within the
standard deviation.

The similarities among these periotics from the Atlantic
and Pacific coasts are not surprising. Kasuya (1973:72) and
Barnes (1976:321-322, 327) have pointed out that periotics
of congeneric species of cetaceans are very similar and that
in some instances closely related modem or fossil species
simply may not be differentiated based on periotics alone.

Among the periotics of K. obscurus, some characters are
variable. These are: ( 1 ) shape and relative size of the posterior
articular facet for the bulla; (2) degree of rugosity of this
articular facet; (3) position of the groove or pit on the medial
side of the anterior process; (4) presence or absence of a
tuberosity on the posterior side of the cochlear portion be-
tween the fenestra rotunda and the orifice of the aquaeductus
cochleae; (5) extent of development of an attenuated groove
in the anterior margin of the internal acoustic meatus at the
cerebral orifice of aquaeductus Fal/opii; and (6) extent of
development of a flat area on the cerebral surface of the body
of the periotic in contrast with a rounded, convex surface.
The only character that we recognize whereby the holotype
periotic of Kentriodon pernix may be separated from this
sample of periotics of K. obscurus is its more circular internal
acoustic meatus. Every other character exists in at least one
of the periotics from the Sharktooth Hill Bonebed that we
refer to K. obscurus.

The only other significantly large, documented sample of
periotics of a species of Kentriodontidae is the series of Lioli-
thax kernensis from the Sharktooth Hill Bonebed that was
described and analyzed by Bames (1978). The periotics from
the bonebed that we now refer to Kentriodon obscurus are
as variable as the sample Bames referred to L. kernensis.
These periotics are not referable to any of the other odon-
tocetes that Kellogg (193 1) named based upon periotics from
the Sharktooth Hill Bonebed.

Among these other species, the problematic odontocete
Platylithax robusta Kellogg, 1931, the primitive sperm whale
Aulophyseter morricei Kellogg, 1927, the platanistoid dol-
phin “ Squalodon" errabundus Kellogg, 1931, and the ken-

triodontid dolphin Liolithax kernensis all have periotics that
are substantially different from those belonging to Kentri-
odon. Periotics of another dolphin, Loxolithax sinuosa Kel-
logg, 1931, differ subtly by being flatter dorsoventrally, hav-
ing a larger cochlear portion, a differently shaped posterior
process, a larger fossa for the head of the malleus, a groove
on the lateral surface of the anterior process, and by lacking
any notable flattening of the cerebral surface. Periotics of the
two species of Lamprolithax Kellogg, 1931, differ from those
of Kentriodon in many of the same ways as do those of
Loxolithax sinuosa. Nannolithax gracilis Kellogg, 1931, has
a smaller periotic with a large anterior process. Oedolithax
mira Kellogg, 1931, has a larger periotic with a deeper cleft
separating the anterior process from the cochlear portion.

cf. Kentriodon sp.

Figures 10-11

cf. Kentriodon Kellogg, 1927. Bames, 1976:326.

REFERRED SPECIMENS. LACM 29549, left periotic,
collected by Richard W. Huddleston in 1969 from LACM
locality 3066, Kem River, Kem County, California; and
UCMP 83605, right periotic, collected by Richard C. Bishop
from UCMP locality V-6953, Kem County, California.

FORMATION AND AGE. Both of these specimens are
from localities stratigraphically below the Sharktooth Hill
Bonebed in the lower part of the Round Mountain Silt. This
part of the formation is early Middle Miocene in age, about
15 to 19 million years old (see Bames, 1976:326). Savage
and Bames (1972:133) have reported Hemingfordian land
mammals from these same strata. Addicott (1972) charac-
terized the age of these beds further as representing part of
the “Temblor” provisional provincial molluscan stage and
the upper part of the Saucesian or the Relizian foraminiferal
stage.

DESCRIPTION AND COMPARISONS. These two
periotics differ from each other in both size and morphology
and may actually represent two different species. Each shares
characters in common with both Kentriodon obscurus and
K. pernix, such as the strong apex on the cerebral surface

16 Contributions in Science, Number 353

Barnes and Mitchell: Kentriodon from California

Figure 10. cf. Kentriodon sp., left periotic, LACM 29549 from
LACM locality 3066; a, cerebral or dorsal view; b, tympanic or
ventral view; both natural size.

posterolateral to the orifice of aquaeductus vestibu/i, small
internal acoustic meatus, and anteromedially projecting an-
terior process— all characters that could be regarded as prim-
itive. The smaller of the two periotics, UCMP 83605, has
an anteroposteriorly elongate cochlear portion as in other
specimens of Kentriodon. The larger periotic, LACM 29549,
has a relatively smaller and more globose cochlear portion,
and in this respect it resembles the holotype periotic of the
primitive kentriodontid Kampholophos serrulus Rensberger,
1969 (see Rensberger, 1969: pi. 4, figs. f-h).

aff. Delphinodon dividum True, 1912

Figure 12

cf. Delphinodon dividum True, 1912. Barnes, 1976:326.

REFERRED SPECIMEN. LACM 41041, left periotic with
stapes, lacking extremity of posterior process, collected by
the late John E. Fitch, about 1970 from LACM locality 6602,
“Barker’s Ranch Faunal Site,” Kern County, California.

FORMATION AND AGE. This specimen is from a lo-
cality in the lower part of the Round Mountain Silt, below
the Sharktooth Hill Bonebed, in roughly the same strata as
the two previously described periotics. Its age is likewise early
Middle Miocene and between approximately 1 5 and 19 mil-
lion years old.

DESCRIPTION AND COMPARISONS. The compari-
sons that we have made are specifically only with Delphi-
nodon dividum. because that is the only species of the genus
for which a periotic is known. The holotype is a specimen
collected from the Calvert Formation and includes a skull,
mandible, and part of the postcranial skeleton. The type
species of the genus, D. mento Cope, 1868, as fixed by Hay
(1902), is known only by teeth.

As noted by Barnes (1976:326) this periotic closely resem-
bles that of Delphinodon dividum. Kentriodon spp. and Del-
phinodon dividum are closely related (cf. Kellogg, 1927a, 1928:
67-69; Slijper, 1936:556), and both have been classified in
the subfamily Kentriodontinae (Barnes, 1978). The periotic
with the holotype of Delphinodon dividum (see True, 1912;
pi. 25, figs. 6, 7) has many characters in common with the
isolated periotic (LACM 41041) from Kern County. Notable
among these are the large fossa for the head of the malleus,
the globose anterior process which is bent medially, and the
small, spherical cochlear portion. The periotic of D. dividum
is somewhat similar in overall shape to periotics of Kentri-
odon obscurus and K. pernix, but differs by not having the

Figure 11. cf. Kentriodon sp., right periotic, UCMP 83605 from
UCMP locality V-6953; a, cerebral or dorsal view; b, tympanic or
ventral view; c, lateral view; all natural size.

cochlear portion as broad anteroposteriorly, by having the
anterior process bent more medially, and by having the coch-
lear portion separated from the anterior process by a much
deeper fissure. The isolated periotic from Kern County
(LACM 41041) differs from the holotype of D. dividum by
being smaller, having a relatively smaller posterior articular
facet for the bulla, and by having a relatively smaller and
narrower internal acoustic meatus. The latter character may
be primitive in comparison with D. dividum. We believe this
isolated periotic from the Round Mountain Silt is congeneric
with Delphinodon dividum and represents an earlier, more
primitive species.

A very closely related, if not identical, species is repre-
sented by a periotic from correlative rocks in Japan. Okazaki
(1976:37-38, text-fig. 6, pi. 11, figs, la-c (where the scale is
incorrect)) has identified that periotic, collected from the late
Early or early Middle Miocene Nataki Formation, as a rhab-
dosteid dolphin, Eurhinodelphis sp. B. We do not concur
with that generic allocation because the periotic described
by Okazaki does not closely resemble periotics found in skulls
of Eurhinodelphis spp. collected from the Calvert Formation
in Maryland and Virginia (USNM collections). Instead, we
believe the periotic from Japan very closely resembles the
periotic of Delphinodon dividum, and the one from California
that we identify here as aff. Delphinodon dividum, although
it apparently is considerably larger than the latter. We con-
clude that the specimen from the Nataki Formation should

a b

Figure 1 2. aff. Delphinodon dividum T rue, 1912, left periotic, LACM
41041 from LACM locality 6602; a, cerebral or dorsal view; b,
tympanic or ventral view; both natural size.

Contributions in Science, Number 353

Barnes and Mitchell: Kentriodon from California 17

Figure 13. Reconstructions of dorsal views of skulls of Kentriodon Kellogg, 1927; a, Kentriodon pernix Kellogg, 1927, based on the holotype,
USNM 8060, and the referred specimen, USNM 10670; b, Kentriodon obscurus (Kellogg, 1931), based on the referred specimen, LACM
21256, with outline of the rostral extremity and brain case from K. pernix; both at different scales, but reduced to the same brain case length
(antorbital notches to condyles), (a from Kellogg, 1927: pis. 2, 6, and Barnes, 1978: fig. 14, c; b from our Fig. 2.)

18 Contributions in Science, Number 353

Barnes and Mitchell: Kentriodon from California

Figure 14. Reconstructions of ventral views of skulls of Kentriodon Kellogg, 1927; a, Kentriodon pernix Kellogg, 1927, based on the holotype,
USNM 8060, and the referred specimen, USNM 10670, with the dentition omitted on one side so that the size and number of alveoli may
be seen; b, Kentriodon obscurus (Kellogg, 1931), based on the referred specimen LACM 21256, with outline of the rostral extremity and brain
case from K. pernix; both at different scales, but reduced to the same brain case length, (a from Kellogg, 1927: pis. 4, 5, 7 and 8; b from our
Fig. 6.)

Contributions in Science, Number 353

Barnes and Mitchell: Kentriodon from California 19

be re-identified as Kentriodontinae, aff. Delphinodon divi-
dum.

DISCUSSION

RELATIVE ABUNDANCE OF
KENT RIODON OBSCURUS IN THE
SHARKTOOTH HILL BONEBED

A list of 1 5 odontocetes in the Sharktooth Hill Bonebed given
by Barnes ( 1976:327, table 3) included 1 1 named species and
four possibly undescribed species. Grypolithax pavida may
now be deleted from that list because it is a synonym of K.
obscurus. Each of the 14 remaining species is recognized by
periotics. Some of the species in polytypic genera that are
known only by periotics may be questionably valid (Barnes,
1976:327). The abundant sperm whale, Aulophyseter mor-
ricei, with quite distinctive periotics, is known by other skel-
etal elements. The platanistoid “ Squalodon" errabundus also
has distinctive periotics and is known from other skeletal
elements, however, it is relatively rare in the bonebed, and
we (unpublished data) have been able to collect or locate only
31 of the highly unique periotics of that species. Similarly,
the distinctive periotics of the kentriodontid Liolithax ker-
nensis total only 42 in number (Barnes, 1978). Only Platy-
lithax robusta. known solely by the holotype, and the other
four unidentified odontocete species are rarer in samples than
Kentriodon obscurus. The total sample of K. obscurus amounts
to only one skull and 31 periotics, and on this evidence the
species ranks as one of the rarest odontocetes in the Shark-
tooth Hill Bonebed. The remaining hundreds of periotics in
museum collections can be identified as belonging to species
of the genera Loxolithax Kellogg, 1931, Oedolithax Kellogg,
1931, Lamprolithax, or Nannolithax Kellogg, 1931.

ZOOGEOGRAPHY

Fossil Kentriodontidae have been reported from the North
Atlantic, the South and North Pacific, and the Paratethys
regions. All the species included by Barnes (1978:26) in this
family were Middle or Late Miocene in age. Fordyce (1980:
328) has subsequently reported a kentriodontid from Late
Oligocene rocks of New Zealand. The fossils from early Mid-
dle Miocene rocks in California identified as cf. Kentriodon
sp. and aff. Delphinodon dividum in this paper constitute the
earliest records for the family in the North Pacific region.
These, along with the sample we regard as Kentriodon ob-
scurus from the Sharktooth Hill Bonebed, the kentriodontids
from Japan, and Late Miocene species from California
(Barnes, 1976) suggest that a considerable diversity of species
in this family inhabited the North Pacific Ocean during the
Miocene. This diversity equals that known for kentriodontids
during the Middle and Late Miocene in the North Atlantic
and Paratethys regions (see Barnes, 1978:26). The family
might have been cosmopolitan during the Miocene, but there
are as yet no kentriodontids recognized from the South At-
lantic realm or from the eastern South Pacific. It would be
premature to speculate on the place of origin of the family.

even though the oldest known occurrence is in the South
Pacific.

Barnes (1976:338) noted that the then available paleon-
tologic literature suggested a pattern of generic cosmopoli-
tanism among larger fossil late Tertiary cetaceans and one
of generic endemism within major northern ocean basins
among the smaller fossil odontocetes. Among the nine cur-
rently recognized genera of Kentriodontidae (cf. Barnes, 1978:
26), more than one-half (i.e. Liolithax Kellogg, 1931, Ken-
triodon. Delphinodon Leidy, 1 869, Pithanodelphis Abel, 1905,
and Lophocetus Cope, 1868) are now known by species in
both the North Atlantic and the North Pacific regions. For
the Kentriodontidae, it now appears that within ocean basins
a pattern of endemism at the species level rather than the
generic level prevailed.

PHYLOGENETIC RELATIONSHIPS

Kentriodon is the most primitive genus yet assigned to the
subfamily Kentriodontidae. For example, Kentriodon lacks
such derived characters as the elevated cranial vertex and
enlarged nasal bones of Pithanodelphis. and the short ros-
trum, bulbous braincase and rounded facial margins of Del-
phinodon dividum. The cranial morphology of Kentriodon
suggests that its origin might have been among more prim-
itive species in the subfamily Kampholophinae. Kentriodon
shares with the kampholophine species Liolithax pappus and
Kampholophos serrulus such primitive characters as a nar-
row, elongate rostrum, high tooth count, low cranial vertex,
and a concave margin of the facial region above the orbit.
Kentriodon is more derived, however, than either of those
species, by having facial surfaces of the frontal and the max-
illa that are spread more over the dorsal opening of the tem-
poral fossa. The extent of this spreading, however, had not
progressed to the stage seen in the derived kentriodontine
genera Delphinodon, Leptodelphis Kirpichnikov, 1954, Mi-
crophocaena Kudrin and Tatarinov, 1965, Pithanodelphis
and Sarmatodelphis Kirpichnikov, 1954.

There are possibly relationships between Kentriodontinae
and primitive modem Delphinidae, such as species in the
subfamily Steninae ( sensu Mead, 1975), and this has been
discussed previously by True (1912), Kellogg (1927), and
Barnes (1978). Similarities exist in overall cranial propor-
tions, shape of the mandible, and size and numbers of teeth,
but kentriodontines are more primitive by having symmet-
rical cranial vertices, less extensive air sinuses and unfused
cervical vertebrae.

The two essentially synchronous species, Kentriodon per-
nix and K. obscurus. have different combinations of both
primitive and derived characters (see Table 1). For example,
K. pernix is more derived than K. obscurus by having the
premaxillary foramina located more posteriorly (Fig. 1 3), and
the posterior part of the palate less convex on either side of
the pterygoid sinuses. Kentriodon obscurus. on the other hand,
is more derived by having shorter postorbital processes of
the frontals, smaller teeth (Fig. 14), medial premaxillary sur-

20 Contributions in Science, Number 353

Barnes and Mitchell: Kentriodon from California

face in front of the nares not inclined, a longer anterior ex-
tension of the palatine bones on the palatal surface and of
the pterygoid air sinuses within the pterygoid hamuli, and a
larger postorbital lobe of the pterygoid air sinus. The derived
character states listed in Table 1 are equally spread between
the two species. The polarity of these characters was deter-
mined by comparisons with species in the more primitive
odontocete families, Squalodontidae and Agorophiidae.

Kentriodon pernix and K. obscurus might have evolved
from a common ancestor, and the discoveries of the older
fossils of kentriodontines in California certainly indicate that
such Kentriodon- like dolphins lived prior to Middle Miocene
time in the North Pacific Ocean.

CONCLUSIONS

A fossil delphinoid genus, Kentriodon Kellogg, 1927, that is
uncommon in northwest Atlantic Middle Miocene rocks,
occurs in contemporaneous rocks on the eastern margin of
the North Pacific Ocean. A few fossils from the North Pacific
margin identified as this genus or related genera of small
dolphins have been briefly cited in previous literature, and
it is shown here that the genus Grypolithax Kellogg, 1931,
originally based on specimens from the Middle Miocene
Sharktooth Hill Bonebed in California, is synonymous with
Kentriodon. Based on isolated fossil periotics, Kellogg (1931)
had named two species in Grypolithax, G. obscura Kellogg,
1931, and G. pavida Kellogg, 1931. These are the only species
that have ever been assigned to Grypolithax and we regard
them as synonymous. G. obscura has page priority, is the
senior synonym of G. pavida, and is the type species of the
genus. Because Grypolithax is not valid, the most appropriate
binomen for the species from the Sharktooth Hill Bonebed
is Kentriodon obscurus (Kellogg, 1931). We refer periotics
and a partial skull to this species. The only other named
species of Kentriodon is the type species, K. pernix Kellogg,
1927, from the Calvert Formation in Maryland.

Among possibly as many as 1 4 species of odontocetes found
in the Sharktooth Hill Bonebed, K. obscurus is not abundant
and is one of the rarest species. Only one of the rarer ones,
Platylithax robusta Kellogg, 1931, has been named previ-
ously in the scientific literature. The other four have not been
named, although Barnes (1976) has called attention to their
presence. Only by increasing the total sample size will we be
able to eventually recognize the rarer species in the Shark-
tooth Hill Local Fauna and learn their anatomy.

Middle Miocene fossils of Kentriodon have now been re-
ported from Maryland and California in the U.S.A., and
apparently also from Japan. The earliest reported kentri-
odontine fossils from the eastern North Pacific region are
two isolated periotics we have identified as cf. Kentriodon
sp. from the early Middle Miocene part of the Round Moun-
tain Silt, lower stratigraphically than the Sharktooth Hill
Bonebed. Kentriodon may have been a cosmopolitan genus
in the Middle Miocene, but there is yet no published fossil
evidence from the southern hemisphere to prove this.

Each of the two known species of Kentriodon has a different

suite of primitive and derived characters, and the two are
probably derived from a common ancestor.

ACKNOWLEDGMENTS

This study was aided by the granting of travel funds to L.G.
Barnes by the Natural History Museum of Los Angeles
County (LACM) Foundation and the Smithsonian Institu-
tion. Some of the artwork was sponsored by the University
of California Museum of Paleontology (UCMP). Curation of
some specimens was aided by National Science Foundation
(NSF) Grant DEB 7202014 to the LACM Foundation for
care of collections. Early fieldwork by E.D. Mitchell was
supported by LACM. Some of the later fieldwork was sup-
ported by NSF Grant EAR 7916508 to the LACM Foun-
dation to facilitate paleoecologic investigations of the Shark-
tooth Hill Bonebed.

We are grateful to the following persons for collecting var-
ious specimens used in this study: Mr. and Mrs. E.L. An-
derson, Shelton P. Applegate, Angela Bennett, Richard C.
Bishop, Dennis L. Clayton, the late John E. Fitch, William
Hawes, Jr., Richard W. Huddleston, Paul Kirkland, Douglas
J. Macdonald, the late Charles Morrice, Michael D. Quarles,
Camm C. Swift, Terry Story, and Howell W. Thomas.

We thank Robert L. Brownell, Jr., R. Ewan Fordyce, Sam-
uel A. McLeod, James G. Mead, David P. Whistler, Frank
C. Whitmore, Jr., and anonymous reviewers for constructive
criticism of the manuscript.

The following persons allowed us to study specimens in
their care: Clayton E. Ray, Frank C. Whitmore, Jr., and
Robert Purdy (USNM), Victor A. Zullo and Peter Rodda
(CAS), and J. Howard Hutchison (UCMP).

We were kindly granted access to collecting sites in Kern
County by Mr. and Mrs. Clint Smoot, Mr. and Mrs. Steven
Smoot, Vincent Bertolucci, the late Bernard Terabino, and
E.H. Shuler and other employees of Getty Oil Company.

David Cook, formerly of UCMP, prepared some of the
line drawings (Fig. 1 1). The photographs (Figs. 2, 4, 6, 8, 9)
were prepared by Lawrence Reynolds, formerly of LACM.
John De Leon of LACM did some of the photographic copy-
ing of the artwork. Other illustrations were prepared by L.G.
Barnes.

LITERATURE CITED

Abel, O. 1905. Les odontocetes du Bolderien (Miocene
superieur) d’Anvers. Memoires du Musee Royal d’His-
toire Naturelle de Belgique 3(2): 1-1 55.

Addicott, W.O. 1972. Provincial middle and late Tertiary
molluscan stages. Temblor Range, California. Pages 1-
26 in E.H. Stinemeyer, ed.. Proceedings of the Pacific
Coast Miocene Biostratigraphic Symposium, Society of
Economic Paleontologists and Mineralogists. 364 pp.
Barnes, L.G. 1976. Outline of eastern North Pacific fossil
cetacean assemblages. Systematic Zoology 25(4):32 1 —
343.

Contributions in Science, Number 353

Barnes and Mitchell: Kentriodon from California 21

. 1978. A review of Lophocetus and Liolithax and

their relationships to the delphinoid family Kentriodon-
tidae (Cetacea: Odontoceti). Natural History Museum
of Los Angeles County Science Bulletin 28:1-35.

Berggren, W., and J.A. Van Couvering. 1974. The late Neo-
gene. Biostratigraphy, geochronology and paleoclima-
tology of the last 15 million years in marine and con-
tinental sequences. Palaeogeography, Palaeoclimatology
and Palaeoecology 16(1 /2):i— xi, 1-216.

Boenninghaus, G. 1904. Das Ohr des Zahnwales, zugleich
ein Beitrag zur Theorie der Schalleitung. Eine biolo-
gische Studie. Zoologische Jahrbiicher. Abteilung fur
Anatomie und Ontogenie der Tiere 1 9: 1 89-360, pis. 1 2-
13.

Clark, W.B., G.B. Shattuck, and W.H. Dali. 1904. The
Miocene Deposits of Maryland. Pages xxi-xxxii in W.B.
Clark, ed., Maryland Geological Survey— Miocene, Bal-
timore, The Johns Hopkins Press.

Cope, E.D. 1868a (for 1867). (Descriptions of Eschrichtius
cephalus, Rhabdosteus latiradix, Squalodon atlanticus
and S. mento.) Proceedings of the Academy of Natural
Sciences, Philadelphia 19:131-132.

. 1868b (for 1867). An addition to the vertebrate

fauna of the Miocene period, with a synopsis of the
extinct Cetacea of the United States. Proceedings of the
Academy of Natural Sciences, Philadelphia 19: 1 38-156.

Denker, A. 1902. Zur Anatomie des Gehororgans der Ce-
tacea. Anatomische Hefte 19(2):423-448, pis. 14-15.

Evemden, J.F., D.E. Savage, G.H. Curtis, and G.T. James.
1964. Potassium-argon dates and the Cenozoic mam-
malian chronology of North America. American Journal
of Science 262:145-198.

Fordyce, R.E. 1980. Whale evolution and Oligocene south-
ern ocean environments. Palaeogeography, Palaeocli-
matology and Palaeoecology 31:319-336.

. 1981. Systematics of the odontocete whale Ago-

rophius pygmaeus and the family Agorophiidae (Mam-
malia: Cetacea). Journal of Paleontology 5 5(5): 1 028—
1045.

Fraser, F.C., and P.E. Purves. 1960. Hearing in cetaceans.
Evolution of the accessory air sacs and the structure and
function of the outer and middle ear in Recent cetaceans.
Bulletin of the British Museum (Natural History), Zo-
ology 7(1): 1-140, frontispiece, pis. 1-53.

Gazin, C.L., and R.L. Collins. 1950. Remains of land mam-
mals from the Miocene of the Chesapeake Bay region.
Smithsonian Miscellaneous Collections 1 16(2): 1-21.

Hay, O.P. 1902. Bibliography and catalogue of the fossil
Vertebrata of North America. Bulletin of the United
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Kasuya, T. 1973. Systematic consideration of Recent
toothed whales based on the morphology of tympano-
periotic bone. Scientific Reports of the Whales Research
Institute 25:1-103, pis. 1-28.

Kellogg, A.R. 1925. The relationships of the Tertiary ce-
taceans of Jugo-Slavia to those of eastern North Amer-
ica. Spomenice U Pocast Prof. Dr. Gorjanovic-Kram-
bergera, Zagreb, pp. 1-8.

. 1927a. Kentriodon pernix, a Miocene porpoise from

Maryland. Proceedings of the United States National
Museum 69( 1 9): 1 —55, pis. 1-14.

. 1927b. Study of the skull of a fossil sperm-whale

from the Temblor Miocene of Southern California. Con-
tributions to Paleontology, Carnegie Institution of
Washington Publications 346:1-24, pis. 1-9.

. 1928. The history of whales— their adaptation to

life in the water. Quarterly Review of Biology 3(1, 2):
29-76, 174-208.

. 1931. Pelagic mammals from the Temblor For-
mation of the Kern River region, California. Proceedings
of the California Academy of Sciences, Ser. 4, 19(12):
217-397.

. 1955. Three Miocene porpoises from the Calvert

Cliffs, Maryland. Proceedings of the United States Na-
tional Museum 105:101-154, pis. 1-21.

Kellogg, A.R., and F.C. Whitmore, Jr. 1957. Mammals.
Pages 1021-1024 in H.S. Ladd, ed.. Treatise on marine
ecology and paleoecology. Volume 2, Paleoecology.
Geological Society of America Memoir 67:i-x, 1-1077.

Kirpichnikov, A. A. 1954. Dva novykh roda del’finov iz
sarmata SSSR. (Two new genera of dolphins from Sar-
matian of U.S.S.R.) Trudy Paleontologicheskogo Insti-
tuta, Akademia Nauk S.S.S.R. 47:181-190, pis. 1-3.

Kudrin, L.N., and K. A. Tatarinov. 1965. O miotzenovykh
del’finakh zapadnoi Ukrainy. (On the Miocene delphin-
ids from the western Ukraine.) Paleontologicheskii
Zhumal 1 965(4):68-74.

Lawrence, B., and W.E. Schevill. 1956. The functional
anatomy of the delphinid nose. Bulletin of the Museum
of Comparative Zoology, Harvard College 1 1 4(4): 1 03—
151.

Leidy, J. 1869. The extinct mammalian fauna of Dakota
and Nebraska, including an account of some allied forms
from other localities, together with a synopsis of the
mammalian remains of North America. Journal of the
Academy of Natural Sciences of Philadelphia, new series
7:1-472.

Mead, J.G. 1975. Anatomy of the external nasal passages
and facial complex in the Delphinidae (Mammalia: Ce-
tacea). Smithsonian Contributions in Zoology 207:i~iv,
1-72.

Miller, G.S., Jr. 1923. The telescoping of the cetacean skull.
Smithsonian Miscellaneous Collections 76(5): 1-71.

Mitchell, E. 1965. History of research at Sharktooth Hill,
Kern County, California. Kern County Historical So-
ciety (Bakersfield, California), Special Publication, vi +
45 pp.

. 1966. The Miocene pinniped Allodesmus. Univer-
sity of California Publications in Geological Sciences 6 1 :
1-46, pis. 1-29.

, and R.H. Tedford. 1973. The Enaliarctinae, a new

group of extinct aquatic Carnivora and a consideration
of the origin of the Otariidae. Bulletin of the American
Museum of Natural History 151(3):201 —284.

Okazaki, Y. 1976. Miocene long-snouted porpoises from

22 Contributions in Science, Number 353

Barnes and Mitchell: Kentriodon from California

the Mizunami Group, central Japan. Bulletin of the Mi-
zunami Fossil Museum 3:25-40, pis. 9-1 1.

Ray, C.E. 1976. Geography of phocid evolution. Systematic
Zoology 25(4):39 1-406.

Rensberger, J.M. 1 969. A new iniid cetacean from the Mio-
cene of California. University of California Publications
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Repenning, C.A., and R.H. Tedford. 1977. Otarioid seals
of the Neogene. United States Geological Survey Profes-
sional Paper 992:i-vi, 1-93, pis. 1-24.

Savage, D.E., and L.G. Barnes. 1972. Miocene vertebrate
geochronology of the west coast of North America. Pages
124-145 in E.H. Stinemeyer, ed., Proceedings of the
Pacific Coast Miocene Biostratigraphic Symposium, So-
ciety of Economic Paleontologists and Mineralogists. 364
PP-

Shikama, T., Y. Hasegawa, and H. Otsuka. 1973. Geolog-
ical range of mammals in the Japanese Neogene. Mem-
oirs of the Geological Society of Japan 8:137-141 (in
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Simpson, G.G., A. Roe, and R.C. Lewontin. 1960. Quan-
titative Zoology. Revised Edition. Harcourt, Brace and
World, Inc., New York, pp. i-vii, 1-440.

Slijper, E.J. 1936. Die Cetaceen. Vergleichend-anatomisch
und systematisch. Capita Zoologica 6:i-xv, 1-590.

. 1958. Walvissen. D.B. Centen’s Uitgeversmaat-

schappij, Amsterdam. 524 pp.

True, F.W. 1912. Description of a new fossil porpoise of
the genus Delphinodon from the Miocene Formation of
Maryland. Journal of the Academy of Natural Sciences
of Philadelphia, Ser. 2, 15:165-194, pis. 17-26.

Weaver, C.E. et al. 1944. Correlation of the marine Ce-
nozoic formations of western North America. Geolog-
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the North American continental Tertiary. Geological So-
ciety of America Bulletin 52:1-48, pi. 1.

Accepted for publication 4 April 1984.

Contributions in Science, Number 353

Barnes and Mitcheil: Kentriodon from California 23

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SYSTEMATICS OF FISSURELLA IN THE PERUVIAN
AND MAGELLANIC FAUNAL PROVINCES
(GASTROPODA: PROSOBRANCHIA)

James H. McLean

Contributions in Science, Number 354
Natural History Museum of Los Angeles County
29 October 1984

ISSN 0459-8113

Natural History Museum of Los Angeles County
900 Exposition Boulevard
Los Angeles, California 90007

CONTENTS

ABSTRACT 1

RESUMEN 1

INTRODUCTION 2

MATERIALS AND METHODS 3

Fieldwork 3

Other Collections Examined 3

Preparation of Specimens 3

Conventions 3

Abbreviations 3

HISTORICAL REVIEW 4

STRUCTURE 5

Internal Anatomy 5

External Anatomy 6

Radula 6

Shell Morphology 8

Shell Structure 8

BIOLOGY AND ECOLOGY 8

Habitat 8

Feeding 8

Reproduction and Growth 9

Epibiotic Associations 9

Epibiotic Scurria parasitica 9

Shell Borers 9

Parasites 1 1

Seastar Predators 11

Vertebrate Predators 1 1

Human Predation and Economic Importance 11

SYSTEMATIC CHARACTERS 12

Size and Shape 12

Sculpture 1 2

Color Pattern 1 2

Interior Margin 1 2

Foramen 1 3

Juvenile Shell 13

External Anatomy 13

Radula 13

NAMES, ALLOCATIONS, AND KEY CHARACTERS 13

Names and Allocations 13

Chart of Key Characters 14

DISTRIBUTION AND ZOOGEOGRAPHY 15

FOSSIL RECORD 15

FORMAT FOR SPECIES ACCOUNTS 16

SYSTEMATICS 17

Family Fissurellidae Fleming, 1822 17

Subfamily Fissurellinae Fleming, 1822 18

Genus Fissurella Bruguiere, 1789 18

Subgenus Fissurella Bruguiere. 1789 19

Group of F. peruviana 19

Fissurella peruviana Lamarck, 1822 21

Group of F. maxima 24

Fissurella maxima Sowerby, 1835 25

Fissurella latimarginata Sowerby, 1835 28

Fissurella cumingi Reeve, 1849 31

Fissurella costata Lesson, 1831 34

Fissurella picta (Gmelin, 1791) 37

Fissurella radiosa Lesson, 1831 43

Fissurella oriens Sowerby, 1835 49

Fissurella nigra Lesson, 1831 52

Group of F. limbata 55

Fissurella limbata Sowerby, 1835 55

Fissurella crassa Lamarck, 1822 58

Fissurella bridgesii Reeve, 1 849 60

Fissurella pulchra Sowerby, 1835 63

LACM LOCALITIES FOR FIGURED SPECIMENS 65

ACKNOWLEDGMENTS 66

LITERATURE CITED 67

SYSTEMATICS OF FISSURELLA IN THE PERUVIAN AND
MAGELLANIC FAUNAL PROVINCES
(GASTROPODA: PROSOBRANCHIA)

James H. McLean1

ABSTRACT. Fifty-eight names have been proposed for the large
and abundant species of Fissurella in the Peruvian and Magellanic
faunal provinces. Fieldwork in Peru and Chile and to a lesser extent
in Argentina, has produced large collections. Following study of these
collections, as well as most of the type specimens, I reduce the
number of species to 1 3, three of which have geographic subspecies.

Peruvian-Magellanic species of Fissurella Bruguiere, 1789, are
members of the nominate subgenus, in which the shell has an inner
layer of crossed lamellar aragonite and a thick outer layer of prismatic
calcite. In contrast, most tropical species of Fissurella, which are in
the subgenus Cremides H. and A. Adams, 1854, have a shell com-
posed entirely of aragonite. The outer layer of calcite is evidently an
adaptation to cold water in the Peruvian-Magellanic species.

Three species groups in Fissurella ( sensu stricto) are recognized.
The group of Fissurella peruviana Lamarck, 1822, is smaller-shelled
and has a thinner calcitic layer than species in the other groups; this
group includes the type species F. nimbosa Linnaeus, 1758, in the
southern Caribbean (the only tropical member of the subgenus) and
F. volcano Reeve, 1849, in California and Baja California.

The group of F. maxima Sowerby, 1 833, is characterized by strong
primary and secondary ribs (at least in juvenile stages) and also
includes F. latimarginata Sowerby, 1835, F. cumingi Reeve, 1849,
and F. costata Lesson, 1831, in the Peruvian Province, and F. picla
(Gmelin, 1791), F. radiosa Lesson, 1831, F. oriens Sowerby, 1835,
and F. nigra Lesson, 1831, in the Magellanic Province. The group
of F. limbata Sowerby, 1835, is characterized by broad primary nbs
and lack of secondary ribs; it includes F. crassa Lamarck, 1 822, F.
bridgesii Reeve, 1849, and F. pulchra Sowerby, 1835, all in the
Peruvian Province.

Geographic subspecies are here recognized for three species broad-
ly distributed in the Magellanic Province: F. picta picta (Gmelin,
1791) in southern Chile, and F. picta lata Sowerby, 1835, in central
Chile; F. radiosa radiosa Lesson, 1831, in southern Chile, F. radiosa
tixierae Metivier, 1969, in the vicinity of the Gulf of San Matias in
Argentina; F. oriens oriens Sowerby, 1835, in southern Chile, and
F. oriens fulvescens Sowerby, 1835, in central Chile.

Three species, F. cumingi, F. bridgesii, and F. pulchra, have been
poorly understood by previous authors and are newly defined here.

Distributions of the Peruvian and Magellanic species overlap in
south-central Chile, where 1 2 of the 1 3 species occur.

This account includes observations on shell epibionts and borers,
and reviews the sparse literature on the biology of these species.

Contributions in Science, Number 354, pp. 1-70
Natural History Museum of Los Angeles County, 1984

RESUMEN. Hasta ahora habian sido propuestos 58 nombres para
las grandes y abundantes especies de Fisssurella de las provincias
biogeograficas Peruana y Magallanica. Trabajos de terrene en Peru
y Chile, y en menor intensidad en Argentina, han proporcionado
importantes colecciones. Realizado el estudio de estas colecciones y
de muchos de los ejemplares tipo, se reduce a 13 el numero de
especies, tres de las cuales poseen subespecies geograficas.

Las especies de Fissurella de las provincias Peruana y Magallanica
son integrantes del subgenero Fissurella Bruguiere, 1789 {sensu stric-
to), en las cuales la concha tiene una capa interna compuesta de
aragonita laminar cruzada y una capa mas externa de calcita pris-
matica. En cambio, muchas especies de Fissurella que pertenecen
al subgenero Cremides H. y A. Adams, 1854, tienen la concha com-
puesta integramente de aragonita. La capa mas externa de calcita es
considerada como una adaptacion de las especies de las provincias
Peruana y Magallanica a aguas frias.

Se reconocen tres grupos de especies. Un primer grupo de Fis-
surella peruviana Lamarck, 1822, de concha mas pequena y cuya
capa de calcita es mas delgada que en las otras especies de los demas
grupos. Este grupo comprende la especies tipo F. nimbosa Linnaeus,
1758, del sur del Caribe (unico miembro tropical del subgenero) y
F. volcano Reeve, 1849, de California y Baja California.

Un segundo grupo de F. maxima Sowerby, 1833, caracterizado
por la presencia de gruesas costillas primarias y secundarias (al me-
nos en los estados juveniles). Comprende tambien a F. latimargi-
nata Sowerby, 1835, F. cumingi Reeve, 1849, y F. costata Lesson,
1 83 1 , de la provincia Peruana y a F. picta (Gmelin, 1 79 1 ), F. radiosa
Lesson, 1831, F. oriens Sowerby, 1835, y F. nigra Lesson, 1831, de
la provincia Magallanica.

El tercer grupo de F. limbata Sowerby, 1835, esta caracterizado
por la presencia de costillas primanas anchas y ausencia de costillas
secundarias. Comprende tambien a F. crassa Lamarck, 1822, F.
bridgesii Reeve, 1849, y F. pulchra Sowerby, 1835, todas de la pro-
vincia Peruana.

Se reconocen las siguientes subespecies geograficas para tres es-
pecies ampliamente distribuidas en la provincia Magallanica: F.
picta picta (Gmelin, 1791) del sur de Chile y F. picta lata Sowerby,
1835, de la zona central de Chile; F. radiosa radiosa Lesson, 1831,

1. Malacology Section, Natural History Museum of Los Angeles
County, 900 Exposition Blvd., Los Angeles, California 90007.

ISSN 0459-81 13

Figure 1. A Fissurella “shell pile,” consisting of large specimens of F. latimarginata, F. cumingi. and F. maxima at Los Molles, Aconcagua
Province, Chile, October 15, 1975. The specimens had presumably been taken by shallow diving in the vicinity and the shells discarded. All
shells were covered with the algal mat, which completely obscures the color pattern.

del sur de Chile y F. radiosa tixierae Metivier, 1 969, de las cercanias
del golfo de San Matias en Argentina; F. oriens oriens Sowerby,
1835, del sur de Chile y F. oriens fulvescens Sowerby, 1 835, de Chile
central.

Las especies F. cumingi, F. bridgesii y F. pulchra, que han sido
escasamente tratadas por autores anteriores, son definidas nueva-
mente.

Se observa una sobreposicion en la distribucion de las especies de
las provincias Peruana y Magallanica en la zona central-sur de Chile,
donde 1 2 de las 13 especies se encuentran presentes.

En el presente trabajo se incluye, ademas, observaciones sobre los
epibiontes y organismos perforadores de las conchas y se revisa la
esparcida literatura que trata sobre la biologia de algunas de estas
especies.

INTRODUCTION

The Fissurella species of the cool waters of Peru, Chile, and
southern Argentina are large and abundant, comprising a
major element of the mollusk fauna of the west coast of South
America. They are extensively used for food and are known
locally as “lapas” (Fig. 1). The importance of the fishery is
second only to that of Concholepas, the “loco,” the large
limpetlike thaidid gastropod of the region. Despite this im-
portance, the taxonomy of the South American species of
Fissurella has been poorly understood.

It has been recognized that a large number of highly vari-
able, sympatric species occur in the region. Some 58 names

for Recent species have been introduced in the literature.
Widely varying estimates of the number of actual species
have been given: Pilsbry (1890) recognized about 20 species;
Ziegenhom and Thiem (1925) treated 1 1 species and three
“varieties”; Riveros-Zuniga (1951) recognized 26 species and
three “varieties”; Dell (1971) listed 1 1 possible species; and
finally Ramirez-Boehme (1974) gave a key to 30 species and
two “varieties.”

The collection of mollusks from Iquique, Chile, reported
upon by Marincovich (1973), and deposited in the Natural
History Museum of Los Angeles County, included five species
of Fissurella. Although this material introduced me to the
subject, fieldwork of my own in Peru in 1972 and 1974, and
in Chile for two months in 1975, and southern Argentina in
1978, enabled me to collect and observe the Fissurella species
from many different localities. I have therefore been able to
observe these mollusks throughout their entire geographic
range from north-central Peru to southern Chile and Argen-
tina.

The Magellanic Province of southern Chile and southern
Argentina is also the center of distribution of another fissu-
rellid group comprising the species Fissurellidea megatrema
Orbigny, 1841, F. patagonica (Strebel, 1907), Pupillaea an-
nulus (Odhner, 1932), and the shell-less Buchanania onchi-
dioides Lesson, 1830. A report on these species has been
published (McLean, 1984b).

2 Contributions in Science, Number 354

McLean; Peruvian and Magellanic Fissurella

In this work I offer a revised classification of the South
American species of Fissurella, based on my field observa-
tions, study of the large collection now housed at the Los
Angeles County Museum of Natural History, and study of
type material borrowed from other museums. The classifi-
cation cannot be exhaustive and does not offer a cladistic
hypothesis of relationships. In the absence of anatomical,
biochemical (electrophoretic), and other characters, that is
beyond the scope of the present work. It is hoped that this
paper will provide a basis for future work on the systematics
and ecology of these species.

MATERIALS AND METHODS

Fieldwork

The collection upon which this report is chiefly based is now
in the Los Angeles County Museum of Natural History. Some
material from miscellaneous sources is represented in the
collection, but the bulk of it resulted from nine major ex-
peditions as follows:

1. Peru: Isla San Lorenzo, Isla Chinchas, and Bahia Inde-
pendencia. Allan Hancock Expeditions, January, 1935,
and February, 1938, intertidal and dredging stations.

2. Chile: Iquique. Louie Marincovich, June-September,
1964, June-July, 1970, intertidal stations.

3. Argentina: Isla de los Estados (E of Tierra del Fuego). R/V
HERO, April, 1971, and October, 1971, intertidal and
dredging stations; collections received from the Smith-
sonian Oceanographic Sorting Center.

4. Peru: Pucasana, Laguna Grande, Isla Chincha Norte, Par-
acas, Asia. James H. McLean, April, 1972, intertidal and
diving stations.

5. Chile: south of Isla de Chiloe. Paul Dayton, on R/V HERO,
October-November, 1972, intertidal and diving stations.

6. Chile and Argentina: Strait of Magellan and Isla de los
Estados. Paul Dayton, on R/V HERO, May, 1973, inter-
tidal and diving stations.

7. Peru: Isla Guanape, Ancon, and Isla San Lorenzo. James
H. McLean, January, 1974, intertidal and diving stations.

8. Chile: Iquique, Antofagasta, Coquimbo, Los Molles,
Montemar, Cartagena, Concepcion, Mehuin, Pargua,
Guabun, Pumalin, Islota Nihuel, Isla Laitec, Puerto
Hambre, Punta Arenas. James H. McLean, October-No-
vember, 1975, intertidal and diving stations.

9. Argentina: Golfo Nuevo and Golfo San Jose. James H.
McLean, on R/V HERO, July 1978, intertidal and dredg-
ing stations.

Other Collections Examined

Upon returning from Chile in 1975, I compared the field-
collected specimens with as many of the types of species
described by nineteenth-century authors as could be located
for me in the British Museum (Natural History) and the Paris
Museum. I have also studied the collections of Fissurella in
the U.S. National Museum of Natural History, Washington,
D.C., the Academy of Natural Sciences, Philadephia, and the
American Museum of Natural History, New York. Other

specimens were received on loan from the Museum of Com-
parative Zoology, Harvard, and the National Museum of
New Zealand, Wellington. After conducting my fieldwork in
Argentina in 1978, I examined the Fissurella material in the
Museo Argentino de Ciencias Naturales, Buenos Aires, and
the Museo Nacional de Historia Natural, Santiago. Records
from these collections enabled further refinements in species
distributions.

Preparation of Specimens

Although most of the field-collected specimens were either
kept dry or the entire specimen preserved in alcohol, without
cleaning the shell, the photographed specimens had to be
cleaned of encrusting organisms. Shells were placed in full
strength laundry bleach, which softens the algal mat and
loosens other encrusting organisms so that the shell can be
scraped clean with a knife or wire brush. Color was restored
with a light application of mineral oil.

Shells of each species were embedded in plaster and cut
with a diamond rock saw for the examination and photog-
raphy of the shell layers. Scanning electron microscopy (SEM)
was used for the examination of shell structure in a fragment
of a small specimen of F. latimarginata. Radulae of large
specimens of each species were air-dried for macrophotog-
raphy; radulae of small specimens were prepared for both
light microscopy and SEM.

Conventions

Figured specimens for each species are arranged by localities
from north to south, including type specimens of nominate
taxa and synonyms. All shell specimens are illustrated with
the anterior at the top; lateral views are those of the left side
of the shell. Measurements for the figured specimens are
given in the captions, not repeated in the text. Measurements
are given in this order: length, width, and height. Unless
otherwise indicated, the figured specimens were collected in
the intertidal zone. Latitude and longitude for the figured
specimens from LACM stations are given in a locality list
following the systematic section.

Abbreviations

Abreviations of institutions mentioned in the text are as
follows:

AHF Allan Hancock Foundation Collection (at LACM)
AMNH American Museum of Natural History, New York

ANSP Academy of Natural Sciences, Philadelphia
BMNH British Museum (Natural History), London
LACM Los Angeles County Museum of Natural History,
Los Angeles

MACN Museo Argentino de Ciencias Naturales, Buenos
Aires

MCZ Museum of Comparative Zoology, Harvard Uni-

versity, Cambridge

MNHN Museo Nacional de Historia Natural, Santiago
MNHNP Museum National d’Histoire Naturelle, Paris
NMNZ National Museum of New Zealand, Wellington

Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella 3

USNM United States National Museum of Natural His-
tory, Washington, D.C.

HISTORICAL REVIEW

The Strait of Magellan was probably the origin of the earliest
collected shells of Fissurella to reach Europe. The first species
to be known was the common Magellanic F. picta, which
was well described and figured in the non-binomial work of
Davila (1767), who called it “Un Lepas rare de Magellan”
(Pilsbry, 1890:114). The name Fissurella picta dates from
J . F. Gmelin (1791), whose knowledge of it came in part from
Davila.

J.B.P.A. de Lamarck (1822) validated F. crassa and F.
peruviana in his “Histoire naturelle des animaux sans ver-
tebres.” Lamarck’s types are preserved at the Geneva Mu-
seum (Mermod, 1950).

G.B. Sowerby (1825) introduced a synonym for F. crassa,
but was later to recognize Lamarck’s unfigured species.

G.P. Deshayes (1830) named two Chilean species in the
“Encyclopedic Methodique,” but one of them was a synonym
of F. peruviana and the other, F. rudis, although prior to F.
cost at a Lesson, 1831, is preoccupied. Types are extant at the
Paris Museum.

R.P. Lesson (1831), in his report on collections from the
South American voyage of the “Coquille,” described four
species of Fissurella without illustrations, all from the south-
ern and southernmost regions of Chile. The names for three
of his species are now in use: F. nigra, F. radiosa and F.
costata. One other, F. obovalis, remains a nomen dubium.
Lesson’s work has frequently been cited as published in 1830,
but the pages that included the Fissurella descriptions are
correctly dated 1831. Although some of Lesson’s types have
been recognized at the Paris Museum, P. Bouchet reports
(personal communication) that he has been unable to locate
any of the Fissurella types.

By 1831, six of the currently recognized species had been
established. Eleven names had then been proposed, but very
few of these taxa had been illustrated.

In 1835, no less than 13 names were introduced by G.B.
Sowerby, based on specimens received from the British col-
lector Hugh Cuming, who had lived in Valparaiso, Chile,
from 1 8 1 9 to 1831. Brief descriptions were given in the Pro-
ceedings of the Zoological Society of London for 1 834. Dating
for these species has frequently been cited as 1834, but the
publication date for the pages involved is 1835. Illustrations
were published simultaneously in the “Conchological Illus-
trations” (Sowerby, 1 835b). Sowerby recognized some of the
species described earlier by French authors; seven of his names
remain useful: F. maxima, F. latimarginata, F. limbata, F.
oriens, F. pulchra, F. lata (here F. picta lata), and F. fulves-
cens (here F. oriens fulvescens). The Sowerby types are pre-
served at the British Museum (Natural History).

By 1835, 1 1 of the 13 species I recognize from Chile had
been named, and the total number of described taxa had
reached 25.

R.A. Philippi (1845, 1845-46) proposed five names, but

none of these has any current utility. One of these, F. alba,
has been used by some authors, but is here regarded as a
synonym of F. oriens. Unfortunately, the present where-
abouts of Philippi’s Fissurella types is unknown.

A. A. Gould (1846) introduced one name, here regarded as
a synonym for F. peruviana. The holotype is in the United
States National Museum.

Lovell Reeve (1849-50), in his monograph of Fissurella
in the Conchologica Iconica, added two more of the species
recognized here, F. cumingi and F. bridgesii, both of which
have been enigmatic until now. However, he also introduced
nine superfluous names, based on further splitting of Cum-
ing’s material. Some were described without locality. Reeve
gave colored illustrations for all the previously recognized
species but did not give any interior views of the shells,
thereby not treating the broad margin, one of the most useful
characters. Also, he did not always figure the same specimen
illustrated by Sowerby, a factor contributing to confusion in
some cases. The Reeve types are housed in the British Mu-
seum (Natural History).

An attempt at summarizing the recognizable species in
Chile was made by L.H. Hupe, 1854, who added Spanish
translations of original descriptions of earlier species, and
included some of his own commentary, but gave no illustra-
tions. Twenty-one species were recognized. One new taxon
was introduced, the renaming of a preoccuppied name of
Philippi.

Philippi (1857) proposed another name now having no
value. In 1860 Philippi briefly treated eight species of Fis-
surella from Paposa (near Antofagasta) in his “Reise durch
die Wueste Atacama . . . .”

G. B. Sowerby II’s treatment of Fissurella in the “Thesau-
rus Conchyliorum” ( 1 862) was scarcely an advance over that
of Reeve. Specimens illustrated were not always those of
Sowerby or Reeve. One additional synonym was named.
Little new information was given, nor was opportunity taken
to reduce the number of names. The figures were smaller
than those given by Reeve, and there were no interior views.

A.T. de Rochebrune and J. Mabille ( 1 885) proposed three
taxa from the southernmost region, none of which were com-
pared to established species; the names are now regarded as
junior synonyms. Two of the three type specimens are housed
in the Paris Museum; the whereabouts of the other is un-
known.

H. A. Pilsbry’s (1890) treatment of Fissurella in the “Man-
ual of Conchology” was an admirable effort at summary and
review. For most taxa he provided English translations of
text by German and French authors and copied original il-
lustrations for all taxa, whether recognized as valid or placed
in synonymy. Synonyms were allocated as far as possible.
Approximately 20 were treated as good species. However,
the specimens available to Pilsbry were limited, and many
questions remained unanswered. Only one unnecessary new
species was introduced, the holotype of which is preserved
at the Academy of Natural Sciences, Philadelphia.

J.C. Melvill and R. Standen (1898, 1907, 1914) listed and
gave notes on mollusks from the Falkland Islands. The Fis-
surella species were briefly treated.

4 Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella

H. Strebel (1907) treated the Magellanic and Patagonian
Fissurel/a. For three of the species that I consider to inhabit
this southern area, he recognized six, but introduced no syn-
onyms. A year later, Strebel ( 1 908) listed F. exquisita from
Paulet Island, Antarctic Peninsula, a record that needs further
verification. Unfortunately, the Strebel Collection was de-
stroyed in World War II (Dance, 1966:302).

W.H. Dali (1909) listed 18 species of Fissurella in his
checklist of mollusks from the Peruvian faunal province.
Those pertaining to the Magellanic area were omitted. Some
of Dali’s records are now clearly erroneous: F. crassa at the
Galapagos Islands and both F. maxima and F. picta at Man-
ta, Ecuador. The records from Manta had previously been
cited by Steams (1891).

A. Ziegenhom and H. Thiem (1925) reported upon a col-
lection made in Chile by L.H. Plate. From a collection of
only 15 specimens, they discussed and illustrated 1 1 species.
They omitted three species that I recognize and treated two
others as “varieties,” but their scheme is the best effort avail-
able in the literature. Some external features of the animal
were mentioned, and good illustrations of the shells were
given, but no interior views. Among the subsequent authors,
only Odhner (1932) and Dell (1971) cited their work.

I. Perez-Farfante (1943) mentioned only F. picta in her
account of Atlantic Fissurellidae. She made this species the
type of her new subgenus, Balboaina, which I here synony-
mize with Fissurella, sensu stricto.

The “Catalogo descriptivo de fisurelidos Chilenos” of F.
Riveros-Zuniga (1951) was compiled almost entirely from
the literature. Text from previous authors was translated into
Spanish. Illustrations were single exterior views, most of which
were copied from other sources. Twenty-six species and three
varieties were recognized. Three of the common Peruvian
Province species were erroneously cited from Fuerte Bulnes,
near Punta Arenas in the Strait of Magellan.

A. R. Carcelles (1950), Carcelles and S.I. Williamson (195 1),
and Carcelles (1953), produced a series of faunal checklists
for the Patagonian, Magellanic, and Antarctic regions, in
which Fissurella species were listed.

Perez-Farfante (1952) proposed the subgenus Carcellesia,
with the new type species F. doellojuradoi, which I regard as
a synonym of F. oriens. The subgeneric name is here regarded
as a synonym of Fissurella, sensu stricto.

B. Metivier ( 1 969) named F. tixierae from the Golfo Nue-
vo, Argentina, a name here treated as a geographic subspecies
of F. radiosa Lesson. The type specimen is preserved at the
Paris Museum.

G.M. Pena (1970) included six species (five that I recog-
nize) in his list of the intertidal mollusks of Peru, and cited
a number of his own collecting localities for each.

R.K. Dell (1971) illustrated many Sowerby and Reeve syn-
types from the Cuming Collection in the British Museum in
his report on mollusks from the Royal Society Expedition to
southern Chile. His collection, however, was not sufficiently
complete to enable a full revision, and his list of 1 1 “possible
species” differs considerably from that adopted here.

L. Marincovich (1973) figured the five most abundant
species from Iquique in northern Chile but did not discuss

their synonymy. The Marincovich collection is preserved at
the Los Angeles County Museum of Natural History.

J. Christiaens (1973) did not treat the Chilean species in
his review of the tropical Fissurella species; however, he
proposed the subgenus Corrina for F. alba Philippi, a species
here placed in the synonymy of F. oriens. Corrina is here
regarded as a synonym of Fissurella, sensu stricto.

J. Ramirez-Boehme (1974) gave a key that included 30
different species (plus two varieties) of Fissurella from Chile.
Some of the taxa recognized in his key have never been
illustrated and have been considered indeterminate by other
authors. He also introduced four synonymous names in Fis-
surella (along with 2 1 “new species” names for acmaeid lim-
pets). All were figured in watercolor. Types are preserved at
the Museo Nacional de Historia Natural, Santiago.

In July, 1978, I distributed copies of a preliminary draft
of this manuscript to a number of Chilean biologists whom
I had met in 1975. That version differed from this primarily
in recommending the replacement of F. rudis Deshayes, 1 830,
for F. costata Lesson, 1831. However, Deshayes’ name is
preoccupied, so the better known name of Lesson is rein-
stated here. I am gratified to see that my present classification
scheme has been adopted by Chilean biologists, and am grate-
ful for the help they have provided in making this account
the more complete.

Recent papers on the biology of Fissurella species are those
ofAcuna(1977), Bretos(1978, 1979, 1980, 1982, 1983), Jara
and Moreno (1984), Moreno and Jaramillo (1983), and Mo-
reno et al. (1984).

STRUCTURE
Internal Anatomy

Anatomy in the Lissurellidae has been treated by Boutan
(1885), Illingworth (1902), Tobler (1902), Ziegenhorn and
Thiem (1925), and Odhner (1932). The latter two accounts
included references to Chilean species of Fissurella. Lretter
and Graham (1962) gave a number of useful drawings of
fissurellid anatomy. The reader is referred to these works for
details.

Although some incidences of hermaphroditism are known
in fissurellids (see Lretter and Graham, 1964), to my knowl-
edge, Fissurella species are gonochoristic, having separate
sexes. There are no apparent external sexually dimorphic
features, although the testis of males is beige-colored and the
ovary of females is bright green. The gonads discharge through
the right kidney, which therefore has a reproductive as well
as an excretory function. The Lissurellidae are unusual among
archaeogastropods in having a highly reduced left kidney,
which is nearly vestigial. In contrast, the left kidney is a
prominent papillary sac in the archaeogastropod families
Pleurotomariidae, Haliotidae, and Trochidae. These families
also differ in having the spiral caecum appendage to the
stomach and a well-developed hypobranchial gland attached
to the mantle skirt, structures that are lacking in the Lissu-
rellidae.

Anatomy in the Lissurellidae is so unlike that of the Pleu-
rotomariidae, Haliotidae, and Trochidae, that the affinity is

Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella 5

Figure 2. External anatomy of Fissurella picta. Two views of same
preserved specimen removed from shell, the mantle skirt cut above
the head to the excurrent siphon and folded back to show the paired
gills in the mantle cavity. Bahia York, Isla de los Estados, Argentina.
LACM 71-177, shell length 75.4 mm. Left, anterior view, showing
radular ribbon protruding from mouth. Right, dorsal view.

distant from these groups. I have argued (McLean, 1984a)
in support of the theory that the Fissurellidae were derived
from the extinct Paleozoic Bellerophontacea. The bilateral
symmetry of the Fissurellidae would therefore not be sec-
ondarily derived, as has been assumed by most authors, but
primitive.

External Anatomy

Structures of the head are the snout, which terminates in a
broad oral disc (Fig. 2), and cephalic tentacles, the eyes at
the bases of the tentacles. The cephalic tentacles extend for-
ward when the animal is active.

The body is attached to the shell by a horseshoe-shaped
shell muscle, which is open anteriorly, corresponding to the
mantle cavity above the head. The animal may be detached
from the shell by severing the shell muscle. Structures within
the mantle cavity (Fig. 2) may then be observed by cutting
the thin mantle roof tissue above the head. This exposes a
pair of large, bipectinate gills, which fill most of the space in
the mantle cavity on either side. The gills are attached by a
long ventral (efferent) membrane and a short dorsal (afferent)
membrane. The anus opens close to the foramen. Water cur-
rents enter above the head, pass the gills, and exit through
the foramen of the shell, sweeping the fecal material out at
the same time. The currents are propelled by bands of cilia
on the gill filaments. Left and right kidney openings are po-
sitioned near the anus.

In living Fissurella , the shell edge is enveloped by the
mantle fold, which secretes and protects the growing edge of

Figure 3. Fissurella oriens, SEM view of juvenile shell 1.9 mm in
length, showing coiled protoconch and early foramen. Islota Nihuel,
Chiloe Province, Chile. LACM 75-42.

the shell and has sensory papillae scattered on its surface.
The mantle fold is color-banded to match the rayed pattern
of the shell. The mantle fold is capable of expanding to cover
the entire shell and foot sides. The foramen is also bordered
by mantle folds that have papillae and a pigment pattern
similar to that of the mantle at the shell margin. The diameter
of the excurrent opening in the mantle skirt varies under
differing conditions of exposure to air or water.

The pigmented side of the foot has a single row of short,
stubby epipodial tentacles, extending anteriorly to the head.
These tentacles are poorly developed. The elaborate mantle
lobes probably have more of a sensory function than do the
epipodial tentacles. In other archaeogastropod families, par-
ticularly the Haliotidae, the epipodium is well developed,
forming several rows of tentacles on a separate fold called
the epipodial lobe.

Radula

The radula consists of rows of chitinous teeth on a long ribbon
(shown projecting through the mouth in Fig. 2). The entire
radular ribbon may be as much as '/j the length of the shell.
The teeth rows are rhipidoglossate, with a narrow rachidian
(central tooth), four pairs of lateral teeth shaped like the
rachidian, a pair of large, four-cusped outer lateral teeth, a
pair of uncusped lateromarginal plates, and a large number
of marginal teeth. The lateromarginal plates separate the large
outer laterals from the “books” of marginal teeth. The fis-
surellid radula is markedly asymmetrical (Hickman, 1981,
1984), with teeth on the left side of the ribbon higher or more
anteriorly placed than those on the right, extending forward
of the rachidian, which itself is asymmetrical. This pro-
nounced asymmetry enables the large outer laterals to inter-
lock like the teeth of a zipper when the ribbon is folded and
retracted at the close of the feeding stroke. The teeth are
folded in the same way while developing in the radular sac.

6 Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella

Figures 4 through 7. Shell structure of Fissurella latimarginata. SEM views of single fragment from young specimen, courtesy H.A. Low-
enstam. LACM 64-16, Iquique, Chile. (4) Interior view of shell fragment, the broad, beveled interior margin (calcitic layer of exterior) of
shell at right (length at margin 2.14 mm), and the smooth interior aragonitic layer in center and left. Fractured area at lower left exposes the
crossed lamellar structure of the aragonitic layer and the platy calcitic layer below, x 30. (5) Enlargement of lower left area of shell fragment,
showing the smooth interior at top, the fractured surface of crossed lamellar aragonite below, and the transition between the latter and the
fractured platy calcitic layer at the lower edge of the frame, x 200. (6) Enlargement of upper left corner of shell fragment. Vertical lines are
the lamellae of the smooth interior aragonitic layer. Curved lines represent the successive positions of the expanding muscle attachment area.
xl20. (7) Enlargement of lower right comer of fragment, showing the smooth growing edge (calcitic layer), the undulations reflecting the
external sculpture of radial ribs. The broad, beveled margin has an irregular surface (for mantle contact) between the smooth edge and the
smooth aragonitic surface at the left, x 80.

Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella 1

The large outer laterals are the functional teeth; the rachidian
and inner laterals are so small that their role is minimal.

Shell Morphology

The apical perforation, or foramen, is the most striking shell
feature. It enlarges as the shell grows; shell material is re-
sorbed by the mantle tissue that surrounds the foramen. Young
shells have a coiled protoconch, but this is obliterated by the
expanding foramen and is generally present only in juvenile
specimens of up to two mm in length (Fig. 3). Although the
postprotoconch stage of Diodora has a selenizone (slitband),
this is lacking altogether in developing stages of Fissurella
(McLean, 1984a).

Prominent features of the shell interior are the horseshoe-
shaped muscle scar, open in front, and the apical callus, a
broad flat area surrounding the foramen. The two round
terminations of the muscle scar are connected by a line that
marks the anterior attachment zone of the mantle skirt.

Shell Structure

Most fissurellids have the shell composed entirely of the
aragonitic form of calcium carbonate (Boggild, 1930;
MacClintock, 1963, 1967). Boggild (1930) noted that the
Chilean Fissurella crassa also has an outer layer composed
of the calcitic form of calcium carbonate. Other species treat-
ed here were not mentioned by Boggild, but all have a similar
two-layered shell (Figs. 4-7). The shell structure of radial ribs
and the pigmentation is confined to the outer layer. This
layer has a waxen, translucent appearance. A periostracum
is lacking.

In the Peruvian and Magellanic species, it is the outer
calcitic layer that comprises the broad, pigmented interior
margin of the shell. This layer is secreted by the mantle lobe
only at the growing edge. In contrast, the opaque white ara-
gonitic inner layer is deposited throughout the interior and
thickens with growth. The interior aragonitic layer is thick
only in the apical region, where it has greatly augmented the
thin calcitic layer of the early shell. Away from the apical
area, the inner layer becomes thinner, its depth only one-
third to one-fifth the thickness of the calcitic layer; it is lacking
altogether at the shell edge (margin).

The prismatic structure of the calcitic layer is not readily
apparent under low magnification; however, the structure of
the opaque white aragonitic layer can be seen under the dis-
secting microscope. It shows a series of lines running parallel
to the shell margin, a typical feature of “concentric cross-
lamellar” shell structure. The lamellae may be seen through-
out the interior of the shell, including the muscle scar and
the apical callus (Fig. 6).

Tropical species of Fissurella have shells composed en-
tirely of aragonite and lack the distinctively colored inner
shell margin. This difference between the tropical species
(Fig. 30) and the cooler-water species is here treated as a
subgeneric distinction.

In molluscan species with both calcite and aragonite de-
posited in separate shell layers, the ratio of calcite to aragonite
deposition varies with temperature. A greater percentage of

calcite deposition takes place at colder latitudes and season-
ally, during winter months (Lowenstam, 1954, 1964; Ver-
meij, 1978). Lowenstam (1954) noted a greater percentage
of calcite deposition in species of Mytilus and Littorina as
latitude increased. This is apparent in specimens of Fissurella
picta from different latitudes. In F. picta from the Strait of
Magellan at 53°37' S (Fig. 145) the aragonitic layer is no-
ticeably thinner than in F. picta from 42°42' S (Fig. 144).
The greater calcitic deposition in cold water helps to explain
why these species are so prolific at high latitudes, where they
reach a much larger size than do their tropical counterparts.
The calcitic layer of the South American species is evidently
an adaptation to cold water.

Calcite is more stable than aragonite, and fossil calcitic
shells are generally better preserved. This is evident in the
specimen of the Pliocene F. concolor Philippi, 1887, from
Antofagasta (Fig. 17). Only the calcitic outer layer remains;
the aragonitic interior is completely missing. However, spec-
imens in old shell piles indicate that when exposed to sub-
aerial weathering, the calcitic layer fractures and separates,
whereas the aragonitic layer tends to remain intact.

BIOLOGY AND ECOLOGY

Habitat

As in other limpet families with large numbers of sympatric
species, each of the Peruvian-Magellanic Fissurella species
has a unique habitat or niche.

All tropical species of Fissurella are limited to the intertidal
zone, but some of the Peruvian-Magellanic species extend
into the subtidal zone. Fissurella peruviana, F. pulchra, and
F. oriens may occur more abundantly in the sublittoral than
in the lower intertidal zone. Fissurella maxima, F. cumingi,
and F. latimarginata occur commonly from the lower inter-
tidal zone to a depth of about 5 m. The intertidal occurrence
of these species is limited to areas protected from strong wave
exposure.

The remaining species are intertidal and do not occur in
the sublittoral zone. The highest occurring species is Fissu-
rella crassa, which is tightly wedged in crevices when exposed
at low tide. Fissurella limbata and F. costata live exposed
to surf in the lower intertidal zone, F. limbata on horizontal
surfaces, and F. costata on vertical surfaces. Fissurella nigra
occurs on the undersides of large rocks in protected tide pools
at mid-tidal to lower intertidal levels.

Fissurella bridgesii has a unique habitat. It occurs on rocks
near sandy areas, unlike the others, which avoid proximity
to sand.

Fissurella picta has a more ubiquitous occurrence. It is rare
at its northern limit, where it is sympatric with other species,
but to the south of the southern limit of most of the other
species it occurs from the mid-tidal to lower intertidal zone
under various conditions of exposure, filling niches that are
occupied by other species in the north.

Feeding

Little is known of the feeding habits of Peruvian and Ma-
gellanic Fissurella. Many genera of fissurellids feed upon

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McLean: Peruvian and Magellanic Fissurella

sponges and detritus (Fretter and Graham, 1976), but Ward
(1966a) has shown that the tropical species F. ( Cremides )
barbadenisis ( Gmelin, 1791) feeds upon algae. Bretos(1978)
indicated that F. crassa feeds upon such green algae as Ulva
and Enteromorpha.

At Mehuin in southern Chile, Fissurella picta lata is a
nocturnal herbivore, feeding upon the red alga Iridaea bory-
ana and the green alga Ulva rigida (Jara and Moreno, 1984;
Moreno and Jaramillo, 1983; Moreno et al., 1984).

Information on the diets of the other South American
species is needed.

Reproduction and Growth

Bretos (1983) treated reproduction in F. maxima, finding
that spawning occurred in late November-December (late
spring) and again in July-August (winter). The breeding cycle
of the tropical F. barbadensis was treated by Ward (1966b).
That species is known to have a pelagic phase of two to three
days duration.

Bretos (1978, 1980) studied the growth rate of F. crassa,
finding that growth is rapid in early spring and late summer,
slower in late spring, autumn, and winter. Harvestable sizes
were reached in 2 to 4 years. Two growth rings were formed
each year and were considered reliable indicators of growth
for the first 6 years, after which growth was slowed and the
rings could not be identified. Fissurella maxima was also
determined to form two growth rings each year (Bretos, 1 982).
Growth in F. latimarginata was studied by Acuna (1977).

The El Nino event of 1982-1983 had a major effect on
the Fissurella populations in northern Chile. According to J.
Tomicic (personal communication), all large Fissurella species
near shore were killed during the austral summer of 1 982—
1983. However, in November, 1983, fast growing juveniles
were reported as abundant.

Epibiotic Associations

Fissurella shells offer a surface for colonization by many
species of algae and invertebrates. Many shells are so en-
crusted that color patterns are obscured (Fig. 1). Only those
species that occur high in the intertidal zone {F. crassa ), or
predominantly on the undersides of large rocks in the lower
intertidal (F. nigra and F. pu/chra), have shells that stay
relatively free of encrusting organisms.

Species of the shallow sublittoral zone (F. latimarginata,
F. cumingi, and F. maxima) generally have a thick algal mat
consisting of dense tufts of finely branched red algae, which
reaches a height of 5 mm above the shell (Figs. 1, 8). This
algal growth is absent on the rocky substrate because grazing
by the black urchin Tetrapygus niger, and presumably by the
Fissurella species, leaves the rock barren except for encrust-
ing coralline algae.

Although the algal mat on the shells of a living Fissurella
would be a source of food for other individuals of Fissurella
as well as the urchins, they evidently do not tolerate grazing
by their own kind or by the urchins, because the algal mat
is usually intact. 1 observed thick algal mats on most shells
seen in the course of diving, those in shell piles on the shore.

and those sold in markets. The chiton Chaetopleura peru-
viana is able to graze successfully on the algal mat of shells
(Fig. 8).

Balanus psittacus is frequently found on subtidal Fissurella
shells, and B. flosculus occurs on shells in exposed intertidal
habitats, particularly on F. costata and F. limbata.

The mussel Semimytilus algosus may form aggregations
on specimens of F. latimarginata (Fig. 74).

Epibiotic growths on Fissurella shells provide a protective
advantage, making it more difficult for boring organisms to
penetrate the shell. Those specimens of F. latimarginata that
have lost the algal mat are usually deeply eroded. The ad-
vantage of epibionts to chamid bivalves was discussed by
Vance (1978).

Epibiotic Scurria parasitica

Most individuals of F. crassa and F. limbata, two species
that only occur in the intertidal zone, have a single (or rarely
two) Scurria parasitica, an acmaeid limpet, attached to “home
scars” on the shell (Figs. 9, 10). This limpet occurs also on
shells of Scurria viridula and the chitons Enoplochiton niger
and Acanthopleura echinata (see Marincovich, 1973) and
rarely on other species of intertidal Fissurella. I am aware of
no studies on its biology.

Lindberg (1976), Dwyer and Lindberg (1981), and Lind-
berg and Dwyer (1983) described the home-scar depression
of the Californian acmaeid Collisella scabra on the shells of
the mussel Mytilus californianus and the acmaeid limpet
Lottia gigantea, noting the similarity of the scar to that formed
by Scurria parasitica on chitons. These epibiotic limpets pro-
duce deep scars on their host shells (Fig. 10), in which there
is an outer depression corresponding to the shell margin, and
an inner depression to correspond to the breadth of the foot.
The outer depression provides a seal that helps to prevent
desiccation and dislodgment. Microscopic examination of
the home scars shows the presence of radular scraping marks,
which indicates that enlargement of the scar is an activity
unassociated with feeding. Lindberg and Dwyer (1983) also
found evidence of shell dissolution by acidic mucopolysac-
charides secreted by the foot and carbonic anhydrase by the
mantle edge.

The feeding range of the limpets studied by Lindberg and
Dwyer (1983) was restricted to the shells of the host mollusks.
This is probably true for S. parasitica on shells of F. limbata
and F. crassa. Scar-bearing Fissurella shells do not have other
encrusting organisms and are always deeply eroded.

Shell Borers

Cirratulid polychaetes of the genus Dodecaceria (identified
both by K. Fauchald and J.C. Castilla) commonly burrow
into Fissurella shells, riddling and weakening them (Fig. 1 1 ).
The polychaete tubes open at the exterior surface, where they
are ordinarily concealed by the algal mat. The inside diameter
of the tube reaches 1.3 mm. The burrows are visible on the
shell interior, but do not break the surface except in gerontic
shells. Gibson (1978) briefly discussed Chilean species of

Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella 9

Figures 8 through 15. Shell epiphytes, shell grazers, shell borers. (8) Dense mat of red algae on exterior of F. latimarginata, with chiton
Chaetopleura peruviana and grazing trails made by the chiton. LACM 75-29, Los Molles, Aconcagua Province, Chile, shell length 106 mm.
(9) Epizoic limpet Scurria parasitica on F. crassa. LACM 90796, Iquique, Chile, shell length 57.5 mm. (10) Scar of .S’, parasitica, showing
inner and outer depression of scar. Same specimen, length of scar 13. 1 mm. (11) Shell of F. latimarginata infested with borings by cirratulid
polychaete Dodecaceria sp., exterior view of cleaned shell showing openings to burrows; interior view showing trace of burrows. LACM 75-
29, Los Molles, Aconcagua Province, Chile, shell length 115 mm. (12) Shell of F. cumingi removed from stomach of clingfish Sicyases
sanguineus, showing breakage pattern caused by this predator. LACM 75-31, Islota Concon, Valparaiso Province, Chile, shell length 50.8
mm. (13) Shell of F. costata with excavated depressions made by vermetid gastropod Dendropoma sp. LACM 75-27, beach-worn specimen,
Bahia El Teniente, Coquimbo Province, Chile, length 50 mm. (14) Shell of F. cumingi with burrows of barnacle Cryptophialus minutus.
MACN 9027-1 1, specimen studied by Tomlinson (1969), locality unknown, shell length 83.7 mm. (15) Exterior surface of F. limbata showing
burrows of Cryptophialus. LACM 75-19, Los Colorados, Antofagasta Province, Chile, length of field 21 mm.

10 Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella

Dodecacaeria. However, the systematics of the group re-
mains poorly known.

K. Fauchald (personal communication) has also identified
the spionid polychaete Polydora sp. as a borer in F. lati-
marginata.

Polychaete burrows in Fissure/la shells provide shelter for
young specimens of the bivalve Hiatella solida, which may
then bore further into the shell, as has been reported in shells
of F. nigra by Gallardo and Osorio (1978).

Acrothoracican barnacles (burrowing barnacles) penetrate
the shells of Chilean Fissurella species (Figs. 14, 15). Tom-
linson (1969:88) reported the species Cryptophialus minutus
Darwin, 1854, in a shell of Fissurella maxima. The specimen
examined by Tomlinson (actually F. cumingi) is illustrated
here (Fig. 14). I have found this pattern of burrows in a
specimen of F. limbata from Antofagasta (Fig. 15) and in F.
latimarginata from a number of localities.

Burrows of a vermetid gastropod, Dendropoma sp., have
been noted on specimens of F. costata (Fig. 1 3), forming one-
whorled depressions nearly flush with the surface of the host
shell, but not penetrating to the interior.

Parasites

Bretos and Jiron (1980) reported that digenetic trematodes
of the genus Proctoeces Odhner, 1911, family Fellodisto-
midae, were present in the gonads of eight species of Fis-
surella in northern Chile. Percentages of infected individuals
in each species ranged from 14% to 97%. The effect of this
parasitism on reproduction in the host species is unknown.

Seastar Predators

The seastar He/iaster helianthus is a voracious predator upon
many species of mollusks in the lower intertidal zone (Paine
and Palmer, 1978; Castilla, 1981). However, Fissurella has
a highly effective escape response. Fissurella respond to ini-
tial contact with Heliaster by first raising the mantle fold
above the edge of the shell, preventing the seastar’s tube feet
from making contact with the shell; thereupon they move
rapidly out of reach. A collector with a seastar in hand may
dislodge tightly wedged specimens without using a tool. The
raising of the mantle to cover most of the shell surface is
similar to the response of the north Pacific fissurellid Diodora
aspera to various seastar predators (Margolin, 1964).

Fissurella costata is the only species that fails to show an
escape response to Heliaster. It remains tightly appressed.
Its foramen, the smallest among the larger species, is evi-
dently too small for penetration by Heliaster.

The overall effect of Heliaster on populations of various
species of Fissurella probably is not significant. Large indi-
viduals can move fast enough to escape and the small, less
motile ones stay out of reach in crevices, or on the undersides
of rocks.

In southern Chile, the asteroid Meyenaster gelatinosus is
a major predator on many mollusks (Dayton et al., 1977).
Fissurella and other gastropods escape predation from this

seastar in surging water by allowing the water motion to move
them away.

Vertebrate Predators

Non-human vertebrate predators that include Fissurella
species in their diets in central Chile are the Chilean sea otter
Lutra felina, the seagull Larus dominicanus, the oyster catch-
er Haematopus ater, and the clingfish Sicyases sanguineus
(see Castilla, 1981).

The Chilean sea otter, the “nutria de mar,” has a restricted
and localized distribution, but where it occurs, the effect of
this carnivore is significant. It feeds upon Sicyases, Concho-
lepas, Fissurella species, the acmaeid limpets Scurria species
and at least three species of crabs. Castilla and Bahamonde
(1979) gave a more complete account of the ecology of Lutra
felina.

According to Castilla (1981), Haematopus ater feeds upon
Concholepas, Scurria species and Fissurella species; Larus
dominicanus feeds upon Concholepas, crabs, herbivorous
snails, Fissurella species, chitons, Scurria species, and mus-
sels.

Sicyases feeds on a wide variety of invertebrates and algae
on vertical walls in the surf-exposed intertidal zone (Paine
and Palmer, 1978). Those authors reported small specimens
of several species of Fissurella, and even one relatively large
specimen of F. cumingi (Fig. 12), in clingfish stomachs. Most
of the Fissurella shells were broken at one end, presumably
by the strong teeth of this predator. Many shells cast up on
beaches are broken in a similar way, suggesting that Sicyases
is a major predator on Fissurella. A study of the breakage
pattern in beach-worn shells would be useful to further doc-
ument the feeding of Sicyases.

Fissurella costata is well adapted to habitats where Si-
cyases occurs. It attaches tightly, making it difficult for the
clingfish to get hold of the shell. Other species of Fissurella
have poor defense against Sicyases because the shell edge
normally is raised and the mantle and foot exposed. Sicyases
may be such an effective predator that it completely removes
other species that stray into its habitat.

Human Predation and Economic Importance

Man is the chief predator upon Fissurella. Large individuals
of all species are used for food throughout Chile and Peru. I
found six species for sale in the municipal market at Iquique:
F. crassa, F. maxima, F. latimarginata, F. cumingi, F. lim-
bata, and F. bridgesii. They are collectively know as “lapas”
and are not sorted by species when sold. Although the fishery
for the lapa is on a small scale compared to that of the “loco,”
Concholepas, it amounts to a significant predation pressure
on the larger-shelled species. Those sold in the market are
kept intact in the shell. However, the shorelines in Chile have
numerous piles of discarded shells (Fig. 1). According to
figures from the Chilean Servicio Nacional de Pesca (SER-
NAP), 451,000 tons of Fissurella species were harvested in
1982 (C.A. Moreno, personal communication).

In populated areas, human predation on Fissurella is sig-
nificant. Moreno et al. (1984) found few specimens of F.

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McLean: Peruvian and Magellanic Fissurella 1 1

picta lata over 4 cm in length in the vicinity of Valdivia.
Much larger sizes occurred in areas where human access was
restricted. Where the Fissurella were experimentally re-
moved from the habitat, there were dramatic increases in the
algal cover of Iridaea boryana, its chief food source. This
alga is also harvested in Chile. Human predation on Fissu-
rella therefore helps to ensure a good harvest of the alga.

Fishermen in Chile know each species by a common, de-
scriptive name (Bretos, personal communication). The names
in use in northern Chile are mentioned in the species ac-
counts.

SYSTEMATIC CHARACTERS

Useful shell characters include: size; outline in dorsal view;
sculpture— the strength and spacing of the radial ribs; color
pattern; interior shell margin — the thickness and pigmenta-
tion pattern; and foramen— the size, shape, and placement.
Other shell characters such as the muscle scar and the internal
callus show few significant differences and are therefore not
treated unless they have unusual features. The organisms that
encrust the surface of shells can provide important ecological
information, but shells must be cleaned in order to see the
sculpture and color pattern.

Juvenile shells are commonly very different from mature
shells and are therefore separately described in this account.

Features of external anatomy such as color of the cephalic
tentacles and development of mantle lobe papillae and tu-
bercles on the foot are specific characters that may enable
identification of species.

The morphology of the large outer lateral tooth of the
radula provides a specific character.

The most useful specific characters are discussed in greater
detail as follows.

Size and Shape

The anterior end of the shell is narrower than the posterior.
The muscle scar opens anteriorly. All shells are illustrated
here with the anterior at the top; the lateral view shows the
left side.

The outline in most species is elongate-oval. Fissurella
costata, F. picta lata, and F. peruviana are generally rounder
than other species. Fissurella latimarginata, F. cumingi, and
F. pulchra are wedge-shaped, having a relatively narrow front
end and tapered sides.

Shells seldom lie flat in one plane. In most species, the
sides are slightly elevated relative to the ends. This is par-
ticularly true of such highly motile species as F. maxima,
and enables a better fit on rounded rock surfaces. Species
that commonly nestle in crevices or have a habitual site of
home attachment may instead have elevated ends for a better
fit. Both extremes are possible in F. nigra and F. oriens. Some
specimens of these species have both elevated sides and el-
evated ends, so that the shell rests on four comers.

Shell height is fairly constant in some species and variable
in others. Species with relatively low shells (length 3.2 to 5.9
times height) include F. crassa, F. bridgesii, and F. pulchra.
Those that vary from low to medium in height (length 2.8

12 Contributions in Science, Number 354

to 4.5 times height) include F. maxima, F. latimarginata, F.
costata, F. radiosa, and F. oriens. Fissurella peruviana varies
from extremely low to high (length 1.5 to 4.8 times height).

In some species, the shell may be steeply conical in young
stages and abruptly become more flattened at later stages.
This commonly happens in F. oriens, F. costata, and F.
limbata.

Fissurella nigra may grow by increasing the shell height
while contracting the length and width, especially in gerontic
specimens. This makes the slopes convex and the shell mar-
gin very thick. This growth form has not been observed in
other species.

Sculpture

Shell sculpture is relatively consistent within most species.
The radial ribs produced in the earliest growth stages are
called the primary ribs and those arising between the primary
ribs at later growth stages are called the secondary ribs. Sec-
ondary ribs attain the size and prominence of the primary
ribs in F. latimarginata, F. cumingi, and F. oriens. Primary
ribs are stronger than the secondary ribs at all growth stages
in F. picta, F. radiosa, F. costata, F. maxima, and F. peru-
viana. Fissurella crassa, F. pulchra, and F. bridgesii generally
have smooth shells in mature stages, although their young
stages have rounded primary ribs, but no secondary ribs.

Color Pattern

Color patterns in all species have definite limits of variation,
extensive in some, limited in others. Most species have a
pattern of dark-colored rays on a lighter ground color. Least
variable in color pattern are F. maxima, F. limbata, and F.
pulchra. Such strongly rayed shells as those of F. picta, F.
radiosa, F. cumingi, and F. oriens have ground colors ranging
from light to dark gray or tan and correspondingly darker
rays. White shells occur only in F. oriens. The most variable
species, having both rayed forms and uniformly colored forms,
are F. peruviana, F. latimarginata, F. radiosa, and F. oriens.

In addition to the radial rays, all species may have con-
centric growth bands of varying color intensity. Changes in
the coloration of growth bands have been correlated with
changes in diet in such herbivorous archaeogastropods as
Haliotis (Olsen, 1968a, 1968b). Changes in supply of food
or a shift in the algal composition of the diet can probably
be correlated with changes in the banding of Fissurella shells.
Concentric color changes are most pronounced in F. picta,
F. radiosa, and F. oriens, the three species that range to the
high southern latitudes where ecological conditions are most
extreme. Bretos (1978, 1980) has shown that there are sea-
sonal growth rings in F. crassa.

Shells exposed to weathering fade. The dark purple or gray
rays change to red, particularly in F. maxima and F. picta
lata.

Interior Margin

The interior margin or border, composed of the calcitic layer
of the shell, generally has several bands or zones, visible also

McLean: Peruvian and Magellanic Fissurella

in cut or broken pieces of the shell. Color differences in the
margin are useful specific characters. The width of the margin
changes with growth. In young, rapidly growing shells, it is
relatively broad; in mature shells it is proportionately nar-
rower, and in old shells it may be nearly obliterated by the
encroachment and thickening of the inner aragonitic layer.
It is consistently narrow in all growth stages of F. radiosa
and F. peruviana. In most species the margin or growing edge
is flat, but in F. maxima it is convex, and in F. crassa the
entire edge is rounded.

In some species the margin of the shell is uniformly pig-
mented across its full width, in others the pigment is con-
centrated near the surface or deeper within the layer. Only
in F. nigra is the outermost zone darker than the inner zone.
In F. latimarginata and F. pulchra the outer edge is lighter
and in F. limbata the outer edge is much lighter and contrasts
sharply with the inner zone. Color rays are confined to the
outermost layers of the margin in F. limbata and F. maxima,
but extend the full width of the margin in F. cumingi and F.
oriens.

Foramen

The relative size and the configuration of the foramen changes
with growth. In young shells it is elongate, broad in the mid-
dle and constricted in two places on the sides. The foramen
can be described as tripartite, and the side walls as bidentate
if the three-lobed aspect is especially conspicuous. In most
species the foramen changes from tripartite in young stages
to oval in mature stages. In some species its size in mature
specimens varies greatly; it may become very large in some
old shells of F. oriens. In F. peruviana, the tripartite aspect
of the foramen is lost at a very early stage, and the foramen
becomes oval. Fissurella costata has a particularly small fo-
ramen at all growth stages. Fissurella limbata is unusual in
retaining an elongate foramen in mature sizes. Fissurella
crassa also retains an elongate foramen that is constricted in
the middle, although the young shells are bidentate like those
of other species.

In most species the position of the foramen is slightly
posterior to the midpoint of the shell, but in F. nigra and F.
radiosa it is more markedly so.

Juvenile Shell

The earliest juvenile shells of all species are more elevated
and conical than later stages. Primary ribs appear at an early
stage. In forms with a rayed pattern, the elevated ribs are
light-colored and the interspaces are dark-rayed. Juveniles
of many species have a pair of broad white rays extending
laterally, more prominently than the other light-colored rays.
This pattern is especially evident in the juveniles of F. la-
timarginata, in which the light rays persist until the shell is
10 or 20 mm in length. In F. nigra, F. crassa, F. peruviana,
F. cumingi , and F. maxima, the two light rays are seen only
in juveniles of less than 5 mm length. Some have character-
istic early colorations unlike the adults. Young F. nigra are
light-colored rather than black; F. oriens, F. maxima, and
F. peruviana are reddish when young; F. limbata has a zigzag

pattern of lines; F. maxima and F. cumingi have speckled
patterns.

External Anatomy

The relative size of the animal in proportion to its shell is a
useful comparison for at least those species at either extreme;
the animals of most species are relatively large and just barely
containable within the shell. The extremes are Fissurella crassa
and F. bridgesii, which have flat shells that cannot contain
the animal, and Fissurella costata and F. peruviana, which
have high conical shells, the animal easily contained within
the shell.

In most species, the cephalic tentacles are dark, reddish
on the inner side and yellowish at the tips. Fissurella nigra
is the only species that shows only shades of gray and black
on the tentacles as well as on the mantle and foot.

The mantle lobe has three edges, here called the inner, the
upper, and the lower. The inner lobe lacks papillae and is in
direct contact with the growing margin of the shell. The upper
lobe extends up over the edge of the shell, and the lower lobe
extends down. The edges of the upper and lower lobes have
finely branched papillae. The papillae of the upper lobe are
generally more strongly developed than those of the lower
edge. The area between, which is greatly expandable, is usu-
ally vertically banded to match the pattern of rays on the
shell. This area may also show dark pigment in concentric
grooves. The edge of the lower lobe of F. latimarginata is a
striking orange color, the only species so marked, making it
readily recognizable.

In all species the side of the foot is rugose or pustular.
Coloration is mottled, the tips of the pustules or tubercles
lighter in color. Overall coloration of the foot is brown or
gray in most species; however, the foot of F. cumingi has a
distinctive strawberry-red color and that of F. costata has a
pale pinkish-brown color.

Epipodial tentacles extend along the foot sides. They are
short and stubby but are slightly more prominent that the
ordinary tubercles on the foot side. They are particularly
prominent in F. oriens.

Radula

There are few specific differences in the rachidian and inner
lateral teeth in Fissurella, although those of F. pulchra (Figs.
266, 267) are somewhat unusual in having longer overhang-
ing cusps. The larger outer laterals, however, show interspe-
cific differences, as will be noted in comparing the illustra-
tions for the radula of F. nimbosa (Figs. 21, 25), F. picta
(Figs. 138, 142, 143), F. peruviana (Figs. 43, 44), F. oriens
(Figs. 193, 194), and F. pulchra (Figs. 266, 267).

NAMES, ALLOCATIONS, AND
KEY CHARACTERS

Names and Allocations

Fifty-eight names have been proposed for Recent species of
Fissurella from the Peruvian Faunal Province, which encom-
passes central Peru to central Chile, and the Magellanic Fau-

Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella 13

Table 1. Names proposed for Recent species of Fissurella from the
Peruvian and Magellanic faunal provinces, with allocations as dis-
cussed in this paper.

affinis Sowerby, 1835 = F. peruviana

alba Philippi, 1845 = F. oriens oriens

arenicola Rochebrune and Mabille, 1885 = F. oriens oriens

atrata Reeve, 1850 = F. picta picta

australis Philippi, 1845 = F. oriens oriens

bella Reeve, 1 849 = F. latimarginata

biradiata Sowerby, 1835 = F. latimarginata

bridgesii Reeve, 1 849

clypeiformis Sowerby, 1825 = F. crassa

clypeus Sowerby, 1835 = F. peruviana

cheullina Ramirez-Boehme, 1974 = F. oriens oriens

chilensis Sowerby, 1835 = F. costata

concinna Philippi, 1845 = F. maxima

costata Lesson, 1831

crassa Lamarck, 1822

cumingi Reeve, 1849

darwinii Reeve, 1 849 = F. radiosa radiosa

depressa Lamarck, 1822 = F. crassa

doellojuradoi Perez-Farfante, 1952 = F. oriens oriens

dozei Rochebrune and Mabille, 1885 = F. radiosa radiosa

exquisita Reeve, 1850 = F. radiosa radiosa

Jlavida Philippi, 1857 = F. oriens oriens

fulvescens Sowerby, 1835 = F. oriens fulvescens

galericulum Reeve, 1 850 = F. latimarginata

grandis Sowerby, 1835 = F. nigra

grisea Reeve, 1 849 = F. radiosa

hedeia Rochebrune and Mabille, 1885 = /•'. oriens oriens

hondurasensis Reeve, 1 849 = F. maxima

lata Sowerby, 1835 = F. picta lata

latimarginata Sowerby, 1835

limbata Sowerby, 1835

maxima Sowerby, 1835

mexicana Sowerby, 1835 = F. oriens oriens

multilineata, limbata var., Ziegenhom and Thiem, 1925 = F. lim-
bata

muricata Reeve, 1850 = F. picta picta
navidensis Ramirez-Boehme, 1974 = F. picta lata
nigra Lesson, 1831

nigra Philippi, 1845, not Lesson, 1831 = F. radiosa radiosa

oblonga Ramirez-Boehme, 1974 = F. oriens oriens

obovalis Lesson, 1831 = ?

occidens Gould, 1846 = F. peruviana

oriens Sowerby, 1835

papudana Ramirez-Boehme, 1974 = F. peruviana
peruviana Lamarck, 1822
philippiana Reeve, 1850 = F. radiosa radiosa
philippii Hupe, 1854 = F. radiosa radiosa
picta Gmelin, 1791

polygona Sowerby II, 1 862 = F radiosa radiosa
pulchra Sowerby, 1835

punctatissima Pilsbry, 1890 = F. latimarginata
radiosa Lesson, 1831

rubra, costata var., Ziegenhom and Thiem, 1925 = F. costata

rudis Deshayes, 1830, not Roeding, 1798 = F. costata

solida Philippi, 1845 = F. maxima

stellata Reeve, 1850 = F. cumingi

subrotunda Deshayes, 1830 = F. peruviana

tixierae Metivier, 1969 = F. radiosa tixierae

violacea Rathke, 1833 = F. nigra

Figure 16. Distribution of Fissurella species in Peru, Chile, and
southern Argentina by degrees south latitude. The identified place
names are major LACM localities, some comprising several stations.
The dots indicate that specimens are represented in the LACM col-
lection. Lines without dots indicate distributions taken from other
sources (see text).

nal Province, which includes southern Chile and southern
Argentina. These names are listed alphabetically in Table 1,
followed by my allocation. In the text that follows, I discuss
the limits of variation for each species, but do not use variety
or form names, even though some may be based upon readily
recognizable variants. Geographic subspecies are discernible
in three species of the Magellanic Faunal Province: F. picta,
F. radiosa, and F. oriens. Trinomial designations are there-
fore used for these three species.

Difficulty in determining the taxa of such early authors as
Lesson (1831) and Philippi (1845, 1857) has been due to a
lack of illlustrations. I have not succeeded in locating the
type specimens of these two authors. Nevertheless, using
evidence from the original descriptions and type localities, I
feel confident of the allocation of all names, except for F.
obovalis Lesson, which is unassigned.

Chart of Key Characters

Attempts to prepare a dichotomuous key have not been suc-
cessful because of the extreme variability of some of the

14 Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella

species. Instead, the key characters are summarized in Ta-
ble 2.

DISTRIBUTION AND ZOOGEOGRAPHY

Distributions of the 13 species of the subgenus Fissurella
known from Peru, Chile, and Argentina are shown by latitude
in Figure 16.

Nine species occur in the warm-temperate Peruvian Prov-
ince, which extends from central Peru to central Chile: F.
peruviana, F. maxima, F. latimarginata, F. cumingi, F. cos-
tata, F. limbaia, F. erassa, F. bridgesii, and / '. pulchra. Their
northern and southern distributional records differ, but all
nine are present between Iquique and Concepcion, Chile (20°
S to 37° S).

Four species are primarily members of the cold-temperate
Magellanic Province, which includes southern Chile and
southern Argentina: F. picta, F. radiosa, F. oriens, and F.
nigra. Their distributions also differ, but all are present be-
tween 43° S and 54° S.

Except for F. radiosa, three of the four Magellanic species
extend north in Chile to overlap with the distributions of the
Peruvian species. The region of overlap is that between Val-
paraiso and Concepcion, from 33° S to 37° S, in which 12 of
the 1 3 species occur. At some point to the south of Concep-
cion, four of the Peruvian Province species (F. maxima, F.
latimarginata, F. pulchra, and F. bridgesii) drop out. I did
not find them at Mehuin, Valdivia Province (39° S). Stuardo
(1964) noted a transition zone between the two provinces
from 38° S to 43° S. The transition zone noted here is there-
fore considerably to the north of that reported by Stuardo.

The number of Fissurella species occurring between Val-
paraiso and Concepcion is significantly greater than the num-
ber known either to the north or the south. Distributions of
species in other families should be considered to determine
whether this transition area has a higher number of species
than either of the two provinces treated separately.

The northernmost occurrence of the Peruvian Province
species of Fissurella is at Isla Guanape, Peru (8°33' S), where
I have found F. latimarginata, F. limbata, F. bridgesii, and
F. peruviana. I found none in January, 1974, in the transi-
tional region between the Peruvian Faunal Province and the
tropical Panamic Faunal Province at the Lobos Afueras Is-
lands in northern Peru.

None of the species is known from the offshore islands of
central Chile, Isla San Felix or Islas Juan Fernandez, either
from literature records or recent expedition material from
ANTON BRUUN cruises. A number of common Chilean
mollusks are known from these islands. The absence of Fis-
surella correlates with the brief planktonic larval stage and
resulting poor colonizing potential of fissurellids and other
archaeogastropod larvae.

Fissurella picta, F. radiosa, and F. oriens are abundant at
the southern limits of their distributions, where they un-
doubtedly extend to Cape Horn. These three species also
occur at the Falkland Islands, which region is included in the
Magellanic Faunal Province. They do not, however, occur
at South Georgia or any of the subantarctic islands east of

the Falklands (Powell, 1951). Strebel (1908:79) reported F.
exquisita [here = F. radiosa] at Paulet Island, Antarctic Pen-
insula, a record that should not be accepted without further
confirmation.

The Magellanic Faunal Province extends north through
the Patagonian region of Argentina to the Gulf of San Matias,
but only one of the Magellanic species, F. radiosa, occurs to
the north of Tierra del Fuego in Argentina. It has a subspecies,
F. radiosa tixierae, in its northernmost extent in Argentina.
Absence of the other three species in Chubut and Santa Cruz
Provinces of Argentina may be due to the vastly different
ecological conditions. In Argentina, the tidal range is ex-
treme, broad tidal flats are exposed, and inshore sediments
often consist of fine beach sand. In southern Chile, the tidal
range is less extensive, inshore waters are deep and clear, and
beaches are few.

FOSSIL RECORD

There is little information in the literature about the fossil
record of Fissurella in Peru and Chile. Ihering (1907) par-
ticularly noted the abundance of Fissurella in the Recent and
the complete lack of the genus in the lower Tertiary of Chile
and Patagonia.

In the most recent report on the mollusks of the Pliocene
and Pleistocene formations of Chile (Herm, 1969), none of
the Fissurella species was given formal systematic treatment.
Herm listed five characteristic Pleistocene species: F. micro-
trema Sowerby, 1 833 [undoubtedly F. peruviana, rather than
the tropical F. microtrema], F. costata, F. erassa, F. lata [F.
picta lata here], and F. concinna [F. maxima here]. In his
list of Pliocene species he noted only: “Fissurella, div. sp.”

One species has been described from fossil material: F.
concolor Philippi, 1887, from Pliocene beds of Mejillones,
north of Antofagasta, Chile. I have received specimens iden-
tified as this species from E. Martinez, collected from two of
the Pliocene localities near Antofagasta detailed by Herm:
the Cerro Costino locality south of Antofagasta, and the Hor-
nito locality north of Antofagasta. Specimens (Fig. 1 7) agree
with Philippi’s description in having three smaller ribs be-
tween each of the larger ribs. The overall shape and profile
is similar to that of F. maxima, but the primary ribs are
stronger than those of F. maxima. The interior aragonitic
layer is missing entirely from the specimens, in agreement
with the principle that calcitic structures are best preserved
in fossils.

A fragmentary specimen of another species from the Cerro
Costino locality has also been received from E. Martinez.
This species (Fig. 18) has some affinity to F. erassa. In the
absence of additional specimens, I am unable to further treat
the fossil record of the group.

Fissurella, sensu stricto, is one of the youngest genera in
the Fissurellidae, traced only to the Pliocene (Herm, 1969).
Except for the Caribbean type species, it is an eastern Pacific
genus, well represented in the Peruvian and Magellanic fau-
nal provinces, and with a single species ranging from Cali-
fornia to Baja California, Mexico (Fig. 19). The type species
may have become established in the Caribbean during the

Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella 15

I’able 2. Chart of key characters.

Size

Height

Mature sculpture

Outline of base

F. peruviana

small

low to

medium ribs.

oval to

25-40 mm

high

strong primaries

elongate

F. maxima

large

medium

strong ribs.

elongate

80-135 mm

strong primaries

oval

F. latimarginata

large

low to

fine ribs

tapered

70-1 1 5 mm

medium

oval

F. cumingi

large

medium

medium ribs

tapered

80-100 mm

oval

F. costata

medium

low to

strong ribs.

oval

50-80 mm

medium

strong primaries

F. picta picta

large

medium

strong ribs.

elongate

65-95 mm

strong primaries

oval

F. picta lata

medium

medium

strong ribs.

oval

50-80 mm

to high

strong primanes

F. rad. radiosa

small

low to

medium nbs,

tapered

40-55 mm

medium

strong primaries

elongate

F. rad. tixierae

small

medium

medium ribs.

tapered

25-45 mm

strong primaries

elongate

F. oriens oriens

medium

low to

fine, broad ribs

elongate

40-70 mm

medium

oval

F. o. fulvescens

medium

low

fine, broad ribs

elongate

45 mm

oval

F. nigra

large

medium

fine, weak ribs

elongate

70-1 10 mm

oval

F. limbata

large

medium

undulations

elongate

60-90 mm

oval

F. crassa

medium

low

undulations

elongate

60-90 mm

oval

F. bridgesii

large

low

irregular striae

tapered

65-90 mm

oval

F. pulchra

medium

low

undulations

tapered

35-75 mm

oval

period in which the Central American seaway provided free
access between the western Atlantic and eastern Pacific dur-
ing the Miocene and early Pliocene (see Woodring, 1965,
1966).

FORMAT FOR SPECIES ACCOUNTS

Description. Shell descriptions treat the following char-
acters in order: the size range (length in mm) of examples
considered to be mature, the relative height, the outline in
dorsal view, whether the sides or ends are raised, strength of
the radial ribs, the color pattern, the shell layers, the interior
margin, and the position and shape of the foramen. Dimen-
sions for shell length, width, and height are given in that
order in the captions for the figured specimens, not duplicated
in the text. For specimens with uneven basal margins, shell
height is the maximum elevation when the shell rests upon
a plane surface.

Juvenile Shell. A separate description.

Mantle and Foot. Anatomical characters include the rel-
ative size of the body and shell, the relative prominence of
papillae on the upper and lower edges of the mantle lobe,
coloration of the cephalic tentacles, the color of the foot-side
and the relative prominence of the foot-side tubercles.

Habitat. The intertidal or subtidal occurrence, conditions
of exposure, and the epibiotic associations.

Distribution. The northernmost and southernmost verified
record, the latitude coordinates for these records, and the
source of the record. The source is the museum catalog num-
ber if the specimen has been examined, or an author and
date, if the record is based on a published account considered
to be correctly identified. This is followed by commentary
about its possible occurrence beyond the verified limits and
corrections of previous records now considered to be inac-
curate.

16 Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella

Table 2. Continued.

Shell color

Shell margin

Mature foramen

Foot side color

red, gray, white

narrow showing

oval

gray-brown mottled

rayed or solid

rays

reddish brown

broad, rays on

oval

dark brown mottled

rays on white

outer edge

solid purple

broad, solid

elongate

black with yellow

gray

reddish brown

oval

outer edge of mantle

reddish rays

broad, rays show

elongate

reddish brown

on dark yellow

on full width

oval

mottled

gray rays on

broad, rays show

elongate

pinkish gray

yellow

on full width

oval, small

mottled

split gray rays

broad, rays strong

elongate

light brown-black

on light ground

on outer edge

mottled

split gray rays

broad, rays strong

elongate

brown-black

on light ground

on outer edge

mottled

gray or reddish

narrow, showing

elongate

gray-brown

rays on white

rays

mottled

gray or reddish

narrow, showing

elongate

gray-brown

rays on white

rays

mottled

red to gray rays

medium width.

elongate

pinkish brown

on light ground

showing rays

oval

mottled

reddish rays on

medium width.

elongate

not seen

dark yellow gr.

showing rays

oval

solid gray,

broad, gray.

elongate oval

gray mottled

rays faint

dark at edge

beveled

purple rays on

broad, purple.

elongate

light gray

dark yellow

white at edge

mottled

solid brown,

brown upturned

very long,

mantle brown banded

faint rays

constricted

foot gray mottled

gray brown,

broad, solid

elongate

brown-black

rays faint

reddish brown

oval

mottled

rays and speckles

broad, reddish

elongate

light pinkish

on pinkish brown

brown

mottled

Number of Lots Examined. The total number of lots and
the number in each of the following museum collections:
LACM, AMNH, ANSP, MACN, MNHN, and USNM. These
counts also provide an indication of the relative abundance
of each species.

Taxonomic History. Indicates whether authors have
understood the species with the limits now recognized, or if
the present treatment is a departure from past classifications.

Abundance and Use. The fishery use and potential of each
species, and the common name, as provided by M. Bretos.

Characteristics and Variability. The first statement under
this heading is a brief description meant to convey the most
characteristic features of the species, followed by an assess-
ment of the variability, and whether variation may be cor-
related with geographic distribution.

Affinity and Comparisons. As assessment of the affinity to
the most closely related species and the chief means of dis-

tinguishing the species from similar forms. Remarks about
affinity are inferences. Future work may support or contradict
these suggestions.

Synonymy and Types. Reasons for the assignment of each
taxon; the type locality, measurements, and repository of type
specimens. Lectotypes are designated for lots consisting of
more than one original specimen.

SYSTEMATICS

Family Fissurellidae Fleming, 1822

I recognize two subfamilies in the Fissurellidae: the Emar-
ginulinae, which I further subdivide into tribes (McLean,
1984b, and other work in progress), and the Fissurellinae.

The Fissurellinae are the youngest members of the family,

Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella 17

Figures 17 and 18. Fossil specimens of Fissurella spp. of Pliocene age, Cerro Costino, 12 km S of Antofagasta, Chile (23°45' S, 70°26' W),
collected by E, Martinez. Specimens have lost the interior aragonitic layer. (17) F. concolor Philippi, 1887. LACM 90797 , 60.6 x 40.8 x 9.5
mm. (18) Fissurella sp., cf. F. crassa, LACM 90798, 25.2 x 24.4 x 8.6 mm (specimen incomplete).

appearing in the Cenozoic; the Emarginulinae arose in the
Mesozoic, with a burst of radiation in the Jurassic.

Subfamily Fissurellinae Fleming, 1822

DIAGNOSIS. Apex of mature shell wholly absorbed by
the foramen; foramen bordered inside by a ring of callus that
is not truncated or excavated posteriorly. Selenizone not
present in juvenile stage. Shell muscle and muscle scar lacking
intumed hooked processes. Rachidian tooth of radula narrow
at the tip, its base broad, no larger than the adjacent laterals;
the massive fifth lateral usually with four cusps; marginals
numerous.

The subfamily Fissurellinae differs from the subfamily
Emarginulinae in radular characters and in the morphology
of the shell muscle. In the Emarginulinae the rachidian tooth
is rhomboidal in shape (whether broad or narrow) and the
enlarged fifth lateral has only two prominent cusps; in the
Fissurellinae the rachidian tooth is narrow at the tip and the
enlarged fifth lateral has four cusps (except three in Ambly-
chilepas, one very small). Also, the muscle scar and corre-
sponding shell muscle of the fissurelline genera lack the hook-
shaped process of such emarginuline genera as Puncturel/a
and Diodora. Early stages of emarginuline genera have a
selenizone that may be lost in mature stages, but the sele-
nizone is not present at any stage in the Fissurellinae.

Other genera in the subfamily Fissurellinae are Ambly-
chilepas Pilsbry, 1890, and Macrochisma Sowerby, 1839,
both of which are limited to the Indo-Pacific, and have an-
imals too large to be contained in the shell.

Genus Fissurella Bruguiere, 1789

DIAGNOSIS. Shell conical, sculpture chiefly radial; fo-
ramen nearly central.

Two subgenera of Fissurella are here recognized: Fissu-
rella, sensu stricto, with a two-layered shell, the outer layer
composed of calcite, the inner layer of aragonite, and Cre-
mides H. and A. Adams, 1854, in which the shell is composed

entirely of aragonite (Fig. 30). This is the first time that the
subgenera have been so defined. As discussed above, it has
not previously been noted that the shell margin is indicative
of a two-layered shell rather than merely a color difference.

Recent authors (e.g., Keen, in Knight et al., 1960) have

Figure 19. Distribution of Fissurella, sensu stricto, in North and
South America.

18 Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella

regarded Fissurella, sensu stricto, as monotypic, and have
placed the tropical species lacking the shell margin in Cre-
mides and the South American species having the shell mar-
gin in Balboaina Perez-Farfante, 1943. In the present treat-
ment, the scope of the subgenus Cremides remains unchanged,
but Balboaina is synonymized with Fissurella , sensu stricto.

Cremides has a fossil record from the Oligocene (Keen, in
Knight et al., 1960), whereas Fissurella, sensu stricto, dates
from the Pliocene.

Subgenus Fissurella Bruguiere, 1789

Fissurella Bruguiere, 1789:xiv (genus without named species).
Type species, by subsequent monotypy of Lamarck, 1799:
Patella nimbosa Linnaeus, 1758.

Balboaina Perez-Farfante, 1943:2. Type species, by original
designation, Fissurella picta (Gmelin, 1791).

Carcellesia Perez-Farfante, 1952:31. Type species, by orig-
inal designation, Fissurella ( Carcellesia ) doellojuradoi Per-
ez-Farfante, 1952 [= F. oriens Sowerby, 1835].

Corrina Christiaens, 1973:93. Type species, by original des-
ignation, Fissurella ( Corrina ) alba Philippi, 1845 [= F.
oriens Sowerby, 1835].

DIAGNOSIS. Shell moderately large, composed of pig-
mented, outer calcitic layer that forms broad margin on inner
side; interior aragonitic layer relatively thin except in apical
area, extending to shell margin only in some fully mature
specimens. Mantle papillae well developed; foot side tuber-
cles well developed, epipodial tentacles short and stubby.

The species of Fissurella, sensu stricto, total 15: the 13
species of Peru and Chile treated in detail here, plus the
Caribbean type species F. nimbosa (Figs. 20-25) and the
Californian F. volcano Reeve, 1 849 (Figs. 26-29). The dis-
tribution of Fissurella, sensu stricto, is shown in Figure 19.

Generic Synonymy. Bruguiere’s introduction of the name
without reference to species has caused some confusion over
the type designation. Although many authors have followed
Pilsbry (1890) in the usage of F. picta as the type species,
more recent authors have followed Wenz (1938) in citing F.
nimbosa, by subsequent monotypy of Lamarck (1799).

Fissurella nimbosa (Figs. 20-25) is unlike all other tropical
species of the genus. Pilsbry (1890) described the interior of
F. nimbosa as having a “black line around the edge.” He
also observed that the species “has more the aspect of the
Chilean species than that of the West Indian,” a comment
overlooked by subsequent authors. Fissurella nimbosa is the
only species with the two-layered shell that has a tropical
distribution. It differs from all the Peruvian and Magellanic
species in having a green suffusion to the interior, a character
shared with the Californian F. volcano (Figs. 26-29).

Perez-Farfante (1943) did not notice that young shells of
F. nimbosa have a well-defined dark margin. The shell she
figured (1943, pi. 1, figs. 1, 2) is mature, with the aragonitic
layer obliterating the dark border. She considered Fissurella
to be a monotypic subgenus consisting only of F. nimbosa
for reasons not clearly stated, but presumably because F.
nimbosa is larger and more conical than most of the tropical
species. She therefore missed the affinity of F. nimbosa with

the Peruvian and Chilean species and proposed the subgenus
Balboaina for those species with the dark border.

In my opinion, F. nimbosa is sufficiently similar to other
species having the two-layered shell that it can not be sep-
arated from them on a subgeneric level. It is premature to
offer a final opinion about the subgeneric division of the
group, and I am, therefore, following a conservative course
in uniting those with the calcitic layer. Further evidence about
the affinity of the species with the calcitic outer layer needs
to be offered from other lines of investigation, for example,
electrophoresis. Until convincing arguments can be ad-
vanced to separate the Chilean species from F. nimbosa and
F. volcano, I am treating Balboaina as a synonym of Fis-
surella, sensu stricto.

Perez-Farfante’s original diagnosis of Balboaina included
the following provisions: “Margin of the shell entirely in one
plane, simple, not crenulated . . . .” Neither of these traits is
true for the majority of the southern species. Those species
having radial ribbing are in fact finely crenulate at the margin,
and most are raised at the sides.

Two other subgeneric names have been proposed: Car-
cellesia Perez-Farfante, 1952, and Corrina Christiaens, 1973.
Both names are based on type species that are here regarded
as synonyms of F. oriens. Carcellesia, type species F. doel-
lojuradoi Perez-Farfante, 1952, was based upon a single aber-
rant example of F. oriens with prominently raised ends. Cor-
rina. tvDe species F. alba Philippi, 1845, was intended to
apply to thin-shelled forms lacking sculpture. I consider F.
alba a synonym of F. oriens, based upon the scarce white-
shelled form of the species. Christiaens also included in Cor-
rina the South African species F. mutabilis Sowerby, 1835,
but that species has no dark margin and is therefore unre-
lated. Fissurella oriens is a readily recognizable species in
which the sculpture is weak, but I find no reason to single it
out as representing a subgenus. Consequently, both names
are relegated to the synonymy of Fissurella, sensu stricto.

Species Groups. Pilsbry (1890) placed the species having
a “distinct dark marginal border inside” in four groups of
species based on shell characters. I recognize a somewhat
similar scheme of three groups within the subgenus Fissu-
rella, sensu stricto, based on the presence or absence of com-
plex radial sculpture and the relative thickness of the calcitic
and aragonitic shell layers. Affinity among the species within
each group is inferred, but this should be tested by future
workers.

1 . Aragonitic layer as thick as calcitic layer

Group of F. peruviana

Aragonitic layer markedly thinner than calcitic layer
2

2. Sculpture of primary and secondary ribs

Group of F. maxima

Sculpture of broad primary ribs only

Group of F. limbata

Group of Fissurella peruviana

Relatively small-shelled species in which the thickness of the
aragonitic layer equals or approximates that of the external

Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella 19

Figures 20 through 30. Shells, radulae, mantle lobes, and cut shells of Fissurella nimbosa (Linnaeus, 1758), F. volcano Reeve, 1849, and F.
( Cremides ) virescens Sowerby, 1835. Figures 20 through 25. F. nimbosa. (20) Cut shell, showing thin, dark calcitic layer. LACM 76-30, Puerte
La Cruz, Venezuela, length of cut edge 20.5 mm. (21) Radular ribbon. Same locality, width of ribbon, 1.4 mm. (22) Shell, Frigate Bay, St.
Christopher, U.S. Virgin Islands. LACM 76-25, 27.0 x 24.9 x 8.1 mm. (23) Shell, Cabo Blanco, Isla Margarita, Venezuela, showing partial
loss of the external calcitic layer and wear obliterating the dark margin in beach-worn shell. LACM 76-28, 37.9 x 26.2 x 14.0 mm. (24)
Mantle edge. Frigate Bay, St. Christopher, U.S. Virgin Islands. LACM 76-25, length 3.5 mm. (25) SEM view of radula. LACM 76-30, Puerte
La Cruz, Venezuela, width of field 0.8 mm. Figures 26 through 29. Fissurella volcano. (26) Cut shell, showing thin calcitic layer. LACM 66-
1, Santo Tomas, Baja California, Mexico, length of cut edge 20 mm. (27) Mantle edge. AHF 1595-47, Rio Santo Tomas, Baja California,

20 Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella

calcitic layer; the shell margin (calcitic layer) narrow at all
growth stages and in mature specimens often obliterated by
encroachment of the aragonitic layer. Radial sculpture of
primary and secondary ribs, which remain pronounced at all
growth stages.

This group differs from both the group of F. maxima and
the group of F. limbata in its relatively smaller size and in
having a thicker aragonitic layer and a relatively thin and
narrow calcitic layer.

In addition to F. peruviana, this group includes the type
species of Fissurella, F. nimbosa (Linnaeus, 1757), of the
tropical Caribbean faunal province (Figs. 20-25), and F. vol-
cano Reeve, 1 849, of the warm temperate Californian Faunal
Province (Figs. 26-29). None of the species extends into the
cold temperate Magellanic Faunal Province.

Although F. radiosa has an unusually narrow margin, as
do the members of this group, it does not have a sufficiently
thick aragonitic layer to suggest that it is related to these
species.

In having a relatively thick aragonitic layer, this group of
species represents the connecting link between the tropical
species lacking the calcitic layer (subgenus Cremides), and
the South American species with thick calcitic layers (sub-
genus Fissurella, sensu stricto).

Fissurella peruviana Lamarck, 1822

Figures 31-50

Fissurella peruviana Lamarck, 1 822, 6(2): 1 5; Orbigny, 1841:
74; Delessert, 1841, pi. 24, fig. 7; Reeve, 1849, pi. 5, figs.
26a-d; Hupe, 1854:241; Philippi, 1860:181; Sowerby II,
1862:185, figs. 38-41; Pilsbry, 1890:155, pi. 33, figs. 41-
45, pi. 42, figs. 57-59; Dali, i 909: 178, 242; Mermod, 1 950:
713, fig. 22; Riveros-Zuniga, 195 1 : 1 30, fig. 35; Pena, 1970:
156; Dell, 1971:190; Christiaens, 1973:86; Ramirez-
Boehme, 1974:31 [key],

Fissurella subrotunda Deshayes, 1830:135; Deshayes in La-
marck, 1836, 7:602; Orbigny, 1841:74 [under F. peru-
viana]; Reeve, 1849, pi. 5 [under F. peruviana}.

Fissurella affinis “Gray,” Sowerby, 1835a: 125; Sowerby,
1835b:4, fig. 44; Sowerby II, 1862:185, figs. 46, 179; Chris-
tiaens, 1973:83.

Fissurella clypeus Sowerby, 1835a: 128; Sowerby, 1835b:4,
fig. 44; Reeve, 1850, fig. 76; Sowerby II, 1862:185, fig. 63;
Pilsbry, 1890:156, pi. 60, fig. 82, pi. 31, fig. 20; McLean
in Keen, 1971:901; Christiaens, 1973:83 [under F. asperel-
la],

Fissurella occidens Gould, 1846:156; Gould, 1852:364, pi.
31, figs. 473a, b; Pilsbry, 1890:155 [under F. peruviana]-,
Johnson, 1964:118 [holotype, USNM 5863],

Fissurella papudana Ramirez-Boehme, 1 974: 1 8, 3 1 [key], pi.
2, figs. 5a, b, c.

SheSL Relatively small (25-40 mm mature length); vari-
able in height from low to high conical; variable in outline
from broadly oval to elongate and uneven; plane of margin
also varying and probably conforming to an attachment site.
Sculpture of fine, often imbricate, radial ribs, primary ribs
remaining strong. Color highly variable, including some that
are solid dark red to reddish gray, some that are faintly rayed,
and some with rays of brown and white; lateral rays of elon-
gate specimens often curving forward. Margin relatively nar-
row at all growth stages, reflecting pattern of rays through
full thickness of calcitic layer. Cut shells show aragonitic layer
slightly thicker than calcitic layer. Foramen elongate and
tripartite in earliest stages but quickly becoming oval in ju-
venile shells and broadly oval in mature shells.

Juvenile Shell. Strongly sculptured, conical; frequently
reddish with two lateral white rays. Mature specimens with
brown and white rays have a red ring in the calcitic layer
surrounding the foramen, a remnant of the juvenile red phase.

Mantle and Foot. Fully retractable in the shell. Mantle
banded to match the rays of the shell. Mantle lobe relatively
narrow, upper and lower edges with branched papillae. Foot
side brown to black, with numerous, projecting, light-tipped
tubercles.

Habitat. Lower intertidal zone to 20 m, but most abundant
in the sublittoral zone. Scattered individuals occurring at low
tide, nestled in crevices on the sides of rocks, the shell outline
moulded to fit the site of attachment. Occurring at greater
depths than any other of the species in northern Chile. At
Mejillones (23°02' S) it was the only species that I saw at 10-
20 m on a sloping rocky bottom dominated by the mussel
Aulacomya ater.

Distribution. Chiclayo, Peru (6°47' S) (LACM, collector
unknown; also reported at Chiclayo by Pena, 1970), to Gua-
bun, northwestern tip of Isla de Chiloe, Chile (41°50' S)
(LACM 75-40, McLean). I have found specimens at most
localities between Isla Guanape, Peru (8°32' S), and Rio Bio-
bio, Concepcion Province, Chile (36°48' S), but did not find
it at Mehuin, Valdivia Province.

Number of Lots Examined. 1 18 (LACM 52, AMNH 11,
ANSP 11, MACN 3, MNHN 17, USNM 24).

Taxonomic History. Fissurella peruviana has been reason-
ably well understood by authors, at least with reference to
its occurrence in Peru. Its presence in Chile has not been
adequately discussed; Riveros-Zuniga (1951) merely listed
previous authors who cited records from Chile. A low form
of this species with irregular outline is common in Chile (Figs.
36, 37, 39-42) and was illustrated by Riveros-Zuniga (1951),
rnisidentified as F. stellata Reeve, 1850. This form was twice
named by Sowerby in 1835 and again by Ramirez-Boehme
in 1974. Christiaens (1973) considered F. affinis Sowerby to
be a good species, but there is little justification for such a

Mexico, length 5 mm. (28) Shell. White’s Point, Los Angeles County, California. LACM 69-37, 28.7 x 20.0 x 11.7 mm. (29) Shell, same
locality, 27.2 x 21.0 x 8.3 mm. (30) Fissurella ( Cremides ) virescens, cut shell, calcitic layer lacking, La Cruz de Juanacaxtl, Nayarit, Mexico.
LACM 71-33, length of cut edge 23.7 mm.

Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella 21

Figures 31 through 42. Fissurella peruviana Lamarck, 1822. Mature shells. (31) 3-5 m, Isla Guanape, Peru. LACM 74-3, 45.1 x 23.5 x
12.8 mm. (32) Holotype, F. occidens Gould. Callao, Peru. USNM 5863, 35 x 27 x 19 mm. (33) Lectotype, F. subrotunda Deshayes. “Peru.”
MNHNP, 31.3 x 27.4 x 16.8 mm. (34) Paracas, lea Province, Peru. LACM 72-79, 31.5 x 24.0 x 17.8 (beach shell). (35) Laguna Grande,
lea Province, Peru. LACM 72-77, 31.2 x 25.9 x 11.5 mm (beach shell). (36) Iquique, Chile. LACM 64-16, 24.0 x 16.7 x 7.6 mm. (37)

22 Contributions in Science, Number 354

MfcLean: Peruvian and Magellanic Fissurella

view, as discussed below. The broad distribution and the
extent of intraspecific variation in F. peruviana have not
previously been understood.

Abundance and Use. Because of its chiefly sublittoral hab-
itat, F. peruviana seems to be uncommon, although beach-
worn shells are common throughout the range. This is the
smallest species of Fissureila in Peru and Chile and therefore
has little importance as a food resource. According to M.
Bretos, the species has no common name in northern Chile,
perhaps because it is considered the juvenile form of such
species as F. maxima.

Characteristics and Variability. Fissureila peruviana is
characterized by its small size, imbricate radial ribs, oval
foramen, narrow margin, and relatively thick aragonitic lay-
er. In lateral profile it varies from low to extremely conical;
in outline it varies from broadly oval to elongate and irreg-
ular. Its color varies from uniformly dark reddish to gray or
rayed with brown and white. It is the most variable species
of Fissureila in the Peruvian Faunal Province.

Specimens from Peru tend to be more conical and more
uniformly colored, whereas those from central Chile tend to
be flatter and are more likely to be rayed with brown and
white. However, I am not convinced that a geographic dis-
tinction can be drawn. I have noticed that the more conical
forms occur on rocks adjacent to sandy bottoms, whereas the
flattened forms with irregular outlines are found on rocky
bottoms away from sand. At Iquique and Antofagasta the
flattened irregular forms occur, but I have found some highly
conical forms associated with the scallop beds north of An-
tofagasta at Bahia Morena. Conical specimens have also been
seen from the Concepcion vicinity. The recognition of geo-
graphic subspecies is therefore not justified.

Affinity and Comparisons. Fissureila peruviana differs from
all other Peruvian and Magellanic species in having a rela-
tively thick aragonitic layer and in having the oval foramen
well developed at an early stage. Yet these differences do not
seem sufficient to regard it as unrelated to those species with
well-defined primary and secondary ribs in the group of Fis-
surella maxima. Of those species, it has the most in common
with F. costata, with which it shares overall shape and the
small, oval foramen. It more closely resembles F. volcano
from California, which is also relatively small, with a narrow
margin and proportionately thick aragonitic layer. Fissureila
peruviana differs from F. volcano in having a more oval
foramen and in lacking the green tinge of the interior and
pink-bordered callus of F. volcano. F. volcano is an intertid-
ally occurring species and F. peruviana is characteristically
sublittoral.

When compared to young specimens of other Peruvian
and Chilean species, F. peruviana can be distinguished by its
narrow margin and oval foramen. Small specimens may be

distinguished from the juveniles of F. maxima in having the
foramen more oval and the margin not rounded. The primary
ribs are not as strongly developed as those in young stages
of F. costata, F. picta lata, or F. radiosa.

Synonymy and Types. Fissureila peruviana Lamarck, 1822,
is one of the three earliest-named species of the region. La-
marck’s specimens were from the “coasts of Peru,” and some
of the specimens were said by Lamarck to be less conical
than others. Unfortunately, the first illustration of Lamarck’s
species (Delessert, 1841) may be a specimen of some other
species, as suggested by Pilsbry ( 1 890), Mermod ( 1 950), and
Christiaens (1973). Mermod (1950) discussed two Lamarck-
ian specimens in the Geneva Museum collection considered
to be original. “Specimen no. 2” discussed by Mermod fits
the present concept of F. peruviana and is inscribed by the
hand of Lamarck, according to Mermod. I am not able to
determine the identity of “specimen no. 1,” which is appar-
ently the specimen figured by Delessert. Inasmuch as La-
marck indicated that there were several specimens, I hereby
designate “specimen no. 2” as the lectotype, which is in
accord with the original intent of the author. The lectotype
(see Mermod, 1950, fig. 22-2) represents the high conical
form of F. peruviana with a nearly circular foramen; length
20, width 25, height 16 mm.

I have examined 8 syntypes and the original mounting
board of F. subrotunda Deshayes, 1830, received on loan
from the Paris Museum. Lengths are 35.6, 31.3, 30.3, 30.0,
25.7, 22.7, and 20.7 mm. The original measurements were
32 mm in length and 28 mm in width; the second largest
specimen is here designated the lectotype (Fig. 33); it is 31.3
mm in length and 27.4 mm in width and is probably the
measured specimen. Type material, from “Perou,” appar-
ently has not previously been illustrated. All specimens are
dark reddish rayed, moderately conical, and with an oval
basal outline. No comparisons were originally made with F.
peruviana; Orbigny (1841) relegated the name to the syn-
onymy of F. peruviana.

Type material of F. affinis Sowerby, 1835, has not been
located. Reeve (1849) regarded it as an elongate, flattened
form of F. peruviana. Sowerby II (1862) doubtfully recog-
nized it as a species “in order to avoid the extreme incon-
venience of including opposite characters under the same
name . . . .” Christiaens (1973) maintained it as a species,
but in my opinion the original figure in the “Conchological
Illustrations” represents the extreme flattened form of F.
peruviana, which occurs throughout the range of the species
in Chile. Several localities were mentioned originally: “In-
sulas Mexillones et Lobos, Iquiqui, and Valparaiso.”

The holotype of F. clypeus Sowerby, 1835 (Fig. 40), was
said to have come from “Sanctam Elenam,” presumably the
Santa Elena Peninsula, Ecuador. This name has baffled sub-

Iquique, Chile. LACM 75-12, 23.6 x 16.4 x 7.4 mm (beach shell). (38) 2-4 m, El Rincon de Mejillones, Antofagasta Province, Chile. LACM
75-23, 32.0 x 29.8 x 15.0 mm. (39) 2-5 m, Antofagasta, Chile. LACM 75-20, 27.0 x 18.8 x 17.6 mm. (40) Holotype, F. clypeus Sowerby.
Locality doubtful. BMNH 197578, 28.0 x 18.9 x 5.8 mm. (41) Holotype, F. papudana Ramirez-Boehme. Papudo, Aconcagua Province,
Chile. MNHN 200374, 36 x 26 x 10 mm. (42) Guabun, Chiloe Province, Chile. LACM 75-40, 27.3 x 19.7 x 7.9 mm (beach shell).

Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissureila 23

Figures 43 through 50. Fissure/Ia peruviana Lamarck, 1822. Radula, mantle lobe, cut shell, juvenile shells, and intact specimens. (43) Radula,
light microscope preparation. Montemar, Valparaiso Province, Chile. LACM 75-30, width of ribbon 0.4 mm. (44) Radula, SEM. 7-20 m,
Mejillones, Chile. LACM 75-21, width of field 1 mm. (45) Mantle edge. 1-2 m, Playa Hermosa, Ancon, Peru. LACM 74-21, length 4 mm.
(46) Cut shell. Los Colorados, Antofagasto, Chile. LACM 75-19, length 21 mm. (47) Juvenile shell. Same locality. 4.5 x 2.7 x 1.5 mm. (48)
Juvenile shell. 3-5 m, Isla Guanape, Peru. LACM 74-3, 9.0 x 5.3 x 2.4 mm. (49) Body of preserved specimen. Antofagasta, Chile. LACM
75-15, shell length 25.1 mm. (50) Living specimen. Same locality, same specimen.

sequent authors, especially because a view of the internal
margin has never been given and the foramen has been fig-
ured to be much longer that it actually is. In my opinion it
is the flattened form of F. peruviana, as was suspected by
Sowerby II ( 1 862), and the shell more likely came from Chile.
The Ecuadorian locality is well to the north of the northern
limit of F. peruviana, and the flattened form of the species
is particularly common in Chile. The foramen of the speci-
men is slightly longer than normal. Pilsbry (1890) copied the
Reeve figure, but his figure 82, said to be a copy of the
Sowerby figure, is not that, which has contributed to the
confusion.

The holotype of F. occidens Gould, 1846, is USNM 5863,
length 35 mm (Fig. 32). There are two paratypes, MCZ 1 55766
(Johnson, 1964). The type locality is Callao, Peru. The ho-
lotype represents the high-conical, reddish rayed form so
abundant in Peru.

Fissurella papudana Ramirez-Boehme, 1974, was de-
scribed without comparisons to other species. The type lo-

cality was Papudo, Aconcagua province, Chile. The holotype
(Fig. 41), MNHN 200374, is an example of the flattened
form of F. peruviana, with a narrow margin, irregular outline,
oval foramen, and brown and white rays.

Group of Fissurella maxima

Relatively large-shelled species in which the thickness of the
exterior calcitic layer of the shell greatly exceeds that of the
interior aragonitic layer. All species have strong radial sculp-
ture, at least in the early stages. Sculpture consists of primary
ribs and weaker secondary ribs that arise between the primary
ribs. Mature shells may retain the distinction between pri-
mary and secondary ribs, or all the ribs may attain a similar
strength, whether coarse or very fine.

There are eight species in the group of F. maxima, four
in the Peruvian Provice and four in the Magellanic Province.
Most of the species in this group are highly variable and
eurytopic, tolerant of a broad range of conditions of intertidal

24 Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella

Figures 51 through 55. Fissurella maxima Sowerby, 1835. Mature shells. (51) Bahia Independencia, Peru. AHF 380-35, 84.0 x 47.7 x 23.0
mm. (52) Lectotype, F. hondurasensis Reeve. Locality unknown. BMNH 1976139, 56.6 x 32.5 x 13.5 mm. (53) Iquique, Chile. LACM 64-
16, 26.0 x 14.4 x 5.8 mm. (54) Los Molles, Aconcagua Province, Chile. LACM 75-28, 73.9 x 47.4 x 18.0 mm. (55) Holotype, F. maxima
Sowerby. Valparaiso, Chile. BMNH 197569, 128.8 x 85.4 x 34.7 mm.

exposure. Most species, particularly those with high vari-
ability, have been ovemamed, the synonyms applying to
variant specimens.

Only one species in this group, F. nigra, is stenotopic, with
a narrow ecological tolerance, and low variability. It also
differs from the other in having primary and secondary ribs
pronounced only in juvenile stages.

Three of the four Magellanic species in this group have
geographic subspecies.

Fissurella maxima Sowerby, 1835

Figures 51-63

Fissurella maxima Sowerby, 1835a: 123; Sowerby, 1 8 3 5b: 3,
fig. 18; Orbigny, 1841:475, pi. 64, figs. 4-7; Philippi, 1845:

Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella 25

Figures 56 through 63. Fissurella maxima Sowerby, 1835. Radula, mantle lobe, cut shell, juvenile shell, and living specimens. (56) Mantle
lobe. Iquique, Chile. LACM 75-12, length 9 mm. (57) Radular teeth, lateral view of large outer lateral teeth. Iquique, Chile. LACM 70-68,
width of ribbon 0.4 mm, shell length 20.5 mm. (58) Radular ribbon, same specimen. Width of field 1.0 mm. (59) Living specimen, anterior
end, showing head with cephalic tentacles and foot side. Iquique, Chile. LACM 75-12. (60) Juvenile specimen. Punta El Lacho, Santiago
Province, Chile. LACM 75-32, 14.0 x 7.8 x 2.9 mm. (61) Radular ribbon, air-dried. Bahia San Juan, Peru. AHF 828-38, width of ribbon
2.5 mm, shell length 58.9 mm. (62) Living specimen on rock substrate, anterior at right. Pozo Toyo, Tarapaca Province, Chile. LACM 75-
10. (63) Cut shell. Montemar, Valparaiso Province, Chile. LACM 75-30, length of cut edge 37 mm.

3, pi. 1, fig. 1; Reeve, 1849, pi. 4, fig. 22; Hupe, 1854:239;
Philippi, 1860:180; Sowerby II, 1862:187, figs. 8, 9; Wat-
son, 1886:33; Pilsbry, 1890:145, pi. 30, figs. 8, 9, pi. 33,
figs. 46, 47; Dali, 1909:242; Ziegenhom and Thiem, 1925:
11, pi. 1, figs. 8a, 8b, 9a, 9b; Pena, 1970:156; Riveros-
Zufiiga, 1951:102, fig. 18; Dell, 1971:188, pi. 4, figs. 1-3;
Marincovich, 1973:18, fig. 31; Ramirez-Boehme, 1974:31
[key].

Fissurella solida Philippi, 1845:142; Carcelles and William-
son, 1951: 256; Dell, 1971:193; Ramirez-Boehme, 1974:
31 [key].

Fissurella concinna Philippi, 1845:143; Philippi, 1846:66,
pi. 2, fig. 5; Reeve, 1850, pi. 15, fig. 112; Hupe, 1854:245;
Sowerby II, 1862:187, figs. 4, 178, 206; Rochebrune and
Mabille, 1889:71; Pilsbry, 1890:146, pi. 32, figs. 32, 33,
pi. 45, figs. 7, 8 [as var. of F. maxima ]; Carcelles and
Williamson, 1951:256; Pena, 1970:156; Dell, 1971:183;
Ramirez-Boehme, 1974:30 [key].

Fissurella hondurasensis Reeve, 1849, pi. 7, fig. 48; Pilsbry,
1890:146, pi. 35, fig. 6.

Shell. Relatively large (80 to 135 mm mature length), low

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McLean: Peruvian and Magellanic Fissurella

to medium in height, outline elongate-oval; sides of shell
elevated. Radial ribs prominent, low and rounded, alternat-
ing in strength, crenulating margin. Ground color pale yellow;
rays dark purple, broad, uninterrupted; primary ribs centered
in each light and dark ray. Margin very broad in growing
shells, narrow in mature shells, rounded at junction with
aragonitic layer; margin zoned; outer zone narrow, showing
color rays; inner zone broader, uniformly pale and translu-
cent. Cut shells show pigment of rays concentrated at surface.
Foramen nearly central, elongate in young shells, oval in
mature shells.

Juvenile Shell. High conical, base elongate, margin round-
ed; ribs rounded, strong; color reddish, with two lateral white
rays and scattered black flecks. After length of 6 mm shell
becomes flatter, the primary ribs nodulous and much more
prominent than the secondary ribs; the light and dark rayed
pattern emerges and the black flecks are lost.

Mantle and Foot. Not fully retractable in shell; cephalic
tentacles brown, tipped with yellow. Mantle lobe broad,
banded with purplish-brown and light gray to match rays on
shell; papillae of upper edge bulbous, finely branched, those
of lower edge smaller. Side of foot brown; tubercles strongly
developed, tips lighter colored.

Habitat. Partially exposed rocky areas at low tide to 4 m.
Some individuals are free of encrusting algae and live on the
under sides of large flat rocks at low tide, but most live in
the open and are covered with a dense low growth of red
algae. When exposed at low tide they are tightly wedged in
crevices. Only rarely do specimens have attached Scurria
parasitica on the shell.

Distribution. Huarmey, Peru (10°06' S) (LACM 70-97, E
del Solar), to Lirquen, Concepcion Province, Chile (36°41'
S) (LACM 72-207, Univ. Concepcion). The southermost range
of F. maxima in Chile is not as extensive as that of most of
the other common species of the Peruvian Faunal Province.
I found no trace of it at Rio Bio-bio, Concepcion Province,
Mehuin, near Valdivia, or Guabun on the northwest tip of
Isla de Chiloe. More extreme records in the literature are
rejected: Dali’s (1909) record from Manta, Ecuador, and Ri-
veros-Zuniga’s (1951) record from Fuerte Bulnes in the Strait
of Magellan. Dell’s (1971) records from Isla de Chiloe are
based upon specimens of F. picta lata.

Number of Lots Examined. 128 (LACM 34, AMNH 25,
ANSP 7, MACN 14, MNHN 30, USNM 18).

Taxonomic History. Fissurella maxima has been recog-
nized by previous authors. Its three synonyms have not been
considered important, although F. concinna Philippi has
sometimes been given status as a narrow “form” of the species.

Abundance and Use. Common throughout its range and
particularly abundant in central and northern Chile. It is one
of the major species used for food, taken by shore collectors
and divers in relatively shallow water. It is called the “lapa
de huiros,” because it frequently occurs near the brown algae
Lessonia, known as “huiros.”

Characteristics and Variability. Always strongly ribbed and
having a consistent color pattern of dark purple rays, the
most characteristic feature is the interior border, which is

uniquely rounded and has two zones. The margin is always
crenulated by the radial ribs, even in the largest specimens.
It is not a highly variable species; color pattern and shell
height are consistent. Width of the shell is the most variable
feature; some specimens may be especially elongate (Fig. 5 1 );
oval specimens are rare. Largest specimens come from shell
piles in central Chile; those from Peru are small. Elongate
specimens are frequently those from Peru. However, I do
not consider the geographic differences of sufficient impor-
tance to warrant recognition of subspecies.

Affinity and Comparisons. Fissurella maxima has features
that represent an extreme; its rounded margin is not shared
by other species. It most resembles F. cumingi, which has a
similarly large mantle and foot, and similar size, height, and
color pattern. Fissurella maxima differs in having a rounded
margin, stronger sculpture, and more pronounced primary
ribbing. It might also be confused with F. picta lata, but it
is lower, more elongate, and has broader ribs than that species.
Juveniles are reddish like those of F. peruviana, but differ in
having an upturned margin, a more elongate foramen, and
are more elongate.

Synonymy and Types. Fissurella maxima was described
by Sowerby, 1835, from a specimen collected by Cuming at
Valparaiso, Chile. The holotype, BMNH 197569 (Fig. 55),
length 128.8 mm, matches the specimen figured by Sowerby
(1835b). Reeve (1849) figured a different specimen; these two
specimens have also been figured by Dell (1971, pi. 4, fig. 1,
holotype; figs. 2, 3, Reeve specimen).

I have not located any type material of F. solida Philippi,
1845, from “Chile.” It has not been illustrated. Philippi’s
description mentions what I interpret as the rounded internal
border of F. maxima and states that he did not know the
young of F. maxima; other features are in accord with F.
maxima, so I am confident that the name should be relegated
to the synonymy of F. maxima.

Type material of F. concinna Philippi, 1845, also from
“Chile,” has not been located. Philippi’s figures have sug-
gested to most authors that it is a small, laterally compressed
form of F. maxima. As with F. solida, Philippi noted the
rounded margin. Stunted, narrow specimens matching his
figure are present in collections; those from central Peru (Fig.
51) may be predominantly narrow, but broader specimens
occur at the same localities and it is doubtful that the name
has any taxonomic utility. Rochebrune and Mabille (1889)
used the name incorrectly for specimens from Tierra del
Fuego (no doubt confusing it with F. radiosa ); this accounts
for the Fuegan records of the “variety” concinna repeated
by subsequent authors.

Fissurella hondurasensis Reeve, 1 849, supposedly (in error)
from “Honduras,” was correctly placed in the synonymy of
F. maxima by Pilsbry (1890). There are four specimens with
the original mounting board now labeled syntypes, BMNH
1976139, lengths 56.6, 36.8, 33.0, and 28.5 mm; the largest
is here figured (Fig. 52) and designated the lectotype. All are
young specimens, more thin-shelled and with narrower mar-
gins than usual, but the dark flecks of the juveniles of F.
maxima are apparent, and I am confident of their identity

Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella 27

with F. maxima. A fifth specimen originally mounted on the
same board is a young specimen of F. pulchra.

Fissure! la latimarginata Sowerby, 1835

Figures 64-79

Fissurella latimarginata Sowerby, 1835a: 126; Sowerby,
1835b:3, fig. 69; Gray, 1839: 148, pi. 39, fig. 8; Reeve, 1849,
pi. 3, fig. 1 9; Hupe, 1 854:242; Philippi, 1 860: 1 80; Sowerby
II, 1862:185, figs. 6, 7, 12; Pilsbry, 1890:153, pi. 32, figs.
36-38; Dali, 1909:242; Ziegenhom and Thiem, 1925:17,
pi. 2, figs. 22a, 22b; Carcelles and Williamson, 1951:255;
Riveros-Zuniga, 1951:125, fig. 33; Pena, 1970:156; Dell,
1971:187, pi. 3, figs. 12-14; Marincovich, 1973:17, fig. 29;
Ramirez-Boehme, 1974:31 [key].

Fissurella biradiata Sowerby, 1 835a: 1 24; Sowerby, 1 835b;3,
figs. 23, 52; Orbigny, 1841:477; Reeve, 1849, pi. 3, fig. 20;
Philippi, 1860:180; Sowerby II, 1862:185, figs. 1-3; Zie-
genhom and Thiem, 1925:17, pi. 2, fig. 23; Dell, 1971:
182, pi. 3, figs. 10, 11.

Fissurella latimarginata var. biradiata, Pilsbry, 1 890: 1 54, pi.
35, fig. 3, pi. 46, figs. 12-14; Riveros-Zuniga, 1951:126,
fig. 34; Ramirez-Boehme, 1974:31 [key],

Fissurella bella Reeve, 1849, pi. 3, fig. 21; Sowerby II, 1862:
185, fig. 25; Pilsbry, 1890:150, pi. 33, fig. 48; Riveros-
Zuniga, 1951:119; Dell, 197 1 : 182, pi. 3, figs. 3, 4; Ramirez-
Boehme, 1974:32 [key],

Fissurella galericulum Reeve, 1850, pi. 11, fig. 77.
Fissurella latimarginata var. galericulum, Ramirez-Boehme,
1974:31 [key],

Fissurella punctatissima Pilsbry, 1890:150, pi. 58, figs. 21-
23; Dali, 1909:124; Riveros-Zuniga, 1951:118, fig. 26; Dell,
1971:192.

Shell. Relatively large (70 to 1 15 mm mature length); low
to moderately high; outline oval, markedly tapered ante-
riorly; base of shell in one plane or slightly elevated on sides.
Sculpture of very fine, sharply raised radial ribs. Color uni-
formly dark purplish red to gray, except for two lighter rays
extending laterally in young shells, fading away in mature
shells; occasional specimens faintly rayed throughout. Mar-
gin very broad and flat in young shells, solid dark red, outer
edge with narrow gray zone. Cut shells showing gray outer
zone in calcitic layer; recent growth of outermost zone in
some mature shells changing from gray to nearly colorless.
Foramen very long and tripartite in young shells, elongate-
oval in mature shells.

Juvenile Shell. Elongate and elevated, radial ribs faint,
nearly black except for two lateral white rays. After shell

length of 5 mm, new growth less conical, fine radial ribs
stronger, and shell lighter in color, not yet showing gray outer
zone to calcitic layer.

Mantle and Foot. Not fully retractable in shell; mantle lobe
usually enveloping and capable of great expansion over edge
of shell. Upper edge with finely branched tongue-shaped pa-
pillae that alternate with shorter papillae. Lower edge with
closely spaced tongue-shaped papillae of lesser size. Mantle
lobe and foot side black except for lower mantle edge, on
which branched papillae are bright yellow-orange. The bright
yellow color of the lower edge makes a very conspicuous ring
that encircles the animal, a coloration not present in other
species. Determination of living specimens is readily made
on this feature alone; in preserved specimens it remains light-
er colored that the rest of the mantle lobe.

Habitat. Lowermost intertidal zone in partially protected
areas, and in the sublittoral to depths of 5 m, living exposed
on the upper surfaces of rocks. At Antofagasta I found that
the sublittoral population occurred along with a few speci-
mens of F. cumingi and still fewer F. maxima. The rocky
substratum there looked barren from urchin grazing, but all
Fissurella shells had a thick algal mat.

Distribution. Chiclayo, Peru (6°47' S) (Pena, 1950), to Rio
Bio-bio, Concepcion Province, Chile (36°48' S) (LACM 75-
35, McLean). I have found it abundantly as far north as Isla
Guanape, Peru (8°32' S), and at all stations throughout the
range. It undoubtedly occurs further south than Concepcion,
but I did not find it at Mehuin, near Valdivia, or at Guabun
at the northwest tip of Isla de Chiloe.

Number of Lots Examined. 82 (LACM 26, AMNH 18,
ANSP 8, MACN 7, MNHN 10, USNM 13).

Taxonomic History. The normal color form of F. lati-
marginata has been understood by authors. Its synonyms are
based upon color forms with the rayed pattern, except for
Pilsbry’s F. punctatissima, which was based upon a gerontic
specimen.

Abundance and Use. Fissurella latimarginata is common
throughout its range. It occurs widely in Peru, where few of
the other species are known. It is one of the most important
food species, more so than any other in northern Chile. The
largest specimens are taken by divers. Its common name is
the “lapa viuda,” which means widow, for its black aspect.

Characteristics and Variability. Fissurella latimarginata
is characterized by its generally uniform gray to reddish-
brown color, fine but persistent ribs, and tapered anterior
end. The lateral white rays that characterize juvenile shells
of so many of the species are likely to persist through later
growth stages in this species. The broad, dark internal border

Figures 64 through 71. Fissurella latimarginata Sowerby, 1835. Mature shells. (64) 3-5 m, Isla Guanape, Peru. LACM 74-3, 41.9 x 27.1
x 1 1.7 mm. (65) Iquique, Chile. LACM 64-16, 87.9 x 64.2 x 20.7 mm. (66) Los Molles, Aconcagua Province, Chile. LACM 75-28, 34.8 x
25.5 x 9.0 mm. (67) Los Molles, Aconcagua Province, Chile. LACM 75-29, 116.5 x 85.5 x 37.5 mm. (68) Lectotype, F. latimarginata
Sowerby. Valparaiso or Iquique, Chile. BMNH 197572, 80.2 x 55.0 x 18.7 mm. (69) Syntype, F. bella Reeve. Cape Horn (probable error).
BMNH 197567, 56.7 x 37.4 x 12.7 mm. (70) Lectotype, F. galericulum Reeve. Locality unknown. BMNH 1976138, 33.0 x 21.5 x 9.5 mm.
(71) Holotype, F. punctatissima Pilsbry. Valparaiso, Chile. ANSP 50262, 88 x 66 x 22 mm.

28 Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella

Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella 29

Figures 72 through 79. Fissurella latimarginata Sowerby, 1835. Preserved specimens, mantle edge, juvenile shells, radula, and cut shell. (72)
Living specimen, showing light colored lower mantle edge against dark foot side. Antofagasta, Chile. LACM 75-20. (73) Mantle lobe. Isla
Guanape, Peru. LACM 74-2, length, 24 mm. (74) Preserved specimen with epizoic mytilids Semimytilus algosus on shell. 1-4 m, Isla San
Lorenzo, Peru. LACM 74-24, 66.2 x 42.5 x 22.0 mm. (75) Juvenile shell. Los Colorados, Antofagasta Province, Chile. LACM 75-19, 16.0 x
10.5 x 3.9 mm. (76) Juvenile shell. Iquique, Chile. LACM 64-16, 13.8 x 3.5 x 3.6 mm. (77) Radula, air-dried. Antofagasta, Chile. LACM
75-20, width of ribbon 3.4 mm, shell length 85.0 mm. (78) Head of preserved specimen, showing mantle lobe and mouth. Isla Guanape, Peru.
LACM 74-2, shell length 70.0 mm. (79) Cut shell, also showing algal mat. Los Molles, Aconcagua Province, Chile. LACM 75-29, length of
cut edge 42 mm.

is characteristic, but it is a feature shared with other species.

Living specimens are always recognizable by the bright yel-
low lower edge to the mantle lobe. Largest specimens seen
were from central Chile; smaller specimens occur at both
extremes of the range. Shell height varies extensively within
populations. Specimens with patterns of rays like those of F.
cumingi (Fig. 61) are uncommon but occur throughout the
range. There are no geographic differences of sufficient im-
portance to warrant recognition of subspecies.

Affinity and Comparisons. Fissurella latimarginata most
resembles F. cumingi, having in common the size, propor-
tions, and tapered anterior end. The unusual pattern of rays
in variant specimens of F. latimarginata is similar to the
normal pattern in F. cumingi. The sculpture of F. latimar-
ginata is finer, sharper, and less beaded than that of F. cum-
ingi; the rare color form of F. latimarginata may always be
distinguished on this difference in sculpture. Shells of F. la-

timarginata may resemble those of F. nigra but are more
sharply ribbed, have a reddish rather than gray interior bor-
der, and lack the inwardly tapered, white bordered foramen
of F. nigra. A similar elaboration of the tongue-shaped pa-
pillae of the mantle lobe is known only in F. bridgesii. Both
have broad margins and are dark colored with an incipient
tendency to produce faintly rayed variants. Fissurella lati-
marginata may always be distinguished by its fine, sharp
ribbing, which is lacking in F. bridgesii.

Synonymy and Types. There are six syntypes of F. lati-
marginata Sowerby, 1835, described from “Valparaiso and
Iquique,” Chile, four from one lot, BMNH 197572, lengths
80.2, 73.8, 61.7, and 51.1 mm, and two in BMNH 197573,
lengths 56.3 and 27.3 mm. Both original mounting boards
are penned with “Valparaiso and Iquique,” so it is not known
from which of the two localities each shell originated. The
shells are clean and uniformly dark red, faintly, or not at all,

30 Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella

showing the lateral white rays. The largest specimen is figured
here (Fig. 68) and designated the lectotype; Dell (1971) fig-
ured the 73.8 mm specimen.

Type material of F. biradiata Sowerby was not received
on loan from the British Museum. The type locality is Val-
paraiso, Chile. Figure 23 of the “Conchological Illustrations”
shows a finely ribbed shell 41 mm long (presumed life-size)
in which the only color pattern consists of the two lateral
white rays. The type figure can therefore be relegated to the
normal form of F. latimarginata without question. Figure
52 of the “Conchological Illustrations” is stated to be a “var.”
of F. biradiata from Iquique; this specimen is the one figured
by Reeve (1849) and Dell (1971). Reeve’s coloration shows
it to be reddish brown and faintly rayed. Until I can examine
the sculpture of that specimen I am unable to decide whether
it is F. cumingi or the rare, rayed form of F. latimarginata,
but the question is of minor importance because the type of
F. biradiata is clearly recognizable as a specimen of F. la-
timarginata.

Fissurella bel/a Reeve, 1849, supposedly from Cape Horn,
is represented by two syntypes, BMNH 197567, lengths 56.7
and 42.5 mm. The smaller specimen was figured by Reeve
(here designated the lectotype) and the larger one by Dell
(1971) and refigured here (Fig. 69). No author has recognized
a species based on this name. Dell suggested an affinity with
F. pulchra, but the absence of flecking rules that out. The
narrowed anterior end and rayed pattern is shared only with
the rare color form of F. latimarginata and normal F. cum-
ingi. The shells appear to be acid-cleaned, the fine radial ribs
are like those of F. latimarginata rather than F. cumingi.
The margin in both shells is narrow, suggesting a degree of
maturity that can be matched with F. latimarginata at ex-
tremes of its distribution, but not with similarly sized F.
cumingi. I therefore identify the syntypes as the rare color
form of F. latimarginata. The Cape Horn locality is therefore
erroneous.

There are three syntypes of F. galericulum Reeve, 1850,
BMNH 1976138, lengths 33.0, 3 1 .0, and 30.5 mm, described
with unknown locality. The largest (here designated the lec-
totype) is illustrated here (Fig. 70). It is clearly the normal
white-rayed young stage of F. latimarginata and has been so
recognized by previous authors.

Fissurella punctatissima Pilsbry, 1890, from Valparaiso,
Chile, was considered by Pilsbry to differ from F. latimar-
ginata in being more elevated and having a narrower internal
margin. Shell proportions of the holotype (ANSP 50262, Fig.
7 1 ) and paratype (ANSP 61923) are within the normal range
of variation and the narrow margin is that of mature ex-
amples of the species. The interior pitting, which suggested
the name, is not unusual in large shells. Dell’s records (1971)
under this name are based upon specimens of F. nigra.

Fissurella cumingi Reeve, 1 849

Figures 80-94

Fissurella cumingi Reeve, 1849, pi. 3, fig. 17; Hupe, 1854:
238; Sowerby II, 1 862: 1 87, figs. 5, 1 32; Carcelles and Wil-
liamson, 1951:256; Dell, 1971:184, pi. 3, figs. 15, 16; Ra-
mirez-Boehme, 1974:32 [key].

Fissurella latimarginata var. cumingi, Pilsbry, 1890:154, pi.

30, fig. 1; Riveros-Zuniga, 1951:28.

Fissurella stellata Reeve, 1850, pi. 12, fig. 80; Hupe, 1854:

245; Sowerby II, 1862:187, fig. 82; Pilsbry, 1890:148, pi.

32, fig. 32; Dali, 1909:242; Riveros-Zuniga, 1951:1 13, fig.

23 [looks like F. peruviana ]; Dell, 1971:193, pi. 4, figs. 7,

8 [not 5, 6]; Ramirez-Boehme, 1974:30 [key].

Shell. Large (80 to 100 mm mature length); height me-
dium; outline oval, tapered anteriorly; sides slightly raised.
Sculpture of fine to medium strength radial ribs, nodulous
or beaded in early stages along growth increments; primary
ribs only slightly more prominent than secondary ribs. Ground
color dark yellowish gray, patterned with dark rays of red-
dish-purple, anteriormost rays often split. Margin very broad
and flat in young shells, showing pattern of rays across the
full width. Cut shells not showing zoning of calcitic layer,
pigmentation of rays of equal intensity throughout layer. Fo-
ramen elongate and tripartite in young shells, elongate-oval
in mature shells.

Juvenile Shell. Oval, conical, with straight slopes, earliest
area white, reddish rays emerging with edges darkly outlined
as if split; lateral white-rayed area prominent. At length of
4 mm scattered brown flecks may be present, forming zigzag
pattern in some specimens; by this size ribs have become
prominent, secondary ribs nearly the size of primary ribs.

Mantle and Foot. Nearly retractable in shell. Cephalic ten-
tacles yellowish on inner surface. Side of foot with strongly
developed tubercles; mantle lobe broad, upper edge with
finely branched papillae, lower edge with more prominent
papillae, banded in light and dark to correspond to rays on
shell. Foot and mantle colored purplish red, lighter or tending
toward pink in individuals with lightly pigmented shells. The
reddish coloration is characteristic and differs from that of
all other species.

Habitat. Lowermost intertidal zone and immediate sub-
tidal zone to 1 5 m, living on the upper surfaces of rocks. At
low tide chiefly in deep tide pools that occur in surf exposed
areas. I obtained living specimens from the municipal market
at Iquique, evidently taken by divers. Living specimens were
collected in lesser numbers along with F. latimarginata at 5
m on a rocky bottom at Antofagasta. Large specimens were
found in shell piles at Los Molles (32°14' S). Intertidally
occurring specimens were common at Montemar and Me-
huin.

Distribution. Matarani, Peru (17°00' S) (AMNH 150892,
B. Marco), to Mehuin, Valdivia Province, Chile (39°23' S)
(LACM 75-36, McLean). The distribution probably extends
farther to the north and south. Its distribution is more south-
ern than that of F. latimarginata and F. maxima, both of
which extend much farther to the north in Peru, but seem
not to be represented at Mehuin, where F. cumingi is com-
mon.

Number of Lots Examined. 4 1 (LACM 16, AMNH 6, ANSP
1, MACN 3, NMHN 10, USNM 5).

Taxonomic History. Fissurella cumingi has not hitherto
been understood and recognized as a separate species. Pre-
vious authors have repeated the early published descriptions

Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella 31

Figures 80 through 87. Fissurella cumingi Reeve, 1849. Mature shells. (80) 17 m, Huayquique, Tarapaca Province, Chile. LACM 90799,
32.5 x 20.5 x 1 1.2 mm. (81) Los Molles, Aconcagua Province, Chile. LACM 75-28, 50.3 x 34.2 x 10.5 mm. (82) Same locality (shell pile).
LACM 75-29, 96.3 x 61.1 x 22.8 mm. (83) Lectotype, F. cumingi Reeve. Valparaiso, Chile. BMNH 197565, 67.2 x 49.1 x 15.2 mm. (84)
Montemar, Valparaiso Province, Chile. LACM 75-30, 49.9 x 34.4 x 12.8 mm. (85) Lectotype, F. stellata Reeve, 1850. 1 1-22 m, Valparaiso,
Chile. BMNH 197549, 16.2 x 10.8 x 3.8 mm. (86) Montemar, Valparaiso Province, Chile. LACM 75-30, 30.0 x 20.4 x 7.9 mm. (87) 75-
36, Mehuin, Valdivia Province, Chile. LACM 75-36, 61.2 x 42.2 x 17.7 mm.

and have considered it to be a color form or “variety” of F.
latimarginata. Confusion over the identity of F. biradiata
Reeve (a synonym of F. latimarginata) and F. bridgesii Reeve
(a good species) has also contributed to the difficulty. Cleaned
specimens found in old collections have usually been mis-
identified as F. maxima; specimens covered with the algal
mat are easily mistaken for F. latimarginata.

Abundance and Use. Fissurella cumingi is not as common
as F. maxima and F. latimarginata. but large specimens can

be found by divers, and the species is exploited for food in
central and northern Chile. I have seen it in the market at
Iquique, and it is a major component in the shell piles in
central Chile. The common name is “lapa frutilla,” meaning
strawberry, which aptly describes the color of the mantle and
foot sides.

Characteristics and Variability. The most characteristic
features of F. cumingi are its tapered front, consistent color
pattern of dark ribs on a relatively dark ground, beaded ribs,

32 Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella

Figures 88 through 94. Fissurella cumingi Reeve, 1849. Radula, cut shell, juvenile shells, living and preserved bodies, mantle lobe. (88)
Radular ribbon, air-dried. 2-5 m, Antofagasta, Chile. LACM 75-20, width of ribbon 2.8 mm. (89) Cut shell. Los Molles, Aconcagua Province,
Chile. LACM 75-29, length of cut edge 38.0 mm. (90) Juvenile shell. Los Molles, Aconcagua Province, Chile. LACM 75-28, 12.8 x 7.8 x
3.4 mm. (91) Juvenile shell. Same locality. LACM 75-28, 9.4 x 5.5 x 2.5 mm. (92) Ventral view of living specimen, head at right. Montemar,
Valparaiso Province, Chile. LACM 75-30. (93) Preserved specimen. 2-5 m, Antofagasta, Chile. LACM 75-20, shell length 73.9 mm. (94)
Mantle lobe. Same locality. LACM 75-20, length 14 mm.

and the unique reddish color of the mantle and foot. Vari-
ation in the species is minimal. The chief variable feature
seems to be the intensity of ground color, which may be pale
to dark yellow. The rayed pattern is consistent; none have
been seen that lack it. Specimens received from M. Bretos
from the subtidal mussel beds at Iquique (Fig. 80) are small,
elevated, and have narrow margins. Those from the south-
ernmost locality collected (Mehuin, Chile) appeared stunted,
and many had narrow shells. Flowever, there seem to be no
geographic differences of sufficient importance to recognize
subspecies.

Affinity and Comparisons. Fissurella cumingi has certain
features in common with both F. maxima and F. latimar-
ginata, suggesting that it is related to both. Its color pattern
combines the rayed pattern of F. maxima, with the addition
of some of the darker ground color of F. latimarginata. Its
margin shows the pattern of rays across the full, flat width,
unlike the solid margin of F. latimarginata and the rounded,
crenulate and zoned margin of F. maxima. It has the same
size and proportions as F. latimarginata, but specimens with
the algal mat may be recognized by the rayed pattern of the

margin. The normal, rayed form of F. cumingi may be dis-
tinguished from the unusual rayed form of F. latimarginata
by the strength of the ribs; they are fine and sharp in F.
latimarginata, coarser and beaded in F. cumingi. Its outline
is more tapered and its ribbing finer than that of either F.
picta lata or F. costata; it does not have the more distinct
primary ribbing of either of these species. Juvenile shells of
F. cumingi have a characteristic color pattern of split rays;
they do not have the rounded margin of F. maxima nor the
primary rays of F. costata or F. picta lata. The reddish color
of the animal is so different from that of any other species
that it may be recognized by this feature.

Synonymy and Types. There are 6 syntypes of F. cumingi
Reeve, 1849, from Quintero, Chile, BMNH 197565, lengths
77.0, 67.2, 62.5, 57.1, 32.7, and 22.5 mm. Reeve (1849) and
Dell (1971) figured the 67.2 mm shell; Dell’s interior view
is of the 77 mm specimen. The 67.3 mm specimen is here
figured and designated the lectotype (Fig. 83).

Fissurella stellata Reeve, 1850, was based on small spec-
imens from “Valparaiso, attached to dead shells at a depth
of from six to twelve fathoms.” There are four syntypes,

Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella 33

BMNH 197549, lengths 16.9, 16.2, 15.0, and 14.5 mm. The
16.2 mm specimen is illustrated and designated the lectotype
(Fig. 85). These specimens are clearly the juveniles of F.
cumingi; they are red-rayed, the ribbing of medium strength
and nodular, the primary and secondary ribs nearly equiv-
alent. The original specified depth is probably accurate; the
species is known from the shallow sublittoral. Other authors
have guessed incorrectly with respect to this name; Riveros-
Zufiiga (1951) figured the brown and white rayed form of F.
peruviana as F. stellata. Dell (1971) mixed the figure numbers
on his plate 4: his figure of a syntype of F. stellata should be
figs. 7, 8, rather than 5, 6.

Fissurella costata Lesson, 1831

Figures 95-108

Fissurella rudis Deshayes, 1830:134; Deshayes in Lamarck,
1836:61; Orbigny, 1841:474 [under F. costata]. Not Pa-
tella rudis Roeding, 1798 [= F. nodosa Bom, 1778],
Fissurella costata Lesson, 1831:41; Sowerby, 1835b:4, fig.
28; Orbigny, 1841:474; Reeve, 1849, pi. 2, fig. 14; Hupe,
1854:243; Philippi, 1860:181; Sowerby II, 1862:187, figs.
15, 205; Pilsbry, 1890:148, pi. 30, fig. 10, pi. 35, fig. 11;
Dali, 1909:177, 241; Ziegenhom and Thiem, 1925:14, pi.
2, fig. 12; Carcelles and Williamson, 1951:255; Riveros-
Zuniga, 1951:108, fig. 21; Dell, 1971:183, pi. 4, figs. 7, 8;
Marincovich, 1973:16, fig. 28; Ramirez-Boehme, 1974:31
[key].

Fissurella chilensis Sowerby, 1835a: 124; Sowerby, 1835b:3,
fig. 36; Orbigny, 1841:474 [under F. costata],

Fissurella costata var. rubra Ziegenhom and Thiem, 1925:
14, pi. 2, figs. 13a, b.

Shell. Medium sized (50 to 80 mm mature length), low to
medium; outline elongate in young shells, broadly oval in
mature shells; base of shell resting flat in one plane. Sculpture
of distinctly raised narrow ribs that crenulate margin; inter-
spaces narrower than ribs. Primary ribs more prominent in
young shells, but in large shells not differing from secondary
ribs. Ground color light yellowish gray; rays gray, often be-
coming faint in later growth stages. Margin broad, flat, show-
ing gray rays across full width but more distinctly at outer
edge. Calcitic layer of cut shells not zoned; rays showing
through full width. Foramen exceptionally small, elongate
and tripartite in young shells, elongate-oval in mature shells.

Juvenile Shell. Conical at earliest stage, becoming flattened
and elongate; primary ribs light yellow, strongly elevated,
becoming nodular after shell reaches length of 5 mm; earliest
rays reddish, changing to black by shell length of 4 mm; black
rays fill interpsaces between primary ribs. With growth, sec-
ondary ribs appear between primary ribs and black rays be-
come gray or disappear altogether.

Mantle and Foot. Fully retractable in shell; shell margin
not raised and mantle not projecting in living specimens.
Cephalic tentacles dark on outer side, yellowish on inside
and at tips. Mantle lobe very narrow, papillae of both edges
small and width little branching. Side of foot light pinkish
brown, surface marbled. Projecting tubercles are not readily
apparent in living specimens but visible in preserved spec-

imens. Elaboration of the mantle lobes is the least pro-
nounced in this species.

Habitat. In northern Chile at Iquique and Antofagasta
Fissurella costata occurs on vertical surfaces of surf-exposed
rock walls at the low tide line, near the holdfasts of the large
brown alga Lessonia, where its habitat is not shared by other
species of Fissurella. In central Chile I found it more abun-
dantly. At Los Molles and Montemar it also occurs on hor-
izontal surfaces in less exposed areas; here it shares the hab-
itat with other species. The shell margin fits the contours of
a habitual site of attachment; when exposed at low tide the
shell margin is in tight contact with the substrate. Unlike the
other species, individuals do not move when touched by
Fleliaster; the small foramen may protect them from access
by this seastar. The shell also seems suited to resist removal
by the clingfish Sicyases because the margin is less raised
that that of other species, which are more subject to predation
by Sicyases. Specimens are rarely seen with attached Scurria
parasitica.

Distribution. Punta Pichalo, Tarapaca Province (19°36' S)
(AMNH 137232, J. Bird), to Guabun, Isla de Chiloe, Chile
(41°50' S) (LACM 75-40, McLean). Dali’s (1909) record from
Mollendo, Peru, was based on a specimen of F. maxima
(USNM 27743). However, the distribution of this species
may extend farther to the north into Peru. In my collecting
only the northwestern tip of Isla de Chiloe was sampled at
Guabun; the southern limit of the species is therefore not
certain. It is apparent, however, that this species has a more
southern center of distribution than most species of the Pe-
ruvian Faunal Province.

Number of Lots Examined. 70 (LACM 23, AMNH 15,
ANSP 3, MACM 9, MNHN 12, USNM 8).

Taxonomic History. Fissurella costata has been reasonably
well understood by recent authors.

Abundance and Use. Fissurella costata is moderately com-
mon throughout its range. In northern Chile it is little used
for food because its habitat is so exposed to surf that it is
usually inaccessible to shore collectors. Beach-worn shells
are common along the shore but fresh shells are seldom seen
in the shell piles. The common name is “lapa senorita,”
because of its resemblance to Scurria viridu/a, which is known
as the “senorita.”

Characteristics and Variability. Fissurella costata is char-
acterized by its relatively small foramen, which is broadly
oval when mature and narrow in young stages, its relatively
low height, and its consistent color pattern of gray rays on a
yellow ground. Its mantle lobe is very narrow, and the foot
color is light pinkish brown. Variations in height and outline
are minor; color variation results from rays that either persist
or fade in later stages. There are no geographic differences
worthy of note. The low variability of this species is corre-
lated with its rather restricted habitat on surf-exposed rocks
in the lower intertidal zone.

Affinity and Comparisons. Fissurella costata seems to be
most closely related to, and difficult to distinguish from, F.
picta lata. Both have similar color patterns, a similar, broadly
oval outline, strong ribs, and primary ribs that remain strong.
Differences are that mantle and foot colors are lighter in F.

34 Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella

Figures 95 through 101. Fissurella costata Lesson, 1831. Mature shells. (95) Punta Jara, Antofagasta Province, Chile. LACM 75-18, 72.3 x
59.0 x 20.4 mm. (96) Los Molles, Aconcagua Province, Chile. LACM 75-28, 66.8 x 54.5 x 23.3 mm. (97) Islota Concon, Valparaiso Province,
Chile. LACM 75-31, 43.3 x 31.8 x 9.9 mm. (98) Lectotype, F. rudis Deshayes. Paita, Peru (probable error). MNHNP, 69.1 x 61.1 x 24.7
mm. (99) Rio Bio-bio, Concepcion Province, Chile. LACM 75-35, 57.7 x 56.4 x 15.3 mm (beach shell). (100) Mehuin, Valdivia Province,
Chile. LACM 33-2, 33.2 x 25.7 x 9.4 mm. (101) Ancud, Chiloe Province, Chile. LACM 62-62, 35.5 x 25.5 x 7.9 mm.

Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella 35

Figures 102 through 108. Fissurella costata Lesson, 1931. Living specimens, juvenile shells, radula, and cut shell. (102) Ventral view, living
specimen. Punta Jara, Antofagasta Province, Chile. LACM 75-18. (103) Living specimens in place. Los Molles, Aconcagua Province, Chile.
LACM 75-28. (104) Juvenile specimen. Same locality. LACM 75-28, 14.2 x 7.9 x 3.1 mm. (105) Juvenile specimen (dead shell). Bahia
Herradura, Coquimbo Province, Chile. LACM 75-25, 10.9 x 7.3 x 2.4 mm. (106) Radula of small specimen. Montemar, Valparaiso Province,
Chile. LACM 75-30, width of field 0.8 mm, shell length 24.5 mm. (107) Mantle lobe. Punta Jara, Antofagasta Province, Chile. LACM 75-
18, length 10 mm. (108) Cut shell. Los Molles, Aconcagua Province, Chile. LACM 75-28, length of cut edge 31 mm.

costata; F. costata is somewhat lower than F. picta lata, al-
though the extremes of variation seem to overlap; F. costata
does not have the tendency to reddish rays nor the penciled
pattern of the rays of F. picta lata. Juveniles of F. costata
are more elongate, flatter, and have more nodulous primary
ribs than those of F. picta lata. In northern Chile where F.
picta lata does not occur, there is no difficulty in recognizing
F. costata as the species that is the most broadly oval and
has the smallest foramen.

Synonymy and Types. Four syntypes of F. rudis Deshayes,
1830, have been examined, received on loan from the Paris
Museum. The lot is accompanied by a cardboard mount; the
shells are 75.8, 69. 1, 53.6, and 5 1.9 mm in length. The largest
of these shells is the polished specimen mentioned in the
original description. The original dimensions were given as

55 mm length and 43 mm width; one of the specimens is
53.6 mm long and 42.8 mm wide. The 69.1 mm shell is
designated the lectotype (Fig. 98). All four specimens are
typical and agree with the thorough and accurate original
description. The original locality was Paita, Peru: “This shell
was first given to us by our friend Lesson, who found it at
Paita, Peru; then later we have found it commercially, like-
wise from Peru.” [Translation.] Later, Deshayes in Lamarck
(1836) changed the locality to “Habite le Chile.” The actual
occurrence of the species in Peru is uncertain, and Paita is
north of the known occurrence of any of the Peruvian species
of Fissurella. This name is preoccupied by Patella rudis Roe-
ding, 1798, a synonym of the Caribbean F. nodosa (Bom,
1778).

Type material of Fissurella costata Lesson, 1831, has never

36 Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella

been illustrated, and specimens are not in the collection of
the Paris Museum, where some of Lesson’s types are known
(Bouchet, personal communication). Lesson’s description is
insufficient to distinguish the species from F. picta lata , which
is closely related and also abundant in the vicinity of Tal-
cahuano, Chile, the type locality. Lesson described the animal
as blackish and the shell margin as bluish with stains of red,
features that would more appropriately apply to F. picta lata.
However, the species is well known under the name F. costata
and no purpose would be served in rejecting this name, even
though there is doubt as to its identity.

Type material for F. chilensis Sowerby, 1 835, has not been
located. It came from “Valparaiso, found on rocks in exposed
situations at low water.” The habitat is accurately described;
F. costata is much more common than F. picta lata at Val-
paraiso, and the original illustration shows a densely ribbed
shell with a very small foramen. Its identity is certain, in
agreement with treatment by previous authors.

Type material of Fissurella costata var. rubra Ziegenhom
and Thiem, 1925, has not been located. The specimen came
from Coquimbo, Chile, and measured 60 x 48 x 15 mm,
distinguished from the typical form in having reddish rays.
Judging from the illustration, it could be either F. costata or
F. picta lata, although I have not seen reddish rayed speci-
mens of F. costata. Lateral profile of the figure is about right
for F. costata; for F. picta lata it would be at the low extreme
of variation; the locality is reasonable for F. costata; to my
knowledge F. picta lata does not occur north of Valparaiso,
where it is uncommon. I therefore favor retaining this taxon
in the synonymy of F. costata, admitting that the other al-
ternative is a possibility.

Fissurella picta ( Gmelin, 1791)

Figures 109-146

Fissurella picta is here considered to have two geographic
subspecies: F. picta picta in the Magellanic region of Chile
and F. picta lata in south-central Chile.

Synonymy for F. picta picta:

Patella picta Gmelin, 1791:3729.

Fissurella picta, Lamarck, 1822:10; Deshayes, 1830:131;
Sowerby, 1 835b: 1 , figs. 4, 26; Lamarck, 1836:559; Orbi-
gny, 1841:472; Reeve, 1849, pi. 1, fig. 6; Hupe, 1854:237;
Sowerby II, 1862:186, figs. 10, 11, 35; Watson, 1886:33;
Rochebrune and Mabille, 1889:70; Pilsbry, 1890:144, pi.
45, figs. 9-11; Melvill and Standen, 1898:102; Strebel,
1907:83, pi. 2, fig. 22; Melvill and Standen, 1907:98; Stre-
bel, 1908:79; Dali, 1909:242; Melvill and Standen, 1914:
115; Ziegenhom and Thiem, 1925:6, pi. 1, figs. 1-4; Car-
celles, 1950:51; Powell, 1951:85; Carcelles and William-
son, 195 1:254; Riveros-Zuniga, 1951:96, fig. 15; Dell, 1971:
191; Ramirez-Boehme, 1974:31 [key].

Fissurella atrata Reeve, 1850, pi. 11, fig. 73; Sowerby II,
1862:186, fig. 71; Pilsbry, 1890:147, pi. 34, fig. 59 [under
F. philippiana\\ Dell, 1971:190, pi. 3, fig. 7 [under F. phi-
lippiana ].

Fissurella muricata Reeve, 1850, pi. 14, fig. 103; Sowerby

II, 1862:106, pi. 4, fig. 68; Pilsbry, 1890:156, pi. 39, fig.

5.

Synonymy for F. picta lata:

Fissurella lata Sowerby, 1835a: 124; Sowerby, 1835b:3, fig.

63; Reeve, 1849, pi. 1, fig. 5; Hupe, 1854:243; Sowerby

II, 1 862: 1 87, fig. 1 3; Pilsbry, 1 890: 1 47, pi. 3 1 , figs. 18,19;

Dali, 1909:241; Ziegenhom and Thiem, 1925:13, pi. 1 , fig.

1 1; Carcelles and Williamson, 1951:255; Riveros-Zuniga,

1951:107; Dell, 1971:187, pi. 4, figs. 12-14; Ramirez-

Boehme, 1974:31 [key].

Fissurella navidensis Ramirez-Boehme, 1974:17, 31 [key].

Shell (F. picta picta). Medium large (65-95 mm mature
length), moderately elevated; outline elongate oval, sides of
shell slightly raised. Sculpture of sharp, narrow radial ribs;
primary ribs remaining stronger than secondary and tertiary
ribs; occasional specimens with weak ribs. Ground color white,
sometimes gray, rayed with black and white; strongest ribs
centered on both light and dark rays; dark rays usually split
into number of fine lines of black, a result of lack of pigment
in grooves between fine ribs. Rays faint on light ground in
some, or black on gray ground and with rays interrupted to
produce concentric patterns of banding. Margin broad and
flat, showing penciled pattern of rays. Cut shells show that
pigment of rays extends through calcitic layer. Foramen elon-
gate in juvenile shells, elongate to oval in mature shells.

Shell ( F . picta lata). Medium large (50-80 mm mature
length), height moderately to strongly elevated; outline elon-
gate-oval to oval, sides of shell slightly raised. Sculpture of
strong radial ribs; primary ribs remaining strong at all growth
stages. Ground color yellowish white, rayed with yellowish
purple; strongest ribs centered on both light and dark rays;
dark rays usually split into number of fine lines coinciding
with secondary and tertiary ribs; pigment tending to be absent
in grooves between ribs. Color pattern fairly uniform, with
dark rays on lighter ground. Margin broad and flat in young
shells, sharply defined, gray, showing pattern of rays and
penciled pattern. Cut shells showing pigment of rays through-
out calcitic layer. Foramen elongate in young shells, oval in
mature shells.

Juvenile Shell. Juveniles of both subspecies are oval and
high, margin broad; strong primary ribs coincide with light
rays, becoming stronger and slightly nodulous in specimens
longer than 5 mm. Secondary ribs develop in interspaces
between primary ribs; dark rays develop in rib interspaces
and show some concentric interruptions in intensity.

Mantle and Foot. Body nearly retractable within shell; ce-
phalic tentacles dark on outer side, reddish on inner side,
and yellowish at tips. Mantle lobe relatively narrow, banded
to correspond to pattern of rays; papillae of both edges of
mantle lobe moderately developed. Side of foot marbled with
light and dark, tubercles lighter tipped; southernmost spec-
imens often lighter overall.

Habitat. Mid-tidal to lower intertidal zone on vertical to
horizontal surfaces and on the sides of loose boulders; wedged
in crevices under more exposed conditions; not extending
into the sublittoral zone. At the north end of the range in the

Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella 37

Figures 109 through 115. Fissurella picta lata Sowerby, 1835. Mature shells. (109) Montemar, Valparaiso Province, Chile. LACM 75-30,
37.9 x 27.8 x 13.4 mm. (110) Holotype, F. navidensis Ramirez-Boehme. Bahia Navidad, Santiago Province, Chile. MNHN 200376, 66 x
51 x 22 mm. (Ill) Rio Bio-bio, Concepcion Province, Chile. LACM 75-35, 24.8 x 17.6 x 8.2 mm. (112) Lectotype, F. lata Sowerby. Isla
Santa Maria, Bahia Concepcion, Chile. BMNH 197571, 83.4 x 64.3 x 31.6 mm. (113) Rio Bio-bio, Concepcion Province, Chile. LACM 75-

35, 37.9 x 28.7 x 8.8 mm. (114) Mehuin, Valdivia Province, Chile. LACM 75-36, 40.4 x 29.2 x 11.7 mm. (115) Same locality. LACM 75-

36, 40.4 x 26.5 x 12.5 mm.

vicinity of Valparaiso, F. picta lata is uncommon; the only
two living specimens that I found occurred on horizontal
surfaces adjacent to F. limbata. At Concepcion and Mehuin
and in the Strait of Magellan where F. picta picta occurs,
specimens were common under all conditions of exposure.
At these localities, such characteristic northern species as F.
latimarginata and F. maxima were scarce or missing, sug-
gesting a correlation between the dominance of F. picta lata
and absence of competition from the other species.

Distribution. Islote Concon, Valparaiso Province, Chile
(32°52' S) (LACM 75-31, McLean), to Tierra del Fuego and
Isla de los Estados, Argentina, probably south to Cape Horn
and east to the Falkland Islands. Fossil specimens are known
from shoreline terraces at Comodora Rivadavia, Chubut
Province, Argentina (45°52' S) (MCZ 28329), but living spec-

imens are unknown on the mainland Patagonian coast north
of Tierra del Fuego. The subspecies F. picta picta occurs in
the Magellanic region of Chile, extending north to the vicinity
of Isla de Chiloe. Populations that occur in the area of over-
lap, chiefly in the vicinity of Isla de Chiloe, are consistent
but may have features that make assignment to either sub-
species arbitrary (see further discussion below).

Number of Lots Examined. F. picta picta: 95 (LACM 19,
AMNH 15, ANSP 5, MACN 33, MNHN 10, USNM 13).
F. picta lata: 64 (LACM 11, AMNH 7, ANSP 2, MACN 5,
MNHN 35, USNM 4).

Taxonomic History. The typical form of Fissurella picta
from the Strait of Magellan has been well known and under-
stood by all authors. Juvenile specimens have been given the
name F. atrata Reeve, but it has otherwise not been burdened

38 Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella

Figures 116 through 122. Fissurella picta lata Sowerby, 1835. Bodies of living and preserved specimens, juvenile shells, radula, mantle lobe,
and cut shell. (116) Living specimen, ventral view, head at right. Rio Bio-bio, Concepcion Province, Chile. LACM 75-35. (117) Juvenile shell.
Guabun, Chiloe Province, Chile. LACM 75-40, 12.6 x 8.6 x 3.3 mm. (118) Radula of small specimen. Rio Bio-bio, Concepcion Province,
Chile. LACM 75-35, width of field 0.9 mm, shell length 27.3 mm. (119) Juvenile shell. Same locality. LACM 75-35, 13.5 x 8.3 x 4.3 mm.
(120) Preserved specimen. Same locality. LACM 75-35, shell length 59.5 mm. (121) Mantle lobe. Island off Mehuin, Valdivia Province, Chile.
LACM 75-37, length 8 mm. (122) Cut shell. Same locality. LACM 75-37, length of cut edge 32 mm.

with excessive numbers of synonyms, as have the other two
common southern species, F. radiosa and F. oriens. Sow-
erby’s Fissurella lata has not previously been a well under-
stood taxon, perhaps because it is rare in the most populated
region of central Chile. It is here for the first time regarded
as a northern subspecies of the well-known F. picta. The
original description of F. lata included the remark: “This
species approaches, in form and colouring, very near to Fiss.
picta, Lam.,’’ an accurate observation not noticed by sub-
sequent authors. In his discussion of this taxon, Riveros-
Zuniga merely quoted previous authors and figured what is
more likely to be a specimen of F. costata. However, the
Ziegenhom and Thiem (1925) figure is a good representation
of F. picta lata.

Abundance and Use. Both subspecies are large enough and
common enough, at least at Concepcion and to the south, to
be important as a food resource. Numerous specimens were
seen in shell piles at Mehuin. Moreno et al. (1984) have given
an account of the fishery and ecology of this species at Me-
huin. I have no information on the utilization of the species
at more southern localities.

Characteristics and Variability. The typical F. picta picta
is large-shelled, with coarse radial ribs and primary ribs that
remain strong; the margin is broad; the dark colored rays are
split into numerous fine lines by grooves that lack pigment
and separate the fine ribs. Variation is extensive, chiefly in
strength of ribbing and color pattern. Variations in outline
of the base are unusual; some oval shells have been seen
(Figs. 127, 128). Sculpture varies from coarse to nearly
smooth. Most specimens are colored with gray rays on a
white ground; variants with dark ground color are common.
There are frequent concentric bands of different color inten-
sity, probably representing seasonal changes in temperature
and food availability.

Fissurella picta lata has a broad outline, as the name im-
plies. Like F. picta picta it is also characterized as a large-
shelled form with coarse ribbing and primary ribs that remain
strong; the color rays are also split into thin lines by deep
grooves in which the color is lacking. The margin is broad
at all growth stages, and the mantle lobe papillae and foot
tubercles are developed as in F. picta picta. As in the typical
subspecies, there is extensive variation in F. picta lata. Al-

Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella 39

Figures 123 through 129. Fissurella picta picta Gmelin, 1791. Mature shells. (123) Pargua, Llanquihue Province, Chile. LACM 75-39, 77.2
x 49.3 x 23. 1 mm. (124) Pumalin, Chiloe Province, Chile. LACM 75-41 , 24.5 x 16.9 x 6.5 mm. (125) Same locality. LACM 75-41, 59.6 x
43.5 x 24.5 mm. (126) Islota Nihuel, Chiloe Province, Chile. LACM 75-44, 39.8 x 26.4 x 16.4 mm. (127) Quellon, Chiloe Province, Chile.
LACM 75-45, 83.5 x 68.8 x 36.8 mm. (128) Pumalin, Chiloe Province, Chile. LACM 75-41, 48.4 x 39.6 x 18.2 mm. (129) Holotype, F.
muricata Reeve. Locality unknown. BMNH 1976144, 25.8 x 18.4 x 13.0 mm.

though most specimens are elevated, the height is variable,
and low forms occur in some populations along with more
elevated specimens. Ground color ranges from light to dark
gray; the rays are usually darker than the ground color but
in some cases only slightly darker.

The major difference between the two subspecies is that
F. picta lata has a more oval outline and usually is more
elevated than F. picta picta. However, these are variable
features within the species as a whole, and specimens occur
at either extreme of the distribution having proportions typ-
ical of the other extreme. Specimens from the area of overlap.

however, are more likely to have the intermediate propor-
tions. Another difference is that of the coloration of the rays:
Fissurella picta picta has dark purple to gray rays, whereas
F. picta lata has rays that more clearly show the purple or
reddish coloration. In both subspecies, however, the rays
change color to reddish if the shells are faded by exposure
to the sun. Shells from shell piles along the shore are much
redder than those of live-collected or beach-worn specimens.

The extreme differences noted in some populations around
Isla de Chiloe seem to be correlated with conditions of ex-
posure to the open coastline on the west, or to a more pro-

40 Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella

Figures 130 through 135. Fissurella picta picta Gmelin, 1791. Mature shells. (130) Puerto el Hambre, Magallanes Province, Chile. LACM
75-49, 58.2 x 40.4 x 18.3 mm. (131) Holotype, F. alrala Reeve. Locality unknown. BMNH 197564, 23.8 x 14.0 x 5.0 mm. (132) Puerto
el Hambre, Magallanes Province, Chile. LACM 75-49, 86.3 x 58.3 x 28.3 mm. (133) Same locality. LACM 75-49, 49.2 x 34.3 x 13.7 mm.
(134) Laredo Bay, Magallanes Province, Chile. USNM 1 18235, 68.6 x 61.7 x 30.0 mm. (135) Fox Bay, Falkland Islands. LACM 90800,
43.3 x 27.3 x 11.5 mm.

tected habitat along the channels to the east, where tidal
extremes are greater than those of the open coast and water
movement is produced by tidal currents. At Guabun, on the
northwest, exposed side of the island (41°50' S, 74°02' W),
the typical purple-rayed, oval, elevated form of F. picta lata
occurs. Not far away at Pargua, on the mainland side of the
Canal de Chacao (41°47' S, 73°28' W), I found the gray-rayed,
low, elongate form typical of F. picta picta (Fig. 123). At
Pumalin on the mainland opposite the southern tip of Isla
de Chiloe (42°42' S, 72°52' W), the population was inter-
mediate between the two extremes, more oval than typical
for F. picta picta, but having no trace of the reddish rays
(Fig. 125). At Isla Laitec off the southeast end of Isla de
Chiloe (43° 1 2' S, 7 3°36' W), shells were proportionately more
elongate, typical of F. picta picta. Based on these four pop-
ulations, I consider that F. picta lata occurs south along the
outer coast of Isla de Chiloe and F. picta picta occurs to the
north along the inner side of Chiloe Island. I do not know,
however, whether populations from the exposed sides of is-

lands to the south of Chiloe Island would also agree with F.
picta lata.

Affinity and Comparisons. Fissurella picta picta most re-
sembles F. radiosa, a smaller-shelled species that is similar
in overall proportion, has sculpture of strong ribbing with
persistently strong primary ribs, and has a similar color pat-
tern. However, F. picta picta reaches nearly twice the size of
F. radiosa, has a much broader margin at all growth stages,
and has a more centrally placed foramen. The penciled effect
of the dark rays of F. picta picta is not seen in F. radiosa.
Fissurella picta picta does not occur sympatrically with any
of the large species of northern Chile. It differs from all of
them sufficiently to require no comparison.

The subspecies F. picta lata does occur sympatrically with
many of the other species. The latter subspecies is most closely
related to, and difficult to distinguish from, F. costata. Fis-
surella picta lata tends to be higher, more darkly rayed, and
to have sculpture with stronger primary ribs and a slightly
larger foramen than that of F. costata. Separation of the two

Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella 41

Figures 136 through 146. Fissurella picta picta Gmelin, 1791. Living and preserved specimens, radulae, juvenile shells, and cut shells. (136)
Ventral view of living specimen. Pumalin, Chiloe Province, Chile. LACM 75-41. (137) Mantle lobe. Puerto el Hambre, Magallanes Province,
Chile. LACM 75-49, length 12 mm. (138) Air-dried radula. Isla Laitec, Chiloe Province, Chile. LACM 75-47, width of ribbon 1.9 mm, shell
length 54.3 mm. (139) Preserved specimen. Isla de Los Estados, Argentina. LACM 71-284, shell length 71.8 mm. (140) Juvenile specimen.
Pumalin, Chiloe Province, Chile. LACM 75-41, 10.9 x 6.7 x 3.6 mm. (141) Juvenile specimen. Puerto Espanol, Bahia Aguirre, Tierra del
Fuego, Argentina. LACM 73-67, 17.4 x 10.6 x 3.6 mm. (142) SEM photo of radula. Width of field 1.0 mm. (143) Radula of small specimen.
Isla Laitec, Chiloe Province, Chile. LACM 75-47, width of field 0.7 mm, shell length 27.9 mm. (144) Cut shell. Pumalin, Gulfo Corcovado,
Chile. LACM 75-41, length of cut 37 mm. (145) Cut shell. Puerto el Hambre, Magallanes Province, Chile. LACM 75-49, length of cut 33.2
mm. (146) Living specimen attached to substrate. Same locality. LACM 75-49.

42 Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella

species on shell characters may be difficult and sometimes
entirely arbitrary, as the range of variation in the two species
seems to overlap. It is easy to distinguish the living animals,
however. The foot and mantle lobe of F. picta lata are gray,
those of F. cost at a a pale pinkish brown.

Synonymy and Types. Fissurel/a picta was known to pre-
Linnaean authors. It was probably the first of the Chilean
species to reach Europe because it is so common in the Strait
of Magellan. Gmelin (1791) is credited with the name; his
knowledge of it came from figures in several previous non-
binomial works. Lamarck and Deshayes referred to it as the
“Fissurelle de Magellan.” Deshayes credited the name picta
to Lamarck, but Sowerby II (1862) and Pilsbry (1890) cor-
rectly credited the authorship to Gmelin. Type material is
unknown.

The holotype of Fissurella atrata Reeve, 1850, is BMNH
197564 (Fig. 131), length 23.8 mm, locality unknown. It is
a small, dark-colored specimen of F. picta, easily recognized
by its relatively broad, dark inner margin.

The holotype of F. muricata Reeve is BMNH 1976144,
locality unknown (Fig. 129). It is a small dark specimen, the
margin sufficiently broad to relate it to F. picta rather than
F. radiosa. Because the interior has not previously been fig-
ured, the dark margin has been missed, which explains why
no author has related it to any of the Peruvian-Magellanic
species.

There are two syntypes of F. lata Sowerby, BMNH 197571,
from Isla Santa Maria, Bahia Concepcion, Chile, dimensions
83.4 mm x 64.3 x 31.6 mm, and 77.0 x 61.3 x 25.5 mm.
The larger specimen, figured by Dell (1971), is figured here
and designated the lectotype (Fig. 1 1 2). Both specimens are
reddish rayed and clearly show the strong primary ribs in
the light interspaces between the rays.

The holotype of F. navidensis Ramirez-Boehme, 1974,
from Bahia Navidad, Santiago Province, Chile (33°56' S,
71°52' W), MNHN 200376 (Fig. 1 10), is a worn specimen
of F. picta lata, with which it was not compared. Its similarity
to F. picta was noted by its author, however, and the differ-
ences described are those that are here used to distinguish
the two subspecies of F. picta.

Fissurella radiosa Lesson, 1831

Figures 147-175

Fissurella radiosa is here considered to have two geographic
subspecies: F. radiosa radiosa in the Magellanic region of
Chile and Argentina, and F. radiosa tixierae in the Golfo
San Matias and Peninsula Valdez region of Argentina.

Synonymy for F. radiosa radiosa:

Fissurella radiosa Lesson, 1831:411; Orbigny, 1841:473;

Pilsbry, 1890:157; Strebel, 1907:85, pi. 1, figs. 4, 5a-d, pi.

9, fig. 6; Melvill and Standen, 1914:115; Carcelles, 1950:

51; Carcelles and Williamson, 1951:254; Riveros-Zuniga,

1951:111; Dell, 1971:1 92; Ramirez-Boehme, 1974:32 [key];

Scarabino, 1977:178, pi. 1, fig. 5.

Fissurella picta var. radiosa, Ziegenhom and Thiem, 1925:

8, 11, pi. 1, fig. 6.

Fissurella nigra Philippi, 1845:60; Philippi, 1846, pi. 2, fig.

22; Reeve, 1849, pi. 6, fig. 37. Not F. nigra Lesson, 1831.

Fissurella darwinii Reeve, 1849, pi. 1. fig. 7; Hupe, 1854:
247; Rochebrune and Mabille, 1 889:74; Pilsbry, 1890:144,
pi. 30, fig. 7, pi. 46, figs. 15-17; Melvill and Standen, 1 898:
102; Strebel, 1907:93; Carcelles, 1950:51; Carcelles and
Williamson, 1951:254; Riveros-Zuniga, 1951:98, fig. 16;
Dell, 1971:185, pi. 4, fig. 4; Ramirez-Boehme, 1974:30
[key].

Fissurella picta var. darwinii, Ziegenhom and Thiem, 1925:
8, 11, pi. 1, fig. 5.

Fissurella grisea Reeve, 1849, pi. 6, fig. 6; Sowerby II, 1862:
184, pi. 239, fig. 85; Pilsbry, 1890:152, pi. 39, fig. 9.
Fissurella exquisita Reeve, 1850, pi. 11, fig. 74; Hupe, 1854:
246; Sowerby II, 1862:186, figs. 32, 128; Rochebrune and
Mabille, 1889:74; Strebel, 1908:78, pi. 5, figs. 74a-c; Car-
celles and Williamson, 1951:256; Riveros-Zuniga, 1951:
112, fig. 22; Metivier, 1969:115, fig. IB [radula]; Dell,
1971:185, pi. 5, figs. 1, 3; Ramirez-Boehme, 1974:30.
Fissurella philippiana Reeve, 1850, errata page; Sowerby II,
1862:186, fig. 30; Pilsbry, 1890:146, pi. 33, fig. 40, pi. 58,
figs. 24-26; Dali, 1909:242; Ziegenhom and Thiem, 1925:
13, pi. 1, figs. 10a, 10b; Carcelles and Williamson, 1951:
253; Riveros-Zuniga, 1951:106, fig. 19; Dell, 1971:190.
Fissurella philippii Hupe, 1854:245 (new name for F. nigra
Philippi, not Lesson).

Fissurella polygona Sowerby II, 1862, fig. 177 (not fig. 137);
Pilsbry, 1890:148, pi. 60, fig. 84; Melvill and Standen,
1898:102; Melvill and Standen, 1914:115; Carcelles and
Williamson, 1951:254; Dell, 1971:192, pi. 4, figs. 9-1 1.
Fissurella dozei Rochebrune and Mabille, 1885:108; Roche-
brune and Mabille, 1889:72, pi. 5, fig. 4; Carcelles, 1950:
51; Carcelles and Williamson, 1951:255; Riveros-Zuniga,
1951:101, fig. 17; Dell, 1971:185.

Synonymy for F. radiosa tixierae:

Fissurella tixierae Metivier, 1969: 1 16, fig. 1 A [radula], pi. 1,
figs. 1-3, 9.

Shell (F. radiosa radiosa). Small to medium sized (40 to
55 mm mature length), low to moderately elevated; outline
elongate oval, somewhat tapered anteriorly; sides of shell
raised. Sculpture of sharply raised, narrow ribs that crenulate
margin; primary ribs remaining stronger and more promi-
nent at all growth stages. Ground color varying from white
to gray or black with gray or reddish gray rays; the pattern
of rays frequently interrupted by concentric changes in color
intensity. Primary ribs coinciding with light rays; in uni-
formly dark shells primary ribs slightly lighter in color. Mar-
gin relatively narrow at all growth stages, not zoned; cut shells
showing ground color or pattern of rays of uniform intensity
throughout calcitic layer. Foramen just anterior of center,
elongate and tripartite at all growth stages.

Shell (F. radiosa tixierae). Small (20 to 45 mm mature
length), moderately to strongly elevated; outline elongate ob-
long, tapered anteriorly; margin more or less in same plane.
Sculpture of narrow ribs that finely crenulate margin. Ground
color white to dark gray or black with gray or reddish gray
rays, often interrupted by concentric changes in intensity.
Primary ribs coinciding with light rays. Margin narrow at all
growth stages, showing pattern of rays. Foramen slightly an-
terior of center, elongate and tripartite at all growth stages.

Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella 43

Figures 147 through 160. Fissurella radiosa radiosa Lesson, 1831. Mature shells. (147) Pumalin, Chiloe Province, Chile. LACM 75-41, 47.0
x 32.0 x 13.1 mm. (148) Same locality. LACM 75-41, 41.9 x 27.4 x 11.8 mm. (149) Same locality. LACM 75-41, 43.5 x 28.4 x 13.7 mm.
(150) “Syntype” [no standing as type] F. phdippiana Reeve. “Chile." BMNH 197562, 42.0 x 22.8 x 8.6. (151) Lectotype, F. darwinii Reeve.
Strait of Magellan, Chile. BMNH 197563, 36.9 x 22.8 x 11.0 mm. (152) Lectotype, F. grisea Reeve. Locality unknown. BMNH 1975140,
30.3 x 19.9 x 9.9 mm. (153) Lectotype, F. exquisita Reeve. Locality uncertain. BMNH 197561, 23.4 x 14.2 x 5.2 mm. (154) Puerto el
Hambre, Magallanes Province, Chile. LACM 75-49, 40.2 x 23.0 x 10.6 mm. (155) Same locality. LACM 75-49, 38.2 x 21.4 x 8.2 mm.
(156) Falkland Islands. USNM 368377, 46.7 x 20.1 x 9.4 mm. (157) Lectotype, F. polygona Sowerby II. Falkland Islands. BMNH 1976151,

44 Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella

Figures 161 through 166. Fissurella radiosa (ixierae Metivier, 1969. Mature shells. (161) San Antonio Oeste, Rio Negro Province, Argentina.
MACN 13361, 48.3 x 31.3 x 1 4.3 mm. (162) Puerto Lobos, Chubut Province, Argentina. MCZ 288334, 24.6 x 14.3 x 8.3 mm. (163)
Holotype, F. tixierae Metivier. Golfo Nuevo, Chubut Province, Argentina. MNHNP, 22.5 x 13.7 x 8.0 mm (beach shell). (164) Puerto
Madryn, Golfo Nuevo, Chubut Province Argentina. LACM 34858, 24.9 x 14.9 x 8.5 mm (beach shell). (165) Punta Cracker, Golfo Nuevo,
Chubut Province, Argentina. LACM 78-90, 27.0 x 18.8 x 8.8 mm. (166) Puerto Madryn, Golfo Nuevo, Chubut Province, Argentina. USNM
152895, 25.5 x 15.6 x 10.6 mm (beach shell).

Juvenile Shell. Elongate, margin narrow, ribs fine and sharp,
primary ribs lighter in color, secondary and tertiary ribs aris-
ing after shell reaches 5 mm in length.

Mantle and Foot. Nearly retractable within shell. Mantle
lobe relatively narrow, banded to correspond to pattern of
rays; black-shelled individuals also banded. Papillae mod-
erately developed, finely branched; side of foot dark, tuber-
cles with lighter tips.

Habitat. Lowermost intertidal zone and offshore to at least
20 m. In 1975 I found it common at Pumalin in the Golfo
Corcovado on the undersides of rocks in an area where the
exposure is limited chiefly to swiftly moving tidal currents;
I also observed it in the sublittoral at Isla 'Falcon. In the

Strait of Magellan it was common at low tide under rocks at
Puerto Hambre. Paul Dayton collected it by diving at Isla
de los Estados in 1973. In 1978 I found F. radiosa tixierae
to be common in the Golfo Nuevo and Golfo San Jose,
Argentina, on undersides of rocks at low tide and dredged
offshore to 20 m.

Distribution. Golfo Corcovado on the east side of Isla Chi-
loe, Chile (northernmost specimens examined from Pumalin,
Chiloe Province, Chile, 42°42' S, 72°52' W, LACM 75-41)
to Tierra del Fuego, probably south to Cape Horn, east to
the Falkland Islands, and north in Argentina to the Golfo
San Matias (northernmost specimens from San Antonio Oeste,
Rio Negro Province, 40°45' S, 64°58' W, MACN 13361, A.

42.4 x 28.9 x 12.3 mm. (158) Puerto Deseado, Santa Cruz Province, Argentina. LACM 34851, 27.5 x 15.7 x 5.8 mm. (159) Puerto San
Julian, Santa Cruz Province, Argentina. AMNH 182640, 45.0 x 26.4 x 12.4 mm. (160) Santa Cruz River, Santa Cruz Province, Argentina.
ANSP 88526, 41.4 x 25.4 x 13.5 mm.

Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella 45

Figures 167 through 175. Fissurella radiosa radiosa Lesson, 1831, and F. radiosa tixierae Metivier, 1969. Radula, cut shell, mantle lobe,
juvenile shell, living and preserved specimens. Figures 167 through 172. F. radiosa radiosa. (167) Radula. Puerto el Hambre, Magallanes
Province, Chile. LACM 75-49, width of field 0.8 mm, shell length 20.8 mm. (168) Cut shell. Pumalin, Chiloe Province, Chile. LACM 75-41,
length of cut edge 18.5 mm. (169) Mantle lobe. Fuerte Bulnes, Magallanes Province, Chile. LACM 75-48, length 1 1 mm. (170) Juvenile shell.
Puerto el Hambre, Magallanes Province, Chile. LACM 75-49, 13.8 x 7.8 x 3.3 mm. (171) Juvenile shell. Pumalin, Chiloe Province, Chile.
LACM 75-41, 10.0 x 6.0 x 2.7 mm. (172) Preserved specimen. Fuerte Bulnes, Magallanes Province, Chile. LACM 75-48, shell length 56.5
mm. Figures 173 through 175. Fissurella radiosa tixierae. (173) Living specimen attached to substrate. Punta Cracker, Golfo Nuevo, Chubut
Province, Argentina. (174) Preserved specimen. Punta Ninfas, Golfo Nuevo, Chubut Province, Argentina. LACM 78-88, shell length 19.3
mm. (175) Ventral-lateral view of living specimen. Same locality. LACM 78-88.

Carcelles). The subspecies F. radiosa tixierae is characteristic
only of the Golfo San Matias and the Golfo Nuevo and Golfo
San Jose; specimens from such localities as Puerto Deseado
and Puerto San Julian, Santa Cruz Province, Argentina, are
consistently larger and lower in profile, identified as F. ra-
diosa radiosa. This is the only species of Fissurella that ranges
throughout the Magellanic Faunal Province in both Chile
and Argentina, and the only one that does not extend into
the region of overlap with the Peruvian Faunal Province in
south-central Chile.

Number of Lots Examined. F. radiosa radiosa: 55 (LACM
14, AMNH 5, ANSP 1, MACN 27, MNHN 1, USNM 7);
F. radiosa tixierae: 28 (LACM 7, AMNH 1, MACN 19,
USNM 1).

Taxonomic History. Fissurella radiosa Lesson, 1831, was
not originally illustrated. Most of the accounts dealing with
this species have consisted of copies and translations of orig-
inal descriptions of its numerous synonyms. Accounts with

additional observations are those of Strebel (1907), who was
the first to recognize the species, Ziegenhom and Thiem
(1925), and Riveros-Zuniga (1951), who recognized a spec-
imen under the name of F. dozei Rochebrune and Mabille.
The species has been misidentified as Lucapinel/a henseli
(Martens, 1 900), from Puerto Deseado, Santa Cruz Province,
Argentina, by Ringuelet et al. (1962).

The Argentinian subspecies F. radiosa tixierae was de-
scribed as a distinct species by Metivier (1969), based on a
single specimen. He also identified F. exquisita Reeve from
the Golfo Nuevo (here considered a synonym of F. radiosa),
apparently not having sufficient specimens to realize that a
single species is represented in the Golfo Nuevo.

Abundance and Use. Fissurella radiosa is common
throughout its range. It is a rather small form occurring to
the south of the populated regions of Chile; I have no infor-
mation as to whether it has been exploited for food.

Characteristics and Variability. The shell of Fissurella ra-

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McLean: Peruvian and Magellanic Fissurella

diosa radiosa is relatively small and elongate, with a narrow
dark margin, the ribbing fine and sharp, and the primary ribs
evident at all growth stages. Shell height varies from low to
moderately high. Color variation includes rayed forms and
some that are uniformly dark. Changes in color often occur
with growth. Some shells are nearly colorless in early stages
and later acquire rays; others are strongly rayed at first and
then lose the rays entirely. Normally rayed specimens may
have growth increments that are uniformly dark. The ribs
can be very evident or so weak that one can barely distinguish
primary ribs from secondary ribs. In the collections at hand
there seem to be more of the weakly sculptured examples
from the vicinity of the Strait of Magellan, whereas the strongly
sculptured specimens are known from more northern local-
ities in Chile and Santa Cruz Province, Argentina. The uni-
formly dark specimens have been seen only at Pumalin, in
the Golfo Corcovado, where they occur with rayed forms.
The largest observed specimen of the typical subspecies is
65 mm in length (LACM 75-42, Isla Talcon, Chile).

The large series of specimens of F. radiosa tixierae that I
collected in the Golfo Nuevo and Golfo San Jose in 1978
are as variable in color as those of the typical subspecies,
including many that are uniformly dark (Figs. 161, 162).
There is such a preponderance of elevated specimens that
the separation of a geographic subspecies based on this fea-
ture is justified. However, some specimens are as low as the
typical subspecies. The largest specimen observed measures
48.3 mm in length (Fig. 161).

Affinity and Comparisons. Fissurella radiosa most resem-
bles F. picta. Fissurella radiosa is smaller, has a narrower
margin, a more anterior foramen, and lacks the penciled
pattern of the rays. Unlike F. picta picta, which has gray rays
only, there are some reddish-rayed forms. In shell mor-
phology F. radiosa approaches F. oriens, which it may re-
semble in size, shape, and range of color possibilities, but it
differs in having a narrower margin, more anterior foramen,
and primary ribs that are raised, slightly nodular, and larger
than the adjacent ribs, in contrast to the nearly smooth aspect
of F. oriens. The papillae of the mantle fold are more strongly
developed in F. radiosa than in F. oriens. Fissurella radiosa
is more elongate than F. peruviana and has an elongate rather
than oval foramen.

Synonymy and Types. Type material of F. radiosa Lesson,
1831, from the Falkland Islands, has never been illustrated
and may not be extant. It is not housed in the Paris Museum
where some of Lesson’s material is now known (Bouchet,
personal communication). Lesson’s description stated: “The
ribs are separated by profound narrow grooves, and they are
arranged with three small ribs between each pair of larger
ones, all over.” This could also apply to F. picta, which was
then well known, but the broad margin of F. picta was not
mentioned; hence, by elimination we are left with F. radiosa.
Strebel ( 1 907) was the first to use the name F. radiosa in the
sense adopted here.

Type material of F. nigra Philippi, 1 845, not Lesson, 1831,
has not been located. The locality was given only as “Chile.”
Philippi’s illustration and description indicate that the shell
was predominantly black, the young stages with lighter rays,

the ribbing strong and unequal, and the margin narrow. Spec-
imens collected at Pumalin, east of Isla de Chiloe, are a good
match (Fig. 149), and the synonymy with F. radiosa is cer-
tain. Both Reeve and Hupe were to offer replacement names
for the preoccupied name of Philippi.

Fissurella darwinii Reeve, 1849, from the “Straits of Ma-
galhaens,” is represented by four syntypes, BMNH 197563,
lengths, 37.0, 36.9, 33.9, and 21.3 mm. The smallest spec-
imen has a margin broader than any in the three larger ones
and is clearly a young F. picta. Dell (1971) figured the second
largest specimen (36.9 mm in length); this is here figured and
designated the lectotype (Fig. 151). The rays are reddish pur-
ple. Because the margin, which has not previously been il-
lustrated, is narrow, I identify the type lot as F. radiosa.
Pilsbry (1890) copied Reeve’s illustration but also identified
and figured a shell with a broader margin — that figure is here
identified as F. picta.

Fissurella grisea Reeve, 1849, described without locality,
is represented by two syntypes, BMNH 1976140, measuring
30.3 and 21.1 mm in length. The largest (Fig. 152) has been
figured by Reeve and Sowerby II (1866) and is here desig-
nated the lectotype. Authors have not recognized this taxon.
The shell is gray and has faint dark rays. The surface of the
shell has evidently been treated with acid and only a trace
of radial sculpture remains. I tentatively assign it to the syn-
onymy of F. radiosa, suggested particularly by the tripartite
foramen.

There are 5 specimens labeled F. exquisita Reeve, 1850,
BMNH 197561. One small specimen is broken; the other
four measure 45.3, 37.8, 34.0, and 23.4 mm in length. The
Reeve locality is “Strait of Magalhaens,” but the original
label reads “Falkland Islands.” The smallest intact specimen
(Fig. 1 53) is a good match for the Reeve figure and was figured
and designated the lectotype by Dell (1971). The largest spec-
imen was also figured by Dell. All specimens are relatively
low, have a narrow margin, are similarly rayed with reddish
brown, and clearly show strong primary ribs.

Fissurella philippiana Reeve, 1850, was a name introduced
on the errata page of the Fissurella monograph: “Sp. 37. For
F. nigra Philippi, read F. philippiana Reeve.” Reeve was
renaming a homonym, although he did not explicitly state
this. The specimens considered syntypes by Dell therefore
have no standing as types. There are four specimens, BMNH
197562, the locality “Chile” in ink on the original mounting
board, although Reeve gave the locality as “Southern Chile.”
The specimens measure 42.0, 38.6, 33.6, and 32.3 mm in
length. They represent the dark form of F. radiosa in which
the primary ribs are prominent and the margin narrow. Dell
(1971) figured the 38.6 mm specimen; the largest is figured
here (Fig. 150). The “syntypes” are very similar to those I
found at Pumalin (Fig. 149). Dali (1909) inexplicably gave
the locality as “Concepcion, Chile,” and this has been re-
peated by subsequent authors. No specimens corresponding
to this locality have been found at the USNM. Concepcion
is well to the north of the known distribution of F. radiosa.

Fissurella philippii Hupe, 1854, was proposed as a replace-
ment name for F. nigra Philippi, not Lesson. Hupe was un-
aware of the similar name proposed by Reeve.

Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella 47

j'ofeiA

Figures 176 through 191. Fissurella oriens fulvescens Sowerby, 1835, and F. oriens oriens Sowerby, 1835. Mature shells. Figures 176 and
177. F. oriens fulvescens. (176) Lectotype, F. fulvescens Sowerby. Valparaiso, Chile. BMNH 197568, 39.7 x 23.0 x 6.8 mm. (177) Islota
Concon, Valparaiso Province, Chile. LACM 75-31, 23.2 x 14.8 x 4.5 mm (beach shell). Figures 178 through 191. F. oriens oriens. (178)
Paralectotype, F. oriens Sowerby. Locality uncertain. BMNH 197575,62.4 x 35.7 x 14.8 mm. (179) Lectotype, F. mexicana Sowerby. Locality
unknown. BMNH 1944593, 38.6 x 22.2 x 8.6 mm. (180) Pargua, Llanquihue Province, Chile. LACM 75-39, 45.8 x 19.6 x 6.8 mm. (181)
Pumalin, Chiloe Province, Chile. LACM 75-41, 58.0 x 32.7 x 12.0 mm. (182) 4-13 m, Islota Nihuel, Chiloe Province, Chile. LACM 75-
43, 41.5 x 24.3 x 8.5 mm. (183) Same locality. LACM 75-42, 49.3 x 33.0 x 18.4 mm. (184) Holotype, F. oblonga Ramirez-Boehme. Punta

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McLean: Peruvian and Magellanic Fissurella

There are two syntypes of F. polygona Sowerby II, 1862,
BMNH 1976151, from the Falkland Islands. The largest
measures 42.4 mm in length and was figured by Dell (1971).
It is here designated the lectotype (Fig. 157). The smaller
specimen measures 22.2 mm in length and has proportions
similar to the larger. Primary ribs are not as pronounced as
shown in the original figure (copied by Pilsbry, 1890); the
rays are reddish brown. The specimens represent the broad,
elevated extreme of F. radiosa, which may be the typical
form, as it has the same general type locality as that of the
senior synonym.

Type material of Fissurella dozei Rochebrune and Mabille,
1885, described from Santa Cruz, Patagonia, was not located
in the Paris Museum by P. Bouchet, although type material
of two other Rochebrune and Mabille taxa is known in the
Paris Museum. The drawing given by the authors in 1 889 is
a fair rendition of F. radiosa with well marked ribs, the figure
showing that there are primary ribs that are slightly more
prominent than the secondary' ribs; the width of the margin
is not mentioned. Puerto Santa Cruz, Santa Cruz Province,
Argentina (50°0T S) is north of the eastern entrance to the
Strait of Magellan. This is a region in which /-. radiosa is
now known to be the only living species of Fissurella. The
synonymy of F. dozei with F. radiosa is therefore reasonably
certain.

Fissurella tixierae Metivier, 1969, was based on one spec-
imen in poor condition, MNHNP uncataloged, from the in-
tertidal zone in the Golfo Nuevo, Argentina (42°56' S, 64°24'
W), length 22.5 mm, width 13.7 mm, height 8 mm (Fig. 163).

Fissurella oriens Sowerby, 1835

Figures 176-199

Fissurella oriens is here considered to have two geographic
subspecies: F. oriens oriens in the Magellanic region of Chile
and F. oriens fulvescens in south-central Chile.

Synonymy for F. oriens oriens:

Fissurella oriens Sowerby, 1835a: 124; Sowerby, 1835b:3, figs.
25, 60; Reeve, 1849, pi. 2, fig. 13; Hupe, 1854:237; Sow-
erby II, 1862:186, figs. 19, 20; Pilsbry, 1890:152, pi. 46,
figs. 18, 19, pi. 34, fig. 58; Melvill and Standen, 1907:97;
Strebel, 1907:88, pi. 1 , figs. 8-14, pi. 2, figs. 1 5-20; Strebel,
1908:78, pi. 6, figs. 97a, b; Dali, 1909:242; Melvill and
Standen, 1914:114; Powell, 1951:85; Carcelles and Wil-
liamson, 1951:254; Riveros-Zuniga, 1951:123, fig. 30; Dell,
1971:185, pi. 5, figs. 3, 4; Ramirez-Boehme, 1974:30 [key].
Fissurella mexicana Sowerby, 1 835b:8, fig. 61; Reeve, 1849,
pi. 6, fig. 40; Sowerby II, 1862:186, figs. 26-28; Pilsbry,

1890:153, pi. 34, fig. 60; Melvill and Standen, 1898:102;
Strebel, 1907:88 [under F. oriens].

Fissurella australis Philippi, 1845:61; Philippi, 1845:142;

Strebel, 1907:88 [under F. oriens],

Fissurella alba Philippi, 1845:61; Philippi, 1845:34, pi. 1,
fig. 4; Hupe, 1854:247; Rochebrune and Mabille, 1889:71;
Pilsbry, 1890:292, pi. 62, figs. 3-5; Strebel, 1907:94, pi. 1,
figs. 1-3, pi. 2, fig. 21; Ziegenhom and Thiem, 1925:14,
pi. 2, figs. 14a, 14b, 15; Carcelles, 1950:51, pi. 1, fig. 11;
Carcelles and Williamson, 1951:254; Riveros-Zuniga, 1951:
100; Dell, 1971:181; Ramirez-Boehme, 1974:30 [key].
Fissurella (Corrina) alba Christiaens, 1973:93, pi. 4, figs. 46,
47.

Fissurella flavida Philippi, 1857:165; Pilsbry, 1 890:292; Stre-
bel, 1907:97; Carcelles and Williamson, 1951:254; Ri-
veros-Zuniga, 1951: 121; Dell, 1971:186; Ramirez-Boehme,
1974:30 [key],

Fissurella hedeia Rochebrune and Mabille, 1885:109;
Rochebrune and Mabille, 1889:72, pi. 5, fig. 3; Carcelles,
1950:51; Carcelles and Williamson, 1951:255; Riveros-
Zuniga, 1951, fig. 32; Dell, 1971:186; Ramirez-Boehme,
1974:30 [key].

Fissurella arenicola Rochebrune and Mabille, 1885:109;
Rochebrune and Mabille, 1889:73, pi. 5, fig. 1; Carcelles,
1950:51; Carcelles and Williamson, 1951:255; Riveros-
Zuniga, 1951:124, fig. 31; Dell, 1971: 182; Ramirez-
Boehme, 1974:30 [key].

Fissurella ( Carcellesia) doellojuradoi Perez- Farfante, 1952:
32, fig. 1; Christiaens, 1973:92, pi. 4, fig. 45.

Fissurella cheullina Ramirez-Boehme, 1974:17, 30 [key], pi.

1, figs. 2a-c.

Fissurella oblonga Ramirez-Boehme, 1974:18, 30 [key], pi.

2, figs. 4a-c.

Synonymy for F. oriens fulvescens:

Fissurella fulvescens Sowerby, 1835a: 127; Sowerby, 1835b:
6, fig. 49; Reeve, 1849, pi. 6, fig. 42; Hupe, 1854:245;
Sowerby II, 1862:184, fig. 36; Pilsbry, 1890:152, pi. 33,
fig. 49; Dali, 1909:241; Riveros-Zuniga, 1951:122, fig. 29;
Dell, 1971:186, pi. 5, fig. 6; Ramirez-Boehme, 1974:30
[key].

Shell (F. oriens oriens). Small to medium sized (mature
length 40-70 mm), low to moderately elevated; outline elon-
gate oval to very elongate, lateral profile variable, ranging
from flat or with either ends or sides raised. Sculpture of fine
radial ribs, ribs very broad and flat at margin, ending in
extremely fine crenulations; under magnification fine con-

Chulao, Chiloe Province, Chile. MNHN 200375, 55.5 x 31.0 x 12.2 mm. (185) Holotype, F. cheullina Ramirez-Boehme. Isla Quellin,
Llanquihue Province, Chile. MNHN 200327, 43 x 23 x 9 mm. (186) Holotype, F. hedeia Rochebrune and Mabille. Punta Arenas, Magallanes
Province, Chile. MNHNP, 29 x 24 x 7 mm. (187) Orange Harbor, Chile. USNM 17328, 37.4 x 22.9 x 8.2 mm. (188) 13-29 m. Strait of
Magellan, Chile. LACM 90801, 47.3 x 29.7 x 10.6 mm. (189) 18-27 rn. Falkland Islands. USNM 368309, 23.2 x 14.7 x 5.0 mm. (190)
Punta Arenas, Magallanes Province, Chile. LACM 75-50, 54.2 x 33.2 x 16.9 mm. (191) Holotype, F. arenicola Rochebrune and Mabille.
Locality uncertain. MNHNP, 42.2 x 26.5 x 14.2 mm.

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McLean: Peruvian and Magellanic Fissurella 49

Figures 192 through 199. Fissure/la oriens oriens Sowerby, 1835. Living specimens, radula, mantle lobe, juvenile shells, and cut shell. (192)
Detached living specimens. 4-13 m, Islota Nihuel, Chiloe Province, Chile. LACM 75-43. (193) SEM view of radular ribbon. Pargua, Llanqihue
Province, Chile. LACM 75-39, width of field 0.7 mm. (194) Radula. Pumalin, Chiloe Province, Chile. LACM 75-41, width of field 0.5 mm.
(195) Mantle lobe. Same locality. LACM 75-41, length 9 mm. (196) Juvenile shell. 11-13 m, Isla Carlos III, Magallanes Province, Chile.
LACM 73-70, 7.8 x 4.5 x 1.9 mm. (197) Juvenile shell. 4-13 m, Islota Nihuel, Chiloe Province, Chile. LACM 75-43, 13.4 x 8.2 x 3.4 mm.
(198) Air-dned radula. Pumalin, Chiloe Province, Chile. LACM 75-41, width of ribbon 3.0 mm, length of shell 48.8 mm. (199) Cut shell. 4-
13 m, Islota Nihuel, Chiloe Province, Chile. LACM 75-42, length of cut edge 17 mm.

centric lamellae visible in early growth stages. Ground color
variable, from colorless to yellow or dark red; rays mostly
reddish or gray, solid or split into two or more broad bands
and frequently with concentric interruptions; gray rays often
changing to reddish or black to gray. Some specimens com-
pletely colorless. Margin relatively narrow, somewhat broad-
er in rapidly growing specimens, not zoned, showing full
pigmentation of rays throughout calcitic layer. Foramen
elongate and tripartite in young shells, elongate-oval in ma-
ture shells.

Shells of the northern subspecies, F. oriens fulvescens,
smaller (maximum length 45 mm), relatively low, ends usu-
ally elevated relative to sides. Color consistent; ground color

yellow orange, rays dark red, rays frequently split into two
adjacent bands, young shells speckled with red. This color-
ation and the speckled pattern of young shells is not matched
by that in any specimens of the typical subspecies.

Juvenile Shell. Elongate oval, more conical than at later
stages, usually reddish overall, with white apical area and
color pattern emerging unevenly; rays arising after shell
reaches about 4 mm in length.

Mantle and Foot. Animal usually large, not retractable in
flattened shell. Cephalic tentacles reddish brown, yellowish
at tips. Mantle lobe rather thin in preserved specimens, edges
with finely branched papillae, banded to correspond to rayed
pattern; side of foot mottled brown to pinkish brown, light

50 Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella

in specimens with colorless shells. Epipodial tentacles es-
pecially prominent.

Habitat. Chiefly sublittoral throughout the range, but ex-
tending up to the lower intertidal zone in places exposed to
currents but not heavy surf. The northern subspecies F. ori-
ens fulvescens is probably limited to the sublittoral zone, for
I was unable to find living specimens at low tide. I observed
the typical subspecies at Mehuin in deep tide pools. I found
it at Pumalin in the Golfo Corcovado, occurring at low tide
in rocky areas free of sand where tidal currents were strong;
at Islote Nihuel in the Golfo Corcovado I found it at a depth
of 10 m on rocks near the sand-rock interface. At Puerto
Hambre in the Strait of Magellan I found it at low water in
sheltered rocky areas. Specimens have been dredged to depths
of 30 m by the R/V HERO near Isla de los Estados, Argen-
tina.

Distribution. Islote Concon, Valparaiso Province, Chile
(32°52'S) (LACM 75-31, McLean), south toTierradel Fuego
and probably Cape Horn, east to Isla de los Estados, Argen-
tina, and the Falkland Islands. Not known living from main-
land Argentina. Pleistocene specimens are known from Co-
modoro Rivadavia, Chubut Province, Argentina (MCZ
288329), as is also true for F. picta. Scarabino’s (1977) record
of the species from the Golfo San Matias, Argentina, is prob-
ably based upon specimens of F. radiosa tixierae. Popula-
tions from the northern end of the range in the vicinity of
Valparaiso and south at least to Concepcion are of the sub-
species F. oriens fulvescens. Those at Mehuin and to the south
are the typical subspecies F. oriens oriens.

Number of Lots Examined. F. oriens oriens: 97(LACM 34,
AMNH 10, ANSP 6, MACN 28, MNHN 6, USNM 13); F.
oriens fulvescens: 4 (LACM).

Taxonomic History. Fissurella oriens Sowerby has been
reasonably well understood by most authors, although the
extent of its variability and the extreme number of synonyms
has not generally been recognized. Most of the synonyms
have not come into general use, with one exception, that of
F. alba Philippi, which was based on white-shelled forms.
The northern subspecies, F. oriens fulvescens Sowerby, is
here recognized for the first time.

Abundance and Use. Fissurella oriens is primarily a species
of the Magellanic Faunal Province, where it is common at
low water and much more abundant in the sublittoral. Al-
though many shells are small, it reaches sizes large enough
to be used for food. I have no information on the extent of
its use.

Characteristics and Variability. The most characteristic
features of F. oriens are the relatively small size and lack of
strong ribbing, the radial sculpture being better described as
consisting of striae or grooves. It is one of the most variable
species in proportions, size of the foramen, and color pattern.
Local populations tend to be consistent, with many similarly
appearing individuals. The ground color of the typical sub-
species ranges from colorless to dark red, but most commonly
has strong rays that may change with growth from gray to
reddish and increase or diminish in intensity. White shelled
specimens have been noticed in scattered populations
throughout the range of the species. Some populations have

shells that are relatively flat, while in others the shells are
more elevated. This is one of the few species in which the
variation includes specimens with either elevated sides or
elevated ends. The margin is usually narrow, though not as
narrow as that of F. radiosa. However, some specimens that
are growing rapidly may have a relatively broad margin. In
some gerontic specimens, the foramen may become very
large and broadly oval; in others it remains narrow and elon-
gate.

Affinity and Comparisons. Fissurella oriens most resem-
bles F. radiosa, a species of similar size in which there is
similar variation in proportions and color. Fissurella oriens
differs chiefly in lacking the strong ribbing of F. radiosa and
in having a more centrally placed foramen. However, some
conspicuously ribbed specimens of F. oriens may so resemble
weakly ribbed specimens of F. radiosa that the only reliable
character to separate them is the presence of primary ribs
larger than the adjacent ribs on F. radiosa, and the absence
of such primary ribs on F. oriens. Strongly rayed specimens
of F. oriens may have a superficial resemblance to uneroded
specimens of F. limbata, but F. oriens lacks the broad two-
zoned margin that is the hallmark of F. limbata.

Synonymy and Types. Eleven names seem to be referable
to F. oriens, the large number of synonyms correlated with
the high variability of the species and the tendency for local
populations to have uniform features.

There are five syntypes of F. oriens Sowerby, 1835, BMNH
197575, lengths 69.3, 62.4, 59.6, 53.7, and 16.7 mm. The
largest specimen, that figured by Reeve ( 1 849) and Dell (1971)
is designated the lectotype; the second largest specimen, a
paralectotype, is figured here (Fig. 1 78). Although Reeve gave
the locality as Valparaiso, Sowerby’s original locality is “In-
sulam Chiloe,” with a “variety” mentioned from Valparaiso.
The specimens are relatively large, rayed in reddish brown,
and represent the normal, elongate, moderately elevated form,
similar to what I have observed from the vicinity of Isla de
Chiloe. The type locality should therefore be limited to Isla
de Chiloe. Sowerby’s original figure in the “Conchological
Illustrations” was of a smaller specimen, probably the 59.6
mm specimen.

Sowerby did not clearly document the above-mentioned
“variety” of F. oriens from Valparaiso. However, in the same
publication (Sowerby, 1835a) he described F. fulvescens from
Valparaiso, which may have been intended as the “variety.”
In 1975 I found beach-worn specimens matching the type
lot on cobble beaches in that vicinity of Chile (Fig. 177). The
type lot consists of four specimens, BMNH 197568, lengths
39.7, 37.0, 33.0, and 27.9 mm. The largest specimen (Fig.
176) was figured originally by Sowerby ( 1835b), later by Reeve
(1849), and more recently by Dell (1971), who designated it
the lectotype. All the specimens are low and elongate, yellow-
orange in ground color, the rays reddish, and the margin
narrow. The specimens are in good condition and were ev-
idently collected alive; according to Sowerby, they were taken
under rocks on the shore.

Fissurella mexicana Sowerby, 1835, was said to come from
“Real Llejos, Mexico,” obviously in error. There are four
specimens in the type lot, BMNH 1966493, lengths 40.6,

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McLean: Peruvian and Magellanic Fissurella 51

40.4, 38.6, and 29.6, the smallest specimen broken and re-
paired. They are elongate, moderately elevated, and rather
thin-shelled, well within the range of variation of F. oriens.
The 38.6 mm specimen is designated the lectotype (Fig. 1 79).

Fissurella australis Philippi, 1845, from “Fretum Magel-
lanicum,” was never illustrated and I have not been able to
locate type material. Philippi described its color and sculp-
ture as similar to that of a young F. oriens , but thinner-shelled
and with extremities raised, the shell resting on the sides.
Inasmuch as specimens of F. oriens with raised ends and
thin shells are well known, the synonymy is reasonably cer-
tain.

Fissurella alba Philippi, also from “Fretum Magellani-
cum,” was illustrated subsequently by Philippi, but again,
type material has not been located. A white, finely sculptured
shell was figured, and Philippi himself indicated that it could
be a variant of F. oriens. This note was, surprisingly, over-
looked by Pilsbry (1890) and the taxon has been accepted
without question by subsequent authors and even made the
type species of a subgenus based upon the character of weak
ribbing. Nevertheless, it is clearly a white-shelled variant of
F. oriens. I have seen white shells from numerous stations
at which rayed forms also occur (see Fig. 188, from a lot of
six specimens, four of which are white and two are rayed).

Fissurella flavida Philippi, 1857, from the “Magellen-
strasse,” has not been figured and I have not located type
material. The description indicates a shell that is solid, el-
liptical, moderately convex, with obscure sculpture, and a
moderately broad margin. The coarse sculpture that char-
acterizes F. picta and F. radiosa were not mentioned. Inas-
much as only three species occur in the region, this name
can be relegated to the synonymy of F. oriens by eliminating
the other possibilities.

The holotype of F. hedeia Rochebrune and Mabille, 1 885,
from Punta Arenas in the Strait of Magellan, was received
on loan from the Paris Museum (Fig. 186). Although not
compared by its authors to other species, it is an elongate,
finely sculptured specimen of F. oriens.

Fissurella arenicola Rochebrune and Mabille, 1885, also
was not compared to other species. Two specimens in the
Paris Museum labeled “type et paratype,” were studied. The
locality inked on the mounting board is “Baie Orange”; the
published locality is “Punta- Arenas Patagoniae, Baie Or-
ange.” The holotype (Fig. 191) is a white-shelled F. oriens,
42.2 mm in length, very worn except at the margin, where
no traces of primary ribs are evident. The paratype is 38.2
mm in length, also white-shelled, but primary ribs are ap-
parent throughout. I therefore identify the paratype as F.
radiosa. Based on the holotype, the name F. arenicola is
placed in the synonymy of F. oriens.

Fissurella ( Carcellesia ) doellojuradoi Perez-Farfante, 1952,
was based on a single specimen from “Tierra del Fuego.”
The specimen, 35 mm in length, was borrowed by its author
from the Museo Argentino de Ciencias Naturales in Buenos
Aires. A new subgenus was based on the single feature of the
raised ends of the specimen, but as discussed above, such
variants are frequently seen in F. oriens, with which it was

not originally compared. The synonymy of this taxon there-
fore seems certain.

Fissurella cheullina Ramirez-Boehme, 1974, was based on
eight specimens from Isla Queullin, Llanquihue Province,
Chile (41°53' S, 72°55' W), holotype MNHN 200377 (Fig.
185). Fissurella oblonga Ramirez-Boehme, 1974, was based
on a single specimen from Punta Chulao, Chiloe Province,
Chile (42°17' S, 72°50' W), holotype MNHN 200375 (Fig.
1 84). Neither taxon was compared by its author to any other
species. Both were described as moderately elevated, elon-
gate, weakly sculptured, and with narrow margins. These
specimens are well within the range of variation known for
F. oriens.

Fissurella nigra Lesson, 1831

Figures 200-2 1 1

Fissurella nigra Lesson, 1831:412; Orbigny, 1841:473; Phi-
lippi, 1846:65, fig. 2; Reeve, 1849, pi. 2, fig. 11; Hupe,
1854:241; Sowerby II, 1862:184, fig. 14; Pilsbry, 1890:
149, pi. 35, figs. 1,2; Dali, 1909: 1 77, 242; Ziegenhom and
Thiem, 1925:15, pi. 2, figs. 16a, b; Carcelles and William-
son, 1951:255; Riveros-Zuniga, 1951:116, fig. 25; Dell,
1971:188, pi. 3, figs. 17-19; Ramirez-Boehme, 1974:31
[key],

Fissurella violacea Rathke, 1833 [ex Eschscholtz manu-
script]^!, pi. 23, fig. 6; Orbigny, 1841:473 [under F. ni-
gra]-, Philippi, 1846:66, pi. 2, fig. 3.

Fissurella grandis Sowerby, 1835a:123; 1835b:3, fig. 48; Or-
bigny, 1841:473 [under F. nigra]-, Philippi, 1846:65, pi. 2,
fig- 1.

Shell. Relatively large (70 to 1 10 mm mature length), mod-
erately elevated; outline elongate oval, base resting flat or
with ends slightly raised, rarely with raised sides. Overall
appearance smooth, sculptured with fine radial ribs that per-
sist to margin. Color black or gray, sometimes pale and show-
ing concentric variation in shading; rays lacking or faint,
consisting of narrow lines slightly darker than adjacent ground.
Margin broad and flat in growing shells, narrow in mature
shells; two-zoned, outer zone black, inner zone translucent
gray; cut shells showing that two zones are nearly equal in
thickness. Foramen anterior to center, elongate and tripartite
in young shells, elongate-oval in mature shells, worn or bev-
eled at apex to reveal aragonitic layer, so that it always ap-
pears white-bordered. Old shells that continue to grow by
increasing height without expanding at base may contract
basal area, forming thick edge and losing distinct zoning of
margin.

Juvenile Shell. Radial ribs fine and sharp; primary and
secondary ribs appearing early. Whitish in earliest stage,
changing abruptly or gradually to black, some showing two
lateral white rays that quickly fade.

Mantle and Foot. Nearly containable in shell. Entire ani-
mal gray; yellow coloration lacking in cephalic tentacles.
Mantle lobe greatly expandable, faintly banded, narrow when
preserved; papillae of upper edge moderately developed, fine-
ly branched; those of lower edge nearly equal in size. Side of

52 Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella

Figures 200 through 204. Fissurella nigra Lesson, 1831. Mature shells. (200) Island off Mehuin, Valdivia Province, Chile. LACM 75-37,
74.7 x 50.2 x 19.5 mm. (201) Lectotype, F. grandis Sowerby, 1835. Locality uncertain. BMNH 1976143, 101.0 x 67.9 x 30.9 mm. (202)
Quellon, Chiloe Province, Chile. LACM 75-45, 79.5 x 51.1 x 30.1 mm. (203) Pumalin, Chiloe Province, Chile. LACM 75-41, 43.8 x 27.5
x 12.4 mm. (204) Isla Bertrand, off Isla Navarino, Magallanes Province, Chile. NMNZ 18409, 78.9 x 56.5 x 25.4 mm.

foot darker than mantle lobe; tubercles small and widely
scattered. This is the only species with a completely gray
animal.

Habitat. Intertidal zone only, on the undersides of rocks
in the mid-tidal to lower intertidal zone. Several large spec-
imens may occur close together on the undersides of large
rocks. Shells are mostly clean, except for scattered incrus-
tations of barnacles, bryozoa, or spirorbid worms. None have

been found with attached Scurria parasitica, as would be
expected if the habitat were more exposed. This is the only
species occurring under rocks in the mid-littoral of south
central and southern Chile; only juveniles of other species of
Fissurella occur in this habitat in northern Chile.

Distribution. Valparaiso, Santiago Province, Chile (33°02'
S) (USNM 48218, Bridges), to Puerto Grandi, Isla Bertrand,
off south side Isla Navarino, Chile (5 5° 1 2' S, 67°02' W) (Dell,

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McLean: Peruvian and Magellanic Fissurella 53

Figures 205 through 211. Fissurella nigra Lesson, 1831. Living specimens, radulae, juvenile shell, cut shell, and mantle lobe. (205) Living
specimen on underside of overturned boulder. Island ofTMehuin, Valdivia Province, Chile. LACM 75-37. (206) Radula. Same locality. LACM
75-37, width of field 0.8 mm, shell length 26.4 mm. (207) Ventral view of living specimen. Rio Bio-bio, Concepcion Province, Chile. LACM
75-35. (208) Juvenile shell. Island off Mehuin, Valdivia Province, Chile. LACM 75-37, 5.5 x 3.5 x 1.5 mm. (209) Air-dried radula. Same
locality. LACM 75-37, width of ribbon 2.6 mm. (210) Cut shell. Same locality. LACM 75-37, length of cut edge 39 mm. (211) Mantle lobe.
Same locality. LACM 75-37, length 10 mm.

1971). The northernmost specimen personally collected is a
single beach-worn juvenile from Punta el Lacho, Santiago
Province, Chile (33°30' S). The species is rare in the vicinity
of Valparaiso, if it now occurs there at all. Dali's ( 1 909) record
from Callao, Peru, is not represented by USNM specimens
and should be discounted. I found this species commonly at
Concepcion and Mehuin, and at all stations in the vicinity
of Isla de Chiloe, both on the exposed outer coast and on
the eastern side where water motion is primarily that of tidal
currents. It is evidently uncommon and sporadic in the south-
ernmost region, for I found no trace of it at Punta Arenas or
Puerto Hambre in the Strait of Magellan. It is unknown from
the Falkland Islands. In addition to the above record of Dell
(1971), the following southern records are known to me:
USNM 1 70205, Port Otway, Chile (46°49' S); MACN 12491,
Canal Smyth (at western end of Strait of Magellan); AMNH
173403, Beagle Canal (south side Tierra del Fuego).

Number of Lots Examined. 67 (LACM 1 1, AMNH 8, ANSP
3, MACN 8, MNHN 33, USNM 4).

Taxonomic History. Fissurella nigra is an easily recognized
species that has been understood by most authors.

Abundance and Use. Common, at least in the northern-
most portion of the range. Its intertidal habitat is accessible,
and it is exploited for food.

Characteristics and Variability. The most characteristic
features of F. nigra are the gray to black surface, the rays, if
present, being faint and split into lines, and the prominent
zoning of the calcitic layer at the margin, the outer zone dark
and the inner zone a light translucent gray. Variation is chiefly
in color pattern. Some specimens change from black to gray
or light brown; such specimens being more likely to show
the fine brown radial pattern. There may be changes in color
intensity with growth, but never as pronounced as that which
occurs in F. picta, F. radiosa, or F. oriens. The size record

54 Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella

for the species, perhaps for the genus as well, is length 135
mm, width 100 mm, height 62 mm (Isla de Chiloe, collected
in 1892, MNHN).

Affinity and Comparisons. Fissurella nigra does not closely
resemble any other species. Although its overall appearance
is smooth, juvenile shells are sharply ribbed, and there are
definite primary and secondary ribs. The presence of these
ribs in the juvenile dissociates it from the smooth-shelled
species in the “group of F. limbata” in which early sculpture
consists only of broad primary ribs. Its closest affinity is
probably with F. radiosa, with which it shares similar pro-
portions, an anteriorly positioned foramen, and the three
series of ribs, although the total number of ribs is greater in
F. nigra. The dark outer zone of F. nigra is unique. There
is only a superficial resemblance of F. nigra to dark gray
specimens of F. latimarginata. Compared to the latter, F.
nigra has fine radial ribs, a dark rather than lighter outer
zone to the margin, and a more anterior and more inwardly
beveled foramen.

Synonymy and Types. Type material of F. nigra Lesson,
1831, is unknown. It is not represented in the Paris Museum
where some of the Lesson collection is now housed. Although
there were no original illustrations, the species is easily rec-
ognized from the description. It is common in the vicinity
of its type locality “Saint-Vincent,” now San Vicente (36°43'
S), near Concepcion, Chile.

Fissurella violacea Rathke, 1833, was described before the
Lesson work was known. The original figure is clear; the type
locality is Concepcion. Type material may be in Leningrad,
where some of the Eschscholtz and Rathke types are extant.

Fissurella grandis Sowerby, 1835, is another early name
evidently introduced before Lesson’s work was known. There
are two specimens in the type lot, BMNH 1976143, lengths,
110.1 and 77.2 mm. The larger specimen was figured by
Sowerby (1835b), Reeve (1849), and Dell (1971). It is here
designated the lectotype (Fig. 201). The original locality was
given as “Valparaiso and Chiloe,” so it is not certain whether
both specimens are from the same place. The correct locality
may be Isla de Chiloe, because the species is rare in the
vicinity of Valparaiso.

Group of Fissurella limbata

Relatively large-shelled species in which the thickness of the
calcitic layer greatly exceeds that of the interior aragonitic
layer. Radial sculpture in the early stage is either absent or
consists of broad, low primary ribs; there are no secondary
ribs. Mature shells may retain the broad primary ribs as low
undulations or be entirely smooth.

Although shells of this group have the thick calcitic layer
of the F. maxima group, they differ from the latter in not
having secondary ribs. Whether this difference in sculpture
has taxonomic significance should be tested at the biochem-
ical level. It is premature to justify a separation at the subge-
neric level.

There are four species in this group: F. limbata. F. crassa.
F. bridgesii, and F. pulchra. The latter three have elongate
shells with low profiles, the animals too large to be retracted
within the shell. These species tend to be more stenotopic

than those of the F. maxima group. Variation in each species
is less extreme, which is correlated with the relatively few
synonyms in the group.

The four species in the group are limited to the Peruvian
Faunal Province, unlike the F. maxima group, in which there
are both Peruvian and Magellanic members.

Fissurella limbata Sowerby, 1835

Figures 2 1 2-224

Fissurella limbata Sowerby, 1835a: 123; Sowerby, 1835b:3,
figs. 42, 66, 74; Orbigny, 1841:474; Reeve, 1849, pi. 2,
figs. 10, 12; Hupe, 1854:239; Sowerby II, 1862:184, figs.
23, 24; Pilsbry, 1890:149, pi. 32, figs. 26-39; Dali, 1909:
242; Ziegenhom and Thiem, 1925; 15, pi. 2, figs. 17-19,
20a, 20b; Riveros-Zuniga, 1951:114, fig. 24; Pena, 1970:
156; Dell, 1971:188, pi. 5, fig. 5; Manncovich, 1973:18,
fig. 30; Ramirez-Boehme, 1974:32 [key].

Fissurella limbata var. multilineata Ziegenhom and Thiem,
1925:16, pi. 2, fig. 21.

Shell. Medium large (60-90 mm mature length), moder-
ately elevated; outline elongate oval; base usually resting flat,
with ends or sides slightly raised. Sculpture of wavy, irregular
radial ribs; overall aspect mostly smooth. Color of uneroded
shells (lacking epibiotic Scurria ) consisting of yellowish ground
and purple rays, rays often split. Calcitic layer zoned, con-
sisting of inner layer of solid dark purple and translucent
outer layer. Eroded shells (those with Scurria) worn to white
aragonitic layer near foramen; area farther away from fora-
men consisting of deep purple portion of calcitic layer, with
pattern of rays evident only near margin. Margin broad in
growing shells, dark purple next to aragonitic interior, outer
edge lighter and revealing pattern of rays. Cut shells showing
outer zone about one-fourth thickness of solid purple zone.
Foramen elongate at all stages, tripartite in young shells,
constricted in middle in mature shells.

Juvenile Shell. Elongate-oval, elevated; primary ribs broad,
weak. Color generally white, marked with concentric zigzag
purple lines coalescing into purple rays; inner purple zone of
calcitic layer apparently lacking in juvenile shells.

Mantle and Foot. Shell edge enveloped by mantle on at-
tached specimens exposed at low tide, body retractable with-
in shell. Mantle lobe light gray, appearing narrow in pre-
served specimens, marked with concentric lines of black, and
only faintly banded to correspond to rays of shell. Papillae
at edges very small, finely branched. Side of foot gray; tu-
bercles small, scattered. Preserved specimens retaining broad
dark ring with lighter edge where the smooth innermost edge
of mantle lobe is in contact with shell. This is the only species
in which a dark zone on the inner lobe shows in preserved
specimens.

Habitat. Mid-intertidal to low-intertidal zones in surf-ex-
posed areas, primarily on horizontal and sloping surfaces,
rather than vertical surfaces. This is the predominant species
on flat areas of exposed reefs. I saw no specimens in the
sublittoral zone at localities where I dived. Most shells have
a single Scurria parasitica, which produces a deeply etched
attachment scar.

Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella 55

Figures 212 through 216. Fissurella limbala Sowerby, 1835. Mature shells. (212) Isla Guanape, Peru. LACM 74-2, 37.0 x 22.9 x 12.1 mm.
(213) Shore opposite Isla Santa Maria, Antofagasta Province, Chile. LACM 75-17, 65.8 x 41.0 x 20.0 mm. (214) Los Molles, Aconcagua
Province, Chile. LACM 75-28, 82.4 x 57.7 x 25.0 mm. (215) Paralectotype, F. limbata Sowerby. Valparaiso, Chile. BMNH 197581, 65.0 x
48.5 x 1 17.8 mm. (216) Lectotype, F. limbata Sowerby. Valparaiso, Chile. BMNH 197582, 60.5 x 40.0 x 17.4 mm.

Distribution. Isla Guanape, Peru (8°30' S) (LACM 74-2,
McLean), to Isla de Chiloe, Chile (AMNH 155914, O. Ruiz).
The exact locality for the southern record is not known, but
it was probably the accessible northwestern tip at approxi-
mately 41°50' S. However, I was unable to find specimens
in that vicinity at Guabun, Isla de Chiloe. The species does
not occur in the sheltered waters of the Golfo Corcovado on
the eastern side of Isla de Chiloe. I discount the record of
Riveros-Zuniga (1951) from Fuerte Bulnes in the Magellan
Strait.

Number of Lots Examined. 100 (LACM 35, AMNH 20,
ANSP 8, MACN 8, MNHN 10, USNM 19).

Taxonomic History. Fissurella limbata has been correctly
interpreted by most authors.

Abundance and Use. This species is common throughout
its range and is exploited for food. Large specimens are sel-

dom seen, because of the accessibility of the habitat at low
tide. It is called the “lapa gaviota” at Iquique. Gaviota is the
name for seagull, a predator on this species.

Characteristics and Variability. The most characteristic
and unusual feature of F. limbata is the zonation of the
calcitic layer, in which the pattern of rays is confined to the
thin, lighter colored surface layer. The greater thickness of
the calcitic layer consists of the dark purple inner zone, which
is much darker than the rays. There is little variation; dif-
ferences in appearance are a result of patterns of wear in
which the outermost rayed layer is lost, leaving a uniform
purple layer. Further wear results in complete loss of the
calcitic layer near the foramen, which exposes the white ara-
gonitic layer. Erosion of the shells is greatly accelerated when
Scurria parasitica is present.

Affinity and Comparisons. Fissurella limbata seems not

56 Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella

WiW 220

222

223

219

217

224

Figures 217 through 224. Fissurella limbata Sowerby, 1835. Living specimens, juvenile shells, radula, cut shell, and mantle lobe. (217)
Ventral view of living specimen. Cumbres Borascosas, Tarapaca Province, Chile. LACM 75-14. (218) Juvenile shell. Iquique, Chile. LACM
64-14, 1 1.8 x 6.8 x 3.0 mm. (219) Air-dried radula. Montemar, Valparaiso Province, Chile. LACM 75-30, width of ribbon 2.4 mm. (220)
Radula of small specimen. Ancon, Lima Province, Peru. LACM 74-21, width of held 0.5 mm, shell length 18.8 mm. (221) Juvenile shell.
Iquique, Chile. LACM 64- 16,5.4 x 3.3 x 1.6 mm. (222) Living specimens with mantle extended next to chiton Enoplochiton niger. Iquique,
Chile. LACM 75-12. (223) Cut shell. Mehuin, Valdivia Province, Chile. LACM 75-36, length 24 mm. (224) Mantle lobe, Islota Concon,
Valparaiso Province, Chile. LACM 75-31, length 12 mm.

closely related to any other species. It has some features in
common with F. crassa— sculpture essentially lacking and a
very elongate foramen. Both are limited to the intertidal zone,
although F. limbata is found at lower levels than is F. crassa.
The margin is broad and flat, unlike the upturned margin of
F. crassa. Also, F. limbata has relatively weak development
of mantle papillae and foot tubercles, whereas both of these
features are strongly developed in F. crassa. It probably has
more in common with F. nigra, with which it shares a rel-
atively smooth shell, similar weak development of mantle

papillae, and foot tubercles. However, the zoning of the cal-
citic layer is reversed; the darkest layer is at the surface in
F. nigra. Fissurella oriens has a rayed pattern similar to that
of F. limbata, but has some radial sculpture and an unzoned
calcitic layer.

Synonymy and Types. Two lots of F. limbata from the
Cuming Collection in the British Museum have been ex-
amined. For each lot the locality “Valparaiso” is written in
ink on the boards. Single specimens from each lot were il-
lustrated by Reeve, 1849, although neither of the two spec-

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McLean: Peruvian and Magellanic Fissurella 57

Figures 225 through 231. Fissurella crassa Lamarck, 1822. Mature shells. (225) Bahia Independencia, lea Province, Peru. AHF 375-35,
15.6 x 9.3 x 3.5 mm. (226) Iquique, Chile. LACM 64- 16, 56.9 x 32.3 x 14.0 mm. (227) Holotype, F. clypeiformis Sowerby . Locality unknown.
BMNH 40.6. 1 .9, 7 1 .4 x 43.4 x 1 5.6 mm. (228) Cumbres Borascosas, Tarapaca Province, Chile. LACM 75-14, 19.8 x 11.4 x 4.6 mm. (229)
Los Molles, Aconcagua Province, Chile. LACM 75-28, 92.0 x 61.0 x 27.4 mm. (230) Vina del Mar, Valparaiso Province, Chile. LACM 66-
46, 45.5 x 19.8 x 7.0 mm. (231) Mehuin, Valdivia Province, Chile. LACM 75-36, 80.3 x 49.0 x 22.1 mm.

imens illustrated by Sowerby in the “Conchological Illustra-
tions” are included in these lots. However, there is no
mistaking the type figure of Sowerby (1835b, figs. 66, 74,
internal and external view) as this species. Of the lots figured
by Reeve, BMNH 197582 has six specimens, length, 60.5
(anterior end chipped), 59.2, 49.2, 48.4, 33.6, and 28.2. The
largest was figured by Reeve (1849) and Dell (1971) and is
here refigured and designated the lectotype (Fig. 216). All
specimens in this lot lack Scurria- made scars and show the
complete normal color pattern of the outer zone of the calcitic
layer. BMNH 197581 has four specimens, length 67.3, 65.0,
63.1, and 61.1 mm. The largest was figured by Reeve, al-
though the scar of a Scurria was omitted; most of the rayed
outer zone of the calcitic layer was removed by the Scurria,
leaving the shell dark purple except for later growth stages.

The 65.0 mm shell is figured here (Fig. 215); it has a Scurria-
made scar but is less eroded.

Fissurella limbata var. multilineata Ziegenhom and Thiem,
1925, was based on two shells from Coquimbo, Chile, the
largest 16.5 mm in length. The figure shows the normal zigzag
pattern of purple lines found in all juveniles (Figs. 218, 221);
the name, therefore, has no systematic value.

Fissurella crassa Lamarck, 1822

Figures 225-237

Fissurella crassa Lamarck, 1822, 6(2): 11; Deshayes, 1830:
134; Sowerby, 1 835b: 1 , figs. 9, 11; Deshayes in Lamarck,
1836, 7:592; Gray, 1839:148, pi. 39, fig. 9; Orbigny, 1841:
472; Reeve, 1849, pi. 1, fig. 4; Hupe, 1854:240; Philippi,

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McLean: Peruvian and Magellanic Fissurella

Figures 232 through 237. Fissurella crassa Lamarck, 1822. Juvenile shell, radula, preserved and living specimens, cut shell, and mantle lobe.
(232) Juvenile shell. Miraflores, Lima Province, Peru. LACM 71-187, 6.9 x 3.5 x 11.6 mm. (233) Radula of small specimen. Cartagena,
Santiago Province, Chile. LACM 75-34, width of field 1.5 mm, shell length 22.1 mm. (234) Living animal submerged in tidepool. Iquique,
Chile. LACM 75-12. (235) Preserved specimen. Mehuin, Valdivia Province, Chile. LACM 75-36, shell length 46.9 mm. (236) Cut shell. Bahia
Moreno, Antofagasta Province, Chile. LACM 75-16, length 35 mm. (237) Mantle edge, Iquique, Chile. LACM 75-12, length 1 1 mm.

1860:181; Sowerby II, 1862:184, figs. 16, 17; Watson, 1886:
32; Pilsbry, 1890:154, pi. 34, figs. 51-53; Dali, 1909:177,
241, pi. 24, figs. 5, 6 [fig. looks like F. limbata\\ Ziegenhom
and Thiem, 1925:18, pi. 2, fig. 24; Carcelles and William-
son, 1951:255; Mermod, 1950:702; Riveros-Zuniga, 1951:
93, fig. 14; Pena, 1970:156; Dell, 1971:184; Marineovich,
1973:17, fig. 27; Ramirez-Boehme, 1974:30 [key],
Fissurella depressa Lamarck, 1822, 6(2): 15; Sowerby, 1835b:
1 (under F. crassa)-, Mermod, 1950:713 [type lost].
Fissurella clypeiformis Sowerby, 1825, app., p. vi; Sowerby,
1835b: 1 [under F. crassa].

Shell. Medium large (60-90 mm mature length), height
low to moderately elevated, elongate oval to very elongate,
some with elevated sides, others with elevated ends or with
both (so that shells rests on four comers). Sculpture smooth
except for early primary ribs that become broad and low,
forming wide marginal crenulations. Color caramel brown,
occasionally with faint rays of darker brown, surface eroded
if epibiotic Scurria is present, clean and uneroded if not.
Margin upwardly rounded at all growth stages. Shell margin
showing light inner zone and darker outer zone. Cut shells
also showing very thin lighter-colored layer, layer not evident

at edge. Foramen elongate and tripartite in young shells, very
elongate and constricted in middle in mature shells, posterior
portion much wider and longer than anterior; foramen in
mature shells beveled inward at ends. Aragonitic layer of
interior between callus and muscle scar pinkish gray through-
out and radially ridged.

Juvenile Shell. Sculpture of strong light-colored primary
ribs that become wide and low, interspaces wide and dark-
colored, two lateral white rays prominent. Shell becomes
dark overall on reaching length of 7 mm. Upwardly rounded
margin begins in earliest stages.

Mantle and Foot. Shell too low to accommodate large an-
imal within it. Cephalic tentacles brown; mantle lobe broad,
banded in light and dark, bands not matched by shell rays
but no doubt corresponding to rayed pattern of early juve-
niles. Papillae of upper edge very large, those of lower edge
much less developed. Side of foot with large, bulbous,white-
tipped tubercles. Papillae project at both ends of foramen.

Habitat. Mid-intertidal zone, in crevices on rocky reefs,
in surf-exposed or partially protected areas, occurring at higher
levels than the other species, tightly wedged in narrow crev-
ices during low tide. The upturned margin may be an ad-
aptation to the rather cramped position of the animal when

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McLean: Peruvian and Magellanic Fissurella 59

exposed at low tide. Most specimens have a single Scurna
parasitica on the shell. Bretos (1978, 1980) has studied growth
in this species.

Distribution. Huarmey, Peru (10°06' S) (AMNH 134571),
to Punta Pulga, Isla de Chiloe, Chile (42°06' S) (Dell, 1971).
Dali ( 1 909) cited the Galapagos Islands, Ecuador, and USNM
59260 is so labeled, but this record is discounted. The south-
ern limit is uncertain. The species is known from the north-
western tip of Isla de Chiioe but may extend farther south.
I did not find it on the eastern side Isla de Chiloe, where
there is little exposure to surf. The record of Riveros-Zuniga
(1951) from Fuerte Bulnes in the Magellan Strait is rejected.

Number of Lots Examined. 88 (LACM 31, AMNH 23,
ANSP 7, MACN 7, MNHN 10, USNM 10).

Taxonomic History. Fissurella crassa has been correctly
interpreted by authors. It differs sufficiently from all other
species that it can not be confused with any of them.

Abundance and Use. Common throughout its range and
widely exploited for food. The upper intertidal habitat is so
accessible that large specimens are seldom seen. It is known
as the “lapa de sol,” because it occurs relatively high, where
it is exposed to the sun.

Characteristics and Variability. The most characteristic
features of F. crassa are the uniformly brown color of the
shell, the lack of radial ribs other than the broad undulations,
the lavender staining of the interior, the great enlargement
of the posterior end of the foramen, and of most importance,
the upturned margin. The foot also has the strongest devel-
opment of tubercles in any of the species. Variation is not
extensive and is limited to rather minor differences in height
and amount of elevation of the sides or ends.

Affinity and Comparisons. The closest affinity of F. crassa
seems to be with F. limbata. In both species the foramen
remains elongate, the sculpture is undulating, and strong ribs
are lacking. As in F. limbata there is a thin outermost zone
to the calcitic layer that is lighter in color. The upturned
margin of F. crassa is unique. Although the margin of F.
maxima is also unusual, in that species only the junction
between the margin and the internal aragonitic layer is round-
ed.

Synonymy and Types. The original description of F. crassa
Lamarck mentioned no locality. Mermod (1950:702), in his
report on types of Lamarck in the Geneva collection, gave
notes on a specimen 73 mm in length, which he considered
to be Lamarck’s original.

The locality for the very briefly described F. depressa La-
marck. 1822, was given as the “Indian Ocean.” Sowerby
( 1 835b: 1 ) stated under F. crassa: “Lamarck’s F. depressa is
only a worn fragment of this species, as Mr. Gray informs
me.” Presumably, Gray had examined the Lamarck collec-
tion. Mermod (1950:713) reported that the type specimen
now is lost.

The holotype of F. clypeiformis Sowerby, 1825, is an un-
worn specimen of F. crassa, BMNH 40.6.1.9, 70.4 mm in
length (Fig. 227). It was described without locality and was

placed in synonymy shortly after publication by Sowerby
( 1 835b) himself.

Fissurella bridgesii Reeve, 1 849

Figures 238-253

Fissurella bridgesii Reeve, 1849, pi. 3, fig. 15; Hupe, 1854:
238; Philippi, 1860:180; Sowerby II, 1862:184, figs. 21,
22, 34; Pilsbry, 1890:151, pi. 30, fig. 3; Dali, 1909:241;
Riveros-Zuniga, 1951:121, fig. 28; Dell, 1971:183, pi. 3,
figs. 8, 9; Ramirez-Boehme, 1974:31 [key].

Shell. Medium large (65-90 mm mature length), low; out-
line elongate-oblong, lateral profile mostly straight, but some
specimens with slightly raised sides or ends. Sculpture of fine
low ribs in young shells; mature shell nearly smooth, with
traces of very broad, irregular radial ribs and irregular radial
striae. Color grayish or reddish brown, with faint lighter and
darker rays, two lateral rays often more prominent; ground
color changing gradually to light gray in large shells. Margin
broad in growing shells, flat and often beveled out and up
(except at front end) so that its edge is not in contact on a
flat surface; margin narrower and rounded in mature shells.
Calcitic layer zoned, inner zone dark reddish brown, changing
to translucent gray at outer surface, outer zone usually lighter.
Foramen unusually large and oval at all growth stages, broad-
ly tripartite in some and showing two projections at sides,
others lacking these projections.

Juvenile Shell. None seen under 20 mm in length; primary
and secondary ribs weak, nearly equal in size at shell length
of 20 mm; foramen large and oval at this shell length. Young
shells have two lateral white rays.

Mantle and Foot. Body too large to retract beneath low
shell; the mantle lobe broad and thin, enveloping shell edge.
Papillae of upper edge tongue-shaped, finely branched, those
of lower lobe similarly shaped but smaller and more nu-
merous. Mantle lobe vertically banded with light and dark
to correspond to weak pattern of rays on shell. Side of foot
marbled; in preserved specimens, dark tubercles are sur-
rounded by lighter areas.

Habitat. Fissurella bridgesii occurs on surf exposed rocks
surrounded by sandy areas in the intertidal zone and the
shallow sublittoral zones, a habitat entirely unlike that of the
others. I missed seeing this species because I did not examine
this habitat in Chile in 1975. However, the occurrence of F.
bridgesii has been detailed by Bretos ( 1 979). Shells are usually
overgrown with an algal mat. Only rarely do shells have
attachment scars of Scurria, indicating that they usually live
deeper than the mid- to upper intertidal zone preferred by
Scurria. The Californian acmaeid limpet Notoacmea fenes-
trata (Reeve, 1855) has a comparable habitat on rocks sur-
rounded by sand.

Distribution. Isla Guanape, Peru (8°32' S) (LACM 74-3,
McLean), to Rio Bio-bio, Concepcion Province, Chile (36°48'
S) (LACM 75-35, McLean). There is but a single record from

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McLean: Peruvian and Magellanic Fissurella

Figures 238 through 248. Fissurel/a bridgesii Reeve, 1849. Mature shells. (238) 3-5 m, Isla Guanape, Peru. LACM 74-3, 66.0 x 46.0 x
15.0 mm. (239) Iquique, Chile. LACM 90802, 22.4 x 15.7 x 5.3 mm. (240) Same locality. LACM 90802, 20.0 x 12.8 x 5.9 mm. (241)
Iquique, Chile. LACM 90803, 65.5 x 41.3 x 14.5 mm. (242) Paposo, Antofagasta Province, Chile. LACM 54764, 44.4 x 28.6 x 9.3 mm.
(243) Quintero, Valparaiso Province, Chile. USNM 48221, 89.3 x 57.0 x 23.5 mm. (244) Paralectotype, F. bridgesii Reeve. Quintero, Val-
paraiso Province, Chile. BMNH 197566, 77.6 x 49.1 x 1 14.3 mm. (245) Lectotype, F. bridgesii Reeve. Quintero, Valparaiso Province, Chile.
BMNH 197566, 68.0 x 47.5 x 13.7 mm. (246) Valparaiso, Chile. USNM 56255, 33.0 x 19.5 x 10.2 mm. (247) Valparaiso, Chile. AMNH
20055, 52.8 x 36.0 x 10.4 mm. (248) Rio Bio-bio, Concepcion Province, Chile. LACM 75-35, 42.4 x 26.5 x 9.0 mm.

Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella 61

249 253

Figures 249 through 253. Fissurella bridgesii Reeve, 1 849. Living and preserved specimens, mantle lobe and radula. (249) Three living
specimens in place. Tocopilla, Antofagasta Province, Chile, photo courtesy A. Viviani. (250) Mantle lobe. Iquique, Chile. LACM 90803, length
15 mm. (251) Mantle lobe. 3-5 m, Isla Guanape, Peru. LACM 74-3, length 20 mm. (252) Preserved specimen. Iquique, Chile. LACM 90803,
shell length 65.5 mm. (253) Air-dried radula. Iquique, Chile. LACM 90804, width 3.5 mm, shell length 74.4 mm.

Peru, the living specimen I collected in 1974 at Isla Guanape
(Fig. 238). I have examined specimens from the following
localities in Chile: Iquique, Paposa, Quintero, Valparaiso,
and Rio Bio-bio, near Concepcion.

Number of Lots Examined. 15 (LACM 8, AMNH 1,MACN
1, MNHN 3, USNM 2). This species is uncommon in the
collections examined.

Taxonomic History. Accounts of F. bridgesii in the liter-
ature prior to the report of Bretos (1979) were copies of the
original description of Reeve (1849). Dell (1971) considered
this taxon a synonym of F. latimarginata. The validity of
the species was confirmed in 1977, when I compared a pre-
served specimen sent to me for identification by M. Bretos
with the syntypes from the British Museum then on loan at
the LACM. Although I was unable to find living examples

of this species during my fieldwork in Chile, puzzling beach-
worn shells were collected at several localities, and I later
discovered that I had purchased live specimens at the market
in Iquique. The specimen I had earlier collected in Peru at
Isla Guanape was then recognized as F. bridgesii. The spe-
cialized habitat of this species accounts for its scarcity in
collections. It may be, however, that Ramirez-Boehme (1974)
had recognized the species, because his key mentions the
characteristic upwardly beveled margin (“bordes laterales re-
fiejados hacia arriba”), a feature not discussed elsewhere in
the literature.

Abundance and Use. Fissurella bridgesii is used for food
at Iquique, and it may have more economic importance than
suggested by the paucity of specimens in collections. Ac-
cording to M. Bretos, it is known to the fishermen as the

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McLean: Peruvian and Magellanic Fissurella

“lapa jerguilla,” or the “lapa de arena” (sand). The jerguilla
is a fish, Aplodactylus punctatus, which has a color pattern
resembling that of the body of F. bridgesii.

Characteristics and Variability. The most characteristic
features of F. bridgesii are the lack of regularly defined radial
sculpture in mature shells, a relatively large foramen, a faintly
rayed pattern on a gray-brown ground, and the broad, flat
margin, which is beveled upward in young, growing speci-
mens. Some specimens have an elongate foramen that is
bidentate on the sides; others of the same size may have a
more oval foramen. Color patterns include faintly rayed spec-
imens and some uniformly gray shells lacking traces of rays.
The northernmost specimen from Isla Guanape, Peru, lacks
rays (Fig. 238).

Affinity and Comparisons. Fissurella bridgesii most resem-
bles F. latimarginata. The normal weakly rayed pattern of
F. bridgesii can be related to the unusual rayed color form
of F. latimarginata. Both have a broad margin and a lighter
outer zone to the calcitic layer, but only F. bridgesii has an
upwardly beveled margin. Both species have similar elabo-
ration of the tongue-shaped papillae of the mantle edge, more
so than do other species. In its light outer shell layer it also
resembles F. limbata, but may be distinguished from that
species in lacking the purple coloration to the shell. Fissurella
bridgesii also resembles F. pulchra in its low profile, weakly
contrasting pattern of rays, and the lighter outermost zone
to the calcitic layer. It differs in lacking the speckled pattern
of F. pulchra and in having a larger foramen and a more
pronounced upward-beveled margin. Although the size of
the foramen varies somewhat in F. bridgesii, it is always
larger than that of similarly sized specimens of F. pulchra.
Before I understood F. bridgesii, my guess was that young
beach-worn specimens, such as the one from Concepcion
(Fig. 248), were most likely to be variants of F. pulchra.

Synonymy and Types. There are four syntype specimens
of F. bridgesii Reeve, 1 849, from Quintero, Valparaiso Prov-
ince, Chile (32°47' S), BMNH 197566, lengths 77.7, 68.0,
61.8, and 46.0 mm. The 68.0 mm specimen was figured by
Reeve (1849), Sowerby II (1862), and Dell (1971), and is
here designated the lectotype (Fig. 245); the larger 77.7 mm
specimen has been figured only by Sowerby II (1862). These
two large specimens (Figs. 244, 245) illustrate two of the
most variable features of the species. They are nearly of the
same width, but one is much more elongate. The larger,
elongate specimen has an oval foramen; the broader speci-
men has a bidentate foramen. The two remaining paralec-
totypes are broad; the 61.8 mm specimen has an oval fora-
men; the 46.0 mm specimen has an elongate foramen with
the bidentate structure nearly imperceptible. Shape of the
foramen is therefore not correlated with shell proportions.

Fissurella pulchra Sowerby, 1835

Figures 254-267

Fissurella pulchra Sowerby, 1835a: 124; Sowerby, 1835b:3,
fig. 24; Reeve, 1849, pi. 2, fig. 9; Hupe, 1854:244; Philippi,
1860:181; Sowerby II, 1862:184, fig. 31; Pilsbry, 1890:
151, pi. 33, fig. 50; Dali, 1909:242; Riveros-Zuniga, 1951:

120, fig. 27; Dell, 1971:191, pi. 3, fig. 5; Ramirez-Boehme,

1974:30 [key].

Shell. Medium-sized (35-75 mm mature length), consis-
tently low; outline elongate oval, tapered anteriorly, sides
slightly raised relative to ends. Radial sculpture in early growth
stages consisting of low, rounded primary ribs, becoming
obsolete when shell reaches length of 20 mm; mature shell
smooth except for faint radial striae. Color grayish lavender
to pink, with alternating lighter and darker rays; entire surface
with fine reddish speckles or zigzag markings especially pro-
nounced near foramen; concentric interruptions to rays
changing color from pink to gray or brown in some. Margin
of moderate width, flat, reddish gray, zoned to make edge
slightly lighter in color; broken shells showing lighter outer
zone of calcitic layer; margin becoming very narrow in large,
full grown shells. Foramen elongate and tripartite in young
shells, only slightly less elongate in mature shells; interior
callus bordered by pink colored ring in attachment region.

Juvenile Shell. Primary ribs rounded, coinciding with
lighter rays, speckled pattern conspicuous. Sides of shell raised
bordering foramen, indicating that earliest stage is more con-
ical.

Mantle and Foot. Not retractable in flattened shell; mantle
lobe normally extending well over shell edge. Cephalic ten-
tacles lavender, yellowish at tips. Mantle lobe pinkish gray
or brown, faintly banded to match rayed pattern, lined con-
centrically with brown. Papillae of upper edge well devel-
oped, those of lower edge more numerous and smaller. Side
of foot same color, tubercles well developed.

Habitat. Low intertidal and sublittoral zones in crevices
or on undersides of large rocks in deep tidepools, protected
from direct exposure to surf. Shells are free of algal incrus-
tations; epibiotic organisms on the shell are bryozoans and
spirorbid polychaetes.

Distribution. Salaverry, Peru (8°14' S) (USNM 368490, W.
Schmitt), to Rio Bio-bio, Concepcion Province, Chile (36°48'
S) (LACM 75-35, McLean). Except for F. bridgesii, this is
the least common species in the Peruvian Faunal Province.
I have found small specimens at most localities in central
Peru. In northern Chile it was rare at Iquique; at Antofagasta
there was no trace of it, even in the beach-worn shell debris.
It was more common in central Chile, where I found living
specimens on undersides of boulders in tidepools at Los
Molles, Aconcagua Province. Beach-worn specimens were
found at other localities in central Chile.

Number of Lots Examined. 21 (LACM 13, AMNH 1,ANSP
3, USNM 4, none at MACN or MNHN).

Taxonomic History. Most authors have merely copied the
original account of F. pulchra, except for Pilsbry ( 1 890), who
emphasized the characteristic speckled pattern. The habitat
is cryptic, and beach-worn shells are sufficiently scarce that
the species has escaped notice in recent years. Riveros-Zuniga
(1951) merely quoted previous authors, and Pena (1970) did
not mention it. The collection from Iquique of Marincovich
(1973) did not include it. Ramirez-Boehme ( 1974) included
it in his key but did not mention the conspicuous speckles.

Abundance and Use. The species is sufficiently uncommon

Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella 63

Figures 254 through 259. Fissurella pulchra Sowerby, 1835. Mature shells. (254) Miraflores, Lima Province, Peru. LACM 71-187, 24.2 x
13.5 x 4. | mm. (255) Pupudo, Aconcagua Province, Chile. LACM 54656, 78.5 x 56.0 x 17.0 mm. (256) Paralectotype, F. pulchra Sowerby.
Valparaiso, Chile. BMNH 197580, 39.6 x 25.8 x 6.6 mm. (257) Lectotype. F. pulchra Sowerby. Valparaiso, Chile. BMNH 197580, 62.2 x
40.2 x 12.8 mm. (258) Cartagena, Santiago Province, Chile. LACM 75-34, 68.5 x 50.8 x 14.3 mm. (259) Rio Bio-bio, Concepcion Province,
Chile. LACM 75-35, 37.5 x 23.3 x 6.6 mm.

to be negligible as a food resource. Large specimens are par-
ticularly uncommon; living specimens that I found were about
half the size of shells in the type lot, although I found some
beach-worn shells that approached the maximum size. The
largest specimen 1 have examined is 78.5 mm in length, from
Papudo, Aconcagua Province, Chile, donated to the LACM
by J. Ramirez-Boehme (Fig. 255). The species has no com-
mon name in northern Chile, according to M. Bretos.

Characteristics and Variability. Fissurella pulchra is aptly
named. It is rayed in pinkish brown and gray and is the only
species in which a speckled pattern persists through all growth
stages. It is one of the least variable species; I have noticed
no unusual color forms. Shells are consistently low and ta-
pered anteriorly. Most shells are relatively smooth, although
the weak primary ribs may be more pronounced in some
specimens.

The radula of F. pulchra (Figs. 266, 267) is the most dis-
tinctive among the Peruvian-Magellanic species. The inner
laterals have longer overhanging cusps than the other species.

and the enlarged outer lateral is unique in having a concave
edge to the second cusp.

Affinity and Comparisons. On shell characters, F. pulchra
surely hts within the group having broad primary ribs and
no secondary ribs. It is closest to F. bridgesii, with which it
shares similar proportions, general overall coloration, and
faint rays. It differs in its speckled pattern, more consistently
narrowed foramen, and pink bordered internal callus. The
margin is less broad than that of F. bridgesii; it is similarly
beveled upward but to a lesser extent than in F. bridgesii.
The tapered anterior end of F. pulchra resembles the con-
dition of the otherwise not closely related F. latimarginata
and F. cumingi.

Synonymy and Types. There are 5 syntypes of F. pulchra
Sowerby, 1835, from Valparaiso, Chile, BMNH 197580,
lengths 70.8 (broken in half), 62.2, 48.7, 39.6, and 24.0 mm.
The largest of these has not been figured. The 62.2 mm
specimen was figured originally by Sowerby (1835b), then by
Reeve (1849) and by Dell (1971); it is here designated the

64 Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella

Figures 260 through 267. Fissurella pulchra Sowerby, 1835. Living specimens, mantle lobe, juvenile shells, radulae. (260) Living specimen
with mantle expanded, in tidepool. Cumbres Borascosas, Tarapaca Province, Chile. LACM 75-14. (261) Same specimen, ventral view. (262)
Living specimen on underside of overturned boulder. Los Molles, Aconcagua Province, Chile. LACM 75-28. (263) Mantle lobe. Cumbres
Borascosas, Tarapaca Province, Chile. LACM 75-14, 8 mm. (264) Juvenile specimen. Los Molles, Tarapaca Province, Chile. LACM 75-28,
5.8 x 3.5 x 1.3 mm. (265) Juvenile shell. Pucusana, Lima Province, Peru. LACM 72-76, 11.7 x 6.7 x 2.0 mm. (266) Radula of small shell.
Same locality. LACM 72-76, 1.5 mm, shell length, 22.1 mm. (267) SEM view of radula. Los Molles, Aconcagua Province, Chile. LACM 75-
28, width of field 0.6 mm.

lectotype. The lectotype and the 39.6 mm paralectotype are
figured here (Figs. 256, 257).

LACM LOCALITIES FOR FIGURED
SPECIMENS

[All collections from intertidal zone, unless otherwise indi-
cated.]

AHF 375-35. Bahia Independencia, lea Province, Peru( 14°14'
S, 76°12.7' W), 13 January 1935.

AHF 380-35. Bahia Independencia, lea Province, Peru (14° 14'
S, 76°08.5' W), 14 January 1935.

AHF 828-38. Bahia San Juan, lea Province, Peru (15°20.7'
S, 75°09.3' W), 8 February 1938.

62-26. Ancud, Chiloe Province, Chile (41°52' S, 73°05' W),
H.C. McMillin, 24 March 1962.

64-16. Iquique (near Ave. Baquedena), Tarapaca Province,
Chile (20°13' S, 70°10' W), L. Marincovich, June through
August, 1964.

66-46. Renaca, Vina del Mar, Valparaiso Province, Chile
(33°06' S, 7 1°50' W), R. Seapy, 24 July 1966.

70- 68. Iquique (10 km S), Tarapaca Province, Chile (20°15'
S, 70°09' W), L. Marincovich, July, 1970.

71- 187. Miraflores, Lima Province, Peru (12°08' S, 77°04.5'
W), T. Bratcher, 24 February 1971.

7 1 -277. Bahia York, Isla de los Estados, Argentina (54°47. 1 1 '
S, 64°1 7.9' W), 5 May 1971.

Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella 65

7 1 -284. Puerto San Juan del Salvamento, Isla de los Estados,
Argentina (54°43.9' S, 63°52' W), 13 May 1971.

72-76. 0-5 m, Pucusana, Lima Province, Peru (12°30' S,
76°49' W), J.H. McLean, 30 March 1972.

72-77. Laguna Granda, lea Province, Peru ( 1 4° 1 8' S, 76°15'
W), J.H. McLean, 31 March 1972 (beach-worn shells).
72-79. Paracas, lea Province, Peru (13°49' S, 76°14.5' W).
J.H. McLean, 2 April 1972 (beach-worn shells).

74-2. NE side, Isla Guanape, La Libertad Province, Peru
(08°32' S, 78°58' W), J.H. McLean, 18 January 1974.

74-3. 3-5 m, NE side, Isla Guanape, La Libertad Province,
Peru (08°32' S, 78°58' W), J.H. McLean, 18 January 1974.
74-21. 1-2 m, Playa Hermosa, Ancon, Lima Province, Peru
(1 1°47' S, 77°1 1.5' W), J.H. McLean, 26 January 1974.

74- 24. \-4 m, Isla San Lorenzo, Lima Province, Peru(12°06.7'
S, 77°13' W), J.H. McLean, 29 January 1974.

75- 10. Pozo Toyo (S of Iquique), Tarapaca Province, Chile
(20°25' S, 70° 10.5' W), J.H. McLean, 29 September and 1
October 1975.

75-12. Iquique (at Marine Laboratory, Universidad del
Norte), Tarapaca Province, Chile (20°1 5.5' S, 70°08' W), J.H.
McLean, 2 October 1975.

75-14. Cumbres Borascosas, Tarapaca Province, Chile
(20°42'S, 70°1 1.5' W), J.H. McLean, 3 October 1975.

75-15. Antofagasta (S end of city), Antofagasta Province,
Chile (23°42' S, 70°27' W), J.H. McLean, 5 and 6 October
1975.

75-16. Bahia Moreno, Antofagasta Province, Chile (23°28'
N, 70°31' W), J.H. McLean, 7 October 1975 (beach shells).
75-17. [Shore opposite] Isla Santa Maria, Antofagasta Prov-
ince, Chile (23°25' S, 70°36' W).

75-18. Punta Jara, Antofagasta Province, Chile (23°49' S,
70°29' W), J. Tomicic, 8 October 1975.

75-19. Los Colorados, Antofagasta Province, Chile (23°29'
N, 70°22' W), J.H. McLean, 9 October 1975.

75-20. 2-5 m, Antofagasta (S end of city), Antofagasta Prov-
ince, Chile (23°42' S, 70°27' W), J.H. McLean, 10 October
1975.

75-21. 7-20 m, El Rincon de Mejillones, Antofagasta Prov-
ince, Chile (23°02' S, 70°31' W), J.H. McLean, 11 October
1975

75-23. 2-4 m, El Rincon de Mejillones, Antofagasta Prov-
ince, Chile (23°05' S, 70°30' W), J. Tomicic, 12 October 1975.
75-25. Bahia Herradura, Coquimbo Province, Chile (29°59'
S, 7 1°22' W), J.H. McLean, 14 October 1975.

75-27. Bahia El Teniente, Coquimbo Province, Chile (30°58'
S, 7 1°39' W), J.H. McLean, 15 October 1975.

75-28. Los Molles, Aconcagua Province, Chile (32°14' S,
71°32' W), J.H. McLean, 16 through 18 October 1975.
75-29. Los Modes, Aconcagua Province, Chile (32°14' S,
71°32' W), J.H. McLean, 16 October 1975 (shed pile).
75-30. Montemar (at Estacion de Biologia Marina), Valpa-
raiso Province, Chile (32°57' S, 71°32' W), J.H. McLean, 19
and 20 October 1975.

75-31. Islota Concon, N of Vina del Mar, Valparaiso Prov-
ince, Chile (32°52' S, 71°33' W), J.H. McLean, 21 and 22
October 1975.

75-34. Cartagena, Santiago Province, Chile (33°33' S, 71°38'
W), J.H. McLean, 23 October 1975.

75-35. Rio Bio-bio, Concepcion Province, Chile (36°48' S,
73°1 1 ' W), J.H. McLean, 29 October 1975.

75-36. Mehuin, Valdivia Province, Chile (39°23' S, 73°14'
W), J.H. McLean, 31 October and 2 November 1975.
75-37. [Island off] Mehuin, Valdivia Province, Chile (39°26'
S, 73°16' W), J.H. McLean, 1 November 1975.

75-39. Pargua, Canal de Chacao, Llanquihue Province, Chile
(41°47' S, 73°28' W), J.H. McLean, 3 November 1975.
75-40. Guabun, Isla de Chiloe, Chiloe Province, Chile (41°50'
S, 74°02' W), J.H. McLean, 4 November 1975.

75-41. Pumalin, Golfo Corcovado, Chiloe Province, Chile
(42°42' S, 72°52' W), J.H. McLean, 4 through 6 November
1975.

75-42. 2-4 m, Punta Estero, Isla Talcon, Golfo Corcovado,
Chiloe Province, Chde (42°46' S, 72°56' W), J.H. McLean,
6 November 1975.

75-43. 4-13 m, Islota Nihuel, Golfo Corcovado, Chiloe
Province, Chile (42°38' S, 72°57' W), J.H. McLean, 7 No-
vember 1975.

75-44. Islota Nihuel, Golfo Corcovado, Chiloe Province, Chile
(42°38' S, 72°57' W), J.H. McLean, 7 November 1975.
75-45. Quellon, Isla de Chiloe, Chiloe Province, Chile (43°09'
S, 73°37' W), J.H. McLean, 8 November 1975.

75-47. Isla Laitec, off SE side Isla de Chiloe, Chiloe Province,
Chile (43°14' S, 73°36' W), 9 November 1975.

75-48. Fuerte Bulnes, Peninsula Brunswick, Magellan Strait,
Magallanes Province, Chile (53°38' S, 70°54.5' W), J.H.
McLean, 16 November 1975.

75-49. Puerto el Hambre, Peninsula Brunswick, Magellan
Strait, Magallanes Province, Chile (53°37' S, 70°56' W), J.H.
McLean, 16 and 19 November 1975.

78-88. Punta Ninfas, Golfo Nuevo, Chubut Province, Ar-
gentina (42°56. 5' S, 64°19.5' W), J.H. McLean, 18 July 1978.
78-90. Punta Cracker, Golfo Nuevo, Chubut Province, Ar-
gentina (42°56.5' S, 64°30' W), J.H. McLean, 19 July 1978.

ACKNOWLEDGMENTS

I am especially indebted to those who made the arrangements
for fieldwork in South America.

In Peru, I was aided by Enrique M. del Solar of Lima, and
Norma Chirichigno and Victor Alamo, of the Instituto del
Mar in Callao. Collecting excursions were arranged out of
Lima in March, 1972, and in January, 1974, 1 was privileged
to join a cruise of the Peruvian Government research vessel
SNP-1 to Isla Guanape and the Islas Lobos Afueras. Other
Peruvian biologists who have been helpful through exchanges
and correspondence include Wenceslao Medina, G. Mario
Pena, and Carlos Paredes Quiroz.

In Chile, I joined the Marine Communities Expedition of
Robert T. Paine of the University of Washington, October-
November, 1975. Expenses were partially underwritten by
the National Science Foundation (DES 75-14378, R.T. Paine,
principal investigator). Other members of the expedition,
Carol Sturgess, Ken Sebens, and Tom Suchanek, all from the
University of Washington, also helped in the collecting.

66 Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella

Arrangements in Chile were made by Juan Carlos Castilla,
Universidad Catolica de Chile, Santiago, who joined us for
part of the expedition, made the facilities of the Universidad
Catolica available to us, and arranged to ship the collections
to me. Others who assisted us were C.A. Viviani and Marta
Bretos, then of Universidad del Norte, Iquique; Jorge To-
rnicic, Universidad de Antofagasta, Antofagasta; Miguel Pa-
dilla and Luis Ramorino, Instituto de Oceanologia, Univer-
sidad de Valparaiso, Valparaiso; Olga Aracena, Universidad
de Concepcion, Concepcion; Hugo Campos, Universidad
Austral de Chile, Valdivia; and Italo Compodonico, Instituto
de la Patagonia, Punta Arenas. Every effort was made to assist
us by our Chilean hosts.

Fieldwork in Argentina in July, 1978, was arranged by
William J. Zinsmeister of Ohio State University, chief sci-
entist on cruise 783 of the R/V HERO to the vicinity of the
Peninsula Valdez.

Collections made by Luie Marincovich in Iquique in 1964
and 1980 were instrumental in introducing me to the Pe-
ruvian-Magellanic fauna.

I also thank Paul Dayton of Scripps Institution of Ocean-
ography for making a very productive effort at my request
to collect mollusks during his research cruises in southern
Chile on the R/V HERO in 1972 and 1973.

Edmundo Martinez, of Antofagasta, kindly provided me
with fossil specimens of Fissurella species from that region
of Chile.

I am particularly indebted to K.M. Way of the British
Museum for arranging the loan of the types of Fissurella
preserved in that institution and providing much necessary
information through correspondence.

Philippe Bouchet and B. Metivier of the Paris Museum
kindly searched for the types and provided loan material of
a number of other species described by early French authors.

In 1978 I visited the Museo Nacional de Historia Natural,
Santiago, where I was assisted by Maria Codoceo and Ni-
baldo Bahamonde. On the same trip I visited the Museo
Argentino de Ciencias Naturales, Buenos Aires, where I was
assisted by Martinez Fontes.

I thank William K. Emerson of the American Museum of
Natural History, Joseph Rosewater and Richard S. Houbrick
of the United States National Museum, and Robert Robert-
son and George M. Davis of Academy of Natural Sciences,
Philadelphia, for making the collections of their institutions
available to me and arranging loans of specimens. Loans were
also arranged by Kenneth J. Boss of the Museum of Com-
parative Zoology, Harvard, and Richard K. Dell of the Na-
tional Museum of New Zealand.

Marta Bretos, now of the Universidad de la Frontera, Te-
muco, Chile, has reviewed the manuscript, sent specimens,
and supplied me with the common names of Fissurella species
as used in northern Chile.

Polychaete shell borers were identified by K. Fauchald of
the U.S. National Museum of Natural History.

C.J. Risso-Dominguez of Buenos Aires provided me with
specimens and information about the occurrence of Fissu-
rella species in Argentina and reviewed an early draft of the
manuscript.

Carole S. Hickman, University of California, Berkeley,
provided the SEM micrographs of radulae; Heinz A. Low-
enstam, California Institute of Technology, Pasadena, con-
tributed the SEM micrographs of the shell of a small speci-
men of Fissurella latimarginata (Figs. 4-7). Edward Wilson
of the LACM section of Invertebrate Paleontology cut spec-
imens for the examination of shell layers.

Photographs of shells and air-dried radulae are the work
of Bertram C. Draper, museum volunteer. Museum photog-
rapher Dick Meier photographed the cut shells. Museum
illustrators Mary Butler and Caryl Maloof assisted in the
preparation of the figures. Museum volunteer Jo-Carol Ram-
saran helped in curatorial tasks and library searches necessary
to the work.

I thank H. Andrade for translating the abstract to Spanish
and Fernando Jara for reviewing the ecological remarks.

Comments and criticisms of various drafts of this paper
have been received from Eugene Coan, Cliff Coney, J. Chris-
tiaens, Myra Keen, Patrick I. LaFollette, David R. Lindberg,
and Robert T. Paine. The submitted manuscript was re-
viewed by William K. Emerson, Barry Roth, and Edward
Wilson.

LITERATURE CITED

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de Fissurella latimarginata (Sowerby, 1834) en Toco-
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Boutan, L. 1885. Recherches sur l’anatomie et le devel-
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Bretos, M. 1978. Growth in the keyhole limpet Fissurella
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. 1979. Observaciones sobre Fissurella bridgesii

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. 1980. Age determination in the keyhole limpet

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. 1982. Biologia de Fissurella maxima Sowerby

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(Mollusca: Archaeogastropoda) in northern Chile. 2.
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Bretos, M., and C. Jiron. 1980. Trematodes in Chilean
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Bruguiere, J. 1789. Histoire naturelle des vers. Paris, v. 1,
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Carcelles, A.R. 1950. Catalogo de los moluscos marinos
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Submitted 25 October 1983; accepted for publication 3 May
1984.

70 Contributions in Science, Number 354

McLean: Peruvian and Magellanic Fissurella

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Number 355
6 December 1984

AN EARLY HEMINGFORDIAN (EARLY MIOCENE)
FOSSIL VERTEBRATE FAUNA FROM BORON,

WESTERN MOJAVE DESERT, CALIFORNIA

David P. Whistler

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AN EARLY HEMINGFORDIAN (EARLY MIOCENE)
FOSSIL VERTEBRATE FAUNA FROM BORON,
WESTERN MOJAVE DESERT, CALIFORNIA

David P. Whistler1

ABSTRACT. A small assemblage of fossil vertebrates has been ob-
tained from the Arkose Member of the Kramer Beds in the upper
part of the Tropico Group at Boron, California. Known as the Boron
Local Fauna (new name), it displays affinities with the Tick Canyon
and Phillips Ranch Local Faunas of southern California and the
fossil assemblages from the Runningwater and Batesland Formations
of Nebraska and South Dakota, respectively, the latter used as the
standard for the earlier part of the Hemingfordian land mammal
age. These correlations and radiometric dates with a range of 18.3
± 0.6 ma to 20.3 ± 0.7 ma from the underlying Saddleback Basalt
support an early Hemingfordian age for the Boron Local Fauna.

Prior to the discovery of this fossil assemblage, considerable un-
certainty existed about the age of scattered occurrences of volcanic
flows and volcaniclastic rocks in the western Mojave Desert which
were restricted and redefined as the Tropico Group by Dibblee
(1958a). Based on lithologic correlations, ages ranging from Middle
Miocene to Early Pliocene had been suggested for the T ropico Group.
The presence of the early Hemingfordian Boron Local Fauna in the
uppermost unit of the Tropico Group limits the age of this entire
sequence to pre-Middle Miocene. The well-preserved assemblage
contains at least 15 vertebrate taxa. Two new species are described,
Cupidinimus boronensis n. sp. (Rodentia, Heteromyidae) and Ale-
tomeryx occidentalis n. sp. (Artiodactyla, Palaeomerycidae). There
are also two lizards which are not diagnostic at the family level, two
snakes, cf. Calamagras sp. (Boidae) and cf. Paracoluber sp. (Colu-
bridae), a lagomorph, Archaeolagus sp. or Hypolagus sp. (Leporidae),
three additional rodents, IMiospermophilus sp. (Sciuridae), Moo-
komys sp. and Trogomys cf. T. rupinimenthae (Heteromyidae) and
five additional artiodactyls, Merychyus cf. M. minimus and IMer-
ychyus sp. (Merycoidodontidae), cf. Hesperocamelus sp. (Camelidae)
and two other camels not identified to genus. Conspicuous by their
absence are eomyid and large geomyoid rodents, horses, dromo-
merycine palaeomerycids, and antilocaprids typical of most Middle
Miocene assemblages.

INTRODUCTION

In March, 1964, the geology staff of the United States Borax
and Chemical Corporation mine at Boron, California, re-
ported uncovering a pocket of fossil bones during operations

in the open pit mine (Figure 1). An initial sample collected
by the Department of Geology at the University of California
at Riverside was recognized as representing a Miocene as-
semblage older than any previously known from the central
Mojave Desert. The fossils were recovered from the Kramer
Beds, sediments which were correlated with other scattered
continental sequences in the Mojave Desert, all of which were
considered to be Late Miocene or Pliocene in age. These
fossil vertebrates, hereafter called the Boron Local Fauna,
demonstrated that the Kramer Beds were older than the rich-
ly fossiliferous deposits of the Barstow Formation which con-
tained the oldest fossil faunas then known from the central
Mojave Desert and necessitated re-thinking the geologic his-
tory of this area. Subsequent discoveries (Woodbume et al.,
1974; Woodbume, Miller, and Tedford, 1982) have led to
recognition of even earlier Miocene assemblages, but the
Boron Local Fauna was the first well-preserved pre-Barsto-
vian assemblage from the Mojave Desert.

The Boron Local Fauna is markedly different in taxonomic
composition from the well-documented faunas of the Bar-
stow Formation and other fairly widespread Barstovian
(Middle Miocene) assemblages in southern California. The
Boron Local Fauna is dominated by four groups (heteromyid
rodents, oreodonts, camels, and palaeomerycids) which, ex-
cept for camels, are rare or absent in Barstovian assemblages
of southern California. In contrast, the Boron Local Fauna
lacks three groups (cricetid rodents, horses, and antiloca-
prids) which are a common element of Barstovian assem-
blages. The Boron Local Fauna also differs from the less well
preserved late Hemingfordian assemblages of southern Cal-
ifornia which are dominated by small merychippine horses,
middle-sized oreodonts, dromomerycines, and antilocaprids,
groups lacking at Boron. The Boron Local Fauna appears to

1 Vertebrate Paleontology Section, Natural History Museum of
Los Angeles County, 900 Exposition Blvd., Los Angeles, California,
90007.

Contributions in Science, Number 355, pp. 1-36
Natural History Museum of Los Angeles County, 1984

ISSN 0459-8113

118

117

Figure 1. Index map showing location of Boron Local Fauna and other southern California Miocene fossil vertebrate assemblages discussed
in this report.

be the first early Hemingfordian fossil assemblage sampled
in southern California.

GEOLOGIC SETTING

The western and central Mojave Desert is a broad, fault-
bounded block underlain by Mesozoic crystalline rocks which
were faulted into many localized basins during the Middle
and Late Cenozoic. Deposition in these basins was domi-

nated by volcanics and volcaniclastics during the first half
of the Miocene. Volcanism subsided during later Miocene
and fluviatile and lacustrine sediments accumulated in the
localized basins which persisted due to down-faulting, a pro-
cess which continues today in some areas. In other areas,
faulting has uplifted and exposed Miocene and younger se-
quences.

Considerable geologic, stratigraphic, and geophysical work
has been done in the general area of Boron, much of it per-

2 Contributions in Science, Number 355

Whistler: Boron Local Fauna

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Contributions in Science, Number 355

Whistler: Boron Local Fauna 3

Figure 2. Stratigraphic nomenclature of geologic units at Boron and vicinity.

taining to the deposition and extent of the borate-rich sed-
iments there (Figure 2). The Tertiary rocks at Boron and
surrounding hills were originally referred to as the Rosamond
Series (Hershey, 1902), a name expanded to include most of
the continental sedimentary and volcanic rocks in the Mo-
jave Desert, including such units as the Ricardo Formation
of Clarendonian age (Merriam, 1919; Dibblee, 1952) and
widely scattered deposits of Hemingfordian and Barstovian
age, usually referred to the Barstow Formation (Merriam,
1919; Bowen, 1954). The first mapping of the borate pro-
ducing beds at Boron was done by Gale (1946). He referred
these beds to the Ricardo Formation and defined a new un-
derlying unit, the Saddleback Basalt, which occurs in surface
exposures at Saddleback Butte, 5 km to the east of Boron.
A similar basaltic flow also occurs as a series of small hills
just north of the mine and is found exposed in several areas
of the open pit. It is also recognized in many of the drill holes
in the general area.

During the 1950’s, Dibblee undertook a regional mapping
program in the western and central Mojave Desert including
the area around Boron. Because of confusion over previous
usage of stratigraphic names, Dibblee (1958a) established a
new unit, the Tropico Group, to include only some of the
Tertiary rocks previously referred to as the Rosamond Series.
He included four formations within the Tropico Group (Gem
Hill Formation, Fiss Fanglomerate, Bissel Formation, and
Saddleback Basalt), but excluded such units as the Ricardo
and Barstow Formations which had been included in the
Rosamond Series by other workers.

Dibblee (1958a, b) did not give any formational designa-
tion to the beds at the open pit mine at Boron but referred
to them only as the “Upper Part of the Tropico Group.”
Following the usage of the geology staff at the Boron mine,
Bowser (1965), in his geochemical studies, used the term
“Kramer Lake Beds” for the sequence above the Saddleback
Basalt. Barnard and Kistler (1966) simplified Bowser’s name
to the Kramer Beds, included the Saddleback Basalt and
published a detailed stratigraphic section which included the
location of the Boron Local Fauna. They subdivided the
Kramer Beds into three members, the Saddleback Basalt,
Shale Member, and Arkose Member in ascending order. The
Saddleback Basalt forms the base of the exposed section in
the open pit mine and is the only member of the Kramer
Beds with natural exposures. This sequence of flows and flow
breccias appears to lie unconformably on pyroclastic rocks
which may, in part, represent an eastern extension of the
Gem Hill Formation, the basal unit of the Tropico Group.

The Shale and Arkose Members of the Kramer Beds over-
lying the Saddleback Basalt include lacustrine and fluviatile
sediments exposed only in the open pit mine and known
otherwise from drilling data (Benda et al., 1960; Dickey,
1957). The seemingly local extent of these units has been
discussed by several authors (Gale, 1946; Bowser, 1964; Bar-
nard and Kistler, 1966). The Shale Member is lacustrine,
composed of dark green to brown, borate-bearing clays, shales,
stratified borate ore bodies, and several thin bentonitic tuff
beds. The Arkose Member, which produced the Boron Local

Fauna, is composed of beds of well-bedded, buff, light tan
and dark reddish brown claystone and micaceous siltstone
and lenses of coarser, cross-bedded, light brown to yellowish
white arkosic sandstone and pebble conglomerate. The Ar-
kose Member contains relatively few volcanic beds, sug-
gesting that the volcanism so commonly represented in the
underlying units of the Tropico Group had subsided.

At the time of his original description, Dibblee (1958a)
believed that much of the Tropico Group was Late Miocene
and Pliocene in age. His conclusion was based on a diatom
flora recovered from a limestone in the Tropico Group west
of Boron and on Gale’s ( 1 946) lithologic correlation of the
Saddleback Basalt with andesite breccia flows in the basal
part of the Ricardo Formation which produced a fossil ver-
tebrate fauna of Clarendonian age (then considered Pliocene).
The discovery of the Boron Local Fauna at first seemed to
refute this latter conclusion, but subsequent work in the type
area of the Ricardo Formation has shown that a correlation
of the basal andesite breccia flows and underlying pyroclastic
rocks with the Tropico Group is reasonable. Field studies
have shown that these volcanic rocks unconformably un-
derlie the fossil-producing rocks of the Ricardo Formation
which are radiometrically dated at a maximum of 10.2 ma
(Evemden et al., 1964). These volcanics have subsequently
been radiometrically dated at 17.1 ma (Loomis et al., 1983)
and thus fall within the age range of the Tropico Group. The
Saddleback Basalt has yielded radiometric dates in a range
from 18.3 ± 0.6 (R.B. Kistler, personal communication) to
20.3 ± 0.7 ma (Armstrong and Higgens, 1973), somewhat
older than, but possibly representing the same volcanic ep-
isode as the andesite flows underlying the Ricardo Forma-
tion.

Dibblee (1958a) recognized lithologic similarities between
the Gem Hill Formation, the basal unit of the Tropico Group,
the Middle Miocene Kinnick Formation in the mountains
north of Tehachapi and the Pick Handle Formation under-
lying the Middle and Late Miocene Barstow Formation in
the Mud and Calico Hills. The presence of the Boron Local
Fauna in the uppermost member of the Tropico Group tends
to support Dibblee’s conclusions, but also shows that this
entire unit is pre-Middle Miocene in age, much older than
he had supposed.

FOSSIL OCCURRENCE

Fossils of the Boron Local Fauna were not observed at the
original site of deposition by the author. They were first
noticed by several mine employees operating heavy equip-
ment removing overburden from the open pit mine. The
mine foreman informed the geology staff who had the re-
maining fossil-bearing matrix dumped in a separate location
from the general mine dump so that the fossils could be
recovered. It is not certain how much of the fossil-bearing
pocket was lost before this action was taken, but 4-5 earth-
mover loads (12-15 cubic meters) were subsequently re-
covered before the pocket stopped producing readily visible
fossil bone.

4 Contributions in Science, Number 355

Whistler: Boron Local Fauna

According to the location provided by mine employees,
the fossil-producing layer was approximately 146 m above
the Saddleback Basalt. Observations made in the pit in 1 964
and from the fossil-bearing matrix indicate the fossils were
restricted to one or more beds of light tan micaceous clay-
stones which locally contain pumice lapilli fragments. These
beds were part of a sequence of fluviatile siltstones and sand-
stones locally cut by channels of coarser arkosic sandstones
and pebble conglomerates. About 3 m above the fossil-pro-
ducing beds were several discontinuous lenses of grayish-
pink bentonitic tuff and yellowish-white, medium-grained
sandy tuff.

Many of the individual bones were fractured and splintered
prior to fossilization, suggesting a period of surface exposure
before burial. On the other hand, some fossils are very well
preserved and several limb fragments were still partially ar-
ticulated. There is no evidence of rodent or carnivore gnaw-
ing on any of the fossil bone. Several matrix blocks contain
small, lenticular concentrations of partially leached small
vertebrate bone, apparently representing disassociated owl
pellets or carnivore coprolites which undoubtedly are the
source of the rich microvertebrate assemblage.

Limited prospecting by the author in the walls of the open
pit mine in 1964 did not produce additional fossils, nor did
extensive prospecting in all exposed fluviatile facies of the
Tropico Group within 15 km of Boron yield further fossil
specimens. The entire collection of fossil vertebrates was
deposited in the Department of Earth Sciences at the Uni-
versity of California at Riverside.

METHODS

The fossils were obtained from the pile of matrix removed
from the open pit mine. Blocks containing natural molds of
bones destroyed during excavation were also collected and
plaster casts were subsequently made of these molds. All
matrix with a lithology similar to known fossil bearing matrix
(about 2700 lb) was sieved under water with screens with an
average mesh opening of 0.7 mm.

Measurements on larger bones were made with a vernier
caliper to the nearest 0. 1 mm and the microvertebrates were
measured with an EPOI optical micrometer to the nearest
0.01 mm. Measurements are taken at their maximum di-
mensions unless otherwise indicated. The anteroposterior
dimension of the P4 of heteromyids was measured with the
posterior surface of the metaloph oriented vertically.

The stereophotographs were taken by the author using an
extension bellows and reversed 50 mm, f 1.4 lens mounted
on a stereophotographic bar. The specimens were coated with
vaporized magnesium oxide prior to photographing.

Radiometric dates have been converted using the new de-
cay and abundance constants given by Dalrhymple (1979).
This has the effect of increasing most later Tertiary dates
published before 1979 by about 2.6 percent.

Metric abbreviations and designations of tooth position
follow standard usage. Other abbreviations used are as fol-
lows:

LACM

Collections of the Natural History Museum of
Los Angeles County

LACM(CIT) Collections formerly at the California Institute
of Technology, now at LACM

UCMP

Collections of the University of California,
Museum of Paleontology, Berkeley

UCR

Collections of the University of California at
Riverside, Department of Geological Sciences

L

Left

R

Right

ma

millions of years before present

FAUNAL LIST

Below is a summary of the fossil vertebrate assemblage of
the Boron Local Fauna as recognized in the following sys-
tematics section.

Class Reptilia
Order Squamata
Suborder Sauria

Family Iguanidae or Xantusiidae

Iguanidae or Xantusiidae sp. indet.

Family undetermined
Sauria, incertae sedis
Suborder Serpentes
Family Boidae

Subfamily Erycinae
cf. Calamagras sp.

Family Colubridae
cf. Paracoluber sp.

Class Mammalia
Order Lagomorpha
Family Leporidae
Subfamily Archaeolaginae
Arachaeolagus sp. or Hypolagus sp.

Order Rodentia

Family Sciuridae

IMiospermophilus sp.

Family Heteromyidae
Subfamily Perognathinae
Mookomys sp.

Trogomys cf. T. rupinimenthae
Subfamily Dipodomyinae
Cupidinimus boronensis n. sp.

Order Artiodactyla

Family Merycoidodontidae
Subfamily Merychinae

Merychyus cf. M. minimus
IMerychyus sp.

Family Camelidae

Subfamily Aepycamelinae
cf. Hesperocamelus sp.

Subfamily undetermined
Camelidae, small species
Camelidae, large species

Contributions in Science, Number 355

Whistler: Boron Local Fauna 5

Family Palaeomerycidae
Subfamily Aletomerycinae
Aletomeryx occidentalis n. sp.

SYSTEMATICS
Class Reptilia
Order Squamata

Lizards and snakes are represented by very fragmentary ma-
terial, but the presence of two lizards and two snakes can be
demonstrated. The lizard identifications are based on the
tooth-bearing bones although some postcranial material is
also present. One of the lizards may represent the only Middle
Miocene record of the family.

At least one boid and one colubrid snake are present. Small
boids are common in the middle and late Tertiary, thus the
Boron occurrence is not unusual. On the other hand, the
colubrid represents one of the earliest records of the family
in North America and extends the geographic range of such
colubrids to west of the Rocky Mountains. Vertebral ter-
minology follows Auffenberg (1963) and taxonomy follows
Holman (1979).

Suborder Sauria

Family Iguanidae or Xantusiidae
Iguanidae sp. indet. or Xantusiidae sp. indet.

MATERIAL. Fragment of right dentary (or splenio-den-
tary), UCR 21 174; anterior left dentary (or splemo-dentary),
UCR 21175; premaxilla fragment, UCR 21176; miscella-
neous unnumbered edentulous maxillary and dentary frag-
ments.

DESCRIPTION. Small lizard, dentary less than 1 cm long
based on estimate from several specimens; Meckelian groove
closed and fused except for anterior opening; posterior teeth
strongly tricuspate with grooves on both lingual and labial
tooth surfaces between main cusp and lateral cusps; anterior
teeth single cusped; teeth of similar width along entire tooth
row; subdental lingual shelf well developed; dentary thin
(dorsoventrally) below subdental lingual shelf; dentary same
thickness below subdental lingual shelf for most of length of
tooth row; premaxilla with 6 teeth.

DISCUSSION. The Boron lizard material is very frag-
mentary and the above description is based on a composite
of several specimens with the single common character of a
fused Meckelian canal, a character restricted to the xantu-
siids, gekkonids, and some iguanids such as Dipsosaurus Hal-
lowell, 1854. The material is too incomplete to precisely
estimate the entire tooth row length or the number of teeth.
The individual tooth bases are relatively large and uncrowd-
ed, typical of xantusiids and some iguanids and in contrast
to gekkonids, which usually have large numbers of small
teeth. No posterior dentary fragments are preserved, thus the
presence or absence of the diagnostic xantusiid character of
separate anterior inferior alveolar and anterior mylohyoid
foramina (terminology of Schatzinger, 1980) entirely en-

closed within the splenio-dentary is not preserved. The teeth
are strongly tricuspate, similar to Paleoxantusia kyrentos
Schatzinger, 1980, but less so than in living Dipsosaurus or
Xantusia riversiana Cope, 1883. The subdental lingual shelf
is well developed as in the extinct Paleoxantusia Hecht, 1956,
and in contrast to most iguanids. The continuously dorso-
ventrally thin dentary below the subdental lingual shelf is
more similar to that of xantusiids than iguanids or gekkonids.

The Boron material does not permit positive identification,
but appears to represent an undescribed species of either
xantusiid or iguanid. Small iguanids are poorly known in the
fossil record but a variety of undescribed material is present
in Middle and Late Miocene deposits of the Mojave Desert
and Great Basin (collections at the University of California,
Berkeley, Museum of Paleontology, Natural History Mu-
seum of Los Angeles County and University of California at
Riverside). Paleoxantusia, an early xantusiid, has been rec-
ognized only from the Paleocene and Eocene of western North
America. Undescribed xantusiid material is known from
Middle and Late Miocene deposits in the Mojave Desert
(specimens in same collections as above) and the Middle
Pleistocene Palm Springs Formation in San Diego and Im-
perial Counties of California (M. Norell, personal commu-
nication; specimens in the Natural History Museum of Los
Angeles County).

Family undetermined
Sauria incertae sedis

DISCUSSION. There is a single dentary fragment, UCR
2 1 1 77, and miscellaneous unnumbered specimens of a small
lizard with an unfused Meckelian canal. The tooth mor-
phology of this (or these) lizard(s) is not preserved. This
material demonstrates the presence of at least one other small
lizard in the fauna.

Suborder Serpentes
Infraorder Henophidia
Family Boidae
Subfamily Erycinae
Ca/amagras Cope, 1873

cf. Calamagras sp.

Figure 3

MATERIAL. Centrum portion of trunk vertebra retaining
right diapophysis and portion of right prezygapophysis, UCR
21178; fragment of cotyle, prezygapophysis, and diapophy-
sis, UCR 21179; centrum fragment, UCR 21180.

DESCRIPTION. Size small; vertebral centrum short (2.35
mm from dorsal edge of cotyle to distal end of condyle);
ventral surface of centrum smooth with no haemal keel; no
subcentral ridges; paradiapophysis subdivided into two dis-
tinct articular surfaces of similar size.

DISCUSSION. Although fragmentary, this material is

6 Contributions in Science, Number 355

Whistler: Boron Local Fauna

clearly referable to the Family Boidae on the basis of the
short vertebral centrum and lack of haemal keel. There are
no neural arches or caudal vertebrae necessary for positive
subfamily assignment, but the small size favors assignment
to the Subfamily Erycinae. Among the fossil North American
erycines, only Calamagras and Tregophis Flolman, 1975,
completely lack a haemal keel. The diagnostic characters of
Tregophis occur in the neural arch, a structure not preserved
in the Boron sample. Tregophis is a peculiar form known
only from one Clarendonian locality in Kansas, and thus it
appears more reasonable to refer the Boron species to Cala-
magras, a wide ranging taxon in the Middle Miocene.

The cf. Calamagras sp. from Boron lacks the haemal keel,
the large, more widely separated paradiapophysial articular
surfaces and subcentral gutter of Charina prebottae Bratt-
strom, 1958, from the Barstow Formation.

The living Lichanura Cope, 1861, also lacks a distinct
haemal keel. The characters of the neural spine and caudal
vertebrae necessary to distinguish Calamagras from Licha-
nura are not preserved in the Boron sample. Lichanura has
not been recognized in the pre-Pleistocene record, and the
Boron species is tentatively referred to Calamagras for this
reason, not on morphological grounds.

Family Colubridae
Subfamily Colubrinae
Paracoluber Holman, 1970

cf. Paracoluber sp.

Figure 4

MATERIAL. Vertebral centrum complete from cotyle to
condyle but lacking neural arch and associated structures,
UCR 21181; five uncataloged posterior centrum fragments
with condyles.

DESCRIPTION. Size small, centrum 3.15 mm long; cen-
trum long compared to width; haemal keel moderately de-
veloped, oblanceolate, posterior end flat, not overlapping
condyle; keel border straight in lateral view.

DISCUSSION. Although fragmentary, this material is
complete enough to permit assignment to the Family Colu-
bridae. These specimens are disinguished from boids, the
common pre-Barstovian snakes in North America, by the
long centrum and relatively narrow, long haemal keel, from
the palaeophids, elaphids, viperids (including crotalids), and
natricine colubrids by the absence of hypapophyses, and from
the xenodontine colubrids (based on Heterodon Latreille,
1802) by the presence of a narrow, relatively high haemal
keel.

The cf. Paracoluber sp. from Boron is distinguished from
small fossil North American colubrines (see Holman, 1979)
by a combination of the long centrum, absence of subcentral
ridges, and structure of the haemal keel. It superficially re-
sembles Nebraskophis Holman, 1973, in length of centrum,
but has a much deeper haemal keel. The Boron species re-
sembles Paroxybelis Auffenberg, 1963, except that the pos-

terior end of the haemal keel is not pointed and does not
extend onto the condyle. It differs from Salvadora pateoli-
neata Holman, 1973, in lacking strong subcentral ridges.
Except for its small size, cf. Paracoluber sp. from Boron
closely resembles Paracoluber storeri Holman, 1970, in
structure of the haemal keel and weak subcentral ridges. Hol-
man (1970:1322) indicates that a paratype of P. storeri is
smaller, but does not give the dimensions.

The incomplete Boron material does not permit species
assignment, but it clearly demonstrates the presence of a
small colubrid in the fauna.

Class Mammalia
Order Lagomorpha

The lagomorphs are the second most common microverte-
brates in the fauna. Because much of the material is from
juveniles, positive identification is difficult. Dental termi-
nology and taxonomy follow Dawson (1958).

Family Leporidae
Subfamily Archaeolaginae

Archaeolagus Dice, 1917 or
Hypo/agus Dice, 1917

Archaeolagus sp. or Hypo/agus sp.

Figures 5-6

MATERIAL. Fragment of left dentary with broken M„
complete M2, alveolus for M3, UCR 10401; isolated, unworn
LM1, UCR 10403; partial LdP4, UCR 10400; isolated, un-
worn LP3 or LdP4, UCR 10402; isolated tooth and postcra-
nial fragments, UCR 10404-10412.

DESCRIPTION. M' with two lophs separated by cement-
filled, lingual reentrant; dP4 bilophed lingually, enamel lim-
ited to lingual side, large, labially curved root centered under
lingual lophs, labial roots missing; UCR 10402 (unworn P3
or dP4) with distinct talonid and trigonid separated by con-
tinuous valley instead of separate lingual and labial reen-
trants; cement only in central part of valley separating talonid
and trigonid; trigonid composed of three separate spurs joined
at center of tooth, labial spur narrow, bordered anterolin-
gually by deep reentrant.

DISCUSSION. Both the isolated M1 and UCR 10402
(unworn P3 or dP4) are from very young individuals, neither
showing wear. Thus, both display distinct, separate cusps in
contrast to the lophodont nature typical of lagomorphs. The
bases of the teeth are open and both are relatively low crowned,
suggesting neither tooth was completely formed. The reen-
trants of UCR 10402 (P3 or dP4) would be persistent in wear
for nearly the entire length of the preserved tooth crown.
With some wear, the tooth would divide into two lophs and
an additional anterolabial reentrant. Such a persistent reen-
trant is more diagnostic of Hypo/agus (Dawson, 1958).

Archaeolagus is typically characterized by the loss of the
lingual reentrant in the P3 with wear, but a very young in-

Contributions in Science, Number 355

Whistler: Boron Local Fauna 7

Figures 3-7. Stereophotographs of cf. Calamagras sp., cf. Paracoluber sp., Archaeolagus sp. or Hypolagus sp., and ? Miospermophilus sp.
Figure 3, cf. Calamagras sp., UCR 21 178, ventral view of vertebral centrum. Figure 4, Paracoluber sp., UCR 21181, ventral view of vertebral
centrum. Figures 5-6, Archaeolagus sp. or Hypolagus sp., UCR 10402, unworn LP, or dP4; (5) occlusal view, (6) labial view. Figure 7,
? Miospermophilus sp., UCR 10399, unworn LdP4 or LP4, occlusal view. Scale bar = 1 mm.

dividual of Archaeolagus acaricolus Dawson, 1958 (speci-
men LACM(CIT) 5176) exhibits a structure in the P3 similar
to UCR 10402. The Boron material appears small, even for
Archaeolagus, but this is probably due to the juvenile state

of most of the specimens. The available material from Boron
is not complete enough for certain assignment to either Hy-
polagus or Archaeolagus. While the size is more consistent
with Archaeolagus, the morphology of UCR 10402, if it is

8 Contributions in Science, Number 355

Whistler: Boron Local Fauna

indeed a P3 in extremely early wear, more nearly resembles
Hypolagus.

Order Rodentia

The rodent fauna is dominated by heteromyids except for a
single small squirrel tooth. Compared with other rodents,
squirrels are generally uncommon in the fossil record, and
although the Boron species appears to have some unique
characters, the single specimen is inadequate to make a spe-
cific assignment. The dental terminology used for the squirrel
follows Black (1963).

At least three heteromyid species representing two subfam-
ilies are present. As recognized by all who work with these
forms, heteromyid taxonomy is in great need of revision.
The Boron species do not help resolve this problem, but they
do indicate intraspecific variation in some characters which
have traditionally been used to separate species or even gen-
era.

The most common species in the microfauna, represented
by more specimens than all other microvertebrates com-
bined, is a new species of dipodomyine heteromyid referred
to Cupidinimus Wood, 1935. This species represents the ear-
liest record of this widespread genus but it is already clearly
derived with respect to species thought to be ancestral to this
and closely related genera. The other heteromyids, one a very
low crowned, bunodont species and the other a relatively
high crowned perognathine, are each represented by only two
teeth but are, nonetheless, identifiable at the generic level.
There are several vials of fragmentary postcranial bones from
small mammals, but none are clearly assignable to any of
the species represented by teeth. Tooth cusp terminology for
these heteromyids is given in Figure 8, taxonomy follows
Wood (1935) and Korth (1979).

Family Sciuridae
Subfamily Sciurinae
Miospermophilus Black, 1963

IMiospermoph ilus sp.

Figure 7

MATERIAL. Unworn, LdP4 or LP4, UCR 10399.

DESCRIPTION. Anteroposterior diameter = 1.65 mm,
width = 1.80 mm; outline sub-triangular; lophs and cusps
high and distinct; metaloph higher than protoloph, connected
to protocone by narrow ridge; connection between protoloph
and protcone stronger than connection between metaloph
and protocone; protocone large, with separate posterolingual
expansion (possibly indicating the position of a separate hy-
pocone) continuous with posterior cingulum; anterior and
posterior cingula low but distinct; anterior cingulum contin-
uous from protocone to termination in large parastyle in
anterolabial comer of tooth; posterior cingulum running from
posterolingual comer of metacone to posteriorly expanded
protocone (hypocone); mesostyle large.

DISCUSSION. The small size, sub-triangular outline, rel-

atively high cusps and lophs and presence of a metaconule
serve to distinguish this species as a ground squirrel or chip-
munk (Black, 1963). The Boron specimen is unusual for
either a chipmunk or a ground squirrel in the retention of
distinct cusps, the presence of a distinct protoconule, large
parastyle, and protocone split into two cusps. All these fea-
tures might simply be attributable to the unworn condition
of this specimen, but they also strongly suggest that it is a
deciduous premolar. The roots are missing, but the specimen
has the hollowed out base typical of both deciduous teeth
and unerupted permanent teeth.

Regardless of permanent versus deciduous designation, the
small size favors assignment of the Boron specimen to one
of three genera, Miospermophilus Black, 1963, Tamias Illi-
ger, 1811, or Ammospermophilus Merriam, 1892, all rep-
resented by Miocene aged species. The size is most consistent
with species of Miospermophilus, which are larger than those
of living and extinct species of Tamias and the Miocene
species of Ammospermophilus.

The dP4 of these small squirrels is poorly known, making
direct comparison difficult. The dP4 of M. bryanti (Wilson,
1960) from the Martin Canyon Quarry in Northeastern Col-
orado (in Black, 1 963: 188) and M. wyomingensis Black, 1963:
192 from the Split Rock Formation of Wyoming are de-
scribed as having a strong metaconule and low, distinct an-
terior and posterior cingula similar to the Boron specimen.
M. bryanti lacks a mesostyle but M. wyomingensis is de-
scribed as having a small mesostyle similar to the Boron
specimen. A partial, isolated dP4 from the Branch Canyon
Formation of California referred to Miospermophilus sp. by
Lindsay (1974: 14) lacks the protoconule and metaconule but
does have a small mesostyle. No described dP4’s of Miosper-
mophilus have the distinct protoconule present in the Boron
specimen, but undescribed specimens from the Thomas Farm
faunas of Florida do have a protoconule (D. Webb and A.
Pratt, personal communication).

The only described dP4 of a fossil species of Tamias (Black,
1963:129) is reported to bear the conules, large mesostyle,
and expanded anterior cingulum found in the Boron speci-
men.

The dP4 of fossil species of Ammospermophilus has not
been described, but specimens of this tooth and the per-
manent P4 in a sample of the living species A. lecurus (Mer-
riam, 1889) from the LACM Mammalogy collections differ
considerably from the Boron specimen. In these, the anterior
cingulum of the dP4 extends only half way along the anter-
olingual comer of the tooth and there is no hint of a parastyle
which is prominent in the Boron IMiospermophilus speci-
men. The metaloph in both the dP4 and P4 is reduced lin-
gually and does not connect to the protocone even in late
wear stages in the modem species examined. A further vari-
ation was found in specimens of the dP4 of the subspecies
A. lecurus peninsulae { Allen, 1893) where the metaloph con-
nects to the posterolingual comer of the lingual end of the
protoloph.

Although similar in most features, the permanent P4 of
species of Miospermophilus, Tamias, and Ammospermophi-
lus is generally even more lophodont, with greater reduction

Contributions in Science, Number 355

Whistler: Boron Local Fauna 9

hypocone
hypostyle

protostyle

anterior cingulum posterior cingulum
RM' Rp‘

Figure 8. Tooth cusp terminology used in heteromyid rodents.

of the metaconule and protoconule than the dP4. The cingula
are also better developed and nearly as high as the protoloph
and metaloph in the P4. The Boron specimen is similar in
size to the P4 of Miospermophilus species, but differs in re-
tention of distinct conules, particularly the protoconule.

Family Heteromyidae
Subfamily Perognathinae
Mookomys Wood, 1931

Mookomys sp.

Figures 9-12, Table 1

MATERIAL. RP4, UCR 10421; LM, or LM2, UCR 10464.

DESCRIPTION. P4 triangular in occlusal outline with four
subequal low rounded cusps; protostylid very small, lower
than protoconid; protostylid and protoconid nearly joined as
single cusp with only a shallow anterior groove separating
cusps on upper part of tooth crown; connection between
protolophid and metalophid central; no posterior cingulum.

UCR 10464 (M, or M2) nearly square in occlusal outline
with six low, separate cusps not distinctly connected until
late wear; tooth bilophodont with wear; central valley deep;
protostylid and hypostylid small, both offset posteriorly rel-
ative to protoconid and hypoconid; protostylid anteropos-
teriorly expanded; no anterior or posterior cingulum; con-
nection of lophids late, central; two roots.

DISCUSSION. The lower premolar, with its reduced pro-
tolophid, is more similar to species of Heliscomys Cope,
1873, than to the common species of Mookomys, M. altiflu-

Table 1. Selected dental measurements of Mookomys sp. and Tro-
gomys cf. T. rupinimenthae. Abbreviations: AP = anteroposterior di-

mension of tooth, WP
loph(id), X = mean.

= width of protoloph(id), WM

= width meta-

UCR

AP X WP X

WM X

Mookomys sp.

P4

10421

0.70 0.36

0.73

M, or M2

10464

0.86 0.78

0.79

Trogomys cf. T. rupinimenthae

P4

21182

1.00

1.15

P4

10419

0.83 0.60

0.85

minis Wood, 1931, and M. formicorum Wood, 1935, both
with distinctly four-cusped lower premolars. The P4 of M.
subtilis Lindsay, 1972, is not known. UCR 10421 lacks the
posterior cingulum diagnostic of Heliscomys species. A tri-
angular P4 with four cusps, but with a very small hypostylid
and a narrow protolophid, would be expected in an early
species of Mookomys if, as is generally accepted, this genus
is a descendant from Heliscomys, which has triangular, three-
cusped P4’s.

The lower molar is typical of species of Mookomys with
a very low crown and low separate rounded cusps which only
join in later wear. The Boron specimen is much smaller than
that of any species of Mookomys except Mookomys subtilis,
even if the specimen is a M2 (which is smaller than M,). It
is closer to size to teeth species of Heliscomys, but it lacks
the characteristic cingula. Mookomys sp. from Boron differs
from the M , of M. subtilis in having a smaller anterior cin-
gulum, less posteriorly offset protostylid and hypostylid, and
lacking a posterior cingulum. In this latter character and the
very low crown height, M. subtilis is more similar to species
of Heliscomys than to species of Mookomys. Lindsay (1972)
does not compare M. subtilis (Barstovian) with Heliscomys,
but it appears that it is more similar to Heliscomys, even
though the latter is restricted to the Oligocene.

Mookomys sp. from Boron probably represents a very small
species of Mookomys, but it could be interpreted as a survivor
species of Heliscomys in the early Hemingfordian.

Trogomys Reeder, 1960
Trogomys rupinimenthae Reeder, 1960

Trogomys cf. T. rupinimenthae

Figures 13-16, Table 1

MATERIAL. LP4, UCR 21182; RP4, UCR 10419.

DESCRIPTION. P4 with transversely expanded proto-
cone but no protostyle; central connection of protoloph to
metaloph; metaloph with three cusps, hypocone and meta-
cone closer together than hypocone and hypostyle; hypostyle
nearly same size as metacone, not anteroposteriorly expand-
ed, connected to hypocone by posterior cingulum; three roots,
separate to base.

10 Contributions in Science, Number 355

Whistler: Boron Local Fauna

Figures 9-16. Stereophotographs of Mookomys sp. and Trogomys cf. T. rupinimenthae. Figures 9-10, Mookomys sp., UCR 10421, RP4; (9)
occlusal view, (10) lingual view. Figures 11-12, Mookomys sp., UCR 10464, LM, or LM2; (11) occlusal view, (12) lingual view. Figures 13-
14, Trogomys cf. T. rupinimenthae. UCR 21 182, LP4; (13) labial view, (14) occlusal view. Figures 15-16, Trogomys cf. T. rupinimenthae,
UCR 10419, RP4; (15) occlusal view, (16) lingual view. Scale bar = 1 mm.

P4 four cusped; no hypostylid; fairly high crowned, clearly
more hyposodont than Perognathus Wied-Neuwied, 1839;
protostylid smaller than protoconid, offset posteriorly; valley
between lophids deep lingually, shallow labially; first con-

nection of lophids labial; sulcus between hypoconid and
metaconid as deep as between metaconid and protoconid;
two roots, separate to base.

DISCUSSION. These premolars are distinctly lower

Contributions in Science, Number 355

Whistler: Boron Local Fauna 11

Figures 17-19. Stereophotographs of Cupidinimus boronensis n. sp., UCR 10413, holotype, right dentary with dP4, M,_2; (17) occlusal view,
(18) labial view, (19) lingual view. Scale bar = 2 mm.

crowned than those of Cupidinimus, higher crowned and
more lophodont than those of Mookomys and somewhat
higher crowned than those of the common Barstovian species,
Perognathus furlongi Gazin, 1930. The diagnostic features

of Trogomys, the asulcate upper incisor and characters of the
molars, are not represented in the Boron sample. The two
Boron teeth are within the size range of P. furlongi, but are
higher crowned, a characteristic of Trogomys. They are com-

12 Contributions in Science, Number 355

Whistler: Boron Local Fauna

Table 2. Selected dental measurements of Cupidinimus boronensis
n. sp. Abbreviations: AP = anteroposterior dimension of tooth, WP
= width protoloph(id), WM = width metaloph(id), X = mean.

UCR

AP

X

WP

X

WM

X

Type M,

10413

0.98

1.19

1.30

m2

0.91

1.15

1.16

dP4

1.42

0.97

P4

10449

1.30

1.26

1.40

1.31

10450

1.31

1.27

10451

1.30

1.25

10452

1.12

1.31

10453

1.20

1.29

10454

1.45

1.37

10456

1.23

1.36

10457

1.13

1.29

10458

1.26

1.29

M1

10459

0.95

0.99

1.37

1.30

1.23

1.16

10460

0.95

1.17

1.10

10461

1.03

1.36

1.25

10462

1.00

1.34

1.20

10463

1.02

1.28

1.16

21183

1.00

1.25

1.00

M2

10465

0.90

0.86

1.23

1.11

1.02

0.99

10467

0.88

1.04

0.92

10468

0.83

1.11

1.04

21184

0.83

1.05

0.98

M3

10466

0.77

0.87

-

dPJ

10471

1.08

1.01

—

10472

1.19

1.15

-

P4

10416

0.95

1.00

0.91

0.88

1.05

1.05

10417

1.11

0.89

1.13

10418

1.02

0.83

1.10

10420

0.99

0.90

0.95

10422

0.94

0.88

1.03

M,

10413

0.98

1.05

1.19

1.17

1.30

1.27

10423

1.04

1.16

1.29

10424

1.06

1.29

1.29

10427

1.09

1.15

1.27

10429

1.02

1.23

1.27

10434

1.15

1.19

1.28

10436

1.08

1.19

1.27

10438

1.06

1.13

1.19

10439

1.00

1.13

1.25

10441

1.09

1.05

1.22

M,

10413

0.91

0.95

1.15

1.19

1.16

1.10

10426

0.90

1.17

1.08

10430

1.03

1.21

1.09

10431

0.89

1.25

1.11

10432

0.90

1.19

—

10433

1.05

1.23

1.10

10435

1.01

1.24

1.07

Table 2. Continued.

UCR

AP

X

WP

X

WM

X

10437

0.87

1.13

1.15

10440

1.04

1.23

1.16

10444

0.96

1.26

1.07

10447

0.88

1.05

1.00

M,

10443

0.74

0.73

1.08

0.92

0.77

0.76

10445

0.74

0.91

0.76

10446

0.72

0.76

0.74

dP4

10414

1.32

-

0.75

parable in size, crown height, and cusp morphology to T.
rupinimenthae from the Arikareean age Tick Canyon For-
mation of southern California.

Subfamily Dipodomyinae
Cupidinimus Wood, 1935

Cupidinimus Wood, 1935:1 18.

Perognathoides W ood, 1935:92.

Prodipodomysl mascallensis Downs, 1956; sample of Shot-
well, 1967:22.

Perognathus saskatchewanensis Storer, 1 970: 1127.

Cupidinimus boronensis new species

Figures 17-39, Table 2

HOLOTYPE. UCR 10413, right dentary with incisor, dP4,

M,_2.

TYPE LOCALITY. UCR locality RV 642 1 , Arkose Mem-
ber of the Kramer Beds of Barnard and Kistler (1966), 146
m above the top of the Saddleback Basalt on the third level
below the surface in the east wall of the open pit mine of the
United States Borax and Chemical Corporation as it existed
in March, 1964, 4.8 km northwest of Boron, Section 23,
T. 1 IN., R.8W., Boron Quadrangle, United States Geological
Survey, 1954, 1:62,500, San Bernardino County, California.

REFERRED MATERIAL. Five RP4’s, UCR 10449, UCR
10450, UCR 10451, UCR 10452, and UCR 10453;five LP4’s,
UCR 10454, UCR 10455, UCR 10456, UCR 10457, and
UCR 10458; two RM"s, UCR 10460 and UCR 21183; four
LM"s, UCR 10459, UCR 10461, UCR 10462, and UCR
10463; two RM2’s, UCR 10465 and UCR 21 184; two LM2’s,
UCR 10467 and UCR 10468; RM3, UCR 10466; LM1 or
LM2, UCR 1 0469; two RdP4’s, UCR 10471 and UCR 10472;
two RP4’s, UCR 10418 and UCR 10420; four LP4’s, UCR
10415, UCR 10416, UCR 10417, and UCR 10422; seven
RM.’s, UCR 10423, UCR 10424, UCR 10427, UCR 10436,
UCR 10438, UCR 10439, and UCR 10441; four LM.’s,
UCR 10428, UCR 10429, UCR 10434, and UCR 10470;
five RM2’s, UCR 10431, UCR 10437, UCR 10440, UCR
10444, and UCR 10447; six LM,’s, UCR 10425, UCR 10430,
UCR 10432, UCR 10433, UCR 10435, and UCR 10442;
RM3, UCR 10445; two LM3’s, UCR 10443 and UCR 10446;

Contributions in Science, Number 355

Whistler: Boron Local Fauna 13

Figures 20-29. Stereophotographs of Cupidinimus boronensis n. sp. lower dentition. Figure 20, UCR 10413, holotype, occlusal view of
dentition only. Figues 21-22, UCR 10416, LP4; (21) occlusal view, (22) labial view. Figure 23, UCR 10422, LP4, occlusal view. Figures 24-
25, UCR 10423, RM,; (24) occlusal view, (25) labial view. Figures 26-27, UCR 10430, LM2; (26) occlusal view, (27) lingual view. Figure 28,
UCR 10445, RM,, occlusal view. Figure 29, UCR 10414, RdP4, occlusal view. Scale bar = 1 mm.

14 Contributions in Science, Number 355

Whistler: Boron Local Fauna

RdP4, UCR 10414; all from type locality, UCR locality RV
6421.

DIAGNOSIS. Teeth typical of dipodomyines by being
higher crowned than those of perognathines and most het-
eromyines; further distinguished from heteromyines by lack-
ing a “J” wear pattern in the P4; distinguished from geo-
myines by smaller size, lower crown height, retention of
cuspidate teeth and central (versus lingual) connection of
lophs in P4; individual teeth larger than those of C. nebras-
kensis Wood, 1935, C. halli (Wood, 1936b), C. cuyamensis
(Wood, 1937), and C. madisonensis (Dorr, 1956), smaller
than C. bidahochiensis (Baskin, 1979); teeth high crowned
but retaining individual cusps until moderate wear; roots
long, separate to base; premolars lacking accessory cusp(id)s;
M33 reduced; dP44 low crowned, with complex cusp pattern;
metaloph of P4 wide compared to width of M1, hypostyle
not greatly elongate and posterior cingulum incomplete; P4
short anteroposteriorly relative to width of protolophid and
length of M,; anterior cingulum not anteriorly expanded in
M, or M2; dP4 with large anteroconid connected to central
mure and lacking hypostylid; in addition to size, differs from
C. halli and C. madisonensis by lack of accessory cusps in
P44, anteroposteriorly short P4, reduced anterior cingula on
M, and M2 and large anteroconid connected to central mure
and lack of hypostylid in dP4; in addition to size, differs from
C. nebraskensis by anteroposteriorly short P4, relatively un-
reduced M2, reduced M33, and strong central mure on dP4.

ETYMOLOGY. Named after the type locality.

DESCRIPTION. P4 with transversely expanded proto-
cone; accessory cusps lacking on protocone except for one
specimen, UCR 10454, with greater transverse expansion of
protoloph suggestive of protostyle; first connection of lophs
central; metaloph three cusped; hypocone slightly posterior
relative to lateral cusps; hypostyle slightly expanded antero-
posteriorly; posterior cingulum small, between postero-
medial surface of hypocone and hypostyle; three roots, one
each under protoloph, metacone, and hypostyle, separate to
base.

M1 six cusped in early wear, cusps less persistent than in
lower molars; anterior cingulum low relative to protocone,
continuous between paracone and protostyle, thickening near
paracone; protoloph wider than metaloph; anterior face of
protoloph flat; small protolophule connecting protocone and
paracone in 2 of 3 specimens with early enough wear to show
character; transverse valley deep and narrow; first union of
lophs lingual, followed by central connection, but no lake
formation; median cusps (protocone and hypocone) posi-
tioned posteriorly relative to lateral cusps, making metaloph
convex posteriorly; no posterior cingulum except for con-
nection between hypocone and hypostyle; one lingual, two
labial roots, separate to base.

M2 smaller than M\ proportionately more reduced than
M2; anterior face convex; posterior face concave; anterior
cingulum reduced to absent; protolophule between protocone

and paracone present in one specimen; first union of lophs
labial, but transverse valley shallower than in M 1 , closed both
lingually and labially; metaloph narrower than protoloph due
to reduction in metacone; metaloph three cusped, joined into
continuous posterior crescent; hypostyle offset relative to hy-
pocone and more anteroposteriorly elongate than in M1; one
lingual, two labial roots, separate to base.

M3 with no anterior cingulum; protostyle elongate antero-
posteriorly; metaloph bearing three cusps joined to posterior
crescent which lacks distinct cusps; roots not preserved in
available sample.

dP4 low crowned; triangular outline; cusps low but distinct;
three lophed; large, transversely expanded anterocone; pro-
toloph with two cusps, larger protocone and smaller para-
cone; metaloph with three cusps, metacone, hypocone, and
anteroposteriorly expanded hypostyle; anterolabial connec-
tion between protoloph and anterocone; no connection be-
tween protoloph and metaloph; hypostyle extends anteriorly
to lingual side of protocone but with no connection to pro-
tocone; three roots, one each under anterocone, metacone,
and protostyle.

P4 with four cusps nearly equal in size; protostylid smaller
than protoconid, closer to hypoconid than protoconid is to
metaconid; protoconid-protostylid connection central; pro-
tolophid narrower than metalophid; first connection of lophs
either slightly lingual of center, between protostylid and hy-
poconid, or central; two roots, separate to base.

M, six cusped; tooth wider than long; protostylid nearly
same size as protoconid, offset posteriorly relative to pro-
toconid, continuous with anterior cingulum which connects
to anteromedial surface of protoconid; metalophid convex
anteriorly, hypolophid straight; transverse valley deep, nar-
row, open labially and lingually, deeper on lingual side; first
union of lophs both between protostylid and hypostylid and
central, probably forming short-lived lake; hypolophid nar-
rower than metalophid due to expanded entoconid; hypo-
stylid smaller than hypoconid; no posterior cingulum; two
roots, separate to base.

M2 six cusped; tooth wider than long; metalophid and
hypolophid same width; metalophid less convex anteriorly
than in M,; protostylid small, smaller than in M„ offset
posteriorly relative to protoconid, continuous with anterior
cingulum which extends only to anterolabial corner of pro-
toconid; hypostylid smaller than on M,; transverse valley
deep, deeper lingually than labially, labial connection deeper
than in M,; first union of lophs more strongly central than
in M„ labial connection late or lacking; hypolophid straight;
two roots, separate to base; two early wear specimens, UCR
1 0442 and 1 0444 displaying variation from above by lacking
distinct hypostylid (but with hypoconid extending labially
toward base, thus forming as broad a hypolophid as an M2
with hypostylid) and with protostylid separated from anterior
cingulum by distinct, shallow groove lost with early wear.

M3 four cusped, lacking stylids; bilophodont in early wear;

Contributions in Science, Number 355

Whistler: Boron Local Fauna 15

4

30

32

•1

tv

31

n 11

33

n

0

35

0 0

34

©

36

W 6

38

w

37

39

Figures 30-39. Stereophotographs of Cupidinimus boronensis n. sp. upper dentition. Figures 30-31, UCR 10450, RP4; (30) occlusal view,
(31) labial view. Figure 32, UCR 10449, RPJ, occlusal view. Figures 33-34, UCR 10460, RM1; (33) labial view, (34) occlusal view. Figure
35, UCR 10467, LM2, occlusal view. Figure 36, UCR 10445, RM3, occlusal view. Figure 37, UCR 10472, RdP\ occlusal view. Figures 38-
39, UCR 10471, RdPJ; (38) lingual view, (39) occlusal view. Scale bar = 1 mm.

16 Contributions in Science, Number 355

Whistler: Boron Local Fauna

very small anterior cingulum in early wear, forming anter-
olabial “comer” on protoconid with wear; metalophid dis-
tinctly wider than hypolophid; central valley narrow, deep;
single rooted.

dP4 very low crowned; elongate oval in occlusal view, nar-
rower anteriorly; complex cusp pattern with three main cen-
tral cusps, metaconid, entoconid, and hypoconid; large an-
terolingually placed anteroconid nearly continuous with high
anterolabial cingulum; anterolabial cingulum and protostylid
as high as central cusps, nearly continuous to mid-labial por-
tion of tooth and connecting back lingually to central mure;
very small cuspule between hypoconid and posterior end of
anterolabial cingulum; low protoconid anterolabial to meta-
conid; no hypostylid; with wear, hypoconid joins a central
mure which runs anteriorly through protoconid to antero-
conid; with wear, metaconid joined by short loph to proto-
conid; entoconid connected by mure either directly to hy-
poconid or through posterior cingulum to hypoconid; small
lingual accessory cuspid between metaconid and entoconid;
large posterior cingulum; two roots, one anterior, one pos-
terior.

DISCUSSION. C. boronensis is assigned to the genus
Cupidinimus following the expanded concept of this genus
proposed by Korth (1979). As previously noted by Reeder
(1956), Lindsay (1972), and Sutton (1977), the genus Cupi-
dinimus, represented by the single species C. nebraskensis,
and at least some of the eight species referred to Perogna-
thoides, are nearly indistinguishable in dental morphology.
Characters which have been used to distinguish these species
were found to be variable as larger samples became available.
Thus, Korth (1979) proposed formal synonymy of Perog-
nathoides and several other species referred to other genera
with Cupidinimus.

C. boronensis is typical of this expanded Cupidinimus
species complex with high-crowned teeth which retain in-
dividual cusps until moderate wear. It is unique in combining
large size, length (anteroposterior) of P4 much less than M,
(a Perognathus- like character), high-crowned teeth, relatively
unreduced M2, reduced M33, and dP4 with central mure and
no hypostylid (C. halli-C. madisonensis characters), with a
lack of accessory cusps on P44, a wide metaloph on P4 com-
pared to M1, a reduced anterior cingulum on M,, and dP4
with large anteroconid connected to central mure (C. ne-
braskensis characters).

C. boronensis is higher crowned than the perognathines
Mookomys, Perognathus, and Trogomys. It is also higher
crowned than most species of the heteromyines Peridiomys
Matthew, 1924, and Proheteromys Wood, 1932. These and
the higher crowned heteromyine Diprionomys Kellogg, 1910,
are further distinguished by the “J” wear pattern in the P4
with the first union of lophs lingually between the protocone
and hypostyle. C. boronensis is smaller, lower crowned, and
lacks the labial connection of protoloph and metaloph in the
P4 of Mojavemys Lindsay, 1972, and other geomyines.

C. boronensis is closest in size to C. halli and C. madi-
sonensis (= C. halli according to Lindsay, 1972, but consid-
ered distinct by Sutton, 1 977, a view I will follow). The molars
are as large as or larger than those of C. halli and C. madi-
sonensis. C. boronensis also differs from these species as fol-
lows: 1) the roots are less fused, 2) the length of P4 is less in
comparison to the width of the metalophid and length of M,,
3) the P4 lacks an anteroconid, and the connection of pro-
toconid and protostylid is central rather than posterior, 4)
the relative shortness of the P4 is similar to the condition in
Perognathus and Trogomys, in contrast to other species of
Cupidinimus in which this is the longest tooth in the lower
tooth row, 5) the anterior cingulum of C. boronensis is small-
er and less angular in both M, and M2, connecting only to
the protostylid, not the protostylid and metaconid, 6) the
hypostylid is lacking in 2 of 12 specimens of M„ 7) the P4
lacks accessory cusps on the protocone, the hypocone is less
posterior relative to lateral cusps, the hypostyle and posterior
cingulum are smaller, and the metalophid is wider relative
to the width of M1, 8) the M1 lacks a posterior cingulum and
the protolophule between protocone and paracone is smaller,
9) compared to specimens referred to C. halli by Lindsay
(1972), the dP4 is much lower crowned with a larger anter-
olabial cingulum, smaller anteroconid, no hypostylid, and a
weaker central mure (except that it is fully connected to an-
teroconid), and 10) the dP4, even though unworn, is lower
crowned, lacks a connection between hypostyle and proto-
cone and has a labial connection between the protoloph and
anterocone.

C. boronensis is 1 8-20 percent larger than C. nebraskensis
from the type area (Korth, 1979) and the sample from the
Barstow Formation (Lindsay, 1972). It resembles the sample
from the Barstow Formation in lacking the anteroconid on
P4, but this character is variably developed in the topotypic
sample from the Valentine Formation. C. boronensis lacks
the relative reduction of the M2 found in C. nebraskensis but
has a proportionally more reduced M3.

C. boronensis differs from species of Cupidinimus previ-
ously referred to Perognathoides, C. cuyamensis, C. klein-
felderi (Storer, 1970), and C. bidahochiensis, by lacking ac-
cessory cusps in the premolars variously developed in all
these and differs further from C. bidahochiensis by much
smaller size. C. boronensis also differs from C. kleinfelderi,
known only from isolated premolars, by lacking an anteriorly
drawn out protolph(id) compared to metaloph(id) on P44.

The types of C. eurekensis (Lindsay, 1972), C. quartus
(Hall, 1930), and C. tertius (Hall, 1930), all previously re-
ferred to Perognathoides, have highly worn teeth which lack
diagnostic characters that would facilitate comparison with
C. boronensis, but C. eurekensis is considerably smaller and
C. quartus and C. tertius are larger.

C. boronensis differs from four other species referred to
Cupidinimus by Korth (1979), a sample of C. cf. C. tertius
from the Avawatz Formation (Wilson, 1939), C. cf. C. cuya-

Contributions in Science, Number 355

Whistler: Boron Local Fauna 17

Table 3. Selected measurements of Merychyus cf. M. minimus and
IMerychyus sp. from the Boron Local Fauna. Abbreviations: AP =
anteroposterior dimension of tooth, TR = width of tooth at anterior
loph(id), CH = crown height at mesostyle in uppers, at metaconid
in lowers, * = approximate.

Skull

Merychyus cf. M. minimus

IMery-
chyus sp.

UCR UCR

10319 10320

UCR

10321

UCR

10322

UCR

10331

Facial length, C to

anterior rim orbit

52.8

51.1

54*

Jugal depth below orbit

13.2

12.8

17.7

Length P'-P4

30.0

30.8

33*

Length M'-M3

38.0

44.7

Length C-M3

74.3

-

P2 AP

7.8

8.0

7.8

TR

6.0

6.8

6.6

P3 AP

8.1

9.6

8.2

9.2

TR

7.6

8.0

7.1

7.5

P4 AP

7.4

7.3

7.2

TR

9.2

8.6

10.1

CH

10.4

7.6

M1 AP

—

11.0

12.8

1 1.5

TR

—

11.1

10.8

1 1*

CH

—

9.1

10.5

-

M2 AP

13.5

15.4

—

13.4

TR

1 1.0

12.5*

13.8

CH

—

12.6

—

-

M3 AP

18.0

21.0

TR

13.0

14.0

CH

14.7

8.9

Length dP2-dP4

25.7

26.8

Length dP2

7.4

Length dP3

8.7

10.1

Length dP4

8.7

9.4

Merychyus cf.

M. minimus

1 Merychyus sp.

UCR

UCR

UCR

UCR

Dentary

10323 1

10325

10332

21270

Mandible depth below P2

22.6

17.1

24.5

Length of symphysis

33.8

22.0

32.3

Length C-M,

79.8

80.1

Length P,-P4

31.9

29.5

Length M,-M3

44.2

47.0

M, AP

12.6

12.9

TR

8.0

8.2

CH

5.5

8.7

M, AP

19.8

23*

21.5

TR

8.9

8.8

8.9

CH

—

17.0

Length dP3-dP4

31.8

mensis from the Valentine Formation (Klinginger, 1968), a
sample referred to as “ Prodipodomys" ? mascallensis (Downs,
1956), from the Quartz Basin in Oregon (in Shotwell, 1967),
and C. saskatchewanensis (Storer, 1975), by lacking acces-
sory cusps on the premolars. C. boronensis is close in size to
the Quartz Basin C. mascallensis and otherwise similar ex-
cept that the connection of the protoloph and metaloph in
P4 is lingual rather than central in C. mascallensis. C. bo-
ronensis is also similar in size to C. saskatchewanensis, but
the latter is lower crowned and the protolophid of its P4 is
relatively anteriorly drawn out as in most species of Cupi-
dinimus.

C. boronensis is conservative among species of Cupidini-
mus in lacking development of accessory cusps on the pre-
molars. It is even larger than Barstovian species, in contrast
to an apparent general trend of size increase from Barstovian
to Hemphillian. Although larger than the common Barsto-
vian C. halli, it is lower crowned, more like the much smaller,
contemporaneous and longer surviving C. nebraskensis. This
combination of characters would suggest that C. boronensis
is not directly ancestral to either the smaller, higher crowned
C. halli or the smaller C. nebraskensis. It may represent a
separate lineage that increased in size early or, by reversal
of the size increase, gave rise to C. nebraskensis or similar
species.

C. boronensis could easily be derived from any perogna-
thine by a significant increase in crown height and a reduction
of cingula. Trogomys was suggested as a likely ancestor to
the C. halli/C. nebraskensis complex in the Barstow For-
mation by Lindsay (1972). C. boronensis is much higher
crowned and has much reduced cingula compared to the type
of Trogomys, and it is only separated from the latter by a
short geologic interval. However, Trogomys is the most likely
candidate among the perognathines.

C. boronensis is the earliest species in the Cupidinimus
species group and the only Hemingfordian representative of
this successful group. In general, Hemingfordian heteromyids
are poorly known, and as such, C. boronensis provides a rare
look at these small rodents in the time span between Ari-
kareean faunas dominated by pleurolicine, primitive-pe-
rognathine, and Proheteromys spp. heteromyid assemblages
and Barstovian faunas dominated by Cupidinimus spp. and
advanced-perognathine heteromyid assemblages.

Order Artiodactyla

The artiodactyls are represented by three families, oreodonts,
camels, and palaeomerycids. More than half the specimens
are juveniles, making positive identifications difficult in some
cases.

The oreodonts, represented by one or more small, sub-
hypsodont species, are more common than the camels. Ore-
odont taxonomy is in need of revision and the material from
Boron sheds little light on this vexing problem. Taxonomy
used follows Schultz and Falkenbach (1947) and Stevens
(1977).

As is typical of Miocene faunas, the Boron Local Fauna
contains at least three camel taxa. Most of the material is

18 Contributions in Science, Number 355

Whistler: Boron Local Fauna

Contributions in Science, Number 355

Whistler: Boron Local Fauna 19

Figures 40-43. Merychyus cf. M. minimus. Figures 40-41, UCR 10319, partial cranium with right facial region, RI1-3 (roots only) and RC-RM3; (40) lateral view, (41) occlusal
view. Figures 42-43, UCR 10323, lower mandible with complete dentition; (42) left lateral view, (43) occlusal view. All actual size.

I

Figures 44-45. Merychyus cf. M. minimus, UCR 10320, juvenile cranium with C, P1, LdP3-4, M1-2, unerupted RP2-4 exposed by removing
deciduous right premolars; (44) dorsal view, (45) occlusal view. All actual size.

poorly preserved. Only two species are represented by partial Webb (1965), and Honey and Taylor (1978); osteological
dentitions (one only by juvenile dentition) and associations terminology follows Webb (1965).

with postcranial elements have been made on size alone. The unique artiodactyl in the Boron Local Fauna is a new

Thus, only one camel is tentatively identified at the generic species of hypsobrachyodont aletomerycine palaeomerycid.

level. The other two cannot even be confidently assigned to Whereas palaeomerycids are relatively uncommon in fossil

subfamily level. The taxonomy follows Macdonald (1949), assemblages of the Great Basin and southern California when

20 Contributions in Science, Number 355

Whistler: Boron Local Fauna

compared to other artiodactyls, this species is the most abun-
dant large animal at Boron and is exceeded in numbers of
specimens only by Cupidinimus boronensis. Palaeomerycid
taxonomy is traditionally based on the orbital horns common
in this group. The new species from Boron is represented by
one horn fragment, but, in this case, the more diagnostic
characters are in the dentition. This new species is notewor-
thy because it is the first well-documented aletomerycine
found west of the Rocky Mountains, thus demonstrating a
broader distribution than previously recognized for these
small, hypsobrachyodont “homed ruminants.” Dental ter-
minology used is given in Figure 50 and taxonomy follows
Frick (1937).

Family Merycoidodontidae
Subfamily Merychyinae
Merychyus Leidy, 1858
Merychyus minimus Peterson, 1906

Merychyus cf. M. minimus

Figures 40-45, Table 3

MATERIAL. Fragment of cranium with right orbit and
facial region, RI1'3 (roots only), and RC-M\ UCR 10319;
juvenile cranium lacking posterior portion behind orbits, with
moderately worn C, P1, dP3~4, M‘~2, and unerupted P2-4 ex-
posed on right side, UCR 10320; juvenile palate with well
worn dl1-3, dC, P1, dP2~4, and slightly worn M1-2, UCR 10321;
partial left maxilla with moderately worn I3, C, P1-3, UCR
10322; isolated LM3, UCR 21185; mandibular fragment with
complete dentition, lacking ascending ramus, angle and con-
dyles, UCR 10323; natural mold of right dentary with con-
dyle, P4-M3, UCR 10324; juvenile mandible with partially
erupted P, slightly worn dC, dP,_3, and M,, UCR 10325;
unworn LM, in maxilla fragment, UCR 10326.

DESCRIPTION. Size (based on length of upper tooth row)
slightly larger than average Merychyus crabilli Schultz and
Falkenbach, 1947, and Merychyus calaminthus Jahns, 1940,
within size range of average Merychyus minimus; skull with
shallow antorbital fossa and narrow lacrimal vacuity; jugal
shallow; orbit large; two infraorbital foramina with posterior
one above P4; dentition sub-hypsodont (ratio of height of
enamel of paracone of M3 to width of M3 = 0.9); molars
appear narrow in relation to length even in late wear giving
teeth less robust appearance than typical in oreodonts; pre-
molars large; ratio of length of P1-4 to length of M1-3 = 0.81
(premolar diameter of Stevens, 1977); premolars lacking
complexity of spurs typical in Merychyus ( Metoreodon ) re-
lictus Matthew and Cook, 1 909; upper premolar spacing closed
but not crowded; lower premolar spacing somewhat open;
anterior margins of P 1-3 slant backward; upper C and P, not
enlarged.

DISCUSSION. Historically, oreodont taxonomy has been
based on samples of complete skulls. The fragmentary ma-
terial from Boron does not permit such an analysis, and the
study is further complicated by the large amount of juvenile
material. Oreodonts appear to be evolutionarily conservative

in dental characters. Even generic and subfamily differences
are difficult to distinguish using only the dentition and there
is little agreement among workers on evolutionary lineages.
Merychyus cf. M. minimus from Boron is assignable to the
genus Merychyus on the basis of the large orbit, presence and
size of facial fossa and prelacrimal vacuity, relatively high-
crowned teeth with large, posteriorly inclined upper pre-
molars, and small upper C and P, (P, is the lower caniniform
tooth in oreodonts).

Based on length of upper dentition, the Boron species is
1 5 percent smaller than M. ( Metoreodon ) relictus but only 3
percent smaller than M. ( Metoreodon ) relictus fletcheri Schultz
and Falkenbach, 1947, from the Barstow Formation. The
subgenus Metoreodon, based on M. ( Metoreodon ) relictus, is
distinguished from the subgenus M. ( Merychyus ) primarily
by its more complex and crowded premolars, P2^, well grooved
externally, P,_3 set obliquely in the jaw, teeth higher crowned,
and several other skull characters not preserved in the Boron
sample. Merychyus cf. M. minimus from Boron has a shallow
lacrimal fossa and fairly high crowned teeth, but it lacks the
premolar complexity and crowding typical of M. ( Metoreo-
don).

The length of the upper dentition of the Boron Merychyus
is 25 percent smaller than Ticholeptus calimontanus (Dough-
erty, 1940), from the Temblor Formation, but it is as high
crowned. The genus Ticholeptus Cope, 1878, although placed
in a different subfamily, is very similar to Merychyus, but
differs by having a proportionally smaller orbit, a deeper
jugal, larger upper canine and P,, and more anterior infraor-
bital foramen above P3-4.

Merychyus cf. M. minimus from Boron is larger and more
hypsodont than M. calaminthus (type series) and material
referred to M. calaminthus from the type area of the Hector
Formation (specimen UCR 10914 — Woodbume et al., 1974),
a specimen from the Hector Formation in the northern Cady
Mountains (specimen UCR 10840— Miller, 1980) and a
specimen from the Orocopia Mountains (specimen LACM
27026 — Woodbume and Whistler, 1973). In overall size, the
Boron material falls between M. calaminthus/M. crabilli and
M. minimus but it is closer to the latter. Stevens (in Wood-
bume etal., 1974:19) has suggested that M. calaminthus and
M. crabilli are the same species, a suggestion she reaffirmed
in a later work (Stevens, 1977:37), but stopped short of pro-
posing a formal synonymy, thus they are considered distinct
in this study.

Stevens (1977) has suggested two evolutionary trends in
Merychyus— decrease in “premolar diameter” (the ratio of
the length of the Pi_4 to the length of M1-3) and an increase
in hypsodonty. The material from Boron (especially UCR
10319) has large premolars that are less reduced than in M.
calaminthus or M. crabilli, but has cheek teeth that are more
hypsodont than even higher crowned individuals of M. min-
imus and Merychyus elegans Leidy, 1858. The jugal depth
(as a ratio of tooth row length) is shallow, more similar to
that of M. calaminthus and M. crabilli than that of M. min-
imus or M. elegans.

Merychyus cf. M. minimus from Boron appears distinct
from M. calaminthus and M. crabilli, but it falls within the

Contributions in Science, Number 355

Whistler: Boron Local Fauna 21

22 Contributions in Science, Number 355

Whistler: Boron Local Fauna

Figures 46-49. IMerychyus sp. Figures 46-47, UCR 10331, adult palate with well worn P2-M3 and part of right side of face; (46) right lateral view, (47) occlusal view. Figures
48-49, UCR 10332, lower mandible with alveolus of I ,_3, partially complete C, P,_3, complete P4-M3; (48) right lateral view, (49) occlusal view. All actual size.

Table 4. Selected measurements of cf. Hesperocamelus sp. and an indeterminate species of small camel from the Boron Local Fauna. Abbre-
viations: AP = anteroposterior dimension of tooth, TR = width of tooth at anterior loph(id), * = approximate.

cf. Hesperocamelus

Camelidae, small sp.

UCR

10366

UCR

10367

UCR

10365

Width palate, anterior end P2

27.3

Length P2-M3

115*

Length dP2-M, 47.0

P2 AP

9.0

M, AP 16.2

TR

L6.1, R6.3

TR 6.2

P3 AP

L16.0, R16.5

Length dP2 5.9

TR

9.8

Length dP, 9.0

M1 AP

19.6

Length dP4 16.8

TR

16.3

M2 AP

L27.7, R27.7

TR

L21.1, R22.0

M3 AP

29.3*

TR

23.5*

M3 AP

29.9

TR

10.0

Width distal radioulna

50.9 (UCR 10384)

35.0 (UCR 10374)

47.7 (UCR 10385)

Width proximal metacarpal

31.3 (UCR 21189)

Width distal tibia

42.0 (UCR 10339)

34.0 (UCR 10371)

42.0 (UCR 10387)

37.0 (UCR 10388)

Width proximal metatarsus

27.9 (UCR 10376)

Length calcaneum

1 17.7 (UCR 21186)

69* (UCR 10379)

Length astragalus

48.0 (UCR 10392)

36.8 (UCR 10378)

considerable range of variation of M. minimus in many char-
acters. Merychyus cf. M. minimus from Boron is probably
distinct from M. minimus, but larger samples and more com-
pletely preserved material are needed before this can be con-
clusively demonstrated.

? Merychyus sp.

Figures 46-49, Table 3

MATERIAL. Adult palate with well-worn P2-M3 and part
of right side of face, UCR 10331; lower mandible bearing
alveolus of I,_3, partially complete C and P,_3, complete P4-
M3, UCR 10332; fragment of left dentary with P4-M,, UCR
10333, fragment of right dentary with M3, UCR 21270.

DESCRIPTION. Size similar to Merychyus cf. M. mini-
mus described above; jugal deep; infraorbital foramen above
posterior root of M1, suggesting a shorter face; upper molars
broad; M3 with split metastyle and prominent posterior cin-
gulum on metaconule; ratio of length of P‘~4 to length of
M'~3 approximately 0.73, considerably smaller than the
Merychyus cf. M. minimus described above; lower premolars
crowded, P3 10 percent larger and more complex than Mer-
ychyus cf. M. minimus described above.

DISCUSSION. Although these three specimens may sim-
ply be larger, shorter faced, more robust individuals of Mer-
ychyus cf. M. minimus described above, they are separated
because of the more robust teeth, deep jugal, and large P3.
Specimen UCR 10322 is a young adult which may account
for the crowding of the lower premolars. The infraorbital
foramen is clearly more posteriorly located than in the three
specimens of Merychyus cf. M. minimus described
above, suggestive of the condition found in species of Bra-
chycrus Matthew, 1901. None of the Boron specimens pre-
serves the nasal region of the cranium which is diagnostic of
Brachycrus. The split metastyle and strong posterior cingu-
lum of M3 and jugal depth are variable characters in ore-
odonts (Lander, personal communication). The jugal, al-
though deep for species of Merychyus, is nowhere near as
deep as in Brachycrus, but is similar to species referred to
Ticholeptus. The Boron specimens are much smaller than
any species of Brachycrus, but only slightly smaller than some
specimens referred to Ticholeptus calimontanus.

These three specimens may be variants of the associated
Merychyus cf. M. minimus, but there is more difference be-
tween them and the Merychyus cf. M. minimus than between
the latter and M. minimus from Nebraska. Thus, they are

Contributions in Science, Number 355

Whistler: Boron Local Fauna 23

considered distinct until larger samples are available to more
clearly demonstrate individual variation. There is some sug-
gestion of convergence with features characteristic of both
Brachycrus and Ticholeptus.

Family Camelidae
Subfamily Aepycamelinae
Hesperocamelus Macdonald, 1 949

cf. Hesperocamelus sp.

Table 4

MATERIAL. Palate with RP‘~2, RM2 and part of RM!,
LP2~3, LM1"3, isolated ? upper incisor, UCR 10366; LM3,
UCR 10367; posterior portion, RM„ UCR 10368; associated
left distal radioulna, scaphoid and lunar, UCR 10384; distal
epiphysis, left radioulna, UCR 10385; right distal tibia, UCR
10388; two left distal tibiae, UCR 10387 and UCR 10389;
left astragalus, UCR 1 0392; left calcaneum, UCR 2 1 1 86; left
proximal metacarpal, UCR 21 187; right proximal metatar-
sal, UCR 21 188; proximal phalanx, UCR 10391.

DESCRIPTION. Size (length of dentition from P1 to M3)
larger than the aepycamelines Oxydactylus Peterson, 1904,
and Paratylopus Matthew, 1904, the camelines Dyseotylo-
pus Stock, 1935, and Priscocamelus Stevens, 1969, and most
protolabines; about same size as the aepycameline Hespero-
camelus alexandrae (Davidson, 1923); smaller than smallest
species of Aepycamelus Macdonald, 1956; rostrum fairly nar-
row at anterior root of P2 but not constricted as in protola-
bines; teeth relatively low crowned with 80-90 percent of
external enamel of paracone of M3 exposed when this tooth
enters occlusion; premolars large in comparison to molars;
P1 large, blade-like, two rooted; P2 long, blade-like, with
weak, discontinuous lingual cingulum; P3 long, parastyle large,
lingual cingulum well developed, with distinct central gap;
M'~3 with weak, but distinct vertical ribs on paracone, ribs
nearly lacking on metacone, metastyle and parastyle strong;
M3 not elongated anteroposteriorly compared to M1; M3 with
strong metastylid and reduced hypoconulid; metacarpals and
metatarsals fused; calcaneum long and slender.

DISCUSSION. Camel taxonomy is usually based on as-
sociated skulls and postcranial material. Tentative associa-
tion of the palate UCR 20366 with some of the postcranial
material provides for a composite picture of this medium
sized camel. The palate is broken anterior to the P2 so that
the degree of palatal constriction and length of the rostrum
are lacking (key characters in distinguishing protolabines from
aepycamelines). Although the palate is fairly narrow between
the anterior roots of the P2, these teeth are not turned inward
anteriorly as is typical in protolabines.

The premolars are unreduced and large in comparison to
the molars, a feature typical of many Early Miocene camels,
but they lack the robustness typical of Miolabis Hay, 1899
(Aepycamelinae). The large, blade-like P' is similar to that
of most protolabines and aepycamelines, but P1 in Hespero-
camelus is typically more caniniform. The P2 in the cf. Hes-

perocamelus sp. from Boron is larger in comparison to the
other cheek teeth than in most protolabines or aepycame-
lines. The P3 is also large. The discontinuous lingual cingulum
of the P3 is a variable character but is similar to the primitive
protolabine Michenia Frick and Taylor, 1971, and Hesper-
ocamelus. The crown of P4 is missing on both sides of the
palate.

The molars are low crowned as is characteristic of aepy-
camelines and most protolabines and in contrast to the higher
crowned cameline Procamelus Leidy, 1858, and the notably
high-crowned stenomylines (Frick and Taylor, 1968). The
vertical ribs on the paracone and metacone are more reduced
than those of the primitive protolabines Michenia and Tany-
mykter Honey and Taylor, 1978, and the primitive aepy-
camelines Oxydactylus and Miolabis. They are more like
those of stenomylines and the more derived aepycamelines
Hesperocamelus and Aepycamelus. However, the strong me-
sostyle is more like that of Michenia, Tanymykter, and Oxy-
dactylus. The M3 is not anteroposteriorly lengthened in re-
lation to the length of the M1 as is common in many derived
protolabines. Two isolated M,’s bear relatively strong meta-
stylids which are typical of most protolabines and aepy-
camelines, otherwise these referred specimens provide little
additional information.

Fragments of a fused metacarpal and fused metatarsal oc-
cur in the fauna, but neither are complete enough to obtain
an estimate of their length. The degree of fusion distinguishes
this Boron material from Miolabis, Paratylopus, most species
of Oxydactylus and all species of Michenia except M. exilis
(Matthew, 1 960) all of which have unfused metapodials. There
is one partial calcaneum which is quite long and slender, a
condition common in stilt-legged (aepycameline) camels.

In most observable characters, cf. Hesperocamelus sp. from
Boron is not clearly distinguishable from early protolabines
( Michenia , Tanymykter) or early aepycamelines ( Oxydac-
tylus). As clearly pointed out by Honey and Taylor (1978:
377), it is difficult to distinguish these taxa on cheek teeth
alone. However, the Boron cf. Hesperocamelus sp. seems to
lack the restricted rostrum characteristic of all protolabines
and it is larger than any species of Oxydactylus. Thus, it is
referred to the larger aepycameline with fused metapodials,
Hesperocamelus. Except for the blade-like P1 and slightly
smaller size, it closely resembles Hesperocamelus alexandrae
from the Barstow Formation.

Subfamily undetermined

Camelidae, small species

Table 4

MATERIAL. Juvenile dentaries with LdP2_4, RdP4, RM,,
and LM,, UCR 10365; proximal right metacarpals III and
IV, UCR 21189; proximal right metatarsal, UCR 10376;
proximal right scapula, UCR 10370; proximal end right ra-
dioulna, UCR 10371; proximal end left radioulna, UCR
10372; proximal articular facet right radioulna, UCR 10373;
distal epiphysis right radioulna, UCR 10374; right lunar UCR
21190; right entocuneiform, UCR 10382; distal left tibia,
UCR 10377; right malleolus, UCR 21191; right juvenile cal-

24 Contributions in Science, Number 355

Whistler: Boron Local Fauna

caneum, UCR 10379; right astragalus, UCR 10378; two right
cuboids, UCR 10393 and UCR 10394; right navicular, UCR
10395; proximal end, proximal phalanx, UCR 10381.

DISCUSSION. This small camel is represented by a pair
ofjuvenile dentaries with the M, the only erupted permanent
tooth and with the dP3_4 moderately worn. The specimen is
broken at the symphysis, but there is an alveolus for a large
P,. Additional small, postcranial camelid elements may also
belong to this taxon. One fragment represents the proximal
ends of metacarpals III and IV which are closely appressed
but unfused; a fragment of the proximal end of the metatarsal
is fused.

The material is too incomplete to permit even a subfamilial
assignment, but clearly demonstrates the presence of a small
camel in the fauna. Based on postcranial elements, this is the
most common camel. This small camel could be any of sev-
eral protolabines that are common in Miocene assemblages
of the western United States or a species of Oxydactylus.
Priscocamelus or Paratylopus, all less common.

Camelidae, large species

DISCUSSION. A large camel is represented by a left distal
femur fragment, UCR 10386, and a left proximal scapular
fragment, UCR 10383. These specimens suggest an animal
about half again as large as the species referred to cf. Hes-
perocamelus sp. They could represent one of several small
species of Aepycamelus which are common in Miocene as-
semblages of the Great Basin.

Family Palaeomerycidae

Subfamily Aletomerycinae
Aletomeryx Lull, 1920

Aletomeryx occidentalis new species

Figures 51-59, Table 5

HOLOTYPE. UCR 10335, partial young adult right den-
tary with P2 (natural cast in part) and complete P3 through
M3.

TYPE LOCALITY. UCR locality RV 6421 (see complete
description under Cupidinimus boronensis above).

REFERRED MATERIAL. Portion of right frontal and
horn base, UCR 10348; partial left dentary with I2, well worn
P2-M2 and roots for I,, I3, and C, UCR 10336; partial right
dentary with little worn P2-M2, UCR 10337; partial sub-
adult right dentary with dP2, unworn P3-M2, UCR 10338;
fragment of right dentary with moderately worn M2_3, UCR
10339; fragment of right dentary with moderately worn M„
UCR 10340; partial right juvenile dentary with symphysis,
early wear dP2_4, M,, UCR 10341; partial right juvenile den-
tary with early wear dP2_.„ M, UCR 10342; partial left ju-
venile dentary with early wear dP2_^, UCR 10343; partial
right maxilla with early wear M 1-3, UCR 10344; partial young
adult right maxilla with alveolus for P1-2, partially complete
P4 and M1-2; UCR 10345; isolated fragment LdP2, UCR
10347; isolated fragment ?RP2, UCR 10398; left proximal
scapula, UCR 10352; two right distal humeri, UCR 10356

mesostyle
metacone
metastYle^^.

metaconule

paracone metacomd parastylid

entostyle

entoconid/ /
entostylid

metaconid

I / paraconid
/ / ...parastylid

protoconid

hypocomd

Figure 50. Tooth cusp terminology used in Aletomeryx.

and UCR 10357; left proximal radius, UCR 10351; distal
half right radioulna, UCR 10349; associated distal end left
radioulna, scaphoid, lunar, unciform, fused magnum-trap-
ezoid, proximal metacarpal, UCR 1 0350; left cuneiform, UCR
21192; associated right tibia lacking proximal articulation,
astragalus, calcaneum, naviculocuboid, entocuneiform, and
proximal fused metatarsals III and IV, UCR 21 193; four left
distal tibiae, UCR 10358, UCR 10359, UCR 10360, and
UCR 10364; right calcaneum, UCR 10361; associated left
naviculocuboid, entocuneiform, fused metatarsals III and IV,
vestiges of metatarsus II and V, UCR 10355; two left na-
viculocuboids, UCR 10362 and UCR 10363; left entocu-
neiform, UCR 10354; proximal phalanx UCR 21 194.

DIAGNOSIS. Slightly larger (1.5 percent) than Aletome-
ryx marslandesis Frick, 1937, the largest species of Aleto-
meryx, but smaller (about 2 percent than the smallest species
of Sinclairomeryx Frick, 1937; horn Aletomeryx- like, situ-
ated over posterior part of orbit with slight forward tilt; den-
tition hypsobrachyodont; P2 large for species of Aletomeryx;
premolars with simple pattern compared to common pa-
laeomerycids such as Dromomeryx Douglass, 1 909, but more
complicated pattern than that of described species of Ale-
tomeryx; lower premolars with relatively straight crests and
lacking expanded metaconid; posterior labial comer (hypo-
stylid) of P3_4 expanded into distinct cuspid that connects to
hypoconid by short crest in moderate wear; labial end of
central crest (metaconid) slightly expanded anterolabially in
P4 but much less developed than in dromomerycines and
giraffids; opening of lingual flexids of P4 not directed poste-
riorly as in dromomerycines; M,_, with well-developed an-
terior cingula and anterolabial buttresses which join in wear;
hypoconulid of M3 formed by closed lingual crescent or cres-
cent in combination with smaller lingual cusp; post-sym-
physial diastema short; limbs lightly constructed, proximal
ends of metatarsals II and V retained as vestiges.

ETYMOLOGY. From occidentalis Latin, of the west, in
reference to the occurrence of this species in far western range
of distribution of the genus.

DESCRIPTION. Orbital Horn. Single broken pedicel,
UCR 10348, situated over posterior portion of orbit, with
slight forward tilt and some suggestion of forward curves;
external central ridge, leading to postorbital process thinner
than in the small species, Aletomeryx gracilis Lull, 1920;
cross section of hom pedicel at base broadly triangular in
outline; internal surface broadly convex, not flat as Sinclai-

Contributions in Science, Number 355

Whistler: Boron Local Fauna 25

Table 5. Selected measurements of Aletomeryx occidentalis n. sp. Abbreviations: AP = anteroposterior dimension of tooth, TR = width of
tooth at anterior loph(id), CH = crown height at paracone in upper molars, at protoconid in lower premolars, at metaconid in lower molars
(figures given only for unworn specimens), * = approximate.

UCR UCR UCR UCR UCR UCR UCR UCR

Dentary 10335 10336 10337 10338 10339 10341 10342 10343

Length P2-M,

75.2

69.0

Length P-P,

24.0

23.9

27.4

25.0

Length M,-M3

48.8

45.7

P2

AP

5.5*

6.4

7.8

7.0

TR

3.0*

3.5

3.7

3.5

P3

AP

8.4

7.5

9.1

8.0

TR

4.8

4.4

4.9

4.7

CH

6.7*

7.7

P<

AP

9.9

10.0

1 1.1

10.1

TR

6.0

5.2

5.0*

4.5

CH

10.2

10.5

M,

AP

1 1.5

9.1

14.0

13.8

13.1

TR

7.0

6.2

6.5

6.5

6.2

CH

8.6

m2

AP

14.2

14.0

16.0

17*

15*

TR

8.8

8.0

8.5

8.0

-

CH

13.1

1 1.7

M,

AP

20.0

21.7

21.0

TR

8.8

8.1

8.4

CH

12.2

Length dP,-dP4

28.0

26.6

27.0

dP2 AP

6*

5.5

5.4

TR

3.1

2.6'

dP, AP

7.0

6.1

8.0

TR

3.6

4.8

4.0

dP4 AP

14.4

14.1

12.5

TR

5.4

5.8

5*

UCR

UCR

UCR

Maxilla

10344

10345

10347

Length P2-M2

#

00

IT)

Length P2-P4
P2 AP
TR

28*

P4 AP

9.1

TR

9.5'

M1 AP

12*

14.1

TR

11.2

M2 AP

14.6

18.0

TR

14.0

14.2

CH

15.0

M3 AP

14.8

TR

14.6

CH

12.0

26 Contributions in Science, Number 355

Whistler: Boron Local Fauna

Table 5. Continued.

Axial skeleton

UCR

10349

UCR

10350

UCR

10356

UCR

10357

Width distal humerus

25.0

23.3

Width distal radioulna

22.8

21.4

Width proximal metacarpal

18.3

UCR

UCR

UCR

UCR

UCR

UCR

UCR

UCR

10355

10358

10360

10361

10362

10363

10364

21193

Width distal tibia

23.8

23.5

23.0

21*

Width naviculocuboid

20.0

19.0

21.4

20.6

Length calcaneum

54.6

51*

Length astragalus

23.8

Width prox. metatarsal

20.0

19.0

romeryx; horn pedicel becomes triangular along posterior
border above base; two foramina separated by a small bridge
of bone present in frontal above orbit; suggestion of third
foramen at frontal-lacrimal suture.

Upper Dentition. Enamel more rugose than in lower den-
tition; styles well developed, but not greatly enlarged; alveo-
lus for P2 in UCR 10345 suggests broader tooth than in other
species of Aletomeryx; alveolus for P3 about same size as in
other species of Aletomeryx or Sinclairomeryx; P2 in UCR
10398 with well developed lingual cingulum; P4 only partially
preserved, but with fossette divided posteriorly; M1-3 with
well-developed styles and accessory folds in fossettes; weak
anterior cingulum; small pillar (entostyle) in M1-2 lacking in
M3.

Upper Deciduous Dentition. dP2 similar to permanent P2
with single high labial crest; continuous lingual cingulum
around anterior end of tooth with shallow groove below high
portion of labial crest (protocone); small spur leading from
center of anterior portion of cingulum to base of protocone.

Dentaries and Lower Dentition. Lower jaws lightly built,
with slight inward curve in region of symphysis; post-sym-
physial diastema proportionally shorter than in smaller species
of Aletomeryx ; molars high crowned with less than half the
entire crown height of M3 exposed when this tooth comes
into occlusion; enamel lightly rugose; premolars, particularly
P2, less reduced than in other described species of Aletomeryx
or Sinclairomeryx.

I , 3 and lower canine all about the same size judging from

roots, all directed anteriorly.

P2 (based on UCR 10327 and UCR 10336) larger, with
more complicated pattern than in previously known Aleto-
meryx species; with well-developed anterior cusp (paraconid)
that persists in wear; paraconid connected by labial crest to
high central cusp (protoconid); separate entoconid and en-
tostylid present in early wear.

P3 with more complicated cusp pattern than P2; paraconid
and parastylid well developed, becoming more separate near
base of crown; protoconid most prominent cusp; entoconid
and entostylid well developed with intervening flexid opening

more labially than in most Aletomeryx species; suggestion of
low postero-labial enlargement (hypostylid) which occludes
only in late stage of overall tooth wear.

P4 with separate and distinct paraconid and parastylid;
parastylid an isolated cusp in early wear; metaconid begin-
ning as isolated cusp but strongly connected to protoconid
by central crest; metastylid absent; posterior portion of P4
distinctly bifurcate with separate entoconid and entostylid;
flexid between entoconid and entostylid opens more poste-
riorly than in P3; posterolabial comer (hypostylid) well de-
veloped to mid height of tooth crown.

M, small in proportion to P4 compared to other species of
Aletomeryx or Sinclairomeryx; parastylid, anterior cingu-
lum, and anterolabial buttress (protostylid) well developed.

M2 with well-developed anterior cingulum bearing distinct
anterolabial cusp or buttress (protostylid) in unworn tooth,
cusp joins cingulum after moderate wear, persists to late
wear; suggestion of metastylid present as inflection in enamel
in UCR 10337 and UCR 10338, but not persistent in wear;
ectostylid developed as small, low pillar; connection between
posterior end of entoconid and posterolingual extension of
hypoconid (= entostylid) delayed until moderate wear, form-
ing a shallow enamel infolding in posterolingual comer of
tooth.

M3 with hypoconulid formed by completely closed labial
loop in the type, UCR 10335, and UCR 10336; UCR 10339
(Figures 47-48) with additional small, lingually projected
spur and small lingual cusp; anterior cingulum not as well
developed as in M, or M2; ectostylids variably developed
between protoconid and hypoconid, hypoconid and hypo-
conulid.

Lower Deciduous Dentition. dP2 very similar to the P2,
except with deeper posterior flexid more persistent with wear;
dP3 with better developed paraconid, parastylid, and bifur-
cation of the anterior cingulum; posterior flexid of dP3 open-
ing more directly lingually than in P3, with small cusp in
anterior labial comer (protostylid) that forms anteriorly
opening flexid after moderate wear; dP4 three lobed, resem-
bling reversed M3, but with stronger ectostylids.

Contributions in Science, Number 355

Whistler: Boron Local Fauna 27

Figures 51-59. Aletomeryx occidentalis n. sp. Figures 51-52, Holotype, UCR 10335, right adult mandible with early wear P2 (in part natural
cast) and complete P3-M3; (51) labial view, (52) occlusal view. Figures 53-54, UCR 10339, RM3; (53) lingual view, (54) occlusal view. Figures
55-56, UCR 10342, juvenile right dentary fragment with early wear RdP2_4, part of M,; (55) labial view, (56) occlusal view. Figure 57, UCR
10348, right horn fragment. Figures 58-59, UCR 10344, RM2-3; (58) labial view, (59) occlusal view. All actual size.

Limbs. Long and lightly built; front and hind metapodials
well fused; principal metatarsal with vestiges of proximal
ends of both metatarsals II and V; vestigial metatarsal II
elongate, triangular, fused to central metatarsal; vestigial
metatarsal V stubby, broad, unfused; vestige of metacarpal

II also present as suggested by flattened, rugose area on meta-
carpal III.

Discussion. The dentition of Aletomeryx occidentalis is
high crowned, more so than most taxa referred to Palaeo-
merycidae. The high-crowned teeth and narrow P,_4 distin-

28 Contributions in Science, Number 355

Whistler: Boron Local Fauna

guish Aletomeryx occidentalis from all the more deer- or
giraffe-like palaeomerycids: Dromomeryx, Rakomeryx Frick,
1937, Cranioceras Matthew, 1918, Barbouromeryx Frick,
1937, Drepanomeryx Sinclair, 1915, and Yumaceras Frick,
1937, of North America, Palaeotneryx Von Meyer, 1834,
Procervulus Gaudry, 1878, Heterocemas Young, 1937, and
Triceromeryx Villalta, Crusafont and Lavocat, 1946, from
Europe and Asia and Climacoceras Maclnnes, 1936, Can-
thumeryx Hamilton, 1973, and Propataeoryx Slromer, 1926,
from Africa. Aletomeryx occidentalis is distinguished from
the smaller, hornless, North American leptomerycine Lep-
tomeryx Leidy, 1853, Pseudoparablastomeryx Frick, 1937,
and Pronodens Koemer, 1940 (see Taylor and Webb, 1976),
from the blastomerycines Blastomeryx Cope, 1877, and Ma-
chaeromeryx Matthew, 1926, and from the palaeomerycid
Longirostromeryx Frick, 1937, by the high-crowned teeth
and presence of orbital horns.

Only three palaeomerycids, Aletomeryx, “ Dyseomeryx,”
and Sinclairomeryx, have high-crowned, antilocaprid-like
teeth. The relatively large premolars, particularly P2, and the
greater complexity of P3_4, with a well-developed metaconid
on P4 and hypostylid on P3_4, are characters derived with
respect to any previously described species of Aletomeryx or
Sinclairomeryx.

There is little clear distinction between species of Aleto-
meryx, “ Dyseomeryx ," and Sinclairomeryx because many
characters overlap. However, the structure of the horn ped-
icel, the development of the anterolabial buttress and anterior
cingulum on M2, and the hypoconulid crescent of M3 favor
assignment of the Boron species to the genus Aletomeryx.
Some smaller specimens of Aletomeryx gracilis approach the
crown height of Aletomeryx occidentalis, but these lack the
lower premolar complexity.

There is also confusion as to the validity of the generic (or
subgeneric) names Aletomeryx, Dyseomeryx, and Sinclai-
romeryx within the Subfamily Aletomerycinae. According to
Stirton (1944) in his sort discussion of the Family Palaeo-
merycidae, the genus Aletomeryx Lull, 1 920, is distinct but
the genera (or subgenera) Dyseomeryx Matthew, 1924, and
Sinclairomeryx Frick, 1937, are synonymous, Dyseomeryx
with priority. He did not thoroughly discuss his reasons for
this synonymy, however. On the other hand, Frick (1937)
stated that Dyseomeryx is in part synonymous with Aleto-
meryx.

Matthew (1924:196) defined the subgenus Dyseomeryx,
choosing Blastomeryx marshi Lull, 1920, as the type, which
then became Blastomeryx {Dyseomeryx) marshi (Lull) as re-
combined by Matthew (1924). But, according to Frick (1937:
152), Blastomeryx {Dyseomeryx) marshi (Lull) actually be-
longs in the genus Aletomeryx, and therefore Blastomeryx
(Dyseomeryx) marshi became Aletomeryx marshi (Lull) as
recombined by Frick (1938). Because Frick (1937) found it
necessary to refer the type of the subgenus (Dyseomeryx) to
Aletomeryx, while he considered other forms referred to (Dy-
seomeryx) generically distinct, he defined the new genus Sin-
clairomeryx, naming as the type species Sinclairomeryx sin-
clairi Frick, 1937. Frick also questionably assigned another
of Matthew’s species, Blastomeryx (Dyseomeryx) riparius

Matthew, 1924, to Sinclairomeryx. The type of this species
is a left maxilla fragment, and certain assignment to Sin-
clairomeryx by Frick would necessitate an associated horn
core, which was not present. Based on comparisons of den-
titions, later workers (Skinner et al., 1977) have indicated 5.
riparius and S. sinclairi are synonymous, with S. riparius
having priority.

Although I have not reviewed the original material, it would
seem that Frick (1937) substantiated his view and I have
accepted the genera Sinclairomeryx (= Dyseomeryx in part)
and Aletomeryx.

AGE AND BIOSTRATIGRAPHY
RADIOMETRIC CORRELATION

There are three radiometric dates on different outcrops of
the Saddleback Basalt: 18.3 ± 0.6 ma, 19.7 ± 0.6 ma (R.B.
Kistler, personal communication), and 20.3 ± 0.7 ma (Arm-
strong and Higgens, 1973). The Saddleback Basalt occurs
146 m below the Boron Local Fauna in the open pit mine
and thus these radiometric dates provide a maximum age
for the fossil assemblage. Based on biostratigraphic corre-
lations (see below), the Boron Local Fauna is distinctly older
than the typical fossil vertebrate assemblages of the Barstow
Formation which have a maximum age of at least 16.3 ±
0.3 ma (Woodbume and Tedford, 1982) and the Early Bar-
stovian Dry Canyon fauna of the Caliente Formation dated
at 16.5 ± 1.3 ma (Turner, 1970, KA 2127). These dates
provide a minimum age for the Boron Local Fauna. On
biostratigraphic grounds, the Boron Local Fauna appears close
to the Phillips Ranch Local Fauna which is underlain by a
17.6 ma date (Evemden et al., 1964, KA 478). All these form
a consistent pattern with the Boron Local Fauna falling be-
tween approximately 16.5 and 18.5 ma.

BIOSTRATIGRAPHIC CORRELATIONS

Generally, correlation of southern Californian Arikareean
and Hemingfordian faunas with the classical sequence in
Nebraska and neighboring states has been difficult because
the fossil assemblages in these widely separated areas usually
have had little in common. The Boron Local Fauna presents
a rare exception to this generality for it contains two taxa
very similar to species in Nebraska, Merychyus cf. M. min-
imus and Aletomeryx occidentalis. Merychyus minimus, re-
stricted to the early Hemingfordian of the central United
States, is a quite variable species and M. cf. M. minimus
from Boron falls within this variation. A. occidentalis, al-
though distinct at the species level and probably representing
a separate lineage from the Aletomeryx/Sinclairomeryx lin-
eage in the central United States, is close to A. marslandensis
from the early Hemingfordian of Nebraska. Taken as a
whole, the Boron Local Fauna is best considered early
Hemingfordian in age, but probably near the transition to
the late Hemingfordian.

Compared to mid-continent assemblages, the Boron Local
Fauna is not particularly diverse, and may lack “key” taxa
which were not preserved or may not have been present. To

Contributions in Science, Number 355

Whistler: Boron Local Fauna 29

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30 Contributions in Science, Number 355

Whistler: Boron Local Fauna

some extent, the Boron Local Fauna represents a unique
combination of taxa, with a rodent ( Trogomys ) previously
thought to be Arikareean, a palaeomerycid genus (Aleto-
meryx) and an oreodont species ( Merychyus cf. M. minimus),
elsewhere restricted to the early Hemingfordian, and a di-
podomyine rodent genus (Cupidinimus) and camel (cf. Hes-
perocamelus sp.) the latter genus previously restricted to the
Barstovian and later.

Absent at Boron are taxa which are generally typical of
late Hemingfordian: small merychippines, middle sized ore-
odonts larger than Merychyus (Brachycrus, Ticholeptus), dro-
momerycids (Bouromeryx Frick, 1937), and antilocaprids.
Conclusions based on absent taxa are always questionable,
but the lack of these taxa found in common association in
other late Hemingfordian faunas of southern California and
Nebraska strongly suggest a real absence in the Boron Local
Fauna, not just an artifact of the record and further supports
a slightly older age than late Hemingfordian. Camel biostra-
tigraphy is not well documented, and thus the cf. Hespero-
camelus sp., a genus previously restricted to Barstovian and
later, is not considered particularly significant.

The microfauna, although seemingly well preserved, lacks
the common Arikareean/Hemingfordian genus Prohetero-
mys and the eomyid/geomyoid/zapodid elements common
in earlier Miocene assemblages. Cupidinimus, the most com-
mon taxon in the Boron Local Fauna and a common rodent
in the earliest Barstovian, is absent in the only other Hem-
ingfordian microvertebrate assemblage in southern Califor-
nia (Vedder Local Fauna; Lindsay, 1 974) and thus may seem
out of place at Boron. However, its absence may be envi-
ronmentally controlled and its presence at Boron merely pro-
vides an earlier link between this common Barstovian taxon
and its presumed ancestor, Trogomys.

The Boron Local Fauna displays greatest similarity with
the central United States fossil assemblages of the Running-
water Formation (Cook, 1965; McKenna, 1965), Rosebud
Formation (Macdonald, 1963, 1970; Macdonald and Hark-
sen, 1968), Batesland Formation (Harksen and Macdonald,
1967; Martin, 1976), and Martin Canyon “beds” (Galbreath,
1953; Wilson, 1 960). These fossil assemblages are considered
typical of early Hemingfordian (Tedford et al., 1973). In
addition to M. minimus and Aletomeryx species, these faunas
also contain taxa similar to the Boron forms such as Ar-
chaeolagus, Hypolagus, Miospermophilus, Mookomys, prim-
itive aepycamelines ( Oxydactylus ) and primitive protola-
bines ( Tanymykter , Protolabis), even though none of these
taxa are limited to the early Hemingfordian.

There is less similarity between the Boron Local Fauna
and the later Hemingfordian fossil assemblages of the Sheep
Creek Formation (Skinner et al., 1977) and the Box Butte
Formation (Galusha, 1975), the latter containing a diversity
of horses, including Merychippus spp., larger oreodonts (Bra-
chycrus, M. ( Metoreodon ), Ticholeptus), Sinclairomeryx spp.
(but no Aletomeryx), dromomerycines, and antilocaprids.

The Boron Local Fauna can also be correlated with south-
ern California fossil assemblages (Figure 60). However, such
correlations present some difficulty because of the seemingly
unusual assemblage of taxa in the Boron Local Fauna. The

Arikareean/Hemingfordian interval is not well represented
in southern California. Most fossil assemblages referred to
this time span are poorly preserved, lack taxonomic diversity
and most correlations with the central United States standard
have been tenuous. In addition, the Boron Local Fauna ap-
pears to sample a restricted time interval not previously seen
in southern California or the southern Great Basin.

Most Arikareean fossil assemblages from southern Cali-
fornia have been derived from sedimentary basins west of
the San Andreas Fault, which, by currently accepted plate
tectonic models, places these basins from 100-300 km to the
southeast at their time of development (see Crowell, 1975).
Whether such a distance would result in significantly different
paleoenvironmental conditions than those east of the San
Andreas Fault is impossible to resolve. Pyroclastic volcanic
deposition dominated in the few basins in existence east of
the San Andreas Fault in the Mojave Desert and only one
area has yielded fossils. By later Hemingfordian, this vol-
canism had subsided considerably, but most of the basins
east of the San Andreas Fault were dominated by coarse
clastic deposition as these volcanics were removed from areas
of higher topographic relief. Fossils in these coarse sediments
are scattered and poorly preserved. The earlier Hemingfor-
dian was not previously sampled (or recognized, if sampled).
The Boron Local Fauna samples this interval for the first
time.

Among those southern California assemblages referred to
the Arikareean, the Boron Local Fauna shows some simi-
larity to the Tick Canyon Local Fauna (Maxson, 1 930; Jahns,
1940) in sharing Trogomys. Merychyus cf. M. minimus from
Boron is derived with respect to M. calaminthus from the
Tick Canyon Local Fauna. The camel referred to Miolabis
californicus Maxson, 1930, from the Tick Canyon Local Fau-
na has unfused metapodials and is considerably larger than
the small camelid from Boron with unfused metapodials.

Merychyus cf. M. minimus from Boron is also derived with
respect to the M. calaminthus from the Late Arikareean Black
Butte Mine Local Fauna in the Hector Formation (Wood-
bume et al., 1974) and the Merychyus cf. M. calaminthus
from the Cady Mountain Local Fauna (Woodbume, Miller,
and Tedford, 1982). The fused, proximal metatarsal from
the small camelid species from Boron is considerably larger
than the Michenia cf. M. agatensis from the latest Arikareean
Logan Mine Local Fauna of the Hector Formation.

The Boron Local Fauna shows little similarity to other
Arikareean assemblages in California such as the Tecuya
Local Fauna (Stock, 1920, 1932a), Kew Quarry (Stock, 1932b,
1935; Wilson, 1934, 1949), and other localities in the upper
Sespe Formation (Wilson, 1949).

Among southern California fossil assemblages referred to
the late Hemingfordian, the Boron Local Fauna closely re-
sembles the poorly documented Phillips Ranch Local Fauna
(localities LACM(CIT) 305, UCMP V2577) in the Kinnick
Formation (or the basal Bopesta Formation, James Quinn,
personal communication). Buwalda (1916) and Buwalda and
Lewis (1955) recognized Merychippus tehachapiensis Bu-
walda and Lewis, 1955 (= M. carrizoensis Dougherty, 1 940—
see Munthe, 1 979), small and large camelids, Moropus Marsh,

Contributions in Science, Number 355

Whistler: Boron Local Fauna 31

1877, Merycodus Leidy, 1854, canids, and felids from this
local fauna. Although not published, the Phillips Ranch Local
Fauna also contains an oreodont, peccary, and palaeomer-
ycid. The oreodont is a small, high-crowned species probably
referable to Merychyus. It is slightly larger than the Boron
oreodonts but otherwise similar to M. cf. M. minimus. The
palaeomerycid (represented only by upper teeth) is a small,
high-crowned species similar in size and morphology to A.
occidentalis or a small species of Rakomeryx. The small cam-
el from Phillips Ranch is intermediate in size between the
cf. Hesperocamelus sp. and the small camel species from
Boron but appears to have fused metatarsals like the cf. Hes-
perocamelus sp. from Boron.

The oreodont from the Late Hemingfordian Red Division
Local Fauna in the type section of the Barstow Formation
in the Mud Hills (Woodbume and Tedford, 1982), Meryc-
hyus ( Metoreodon ) relictus fletcheri Schultz and Falkenbach,
1 947, is based on a heavily worn, poorly preserved specimen.
About all that can be said is that this individual is larger than
the oreodonts from Boron. The Red Division Local Fauna
also produces Merychippus carrizoensis which is absent at
Boron.

Among other late Hemingfordian fossil assemblages in
southern California, the Boron Local Fauna shares no taxa
with the Alvord Mountain faunas from the lower member
of the Barstow Formation (localities D“319,” D321) (Byers,
1960; Lewis, 1968), the Yermo Hills Local Fauna from sed-
imentary rocks in the Yermo Hills also referred to the Bar-
stow Formation (Woodbume, Miller, and Tedford, 1982),
the fauna of the basal Punchbowl Formation in the Cajon
Valley (Woodbume and Golz, 1972), Hidden Treasure Spring
Local Fauna of the Caliente Formation in the upper Cuyama
Valley (Dougherty, 1940b; Repenning and Vedder, 1961;
James, 1 963), the Vedder Local Fauna of the Branch Canyon
Formation (Hutchison and Lindsay, 1974; Lindsay, 1974;
Munthe, 1979) or the Upper Cady Mountain Local Fauna
in the northern Cady Mountains (Miller, 1980; Woodbume,
Miller, and Tedford, 1982). All these have been referred to
the late Hemingfordian and all contain the small Merychip-
pus carrizoensis, middle sized oreodonts (except the Cady
Mountain Local Fauna) referred either to Ticholeptus cali-
montanus, Merychyus ( Metoreodon ) relictus or Brachyus bu-
waldi, and small camelids (except the Alvord Mountain area),
often referred to Protolabinae. Although the taxonomic as-
signments may be erroneous, all these oreodonts are derived
with respect to those from Boron. The only palaeomerycids
in any of these faunas are brachyodont dromomerycines.

The Vedder Local Fauna is the only one of these late Hem-
ingfordian assemblages to also contain microvertebrates ex-
cept for 5 isolated specimens of Proheteromys sulculus from
the Cady Mountain Local Fauna. The Vedder Local Fauna
shares only one taxon ( Mookomys ) at the generic level with
the Boron Local Fauna, but the Boron species is distinctly
smaller. The Boron Local Fauna lacks the squirrels, eomyids,
and Proheteromys species found in the Vedder Local Fauna,
and the P. sulculus found in the Cady Mountain Local Fauna.
On the other hand, the Vedder Local Fauna lacks Trogomys
and Cupidinimus of the Boron Local Fauna, even though the

latter is a common element in earliest Barstovian and later
Miocene faunas. The Vedder Local Fauna is from near shore
or possibly even marine rocks, and perhaps the absence of
Cupidinimus is environmentally controlled.

Taken on balance, the Boron Local Fauna is best consid-
ered early Hemingfordian, a view consistent with the avail-
able radiometric data and biostratigraphic correlations with
the reference fossil assemblages in Southern California and
the central United States.

CONCLUSIONS

Based on the above discussions, the Boron Local Fauna:

1 . is the first diverse early Hemingfordian fossil vertebrate
assemblage from southern California,

2. is the only diverse Hemingfordian microvertebrate as-
semblage in southern California east of the San Andreas
Fault,

3. contains two new mammalian species, Cupidinimus bo-
ronensis n. sp. (Mammalia, Rodentia, Heteromyidae)
and Aletomeryx occidentalis n. sp. (Mammalia, Artio-
dactyla, Palaeomerycidae),

4. contains one of the earliest North American records of
the Family Colubridae (Reptilia, Squamata) and the first
Hemingfordian record west of the Rocky Mountains,

5. contains the earliest species of the genus Cupidinimus,

6. contains the second occurrence of Trogomys (Mam-
malia, Rodentia, Heteromyidae), extending the range of
this genus from the Arikareean into the Hemingfordian,

7. contains the first association of Cupidinimus and Tro-
gomys, the assumed ancestor of Cupidinimus,

8. contains the first Hemingfordian record of a Great Plains
species of merychyine oreodont, Merychyus minimus
(Mammalia, Artiodactyla, Merycoidodontidae), west of
the Rocky Mountains,

9. contains the oldest record of the genus Hesperocamelus
(Mammalia, Artiodactyla, Camelidae),

10. contains the first aletomerycine palaeomerycid, Aleto-
meryx occidentalis, west of the Rocky Mountains,

1 1 . provides a correlative link to the classical early Hem-
ingfordian assemblages of the mid-continent because of
the occurrence of Merychyus cf. M. minimus and Ale-
tomeryx (both common mid-continent taxa), and

12. provides a minimum age of Early Miocene for the Tro-
pico Group.

ACKNOWLEDGMENTS

I would like to thank the geology staff of the United States
Borax and Chemical Corporation, particularly Robert Kis-
tler, Sig Muessig, James Minette, and Ralph Barnard who
provided assistance in the collection of the vertebrate fossils
and valuable insight into the deposits at Boron. I am indebted
to Richard H. Tedford whose guidance and encouragement
enhanced the original study of this fauna in 1964-1965. Rob-
ert Reynolds volunteered considerable time in preparing some
of the specimens. Comparative collections at the American

32 Contributions in Science, Number 355

Whistler: Boron Local Fauna

Museum of Natural History and the University of California
at Berkeley were made available through assistance from
Richard H. Tedford and J. Howard Hutchison, respectively.
Discussions with Bruce Lander aided the analysis of the ore-
odonts. John M. Harris, William A. Akersten, and Lawrence
G. Barnes have provided valuable review comments. The
illustrations of artiodactyls are by Karoly Fogassy of the Uni-
versity of California at Riverside. Drafting of Figures 1, 2,
and 60 is by Mary Butler. Partial support was provided by
the Hewett Fund of the Department of Geological Sciences,
University of California at Riverside.

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Miocene foraminiferal stages. Geological Society of
America Special Paper 124:91-129.

Webb, S.D. 1965. The osteology of Camelops. Bulletin,
Los Angeles County Museum of Natural History 1:1-
54.

Webb, S.D., and B.E. Taylor. 1980. The phylogeny of horn-
less ruminants and a description of the cranium of Ar-
chaeomeryx. Bulletin, American Museum of Natural
History 167(3): 12 1-1 57.

Wilson, R.W. 1934. Two rodents and a lagomorph from
the Sespe of the Las Posas Hills, California. Carnegie
Institution Washington Publications, Contributions in
Paleontology 453: 1 1-17.

. 1939. Rodents and lagomorphs of the Late Tertiary

Avawatz fauna, California. Carnegie Institution Wash-
ington Publications, Contributions in Paleontology 514:
31-38.

. 1 949. Rodents and lagomorphs of the upper Sespe.

Carnegie Institution Washington Publications, Contri-
butions in Paleontology 584:51-65.

. 1960. Early Miocene rodents and insectivores from

northeastern Colorado. University of Kansas Paleon-
tological Contributions, Vertebrata 7:1-92.

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American Museum Novitates 501:1-19.

. 1932. New Miocene rodents from the Miocene of

Florida. Florida State Geological Survey Bulletin 10:43-
51.

. 1935. Evolution and relationship of the heteromyid

rodents with new forms from the Tertiary of western
North America. Annals Carnegie Museum 24:73-262.

. 1936a. Geomyid rodents from the middle Tertiary.

American Museum of Natural History Novitates 866:
1-23.

. 1936b. Fossil heteromyid rodents in the collections

of the University of California. American Journal of
Science, Fifth Series 32(188):1 12—1 19.

. 1937. Additional material from the Tertiary of the

Cuyama basin of California. American Journal of Sci-
ence 33:29-43.

Woodbume, M.O., and D.J. Golz. 1972. Stratigraphy of
the Punchbowl Formation, Cajon Valley, southern Cal-
ifornia. University of California Publications Geological
Sciences 92: 1-57.

Woodbume, M.O., and R.H. Tedford. 1982. Litho- and
biostratigraphy of the Barstow Formation, Mojave Des-
ert, California. In Guidebook, Geologic Excursions in the
California Desert. Cordilleran Section, 78th Annual
Meeting, Geological Society of America:47-64.

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Miocene oreodont (Merychyinae, Mammalia) from the
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jave Desert, California. Journal of Paleontology 48(1):
6-26.

Accepted 25 May 1984.

36 Contributions in Science, Number 355

Whistler: Boron Local Fauna

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