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HARVARD UNIVERSITY

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LIBRARY

OF THE

Museum of Comparative Zoology

O urr-

TRANSACTIONS

OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY

VOLUME 12

Printed from the
W. W. Whitney Publication Endowment

&M*kc^r'

0

SAN DIEGO, CALIFORNIA

Printed for the Society

1954-1962

COMMITTEE ON PUBLICATION

Joshua L. Baily, Jr.

Carl L. Hubbs

Laurence M. Klauber

Charles C. Haines

mk. ma?.

LIBRARV

DEC 3 11962

HARVARD
WIVERSITY

CONTENTS

1. A new form of Perognathus fortnosus from Baja California,
Mexico. By Laurence M. Huey. March 1, 1954 1-2

2. A preliminary revision of the genus Pselaptrichus (Coleoptera:
Pselaphidae) . By Gordon A. Marsh and Robert O. Schuster.

June 1, 1954 3-28

3. The fossil pit-vipers (Reptilia: Crotalidae) of North Anerica.

By Bayard H. Brattstrom. July 1, 1954 31-46

4. Analysis of the herpetofauna of Baja California, Mexico. By
Charles H. Lowe, Jr., and Kenneth S. Norris. September 10, 1954. 47-64

5. Snakes of the islands in the Gulf of California, Mexico. By Frank

S. Cliff. October 1, 1954 67-97

6. A new race of Dipodomys and a new race of Thomomys from
Arizona. By Laurence M. Huey. February 10, 1955 99-101

7. A revision of the Pacific Coast Phytosi, with a review of the
foreign genera (Coleoptera: Staphylinidae). By Ian Moore.

March 1, 1956 103-151

8. Marine Pleistocene invertebrates from near Puerto Penasco, So-

nora, Mexico. By Leo George Hertlein. June 7, 1956 153-175

9. Additional notes on the Pliocene and Pleistocene fauna of the
Turtle Bay area, Baja California, Mexico. By E. P. Chace.

June 11, 1956 177-179

10. Upper Pleistocene Mollusca from Potrero Canyon, Pacific Pali-
sades, California. By James W. Valentine. July 2, 1956 181-205

11. Notes on some intertidal Coleoptera, with descriptions of the early
stages (Carabidae, Staphylinidae, Malachiidae) . By Ian Moore.
August 24, 1956 207-229

12. A new species of Mcgathymus from Baja California, Mexico
(Lepidoptera: Megathymidae) . By Charles F. Harbison.

March 15, 1957 231-261

13. Notes on the metamorphosis of an Agave-boring butterfly from

Baja California, Mexico. By John Adams Comstock. May 1, 1957. 263-275

14. Fish remains from aboriginal sites in the Punta Penasco region

of Sonora, Mexico. By W. I. Follett. July 12, 1957 279-286

15. A new race of wood rat (Neoto?na) from the Gulf side of central
Baja California, Mexico. By Laurence M. Huey.

September 25, 1957 287-288

16. Late Pleistocene faunas from the northwestern coast of Baja Cali-
fornia, Mexico. By James W. Valentine. September 25, 1957 289-308

17. Type material of Eucalodium orcutti Dall (Gastropoda: Pul-
monata) from Oaxaca, Mexico. By Robert J. Drake.

September 25, 1957 309-3 10

18. The North American species of Hesperus Fauvel, with descrip-
tions of two new species (Coleoptera: Staphylinidae) . By Ian
Moore. October 16, 1958 311-318

19. The marine molluscan fauna of Guadalupe Island, Mexico. By

E. P. Chace. October 16, 1958 319-332

20. A new mollusk from San Felipe, Baja California. By E. P. Chace.
October 16, 1958 333-334

21. Pleistocene mollusks from Tecolote Creek, San Diego, California.

By William K. Emerson and Emery P. Chace. May 27, 1959 335-346

22. Type localities of vascular plants in San Diego County, Cali-
fornia.. By Ethel Bailey Higgins. July 22, 1959 347-406

23. A new race of pocket gopher (Thomomys) from San Fernando
Mission, Baja California, Mexico. By Laurence M. Huey.

February 1, 1960 407-408

24. Two new races of Perognathus spinatus from Baja California,
Mexico. By Laurence M. Huey. February 1, 1960 409-412

25. Comments on the pocket mouse. Perognathus fallax, with de-
scriptions of two new races from Baja California, Mexico. By
Laurence M. Huey. February 1, 1960 413-419

26. Inherent and applied camouflage in the subfamily Geometrinae
(Lepidoptera) , including three new life history studies. By John
Adams Comstock. December 1, 1960 421-439

27. Differentiation of the southwestern tortoises (genus Gopherus) ,
with notes on their habits. By Chapman Grant.

November 15, 1960 441-448

28. Marine mollusks from Los Angeles Bay, Gulf of California. By

James H. McLean. August 15, 1961 449-476

29. Two new species of broad-faced, five-toed kangaroo rats (genus
Dipodomys) . By Laurence M. Huey. August 30, 1962 477-480

/

TRANSACTIONS

OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY

Vol. 12, No. 1, pp. 1-2

March 1, 1954

A NEW FORM OF PEROGNATHUS FORMOSUS
FROM BAJA CALIFORNIA, MEXICO

BY

Laurence M. Huey

Curator of Birds and Mammals, San Diego Society of Natural History

A study of accumulated specimens of Perognathus formosus
in the collection of the San Diego Society of Natural History
reveals an undescribed race of this species from the central, arid
section of the peninsula of Baja California. It may be known as

Perognathus formosus infolatus1 subsp. nov.

San Francisquito Pocket Mouse

Type. — From 7 miles west of San Francisquito Bay, lat. 28°30' N.,
Gulf of California, Baja California, Mexico; No. 15664 collection of the
San Diego Society of Natural History; adult male; collected by Laurence
M. Huey, April 3, 1947.

Measurements of Type. — Total length 187; tail, 104; hind foot, 25;
ear, 6. Skull: Occipitonasal Jength, 27.5; mastoid breadth, 14.6; inter-
orbital constriction, 6.6; nasals, 10.2; alveolar length of upper molar
series, 3.8.

1 Lat. Bleached

ys. ccmp. zool

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MAR l 5 1954

HARVARD
U^VERSiTY

San Diego Society of Natural History

Diagnosis and Comparisons. — In color, Perognathus formosus info/a-
tits is the most pallid, dorsally, of all the known races of Perognathus
formosus. In this character it even exceeds the pallor of the ashen-colored
Perognathus formosus cinerascens which inhabits the extremely arid, stony
areas northward near San Felipe, Baja California, Mexico. It is larger in
size than P. f. cinerascens, and has a proportionately greater cranium and
larger, more inflated mastoid bullae.

Range. — This newly described race is known from 7 miles west of
San Francisquio Bay and Barril, situated not far distant on the Gulf of
California coast (lat. 28°20' N) and from 3 specimens 2 from 3 miles
west of El Marmol, and 1 from 3 miles south of El Marmol, lat. 30°. A
single specimen taken on the barren rocky area northeast of El Marmol
is referable to P. f. cinerascens, though not typical of the latter subspecies.
A change in the character of the region is noticeable in the vicinity of
El Marmol and is well demonstrated in the determination of these speci-
mens. The capture of Perognathus formosus at Barril (28°20' N) marks
the southernmost recorded locality for this species so far.

Specimens examined. — Perognathus formosus formosus, 29 from
Saint George, Washington County, Utah (type locality). Perognathus
formosus mohavensis, 78 from Mohave Desert Region, California (in-
cluding type). Perognathus formosus mesembrinus, 18 from eastern San
Diego County and western Imperial County, California. Perognathus
formosus cinerascens, 8 from San Felipe (type locality) ; 1 from northeast
of El Marmol, Baja California, Mexico. Perognathus formosus infolatus,
4 from Barril; 2 from 7 miles west of San Francisquito Bay (type local-
ity) ; 1 from 3 miles south of El Marmol, and 2 from 3 miles west of
El Marmol, Baja California, Mexico.

TRANSACTIONS

OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY

Volume XII, No. 2, pp. 3-28, plates 1-3, map

A PRELIMINARY REVISION OF THE
GENUS PSELAPTRICHUS

(COLEOPTERA:PSELAPHIDAE)

Gordon A. Marsh and Robert O. Schuster

Department of Entomology and Parasitology

University of California

M%. CQMP. ZOQL
UBRARY

JUN 2 2 1954

HARVARD
UNIVERSITY

SAN DIEGO, CALIFORNIA
Printed for the Society

June 1, 1954

COMMITTEE ON PUBLICATION

Joshua L. Baily

Carl L. Hubbs

Laurence M. Klauber

Charles C. Haines

,.}

Marsh-Schuster — Genus Pselaptrichus

San Diego iocitu or Natural History

INTRODUCTION

The genus Pselaptrkhus represents a small and poorly known group
of bythinine beetles endemic to the montane regions of the western United
States, and, in particular, the coniferous forests of the Pacific Coast as
defined by Cain (1944, p. 110).

In such areas they usually occur in the litter and duff layers of the
dark, humid forest floor. Because of such a secluded habitat and secretive
nature, they were once considered to be rare as is indicated by the paucity
of specimens in the major collections of the West. However, upon the
exploitation of such habitats through the utilization of adequate sampling
and collecting devices, it has been found, as is the case with so many
specialized and "rare" organisms, that they are actually quite common and
form an integral part of the western pselaphid fauna.

Prior to this paper, the genus was represented in literature by three
species, two of which were recently named by Park (1953). The type
species, P. tuberculipalpus, was described by Brendel (1889). Raffray
(1908) integrated the genus with the world pselaphid fauna, while
Bowman (1934) included the genus in his compilation of the Pselaphidae
of North America.

This paper is designed to combine all previous information with the
new material, to clarify the identity of the type species, to describe seven
new species, to provide a key for the integration and recognition of all the
species, and, where possible, to indicate relationships as well as variation
at the specific level.

PROCEDURE

The specimens which constitute the material basis for this paper were
collected during the first nine months of 1953 by the authors in conjunc-
tion with the California Insect Survey (C. I. S. ), by C. D. MacNeill, E. E.
Gilbert, and by other students of the Department of Entomology of the
University of California whose names appear in the text. Additional
specimens were loaned through the courtesy of H. B. Leech, California
Academy of Sciences (C. A. S.), and Orlando Park, Northwestern
University (N. U.).

Collecting methods, for the most part, were divisible into two separate
activities: location of sampling sites by random sifting, and the accumu-
lation of ground litter in paper sacks for subsequent treatment by Berlese
funnels in the laboratory.

Whenever possible, specimens were mounted on both points and slides
for study. Individuals to be placed on slides were cleared in warm lacto-
phenol and mounted in either balsam cr Hoyer's medium. Genitalia were
removed from partially cleared specimens and mounted separately.

The descriptions of all species have been made from pointed specimens
while using a binocular microscope with a magnification of 70 X, and, in
certain instances, individual characters were examined on small specimens

Marsh-Schuster — Genus Pselaptrichus

while using 105 X. Slide mounts of specimens from the type locality were
used to describe mandibular dentition.

The holotype, allotype, or both, as well as some of the paratypes, are
deposited in the entomological collection of the California Academy of
Sciences; the remaining paratypes in the collection of the California
Insect Survey, University of California, unless otherwise indicated in
the text.

The illustrations have been drawn by the authors with the aid of an
ocular grid using both slide and pointed material.

MORPHOLOGY

The following discussion is based on those characters which, under
the present classification, exhibit a greater degree of interspecific varia-
tion and have a certain value in species discernment. The first section
deals only with those characters common to both sexes. This is followed
by a discussion of secondary sexual characters.

EYES. The number of facets is one of the most variable characters in
the genus, varying from four to thirty-six, with the majority of species
having under twelve. This variation is particularly evident in some males
where not only the number but the shape is distinctive. Those species with
a small number of facets have round or subcircular eyes. As the facet
number increases, the eyes become dorsoventrally elongate forming an in-
verted triangle as in roth/ and ornatus, or become subreniform as in
oculatus.

VERTEXAL FOVEAE. The position of the foveae in relation to the
eyes has not been used in previous classifications. However, their position
merits recognition as an added character in distinguishing between certain
species and is particularly useful when dealing with female specimens. If
an imaginary transverse line is constructed through the middle of the eyes,
the foveae will lie anterior, posterior, or directly on such a line providing
the head is oriented in such a manner as to be horizontally parallel to the
plane of focus.

TEMPORA. In the majority of species the tempora are simply rounded
or rounded angulate in dorsal outline (PI. 2, figs. 9, 14), but in cavatus
they form a nearly straight line to the cervix.

CERVICO-OCCIPITAL CARINA. Three species groups are segreg-
ate on the basis of this carina: those in which it does not attain the
posterior border of the vertexal foveae; those in which it just attains but
does not surpass the border; and those in which the carina distinctly
surpasses the posterior border.

FRONTAL PROLONGATION. The median sulcus and its relation
to the lateral convexities provide one means of associating males and
females as this character is constant for both sexes. The sulcus may be a
narrow, vestigal, canaliculate impression which is subequal in width to
the lateral margins as in carinatus. or a broad, flat sulcus occupying a
majority of the dorsal surface of the frontal prolongation, this impres-
sion being wider than the lateral margins as in rothi. The sulcal margins
may be evenly convex, obliquely flattened forming lateral carinae, or

8 San Diego Society of Natural History

completely flattened and highly rugulose. They may also converge apically
as in ornatus or diverge apically as in rothi.

MAXILLARY PALPI. Unfortunately little aid in classification can be
obtained by using this structure to differentiate closely related species.
There are, however, three basic forms of the fourth segment with minor
variations of each. In two species, this segment is elongate, securiform
and approximately three times longer than broad (PI. 3, fig. 17). In all
other species, this segment is approximately two times longer than broad
but is of two moderately distinct forms. In some species the inner or
upper margins forming the declivities are rounded angulate (Pi. 3, fig-
16). In others, the declivity of the upper apical margin is gradually
rounded to the terminal palpal cone (PL 3, figs. 15, 18).

ELYTRA. This structure is comparable in most of the species, and yet,
provides the basis for subgeneric distinction. In nine of the ten species
each elytron has two antebasal foveae and a subhumeral foveae which is
connected to the margin by an entire, oblique epipleural sulcus. In curwsus,
both the subhumeral fovea and sulcus are lacking. Because of the absence
of these characters curiosus was designated as the type species of the
subgenus V estitrichus Park.

ABDOMEN. The ratios of the abdominal tergites are similar in those
species which are assumed to be closely related. Also, the ratio of the
first tergite is inversely proportional to the total size of the species under
consideration. Abdominal foveae are evident on the second sternite in all
species. However, in ornatus, cavatus and gibbosus the weak apodemal
invaginations at the lateral margins of both sternites and tergites which
are common to those species from the central coast of California have
become well developed, forming distinct pubescent foveae which are par-
tially covered by the apical margin of each segment and are not readily
visible on pointed specimens.

SECONDARY SEXUAL CHARACTERS. An accurate determination
of the sexes in the genus Pselaptrichus is necessary not only from the
practical value of the species determination, but is of considerable impor-
tance in the study of interspecific relationships. In previous classifications,
the sexes were readily segregable on the basis of two morphological
characters unique to the males; a preapical protibial notch or spinoid
process, an apical metatibial spine, or both. With the description of
several new species these two characters are insufficient to provide a
positive distinction. Consequently, a combination of three anatomical
landmarks as noted in the first couplet of the species key is necessary
to realize a correct diagnosis.

VENTRAL SURFACE OF HEAD. In the majority of species, the
ventral surface of the male head is structurally complex (PI. 2, figs. 10-
14). These structural modifications may take the form of a median
carinoid process or an excavation which is occupied by various protuber-
ances, transverse carinae, and hairs. The females of the above species lack
these gross structures but have modifications of the ventral apical declivity.
These modifications include a median excavation varying in size and form,
and the presence or absence of a median carina, callosity, or flat deltoid
impression. Both males and females of tuber culipalpus, rectus and spinosus

Marsh-Schuster — Genus Pselaptrichus 9

have a simple tumid surface without any obvious structural modifications
(PI. 2, fig. 9).

TIBIAE. The protibia of all males, with the exception of two species,
possess either a spinoid process or a notch varying in form and position
(PI. 3, figs. 19, 20). The two exceptions, rectus and carinatus, have simple
protibia which resemble the females in respect to this character (PI. 3,
fig. 21).

The presence of a distal metatibial spine in the males is considered to
be the most valid character differentiating the sexes, though not without
exception, since carinatus exhibits a complete loss of this structure.

ANTENNAE. The form of this structure is exceedingly similar in both
sexes of most species and offers little in the way of diagnostic characters.
The fourth through eighth segments are rounded hexagonal and subequal
in width. However, the seventh and eighth segments of cavatus and
gibbosus are dorso-ventrally compressed and longer than wide. This is
noticeable in lateral aspect, appearing as a short ventral arcuation inter-
rupting the segmental continuity.

SEX-LIMITED CHARACTERS. The eyes of the male population
samples of spinosus and rectus normally have seven facets which show
little intraspecific variation, and yet, two specimens of spinosus and one
of rectus have twenty-five facets. Correlated with this differential number
of facets is the development of presumably functional wings and associated
thoracic structures which are normally lacking. Since other diagnostic
characters such as the genitalia and protibia are identical, and since this
dimorphism has been observed only in male specimens, it is assumed <Lat
these are sex-limited characters.

Genus Pselaptrichus Brendel

Pselaptrichus Brendel, 1889, Entom. Amcr., 5: 194; Brendel and
Wickham, 1890, Bull. Lab. Nat. Hist. St. Univ. Iowa, p. 242;
Raffray, 1908, Genera Insectorum, fasc. 64, p. 285; Bowman, 1934,
Pselaphidae of North America, Pittsburgh, privately published, p.
117; Park, 1953, Chicago Acad. Sci., Bull., 9: 255-256.

ADULT. Head pyriform, narrower than prothorax, widest across eyes,
attenuate apically, forming frontal prolongation ; prolongation porrect at
apex, bearing large, approximate antennal acetabulae; disk with two
nude vertexal foveae between eyes connected to frontal sulcus by oblique
impressions; occiput with fine median carina of variable extent; clypeus
semicircular, weakly carinate medially; labrum small, transverse, lateral
angles rounded; mandibles expanded laterally at base, scrobes setate, inner
ramus toothed ; antennae eleven segmented ; first segment elongate,
cylindro-arcuate; second segment ovoid, subequal in width; third segment
small, obconical ; segments four through eight usually rounded-hexagonal,
subequal in width; nine through eleven forming weak club; eleven obconi-
cal, larger than preceding; maxillary palpi four segmented, elongate;
first segment small ; second segment elongate, sinuate-clavate, coarsely
tuberculate; third segment small, quadrate-tuberculate; fourth segment
large, securiform, microgranulate, densely pubescent, as long or longer
than second; eyes normally small, individual facets large; ventral surface
of herd simple or with various structural modifications. Pronotum sub-

10 San Diego Society of Natural History

cordiform, widest at apical one-third; disk with transverse, biarcuate
antebasal sulcus connecting pubescent lateral foveae; procoxae confluent;
mesocoxae approximate; metacoxae distant, inner proximal one-half of
profemur and trochanter tuberculate; protibia simple or modified; tarsi
three segmented ; first minute, second elongate, longer than third ; one
tarsal claw or claw and accessory spine. Elytra widest at apex, narrowed
to base; sutural striae evident; base of each elytron with two, pubescent,
antebasal foveae; subhumeral foveae usually present, connected to lateral
margin by oblique linear impression. Abdomen with five visible tergites
and six sternites; tergites one through three margined laterally.
TYPE. — Pselaptrichus tuberculipalpus Brendel (monotypic).
This genus was distinct among the North American Bythinini on the
basis of the fourth palpal segment which is relatively broad and about
twice as long as wide. However, one new species described herein has the
fourth segment three times longer than wide, thus encompassing a
character often used to distinguish the eastern genus Bythinopsis.

The genus was recently divided into two sections by Park (1953);
subgenus Pselaptrichus sensu Brendel (1889), containing the type species
tuberculipalpus Brendel and rot hi Park; subgenus Vestitrichus Park, con-
taining curiosus Park. The seven new species described in this paper be-
long to the former section.

KEY TO PSELAPTRICHUS
1. Ventral surface of head with obvious structural modifica-

tion; or protibia excavated on inner surface; or metatibia

spined apically male 2

-All of above characters absent female 12

2.(1) Ventral surface of head simple 3

-Ventral surface of head with obvious structural modification 5
3.(2) Protibia with small notch at apical one fifth (PI. 3, fig.
19); about 1.5 mm.; Alameda Co., Contra Costa Co.,

Calif tuberculipalpus

-Protibia simple or with prominent spinose projection 4

4.(3) Protibia simple; about 1.5 mm.; Marin Co., Calif rectus

-Protibia with prominent spinose projection (PI. 3, fig. 20) ;

about 1.5 mm.; Santa Cruz Co., Calif spbiosus

5.(2) Ventral surface of head with prominent median longitudinal

carinoid process 6

-Ventral surface of head otherwise modified 7

6.(5) Elytron with subhumeral fovea and epipleural sulcus; pro-
tibia simple (PI. 3, fig. 21) lacking apical metatibial spine;

1.3 mm.; Humboldt Co., Calif carinatus

-Without the above combination of characters 11

7.(5) Eyes large, subreniform, over 30 facets; protibial notch bi-
sected by two long setae (PI. 3, fig. 22) ; winged; 1.7 mm.;

Central Sierra Nevada oculatus

-Eyes small, less than 25 facets; protibial notch with at most

short setae 8

8.(7) Antennal segments III through VIII similar in form and

size 9

-Antennal segments VII and VIII conspicuously narrower in
lateral outline 10

Marsh-Schuster — Genus Pselaptrichus 11

9.(8) Large species; total length 2.3 mm.; Oregon rothi

-Small species; total length 1.5 mm.; Mendocino Co.,
Calif ornatus

10.(8) Tempora straight in dorsal outline (PI. 2, fig. 11); total

length 1.8 mm.; Humboldt Co., Calif cavatus

-Tempora angulate rounded in dorsal outline (PI. 2, fig.
14) ; total length 1.8 mm.; Humboldt Co., Calif gibbosus

11.(6) Elytron without subhumeral fovea or epipleural sulcus;
protibia excavated on inner surface; metatibia spined api-
cally; total length 1.6 mm.; Idaho curiosus

12.(1) Ventral surface of head shining, lacking small irregular
punctures bearing fine hairs; gena with 2 oblique rows of
setae (PI. 2, fig. 9); total length about 1.5 mm.

Alameda Co., Contra Costa Co., Calif tuberculipalpus

Marin Co., Calif rectus

Santa Cruz Co., Calif spinosus

-Ventral surface of head shining, with irregular punctures
bearing fine hairs 13

13.(12) Ventral surface of head subopaque, microgranulate, with
few small irregular punctures bearing fine hairs; lateral
borders of frontal prolongation longitudinally depressed,
bicarinate; total length 1.3 mm.; Humboldt Co., Calif.

carinatus
-Ventral surface of head punctate, bearing many fine hairs;
total length over 1.5 mm 14

14.(13) Ventral surface of head with apical declivity weakly im-
pressed with a small circular median pit; eyes of 12 or 13
facets; total length 1.7 mm:; Central Sierra Nevada.. oculatus
-Without the above characters 15

15.(14) Ventral surface of head with apical declivity strongly exca-
vate with a fine median carina; total length 2.3 mm.;

Oregon rothi

-Ventral surface of head with apical declivity not carinate;
total length less than 2.0 mm 16

16.(15) Ventral surface of head with apical declivity weakly exca-
vate; total length 1.5 mm.; Mendocino Co., Calif ornatus

-Ventral surface" of head with 2pical declivity otherwise;
larger species - 17

17.(16) Tempora straight in dorsal outline (PI. 2, fig. 11) ; ventral
surface of head with apical declivity strongly excavate; with
a small median longitudinal callosity; total length 1.8 mm.;

Humboldt Co., Calif cavatus

-Tempora angulate rounded in dorsal outline (PI. 2, fig.
14); ventral surface of head with apical declivity moder-
ately excavate, forming a flat, median, glabrous, triangular
impression; total length 1.8 mm.; Humboldt Co., Calif.

gibbosus

12 San Diego Society of Natural History

Pselaptrichus tuberculipalpus Brendel

(Figs. 1, 15, 19)

Pselaptrichus tuberculipalpus Brendel, 1889, Entom. Amer., 5: 194.

Male. Head 0.28±0.01 mm. long X 0.28±0.01 mm. wide; pronotum
0.33±0.02 X 0.35±0.02; elytra 0.48±0.02 X 0.55±0.04; total length
1.47±0.01 mm.1

Integument shining, subimpunctate, rufotestaceus; vestiture coarse,
moderately lon.<*, decumbent and sparse; transverse on head and prothorax,
inserted laterally, extending medially; longitudinally parallel on elytra;
shorter on venter. Head pyriform in dorsal outline, as wide as long; eyes
small, with either five or six facets, subcircular to deltoid in outline;
tempora evenly rounded to neck, moderately short, subequal to twice eye
width; median cervico-occipital carina present, not prominent nor attaining
posterior border of vertexal foveae; two, deep, nude vertexal foveae
located anterior to transverse line drawn between middle of eyes; distance
between foveae equal to that between fovea and eye; foveae connected
to narrow frontal sulcus by straight shallow impressions forming inverted
Y; dorsal surface of head not noticeably flattened, convex across cervico-
temporal region where median carina delimits two distinct lateral callosi-
ties; frons sharply declivous in lateral outline, forming weak median
carina extending to broad, semicircular clypeal margin; inner ramus of
mandibles with 7 teeth; labrum large, medially bidentate, dentition cor-
responding to two blunt, subapical projections; ventral surface of head
simple, without evident secondary sexual modifications; maxillary palpi
as illustrated; fourth segment securiform, two times longer than broad;
declivity of inner apical angle rounded angulate; antennae eleven seg-
mented; segment T cylindro-arcuate when viewed from above, as long as
segments three through eight combined; II one third as long, slightly
narrower, elongate oval; III through VIII subequal in width: III obconical,
slightly longer than four through eight which are equal in length; IV
through VIII subquadrate, rounded hexagonal; IX through XI forming
weak club; IX transverse; X transverse, slightly wider than ninth; XI
slightly longer than eighth, ninth, and tenth combined, wider than tenth,
base ovate, apex subcorneal. Pronotum as long as wide, slightly wider than
head; apical and basal margins subtruncate; apical margin subequal in
width to base; lateral margin rounded, widest at apical one third, subsin-
uate to lateral apical angle, straight to lateral basal angle; disk weakly
arcuate-convex in transverse section with two, oblique, pubescent foveae
near lateral basal angles connected by biarcuate antebasal sulcus; medial
portion of arcuation with small, obtuse, denticulation at inner edge.
Elytra wider than long, as wide as pronotum across humeri; subimpunctate;
disk somewhat flattened; each elytron bearing two, large, deep pubescent,
antebasal foveae separated by width of one fovea; medial fovea inserted
at base of entire parallel sutural stria; lateral fovea at base of shallow
longitudinal impression which extends one fourth of elytral length; sub-
humeral foveae moderately large, pubescent, connected by oblique line to
lateral apical margin forming entire epipleural sulcus; area between sub-
humeral and lateral foveae convex, forming subprominent humeral angle.

JThe measurements and their deviations were based on eighteen male
and twenty-six female specimens.

Marsh-Schuster — Genus Pselaptrichus 13

Abdomen slightly deflexed in lateral outline, with five visible tergites
and six sternites; ratio of median tergite lengths 1.6/1.2/1.0/1.0/1.0;
sixth sternite slightly concave medially. Prosternum convex anterior to
procoxae; protibiae arcuate in outline with notch at distal one third on
mesial face as illustrated; upper margin of notch with blunt tooth at inner
two thirds; pro and mesotibiae with small apical spur, metatibia with
inner apical spine. Genitalia complex as illustrated.

Female. Head 0.28±0.02 mm. long X 0.26±0.02 mm. wide; prono-
tum 0.35±0.03 X 0.36±0.02; elytra 0.51±0.045 X 0.59±0.04; total
length 1.54±0.11 mm.

As in the male but differing in the following respects: eyes vestigial,
of five facets, three setae on each gena parallel, not obliquely converging;
prothoracic tibia simple, without inner distal notch; metathoracic tibia
lacking distal spine but with small movable spur.

Type locality. — Alameda County, California.

Type deposition. — Philadelphia Academy of Natural Sciences.

Recorded distribution. — Alameda County, California (Brendel, 1889).

New records. — California. — ALAMEDA CO.: 2tf, no date (Fuchs,
CAS); Oakland. 4 J1, 5 9, 1-28-53 R. O. Schuster (CIS); 1 d\ 1$,
1-30-53 R. O. Schuster (CIS); 4d\ 3$, II-5-53 R. O. Schuster (CIS);
lr?, 1 9, H-8-53 R. O. Schuster (CIS); 2J\ 11-12-53 R. O. Schuster
(CIS); 3r?, 5 9. H-13-53 W. C. Bentinck (CIS); 39, X-17-53 R. O.
Schuster (CIS); CONTRA COSTA CO.; Ml. Diablo. 2J, 6 9 , 11-15-53
G. A. Marsh (CIS); 4^,49, 11-23-53 R. O. Schuster (CIS); Redwood
Park, 2c? , 3 9, V-18-53 E. E. Gilbert, R. O. Schuster (CIS); 1 d\ V-
28-53 E. E. Gilbert (CIS).

The male of tuberculipalpus differs from all the other species in the
genus by its simple, tumid, ventral surface of the head, a pre-apical notch
on the inner surface of the protibia, and its small size. The female is
indistinguishable from the two most closely related species, spinosus and
rectus, which occupy areas immediately to the south and north of tuber-
culipalpus (see map).

Pselaptrichus rectus new species
(Fig- 2)

Male. Head 0.29 mm. long X 0.26 mm. wide; pronotum 0.32 X 0.34;
elytra 0.49 X 0.54; total length 1.56 mm.

Integument shining, subimpunctate, rufotestaceus; vestiture coarse,
moderately long, decumbent and sparse; transverse on head and prothorax,
inserted laterally, extending medially; longitudinally parallel on elytra;
shorter on venter. Head pyriform in dorsal outline, longer than wide;
eyes small, with six facets, subcircular in outline; tempora evenly rounded
in outline, moderately short, subequal to twice eye width; median cervico-
occipital carina present, not prominent nor attaining posterior border of
vertexal foveae; two, deep, nude vertexal foveae located anterior to
transverse line drawn between middle of eyes; distance between foveae
equal to that between fovea and eye; foveae connected to narrow frontal
sulcus by straight shallow impressions forming inverted Y; dorsal surface
of head not noticeably flattened, convex across cervico-temporal region
where median carina delimits two distinct lateral callosities; frons sharply
declivous in lateral outline, forming weak median carina extending to
broad, semicircular clypeal margin; inner ramus of mandibles with 6

14 San Diego Society of Natural History

teeth; labrum large, medially bidentate, dentition corresponding to two
blunt, subapical projections; ventral surface of head simple, without evi-
dent secondary sexual modifications; maxillary palpi as in tuberculipalpus;
fourth segment securiform, two times longer than broad; declivity of
inner apical angle rounded angulate; antennae eleven segmented; seg-
ment I cylindro-arcuate when viewed from above, as long as segments
three through eight combined; II one third as long, slightly narrower,
elongate oval; III through VIII subequal in width; III obconical, slightly
longer than four through eight which are equal in length; IV through

VIII subquadrate, rounded hexagonal; IX through XI forming weak club;

IX transverse; X transverse, slightly wider than ninth; XI slightly longer
than eighth, ninth, and tenth combined, wider than tenth, base ovate, apex
subconical. Pronotum wider than long, slightly wider than head; apical
and basal margins subtruncate; apical margin subequal in width to base;
lateral margin rounded, widest at apical one third, subsinuate to lateral
apical angle; disk weakly arcuate-convex in transverse section with two,
oblique, pubescent foveae near later? ' basal angles connected by biarcuate
antebasal sulcus; medial portion of arcuation with small, obtuse, denticu-
lation at inner edge. Elytra wider than long, as wide as pronotum across
humeri; subimpunctate; disk somewhat flattened; each elytron bearing two,
large, deep, pubescent, antebasal foveae separated by width of one fovea;
medial fovea inserted at base of entire parallel sutural stria; lateral fovea
at base of shallow longitudinal impression which extends one fourth of
elytral length; subhumeral foveae moderately large, pubescent, connected
by oblique line to lateral apical margin forming entire epipleural sulcus;
area between subhumeral and lateral foveae convex, forming subprominent
humeral angle. Abdomen slightly deflexed in lateral outline, with five
visible tergites and six sternites; ratio of median tergite lengths 1.6/1.2/
1.0/1.2/1.0; sixth sternite slightly concave medially. Prosternum convex
anterior to procoxae; protibia nearly straight in outline, without a notch or
spinoid process at distal one third on mesial face, simple; metatibia with
inner apical spine. Genitalia complex as illustrated diagrammatically.

Female. Head 0.26 mm. long X 0.24 mm. wide; pronotum 0.31 X
0.33; elytra 0.46 X 0.53; total len^h 1.47 mm.

As in the male but differing in the following respects: eyes vestigial,
of four facets; protibia simple as in the male; metathoracic tibia lacking
distal spine.

Type materials. Holotype male and allotype female deposited in the
entomological collection of the California Academy of Sciences, taken at
Alpine Lake, Marin County, California, June 18, 1953 (C. D. MacNeill,
R. O. Schuster). Two additional paratypes were collected at the same
locality and date and are deposited in the collection of the California
Insect Survey, University of California. Two specimens not designated as
paratvpic are from Mill Valley, Marin County, California, June 14, 1952
(H. B. Leech).

P. rectus is the second of three closely related species in which the
females prove to be indistinguishable. The males however, unlike tuber-
culipalpus and spinosus, have the protibia simple, nearly straight, and
without a trace of either a notch or a spinoid process. This, plus the lack
of any structural modifications of the ventral surface of the head, will
distinguish this species from any other in the genus. The protibia of

Marsh-Schuster — Genus Pselaptrichus 15

carinatus from Humboldt County, California, also has a simple protibia,
but rectus has a metatibial spine which is lacking in carinatus.

Pselaptrichus spinosus new species
(Figs. 3, 9, 20)

Pselaptrichus tuber culipalpus, Park, 1953, Bull. Chicago Acad. Sci.,
9: 261 (nee Brendel, 1889)

Male. Head 0.31 mm. long X 0.28 mm. wide; pronotum 0.33 X 0.33;
elytra 0.51 X 0.58; total length 1.48 mm.

Integument shining, subimpunctate, rufotestaceus; vestiture coarse,
moderately long, decumbent and sparse; transverse on head and prothorax,
inserted laterally, extending medially; longitudinally parallel on elytra;
shorter on venter. Head pyriform in dorsal outline, longer than wide;
eyes small, with either seven or eight facets, subcircular in outline; tem-
pora evenly rounded to neck, moderately short, subequal to twice eye
width; median cervico-occipital carina present, not prominent nor attain-
ing posterior border of vertexal foveae; two, deep, nude vertexal foveae
located anterior to transverse line drawn between middle of eyes; distance
between foveae equal to that between fovea and eye; foveae connected to
narrow frontal sulcus by straight shallow impressions forming inverted Y;
dorsal surface of head not noticeably flattened, convex across cervico-
temporal region where median carina delimits two distinct lateral
callosities; frons sharply declivous in lateral outline, forming weak median
carina extending to broad, subtruncate clypeal margin; inner ramus of
mandibles with five blunt teeth; labrum large, medially bidentate, dentition
corresponding to two blunt, subapical projections; ventral surface of head
as illustrated, without evident secondary sexual modifications; area around
tentorial pits somewhat coriaceous; maxillary palpi with fourth segment
securiform, two times longer than broad; declivity of inner apical angle
rounded angulate; antennae eleven segmented; segment I cylindro-arcuate
when viewed from above, as long as segments three through eight com-
bined; II one third as long, slightly narrower, elongate oval; III through
VIII subequal in width; III obconical, slightly longer than four through
eight which are equal in length; IV through VIII subquadrate, rounded
hexagonal; IX through XI forming weak club; IX transverse, slightly
wider than eighth; X transverse, slightly wider than ninth; XI slightly
longer than eighth, ninth, and tenth combined, wider than tenth, base
ovate, apex subconical. Pronotum as long as wide, slightly wider than
head; apical and basal margins subtruncate; apical margin subequal in
width to base; lateral margin rounded, widest at apical one third, sub-
sinuate to lateral apical angle, straight to lateral basal angle; disk weakly
arcuate-convex in transverse section with two, oblique, pubescent foveae
near lateral basal angles connected by biarcuate antebasal sulcus; medial
portion of arcuation with small, obtuse denticulation at inner edge. Elytra
wider than long, as wide as pronotum across humeri; subimpunctate; disk
somewhat flattened; each elytron bearing two, large, deep, pubescent, ante-
basal foveae separated by width of one fovea; medial fovea inserted at
base of entire parallel sutural stria; lateral fovea at base of shallow longi-
tudinal impression which extends one fourth of elytral length; subhumeral
foveae moderately large, pubescent, connected by oblique line to lateral
apical margin forming entire epipleural sulcus; area between subhumeral

16 San Diego Society of Natural History

and lateral foveae convex, forming subprominent humeral angles.
Abdomen slightly deflexed in lateral outline, with five visible tergites and
six sternites; ratio of median tergite lengths 1.8/1.3/1.0/1.3/1.0; sixth
sternite slightly concave medially. Presternum convex anterior to procoxae;
protibia strongly arcuate in outline with large spinoid process on inner
face as illustrated; metatibia with inner apical spine. Genitalia complex
as illustrated diagrammatically.

Female. Head 0.29 mm. long X 0.25 mm. wide; pronotum 0.34 X
0.34; eyltra 0.53 X 0.61; total length 1.4 mm.

As in the male but differing in the following respects: eyes vestigial,
of six facets; prothoracic tibia simple, without inner spinoid process;
metathoracic tibia lacking distal spine.

Type material. Holotype male and allotype female deposited in the
entomological collection of the California Academy of Sciences, taken in
redwood litter at Ben Lomond, Santa Cruz County, California, July 5,
1953 (C. D. MacNeill). Thirteen additional paratypes were collected at
the same locality and date; twenty-one paratypes at the same locality in
both redwood and Douglas Fir litter, June 21, 1953 (C. D. MacNeill);
three paratypes, Santa Cruz County. California, July, 1906 (F. E. Blais-
dell); six paratypes, Santa Cruz Mountains, California (Koebele); two
paratypes, Santa Cruz County, California (Nunenmacher) ; ten paratypes
deposited in the entomological collection of the California Academy of
Sciences; thirty-five paratypes in the collection of the California Insect
Survey, University of California; two paratypes in the Park collection,
Northwestern University. Two specimens not designated as paratypic are
from Ben Lomond, Santa Cruz County, California, June 21, 1953 (C. D.
MacNeill).

Park, 1953, in describing rot hi and cur'wsus, utilized specimens from
Santa Cruz County, California, as typical of tiiberculipalpus for the key
and species comparisons. Upon examining specimens from the type locality
as well as the above mentioned locality, a discrepancy in the configuration
of the male protibia was noted. It was found, after a critical examination
of the genitalia and a comparison of materials with the type through the
kindness of Dr. James A. G. Rehn, that tiiberculipalpus of Park repre-
sents a new and distinct species herein described as spinosus.

This interesting species differs at once from tiiberculipalpus by having
a well developed spinoid process on the protibia of the males rather than
a simple notch. Also, the lateral lobes of the male genital armature have
three distinct setae while tiiberculipalpus has five. At present, the females
of these two species are morphologically inseparable.

Occasionally, single male specimens may occur with large eyes and
fully developed wings as previously mentioned in the discussion on sex-
limited characters. Until further evidence is obtained concerning such
specimens, they are to be considered as conspecific with spinosus.

Pselaptrichus carinatus new species
(Figs. 4, 10, 21)

Male. Head 0.25 mm. long X 0.27 mm. wide; pronotum 0.30 X 0.36;
elytra 0.47 X 0.52; total length 1.33 mm.

Integument shining, subimpunctate, testaceus; vestiture long, very fine,
reflexed, of equal density on entire dorsum; transverse on head and prono-

Marsh-Schuster — Genus Pselaptrichus 17

turn, inserted laterally, extending medially; not parallel on elytra but
converging toward median suture; shorter on venter. Head pyriform in
dorsal outline, wider than long; eyes small, of either six or seven facets,
subcircular in outline; tempora evenly rounded to neck, moderately short,
equal to twice eye width; median cervico-occipital carina present, sub-
prominent, just attaining posterior border of vertexal foveae; two, shallow,
nude, vertexal foveae located slightly anterior to transverse line drawn
through middle of eyes; distance between foveae equal to that between
fovea and. eye; foveae connected to narrow frontal sulcus by straight, some-
what deep impressions forming inverted Y; oblique impressions partially
interrupted at apex by flat ridge giving bisulcate appearance to median
sulcus; dorsal surface of head noticeably flattened, slightly convex across
cervico-temporal region; frons sharply declivous in lateral outline, forming
weak median carina extending to narrow elliptical clypeal margin; lab-
rum large, subtruncate, weakly bidentate medially; inner ramus of
mandibles with seven teeth; ventral surface of head complex as illustrated,
with an obliquely truncate, medial carinoid process bisetose at both apices;
maxillary palpi with fourth segment securiform, two times longer than
broad; declivity of inner apical angle obliquely subtruncate to palpal
cone; antennae eleven segmented; segment I cylindro-arcuate when viewed
from above, as long as segments three through eight combined; II one
third as long, slightly narrower, rounded rectangular; III through VIII
subequal in width; III obconical, slightly longer than four; IV through
VIII equal in length, subquadrate, rounded hexagonal; IX through XI
forming weak club; IX transverse, X transverse, slightly wider than
ninth; XI subconical, wider than tenth, slightly longer than seventh
through tenth combined. Pronotum wider than long, slightly wider than
head; apical and basal margins subtruncate; apical margin narrower than
base; lateral margin widest at apical one third, evenly rounded, subsinuate
to lateral basal angle; disk weakly arcuate-convex in transverse section
with two, oblique, pubescent foveae near lateral basal angles; foveae con-
nected by biarcuate antebasal sulcus; medial portion of arcuation with
small, obtuse denticulation at inner edge. Elytra wider than long, as wide
as pronotum across humeri; subimpunctate; each elytron bearing two, large,
deep, pubescent antebasal foveae separated by width of one fovea; median
foveae inserted at base of entire parallel, sutural striae which rapidly
converge at apical one fifth; lateral fovea at base of shallow longitudinal
impression; subhumeral fovea large, pubescent, connected to lateral apical
margin by line forming entire epipleural sulcus; area between subhumeral
and lateral foveae evenly rounded, not noticeably convex, forming humeral
angles. Abdomen with five visible tergites and six sternites; tergites
broadly convex, reflexed in lateral outline; ratio of median tergite
lengths 1.6/1.2/1.0/1.0/1.0; anterior one half of prosternum convex,
basal one half flattened; protibia straight, simple as illustrated, without
distal notch on inner surface; metatibia subarcuate, lacking inner distal
spine. Genitalia complex as illustrated.

Female. Head 0.25 mm. long X 0.27 mm. wide; pronotum 0.29 X
0.35; elytra 0.45 X 0.53; total length 1.29 mm.

As in the male but differing in the following respects: eyes vestigial,
of four or five facets, semicircular in outline; ventral surface of head

18 San Diego Society of Natural History

simple, without a median carinoid process, with few scattered punctures
bearing fine hairs; sixth sternite evenly convex.

Type material. Holotype male and allotype female deposited in the
entomological collection of the California Academy of Sciences, taken in
redwood litter eighteen miles south of Klamath, Del Norte County,
California, September 19, 1953 (E. E. Gilbert, R. O. Schuster). Ninety-
six additional paratypes were collected at the same locality and date;
twenty-two paratypes at the same locality, June 26, 1953 (G. A. Marsh,
R. O. Schuster); one paratype at the same locality, June 26, 1953 (J. D.
Lattin); thirty paratypes desposited in the California Academy of Sci-
ences; eighty-eight paratypes in the California Insect Survey collection,
University of California.

This species differs from all the other species in the genus by having a
complete lack of the apical metatibial spine, a narrow canaliculate frontal
sulcus in which the lateral margins of the frontal prolongation are bicar-
inate, and its very small size.

The median carinoid process of the ventral surface of the head is
highly suggestive of curiosus from which it may be readily separated by
having a well developed subhumeral fovea and no evidence of a protibial
notch.

The female is most readily comparable to tuberculipalpus and its con-
geners but can be distinguished on the basis of the above mentioned
configuration of the frontal prolongation.

Thus far, this species is known from only the type locality. Since
numerous samples were taken to the south, it would suggest that the
range of carinatus extends north to southern Oregon providing this species
dispersal corresponds to others found in northern California.

Pselaptrichus oculatus new species
(Figs. 5, 12, 22)

Male. Head 0.33 mm. long X 0.34 mm. wide; pronotum 0.40 X 0.40;
elytra 0.67 X 0.72; total length 1.74 mm.

Integument shining, subimpunctate, rufotestaceus; vestiture fine, long,
suberect, somewhat sparse; transverse on head and pronotum, inserted
laterally, extending medially; longitudinally parallel on elytra; shorter on
venter. Head pyriform in dorsal outline, slightly wider than long, nar-
rowed apically to form wide, short, subtruncate frontal prolongation
bearing medial sulcus; eyes large, of thirty-three to thirty-five facets, sub-
reniform in outline; tempora rounded angulate, rapidly descending to
neck, equal to eye width; median cervico-occipital carina present, promi-
nent, attaining or slightly surpassing posterior border of vertexal foveae;
two, deep, nude vertexal foveae located on transverse line drawn between
middle of eyes; distance between foveae subequal to that between fovea
and eye; foveae connected to wide frontal sulcus by weakly arcuate, shal-
low impressions forming inverted Y; medial sulcus flat, shining, glabrous,
base wider than apex; lateral convex margins flattened, rugose; dorsal
surface of head flattened though moderately convex between foveae and
across cervico-temporal region where median carina delimits two weak
lateral callosities; frons sharply declivious in lateral outline, forming weak
median carina extending to wide, elliptical clypeal margin; labrum large,
subtruncate, weakly bidentate medially; ventral surface of head complex as

Marsh-Schuster — Genus Pselaptrichus 19

illustrated; fourth segment of maxillary palpi securiform, two times longer
than broad; declivity of inner apical angle rounded angulate; antennae
eleven segmented; segment I cylindro-arcuate when viewed from above,
subequal to segments three through eight combined; II one half as long,
slightly narrower, oval; III through VIII subequal in width; III obconical,
slightly longer than following which are equal in length; IV through VIII
subquadrate, rounded hexagonal; IX through XI forming weak club; IX
transverse; X transverse, slightly wider than ninth; XI wider than tenth,
base truncate, apex subconical. Pronotum as long as wide, slightly wider
than head; apical and basal margins subtruncate; lateral margin evenly
rounded, widest at apical one third; disk evenly convex in transverse
section, with two oblique, pubescent foveae at lateral basal angles; foveae
connected by biarcuate antebasal sulcus; medial portion of arcuation com-
pletely interrupted by short, longitudinal carina. Elytra wider than long,
as wide as pronotum across humeri; punctulate at base; each elytron bear-
ing two, large, deep, pubescent antebasal foveae; median fovea originating
at base of entire sutural stria; lateral fovea at base of shallow longitudinal
impression; subhumeral fovea moderately large, pubescent, connected by
oblique line to form entire epipleural sulcus; area between lateral and
subhumeral foveae evenly convex, forming well pronounced humeral
angles; sutural striae narrow, slightly converging at base and apex.
Abdomen with five visible tergites and six sternites; broadly convex dor-
sally; lateral margin strongly reflexed toward apex; ratio of median
tergite lengths 1.3/1.0/1.0/1.57/1.7; sixth sternite transversely concave.
Prosternum weakly concave before coxae; protibia weakly arcuate in out-
line, notched internally at apical one fifth as illustrated; notch rounded,
upper lateral angle forming weak spine, upper margin bisetose medially;
metatibia with prominent spine. Genitalia complex as illustrated.

Female. Head 0.33 mm. long X 0.28 mm. wide; pronotum 0.39 X
0.40; elytra 0.58 X 0.66; total length 1.66 mm.

As in the male but differing in the following respects: eyes of nine
facets, oblong in outline; ventral surface of head simple, tumid, apical
declivity with small median pit; wingless; protibia without subapical
notch; metatibia lacking inner apical spine.

Type materials. Holotype male, allotype female, and one paratype
deposited in the entomological collection of the California Academy of
Sciences. Holotype from Wawona, Mariposa County, California, June
(A. Fenyes); allotype from Sugar Pine, Fresno County, California (A.
Fenves). One dissected male paratype from Wawona deposited in the
California Insect Survey collection, University of California.

This species differs from typical specimens of all the other species in
the genus by having large, subreniform eyes composed of over thirty
facets, wings, and the characteristic ventral surface of the head.

The female exhibits a similarity of gross structure to rothi. but can be
distinguished on the basis of the medial sulcus of the frontal prolongation
which is considerably narrower than in rothi. Also, the apical declivity
of the ventral surface of the head has a median pit rather than an exca-
vation with a fine median carina as in rothi.

Thus far, this is the only species found in the Sierra Nevada range.
Since those species which exhibit morphological similarities are from
Oregon and northern coastal California, it would seem possible that

20 San Diego Society of Natural History

several other species will be found extending north into the southern
Cascades.

Pselaptrichus rothi Park

Pselaptrichus rothi Park, 1953, Chicago Acad. Sci., Bull. 9: 256-257,
(PI. IB; 2AF; 3C).

Male. Sulcus of frontal prolongation wider than long; eyes of fourteen
facets, dorso-ventrally triangular in outline; tempora rounded angulate;
antennal segments III through VIII similar in size and form; ventral
surface of head with evident structural modification; protibia distally
excavate at inner distal surface; metatibia spined distally; total length
2.3 mm.

Female. Eyes subcircular, of eight facets; ventral surface of head simply
tumid; apical declivity excavate, bearing a fine median carina; protibia
simple; metatibia not distally spined.

Type locality. McDonald Forest, eight miles north of Corvallis,
Oregon.

Type deposition. California Academy of Sciences.

Recorded distribution. OREGON: McDonald Forest, eight miles north
of Corvallis; Cascadia (Park, 1953).

At present, rothi is the only species found in Oregon. Its closest affi-
nities are with gibbosus and ornatiis from northern California. Of par-
ticular interest is the similarity in the curious form of the eyes which are
dorsoventrally triangular. The male of rothi may be distinguished by
its larger size, the median sulcus of the head which is broader than long,
and the tuft of hairs inserted at the apex of the tuberosity on the ventral
surface of the head.

The females can be recognized by a longitudinally carinate excavation
of the apical declivity on the ventral surface of the head.

Pselaptrichus ornatus new species
(Figs. 6, 13, 18, 23)

Male. Head 0.27 mm. long X 0.31 mm. wide; pronotum 0.36 X 0.38;
elytra 0.55 X 0.58; total length 1.44 mm.

Integument shining, subimpunctate, rufotestaceus; vestiture fine, long,
reflexed; transverse on head and pronotum, parallel on sides of elytra,
converging medially; shorter on venter. Head pyriform in dorsal outline,
longer than wide; frontal prolongation short, broad, subtruncate apically;
eyes with seventeen facets, comma shaped; tempora evenly rounded to
neck, one and one half times eye width; median cervico-occipital carina
prominent, slightly surpassing posterior border of vertexal foveae; two,
small, shallow, nude vertexal foveae located slightly anterior to transverse
line drawn between middle of eyes; foveae connected to median sulcus of
frontal prolongation by two moderately deep, oblique impressions forming
inverted Y; dorsal surface of head flattened anteriorly, rounded convex
across cervico-temporal region; sulcus of frontal prolongation wide, flat,
expanded apically, wider than lateral convex margins which are glabrous
basally, rugosely carinate apically; frons sharply declivous in lateral out-
line, forming median carina extending to narrowly elliptical clypeal
margin; labrum large, subtruncate, lateral apical margins rounded, sides
strongly convergent to base; ventral surface of head complex as illus-
trated; fourth segment of maxillary palpi two times longer than broad;

Marsh-Schuster — Genus Pselaptrichus 21

declivity of inner basal and apical angles evenly rounded as illustrated;
antennae eleven segmented; segment I cylindro-arcuate when viewed from
above, as long as segments three through six combined; II one third as
long, slightly narrower, obovate; III through VIII subequal in width;
III obcordate, slightly longer than four through eight which are equal in
length; IV through VIII subquadrate, rounded hexagonal; IX through XI
forming weak club; IX transverse; X transverse, slightly wider than
ninth; XI slightly longer than eighth, ninth, and tenth combined, sub-
conical. Pronotum slightly wider than long, noticeably wider than head,
widest at apical one-third; apical and basal margins subtruncate; lateral
margins evenly rounded to apical angle, weakly arcuate to base; disk
evenly convex in transverse section, with two, oblique, pubescent foveae
near lateral basal angles; foveae connected by biarcuate antebasal sulcus;
median basal carina extending to, but not completely interrupting ante-
basal sulcus. Elytra subimpunctate, wider than long, slightly wider than
prothorax across humeri; each elytron with two, large, deep, pubescent,
foveae separated by width of one fovea; median fovea originating at base
of narrow, entire, sutural stria; lateral fovea at inner basal angle of humeri;
subhumeral fovea large, pubescent, connected by oblique line to form
entire epipleural sulcus. Abdomen moderately deflexed in lateral outline
with live visible tergites and six sternites; ratio of median tergite lengths
1.6/1.3/1.0/1.4/1.4. Protibia arcuate with notch at inner distal one
third as illustrated; metatibia with inner apical spine. Genitalia complex
as illustrated.

Female. Head 0.27 mm. long X 0.28 mm. wide; pronotum 0.37 X
0.40; elytra 0.56 X 0.61; total length 1.5 mm.

As in the male but differing in the following respects: eyes of seven
facets; ventral surface of head simple, tumid, apical declivity weakly
excavate; protibia without notch; metatibia lacking inner apical spine.

Type materials. Holotype male, allotype female, and two paratypes
deposited in the entomological collection of the California Academy of
Sciences, taken in redwood litter at Hartsook Grove, Mendocino County,
California, September 19, 1953 (E. E. Gilbert, R. O. Schuster). Five
additional paratypes collected at the same locality and date are deposited
in the California Insect Survey collection, University of California. Three
specimens not designated as paratypic are from one mile south of Dyer-
ville, Humboldt County, California, August 13, 1953 (G. A. Marsh,
R. O. Schuster) and September 19, 1953 (E. E. Gilbert, R. O. Schuster).

The males of this species are closely related to roth/ and gibbosus but
are easily distinguished from the former by its smaller size and from the
latter by the antennae, segments three through eight being subequal in
width in ornatus.

In the females, the apical declivity of the ventral surface of the head
is but weakly excavate and lacks the triangular impression of gibbosus
and the median carina of rothi.

Pselaptrichus cavatus new species
(Figs. 7, 11, 17, 24)
Male. Head 0.40 mm. long X 0.33 mm. wide; pronotum 0.44 X 0.45;
elytra 0.69 X 0.68; total length 1.78 mm.

Integument shining, subimpunctate, rufotestaceus; vestiture fine, short,

22 San Diego Society of Natural History

decumbent, sparse, tending to become deciduous; transverse on head and
pronotum; elytral pubescence longitudinally parallel; shorter on venter.
Head elongate pyriform in dorsal outline, longer than wide; with con-
spicuous narrow, subtruncate frontal prolongation; eyes small, of ten
facets, dorsoventrally conical in outline; tempora straight to neck, long,
equal to three times eye width; median cervico-occipital carina subpromi-
nent, surpassing posterior border of vertexal foveae; two, shallow, nude,
vertexal foveae located posterior to transverse line drawn through middle
of eyes; distance between foveae equal to that between fovea and eye;
oblique lateral sulcal connections to broad shallow median sulcus of
frontal prolongation partially interrupted by converging basal lateral angles
of prolongation anterior to fovea, forming weak inverted Y; dorsal surface
of head noticeably flattened except for weak convexity between vertexal
foveae; margin of foveae not continuous, anterior one half open; two,
minute, shallow, subcircular impressions obliquely posterior to ventral
margin of eyes; each bearing a long seta; frons sharply declivous in
lateral outline, forming a median carina extending to narrowly elliptical
clypeal margin; labrum large, subtruncate, lateral margins strongly sinuate
to base; apical angles evenly rounded; inner ramus of mandibles with
eight teeth; ventral surface of head complex as illustrated; maxillary palpi
as illustrated; fourth segment three times longer than broad; declivity of
inner apical angles obsolete, gradually attenuate to palpal cone; antennae
eleven segmented; segment I cylindro-arcuate when viewed dorsally, as
long as segments three through nine combined; II oval, one fourth as long,
subequal in width; III through VI subequal in width; III obconical,
slightly longer than four; VII dorso-ventrally compressed, longer than
wide, rounded rectangular; VIII similar to four through six; IX through
XI forming weak club; IX transverse; X transverse, slightly wider than
ninth; XI subequal to nine and ten, subconical. Pronotum as long as
wide, slightly wider than head, widest at apical one third; evenly rounded
to lateral apical angle, nearly straight to base; apical and basal margins
subtruncate; disk weakly arcuate convex in transverse section with two,
oblique, pubescent foveae near lateral basal angles; foveae connected by
biarcuate antebasal sulcus; median portion of arcuation with small, obtuse,
medial denticulation at inner edge completely interrupting antebasal
sulcus. Elytra subimpunctate; wider than long, as wide as pronotum across
humeri; each elytron bearing two, large, deep, pubescent antebasal foveae
separated by width of one fovea; median fovea inserted at base of entire
sutural stria; shallow impression at base of lateral fovea obsolete, not
extending apically; subhumeral fovea large, pubescent, connected by
oblique line to lateral margin forming entire epipleural sulcus. Abdomen
strongly deflexed in lateral outline; with five visible tergites and six
sternites; ratio of median tergite lengths 1.3/1.14/1.0/1.57/1.4; prester-
num flattened anterior to coxae, broadly emarginate along apical margin:
protibia slightly arcuate to distal one fifth, with a shallow U shaped
notch on inner surface; metatibia with large prominent, apical spine.
Genitalia complex as illustrated.

Female. Head 0.39 mm. long X 0.33 mm. wide; pronotum 0.44 X
0.45; elytra 0.65 X 0.70; total length 1.78 mm.

As in the male but differing in the following respects: eyes of five
facets, subcircular in outline; seventh antennal segment not elongate,

Marsh-Schuster — Genus Pselaptrichus 23

identical with sixth; ventral surface of head simple, apical declivity
excavate with median longitudinal callosity; protibia simple, without notch
on inner apical surface; metatibia without apical spine.

Type materials. Holotype male and allotype female deposited in the
entomological collection of the California Academy of Sciences, taken in
redwood litter at Freshwater, Humboldt County, California, August 13,
1953 (G. A. Marsh, R. O. Schuster). Four paratypes were collected at
the same locality and date; ten paratypes in redwood litter eighteen miles
south of Klamath, Del Norte County, California, September 19, 1953
(E. E. Gilbert, R. O. Schuster) ; twelve paratypes in redwood litter one
and one half miles south of Dyerville, Humboldt County, California,
August 13, 1953 (G. A. Marsh, R. O. Schuster) and September 19, 1953
(E. E. Gilbert, R. O. Schuster); one paratype at Green Point Ranch,
Humboldt County, California, June 11, 1919 (F. E. Blaisdell). Seven
paratypes deposited in the collection of the California Academy of
Sciences; twenty-two in the California Insect Survey collection, University
of California.

Although gibbosiis is of comparable size, cavatus can be readily recog-
nized from this or any other species in the genus by the fourth segment
of the maxillary palpus which is fully three times as long as wide, or by
the tempora which are nearly straight to the cervex. At present, this
species cannot be compared with any other in the genus.

Pselaptrichus gibbosus new species
(Figs. 8, 14, 25)

Male. Head 0.38 mm. long X 0.33 mm. wide; pronotum 0.42 X
0.43; elytra 0.61 X 0.65; total length 1.67 mm.

Integument shining, subimpunctate, rufotestaceus; vestiture long, fine,
dense, sparser on head and prothorax and transverse, inserted . laterally,
extending medially, decumbent; elytral pubescence longitudinally parallel
at sides, converging medially toward sutural striae; shorter on venter.
Head pyriform, slightly longer than wide in dorsal outline; frontal
prolongation one half length of head, subtruncate apically; eyes small, not
vestigial, with twelve facets, dorsoventrally elongate triangular; tempora
rounded angulate to neck, equal to two times eye width; median cervico-
occipital carina weak, not attaining line drawn between posterior border
of vertexal foveae; two, deep, nude, vertexal foveae located on a line
drawn through middle of eyes; foveae connected to median sulcus of
frontal prolongation by shallow, oblique impressions forming inverted Y;
dorsal surface of head noticeably flattened except for weakly convex area
across cervico-temporal region; medial sulcus shallow, narrow, subequal
to width of either lateral areas which are flattened and rugosely im-
pressed; frons sharply declivous in lateral outline, forming median carina
extending to moderately elliptical clypeal margin; labrum large, apex sub-
truncate, weakly bidentate medially; ventral surface of head complex as
illustrated; fourth segment of maxillary palpi two times longer than wide;
declivity of inner apical angle rounded angulate; antennae eleven seg-
mented; segment I cylindro-arcuate when viewed dorsally, as long as
segments three through eight combined; II obovate, one third as long,
subequal in width; III obcordate, slightly longer than four; IV and V
rounded hexagonal, subequal in width; VI elongate, longer than pre-

24 San Diego Society of Natural History

ceding; VII subequal to fourth and fifth, elongate rectangular; VIII
rounded hexagonal; IX through XI forming weak club; IX transverse;
X transverse, wider than ninth; XI subconical, equal to ninth and tenth
combined. Pronotum as long as wide, widest at apical one third; sinuate
to lateral apical angle, nearly straight to base; apical and basal margins
subtruncate; disk weakly arcuate-convex in transverse section, with two,
oblique, pubescent foveae near lateral basal angles; foveae connected by
biarcuate antebasal sulcus with medial denticulation at inner edge com-
pletely interrupting sulcus. Elytra subimpunctate; as long as wide, as wide
as prothorax across humeri; each elytron bearing two moderately large,
deep, pubescent, antebasal foveae separated by width of one fovea; median
fovea originating at base of entire sutural stria; shallow impression at base
of lateral fovea obscure; subhumeral fovea moderately large, deep, con-
nected by oblique line to lateral margin, forming entire epipleural sulcus.
Abdomen strongly deflexed in lateral outline with five visible tergites
and six sternites; ratio of median tergite lengths 1.3/1.4/1.0/1.14/1.2;
sixth sternite weakly concave; prosternum weakly concave; protibia nearly
straight, bearing sinuate notch at inner one fifth as illustrated; metatibia
with a small vestigial spine. Genitalia complex as illustrated.

Female. Head 0.36 mm. long X 0.30 mm. wide; pronotum 0.40 X
0.45; elytra 0.63 X 0.66; total length 1.83 mm.

As in the male but differing in the following respects: ventral surface
of head simple, microgranulate and hirsute, slightly tumid apically; apical
declivity with glabrous triangular impression; antennae lacking segmental
modification; eyes small, of six facets; sixth sternite flattened; protibia
simple, without notch at inner distal one fifth; metatibial spine absent.

Type material. Holotype male, allotype female, and five paratypes
deposited in the entomological collection of the California Academy of
Sciences, taken in Umbellular'ia califomica litter fourteen miles east of
Blue Lake, Humboldt County, California, September 19, 1953 (E. E.
Gilbert, R. O. Schuster). Ten additional paratypes were collected at the
same locality and date; ten paratypes in redwood litter eighteen miles
south of Klamath, Del Norte County, California, August 13, 1953
(G. A. Marsh, R. O. Schuster); and September 19, 1953 (E. E. Gilbert,
R. O. Schuster) ; one paratype from Green Point Ranch, Humboldt
County, California, June 11, 1919 (F. E. Blaisdell); twenty-one paratypes
deposited in the collection of the California Insect Survey, University of
California.

The affinities of this species remain uncertain because individual
characters are found in common with several species, all of which have
characteristic modifications of the ventral surface of the head. For exam-
ple, the general facies of the male of gibbosus is similar to that of ornatus
and roth/ but differs from these species in having the sixth and seventh
segments of the antennae longer than wide. This antennal character is
present in the male of cavatus, from which gibbosus is easily distinguished
by the rounded angulate tempora. The females of gibbosus can be distin-
guished by the glabrous triangular impression on the apical declivity of
the ventral surface of the head.

Marsh-Schuster — Genus Pselaptrichus 25

Pselaptrichus curiosus Park

Pselaptrichus curiosus Park, 1953, Chicago Acad. Sci., Bull., 9(3):
258-259, (pi. 1C; 2B).

Male. Eyes of five facets, triangular in outline; ventral surface of
head with high, median carinoid process; antennal segments III through
VIII similar in size and form; subhumeral foveae of elytra absent;
protibia excavate at inner distal margin; metatibia spined apically; total
length 1.62 mm. (fide Park).

Female. Unknown.

Type locality. Avery, Shoshone County, Idaho.

Type deposition. Northwestern University, Park collection.

Recorded distribution. IDAHO: Avery (Park, 1953 a).

The authors have not seen specimens of curiosus, but there can be little
doubt as to the validity of this interesting species since it is the only one
which lacks the subhumeral foveae.

Selected Bibliography
Bowman, John R.

1934. The Pselaphidae of North America. Pittsburgh, privately
published, p. 1-149.

Brendel, Emil

1889. Description of new Scydmaenidae and Pselaphidae. Entom.
Amer., 5: 193-197.

Brendel, Emil, and H. F. Wickham

1890. The Pselaphidae of North America. Bull. Lab. Nat. Hist.
St. Univ. Iowa, 1: 216-304; 2: 1-84, pi. 6-12.

Cain, Stanley A.

1944. Foundations of plant Geography. Harper and Brothers, New
York, xvi 556 pp.

Park, Orlando

1953. a. New or little known pselaphid beetles of the United States,
with observations on taxonomy and evolution of the family
Pselaphidae. Chicago Acad. Sci., Bull., 9(3): p. 249-283,
pi. 1-5.

1953. b. Discrimination "of Genera of Pselaphid Beetles of the
United States. Chicago Acad. Sci., Bull., 9(16): p. 299-331,
pi. 1-5.

Raffray, Achille

1908. Pselaphidae. Genera Insectorum 64th Fascicle, P. Wytsmann,
ed. Bruxelles, p. 1-487, pi. 1-9.

26

San Diego Society of Natural History

tuberculipalpus

2 rectus

spinosus

4 carinatus

5 oculatus 6 ornatus

Plate I

Male genitalia

1. Pselaptrichus tuberculipalpus Brendel, 0.15 mm. long, 0.15 wide.

2. P. rectus new species, 0.15 mm. long, 0.15 mm. wide.

3. P. spinosus new species, 0.14 mm. long, 0.14 mm. wide.

4. P. carinatus new species, 0.17 mm. long, 0.17 mm. wide.

5. P. oculatus new species, 0.38 mm. long, 0.32 mm. wide.

6. P. ornatus new species, 0.16 mm. long, 0.14 mm. wide.

Marsh-Schuster — Genus Pselaptrichus

27

7 cavatus

8 gibbosus

9 spinosus

10 carinatus

1 1 cavatus

12 oculatus

13 ornatus

Plate II

Male genitalia

14 gibbosus

7. P. cavatus new species, 0.31 mm. long, 0.37 mm. wide.

8. P. gibbosus new species, 0.26 mm. long, 0.21 mm. wide.

Male head

9. P. spinosus new species, ventral surface.

10. P. carinatus new species, lateral aspect.

11. P. cavatus new species, ventral surface.

12. P. oculatus new species, ventral surface.

13. P. ornatus new species, ventral surface.

14. P. gibbosus new species, ventral surface.

28

San Diego Society of Natural History

15 tuberculipalpus

16 carinatus

17

cavatus

18

ornatus

23 ornatus

20 spinosus

24 cavatus

21 carinatus

22 oculatus PLATE III 25 gibbosus

Right maxillary palpi, ventral aspect

15. P. tuberculipalpus Brendel. 17. P. cavatus new species.

16. P. carinatus new species. 18. P. ornatus new species.

Male prothoracic tibia
19- P. tuberculipalpus Brendel. 23. P. ornatus new species.

20. P. spinosus new species. 24. P. cavatus new species.

21. P. carinatus new species. 25. P. gibbosus new species.

22. P. oculatus new species.

TRANSACTIONS

OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY

Volume XII, No. 3, pp. 31-46

THE FOSSIL PIT- VIPERS
(REPTILIA: CROTALIDAE) OF NORTH AMERICA

BY

Bayard H. Brattstrom

Department of Zoology
University of California, Los Angeles

SAN DIEGO, CALIFORNIA

Printed for the Society

July 1, 1954

mi COMP. ZOOL

JUL 1 6 1954

UNIVERSITY

COMMITTEE ON PUBLICATION

Joshua L. Baily

Carl L. Hubbs

Laurence M. Klauber

Charles C. Haines

1

INTRODUCTION

■ ■ '

In 1920, Hay referred several fossil rattlesnake vertebrae to existent
species. Later Gilmore (1938) could find no differences between the
vertebrae of recent species of rattlesnakes, much less between the vertebrae
of the various genera of the Crotalidae. He therefore referred all the
fossil crotalids to Crotalus sp. except for the large vertebrae from the
Pleistocene of Florida which, on the basis of size and distribution, he
referred to the recent species, Crotalus adamanteus.

For several years the writer has been studying the comparative osteol-
ogy of the rattlesnakes and their relatives and has found that not only the
genera, but the species as well, can be distinguished on the basis of skull
and vertebral characteristics. The results of this study will be presented
elsewhere, with reasons for reaching certain conclusions as to identification
of species (Brattstrom, MS.). During the study here reported the fossil
crotalids in the various museums in the country have been examined. So
that some of the names presented herein might be made available, this
short summary of our current knowledge of the fossil rattlesnakes, copper-
heads, and moccasins of North America is presented.

The Crotalidae or "Pit-Vipers" are a family of solenoglyph or mov-
able front-fanged, poisonous snakes showing remarkable parallel and con-
vergent evolution with the Old World family of true vipers (Viperidae).
The Crotalidae differ from the Viperidae in many characters, the most
important of which is the heat-sensitive facial or loreal pit lying in a
depression that accommodates this pit in the maxillary bone.

The Crotalidae include six genera {Crotalus, Sistr tints, Agk'tstrodon,
Lachesis, Bothrops, and Trimeresttrtts) , of which Crotalus, Sistrttrus, and
Agk'tstrodon are the Recent North American representatives. The extinct
genera Neurodromicus Cope, 1873 (Miocene-Oligocene) ; Helagras Cope,
1883 (Paleocene); and Coniopbis Marsh, 1892 (Upper Cretaceous)
were assigned to the Crotalidae by Hay (1902). These genera, except
Coniopbis, were placed in the broad family Viperidae (considering the
Crotalidae as a subfamily) by Williston (1925). Gilmore (1938) re-
moved Coniopbis and Helagras from the Crotalidae and placed them in
Incertae Sedis, but retained Neurodromicus in the family. Vanzolini
(1952) has recently shown, and the writer concurs, that Neurodromicus
should be placed in the Boidae. The oldest known crotalids are Crotalus
viridis and Agk'tstrodon contortrix from the ?Lower Pliocene of Drift-
wood Creek, Nebraska.

The writer has referred the skeletal remains of rattlesnakes from the
asphalt pits and limestone caves of California to existing species (Bratt-
strom, 1953a, 1953b). One fossil from Potter Creek Cave, California,
however, differed from recent species and was described as Crotalus pot-
terensis (Brattstrom, 1953b). As other Pleistocene material containing
rattlesnakes was examined, short papers were published on the herpeto-
fauna of some of the sites. This included some material from Florida
(Brattstrom, 1953c) and from Gypsum Cave, Nevada (Brattstrom,
1954). Many identifications were made on fossil material in the various
museums and these are now reported. In addition, one new species and
one new subspecies are described herein.

34 San Diego Society of Natural History

The large number of vertebrae of Crotalus adamanteus from many
Pleistocene sites in Florida makes possible a statistical comparison with
Recent material. Differences are indicated that show a gradient, or inter-
gradation in time. This problem has faced others (Howard, 1946; Col-
bert and Hooijer, 1953) and they solved the problem by considering the
Pleistocene form as a subspecies of the Recent form. Though many of the
characteristics are obvious and qualitative, to demonstrate further the dif-
ferences, the method of ratio of log- differences of a series of analogous
measurements was used. This method was developed by Simpson (1941)
and used to good purpose by Colbert and Hooijer (1953) in showing
differences between Pleistocene and Recent subspecies of mammals from
China. It has proven to be very useful in distinguishing closely related
forms on the basis of vertebrae (Brattstrom, MS.).

Brattstrom — The Fossil Pit- Vipers 35

SYSTEMATIC LIST

Agk'istrodon contortrix (Linnaeus)
Nebraska, Driftwood Creek, Hitchcock County, PLower Pliocene.

A.M.N.H. 7182, 2 vertebrae; A.M.N.H. 7185, 29 vertebrae.

These 31 vertebrae, representing the first record of fossil copper-
heads, do not differ from vertebrae of Recent specimens.

Agk'istrodon piscivorus (Lacepede)
Florida, Seminole, Pinellas County, Late Pleistocene. A.M.N.H. 7181,
4 vertebrae.

Florida, Melbourne, Brevard County, Late Pleistocene. M.C.Z. 2111, 10
vertebrae.

Florida, Crystal Springs, 28 miles N.E. of Tampa, Pasco County, M.C.Z.
2116, 3 vertebrae.

Crotalus adamanteus pleistofloridensis. new subspecies

TYPE: A.M.N.H. Department of Vertebrate Paleontology, number 6779
(one mid-thoracic vertebra).

LOCALITY AND AGE: Seminole, Pinellas County, Florida, Late Pleis-
tocene.

DIAGNOSIS: A Crotalus differing from C. a. adamanteus in being
larger and in having a higher neural spine, a longer hypapophysis, the
parapophysis longer and bent laterally, and the process of prezygapophysis
wider that the process of the postzygapophysis.

DESCRIPTION OF TYPE: The holotype has a tall neural spine and an
elongate hypapophysis (Figure 2). Neural spine widest at the top, zygo-
sphene indented slightly at the middle, prezygapophysis turned up. slightly.
Centrum cup and ball slightly wider than high, lower part of the para-
pophysis (prepophysis) longer than wide, the hypapophysis elongate and
flat in its posterior part, widest (anteroposteriorly) at its base. Neural
canal a little higher than wide, zygantrum deep and narrow with the re-
sult that the posterior part of the postzygapophysis process is solid. Area
of vertebra between the process of the pre- and postzygapophysis straight,
not curved.

Comparative measurements of all known specimens of C. adamanteus
pleistofloridensis are presented in Figure 1. Measurements in millimeters
of the type (and paratype) are as follows: Height, 36.9 (35.2); width,
14.8 (15.8); length at centrum from ball to cup, 11.8 (12.8); height of
neural spine, 12.1 (10.0); width of neural spine, 8.7 (9.0); height of
centrum, 6.4 (6.8); width of centrum, 7.0 (7.5); width across prezyga-
pophysis, 25.0+ (27.3); width across postzygapophysis, 24.2 (24.2+);
width of prezygapophysis process, 4.5 (4.6); width of postzygapophysis
process, 4.1 (4.0); length of hyapophysis, 18.8 (19.5); width of hyapo-
physis, 4.1 (4.3); height of zygosphene from top of cup, 7.0 (7.0);
width across zygosphene, 10.2 (10.9); length of anterior parapophysis,
5.0 (5.6); width of anterior parapophysis 3.4 (3.4).
REFERRED MATERIAL: (All from the Pleistocene of Florida)
A.M.N.H. 7170, Paratype, Seminole, Pinellas County, 1 vertebra.

36 San Diego Society of Natural History

A.M.N.H. 6776, Seminole, Pinellas County, many vertebrae.

A.M.N.H. 6778, "Allen Cave," Lecanto, Citrus County, 6 vertebrae.

A.M.N.H. 7514, Florida, locality unknown, 17 vertebrae.

U.C. 28910, Seminole, Pinellas County, 2 vertebrae.

U.S.N.M. 13678, 2 miles W. of Melbourne, Brevard County, 19 verte-
brae.

U.S.N.M. 13679, near Melbourne, Brevard County, 5 vertebrae.

U.S.N.M. 13677, near Melbourne, Brevard County, 3 vertebrae.

U.S.N.M. 11220, 2 miles W. of Melbourne, Brevard County, 3 vertebrae.

U.S.N.M. 11333, N. side Canal, Vero Beach, St. Lucie County, 46 ver-
tebrae (figured in Gilmore, 1938, pg. 74, Fig. 31).

U.S.N.M. 11856, Melbourne, Brevard County, 12 vertebrae.

U. Mich. 1009-1010, Seminole, Pinellas County, 2 vertebrae.

M.C.Z. 2101, Melbourne, Brevard County, many vertebrae.

M.C.Z. 2112-2113, Vero Beach, St. Lucie County, 23 vertebrae.

F.G.S. V-471, Florida, locality unknown, 1 vertebra.

F.G.S. V-472, Stratum, Vero Beach, St. Lucie County, 1 vertebra.

F.G.S. V-2455, V-3472, V-2413, V-3492, V-1630, Stratum 3, Vero
Beach, St. Lucie County, 10 vertebrae.

DISCUSSION:

C. a. pleistofloridensis is most closely related to the recent form C.
a. adamanteus Beauvois from which it differs in the characters presented
in the diagnosis above, but especially in its large size. The subspecies
C. a. pleistofloridensis is extinct though it is survived directly by C. a.
adamanteus. The extinct subspecies occurs in the many Pleistocene locali-
ties mentioned above and in association with an extinct, still larger species
(described below).

Photographs of the type, paratype, and several series of C. a. pleisto-
floridensis are shown in Figure 2. A vertebra of this subspecies was fig-
ured in Gilmore (1938), as C. adamanteus, (Fig. 31, pg. 74) from Vero
Beach, Florida.

Crotalus atrox Baird and Girard
Nevada, Gypsum Cave, Late Pleistocene-Recent (8,000 to 10,000 years)

C.I.T. 94 vertebrae.
Texas, Bulverde, Bexar County, Middle Pleistocene, U.S.N.M. 9223, 10
vertebrae, 1 dentary, 1 maxilla and fang, 15 ribs.
Crotalus atrox was first found as a fossil by Hay (1920), from Texas.
A vertebra from this collection was figured by Hay (1920, pi. 10, Fig.
2.), upside down. It was also figured by Gilmore (1938, pg. 73, fig. 30).
The length of the fang from Texas is 11.0 mm.

Crotalus giganfeus, new species
TYPE: A.M.N.H. Department of Vertebrate Paleontology number 6772
(one vertebrae) .

TYPE LOCALITY AND AGE: "Allen Cave," Lecanto, Citrus County,
Florida, Pleistocene.

DIAGNOSIS: A large rattlesnake related to C. adamanteus but differing
from it in the large size of the vertebra, centrum, hypapophysis, and
parapophysis, and in having the hypapophysis widest (anteroposteriorly)
at its tip and not its base.

Brattstrom — The Fossil Pit-Vipers 37

DESCRIPTION OF TYPE: A large vertebra measuring 41.2 mm. high,
17.7 mm. wide, and 13.3 mm. long at the centrum from cup to ball (Fig-
ure 2). Neural spine relatively wide, front of zygosphene even with the
anterior edge of neural spine. Top of the zygosphene slanting ventrally.
Centrum round, parapophysis almost as wide as long, width across pre-
zygapophysis greater than width across postzygapophysis, process of pre-
zygapophysis wider than process of postzygapophysis. Hypapophysis
longer than height of neural spine and widely expanded at its tip. The
tip of the left prezygapophysis is broken in the type. Measurements of
the type (in millimeters) are as follows: height and width of neural
spine, 13.7 and 11.4; height and width of centrum, 7.5 and 5.6; width
across pre- and postzygapophyses 29.6 and 29.2; width of process of pre-
and postzygapophysis, 5.9 and 5.1; length and width of hypapophysis,
22.9 and 5.0; height of zygosphene above centrum cup, 7.9; width across
zygosphene, 11.5; length and width of anterior parapophyses, 6.9 and 5.2.
PARATYPE: Besides the type there is a fragment of a vertebra
(A.M.N.H. 7171) that is designated as the paratype. This fragment con-
tains the left side of the vertebra and the centrum at its posterior end.
This centrum is also round (height and width, 7.5 and 7.5 mm.).

DISCUSSION: The type and paratype are figured in Figure 2, and their
measurements are compared with those of Recent C. a. adamant eus and
Pleistocene C. a. pleistofloridensis in Figure 1, on a Ratio of log-differ-
ence scale.

C. giganteus most closely resembles C. a. pleistofloridensis, from
which it differs in its much larger size and in having the hypapophysis
widest (anteroposteriorly) at its tip. Crotalus giganteus is found in asso-
ciation with (and thus probably co-existing with) C. a. pleistofloridensis
and Lampropeltis getulus in "Allen Cave." Recent C. a. adamant eus at-
tains a maximum recorded size of 7 feet, 4 inches. Judging from the large
size of C. a. pleistofloridensis and the much larger C. giganteus vertebrae,
it is possible to speculate as to the maximum sizes of these forms. It is
possible that C. a. pleistofloridensis attained a length of 9 or 10 feet, and
12 feet was probably the maximum length for C. giganteus. The width
of giganteus was probably similar to that of the modern boa constrictor.
The species is extinct.

Crotalus horridus Linnaeus

Arkansas, Conard Fissure, Pleistocene. A.M.N.H. 6404, many vertebrae
and the posterior half of a lower jaw.

Georgia, Near Cartersville, Pleistocene, U.S.N.M. 4887, 53 vertebrae.

Maryland, Cavetown, Middle Pleistocene, U.S.N.M. 9157, 2 vertebrae.

Maryland, Cumberland Cave, Pleistocene, U.S.N.M. 13772, 11 vertebrae.

Massachusetts, , Pleistocene, A.M.N.H. many bones, Hecht, MS.

Pennsylvania, Carlisle Cave, Pleistocene-Recent, U.S.N.M. 102698, 4 ver-
tebrae.

Tennessee, Big Pigeon River, near Newport, Cocke County, Pleistocene,

U.S.N.M. 13686, 13 vertebrae.

The only previously known fossil timber rattlesnake was the C. hor-
ridus discussed by Hay (1920) from Cavetown, Maryland. The material

38 San Diego Society of Natural History

from Massachusetts will be discussed by Hecht (MS), though I have
examined the material and approve its identification as C. horridus. All
the fossil material falls within the variation demonstrated by C. horridus.

Crotdus cf. lepidus Kennicott
Arizona, San Pedro Valley, Cochise County, Early Pleistocene. C.I.T. (no
number), 1 vertebra.

The one small vertebra from the Pliocene of Arizona is probably C.
lepidus, as it agrees with it in all characters. The fossil is so fragmentary,
however, that it can only tentatively be referred to this species.

Crotdus mite belli (Cope)
Nevada, Gypsum Cave, Late Pleistocene-Recent (8,000-10,000 years),
C.I.T. (no number), 55 vertebrae plus one entire skeleton.

Crotdus potterensis Brattstrom
California, Shasta County, Potter Creek Cave, Late Pleistocene, U.C. 3129,

2 vertebrae, a part of a pterygoid, fragments of ribs and three other

fragments of bones.

No other material has come to light since the recent original descrip-
tion of this extinct species.

Crotdus scutulatus (Kennicott)
Mexico, Zumpango, Mexico, Pleistocene. C.I.T. Locality 310, C.I.T. (no

number), 2 vertebrae.

These two mid-thoracic vertebrae do not differ from recent specimens
of this species.

Crotdus riridis (Rafinesque)
California, Rancho La Brea, Los Angeles County, Late Pleistocene,

L.A.C.M. Pits A, B, 81, U.C. locality 2052, many vertebrae.
California, McKittrick Asphalt, Kern County, Late Pleistocene, U.C.
35114, 35122, 35123, 35124, 35125, 35126, 35127, 35128, 20 ver-
tebrae.
California, Hawver Cave, El Dorado County, Late Pleistocene. U.C. (no

number), 5 vertebrae.
Iowa, Andrews Stone Quarry, above Sioux City, Pleistocene. U.S.N.M.

13655, 7 vertebrae.
Nebraska, Driftwood Creek, Hitchcock County, PLower Pliocene.
A.M.N.H. 1639, many vertebrae, 2 fangs (length 7.0 and 6.8 mm.
long), A.M.N.H. 1638, 2 vertebrae, A.M.N.H. 1637, many verte-
brae.
Nevada, Gypsum Cave, Late Pleistocene-Recent (8,000-10,000 years).
C.I.T. (no number), 17 vertebrae.

The California material of this widespread rattlesnake has been dis-
cussed (Brattstrom, 1953a, 1953b) as was the Gypsum Cave fossils
(1954). As comparative measurements of C. viridis were given previ-
ously, the summary of the measurements made on the Gypsum Cave,
Nebraska, and Iowa material is presented herein (Table I). The material
from all three of these sites averages slightly larger than those from
Rancho La Brea, though the maximum measurements are not larger than
for the La Brea material.

Brattstrom — The Fossil Pit-Vipers 39

AGE (12BBCAPS)

In the following list the North American Crotalids are listed by the
age from which they have been recorded. An asterisk indicates that the
species or subspecies is extinct.

PLIOCENE

Agkistrodon contortrix
Crotalus viridis

PLEISTOCENE

Agkistrodon piscivorus
* Crotalus adamant eus pleistofloridensis

Crotalus atrox
* Crotalus giganteus

Crotalus horridus

Crotalus cf. lepidus

Crotalus mitchelli
* Crotalus potterensis
* Crotalus scut ul at us

Crotalus viridis

40 San Diego Society of Natural History

LOCALITIES

In the following list the North American Crotalids are listed as to
the fossil localities from which they have been found.

ARIZONA

San Pedro Valley, Early Pleistocene
Crotalus cf. lepidus

ARKANSAS

Conard Fissure, Pleistocene

Crotalus horrid us

CALIFORNIA

Rancho La Brea, Late Pleistocene

Crotalus viridis
Hawver Cave, Late Pleistocene

Crotalus viridis
McKittrick Asphalt, Late Pleistocene

Crotalus viridis
Potter Creek Cave, Late Pleistocene

Crotalus potterensis

FLORIDA

"Allen Cave," Lecanto, Pleistocene

Crotalus giganteus

Crotalus adamanteus pleistojloridensis
Melbourne, Late Pleistocene

Agkistrodon piscivorus

Crotalus adamanteus pleistojloridensis
Vero Beach, Late Pleistocene

Crotalus adamanteus pleistojloridensis
Seminole, Late Pleistocene

Agkistrodon piscivorus

Crotalus adamanteus pleistojloridensis
Crystal Springs, Late Pleistocene

Agkistrodon piscivorus
No Locality, Pleistocene

Crotalus adamanteus pleistojloridensis

GEORGIA

Cartersville, Pleistocene

Crotalus horrid us

IOWA

Andrews Strong Quarry, Sioux City, Pleistocene

Crotalus viridis

MARYLAND

Cavetown, Pleistocene

Crotalus horridus
Cumberland Cave, Pleistocene

Crotalus horridus

Brattstrom — The Fossil Pit-Vipers 41

MASSACHUSETTS

— , Pleistocene, Hecht, MS.
Crotalus horridus

NEBRASKA

Driftwood Creek, PLower Pliocene
Agkistrodon contortrix
Crotalus viridis

NEVADA

Gypsum Cave, Late Pleistocene-Recent

Crotalus atrox
Crotalus mitchelli
Crotalus viridis

PENNSYLVANIA

Carlisle Cave, Pleistocene-Recent
Crotalus horridus

TENNESSEE

Big Pigeon River, Pleistocene
Crotalus horridus

TEXAS

Bulverde, Pleistocene

Crotalus atrox

MEXICO

Zumpango, Pleistocene

Crotalus scutulatus

42 San Diego Society of Natural History

ACKNOWLEDGMENTS

The writer wishes to thank all who lent him fossil and recent skele-
tons, as well as those who have offered many suggestions, criticisms, and
encouragements (a full list of these people will be presented elsewhere).
I wish especially to thank, for the use of fossils, the following (the initials
indicate the abbreviations of institutions used in this paper) : Dr. Charles
L. Camp, Museum of Paleontology, University of California, Berkeley
(U.C. ); Dr. Hildegarde Howard, Los Angeles County Museum
(L.A.C.M. ); Dr. Ernest Williams, Museum of Comparative Zoology,
Harvard University (M.C.Z. ); Dr. Edwin H. Colbert, Department of
Paleontology, American Museum of Natural History (A.M.N.H.); Dr.
Doris M. Cochran, Department of Herpetology and Dr. D. H. Dunkle,
Department of Vertebrate Paleontology, U. S. National Museum
(U.S.N.M.); Mr. William Otto, Department of Vertebrate Paleontology,
California Institute of Technology (C.I.T.); and Dr. Gordon Gunter
and Dr. H. H. Winters, Florida Geological Survey (F.G.S.).

The writer wishes to also thank Dr. Clark P. Read, Department of
Zoology, University of California, Los Angeles, and Dr. Laurence M.
Klauber, San Diego Society of Natural History, for their suggestions and
criticisms.

Literature Cited

Brattstrom, Bayard H.

1953a. The amphibians and reptiles from Rancho La Brea. Trans.
San Diego Soc. Nat. Hist. 11 (14): 365-392.

1953b. Records of Pleistocene reptiles from California. Copeia 3:

174-179.
1953c. Records of Pleistocene reptiles and amphibians from Florida.

Quart. Jour. Fla. Acad. Sci. 16 (4): 243-248.

1954. Amphibians and reptiles from Gypsum Cave, Nevada. Bull.
So. Calif. Acad. Sci. 53 (1): 8-12.

Colbert, Edwin H. and D. A. Hooijer

1953. Pleistocene mammals from the limestone fissures of Szechwan,
China. Bull. Amer. Mus. Nat. Hist. 102 (1): 1-134.

Gilmore, Charles W.

1938. Fossil snakes of North America. Sp. Pap. Geol. Soc. Amer.
9: 1-96.

Hay, O. P.

1902. Bibliography and catalogue of the fossil Vertebrata of North
America. U. S. Geol. Surv. Bull. 197: 478-481.

1920. Description of some Pleistocene vertebrates found in the
United States. Proc. U. S. Nat. Mus. 58: 83-146.

Brattstrom — The Fossil Pit-Vipers 43

Howard, Hildegarde

1947. An ancestral golden eagle raises a question in taxonomy.
Auk 64: 287-291.

Simpson, G. G.

1941. Large Pleistocene felines of North America. Amer. Mus.
Novitates, 1136: 1-27.

Vanzolini, P. E.

1952. Fossil snakes and lizards from the Lower Miocene of Florida.
Jour. Paleo. 26 (3): 452-457.

Williston, S. W.

1925. Osteology of the Reptiles. Harvard Univ. Press, 298 pp.

44

San Diego Society of Natural History

Table 1

SUMMARY OF MEASUREMENTS (IN TENTHS OF MM.)
OF FOSSIL Crotalus viridis

Measurement

Locality

Number of specimens, range (in parentheses).

and average of measurements

IOWA

NEBRASKA

NEVADA

VERTEBRAE

Height
Width
Length at centrum

1(138)

7(45-54)49
7(60)60

4(70-113)97
12(33-63)49
12(40-64)53

2(79-113)96

2(49-74)62

2(55-62)59

NEURAL SPINE

Height
Width

3(30-49)41
3(38-47)56

7(19-41)30
7(24-47)38

2(19-28)24
2(43-46)45

PREZYGAPOPHYSES

Width across

5(85-98)91

10(58-117)80

2(87-126)107

Width of process

1(20)

7(10-20)17

2(20-23)22

POSTZYGAPOPHYSES

Width across
Width of process

4(84-87)85
1(20)

12(63-106)81
7(10-22)18

2(88-119)104
1(20)

CENTRUM

Height
Width

6(27-32)30
6(28-32)30

8(20-33)29
8(21-34)30

1(30)
1(33)

ZYGOSPHENE

Width across
Height from centrum
Width of process

5(37-40)39
5(25-30)28
5(10-20)20

7(27-48)41
7(19-30)28
7(10-21)18

1(50)
1(29)
1(20)

Brattstrom — The Fossil Pit-Vipers

45

-35 -30 -25 -20 -15 -10 -05 0 05 10 15 20 25 .30 .35 40 45 ,50

VERTEBRAE HEIGHT

VERTEBRAE WIDTH

C,.

/ BH

WIDTH ACROSS POSTZYGAPOPHYSIS \ B j_'

C >

WIDTH ACROSS PREZYGAPOPHYSIS

LENGTH OF CENTRUM

NEURAL SPINE HEIGHT

ZYGOSPHENE HEIGHT

ZYGOSPHENE WIDTH

PARAPOPHYSIS WIDTH

C *,-'

PARAPOPHYSIS LENGTH vv

C

CENTRUM HEIGHT
HYPAPOPHYSIS LENGTH

BH

A( -/—

Al ^r

C>

Al T

Bl f-

\AH

O

Bt-

ftt-

C

B»-

Figure 1. Ratio diagram (on a log-difference scale) showing comparison
of certain osteological characters in Crotalus giganteus and two subspecies
of C. adamanteus. A. Recent C. a. adamanteus, summary of many speci-
mens. Standard (.0) is largest vertebra of a good-sized C. a. adamanteus,
U.S.N.M. 29419. B. All Pleistocene material of C. a. pleistofloridensis;
dashed line passes through average. C. Type of C. giganteus. ■ A strong
overlap is evident in A and B, with very little in C.

46

San Diego Society of Natural History

Figure 2. Thoracic vertebrae of Crotalus adamanteus pleistofloridensis
and C. giganteus. 1 and 4, type and paratype of C. a. pleistofloridensis,
Seminole, Florida. 5, 7, 8, 10, and 11, C. a. pleistofloridensis, part of
syntypic series, Seminole, Florida. 6, 9, and 12, C. a. pleistofloridensis,
A.M.N. H. 7514, Florida, no other data. 2 and 3, type and paratype of
C. giganteus.

TRANSACTIONS

OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY

Volume XII, No. 4, pp. 47-64, plates 4-5, figs. 1-2

ANALYSIS OF THE HERPETOFAUNA OF
BAJA CALIFORNIA, MEXICO

BY

Charles H. Lowe, Jr. and Kenneth S. Norris

SAN DIEGO, CALIFORNIA

Printed for the Society

September 10, 1954

MUS. COM

LIBR

SEP 3 i

HARY

UNIVt!

COMMITTEE ON PUBLICATION

Joshua L. Baily

Carl L. Hubbs

Laurence M. Klauber

Charles C. Haines

Lowe-Norris — Herpetofauna of Baja California 49

ANALYSIS OF THE HERPETOFAUNA OF
BAJA CALIFORNIA, MEXICO

BY

Charles H. Lowe, Jr. and Kenneth S. Norris
I. INTRODUCTION

This study is the first of a series concerning the biogeography and
systematics of the herpetofauna of Baja California, Mexico, and adjacent
islands. Field work in this connection was begun at the close of World
War II. In the summer of 1949, studies in Baja California were extended
to the Cape San Lucas region, from sea level to the 7,000 foot crest of
the Sierra Laguna. At this writing, work is continuing, both in the labora-
tory and field.

The patterns of ecologic and geographic distribution shown by the
reptiles and amphibians of Baja California and adjacent regions can be
correlated with the recently defined history of Tertiary environments of
Western North America. This environmental history has been illuminated
largely through advances in the field of paleobotany during the last
decade. We propose to define the variation and ecologic distribution of
the amphibians and reptiles of the peninsula and adjacent islands as well
as available material well permit and to attempt the correlation of distri-
bution patterns with the environmental history.

Among the new forms which have come to light in the course of
our field work is a new northern coastal subspecies of Crotalus enyo, the
Baja California Rattlesnake. This form is described, its ecology discussed,
and pertinent new scale terminology is proposed.

II. ANALYSIS OF SUBSPECIFIC

DIFFERENTIATION IN CROTALUS ENYO,

THE BAJA CALIFORNIA RATTLESNAKE

Scale Terminology

There is, as yet, no adequate terminology for the somewhat irregular
scales on the side of the head of Crotalus. C. enyo is not unique in hav-
ing small scales present in thcee areas on the side of the head: (1)
bordering the upper loreal, (2) at the upper anterior corner of the eye,
and (3) about the loreal pit. Current problems relative to these scales
have been obscured by the lack of an accepted precise terminology. This
problem has been discussed with Laurence M. Klauber, and the new
names for scales as used herein are the result of our mutual agreements
(see Klauber, 1952:9, footnote).

San Diego Society of Natural History

O C. ENYO FURVUS

C. ENYO ENYO

Fig. 1. Distribution of the subspecies of Crotalus enyo, the Baja California Rattlesnake.

Lowe-Norris — Herpetofauna of Baja California 51

12 3 4 5

13 12 U 10

Fig. 2. Scales on the anterior lateral surface of the head of Cro talus, as exem-
plified by C. enyo. 1, lower loreal (subloreal) ; 2, upper loreal (supraloreal) ;
3, lacunal; 4, postloreal; 5, postsupraloreal ; 6, preciliary; 7, orbit; 8, upper pre-
ocular; 9, lower preocular; 10, postfoveals; 11, subfoveals; 12, second supralabial;
13, prefoveals, (there are five in this example, as usual, they occupy a triangular
area); 14, rostral; 15, prenasal ; 16, postnasal.

The following terms for head scalation are suggested (see Fig. 2) :

(1) prefoveal(s), subfoveal(s). and postfoveal(s). for the small
scales at the anterior, lower, and posterior borders of the pit. The scales
lining the pit (and small extensions of these scales along its outer bor-
der) are termed lacunals (see below) . Two of these terms (subfoveal
and postfoveal) are restricted in the sense that they apply only to the
border roiv of scales about the pit (Fig. 2). The prefoveals comprise all
small scales in the small area anterior to the subfoveals which are bor-
dered by the following scales: supralabial (s), subfoveal, lacunal, loreal
or lower loreal, and nasal. Where prefoveals and subfoveals are to be
distinguished on the one hand, and postfoveals and subfoveals on the
other, is a matter of reference to a line drawn at 45 degrees to the hori-
zontal, through the center of the pit. This is analogous to the situation in
segregating suboculars and postoculars. The term prefoveal may prove to
be the most useful of the three; the terms subfoveal and postfoveal may
be used where they clarify description, as in C. enyo.

(2) lacunal(s), for the relatively large scale (s) forming the inner
border of the pit. These scales are reduced in size when foveal scales,
particularly subfoveals, are present.

(3) preciliary(s), for the small scale in the upper anterior corner
of the eye. This scale is just below the contact of the postsupraloreal and
the supraocular and is a derivative from the upper preocular scale, that
is split off of its upper posterior corner. It is not consistently present in
C. enyo. The postsupraloreal is more consistently present.

(4) presupraloreal(s) and postsupraloreal(s), for the small scales
which may border the supraloreal and which are, for the most part, below
the canthus. Such distinction is useful in some species, including C. enyo.
but becomes arbitrary and confusing in others, such as C. molossus, that
have numerous scales in the loreal region. The presupraloreal(s) is an-
terior to the upper loreal (=supraloreal) , between that scale and the
postnasal. The postsupraloreal(s) is located posterior to the upper loreal,
between it and the supraocular; it is anterior to, and slightly above, the
preciliary, when that scale is developed. These scales are present in a few

52 San Diego Society of Natural History

species including C. enyo which characteristically has a single postsupra-
loreal but lacks a presupraloreal.

(5) postloreal(s), for the small irregularly occurring scale (s) inter-
posed between the loreal(s) and preocular(s), as in C. enyo. By use of
the proposed terms "postloreal" and "preciliary" in forms like C. enyo,
there is no disturbance of the usual nomenclatorial designations of the
regular upper and lower preoculars and upper and lower loreals. This
distinction also is less applicable to forms, like C. molossus, that have
numerous loreals.

Crotalus enyo furvus, subsp. nov.1

Dusky Baja California Rattlesnake

Holotype. — Adult male, No. 55388, Museum of Vertebrate Zoology,
collected alive by Kenneth S. Norris and Charles H. Lowe, Jr., July 21,
1949, 10.9 miles (by road) north of El Rosario, along the main road
on the coastwise terrace near the foot of a bold Cretaceous escarpment,
Baja California Norte, Mexico. The type is deposited in the Museum of
Vertebrate Zoology, University of California, Berkeley, California. (Fig.

3)-

Diagnosis. — A distinctive subspecies distinguished from the pallid
C. e. enyo by a very dark brown ground color and darker blotches and
spotting; proximal rattle segment black; 10-12 (11.0) posterior body
crossbands; the lacunal(s) wholly or partly separated, by 1-3 subfoveals,
from the supralabial row; 3 scales (average) in minimum count between
orbit and supralabials.

Measurements (taken on the fresh specimen) and scalation of holo-
type.— Snout-vent length, 606 mm.; tail length, 60 mm.; ratio of tail
to total length, .090; head length, 30.6 mm., contained 21.8 times in over-
all length; head width, 21.7 mm.; ratio of head length to head width,
1.41.

Scale rows, 25-25-21, with 1 1 at middle of tail; scales are strongly
keeled, with first and occasionally second rows smooth on anterior and
middle body, with all rows keeled an extreme posterior body and tail.
Ventrals, 162; subcaudals, 29, of which the last 4 are divided. Anal
entire. Supralabials, 13 — 13; infralabials, 13 — 14. First infralabials undi-
vided; first two in contact with genials on either side; mental triangular;
neither intergenials nor submentals present. Rostral wider than high and
in contact with supralabials. Internasals, 2. Minimum scale rows between
supraoculars, 5. Nasals, 2-2. Loreals, 2, of approximately the same size,
or larger than, upper. A postsupraloreal scale between supraocular and
supraloreal. A single postloreal scale. Canthals, 3. Preoculars, 2, with
suprapreocular larger. A preciliary scale in the upper anterior corner of
eye. Minimum scale rows from supralabials to orbit, 3. Total scales in
orbit, 9. Prefoveals, 6-4, subfoveals, 3-3, and postfoveals, 2-2, forming
a complete row separating the supralabials from the lacunal on the outer
margin of the pit floor. Rattle with 14 segments, broken near former tip.

1 The name (L. furvus, dark) refers to the dark or dusky coloration of this
race.

Lowe-Norris — Herpetofauna of Baja California 53

Coloration of holotype (in life). — The principal ground color of the
upper surfaces is dark brown (PI. 15, near C 6)2 on the middle third of
the body. Anteriorly, the ground color is slightly darker. Posteriorly, the
ground color gradually lightens to tan (14, H 8) just anterior to the
vent. Everywhere the ground color is somewhat darker than the primary
dorsal row, or series, of blotches, which are rich brown (15, A 12) and
are edged with black. The blotch outlines do not adhere to scale rows.
The black-edging scales and black scales elsewhere are often tipped with
gray (29, A 1). The dorsal blotches of the primary series gradually
change backward from longitudinal rectangles on the neck through hexa-
gons and crude diamonds to crossbands on the posterior body and tail.
Dorsally, the brownish posterior body crossbands (15, L 12) are faintly
edged with dark brown to black; laterally, they are darker brown (15,
A 17). A secondary series of smaller blotches, on the lower lateral sur-
face, parallels the primary series. These are variously positioned on the
first six scale rows. On the posterior body, they coalesce with the dorsal
series to form the afore-mentioned crossbands. Anteriorly, however, they
are well separated from the dorsal blotches; the first 5 or 6 on the neck
are brown, bordered with black as in the dorsal primary series. Farther
back, they take the form of small black spots, involving 2 to 5 scales,
until they coalesce with the primary series where they are brown. Still
smaller and lighter spots on the first 3 scale rows form a tertiary series
and alternate with the elements of the secondary series. They vary in
color from black to brown (anterioposteriorly) . Faint traces of other
blotches, appearing as slight darkenings of the ground color, occur above
the tertiary spots on the mid-lateral surfaces, alternate with the spots of
the primary and secondary series. These faint blotches in the adult may
be traces of more prominent juvenile markings which fade in ontogeny.

The ground color of the upper surfaces of the head posterior to the
eyes is the same as that of the neck and anterior body. On the anterior
portions, the ground color is an even darker brown. A pair of parallel
longitudinal marks begins at the dark brown (8, J 10) supraoculars and
continues posteriorly onto the neck, where they become confluent just
anterior to the first dorsal blotch. They do not join with this blotch. The
brown color (15, A 1) and black bordering of these marks is approxi-
mately the same as that of the dorsal blotches. A darker lateral head
stripe of rich olive brown (15, H 7), stippled with blackish brown and
faintly bordered with black, arises at the ventroposterior corner of the
eye, passes backward above the commissure and turns below it posteriorly.
No preocular light line is present. A light tan (12, C 5) supraocular cross
mark on the anterior half of this scale widens mesially, curving both for-
ward and inward from the outer edge.

The color and pattern of the proximal two-thirds of the tail approxi-
mates that of the extreme posterior body. There are 5 light brown cross-
bands on the tail. The distal third of the tail is a uniform dark blackish
brown. The proximal rattle segment is solid black; the second segment

2 Maerz and Paul (1930) color determinations.

54 San Diego Society of Natural History

is black tinged slightly with brown. The remaining segments are light
brown.

Ventral coloration at mid-body is yellowish (10, C 2 to 10, E 2),
with light brown stippling, becoming lighter anteriorly and evenly grad-
ing to white on the neck and ventral surface of the head. The mental
and first 4 infralabials are a darker blackish brown than the remaining
infralabials. The genials and other ventral scales of the anterior portion
of the head scales are stippled with light brown over the white ground
color. Occasional brown stippling occurs on the remainder of the gular
area. The color of the iris is light brown (13, C 5).

Intrasub specific and intersubspeafic variation. — Individual differences
are slight in the 4 available specimens of C. e. furvus. Certain of these
differences are set forth in Table 1, in which comparison is also made
with C. e. enyo. The largest specimen of f units, a badly damaged male
DOR (KSN 1337), measures approximately 580 mm. from snout to
vent.

Like many other immature rattlesnakes, a female of e. furvus
(LMK 41087), studied shortly after preservation in alcohol, was found
to be lighter in ground color and bolder in pattern. The mediolateral
blotches that alternate with those of the primary and secondary series are
clearly marked and less obscured than in the adult. These marks are absent
from the neck and posterior quarter of the body in the immature. In
color they are darker brown than the ground color and partly edged
with black in the same manner as the dorsal primary series of blotches.
In this small specimen there are two prominent black spots on each infra-
labial row ; one is on the third infralabial scale, and the other, of approxi-
mately the same size, borders the ninth and tenth infralabials. The dorsal
head marks of this specimen are unlike the type and more like e, enyo
in being broken at the middle of the head and not confluent posteriorly.
In addition, this specimen does not have a preciliary scale, and in con-
trast to the remainder of the topotypic series, the upper and lower loreals
are of approximately the same size on one side, but with the upper still
slightly larger on the other side. Nevertheless, it appears that a difference
in the size of the loreals may prove to be another diagnostic scale differ-
ence between e. enyo and e. furvus.

In e. enyo. the third and fourth supralabials are often in contact
with the lacunal scale. In e. furvus, the supralabials are completely or
nearly completely separated from the lacunal (s) by one or more small
foveals (see Fig. 2). Many of these small scales (particularly the sub-
foveals), when present, are developed at the expense of the material
which would otherwise form a larger lacunal scale(s).

When a larger series of e. furvus is available, the following char-
acters may prove to be clinal: the number and relative size of canthals,
relative size of the upper and lower loreals, number of scales between
supralabials and orbit, number of scales in orbit, number of prefoveals
and subfoveals, and number of infralabials in contact with genials. The
geographic differences in color (Table 2) may intergrade sharply, in
step-cline fashion. The color features of both e. furvus and e. enyo appear

Lowe-Norris — Hfrpetofauna of Baja California 55

to be correlated with color features of their habitats. The known ranges
of the subspecies of C. enyo (Fig. 1) are separated by about 60 miles
and no intergrades have yet been taken. The northernmost specimens of
e. enyo were collected by L. M. Huey in 1930 from Jaraguay and from
near Cataviha (Klauber, 1931). These two northerly specimens of e. enyo
extended the known range of the species about 270 miles north of the
northernmost previously known record, at Mulege. The type specimen
of e. furvus, collected near El Rosario in 1949, again extended the known
range of the species an additional jump to the north — in this case 60
miles, nearly 20 years later and in a relatively well traversed area of
Baja California. The third specimen of e. furvus, from Punta Camalu,
extended the range northward an additional 45 miles; the fourth speci-
men slightly farther. With additional collecting, the present gaps will
almost certainly narrow, and intergrades between the two subspecies may
be expected. The new form is accorded subspecific status because the
discovery of intergradation is anticipated and because, even though geo-
graphic intergrades are not discovered or do not exist, the differences
will not prove sufficiently consistent to warrant specific separation. The
differences, furthermore, appear to represent the low level of morpholo-
gical difference usually associated with subspecific differentiation.

Locality Records. — In addition to the type, 3 specimens (paratypes)
of C. e. furvus are available. These are (1) an immature female, LMK
41087, collected (LOR) by Charles E. Shaw and Richard Schwenkmeyer,
April 2, 1950, 2 miles north of El Rosario, near the type locality; (2)
an adult male, KSN 1337, collected (DOR) by Kenneth S. Norris,
August 30, 1951, at Punta Camalu; and (3) an immature female, KSN
1454, collected (DOR) by K. S. Norris and Arthur Lockley, May 24,
1952, 9.5 miles south of San Telmo River valley (on main road), all in
Baja California Norte, Mexico.

The localities at which C. enyo has been collected are the following:3

CROTALUS ENYO ENYO

Southern Cape Region, Baja California, Mexico
Cape San Lucas (type locality)
La Paz

San Jose del Cabo
San Bartolo
Miraflores

6 miles south of Miraflores
Santa Anita
San Antonio

Sierra de la Laguna, opposite Todos Santos
San Pedro
Todos Santos
Sierra de la Laguna
La Rivera
Eureka
San Pedro Mountains

3 Localities are from published data (Klauber, 1931; Gloyd, 1940), from
Klauber (in litt.), and localities recorded during our studies to date.

56 San Diego Society of Natural History

Central Baja California Region
Mulege
San Ignacio

29 miles south of Punta Prieta
Almejas Bay, Santa Margarita Island
Los Angeles Bay

10 miles west of Los Angeles Bay
2 miles south of Los Angeles Corral
1.6 miles west of San Ignacio

Northern Baja California Region
Jaraguay
10 miles north of Catavina

Gulf of California Islands
Isla Partida
San Francisco Island
Carmen Island

CROTALUS ENYO FURVUS

Northern Baja California Region

10.9 miles north of El Rosario, along main road (type

locality)

2 miles north of El Rosario, along main road

Punta Camalu

9.5 miles south of San Telmo River valley, along main

road
Ecology and Distribution. — The area from which the four speci-
mens of C. enyo furvus have been taken is relatively limited in extent.
This is the coastal strip from 2 miles north of El Rosario northward to
Punta Camalu, 65 miles north of EI Rosario. Almost certainly this does
not represent the total range of the animal. The plant associations and
climatic conditions typical of the localities at which the snake has been
found to date extend both north and south of the present known locali-
ties. A tentative and conservative estimate of the probable range of the
subspecies, the evidence for which is discussed below, is the western
coastal area of northern Baja California from Punta San Antonio (or
possibly Punta Canoas farther to the south) northward to Cabo Colnett.
This range includes the townsites of El Rosario and San Quintin.

At San Quintin, in 1949 and prior to the finding of the Camalu
(northernmost) specimen, the authors and Mrs. Lowe were the guests
of Mr. Frank Frymier, a resident American farming on the flat plain.
He described two kinds of rattlesnakes from the plain, a large red one
(obviously C. r. ruber) and a smaller "gray" one (C. enyo furvus). Both
were adequately described from his firsthand knowledge of the snakes.
(The Camalu and near San Telmo River valley specimens, more recently
collected, confirm our opinion that C. enyo furvus is an inhabitant of the
Sin Quintin area and northward.) Mr. Frymier said that many of the
small gray rattlesnakes were killed during brush clearing operations. A
large drag was pulled over the brush, rolling it away from the "round.
ITe stated the red snake and the smaller gray" one were present in about

Lowe-Norris — Hhrpetofauna of Baja California 57

equal numbers. When shown living C. r. ruber and C. enyo furvus, he
identified them as being the two rattlesnakes present on the plain.

The San Quintin Plain extends for several miles along the coast.
Beyond the plain to the north, to a few miles north of Punta Cabras
which is north of Cabo Colnett, and to the south at least to Punta San
Antonio (south of El Rosario), there is a continuation of the same open,
low-growing scrubby vegetation on the coastly terraces. South of El
Rosario. the coast rises more abruptly to foothills than it does to the
north. Similarly, north of Punta Cabras the country is rugged and moun-
tainous along the coast, with narrow coastal shelves and numerous high
sea cliffs. Rather marked environmental changes occur over relatively
short distances to the northward, eastward, and southward of the coastal
area just described. C. e. furvus may also occur on the higher coastal ter-
races back of the San Quintin Plain.

At the type locality (Fig. 3B), 10.9 miles north of Rosario (along
the main road), which is south and slightly higher in elevation than the
San Quintin Plain, the habitat is a marine terrace extending inland from
sea bluffs to nearby low hills. The area is covered by low shrubby vege-
tation. Euphorbia misera is a conspicuous plant. It is common both to
the north and south along the coastal plains and terraces. Agave and
Opunt'ia are conspicuous but less common.

It is of interest to note that the extreme latitudinal range of the
endemic Crotahis enyo from Cape San Lucas northward is nearly within
that of the giant cactus, Cereus pr'inglei. the "Cardon" or "Cardon
Grande." The northernmost Cardons occur in the vicinity of Hamilton
Ranch and on the San Quintin Plain. Another conspicuous cactus, the
Senita (Cereus schott'i), reaches its northern limit in Baja California in
this general vicinity.

At San Quintin, average rainfall is approximately 5 inches an-
nually, and increases northward to approximately 9 inches at Ensenada
(Beal, 1948). Southward, rainfall decreases rapidly, though coastal fogs
are of common occurrence on the Vizcaino Desert and farther south.
The terraces and plains in the general vicinity of San Quintin and El
Rosario are areas characterized by a markedly more mesic climate and
biota than the harsh environments both southward and in the interior.

As has been pointed out. the manner in which the two subspecies
of C. enyo differ most conspicuously is in the degree of difference in
amount of melanin. C. enyo enyo is light colored, and C. enyo furvus is
dark colored. The darker form occurs on darker soils and in an environ-
ment of higher moisture levels and less extreme diurnal and seasonal
temperature fluctuation. All these factors are in contrast with those in
the habitat of C. enyo enyo.

Little has been recorded of the behavior of C. enyo. Klauber (1931)
states: "Both island specimens (San Francisco and Carmen Islands)
contained mammal remains, and observations on captive specimens and
character of the eye are indicative of largely nocturnal habits. A speci-
men 608 mm. in length collected March 22, contained eggs."

The type was captured by tracking it at noonday to the mouth of a
small rodent burrow where the snake was found resting with part of its

58 San Diego Society of Natural History

coils in the sunlight. When first observed, it remained motionless and
did not rattle. It blended perfectly with the dark soil background and
would not have been seen at all had not its track betrayed its presence.
The snake was not extremely vicious, nor was it as docile as are most
individuals of C. r. ruber upon capture.

A specimen of C. enyo enyo, captured alive 1.6 miles west of San
Ignacio, was very difficult to see lying across the rocky road. Its light
background color, matching effectively, concealed the animal. When this
specimen was shown to Mexicans at San Ignacio, they told us it was
most commonly found in rocky areas. C. enyo, in this more southerly
area, appears to occupy a markedly different microenvironment than
does C. e. furvus to the north. On June 21, 1949, at dusk, the following
temperatures were recorded for the adult specimen from near San
Ignacio: cloacal temperature, 29.7°C; air temperature (1 cm. above the
ground), 27.5°C; soil surface, 32.1°C.

Acknowledgments

Our studies of the herpetofauna of Baja California have been
greatly facilitated by Mr. C. M. Goethe of Sacramento, California, Mr.
Joseph R. Slevin of the California Academy of Sciences, Dr. Laurence
M. Klauber of the San Diego Society of Natural History, and Drs. Ray-
mond B. Cowles and the late Adriaan J. van Rossem of the University
of California, Los Angeles. We are especially indebted to Dr. Cowles
and Mr. Goethe for assistance in our field work conducted in Baja Cali-
fornia during 1949, during which time Crotalus enyo furvus was first
found.

Drs. Howard K. Gloyd, Laurence M. Klauber, and Carl L. Hubbs
have read manuscript and gave many helpful suggestions that have been
incorporated. Fig. 2 was drawn by Mr. Donald B. Sayner of the Uni-
versity of Arizona.

Lowe-Norris — Herpetofauna of Baja California 59

Literature Cited

Beal, C. H.

1948. Reconnaissance of the geology and oil possibilities of Baja
California, Mexico. Geol. Soc. Amer., Memoir Series, No. 31,
x -f 138 pp., 11 pis.

Glovd, H. K.

1940. The rattlesnakes, genera Sistrurus and Crotalus. A study in
zoogeography and evolution. Chicago Acad. Sci., Special Publ.,
No. 4, tables I-XXII, 6 figs., 22 maps, 31 pis.

Klauber, L. M.

1931. Crotalus tigris and Crotalus enyo, two little known rattle-
snakes of the Southwest. Trans. San Diego Soc. Nat. Hist.,
Vol. 6, No. 24, pp. 353-370, 2 figs., 1 map.

1952. Taxonomic studies of the rattlesnakes of mainland Mexico.
Bull. Zool. Soc. San Diego, No. 26:1-143.

Maerz. A., and Paul, M. R.

1930. A dictionary of color. New York, McGraw Hill Co., Inc.

60

San Diego Society of Natural History

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KS-N 7&6> ■ to ■ 9 tt, ,/cs.
California. d<z( "Ylor-ta

tfen-netii S Warns. a~noi
(?har/es N Louje Jr.

Plate 4. Type specimen of Crotalus enyo furvus.

64

San Diego Society of Natural History

Plate 5. Type locality of C. e. furvus, 10.9 miles north of El Rosario, along main road, Baja
California Norte, Mexico. From low coastal terrace, looking inland toward escarpment of higher
terrace, with eroding ledges of Cretaceous bed.

TRANSACTIONS

OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY

Volume XII, No. 5, pp. 67-98, plates 6-7, figs. 1-4

SNAKES OF THE ISLANDS IN THE GULF OF
CALIFORNIA, MEXICO

BY

Frank S. Cliff

Natural History Museum. Stanford University, California

SAN DIEGO, CALIFORNIA

Printed for the Society

October 1, 1954

mi. m?. zi

mitm

DCT 1 8 19!
HARVARD

COMMITTEE ON PUBLICATION

Joshua L. Baily

Carl L. Hubbs

Laurence M. Klauber

Charles C. Haines

SNAKES OF THE ISLANDS IN THE GULF O^
CALIFORNIA, MEXICO

BY

Frank S. Cliff

Natural History Museum. Stanford University, California

mi zm

oci 1 8

In connection with a general study of the herpetofauna and zoogeog-
raphy of the islands in the Gulf of California, several new species and
unrecorded occurrences of snakes have come to light. As an aid to other
workers, it seems advisable to present the new data in advance of the
larger work, and to integrate these data in a revised synopsis of the snakes
of the islands.

Much new material has been collected on the islands and deposited in
the Stanford Natural History Museum through the generosity of Mr.
Joseph W. Sefton, Jr., and the Sefton Foundation of San Diego, Califor-
nia. The Foundation's fine research ship Orca was made available to a
group of biologists, primarily from Stanford University, for two successive
spring trips in the Gulf. The 1952 trip lasted three months and the 1953
trip six weeks. Previous to the Orca expeditions, only two major expedi-
tions had visited the islands, that of the United States Bureau of Fisheries
Steamer Albatross in 1911 and that of the California Academy of Sciences
expedition in 1921. Subsequent to these two expeditions, few specimens
have been taken from the islands, largely the result of incidental collect-
ing by persons engaged in other work.

Twenty-nine species and subspecies of snakes are now known to occur
on the islands (see Table 1). Twelve of the 29 species and subspecies,
or about 40 per cent, are endemic to single islands.

In studies of island populations, the question of what constitutes a
species and what constitutes a subspecies is immediately raised.. Obviously,
there is no completely free exchange of genes or interbreeding between
populations on the islands and the mainland, or between different islands.
Inasmuch as no experimental work has been done, the only criterion for
specific or subspecific recognition that can be used in these instances is
the degree and constancy of morphological differences between popula-
tions. If the island population is widely different from that encountered
on the mainland or adjacent islands and is consistent in these differences,
the insular population is best termed a species. However, if the island
population is only slightly different and its differential characteristics not
entirely constant, it is best known as a subspecies.

The plant associations on the islands are quite similar to those found
on the adjacent mainland. The flora of the islands has been indicated as
only two per cent endemic (Johnston, 1924). Collecting on the islands
is quite different from collecting on the mainland, because, on the islands,
the reptiles occur in much greater densities. This is especially true of the
lizards. The vegetation of most of the islands is denser than that of the
mainland. This ecological factor may be the cause of the higher density
of the reptile population.

More collecting on the islands will undoubtedly disclose new records
and possibly new species of snakes. Crotalus and Hypsiglena probably
occur on all of the larger islands. Burrowing and nocturnal snakes are
also to be expected, especially representatives of the genera Leptotyphlops,

70 San Diego Society of Natural History

Lampropeltis, Chilomeniscus, and Sonora. It is strange that no Pituophis
has been taken on any of the islands.

In the text and on the map (fig. 1), the islands are assigned the most
generally accepted names. Two islands in the Gulf of California are
called Partida. The southern Partida lies just to the north of Espiritu
Santo Island and at low tide is separated from it by a water gap of only
ten feet. These islands are a biological unit and are included in this report
as such, under the name of the larger island, Espiritu Santo. The northern
Partida (Partida del Norte) lies between Angel de la Guarda Island and
Raza Island. San Jose Island is sometimes written as San Josef, but the
former name is now generally accepted. Cerralvo Island appears as Cer-
albo or Ceralvo on some maps, but the majority of modern maps now
designate the island as Cerravlo.

The small islands north of San Luis Bay are named San Luis Island
(or Islands) on many published charts and other maps, but locally, the
name Isla San Luis is applied to the island in San Luis Gonzaga Bay, and
that seems to be the correct usage. The islands north of the bay are
variously named. The group as a whole is known locally as Islas Las
Encantadas, though some persons recognize as a "correct name" Islas
Salvatierra. The largest and southernmost, to which the name San Luis
has been wrongly applied, is more properly known as Isla Encantada
Grande. Just northwest is a much smaller islet, Isla Poma. Ten miles
northwest of Encantada Grande lies Isla El Muerto, the only island on
which snakes have been found. Between Encantada Grande and El
Muerto are Isla Choyudo (or Cholludo) and Isla Coloradito. Northwest
of El Muerto, and very close to the mainland shore, is the smallest island
of the group, Isla Huerfanito. These names seem to be in universal local
use, though they conflict with names applied on the few maps that assign
any name other than San Luis to any of these islands. The conflicting
names applied to these islands are discussed by Klauber (1949, p. 96).
He lists, in addition to the islands mentioned here, an island between
Encantada Grande and El Muerto. Dr. Carl L. Hubbs, Scripps Institution
of Oceanography, who has supplied data on the proper names of these
islands, informs me that there is no island between La Encantada and
Porno and consequently there are only six islands in the group, rather
than seven as listed by Klauber.

Abbreviations have been used to designate the museums where speci-
mens are deposited, as follows: SU, Natural History Museum, Stanford
University; CAS, California Academy of Sciences; LMK, private collection
of L. M. Klauber; MVZ, Museum of Vertebrate Zoology at the University
of California, Berkeley; MCZ, Museum of Comparative Zoology at Har-
vard University; USNM, United States National Museum.

Lichanura roseofusca gracia Klauber, 1931
A single dried example of this subspecies is known from Mejia Island.
This single example (CAS 50804) was found under a rock by J. R. Slevin
on June 28, 1921.

Scale counts are impossible to make with certainty because of the dried
and brittle condition of the specimen. Ventral and caudal counts are
likewise impossible to ascertain, since the snake is broken into several
pieces and portions are missing. I am assigning this specimen to the
subspecies gracia because the longitudinal stripe is present, it has even
edges, and it is red brown. On geographical grounds, also, this subspecies

Cliff — Snakes of the Gulf Islands 71

would be expected on a northern island, as it occurs on the adjacent main-
land of Baja California.

Chilomeniscus ductus Cope, 1861

A single male (SU 14035) referable to this species was collected on
San Jose Island near Punta Salina on April 11, 1952. It constitutes a new
insular record.

It is a young specimen only 152 mm. from snout to vent, tail length
27 mm. Ventrals number 109, caudals 25, supralabials 7, and infralabials
8.

The head scales are typical of mainland examples of the species and
consist of: a rostral twice as wide as high, paired nasal-internasals which
are separated medially by the posterior apex of the rostral, paired pre-
frontals which are in contact medially, and a shield-shaped frontal which
is bordered by 2 parietals posteriorly. On either side there is 1 preocular,
2 postoculars and no loreal. The third and fourth supralabials contact the
eye.

In coloration, this example is similar to the endemic species from
Espiritu Santo Island, Chilomeniscus punctatissimus. The 32 brown bands
are slightly wider than the interspaces. The interspaces contain a series
of dots; these dots are about half a dorsal scale in size and are of the
same color as the bands. The head scales anterior to the first body band
are gray-olive and are suffused with light brown. None of the bands
encircle the body; the majority barely touch the ventrals. Five of the
bands are on the tail. However, in both scjuamation and coloration, this
example falls well within the limits of variation for C. ductus.

Figure 2. Outline Drawing of the Head of Chilomeniscus savagei. Holo-
type (SU 14034).

Holotype (SU 14034). Collected under debris on the sand dunes on
the southwest coast of Cerralvo Island by Jay M. Savage, April 3, 1952.
The snake is named for the collector whose numerous suggestions and
thorough knowledge of the reptile and amphibian fauna of Baja Cali-
fornia have aided me on numerous occasions.

Diagnosis : This distinctive insular species differs from all other species
of the genus in having a higher number of ventrals and unique color
pattern, and from all species except C. punctatissimus from Espiritu Santo
(Partida portion), in having the prefrontal reduced in size and the frontal
contacting the fused nasal-internasal.

Description of type: A mature female, 234 mm. in length from snout
to anus, length of tail 34 mm., head length 10.3 mm., head width 6.0

72 San Diego Society of Natural History

mm., greatest width of body 8.0 mm., greatest eye diameter, 1.1 mm.

Head not distinct from body; short tail tapering to a point. The upper
jaw protrudes and the rostral is prominent. The pupil is round. The
scales are smooth, with no apical pits; they are arranged in 15-13-13 rows,
with 6 scale rows at the middle of the tail length. The ventrals number
134 and the paired caudals 27. The anal plate is divided. There are 7-7
supralabials and 8-8 infralabials. The supralabials increase in size posteri-
orly; the last is the largest; the third and fourth contact the eye.

The rostral is about twice as wide as high and extends posteriorly to
a blunt angle partly separating the nasal-internasal. The inferior surface
of the rostral has a central depression near its posterior margin.

The scales on the crown comprise paired nasal-internasals, paired pre-
frontals, paired supraoculars, paired parietals, and a single median frontal.
The nasal-internasals meet medially and meet the first supralabial, pre-
frontal, and rostral broadly; they barely touch the second supralabial. The
valvular nostril is pierced near the lateral edge of the nasal-internasal.
The prefrontals are widely separated medially by the forward extension
of the frontal and also meet the preocular, supraocular, second supralabial,
and nasal-internasal. The frontal is shield-shaped and is bordered posteri-
orly by the 2 large parietals. There are two postoculars, the upper of
which is the larger. The temporals are 1-2. There is no loreal.

There are two pairs of chin shields. The anterior comes to a point
medially, the posterior to a point laterally. The anterior is slightly larger.
The first supralabial extends laterally to separate the mental completely
from the chin shield. The anterior chin shield is bordered laterally by the
first three infralabials. Four scales separate the posterior chin shields from
the most anterior ventral.

Coloration : The very simple pattern consists of 29 body rings of a
light brown color about 3 scale rows wide. The light interspaces are
narrower, about 2 scale rows wide. The most anterior dark ring extends
from the edge of the second preocular on each side to the posterior edge
of the parietals. The posteriormost ring surrounds the end of the tail.
The dark body rings are continued onto the ventral surface, where all but
a few of the anterior merge into one another. Five of the rings are on
the tail. In life, there is a pinkish-orange suffusion in the flesh-white
interspaces, which includes the vertebral scale row in addition to from 2 to
3V2 scales on each side. This suffusion is bleached out in preservative.
The snout is white. In life, the venter between the brown rings is a
delicate pink, but in the preserved state this color disappears and the
venter is an immaculate flesh color. The throat is slightly suffused with
brown. The rostral, nasal-internasal and anterior portions of the pre-
frontals and frontal are white in life, but turn yellowish in preservative.

The paratype (SU 16062) is an adult female with an incomplete tail,
collected by John P. Figg-Hoblyn in the sand dunes near the place where
the holotype was collected, on Cerralvo Island, but a year later, on March
20, 1953. The paratype has 136 ventrals, 7 supralabials, 8 infralabials
and has the same details of head squamation as the type. It differs in
that none of the body blotches encircle the body. Most, however, extend
onto the ventrals. The first and only blotch on the tail completely encircles
it.

Cliff — Snakes of the Gulf Islands 73

Relationships: In color pattern, this insular species resembles Chilo-
meniscus ductus from Arizona, Sonora, and the mid-peninsula region of
Baja California. It shares its distinctive arrangement of head scales only
with the species from Espiritu Santo Island. In addition, the Cerralvo
specimens have a higher number of ventrals than any other species. It is
problematical whether the two species on Cerralvo and Espiritu Santo
islands are insular relics of an older stock of Chilomeniscus, or whether
the evolution of these species has been effected solely through isolation
in an insular environment. It does seem evident, however, that the two
insular species are closely related and, since Cerralvo harbors many dis-
tinctive species, it would appear likely that Espiritu Santo Island has
derived its species of Chilomeniscus from Cerralvo.

Chilomeniscus punctatissimus Van Denburgh and Slevin, 1921

This insular species, based on only one specimen from Partida, Espiritu
Santo Island, was placed in the synonymy of Chilomeniscus ductus by
Smith (1945, p. 34). Inspection of the type and comparison with main-
land and insular examples of the genus warrant the specific recognition
of this single example.

The holotype of the species is CAS 49516, an adult female collected
on Isla Partida by Chamberlin, May 31, 1921.

The most important characters separating this species from C. cinctus
lie in the arrangement of the scales on top of the head. As in Chilo-
meniscus savagei, the prefrontals are reduced in size, allowing the frontal
to meet the fused nasal-internasal. On each side, a small scale is inter-
posed laterally to the prefrontal and above the first supraocular, which
forms the posterior border of the nasal opening, but I believe this to be
of very little taxonomic importance.

Van Denburgh and Slevin recognized this species primarily on the
basis of the spotting that occurs on the light interspaces between the body
blotches — a character of little importance in this genus, as pointed out by
Linsdale (1932, p. 382).

A brief redescription of the species seems desirable, with particular
emphasis on head squamation. The scales on the top of the head comprise
paired nasal-internasals, prefrontals and supraoculars, a single rostral, and
a frontal. The rostral is about twice as wide as high and extends far
enough posteriorly almost to interrupt the contact of the internasals. The
prefrontals are prevented from touching one another medially by the
forward extension of the frontal, which extends anteriorly to meet the
internasal. The small single preocular touches the small postnasal; there
is no loreal. There are 2 postoculars. The supralabials number 7-7, the
infralabials 8-8. The third and fourth supralabials border the eye. The
dorsal body scales are in 13 rows; they are smooth with no apical pits.
The anal plate is divided. There are 120 ventrals and 23 caudals.

Coloration: There are 39 dark brown bands on the body, including
both the band on the head that extends from the anterior margin of the
eye to the middle of the last supralabial and the dark tip of the tail. The
interspaces are slightly wider than the bands. Within the light interspaces
there are from 10 to 15 light tan spots which may occur on any of the
dorsal scale rows except the 2 lateral rows on each side. The spots are

74 San Diego Society of Natural History

most intense near the vertebral scale row and are about half as large as
the dorsal scales.

Relationships : As discussed under C. savagei, this species is considered
to be closely related to the Cerralvo species and possibly derived from it.
Hypsiglena torquata gularis Tanner, 1954

The two females of this genus from Partida Island have recently been
considered to represent a distinct insular subspecies. The type is CAS
51009, and the paratype 51010. Ventrals number 185-183, caudals 43-
43, scale rows at midbody 17-19, and body blotches 68-68.

This subspecies is most closely related to H. t. venusta of the mid-
peninsula, from which it is said to differ in having larger dorsal blotches,
one or more pairs of enlarged gulars, fewer caudals, and a proportionately
shorter tail.

Hypsiglena torquata ochrorhyncha Cope, I860

This subspecies, which occurs in the Cape region of Baja California,
is found on San Jose Island. Four specimens are known from this island:
CAS 57398 and 57392, and LMK 3816 and 3815. The first three are
males. Ventrals number 176-1 78-168-1 76,1 caudals 46-50-50-47, scale
rows at midbody 17-17-17-17 and body blotches 52-50-65-66.

This subspecies is distinguished from H. t. venusta in having fewer
and correspondingly larger dorsal blotches and in having fewer ventral
and caudal plates. The body blotches are undivided dorsally, and there
is no median light stripe.

Examination of more specimens of this subspecies will probably lead
to the distinguishing of a Cape subspecies from the populations in Arizona
and Sonora. A trend is evident in ventral counts, but no characters that
would justify nomenclatorial separation have as yet been discovered.

1Counts listed in same order as register numbers of the specimens.
Hypsiglena torquata tortugaensis W. Tanner, 1944

This subspecies is known from only two male specimens from Tortuga
Island. The type is CAS 51460 and paratype LMK 4074.

The insular subspecies may be separated from the mainland forms,
ochrorhyncha and venusta, by a higher number of ventrals and caudals,
as shown in the following table:

Male

Male

ventrals

caudals

ochrorhyncha

166-188

46-66

venusta

176-184

54-56

tortugaensis

187-190

57-59

The other distinguishing characteristics of this insular subspecies are
an elongate narrow median stripe on the neck and divided lateral neck
stripes.

Tanner (1944, p. 70) believes that this insular subspecies is more
closely related to venusta than to ochrorhyncha.

Hypsiglena torquata venusta Mocquard, 1899

The mid-peninsula subspecies of Hypsiglena is known from the
following Gulf of California islands: Carmen, San Marcos, San Esteban,

Cliff — Snakes of the Gulf Islands 75

and San Francisco. The specimens from the last two islands constitute
new insular records.

This subspecies is distinguished from ochrorhyncha in having a higher
number of dorsal blotches, in having these blotches of small size, and in
having the blotches alternating or divided to produce two rows instead
of one, with a resultant light median dorsal stripe narrower than one scale
in width.

The single female from San Esteban Island (SU 16066) has 179
ventrals, 29 caudals (tail incomplete), 21 scale rows at midbody, and 74
body blotches. On the posterior quarter of the body, the blotches coalesce.

The two females from San Francisco Island (SU 16067 and 14039)
have 183-180 ventrals, 46-44 caudals, 21-21 scale rows at midbody, and
76-81 body blotches.

There are also two females from Carmen Island (CAS 51814, and
MCZ 31583). The counts are as follows: ventrals 174-182, caudals
50-51, 19-19 scale rows at midbody, and 95-95 body blotches.

The male from San Marcos (CAS 51462) has 184 ventrals, 54 cau-
dals, 19 scale rows at midbody, and 74 body blotches.

Lampropeltis catalinensis Van Denburgh and Slevin, 1921

This beautiful and distinctive insular species is known from a single
adult male (CAS 50514) which was dug from the center of a decaying
cactus stump on Santa Catalina Island. Although a thorough search for
this species was made on both the occasions on which the author was on
this island, no additional specimens were collected. A shed skin was
found near a small spring which may have been from a Lampropeltis,
but no live animals were seen or captured.

Since Van Denburgh (1922, p. 769) has presented a detailed account,
only a short description will be presented here.

Ventrals number 228, divided caudals 63, scale rows at midbody 23,
supralabials 8, infralabials 9. Color and color pattern are the distinguish-
ing peculiarities. A purple longitudinal dorsal band, about 5 scale rows
wide, extends from the head to the end of the tail. Along this mid-dorsal
line, at intervals of 3 to 4 scales, are light yellow spots on a single scale.
The lateral dorsal scales are purple with light yellow centers. The head
above has the same color as the dorsal band and is marked with light
yellow suffusions on the crown and sides. The ventrals and caudals are
black, with extensive white marbling in an irregular pattern. The under
surface of the head is light yellow, with the chin shields, gulars, and first
several ventrals edged with black or purple.

The only other Lampropeltis with a color pattern near that of cata-
linensis is L. nitida from the Cape region of Baja California, which has
a poorly denned light mid-dorsal stripe, no uniform spotting laterally,
and the ventral surface uniformly stippled with brown. Van Denburgh
(loc. cit.) states that the snake was canary yellow, purple, and black
before preservation.

Masticophis barbouri (Van Denburgh and Slevin), 1921
The second and third examples of this insular species were collected
on Espiritu Santo Island. SU 14040 was collected by John P. Figg-Hoblyn
on April 5, 1952. It is an adult male, 872 mm. from snout to tip of tail,

76 San Diego Society of Natural History

head length 24 mm., tail length 176 mm. (incomplete). The dorsal scale
row formula is 19-17-13. There are 194 ventrals and 68 caudals (in-
complete). There are 8 supralabials and 9 infralabials. SU 16068 was
collected by Figg-Hoblyn on March 23, 1953. It is also an adult male,
701 mm. from snout to tip of tail, head length 21 mm., tail length
226 mm. The dorsal scale row formula is 19-17-13. There are 195
ventrals, 141 caudals (tail presumably complete), 8 supralabials, and 9
infralabials.

In details of coloration, these specimens agree closely with the type,
which is an adult female larger than either of the two recently acquired
specimens. The color above is a uniform olive-brown extending onto the
ventrals. Along each side, a single white stripe is located on the medial
half of the fourth scale row and on the lateral half of the third scale row,
and fades out at the anal plate. This stripe is edged narrowly with black.
Anteriorly, this stripe is dilated at intervals of 4 or more scales. The
sides of the head are spotted with white; there are two spots posterior to
the eye, and a white line extends anteriorly from the eye to the tip of the
snout. The upper quarter of the supralabials is blackish-brown. The
remainder of the supralabials and the infralabials, chin, and neck are
white, lightly flecked with darker spots, which appear blue under the
microscope. The anteriormost ventrals are suffused with salmon and bear
a few spots of blue. The remainder of the ventral surface is an immacu-
late white.

This species is closely related to Masticophis lateralis and also to M.
aurigulus, but I disagree with Ortenburger (1928, p. 68), who assumes
it to be an annectant between lateralis and aurigulus. I consider it to be
a distinct species derived from lateralis by isolation in an insular environ-
ment.

Masticophis bilineatus Jan, 1863

This species, previously unknown from the Gulf of California, was
found on two islands, Tiburon and San Esteban.

Hensley (1940, p. 272) described a subspecies, Masticophis bilineatus
I'meolatus, from 4 specimens collected in the north branch of Alamo
Canyon, Ajo Mountains, 12.9 miles south and 5 miles east of the Ajo-
Tucson-Sonoyta Junction in Organ Pipe Cactus National Monument, Pima
County, Arizona, May 23, 1949. He used four characteristics to differ-
entiate his race from the common Arizona form:

Masticophis bilineatus bilineatus: (1) Dorsolateral light stripe com-
prises the upper half of the third and lower half of the fourth scale row.
(2) Dorsolateral light stripe originates on fourth scale row posterior to
last supralabial. (3) Chin spotting is very light or usually not evident.
(4) The pattern of dots on the edge of the ventrals is not evident enough
to warrant calling them a definite stripe.

M. b. lineolatus: (1) Dorsolateral light stripe is narrower, compris-
ing only the upper quarter of third scale row and lower quarter of fourth
scale row. (2) Dorsolateral light line originates 7.75 scale rows posterior
to the last supralabial. (3) Chin spotting is pronounced and blotches
occur in the neck region and on the anterior portion of the ventrals.
(4) There is a definite dark stripe on the lateral edges of the ventrals.

Cliff — Snakfs of the Gulf Islands

77

The four examples from Isla San Esteban (SU 14041-43 and 16069)
and Tiburon (SU 14044) are like Hensley's subspecies in these four
characteristics. An examination shows coastal Sonora material to be inter-
mediate in these characteristics. I hesitate to refer these insular examples
to Hensley's subspecies until further specimens are collected and a more
substantial geographical distribution pattern is outlined.

Data on the insular specimens examined are as follows:
San Esteban:

208 ventrals, 139 caudals, dorsolateral line originates

on 7th scale posterior to
last supralabial.
204 137 ... on 7th scale

206 142 ... on 6th scale

197 incomplete ... on 6th scale

14041 (male)

14042 (female)

14043 (female)
16069 (female)
Tiburon :

14044 (male)

206

139

on 7th scale

Masticophis flagelhim pic ens (Cope), 1892

This subspecies is now known to occur on eight islands in the Gulf
of California: Cerralvo, Espiritu Santo, San Jose, Monserrate, Carmen,
Coronado, Ildefonso, and Tiburon. The three examples of this race (SU
14062-64) collected on Cerralvo constitute a new insular record.

The three specimens from Cerralvo are very dark brown; almost black
dorsally, with no white flecks or dots. The top of the head is somewhat
lighter brown. The supralabials and lateral scales anterior to the eye are
of a lighter olive-brown and have suffusions of cream. The infralabials
have small patches of white. The dark dorsal coloration extends to the
edge of the ventrals, which are cream colored, except in the neck and chin
regions which are heavily suffused with the dorsal ground color.

Klauber (1942, p. 93) states that the three specimens from Espiritu
Santo are "dark brown to black with light longitudinal dashes. One is
as dark as any from Tucson."

Five specimens from Isla San Jose were available for comparison (SU
14046-48 and CAS 52548-49). The dorsal ground color is yellow. There
is a dark gray suffusion on the neck posterior to the parietals, which
extends to the 1 2th— 1 5th ventral and contains 4-5 irregular narrow light
bands. The anterior third to half of the body length has dark punctations
on the dorsal scale tips (or on their edges) which appear as wavy thin
bands. There are gray-black blotchings of no definite pattern on the labials
and under the throat. Anteriorly there are four rows of spots on the
ventrals. The paired medial rows of these spots are regular and are found
on each ventral on the anterior third of the body, fading out posteriorly.
The lateral rows of spots are larger and extend farther back, but occa-
sionally skip a ventral.

Five specimens from Monserrate Island were available for study (CAS
52178-52182). The dorsal ground color is dark brown, becoming lighter
toward the tail. The side of the head is blotched with cream and the
crown is olive-brown. There are numerous yellow markings on the edges
of the scales posteriorly. The ventral surface varies from yellow to cream.
A very irregular series of brown blotches on the ventrals extends posteri-

78 San Diego Society of Natural History

orly for about half the body length. The neck and chin region are mottled
with extensive suffusions of gray brown. CAS 52180, a young specimen,
is lighter than the rest and has a more regular pattern.

Klauber (1942, p. 93) states that the one known example from
Carmen is "a yellowish specimen with punctated head."

There is but one known example of this race from the tiny island of
Ildefonso (CAS 51717). It is colored very similarly to Monserrate
examples, except that the ventral surface is very heavily suffused with
brown.

Thirteen specimens were collected on Isla Coronados in four hours,
on a cloudy day (SU 14049-61). They were seen in the branches of
bushes and lying motionless out in the open. These snakes are of the
dark calor phase. All thirteen are very similar in color pattern, but the
young are somewhat the lighter. There is a narrow white edging on some
of the lateral dorsal scales. The ventral surface is cream, suffused with
large gray spots in the neck and chin region. The spotting changes from
large spots anteriorly to scattered flecks posteriorly. SU 14056 is almost
as dark as specimens from Cerralvo.

Klauber (1942, p. 93) states that on Tiburon Island both the light
and dark phases of the whip snake are found. CAS 53245 from this
island is extremely melanistic. There are no light marks on the labials or
on the sides of the head and the ventral surface is suffused with dark as
far as the anal plate. Even the under surface of the tail is tinged with
dark. The dorsal coloration gradually lightens posteriorly and the tail is
brown with darker suffusions on part of almost every scale.

In ventral and caudal counts, and in head scales, all insular specimens
fall within the limits of variation ascribed to the wide-ranging mainland
population. It is interesting to note that on each of the islands (disregard-
ing Carmen and Ildefonso, from each of which we have only one speci-
men) a particular color phase has become established. Tiburon, a large
coastal island, is the only exception. Since intermediates are found stabil-
ized on islands, it seems possible that color is dependent upon stabilization
of not only one pair of genes, but also upon one or more pairs of modify-
ing genes as well.

Phyllorhynchus arenicola Savage and Cliff, 1954
This interesting species is endemic to Monserrate Island. It is a
member of the decurtatus section of the genus, readily distinguished from
all other forms of Phyllorhynchus by the following combination of char-
acteristics: (1) Dorsal body blotches 30-32 in number, at least twice as
wide as light interspaces when measured longitudinally near midbody; at
least one-fourth of these blotches partially split by a light medial area.
(2) Four to 6 lateral scale rows suffused with brown; this suffusion not
extending onto back between dorsal blotches. (3) Lateral spots large,
distinct. (4) Ventrals 164 in the male, 172 in the female. (5) Caudals
39 in the male, 31 in the female.

Salvadora hexalepis hexalepis (Cope), 1867
Two specimens of this subspecies are recorded from Tiburon Island.
Although the island lies near the proposed line of intergradation between

Cliff — Snakes of thf Gulf Islands 79

hexalepis and deserticola, Bogcrt (1945, p. 13, fig. 10) places these two
examples as members of the typical subspecies.

Salvadora hexalepis klauberi Bogert, 1945

The first known example of this subspecies to be reported on any
Gulf island was collected on San Jose Island by Jon Lindbergh on April
11, 1952 (SU 14016). It is an adult male with 197 ventrals and 96
caudals. This snake seems referable to S. h. klauberi on the basis of the
fifth and sixth supralabials reaching the eye, the two loreals, and the
ventral and caudal counts.

Our specimen agrees with typical klauberi in coloration except that
the lower lateral stripe is not regularly present on the fourth scale row
anteriorly (well demarcated on third and fourth scale rows in peninsular
examples) and the belly is yellow (usually white or only tinged with
yellow in peninsula examples).

Mr. Charles M. Bogert has examined the specimen and states (per-
sonal communication) : "Except for the abnormal, small triangular scale
wedged in between the last and penultimate supralabials on each side,
and an extra labial below the loreals on the right side, the specimen is
a rather typical S. h. klauberi. These extra scales in the labial series . . .
have no taxonomic importance."

Sonora mosaueri Stickel, 1938

An adult male of this species (SU 14038) was removed from the
stomach of a Masticophis flagellum piceus collected on the flats of the
southwest corner of San Jose Island. This snake is the first of the genus
to be reported from any island in the Gulf of California. The specimen
was only slightly digested and it was still possible to ascertain all charac-
teristics necessary for specific placement. The combined ventral and caudal
counts exceed by one the previous upper limit of the range of the species.
The scale rows are 15-14-13, ventrals 156, and caudals 49.

In addition to the insular example recorded above, one other specimen
(SU 14019) of S. mosaueri was collected at Bahia San Carlos on the
mainland. Bahia San Carlos2 is on the Gulf side of the peninsula and
lies about 3 miles south of Punta San Telmo at about 25° 30' N. latitude
and 111° 1' W. longitude. This snake is an adult male with scale rows
15-14—13, 150 ventrals, and 47 caudals. This species has previously
been known only from the vicinity of Comondii, and this example extends
the range approximately 100 miles southward.

The coloration in both examples agrees with that given by Stickel
(1938, p. 189). The dorsal color is slightly darker brown in the main-
land snake and the terminal spots on each scale are less prominent.

The counts on both these examples further support the distinctiveness
of S. mosaueri, which is separated from all other members of the genus
by the number of scale rows (15-14-13), of ventrals (males, 150-156;
female, 164), and of caudals (males, 43-49; female, 39), and by the
uniform coloration. Additional material may be expected from the main-
land as far south as the coast opposite San Jose Island.

2There is another Bahia San Carlos at about 27° 48' N. latitude.

80 San Diego Society of Natural History

Micruroides euryxanthus (Kennicott), I860
A single example of this Arizona and Sonora elapid was recorded by
Streets (1877, p. 20) from Tiburon Island. This specimen is now USNM
8566.

Dr. Doris M. Cochran of the National Museum has kindly examined
the specimen and has forwarded the information presented here. Ventrals
241, caudals 30, supralabials 7, and infralabials 6. The specimen is very
faded and its color pattern now consists of black, white and gray rings.
In life, the white rings were yellow and the gray rings were red. The
black rings are the widest, the white rings about one quarter as wide,
and the gray rings about one-half the width of the black rings.

Cro talus atrox Baird and Girard, 1853

Previous to this report, C. atrox was known from only one Gulf
island, the coastal island of Tiburon. The specimens from that island, as
might be expected, are identical with those from the mainland of Sonora.
In addition, one specimen referable to this species and constituting a new
insular record was captured on San Pedro Martir Island (SU 14029) on
May 4, 1952, by Dr. Reid Moran.

It is a gravid female having 185 ventrals, 22 caudals, 25 scale rows
at midbody, and 5 scale rows between the supraoculars. There are 17-16
infralabials and 14-15 supralabials. The postnasal meets the upper
preocular and the prenasal does not contact the supralabial. In coloration,
especially in punctations, it is similar to atrox in Sonora, except that the
posterior dorsal body blotches (the last eight) are extremely faint and
only discernable when the specimen is held under water. These dorsal
blotches number 34.

In all diagnostic characteristics, the snake seems to fit well within
the limits of variation ascribed to C. atrox by Klauber (1952, table 1)
except in the relationship of the head dimensions to the standard body
length. This relationship indicates dwarfing. This specimen has a body
length of 830 mm., a head length of 30 mm., and a head width of 22 mm.
Following Klauber in his statistical studies on Crotalits, we find that for
an atrox 830 mm. long we should expect a head length of 38 mm. This
insular example thus falls well outside the 99 per cent confidence limits
for head length as indicated by Klauber (1938, p. 24). If more speci-
mens are collected from San Pedro Martir Island, this insular atrox may
well prove to be a dwarfed race.

Crolalus catalinensis, new species

Holotype (SU 15631). (Fig. 3.) Collected on Isla Santa Catalina in
the Gulf of California, Mexico, by Bruce Firstman, Dr. John C. Briggs,
and the author, on March 27, 1953, as it lay coiled under the shade of a
large cactus.

Diagnosis: This species is allied to the Crotalits atrox group, but
differs from others of that group in being a stunted form, with a distinc-
tive color pattern and oddly-shaped internasals. It can be distinguished
from C. atrox by the excessively large postcanthal, which extends laterally
to separate the preocular from the postnasal, from C. ruber ruber and
C. ruber lucasensis in not having the first infralabials divided transversely,
and from C. tortugensis both in having the dorsal blotches surrounded by

Cliff — Snakes of thf Gulf Islands 81

well defined light scales and in having fewer punctations on the dorsal
ground color.

Description of holotype: A sexually mature female. Over-all length
701 mm., length of tail 46 mm., head length 29 mm., head width 22 mm.,
width of proximal rattle 6.0 mm.

The scale rows number 27-25-21, with 13 rows at the center of the
tail and 11 scales bordering the rattle. The ventrals number 182 and the
subcaudals 20. The anal is entire. There are 13-15 supralabials and
14-14 infralabials. The second, third, and fourth supralabials are con-
siderably reduced in size, and the fifth is slightly larger than any of the
other supralabials. The rostral is very slightly higher than wide. The
scales on the crown of the head comprise two irregularly kidney-shaped
internasals that are about three times as long as wide. The anterior
canthals are small and fit into the concavity of the kidney-shaped inter-
nasals. They meet medially and are longer than wide. The posterior
canthals (prefrontals) are separated medially by two pairs of small scales
and are roughly triangular in shape, being rounded anteriorly at the apex.
The supraoculars are the largest of the head scales and are bordered
medially by nine small scales posterior to the posterior canthal (pre-
frontal). The prenasals are larger than the postnasals, higher than long
and in broad contact with the first supralabial on both sides. There is but
one loreal on each side, but the posterior canthal extends laterally, pre-
venting contact between postnasal and upper preocular. One small scale
separates the postnasal from the supralabials. The upper preocular is the
larger; the lower preocular is crescentic. Both preoculars contact with the
loreal. Nine scales border the eye, including the supraocular and both
preoculars. The triangular mental is longer than wide; the first infralabials
are divided; the genials are long and pointed posteriorly. There are
neither intergenials nor submentals. The second and third infralabials
are noticeably longer, but not wider than the remaining infralabials.

The head is light gray-brown, punctated with brown above, with a
faint light streak through the center of the supraoculars. The ocular light
stripe extends from the third supralabial (anterior to the commissure) to
the anterior edge of the eye and is not sharply defined. The sides of the
head are of the same color as the crown, with the exception that the scales
posterior to the eye are more heavily punctated with brown.

The dorsal ground color approximates that of the head. The 38 dorsal
body blotches are medium brown in their centers and are bordered by a
single row of scales of darker -brown. These darker scales at the margin
of the blotch are bordered by 2 rows of cream-colored scales that separate
successive blotches. Both the dark and light scale margins become indis-
tinct on the posterior quarter of the body. Anteriorly the blotches are
elongate. In the center of the body, they are of a rounded diamond shape
and extend farther laterally. The bordering scales lose their darker hue
posteriorly. The blotches are about 7 scale rows wide and 4 scales long
in the center of the body. Below the dorsal blotches on either side there
is a row of smaller blotches. One of these smaller blotches (slightly the
larger) falls below the lateral point of each dorsal blotch, the other
between each pair of dorsal blotches. The posterior 9 dorsal blotches
coalesce with the larger lateral blotches.

82 San Diego Society of Natural History

The venter is buff, lightly suffused with gray laterally. The chin and
neck region are immaculate. Dorsally, the tail has 5 black rings, which
are 4 to 5 scale rows wide medially and taper to a point laterally, except
for the posteriormost ring. This is only one and one-half scale rows wide
and borders the rattle. The interspaces between the rings are slightly
narrower than the dark rings, at least medially; are somewhat darker
than the ground color of the body, and are more heavily punctated with
brown. The under surface of the tail is darker and more heavily punctated
than the venter and in its center has a suffusion of brown-black puncta-
tions which forms an indefinite ventral medial stripe. The anterior rattle
matrix is black. Only the button is present, no rattles have developed.

Relationship: One would expect this new rattlesnake to be closely
related to either C. ruber ruber or to C. ruber lucasensis, simply because
it occurs on Santa Catalina Island which is nearly opposite the intergrading
area of these two subspecies on the mainland of Lower California. If
color pattern alone were considered, the new snake would appear most
closely related to C. r. lucasensis. In having the canthal separating the
preocular and postnasal, it is typical of C. ruber ruber. In having the
first infralabial undivided, it is characteristic of neither species. However,
it seems likely that this snake is most closely related to C. ruber, and has
evolved its differences in response to insular isolation.

The ovaries of this specimen were examined by Dr. Malcolm Miller
of the Department of Anatomy of Stanford Medical School who reports
that the animal did not ovulate this year, but was sexually mature. Since
the specimen is only 701 mm. long, this indicates that the animal is
dwarfed.

Crotalus enyo cerralvensis, new subspecies
Holotype (SU 14021). (Fig. 4.) Collected on Isla Cerralvo in the
Gulf of California on April 3, 1952, by John Figg-Hoblyn, Jay M.
Savage, and the author, as it lay coiled in the shade of a bush in the
dunes on the southwest coast.

Paratype (CAS 84834). Collected by Dr. G. Dallas Hanna, March
22, 1953. It was buried in a talus slope near the remains of the old
Ruffo Ranch on the western coast of Cerralvo Island.

Diagnosis: This subspecies is closely related to, and obviously derived
from, the mainland Crotalus enyo. from which it can be distinguished in
having a higher number of ventral and caudal scales and a proportionately
smaller head.

Description of holotype: A mature male, over-all length 765 mm.,
tail length 72 mm., head length 26.5 mm., head width 17 mm., width of
proximal rattle 9.5 mm.

The scale rows are 27-23-19, with 11 rows at the center of the
tail; 10 scales border the rattle. The ventrals number 167 and the sub-
caudals 31; the anal plate is entire. There are 13-13 supralabials and
12-14 infralabials. The fourth supralabial is the only noticeably enlarged
labial. The rostral is equal in height and width. The scales on the crown
are-ridged and knobby and include two rounded triangular-shaped inter-
nasals. Posterior to the internasals, a single round canthal meets the

Cliff — Snakes of the Gulf Islands 83

supraocular. The supraoculars are the largest scales on the top of the
head. The prenasals are larger than the postnasals, square in form, and
contact the first supralabial on either side. There are three loreals on
either side. The most dorsal loreal lies caudad to the remaining loreals,
which separate the preoculars from the prenasal, but does not interfere
with the contact of the canthal with the supraocular. The upper 2 loreals
are smaller than the lower loreal. Two small scales on either side separate
the postnasal from the first supralabial. The upper preocular is larger
than the lower preocular. The lower preocular contacts only the lower
loreal, but the upper preocular meets all three loreals. Eight scales border
the orbit. The triangular mental is slightly wider than long. The first
infralabials are undivided. The 2 genials are touched by 3 infralabials
on one side and by four on the other, and converge toward each other to
form a point medially. There are neither intergenials nor submentals.
All dorsal body scales are keeled except for the rows bordering the
ventrals.

The ground color of the head is a heavily punctate gray above. From
the first dorsal body blotch arise two brown diverging fingers that extend
to the posterior border of the postocular. The posterior half of the supra-
ocular is olive; the anterior half is light gray bordered by a narrow stripe
of dark brown. The postocular light stripe extends from the upper corner
of the eye, passing above the commissure to fuse with the light ground
color below the first body blotch. This stripe is limited above by the finger
of the first dorsal blotch and below by the gray-brown color that begins
at the angle of the jaws and extends forward to include the upper labials;
it continues beneath the eye to the posterior edge of the sensory pit. The
preoculars are less punctate and appear to be of approximately the same
color as the' postocular light stripe.

The body pattern consists of 34 dorsal blotches, of which the first 10
are rectangular with light areas in their centers, and the remainder are
rounded diamonds without light centers. A secondary series of black
spots appears laterally on each side. The dorsal blotches are outlined in
black, and surrounded by a light area about one to two scale rows wide.
This light area extends to the lateral apex and diverges around the
secondary series of blotches that are opposite the dorsal blotches. The
lateral scales enclosed by the light scales are heavily punctated and give
the appearance of another lateral series of blotches. Posteriorly, the
secondary series of spots merges with the dorsal blotches.

The ventral surface is buff, heavily dotted with gray and brown. The
chin and neck region is less spotted. The lower labials are heavily suffused
with gray.

The dorsal surface of the tail has six indistinct light brown tail rings.
The ventral surface is punctated like that of the venter of the body. The
anterior rattle matrix is black; the rattle string is incomplete.

Paratype: The single paratype is a female, over-all length 681 mm.,
tail length 44 mm., head length 21 mm., head width 16 mm. It has 25
scale rows at midbody, 181 ventrals and 23 subcaudals.

In coloration, it is quite similar to the type, being slightly darker in
ground color. Unfortunately, the head was injured in its capture and the

Mainland

Cerralvo

157-168

167

164-177

181

22-28

31

18-23

23

22.5-25.8

28.9-32.5

84 San Diego Society of Natural History

details of head squamation are difficult to ascertain. It does differ from
the type in having only 2 loreals.

Comparisons: This island subspecies differs as follows from Crotalus
enyo enyo:

Ventrals, males

Ventrals, females

Caudals, males

Caudals, females

Ratio of head to body length

Crotalus enyo enyo Cope, 1861

This beautiful rattlesnake is known from the following islands in the
Gulf of California: Espiritu Santo, including Partida portion; San Fran-
cisco, and Carmen.

The Espiritu Santo examples were taken on successive years. SU 14022
is a male and has 23 dorsal scale rows at midbody, 160 ventrals, and 26
caudals. The total length is 751 mm., the head length 28 mm. SU 15630
is also from Espiritu Santo and is a female with 23 dorsal scale rows at
midbody, 167 ventrals, and 21 caudals. The total length is 581 mm., the
head length 25 mm.

Our single San Francisco Island example (SU 15629) is an immature
female only 349 mm. in total length with a head length of 19 mm. It
has 23 scale rows at midbody, 166 ventrals, and 21 caudals. There are
13 supra- and 13 infralabials.

Two specimens were collected near a deserted ranch building on
Carmen Island. The larger (SU 16532) was collected by the author,
under a piece of tin roofing that was exposed to the sun. The heat under
the tin roofing was extreme and it was surprising to encounter a rattle-
snake there. This specimen, a female, had 23 scale rows at midbody, 173
ventrals, and 22 caudals. The total length is 653 mm., the head length
26 mm. SU 15633, a small immature female, has 25 dorsal scale rows
at midbody, 171 ventrals, and 22 caudals. The total length is 338 mm.,
the head length 18 mm.

Crotalus Mitchell i mitchelli Cope, 1861
This subspecies has been noted by Klauber (1936, p. 176) to occur
on Cerralvo, Espiritu Santo, and San Jose islands. He notes that the single
specimens examined from San Jose and Cerralvo are quite typical of C. m.
mitchelli and that the three examined from Espiritu Santo are somewhat
intermediate between the two mainland forms, C. m. mitchelli and Cro-
talus in. pyrrhus, but somewhat closer to mitchelli.

The author has seen two other examples of this subspecies from Gulf
islands. An adult male from San Jose Island (SU 14030) has 166
ventrals, 27 caudals, 18-17 supralabials, 16-15 infralabials, 36 body
blotches and 25 scale rows at midbody.

The other example of this subspecies (SU 14031) constitutes a new
record for Carmen Island. It was collected in the vicinity of Puerta
Ballendra, April 18, 1953, by John P. Figg-Hoblyn and Jon Lindbergh.
It is an adult female which has 175 ventrals, 19 caudals, 16-17 supra-
labials, 15-14 infralabials, 39 body blotches, and 27 scale rows at midbody.

Cliff — Snakes of the Gulf Islands 85

It is 690 mm. in body length, has a tail length of 44 mm., a head length
of 22 mm., and a head width of 28 mm.

Crotalus mitchelli muerlensis Klauber, 1949
This dwarfed insular race was recently described by Klauber (1949,
p. 97) from 19 specimens. It is endemic to El Muerto Island, which lies
near Encantada Grande (wrongly called San Luis on charts) at Lat.
30° N. El Muerto is the fifth of the 6 small islands extending northward
from Encantada Grande and seems to be the only island of the Encantada
group that is inhabited by rattlesnakes.

This race is closely related to, and undoubtedly derived from, C. rn.
pyrrhus from the adjacent mainland. It differs from pyrrhus in its small
size (largest specimen only 637 mm.), in having 24 or fewer scale rows
at midbody (pyrrhus usually has 25 or more), and in having fewer body
blotches (32-39, average 35.7).

Crotalus mitchelli pyrrhus (Cope), 1866

Five specimens of this subspecies are known from Angel de la Guarda
Island, according to Klauber (1936, p. 176). Head proportions separate
C. m. mitchelli from C. m. pyrrhus. and these specimens are typical of
pyrrhus. In ventrals, Klauber notes that the insular representatives have
slightly higher counts than specimens from the mainland. It is of interest
to note that the specimens from Angel de la Guarda are very large, which
is very unusual in an island population. This island, however, is one of
the largest in the Gulf. Klauber also states that if a large series were
available, differences in the ventral counts might be significant enough to
warrant subspecific recognition of the insular population.

Crotalus molossus estebanensis Klauber, 1949
The third specimen of the San Esteban rattlesnake (SU 14020) was
collected by Jay M. Savage and the author as it was seen escaping toward
a talus slope at the mouth of a narrow canyon on the southeastern end of
San Esteban Island on May 6, 1952. This subspecies was previously
known from only the type specimen (LMK 26792) and a single paratype
(USNM 64586).

Klauber (1949, p. 104) differentiates this subspecies from the nearby
mainland form on differences in pattern, body proportions, and on a
higher number of dorsal blotches.

The Stanford example is an adult male, 890 mm. in body length, with
tail length 61 mm., and head length 41 mm. It has six rattles. There are
187 ventrals and 25 caudals. The scale rows are 33-27-21. The scales on
the crown consist of two large triangular-shaped internasals which are
almost broken to form a pair of canthals, and 2 prefrontals that are
wider than long. Three large irregular scales, behind the prefrontals,
extend between the supraoculars. The supraoculars are undivided. The
other scales on the top of the head are quite small except for a pair of
elongate obliquely situated scales, which border the posterior medial
surface of the supraoculars and are about the size of 3 of the other scales
in that region placed end to end. The rostral is wider than high. The
prenasal barely touches the supralabial on one side and is excluded from
contact on the other by the small scales anterior to the pit; these scales

86 San Diego Society of Natural History

number 7 on one side and 8 on the other. On each side, the 2 loreals
separate the postnasal from the preoculars. A small scale on each lateral
edge of the prefrontals separates the upper preocular from the prefrontal
and the supraocular from the upper loreal. The upper preocular is the
larger. Including the supraocular and preoculars, 10 scales border the
orbit on one side and 9 on the other. A minimum of 3 scale rows
separates the orbit from the supralabials. The supralabials number 18-19,
the infralabials 16-16. There are no intergenials or submentals and the
first infralabial is undivided. Three infralabials contact the genials on
each side.

The head has a prominent dark brown patch on the crown in the
internasal-prefrontal region. A faint ocular stripe beneath the eye fuses
with the lighter supralabials about 3 scales anterior to the commissure.
The more posterior ocular stripe is almost obsolete.

The dorsal pattern consists of 39 olive-brown blotches which have no
light areas in their centers and are separated dorsally by only one row of
light gray scales. The narrow blotches extend laterally to the edge of the
ventrals. The light borders of the dorsal blotches divide laterally and
form lateral light-ringed circles which are more evident than the dorsal
blotches, but become obscured anteriorly and posteriorly as do the dorsal
blotches. Neither the circles nor the blotches can be distinguished unless
the specimen is held beneath water. The 3 distinct tail rings are brownish-
black anteriorly. Posterior to these, the tail is uniform black-brown.
Below, the tail is spotted with gray anteriorly, but, posteriorly, changes
to the brownish-black of the dorsal surface. The ventral surface of the
body is cream, with brown punctations extending laterally.

In life, this rattlesnake appeared to be an irridescent dark green when
viewed from a distance. In preservative, it lost this sheen and appears
brownish-olive.

The main difference between this specimen and those seen by Klauber
is that its crown is darkened, as in mainland C. molossus molossus. The
best key character to separate the island subspecies from the typical sub-
species is, therefore, the higher number of dorsal blotches.

Crotalus molossus molossus Baird and Girard, 1853
A single male example of this subspecies is known from Tiburon
Island (MVZ 36490). It was collected by Charles G. Sibley on Novem-
ber 10, 1941.

This is an extremely faded specimen, with the ground color light tan.
Due to this fading, it was impossible to count accurately the number of
dorsal blotches, even when the specimen was held beneath water. An
approximation of the number of blotches was attempted by using the
lateral extensions of the dorsal blotch as a guide. This resulted in an
estimate of 32-36 blotches. The head is extremely faded, also, and there
is no indication of a darkening on the crown.

Ventrals number 184, caudals 25, supralabials 17-16, infralabials
16-16. Dorsal scale formula is 31-26—20.

The tail is dark brown with indications of 6 obscure darker rings.
This suffusion of dark brown is extended onto the ventral surface of the
tail. The ventrals and chin region are immaculate white.

Cliff — Snakes of the Gulf Islands 87

Crotalus ruber lucasensis Van Denburgh, 1920

The San Lucan rattlesnake is known to occur on only one island in
the Gulf of California, San Jose Island. Two males were collected by
members of the first Orca expedition (SU 14023-24). Both are large
and fully mature and have 192-190 ventrals, 23-22 caudals, 34-32 dorsal
blotches, and 27 scale rows at midbody. Each has 2 loreal scales in front
of the preoculars. These specimens are not typical of mainland C. r.
lucasensis and lean somewhat toward C. r. ruber, at least in the high
number of ventrals and body blotches. In having 2 loreals and 27 scale
rows at midbody, however, they are characteristic of C. r. lucasensis. In
coloration, they are definitely closer to lucasensis and on this character I
am assigning them to this subspecies. C. ruber ruber tends toward red or
red-brown, while lucasensis is straw-colored, with white scales bordering
the blotches in the center of the body. The anteriormost and posterior-
most body blotches are poorly defined and can be counted only with
difficulty. The tail is light gray with narrow black rings 2 or 3 scales in
width. The anterior rattle matrix is black.

Both specimens were collected in the early evening and were quite
docile, not bothering to rattle even when approached closely. Klauber
has seen 3 additional specimens from this island and assigned them to
C. r. lucasensis.

Crotalus ruber ruber Cope, 1892

This subspecies is known to occur on the following Gulf islands:
Monserrate, San Marcos, San Lorenzo, Angel de la Guarda, and Pond.
Klauber has examined specimens from all of these islands, and none
differs significantly from mainland representatives of this subspecies.

This rattlesnake was very common on Monserrate, and although only
two were collected (SU 14025-26), tracks were very numerous in the
dunes and on the floors of rocky canyons. Both specimens were lying in
the shade of rocks and even though they were approached within a few
teet, they did not coil or rattle.

One specimen was collected on San Marcos (SU 14027) as it lay
coiled under a bush.

The two specimens from Monserrate are males. SU 14025 has 192
ventrals, 23 caudals, and 27 scale rows at midbody. It is quite noticeably
faded in pattern anteriorly and is more brown than red. SU 14026 has
190 ventrals, 23 caudals, and 27 scale rows at midbody; it is a small
specimen and is not faded anteriorly.

SU 14027 from San Marcos has 189 ventrals, 27 scale rows at mid-
body, and 20 caudals; it is darker than the other 2 specimens.
Crotalus tortugensis Van Denburgh and Slevin, 1921

Crotalus tortugensis is a well-known endemic on Tortuga Island. It
is a member of the atrox group, and is definitely most closely related to
atrox. rather than to ruber or exsul.

As Klauber points out (1930, p. 10), the postnasal-preocular relation-
ship and the head size are the most important characteristics that distin-
guish atrox from tortugensis. Of the atrox specimens, 94 per cent have
contact between the postnasal and preocular, or such contact is prevented
by an upper loreal; 3 per cent have such contact prevented by a lower
loreal-canthal contact, and 3 per cent are intermediate or borderline cases.

88 San Diego Society of Natural History

In tortugensis, on the other hand, an exactly opposite ratio exists, only
3 per cent have a postnasal-preocular contact; 94 per cent have the contact
prevented by a lower loreal-canthal contact; and again, 3 per cent are
borderline intermediates. The ratio of the head length to body length
is smaller in tortugensis than in atrox and indicates dwarfing.

Distinctive characteristics: Dorsal coloration is gray and punctated
with brown. Dorsal blotches vary from 32 to 40 (averaging 37). They
are purplish brown, poorly denned and with light borders only medially.
Head markings not well defined. Scale rows 27, occasionally 25; ventrals
in males 180-190 (average 184), in females 183-189 (average 186);
caudals in males 22-25 (average 24), in females 16-19 (average 18).

Acknowledgments

I am indebted to a large number of persons who have assisted me in
the preparation of this report. To the Sefton Foundation of San Diego,
I am especially grateful. The Foundation made its fine research vessel
Ore a available for two successive spring expeditions into the Gulf of
California.

All of the scientific personnel on the two trips into the Gulf of
California assisted at various times in the collection of reptiles. Those
engaged primarily in herpetological collecting were Joseph Ball of the
San Diego Zoo, Joseph R. Slevin of the California Academy of Sciences,
and Jay M. Savage, John P. Figg-Hoblyn, and Alan Leviton of the
Stanford Natural History Museum. I wish to thank all these persons for
their excellent collections and hard work.

Mr. Slevin and Mr. Savage, who are both well acquainted with the
herpetofauna of Baja California and Sonora, offered very helpful com-
ments in the field and provided much-appreciated advice and aid in the
preparation of this paper. Mr. Slevin, the Curator of the herpetological
collections of the California Academy of Sciences, has allowed me to
borrow and examine specimens from the excellent collections of the
Academy expedition of 1921. In addition, I would like to thank the
following persons and institutions for loan of material or for information
regarding specimens deposited in the institutions: Dr. Doris M. Cochran,
United States National Museum; Dr. Laurence M. Klauber, San Diego
Society of Natural History; Mr. Charles M. Bogert, American Museum
of Natural History; Dr. Wilmer W. Tanner, Brigham Young Univer-
sity; Mr. Benjamin Shreve, Museum of Comparative Zoology, Harvard
University.

Dr. George S. Myers of Stanford University has read the entire
manuscript and offered many useful comments and suggestions. Dr. John
C. Briggs of Stanford drew the outline of the head of Chilomenisats
sava^ei. Mr. Stanley Weitzman of Stanford took the two photographs
of Crotalus. This paper was prepared in the Natural History Museum of
Stanford University.

Cliff — Snakes of thh Gulf Islands 89

Bibliography

Anonymous

1937. Sailing Directions for the West Coasts of Mexico and Central
America. Eighth ed., Hydrographic Office, Washington, pp.
vii -f 430.

Daird, Spencer F., and Girard, Charles

1853. Catalogue of North American Reptiles in the Museum of the
Smithsonian Institution. Part 1 — Serpents, pp. xvi -\- 172.

Bogert, Charles M.

1931. A Study of the Genus Sa.lva.dora, the Patch-nosed Snakes.

Publ. Univ. Calif. Los Angeles, Biol. Sci., Vol. 1, No. 10,

pp. 177-236.
1945. Two Additional Races of the Patch-nosed Snake, Salvadora

hexalepis. Amer. Mus. Nov., No. 1285, pp. 1-14.

Cope, Edward D.

1861. Contributions to the Ophiology of Lower California, Mexico,
and Central America. Proc. Acad. Nat. Sci. Phila., 1861,
pp. 292-306.

1866. On Reptilia and Batrachia of the Sonoran Province of the
Nearctic Region. Proc. Acad. Nat. Sci. Phila., 1866, pp. 300-
314.

1867. A Collection of Reptiles from Owen's Valley, California.
Proc. Acad. Nat. Sci. Phila., 1867, p. 85.

1892. A Critical Review of the Characters and Variations of the
Snakes of North America. Proc. U. S. Nat. Mus., Vol. 14,
pp. 589-694.

Hensley, M. Max

1950. Results of a Herpetological Reconnaissance in Extreme South-
western Arizona and Adjacent Sonora with a Description of
a New Species of the Sonoran Whipsnake Masticophis bil'tn-
eatus. Trans. Kan. Acad. Sci., Vol. 53, No. 2, pp. 270-288.

Jan, Georg

1863. Elenco sistimatico degli ondi. Milan, pp. 1-143.

Johnston, Ivan M.

1924. Expedition of the California Academy of Sciences to the
Gulf of California in 1920. The Botany (The Vascular
Plants). Proc. Calif. Acad. Sci., Ser. 4, Vol. 12, No. 30,
pp. 951-1218, map.

Klauber, L. M.

1930. Differential Characteristics of Southwestern Rattlesnakes Al-
lied to Crotalus atrox. Bull. Zool. Soc. San Diego, No. 6,
pp. 1-72.

1931. A New Subspecies of the California Boa with Notes on the
Genus Lichanura. Trans. San Diego Soc. Nat. Hist., Vol. 6,
No. 20, pp. 305-318.

1936. Crotalus mitchellii, the Speckled Rattlesnake. Trans. San
Diego Soc. Nat. Hist., Vol. 8, No. 19, pp. 149-184.

90 San Diego Society of Natural History

1938. A Statistical Study of the Rattlesnake. V. Head Dimensions.

Occ. Papers San Diego Soc. Nat. Hist., No. 4, pp. 1-53.
1942. The Status of the Black Whip Snake. Copeia, 1942, No. 2,

pp. 88-97.
1949. Some New and Revived Subspecies of Rattlesnakes. Trans.

San Diego Soc. Nat. Hist., Vol. 11, No. 6, pp. 61-1 16,

pi. 4-6. '
Linsdale, Jean M.

1932. Amphibians and Reptiles from Lower California. Univ.

Cal. Publ. Zoology, Vol. 38, No. 6, pp. 345-386.
Moccjuard, F.

1899. Contribution a la faune herpetologique de la Basse-Californie.

Nouv. Arch. Mus. Nat. Hist. Paris, Ser. 4, Vol. 1, pp. 297-

344.
Nelson, E. W.

1921. Lower California and Its Natural Resources, Mem. Nat.
Acad. Sci., Vol. 16, pp. 1-194, pis. 1-35.

Ortenburger, Arthur Irving

1928. The Whip Snakes and Racers. Memoirs. Univ. Mich.
Museums, Vol. 1, pp. xviii -\- 247.
Savage, Jay M., and Cliff, Frank S.

1954. A New Snake, Pbyllorhynchus arenicola, from the Gulf of
Caliform'a, Mexico. Proc. Biol. Soc. Wash., Vol. 67, pp.
69-76.
Schmidt, K. P.

1922. The Amphibians and Reptiles of Lower California. Bull.
Am. Mus. Nat. Hist., Vol. 46, Art. 11, pp. 607-707.

Smith, Hobart M., and Taylor, Edward H.

1945. An Annotated Checklist and Key to the Snakes of Mexico.
Bull. 187, U. S. Nat. Mus., pp. iv -\- 239.
Stickel, William H.

1938. The Snakes of the Genus Sonora in the United States and
Lower California. Copeia, 1938, No. 4, pp. 182-190.
Streets, Thomas H.

1877. Contributions to the Natural History of the Hawaiian and
Fanning Islands and Lower California. Bull. 7, U. S. Nat.
Mus., pp. 1-172.
Tanner, Wilmer W.

1944. A Taxonomic Study of the Genus Hypsiglena. The Great

Basin Naturalist, Vol. V, Nos. 3 & 4, pp. 25-92.
1954. Additional Note on the Genus Hypsiglena with a Description
of a New Subspecies. Herpetologica, Vol. 10, Pt. 1, pp.
54-56.
Van Denburgh, J.

1920. Description of a New Species of Rattlesnake from Lower
California. Proc. Calif. Acad. Sci., Ser. 4, Vol. 10, No. 2,
pp. 29-30.
1922. The Reptiles of Western North America. Occ. Pap. Calif.
Acad. Sci., No. 10, Vol. 1, Lizards; Vol. 2, Snakes and
Turtles, pp. 1-1028.

Cliff — Snakes of the Gulf Islands 91

Van Denburgh, J., and Slevin, J. R.

1921a. Preliminary Diagnosis of New Species of Reptiles from
Islands in the Gulf of California, Mexico. Proc. Calif.
Acad. Sci., Ser. 4, Vol. 11, No. 6, pp. 95-98.

1921b. Preliminary Diagnoses of More New Species of Reptiles
from Islands in the Gulf of California, Mexico. Proc. Calif.
Acad. Sci., Ser. 4, Vol. 11, No. 17, pp. 395-398.

92 San Diego Society of Natural History

Artificial Key to the Snakes on the Islands in the
Gulf of California, Mexico
la. A rattle or large horny button on tail; a sensory loreal pit between
nostril and eye.
2a. Prenasals separated from rostral by a row of small scales or granules.
3a. Last supralabial twice as long as those before it; head of adult
contained in over-all body length more than 24 times.

Crotalus mitcbelli mitchelli

3b. Last supralabial not conspicuously enlarged; head of adult con-
tained in over-all body length less than 24 times.
4a. Scale rows usually 23 or fewer; size small, less than 650 mm.

Crotalus mitchelli muertensis

4b. Scale rows usually 25 or more; adults larger than 650 mm.

Crotalus mitchelli pyrrhus

2b. Prenasals in contact with the rostral; no granules between rostral
and prenasal.
5a. Tail with alternating black and ash-gray rings.
6a. First infralabial usually divided.
7a. A single loreal; scale rows 28 or more; body color red or red-
brown Crotalus ruber ruber

7b. Two or more loreals; scale rows 27 or fewer; body color yellow,

brown, or olive-brown Crotalus ruber lucasensis

6b. First infralabial usually not divided.
8a. Upper preocular usually not in contact with the postnasal, and
no upper loreal present.
9a. Light scales surrounding the dorsal blotches; very few puncta-

tions on dorsal ground color Crotalus catalinensis

9b. Light scales evident only on the mid-dorsal scale rows surround-
ing blotches; very heavily punctated with darker spots and flecks

on dorsal ground color Crotalus tortugensis

8b. Upper preocular usually in contact with the postnasal, or such
contact prevented by an upper loreal. . . . Crotalus atrox
5b. No alternating black and ash-gray rings on the tail.
10a. Scales on crown knobby with a definite keel in their centers.
11a. Head contained in body length less than 26 times.

Crotalus enyo enyo

lib. Head contained in body length more than 28 times.

Crotalus enyo cerralvensis

10b. Scales on crown flat, though their edges overlap.
12a. Body blotches more than 37. . Crotalus molossus estebanensis
12b. Body blotches fewer than 38. . Crotalus molossus molossus
lb. No rattle or button on tail; no pit between nostril and eye.
13a. No elongate chin shields between lower labials.

Lichamira roseofusca

1 3b. One or more pairs of elongate chin shields.
14a. Internasals fused with nasals; rostral enlarged.
15a. Prefrontals in contact medially. . . . Chilomeniscus cinctus
15b. Prefrontals widely separated medially.
16a. Ventrals number 130 or more. . . . Chilomeniscus savagei
16b. Ventrals number 121 or fewer. Chilomeniscus punctatissimus

Cliff — Snakes of the Gulf Islands 93

14b. Internasals and nasals present; rostral enlarged or not.
17a. Rostral plate with free lateral edges, conspicuously enlarged.
18a. A series of body blotches across dorsum; no longitudinal stripes.

Phyllorhynchus arenicola

18b. No series of body blotches across dorsum; a pattern of longi-
tudinal stripes.
19a. Two supralabials reaching the eye.

Salvadora hexalepis klauberi

19b. One supralabial reaching the eye.

Salvadora hexalepis hexalepis

17b. Rostral plate not conspicuously enlarged.
20a. A pattern of orange, red, and black rings on head, body, and

tail Micruroides euryxanthus

20b. No pattern of rings on body, head or tail; stripes present or
not.
21a. Scale rows at midbody 23. . . . Lampropeltis catalinensis
21b. Scale rows at midbody 19 or fewer.

22a. Scale rows at midbody 14 Sonora mosaueri

22b. Scale rows at midbody more than 15.
23a. A pattern of small brown blotches on dorsum, with a large
blotch or blotches on the nape.
24a. Two rows of blotches mid-dorsally, separated by a median
stripe a scale or less in width.

Hypsiglena torquata venusta

24b. Dorsal row of blotches single; no median light stripe.
25a. A narrow median nuchal blotch 1-4 scales wide.

Hypsiglena torquata tortugaensis

25b. Median nuchal blotch much wider than 4 scales.
26a. Gulars posterior to chin shields enlarged; 43 caudals in
the two known females. . Hypsiglena torquata gularis
26b. Gulars not enlarged; caudals 42-54 in females

Hypsiglena torquata ochrorhyncha

23b. No pattern of blotches on dorsum and no well-defined
blotch on the nape or lateral to the nape.
27a. No well-defined longitudinal light stripes on body.

Masticophis flagellum piceus

27b. Well-defined longitudinal light stripes on lateral body
scales.
28a. Pattern consisting of one or more dark lateral stripes.

Masticophis bilineatus

28b. Pattern consisting of a single light lateral stripe interrupt-
ed at intervals of 4-7 scales. . . Masticophis barbouri

94

San Diego Society of Natural History

Distribution of Snakes on Islands in the Gulf of California, Mexico
(X = occurrence, E = endemic)

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LICHANURA ROSEOFUSCA GRACIA

CHILOMENISCUS CINCTUS

CHILOMENISCUS SAVAGEI

■■

• ■

CHILOMENISCUS PUNCTATISSIMUS

HYPSIGLENA TORQUATA GULARIS

HVPSIGLENA TORQUATA OCHRORHYNCHA

X ••

HYPSIGLENA TORQUATA TORTUGAENSIS

X

HYPSIGLENA TORQUATA VENUSTA

X

X

.. ..

X

LAMPROPELTIS CATALINENSIS

E

••

MASTICOPHIS BARBOURI

MASTICOPHIS BILINEATUS

V

X

MASTICOPHIS FLAGELLUM PICEUS

X

X X

• X

V V

X

PHYLLORHYNCHUS ARENICOLA

SALVADORA HEXALEPIS HEXALEPIS

X

SALVADORA HEXALEPIS KLAUBERI

SONORA MOSAUERI

MICRUROIDES EURYXANTHUS

X

CROTALUS ATROX

X

CROTALIIS CATALINENSIS

E

• •

CROTALUS ENYO CERRALVENSIS

••

CROTALUS ENYO ENYO

X

X

CROTALUS MITCHELLI MITCHELLI

X

X ••

•• X

X ••

CROTALUS MITCHELLI MUERTENSIS

CROTALUS MITCHELLI PYRRHUS

X

CROTALUS MOLOSSUS ESTEBANENSIS

F

CROTALUS MOLOSSUS MOLOSSUS

X

CROTALUS RUBER LUCASENSIS

CROTALUS RUBER RUBER

X

V

x ••

•• X

X ••

CROTALUS TORTUGENSIS

E

Cliff — Snakes of the Gulf Islands

95

Fig. 1. Gulf of California, showing principal islands.
Based on Charts No. 620, 621, 1006, U. S. Hydrographic Office.

96

San Diego Society of Natural History

Plate 6, Fig. 3. Photograph of Crotalus enyo cerralvensis. Holotype (SU
14021).

Cliff — Snakes of thl Gulf Islands

Hi 4 .

' ' : ft V;

^^* , ► 'I*?*' '■' "\mL*+r~

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"jt&*"*fc-«

Plate 7, Fig. 4. Photograph of Crotalus catalinensis. Holotype (SU 15631.)

TRANSACTIONS

OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY

Volume 12, No. 6, pp. 99-102 February 10, 195 5

A NEW RACE OF DIPODOMYS AND A NEW
RACE OF THOMOMYS FROM ARIZONA

BY

Laurence M. Huey

Curator of Birth ami Mammals, San Diego Society of Natural History

Over a period of years there has been accumulated in
the mammal collection of the San Diego Society of Natural
History a series of specimens of Dipodomys deserti from
southcentral Arizona and of Thomomys bottae from desert
ranges in western Arizona bordering the Colorado River.
The study of this material reveals sufficient characters in
two geographic populations to indicate them as worthy of de-
scriptions and new names. The Dipodomys may be known as

Dipodomys deserti arizonae subsp. nov.
Arizona Desert Kangaroo Rat

Type. — From 3 miles southeast of Picacho, Pinal County, Arizona;
No. 12 5 32 Collection of the San Diego Society of Natural History; adult
male, collected by Laurence M. Huey, May 14, 1937.

Characters. — Compared with Dipodomys deserti deserti, D. d. arizonae
is grayish in color instead of buffy; it has a well marked undertail stripe
in most specimens examined. The skull is smaller, with noticeably more
rounded, inflated bullae and with a more slender rostrum. The maxillary
arches of D. d. arizonae are more sharply angled with the axis of the skull
than in either Dipodomys deserti sonoriensis or D. d deserti. Compared
with D. d. sonoriensis, D. d. arizonae is lighter in color dorsally, with a
tinge of buffy suffusion on the sides that is not present on the specimens
of D. d. sonoriensis examined. The skull of D. d. arizonae differs from that
of D. d. sonoriensis in having more rounded and massive bullae, heavier
molar teeth, and wider maxillary arches.

Measurements of type. — Total length, 345; tail, 193; hind foot, 52;
ear, 14. Skull: Greatest length, 45.3; width across bullae, 30.8; spread

PS. C^P. ZSOL

uimm

ft 9 * 3 1955

i- sri" i r^n

100 San Diego Society of Natural History

of maxillary arches, 2 3.7; greatest length of nasals, 16.2; width of
maxillary arch at middle, 6.5.

Remarks. — Specimens of the race herewith named were first reported
in 1937, from the Picacho area by Willett (Jour. Mamm. 18: 101).
Since that year the writer has, at every opportunity, collected specimens
of this species. Prior to and during this period, 139 skins and skulls have
been accumulated from localities in Death Valley, Inyo County, California,
and from southern Nevada southward to San Felipe, Baja California,
Mexico, on the western side of the Colorado River, and from southwestern
Arizona to Kino Bay, Sonora, Mexico, on the eastern side of the river.
This assemblage has revealed interesting range boundaries and variations
in several characters of this large Kangaroo rat. Oddly enough, the largest
development in size has been found in the population living near Welton
in the extremely arid southwestern section of Arizona. The further
collection of specimens of comparable age may reveal this population
to differ from the population of the Mojave desert region of California
from whence the nominate race was named.

Range and specimens examined. — D. d. arizonae: — Arizona: 3 miles
southeast of Picacho, Pinal Co., 18 (Type locality); 11 miles west of
Casa Grande, Pinal Co., 11; 10 miles south of Gila Bend, Maricopa Co., 3
(not typical); 7 miles east of Papago Well, Pima Co., 2 (not typical).

D. d. sonoriensis — Mexico: Kino Bay, Sonora, 10.

D. d. desert/ — Mexico: Punta Penascosa, Sonora, 17. Arizona: Old
quarry near Yuma, 1; 6 miles east of Yuma, 1; south end of Tule Desert,
Yuma Co., 1; Quitovaquito, Pima Co., 1; Welton, Yuma Co., 20.
Nevada: Coyote Spring, Lincoln Co., 1. California: Mesquite Flat, 8 miles
north of Stovepipe Well, Death Valley, 2; Mesquite Spring, north end
of Death Valley, 6; 4 miles north of Keeler, Inyo Co., 6; Mojave River,
San Bernardino Co., 1 (topotype) ; Whitewater, San Gorgonio Pass,
Riverside Co., 1; Palm Springs, Riverside Co., 2; Borego Valley, San
Diego Co., 4; La Puerta Valley, San Diego Co., 10; Carrizo Creek,
San Diego Co., 3; Laguna, Colorado Desert (Imperial Co.), 2; Coyote
Wells, Colorado Desert (Imperial Co.), 1; 1 mile west of Pilot Knob,
Imperial Co., 1. Mexico: De Mara's Well, Laguna Salada, Baja California,
5; 40 miles north of San Felipe, Baja California, 1; 30 miles north of
San Felipe, Baja California, 1 ; San Felipe, Baja California, 7.

The Thomomys may be known as

Thomomys bottae cedrinus subsp. nov.
Chemehuevis Mountain Pocket Gopher

Type. — From summit of Crossman Peak, (Juniper — Pifion Belt),
Chemehuevis Mountains, Mohave County, Arizona; No. 13161 Collection
of the San Diego Society of Natural History; adult female; collected by
Laurence M. Huey, April 27, 193 8.

Huey — New Race of Dipodomys and Thomomys 101

Characters. — Compared with Thomomys bottae desertorwm, its near-
est comparatum, which abounds in the valley floor regions to the north
and east of the Chemehuevis Mountains, T . b. cedrinus is smaller in size
and more brightly colored dorsally. The upper parts are bright deeply
shaded buff, which extends well down onto the sides. There is a sprinkling
of blackish hairs on the upper median surface. The undercoat is dark
bluish-gray. The muzzle and fairly large auricular areas are black. The
ears are small. The skull differs from that of T. h. dcscrtonim in being
slightly smaller and lighter boned, with zygomatic arches less sharply
angled from the axis of the skull; they broaden in spread anteriorly. The
rostrum is more slender and auditory bullae are larger and more roundly
inflated.

Measurements. — Type: Total length, 182; tail, 53; hind foot, 24;
ear, 4. Skull: Greatest length, 34.0; spread of maxillary arches 20.2;
length of nasals, 12.1; interorbital constriction, 6.7; alveolar length of
upper molar series, 8.1.

Range. — So far as is now known, from the summit and north slope
of Crossman Peak, Chemehuevis Mountains, to the vicinity of the Lucky
Star Mine on the lower slopes of the range, all within the Juniper-
Pirion Belt.

Remarks. — This newly named race is a dwarf mountain-top gopher.
It is found living in shallow rocky soils within the pinon-juniper area of
this desert mountain range.

Specimens examined. — T. b. cedrinus: Summit of Crossman Peak, 4
(including type) ; Lucky Star Mine, 4.

T. b. desertorum: 1 mile south of Yucca, Mohave Co., 1; 3 miles
south of Yucca, Mohave Co., 2; 12 miles south of Yucca, Mohave Co.,
3 (all collected in a sandy, desert valley — cactus — yucca association).

Thomomys bottae desitus: Wickiup (Big Sandy River), Mohave
Co., 13.

Thomomys bottae hualpaiensis: Democrat Mine, Hualpai Mountains,
Mohave Co., 14.

102 San Diego Society of Natural History

TRANSACTIONS

OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY

Volume XII. No. 7, pp. 103-152. Plates 8-11

A REVISION OF THE PACIFIC COAST PHYTOSI

WITH A REVIEW OF THE FOREIGN GENERA

(COLEOPTERA: STAPHYLINIDAE)

BY

Ian Moore

BUS. «. Z00£

El Cajoir, California

LIMY

MAR 2 3 1956

-

wmim

1

SAN DIEGO, CALIFORNIA

Printed for the Society

March 1, 1956

COMMITTEE ON PUBLICATION

Laurence M. Klauber, Chairman
Joshua L. Baily Charles C. Haines

Carl L. Hubbs
John A. Comstock, Editor

MAR 2 ,

Table of Contents

Page .
Introduction 107

Acknowledgments 108

Material Examined 108

Subtribe Phytosi 108

Ecology 110

Characters of Generic Value 110

Key to the Genera of the Subtribe Phytosi 115

Systematic Treatment of the Pacific Coast Genera 116

Liparocepbalus 116

Diaulota 119

Amblopusa 127

Bryobiota ..129

Bryothinusa •. 131

Tbinusa 132

The Foreign Genera 135

Pbytosus 135

Cameronium -. 137

Baeostetbus 137

Antarctophytosus 139

Bibliography 140

Index of Figures 145

A REVISION OF THE PACIFIC COAST PHYTOSI

WITH A REVIEW OF THE FOREIGN GENERA

(COLEOPTERA: STAPHYLINIDAE)

by Ian Moore

INTRODUCTION

The present study is an effort to assemble in one paper as much as could
be discovered about one small group of genera of the Aleocharinae, a poorly
organized subfamily of staphylinid beetles. The very large number of mi-
nute insects which are grouped together in this subfamily are represented in
a great variety of environmental situations by an almost innumerable num-
ber of individuals. In spite of their abundance, the members of this sub-
family are probably the least known of any large group of insects. Black-
welder stated that "Many thousands of species have been described, but
comparatively few generic revisions have been published, and no adequate
key to the genera has been proposed," and characterizes the situation as
". . . what appears now to be a hopeless chaos. . ."* From the foregoing
remarks it would seem that any effort to assemble material and redescribe
even a small number of these genera would be of some help toward a later
reclassification of the subfamily. In the present study, this appears to be
particularly true, because a number of characters have been discovered in
the material at hand that are not recorded in the literature.

Whether the association of the genera treated here is a true phylogenetic
one is open to question, and is beyond the scope of the present paper. This
problem can be attacked only in a much more comprehensive study than
the present available collection and library permit.

The ten genera discussed in this paper have in the past been associated
by various authors in the subtribe Phytosi of the tribe Bolitocharini of the
subfamily Aleocharinae. I was first introduced to this interesting group
of marine insects by the late Edwin C. Van Dyke during a collecting trip
to Moss Beach, California, in the fall of 1950. Later, while attempting to
identify some species of Diaulota from Southern California, considerable
difficulty was encountered which led eventually to the assembling of the
material which forms the basis for this review of the group. Due to lack
of material, it is not possible to present a complete revision of the foreign
genera. However, for comparative purposes, these genera are included and
some discussion of the species is attempted.

*Blackwelder, R. E. 1943. Monograph of the West Indian beetles of the family Staphy-
linidae. U. S. Nat. Mus. Bull. 182, pp. 1-658.

108 San Diego Society of Natural History

ACKNOWLEDGMENTS

The opportunity to examine the material in the collections of the Cali-
fornia Academy of Sciences at San Francisco was generously extended by
Hugh B. Leech, Curator of Coleoptera of that institution. Through the
kindness of C. M. F. Von Hayek and E. B. Britton, I have been able to
borrow a number of specimens from the British Museum. I wish particularly
to acknowledge my indebtedness to Mildred H. Meeder, librarian of the
San Diego Scientific Library, for very considerable aid in obtaining publica-
tions and photostats of publications not available in San Diego, and to
Arthur H. Fischer, Director, and Charles Harbison, Curator of Insects of
the San Diego Museum of Natural History, for allowing me unlimited use
of the facilities of that institution. Richard E. Blackwelder of the United
States National Museum has kindly compared specimens of one species with
Casey's type and has given other aid. Samuel G. Harter has been of very con-
siderable help and a very congenial companion on several collecting trips,
on one of which he found the first specimens of one of the new species de-
scribed here. Helen Moore has spent a great deal of time typing the manu-
script. Above all else, I will always be indebted to the late Edwin C. Van
Dyke for encouragement and assistance on many occasions.

MATERIAL EXAMINED

The very excellent series lent to me by Leech from the collections of the
California Academy of Sciences contains all the material in this group from
the Fenyes, Van Dyke, Blaisdell and Martin collections, as well as some
slide preparations by E. S. Ross and by F. L. Rodgers. This material has
been augmented by a fairly large series of a number of the California species
collected by myself since 1950 and, except for the types and certain desig-
nated paratypes, at present in my own collection. Additional specimens
which were lent by the British Museum brought the total number of speci-
mens studied to about one thousand.

SUBTRIBE PHYTOSI

Pbytos7is was the first described of the ten genera now placed in this
subtribe. It was first recorded by Curtis from England in 1838. A number
of species of Phytosns have since been described from widespread localities
in the northern hemisphere. None is known from the Pacific Coast of North
America.* Two species of Cameronium are known from the east coast of

*The name Phytosus opacus LeConte is listed in the Leng catalogue and is retained in
the supplements, although the species is also listed there, correctly as Potitamalofa opaca
(LeConte). This error has been copied in the supplement to the Coleopterorum Catalogus.

Moore — Pacific Coast Phytosi 109

Africa. The genera Antarctophytosus Waterhouse and Baeostethns Broun
are known only from the antarctic area. Three species of Liparocephalus
Maklin have been described from the north Pacific. The remaining four
genera described by Casey are all known only from the pacific coast of
North America. The species of this subtribe have in common the fact that
they are all closely associated with the marine environment, either living
on the seashore or in the intertidal belt. The following characters are com-
mon to all the genera examined, but have not been verified in Baeostethus.

1. Middle coxae contiguous, the mesosternal and metasternal processes
acute and their apices separated from each other by about one-third of the
length of the coxal cavities.

2. Labial palpi either two- or three-segmented, short, never as long
as the mandibles.

3. Mandibles simple at tip.

4. Tarsi short. Anterior and middle tarsi four-segmented, posterior
tarsi five-segmented. Last tarsal segment as long at least as the preceding
two together. First segment usually no longer than the second but never
longer than the second and third together.

The character of the contiguous middle coxal cavities appears to
separate these genera from most other bolitocharids.* Not all authors define
this character in the same manner. I consider the cavities separated only if
the mesosternal process meets or nearly meets the metasternal process or if
the apices of these processes are separated only by a distinct intercoxal plate.
In the Phytosi both processes are acute and distinctly separated from one
another by about one-third of the length of the coxal cavities. The two
coxal cavities are in reality divided only by a sharp ridge which is usually
not visible.

Fenyes did not follow previous workers but divided the tribe in a dif-
ferent way so that these genera fell into two separate subtribes, the Liparo-
cephali (including Actocharis, Amblopusa, Antarctophytosus, Diaulota,
and Liparocephalus) and the Phytosf (including Arena, Baeostethus, Bryo-
biota, Bryothimisa, Phytosus, Poly pea, and Thinusa) . He based this division
on the supposed segmentation of the labial palpi. The inclusion of Actocharis
and Polypea is not in accordance with the classification of Bernhauer and
Scheerpeltz, who placed the former with the Oxytelinae and the latter
with the Mylaenini. As these two genera have not been seen, no attempt is

*The only exceptions known to me are certain species of Placusa, which can be sep-
arated by the structure of the posterior tarsi.

110 San Diego Society of Natural History

made here to determine their true relationships. Cameron has demonstrated
that Arena belongs with the Myrmedoniini.

ECOLOGY

It is of interest that at least four of the Pacific Coast genera of this
subtribe, Amblopusa, Bryothinusa, Liparocephalus, and Diaulota, are among
the very few insects which inhabit the intertidal belt of the ocean. The
remaining genera apparently occur only on the seashore, largely in decay-
ing seaweed. Detailed illustrations of the larvae of Liparocephalus and
Diaulota were published by Chamberlin and Ferris and by Saunders. Keen
gave some information about the habits of Liparocephalus at the Queen
Charlotte Islands. Nothing is known of the early stages of Bryobiota,
Thinusa, Bryothinusa, Amblopusa and Baeostethus, but the larva of Phyto-
sus was described by Fauvel and that of Antarctophytosns by Enderlein.

CHARACTERS OF GENERIC VALUE

To anyone reviewing the literature of this group of insects, it will be
apparent that of the few characters which have been given generic value,
several have been interpreted in different ways by different workers. In order
to avoid confusion in the use of this paper, it seems worthwhile to discuss
these characters and explain their use by the present author.

Labrum. — The anterior margin of the labrum may be rounded, arcuate,
truncate, emarginate, or bilobed. When it is rounded, it is evenly so; when
it is arcuate, the sides turn rather abruptly back from the apex. It is con-
sidered truncate if there is a considerable straight central edge, even if the
outer angles are slightly rounded. It is emarginate if the outer angles are
produced anteriorly to the central edge. When it is bilobed, the central edge
is acutely notched in the middle. In Thinusa, this notch is filled with a dark
membrane on a slightly more ventral plane.

Mandibles, — The mandibles may be in one plane, or they may be arched
gradually downward in the apical half, a condition I have referred to as
"curved ventrally." Often, on one or on both mandibles, there is a median
tooth at the inner edge about halfway between the base and the apex. In
Amblopusa, Bryothinusa, and Liparocephalus there are also serrations be-
tween the median tooth and the tip. Baeostethus is said to have three inner
teeth on each mandible.

Maxillae. — The very striking variations in the inner and outer lobes
of the maxillae are probably constant generically. When better known, these
parts may prove to be quite useful in the classification of the genera. Un-
fortunately, these characters could not be fully investigated in the present

Moore — Pacific Coast Phytosi 111

study, because some of the genera have been seen only in loan material which
could not be dissected.

Labial palpi. — It has been pointed out by several authors that the num-
ber of segments of the labial palpi in many groups of Staphylinidae is an
inconstant and unreliable character, variable even within some species. How-
ever, this character has been used extensively in separating genera and even
higher groups. Within the Phytosi, as pointed out by Chamberlin and Ferris,
the genus Diaulota shows considerable variation: some specimens of a species
have only 2 segments, whereas in others the basal segment is divided into
two, making 3 segments in all. I have observed that in the genera in which
the labial palpi are always three-segmented, the basal segment is always the
widest as well as the longest. In the genera in which either condition may
exist, the three-segmented condition is brought about by a division of the
basal segment of the two-segmented palpus, so that segments one and two
are of nearly equal width, and segment one is always the shorter. If the
relative widths and lengths of the segments are recorded, a great deal of
confusion can be avoided. The labial palpi of Antarctophytosus, Bryothinusa
and Liparocephalm are similar in structure to those of Diaulota. Although I
have observed only the three-segmented condition in Antarctophytosus, Dr.
Cameron describes the palpi as two-segmented in Paraphytosus, a synonym
of that genus. In Liparocephalus I know only the two-segmented condition
but have seen both types in Bryothinusa. In the remaining genera the labial
palpi are apparently always three-segmented. Casey described Amblopusa as
having two-segmented labial palpi, but his descriptions of the widths and
lengths of the segments leads me to believe that the third segment was
broken off his specimen (see accompanying illustration of A. borealis Casey
from the Fenyes Collection) .

Ligula. — The ligula is often difficult or impossible to observe in
museum specimens. In most of the phytosids it is moderately developed and
slender. In some species of Diaulota it is so small that it might be considered
wanting. In Thinusa and Cameronium the ligula is bifid (perfectly Y-
shaped), a condition found in many other aleocharinids but unknown in
the other phytosids.

Mentum. — The mentum is trapezoidal, widest at its base, and either
truncate anteriorly or moderately or deeply emarginate.

Eyes. — The eyes are never prominent, seldom interrupting the side
margin of the head. They vary greatly in size. They are minute in Am-
blopusa and Baeostethus (with as few as 3 separate facets in Amblopusa
brevipes Casey). In Thinusa they occupy more than half the side of the

112 San Diego Society of Natural History

head. In most of the genera setae are interspersed with the facets. In others,
Liparocephalus for example, there are no such setae.

Antennae. — The antennae present some differences in the relative
lengths and widths of the segments. The most useful of these characters
are the very short third segment in Amblopusa and the very long tenth
segment in Bryothinusa. The relative length of the antennae, as compared
with that of the head, pronotum, and elytra, is often used to separate
species and sometimes even genera. Because the antennae do not usually lie
straight back over these parts, and it is very difficult to so arrange them,
such estimates can be inaccurate and should therefore be used with care.

Infraorbital ridge. — This ridge is quite variable in the Phytosi, and,
although apparently always lacking in some genera and always well de-
veloped in others, in some species of Diaidota it may be either entirely lack-
ing or moderately developed basally. The phrase "temples margined be-
neath" is often used to refer to the presence of this ridge.

Pronotum. — The differences in the shape of the pronotum are largely
only specific in value and often not that. In the center of the disc Bryobiota
has a longitudinal impression that is not found in the other genera.

Hypomera. — The hypomera is always visible from the side in the
Phytosi but in Liparocephalus it is extremely small and hard to see. Casey
used its shape for separating some genera, but I am unable to make such an
application.

Prosternum. — This is difficult to observe in museum specimens. The
small differences I have been able to observe do not seem to have much
generic value.

Elytra. — The numerous minor variations in shape and size seem to be
largely specific differences. In Thinusa the outer edge is much longer than
the suture.

Tibiae. — In most of the genera, the tibiae are clothed by a long sparse
pubescence. In Phytosus (Pbytosus) , Phytosus (Actostis) , and Thinusa the
anterior and middle tibiae possess, in addition to the usual pubescence, sev-
eral series of long setae. The setae, although usually referred to as spines,
are not direct extensions of the chitinous integuments and each has a dis-
tinct articulation at the base. They are several times the diameter of the
pubescence which is interspersed with them.

Tarsi. — Any discussion of the tarsi brings up the problem of the major
classification of the Aleocharinae. The present classification is based largely
on the number of segments possessed by the tarsi. In the tribe Bolitocharini,
of which the Phytosi is treated as constituting a subtribe, the tarsal seg-

Moore — Pacific Coast Phytosi 113

merits are supposed to number 4 on the anterior and middle legs and 5
on the posterior. By many authors, this tarsal classification is considered
not to indicate true relationships but merely to be a matter of convenience.
Actually, it is often a matter of great inconvenience. The very small size
of the insects, coupled with the peculiar structure and often dense vestiture
of the tarsi, makes the counting of the segments difficult and often very
doubtful. Many genera have been switched back and forth from one tribe to
another by different students because of the different counts arrived at.
This situation undoubtedly accounts for some of the confusion surrounding
the classification of the subfamily. It seems likely that a truly phylogenetic
classification will be arrived at only when other characters are utilized in
conjunction with the tarsal structure. Another difficulty is indicated by the
observation of Chamberlin and Ferris that in one species of Diaulota the
posterior tarsi in the majority of specimens studied are reduced to 4 seg-
ments. They state that "The proportions of the segments and the arrange-
ment of the setae indicate that the four-segmented condition arises from
a fusion of the normal fourth and fifth segments." The following charac-
teristics seem to be of value in distinguishing the genera of the Phytosi. The
anterior tarsi are short and compact in Amblopusa. In Baeostethus they
are said to be so compact and so densely setose that the basal segments are
not readily visible. The unique spatulate plantar setae of the posterior tarsi
of Liparocephahis have been pointed out by Chamberlin and Ferris. Varia-
tions in the relative lengths of the tarsal segments have been used to sep-
arate some genera of the tribe. In Thinusa certain such variations were given
specific value by Casey, but according to Fenyes' synonomy of that genus,
these variations would be intraspecific.

Metasternum. — In Phytosus and some of the other genera the meta-
sternum is long, so that there is a considerable separation of the middle from
the hind coxae. In Diaulota, Liparocephahis, and Baeostethus the metaster-
num is so reduced in length that the coxae are placed together in a compact
group. This is a very distinctive character which indicates, along with other
structural similarities, the close relationship between Diaulota and Liparo-
cephahis.

Basal impressions of the tergites. — In the key that follows, consider-
able use has been made of the presence or absence of transverse impressions
near the bases of certain tergites. The number of tergites having such im-
pressions is fairly constant within the limits of the genera, with some ex-
ceptions (the two known exceptions in this subtribe are Thinusa and Diau-
lota). My understanding of this character is as follows. A tergite is said to

114 San Diego Society of Natural History

be impressed at the base if, at the anterior margin, there is an unbounded
transverse groove which extends nearly to the lateral edge. The impression
will be just posterior to a raised line (the antecostal suture) which delin-
eates the portion of the tergite (the acrotergite) that normally is beneath
the free posterior edge of the preceding tergite. The impression should not
be confused with the antecostal suture or with the acrotergite. When pres-
ent, the impressions are visible only in the proper light, so that often a speci-
men must be rotated in the light in order to cast a shadow in the impression.
The impressions are not ordinarily visible in slide preparations. When pres-
ent, it seems to be invariable that all visible anterior tergites will be im-
pressed up to a certain number, so that if, for example, tergite number
three is impressed, tergites number one and two will also be impressed. These
basal impressions seem to be of wide occurrence in the Aleocharinae. The
number of segments impressed has been used extensively in classification and
is usually given generic significance. However, sometimes the last impressed
tergite is so faintly impressed as to be easily overlooked in a count and the
number is subject to variation within some species. In Diaulota densissima
Casey the fourth tergite is definitely not impressed in some specimens,
whereas in others there is a faint impression. Under these circumstances this
character even though very convenient and often easily used, must be em-
ployed with care.

Constriction of the sternites — This character is difficult to observe,
since it is very similar to the basal impressions of the tergites. The first four
visible sternites are constricted on Phytosus, Amblopusa, and Bryobiota. The
first two of these genera have five impressed tergites, and the last one four
so impressed. When the insect is viewed from the side, it will be seen that
the constriction of the base of the sternite is exactly opposite the impres-
sion on the tergite, so that together they form, around the segment, a ring
that is interrupted only by the paratergites. In Amblopusa the paratergites
are obviously affected also. These two structural modifications very likely
have a common cause or function, possibly allowing greater flexibility of
the abdomen. I have not observed this constriction in any of the remaining
seven genera, although some have strong basal impressions of the tergites.

Sexual characters. — External differences between the sexes are very
weak in this group of insects. No differences have been observed in Bryothi-
nusa, Bryobiota and Thinusa. In Diaulota and Liparocephalus a modification
of the apical margin of the sixth sternite offers some characters of specific
value. In the female of Liparocephalus brevipennis Maklin I have discovered
some very striking modifications of the fifth and sixth tergites. These are

Moore — Pacific Coast Phytosi 115

shown in the drawing of that species. Various degrees of enlargement of the
head in males of some species of Diaulota and Liparocephalus are apparently
intraspecific variations.

The following key is presented in the hope that it may be of aid to
students, in spite of the fact that characters could not always be chosen that
are most readily visible in museum specimens. The fact that Baeostethus
has not been seen and that certain characters have not been recorded in the
original description of it has added to the difficulty of composing the key.
Only two subgenera of Phytosus have been included. The characterization
of Phytosus s. s. and Actosus are based entirely on P. (Phytosus) spinifer
Curtis, P. (Actosus) nigriventris (Chevrolat) and P. (Actosus) balticus
Kraatz. The other two proposed subgenera of Phytosus, Euphytosus Bern-
hauer and Scheerpeltz and Anopsrsus Bernhauer have not been seen and
probably do not belong here. The former is too incompletely described to
permit its inclusion. If Anopsisus is actually a Diglotta, as Koch says it
probably is, then the original description is certainly very poor.

KEY TO THE GENERA OF THE SUBTRIBE PHYTOSI

A. Anterior and middle tibiae spinose externally.

B. Fifth tergite impressed at base; ligula simple.

C. Elytra considerably longer than pronotum; metasternum

long Phytosus (Phytosus)

CC. Elytra not or very little longer than pronotum; metasternum

short Phytosus (Actosus)

BB. Fifth tergite not impressed at base; ligula bifid; metasternum short Thinusa

AA. Tibiae not spinose externally; ligula simple.

D. Anterior tarsi short, compact, so densely setose that the basal segments are not

visible; mandibles with three internal teeth Baeostethus

DD. Anterior tarsi usually as long as middle tarsi, not densely setose.
E. No tergites impressed at base.

F. Right mandible very minutely serrate between median tooth and tip, left

simple; mentum truncate; body slender, parallel Bryothinusa

FF. Both mandibles coarsely serrate between median tooth and tip; mentum

emarginate; abdomen inflated, sides arcuate Liparocephalus

EE. At least three tergites impressed at base.

G. Fifth tergite impressed at base Amblopusa

GG. Fifth tergite not impressed at base.

H. Fourth tergite strongly impressed at base Antarctophytosus

HH. Fourth tergite not (or very faintly) impressed at base.

I. Base of head longitudinally impressed Bryobiota

II. Base of head not longitudinally impressed.

J. Metasternum long, hind coxae distant from

middle coxae Cameroniutn

JJ. Metasternum short, hind and middle coxae

very approximate Diaulota

116 San Diego Society of Natural History

LIPAROCEPHALUS Maklin

Liparocephalus Maklin, 1853, p. 191; Casey, 1893, p. 353; Fenyes, 1918, p. 106; Cham-
berlin and Ferris, 1929, p. 143; Blackwelder, 1936 (various figures to parts of adult).

Generitype. — Liparocepbalus brevipennh Maklin.

Diagnosis. — Body broad, inflated, somewhat convex. Head variable.
Labrum wide, truncate or very shallowly emarginate. Mandibles with coarse
serrations between median tooth and tip, not curved ventrally at tip. Men-
turn deeply emarginate in central third, the emargination straight at bottom.
Third segment of maxillary palpi subconical, widest at apex; fourth narrow
and short. Inner lobe of maxilla broad at base, pointed at apex, densely setose
within; outer lobe broad with a dense brush of setae internally at tip. Labial
palpi two-segmented; first segment four times as long as wide; second half
as wide as first and little more than half as long; palpi separated at base by
width of first segment. Ligula simple, slender, a little less than half as long
as first segment of labial palpi. Eyes moderately large, occupying about one-
third of side of head; hairless between facets. Antennae with all segments
elongate. Pronotum variable. Anterior margin of prosternum straight.
Hypomera very small but visible from the side. Elytra short, not more than
half the length of pronotum; internal apical angles rounded. Tibiae pubes-
cent. Plantar setae of posterior tarsi spatulate. Abdomen inflated, arcuate
at sides; without impressions at base of tergites; sternites not constricted at
base.

Remarks. — This genus is recognizable at a glance by the inflation of
the abdomen. It differs from all but Amblopusa and Bryothinusa in the ser-
rate mandibles, but it is indicated by a number of more important characters
to be most closely allied to Diaulota. The spatulate plantar setae of the hind
tarsi have been observed in none of the other genera. Three species have been
described; brevipennh Maklin from Alaska, cordicollis LeConte from Cali-
fornia and takunagai Sakaguti from Japan. The latter is apparently very
similar to cordicollis but is said to differ in the non-dehiscent sutural margin
of the elytra. Judging from an illustration given by Sakaguti it also differs
from our species in the male secondary sexual characters. Cordicollis has
been variously considered a full species, a subspecies of brevipennis, and a
synonym of that species. In the material examined by me I find two quite
distinct species, one from Alaska and the other ranging from British Colum-
bia south to San Luis Obispo County, California. I believe these to be the
two previously described North American species, although my determina-
tions do not agree with those of most former students. In former keys, color

Moore — Pacific Coast Phytosi 117

has been used to separate the species. My series shows that the color is quite
variable and is completely unreliable in separating the species. The size
and shape of the head is also extremely variable. The secondary sexual
characters provide such positive means of identification in both sexes that
these have been chosen for the new key.

KEY TO THE NORTH AMERICAN SPECIES
OF LIPAROCEPHALUS MAKLIN

A. Median lobe of sixth sternite of male shorter than lateral lobes; apical margin of fifth

tergite of female broadly emarginate brevipennis

AA. Median lobe of sixth sternite of male longer than lateral lobes; apical margin of fifth
tergite of female straight cordicollis

Liparocephalus brevipennis Maklin

Liparocephalus brevipennis Maklin, 185 3, p. 192.

Description. — Densely piceous to dark reddish brown; integuments
densely granulose; clothed in a short dense pubescence which is densest on
the head; distinctly and rather closely punctured throughout. Head about
as wide as long; narrower than pronotum. Eyes moderate in size, occupy-
ing not more than one-third of side of head. Temples margined beneath the
eyes. Antennae about as long as head and pronotum; segments one to eight
elongate, nine and ten very little longer than wide, eleventh elongate. Lab-
rum broadly and feebly emarginate in a continuous arc. Pronotum a little
wider than long, widest at apical third; sides arcuate to hind angles which
are not sinuate; base narrower than apex. Each elytron about as wide as
long and about half the length of pronotum; internal apical angles broadly
rounded. Abdomen broadly inflated, with arcuate sides; no tergites im-
pressed at base; sternites not constricted. Male with sixth sternite pro-
duced apically in three rounded lobes, the median lobe shortest; fifth ter-
gite straight at apex. Female with sixth sternite arcuate apically; apical
margin of fifth tergite strongly emarginate with a small, broad arcuate lobe
in the center of the emargination;, sixth tergite with a longitudinal ridge
for its full length. Length about 3.5 mm.

Type locality. — "In insula Chtaguluk ad castellum Constantinovsk,"
Alaska. J. B. Tompkins of the Bancroft Library at Berkeley, California,
has kindly located that island for me under the present name of Hinchin-
brook Island in Prince William Sound in southern Alaska.

Additional published records. — Unalaska, Alaska. Other records are
doubtful because most identifications have been based on color.

118 San Diego Society of Natural History

Material examined. — Nine specimens of both sexes in the Blaisdell Col-
lection (C.A.S.) from Dutch Harbor, Unalaska, Alaska, and five from the
same series in the Fenyes Collection.

Remarks. — This species is quite distinct from the one I identify as
cordicollis LeConte and can easily be separated from it by the secondary
sexual characters of both sexes. The specimens collected by Blaisdell at
Dutch Harbor are the only ones I have seen. If this is not brevipennis, it is
unnamed.

Liparocephalus cordicollis LeConte

Liparocephalus cordicollis LeConte, 1880, p. 177; Chamberlin and Ferris, 1929, p. 154
(figures of adult, part).

Liparocephalus brevipennis Keen, 1897, Fig. 35; Saunders, 1928, p. 542 (figures of adult
and larva), Chamberlin and Ferris, 1929, p. 143.

Description. — Densely piceous to dark brown; integuments densely
granulose throughout; clothed in a short dense pubescence, which is less
apparent on the head; distinctly and rather closely punctured throughout.
Head broader than or as broad as long, subquadrate to round; sides behind
the eyes subparallel and straight for most of the length or gently arcuate.
Eyes moderate, occupying about one-fourth of side of head. Antennae about
as long as head and pronotum; all segments elongate. Labrum truncate.
Mentum emarginate for central third; the emargination straight at bottom.
Temples bordered beneath the eyes only at base of head. Pronotum a little
wider than long, widest at apical third; sides arcuate to basal angles where
they sometimes become sinuate; base narrower than apex. Each elytron
about as long as wide and about half as long as pronotum. Abdomen broadly
inflated, with sides arcuate; tergites not impressed; sternites not constricted.
Male with sixth sternite produced apically in three lobes, the median lobe
longest. Female with sixth sternite arcuate. Length about 4.0 mm.

Type locality. — Mendocino, California.

Recorded distribution. — Massett, Queen Charlotte Island, British Co-
lumbia, to San Mateo County, California.

Material examined. — I have examined 156 specimens: 17 from Massett,
Queen Charlotte Island, British Columbia (Fenyes Collection, C.A.S.) ; 1
from Torino, Vancouver Island, British Columbia, collected by Spencer
(C.A.S.) ; 4 from Indian River, British Columbia, H. B. Leech (C.A.S.) ; 8
from Vancouver, British Columbia, H. B. Leech (C.A.S.) ; 7 from Cannon
Beach, Oregon, E. C. Van Dyke (C.A.S.) ; 2 from DePoe Bay, Oregon, E. S.
Ross (C.A.S.) ; 1 from Bodega Bay, California, C. H. Hanna (C.A.S.) ; 1

Moore — Pacific Coast Phytosi 119

from San Francisco, California, F. L. Rodgers (C.A.S.) ; 7 from Moss Beach,
San Mateo County, California, F. E. Blaisdell (Fenyes Collection, C.A.S. ),
and 4 from the same locality, Ian Moore; 4 from Pigeon Point, San Mateo
County, California, Ian Moore; 99 from Pacific Grove, Monterey County,
California, Ian Moore; 1 from Piedras Blancas Point, San Luis Obispo Coun-
ty, California, Ian Moore. My specimens were all collected in October and
the others from May to July.

Remarks. — It is possible that all records for this genus south of Alaska
are based on this species. The great variability in color and in the shape and
size of the head leads me to believe that most identifications as brevipennis
are incorrect. Chamberlin and Ferris describe what they believe to be two
species from Moss Beach based on "obvious differences" in the male genitalia.
My own experience is that the male genitalia of cordicollis will appear simi-
lar to either drawing given by them if viewed from different angles.

DIAULOTA Casey

Diaulota Casey, 1893, p. 354; Fenyes, 1918, p. 105; Chamberlin and Ferris, 1929, p. 155.

Generitype. — Diaulota densissima Casey.

Diagnosis. — Body elongate, subparallel with slightly inflated abdomen,
subdepressed. Head variable. Labrum rounded. Mandibles asymmetrical, each
with a median tooth, not curved ventrally. Mentum broadly, shallowly
emarginate. Maxillary palpi with third segment subovoid; fourth segment
moderately elongate, slender. Inner lobe of maxilla narrowed toward the
tip, strongly setose within; outer lobe with dense brush of setae at tip. Labial
palpi two- or three-segmented; first segment, or first two conjointly, four
times as long as wide; last segment half as long and half as wide as first, or
first two together; first segment, when palpus three-segmented, only half
as long as second. Ligula minute, hardly visible between the palpi. Eyes
small, occupying less than one-third of side of head. Antennae with first
three segments elongate, ninth and tenth transverse. Pronotum variable.
Anterior margin of prosternum arcuate. Elytra short, not more than half
the length of pronotum. Tibiae pubescent. Tarsi with basal segments short,
equal to second; last segment elongate. Abdomen slightly inflated posterior-
ly; first three or four visible tergites impressed at base; sternites not con-
stricted basally.

Remarks. — As will be pointed out in the remarks following Amblo-
pusa, that genus is quite distinct from Diaulota. Diaulota seems most closely

120 San Diego Society of Natural History

allied to Liparocephalus in the very short metasternum, the short elytra, and
particularly in the very close similarity of the labial palpi and the maxillae.
It differs from Liparocephalus in the form of the mandibles, the impressed
tergites, the very short ligula and many other characters, as pointed out by
Chamberlin and Ferris. Until now, only one species of Dianlota has been
known, D. densissima Casey, with its synonym, D. insolita Casey. There are
in my collection four other species, which are described now.

KEY TO THE SPECIES OF DIAULOTA CASEY

A. Head densely piceous.

B. Each elytron as long as wide; in each sex, head narrower than pronotum and anten-
nae one-third longer than head densissima

BB. Each elytron wider than long; in male, head as wide as pronotum and antennae al-
most as long as head and pronotum fulviventris

AA. Head transluscent, ferruginous.

C. Head small, elongate, narrower than pronotum, narrowest near apex harteri

CO Head large, as wide as or wider than pronotum.

D. Each elytron shorter than wide; third antennal segment as long as second, very

long and slender megacephala

DD. Each elytron longer than wide; third antennal segment shorter than second,
stout vandykei

Diaulota densissima Casey

Dianlota densissima Casey, 1893, p. 354; Saunders, 1928, p. 542 (figures of adult and larva) ;

Chamberlin and Ferris, 1929, p. 157 (figures of adult and larva).
Diaulota insolita Casey, 1893, p. 355.

Description. — Body densely piceous above, beneath sometimes faintly
paler; densely granulose, not shining; clothed in a short dense pubescence;
finely densely punctured throughout, more coarsely on the abdomen. Head
narrower than pronotum in both sexes, sides evenly arcuate. Antennae one-
third longer than head in each sex. Eyes of about 3 5 facets. Temples strongly
margined beneath the eyes. Pronotum a little longer than wide, widest
near apical third; sides evenly arcuate to base and hardly sinuate; base defi-
nitely narrower than apex. Elytra nearly as long as wide and nearly half
as long as pronotum. Abdomen subparallel but inflated slightly pos-
teriorly. First three tergites impressed at base, fourth sometimes with a
very faint suggestion of an impression. Male with sixth sternite produced
in the middle in a rounded triangular lobe and at the sides produced nearly
as far posteriorly, the intervening areas deeply arcuate. Female with the
sixth sternite arcuate posteriorly. Length 2.0-2.9 mm.

Type locality. — Mainland opposite Fort Wrangel, Alaska.

Moore — Pacific Coast Phytosi 121

Additional published records. — Moss Beach, San Mateo County, Cali-
fornia; Nanaimo, Vancouver Island, British Columbia; Yakutat, Alaska.
The male, a unique, was described as D. insolita by Casey from Queen Char-
lotte Islands, British Columbia.

Material examined. — Two females from Dutch Harbor, Unalaska
(C.A.S.); 1 male and 1 female from Massett, Queen Charlotte Island,
British Columbia (C.A.S.) ; 2 specimens in the British Museum from Metla-
katla, "British Columbia" [Metlakatla is in southeastern Alaska], taken by
J. H. Keen, labeled D. insolita; 6 females from Moss Beach, San Mateo
County, California (C.A.S.) ; 1 female and 5 males from Moss Beach (Ian
Moore); 7 females and 11 males from Pigeon Point, San Mateo County,
California (Ian Moore); 110 females and 89 males from Pacific Grove,
Monterey County, California (Ian Moore) . My specimens were all taken in
November, 1954.

Diaulota fulviventris new species

Description of male holotype. — Densely piceous above, dark fulvous
beneath; integuments densely granulose; clothed in a short, fine, dense
pubescence throughout; very finely, densely punctured, more coarsely on
the abdomen. Head round, as wide as pronotum; sides evenly arcuate.
Antennae nearly as long as head and pronotum; first three segments elongate,
each narrower and shorter than the preceding, fourth to seventh ovoid,
eighth to tenth transverse, eleventh obconical. Eyes of about 3 5 facets.
Temples strongly margined beneath the eyes. Pronotum as wide as long;
sides evenly arcuate to hind angles which are slightly sinuate. Elytra con-
siderably shorter than wide and very much less than half the length of
pronotum. Abdomen somewhat inflated posteriorly; apex of sixth sternite
produced medially in a small, rounded, triangular lobe; apical edge lateral to
lobe hardly sinuate. Lateral lobe of Genitalia with enlarged apex produced
at right angle to a short narrow stalk; median lobe with a small hook-like
process at extremity. Length 1.7 mm.

Allotype (female). — Head elongate, narrower than pronotum. An-
tennae one-third longer than head. Abdomen somewhat inflated but sides
more arcuate posteriorly; apex of sixth sternite arcuate posteriorly.

Type locality. — La Jolla Shores, San Diego, California.

Types. — Holotype, male, allotype, female, paratypes, 6 males and 10
females taken at the type locality November 9, 1950 by Samuel G. Harter
and Ian Moore, and 7 males and 14 females taken at the same locality on
November 22, 1950 by Ian Moore. All specimens were collected in fine well-

122 San Diego Society of Natural History

drained cracks in rocks which had to be pried apart with a wrecking bar, be-
tween +4 and +6 foot tide levels in association with Thalassotrechus bar-
barae (Horn) at the higher level and with the two following species at the
lower level.

Disposition of types. — Holotype, allotype, and 4 paratypes deposited
in the collection of the California Academy of Sciences. Two paratypes are
deposited with each of the following: San Diego Museum of Natural His-
tory; American Museum of Natural History; Museum of Comparative Zoo-
logy; Canadian National Collection; Chicago Natural History Museum;
British Museum (Natural History) ; United States National Museum. The
remaining paratypes are at present retained in my collection.

Additional material. — Thirteen specimens from the type locality taken
in September and October, 195 3; 16 males and 37 females from Sunset
Cliffs, San Diego, California, November 21, 1950; 7 males and 4 females
from Pigeon Point, San Mateo County, California, October, 1954; 6 males
and 1 female from Shell Beach, San Luis Obispo County, California, Octo-
ber, 1950; 4 males and 5 females from Descanso Bay, Baja California, Mexi-
co, in April, 1951, and the others in October, 195 3; 7 males and 8 females
from Sauzal, Baja California, Mexico, May, 1951; 1 male and 1 female from
Punta Morro, north of Ensenada, Baja California, in April, 1951, and 2
males and 4 females from the same locality in November, 1954. Larvae that
probably belong to this species were taken on various dates in the spring
and fall at San Diego and at Punta Morro.

Remarks. — This species closely resembles D. densissima and might have
been considered that species had it not been for the differences in the male
genitalia. The females of the two species are so similar that only a very care-
ful examination will distinguish them. The male of fulviventris can be sep-
arated from that of densissima by a number of characters such as the wider
head, longer antennae, and differences in the apical margin of the sixth
sternite. There are some noticeable variations in series taken from different
parts of the range, but the male genitalia show no differences in specimens
from Shell Beach, San Diego, and Punta Morro. The specimens from Pigeon
Point and Shell Beach are darker beneath, those from Sauzal very much
lighter, both above and beneath and in addition have more delicate an-
tennae. These differences appear in the majority of the specimens but not in
all, and possibly indicate enough differentiation to warrant the recognition
of a slightly different "race" to the south.

Moore — Pacific Coast Phytosi 123

Diaulota harteri new species

Description of male holotype. — Testaceous, with fourth dorsal seg-
ment a little darker; integuments finely and densely granulose except on
the abdomen; pubescence fine, short, dense; very finely, rather closely
punctured throughout; the abdomen with a network of fine lines connect-
ing the punctures. Head distinctly longer than wide, widest at posterior
fourth, very distinctly narrower than apex of pronotum; sides of head be-
hind the eyes distinctly sinuate. Eyes of about 16 facets. Antennae nearly
as long as head and pronotum; first three segments elongate, decreasing in
length, third narrower than second, fourth to sixth hardly elongate, seventh
and eighth round, ninth and tenth transverse, eleventh elongate, obconical.
Temples strongly margined below the eyes. Pronotum about as wide as
long, widest at apical third; sides evenly arcuate and slightly sinuate at hind
angles; base narrower than apex. Elytra conjointly about as wide as prono-
tum and about half as long, each elytron nearly square except for the broadly
rounded humeral angle. Abdomen gradually expanded posteriorly to the
fifth segment; apex of sixth sternite produced in its central third in a broad,
pointed triangle; margins from the base of the triangle to the sides, straight.
Length 1.8 mm.

Allotype (female). — Similar to male, but with sixth sternite rounded
posteriorly.

Type locality. — Descanso Bay, Baja California, Mexico.

Types. — Holotype, male, allotype, female, and 26 paratypes taken at
the type locality on October 11, 195 3.

Disposition of types. — Holotype, allotype, and 4 paratypes deposited
in the collection of the California Academy of Sciences. Two para-
types are deposited with each of the following: San Diego Museum of Nat-
ural History; American Museum of Natural History; Museum of Compara-
tive Zoology; Canadian National Collection; Chicago Natural History
Museum; British Museum (Natural History) ; United States National
Museum. The remaining paratypes are at present retained in my collection.

Additional material. — Seventeen specimens were taken at La Jolla
Shores, San Diego, California, in September, October and November. All
were collected from very narrow, well-drained cracks in rocks between +4
foot tide level and mean low water, in association with the following species.
It has not been determined if they extend below mean low water, but at that
point they would occasionally be submerged for as long as 16 hours at a

124 San Diego Society of Natural History

time. It seems likely that a film or bubble of air is retained in the cracks
where they are found.

Remarks. — This species is easily distinguished from densissima and
fulviventris by its very pale, transluscent, yellow or red integuments and
from megacephala by the very narrow head. The color is variable. The type
is the palest specimen seen. The usual color is ferruginous with one to three
abdominal segments black. This species is named for Samuel G. Harter,
who assisted on numerous collecting trips and who collected the first speci-
mens of Diaulota fulviventris.

Diaulota megacephala new species

Description of male bolotype. — Ferruginous, with fourth and part of
fifth abdominal segments and eleventh antennal segment black; integuments
transluscent, minutely and densely granulose except on the abdomen, which
is rather distinctly reticulate; pubescence short, fine and dense; punctura-
tion fine, rather dense. Head trapezoidal, very large, wider and longer than
pronotum, wider than long, widest at the anterior margin; the sides slightly
and evenly arcuate to the base, which is as wide as apex of pronotum. Eyes
of about 22 facets, occupying about one-sixth of side of head. Antennae
nearly as long as head and pronotum; segments one to seven elongate, one
longest and widest, more than twice as long as wide, two very little nar-
rower, about twice as long as wide, three as long as two and very slender,
little more than half as wide as two, four barely wider than three and twice
as long as wide, five to ten increasing gradually in length and width, tenth
about as long as wide, eleventh longer than wide, obconical. Temples not
margined beneath the eyes. Pronotum a little wider than long, widest at
apical third; sides arcuate from apex to near basal angles; basal angles slight-
ly sinuate; base distinctly narrower than apex. Elytra almost as long as
wide, conjointly as wide as pronotum. Abdomen inflated posteriorly; first
three visible tergites deeply, fourth shallowly impressed at base; sixth ster-
nite produced medially in a rounded triangular lobe with the apical margin
slightly sinuate to the side. Lateral lobe of male genitalia slender, with the
expanded tip produced at a little more than a right angle. Length 2.4 mm.
Female unknown.

Type locality. — Descanso Bay, Baja California, Mexico.

Types. — Holotype, male, and 29 paratypes, apparently all males, from
the type locality, on October 11, 195 3.

Disposition of types. — The holotype and four paratypes deposited in
the collection of the California Academy of Sciences. Two paratypes are

Moore — Pacific Coast Phytosi 12 5

deposited with each of the following: San Diego Museum of Natural His-
tory; American Museum of Natural History; Museum of Comparative
Zoology; Canadian National Collection; Chicago Natural History Museum;
British Museum (Natural History) ; United States National Museum. The
remaining paratypes are at present retained in my collection.

Additional material. — Thirteen specimens from La Jolla Shores, San
Diego, California, in October and November, all taken in company with D.
harteri under the same circumstances.

Remarks. — It is assumed that all the specimens taken are males because
of the condition of the sixth sternite. The discovery of a single male of
D. harteri disposes of the likelihood that the two species are opposite sexes
of one species. Megacephala is distinct from all other species but vandykei
by its very large, trapezoidal head. It can be distinguished from vandykei
by the very long, slender third antennal segment and by the shorter elytra.
The head varies in size from slightly larger than the pronotum to very much
larger (as in the holotype). The holotype shows no intraorbital ridge but
some other specimens have a trace of it basally. The color is about as
variable as in harteri. The antennae are the most slender of the genus.

Diaulota vandykei, new species

Diaiilota brevipes Chamberlin and Ferris, 1929, p. 1 5 5 (not Amblopusa brevipes Casey).

Description of male holotype. — Dark ferruginous, with elytra and
outer antennal segments tinged with piceous; abdominal segments one to
four and part of five dense-piceous; integuments minutely and densely
granulose except on abdomen, which is densely reticulate; pubescence short,
fine, and dense; puncturation fine and very dense, much denser on the ab-
domen than in megacephala. Head nearly round, very large, wider and long-
er than pronotum, about as wide as long, widest near the middle; sides evenly
arcuate from behind the eyes to base, which is as wide as base of pronotum.
Eyes of about 20 facets, occupying about one-eighth of side of head. An-
tennae about as long as head plus one-half of pronotum; segments one to
three elongate, segment one mor,e than twice as long as wide, two wider
than one and very little longer than wide, three very little more than half
as wide as two and distinctly shorter than two, a little longer than wide,
four to six oval, increasing very gradually in size, seven to ten transverse,
ten half again as wide as long, eleven elongate, obconical. Temples faintly
margined beneath the eyes at base only. Pronotum very little wider than
long, widest near apical third, sides arcuate from apex to base, base distinctly
narrower than apex. Each elytron a little longer than wide, elytra con-
jointly as wide as pronotum. Abdomen slightly inflated posteriorly, first

126 San Diego Society of Natural History

three visible tergites deeply, fourth shallowly impressed at base; sixth ster-
nite produced medially in a rounded triangular lobe, sides not sinuate.
Length 2.8 mm.

Allotype (female). — Similar to male but head smaller, about as wide
as pronotum; sixth sternite rounded posteriorly.

Type locality. — Pacific Grove, Monterey County, California.

Types. — Holotype, male, allotype, female, and 2 5 paratypes (12 males
and 13 females) taken at the type locality on November 24 and 28, 1954,
and 10 paratypes (5 males and 5 females) taken at Pigeon Point, San Mateo
County, California, on November 2 5, 1954. All specimens were collected
from cracks in the rocks in company with Diaulota densissima and Liparoce-
phalus cordicollis at or near the +3 to +4 foot tide level on the exposed
reefs. The specimens were all taken just above the line of dense seaweed,
as no suitable place to search for them could be found below this mark.

Disposition of types. — The holotype, allotype, and four paratypes de-
posited in the collection of the California Academy of Sciences. Two para-
types are deposited with each of the following: San Diego Museum of Nat-
ural History; American Museum of Natural History; Museum of Com-
parative Zoology; Canadian National Collection; Chicago Natural History
Museum; British Museum (Natural History) ; United States National
Museum. The remaining paratypes are at present retained in my collection.

Additional material. — More than 100 specimens, males, females and
larvae, of this species (or one which is very similar) were taken from the
reef at about +2 tide level at Shell Beach, San Luis Obispo County, Cali-
fornia. This series is quite variable in the shape and size of the head of the
male and is much lighter in color than the type series. The abdominal punc-
tuation is not quite so dense, particularly beneath. However, I am unable
to find any consistent character that will separate the tv/o series with cer-
tainty. The antennae, the secondary sexual characters, and other characters
are very similar.

Remarks. — This species is more likely to be confused with megacephala
than with any other. It can readily be separated from that species by the
very thick second antennal segment and by the much shorter and stouter
third antennal segment, as well as by the longer elytra. This is the species
that Chamberlin and Ferris identified as Amblopusa brevipes Casey, an
identification that led them to synonomize the two genera. It differs from
that species, however, in many fundamental structural characters, as ex-
plained in the remarks following the description of Amblopusa. This species
is named in memory of the late Edwin C. Van Dyke.

Moore — Pacific Coast Phytosi 127

AMBLOPUSA Casey

Amblopusa Casey, 1893, p. 355; Fenyes, 1918, p. 104.
Amblyopusa Eichelbaum, 1909, p. 209 (emendation).

Generitypc. — Amblopusa brevipes Casey.

Diagnosis. — Body elongate, parallel, depressed. Head ovoid. Labrum
rounded. Mandibles symmetrical, with 2 or 3 serrations between median
tooth and tip; not curved ventrally at tip. Mentum emarginate, the emar-
gination flat at bottom. Maxillary palpi with third segment distinctly
ovoid, fourth long and slender. Inner lobe of maxilla slender, with a long
blunt tooth at tip, 3 or 4 shorter teeth close to the tip, and 2 larger median
teeth well separated from each other; outer lobe slender with a brush of
long setae at the tip and another toward the middle. Labial palpi distinctly
three-segmented; basal segment twice as long as wide; second a little nar-
rower than first, a little longer than wide; third narrower than second and
as long as first, four times as long as wide. Ligula simple, a little longer than
first segment of labial palpi, truncate at tip. Eyes minute, with from 3 to
18 facets. Antennae with first two segments elongate, third to seventh
slightly elongate, eighth to tenth ovoid. Pronotum faintly impressed longi-
tudinally. Anterior margin of prosternum straight. Elytra more than half
as long as pronotum. Tibiae pubescent. Anterior tarsi short and compact.
Abdomen with first five tergites impressed at base; first four sternites con-
stricted at base.

Remarks. — Chamberlin and Ferris stated that in their opinion, based
on the identification of the species at hand, Amblopusa is congeneric with
Diaulota. Examination of specimens of Amblopusa brevipes from British
Columbia reveals that the specimens which they identified as that species
were actually Diaulota vandykei, described here. It is my opinion that the
two genera are quite distinct from one another. Amblopusa differs from
Diaulota in many characters, including the very small eyes, the structure
of the labial palpi (Casey's specimen obviously had the last segments miss-
ing), the serrate mandibles, the longer elytra, the longer metasternum, the
structure of the maxillae, the impressed fourth and fifth tergites and the
constricted sternites. Casey had described three species of Amblopusa, one
of which, A. pallida, has been reduced to a synonym of A. brevipes by
Fenyes. The two species appear to be quite similar but probably are distinct.

KEY TO THE SPECIES OF AMBLOPUSA CASEY

A. Eyes of 3 to 5 facets, elytra three fourths as long as pronotum brevipes

AA. Eyes of 12 to 18 facets, elytra two-thirds as long as pronotum borealis

128 San Diego Society of Natural History

Amblopusa brevipes Casey

Amblopusa brevipes Casey, 1893, p. 356 (not Chamberlin and Ferris, 1929).
Amblopusa pallida Casey, 1911, p. 212.

Description. — Ferruginous, with the legs paler; integuments very
minutely and densely granulose, except on abdomen, which is very minutely
reticulate, shining; pubescence moderately long and fine, rather sparse, par-
ticularly on the abdomen; puncturation fine, not dense, more noticeable
on the elytra. Head oval, very little wider than pronotum, a little longer
than wide, widest near basal third, sides evenly arcuate. Eyes minute, of 3
to 5 facets, near the insertion of the mandibles. Pronotum roughly trape-
zoidal, a little longer than wide, with sides rounded at the anterior fifth
which is the widest point, and thence nearly straight to the obtuse basal
angles; base and apex both arcuate; base narrower than apex. Elytra con-
jointly nearly square, not quite as wide as pronotum and about three-
fourths as long, outer edges slightly arcuate, particularly at the humeral
angles. Abdomen nearly parallel, as wide as pronotum; first five visible
tergites strongly impressed at base; first four sternites somewhat constricted
at base. Male with sixth sternite produced medially in a very short broad
rounded triangular lobe. Female with sixth sternite rounded. Length
2.4 mm.

Type locality. — Fort Wrangel, Alaska.

Additional published record. — Metlakatla, "British Columbia" [Met-
lakatla is in Alaska].

Material examined. — I have seen 4 specimens from Massett, British Co-
lumbia, and 1 from Fort Wrangel, Alaska, all in the Fenyes collection
(C.A.S.). The last specimen is labeled "Dupl. No. 131 Casey 1891."

Remarks. — This species differs from A. borealis in the smaller eyes and
longer elytra. A previous record of this species from Moss Beach, California,
was based on Diaulofa vandykei.

Amblopusa borealis Casey

Amblopusa borealis Casey, 1906, p. 3 54.

Description. — Ferruginous; the abdomen sometimes partially piceous;
integuments very minutely and densely granulose except on abdomen,
which is very minutely reticulate, shining; pubescence moderately long and
fine, rather sparse, particularly on the abdomen; puncturation fine, not
dense, most noticeable on the elytra. Head round, wider than pronotum,
widest near the middle; vertex nearly flat; eyes of 12 to 18 facets located
at anterior fourth. Pronotum roughly trapezoidal, a little longer than wide;

Moore — Pacific Coast Phytosi 129

sides rounded at anterior fifth, which is the widest point, and thence nearly
straight, but slightly sinuate just before the obtuse basal angles; base and
apex both arcuate; base narrower than apex. Elytra conjointly wider than
long; two-thirds as long as pronotum; apex very little wider than base;
sides slightly arcuate, particularly at the humeral angles. Abdomen nearly
parallel, as wide as pronotum; first five visible tergites strongly impressed
at base; first four sternites strongly constricted at base. Male with sixth
sternite produced medially in a short broad roaded triangular lobe. Female
with sixth sternite rounded. Length 3.3 mm.

Type locality. — Massett, Queen Charlotte Island, British Columbia.

Recorded distribution. — Metlakatla, "British Columbia" [Metlakatla
is in Alaska].

Material examined. — I have seen 6 specimens from Massett, Queen
Charlotte Island, British Columbia, and 1 collected by E. C. Van Dyke at
Dutch Harbor, Unalaska, Alaska. These are in the Fenyes collection
(C.A.S.) . I collected 2 specimens at Pacific Grove, Monterey County, Cali-
fornia, in November, in company with Diaulota vandykei, D. densissima,
and Liparocephalus cordicollis.

Remarks. — Differs from brevipes in the larger eyes and shorter elytra,
as well as in the shape and size of the head.

BRYOBIOTA Casey

Bryobiota Casey, 1893, p. 367; Fenyes, 1920, p. 130.

Generitype. — Bryobiota bicolor (Casey) .

Diagnosis. — Body elongate, parallel, subdepressed. Head subquadrate,
deeply impressed longitudinally along the midline, particularly at the base.
Labrum truncate. Mandibles asymmetrical, left simple, right with a small
blunt median tooth, not curved ventrally at tip. Mentum broadly, shallow -
ly, arcuately emarginate. Third segment of maxillary palpi oval; fourth
narrow, elongate. Inner lobe of maxillae spinose apically with a setose mem-
branous expansion at base; outer lobe one-fourth as broad as long, with an
expanded membranous ball at tip. Labial palpi three-segmented; basal seg-
ment twice as long as wide, second a little narrower and two-thirds as long,
third half as wide as first and as long as second. Ligula simple, narrow, as
long as basal segment of labial palpus. Eyes very small, occupying less than
one-fifth of side of head. Antennae with first five segments elongate, tenth
slightly transverse. Pronotum ovoid. Anterior margin of prosternum

130 San Diego Society of Natural History

slightly arcuate anteriorly. Elytra as long as pronotum. Tibiae pubescent.
Posterior tarsi with basal segment longer than second. Abdomen with first
four tergites impressed at base; first four sternites constricted at base.

Remarks. — This genus shares a number of characters with Antarcto-
phytosus and with Amblopusa. It is easily recognized by the longitudinal
impression of the head and pronotum as well as by the rather quadrate head.
The generitype is the only species known.

Bryobiota bicolor (Casey)

Pbytosus bicolor Casey, 1885, p. 311.
Bryobiota bicolor (Casey), 1893, p. 368.

Description. — Ferruginous, with the first five abdominal segments
black; integuments very finely granulose, shining; pubescence short, moder-
ately sparse; moderately shallowly punctured throughout. Head a very little
wider than pronotum, subquadrate; sides nearly straight and nearly parallel
from the eyes to a sharply rounded hind angle, thence almost straight in to
the neck, which is half as wide as head. Head barely widest at hind angle;
divided longitudinally by a rather strong impression which is deepest pos-
teriorly, giving on the dorsal posterior surface the appearance of low mound-
like elevations. Eyes small, inserted at the anterior angles of the head.
Pronotum ovoid, a little longer than wide, widest near anterior third;
sides arcuate; hind angles not sinuate; anterior and posterior margins evenly,
gently arcuate; base narrower than apex. Elytra a little wider and about
as long as pronotum, conjointly about as wide as long; sides evenly arcuate;
apices nearly straight; hind angles hardly rounded. Abdomen nearly paral-
lel but with fifth segment the widest; 4 tergites impressed at base, impres-
sions strong. No sexual differences have been observed. Length about
2.4 mm.

Type locality. — San Diego, California.

Additional published records. — San Diego, California. Scheerpeltz,
1934, p. 15 57, gave British Columbia as a locality record without citation
to any authority. I am unable to locate this record.

Material examined. — Seven specimens from San Diego County, Cali-
fornia, collected by Blaisdell (C.A.S.) ; 1 from San Diego, California, A.
Fenyes (Brit. Mus.); 5 from San Diego, California, Fenyes (C.A.S.) ; 1
taken by myself at San Diego, California; 1 from Moss Beach, San Mateo
County, California, by J. O. Martin (C.A.S.).

Remarks. — In 188 5 Casey appended the following note to his descrip-
tion of the species. "This species is extremely abundant under the densely

Moore — Pacific Coast Phytosi 131

packed seaweed thrown up on the shores of the inner harbor in the spring
of the year; occurring with it and also in great abundance, were Cafius
(Remus) decipiens Lee, Motschulskium sinuaticolle Matth., and Phyco-
coetes testaceus Lee, and in less number, Cafius (Remus) opacus Lee." I
have collected in the area of San Diego Bay for many years and have never
seen masses of seaweed cast up on the shores of the inner harbor. I have never
taken Cafius decipiens LeConte, only one specimen of Cafius opacus LeConte,
and only one specimen of Bryobiota bicolor (Casey). The conditions in the
bay are so changed since 188 5 that it is unlikely that these species will ever
be found there in abundance again.

BRYOTHINUSA Casey

Bryothinusa Casey, 1904, p. 312; Fenyes, 1920, p. 131.

Generitype. — Bryothinusa catalinae Casey.

Diagnosis. — Body elongate, parallel, depressed. Head oval, shallowly
concave centrally. Labrum transverse, with anterior margin gently arcuate.
Mandibles asymmetrical, the right with a median tooth and with very
minute serrations between the median tooth and tip, the left simple; not
curved ventrally. Mentum broadly trapezoidal, truncate. Second and third
segments of maxillary palpi ovoid, fourth shorter than width of third. Inner
lobe of maxillae slender, acutely pointed and hooked at tip with a row of
evenly placed, short, stout, pointed setae at inner margin; outer lobe as long
as inner, very slender, particularly at apical fourth, with tip hooked and
finely pointed, and inner edge simple. Labial palpi either two- or three-
segmented; when three-segmented, first segment shorter than second and
wider only at base, and third segment narrower than second and a little
longer than first. Ligula short, no longer than width of basal segment of
labial palpus; simple. Eyes small, occupying about one-fourth of side of
head; composed of about 20 facets interspersed with short hairs. Antennae
long, with all segments elongate; first segment widest, third narrowest and
a little longer than wide, the remainder each a little longer and a little wider
than the preceding, the tenth distinctly elongate, eleventh more than twice
as long as wide and gradually pointed at apex, nearly equal to first in size.
Pronotum transverse, quadrate, widest at apical fifth. Anterior margin of
prosternum slightly emarginate in a gentle arc. Elytra about as long and as
wide as pronotum, longest at outer edge. Tibiae pubescent. Posterior tarsi
with basal segment longer than second. Abdominal segments not impressed
at base; sternites not constricted.

Remarks. — Bryothinusa is at once distinguished from all other phyto-

132 San Diego Society of Natural History

sids by the very elongate antennal segments and the very slender, curved,
simple outer lobe of the maxillae. Only one species is known, the generitype.
This genus would be placed in the Myllaenini by those following pres-
ent classification, but it probably is not closely related to members of that
tribe. It may be closely related to Polypea Fauvel, which, as has already
been pointed out, was placed in the Myllaenini by Bernhauer and Scheerpeltz
and in the Phytosi by Fenyes. It appears to be the most discordant element
in the present group.

Bryothinusa catalinae Casey

Bryothinusa catalinae Casey, 1904, p. 312.

Description. — Fulvous, with abdomen slightly darker; integuments
clothed in a dense short pubescence; finely, densely punctate, the abdomen
with a network of fine lines around the punctures. Head nearly round,
vertex broadly concave. Pronotum subquadrate, widest at anterior third,
sides evenly arcuate, base slightly narrower than apex. Elytra about as
wide and about as long as pronotum; apex slightly sinuate at outer angle.
Abdomen nearly parallel. No sexual differences observed. Length
about 2.0 mm.

Type locality. — Catalina Island, California.

Recorded distribution. — Catalina Island, California.

Material examined. — I have seen 2 specimens in the Fenyes collection
from Catalina Island, California, one collected by Fall and the other by
Baker, as well as 2 specimens from San Diego, California, in the Fenyes col-
lection (C.A.S.). Nine specimens were taken together from a crack in a
large wet stone near high tide mark on a stony beach at La Jolla, California,
in September, 195 3. They were much more agile than any other phytosids
collected by me, running rapidly downward when the stone was split open.
I believe that several of the colony escaped. Two of these specimens were
kindly compared with Casey's type by R. E. Blackwelder.

THINUSA Casey

Tbinusa Casey, 1893, p. 371; Fenyes, 1920, p. 134.

Generitype. — Thinusa maritima (Casey).

Diagnosis. — Body elongate, parallel, somewhat depressed. Head oval,
slightly pointed anteriorly. Labrum bilobed, with the central emargination
filled with a dark, depressed membrane. Mandibles curved ventrally at tip;
right mandible with a median tooth, left simple. Mentum truncate. Maxil-
lary palpi with third segment evenly conical, narrow, widest at apex; fourth

Moore — Pacific Coast Phytosi 1 3 3

segment very slender, and longer than in Phytosus. Inner lobe of maxillae
spinose internally, with a membranous expansion at the base through which
run several long spines; outer lobe with a membranous finger-like process
at tip. Labial palpi three-segmented, the basal segment longest, second
shortest; segments decreasing in width apically. Ligula deeply bifid, the
two processes forming a Y; half as long as first segment of labial palpi. Eyes
large, occupying more than one-half the side of head. Antennae with first
3 segments elongate, sixth to tenth transverse, eleventh with a brush of
short setae at tip. Anterior margin of prosternum straight. Elytra extend-
ing more posteriorly at the sides than at the suture, so that the apex, al-
though straight, is oblique. Anterior and middle tibiae externally with long
spines, which are interspersed with pubescence. Posterior tarsi with first
segment variable, either shorter, equal to or longer than second but never
longer than second and third together. Abdomen with either 3 or 4 seg-
ments impressed at base; sternites not constricted.

Remarks. — Casey stated that Thintisa so closely resembles Actosus that
it might be considered a subgenus of it. He apparently did not observe the
bifid ligula, by which it differs from all other phytosids except Cameronium,
Besides the spinose tibiae, Thinusa resembles Phytosus in the form of the
maxillae and in the pubescence on the pronotum, which streams out later-
ally from the mid-line in a most striking manner. It is an interesting and
puzzling fact that several very abundant species of the tribe Myrmedoniini
which are found in the seaweed in association with Thinusa so closely re-
semble it in many structural characters (i.e. the spinose tibiae, the arrange-
ment of the pubescence; the shape of head and size of eyes, the structure
of the mouth parts) that it is a great task to collect Thinusa and to segregate
the few specimens from their many cousins. The similarities are so great as
to make one wonder about our tarsal system of classification which separates
these forms so widely.

Of the 6 species of Thinusa that Casey described from California,
Fenyes considered only maritima and fletcheri valid. One of the main char-
acters mentioned by Casey for the separation of the species was the relative
length of the basal segment of the posterior tarsi. According to Fenyes'
synonomy, this character would be variable intraspecifically. All the speci-
mens examined by me have the first tarsal segment a little longer than the
second; but, because of the overlapping structure of the segments, the first
may appear shorter if viewed other than directly from the side. Casey's
original description of maritima states that the first and second segments are
equal; but following his description of obscura he states that the first seg-
ment of that species is longer, not shorter, than the second, as in maritima.

134 San Diego Society of Natural History

I believe the synonomy given by Fenyes to be correct. The rather good series
I have seen from various localities indicates that there is considerable varia-
tions in the characters chosen by Casey. Even fletcheri and maritima are
difficult to separate with certainty and future studies may show that the
former is merely a geographic race of maritima.

KEY TO THE SPECIES OF THINUSA CASEY

A. Abdomen shining, very finely reticulate maritima

AA. Abdomen dull, densely granulose fletcheri

Thinusa maritima (Casey)

Phytosus maritimus Casey, 1885, p. 312.
Thinusa maritima Casey, 1893, p. 371.
Thinusa obscura Casey, 1906, p. 3 54.

Description. — Ferruginous, head brown; abdomen sometimes piceous;
integuments densely and finely granulose, except on abdomen, which is
finely reticulate; clothed in a moderately long, even, pale pubescence, which
is sparse on the abdomen; moderately punctured throughout. Head about
as wide as long, pointed anteriorly, widest just behind the large eyes. An-
tennae not as long as head and pronotum. Pronotum widest at anterior
third, sides arcuate in front and straighter to the slightly rounded posterior
angles. Elytra three-fourths as long as pronotum at sides, two-thirds as
long at suture, conjointly not as wide as pronotum. Abdomen narrower
than pronotum; sides straight and nearly parallel. No sexual differences
observed. Length 2.4 mm.

Type locality. — Oakland, Alameda County, California.

Additional published records. — Santa Barbara, Santa Barbara County,
California (as T. obscura Casey). This species was reported from four lo-
calities in New Jersey by Morse, 1909, but I believe on the basis of a mis-
identification, or, more likely, on an error in transcription, as it is listed as
"Thinusa maritima Casey (Polystoma) ."

Material examined. — One specimen from Ilwaco, Washington, col-
lected by Hubbard and Schwarz (Fenyes collection, C.A.S.); 6 collected
in May at San Francisco, California, by Blaisdell (C.A.S.) ; 15 from Rock-
away Beach, San Mateo County, California, in October (Ian Moore) ; 6
from Redondo Beach, Los Angeles County, California, by Fenyes (5
C.A.S. ; 1 Brit. Mus.) ; 3 from Balboa, Orange County, California (Fenyes,
C.A.S.) ; 5 from La Jolla in San Diego, California, in November (Ian
Moore).

Moore — Pacific Coast Phytosi 13 5

Remarks. — If obscura is actually distinct from maritima, the above
specimens must all be the former and maritima not seen by me. There is
considerable variation in color, in shape and size of the antennal segments,
particularly the eleventh, and in some other characters.

Thinusa f letcheri Casey

Thinusa fletcheri Casey, 1906, p. 3 J 3.
Thinusa diver gens Casey, 1911, p. 213.
Thinusa nigra Casey, 1911, p. 214.
Thinusa rohusiula Casey, 1911, p. 214.

Description. — Dark brown, with abdomen piceous; integuments dense-
ly granulose throughout; clothed in a moderately long, pale pubescence
which is sparser on the abdomen; moderately punctured throughout. Head
a little wider than long, hardly pointed in front, widest just behind the
eyes; antennae not as long as head and pronotum. Pronotum a little wider
than long, widest at anterior third, sides arcuate anteriorly and slightly
sinuate before the hind angles. Elytra two-thirds as long as pronotum, a
little longer at the sides. Abdomen parallel, almost as wide as pronotum.
No sexual differences observed. Length 3.0 mm.

Type locality. — Massett, Queen Charlotte Island, British Columbia.

Recorded distribution. — British Columbia.

Material examined. — I have seen 2 specimens from the type locality
(Fenyes collection, C.A.S.) and 3 from Nazan Bay, Atka, Aleutians, Alaska,
collected by E. C. Van Dyke in August (Fenyes collection, C.A.S.) .

Remarks. — Although this may eventually prove to be only a geographic
race of maritima, the specimens examined are quite noticeably larger, stout-
er, darker, and with a duller, more strongly granulose surface. I can find no
other consistent characters to separate the species.

PHYTOSUS Curtis

Phytosus Curtis, 1838, pi. 718; Erichson, 1840, p. 177; Kraatz, 185 3, p. 2 57, t. 3; Kraatz,

1856, p. 41; Kraatz, 1857, t. 1; Jacguelin DuVal, 1856, p. 5, pi. 3; Fauvel, 1862, p. 82;

Thomson, 1859, p. 207; Mulsant and Rey, 1872, p. 381; Ganglbauer, 1895, p. 285;

Fenyes, 1920, p. 131; Bernhauer, 1941, p. 96.
Paraphytosus Bernhauer, 1922, p. 236 (not Cameron, 1917).
Phytosus Kiessenwetter, 18 50, p. 385. ,

Subgenus Actosus Mulsant and Rey, 1872, p. 391.

Subgenus Euphytosus Bernhauer and Scheerpeltz, 1926, p. 5 52.

Subgenus Anopsisus Bernhauer, 1929, p. 187.

Generitypes. — of Phytosus, Phytosus spinifer Curtis; of Actosus, Acto-
sus nigriventris (Chevrolat) ; of Euphytosus, Euphytosus schenklingi
(Bernhauer) ; of Anopsisus, Anopsisus micro phthlalmus Bernhauer.

136 San Diego Society of Natural History

Diagnosis. — Body elongate, parallel, somewhat depressed. Head oval.
Labrum variable, truncate or slightly emarginate. Mandibles each with a
median tooth, with the tip curved ventrally. Mentum truncate. Maxillary
palpi with second segment ovoid, larger than first. Outer lobe of maxillae
with a membranous finger-like process at tip; inner lobe setose internally at
apex. Labial palpi distinctly three-segmented; the segments of about equal
length; the last segment narrowest. Ligula simple, slender, little more than
half as long as first segment of labial palpi. Eyes moderate in size, but less
than one-half the length of side of head. Antennae with first three seg-
ments elongate, seventh to tenth transverse, eleventh elongate obconical.
Anterior margin of prosternum straight. Anterior and middle tibiae with
several series of long thick setae externally which are interspersed with
the pubescence. Elytra in P. spinifer longest observed in the subtribe, con-
siderably longer than pronotum; in nigriventris not as long as pronotum.
Abdomen with first four tergites impressed at base; first four sternites con-
stricted at base.

Remarks. — The diagnosis is based entirely on Phytosus (Pbytosus)
spinifer Curtis, the generitype, Phytosus (Actosns) nigriventris (Chevro-
lat), and Phytosus (Actosns) balticns Kraatz. Phytosus, the first genus
described in the subtribe, has been a catch-all for numerous doubtful sea-
shore species, some of which were later removed to other genera. There still
remain in the genus several species which will have to be re-examined before
an adequate diagnosis of the genus can be given. I have seen too few of
the species listed for the genus to be able to give it a satisfactory evaluation.
The descriptions of most of these species are inadequate. To judge from the
descriptions, two of the subgenera, Anopsisus Bernhauer and Euphytosus
Bernhauer and Scheerpeltz (= Paraphytosus Bernhauer, not Cameron),
probably do not belong here. The former, according to Koch, is probably a
Diglotta. The latter probably does not belong to this subtribe either.

Material examined. — Phytosus spinifer Curtis: 16 specimens from
France (Fenyes, C.A.S.), 1 from Spain (Fenyes, C.A.S.), 1 from Crete
(Fenyes, C.A.S.), 1 from Senegal (Brit. Mus.), 3 from the Canary Islands,
labeled P. dimidiatus Wollaston (Brit. Mus.) and 2 which appear to be this
species labeled "Phytosus littoralis Horn" from "Gaudel I." (Fenyes C.A.S.) .
Phytosus nigriventris (Chevrolat) : 9 from France (Fenyes, C.A.S.) . Phy-
tosus balticus Kraatz: 5 from France, 1 from Crete and 2 from Tunis
(Fenyes, C.A.S.). Phytosus fenyesi Bernhauer: one specimen in very poor
condition from Senegal, identified by Bernhauer (Fenyes, C.A.S.).

Moore — Pacific Coast Phytosi 137

CAMERONIUM Koch

Cameronium Koch, 193 6, p. 202.

Generitype. — Cameronium obockianum (Fauvel).

Diagnosis. — Body elongate, parallel, subdepressed. Head oval. Infra-
orbital ridge entire. Labrum rounded at apex. Mandibles not curved vent-
rally; simple. Third segment of maxillary palpi oval; fourth narrow, short.
Inner lobe of maxillae with fine, evenly spaced, internal spines at apical half
and a group of long setae at base; outer lobe a little longer than inner with
a brush of setae at tip. Labial palpi apparently two-segmented. Ligula short,
bifid at tip in the form of a blunt Y. Eyes not prominent, occupying a little
less than half the side of head, with setae between the facets. Antennae
with first three segments elongate, tenth transverse. Pronotum quadrate.
Elytra conjointly nearly square, a little wider and considerably longer than
pronotum. Tibiae pubescent with three or four very short, stout setae on
the anterior pair. Posterior tarsus with the first four segments subequal,
fifth as long as the preceding two together. Abdomen with the first three
tergites impressed at base, the first three sternites faintly constricted.

Remarks. — The type species was originally described as a Phytosus
which it resembles rather closely, but differs in the lack of long spines on
the front and middle tibiae. The bifid ligula it shares with Thinusa, which,
however, has spinose tibiae. My description of the inner and outer lobes of
the maxillae differs somewhat from the figure given by Koch. These parts,
often difficult to see, were very clearly visible on one specimen I examined.

Material examined. — I have seen seven specimens of Cameronium
obockianum (Fauvel) from the type locality, Perim, at the mouth of the
Red Sea, collected by Fauvel (Fenyes Coll., C.A.S.). Another species, C.
flavipenne Cam. which was described from Zanzibar is said to be smaller,
more shining and differently colored.

BAEOSTETHUS Broun

Baeostethus Broun, 1909, p. 96; Fenyes, 1920, p. 130.

Generitype. — Baeosfetbus cbiltoni Broun.

No specimens of this genus have been seen, so the original description
is repeated here. Only one species, the generitype, is known.

"Body very elongate. Head subrotundate, with a short narrow muzzle.
Thorax cordate-quadrate. Elytra very short. Hind-body very elongate.
Eyes minute. Mentum very large, slightly emarginate in front. Labial

138 San Diego Society of Natural History

palpi rather short; basal 2 joints cylindric, equally elongate; 3rd slender
and nearly the length of the penultimate. Maxillary palpi setose; basal joint
small; 2nd stout and elongate, gradually thickened; 3rd inserted at apex
of the preceding one but so as to be at a right angle to it, rather longer than
2nd, gradually incrassate towards and truncate at the apex; 4th joint small,
aciculate. Mandibles stout, rather short, acutely curvate at the extremity,
with 3 inner teeth. Antennae inserted at the sides of the forehead, in
front of the eyes; basal 3 joints stout and elongate, narrowed towards the
base; 2nd articulation a little shorter than 1st, but slightly longer than 3rd;
4th oblong; 5th and 6th oviform; 7th and 8th slightly broader than pre-
ceding one; 9th and 10th subquadrate; 11th oblong-oval. Tarsi filiform,
the posterior pentamerous, intermediate quadriarticulate, the anterior seem-
ingly also 4-jointed but so short and compact and thickly setose that the
basal joints cannot be distinguished separately. Claws elongate, simple. All
the coxae elongate, prominent, and contiguous. Prosternum corneous
across the middle, membranous elsewhere. The ligula appears to be simple
and aciculate.

"Notwithstanding the elongation of the body, the metasternum is so
excessively reduced that the intermediate and posterior coxae are in actual
contact. This character of itself is distinctive.

"Baeostethus chiltoni, sp. nov.

"Subopaque, finely pubescent; head and elytra obscure infuscate red;
thorax, legs and antennae fusco- testaceous; hind-body fuscous or nigres-
cent, the segments with a short pallid basal membrane. Head broadly
rounded, somewhat depressed on the middle, closely and very minutely
punctate, with 2 small indistinct median foveae. Forehead rather abrupt-
ly narrowed, short, medially convex, nearly smooth and shining, with a
setigerous fovea at each side, truncate and with a short grey membrane in
front. Labrum prominent, rounded and bearing fine yellow setae in front.
Eyes minute, situated at the sides in front, depressed, hardly discernible.
There is no neck. Thorax widest in front, gradually narrowed backwards;
base truncate, apex feebly and broadly curvate; it is without definite lat-
eral margins; the angles are nearly rectangular; there is a feebly median im-
pression behind; its surface is finely and closely punctured, and bears slender
greyish and infuscate pubescence. Scutellum large and broad. Elytra ab-
breviated, shorter than thorax, each strongly rounded and finely margined
at the base so as to be oblique towards the suture, apices subtruncate yet
almost oblique inwardly, their sides curvedly narrowed towards the base;
their surface dull, the sculpture concealed by the pubescence, but consist-

Moore — Pacific Coast Phytosi 139

ing apparently of very minute distant granules. Hind-body very elongate,
broadly marginated, the basal 5 segments transversal, each however becom-
ing rather longer than its predecessor, 6th with short styles, 7th narrow and
testaceous, all finely and moderately closely punctured and pubescent. Legs
slender. Femora and tibiae ciliated with fine greyish setae.

"Length, iy2 lines; breadth, quite % line.

"Campbell Island.

"Named in honour of Professor Chilton, to whom we are indebted for
the discovery of this and some other species."

Judging from the above description, this genus would appear to re-
semble Amblopusa in most characters except for the very short metaster-
num (a character which it shares with Diaulota and Liparocephalus) and
the densely setose anterior tarsae. The mention of three internal teeth on
the mandible could be interpreted to mean the same as serrations between
the median tooth and the tip, particularly if only the end of the mandible
was visible in the specimen examined. No mention is made of dorsal im-
pressions on the tergites.

ANTARCTOPHYTOSUS Enderlein

Antarctophytostis Enderlein, 1909, p. 377; Fenyes, 1918, p. 10 5.
Parapbytosus Cameron, 1917, p. 12 5.
Austromalota Brethes, 1925, p. 170.

Generitype. — Antarctophytosus atriceps (Waterhouse) .

Diagnosis. — Body elongate, subparallel, depressed. Head oval; the
clypeal area depressed on each side but not on the midline, so that there is a
broadly triangular raised area with its apex at the anterior margin of the
head and its base just anterior to the anterior margins of the eyes. Infra-
orbital ridge very strong. Labrum broadly rounded. Mandibles not curved
ventrally; asymmetrical, the left simple, the right with a very short median
tooth. Mentum rather deeply, arcuately emarginate in its central two thirds.
Third segment of maxillary palpi slightly ovoid; fourth narrow, elongate.
Inner lobe of maxillae spinose internally. Labial palpi two- or three-seg-
mented, first a little longer than wide, second as wide as first and twice as
long, third narrower than second and nearly as long. Ligula, slender, simple;
twice as long as first segment of labial palpi. Eyes flat, not interrupting the
contour of head; large, occupying more than one-third of the side of head.
Antennae with the first three segments elongate, third shorter than second,
segments nine and ten transverse. Pronotum ovoid; widest at anterior
fourth; sides evenly arcuate; hind angles broadly rounded into the arcuate

140 San Diego Society of Natural History

base. Anterior margin of prosternum bisinuate. Elytra about as long and as
wide as pronotum; apices emarginate at outer angle. Tibiae setose. Posterior
tarsi with first four segments subequal, fifth as long as preceding two to-
gether. Abdomen with first four tergites impressed at base; sternites not
constricted.

Remarks. — Rather closely resembles Bryobiota in many characters but
not in appearance. Can be distinguished at once from Bryobiota by the
evenly convex dorsal surface of the basal half of the head.

Material examined. — I have seen 2 specimens of A. atriceps (Water-
house) , 1 from Kerguelen Island in the Antarctic-Indian Ocean determined
by Jeannel and 1 from Cape of Good Hope; a specimen of A. darwini Water-
house from the Faulkland Islands, determined by Dr. Cameron, and two
specimens of A. darwini from the Faulkland Islands (in the Fenyes col-
lection). I am unable to distinguish between the two species.

BIBLIOGRAPHY

The following is a selected bibliography intended to include all the
important references to the genera and subgenera of the Phytosi and to the
species of the Pacific Coast of North America. Catalogues, local lists, man-
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been seen, several were seen only in photostats of the pertinent pages.

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Koleopt. Rdsch., vol. 26, pp. 95-96.
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Moore — Pacific Coast Phytosi 141

Blackwelder, Richard E.

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1952. The generic names of the beetle family Staphylinidae with an
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1925. Un coleoptere et un diptere nouveaux de la Georgie du Sud.
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142 San Diego Society of Natural History

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359.
18 80. Short studies of North American Coleoptera. Trans. Amer.

Ent. Soc, vol. 8, pp. 163-218.
Maklin, Fredrico Guilielmo.

18 5 3. In Mannerheim, Dritter Nachtrag zu Kaefer-Fauna der Nord-

Amerikanischen Laender des Russischen Reiches. Bull. Soc.

Imp. Nat. Moscou, vol. 26, No. 3, pp. 95-269.
Morse, Silas R.

1909. The insects of New Jersey. Ann. Report of the N. J. State

Museum, p. 1-888.

144 San Diego Society of Natural History

Motschulsky, T. Victor von.

:-"1860. Enumeration des nouvelles especes de coleopteres raportees
de ses voyages. Bull. Soc. Imp. Nat. Moscou, vol. 3 3, pp.
539-588.
Mulsant, M. E., and Rey, Claudius.

::"1870. Description d'un genre nouveaux de l'ordre des coleopteres,
tribu des brachelytres, famille des aleochariens. Oposc. Ent.,
vol. 14, pp. 194-199.
1872. Tribu des brevipennes: Famille des aleochariens: Huitieme
Branche: Bolitocharaires. Ann. Soc. Linn. Lyon, ser. 2, vol. 19,
pp. 91-413, 426.
Ross, Edward S.

1953. Insects close up. Univ. Calif. Press, pp. 1-80 (figure lower left.,
p. 46, is an excellent photograph of Liparocephahis cordicollis
LeConte).
Sakaguti, Kohei.

1944. A new intertidal rove-beetle from the Pacific Coast of Japan.
Trans. Kansai Ent. Soc, vol. 14, pp. 20-21, pi. 1, fig. 1-2.
Saunders, L. G.

1929. Some marine insects of the Pacific Coast of Canada. Ann. Ent.
Soc. Amer., vol. 21, pp. 521-545, 1 fig.

Scheerpeltz, Otto.

1934. Coleopterorum catalogus . . . . , pars 130, Staphylinidae VIII,
supplementum II, pp. 1501-1831. Berlin.
Schwarz, E. A.

1910. Coleoptera of the expedition. Harriman Alaska Ser., Smith.
Inst., vol. 8, pp. 169-185.
Thomson, Carl Gustaf.

*18 59. Skandinaviens Coleoptera . . . . , vol. 1, 290 pp. Lund.
Waterhouse, Charles Owen.

1875. On the Coleoptera of Kerguelen's Island. Ent. Monthly Mag.,
vol. 12, pp. 54-57.
Waterhouse, Frederick H.

1879. Descriptions of new Coleoptera of geographical interest, col-
lected by Charles Darwin, Esq. Jr. Linn. Soc. London, vol. 14,
pp. 530-534.
Wendeler, Hans.

1930. Neue exotische Staphyliniden (17. Beitrag zur Kenntniss der
Staphyliniden) . Neue Beitr. syst. Insektenk, vol. 4, pp. 181-
192, 248-252.

Moore — Pacific Coast Phytosi 145

Wollaston, Thomas Vernon.

18 57. Catalogue of the Coleopterous insects of Madeira in the col-
lection of the British Museum, 648 pp. London.

1864. Catalogue of the coleopterous insects of the Canaries in the
British Museum, 648 pp. London.

1865. Coleoptera Atlantidum . . . . , 526— |— 140 pp. London.

Explanation of Figures
Plate 8

Figure 1. Liparoccphalus brevipennis M'ikVm, apical margin of labrum.

Figure 2. Liparoccphalus brevipennis Maklin, labial palpi and ligula.

Figure 3. Liparoccphalus brevipennis Maklin, apical margin of mentum.

Figure 4. Liparoccphalus brevipennis Maklin, outline of the body of fe-
male, dorsal view.

Figure 5. Liparocephalus cordicollis LeConte, apical margin of labrum.

Figure 6. Liparocephalus cordicollis LeConte, labial palpi and ligula.

Figure 7. Liparoccphalus cordicollis LeConte, apical margin of mentum.

Figure 8. Liparocephalus cordicollis LeConte, maxilla.

Figure 9. Liparocephalus cordicollis LeConte, outline of body of male,
dorsal view.

Figure 10. Diaulota densissima Casey, apical margin of labrum.

Figure 1 1 . Diaulota densissima Casey, labial palpi.

Figure 12. Diaulota densissima Casey, apical margin of mentum.

Figure 13. Diaulota densissima Casey, apex of abdomen of male, ventral
view.

Figure 14. Diaulota densissima Casey, outline of body of female, dorsal
view.

Figure 1 5. Diaulota fnlviventris Moore, apical margin of labrum, male.

Figure 1 6. Diaulota fulviventris Moore, apical margin of mentum, male.

Figure 17. Diaulota fulviventris Moore, apex of abdomen of male, ventral
view.

Figure 18. Diaulota fulviventris Moore, apex of abdomen of male with
genitalia extruded, lateral view.

Figure 19. Diaulota fulviventris Moore, labial palpi, male.

Figure 20. Diaulota fulviventris Moore, labial palpi, male.

Plate 9

Figure 21. Diaulota fulviventris Moore, apical margin of labrum, female.
Figure 22. Diaulota fulviventris Moore, labial palpi, female.

146 San Diego Society of Natural History

Figure 23. Diaidota fulviventris Moore, apical margin of mentum, female.

Figure 24. Diaulota fulviventris Moore, maxilla.

Figure 2 5. Diaulota fulviventris Moore, outline of body of female, dorsal

view.
Figure 26. Diaulota hartcri Moore, anterior margin of labrum.
Figure 27. Diaulota harteri Moore, labial palpi.
Figure 28. Diaulota harteri Moore, apical margin of mentum.
Figure 29. Diaulota harteri Moore, apex of abdomen of male, ventral view.
Figure 30. Diaulota harteri Moore, apex of abdomen of female, ventral

view.
Figure 31. Diaulota harteri Moore, outline of body of female, dorsal view.
Figure 32. Diaulota megacephala Moore, apex of labrum.
Figure 3 3. Diaulota megacephala Moore, labial palpi and ligula.
Figure 34. Diaulota megacephala Moore, apex of mentum.
Figure 3 5. Diaulota megacephala Moore, apex of abdomen of male, ventral

view.
Figure 36. Diaulota megacephala Moore, lateral lobe of male genitalia.
Figure 37. Diaulota megacephala Moore, first three antennal segments.
Figure 3 8. Diaulota megacephala Moore, outline of body of male, dorsal

view.
Figure 39. Diaulota vandykei Moore, apical margin of labrum.
Figure 40. Diaulota vandykei Moore, labial palpi.
Figure 41. Diaulota vandykei Moore, apical margin of mentum.
Figure 42. Diaulota vandykei Moore, apex of abdomen of male, ventral

view.
Figure 43. Diaulota vandykei Moore, first three antennal segments.
Figure 44. Diaulota vandykei Moore, outline of body of male, dorsal view.

Plate 10

Figure 45. Amhlopusa brevipes Casey, apical margin of labrum.

Figure 46. Amhlopusa brevipes Casey, apical margin of mentum.

Figure 47. Amblopusa brevipes Casey, labial palpi and ligula.

Figure 48. Amblopusa brevipes Casey, apical margin of sixth sternite of

male.

Figure 49. Amblopusa brevipes Casey, outline of body, dorsal view.

Figure 5 0. Amblopusa borealis Casey, apical margin of labrum.

Figure 51. Amblopusa borealis Casey, labial palpi and ligula.

Figure 52. Amblopusa borealis Casey, inner and outer lobes of maxilla.

Moore — Pacific Coast Phytosi 147

Figure 5 3. Amblopusa borealis Casey, apical margin of sixth sternite of

male.

Figure 54. Amblopusa borealis Casey, outline of body, dorsal view.

Figure 5 5. Bryobiota bicolor (Casey) , apical margin of labrum.

Figure 56. Bryobiota bicolor (Casey) , labial palpi and ligula.

Figure 57. Bryobiota bicolor (Casey) , apical margin of mentum.

Figure 5 8. Bryobiota bicolor (Casey), inner and outer lobes of maxilla.

Figure 59. Bryobiota bicolor (Casey) , outline of body, dorsal view.

Figure 60. Bryothinusa catalinae Casey, apical margin of labrum.

Figure 61. Bryothinusa catalinae Casey, labial palpi and ligula.

Figure 62. Bryothinusa catalinae Casey, apical margin of mentum.

Figure 63. Bryothinusa catalinae Casey, ends of mandibles.

Figure 64. Bryothinusa catalinae Casey, ends of outer and inner lobes of

maxilla.

Figure 6 5. Bryothinusa catalinae Casey, outline of body, dorsal view.

Plate 11

Figure 66. Thinusa maritima (Casey) , apical margin of labrum.
Figure 67. Thinusa maritima (Casey) , labial palpi and ligula.
Figure 68. Thinusa maritima (Casey) , apical margin of mentum.
Figure 69. Thinusa mariti ma (Casey) , maxilla.

Figure 70. Thinusa maritima (Casey) , outline of body, dorsal view.
Figure 71. Phytosus (Phytosus) spinifer Curtis, apical margin of labrum.
Figure 72. Phytosus (Phytosus) spinifer Curtis, labial palpi and ligula.
Figure 73. Phytosus (Phytosus) spinifer Curtis, apical margin of mentum.
Figure 74. Phytosus (Phytosus) spinifer Curtis, outline of body, dorsal

view.
Figure 75. Phytosus (Actosus) nigriventris (Chevrolat) , outline of body,

dorsal view.
Figure 76. Phytosus (Actosus) balticus Kraatz, outline of body, dorsal

view.
Figure 77. Antarctophytosus atriceps (Waterhouse) , apical margin of

labrum.
Figure 78. Antarctophytosus atriceps (Waterhouse), labial palpi and

ligula.
Figure 79. Antarctophytosus atriceps (Waterhouse), apical margin of

mentum.
Figure 80. Antarctophytosus atriceps (Waterhouse), outline of body.

dorsal view.

148

Moore — Pacific Coast Phytosi

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FOR EXPLANATION OF FIGURES SEE PAGE 145

Moore — Pacific Coast Phytosi

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FOR EXPLANATION OF FIGURES SEE PAGES 145 - 146

150

Moore — Pacific Coast Phytosi

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Moore — Pacific Coast Phytosi

151

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PLATE 11

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FOR EXPLANATION OF FIGURES SEE PAGE 147

TRANSACTIONS

OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY

Volume XII. No. 8, pp. 154-176, plate 12, 2 maps

MARINE PLEISTOCENE
INVERTEBRATES

FROM NEAR

PUERTO PENASCO, SONORA, MEXICO

BY

Leo George Hertlein

Associate Curator, Department of Geology, California
Academy of Sciences

AND

William K. Emerson

Museum Paleontologist, University of California Museum
of Paleontology

mus. m?.

LIBRARY

JUL 2

MRVARf
VNIVERSI1

SAN DIEGO, CALIFORNIA

Printed for the Society

June, 7, 1956

COMMITTEE ON PUBLICATION

Laurence M. Klauber, Chairman

Joshua L. Baily Charles C. Haines

Carl L. Hubbs

John A. Comstock, Editor

TABLE OF CONTENTS

Introduction 157

General description of area 158

Punta Penasco area 161

Punta la Cholla area 162

Descriptions of collecting stations 164

List of species from the Pleistocene near Puerto Penasco,

Sonora, Mexico 165

Summary - — - - 170

Bibliography 171

List of Illustrations

Index map of the Puerto Penasco area and adjacent region 159

Sketch map showing the location of the principal collecting stations 160

Plate 12. Figure 1. Terrace along beach at Punta la Cholla.
Figure 2. Cliff face of terrace.
Figure 3. Beach along north side of Punta la Cholla 174

LIBRAE

1956

HfSVJLRH

WIVt&S'iTY

MARINE PLEISTOCENE
INVERTEBRATES
FROM NEAR
PUERTO PENASCO, SONORA, MEXICO1

BY

Leo George Hertlein2 and William K. Emerson*

INTRODUCTION

This paper is based on a study of a collection containing 63 identi-
fied species and subspecies of metazoan invertebrates, mostly moliusks,
taken from low terrace deposits in the Puerto Penasco region of north-
western Sonora, Mexico. It is the first general discussion of a Pleisto-
cene faunule from that area to appear in the literature. Paleoecological
interpretations are presented in an attempt to evaluate this faunal
assemblage in terms of the known late Cenozoic distribution of the
component species.

The occurrence of fossiliferous terrace deposits along the northern
coast of Sonora, near Puerto Penasco, was briefly mentioned in a paper
by Lowe (1934: 3) concerning the Recent moliusks and in one by
Gifford (1946: 215) on the archaeology of the area. Beal (1948: 30)
observed low terraces at an elevation of about 20 feet above sea level
along most of the Sonoran coast from Guaymas (Lat. 27° 55' N.)
to the head of the Gulf of California and on some of the islands off
the coast of northern Sonora. He suggested that correlation of these
terraces with similar terrace deposits on the coast of Baja California
and the adjacent islands would be of considerable interest and would
aid in the interpretation of the diastrophic history of the Gulf of Cali-
fornia. Durham (1950) recently described the invertebrate fossils from
several terrace deposits on the east coast of Baja California and of
adjacent islands. In a paper dealing with high sea levels of the Sonoran
shore, Ives (1951 ) discussed the occurrence of the "Chione cancellata
beds" in the vicinity of Punta Penasco.

1 Contribution from the Department of Geology, California Academy of Sciences, and from

the Museum of Paleontology, University of California.
^Associate Curator, Department of Geology, California Academy of Sciences.
3Museum Paleontologist, University of California Museum of Paleontology.
4The species cited by Ives is probably referable to Chione calijorniensis Broderip.

158 San Diego Society of Natural History

A small number of fossil mollusks were collected by Professor
Gifford (1946: 215) in 1944 and 1945 from raised beaches on the
north side of Punta Penasco. An examination of these specimens led
the senior author to assign a late Pleistocene age to them. A few addi-
tinal specimens, obtained by Professor Gifford in 1948 and 1950 and
presented to the University of California Museum of Paleontology,
were examined by the junior author; this examination prompted him,
accompanied by Arthur P. Loring, Jr., and Jerry A. Powell, to visit the
area in December, 1954. The large collection of fossils and the acquisi-
tion of field data resulting from this trip, together with the specimens
and information provided by Professor Gifford, form the basis for this
paper. The senior author has collaborated in the preparation of the
manuscript and has identified the mollusks.

The writers wish to acknowledge the very helpful assistance received
in the field from Messrs. Loring and Powell and the many courtesies
extended by Professor Gifford. Acknowledgment also is due to Dr.
Robert J. Menzies of the Scripps Institution of Oceanography for infor-
mation concerning the species of fossil crab, and to Dr. John D. Soule
of the Allan Hancock Foundation, University of Southern California,
for kindly identifying the species of Bryozoa. Dr. John S. Garth of the
latter institution furnished information concerning the distribution of
decapod Crustacea in the Gulf of California region, and he also identified
the single species of fossil crab in the present collection.

GENERAL DESCRIPTION OF AREA

Fossils were collected at Punta Penasco (Lat. 31° 18' N.) and at
Punta La Cholla (Lat. 31° 20' N.), prominent headlands situated
between Bahia de San Jorge on the south and Bahia de Adair on the
north (figs. 1, 2). South of Punta La Cholla, the coast line follows a
south-southeast direction as a sandy beach and then recedes somewhat
to form a small bay, Bahia Punta Penasco (Rocky Point Bay) , on the
north side of Punta Penasco. The village of Puerto Penasco is located

5F. Shreve remarked on the origin of the beach sands of this portion of the Gulf of
California in his paper dealing with the distribution of the adjacent sandy areas on the
mainland (see Yearbook Assoc. Pac. Coast Geogr., Vol. 4, pp. 11-12, 1938).

Puerto Penasco Fossils. Hertlein & Emerson

159

Figure 1. Index map showing the general location of the Puerto Penasco
area and the adjacent region.

160

San Diego Society of Natural History

Figure 2. Sketch map showing the location of the principal collecting sta-
tions mentioned in the present paper.

Puerto Penasco Fossils. Hertlein & Emerson 161

on the south shore of the bay and is served by the Ferrocarril Fuentes
Brotantes Punta Penasco railroad, a spur line of which runs out to the
headland along the south shore of Rocky Point Bay. A paved highway
connects Puerto Penasco with the town of Sonoyta, Sonora, and extends
north, where it crosses the international boundary into Arizona. Punta
La Cholla may be reached from Punta Penasco by a graded dirt road.

PUNTA PEISrASCO AREA

Punta Penasco, or Rocky Point, is a volcanic headland of weathered
basaltic blocks and scattered pumice fragments forming a low hill which
attains a maximum elevation of 226 feet8. Remnants of a terrace deposit
abutting against the north side of the headland reach a maximum
elevation of about 20 feet. At the edge of the village, a highway cut
exposes fossiliferous sandstone 1 to 6 feet in thickness which is overlain
by 1 to 10 feet of sparsely fossiliferous cobble conglomerate, Locality
B-1397 . This very fossiliferous sand grades locally into a well cemented
coquina containing small rocks and pebbles. Between the railroad spur
tracks and the shore of the bay, Professor Gifford found a poorly con-
solidated fossiliferous sand exposed in the bank of the railroad cut,
Locality A-4878. In this vicinity, fossiliferous sands also were located
by him in a cut near an oil storage tank at an elevation of about 10 feet
above high tide level, Locality A-6602, and at an elevation about 10
feet stratigraphically higher and approximately 100 yards south of this
outcrop, Locality A-6603. The last two exposures, which were not found
by the junior author and party in December, 1954, may have been des-
troyed by building activity in the area.

On the southwest side of Punta Penasco, a wave-cut bench was
noted about 10 feet above sea level. Wave action has eroded nearly
all of the terrace sediments off the bench platform and only small patches
of well indurated fossiliferous sand remain in the tidal zone, Locality

"Anonymous, 1938, page 170.

7Locality numbers refer to the University of California Museum of Paleontology
unless otherwise stated.

162 San Diego Society of Natural History

B-1400. The sandstone grades locally into a coquina and overlies large
blocks of basalt. The fossils at this locality are very poorly preserved
due to the permeation and leaching action of water.

It is quite probable that marine waters at one time extended farther
inland than is indicated by the extent of the fossiliferous sediment in
this area. Hornaday (1908: 270) was of the opinion that in com-
paratively recent times an arm of the Gulf of California extended up
the Sonoyta valley as far as the Playa Agua Dulce. He based this
opinion partly on the occurrence at places of marine shells, citing
as examples the occurrence of Cardium elatum and "Pectunculus"
[=Glycymeris~\ gigantea near the Playa Salada below Agua Dulce.
However, as pointed out by Ives (1935: 337), evidence of former
extension of marine waters based upon isolated occurrences of marine
shells should be carefully considered, especially in regions where mollusks
were used for food and as articles of trade by the aborigines. The
present collection is composed of shells collected in place and nearly all
contain some of the cemented matrix.

PUNTA LA CHOLLA AREA

Situated approximately five miles west-northwestward of Punta
Penasco is Punta La Cholla, or Rocky Bluff, a headland composed of
a series of low granitic hills, the highest of which is 408 feet in elevation.
On the north side of the point lies a small bight which is known locally
as Bahia de Cholla. Overlying the granitic rocks exposed in the inter-
tidal zone just inside of the entrance to the bight are isolated ledges of
well indurated sandstone containing granitic pebbles, rocks up to three
inches in diameter and occasional fossils, Locality B-1394 (plate 12,
figure 3). Along the west side of the headland, numerous undercut
and partially slumped ledges of fossiliferous sandstone are exposed in
the tidal zone. Fossils were collected from well-indurated sandstone
outcropping in the tidal zone on the south side of the headland, immedi-
ately north of the large sand beach, Locality B-1398.

"Strata in southwestern Yuma County, Arizona (Lat. 33° 10' N), containing
unidentified species of barnacles as well as Corbicula(?) and other brackish water fossils
are considered to be of late Tertiary age [see E. D. Wilson, Science, New Ser., Vol. 74,
No. 1927, pp. 567-568, December 4, 1931; Arizona Bur. Mines, Bull. 134, Geol. Ser.
No. 7 (Univ. Arizona, Bull., Vol. 4, No. 2), pp. 31, 32, February 15, 1933].

'Anonymous, 1938, page 170.

Puerto Penasco Fossils. Hertlein & Emerson 163

With the exception of sandstone remnants in the tidal and infra-
tidal zones of the beach, erosion has removed the terrace sediments
from the southern and most of the western portions of the headland.
The terrace is preserved, however, along a 200 yard stretch of the north-
west coast, at a maximum elevation of 20 to 25 feet and abuts against
the headland a few hundred feet from the shore line (plate 12, figure l) .
The cliff face exposes a basal 4 to 10 feet of poorly sorted, very fossil-
iferous sand with interbedded pebbles and boulders up to two feet in
diameter, Locality B-1395 (plate 12, figure 2). The conglomerate is
covered by a thin veneer of soil in which Recent mollusks of apparent
kitchen midden origin are extremely common. The sandstone is weakly
cemented above the spray zone of the surf, and locally contains large
pockets of coquina. This locality proved to be the most fossiliferous
encountered; however, because of selective leaching of the fossils in the
surface exposures, many of the faunal elements are represented by
internal molds. In contrast, several groups, including the oysters,
pectens, thaisids, tegulids and barnacles, are well preserved. Durham
(1950: 4) commented on the prevalence of differential leaching of
fossils in some of the late Tertiary and Pleistocene formations of the
east coast of Baja California and concluded, on the basis of data from
Clarke and Wheeler (1917), that the presence of a relatively high
MgCOs content in the fossils may be a primary prerequisite for their
preservation. Similar differential leaching of fossil shells occurring
under desert conditions at Puerto San Bartolome (Turtle Bay) on the
west coast of Baja California was mentioned by Jordan and Hertlein
(1926: 414).

10R. L. Ives (1951) mentioned the occurrences of two well-defined high sea level
terraces along this portion of the Sonoran coast, one at 75 feet and one at 115 feet in
elevation, the latter dated as possibly corresponding to the Sangamon or Yarmouth
stages, Pleistocene. He also mentioned that other terraces occur along this coast varying
from heights of 250 feet above present sea level to perhaps 30 feet below it. He sug-
gested the possibility that some of these may be detached, possibly faulted, portions of
the main wave cut bench occurring at an elevation of 115 feet.

During the field work upon which the present paper is based, emphasis was directed
toward tracing fossiliferous strata and collecting the embedded fossils. The only terrace
which received particular attention from the collectors was the one at 20 to 25 feet in
elevation. No fossiliferous exposures were noted at elevations greater than 25 feet above
sea level.

"Gifford (1946, page 216) listed eight species of mollusks, all of which are living
in the adjacent waters, from kitchen middens on the north side of Punta La Cholla.

164 San Diego Society of Natural History

DESCRIPTIONS OF COLLECTING STATIONS

B-1394 Isolated ledges of well-indurated fossiliferous sandstone con-
taining granitic pebbles and rocks overlying granitic rocks exposed
in the intertidal zone just inside the entrance to the bight at Punta
La Cholla, Sonora, Mexico. W. K. Emerson and A. P. Loring, Jr.,
collectors; Dec. 19, 1954.

B-1395 Poorly-cemented fossiliferous sandstone, 4 to 10 feet in thick-
ness and overlain by 2 to 10 feet of conglomerate, outcropping in
terrace face exposed as sea cliff along a 200 yard stretch of the
northwest side of Punta La Cholla, Sonora, Mexico. Principal
fossil locality. W. K. Emerson, A. P. Loring, Jr., and J. A. Powell,
collectors; December 19-20, 1954.

B-1397 Fossiliferous sandstone 1 to 6 feet in thickness exposed in
highway cut at edge of village of Punta Penasco, Sonora, Mexico;
W. K. Emerson, A. P. Loring, Jr., and J. A. Powell, collectors;
December 21, 1954.

B-1398 Well-indurated fossiliferous sandstone outcropping in inter-
tidal zone on the south side of the headland immediately north of
a large sandy beach at Punta La Cholla, Sonora, Mexico. W. K.
Emerson and A. P. Loring, Jr., collectors; December 21, 1954.

B-1400 Well-indurated fossiliferous sand overlying blocks of basalt
on terrace on the southwest side of Punta Penasco, Sonora,
Mexico. W. K. Emerson and A. P. Loring, Jr. collectors; Decem-
ber 21, 1954.

Collecting stations at Punta Penasco, near Puerto Penasco, Muni-
cipio de Caborca, Sonora, Mexico, at edge of town, between railroad
spur tracks and Rocky Point Bay, Prof. Edward W. Gifford, collector :

A-4878 Fossiliferous stratum exposed in railroad cut at track bed
level; rock type: poorly consolidated sands. December, 1948.

A-6602 Fossiliferous stratum exposed at cut along oil storage tank,
approximately 10 feet above high tide level; rock type: unconsoli-
dated sands. March 3, 1940.

A-6603 Fossiliferous stratum exposed approximately 100 yards south
of Loc. A-6602 and about 10 feet higher stratigraphically than
Loc. A-6602; rock type: locally consolidated sands, grading into
a coquina. March 3, 1950.

Puerto Penasco Fossils. Hertlein & Emerson

165

LIST OF SPECIES FROM THE PLEISTOCENE NEAR
PUERTO PENASCO, SONORA, MEXICO

PELECYPODA

Anemia peruviana d'Orbigny

Apclymetis ci. A. cognata

Pilsbry & Vanatta

Area gradata Broderip & Sowerby

Area solida Sowerby

Braehidontes multiformis Carpenter15..

Cardium datum Sowerby

Cardium procerum Sowerby ...

Chama jrondosa mexicana Carpenter...

Chione ealijorniensis Broderip16

Chione gnidia Broderip & Sowerby

Codakia distinguenda Tryon

Corbula nasuta Sowerby

Diplodonta orbella Gould

Donax gracilis Hanley

Dosinia ponderosa Gray

Glans affinis calijornica Deshayes

Glycymeris maculata Broderip

Glycymeris multieostata Sowerby

Hormomya adamsiana Danker

Isognomon chemnitzianum d'Orbigny.

Lithophaga sp. (impression)

Lueina lampra Dall

Megapitaria sqiutlida Sowerby

Modiolus eapax Conrad

Ostrea angelica Rochebrune

Ostrea conchaphila Carpenter

Ostrea palmula Carpenter

Peeten circularis Sowerby

Pecten vogdesi Arnold

Protothaca grata Say

Semele flavescens Gould

Semele guaymasensis Pilsbry & Lowe..
Tagelus californianus Conrad

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x

x
x

X

X

X
X
X
X
X
X

X
X

x
x

x

X
X
X
X
X

12Kino Bay, Sonora, Mexico.
13Tepoca Bay, Sonora, Mexico.

14

15

Georges Island, Gulf of California.

A single, somewhat eroded valve is, with some doubt, referred to this species.
18AIso from Pleistocene at Punta Penasco, Sonora, Mexico. Loc. 32112 (C.A.S. ).

166

San Diego Society of Natural History

GASTROPODA

Acanthina angelica I. S. Oldroyd

Acanthina muTicata tuber culata Gray ..

CA
2

c*
CQ

X
X

5

X
X
X

CO

o

5

o
o
"*•

CQ

CO
CO

<

s

<

o

<

X

Recent at Pta.

Penasco or ad-
jacent region
(12,13,14)

X

X

Acmaea mesoleuca Menke

X

Aletes sp

X

cf.

X
X

C onus regularis Sowcrby

X

X

Crepidula sp

Cypraea ci C. annettae Dall

X

X
X

X

Murex ci. M. erythrostomus Swainson.
A4urex radix nigritus Philippi

X

X

cf.

X
X
X
X
X

cf.

?

X
X

Nassarius leucops Pilsbry 6C Lowe

X

Nerita scabricosta Lamarck

X

X

Oliva incrassata Solander

X

X

X

Oliva spicata Bolten

X

Olivella ci. 0. dama Mawe

X

Polinices reclusianus Deshayes

X

X

X

Polimccs uber Valenciennes

X
X

X

Pyrazus humboldti Valenciennes

Solenosteira ci. S. pallida
Broderip & Sowerby..

X
X
X
X
X

X

Strombus galeatus Swainson

X

Tegula cf. T. mariana Dall

X

Tegula rubroflammulata Koch

X

X

12
X

Tegula rugosa A. Adams

X

Thais haemastoma biserialis Blainville..

X

IS

X

Turbo fluctuosus \C^ood

X

X .

X

X

Turritella gonostoma Valenciennes

ECHINOIDEA
Encobe so

X

X

x

X

11
X

"Recent range: Point Conception, California, to Mazatlan, Mexico, and south to
Peril and the Galapagos Islands in 2 to 78 fathoms (Osburn, 1950: 54).

18Recent range: known only from Mazatlan, Mexico, beach (type locality), and
Panama (Osburn, 1950: 104).

"Recent range: West America: San Felipe, Baja California, to Tepoca Bay,
Sonora, Mexico, to the island of La Plata, Ecuador, in 2 to 45 fathoms. Cape Blanco,
west Africa (type locality). (Osburn, 1950: 31).

20Recent range: Angel de la Guardia Island in the Gulf of California to Chile and
the Galapagos Islands. Originally described by Busk from Chiloe Island, Chile, and
later cited by him as occurring at Mazatlan, Mexico. It occurs commonly in the Galapagos
Islands, according to Osburn (1952: 467).

21Recent range: Magdalena Bay to San Felipe, Baja California, and south to Las
Vacas, near Capon, Peru, and the Galapagos Islands. Type locality: Mazatlan, Mexico
(Rathbun, 1930: 550).

Puerto Penasco Fossils. Hertlein & Emerson

167

BRYOZOA

Antropora tincta Hastings17

Caleschara mexlcana Osburn

ON

3

XA
ON
rr\

CO

X
X .

X .
X .

X .

X
X

X

CQ

00

c6

o
o

c6

00

00

<

X

O
VO

<

O

o

<

Recent at Pta.

Penasco or ad-
jacent region
(12,13,14)

Conopeum commensale

Kirkpatrick & Metzelaar19

Hippopodinella adpressa Busk"0

CRUSTACEA
Decapoda
Eriphia squamata Stimpson21

Cirripedia
Balanus cf. B. amphitrite Darwin var.
Balanus sp

X

X

Tetraclita squamosa cf. T. s.

stalactifera form confinis Pilsbry

VERTEBRATA
Fish vertebra

X

X

X

.:...

14
X

This list contains 56 species and subspecies of identified mollusks,
one echinoid, 4 bryozoans, 2 barnacles, 1 crab, and one fish vertebra.
The mollusks, because of their large number and known ranges, furnish
the basis for most of the present discussion of this Pleistocene faunule.

Of the total of 56 species and subspecies of mollusks, five, because
of imperfect preservation, are not identified with certainty, but are com-
pared to known species. Three others are cited only as to genera.

All except four of the total assemblage are known to occur at the
present time in the Puerto Penasco area or in adjacent waters (Lowe,
1935; Keen, 1947). The four species which have not been recorded
from the region north of Angel de la Guardia Island do occur in the
Gulf south of that island, and it is quite probable that all these species
may be found to occur with the others at or near Punta Penasco. All
the non-molluscan species in this faunule also have been recorded living
in the Gulf of California. The general assemblage consists of rock
dwellers and inhabitants of sandy beaches representing near-shore con-
ditions similar to those existing in that region at the present time.

Cooper, in 1895 (p. 37), mentioned: "From small collections
hitherto made in the northern end of the Gulf, quoted by Carpenter or
Stearns, it appears that the species found there are more largely of the
temperate fauna, many of them being identical with those from the
same latitude on the west coast of the peninsula. This seems to indicate

168 San Diego Society of Natural History

that the dividing ridge, now 3,000 feet or more in altitude, was crossed
by one or more channels within geologically recent times." Steinbeck
and Ricketts (1941: see pp. 177-179, 306, 307) stated (p. 307) : "The
faunal differences between the northern and southern portions of the
Gulf are well marked in most groups." However, on the preceding
page the latter authors stated, concerning the fauna of the Gulf, that
with the exceptions of the sponges and tunicates, "there are but slight
affiliations with animals of the north temperate zones . . . Garth
(oral communication)* reported a decided difference in the decapod
crab fauna in the northern and southern portions of the Gulf of
California. These and other observations have led to at least two theories
to account for such a supposed distribution. One of these assumes
submergence of portions of Baja California during at least a portion of
Pleistocene time, thus permitting free movement of marine animals
between the waters of the open Pacific and the Gulf. The other theory
assumes that cooling of marine waters during early Pleistocene time
was sufficient to enable mollusks and other animals from southern
California to migrate around Cape San Lucas and then move northward
to the waters at the head of the Gulf of California where they became
isolated on account of the later rise in temperature of the waters about
the southern portion of the peninsula.

Wittich (1920: 123) mentioned the presence of "subfossil" marine
shells in the peninsula at elevations of 1,052 meters above sea level, and
Beal (1948: 28-33) mentioned observing such occurrences to heights
of nearly 2,000 feet." Beal, after a study of the evidence involved, con-
cluded that it was probable that at least portions of the peninsula were
submerged during Pleistocene time from 1000 to 1800 feet. Further
investigation will be necessary before detailed information is available
concerning such possible extent and pathway of marine waters across
the peninsula during Pleistocene time.

It is true that there are certain species of mollusks in the waters at
the head of the Gulf of California whose affinities are believed to lie
with species on the Pacific coast of southern California and northern

21"The shells were occasionally in the form of terrace deposits, though at higher
elevations they were usually found loose, and scattered over such wide areas and in such
quantities as to indicate strongly that they were not carried there by man." (Beal,
1948:28). It is possible that some of the latter may prove to be the remains of
aboriginal kitchen middens.

*A publication by Garth (1955) containing a discussion of this subject appeared
after the present paper was submitted for publication. A reference to his paper is cited
in the bibliography.

Puerto Penasco Fossils. Hertlein & Emerson 169

Baja California. Such species include Panope globosa Dall, Acmaea
dalliana Pilsbry, Acanthina angelica I. S. Oldroyd and Calliostoma
gemmuloides Lowe. However, the geographic and geologic ranges of
many such species are imperfectly known at the present time. Further-
more, some species which occur in those waters which may appear to
lend a temperate aspect to this faunal assemblage, such as Cardium
datum Sowerby, Chione californiensis Broderip, Chione fluctifraga
Sowerby, Modiolus capax Conrad, Tagelus californianus Conrad,
Crucibulum spinosum Sowerby, Marginella calif ornica Tomlin, Polinices
reclusianus Deshayes and Ischnochiton "acrior Carpenter,"* are in fact
subtropical species which range north to southern California. These
species range farther south of Cedros Island (a general dividing point
between subtropical and warm temperate marine mollusks) than they
do north of it. Also, most of them attain a larger size in the warmer,
more southern waters.

Steinbeck and Ricketts remarked on the occurrence of the common
California shore crab, Pachygrapsus crassipes Randall, in the waters of
the northern portion of the Gulf of California. This occurrence, how-
ever, is not especially significant because this species also is known to
occur at various localities22 in the southern portion of the Gulf.

The knowledge of the diastrophic and faunal history of Baja
California during the Pleistocene is very incomplete at the present time.
It is possible that further investigations may reveal the presence of
cool-water molluscan faunas of Pleistocene age south of Cedros Island
and in the Gulf of California, but at the present time none are known.
It has already been pointed out that, after a trip to Magdalena Bay
in 1925, E. K. Jordan (1936: 109) was convinced that only one faunal
zone (not two, one representing warm the other cool water conditions as
suggested by him in 1924), occurs at that locality and is of late Pleis-
tocene age. He concluded that the assemblage of fossils at that locality
is one which lived under comparatively warm-water conditions at least
no cooler than those now prevailing at Magdalena Bay.

*According to Berry, records attributed to Ischnochiton acrior Carpenter from the
Gulf of California are applicable to Stenoplax (Sienoradsia) magdalenensis (Hinds,
1844) and those from southern California to Stenoplax (Stenoradsia) heathiana S. S.
Berry (see Proc. Malacol. Soc. London, Vol. 26, Pt. 6, pp. 161-166, pis. 4, 5, figs.
1-6 in text, January, 1946).

"Dr. John S. Garth has informed us that this species also has been collected at the
following localities in the Gulf of California: Agua Verde Bay, Puerto Escondido,
Willards Point, Angel de la Guardia Island, Pond Island, and Ensenada de San
Francisco, Sonora.

170 San Diego Society of Natural History

The fossil mollusks from the Puerto Pefiasco area show no marked
differences from those now living there, nor do they differ from those
occurring in that portion of the Gulf south of Angel de la Guardia and
Tiburon Islands. Many of these species range south to Panama and
even to northern Peru. There is no evidence indicated by these fossils
to suggest any essential differences in climatic conditions existing at the
time of deposition of the enclosing sediments with conditions now pre-
vailing at that locality.

Many of the species of fossil mollusks from Punta Penasco lived
in the Gulf of California region during Pliocene and Pleistocene times.
At least 26 have been recorded from beds of Pliocene age along the
east coast of Baja California or on islands in the Gulf, and 41 from
beds of Pleistocene age. In common with other molluscan assemblages
of Pleistocene age from west Mexico, 7 occur at San Quintin, 41 at
Magdalena Bay, 10 at the Tres Marias Islands, and 15 in the Colotepec
formation in Oaxaca. Undoubtedly the large number of species in
common with the Pleistocene fauna at Magdalena Bay reflects similar
depositional environments and also is due in part to the very large
number of species recorded from those deposits.

The two species of barnacles cited in the present paper are known
to occur in the northern portion of the Gulf at the present time. Four
species of Bryozoa are present in this collection. One is known to range
from San Felipe, Baja California, near the head of the Gulf of Cali-
fornia, to Ecuador, another from Mazatlan, Sinaloa, Mexico, to Pan-
ama, the third from Point Conception, California, to Mazatlan, Mexico,
and south to Pern and the Galapagos Islands, and the fourth from
Angel de la Guardia Island in the Gulf of California to Chile and the
Galapagos Islands. The single species of crab, identified by Dr. John
S. Garth as Eriphia squamata Stimpson, has been recorded as ranging
at the present time from Magdalena Bay to San Felipe, Baja California,
and south to Las Vacas, near Capon, northern Peru.

SUMMARY

The present paper is based on field observations and study of
collections of invertebrate fossils from the vicinity of Puerto Penasco,
Sonora, Mexico, northeastern shore of the Gulf of California. These
fossils occur in moderately indurated sands, varying from 3 to 20 feet
in thickness, locally grading into deposits of coquina. These sediments

Puerto Penasco Fossils. Hertlein & Emerson 171

are generally flat-lying or only slightly inclined and lie disconformably
upon blocks of basalt at Punta Penasco and upon quartz diorite at Punta
La Cholla.

The present study leads to the conclusion that the fossil mollusks,
barnacles and bryozoans from Puerto Penasco, near the head of the
Gulf of California, are of late Pleistocene age. All of the species of
mollusks have been found living at the present time in the Gulf of
California, and all but five of these mollusks occur in the vicinity of
Punta Penasco. The single species of fossil crab has been recorded
living in the waters between San Felipe, Baja California, and Peru.
The three species of Bryozoa in the collection have been recorded living
in the Gulf of California at the present time, and all range south as far
as Panama or Ecuador, one from Point Conception, California, to Peru
and the Galapagos Islands. There are no elements in the fossil fauna
which would indicate essential differences of paleoecological or paleo-
temperature conditions between the Pleistocene depositional environ-
ment and the conditions existing at the present time in the vicinity of
Puerto Penasco.

BIBLIOGRAPHY

Anonymous

1938. Sailing Directions for the West Coasts of Mexico and
Central America. Hydrographic Office, No. 84 (Wash-
ington D. C), Ed. 8, 1937, pp. I-VII, 1-430, 2 charts.

Beal, C. H.

1948. Reconnaissance of the Geology and Oil Possibilities of Baja
California, Mexico. Geol. Soc. Amer., Mem. 31, pp. I-X,
1-138, pis. 1-11, 1 map, December 1.

Cardwell, L.

1947. Deep-Sea Fishin' Hole. Arizona Highways, Vol. 23, No.
11, pp. 32-35, 5 illustrations (halftone), November. [Popu-
lar account].

Clarke, F. W., and Wheeler, W. C

1917. The Inorganic Constituents of Marine Invertebrates. U. S.
Geol. Surv., Prof. Paper 102, pp. 1-56.

172 San Diego Society of Natural History

Cooper, J. G.

1895. Catalogue of Marine Shells, collected chiefly on the Eastern
Shore of Lower California for the California Academy of
Sciences during 1891-2. Proc. Calif. Acad. Sci., Ser. 2,
Vol. 5, pp. 34-48, May 21.

Durham, J. W.

1950. 1940 E. W. Scripps Cruise to the Gulf of California, Part
2, Megascopic Paleontology and Marine Stratigraphy.
Geol. Soc. Amer., Mem. 43, pp. I-VIII, 1-216, pis. 1-48,
10 tables, August 10.

Garth, J. S.

1955. The Case for a Warm-Temperate Marine Fauna on the
West Coast of North America. Essays in the Natural
Sciences in honor of Captain Allan Hancock on the Oc-
casion of his Birthday July 26, 1955. (Los Angeles : Univ.
Southern California Press), pp. 19-27, November 8.
Gifford, E. W.

1946. Archaeology in the Punta Penasco Region, Sonora. Amer.
Antiquity, Vol. 11, No. 4, pp. 215-221, 1 fig., April.

Hornaday, W. T.

1908. Camp-Fires on Desert and Lava. (Chas. Scribner's Sons:
New York), pp. I-XIX, 1-366, 12 illustrations, 2 maps.

Ives, R. L.

1935. Recent Vulcanism in Northwestern Mexico. Pan- Amer.
Geol., Vol. 63, No. 5, pp. 335-338, pi. 32, 1 fig., June.

1951. High Sea Levels of the Sonoran Shore. Amer. Jour. Sci.,
Vol. 249, pp. 215-223, pis. 1-2, figs. 1-3, March.

Jordan, E. K.

1936. The Pleistocene Fauna of Magdalena Bay, Lower Cali-
fornia (Introduction by Leo George Hertlein). Contrib.
Dept. Geol. Stanford Univ., Vol. 1, No. 4, pp. 103-174,
pis. 17-19, November 13.

Jordan, E. K., and Hertlein, L. G.

1926. Expedition to the Revillagigedo Islands, Mexico, in 1925,

VII. Contribution to the Geology and Paleontology of the

Tertiary of Cedros Island and Adjacent parts of Lower

California. Proc. Calif. Acad. Sci., 4th Ser., Vol. 15, No.

14, pp. 409-464, pis. 27-34, 1 fig. in text, July 22.

Puerto Penasco Fossils. Hertlein & Emerson 173

Keen, A. M.

1947. Species of Mollusks collected by the Stanford Expedition,
1941, to be added to the List published by H. N. Lowe,
1935. Minutes Conch. Club South. Calif., No. 75, pp. 3^1,
December. [This list appeared without indication of author-
ship, an oversight of the editor, according to A. M. Keen
(written communication), September, 1954}.

Lowe, H. N.

1934. On the Sonoran Side of the Gulf. Nautilus, Vol. 48, No.
1, pp. 1-4, July 10; No. 2, pp. 43-46, October 15.

1935. New Marine Mollusca from West Mexico, together with
a List of Shells collected at Punta Penasco, Sonora, Mexico.
Trans. San Diego Soc. Nat. Hist., Vol. 8, No. 6, pp.
15-32, pis. 1-4, March 21.

OSBURN, R. C.

1950. Bryozoa of the Pacific Coast of America. Part 1, Cheilo-
stomata-Anasca. Allan Hancock Pac. Exped., Vol. 14, No.
1, pp. 1-269, pis. 1-29, June 20.

1952. Bryozoa of the Pacific Coast of America. Part 2, Cheilo-
stomata-Ascophora. Allan Hancock Pac. Exped., Vol. 14,
No. 2, pp. 271-611, pis. 30-64, March 20.

Rathbun, M. J.

1930. The Cancroid Crabs of America of the Families Euryalidae,
Portunidae, Ateiecyclidae, Cancridae and Xanthidae. U. S.
Nat. Mus., Bull. 152, pp. I-XVI, 1-609, pis. 1-230, text
figs. 1-85, May 29.

Steinbeck, J., and Ricketts, EL F.

1941. Sea of Cortez. (Viking Press : New York) , pp. I-X, 1-598,
pis. 1-40, December.

Wittich, E.

1920. La Emersion Moderna de la Costa Occidental de la Baja
California. Mem. y Rev. Soc. Cient. "Antonio Alzate",
Vol. 35, Nos. 3-4, pp. 121-144, pis. 14-23, 1 fig. in text,
August.

174

San Diego Society of Natural History

Explanation of Plate 12

Figure 1. Terrace along beach at Punta La Cholla, showing basal sandstone
overlain by a conglomerate, Locality B-1395.

Figure 2. Qiff face of terrace material composed of fossiliferous, poorly
sorted sandstone containing interbedded pebbles and rocks and
overlain by a conglomerate composed of granitic rocks and
boulders, Locality B-1395.

Figure 3. Beach along north side of Punta La Cholla, inside entrance to
Bahia La Cholla, showing residual ledges of fossiliferous sand-
stone overlying granitic rocks exposed in the tidal zone, Locality
B-1394.

Puerto Penasco Fossils. Hertlein & Emerson

175

Plate 12

TRANSACTIONS

OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY
Volume XII. No. 9, pp. 177-180 June 11, 1956

ADDITIONAL NOTES ON THE PLIOCENE AND

PLEISTOCENE FAUNA OF THE HauTcS^

TURTLE BAY AREA, \ LIBRARY

BAJA CALIFORNIA, MEXICO ... ^

by I mm

BY | WMERSI

E. P. CHACE

Curator of Conchology, San Diego Society of Natural History

During July of 1954 the research ship ORCA, of the J. W. Sefton
Foundation, made a trip to Port San Bartolome (Turtle Bay), Baja
California, and I was included in the group aboard as conchologist.
The ship was in Turtle Bay from July 15 to 18.

On the first morning, I landed on the beach at the southeast section
of the bay, where I sought marine shells, but found few. I then made
a trip into the sand dunes back of the beach, looking for land snails.
There I found many dead shells of Micrarionta levis (Pfr.), but no
live specimens. Some fragments of fossil pectens were also observed,
and, following that indication of fossils, I located a small exposure of
Pliocene shells on the side of a low ridge, and collected a few pounds
of material.

In 1926 Jordan and Hertlein reported on an exposure of fos-
siliferous Pliocene strata southeast of the bay, and the spot which I
found is probably the extreme western end of that exposure.

Jordan, E. K., and Hertlein, L. G. Contributions to the Geology and Paleontology of
the Tertiary of Cedros Island and Adjacent Parts of Lower California. Proc. Calif.
Acad. Sci., (ser. 4) vol. 15, no. 14, pp. 409-464, pis. 27-34, 1926.

178

San Diego Society of Natural History

In 1933 Dr. Hertlein published an additional report on the Plio-
cene fauna of this region".

The following species are represented in the material taken from
the small Pliocene exposure herein mentioned.

PELECYPODA

Ostrea megodon Hani.
Ostrea vespertina Conr.
Pecten bellus hempbillii Dall
Pecten callidus Hertl.

Pecten cf. P. cerrosensis Gabb
Pecten cf. P. cristobalensis Hertl.
Pecten cf. P. bake'i Hertl.

ECHINODERMATA

Dendraster ashleyi Arn.
Dendraster gibbsi bumilis Kew

On July 16 a landing was made at the southwest corner of the
bay, and here an apparently unreported exposure of Pleistocene fossils
was found. This second locale was at the south end of the hills on
the peninsula between the bay and the ocean.

Fossils were collected along the side of the hill for several hundred
feet. The exposure is about one-fourth mile from the tide line, the top
being about 50 feet above it. The total thickness was not determined.

The species found were:

PELECYPODA

Area reeveana D'Orb.
Area solida Sby.
Area gradata Brod. & Sby.
Ostrea cumingiana Dkr.
Pecten circularis Sby.
Anomia peruviana D'Orb.
Pseudochama exogyra (Conr.)
Lucina nuttalli Conr.
Diplodonta orbella (Gld.)

Tracbycardium procerum (Sby.)
Chione californiensis (Brod.)
Cbione gnidia Brod. & Sby.
Megapitaria squalida (Sby.)
Prototbaca grata (Say)
Apolymetis biangulata (Cpr.)
Semele flavescens (Gld.)
Crassinella cf. C. varians (Cpr.)
Pholad sp. ? (a fragment)

"Hertlein, Leo George. Additions to the Pliocene Fauna of Turtle Bay, Lower
California, with a Note on the Miocene Diatomite. Jour. Paleo. vol. 7, no. 4, pp. 438-
441, December, 1933.

Fossils of The Turtle Bay Area. E. P. Chace

179

GASTROPODA

Terebra armillata Hds.
Conus californicus Hds.
Conus fergusoni Sby.
Conus perplexus Sby.
Conus purpurascens Brod.
Turricula maculosa (Sby.)
Cancellaria cassidiformis Sby.
Oliva incrassata (Sol.)
Oiiva davisac Durham
Olivella biplicata (Sby.)
Macron aethiops (Rve.)
Nassarius tegulus (Rve.)
Purpura f estiva (Hds.)
Eupleura triquetra (Rve.)
Thais biserialis (Blv.)
Acanthina lugubre (Sby.)

Vitidaria salebrosa (King)
Cypraea arabicula Lam.
Cypraea annettae Dall
Bursa calif ornica (Hds.)
Strombus granulatus Swain.
Truncatella calif ornica Pfr.
Hipponix antiquatus (Linn.)
Crepidula onyx Sby.
Crucibulum imbricatum (Sby.)
Crucibulum spinosum (Sby.)
Polinices recluzianus (Desh.)
Lottia gigantea Gray
Astraea undosa (Wood)
Tegula aureotincta (Fbs.)
Tegula gallina (Fbs.)
Turbo fluctuosus Wood

ARTHROPODA

Tetraclita cf. T. squamosa rubescens Darwin

Many of the shells had weathered out of a stratum of sandy marl
and had rolled, unbroken, down the slope. More time and some digging
might have turned up other species. All the species on this list are
members of the recent fauna of Baja California; 25 of them with ranges
extending from nearby Cedros Island south; 4 of them with ranges
extending north from Cedros Island, and the other 15 species are found
both above and below Cedros Island. Apparently climatic conditions
when this Pleistocene deposit was formed were similar to those ex-
isting today.

About one mile east of the village a large outcrop of rock juts into
the bay. I visited this point to collect recent marine shells and there I
observed some large blocks of rock containing fossil shells which had
fallen from a point higher up. Time did not permit breaking up this
rock to obtain specimens, and no material was brought in. This loca-
tion awaits investigation at some future date.

I wish to extend thanks to Dr. L. G. Hertlein for his valued as-
sistance in the identification of the material listed in this report; also
to Dr. John A. Comstock for helpful suggestions in the preparation of
this paper.

wus. cw„ mi
JUL 2 195C

HARVARD

HARVARD
UNIVERSITY

TRANSACTIONS

OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY
Volume XII. No. 10, pp. 181-205, 1 map, plate 13

UPPER PLEISTOCENE MOLLUSCA

FROM POTRERO CANYON,
PACIFIC PALISADES, CALIFORNIA

BY

James W. Valentine

Department of Geology
University of California, Los Angeles

SAN DIEGO, CALIFORNIA

Printed for the Society

July 2, 1956

COMMITTEE ON PUBLICATION

Laurence M. Klauber, Chairman
Joshua L. Baily Charles C. Haines

Carl L. Hubbs
John A. Comstock, Editor

•■mruti

JUL 2 5 195S

wmasrr

TABLE OF CONTENTS

Index map of Pacific Palisades 184

Introduction 183

Previous work 185

Present work 187

Acknowledgments 187

Stratigraphy of trie terrace deposits 188

Paleoecology 189

Habitat 189

Temperature 190

Inferred depositional environment 192

Age and faunal affinities 192

Checklist of fossils 194

Key 194

Systematic checklist v 194

References 202

Plate 13 205

Figure 1. Index map of a portion of Pacific Palisades, California, showing
the locality from which the Clark collection was recovered (UCLA
Loc. no. 3225) and its relation to the shoreline during maximum
sea stand on the Dume terrace platform.

UPPER PLEISTOCENE MOLLUSCA

FROM POTRERO CANYON,
PACIFIC PALISADES, CALIFORNIA

BY

James W. Valentine

INTRODUCTION

Fossiliferous marine terrace sands of Upper Pleistocene age are
exposed near the head of Potrero Canyon, Pacific Palisades, California
(figure 1 ) . Tens of thousands of mollusks from this deposit, collected
over a period of years by Dr. F. C. Clark, were acquired by the Depart-
ment of Geology, University of California, Los Angeles, in January,
1943. The preparation of homesites and other construction in Potrero
Canyon has caused removal or burial of the most fossiliferous portions
of the terrace sands, so that Clark's original locality is destroyed, and
his collection cannot now be duplicated. It is therefore desirable to
record this large assemblage, which contains 128 species and varieties
previously unreported from the Upper Pleistocene of Potrero Canyon.

Previous Work

Ralph Arnold was apparently the first to assign strata at Potrero
Canyon to the Upper Pleistocene; he referred to the soft, rudely
stratified nonmarine deposits forming much of the Palisades at Potrero
Canyon and southward as probable equivalents of the Upper San Pedro
series, "from lithological and stratigraphical reasons" (1903, p. 56).
This correlation cannot be validated at present, although it is likely
that some of the nonmarine terrace cover is of Upper Pleistocene age.

Marine deposits that are now considered Upper Pleistocene were
first reported from the Santa Monica Mountains in 1904 by Rivers,
who mentions "Quaternary beach gravels" as capping strata that he
called Pliocene. Rivers described two mollusks, Carolina telemus var.
tricuspids and Strombiformis raymondi, from the Upper Pleistocene.
It is probable that these came from Potrero Canyon.

186 San Diego Society of Natural History

Following Rivers' paper, several records of species from Upper
Pleistocene deposits at "Santa Monica" appear in the literature. These
forms are with but one exception (Anachis lineolata) present in the
Clark collection, and are presumed to be from Potrero Canyon. The
earliest of these papers is by Raymond (1906), who recorded varieties
of Megasurcula carpenteriana from the "Pleistocene of Santa Monica",
attributing the collection to Rivers. Bartsch described Alabina monicensis
from the "Upper San Pedro series at Santa Monica, California" in
1911, and in 1917 described a new species of Balcis (monicensis) and
recorded three additional species of this genus (micans, oldroydi, and
rutila) from the same locality. The first chitons were reported from
"Pleistocene deposits of the Santa Monica Hills" by Chace (1917),
who recorded 7 identified forms collected by Clark. More gastropods
were described from the "Upper Pleistocene of Santa Monica" by T. S.
Oldroyd in 1921 (Anachis minuta = A. lineolata, and Epitonium
clarki), and a record of Acteocina infrequens by Dall (1922), from
an unspecified locality in Santa Monica, may belong here.

The earliest specific reference to an Upper Pleistocene fossil
locality in Potrero Canyon appears to have been made by Berry (1922,
p. 412, as "Long Wharf Canyon"). Working with material supplied
by Clark, he revised and expanded Chace's list of chitons to 9 species.
It remained however for Hoots to locate accurately the Potrero Canyon
exposures on a map (1931, pi. 16), accompanied by the most complete
description of the fauna yet published (Woodring in Hoots, 1931, pp.
121-122). Woodring listed 121 forms, of which 89 were specifically
identified. The fauna was interpreted as a warm-water assemblage, cor-
relative with the Upper San Pedro of Deadman's Island (Palos Verdes
sand) . Differences between the San Pedro and Potrero Canyon Upper
Pleistocene assemblages were ascribed to biofacies differences.

A considerable number of species from the Upper Pleistocene of
"Santa Monica", presumably Potrero Canyon, were included among
the Pleistocene records compiled in Grant and Gale's famous catalogue
(l93l). These records are based on specimens in the Oldroyd and
Clark collections (now at Stanford University and the University of
California, Los Angeles, respectively). More recently, Woodring has
re-emphasized the southern aspect of the Potrero Canyon fauna while
affirming his earlier correlations (in Woodring, Bramlette, and Kew,
1946, p. 106).

Potrero Canyon Pleistocene. J. W. Valentine 187

Present Work

Preparation and identification of the F. C. Clark collection at
U.C.L.A. was begun by Mr. E. H. Quayle in 1944, but was put aside
due to the press of other duties. This work was completed by the
writer in 1955 during the course of studies of Pleistocene molluscan
faunas. The material received from Dr. Clark included Pleistocene
mollusks from localities other than Potrero Canyon, which had un-
fortunately become mixed with a few of the Potrero Canyon fossils.
Material so contaminated had been preserved separately by the direction
of Professor W. P. Popenoe. Attempts to identify the source of this
mixed material proved fruitless, as much of it had been cleaned of
matrix. It has accordingly been excluded from the present study.

There remain in the Clark collection 227 species and varieties of
mollusks that are unquestionably from the Upper Pleistocene deposits
in Potrero Canyon (U.C.L.A. Locality no. 3225). The fauna is
herein listed, together with all known published records of fossils from
the same locality. In listing previous records, the nomenclature has
been revised to accord with present usage. The chitons were identified
by Dr. S. Stillman Berry during the preparation of his monograph
(Berry, 1922). A total of 262 molluscan species and varieties are now
reported from Potrero Canyon Upper Pleistocene.

Species or varieties not previously reported as fossils are Acteocina
magdalenensis, Sulcoretusa xystrwn (plate 13, fig. 10), Hanetia elegans
(plate 13, figs. 7, 8), Odostomia jarallonensis, Odostomia gravida, Ala-
bind tenuisculpta var. phalacra, and Polinices draconis.

Acknowledgments

Professors U. S. Grant and W. P. Popenoe, University of Cali-
fornia, Los Angeles, and Dr. John T. McGill, Geologist, U. S.
Geological Survey, aided the ..preparation of this paper by careful
criticism and discussion. The Clark collection was made available to
the writer through the courtesy of Mr. Takeo Susuki, Junior Research
Geologist, University of California, Los Angeles. Dr. Leo G. Hertlein,
California Academy of Sciences, kindly furnished comparative material
and ecologic data on certain mollusks. The index map was drafted
by Mr. Ronald H. Arntson, University of California, Los Angeles,
and Mr. Armour C. Winslow, Louisiana State University, aided in the
preparation of the plate.

188 San Diego Society of Natural History

STRATIGRAPHY OF THE TERRACE DEPOSITS

Remnants of a marine platform of abrasion, included by Davis in
his Dume terrace (1931, p. 1098 ff.), are preserved beneath an alluvial
cover in the Pacific Palisades region. The platform surface is eroded
on tilted strata of considerable structural complexity, which range in
age from Paleocene to Lower Pleistocene. Near the head of Potrero
Canyon, about 0.1 mile south of U.C.L.A. Locality no. 3225 (figure
l), this surface is exposed in stream and road cuts. It is underlain
here by gently tilted, gray-green, massive siltstones mapped as Pliocene
by Hoots (1931, pi. 16), and overlain by 1 to 6 feet of essentially
horizontal, iron-stained, crudely bedded, loosely consolidated, fine to
medium grained, angular quartz sands. The sands, where exposed, are
largely unf ossilif erous ; only a few scattered sponge spicules and small,
badly worn shell fragments could be found. About 50 feet of alluvium
overlie the sands.

The surface of the terrace platform is somewhat irregular, as is
well shown in Davis' photographs (1933, pi. 55). In Potrero Canyon,
at the locality mentioned above, a shallow depression in the platform
surface is exposed on the east bank of the stream. Very fine, highly
f ossilif erous sand fills this depression; it is finer than the sands above
but coarser than the underlying siltstones. The molluscan fauna is
similar to that in the Clark collection; in addition, foraminifera, ostra-
codes, and the minute skeletal remains of various other groups of
marine invertebrates are present.

Hoots reports that the locality from which the Clark collection
was recovered (locality 61 of Hoots, 1931, p. 121 ) is in a fine white
and brown sand resting directly upon the Pliocene siltstones. The
matrix of the mollusks in the Clark collection agrees well texturally with
the very fine fossiliferous sand exposed in the depression, and contains
a similar microfauna. It seems likely therefore that the rich Upper
Pleistocene fauna has come from a unit lying between the Dume plat-
form surface and the barren, fine to medium grained sand exposed in
Potrero Canyon today. The fossiliferous unit was apparently stripped
off much of the terrace before the overlying sands were deposited.

Among the most abundant foraminifera from the fossiliferous
unit are Bolivina interjuncta Cushman, Uvigerina peregrina Cushman,
s. L, Cassidulina californica Cushman and Hughes, Cassidulina trans-
lucens Cushman and Hughes, and Anomalina c(. A. schmitti Cushman

Potrero Canyon Pleistocene. J. W. Valentine 189

and Wickendon. With the exception of the Anomalina, all these species
are reported from formations beveled by the terrace platform in the
Pacific Palisades region (Goudkoff et al. in Hoots, 1931, pp. 117-118),
and as they live today in depths much too great for waters on the Dume
terrace (see below; Natland, 1933) they have doubtless been reworked
from the underlying strata. The Anomalina may be conspecific with
a form that is now abundant in protected shallow water off Catalina
Island, but that lives in moderately deep water as well (Natland, 1933).
It may or may not represent an Upper Pleistocene microfauna.

PALEOECOLOGY

Habitat

A summary of the habitats of Recent representatives of Potrero
Canyon species is given in table 1. Most data on the ecology of
Pacific Coast Mollusca are recorded in rather general terms, and
therefore broad habitat categories have been used. Some of these
categories require definition.

Under "protected shallow water" are tallied species which live in
embayments or in quiet water along protected outer coasts, and which
are commonly found above 5 fathoms; most of them are subtidal, while
many range also into shallow offshore waters. "Ubiquitous" forms are
found in such a variety of environments that they cannot be used as
ecologic indicators in this study; few of them range into moderately
deep water, however. "Offshore, shallow water" forms live in waters
at least as shallow as 20 fathoms, but are found below most heavy

Table 1. Present habitats of species and varieties in the Potrero
Canyon fauna.

Species and Per Cent

Habitat Varieties

Protected shallow water 68 33.0

Ubiquitous 61 29.6

Offshore, shallow water 42 20.4

Exposed rocky shores 19 9.2

Offshore, moderately deep water 11 5.3

Exposed sandy shores 3 1.5

Pelagic; open ocean 2 1.0

Total for which data available 206 100.0

190 San Diego Society of Natural History

wave action; they are not recorded from very shallow, subtidal zones.
The "offshore, moderately deep" category includes species known to
be living only at depths greater than 20 fathoms.

From the table, it is clear that most of the species prefer quiet
shallow waters. _ The habitats from which "ubiquitous" species were
derived cannot be known, but if the same proportions hold for them
as hold among the rest of the fauna, fully 76.4 per cent of the species
tallied may represent shallow water habitats free from strong wave
action. The great majority of individuals as well as of species have
been drawn from these habitats. Of the 42 "superabundant" species
for which ecologic data are available, 23 live today in protected shallow
water, 2 offshore in shallow water, and 1 each in rocky exposed shore
and moderately deep water habitats. The remaining 15 are "ubiquitous".

Temperature

Using the present distribution of species and of water temperatures
as a datum, it seems possible to estimate the approximate magnitude
and range of water temperatures represented by the Potrero Canyon
fauna. The present geographic ranges of about 75 per cent of the
species includes the latitude of Potrero Canyon; all of the remaining
25 per cent (59 species) live today only south of the Potrero Canyon
area. Northern end-points of the ranges of these species are found in
the following areas: Santa Monica, 10 species; Redondo Beach, 11
species; San Pedro-Long Beach, 13 species; Newport Beach, 1 species;
Catalina Island, 4 species; La Jolla-San Diego, 5 species; and Baja
California, 15 species.

Thus, two-thirds of the south-ranging species have the northern
end-points of their present ranges within 25 or 30 miles of Potrero
Canyon, while one-third do not now live within 100 to 900 or more
miles of the region. Among the most southerly species are Trachy-
cardium procerum (Scammon's Lagoon to Peru), Chione gnidia
(Cedros Island to Panama), "Chione" picta (Magdalena Bay to
Panama), Dosinia ponderosa (Scammon's Lagoon to Peru), Mulinia
pallida var. modesta (Baja California and the Gulf of California),

1Species listed as "superabundant" in the Checklist of Species, to follow, are repre-
sented by 256 or more specimens in the Clark collection.

Potrero Canyon Pleistocene. J. W. Valentine 191

Bivetopsid bullata (plate 13, figs. 1, 2; Cedros Island to Panama),
Hanetia elegans, previously unreported fossil" (plate 13, figs. 7, 8;
Cape San Lucas to Panama), "Ndssd" cerritensis (Point Abreojos to
Guaymas), Centrifugd leednd (Guadalupe Island to Cedros Island),
Tricolid vdriegdtd (Magdalena Bay to Cape San Lucas), and Ischno-
chiton dcrior (Cedros Island to Cape San Lucas) .

Although no species are reported fossil at Potrero Canyon that
live only to the north, several are essentially northward-ranging types.
Among these are Bornid retiferd (Monterey to the Santa Barbara Is-
lands), Mitromorphd interfossd (Alaska to Catalina Island), Ocinebrd
barbdrensis (British Columbia to San Pedro), and Epitonium sdwinde
(British Columbia to Catalina Island).

Mean annual near-shore sea-surface temperatures range from some-
thing below 61° F. in Santa Monica Bay to about 68° or 69° F. near
Magdalena Bay and about 75° F. near Cape San Lucas (Hertlein and
Grant, 1944, p. 72). Although northern and southern species in the
Potrero Canyon assemblage may live today in waters warmer or cooler
than mean annual temperatures can indicate, their present distribution
strongly suggests that the range of water temperature off the Dume
terrace coast was greater than at present. Waters no warmer than
those in Santa Monica Bay today, together with waters perhaps, as a
conservative guess, 8° F. warmer, seem to have been present con-
temporaneously, or perhaps in alternate seasons, during the sea-stand
on the Dume terrace platform.

A mechanism which might account for this condition has recently
been postulated (Valentine, 1955) : shallow quiet waters were warmed
at some time in the Upper Pleistocene either by increasing heat from
the sun or by the introduction of warm southern water in counter-
currents; at the same time, shallow waters in areas where upwelling
occurred remained relatively cool. As it seems likely that the tempera-
ture of upwelling water was lower than at present during and shortly
after glacial ages, relatively warm and cold water species may have
been able to live in close proximity. Emerson (1956) has also discussed
the effect of upwelling on molluscan faunas, and its significance in
Pleistocene paleoecology.

2Gratitude is expressed to George P. Kanakoff, Los Angeles County Museum, for
identification of this species. Mr. Kanakoff has discovered Hanetia elegans in Upper
Pleistocene deposits at Playa Del Rey, California.

192 San Diego Society of Natural History

INFERRED DEPOSITIONAL ENVIRONMENT
The position of the shoreline at high sea stand on the Dume ter-
race, indicated in figure 1, is an approximation based on the slope of
the platform surface as extrapolated from localities where it is exposed,
and on such evidence as is afforded by the present topography. No
exposure of the shoreline angle was found. Although the precise posi-
tion and character of the Dume shore cannot be determined, this ap-
proximation will serve as a basis for limiting the depth at which the
fossils could have been deposited; the difference in elevation between
the fossil locality and the shoreline angle is judged to be no more than
30 feet.

Conditions at and near the site of deposition may be limited by
combining the geographic and faunal evidence. The fossil locality lay
no farther than l/> mile from shore and under a probable maximum of
5 fathoms of water. Silt and fine sand composed the substratum. These
conditions seem to be reflected in the ecologic composition of the
fossil fauna, for Recent representatives of most of the species prefer or
tolerate such an environment. Trie waters near the fossil locality were
probably fairly quiet, and not consistently disturbed by wave action while
the assemblage lived. Perhaps an offshore bar or spit protected this
part of the platform at times from most wave action; Davis (1933,
p. 1103) mentions this possibility in another connection. Moderately
deep water or open ocean types must have been swept inshore by oc-
casional storm waves, while exposed sandy beach and rocky shore forms
may have been transported offshore through the action of currents and
gravity.

AGE AND FAUNAL AFFINITIES

As noted above, the Dume terrace bevels tilted strata of Lower
Pleistocene age, and the marine terrace deposits at Potrero Canyon
contain a fauna characterized by many species that live today along
the nearby coast, together with several markedly southern forms. Similar
faunas are commonly found in strata assigned to the Upper Pleistocene
of Southern California. The Potrero Canyon fauna may therefore be
considered as Upper Pleistocene on both stratigraphic and faunal
grounds.

Similarities between the Upper Pleistocene fauna described by
Willett from Playa Del Rey (1937) and the Potrero Canyon as-
semblage are striking, as Woodring has noted (in Hoots, 1931, p. 122,
and in Woodring, Bramlette, and Kew, 1946, p. 106). The two

Potrero Canyon Pleistocene. J. W. Valentine 193

faunas have 194 species and varieties in common, while 96 forms from
Playa Del Rey are not reported at Potrero Canyon, and 68 of the
Potrero Canyon forms are not known from Playa Del Rey. The Playa
Del Rey assemblage is slightly the larger (2903 vs. 262 species and
varieties). Differences between the two faunas are much smaller than
these figures indicate. Of the forms not found in both faunas, only 20
from Playa Del Rey and 17 from Potrero Canyon are represented by
more than 10 specimens in the very large collections available from
each locality. Relative abundances of species in each assemblage are
remarkably alike. Several species common to the two localities are
not reported elsewhere in the Pleistocene of Southern California. These
include Diacria trispinosa (plate 13, figs. 3, 4), Bivetopsia bullata
(plate 13, figs. 1, 2), Engina strongi (plate 13, fig. 9), Hanetia elegans
(plate 13, figs. 7, 8), and Calliostoma gloriosum (plate 13, figs. 5, 6).
Several species from Playa Del Rey that are unknown at Potrero Can-
yon are tidal marsh or mud flat forms, a biotope not represented at the
latter locality. With this exception, both faunas have clearly been drawn
from similar habitats within nearly the same range of ecologic condi-
tions. Both faunas have essentially the same thermal significance as
well.

The bearing of the Potrero Canyon assemblage on the age of
the Rancho La Brea vertebrate fauna has been discussed by Grant and
Sheppard (1939, p. 308), who point out that the Rancho La Brea de-
posits seem to be contained in the distal end of the Hollywood fan,
one of a series of alluvial fans that blanket the northern margin of
the Los Angeles basin. These fans appear to be roughly contem-
poraneous with nonmarine terrace cover at Pacific Palisades. The
vertebrate fauna is thus probably Upper Pleistocene, and somewhat
younger than the Potrero Canyon assemblage.

Precise correlation of the Dume terrace with individual terraces
elsewhere in Southern California cannot yet be made, but the correlations
suggested by Woodring (loc. cit.) are doubtless correct. That is, the
Potrero Canyon assemblage belongs within a period of uncertain dura-
tion in the Upper Pleistocene that is characterized faunally by the
presence of a prominent southern molluscan element in protected
habitat biofacies; to this period of time also belong the lowest terrace
in the Palos Verdes Hills and possibly several of the earlier terraces,
the Playa Del Rey deposits, and their correlatives.

3The figures given for the Playa Del Rey assemblage differ slightly from Willett's,
owing to the necessity for "splitting" or "lumping" a few forms in order to obtain
uniform taxonomic treatment for both faunas.

194

San Diego Society of Natural History

CHECKLIST OF FOSSILS

Key
Column 1, Clark collection:

V — Very rare, less than 5 specimens.

R — Rare, from 5 to 16 specimens.

C — Common, from 17 to 64 specimens.

A — Abundant, from 65 to 256 specimens.

S — Superabundant, more than 256 specimens.
Column 2, Woodring in Hoots, 1941 :

X — Reported present.
Column 3, Grant and Gale, 1931:

Clk — Attributed to the Clark collection.

Old — Attributed to the Oldroyd collection.
Column 4, other authorities:

B-l— Reported by Bartsch, 1911.

B-2— Reported by Bartsch, 1917.

Ber — Reported by Berry, 1922.

Cha — Reported by Chace, 1917.

Dal— Reported by Dall, 1922.

Old— Reported by Oldroyd, 1921.

Ray — Reported by Raymond, 1906.

Riv — Reported by Rivers, 1904.

Wdg — Reported by Woodring, 1946.
Column 5, present geographic distribution:

P — Present range includes the latitude of Potrero Canyon.

S — Known to be living only south of Potrero Canyon.

E — Not known to be living.

SYSTEMATIC CHECKLIST

1

SPECIES and VARIETIES

°3

PELECYPODA

Nucula suprastriata Arnold S

Nuculana taphria (Dall) S

Yolida cooperi Gabb S

Ostrea lunda Carpenter S

Leptopecten latiauratus (Conrad) S

Leptopecten latiauratus var. monotimeris

(Conrad) S

2

3

4

rr\

C ~

!r 'S

Wood.
Hoots

Gram
Gale,

4> 0

■fa •£

c

X

X ...

Old

X ...

c

<-. o

c -c

U 3

P
P
P
P
P

Potrero Canyon Pleistocene. J. W. Valentine

195

SYSTEMATIC CHECKLIST

1

SPECIES and VARIETIES

u-s

Lima hemphilli Hertlein & Strong V

Anomia peruviana d'Orbigny C

Pododesmus macroschisma (Deshayes) R

Modiolus jornicatus Carpenter V

Lithophaga plumula (Hanley) C

Periploma planiusculum Sowerby C

Pandora punctata Conrad C

Crassinella branneri (Arnold) R

Crassinella nuculiformis Berry V

Chama pellucida Broderip C

Pseudcchama exogyra (Conrad) S

Epilucina calif ornica (Conrad) V

Here excavata (Carpenter) C

Lucinisca nuttallii (Conrad) S

Parvilucina tenuisculpta (Carpenter) C

Diplodonta orbella (Gould)

Kellia laperousii (Deshayes) V

Aligena cerritensis Arnold V

Mysella aleutica (Dall) V

Bornia retifera Dall V

Laevicardium datum (Sowerby) R

Trachycardium procerum (Sowerby) C

Trachycardium quadrigenarium (Conrad) A

Trigoniocardia biangulata (Sowerby) A

Chione gnidia (Broderip & Sowerby)

Chione undatella (Sowerby)

Chione ci. C. undatella (Sowerby) V

Chione undatella var. simillima (Sowerby) ....A

"Chione" picta Dall R

Protothaca tenerrima (Carpenter)^

Saxidomus nuttalli Conrad. A

Transenella tantilla (Gould)

Amiantis callosa (Conrad) , A

Dosinia ponderosa (Gray) V

Cooperella subdiaphana (Carpenter) aV

Tellina buttoni Dall ..V

Tellina idae Dall S

Apolymetis biangulata (Carpenter) C

Macoma indentata Carpenter S

Woodring
in Hoots, 1931 K>

3

?!

Other
Authority ■*■

X ...

X ....

X ...

X ....

X ....

Old'

X ...

X

X ...

Old

Wdg

X ...

X ...

X ...

X ...

Wdg

X ....

X ...

X

Clk

c

fi s

ai 3
"» _o

<u rs

^ is

S
P

s
p
p
p
p
p
s

E
P
P
P
S
P
P
P
P

s
p
p
s
s
p
s
s
p

9

s
s
p
p
p
s
s
p
p
s
p
p

196

San Diego Society of Natural History

SYSTEMATIC CHECKLIST

I

SPECIES and VARIETIES

U;

Macoma nasuta (Conrad) S

Macotna secta (Conrad) S

Macoma yoldiformis Carpenter V

Semele decisa (Conrad) R

Semele rubropicta Dall V

Donax gouldii Dall V

Psammobia edentula (Gabb) - R

Tagelus subtcres (Conrad) V

Solen sicariui Gould. A

Siliqua lucida (Conrad) R

Ensis californicus Dall A

Mactra californica Conrad. A

Spisula catilliformis Conrad. V

Spisula " dolabriformis Conrad" R

Spisula hemphtlli (Dall) A

Mulinia pallida var. modestd Dall V

Schizothaerus nuttallii (Conrad) A

Platyodon cancellata (Conrad) R

Cryptomya californica (Conrad) S

"Ccrbtda" luteola Carpenter R

Panope generosa Gould C

Hiatella arctica (Linnaeus) V

Zirjaea pilsbryi Lowe C

SCAPHOPODA

Dentalium neohexagonum Sharp 6C Pilsbry S

Cadulus fusiformis Pilsbry & Sharp A

GASTROPODA

Carolina telemus var. tricuspida (Rivers)

Diacria trispinosa (Lesueur) V

Acteon punctocaelatus (Carpenter) C

Acteon trash Stearns R

Acteocina culcitella (Gould) S

Acteocina incidta (Gould) S

Acteocina infrequent (C. B. Adams)

Acteocina magdalenensis Dall V

Sulcoretusa xystrum (Dall) V

Coleophysis carinata (Carpenter)

.If2

X

X
X
X

X

X

X

X
X

X
X

X
X

X
X
X

o

c ~"
n .-

Old

uC

I J

Old

Clk

Old
Old
Clk
Old

Old

Riv

Dal

c

w O
C "C

V 3

■3

P
P
P

s
p
p
s
p
p
p
p
p
p
p
s
s
p
p
p
p
p
p
p

p
p

p
p
p
s
p

p
s
s
s
p

Potrero Canyon Pleistocene. J. W. Valentine

197

SYSTEMATIC CHECKLIST

SPECIES and VARIETIES

c
o

re il

U-3
U

Coleophysis harpa (Dall) S

Volvulella cylindricd (Carpenter) S

Cylichna attonsa Carpenter S

Bulla cf. B. punctulata A. Adams V

Williamia peltoides (Carpenter) R

Terebra pedroana Dall S

Conus californicus Hinds S

Megasurcula car pent eriarut (Gabb) R

Megdsurcula carpenteriana var.

tryoniand (Gabb) A

Antiplanes perversa (Gabb) V

Pseudomeidtoma peniallata var.

semtinfldtd Grant 8C Gale A

Moniliopsis incisd (Carpenter) ?

Moniliopsis incisa var. jancherde (Dall) S

Moniliopsis incisa var. ophioderma (Dall) S

Clavus hemphilli (Steams) S

Kurtzia arteagd var. roperi (Dall) C

"Mdngelid" cetoldcd Dall S

"Mangelia" variegdtd Carpenter S

Mitromorphd cf. M. interjossd Carpenter

Bivetopsia bullata (Sowerby) R

Olivella baetica Carpenter S

Olivella btplicatd (Sowerby) S

Olivelld pedrodnd (Conrad)

Hyalina cdlifornicd (Tomlin) R

Cystiscus reguldris (Carpenter)

Cystiscus subtrigond (Carpenter) V

Mitrd fultoni Smith ^ R

Mitrd idde Melvill R

"Fusinus" luteopictus Dall C

Barbdrofusus bdrbarensis (Trask) R

Hdrfordia rn.onk.Sde (Dall)

Kelletia kelletii (Forbes) V

Engind strongi Pilsbry 8C Lowe V

Hanetia elegans Dall V

"Ndssa" cerritensis Arnold C

"Nassd" delosi Woodring S

"Nassa" jossata (Gould) S

2

if

fix

3

c

Other
Authority ^

X
X

Old
Clk

Old
Clk

X
X
X .

Ray
Ray

X

?

X

Clk

X

X

Old

X

X .

Clk

X

Old

X

Old

Wdg

X
X .

c
w o
c '5

4) 3

P
P
P

P
P
P
P

P
P

E
P
P
P
P
P
E
P

S

P
P
P

s
p
p
s
p
p
p
p
p
s
s
s

E
P

198

San Diego Society of Natural History

SYSTEMATIC CHECKLIST

SPECIES and VARIETIES

c
o

U

"Nassa" mendica Gould

"Nassa" mendica var. cooperi Forbes C

"Nassa" perpinguis (Hinds) S

"Nassa" tegula Reeve V

Anachis lineolata (Reeve)

Mitrella carinata (Hinds) A

Mitrella carinata var. gausapata (Gould) S

Mitrella gouldi (Carpenter) V

Mitrella tuberosa (Carpenter) C

Amphissa versicolor Dall C

Pteropurpura trialatus (Sowerby) R

Pterynotus petri (Dall) C

Centrifu°a leeana (Dall) C

Jaton festivus (Hinds) S

Maxwellia gemma (Sowerby) A

Ocinebra barbarensis (Gabb)

Ocinebra circumtexta (Stearns) V

Ocinebra foveolata (Hinds) V

Ocinebra interfossa Carpenter V

Ocinebra poulsoni Carpenter S

Stramonita biserialis (Blainville) A

Nucella emarginata var. ostrina (Gould) R

Acanthina lugubris (Sowerby)

Acanthina paucilirata (Steams)

Acanthina spirata (Blainville) A

Boreotrophon stuarti (Smith)

Forreria belcheri (Hinds) C

Epitonium bellastriatum (Carpenter) A

Epitonium clarki T. S. Oldroyd S

Epitonium cooperi Strong A

Epitonium insculptum (Carpenter) V

Epitonium sawinae Dall A

Epitonium tinctum (Carpenter)

Balcis micans (Carpenter) C

Balcis monicensis (Bartsch) R

Balcis oldroydi (Bartsch) V

Balcis rutila (Carpenter) C

• Balcis thersites (Carpenter) R

2

If

X

3

o
c

o

Old
Clk

Clk

4

oj O

Old

X

Cik
Gk
Old

X

X

Old

X .

X .

Wdg

Wdg

X .

X .

X

Old

Old

X .

Clk

B-2
B-2
B-2
B-2

c

i_, o
C 'C
en 3

8-o
a, g

S

P
P
P
P
S
P
P

P
P
P
P
P
S

p
p
p
p
p
p
p
s
p
s
s
p
p
p
p

E
P
E
P
P
P
E
P
P
P

Potrero Canyon Pleistocene. J. W. Valentine

199

SYSTEMATIC CHECKLIST

SPECIES and VARIETIES

c
o

re £

u3

Strombiformis raymondi (Rivers) R

Turbonilla almo Dall dc Bartsch A

Turbonilla asser Dall & Bartsch. R

Turbonilla canfieldi Dall & Bartsch C

Turbonilla carpenteri Dall & Bartsch. V

Turbonilla keepi Dall 6C Bartsch A

Turbonilla laminata (Carpenter) R

Turbonilla lowei Dall & Bartsch A

Turbonilla lowei var. pedroana

Dall & Bartsch C

Turbonilla pentalopha Dall & Bartsch V

Turbonilla regina Dall & Bartsch S

Turbonilla stylina (Carpenter) R

Turbonilla tenuicula (Gould) C

Turbonilla torquata (Gould) R

Turbonilla tridentata (Carpenter) R

Turbonilla weldi Dall & Bartsch. A

Odostomia cf. O. californica Dall & Bartsch....V

Odostomia donilla Dall & Bartsch C

Odostomia eugena Dall & Bartsch. R

Odostomia farallonensis Dall & Bartsch- C

Odostomia gravida Gould A

Odostomia belena Bartsch C

Odostomia cf. O. minutissima Dall & Bartsch..A

Odostomia nemo Dall & Bartsch S

Odostomia cf. O. phanella Dall & Bartsch V

Burj-d californica (Hinds) A

Pusula calif ornianus (Gray) V

Pusula solandri (Sowerby) ^

Erato columbella Menke V

Alabina monicensis Bartsch

Alabina tenuisculpta var.

diegensis Bartsch V

Alabina tenuisculpta var. phalacra Bartsch R

Bittium attenuatum Carpenter

Bittium rugatum Carpenter V

Bittium cf. B. rugatum var.

giganteum Bartsch

Bittium rugatum var. larum Bartsch R

2

rr\

C *"'

IJ
pi

.s

3

c

E *

4

4) O

-S-£

X? w

O 3
<

Riv

X

Old

Old
Old

X

B-1

X

X

c

u O
C "C

§J

fv C

s
p
p
p
s
s
s
p

p
s
p
p
p
p
p
s

9

s
s
p
p
p

s

p
p
s

p

E

s
s
p
s

E
.S

200

San Diego Society of Natural History

SYSTEMATIC CHECKLIST

1

SPECIES and VARIETIES

c
o

D

Seila montereyensis Bartsch R

Cerithiopsis antefilosa Bartsch R

Cerithiopsis arnoldi var. fossilis Bartsch

Cerithiopsis cosmia Bartsch R

Cerithiopsis pedroana (Bartsch) R

Cerithiopsis pedroana var. fatua Bartsch

Triphora pedroana (Bartsch) R

Metaxia diadema Bartsch

Alvania acutelirata (Carpenter) V

Rissoina pleistocena Bartsch C

Turritella cooperi Carpenter A

Aletes squamigerus Carpenter V

Spiroglyphus lituella (Morch) R

Caecum californicum Dall V

Micranellum crebricinctum (Carpenter) S

Fartulum hemphilli Bartsch R

Fartulum occidentale Bartsch C

Littorina planaxis Philippi V

Littorina scutulata Gould C

Lacuna carinata Gould V

Lacuna carinata var. aurantiaca Carpenter R

Lacuna unifasciata Carpenter S

Alaba catalinensis Bartsch V

Alaba jeannettae Bartsch V

Hipponix antiquatus (Linnaeus)

Hipponix lumens Carpenter V

Crepidula adunca Sowerby

Crepidula excavata (Broderip) S

Crepidula nummaria Gould C

Crepidula onyx Sowerby A

Crepipatella lingulata (Gould) C

Crucibulum spinosum (Sowerby) V

Calyptraea contorta Carpenter C

Polinices draconis Dall R

Neverita reclusiana (Deshayes) A

Neverita reclusiana var. alia Arnold S

Neverita reclusiana var. imperforata Dall S

Euspira lewisii (Gould) V

Sinum debile (Gould)

Sinum cf. S. debile (Gould) V

2

J j

X

3

c

ak

Other
Authority 4*.

X ..

X .

Clk

X .

X ..

X ..

Clk

X ..

X ..

Clk

Clk

X

Clk

ak
ak

Clk
Old

X

X ..

Old

X

Clk

X

X ..

Clk

c

« .2

c n
<u 3

A4 h

S

s

E
P
S
E
S
P
S
E
P
P
P
P
P
S
P
P
P
P
P
P
S
S
P
P
P
P
P
P
P
P
P
P
P
P
P
P
S

Potrero Canyon Pleistocene. J. W. Valentine

201

SYSTEMATIC CHECKLIST

SPECIES and VARIETIES

1

c

D3

Sinum scopulosum (Conrad) S

Acmaea insessa (Hinds)

Tricolia compta (Gould) C

Tricolia pulloides (Carpenter) S

Tricolia variegata (Carpenter)

Pomaulax undosus (Wood) A

Homalopoma carpenteri (Pilsbry) C

Norrisia norrisi (Sowerby) A

Tegula aureotincta (Forbes) S

Tegula gallina (Forbes) A

Tegula ligulata (Menke) A

Calliostoma annulatum (Humphreys)

Calliostoma doliarium (Holten) C

Calliostoma gemmulatum Carpenter C

Calliostoma gloriosum Dall V

Calliostoma cf. C. splendens Carpenter

Calliostoma supragranosum Carpenter V

Calliostoma tricolor Gabb S

Turcica caffea Gabb V

Pupillaria optabilis (Carpenter) A

Vitrinella? sp V

Skenea calif ornica (Bartsch) V

Diodora aspera Eschscholtz

Megatebennus bimaculatus (Dall) V

AMPHINEURA

Leptochiton clarki Berry

Mapalia acuta (Carpenter) R

Acanthochitona avicula (Carpenter)

Ischnochiton acrior Carpenter R

Ischnocbiton conspicuus Carpenter...." V

Ischnochiton pectinulatus Carpenter R

Ischnochiton sanctaemonicae Berry A

Callistochiton crassicostatus Pilsbry C

Callistochiton palmulatus var.

■mirabilis Pilsbry A

2

CO

in <*'

3

c

ID *

4
b

X ...

X ...

X ...

X ...

X

X

X

Old

X ...

X

Old

Ber

Ber

Ber

Ber

Cha

Ber

Cha

Ber

Ber

Cha

Ber

Cha

Ber
Cha

X

§1

P
P
P
P

s
P
P
p
p
p
p
p
p
p
p

p
p
p
p
s
p
p
p

E
P
P
S

P

E

202 San Diego Society of Natural History

REFERENCES
Arnold, Ralph

1903. The paleontology and stratigraphy of the Pliocene and
Pleistocene of San Pedro, California. Calif. Acad. Sci.,
Mem., vol. 3, 420 pp., 37 pis.

Bartsch, Paul

1911. The Recent and fossil mollusks of the genus Alabina from
the west coast of America. U.S. Nat. Mus., Proc, vol. 39,
pp. 409-418, pis. 61-62.

1917. A monograph of west American melanellid mollusks. U.S.
Nat. Mus., Proc, vol. 53, pp. 295-356, pis. 34-49.

Berry, S. S.

1922. Fossil chitons of western North America. Calif. Acad. Sci.,
Proc, ser. 4, vol. 11, pp. 399-526, pis. 1-16.

Chace, E.

1917. Fossil chitons. Lorquinia, vol. 2, no. 4, pp. 30-31.
Dall, W. H.

1922. Note on Acteoc'ina. Nautilus, vol 35, p. 76.
Davis, W. N.

1933. Glacial epochs of the Santa Monica Mountains, California.
Geol. Soc Amer., Bull., vol. 44, pp. 1041-1133, pis. 40-65.
Emerson, W. K.

1952. The influence of upwelling on the distribution of marine
floras and faunas of the West Coast of Baja California,
Mexico. (Abstract). Amer. Malacol. Union, Ann. Rept.
for 1952, pp. 32-33.

1956. Upwelling and associated marine life along Pacific Baja
California, Mexico. Jour. Paleo., vol. 30, in press.
Grant, U. S., IV, and Gale, H. R.

1931. Catalogue of the marine Pliocene and Pleistocene Mol-
lusca of California and adjacent regions, etc. San Diego
Soc Nat. Hist, Mem., vol. 1, 1036 pp., 32 pis.

, AND SHEPPARD, W. E.

1939. Some recent changes of elevation in the Los Angeles basin
of southern California, and their possible significance.
Seismol. Soc. Amer., Bull., vol. 29, pp. 299-326.

Potrero Canyon Pleistocene. J. W. Valentine 203

Hertlein, L. G., and Grant, U. S., IV

1944. The geology and paleontology of the marine Pliocene of
San Diego, California, Part 1, Geology. San Diego Soc.
Nat. Hist., Mem., vol. 2, 72 pp., 18 pis.

Hoots, H. W.

1931. Geology of the eastern part of the Santa Monica Moun-
tains, Los Angeles County, California. U. S. Geol. Surv.,
Prof. Paper 165-C, pp. 83-134, pis. 16-34.

Natland, M. L.

1933. The temperature and depth-distribution of some Recent
and fossil foraminifera in the southern California region.
Univ. California, Scripps Inst. Oceanography, Tech. Ser.,
Bull., vol. 3, no. 10, pp. 225-230.

Oldroyd, T. S.

1921. New Pleistocene mollusks from California. Nautilus, vol.
34, pp. 114-116, pi. 5, figs. 8-13.

Raymond, W. J.

1906. The west American species of Pleurotoma, subgenus Genota.
Nautilus, vol. 20, pp. 37-39, pi. 2.

Rivers, J. J.

1904. Descriptions of some undescribed fossil shells of Pleistocene
and Pliocene formations of the Santa Monica range. So.
Calif. Acad. Sci., Bull., vol. 3, pp. 69-72.

Valentine, J. W.

1955. Upwelling and thermally anomalous Pacific Coast Pleisto-
cene molluscan faunas. Amer. Jour. Sci., vol. 253, pp.
462-474.

Willett, George

1937. An Upper Pleistocene fauna from the Baldwin Hills, Los
Angeles County, California. San Diego Soc. Nat. Hist.,
Trans., vol. 8, pp. 379-406, pis. 25-26.

Woodring, W. P., Bramlette, M. N., and Kew, W. S. W.

1946. Geology and paleontology of Palos Verdes Hills, Cali-
fornia. U. S. Geol. Surv., Prof. Paper 207, 145 pp., 37 pis.

204 San Diego Society of Natural History

EXPLANATION OF PLATE 13

Figs. 1, 2. Bivetopsia bullata (Sowerby), approximately natural size.
Hypotype, U.C.L.A. coll., no. 27480. Length, 45 mm.,
diameter of last whorl, 34 mm. U.C.L.A. Loc. no. 3225.

Figs. 3, 4. Diacria trispinosa (Lesueur), approximately X 51^. Hypo-
type, U.C.L.A. coll., no. 27381. Length, incomplete, 7.6
mm., width, incomplete, 6.5 mm. U.C.L.A. Loc. no. 3225.

Figs. 5, 6. Calliostoma gloriosum Dall, approximately X 2. Hypo-
type, U.C.L.A. coll., no. 27482. Length, 17 mm., di-
ameter of last whorl, 13 mm. U.C.L.A. Loc. no. 3225.

Figs. 7, 8. Hanetia elegans Dall, approximately natural size. Hypo-
type, U.C.L.A. coll., no. 27483. Length, 41 mm., diameter
of last whorl, 26 mm. U.C.L.A. Loc. no. 3225.

Fig. 9. Engina strongi Pilsbry & Lowe, approximately X 2. Hypo-

type, U.C.L.A. coll., no. 27484. Length, 14 mm., diameter
of last whorl, 8 mm. U.C.L.A. Loc. no. 3225.

Fig. 10. Sulcoretusa xystrum (Dall), approximately X 15l/>.
Hypotype, U.C.L.A. coll., no. 27485. Length, 1.7 mm.,
greatest diameter, 0.8 mm. U.C.L.A. Loc. no. 3225.

Potrero Canyon Pleistocene. J. W. Valentine

205

2

/

Ik

y

^15

Plate 13

TRANSACTIONS

OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY

Volume XII. No. 11, pp. 207-230. Plates 14-17

NOTES ON SOME INTERTIDAL COLEOPTERA

WITH DESCRIPTIONS OF THE EARLY STAGES

(CARABIDAE, STAPHYLINIDAE,

MALACHIIDAE)

BY

Ian Moore

Research Associate in Co! copter a
San Diego Museum of Natural History

MUS. COMP. ZOOL
LIBRARY

SEP 1 3 1956

UNIVERSITY

SAN DIEGO, CALIFORNIA

Printed for the Society

August 24, 1956

COMMITTEE ON PUBLICATION

Laurence M. Klauber, Chairman
Joshua L. Baily Charles C. Haines

Carl L. Hubbs
John A. Comstock, Editor

Table of Contents

Page
Introduction 211

Table of seashore zones 212

Genus Thalassotrechns Van Dyke 213

T. barbarae barbarae (Horn) 214

T. barbarae nigripennis Van Dyke 214

Larva of T. barbarae (Horn) 215

Key to the known larvae of the Phytosi 216

Phytosus 216

Antarctophytosns 216

Liparocephalus 216

Diaidota 217

Key to the known larvae of Diaulota 217

D. dentissima Casey 217

D. fulviventris Moore 218

D. vandykei Moore 219

D. megacephala Moore or D. hartcri Moore 219

Genus Endeodes LeConte 220

E. collaris (LeConte) 220

E. rugiceps Blackwelder 221

E. insularis Blackwelder 222

Bibliography 223

Explanation of figures, Plates 14-17 224

Plate 14 226

Plate 15 227

Plate 16 228

Plate 17 229

vJI_l

NOTES ON SOME INTERTIDAL COLEOPTERA

WITH DESCRIPTIONS OF THE EARLY STAGES

(CARABIDAE, STAPHYLINIDAE,

MALACHIIDAE)

By Ian Moore

Research Associate in Coleoptera
San Diego Museum of Natural History

INTRODUCTION

The rich and varied insect fauna of the Southern California seashore
has been little investigated except for the describing and naming of species.
The several distinctly different ecological situations in this realm, each of
which supports its own assemblage of insect species, can be readily located
in relation to each other according to the type of shore and to the range
of the tide. The beetle fauna falls naturally into three main zones, with
almost no overlapping of species between them. The three zones are deter-
mined by the type of shore: whether it is (1) a mud flat of a bay or
estuary, (2) a sandy beach, or (3) a rocky headland with a submerged
reef. The beetle fauna of the mud flats is the least apparent of the three
and is little known on the California coast. The sandy beaches support a
very extensive insect fauna, of which Coleoptera comprise a major part.
Of the three quite obvious horizontal subdivisions of this zone, each has
its own group of species, a few of which are common to more than one
subdivision. In the lowest subdivision, that of the fresh seaweed and wet
sand, which is characterized by the presence of numerous amphipods, will
be found Thinopinus pictus LeConte and Dyschirius marinus (LeConte).
The next subzone is a narrow belt of patches of decaying seaweed, which
is probably reached by only one or two high tides a month. Here are found
a large number of species of beetles that feed largely on the larvae of the
kelp flies. Most, but not all, of the genera found here are restricted to the
sea beach. They include Cafiiis, Bryonomits, Bledins, Aleochara, Pbaleria,
and Cercyon. Above this subzone is an area of patches of dry seaweed that
are probably left there by a major storm or an exceptionally high tide. The
genera, which again are largely restricted to this subzone, and some of
which are representatives of families more commonly associated with dried
cereals, include Epantius, Phyconomus, Catorama, and Endeodes. The up-
per limit of this zone marks the edge of the intertidal area and the be-
ginning of the sand-dune area, the fauna of which more closely resembles
that of the desert fifty or more miles to the east. The last major zone,
that of the rocky headlands and reefs, supports a fauna of beetles capable
of living submerged in seawater for long periods. Two species of Diaulofa

212

San Diego Society of Natural History

TABLE 1

ZONES OF THE SOUTHERN CALIFORNIA SEASHORE

IN WHICH COLEOPTERA ARE FOUND, WITH SOME

CHARACTERISTIC GENERA

1. MUD FLATS 2. SANDY BEACHES 3. ROCKY HEADLANDS

AND REEFS

B. Area of sea-
lettuce
Cicindela
Tachys
Bembidion
Carpelimus

A. Area of dry
seaweed

Endeodes

Catorama

Epantius

Amblydertts

Ant hi cm

Phyconomus

Phaleria

B. Area of decay-
ing seaweed

Cercyon

Saprinus

Bledius

Bryonomus

Cafius

Hadrotes

Pontamalota

Thinusa

Bryobiota

Tarphiota

Aleochara

Emplenota

Motschulskium

Phaleria

Phycocoetes

Emphyastes

C. Area of fresh
seaweed

Cicindela
Dyschirius
Thinopinus
{Bryonomus
occasionally)

A. Area of spray
(Species from
Zone 2B often
take refuge
here in cracks
in the rocks.)

B. Area of high
tides

Thalassotrechus
Diglotta
Bryothinusa
Diatdota

(Also Ochthebias,
Amblopusa, Lip-
arocephalus and
Endeodes in Cen-
tral California)

C. Area of red
seaweed down
to mean low
water

Diaulota

Moore — Intertidal Coleoptera 213

are found as far down as mean low water. Other species are found within
a foot or two of average high water and some species are found more com-
monly within the spray zone just above high water. All are found most
often in fine, well-drained cracks in the rocks. Table 1 indicates the rela-
tion of the various zones to each other as well as some of the commoner
forms of Coleoptera to be found in each subzone.

This paper deals with the beetles of the rocky shores of Southern
California, with particular emphasis on the descriptions of the larvae of
these forms and their near relatives of central California.

CARABIDAE
THALASSOTRECHUS Van Dyke

This genus, described in 1918 by Van Dyke, was based on Trechus
barbarae Horn1 from Santa Barbara, California. At the time that Van Dyke
described the genus, he also described another species, Thalassotrechus
uigripennis Van Dyke from Moss Beach, San Mateo County, California.
In 1951 I sent some examples of the genus to Van Dyke for identification
and he wrote saying that all the southern specimens were barbarae and all
the northern nigripcnnis. He stated that there was very little difference
between the two and that he had actually considered tiigripemiis, when he
described it, to be a subspecies of barbarae. My own series, now over one
hundred specimens, substantiates this view.2 There is no consistent ob-
servable structural difference between the two forms, but if the specimens
are divided into two groups, one from the north of Point Conception and
one to the southward, certain characters stand out prominently. Nigripen-
n'ts is distinctly larger, darker, and more shining. In a long series of larvae,
I can find no structural difference, but most of the specimens of nigri-
pennh are again definitely darker. Although Van Dyke considered certain
specimens that I had collected at Gaviota, Santa Barbara County, Cali-
fornia, as belonging to the northern subspecies, I now find that they are
much closer to the southern series. The dividing point between the two
subspecies is somewhere between Point Conception and the Big Sur region
of Monterey County. The region just north of Point Conception seems

1 Anatrcchus Casey, 1918, Memoirs on the Coleoptera, VIII, p. 411, was based on the
same species. According to Casey, 1920, p. 185, it appeared about a month after Van
Dyke's paper and so is a synonym of Thalassotrechus.

-Since the above was written, I have received from Dr. P. J. Darlington a copy of his
The American Patrohini, Ent. Amer., vol. 18, No. 4, pp. 135-183, 1938, in which he
discusses Thalassotrechus in some detail in a long footnote on pages 143-144, and states
that the two forms are best considered subspecies. This suggested change has not yet been
recorded in the Leng supplements.

214 San Diego Society of Natural History

to be a barrier to the intertidal Coleoptera. Only one species of the genus
Endeodes is known to occur on both sides of this area, the species of
Diaulota are distinctly separated near this region, and Liparoccphalus cordi-
collis LeConte and several other northern species are not known south of
San Luis Obispo County. These observations are in accord with those
of marine zoologists, many of whom consider Point Conception a divid-
ing line between two faunal areas.

The following is the distribution of the two subspecies as represented in
my collection:

Thalassotrechus barbarae barbarae (Horn)

Punta Morro, north of Ensenada, Baja California, Mexico: adults and
larvae in November, larvae in April.

La Mision de San Miguel, Baja California: adults in November.

San Diego, California: adults in October, larvae in March, April, and
November.

Gaviota, Santa Barbara County, California: adults in October and No-
vember.

Thalassotrechus barbarae nigripennis Van Dyke

Pacific Grove, Monterey County, California: adults and larvae in October.
Moss Beach, San Mateo County, California: adults and larvae in October.
Van Dyke reported it also from Marin County, California.

It must not be assumed from the foregoing dates that these forms are
necessarily found only in the spring and fall. The main reason for these
capture dates is probably that the very favorable spring and fall tides lead
one to search for intertidal insects at these times. On the summer trips
time is spent more commonly on the beach when the kelp fauna is richest
and when low tides are usually at an unfavorable time of day. Thalasso-
trechus is not particularly common, but when it is found, several speci-
mens are often together. It is encountered near the upper reaches of the
intertidal area in well-drained cracks in the rocks, in association with
Liparocephalus cordicollis LeConte in central California, and with Diaulota
fulviventris Moore in southern California and Baja California. As when
collecting other intertidal rock-dwelling insects, the cracked rocks must
be split open with a wrecking bar or some other implement to expose the
insects. When uncovered, the adults run for shelter rather rapidly, but by
no means as fast as most small terrestrial carabids. The larvae are more
sluggish than the adults and probably feed on the larvae of chironomid
flies, which are often found with them.

Moore — Intertidal Coleoptera 2 1 5

Larva of Thalassotrechus barbarae Horn

Body elongate, cylindrical, somewhat depressed. Color testaceous to
piceous, head and pronotum often lighter than abdomen. Integuments very
shining, heavily chitinized, glabrous except for rather sparsely placed,
long, strong setae. Head quadrate, about as wide as long. Eyes of six ocelli
on each side near the anterior angles; anterior three ocelli largest. Coronal
suture meeting frontal sutures at angles of 120 degrees. Frontal sutures
biarcuate, the two arcs meeting in an inward pointing cusp. Mandibles
falcate, slender, as long as head, with a small tooth at the middle of the
inner margin. Antennae half as long as mandible; the last and penultimate
segments each strongly geniculate; the penultimate bearing a small acorn-
shaped seta at middle of its strongly diagonally beveled apex and one long
seta on each side of apex; last segment with three long, strong setae at
apex. Maxilla with lacina not separate from the stipes, galea with segments
of equal length, the first segment with a long seta at the tip, second seg-
ment geniculate to first. Labial palpi two-segmented, first segment three
times as long as wide, second one-third as wide as first and one-third as
long. Ligula very small, one-fifth as long as first segment of labial palpi,
semi-quadrate with a very long, stout seta at tip and a small seta on each
side of tip. Thorax with first segment at widest part of body, a little
longer than second or third, subquadrate but arcuate at posterior margin;
second and third segments subquadrate, each considerably wider than long.
Abdomen long, slender, tapering toward the tip; the tergal plates sepa-
rated from the pleural plates by wide areas of membrane; tergal plates
with distinct longitudinal suture. Urogomphus long, slender, not seg-
mented, with three long spines at outer edge and three more at tip. Pseudo-
pode two-thirds as long as urogomphus. Legs pale, not strong, bearing
one sharp claw. Length 7.5 mm.

STAPHYLINIDAE
SUBFAMILY ALEOCHARINAE

The larvae of only four of the ten genera at present placed in the
subtribe Phytosi of the tribe Bolitocharini have been discussed in the
literature. As the larvae of Diaulota have never been described in detail,
and larvae of four of the five known species are at hand, it seems worth-
while to describe these and to compare them with the known larvae of the
other genera.

216 San Diego Society of Natural History

KEY TO THE KNOWN LARVAE OF THE GENERA
OF THE PHYTOSI

A. Apex of ninth abdominal segment truncate Pbyfosm

AA. Apex of ninth abdominal segment produced in two prominent appendages.

B. Urogomphus two-segmented Antarctophytosus

BB. Urogomphus less than two-segmented

C. Antennae three-segmented Liparocephalus

CC. Antennae four-segmented Diaulota

PHYTOSUS Curtis

The larva of Pbytosus nigriventris (Chevrolat) was described and
figured by Fauvel in 1862. According to him, the antennae are of four
segments; the first and last are small, and the supplementary segment (a
modified seta of the second) is unusually long (it is figured as being about
half as long as the third segment). The mandibles have no visible tooth
internally. The eyes are of a single ocellus on each side. The maxillary
palpi are of three segments; the first is quite small, the second several times
as long, and the third long, setiform and sharply pointed, so that the entire
palpus has the aspect of a long spine. He called particular attention to the
enlarged middle and posterior femora. The eighth abdominal segment has
a noticeable tubercle above, in the center of the apical margin. The ninth
segment is apparently truncate without the prolongations (urogomphi)
typical of Diaulota.

ANTARCTOPHYTOSUS Waterhouse

I am indebted to Renaud Paulian for bringing to my attention the
fact that in a previous paper on the Phytosi I overlooked that, in 1941,
he described the true larva of Antarctophytosus and showed that Ender-
lein's earlier description was based on a misidentification. According to
Paulian, the antennae are three-segmented with the modified seta not as
long as segment three. The mandibles are bidentate at the apex. The three-
segmented maxillary palpi have the last segment acuminate. The labial
palpi are two-segmented. The urogomphi are two-segmented with the
first segment shorter and stouter than the second.

LIPAROCEPHALUS Maklin

Saunders in 1928, and Chamberlin and Ferris in considerably greater
detail in 1929, have published figures of the larvae of Liparocephalus cordi-
collis LeConte (as L. brevipennis Maklin). I have specimens of the larvae
from Pigeon Point, San Mateo County, California, before me, so any dif-

Moore — Intertidal Coleoptera 217

ference between the following brief description and former ones should
be construed as my own opinion. The antennae are of three segments, the
first short and wide, the second long and stout, the third slender but
longer than the modified setae which with it occupies the apex of the sec-
ond segment. The mandibles are curved, with a median tooth internally
and with serrations basad from the median tooth. The eyes are of
a single facet on each side. The maxillary palpi are three-segmented and
the labial palpi two-segmented. The eighth abdominal tergite is produced
medially near its apex in a large, dark, very prominent tubercle. The uro-
gomphi are produced on each side of the apex of the ninth segment as a
short, stout process to which is articulated a long, slender, stalk-like
process, bearing at its tip a long seta.

DIAULOTA Casey

The larva of this genus has been figured in toto by Saunders in 1928,
and some parts have been shown in greater detail by Chamberlin and Fer-
ris in 1929. The antenna is of four segments, with the modified seta at the
apex of the second, either longer or shorter than the third segment, de-
pending on the species. The mandibles bear an internal tooth. In one species
(D. fnlviventris Moore) there are serrations between the median tooth and
the tip. The eyes consist of a single facet on each side. The maxillary palpi
are of three segments and the labial palpi of two. The tubercle at the
dorsal apex of the eighth segment is not as pronounced as in Liparocep-
balus. The urogomphi are very large, variable specifically, and usually with
a very minute segment articulated at the point.

The determinations as to species were made largely by association and
elimination, but I am reasonably certain that they are correct.

KEY TO THE KNOWN LARVAE OF THE SPECIES OF DIAULOTA

A. Mandibles serrate internally between median tooth and tip fulviventris

AA. Mandibles simple internally between median tooth and tip.

B. Urogomphi bent sharply upward at tip densissima

BB. Urogomphi straight or nearly -so.

C. Modified seta of second antennal segment at least as long as third .... vandykci
CC. Modified seta of second antennal segment much shorter

than third megacephala

Larva of Diaulota densissima Casey

Body elongate and subparallel, a little wider toward apex of abdomen.
Integuments well chitinized, shining, sparsely punctured, and glabrous
except for numerous setae, which are more abundant toward the sides.
Color dark grey, with eighth abdominal segment black and ninth reddish;

218 San Diego Society of Natural History

under parts very pale. Head nearly round, as long as wide. Mandibles
without serrations between median tooth and tip; median tooth very
large. Antenna with first segment shorter than wide; second rectangular,
over twice as long as wide, with a seta on each side near apex; third seg-
ment less than half as wide as second, about twice as long as wide, distinct-
ly shorter than acorn-like seta at apex of second, with two small setae
approximated near apex; fourth segment half as wide as third and about
as long as wide, with two apical setae. Pronotum nearly square, widest
at apex; sides nearly straight; about as wide as head. Second thoracic seg-
ment about half as long as pronotum and a little wider; third a little
shorter and a little wider. Abdomen with first segment about as wide as
thorax but somewhat shorter than third thoracic segment, second to
seventh each a little longer than preceding, eighth longer than seventh
but narrower, with the apex produced posteriorly in the center in a small
wide lobe. Urogomphi bend sharply upward at tip. Legs pale, shining
with a few setae. Length 1.8 mm.

This description is based on a specimen taken at Pigeon Point, San
Mateo County, California, in October, in company with adults. This
agrees well with the larva chosen by Chamberlin and Ferris as representa-
tive of this species.

The very long, modified seta of the second antennal segment and
the sharply upturned urogomphus readily separate this species from the
others.

Larva of Diaulota fulviventris Moore

Body elongate,' subparallel, a little wider posteriorly. Integuments well
chitinized, shining, sparsely punctured, and glabrous, except for numerous
setae, particularly toward the side. Color black to reddish brown, paler
below, with urogomphi testaceous. Head nearly round, a little narrowed
in front. Mandibles with serrations between median tooth and tip. An-
tennae with first segment wider than long; second narrower, more than
twice as long as wide, with two setae, one near the middle and the other
near the apex; third segment half as wide as second, about twice as long
as wide; fourth very small; modified seta at apex of second segment about
half as long as third segment. Pronotum considerably wider than head,
nearly twice as wide as long, second and third segments shorter but as
wide as first. Abdomen very similar to that of D. densissima Casey.
Length 1.6 mm.

Four specimens from Punta Morro, north of Ensenada, Baja Cali-

Moore — Intertidal Coleoptera 219

fornia, Mexico, two in April and two in November, and nine specimens
from La Jolla, San Diego, California, in September and November, all
taken in company of the adults.

The larva of this species differs from all the other larvae by having
serrations between the median tooth and tip of the mandibles. The larvae
of this species and those of D. dcnsissima Casey are more easily separated
than are the adults of the two.

Larva of Diaulota vandykei Moore

Elongate, subparallel, but widest near the abdominal apex. Integu-
ments shining but very minutely roughened. Color yellow to reddish
brown, with eighth abdominal segment sometimes black. Head somewhat
quadrate as in the adult. Mandibles without serrations between median
tooth and tip. Antennae with first segment about as long as wide; second
narrow, twice as long as wide, with a small median beanlike seta, on each
side of which is a long seta; third segment three times as long as wide,
with a seta on each side of apex; fourth short, modified seta at apex of
second segment about as long as third segment. Pronotum subquadrate,
a little wider than long; sides evenly arcuate. Second thoracic segment
less than half as long as pronotum, a little wider; third about equal to
second. Abdomen with eighth segment broad. Urogomphus curved up-
ward slightly at tip. Length 1.9 mm.

Seven specimens from Shell Beach, San Luis Obispo County, Cali-
fornia, were taken in company with over one hundred adults from a
reef exposed near mean low water. Although these are not from the type
locality and differ somewhat from the type series, I believe that they be-
long to this species.

The length of the modified antennal seta readily distinguishes this
species.

Larva of Diaulota megacephala Moore
or Diaulota harteri Moore

Elongate, wider posteriorly. Integuments smooth, shining, and gla-
brous, except for scattered setae. Color pale yellow to ferruginous, with
a small black spot near the center of the eighth tergite. Head ovoid, wider
than long. Mandibles without serrations between median tooth and tip.
Antenna with first segment short; second twice as long as wide, a seta
each side of apex; third less than half as wide as second, more than twice
as long as wide, with a seta at apex; fourth very small, with two small

220 San Diego Society of Natural History

setae at apex; modified seta at apex of second segment as wide as third
segment but only two-thirds as long. Pronotum subquadrate, wider than
long, wider than head. Second and third segments about as wide as prono-
tum and about half as long. Abdomen with eighth segment produced
in the center of the apical margin in a small, black, truncate lobe.
Urogomphus long, slender, sharply pointed, and not curved upward at
the tip. No minute articulation at tip of urogomphus. Length 1.4 mm.

Eight specimens taken at La Jolla, San Diego, California, in company
with adults of both species which are usually found together. Because of
the larger head it seems more likely that this is the larva of megacephala
than of barter/. The former is the commoner species.

This larva is easily known by the simple tips of the mandibles and by
the short antennal seta, as well as by the straight urogomphus.

MALACHIIDAE
ENDEODES LeConte

Of the five known species of Endeodes, I have examples of the larvae
of the three semi-intertidal species. The larvae of the two southern species
are unknown but apparently are not intertidal rock-dwellers; the adults
are seashore inhabitants in the dry kelp above high tide. I believe no larva
of this genus has as yet been described. The three closely related species
discussed here are all found on the sea beaches of central California. Some-
times the adults are found under debris on the sandy beaches, but more
commonly I have found adults and larvae together in damp cracks in the
rocks just at, or just above, the high-tide mark.

Larva of Endeodes collaris (LeConte)

Body fusiform, roughly three and one-half times as long as wide.
Color dark sand to piceous, with the head often ferruginous, the legs and
undersides lighter, and the ninth abdominal segment entirely black above.
Sometimes a triangular black spot in center of head, a large oblong central
black spot and two small lateral black spots on pronotum, a small black
spot on each side of second and third thoracic segments. First to eighth
abdominal segments and usually all three thoracic segments with a pale
circle on each side, in the center of which is a minute black spot. These
pale circles are hardly apparent on the thorax of dark specimens. Head
nearly square, shining, finely granulose throughout, sparsely and very
finely pubescent with a few long setae near the ocelli. Eyes of four ocelli
near the anterior angle, with the three front ocelli in a row. Coronal suture

Moore — Intertidal Coleoptera 221

meeting the straight frontal sutures at an angle of 120°. Mandibles deeply
notched at tip to form two teeth, one dorsal to the other and with a small
median tooth at inner edge. Antennae very short, retractable in a telescop-
ing manner; first segment a little longer than wide; second narrower, half
as long as wide, with three setae near apex; third segment less than half
as wide as second and twice as long, with two setae remote from apex;
the large acorn-shaped setae of second segment wider than third segment
and nearly as long. Labrum distinct, apically arcuate. Maxillary palpi short
and robust; each segment wider than long except the last, which is more
than twice as long as wide. Labial palpi two-segmented; first segment
twice as long as wide; second narrower but of same proportions. Ligula
about as long as first segment of labial palpi, wide and broadly rounded
at apex. Thorax not strongly chitinized, rather densely and minutely
punctured throughout, and clothed with a very fine, pale pubescence; a
strong lateral seta on each side of each segment; third segment widest,
almost twice as wide as head. Pronotum subquadrate to oval, as long as
head, half again as wide as long; second segment wider than first, more
than twice as wide as long; third segment about as long as second and a
little wider. Abdomen not chitinized, with first eight segments gradually
narrower toward apex; first segment a little shorter than the others, which
are subequal in length. Surface finely, not densely punctured, and finely,
transversely rugulose, finely pubescent with a few moderate setae. Ninth
segment heavily chitinized, finely granulose, finely pubescent with num-
erous strong setae. Urogomphus not segmented, twice as long as wide,
pointed; apex with a very small upturned hook. Legs slender, pale, sparsely
pubescent and shining, with a single claw at apex. Length 5 mm.

Three specimens were taken at Santa Cruz, California, in September,
in company with five adults. Five taken in October, at Pacific Grove,
Monterey County, California, in cracks in the rocks just above high
tide and just above the area in which Liparocepbaliis and Diaulota are
found in this region. The rocks were covered with a heavy coating of
bird droppings.

Larva of Endeodes rugiceps Blackwelder

Very similar to the larva of E. collans (LeConte) but lighter in
color and with the urogomphus ferruginous above and below, at least at its
apical half.

Seven larvae from Pacific Grove, Monterey County, California; one
adult and twenty-five larvae from Shell Beach, San Luis Obispo County,

222 San Diego Society of Natural History

California. At Shell Beach a point of soft sandstone dividing two small
coves terminates in a precipitous cliff which, except at low tide, is mois-
tened by the spray of a heavy surf. All the specimens were found in a
horizontal crack at the top of a ledge which was about halfway down
the cliff and about five feet above extreme high water. As pieces of stone
were pried away from the ledge, the lower surface of the crack was ex-
posed, revealing several larvae in an area about a foot square. Each larva
was in a slightly oblong cell bounded by loose debris. The cell was
about twice the length of the insect and about one-sixteenth of an inch
thick (the thickness of the crack). From each cell ran a poorly defined
pathway an inch or two long leading out to other areas of the crack.
This pathway, which was about the width of the insect, was bordered
by the same loose debris as the cell. It was most sharply defined where it
joined the cell, becoming gradually more indistinct until it disappeared.
The cells were separated from each other by several inches.

Larva of Endeodes insularis Blackwelder
A single larva and a single pupa were taken in company with several
adults at Gaviota State Park, Santa Barbara County, California, and are
assigned to this species because of their association with the adults. They
were taken in company with Thalassotrechus barbarae (Horn) at different
spots in the cracks of a tilted sedimentary deposit which would be covered
by the high tide. This larva is inseparable from those of E. riigiceps Black-
welder and may belong to that species rather than this one.

Pupa of Endeodes insularis (?) Blackwelder

Body fusiform, strongly flattened dorsoventrally; head deflexed. Pale,
testaceous throughout, transluscent; integuments soft and flexible, sparsely
pubescent, and with numerous large setae. Nearly-formed body of adult
clearly distinguishable within. Head, thorax, and abdomen each forming
a close fitting, clearly demarked cover for those segments of adult. Each
leg and each antenna with separate sack which closely approximates shape
of these parts in adult except for being considerably stouter. Antennal
sack clearly segmented and about twice as wide as the adult antenna, which
can be seen lying loosely within. Length. 3.5 mm.

It is not possible to identify this pupa with certainty as that of E.
insularis Blackwelder, but the association with the adults makes such an
identification very likely correct.

Moore — Intertidal Coleoptera 22 3

BIBLIOGRAPHY

Bernhauer, Max and Scheerpeltz, Otto.

1926. Coleopterorum catalogus, pars 82, Staphylinidae VI, pp. 499-
988. Berlin.

Blackwelder, Richard Eliot.

1932. The genus Endcodes LeConte. Pan.-Pac. Ent., vol. 8, pp. 128-
136, 3 figs.

Boving, Adam G. and Craighead, F. C.

1930. An illustrated synopsis of the principal larval forms of the
order Coleoptera. Ent. Am., vol. 11, pp. 1-3 51.

Casey, Thomas Lincoln.

1893. Coleopterological notices, V. Ann. New York Acad. Sci., vol.
7, pp. 281-606.

Chamberlin, J. S., and Ferris, G. F.

1929. On Liparocepbalus and allied genera. Pan.-Pac. Ent., Vol. 5,
pp. 137-143, 153-162.

Chevrolat, Louis Alexandre Auguste.

1843. Description d'une nouvelle espece du genre Myrmedonia.
Revue Zoologique, vol. 6, pp. 42-43.

Enderlein, Giinther.

1909. Die Insekten des Antarktischen Gebietes. Deutsche Siidpolar-
Exp., vol. 10, Zool. II, (1908) 1909, pp. 361-518.

Fauvel, Albert.

1862. Notice sur quelque aleochariens nouveaux ou peu connus et
description de larves de Pbyfosus et Lep/usa. Ann. Soc. Ent.
France, ser. 4, vol. 2, pp. 81-94.

Fenyes, Adelbert.

1918. Coleoptera: Fam. Staphylinidae, subfam. Aleocharinae. Gen-
era Insectorum, fasc."173a, pp. 1-110.

1920. Coleoptera: Fam. Staphylinidae, subfam. Aleocharinae. Gen-
era Insectorum, fasc. 173b, pp. 111-414.

Horn, George Henry.

1872. Synopsis of the Malachiidae of the United States. Trans. Am.

Ent. Soc, vol. 4, pp. 109-127.
1892. Random studies in North American Coleoptera. Trans. Am.

Ent. Soc, vol. 19, pp. 40-48.

224 San Diego Society of Natural History

LeConte, John Lawrence.

18 52. Catalogue of the Melyrides of the United States with descrip-
tion of new species. Proc. Acad. Nat. Sci. Phil., vol. 6, pp.
163-171.

1860. Description of some genera and species of Coleoptera from
the vicinity of the southern boundary of the U. S. Arcanae
Naturae, vol. 3, pp. 121-128.

18 80. Short studies of North American Coleoptera. Trans. Am.

Ent. Soc, vol. 8, pp. 163-218.
Moore, Ian.

19 56. A revision of the Pacific Coast Phytosi with a review of the

foreign genera. Trans. San Diego Soc. Nat. Hist., vol. XII,
No. 7, pp. 103-152.
Paulian, Renaud.

1941. Les premiers etats des staphylinoidea. Mem. Mus. Nat. Hist.
Nat., vol. 15, pp. 1-361, 1367 figs., 3 pis.
Saunders, L. G.

1929. Some marine insects of the Pacific Coast of Canada. Ann. Ent.
Soc. Am., vol. 21, pp. 521-545.
Schaeffer, Charles.

1901. Synopsis of the species of Trechus, with description of a new
species. Bull. Am. Mus. Nat. Hist., vol. 14, pp. 209-212.
Van Dyke, Edwin Cooper.

1918. New inter- tidal rock-dwelling Coleoptera from California.
Ent. News, vol. 29, pp. 303-308.

Explanation of Figures
Plate 14

Figure 1. Urogomphi and pseudopode of larva of Thalassotrcchus bar-
bar ae (Horn).

Figure 2. Ocelli of larva of Thalassotrcchus bar barac (Horn).

Figure 3. Larva of Thalassotrcchus barbarac (Horn), dorsal view.

Figure 4. Labium and maxilla of larvae of Thalassotrcchus barbarac
(Horn).

Figure 5. Mandible of larva of Thalassotrcchus barbarae (Horn).

Figure 6. Posterior leg of larva of Thalassotrcchus barbarac (Horn).

Figure 7. Antenna of larva of Thalassotrcchus barbarae (Horn).

Moore — Intertidal Coleoptera 22 5

Plate 15

Figure 8. Antenna of larva of Diaulofa densissima Casey.
Figure 9. Mandible of larva of Diaulofa densissima Casey.
Figure 10. Outline of body of larva of Diaulofa densissima. Casey, dorsal

view.
Figure 11. Ninth abdominal segment of larva of Diaulofa densissima

Casey, lateral view.
Figure 12. Antenna of larva of Diaulofa fulviventris Moore.
Figure 13. Mandible of larva of Diaulofa fulviventris Moore.
Figure 14. Larva of Diaulofa fulviventris Moore, dorsal view.
Figure 15. Ninth abdominal segment of larva of Diaulofa fulviventris

Moore, lateral view.

Plate 16

Figure 16. Antenna of larva of Diaulofa vandykei Moore.

Figure 17. Mandible of larva of Diaulofa vandykei Moore.

Figure 18. Outline of body of larva of Diaulofa vandykei Moore, dorsal
view.

Figure 19. Ninth abdominal segment of larva of Diaulofa vandykei
Moore, lateral view.

Figure 20. Antenna of larva of Diaulofa megacephala Moore or of Diau-
lofa harteri Moore.

Figure 21. Mandible of larva of Diaulofa megacephala Moore or of Diau-
lofa harteri Moore.

Figure 22. Outline of body of larva of Diaulofa megacephala Moore or
of Diaulofa harteri Moore.

Figure 23. Ninth abdominal segment of larva of Diaulofa megacephala
Moore or of Diaulofa harteri Moore.

Plate 17

Figure 24. Larva of Endeodes collaris (LeConte), dorsal view.

Figure 2 5. Antenna of larva of Endeodes collaris (LeConte).

Figure 26. Labium and maxillary palpus of larva of Endeodes collaris

(LeConte) .

Figure 27. Mandible of larva of Endeodes collaris (LeConte).

Figure 28. Pupa of Endeodes insularis (?) Blackwelder, lateral view.

Figure 29. Pupa of Endeodes insularis (?) Blackwelder, ventral view.

Figure 30. Pupa of Endeodes insularis (?) Blackwelder, dorsal view.

226

Moore — Intertidal Coleoptera

PLATE 14

FOR EXPLANATION OF FIGURES SEE PAGE 224

Moore — Intertidal Coleoptera

227

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Moore — Intertidal Coleoptera

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TRANSACTIONS j jj|jfe

OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY

Volume XII, No. 12, pp. 231-262 Plates 18-21

A NEW SPECIES OF MEGATHYMUS
FROM BAJA CALIFORNIA, MEXICO

(Lepidoptera : Megathymidae)
BY

Charles F. Harbison

Curator of Entomology
San Diego Society of Natural History

SAN DIEGO, CALIFORNIA
Printed for the Society

March 15, 1957

TABLE OF CONTENTS £ WflffilFf \

Frontispiece, Plate 18 !T.....'.7234

Introduction 235

Acknowledgments 237

Plant Association 238

Laboratory Procedure _ 239

Megathymus corns toe ki, new species

Type Specimens _ 241

Holotype Male 242

The head and its appendages _ 242

The thorax and its appendages 243

Wings of holotype 243

Upper surface 243

Under surface 244

Legs of holotype 245

Tarsi 245

The abdomen and its appendages 246

Allotype Female 246

The head and its appendages 246

The thorax and its appendages 247

Wings of allotype 247

Upper surface 247

Under surface 248

Legs of allotype 248

The abdomen and its appendages 249

Study of the Paratype series 250

Appendages of the head 250

Antenna 250

Labial palpus 250

The thorax and its appendages 250

The wings 251

Tarsus 252

Genitalia 252

Comparison of the new species with related species 252

Megathymus stepbensi Skinner 253

Megathymus mariae Barnes and Benjamin 254

Megathymus polingi Skinner 255

Summary 256

Bibliography 256

Explanation of figures, Plates 19 - 20 258

Plate 19 259

Plate 20 260

Plate 21 261

Upper Panel

Agave shawii Engelmann {orcuttiana Trelease) as it grows on the hills east of San
Simon, Baja California. This is the host plant of Megathymus comstocki. The photo-
graphs from which the cuts were made, were reproduced from kodachromes taken by
the author in the type locality of the new species.

Lower Panel

Typical group of A. shawii in the type locality of Megathymus comstocki- The
relatively large size of the agave will be particularly noted. E. P. Chace at right, in
readiness for collecting.

A NEW SPECIES OF MEGATHYMUS
FROM BAJA CALIFORNIA, MEXICO

(Lepidoptera : Megathymidae)
BY

Charles F. Harbison

Curator of Entomology
San Diego Society of Natural History

INTRODUCTION

In the literature covering the Lepidoptera of Baja California,
Mexico, there is little information on the megathymids. No Megathy-
mus is listed in the Rindge 1948 paper, "Rhopalocera of Lower
California." Dr. Cockerell's 1941 article, "Observations on Plants
and Insects in Northwestern Baja California, Mexico, with Descrip-
tions of New Bees," does not list any species of these interesting
butterflies. In Dr. Hoffmann's, "Catalogo de Lepidopteros Mexicanos"
of 1941, two species, M. yucca navajo Skinner and M. stephensi Skinner
are given as occurring in Baja California. No exact locality is mentioned
for either. I believe that the species are listed because their known
occurrence in adjacent San Diego County, California, suggested a
probable extension of range south of the international boundary.

In late August of 1953, the new megathymid described in this
paper was discovered during a collecting trip with Mr. Laurence M.
Huey, Curator of Birds and Mammals, San Diego Society of Natural
History, Mr. Huey's wife, Mrs. Eva Huey, and Mr. Wesley Farmer
of the Museum's Activities Department. We camped about two miles
northeast of San Simon on the north side of San Simon (Santa
Maria) River Valley. Dr. Carl L. Hubbs of Scripps Institution of
Oceanography, who has often visited the region states that "the
locality is about six miles east-southeast of the present village of San
Quintin, which is located at the old mill, approximately three miles
northwest of the old, now abandoned village of San Quintin." Both
localities are on Bahia San Quintin. (Lat 30° 23' N).

Mr. Farmer and the author succeeded, after much effort, in
collecting a series of ten males and two females. The butterflies are
strong and rather erratic flyers and to net one requires considerable
stalking and agility. The specimens captured were in very poor shape
by the time they were quieted by the cyanide killing-jar fumes. Anxious
to obtain a longer series and more perfect specimens before describing

236 San Diego Society of Natural History

this new insect, the author made two return trips to the vicinity, one in
December, 1953, and the other on August 11, 1954. He planned to
collect the immature stages in the host plant and to rear the imagines.
These could be killed before they had damaged themselves. No
larvae or pupae were obtained on either trip. The adult butterflies
were not expected to be on the wing on the December visit but it
was hoped that some could be captured on the August 11th trip.
Apparently this was too early. On this latter trip, the author was
accompanied by Mr. Jerry Powell and Mr. F. James Rohlf, both
enthusiastic entomologists. If the butterflies had been on the wing,
some certainly would have been seen.

Early in September, 1955, the author, accompanied by a group
of his entomological friends, returned again to the vicinity of the
first collection of the new species. On September 6, his cousin, Mrs.
Marion Beckler, an author of children's books, accompanied him down
to Colonia Guerrero where they met two other members of the
expedition, Mr. Ian Moore, Research Associate in Coleoptera at the
San Diego Natural History Museum, and Mr. Joe McKenney, a
pre-medical student at Stanford University. Mr. McKenney had
succeeded in netting one of the butterflies at Camalu, several miles
north of Colonia Guerrero and almost thirty miles north of the
original locality. The next day, these four went south to Laguna
Santa Maria where they spent part of two days and one night
collecting. No megathymids were taken and none were expected.

On September 8, the four members of the expedition returned
to San Simon, up the river valley at the base of the low, flat-topped
hills, about three miles northeast of the ponds and beach. Mr.
Moore then headed north to the United States. The three remaining
members of the early group, while exploring the hillsides and flat
mesas above the store at San Simon, captured several specimens of
the new butterfly. That evening the party was joined by Dr. Francis
X. Williams, Research Associate in Hymenoptera at the Museum,
his wife, Mrs. Louisa C. Williams, Dr. John Adams Comstock, a
Director of the Society of Natural History, the Museum's Editor
of publications, and an experienced collector of Megathymus larvae
and pupae, and Mr. E. P. Chace, Curator of Conchology and Marine
Invertebrates of the San Diego Society of Natural History. The
party camped about two miles northeast of San Simon on the north
side of the broad San Simon River valley near an old eucalyptus tree
on the floor of a branch canon.

A New Megathymus. C. F. Harbison 237

On the next day the entomologists proceeded to work the agaves
growing north of the camp site, for larvae and pupae. We found
that with an ax and machete, we could cut the long horizontal trunk
of each plant below the green basal leaves. We then proceeded to tip
the plant over so that the sharp terminal spines on the leaves were
forced into the ground below. This anchored the plant in an up-side-
down position. The entomologists and the conchologist then removed
one leaf at a time from the stalk, examining each leaf, in turn, for
indication of megathymid infestation. The infested leaves were placed
in pillow cages made of copper screen-wire cloth. These were sealed
by folding over the screen at the open end and running a piece of
wire through the fold to keep it closed. The infested leaves were
brought back to the United States under special permit from both
Mexican and American Governments and the immature stages reared
through to imago by Dr. Comstock. Later in the day Mr. Paul Arnaud,
a staff member of the California Bureau of Plant Quarantine, based
in San Diego, and Mr. Arthur Lee of the San Diego City and
County Health Department joined the party. The combined group
collected a fine series of specimens constituting most of the types
of the new species.

Acknowledgments

The author wishes to express his appreciation to his many friends
for the assistance given. Mr. Lee Passmore, local naturalist and
nature photographer, made a series of kodachromes of the adult
insect. Mr. T. W. Pace, a commercial photographer, took the black-
and-white photographs that illustrate several type specimens (Plate 21 ).
Dr. H. Avery Freeman of Garland, Texas, gave valued advice and
furnished a male and female paratype of M. maculosa Freeman. On
three separate visits, Mr. Lloyd M. Martin, Associate Curator of the
Department of Entomology in the Los Angeles County Museum,
made available for study the entire collection of his institution, and
loaned the author two males and two females of Megathymus mar'iae
Barnes and Benjamin and one female of M. polingi Skinner. He also
permitted the author to make slides of the genitalia of these specimens.
His advice and suggestions were very valuable. Mr. Fred Thorne
of the San Diego County Department of Agriculture made important
suggestions and gave the author some megathymids. Mr. Wesley
Farmer collected most of the original series of the new species in 1953.

238 San Diego Society of Natural History

To Mr. Joe McKenney is due our particular thanks for the long
series of passably good material of the new species he collected and
submitted. Some of these were given to the Society; in fact, the
holotype male was supplied through the courtesy of Mr. McKenney.
Mr. E. P. Chace worked arduously with the author and Dr. Comstock
in digging the immature insects out of the century plants. Mrs.
Mildred H. Meeder, the Librarian of the San Diego Society of
Natural History, has helped by obtaining needed publications. To
Colonel Arthur F. Fischer, who was Director of the San Diego
Natural History Museum while the study was being made, the
author wishes to extend thanks for his continuous interest. To Dr.
John Adams Comstock is due special thanks for his interest and for
his enthusiastic help in the field. The new species is named in honor
of this outstandng lepidopterist.

PLANT ASSOCIATION

Both north and south of the broad, sandy Arroyo San Simon
are high, flat-topped rocky mesas covered with extensive growths
of a century plant or maguey, which is the host plant of the new
butterfly. We have heretofore been designating this plant Agave
orcuttiana Trelease, the type of which was collected at San Quintin in
1886 by Charles Russell Orcutt, for whom the species was named by
Dr. William Trelease in his monograph on "The Agaves of Lower
California." This name was considered valid by Alwin Berger in
his "Die Agaven" in 1915.= Dr. L. H. and Ethel Zoe Bailey listed
Agave orcuttiana Trelease, in their 1941 "Hortus Second." In Dr.
Bailey's 1935 "Hortus" there is no listing of A. orcuttiana but in his
1944 reprinting of his "Encyclopedia of Horticulture" he gave A.
orcuttiana under his heading, Agave shawii, as a related species. Dr.
Ivan M. Johnston in his 1924 report on the botany of "the Expedition
of the California Academy of Sciences to the Gulf of California in
1921", stated that in his opinion " 'Agave shawii Engelmann includes
A. sebastiana Greene, A. orcuttiana Trelease, A. pachyacantha Tre-
lease, and A. goldmaniana Trelease." Dr. Reid Moran and Dr. George
E. Lindsay, in their 1951 paper on "San Benito Island"4 stated that,
"in view of the intergradation that apparently exists, some of these
species seem rather weak." In conformity with these later opinions we are

1 Rep. Mo. Bot. Gard. 22 : 47, 1912

2 Gustav Fischer, Jena, 1915. p. 17 1.

3 Proc. Calif. Acad. Sciences 12 : 1003. 1924
* Desert Plant Life, 23 : 83. 1951

A New Megathymus. C. F. Harbison 239

considering the species of agave occurring at and near San Quintin
as only a non-specific variation of a widely ranging species, Agave
shawii Engelmann.

The high mesa country north of the San Simon River Valley is
dissected by many steep-sided gullies and small canons. These drain
southeastward and enter the main valley almost at right angles.
Many of these canons have a number of forks as they penetrate the
great mesa and drain it during the infrequent rains. By following up one
of these forks to its source one finds himself on a flat-topped, rock-
covered plateau that extends almost level for miles, and is covered
with little vegetation.

The agave is the dominant plant of the canon sides and hill tops
near the large valley. (See plate 18.) It seems to prefer to grow
in rocky ground along with PITAHAYA agria, Machaerocereus gummosus
(Englemann) Britton and Rose, and COCHAL, Myrtillocactus cochal
(Orcutt) Britton and Rose. GRAY bur-shrub., Franseria chenopo-
diifolia Bentham, and the low growing California box-thorn, Lycium
californicum Nuttall also clothe the steep rocky sides of the branch
canons. On the flat-topped mesas near the main valley are numerous
specimens of the BRANDEGEE WREATH CACTUS, N eomammillaria bmnde-
geei (Coulter) Britton and Rose, and a low, flat barrel, ford bisnaga,
Ferocactus fordii (Orcutt) Britton and Rose. Here also is found
parry BUCKEYE, Aescidus parryi A. Gray, several species of CHOLLA,
Opuntia spp., SAWTOOTH GOLDENBUSH, Haplopappus squarrosus
Hooker and Arnott, and CLIFF SPURGE, Euphorbia misera Bentham.

The floor of the side canon is covered with large shrubs of rich
box-thorn, Lycium richii A. Gray, LAUREL SUMAC, Rhus laurina
Nuttall, palmer goldenbush, Haplopappus palmeri A. Gray, desert
fragrance, Hymenoclea monogyra Torrey and Gray, and shad-scale,
A triplex canescens (Pursh) Nuttall. Also, not far south of camp are
several fine clumps of rush milkweed, Asclepias subulata Decaisne,
and that very beautiful cactus of Baja California, CABEZA VIEJA, Lopho-
cereus schottii (Engelmann) Britton and Rose.

Laboratory Procedure

Briefly, the laboratory procedure used in this study of the new
species is as follows. The antenna, palpus, and legs of one side of the
holotype were removed and studied and drawings made thereof.

240 San Diego Society of Natural History

The antenna was drawn with scales in place and then glued back on
the head. An antenna of one of the paratypes was removed and
soaked for twenty-four hours in each of the following: 10 per cent
sodium hydroxide solution, water, 95 per cent alcohol and, finally,
xylene. Later it was mounted on a microscope slide in clear Canada
balsam. This procedure removed the scales and slightly distorted
each of the segments, but every joint in the antenna became exposed
and easy to count and study. The same method was used on the
labial palpus and in the study of male and female genitalia. A right
palpus was removed from one of the paratypes and, after descaling
dry, was studied to see what effect the soaking in solutions had
produced on this appendage of the holotype, and then was drawn
from three aspects.

The head was removed from one of the paratypes, making it
possible to lift the patagium from the prothorax. This was descaled
after the arrangement of scales had been studied. The tegula was
removed from the mesothorax at base of the fore wings and the
arrangement of the scales noted. It was then descaled.

The legs removed from the holotype were studied with scales in
place. They were then descaled and studied from both the upper
and lower aspects in the dry state. (Plate 19.) The claw-bearing
segment of the tarsus from one of the paratypes was mounted on a
slide after it had been expanded by the same method used with the
palpus (Plate 19). The wings from the right side of one of the
paratypes were removed, descaled dry, and then mounted between
two glass slides. Later these were projected by using a two by two
inch slide projector with the mirror of the camera lucida reversed,
so that they appeared projected downward on drawing paper on the
surface of the table and could be accurately traced, after which the
light maculations of holotype male were drawn in, free hand, on this
drawing in approximate position.

The genitalia were removed from the holotype male, the allotype
female, 10 male and 4 additional female paratypes. After being carried
through the solutions as used with the labial palpus, they were mounted
on slides in balsam. Well slides were used for the male genitalia.
Later, camera lucida drawings were made. Details were studied directly
under the high power of the compound microscope and added to
the camera lucida drawings.

A New Megathymus. C. F. Harbison

241

Megathymus comstocki, new species
Plate 21, figures 1 to 8.

Type Specimens
The series examined consisted of forty-two specimens: holotype
male, allotype female and forty additional paratypes, of which twenty-
three were males and seventeen were females.
Table 1. Data on Type Series of Megathymus comstocki
All from Baja California, Mexico; collected, except as indicated,
in 1955. All specimens other than the holotype are paratypes.

Locality

2 miles NE.
of San Simon

Collector

McKenney

Date
Sept. 10

Type Designation
and specimen No.

Holotype

Males
1

Females

Do.

Comstock

Sept. 9

1 Allotype (9 1)

—

1

Do.

Harbison

Sept. 9

9 9 2-5

—

4

Do.

McKenney

Sept. 9

5 5 14, 17, 19 - 22,
9 11.

6

1

Do.

Comstock

Sept. 9

1 98

—

1

Do.

Comstock

Sept. 9

9 15 (malformed)

—

1

Do.

Comstock

Sept. 9

99

—

1

Do.

Williams

Sept. 9

c?23;
9 18

1

1

San Simon

Harbison

Sept. 8

5 5 2, 3

2

— :

Do.

McKenney

Sept. 8

<? $ 13, 15, 16, 18
9 9 10, 12, 13, 16, 17

4

5

Mesa S. of
San Simon

Harbison

Aug. 31,
1953

5 5 1, 6

2

Do.

Farmer

Do.

5 5 4, 5, 7- 12;

9 9 6, 7

8

2

Camalu

McKenney

Sept. 6

9 14

—

1

Holotype male* and allotype female — are in the type collection of the
San Diego Society of Natural History. Paratypes (see table 1) have been
deposited thus: American Museum of Natural History, 5 22, 9 3; Cali-
fornia Academy of Sciences, 5 23*, 9 18*; Los Angeles County Museum,
5 21, 9 17; National University of Mexico, Instituto de Biologia, 5 20, 9 5;
Dr. J. A. Comstock, 5 18, 9 8, 9 9*; Dr. H. A. Freeman, 5 19, 9 16;
Mr. M. J. McKenney, 5 13, 5 14, 5 15, 5 16, 5 17, 9 10, 9 11, 9 12,
9 13, 9 14. Paratypes 5 1*, 5 2*, $3, 5 4, 5 5*, 5 6*. 5 7*, $8*,
5 9*, 5 10*, 5 11*, 5 12, 9 2, 9 4, 9 6*, 9 7*, 9 15* have been re-
tained by the San Diego Society of Natural History.

1 Pupa collected in host plant; emerged Sept. 12, 1955
Pupa collected in host plant; emerged Sept. 27, 1955
*Microscope slide made of genitalia; should be kept with mounted specimen.

242 San Diego Society of Natural History

HOLOTYPE MALE

The Head and its Appendages
The head, including the compound eyes, is as wide as the mesono-
tum between the attachments of the forewings. The eyes are large:
their length as viewed from above is about equal to the shortest dis-
tance between their bases across the vertex of the head just above the
attachment of the antenna. The eye is beveled to the attachment
to the head and lacks ommatidia in this smooth, shining black,
scaleless, ringlike area. Some of the scales of the head extend
laterally over this area so that, in life, the beveled area is not easily
seen. The seeing surface of the eye is evenly rounded and is com-
posed of numerous blackish ommatidia. The bases of the antennae
are far apart; in fact, as viewed from above, the outer side of the
basal segment rests against the beveled edge of the compound eye.
The antenna has 35 segments. The basal segment bears a comblike
structure of 4 stiff black bristles that extend out laterally over the
compound eye. It is about as long as wide. Succeeding segments
lengthen gradually until they become about twice as long as wide
just before the club. The club begins with joint 21, reaches its
widest part, as viewed from above, at joint 26 and then tapers quite
abruptly to its end. The segments near the head have a narrow
basal ring of white scales that cover about one-sixth the length of
the segment when viewed from above. This ring becomes much
wider beneath, covering from one-half to five-sixths of the length of
the segment. The part of each segment that is not covered with white
scales is clothed with dark brown scales. The white scales are more
extensive in the club. The central, lower surface of the club is covered
almost completely with white scales, but at the tip there is a heavy
admixture of brown. The last segment is completely covered with
these brown scales.

Attached to the underside of the head is the well-developed
light brown, and heavily chitinized proboscis which, if extended, would
reach the base of the abdomen. It is tightly coiled like a watch spring
when the insect is not feeding. Striae transversely encircling the
proboscis almost suggest a fine annular structure. On each side of
the proboscis and fastened to the lower side of the head capsule is
the three-jointed labial palpus. The first joint of this structure is
small, and fastens the appendage to the head capsule. It extends
downward and forward from the point of attachment, and then bends
upward. The second joint is about two and one-half times the

A New Megathymus. C. F. Harbison 243

length of the first segment and stands erect above it on either side of
the coiled proboscis. The inner surface of this appendage is flat-
tened so as to fit snugly in place against the proboscis. The outer
surface is evenly curved and rests against the beveled area of the
compound eye. Scales on the inner surface of the second joint of
the palpus are flattened and white, and each usually has three or
four sharp teeth at the distal edge. They are appressed to the surface
of the joint and their distal free ends project upward. The segment
viewed from the front is clothed with toothed scales, in an admixture
of white and brown. The side of the palpus that rests against the
compound eye is covered with flattened, toothed white scales and
long dark brown hairlike scales. The very small third segment is
about one-seventh as long as the second segment and is clothed
densely with more or less erect scales. The vertex of the head also
is covered with an admixture of vertical scales.

The Thorax and its Appendages
The pronotum is covered with brownish scales that tend to
have their free edges rearward. The mesonotum bears a pair of
platelike structures, (the tegulae), between the forewings at their
attachment. These are densely covered with large scales, which have
their distal free end caudad. Many of these scales are very long and
hairlike. They are light brown or yellowish white. The under surface
of the thoracic segments is clothed with an admixture of linear and
hairlike white scales that tend to have their free ends caudad.

Wings of Holotype
Upper Surface
Primaries. — The forewings are mainly covered with dark brown
appressed overlapping scales. The free end of each scale lies toward
the outer margin of the wing. The scales cover the entire surface
of the wings. They densely clothe the very heavily chitinized veins
as well as the membranous surface between. The exposed length of
surface of each scale is usually about twice the width. There is
considerable variation, especially toward edge of wing and just inside
the fringing scales, where usually the exposed length is about equal
to the width. Near the base of the wing between the fork of vein Q12
and A2 and near vein A3 are numerous long hairlike yellow scales
similar to those covering adjacent parts of the thorax and abdomen.
The light maculations on the primary are composed of yellowish
white scales similar in shape to the brown ones covering the main
surface of the wing.

244 San Diego Society of Natural History

A small cell spot occurs just caudad to the fork of R2. Of the
four subapical spots, the first is located near the margin of the wing
between R2 and R3, the second lies diagonally behind this, between
R3 and R4; the third directly behind the second, between R4 and
Rs, some distance internal to the apex of the wing; and the fourth and
largest is diagonally in line with subapical spots 1 and 2; it lies between
veins R5 and Mi. The rather small and indistinct extra-discal spots
lie respectively between Mi and M2 and between M2 and Ma. They
are parallel to the outer margin of the wing and are nearer to it than
are the spots of the discal band. The discal band is made up of four dis-
tinct spots. The first lies between Ms and Cui and is the largest. It is
rounded and occupies about one-half of the distance between veins.
The second discal spot lies caudad of the first, between Cui and Cu2.
Two very small spots lie between Q12 and A2. These actually seem
to represent one spot divided by the fold that represents vein Ai. The
fringes are alternately dark brown and white. The scales making up
the fringe are about four times as long as wide, with the outer edges
toothed.

Secondaries. — The entire upper surface is almost completely
covered with small dark brown scales. In addition, the bases of the
wings, out as far as the maculations, are covered with an admixture of
dark brown and yellowish hairlike scales. The maculations are formed
of small yellowish white flattened scales similar in shape to the small
brown scales. Between Sc and R and nearer to the base of the wing than
any other spot is a large indistinct maculation. Diagonally caudad
of this spot, between R and Mi, is a second indistinct light area. A
discal band of five spots occurs some distance internal to the margin
of the wing. The last of these spots, between Q12 and A2 is actually
a double spot. None of the spots occupies the entire space between
veins. The fringe along the edge of wing is made up of white and
dark scales. The dark areas are located where the principal veins
reach the edge of the wing.

Under Surface

Primaries. — Most of the under surface of the wing is covered
with small dark brown appressed scales, having their free tips toward
the outer margin of the wing. The light maculations show more
plainly on the lower side than on the upper. The cell spot is obsolete.
The four subapical spots are very distinct and occupy the same
position that they do on the upper surface. This is also true of the
two extra-discal spots and the three spots that make up the discal

A New Megathymus. C. F. Harbison 245

band. The last spot of this band is distinctly divided through the
middle. The tip of the wing has an over-scaling, of toothed white
scales, that produces a frosted appearance. The fringe along the
outer margin of the wing is composed of two layers of scales.

Secondaries. — The under surface shows the same discal band of
five yellowish white spots extending from vein Mi to vein A2. Internal
to the discal band are faint light patches, one between R and Mi, and
a second between Sc and R. Another light patch occurs between
Cu2 and A2, caudad of the fork of the Cu2 vein. The lower surface
is overlaid with toothed white scales and hairlike scales. This gives
the under surface a frosted appearance. The double-row nature of
the black and white fringe shows plainly along the margin of the wing.

Expanse of wings of holotype male, 40.2 mm.

Legs of Holotype

The femur of the foreleg is long and laterally flattened; white
and gray overlapping scales densely cover the top edge and also the
two sides and are markedly appressed to the surface. A number
of hairlike scales occur along the outer lower edge of the femur.
Jointed to the distal part of the femur and, at rest, bent backward,
jack-knife fashion, is the shorter tibia. The tarsus is five jointed.

The femur of the second leg is longer than that of the foreleg
and about equal to the length of the next segment, the tibia. This
segment is laterally flattened and is covered with hairlike scales on
the inner surface next to the body. This joint, at rest, extends caudally.
The tibia is clothed with gray and white scales. There is a five
jointed tarsus.

The very short, laterally flattened femur of the third leg is
clothed on the inner side, next the body, with dark hairlike appressed
scales that extend downward beyond the lower edge of the segment,
forming a fringe wider then the femur.

On its outer side are flattened whitish scales. The tibia is
much longer than the femur, almost as long as the femur of leg two.
The tibia is clothed with flattened and hairlike scales, extending
downward over the basal segment of the tarsus. The inner side
bears a dense covering of flattened scales.

Tarsi
The five-segmented tarsus is covered with flat imbricated scales.
The final row of scales on the last tarsal segment extends over the
claws and actually hides from view the truncate pulvillus. Stiff spines

246 San Diego Society of Natural History

arm the inner and lower side of the tarsus. Scattered spines on the
upper side are visible only when the imbricated scales are removed.

The Abdomen and its Appendages
The basal segments are complete rings. Segments VIII, IX
and X are modified into the male genitalia and are covered with
flattened, toothed, light brown scales overlaid by long yellowish
hairlike scales. The dense covering of scales and their extension rear-
ward makes it difficult to see the male genitalia until the scales are
removed. Segment IX is telescoped forward into the ringlike seg-
ment VIII, and is composed of the heavily chitinized tegumen, the
narrow ringlike vinculum, and the bladelike, forward-pointing saccus.
The notum of segment X is the uncus and forms two downward-bent
hooks. Viewed from beneath, it is very broad and heavily chitinized
at the tip. Its tip is nearly half the breadth between the wide part
of the uncus, about two-thirds of the way rostrad to its base. This
breadth is about one-third of the length of the segment. It is slightly
bifid. The sternite of this segment X constitutes the upward bent
hooks, the gnathos. These are rounded basally and bent up with an
even curve on either side of the anal opening. Fastened to the
vinculum are two clasping organs, which are thought to represent
abdominal legs. These valvae, harpes, or claspers (depending on the
authority followed) are slightly more than three times longer than
broad. They have two basic parts, the lobe above and the blade
below. The blade extends into a sharp up-pointed end. This is the
cucullus of the valva. Various teeth occur along the edge, from
the pointed tip rostrad. The upper edge of the lobe is the costa of
the valva, and the lower edge of the blade, forming a heavily chitinized
brace-like thickening, is the sacculus. A large process, the proharpe
(using the Barnes and McDunnough term) is rostrad from the
tip of the blade at a point where the outer part of the lobe touches
the blade. This structure is heavily spined and has a basal fork that
is very distinctive in the new species. The aedeagus is held in place
in the cavity between the valvae by an annulus, a transverse plate with
encircling upward-pointing lateral arms. The aedeagus is a chitinized
tube-like structure with a flaring tip. This, the vesica, has, on each
side, ten cornuti.

ALLOTYPE FEMALE

The Head and its Appendages
The head, including the compound eyes, is about two-thirds the
width of the mesonotum between the wing bases. The compound

A New Megathymus. C. F. Harbison 247

eyes are smaller than in the male. Their length, when they are viewed
from above, is about one-third less than the distance between their
bases, across the vertex of the head above the attachment of the
antennae. The flattened linear scales on the front below the antennae
stand more or less erect. The toothed brownish scales of the top and
back of head tend to He forward thickly over the base of the antennae,
so that it is impossible to see the basal joint without their removal.
The black spinelike scales forming the so-called eyebrow are well
developed in the female. The antenna is ringed narrowly with white
at base of each joint, becoming wider on inferior surface. The under
side of club is white; the upper side, brown. The labial palpus is
clothed densely with white, sharply-toothed, upright scales. The
proboscis seems to be similar to that of the male.

The Thorax and its Appendages
The pronotum is covered with a dense thatch of flat, toothed
scales, each of which tends to have its free end appressed to the surface
of the segment. The mesothorax bears two large platelike structures,
the tegulae. These are covered with light brown acicular scales. Mixed
with these are a few darker, flat scales. The number and length of the
hairlike scales increase on the metanotum.

WINGS OF ALLOTYPE

Upper Surface
Primaries. — Most of the dark brown scales covering the wing are
flat and have striae running their length. Scales of this type cover most
of both surfaces of the wing, and are basic. Hairlike scales of light
golden color cover the base of the forewing. The maculations are
similar to those of the male but are much more extensive and of a
richer, brighter color. The four-sided cell spot is caudad from the
costal edge of the wing a distance equal its width. Its front edge
is straight and parallel to this margin. The side toward the apex
of the wing is concave to a point. The rear margin of this spot bows
downward. The fourth margin is concave to its juncture with the
straight front edge. The first spot of the subapical band is very in-
distinct. The second, third, and fourth are about equal in size and
are arranged diagonally across the wing tip. The extra-discal spots of
the female are about four times the size of those of the holotype male
and are parallel to the outer margin of the wing. The first spot in the
discal band between M3 and Cui is nearly rectangular and in width
covers most of the interspace between these veins. It is twice as long
as wide and of the two ends the outer is convex, the inner concave.

248 San Diego Society of Natural History

Spot two of the discal band is five-sided. The straight front edge
of this spot is of about the same length as the rear edge of spot one.
The outer edge of spot two becomes slightly diagonal outward. The
rear side is nearly straight. The fourth side runs diagonally toward
the body of the insect to contact with the fifth side. The fifth side
is slightly concave. This spot is about twice as long as broad. Spot
three of the discal band lies between Cu? and A2. It is very irregularly
four-sided. The front edge of this spot on the right forewing is
nearly straight; the outer side tends to be undulate; the rear side is
straight; the inner or fourth side is again undulate. In the left forewing
as a variation, the undulations of the two sides are carried to a
point, where the spot forms a very thick letter Z reversed. The spot
is twice as long as wide, crossing the entire interspace between the
veins. The fringe along the outer edge of the wing is made up of
dark and light, flattened, toothed scales. Each scale is about four
times as long as wide.

Secondaries. — Dark brown scales are basic as in the primaries.
There is a much more extensive area of long golden hairlike scales, ex-
tending almost to the row of discal band spots. Between R and Mi is an
orange-yellow spot, indistinct because overlaid by long brown hairlike
scales. Diagonally behind this, between Mi and M2, is the first
spot of the discal band. This is of about the same size as in the
holotype male. There appear to be six of these discal spots but I
believe that the fifth spot is divided in two parts. On the underside
of the wing these two spots tend to be fused. The black and white
fringe along the edge of the wing is a very striking feature.

Under Surface

The primaries have the same maculations as on the upper surface.
The cell spot tends to be double, comprising a very small spot,
followed by a large light maculation. The apex of wing bears an
overlay of white scales. The secondaries beneath have an extensive
overlay of white scales that more or less hide the maculations. Spots
equivalent to those of the upper surface can be found if carefully
sought but they average smaller. Discal spot five appears as one large
spot, thus differing from its equivalent on the upper surface, where
it appears to be double. The fringe along the outer edge of the wing
is alternately white and dark.

Expanse of wings of allotype female, 53.3 mm.

Legs of Allotype
The legs are similar to those of the holotype male.

A New Megathymus. C. F. Harbison 249

The Abdomen and its Appendages
The abdomen of the allotype female is much longer than that
of the holotype male and is distended with eggs. It is covered
densely with overlapping scales. The female genitalia* comprise
vaginal and postvaginal plates and the ovipositor. The ostium or orifice
of the bursa copulatrix is on the caudal (posterior) edge of the
vaginal place. Mating with the male is through this opening. The
vaginal plate, measured medially from its posterior edge above the
ostium to its anterior margin between the rounded inner end of the
lateral fold, about equals two-thirds the distance between the posterior
points of that structure. This lateral fold forms a nearly straight
inner edge. Its outer edge is decidedly bowed outwardly. The
posterior end is about in line with the anterior margin of the ostium
or opening of the bursa, and diverges outwardly. The anterior or
rostrad end of the lateral fold is rounded and converges medially.
The distance between the rounded anterior ends of the lateral fold
is about one-half of the distance between the pointed posterior ends.

A convex lens-shaped sclerite rests medially with its inner side
against the pointed end of the lateral fold. The outer point of the
sclerite is diagonally outward. The inner point is directed inward
and reaches the level of the posterior margin of the medially placed
orifice. Directly posterior to this are the structures called by Barnes
and McDunnough the alae.

From the posterior margin of the orifice a ribbon-like band
curves diagonally forward and outward until it reaches the level of
the anterior edge of the orifice. This brings it about half way to
the pointed end of the lateral fold. Here it apparently is turned up
and over. It extends rearward to the inner point of the convex lens-
shaped sclerite. Because of the apparent twisting of this band in the
middle, an isosceles-triangle-shaped dark spot appears at the point of
the twist, with its apex pointing rearward. The band again curves over
on itself and ends in a flaring, .only weakly chitinized funnel-shaped
structure.

The anterior margin of the plate between the rounded ends of
the lateral fold is sinuous. The medial part of this edge is bowed
forward. On each side of this, and connected to the broad end of
the lateral fold, the edge is bowed rearward. The outer curved edge
of the lateral fold is margined with long hairs that extend outward

*The author accepts the terms for the female genitalia used by Barnes and
McDunnough in "Revision of the Megathymidae," 1912, rather than those suggested
by A. Diakinoff in "Lepidopterist News," 1954.

250 San Diego Society of Natural History

at right angles to the edge a distance of about half the width of the
fold. The general surface of the plate between the folds is covered
with many short bristle-like spines. These are placed in definite
rows and form a beautiful pattern.

The oviporus is on the postvaginal plate. Through this opening
the eggs are extruded. The postvaginal plate has a heavily chitinized
fold along its anterior edge. This fold is bluntly crescent-shaped
and is covered with short bristles. Behind this is a wing-shaped
structure, which is emarginate medially on the posterior edge. It then
bows outward and gradually tapers off to a point laterally. The
oviporus is in the emarginate middle posterior part of the plate,
which is covered with various types of spines. On either side of
this area the covering spines become coarser and more numerous.
Beginning with this area, medial to the tapered outer part of the
plate, the spines become very heavy and several may originate from
a single spot on the surface. The spines lie with their unattached
tips pointing caudally.

The lobes of the ovipositor at the end of the abdomen are re-
tracted into a cavity by the telescoping of several segments. They
are ear-shaped and are covered with long bristle-like spines on their
outer surfaces. In the allotype slide they are flattened out. Usually,
on the slide they stand in vertical position, suggesting the ears of an
alert rabbit.

STUDY OF THE PARATYPE SERIES

Appendages of the Head

Antenna
Certain structures were studied on paratypes that could not be
examined on the holotype without complete dismemberment. An
antenna was removed from a paratype, descaled and then run through
sodium hydroxide, water, alcohol, and xylene. This expanded the
segments. The antenna is composed of 35 segments, 20 in the stalk
and 15 in the club. The basal 7 segments of the club are each
progressively larger. The remaining 8 segments are each progressively
smaller. (Plate 19, fig. IB.)

Labial Palpus
The labial palpus of a paratype is shown in several aspects on
Plate 19, fig. 2B, 2C, D. Note that the basal segment is as broad as
the second and that the ultimate segment is very small.

The Thorax and its Appendages
The head of the paratype was removed and the prothorax

A New Megathymus. C. F. Harbison 251

examined. On the upper part of the pronotum, covered with upright
scales, are two reniform structures, the patagia, which are rather
thin vertical extensions of the integument. They are readily removed
and descaled (Plate 19. fig. 3). Another structure of interest is the
platelike outgrowth of the mesonotum at the base of each primary.
This structure, the tegula, has a long, rounded, bladelike extension
caudad and a down-curved fore part that forms a shoulder cover.
This plate is clothed heavily with long, grayish-brown flattened scales
and numerous long hairlike scales. Denuded of scales it is illustrated
on Plate 19, fig. 4. At present these structures are not considered as
diagnostic in a study of the species of this genus, but on comparison,
long series of slides of each species might show some significant
differences.

The Wings

The descaled wings of a paratype did not show any distinctive
venation. The maculations of the male and female wings vary somewhat
in the series of specimens before the author. This is especially true
of the cell spot of the primaries. In some specimens this is large
and extensive; in others it is indistinct. The two extra-discal spots
vary in size and shape. The discal band may contain four distinctly
separate spots, or the last two (caudad) spots may be fused to
form one. Only three paratypes in the series of twenty-three males
show a distinct spot between A2 and Cu2, about halfway from the base
of the wing to its outer margin. If it were not for these exceptions and
for the fact that about three-fourths of the females examined show
the spot, this would be a good character for separating this from
related species. The secondaries have a discal band of five spots.
The last spot tends to be double in some of the paratypes.

In the wing maculations of the female paratypes all spots are
more extensive than in the males, but vary decidedly as to size and
shape. The cell spot is seen in all eighteen female paratypes. The
greatest difference is in the size and shape of the spots composing the
discal band of the primaries. Each spot reaches almost completely
across the interspace between the veins. In the allotype female
(paratype ? l) spots 1 and 2 of the discal band are much broader
than long. Spot 3 tends to be shaped like the letter Z. In some of
the females, spots 1 and 2 become almost square. Spot 3 varies to
almost the double condition found in the holotype male. The spots
in the discal band on the secondaries vary from six to five. In twelve

252 San Diego Society of Natural History

of the eighteen paratype females there is a spot between A2 and Q12
about half way from base to outer margin.

Tarsus

The tarsus (Plate 19, fig. 6A) is quite distinctive. The pulvillus
is truncate and slightly emarginate. It is dark at the tip. The
paronychium is a broad ribbon-like structure with ciliated edges
two-thirds the length of the claws. The claws are only slightly curved
at the tip.

Genitalia
(Plate 20)

To determine the variation in the paratypes, slides were prepared
of the genitalia of nine males and four additional females. The uncus
of each paratype was studied from below before it was mounted
permanently on a slide. The genitalia of paratypes $ 7 and $ 1 1 were
placed so that the uncus and tegumen would remain with ventral
surface exposed. The other seven slides were assembled with the
uncus showing the lateral aspect. In the holotype male, after the
underside of the uncus had been studied and drawn, this part was
laid on its left side.

Little variation was found. The valvae vary slightly in shape.
In some paratypes the cucullus is drawn out into an upcurving point,
in others it is more blunt. The proharpe tends to show a basal arm
or branch on its rostrad side where it is attached to the blade. Spines
similar to those on the main proharpe are found on the basal arm
and also point with free end upward. The branching of proharpe is
characteristic and uniform in this species.

The genitalia of the females are less uniform than those of the
males. The author made a slide of the genitalia of the allotype female,
a specimen collected as a pupa and reared. Slides were also made of
three other female paratypes, collected in the field as adults. These
had probably already mated. The allotype had had no opportunity
to mate as no males were reared; therefore the author suspected
that the observed differences might be attributed to her virginity.
A second slide of a reared, unmated female was made late in the
study. This latter slide did not agree completely either with the
allotype or with any of the other paratypes.

COMPARISON OF THE NEW SPECIES WITH
RELATED SPECIES

The new species of megathymid was compared with other members
of the agave-feeding section of the family. When viewed from below,

A New Megathymus. C. F. Harbison 253

the male unci of M. stephensi, M. mariae, M. polingi, and M. comstocki
are equally broad at tip, about one-fourth broader than in M.
neumoegeni, as indicated in the illustration in the Barnes and Mc-
Dunnough monograph. M. stephensi apparently should not be con-
sidered a subspecies of M. neumoegeni.

Megathymus stephensi Skinner
One might expect that the nearest relative of the new Megathymus
would be M. stephensi, another agave feeder found in the desert
region of San Diego County just north of Baja California, but the
maculations are quite different and the male and female genitalia
do not indicate a close relationship. Also the host plants of the
immature insects are very distinct.

In the male of M. stephensi spot 3 of the discal band is not
double in the forewing and the spot between A2 and Cu2, midway
between the base of the wing and outer wing margin, is very distinct.
On the secondaries, the discal band is much more irregularly placed
than in the new species. The author checked the number of spots in
the series of the San Diego Natural History Museum's collection
and found that there are five in M. stephensi. In the original
description, Skinner stated that "on underside secondaries . . . there
is a large coalesced spot near the center of the wing and eight spots
parallel to the margin." He evidently started with the spot inward
from the costal margin and near the base of the wing and counted
three spots before reaching the upper spot of the discal band. There
are five spots in the discal band; adding these three, gives a total
of eight. On the underside of the secondaries of M. stephensi there
is a spot internal to the last discal, about halfway to the base of the
wing. Forward of this, near the center of the wing, is an additional
spot. Thus M. stephensi may show ten light maculations on the
underside of the secondaries.

The new species is smaller in wing expanse; the largest male
measures 46.5 mm; the smallest male of M. stephensi in the collection
of the Museum, 50.0 mm.

Slides were made of the genitalia of two males and four females
of M. stephensi taken near the type locality and these were compared
with the slides of M. comstocki. The cucullus of the blade of the
valva in these is more abruptly turned up and the space along the
upper edge to the base of the proharpe is more irregularly spined.
The proharpe of M. stephensi is not broad at the base as it is in the
new species nor is it narrow at the base as in M. neumoegeni, but is

254 San Diego Society of Natural History

intermediate in width. There is no cephalic branch or even a tendency
to branch in M. stephensi.

In the female genitalia of M. stephensi a heavily chitinized
triangular extension runs diagonally from a plate located beside the
ostium. The point of this triangle is extended to meet the inner side
of the lateral fold slightly posterior to its middle. In the new species,
the allotype slide shows the triangular area to be very small and it
does not point forward toward the lateral fold; its apex is to the rear.
The lateral fold of the Stephens' megathymid is sometimes extended
around the edge of the plate so that the anterior point is directed
rearward, a condition not observed in the new species.

Megathymus mariae Barnes and Benjamin

In respect to the light spots, M. mariae is intermediate between
M. polingi and the new species. M. mariae has discal spot 3 wider
than spots 2 and 1. In this character it differs from both M. polingi
and M. comstocki, in which spot 3 is narrower than the two spots
above. There is no tendency for this spot to become paired. In the
new species, spot 3 is small and often divided by the very faint Ai vein
into two spots in the male. In the female this spot tends to be
Z-shaped. The discal band of the secondaries in M. mariae is made
up of five definite spots. In the new species there tend to be six.

The male genitalia of M. mariae are much more like those of
M. stephensi than those of the new species. The cucullus of the blade
of the valva is truncate and the space along upper edge to the base
of the proharpe is even more irregularly spined than M. stephensi.
The teeth are large and heavily chitinized. The upturned end of the
cucullus is extremely heavily spined in M. mariae. The proharpe is
also more heavily spined than in M. stephensi or M. polingi. Its
basal breadth is about the same as in M. stephensi, much narrower
than in the new species. It shows little tendency to become branched
on its inner side as in M. comstocki-

In the female genitalia of M. mariae a heavily chitinized extension,
at the level of the ostium, curves outward and then forward toward
but not to the lateral fold, which is long and narrow. At its rostrad
end, where the two pointed ends approach each other, the fold is bent
so that these points actually are directed backward. This condition
is quite different from that of M. comstocki, in which this part of
the band is nearly straight and truncated. The margin of the plate
between the curved ends of the lateral fold is deeply concave and

A New Megathymus. C. F. Harbison 255

the edge is heavily chitinized. This margin in the new species is
convex in the middle and concave on the two sides.

Megathymus polingi Skinner

On the basis of the study of the genitalia the author believes that
the new species is more closely related to M. polingi Skinner than
to the previously mentioned species.

However, the maculations of male of M. polingi are much
larger than those of the male of M. comstocki. In M. polingi spot 3
of the discal band is large and single, completely crossing the interspace
between A2 and Cu2. This spot is slightly narrower than spot 2.
The spot in the interspace between A2 and Cu2, about half-way out
from base of wing to the margin, is very distinct. This is often
lacking in M. comstocki. The five definite spots in the discal band
of the secondaries in M. polingi are larger than those of M. comstocki.

In the female of M. polingi the spots on the forewings are very
large and entirely cross the interspaces between veins. This makes
the wings truly banded. On the primaries of M. polingi the cell
spot is broadly connected with spot 1 of the discal band and the two
extra-discal spots form a band that connects broadly with spot 1 of
the discal band caudadly and narrowly with subapical spots cephaladly.
The subapical spots form a band to the costal margin of the wing.
Thus the forewings appear to be crossed by a wide band of light color
that is forked in its upper part, with the right fork bent inward to
the costal margin of the wing when the right forewing was examined.
The band between A2 and Cu2 is narrower than it is between Cu2
and Cui and between Cui and Ms. In M. comstocki females the
light scales are in separate spots rather than bands. None of the
light scales are on the veins. Discal spot 3, between A2 and Cu2,
is the narrowest of the three. Spot 1 does not come close to the
cell spot.

The male genitalia of M. polingi is most like those of the new
species. The cucullus of the blade of the valva is evenly curved upward
and similarly spined. The spines along this edge are more regular and
smaller in M. comstocki than in M. polingi. The base of the proharpe
is broad and there is a tendency toward the development of a second
branch at the inner base in M. polingi. There is an even greater
tendency for the development of this branch in the new species.

The female genitalia of M. polingi have a heavily chitinized
triangular extension from the plate beside the ostium. Its point is
directed toward the upper part of the lateral fold but does not reach

256 San Diego Society of Natural History

its inner margin. The slide of the female M. polingi prepared from
material loaned to the author by the Los Angeles County Museum
shows this condition more plainly than does the illustration in the
Barnes and McDunnough monograph. In the allotype of the new
species the dark triangle does not point forward. The lateral fold in
M. polingi is long and ends in a point at the posterior end. The
cephalad end tends to be truncate. The fore edge of the vaginal plate
on the margin where the lateral folds approach each other is strongly
concave. In the new species the fold is similar but the edge of the plate
between the truncated ends of the fold is convex in the median line,
with an abrupt concavity on either side.

SUMMARY

The main purpose of this paper is to name a megathymid, be-
lieved to be new to science, Megathymus comstocki, captured first
on the mesa south of the broad San Simon (Santa Maria) River
valley, Baja California, Mexico, on August 31, 1953. Several return
trips were made to the vicinity to collect a larger and more perfect
series of this insect during late 1953, mid- August of 1954 and finally
in early September, 1955. A series of 42 specimens was ultimately
obtained. The holotype male and allotype female were studied in
much detail and diagnostic structures illustrated, both photographically
and by drawings. The paratypes of the new species were studied to
determine variation. Other megathymids of the agave-feeding group
were compared with this new species. Among the known species of the
genus, Megathymus polingi Skinner appears to be the most nearly
related.

A distinctive feature of the new species is the shape of spot 3
of the discal band in forewings, which tends to be very small and
double in the male. The proharpe of this sex tends to have a basal
arm, and is broad at base. In the female, spot 3 of the discal band
of forewings is shaped like the letter Z. The structures on the two
sides of the ostium in the allotype female are distinctive in that the
triangular, heavily chitinized part has its apex pointing rearward.

BIBLIOGRAPHY

Barnes, William and F. H. Benjamin

1924. Notes and New Species. Contributions to the Natural
History of the Lepidoptera of North America. Decatur,
Illinois. The Reveiw Press. Vol. 5, no. 3, pp. 100-101.

A New Megathymus. C. F. Harbison 257

Barnes, William and James H. McDunnough

1912. Revision of the Megathymus. Contributions to the Natural
History of the Lepidoptera of North America. Decatur,
Illinois. The Review Press. Vol. 1, no. 3, 56 pp., 1 fig.,
6 pis.

Bell, Ernest L.

1938. A New Genus and Five New Species of Neotropical
Hesperidae. American Museum Novitates, no. 1013, pp. 1
to 11, figs. 1 to 6.

COCKERELL, T. D. A.

1941. Observations on Plants and Insects in Northwestern Baja
California, Mexico, with Descriptions of New Bees. Tran-
sactions, San Diego Society of Natural History, Vol. 9,
pp. 337 to 352.

Freeman, Hugh A.

1951. Notes on the Agave Feeders of the Genus Megathymus.
Field and Laboratory, Southern Methodist University, Vol.
19, pp. 25 to 32.

1951. Ecological and Systematic Study of the Hesperioidea of
Texas. Studies, no. 6, Southern Methodist University, pp.
1 to 68.

1952. Two new species of Megathymus from Texas and Mexico.
American Museum Novitates, no. 1593, pp. 1 to 9, figs.
1 to 13.

Hoffmann, Carlo C.

1941. Catalogo Sistematico y Zoogeografico de los Lepidopteros
Mexicanos. Segunda Parte — Hesperioidea. Anales Insti-
tute de Biologia, Mexico, Vol. 12, pp. 237 to 294.

Rindge, Frederick H.

1948. Contributions Towards Knowledge of the Insect Fauna of
Lower California. Rhopalocera of Lower California. Pro-
ceedings of California Academy of Sciences, Vol. 24, no.
8, pp. 289 to 312.

Skinner, Henry

1905. A New Megathymus from Arizona. Entomological News,

Vol. 16, pp. 232.
1912. Two New Butterflies. Entomological News, Vol. 23,

p. 126.

258 San Diego Society of Natural History

Explanation of Figures on
Plates 19 and 20

Plate 19

Figures 1 to 6. Appendages of head and thorax.

1 A. Antenna of Holotype, upper and lower surface, scales in place.

1 B. Antenna of Paratype denuded and distended.

2 A. Labial Palpus of Holotype denuded and distended, first joint

separated.

2 B. C. D. Labial Palpus of Paratype denuded but dry, front, inner
and outer surface.

3. Patagium of Paratype romoved from pro thorax and denuded of
scales.

4. Tegula of Paratype removed from mesothorax and denuded of
scales.

5. Wings of Paratype descaled and projected. Maculations of Holo-
type placed in relative position.

6 A. Claw segment of a paratype distended and denuded.

6 B. C. Legs of Holotype denuded dry, outer and inner surface
Venation of wings, both Comstock-Needham (abbreviations) and
British method (small numbers) used.

A. — Anal Vein; Cu. — Cubitus; Epi. — Epiphysis; M. — Media; Par. —
paronychia; Pul. — pulvillus; R. — Radius; 8. — Subcosta; Seg. 5 — Segment
5 of tarsus with attached claws and other parts; Sp. — Spur on Tibia.

Plate 20

Figures 1 to 3. Appendages of the abdomen. The genitalia.

1 A, B, C. Genitalia of Holotype male, lateral view from left side
showing inner surface of right valva. Roman numbers refer to
abdominal segment.

1 D. Annulus, front aspect.

2 Uncus and associated parts viewed from below.

3 Genitalia of Allotype female. Vaginal plate, post vaginal plate
and ovipositor.

A — annellus; Ae — aedaegus; B — blade of valva; C — costa of valva; Cu —
cucullus of valva; G — gnathos; L — lateral fold; Lo — lobe of valva;
Opr — lobe of ovipositor; Pvp — Post vaginal plate; P — proharpe; S —
saccus; Si — sacculus of valva; T — tegumen; U — uncus; Vp> — vaginal
plate; Vo — vaginal oriface; Ve — vesica; Vi — vinculum.

A New Megathymus. C. F. Harbison

259

Plate 19

For explanation of figures see page 258.

260

San Diego Society of Natural History

Plate 20

For explanation of figures see page 258.

A New Megathymus. C. F. Harbison

261

<>"' \

Plate 21

Figures 1 to 8. Adults, upper and under surfaces..

1, 2. Megathymus comstocki Harbison, holotype male.
3, 4. Megathymus comstocki Harbison, paratype male.
5, 6. Megathymus comstocki Harbison, allotype female.
7, 8. Megathymus comstocki Harbison, paratype female.

TRANSACTIONS

inwui u,r • fcUUk

MAY 2 3 1957

HARVARD

UNIVERSITY

OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY

Volume XII, No. 13, pp. 263 - 276

Plate 22

NOTES ON THE METAMORPHOSIS
OF AN AGAVE-BORING BUTTERFLY
FROM BAJA CALIFORNIA, MEXICO

BY

John Adams Comstock

Fellow of the San Diego Society
of Natural History

SAN DIEGO, CALIFORNIA

Printed for the Society

May 1, 1957

MUS. COMP. ZOOL
LIBRARY

MAY 2 31957

HARVARD
UNIVERSITY

Table of Contents

page
Introduction 267

Acknowledgments 267

Larval Habits of Megathymus comstocki 268

The Ovum, and Oviposition 269

Collecting Terrain and Technique 269

Larva of Megathymus corns toch 271

Pupa of Megathymus corns toch 272

Comparisons 272

Parasites 273

References 274

Plate 22: Larva and pupa of Megathymus comstock' 275

NOTES ON THE METAMORPHOSIS
OF AN AGAVE-BORING BUTTERFLY
FROM BAJA CALIFORNIA, MEXICO

BY

John Adams Comstock

INTRODUCTION

The recent publication by Charles F. Harbison (1957) of a new
species of agave-boring skipper, Megathymus comstock}, has established
the first authentic record of an indigenous species of this interesting
genus from Baja California.

The writer had the pleasure of participating in a field study of the
species with a group of naturalists, working in the area two miles
northeast of San Simon, Baja California, under the aegis of the San
Diego Society of Natural History, September 9 and 10, 1955.

A record of that trip, with a description of the physical and ecological
features of the type locality of the new species, was adequately set
forth in Harbison's paper.

My interest and effort centered largely around the factors concerned
in the life history of this butterfly, and of other lepidoptera in the area.
Having previously (1934) investigated and published on the meta-
morphosis of Megathymus stephensi Skinner, and more recently (1956)
on Megathymus evansi Freeman, it was my hope that a similar investi-
gation of the habits of the new species might bear fruit. Efforts along
that line had previously been made by others, without results.

Acknowledgments

I am greatly indebted to the several collaborators who had previously
carried on studies in the Genus Megathymus, and who have generously
given of their knowledge, experience, and published records. This
particularly includes H. Avery Freeman of Garland, Texas; the late
Brigadier W. H. Evans of the Department of Entomology, British
Museum, London; Don B. Stallings of Caldwell, Kansas; and Lloyd
M. Martin, Associate Curator of Entomology, Los Angeles County
Museum.

Colonel Arthur F. Fischer, former Director of the San Diego Museum
of Natural History, was most helpful in making available the facilities
of that institution, and lending support and encouragement to the
members of his staff engaged in research.

268 San Diego Society of Natural History

To Charles F. Harbison, Curator of Entomology, and indefatigable
staff worker in the San Diego Museum of Natural History, I express
appreciation for his dedication of the new species to one who is more
grateful than worthy of the honor.

Larval Habits of Megathymus comstocki

Megathymus cotnstocki lives as a larva in the interior of the leaves
of the century plant, known by the natives as "maguey", and to the
botanist as Agave. The dominant agave in the San Simon area is one
that was given the name Agave orcuttiana by Trelease, the type locality
being San Quintin, Baja California. A number of authorities on the
century plants are inclined to the opinion that A. orcuttiana is synony-
mous with Agave shawii Engelmann, or at best only a subspecies of
the latter.

In this connection, Stallings and Turner (1956) report that in the
Redington — Tucson area of Arizona Megathymus polingi Skinner
infests Agave schottii Engelmann, which is "the smallest known species
(of true Agave) in the U.S.A." A glance at Plate 18 of Harbison's
paper shows Agave shawii (orcuttiana) to be a large, robust, and heavy
species, occurring frequently in large dense clusters. This marked dif-
ference in food plants serves further to set apart Megathymus com-
stocki from its nearest relative, M. polingi.

Locating the larval chambers of M. comstocki in these plants proved,
at first, to be more difficult than was anticipated, owing to the fact that
they are at a lower and less exposed level than the larval chambers of
Megathymus stephensi or M. evansi. The burrows of these species occur
at a level well within the range of visibility, and the entrances are
typically on the upper surfaces of the leaves. In consequence, the
'doorway' to the larval tunnel is usually visible, and, in addition, the
extruded frass pushed out by the larva forms a telltale accumulation that
is plainly discernible. The entrance to the burrow of M. comstocki is
on the underside of the leaf, below the range of visibility.

It probably takes considerable time for the young larva of M. com-
stocki to channel its way down through the leaf to a location near its
base, where it eventually cuts out the chamber in which it spends the
greater part of its larval span, and in which it finally pupates. It was
particularly noted that the channel and chamber were contained in a
single leaf. This is in marked contrast to the habit described by Stallings
and Turner (1956) for Megathymus polingi. That species extends its

Comstock — Metamorphosis of A Megathymus 269

burrow and chamber through more than one leaf, and into the main
stalk or heart of the plant.

During its larval span M. comstocki extrudes its frass and exuviae,
as do M. stephens'i and polingi, and after each sanitary performance,
the exit is sealed and the burrow is thus closed to unwelcome intruders.
Even this precaution does not prevent the attacks of dipterous and
hymenopterous parasites, many of which were found in the pupal
chambers.

The Ovum, and Oviposition
In the short time at our disposal, it was impossible to obtain informa-
tion on the ovum, or on the method of ovipositing. It is however
assumed that, as in certain other species, the female takes a position on
the plant, and snaps or 'flicks' off her eggs, one at a time. These
probably fall close enough to the plant so that the young larva can seek
a leaf and burrow in near its tip. To demonstrate this assumption with
certainty would require observation in the field over a considerable
period of time. The method of ovipositing of the several yucca-feeding
megathymids is entirely different. The writer, in association with the
late Charles M. Dammers, (1934) established Megathymus yuccae
martini Stallings & Turner, affixes its egg solidly to the tip of a young
and succulent yucca leaf, and that the newly hatched larva burrows im-
mediately into the leaf, and thence down into the root.

Collecting Terrain and Technique

As previously reported by Harbison in his careful and thorough de-
scription of the new species, the type locality is a small side canyon of
Santa Maria Valley, about two miles northeast of San Simon, Baja
California Norte. The surrounding hills are heavily clothed with sub-
tropical desert vegetation, in which Agave shawii (orcuttiana) and
numerous cacti dominate. This is a rough and difficult terrain in
which to collect.

This agave is a compact unyielding plant, frequently growing in
crowded clumps, closely appressed to its fellows, seldom erect and
solitary. Its associates are predominantly spine-armored or thorny.
Its own sharp, recurved teeth along the margins of each leaf necessitate
the use of heavy gloves.

We were unable to locate the larval chambers until we had severed
the plant close to the ground, turned it top end down, and peeled off the
leaves, one by one, as a person does with an artichoke. The first leaves

270 San Diego Society of Natural History

to be thus removed were those that were dried and shriveled. In these
we discovered old and previously evacuated larval chambers.

As this peeling process proceeded downward on the upturned plant,
and the leaves began to show some green at their bases, the larval
burrows evidenced more recent occupancy. A few contained live pupae,
and several held shriveled larvae with puparia of a dipterous parasite.

Mature larvae were encountered in green leaves immediately proximal
to the desiccated layer. From that point onward toward the tip of the
plant, most of the leaves were sterile.

This process is long and tedious, and the total larvae plus pupae
obtained in the two days was only ten. A considerable number of
chambers containing empty pupal cases showed that we were somewhat
late in the season.

This collecting procedure is apparently the only one that gives
successful results with M. comstocki. Obviously it means destruction
of each agave examined, but by selective cutting in large masses the
remaining plants may be benefited.

The method employed in securing larvae of M. stephensi from Agdve
deserti Engelmann is in strong contrast to that described above. The
technique consists in using a crowbar to pry out a few leaves adjacent
to the one suspected of containing a larva or pupa, then removing the
desired leaf by hand. Plants thus handled soon repair the slight damage.

As with all megathymids, netting the imagines of M. comstock*
requires dexterity, and the results are seldom satisfactory. The insect
is a rapid and erratic flyer and frequently beats itself into a ragged
condition before it can be transferred to the cyanide jar.

In the rugged terrain that is its home, the thorny bushes cause
casualties to the net, and one must always move with caution, to avoid
dermal punctures with cactus spines or thorns. Rattlesnakes should
also be kept in mind, although, in this particular locality, only one
was observed during our stay.

Permits had previously been obtained by the authorities of the San
Diego Museum of Natural History for passing the living material
through the inspection station at the border on our return. The viable
specimens were thereafter held in the laboratory under careful safe-
guards against escape.

From the eight mature larvae and two pupae, only three imagines
emerged, one of which was deformed. The remaining larvae were

Comstock — Metamorphosis of A Megathymus 271

dormant for a considerable period of time before giving forth hymen-
opterous parasites.

Of the two females thus obtained in good condition, one served as
the allotype (No. l), and the other as paratype (No. 8) in Harbison's
type series.

Notes were made of the larva and pupa, and drawings prepared, as
will be noted on our figure (Plate 22).

Larva of Megathymus comstocki
(Plate 22, fig. l)

Mature Larva: Length, 33mm. for the single example measured.
The subcylindrical body tapers toward the head for the first three seg-
ments, and gradually narrows toward the cauda on the last six seg-
ments. It is characteristically grub-like in appearance.

Head: hemispherical, small, light tan in color, and covered with
dull yellow hairs. The mouth parts are brown, and the ocelli con-
colorous with the head.

Body: ground color bluish gray, the segmental junctures shading
to gray, and the caudal segments tinged with brown.

A conspicuous dark brown scutellum on the first segment arches
across the neck, uninterrupted in the median line.

There is a submedian longitudinal row of black points, one to a seg-
ment on each side. Each point bears a short black seta. A second
row of similar character parallels this laterally, but is less conspicuous,
and bears shorter setae.

There is a middorsal longitudinal pulsating green line.

The spiracles are rimmed narrowly with black. The true legs are
concolorous with the body, except that their tips are tinged with light
brown. The prolegs are likewise colored as is the body, and bear light
gray crochets.

The body is thickly sprinkled with minute yellowish hairs, discernible
only under a lens.

The body surfaces produce a flocculent white powder shortly before
pupation, but not to the profuse degree that was observed in Megathy-
mus stephensi.

272 San Diego Society of Natural History

Pupa of Megathymus comstockl
(Plate 22, figs. 2-3)

Measurements: Length, 28 mm. Greatest width through center,
7 mm.

The fusiform body tapers gradually toward the cauda and more
abruptly toward the head.

Head: Rounded, with the central cephalic elements protruding
forward in two lobes, one on each side of the median line.

The prominent eyes bulge laterally. Their color is slightly darker than
the dark brown of the head.

The leg sheaths and antennae are light brown, and the wing cases
yellow-tan. The maxillae extend to a point approximately 2 mm.
cephalad of the wing margins, and the antennae are still shorter by
about 1^/2 mm.

The abdominal segments are dark brown, tinged with olive, and
the spiracles are concolorous with the body.

Other structural features are adequately shown in the illustration,
Plate 22, figures 2 and 3.

Comparisons

It has been generally assumed that any Megathymus occurring below
the California — Mexican border, with a range more or less contiguous
to that of the San Diego County species, would likely prove to be a
close relative of Megathymus stephensi. The following table of com-
parisons, together with the genitalic differences demonstrated by Harbi-
son, prove that such is not the case.

COMPARSIONS OF THE LARVAE

M. comstocki M. stephensi

Shorter (33 mm.) Longer (40 mm.)

Slender Robust

Tapering gradually towards cauda Tapering abruptly towards cauda

Hair covering of head relatively long Hair covering of head short

Ocelli concolorous with head Ocelli dark brown

Body color bluish gray Body color soiled ivory

Body pile yellowish Body pile brownish

Crochets light gray Crochets black

Comstock — Metamorphosis of A Megathymus 273

Comparisons of the Pupae

M. comstocki M. stephensi

Subfusiform Subcylindrical

Curved ventrally Straight

Tapering caudally on last Tapering caudally only

seven segments on last four segments

Head irregularly rounded Head regularly rounded

Color of head dark brown Color of head amber

Abdominal segments dark brown, Abdominal segments

tinged with olive greenish-yellow

Cremaster long, flat, Cremaster short,

recurved relatively straight

Harbison's thorough analysis of his new species, and his figures
of genitalic armature, indicate that it is distinct from all others in the
genus, its closest relative being Megathymus polingi Skinner, described
originally (1905) from "Southern Arizona."

Stallings' and Turner's recording of Agave schottii as the food plant
of M. polingi is significant, as it is well established that nearly every
agave-feeder lives in a separate species of agave.

(Freeman had previously reported M. polingi as feeding in Agave
Palmeri Engelmann (1951 ) but he recently (in litt.) corrected this to
A. schottii.)

Parasites

As previously stated, a number of puparia of dipterous parasites
were found in the larval chambers, along with the shriveled larvae of
M. comstocki, from which they had emerged. These were left with
Paul Arnot (a member of our party on the San Simon trip). As yet,
no report on these has been received.

In addition, several hymenopterous parasites emerged from the seven
surviving larvae that were under observation in our laboratory. These
were submitted to P. H. Timberlake, of the Citrus Experiment Station,
University of California, at Riverside, who determined them as Apan-
teles megathymi Riley, with the comment that "This parasite was de-
scribed from North Carolina, but it has been recorded from Arizona
and California."

274 San Diego Society of Natural History

References

Comstock, John A.

1956. Notes on Metamorphoses of the Giant Skippers (Mega-
thyminae) and the Life History of an Arizona Species. Bull.
So. Calif. Acad. Sci. 55 (l) : pp. 19-27.

Comstock, John A. and Charles M. Dammers

1934. The Metamorphoses of Three California Diurnals. Bull.
So. Calif. Acad. Sci. XXXIII (2) : pp. 81-86.

1934. Ibid. Megathymus yucca navajo. pp. 87-92.

Freeman, H. Avery

1951. Notes on the Agave Feeders of the Genus Megathymus.
Field and Lab. XIX (l) : p. 29.

Harbison, Charles F.

1957. A new Species of Megathymus from Baja California. Trans.
San Diego Soc. Nat. Hist. XII (12) : pp. 231-262.

Skinner, Henry

1905. A New Megathymus from Arizona. Ent. News, XVI (7) :
p. 232.

Stallings, Don B. and J. R. Turner

1956. Notes on Megathymus polingi (Megathymidae) Lepidopt.
News, 10 (3-4) : p. 109.

Comstock — Metamorphosis of A Megathymus

275

Plate 22

Larva and pupa of Megathymus corns tocfo Harbison

1. Larva, dorsal aspect. 2. Pupa, ventral aspect. 3. Pupa, lateral aspect. All
figures enlarged approximately X 3. Reproduced from a painting by the author.

TRANSACTIONS

OF THE

_SAN DIEGO SOCIETY OF NATURAL HISTORY
Vol. XII, No. 14, pp. 279-286, figs. 1-4 July 12, 1957

FISH REMAINS FROM ABORIGINAL SITES IN THE
PUNTA PENASCO REGION OF SONORA, MEXICO

BY

W. I. FOLLETT

Curator of Ichthyology,
California Academy of Sciences

CONTENTS

Nomenclature 279

Fishes Represented 280

Fish Remains, by Site and Depth 282

Acknowledgments 283

Literature Cited 283

It has been my privilege to examine a collection of fish remains obtained
by Professor E. W. Gifford, of the University of California, from two aborigi-
nal sites in the Punta Pehasco region of northwestern Sonora, Mexico. One of
these sites is near Punta La Cholla, some five miles west and north of Punta
Pehasco. The other site, El Esterito, is near the mouth of the Sonoita River,
approximately ten miles south and east of Punta Pehasco. These aboriginal
sites have been described by Gifford (1946, pp. 216-218, fig. 29), and have
been compared by Schenck and Gifford (1952, p. 265) with similar sites at San
Felipe, Baja California, on the opposite shore of the Gulf of California. Photo-
graphs and maps of Punta La Cholla and vicinity were published by Hertlein
and Emerson (1956, figs. 1-2; pi. 12, figs. 1-3).

NOMENCLATURE

The ichthyological nomenclature employed in this paper generally follows
that of Hubbs and Follett (MS.). The osteological nomenclature follows
the later usage of Starks (1901, 1923, 1930).

280 San Diego Society of Natural History

FISHES REPRESENTED

The collection, taken from the sites and at the depths noted on page 282,
consists of 41 fish remains. Of these, the 38 that are identifiable represent
the nine species listed below. All are commonly taken, at one season or an-
other, in comparatively shallow water near the head of the Gulf of California.

TRIAKIDIDAE — Smoothhounds

Mustelus lunulatus Jordan and Gilbert (?). — The sicklefin smoothhound
is edible and is sold in the markets, according to Berdegue (1956, p. 105), who
stated that the vernacular name "tiburon mamon" ("suckling shark"), applied
to this species, refers to its weak dentition and small size. We saw one of
these sharks captured on hook and line from a rocky promontory at Punta
San Felipe. The 681/2-inch specimen recorded by Norris (1923, p. 1) is the
largest of which I have found mention. A figure of this species was published
by Kumada and Hiyama (1937, pi. 47). Material (referred with some
doubt to this species) : seven complete centra, possibly all from the same
fish, which was approximately 30 inches long. Because of the lack of adequate
comparative material, the identification is presumptive.

CARCHARHINIDAE — Requiem Sharks

Scoliodon longurio (Jordan and Gilbert) . — The Pacific sharpnose shark
is used as food to a limited extent in Panama, according to Meek and Hilde-
brand (1923, p. 53). The 421/2-inch specimen, weighing nine pounds, recorded
by Beebe and Tee- Van (1941, p. 112), is the largest I have found mentioned.
Walford (1944, p. 5) stated that this species was known as "puro" ("cigar";
doubtless in reference to its shape) . A figure of this shark was published by
Hubbs and McHugh (1950, fig. 3). Material: one complete centrum,
from a fish approximately 16 inches long.

SERRANIDAE — Basses

Mycteroperca jordani (Jenkins and Evermann) . — The Gulf grouper,
"baya," is a valuable food fish, according to Evermann and Jenkins (1891,
p. 143). It was stated by Jordan and Evermann (1896, p. 1177) to be common
in bays and sheltered waters of the Gulf, but not about rocks. Walford (1937,
p. 105) reported the largest specimen caught by his party as measuring four
feet eight inches in length, although elsewhere (1936, p. 8) he referred to a
specimen that measured between five and six feet. Figures of both young and
adult were published by Walford (1937, pi. 12, fig. a). Material: one
nearly complete vertebra, the 1st precaudal, from a fish approximately 15
inches long. The identification is made on the basis of agreement with a
235-mm. specimen of Mycteroperca jordani from Guaymas, and of disagree-
ment with material of Epinepbelus analogus Gill and of Paralabrax maculato-
jasciatus (Girard) .

JUL 6 1 13

HRVARD
■HIERSffi

Follett — Fish Remains from Sonora 281

MUGILIDAE — Mullets

Mugil cephalus Linnaeus. — The striped mullet, "lisa" (Beltran 1934,
p. 7), is exceedingly abundant in the Gulf. At times during our stay at San
Felipe, scarcely a second would elapse when a leaping mullet could not be seen
in one direction or another. Berdegue (1956, p. 127), who commented on its
extensive use for food, stated that the Yaqui fishermen take it with the spear.
Since this species feeds on minute algae and detritus (Dill 1944, p. 141), it is
seldom taken on hook and line. Furthermore, it is so agile that it will usually
leap over a seine, although according to Thompson and Bryant (1920, p. 60)
it has been captured by weirs placed in shallow water. Roedel (1953, fig. 76)
published a photograph of this species. Material: one incomplete basioc-
cipital; one nearly complete left hyomandibular (fig. 1) ; nine incomplete caudal
vertebrae, appearing to represent the 3rd, 4th, 6th, 8th, 10th, 11th (two), 12th,
and 13th; one complete hypural. These elements are from fish ranging approxi-
mately from 15 to 20 inches in length.

SPARIDAE — Porgies

Calamus taurinus (Jenyns) (?). — The Pacific porgy, "mojarron chino"
or "mojarra garabata," is an important food fish, according to Berdegue (1956,
p. 251), who stated that it attains a length of approximately 18 inches. A
colored plate of this species was published by Kumada and Hiyama (1937,
pi. 33) and another, by Walford (1937, pi. 68). Material: one incomplete
vertebra, questionably the 1st caudal, from a fish approximately 15 inches
long. The identification is queried because of minor inconsistencies observed
on comparing material.

SCIAENIDAE — Croakers

Cynosaon xanthulus Jordan and Gilbert. — The orangemouth corvina was
one of the best food fishes that we encountered at San Felipe. We took it occa-
sionally on hook and line while fishing from a sand beach, although we caught
it more frequently from a skiff a short distance offshore. Jordan and Ever-
mann (1898, p. 1410) stated that it attains a length of three feet. A photo-
graph of this species was published by Walford (1937, pi. 20, fig. d).
Material: one incomplete vertebra (fig. 2), the 7th (?) caudal, from a fish
more than 30 inches long.

Cynosaon macdonaldi Gilbert. ^ — The totuava, also spelled "totoaba"
(Beltran 1934, p 12; Berdegue 1956, p. 235), attains a weight of at least 225
pounds (Walford 1937, p. 132). It was reported by Gilbert (1890, p. 65)
as being very abundant along the entire eastern shore of the Gulf and as
feeding in shallow water, where it was easily approached and speared. Naka-
shima (1916, pp. 85, 86) stated that this species often comes into very shallow
water; that it is not very shy; and that when one is hooked or speared, the
others crowd around as if in curiosity. The wasteful exploitation of this food
fish since the advent of the white man has been described by Chute (1928,
p. 275). A photograph of the spotted juvenile stage was published by Berdegue

282 San Diego Society of Natural History

(1956, fig. 168), and an excellent color photograph of the adult, by Walford
(1937, pi. 65). Material: one fragmentary left and one fragmentary right
epiotic; one incomplete left quadrate (fig. 3); one fragmentary right maxillary;
one fragmentary right ceratohyal; four vertebrae (two nearly complete, the
3rd and 4th precaudal, probably from the same fish; one fragment, the 6th
precaudal; one fragment, the 6th (?) caudal); two fragmentary fin rays (dorsal
and caudal); one fragment, questionably epural. These elements are from fish
ranging approximately from 20 to 60 inches in length.

GIRELLIDAE — Nibblers

Girella simplicidens (Steindachner) . ■ — The Gulf opaleye occurs about
rocky points, its young entering tide pools, as at Punta San Felipe, where we
found it abundant. It is edible, though not esteemed. Osburn and Nichols
(1916, p. 166) recorded a specimen 15 inches in length and published a figure
of this species. Material: one incomplete vertebra, the 1st (?) caudal, from
a fish approximately 9 inches long.

BALISTIDAE — Triggerfishes

V errunculus polylepis (Steindachner) . — • That the fine-scale triggerfish
is used as food was indicated at San Felipe, where on two occasions the remains
of adults, with the flesh sliced from each side, floated past our camp. We caught
a number of small specimens by hook-and-Iine fishing from rocky points. This
species was stated by Jordan and Evermann (1898, p. 1700) to reach a length
of two feet. It is known as "pez puerco," according to Beltran (1934, p. 10).
Photographs of this fish and of its trigger mechanism were published by
Clothier (1939, figs. 86-89). Material: two incomplete vertebrae, the 1st
precaudal (fig. 4), from a fish approximately 14 inches long, and the 4th
precaudal, from one of approximately 10 inches.

FISH REMAINS, BY SITE AND DEPTH

The numeral heading each paragraph is a catalog number of the Museum
of Anthropology, University of California, Berkeley.

3-15177. La Cholla Bay, approximately one-fourth mile north of "Site 1"
as indicated by Gifford (1946, fig. 29), on face of sand dunes near the slough;
depth 13 to 26 inches. — Mugil cephalus: left hyomandibular (fig. 1). Cyno-
scion macdonaldi: two epiotic fragments (left and right); left quadrate (fig.
3); right-maxillary fragment; right-ceratohyal fragment; 6th (?) caudal vertebra,
fragmentary. Unidentified material: two fragments.

3-15178. Same site and depth as preceding. — Cynoscion xanthulus:
7th (?) caudal vertebra (fig. 2).

3-15179. Same site and depth as preceding. — Mugil cephalus: basioc-
cipital; five vertebrae, appearing to represent the 6th, 8th, 11th, 12th, and
13th caudal. Calamus taurinus (?) : 1st (?) caudal vertebra. Cynoscion
macdonaldi: two vertebrae, the 3rd and 4th precaudal. V errunculus polylepis:
4th precaudal vertebra.

Follett — Fish Remains from Sonora 283

3-15184. La Cholla Bay, approximately one-fourth mile north of "Site 1"
as indicated by Gifford (1946, fig. 29), on hummock No. 4 near the slough;
depth 20 to 30 inches. — Scoliodon longnrio: centrum. Mugd cephalns:
three vertebrae, appearing to represent the 4th, 10th, and 11th caudal; hypural.
Cynoscion macdonaldi (?) : 6th precaudal vertebra, fragmentary. Girella
simplicidens: 1st (?) caudal vertebra. Verrunculus polylepis: 1st precaudal
vertebra (fig. 4). Unidentified fragment: rim of vertebra.

3-15231. El Esterito, at "Site 3" as indicated by Gifford (1946, fig. 29);
depth 0 to 6 inches. — Mustelns lunulatus (?) : seven centra. Mugil cephalus:
3rd (?) caudal vertebra.

3-15246. Same site as preceding; depth 6 to 12 inches. — Cynoscion
macdonaldi: dorsal-ray fragment; epural (?) fragment; caudal-ray fragment.

3-15281. Same site as preceding; depth 18 to 24 inches. — Mycteroperca
jordani: 1st precaudal vertebra.

ACKNOWLEDGMENTS

For the opportunity to report on this collection of fish remains, I wish
to acknowledge my indebtedness to E. W. Gifford. I wish also to express
my appreciation to my wife, Evelyn Follett, and especially to Richard S.
Croker and his wife, Annie Croker, for extensive assistance, during a week's
stay at San Felipe, in collecting most of the comparative material used in
identifying these remains. My thanks are due to Lillian Dempster and to
Carl L. Hubbs for aid in the preparation of the manuscript.

LITERATURE CITED

Beebe, William, and John Tee-Van

1941. Eastern Pacific expeditions of the New York Zoological Society.
XXV. Fishes from the tropical eastern Pacific. (From Cedros
Island, Lower California, south to the Galapagos Islands and
northern Peru.) Part 2. Sharks. Zoologica 26: 93-122, figs. 1-34,
pis. 1-2.

Beltran, Enrique

1934. Lista de peces mexicanos. 1-13 (processed). Instituto Biotecnico,
Secretaria de Agricultura y Fomento, Mexico, D. F.

Berdegue A., Julio

1956. Peces de importancia comercial en la costa nor-occidental de
Mexico. 1-345, figs. 1-206, 2 maps. Comision Para el Fomento
de la Piscicultura Rural, Secretaria de Marina, Mexico, D. F.

Chute, Geo. Roger

1928. The totuava fishery of the California Gulf. Trans-desert trucking
of Mexican-caught fish. Calif. Fish & Game 14: 275-281, figs. 81-88

284 San Diego Society of Natural History

Clothier, Charles R.

1939. The trigger mechanism of a trigger fish (Capriscus poiylepis) .
Calif. Fish & Game 25: 233-236, figs. 86-89.

Dill, William A.

1944. The fishery of the lower Colorado River. Calif. Fish & Game
30: 109-211, figs. 45-82.

Evermann, Barton W., and Oliver P. Jenkins

1891. Report upon a collection of fishes made at Guaymas, Sonora,
Mexico, with descriptions of new species. Proc. U. S. Nat. Mus.
14: 121-165, pis. 1-2.

GlFFORD, E. W.

1946. Archaeology in the Punta Penasco region, Sonora. Am. Antiquity
11: 215-221, fig. 29.

Gilbert, Charles H.

1890. Scientific results of explorations by the U. S. Fish Commission
steamer Albatross. XII. — A preliminary report on the fishes
collected by the steamer Albatross on the Pacific coast of North
America during the year 1889, with descriptions of twelve new
genera and ninety-two new species. Proc. U. S. Nat. Mus. 13:
49-126.

Hertlein, Leo George, and William K. Emerson

1956. Marine Pleistocene invertebrates from near Puerto Penasco, Sonora,
Mexico. Trans. San Diego Soc. Nat. Hist. 12: 154-176, pi. 12,
2 maps.

Hubbs, Carl L., and W. I. Follett

MS. List of the fishes of California.

Hubbs, Carl L., and J. L. McHugh

1950. Pacific sharpnose shark (Scoliodon longurio) in California and
Baja California. Calif. Fish & Game 36: 7-11, figs. 3-4.

Jordan, David Starr, and Barton Warren Evermann

1896. The fishes of North and Middle America: a descriptive catalogue
of the species of fish-like vertebrates found in the waters of North
America, north of the Isthmus of Panama. Part 1. Bull. U. S.
Nat. Mus. 47: i-lx+ 1-1240.

1898. The fishes of North and Middle America ... Part 2. Bull. U. S.
Nat. Mus. 47: i-xxx+ 1241-2183.

Kumada, Tosio, and Yosio HlYAMA

1937. Marine fishes of the Pacific coast of Mexico. 1-75, pis. 1-102
(1-43 colored). The Nissan Fisheries Institute & Co., Ltd.,
Odawara, Japan.

Meek, Seth E., and Samuel F. Hildebrand

1923. The marine fishes of Panama. Part I. Field Mus. Nat. Hist. Publ.
Zool. 15: i-xi+ 1-330, pis. 1-24.

Follett — Fish Remains from Sonora 285

Nakashima, Eiichiro

1916. Notes on the totuava (Cynoscion macdonaldi Gilbert) . Copeia
37: 85-86.

Norris, H. W.

1923. The occurrence of Mustelus lunulatus on the California coast.
Copeia 114: 1-2.

Osburn, Raymond C, and John Treadwell Nichols

1916. Shore fishes collected by the 'Albatross' expedition in Lower
California with descriptions of new species. Bull. Am. Mus. Nat.
Hist. 35: 139-181, figs. 1-15.

Roedel, Phil M.

1953. Common ocean fishes of the California coast. Calif. Dept. Fish &
Game, Fish Bull. 91: 1-184, figs. 1-175, colored frontispiece.

Schenck, W. Egbert, and E. W. Gifford

1952. Archaeological sites on opposite shores of the Gulf of California.
Am. Antiquity 17: 265.

Starks, Edwin Chapin

1901. Synonymy of the fish skeleton. Proc. Wash. Acad. 3: 507-539,
figs. 45-46, pis. 63-65.

1923. The osteology and relationships of the uranoscopoid fishes. Stan-
ford Univ. Publ. Biol. Sci. 3: 255-290, pis. 1-5.

1930. The primary shoulder girdle of the bony fishes. Stanford Univ.
Publ. Biol. Sci. 6: 147-239, figs. 1-38.

Thompson, Will F., and Harold C. Bryant

1920. The mullet fisheries of Salton Sea. Calif. Fish & Game 6: 60-63,
figs. 23-25.

Walford, Lionel A.

1936. Contributions from the Fleischmann expedition along the west
coast of Mexico. II. The groupers (Mycteroperca) of the Pacific
coast of the Americas. Occ. Pap. Santa Barbara Mus. Nat. Hist.
4: 5-8.

1937. Marine game fishes of the Pacific coast from Alaska to the
Equator. i-xxix+ 1-205, figs. 1-56, pis. 1-69 (33-69 colored),
colored frontispiece, 1 map. Univ. Calif. Press, Berkeley.

1944. Observations on the shark fishery in the central part of the Gulf
of California, with records of vitamin potency of liver oils and
with keys to the identification of commercially important sharks.
Fishery Market News 6 (6) : 2-7, figs. 1-10.

286

San Diego Society of Natural History

Fig. 1. Mugil cephalus, left hyomandibular; height 25 mm.; UCMA 3-15177.
Fig. 2. Cynoscion xanthulus, 7th (?) caudal vertebra; length 30 mm.; UCMA
3-15178. Fig. 3. Cynoscion tnacdonaldi, left quadrate; height 26 mm.; UCMA
3-15177. Fig. 4. Verrunculus polylepis, 1st precaudal vertebra; height 14 mm.;
UCMA 3-15184.

TRANSACTIONS

OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY
Vol. XII, No. 15, pp. 287, 288 September 25, 1957

A NEW RACE OF WOOD RAT (NEOTOMA)

FROM THE GULF SIDE
OF CENTRAL BAJA CALIFORNIA, MEXICO

BY

Laurence M. Huey

Curator of Birds and Mammals, San Diego Society of Natural History

Study of the chain of races of Neotoma lepida, from southern California
down through the peninsula of Baja California, reveals the presence of an
undescribed race on the coastal Gulf slope of the central section of this long
peninsula. This race may be known as

Neotoma lepida aridicola subsp. nov.
San Francisquito White-footed Wood Rat.

Type.— From El Barril (near 28° 20' N), Gulf of California, Baja
California, Mexico; No. 15595, collection of the San Diego Society of Natural
History; adult male; collected by Laurence M. Huey. March 27, 1947.

Characters. — Neotoma lepida aridicola is similar in body size to Neo-
toma lepida felipensis, the race living along the Gulf coast to the north in the
vicinity of San Felipe, but its buffy color appears richer and brighter when the
two forms are compared in series. Cranially, N. 1. aridicola differs from N. I.
felipensis in several characters. The rostrum is narrower and more slender
and the braincase is more rounded when viewed dorsally. Posteriorly, the
braincase is narrower and more vaulted. The ventral surface of the skull of
N. I. aridicola shows several outstanding differences when compared with
N. I. felipensis. The molars are smaller and the series from m1 to m3 tapers
more acutely. The pterygoids are more nearly parallel than those of N. I.
felipensis, and the auditory bullae are decidedly smaller and rounder.

Compared with Neotoma lepida molagrandis, the race that occupies the
opposite side of the peninsula on the Pacific Ocean slope, N. I. aridicola is
immediately set apart by its very light-colored pelage and its smaller size. In
cranial characters N . I. aridicola differs as much from N. I. molagrandis as it
does from N. I. felipensis; the bullae are smaller and more rounded, and the
molariform teeth are smaller.

The light-colored desert race N. I. aridicola differs conspicuously in color
from the dark-pelaged Neotoma lepida ravida, the race found in the black
lava mountainous district to the south. Cranial differences are not so pro-
nounced as between the two former races, though the molariform teeth and
auditory bullae differ slightly.

OCT 2 4 19

K-. '-3
ofJY

288 San Diego Society of Natural History

Measurements of Type. — Total length, 325; tail, 145; hindfoot, 31;
ear, 28. Skull: Greatest length, 41.2; zygomatic breadth, 21.1; interorbital
breadth, 5.3; greatest length of nasals, 16.2; length of palatal bridge, 7.4; alveolar
length of upper molar series, 7.3.

Range. — So far as known, the desert slopes on the Gulf side of the
peninsula in the region from San Francisquito Bay to El Barril. Further ex-
ploration of the arid Gulf slopes will probably show that this race lives con-
siderably farther both to the north and to the south.

Remarks. — Studies of the various species of small land mammals living
on the peninsula of Baja California reveal links in the racial chains yet to be
described. These are to be found in the remote regions which are difficult of
access in this interesting land. As these areas are explored and carefully trapped,
novelties will be collected, such as the wood rat described herein.

Following is a list, with type localities, of the races of the wood rats of the
Neotoma lepida group now known to occur on the peninsula, exclusive of its
islands.

Race Type locality

Neotoma lepida intermedia Dulzura, San Diego Co., California.

Neotoma lepida gilva Banning, Riverside Co., California.

Neotoma lepida jelipensis San Felipe, Baja California, Mexico.

Neotoma lepida egressa 1 mi. e. of El Rosario, Baja California, Mexico.

Neotoma lepida niolagrandis Santo Domingo Landing (3 mi. inland from

beach; near 28° 15' N), Baja California,
Mexico.

Neotoma lepida andicola El Barril (near 28° 20' N) , Gulf of Cali-

fornia, Baja California, Mexico.

Neotoma lepida ravida Comondu, Baja California, Mexico, alt. 700'

Neotoma lepida pretiosa Matancita (= Soledad), 50 mi. n. cf Mag-

dalena Bay, Baja California. Mexico.

Neotoma lepida notia La Laguna, Sierra de la Victoria, southern

Baja California, Mexico.

Neotoma lepida arenacea San Jose del Cabo, Baja California, Mexico.

Specimens examined. — Neotoma lepida jelipensis. Baja California, Mex-
ico: San Felipe (type locality), 15.

Neotoma lepida egressa. Baja California, Mexico: 7 mi. e. of San Quintin,
3; 10 mi e. of San Quintin, 5; Santa Maria near San Quintin, 7; San Quin-
tin, 4; Aguaita. 1; 15 mi. nw. of San Fernando, 5; Santa Catarina Landing 1.

Neotoma lepida ravida. Baja California, Mexico: south end of Concep-
tion Bay, 7; Comondu (type locality), 5.

Neotoma lepida andicola. Baja California, Mexico: El Barril (near 28°
20' N), Gulf of California (type locality), 11; 7 mi. w. of San Francisquito
Bay, 2

Neotoma lepida niolagrandis. Baja California, Mexico: Santo Domingo
Landing (3 mi. inland from beach; near 28° 15' N; type locality), 2; Punta
Prieta, 2; Mesquital, 2; Calmalli, 1: Santa Gertrudis, 1; 12 mi. e. of El Arco,
1; San Ignacio, 3.

TRANSACTIONS

L...

0CT24
liill.uo,

OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY

Volume XII, No. 16, pp. 289-308, figs. 1-6

LATE PLEISTOCENE FAUNAS FROM

THE NORTHWESTERN COAST OF

BAJA CALIFORNIA, MEXICO

BY

James W. Valentine

Department of Geology,
University of California, Los Angeles

SAN DIEGO, CALIFORNIA

Printed for the Society

September 25, 1957

Border Locality +

32° 30'

lNTE_RNiTlO_NA_L.
BOUNDARY

U.C.L.A. Loc No 3161

UCLA Loc No 3160

o

U C LA Loc No 2720
UCLA Loc No 271 9

U C LA Loc No 2715

UCLA. Loc No 2718 "^NpjWi C LA Loc No 3162
UCLA Loc No 2717

Area included
in this map

Rio

San Miguel

0 12 3 4

Fig. 1. Index map of northwestern Baja California, showing fossil
localities between the International Boundary and Rio San Miguel recorded here.

Base maps: U. S. Navy Hydrographic Office Chart no. 1149, San Diego to Bahia
San Quintin, ed. 45, 1948; U. S. Army Air Force Preliminary Base Map, (404C)
San Diego, 1945.

mil.

OCT 2 4

LATE PLEISTOCENE FAUNAS FROM

THE NORTHWESTERN COAST OF

BAJA CALIFORNIA, MEXICO

BY

James W. Valentine
CONTENTS

Introduction 291

Previous Work 292

Description of the Area

General Features 293

Local Features 293

Composition of the Faunas

General Features 294

Specific Composition of Faunas 295

Habitats and Suggested Depositional Environments 301

Geographic Ranges and Suggested Temperatures 302

Climatic Significance of the Faunas 302

Description of Fossil Localities 303

Acknowledgments 304

Literature Cited 305

m

INTRODUCTION

The Council of the Eighteenth International Geological Congress (Anony-
mous 1950) recommended that the base of the Pleistocene in its type-area in
Italy be drawn at the base of the Calabrian (marine) and Villefranchian (con-
tinental) formations. It was noted in the recommendation that this boundary
coincides with the earliest indication of climatic deterioration in the late Cenozoic
section of Italy. In order to locate this boundary in marine sediments of other
regions, it seems necessary to determine the major features of Pleistocene oceanic
circulation patterns in each area and their thermal significance. In view of the
great provinciality of Pleistocene faunas, such a climatic approach to Pleistocene
correlation, particularly on the interprovincial and intercontinental scales, is
promising. Reconstruction of the history of marine Pleistocene circulation in
the eastern Pacific should greatly aid the establishment of stage boundaries here.

In the hope of bringing more evidence to bear upon this problem, a study
of Pleistocene invertebrate faunas has been undertaken, particularly those from
marine terraces along the coast of southern California and Baja California.
These terrace faunas provide especially favorable material for paleoecologic
studies, inasmuch as representatives of nearly all the species are living today
in the eastern Pacific. Thus reconstructions of their ancient environments may

292 San Diego Society of Natural History

be based directly upon their present distribution and habits. Furthermore, the
paleogeographic settings of many terrace fossil localities are suggested by
present topography. The configuration of a terrace shoreline is commonly in-
dicated by the position of ancient sea-cut cliffs at the landward margins of
the terrace. The difference in altitude between a fossil locality and this shoreline
approximates the maximum depth of water under which the fauna could have
been deposited.

A large number of invertebrate assemblages, chiefly of mollusks, has been
described from Upper Pleistocene marine terraces in southern California. The
majority of species in these faunas live today in shallow-water environments
such as were doubtless present near the fossil localities. To interpret the
environment of such assemblages of Recent types in known geographic situa-
tions is to approach paleoecology in its simplest aspect. Nevertheless, the
faunas regularly contain a few species that do not live at present in association
or near-association with the bulk of the fauna. Except for cases of probable
reworking, nearly all the ecologically anomalous forms represent either (1)
locally extinct species that live in southern, warmer water or northern, cooler
water or (2) locally deep-water types that live in shallow water only to the
north. Cool-water types and warm-water types are found together at some
localities. This situation has been well outlined by Woodring (in Woodring,
Bramlette, and Kew 1946) .

The present report, which is part of a larger work in progress, records
faunas collected in 1952 from ten localities on the lowest remaining terrace
between the International Boundary and Rio San Miguel (fig. 1). The com-
position of these faunas was outlined in a preliminary paper (Valentine 1954) .

Previous Work

Dall apparently was responsible for the earliest record of Pleistocene fossils
from Baja California (1890, p. 145). A single species, Eupleura muriciformis,
was reported from the "Pacific coast of Lower California."

Although no Pleistocene faunas have previously been described from the
area under consideration, Emerson and Addicott (1953) enumerated a ter-
race fauna from just north of the International Boundary at the "Border
locality" (fig. 1). The Border fauna, collected evidently from the same
terrace as the fossils recorded here, was interpreted as indicating quiet shallow
water with temperatures much like those of today.

A fauna recently described from terrace deposits at Punta China (Emerson
1956) represents the nearest recorded Pleistocene fossils to the south of Rio
San Miguel. The Punta China assemblage includes several species living only
well to the north at present and was interpreted as indicating cooler water, at
least locally, than is found near Punta China today. Records by Dall (1900,
p. 1001; 1903, pp. 1245, 1258) of Pleistocene shells from Todos Santos Bay
presumably are based upon Indian midden material.

Valentine — Pleistocene Faunas from Baja California 293

DESCRIPTION OF THE AREA

General Features

The coast between the International Boundary and Rio San Miguel (fig. 1)
is chiefly one of steep irregular or crescentic beaches, separated by rocky points
and backed by sea-cut cliffs. In the Punta Descanso region and northward,
rocks exposed in the cliffs are black to dark reddish vesicular basalts interbedded
with light colored tuffs and tuffaceous sandstones. No fossils are reported from
these rocks. Hertlein and Grant (1944, p. 60) suggested that these volcanics
overlie the San Diego formation, and are therefore of "late San Diego age"
(Upper? Pliocene). They were mapped by Beale (1948, p. 74, pi. 1) as the
"Comondu formation," a term he used to include "all volcanic rocks and
terrestrial sediments of Tertiary age" in Baja California, and were assigned to
the Middle or Upper Miocene on a reconnaissance map published by Anderson
(1950, pi. 1).

From the Punta Descanso area southward to Rio San Miguel, rocks ex-
posed in the sea cliffs have been mapped as Cretaceous (Beale 1948, pi. 1,
"Rosario formation"; Anderson 1950, pi. 1). The basis for the age assignment
is not known; some of the rocks are massive tuffs and sandstones with inter-
bedded basalts, and are not unlike the "Comondu" rocks farther north.

A terrace platform, eroded on "Comondu" and "Rosario" rocks, is exposed
in the sea cliffs at altitudes chiefly between 25 and 45 feet. The platform surface
slopes gently seaward, and differences in the altitude of the terrace platform at
the sea cliffs are due largely to variations in the distance between the cliffs and
the shoreline angle at the rear of the terrace. Wherever the position of the
shoreline angle could be determined, it was judged to be at about 75 feet in
altitude. Although the terrace is interrupted by several stream valleys, terrace
segments north and south of the valleys extend for many miles at essentially
the same altitudes and are presumably correlative. If so, this terrace extends
from the San Diego region (as the Nestor terrace of Ellis, in Ellis and Lee 1919,
pi. 6) at least as far south as Rio San Miguel, a distance of approximately 45
miles. The configuration of the terrace shoreline suggests that the coast was
formerly one of alternating rocky shores and sandy beaches, much as it is
today. The ten fossil assemblages recorded below were collected from marine
deposits preserved locally on this terrace platform.

Local Features

A notable exposure of Pleistocene sediments occurs on the north side
of Punta Descanso (figs. 2, 3), where about 3 feet of richly fossiliferous
pebbly sand, containing lenses of basalt cobbles at the base, lies on a basalt
terrace platform (locality 2715). Overlying the sand is a second fossiliferous
deposit, 3 feet thick, with a matrix of alluvium and comminuted shell (locality
2716). The upper unit evidently represents either (1) a non-marine deposit
containing abundant reworked shell material eroded from a shoreward part
of the underlying marine stratum or (2) a beach deposit with which alluvial
material has mingled. The fossiliferous deposits are overlapped by an alluvial
fan.

294 San Diego Society of Natural History

At Rio Monro (locality 3162) the lowest terrace platform is clearly shown
in a road cut (figs. 4, 5) . This locality is near the rear of the terrace; remnants
of the next higher terrace may be seen above the cut in fig. 4. Between 2 and
3 feet of a fossiliferous, poorly sorted sand, grit, and rubble conglomerate rest
on the platform surface. Conglomeratic fragments of pebble to boulder sizes
are well- to sub-rounded. The conglomerate is overlain by 4 to 5 feet of
alluvium. The platform rocks are basalts, and the matrix of the fossiliferous
stratum consists in large measure of weathered basaltic material. This deposit
is interpreted as talus rubble washed or wasted from the former sea cliff, together
with alluvium and talus material from higher terraces, intermingled with
marine sediments.

Small lenses of fossil mollusks occur on the low terrace between the
fossil localities reported here. These faunas are small, the shells usually well
worn and broken; all species identified are found also in the assemblages listed
below. The fauna of one of these fossiliferous lenses has been included in the
list of species; the locality (locality 3569, 4J/2 miles south of Rio Morro) is
illustrated in fig. 6. It typifies the occurrence of small fossiliferous lenses.
Between Rio Morro and Rio San Miguel, parts of the terrace are dissected into
badlands by numerous steep-sided gullies, and marine sediments that locally
contain a few shells are exposed in several of the deeper gullies near the present
sea cliff.

Shells of mollusks used for food by the early Indians of Baja California
are found throughout the area, usually within and upon alluvial fans mantling
the marine terraces. The shells may be scattered and rare or in heaps or layers
commonly termed Indian middens. At many localities, middens are so thick
and extensive that they may readily be mapped. Only a few molluscan species
are present in most of these middens, chiefly Mytilus calif ornianus, Septifer
bijurcatus, Lottia gigantea, Haliotis cracherodi, and Fissurella volcano. In col-
lecting Pleistocene fossils in this region, it is necessary to exercise great care
to insure against contamination by the closely associated midden shells.

COMPOSITION OF THE FAUNAS
General Features

A total of 230 molluscan species and subspecies are found in the collections,
of which 23 are of more or less uncertain identity. Gastropods number 163,
pelecypods 54, chitons 12, and scaphopods 1. The numbers of molluscan species
found at each locality are tallied by classes in table 1. The greater number
of gastropods relative to other classes at localities 2715, 2716, 2718, 2717,
and 3162 is due to the presence of many small to minute species in these faunas.
Six species are particularly abundant and widespread; these are Mytilus cali-
fornianus, Protothaca staminea, Olivella biplicata, "Nassa" fossata, Littorina
scutulata, and Tegula funebralis.

Remains of invertebrates of groups other than mollusks have not been
identified, except for test fragments of the sea-urchin S trongylocentrotus pur-
puratus from localities 2715 and 2716. Barnacles (Balanus and Tetraclita),

Valentine — Pleistocene Faunas from Baja California 295

Table 1. Composition of the faunas by classes, and present ranges of their

species and subspecies

U.C.L.A. LOCALITY NUMBER

3161

3160 2720 2719 2715 2716 2718 2717 3162 3569

Identified species
and subspecies

47

60

32

39

107

96

107

110

68

16

Species or

subspecies of

uncertain identity
Pelecypoda
Scaphopoda
Gastropoda
Amphineura
Species or

1

14
1

33
0

1
18

1

42

0

1

12
0

21
0

6

17

0

26

2

7
24

0
84

6

4
22

0
71

5

7
20

1
90

3

10
28

1
87

4

4
13

0
59

0

4
7
0
12
1

subspecies living
only to the north

1

5

2

4

5

6

7

7

3

1

Species or

subspecies living
only to the south

1

1

1

1

3

2

1

1

1

0

crab-claws, and worm-tubes are common at most localities. No foraminifera
were found.

The following forms have not previously been recorded as fossil: Lasaea
subviridus, Odostomia trachis, O. virginalis, Cerithiopsis carpenteri, Triphora
hempbilli, Nodulus kelseyi, Amphithalamus tenuis, Rissoina cleo, Caecum
licalum, and Acmaea persona strigatella.

Only a single form that is not known to be living, Pseudomelatoma pent-
cillata semilnjlata, is found in the collections.

Specific Composition of Faunas

Table 2 lists by localities all species of mollusks collected. Each symbol
in table 2 indicates the abundance of a species relative to the entire fauna
collected at the locality. This method permits specific comparison between
faunas irrespective of size. The following symbols are used :

Representation per 1000
Symbol Meaning specimens collected

V Very rare 2 or fewer

R Rare 3 to 8

C Common 9 to 32

A Abundant 33 to 128

S Superabundant 129 or more

To obtain as many species as possible, somewhat selective collections were
made at all localities. In addition, at richly fossiliferous localities, bulk collec-
tions were made to provide abundance data. Nevertheless, the abundance
values in the table are biased in favor of scarce forms.

296 San Diego Society of Natural History

Table 2. List of species, showing abundance in each fauna
Meaning of symbols: V, very rare; R, rare; C, common; A, abundant; S, super-
abundant. For further explanation, see page 295.

u.c.l.a. locality number

Pelecypoda

Nuculana taphria (Dall) V

Glycymeris aff. G. profunda (Dall) C .. R .. .. R

Glycymeris subobsoleta (Carpenter) R R .. C

Barbatia bailyi (Bartsch) R V

Ostrea lurida Carpenter V V .. ..

Ostrea megodon Hanley R

Leptopecten latiauratus (Conrad) V v R ..

Leptopecten latiauratus monotimeris (Conrad) . R R

Leptopecten ? sp C

Pecten diegensis Dall - R

Hinnites giganteus (Gray) C v V v V ..

Pododesmus macroschisma (Deshayes) C .. .. R .. V R V .. ..

Mytilus californianus Conrad ACAARCVVRC

Septifer bifurcatus (Conrad) R .. .. R R C V R R ..

Modiolus jornicatus Carpenter v C .. R ~ ~

Glans subquadrata (Carpenter) v .. .. R C v C R ..

Milneria kelseyi Dall ■■ C .. .. V V ..

Chama pellucida Broderip V V R v .. ..

Pseudochama exogyra (Conrad) R R R C R ..

Epilucina californica (Conrad) A R R C .. ..

Here excavata (Carpenter) R

Lucinisca nuttalli (Conrad) R R R R - - R V .. ..

Diplodonta orbella (Gould) R V .. C .. ..

Kellia laperousi (Deshayes) v V v v .. ..

Lasaea cistula Keen C R ..

Lasaea subviridus Dall C V .. R R ..

T r achy car dium cf. T. quadragenarium

(Conrad) R

Chione undatella simillima (Sowerby) .. V .. R .. .. V

"Chione" picta Dall V v

Protothaca staminea (Conrad) ACRARCCCRA

Irus lamellijer (Conrad) v v

Transenella tantilla (Gould) C v v c .. -

Tivela stultorum (Mawe) R C C C .. C

Saxidomus nuttalli Conrad v

A miantis callosa (Conrad) R V

Petricola californiensis Pilsbry and Lowe c

Petricola carditoides (Conrad) C C R C .. - V V V ..

Tellina bodegensis Hinds R V A

Macoma nasuta (Conrad) R V

Valentine — Pleistocene Faunas from Baja California 297

Table 2. — Continued

,

O

O

On

ir\

\o

00

r-s

Csl

Os

vO

\T>

<N

»— i

•— t

^■H

i—i

v©

VO

i-H

t^.

r-.

r-^

I~x

r-

rv.

irs

ro

fA

(N

fN

fN

<N

<N

r-4

ro

m

Macoma secta (Conrad)

Semele rupicola Dall

Cumingia californica Conrad

Donax gouldi Dall

Sanguinolaria nuttalli Conrad

Siliqua lucida (Conrad)

Siliqua patula (Dixon)

Spisula "dolabriformis Conrad" -

Spisula hemphilli Dall

Schizothaerus nuttalli (Conrad) A

Cryptomya californica (Conrad)

"Corbula" luteola Carpenter

Hiatella arctica (Linnaeus)

Z.irjaea pilsbryi Lowe

Pholadidea penita (Conrad) a

Scaphopoda
Dentalium neohexagonurn Pilsbry and Sharp... C
Gastropoda

Acteocina culcitella (Gould)

Trimusculus reticulata (Sowerby) C

Terebra pedroana Dall

Conus calijornicus Hinds C

Megasurcula carpenteriana (Gabb) C

Megasurcula carpenteriana tryoniana (Gabb) .... R
Pseudomelatoma penicillata semiinflata

Grant and Gale

Ophiodermella incisa fancherae (Dall) -

Ophiodermella ophioderma (Dall)

Clavus ci C. empyrosia (Dall)

Clavus hemphilli (Stearns)

"Mangelia" interlirata Stearns

"Mangelia" variegata Carpenter

Mitromorpha aspera (Carpenter) ...v

Mitromorpha jilosa (Carpenter)

Cancellaria cooperi Gabb

Olivella baetica Carpenter

Olivella biplicata (Sowerby) A

Olivella pedroana (Conrad) A

Hyalina californica (Tomlin)

Cystiscus jewetti (Carpenter)

Cystiscus regularis (Carpenter)

Cystiscus subtrigona (Carpenter) ..

Cypraeolina pyriformis (Carpenter)

Mitra catalinae (Dall)

.. R ..

.. .. V

.. C V

C .. ..

V
V

A A .... V R
R .. R V V V
.. .. R V .. V
.. .. R V V ..

R

R R? .. .. V R

V C

V ..

.. C?

c c

.. .. V .. V R

C A V C V ..

R R R

C .. C A S A

C R

C C
C ..

R
R

R ..

C V?

R V?

V ..
R V?

V ..

V R

V V

V ..

A

s

SARCSACS

c

V

..

..

V

—

R

V

..

..

..

—

V

..

V

..

..

—

..

V

V

..

..

..

R

V

V

V

V

._

R

R

V

V

298

San Diego Society of Natural History

Table 2. — Continued

-HoooNi/^^oor-NjNOv

Mitra idae Melvill

"Fusinus" luteopictus Dall

Harjordia cf. H. monksae (Dall)

Kelletia kelleti (Forbes) R

Macron lividus (A. Adams) ~

"Nassa" delosi Woodring

"Nassa" fossata (Gould) A

"Nassa" mendica Gould

"Nassa" mendica cooperi Forbes R

"Nassa" perpinguis Hinds R

Mitrella carinata (Hinds) R

Mitrella cannata gausapata (Gould)

Amphissa cf. A. undata (Carpenter)

Amphissa versicolor Dall

Aesopus cf. A. myrmecoon Dall

Ceratostoma nuttalli (Conrad)

Jaton festivus (Hinds)

Maxwellia gemma (Sowerby)

Maxwellia santarosana (Dall)

Ocinebra foveolata (Hinds) -

Ocinebra gracillima Stearns

Ocinebra interjossa Carpenter

Ocinebra interjossa beta (Dall)

Ocinebra poulsoni Carpenter

Nucella emarginata ostrina (Gould) R

Stramonita biserialis (Blainville)

Acanthina lugubris (Sowerby) R

Acanthina paucilirata (Stearns) -

Acanthina spirata (Blainville) C

Opalia chacei Strong

Opalia inscidpta Carpenter

Epitonium indianorum (Carpenter)

Epitonium tinctum (Carpenter)

Balds rutila (Carpenter)

Balds tbersites (Carpenter)

Turbonilla almo Dall and Bartsch

Turbonilla aresta Dall and Bartsch

Turbonilla buttoni Dall and Bartsch

Turbonilla tenuicula (Gould)

Turbonilla spp

Odostomia cf. O. donilla Dall and Bartsch

Odostomia cf. O. io Dall and Bartsch

Odostomia phanea Dall and Bartsch

Odostomia cf. O. phanella Dall and Bartsch

V

R
A
C
R
A
R
C
V
V

R
R

A

V R

V ..

C R

V .. V

s S V V c c

.. V V C R - -

C .. .. A R .. C

.. V V C C C C

R C C A C R -

.. C V R C R ..

.. V

.. .. V

R

.... V V

R

V V .. ..

V ..

.. R C V R R ..

.. V .. V .. V ..

V

R V R

.. .. V

.. R C C R C ..

C C C V C V C

A V .. C R .. C

.. V .. V .. ..

.. V

C R V V V?

V

V V .. R ..

V

.. .. V .. ..

.. .. V .. ..

V R V R V
R V .. V ..

.. V

V ..

.. V

V ..

Valentine — Pleistocene Faunas from Baja California 299

Table 2.— Continued

,

o

O

Ov

•TN

\©

oo

l\

rsi

Os

so

so

fS

l— H

f-H

»— t

*— i

SO

VO

»-H

1—4

l\

r^

IV

r\

r-s

r-s

t-H

u-\

ro

r<\

<N

<N

<N

<N

r-i

<N

CA

rA

Odostomia trachis Dall and Bartsch

Odostomia virginalis Dall and Bartsch....

Odostomia spp

Bwri'd calif ornica (Hinds)

Z^onaria spadicea (Swainson)

Pusula calif orniana (Gray)

Erato columbella Menke

Erato vitellina Hinds

Bittium interfossa Carpenter

Bittium purpureum Carpenter

Seila montereyensis Bartsch

Ceritbiopsis antefilosa Bartsch

Cerithiopsis cf. C. antemunda Bartsch

Cerithiopsis carpenteri Bartsch

Cerithiopsis diegensis Bartsch

Cerithiopsis spp

Triphora hemphilli (Bartsch)

Triphora cf. T. pedroana (Bartsch)

Nodulus kelseyi Bartsch

Amphithalamus tenuis Bartsch

Alvania acutelirata (Carpenter)

Alvania purpurea Dall

Rissoina cleo Bartsch

Turritella cooperi Carpenter

Aletes squamigerus Carpenter

Spiroglyphus lituella (Morch)?

Petaloconchus cf. P. complicatus Dall

Caecum calif ornicum Dall

Caecum licalum Bartsch

Fartulum hemphilli Bartsch

Fartulum occidentale Bartsch

Fartulum orcutti (Dall)

Littorina planaxis Philippi *

Littorina scutulata Gould

Lacuna carinata Gould

Lacuna unifasciata Carpenter

Barleeia haliotiphila Carpenter

Barleeia oldroydi Bartsch

Truncatella stimpsoni Stearns

Assiminea translucens (Carpenter)

Hipponix antiquatus (Linnaeus)

Hipponix tumens Carpenter

Crepidula adunca Sowerby

Crepidula coei Berry

v
R

C C A A

V .. V ..

V V C ..
R V .. ..
C R -. -

V V

R R R V .. ..

V V V

R

'.. V R V? ..

V R V V .. ..

.. V .. ..

V

V

R V .. ..

R V V V .. ..

V

V V

R

R

V .. ..

C V ..

R

.. R R?

C- .. RRCCCCC

R R R R R ..

V ..

C .... V V ..

V .. ..

A .... V R ..

R .. .. R .. ..

A .. V V A ..

C.. CRAARRA..
CCAARSASS-

R .. C C .. C

R R R V V ..

C .. R C R ..

V .... R .. ..

R

C R .. .. R ..

A .. .. C C C C C C ..

V R C V R ..

.. R R R V ..

R R V R R V ..

300

San Diego Society of Natural History

Table 2. — Continued

o

o

Ov

IT\

vO

00

r^

<N

0\

o

VO

rg

1-H

■— i

i— i

*— (

i— 1

Vf)

VO

»-H

1^

1^

i-^

IV

rv

r^

»— 1

IT\

H"\

CTl

<N

<N

<N

<N

«N

<N

cr\

fA

Crepidula nummaria Gould

Crepidula onyx Sowerby

Crepipatella lingulata (Gould)

Neverita reclusiana alta Arnold

Neverita ? sp

Euspira lewisi (Gould)

Velutina laevigata (Linnaeus)

Acmaea asmi (Middendorff)

Acmaea digitalis Eschscholtz

Acmaea insessa (Hinds)

Acmaea limatula Carpenter

Acmaea mitra Eschscholtz

Acmaea paleacea Gould

Acmaea pelta Eschscholtz

Acmaea pelta navicelloides Dall

Acmaea cf. A. persona Eschscholtz

Acmaea persona strigatella Carpenter

Acmaea scabra (Gould)

Acmaea scutum Eschscholtz

Lottia gigantea (Gray)

Tricolia cf . T. compta (Gould)

Tricolia pulloides (Carpenter)

Pomaulax undosus (Wood)

Homalopoma carpenteri (Pilsbry)

Norrisia norrisi (Sowerby)

Halistylus pupoideus (Carpenter)

Tegula aureotincta (Forbes)

Tegula brunnea (Philippi)

Tegula brunnea fluctuata (Dall)

Tegula funebralis (A. Adams)

Tegula gallina (Forbes)

Tegula gallina multifilosa (Stearns)

Tegula ligulata (Menke)

Tegula marcida (Gould)

Tegula marcida subsp. nov.?

Calliostoma doliarium (Holten)

Calliostoma gemmulatum Carpenter

Calliostoma ligatum (Gould)

Pupillaria parcipictus (Carpenter)

Vitrinella oldroydi Bartsch

Pseudorotella cf. P. bibbiana (Dall)

Pseudorotella invallata (Carpenter)

Pseudorotella supravallata (Carpenter)....
Haliotis cracherodi Leach

R .. .. R C

R

V

R C

R R R R

.. R V? ..

C R R R

.. A C V

c r a cr ..

V

c

C C R A ..

.. R .. R V

R R

R V

V

C V

.. R .... V
.. R .... A
R V .. V? ..

V R

C
C
C

R
V
V

c

V

c
c

V

R
V
V

R
R

R

R
V

V
R
R
V

V

.. .. c
.. .. c

c

V

c

R

V
R

R
V
R
V
V

S C R C C

C

R

C A

R R V

.. .. V

.. .. V

.. .. V

.. .. R

.. .. C

.. .. R

R
C
V

c
c

V

R

V V ..

c c c

c

R

.. V

.. c

V V

A A

R .. V C

.. V .. ..

.. V .. ..

.. R V ..

.. .. V ..

.. V V ..

V .. .. R

V V R C

Valentine — Pleistocene Faunas from Baja California 301

Table 2-

—Continued

O VO rs| •— • • •— 1 -H _ 1 ,—1 NO ^O

Haliotis rufescens Swainson v .

Halwtis sp R

Fissurella volcano Reeve C R C R C C

Megatebennus bimaculata (Dall) V .. C R R .. ..

Lucapinella callomarginata Dall V

Diodora aspera (Eschscholtz) R V R v v

Amphineura

Tonicella ci. T. lineata (Wood) R .. v ..

Nuttallina calif ornica (Reeve) V

Mopalia ciliata (Sowerby) v .. ..

Mopalia hind si (Sowerbv) R

Mo palia muscosa (Gould) R V v v

Mopalia ? sp

Cryptochiton stelleri (Middendorff) R

Stenoplax acrior (Carpenter) V .... v

Stenoplax conspicnus (Carpenter) V

Lepidozona pectimdatus (Carpenter) V V .. ..

Callistochiton a as sico status Pilsbry v R v

Callistochiton palmidatus mirabilis Pilsbry V .. v

Habitats and Suggested Depositional Environments
Faunas from seven of the localities are so similar in their ecologic charac-
ter that they may be discussed as a unit. The fauna at locality 3569, with 16
identified species, is too small for comparison; and the faunas from localities
2716 and 3162, discussed below, differ only quantitatively from the other
assemblages in their ecologic character.

In the seven similar faunas, between 40 and 50 per cent of the species at
each locality live today most commonly in shallow water along exposed coasts.
Many are essentially intertidal types usually living on rocks washed or sprayed
by waves; Mytilus calif ornianus, Nucella emarginata ostrina, Littorina planaxis,
L. scutulata, Lottia gigantea, several species of Aonaea and Tegula, and several
of the chitons represent this group. A few other forms live intertidally or
subtidally in sandy areas along exposed coasts, as Tivela stultorum, Amiantis
callosa, and Donax gouldi.

In these seven faunas, about 20 to 30 per cent of the species have a wide
ecological range today and are not uncommon along exposed rocky coasts in shal-
low water. These include Leptopecten latiauratus , Macoma secta, Conus cali-
fornicus, Olivella biplicata, and "Nassa" perpinguis. The remaining species,
about 30 per cent of the fauna at each locality, are those that live most commonly
in shallow waters of embayments or offshore below most wave action along
exposed coasts. Among this group are Nuculana taphria, Macoma nasuta,
Spisula hemphilli, Ocinebra poulsoni, Balds thersites, Bursa californica, and
Euspira lewisi.

302 San Diego Society of Natural History

The collections from locality 2716, the upper unit at Punta Descanso, and
from locality 3162, near the rear of the terrace at Rio Monro, contain fewer
quiet- water species (10 to 20 per cent) and more exposed-rocky-shore species
(60 to 70 per cent) than the faunas discussed above. Nearly the entire fauna
from locality 3569 is composed of shallow- water exposed coast and ecologically
ubiquitous species.

The preponderance of shore types at all localities suggests deposition
along a coast much like the present one, with alternating short rocky and sandy
stretches. Many specimens of quiet-water species, which probably were trans-
ported shoreward by storm waves, are broken and worn. All faunas probably
were deposited in less than 10 fathoms of water; localities 2716 and 3162 may
represent shore deposits.

Geographic Ranges and Suggested Temperatures

A total of 22 species and subspecies, from all localities combined, are not
known to live at present at the latitude of their Pleistocene occurrence. Of
these, 19 live only to the north and 3 live only to the south. They are listed
in table 3, together with the nearest known living occurrence of each. A sum-
mary of their fossil occurrence by localities is given in table 1.

The presence of a small northern shallow-water element at all localities sug-
gests that sea-surface temperatures were locally cooler than at present along
northwestern Baja California. However, the rare southern shallow-water ele-
ment suggests that water slightly warmer than at present was available at
least intermittently in the area.

CLIMATIC SIGNIFICANCE OF THE FAUNAS

A striking feature of late Pleistocene faunas from southern California
and Baja California is the contrast in thermal significance (as interpreted from
the present ranges of species) between exposed and protected shallow- water
faunas (Valentine 1954) . Assemblages dominated by exposed-coast or off-
shore species, such as those recorded here, commonly include northern, cool-
water species, whereas southward-ranging species are rare or absent. In contrast,
assemblages dominated by protected shallow-water species, especially assemblages
from ancient embayments, include many more warm-water than cool-water
species. Emerson (1956) reviewed several late Pleistocene faunas to illustrate
this point. This relation of temperature to habitat, particularly to degree of
protection or exposure, formed the basis of a recently suggested model of
Pleistocene marine climates in the region (Valentine 1955). It was proposed
that increased temperature contrasts — indicated by cool, exposed and warm,
protected faunas that are essentially contemporaneous — might be explained as
a result of increased oceanic and atmospheric circulation during glacial times.
The cool-water elements in exposed-coast faunas were attributed to increased
flow and lower temperatures of upwelling waters. Higher temperatures neces-
sary to support southern species may have been due to a general warming of
the ocean or to the introduction of warm southern water by northward-flowing
currents.

Valentine — Pleistocene Faunas from Baja California 303

Table 3. Nearest living occurrence of locally extinct species
Most range and habitat data are from Berry (1922), Keen (1937), Burch et al.
(1944-46), and U. C. L. A. collections.

Sp

ecies

Nearest known occurrence

Northern species
Glycymeris profunda (Dall)
Irus lamellifer (Conrad)
Siliqua patula (Dixon)
Mitromorpha aspera (Carpenter)
Cancellaria cooperi Gabb
Mitra idae Melvill
"Nassa" delosi Woodring
Ocinebra interfossa beta (Dall)
Opalia chacei Strong
Odostomia phanea Dall & Bartsch
Lacuna carinata Gould
Barleeia oldroydi Bartsch
Velutina laevigata (Linnaeus)
Tegula brunnea (Philippi)
Tegula brunnea fluctuata (Dall)
Calliostoma doliarium (Holten)
Callio stoma ligatum (Gould)
Cryptochiton stelleri (Middendorff)
Callistochiton palmulatus mirabilis Pilsbry

Southern species
Ostrea megodon Hartley
"Chione" picta Dall
Acanthina lugubris (Sowerby)

Off Redondo Beach
San Diego
Pismo Beach
Coronado Islands
Coronado Islands
Cortez Bank
Balboa Beach1
So. Coronado Is.
San Diego
San Diego
San Diego
Coronado Islands
Cayucos

Santa Barbara Is.
San Nicolas Is.
San Diego
San Diego
Channel Islands
San Diego

Scammons Lagoon2
Magdalena Bay
Todos Santos Bay

^Record by E. P. Chace, personal communication, 1956.

2 Or perhaps farther south. See Woodring, Bramlette, and Kew 1946, p. 81.

DESCRIPTION OF FOSSIL LOCALITIES

Locality numbers in the following paragraphs are those of the Department
of Geology, University of California, Los Angeles.

The two base maps available (fig. 1) are very generalized and do not
always agree. Localities may most easily be found by reference to local coastal
topography. The kilometer signs, although quite useful today, may be moved
at a future date.

Locality no. 2715. — Unconsolidated pebble conglomerate with sand mat-
rix, 3 feet thick, extending about 40 yards along the sea cliff approximately 5
miles south of the radio tower at Rosarito Beach, and about 0.3 mile north of
kilometer 35 sign on Highway 1, Baja California, Mexico. Collected by Cun-
ningham and Valentine, February, 1952.

Locality no. 2716. — Unconsolidated rubble with matrix of alluvium and
broken shell, 3 feet thick, directly overlying pebbly sand at locality 2715.
Collected by Cunningham and Valentine, February, 1952.

304 San Diego Society of Natural History

Locality no. 2717. — Sandy rubble exposed in sea cliff approximately
6.4 miles south of the radio tower at Rosarito Beach, and about 0.3 mile south
of kilometer 36 sign on Highway 1, Baja California, Mexico. Collected by
Cunningham and Valentine, February, 1952.

Locality no. 2718. — Sandy rubble exposed in sea cliff approximately
5.7 miles south of the radio tower at Rosarito Beach, at kilometer 36 sign on
Highway 1, Baja California, Mexico. Collected by Cunningham and Valentine,
February, 1952.

Locality no. 2719. — Unconsolidated pebble conglomerate exposed in sea
cliff approximately 2.2 miles south of the radio tower at Rosarito Beach, at
kilometer 30 sign on Highway 1, Baja California, Mexico. Collected by Valen-
tine, December, 1952.

Locality no. 2720. — Sandy rubble exposed in sea cliff at the second
small re-entrant in first sea cliffs south of the radio tower at Rosarito Beach,
Baja California, Mexico. Collected by Valentine, December, 1952.

Locality no. 3160. — Unconsolidated cobble conglomerate exposed in
low sea cliff approximately 5.2 miles north of the radio tower at Rosarito
Beach, Baja California, Mexico. Bearing from locality to south end of South
Coronado Island, 263° true. Collected by Cunningham and Valentine, January
1952.

Locality no. 3161. — Unconsolidated cobble conglomerate exposed in sea
cliff approximately 8 miles north of the radio tower at Rosarito Beach, Baja
California, Mexico. Bearing from locality to south end of South Coronado
Island, 238° true. Collected by Cunningham and Valentine, January, 1952.

Locality no. 3162. — Unconsolidated pebble conglomerate exposed in
road cut of Highway 1, 50 yards south of the bridge across Rio Morro, Baja
California, Mexico. Collected by Valentine, November, 1952.

Locality no. 3569. — Unconsolidated cobble conglomerate exposed in
gully 20 feet inland from sea cliff, approximately 4V2 miles south of the mouth
of Rio Morro, and about 150 yards north of kilometer 45 sign on Highway 1,
Baja California, Mexico. Collected by Valentine, November, 1952.

ACKNOWLEDGMENTS

It is a pleasure to acknowledge the many courtesies extended by Pro-
fessors Daniel I. Axelrod and U. S. Grant, IV, University of California, Los
Angeles, and by Professor Carl L. Hubbs, Scripps Institution of Oceanography,
La Jolla, during the course of this study. Gratitude is also expressed to Mr.
A. A. Allanson, Dr. G. W. Brainerd, Mr. J. S. Cunningham, Dr. Leo G.
Hertlein, Dr. C. A. Nelson, Mr. Peter U. Rodda, and Mrs. L. E. Saul for
their aid during the preparation of this report. The index map was drafted
by Mrs. Opal L. Kurtz.

Acknowledgment is due the Humble Oil and Refining Company for
financial support of this investigation.

Valentine — Pleistocene Faunas from Baja California 305

LITERATURE CITED

Anderson, Charles A.

1950 1940 E. W. Scripps cruise to the Gulf of California. Part 1. Geo-
logy of islands and neighboring land areas. Geol. Soc. Am. Mem.
43 (1): 1-53, figs. 1-19, pis. 1-3.

Anonymous

1950 Recommendations of Commission appointed to advise on the defini-
tion of the Pliocene- Pleistocene boundary. Int. Geol. Congr. Rep.
18th Session (9) : 6.

Beale, Carl H.

1948 Reconnaissance of the geology and oil possibilities of Baja California,
Mexico. Geol. Soc. Am. Mem. 31: 1-138, pis. 1-11.

Berry, S. Stillman

1922 Fossil chitons of western North America. Proc. Calif. Acad. ser.
4. 11: 399-526, figs. 1-11, pis. 1-16.

Burch, John Q. [ed.] et al.
1944-46 Distributional list of the west American marine mollusks from San
Diego, California to the Polar Sea. Min. Conch. Club So. Calif.
33-63: pagination by issue, pis. 1-3.

Dall, William Healey

1890 Contributions to the Tertiary fauna of Florida, with especial refer-
ence to the Miocene Silex-beds of Tampa and the Pliocene beds of
the Caloosahatchie River. Part 1. Pulmonate, opisthobranchiate and
orthodont gastropods. Trans. Wagner Free Inst. Sci. 3: 1-200,
pis. 1-12.

1900 Idem. Part 5. Teleodesmacea : Solen to Diplodonta. Ibid. 3:
949-1218, pis. 36-47.

1903 Idem. Part 6. Concluding the work. Ibid. 3: 1219-1654, pis. 48-60

Ellis, Arthur J., and Charles H. Lee

1919 Geology and ground waters of the western part of San Diego
County, California. U.S. Geol. Surv. Water-Sup. Pap. 446: 1-321,
pis. 1-47.

Emerson, William K.

1956 Pleistocene invertebrates from Punta China, Baja California, Mexi-
co, with remarks on the composition of the Pacific Coast Quater-
nary faunas. Bull. Am. Mus. Nat. Hist. Ill: 313-342, pis. 22-23.

■ , and Warren O. Addicott

1953 A Pleistocene invertebrate fauna from the southwest corner of
San Diego County, California. Trans. San Diego Soc. Nat. Hist.
11: 429-444.

306

San Diego Society of Natural History

Hertlein, Leo George, and U. S. Grant, IV

1944 The geology and paleontology of the marine Pliocene of San Diego,
California. Part 1, Geology. Mem. San Diego Soc. Nat. Hist.
2: 1-72, figs. 1-6, pis. 1-18.

Keen, A. Myra

1937 An abridged check list and bibliography of west North American
marine Mollusca. 1-87, figs. 1-3. Stanford Univ. Press.

Valentine, James W.

1954 Ecologic requirements and depositional environments of Pleistocene
molluscan faunas from southern and Baja California [abst.] Jour.
Paleo. 28: 881.

1955 Upwelling and thermally anomalous Pacific Coast Pleistocene mol-
luscan faunas. Amer. Jour. Sci. 253: 462-474, figs. 1-3.

Woodring, W. P., M. N. Bramlette, and W. S. W. Kew

1946 Geology and paleontology of Palos Verdes Hills, California. U. S.
Geol. Surv. Prof. Pap. 207: 1-145, figs. 1-16, pis. 1-37.

Fig. 2. Fossiliferous terrace deposits on the north side of
Punta Descanso (localities 2715, 2716), looking southward.

Marine conglomerates and pebbly sands overlie Pliocene (?) basalt, on which
the terrace platform is eroded, and are in turn overlain by fossiliferous alluvial rubble.
A narrow bench is developed on the slightly more resistant sands.

Valentine — Pleistocene Faunas from Baja California 307

ftp • -;. r*y*.*y&-> *^

Fig. 3. Terrace deposits in upper central portion of fig. 2.

Marine sediments (by the mattock) have a light-colored sandy matrix, whereas the
matrix of the upper unit (by the hammer) is dark unsorted alluvium and rubble.

-v

Q/j

■- y~.

'->*

sat -'■• wfig

-SS

I M

Fig. 4. Terrace platform overlain by unsorted fossiliferous deposit,
exposed in road cut just south of Rio Morro (locality 3162).

308

San Diego Society of Natural History

Fig. 5. Detail of fossiliferous horizon in fig. 4, show-
ing massive unsorted sediments overlying the basalt.

Fig. 6. Fossiliferous lens of marine terrace sediment, left foreground, exposed in
gully 20 feet inland from sea cliff, about 41/2 miles south of Rio Morro
(locality 3569).

OCI 2

univei

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OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY

Vol. XII, No. 17, pp. 309, 310, fig. 1 September 25, 1957

TYPE MATERIAL OF

EUCALODIUM ORCUTTI DALL

(GASTROPODA: PULMONATA)

FROM OAXACA, MEXICO

BY

Robert J. Drake

Department of Zoology, University of Arizona

Eucalodium (Atusospira) orcutti was described by Dall (1910) from
specimens collected by Charles Russell Orcutt "at some hot springs near Rio
Verde, Oaxaca." The type is No. 212319 in the United States National
Museum. Orcutt (1915) told of making botanical collections in northern and
eastern Oaxaca in February and April of 1910. It is probable that the speci-
mens were collected near the river and town of Rio Verde in the eastern part
of the state, along the Isthmian Railroad. There is another Rio Verde in
western Oaxaca but there is no indication that he visited that area, at least in
that year.

After Orcutt's death, his large cabinet of shells was deposited in the San
Diego Museum of Natural History. The collections include, as No. 7615,
ten specimens of Eucalodium orcutti from "Rio Verde, Oaxaca." Since Dall
cited specimens in the Orcutt collection in addition to the type, and since there
are no other specimens of the species in the Museum collection, it is probable,
despite the less precise locality designation, that this set is part of the type
collection.

Since Eucalodium orcutti apparently has not been illustrated, two speci-
mens are shown in fig. 1. For the larger shell illustrated, which is the largest
in the lot, measurements are as follows: height, 54 mm.; width of the spire,
4 mm.; greatest width, 12 mm.; height of the aperture, 7 mm.; width of the
aperture, 8 mm. There are 2OV2 whorls, with the decollation at 9V2 whorls. Of
the other nine shells in the lot, seven are decollated at 9Vz whorls, one at 8V2
whorls, and one at IOV2 whorls.

Charles Russell Orcutt (1864-1929) was a professional collector of natural
history materials (Jepson 1929). From the early 1880's to the 1920's he
collected widely in the southwestern United States and in Mexico. The extent
of his contribution to the malacology of Mexico has never been assessed.

Study to locate museum collections of Mexican nonmarine mollusks was
supported by American Philosophical Society grants (Penrose Fund) in 1951

310

San Diego Society of Natural History

and 1952. Research at the San Diego Museum of Natural History was made
possible by the Bache Fund of the National Academy of Sciences in 1954.
That study included the urocoptid portions of the Orcutt and H. N. Lowe
collections permanently deposited there. Help with references and localities
was kindly given by Drs. Reid Moran and Carl L. Hubbs.

LITERATURE CITED

Dall, William Healey

1910 Two new Mexican land shells. The Nautilus 24: 34-36.
Jepson, Willis Linn

1929 Charles Russell Orcutt, natural history collector. Madrono 1 :
273, 274.

Orcutt, Charles Russell

1915 A botanical study of Mexico. West Am. Sci. 19: 2-6.

Fig. 1. Eucalodium orcutti Dall

Shells from Rio Verde, Oaxaca; No. 7615 in the collection of the San Diego Museum
of Natural History; approximately Xla/2. Photograph by Robert J. Drake.

; .

TRANSACTIONS

OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY
Vol. XII, No. 18, pp. 311-318, figs. 1-4 October 16, 1958

THE NORTH AMERICAN SPECIES OF HESPERUS FAUVEL,

WITH DESCRIPTIONS OF TWO NEW SPECIES

(COLEOPTERA; STAPHYLINIDAE)

BY

Ian Moore

Associate in Entomology, San Diego Museum of Natural History

The genus Hesperus was based by Fauvel (1874) on the European species
Stapbylinus rujipennis Gravenhorst. He included in the genus two conspicuous
and well-known eastern American species, Stapbylinus baltimorensis Graven-
horst (1802) and S. apicialis Say (1834), both of which had previously been
removed to Philontbus. It is surprising at this late date to find two new large
and even more conspicuous species in the North American fauna, one from
Ohio and the other from Arizona. This discovery would seem to indicate
either that the new species are quite rare or that the American staphylinid
fauna is poorly known. The two new species are so distinctive that, although
each is represented in our material by only a single specimen, I do not hesitate
to describe them.

Hesperus is a widespread genus with many tropical species but only a few
in temperate regions. Nearly one hundred species have been described from
tropical Africa, Asia, the East Indies, and Australia. Of the four palearctic
species, only one is found in Europe. Although only six species are known
from tropical America, two of these from Guatemala and four from South
America, it is likely that many more will be found when these regions are fully
explored. In 1936 Scheerpeltz published a key to the six Neotropical and two
Nearctic species.

The genus Hesperus can be distinguished from all other genera of the
North American Staphylininae except Belonuchus by the elongate, slender
palpi, with the last segment of the maxillary palpus at least five times as long
as wide and slightly truncate at the tip. Hesperus differs from Belonuchus
in lacking the row of strong setae (often called "spines") that in Belonuchus
usually arm the posterior femora internally. The North American species of
Hesperus further differ from those of Belonuchus in having the first four seg-
ments of the anterior tarsus strongly dilated in each sex, whereas in the latter
genus the anterior tarsus is slender (in females) or only slightly dilated (in
males) .

312 San Diego Society of Natural History

Hesperus arizonicus sp. nov.

Description of holotype (male) . — Color largely ferruginous; fourth
to tenth antennomeres and apex of fifth abdominal segment dusky; most of
basal half of each elytron, except for the suture and a narrow basal strip,
covered by a large yellow spot; entire sixth and bases of first five abdominal
segments yellow. Head much broader than long, slightly broader than pro-
notum; dorsal surface with punctures coarse and very close, generally separated
by less than one-half of their width except in a small central area anteriorly,
almost coalescing at the sides, the ground sculpture a very feeble micro-reticula-
tion; ventral surface impunctate, but with a feeble micro-reticulation throughout.
Antennae shorter than head and pronotum, the first segment as long as next
three together, the second twice as long as wide, the third three times as long
as wide and 1.5 times as long as second, the remaining segments gradually
shorter but narrower after the ninth so that none of them is transverse. Pro-
notum subquadrate, a little longer than wide, the apex gently arcuate, the
anterior angles rounded, the sides thence gently arcuate and slightly sinuate
before the rounded posterior angles, the base arcuate; surface as coarsely punc-
tate as the head but more sparsely, the punctures generally separated by about
their width, a central strip impunctate and impressed in a broad irregular groove
throughout most of the length, the ground sculpture a feeble micro-reticulation.
Elytra quadrate, wider and longer than pronotum, the humeri broadly rounded,
the sides straight and hardly divergent to the narrowly rounded apical angles,
the apex conjointly emarginate in a gentle arc in central three-fifths; surface
finely, rather roughly, and moderately closely punctate. Abdomen narrower
than elytra, tapering only slightly to apex; dorsal and ventral surfaces about
as finely punctate as elytra but more sparsely and not so roughly; without
ground sculpture; last sternite emarginate in its central third in a gentle arc
with rounded outer angles, the emargination wider than deep. Length 15 mm.

Female. — Unknown.

Type. — The unique type collected at Patagonia, Arizona, on January 4,
1938, by Dr. E. C. Van Dyke. It is deposited in the California Academy of
Sciences, San Francisco.

Diagnosis. — Among the North American species this large colorful
insect is very distinct in its markings and color, and particularly in the puncta-
tion of the head, which is most like that of H. stehr'i, but with even larger and
more crowded punctures. This species differs from the ferruginous Neotropical
species by the yellow markings of the elytra and abdomen and by its larger size.

Moore — North American Hesperus 313

Hesperus stehri sp. nov.

Description of holotype (male) . — Color black except eleventh anten-
nomere and tarsi, which are dark rufous, and apex of abdomen, which is bright
rufous. Head quadrate, broader than long, wider than pronotum, densely
punctate dorsally except in a very narrow central strip, the punctures of disk
separated by little more than their diameters, becoming closer toward the apex
and sides until they nearly coalesce in places; dorsally without ground sculpture;
ventrally with a few scattered punctures and a ground sculpture of very fine
diagonal lines near the gular sutures. Antennae shorter than head and pro-
notum, the first segment widest, almost as long as next three together, the second
about twice as long as wide, the third not quite twice as long as second and
about as wide, the fourth and fifth slightly elongate, the sixth to tenth about
as wide as long, the eleventh very little longer than wide. Pronotum about as
wide as long, narrower than but about as long as head; apex nearly straight,
but slightly sinuate before anterior angles, the angles narrowly rounded, the
sides arcuate in anterior half, thence sinuate to rounded posterior angles, the
base slightly arcuate; punctures sparse and fine except in a narrow impunctate
central line, closer toward sides; without ground sculpture. Elytra about as
long as wide conjointly, the sides straight from broadly rounded humeri to
narrowly rounded apical angles, the apex shallowly emarginate conjointly in
central three-fifths; punctures fine and a little closer than those of pronotum;
without ground sculpture. Abdomen narrower than elytra, tapering to apex
which is about one-half the basal width; punctures dorsally about as close as on
elytra but coarser at base; ventrally a little more closely punctured; without
ground sculpture; last sternite apically emarginate in its central fourth, the
emargination evenly arcuate with rounded outer angles, about as wide as deep.
Length 15 mm.

Female. — Unknown.

Type. — A single specimen collected by Dr. William C. Stehr at Ash
Cave, Hocking County, Ohio on May 5, 1934. With the kind permission of
Dr. Stehr, the type has been deposited in the California Academy of Sciences,
San Francisco.

Diagnosis. — This species is colored like H. apicialis but is larger, with
a wider head, differently shaped pronotum, relatively wider elytra, and narrower
abdomen. It is very distinct in the close set punctures of the head and the lack
of ground sculpture of the head and pronotum. It differs from all of the
Neotropical species in its larger size and in its color. It is a pleasure to name
it for Dr. William C. Stehr, who collected it.

3 14 San Diego Society of Natural History

Hesperus baltimorensis (Gravenhorst)

Staphylinus baltimorensis Gravenhorst 1802, p. 163; 1806, pp. 112, 122; Say
1834, p. 451.

Philonthus baltimorensis Erichson 1840, p. 503; Gemminger 8C Harold 1868,
p. 585; Horn 1884, p. 220.

Hesperus baltimorensis Fauvel 1874a, p. 200; 1874b, p. 426; Blatchley 1910,
p. 390; Bernhauer & Schubert 1914, p. 364; Scheerpeltz 1936, p. 490.

Description. — Color: head and thorax black; elytra, abdomen and legs
dark rufous. Head subquadrate with the basal angles so broadly rounded as
to be obsolete; dorsally very sparsely punctate except on the impunctate central
area, the punctures separated by four to five times their diameters; ground
sculpture of fine obsolescent strigulose lines; ventrally almost impunctate, with
somewhat stronger ground sculpture. Antennae shorter than head and pro-
notum, the first segment a little longer than second and third together, the
second almost twice as long as wide, the third 1.5 times as long as second, the
fourth to sixth quadrate, the seventh to tenth transverse, the eleventh a little
longer than wide. Pronotum subquadrate, narrower and longer than head, one-
third longer than wide, the anterior angles narrowly rounded, the sides gently
arcuate and very slightly sinuate before the narrowly rounded basal angles, the
base strongly arcuate; punctures sparse and not coarse; ground sculpture almost
effaced. Elytra wider and slightly longer than pronotum, conjointly about as
long as wide, the sides straight from the broadly rounded humeri to the rounded
apical angles, the apex conjointly emarginate in a gentle arc in the central three-
fifths; surface rather roughly punctate, a little more coarsely so than the
pronotum and twice as closely, without ground sculpture. Abdomen narrower
at base than elytra, tapering to apex to about half the basal width, the puncta-
tion as on elytra; ventral surface similar to dorsal. Length 10 to 13 mm.

Male. - — ■ Last sternite apically emarginate in its central third, the emargina-
tion evenly arcuate with narrowly rounded outer angles, about as deep as wide.
Head a little wider than long, a little wider than pronotum.

Female. — Last sternite entire. Head not wider than long, not wider than
pronotum.

Type. — "America septentr." Probably in the Zoologische Museum,
Berlin.

Distribution. — Has been recorded from the middle and southern states
and from Indiana. I have seen specimens from New York, Alabama and
Missouri.

Diagnosis. — This species differs in color from the other North American
species. The head is even more sparsely punctate than that of H. apicialis and
the antennae are more robust.

Moore — North American Hesperus 3 1 5

Hesperus apicialis (Say)

Staphylinus apicialis Say 1834, p. 451; 1869, p. 566.

Philonthus haematurus Erichson 1840, p. 504.

Philonthus apicalis Erichson 1840, p. 931; Horn 1884, p. 220.

Philonthus apicialis Gemminger 6C Harold 1868, p. 585.

Hesperus apicalis Fauvel 1874a, p. 200; 1874b, p. 426; Blatchley 1910, p. 390;

Leng 1920, p. 108; Scheerpeltz 1936, p. 490.
Hesperus apicialis Bernhauer & Schubert 1914, p. 364. [In this work Horn is

erroneously credited with the spelling "apicialis".']

Description. — Color black except for eleventh antennomere and tarsi,
which are dark rufous, and last two or three abdominal segments, which are
dark or bright rufous. Head subquadrate, with posterior angles very broadly
rounded, the disk broader than long, slightly wider than pronotum; sparsely
punctate dorsally, the punctures not very coarse, separated by about three times
their diameter, a little more closely punctate toward sides; with a ground sculp-
ture of very fine obsolescent strigulose lines; ventral surface much more sparsely
punctate, but ground sculpture much stronger, particularly toward gular sutures..
Antennae shorter than head and pronotum, the first segment thickest, as long
as next two combined, the second shorter than third, twice as long as wide, the
third three times as long as wide, as long as next two combined, the fourth to
eighth just perceptibly elongate, the ninth to eleventh more slender and hence
relatively more elongate. Pronotum one-third longer than wide, longer and
narrower than head, the anterior angles broadly rounded into the sides, thence
very gradually arcuate to the broadly rounded posterior angles, the base evenly
arcuate, the surface punctation and sculpture very similar to those of dorsal
surface of head, with a central impunctate strip the entire length. Elytra con-
jointly a little wider than head, a Iitde longer than wide, the humeral angles
broadly rounded, the sides nearly straight and slightly diverging to near the
rounded apical angles, the apices shallowly emarginate conjointly in central
three-fifths; surface twice as densely punctate as head but somewhat more finely
so, the punctures becoming a little closer and somewhat aspirate laterally, with-
out ground sculpture. Abdomen narrower than elytra, gradually tapering to
apex, which is about one-half as wide as base; dorsal surface with punctures of
basal segments closer than those of head but sparser than those of elytra, grad-
ually sparser apically until no closer than on head, the punctures aspirate and
elongate; ground sculpture lacking; ventral surface similarly punctate except for
closer, coarser punctation at bases of anterior segments, with traces of strigulose
ground sculpture basally. Length 9 to 1 1 mm.

Male. — Last sternite apically emarginate in its central fourth, the emargi-
nation evenly arcuate, with rounded outer angles, about as deep as wide. Head
wider than long, slightly wider than pronotum.

Female. — Last sternite entire. Head hardly wider than pronotum, very
little wider than long.

316 San Diego Society of Natural History

Type. — "Inhab. United States." The holotype has been lost.

Distribution. — This species has been recorded from Canada to Georgia
and from Indiana. I have seen specimens from Ohio, Missouri and Alabama.

Diagnosis. — In color this species resembles only H. stebri among the
North American species. H. stehri is larger and has a much more closely
punctate head without ground sculpture.

It should be noted that Erichson (1840, p. 931) misspelled the name of
this species and that his misspelling has been used by almost all subsequent
workers.

Key to the North American Species of Hesperus

la. Head closely punctate, the punctures mostly separated by their own width
or less.

2a. Head, pronotum, and elytra ferruginous H. arizonicus

2b. Head, pronotum, and elytra black H. stehri

lb. Head sparsely punctate, the punctures mostly separated by more than twice
their width.

3a. Elytra and abdomen rufous H. baltimorensis

3b. Elytra and base of abdomen black H. apicialis

Acknowledgments

In connection with this paper, I am particularly indebted to the following:
Mr. Hugh B. Leach of the California Academy of Sciences for the opportunity
to study material in their collections (from which came one of the new species) ,
and many other favors; Dr. William C. Stehr of the University of Ohio for
allowing me to study material collected by him, among which was the other new
species; Dr. George E. Lindsay, Director, Mr. Charles F. Harbison, Curator of
Insects, and Mrs. Mildred Meeder, Librarian, San Diego Museum of Natural
History, for use of the facilities of that institution and for encouragement and
assistance on many occasions.

Literature Cited

Bernhauer, Max, and Karl Schubert

1914 Coleopterorum catalogus . . . pars 57, Staphylinidae IV. 289-408.
Berlin.

Blatchley, Willis Stanley

1910 An illustrated descriptive catalogue of the Coleoptera or beetles . . .
[of} Indiana. 1-1386. Nature Publishing Co., Indianapolis.

Erichson, Wilhelm Ferdinand

1840 Genera et species staphylinorum insectorum coleopterorum familiae
[part 2]. 401-954. Berlin.

Moore — North American Hesperus 3 17

Fauvel, Charles Alphonse Albert

1874a Faune gallo-rhenane ou species des insectes qui habitant la France,
la Belgique, la Hollande, le Luxembourg, la Prusse rhenane, la
Nassau et la Valais avec tableaux synoptiques et planches gravees.
Bull. Soc. Linn. Normandie ser. 2. 8:167-340.

1874b The same issued separately at Caen as "vol. 3, livre 5": 391-544.

Gemminger, Max, and Edgar von Harold

1868 Catalogus coleopterorum hucusque descriptorum synonymicus et
systematicus. 2: 425-752. Monachii.

Gravenhorst, Johann Ludwig Christian

1802 Coleoptera microptera brunsvicensia nee non exoticorum quotquot
exstant in collectionibus entomologorum brunsvicensium in genera
familias et species distribuit. i-lxvi, 1-206. Brunsvigae.

1806 Monographia coleopterorum micropterorum. 1-236. Gottingae.

Horn, George Henry

1884 Synopsis of the Philonthi of boreal America. Trans. Am. Ent. Soc.
11: 177-244.

Leng, Charles William

1920 Catalogue of the Coleoptera of North America, north of Mexico,
i-x, 1-470. Mount Vernon.

Say, Thomas

1834 Descriptions of new North American insects, and observations on
some already described. Trans. Am. Philos. Soc. 4: 409-470.

1869 American entomology. A description of the insects of North
America. Edited by John Lawrence LeConte. 2: i-iv, 1-814. New
York.

SCHEERPELTZ, OTTO

1936 Die von Prof. Dr. H. Eidmann gelegentlich seiner im Jahre 1933
nach Brasilien unternommenen Studienreise aufgesammelten Staphy-
liniden. I. Die in dem Nestern von Atta sexdens L. aufgefundenen
Staphyliniden, nebst einigen Bemerkungen iiber die Gattung Scari-
phaeus Er. Arch. Naturges. ser. 2. 5:483-540.

318

San Diego Society of Natural History

H. STEHRI

H BALTIMORENSIS

H. APICIALIS

i;

TRANSACTIONS

OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY

Volume XII, No. 19, pp. 319-332, fig. 1

THE MARINE MOLLUSCAN FAUNA OF

GUADALUPE ISLAND, MEXICO

Volume XII, No. 20, pp. 333, 334, fig. 1

A NEW MOLLUSK FROM SAN FELIPE,
BAJA CALIFORNIA

BY

E. P. CHACE

Curator of Conchology, San Diego Museum of Natural History

SAN DIEGO, CALIFORNIA

Printed for the Society

October 16, 1958

Fig. 1. Ocenebra seftoni sp. nov. Holotype (left) and paratype.
Millimeter scale at right.

THE MARINE MOLLUSCAN FAUNA OF
GUADALUPE ISLAND, MEXICO

BY

E. P. Chace

CONTENTS

Introduction 321

Faunal Relationships 322

List of Mollusks and Brachiopods 323

A New Species of Gastropod 331

Literature Cited 332

INTRODUCTION

Guadalupe Island is 250 statute miles south-southwest of San Diego and
162 miles off the coast of Baja California. The island is 22 miles long and
4 to 6 miles wide. The highest point, 4257 feet above sea level, rises some
16,000 feet above the floor of the ocean. Thus the island is surrounded by
deep water.

The part of the island above sea level is volcanic in origin. Strong (1954)
suggested from the character of the lava and the amount of disintegration that
its geologic age might be Pliocene. Johnson (1953) reported a small fossil
marine fauna probably of Upper Tertiary or Quaternary age. In 1957 Dr.
Carl L. Hubbs found exposures of Pleistocene fossils near the high-tide line
at several points around the island, and in December of that year he and the
writer collected about 50 pounds of shells and matrix at three localities. This
material, representing a warm- water fauna, is under study.

Strong and Hanna (1930) listed 87 species of marine mollusks from
Guadalupe Island, collected mainly by an expedition of the California Academy
of Sciences in 1925. Strong (1954) increased the number to 116 species on
the basis of a collection of 82 species by M. Woodbridge Williams on an
expedition of the Scripps Institution of Oceanography in 1946. Pilsbry (1927)
treated the land and freshwater mollusks of the island.

Since 1945 Dr. Hubbs and his associates from the Scripps Institution
have collected mollusks on several trips to the island. Some of this material
was collected in the littoral zone, some was dredged, and some was brought
up by divers. Most of this material has come to the San Diego Museum of
Natural History, and it has now been identified except for some microscopic
shells. Since this material includes 77 species not reported from the island
before, it seems worth while to publish a new list.

322 San Diego Society of Natural History

FAUNAL RELATIONSHIPS

Since there are no protected bays on Guadalupe Island, the only littoral
species are those of exposed habitats, such as Littorina, Acmaea, Lottia, and
Haliotis. Most of the species listed are from deeper water.

Many species are represented in the collections by only one or two speci-
mens each. Several species were found only in one dredge haul, which happened
to hit a mud pocket; these include Lima subauriculata, Leiomya scaber, Cuspi-
daria pectinata, Thracia squamosa, and Crenella megas. And each big collection
has added several species to the list. These facts suggest that sampling has
been somewhat haphazard and that further collecting may add many species.
Nevertheless, since eight collecting trips have yielded only 193 species of
mollusks, the marine molluscan fauna of Guadalupe Island may be character-
ized as scanty.

Of the 193 species of marine mollusks known from Guadalupe Island,
149 species, or 77.2 percent, belong to the Southern California Fauna, though
some of these extend as far south as Cedros Island or beyond. Twenty-one of
these, or 10.9 percent, are not otherwise known south of San Diego. Thirty-
four species or 17.6 percent, belong to the Panamic Fauna, occurring on the
mainland between San Diego and Panama but apparently not north of San
Diego. Three of these are known only from Guadalupe Island and Panama:
Bursa calcipicta Dall, Liotia heitni Strong 8C Hanna, and Crenella megas Dall.
(Distributional data are mostly from Keen [1937].)

As with the other terrestrial and marine biota of Guadalupe Island, the
mollusks show some endemism. Ten species, or 5.2 percent of the marine
molluscan fauna, are not known to occur elsewhere. These are:
Ocenebra seftoni Chace Rissoina willetti Strong

Diastema slevini Strong Rissoella bakeri Strong

Alabina jordani Strong Astraea guadalupeana Berry

Rissoina guadalupensis Strong Haliotis californiensis Swainson

Rissoina lowei Strong Glycymeris guadalupensis Strong

Only two species of brachiopods have been collected at Guadalupe Island.
Both occur also in southern California.

Scharff (1911) believed that Guadalupe Island and the other islands off
the west coast of North America were remnants of a large land mass reaching
in the distant past from Alaska to Peru and extending a considerable distance
west of the present coastline. He claimed that the flora and fauna of the
islands supported this hypothesis. It is now generally believed, however, that
Guadalupe Island is an oceanic island and that its biota has resulted from
fortuitous dispersal. Since many of the molluscs are kelp dwellers or are as-
sociated with kelp, Strong and Hanna (1930) believed that much of the
molluscan fauna had arrived on drifting kelp from the islands of southern
California.

Chace — Marine Mollusks of Guadalupe Island 323

LIST OF MOLLUSKS AND BRACHIOPODS

With one or two exceptions, the systematic arrangement of this list is
that of Strong (1954). Trips on which the various species were collected are
indicated as follows — S: California Academy of Sciences in 1925; W: M.
Woodbridge Williams in 1946; 50, 54, 55, 56, and 57: Scripps Institution in
1950, 1954, 1955, 1956, and 1957. Notes as to abundance are based only on
the collections of 1950 to 1957.

Gastropoda
Umbraculidae

Tylodina fungina Gabb (W, 57) . Two specimens in dredgings.
Umbraculum ovale Carpenter (57) . Two specimens in dredgings.

ACTEONIDAE

Acteon punctocaelata (Carpenter) (54). Several specimens.

ACTEOCINIDAE

Acteocina angustior Baker 8C Hanna (S, W, 57). A few specimens.
Acteocina magdalenensis Dall (50, 57). A few specimens each trip.
Acteocina planata (Carpenter) (54). Nine specimens in dredgings.
Retusa harpa (Dall) (S, 50). Seven specimens in dredgings.

SCAPHANDRIDAE

Cylichna attonsa Carpenter (50) . Eight specimens.

Akeratidae
Haminoea angelensis Baker & Hanna (54). One specimen.

SlPHONARIIDAE

Williamia peltoides (Carpenter) (S, W, 55). One specimen.

Conidae
Conus calif ornicus Hinds (50, 54, 57) . Apparently fairly common.

TURRIDAE

Kurtzia beta (Dall) (S).

Mangelia barbarensis Oldroyd (54) . Fifteen specimens in dredgings.

Mangelia interlirata Stearns (W).

Mitromorpha crassaspera Grant & Gale (50). Three specimens.

Mitromorpha filosa (Carpenter) (W, 50, 54).

Philbertia crystallina (Gabb) (50) .

324 San Diego Society of Natural History

Marginellidae

Cypreolina pyriformis (Carpenter) (S, W, 50, 54) .

Cystiscus jewettii (Carpenter) (S, W, 57) .

Cystiscus minor (C. B. Adams) (54). One specimen in dredgings.

Cystiscus polttulus (Dall) (S, W, 54) .

Hyalina calif ornica (Tomlin) (S, W, 50, 57) .

MlTRIDAE

Mitra catalinae Dall (S) .

Mitra fultoni E. A. Smith (54) . One specimen in dredgings.

Fasciolariidae
Fusinus kobelti (Dall) (54, 57). One specimen each trip.

Buccinidae
Cantharus lugubris (C. B. Adams) (54). Two specimens.

Nassariidae
Nassarius insculptus (Carpenter) (W, 50, 54, 57).

COLUMBELLIDAE

Aesopus arestus Dall (S) .

Aesopus eurytoides (Carpenter) (W, 50). Three specimens in dredgings.

Aesopus sanctus Dall (W).

Anachis subturrita Carpenter (S, W) .

Mitrella carinata (Hinds) (50). Five specimens.

Parametaria duponti (Kiener) (50) . One very small specimen.

MURICIDAE

Ceratostoma nuttallii (Conrad) (50, 57). Eight specimens in the two trips.

Maxwellid gemma (Sowerby) (50, 57). One specimen each trip.

Ocenebra gracdlima (Stearns) (W) .

Ocenebra poulsoni (Carpenter) (W) .

Ocenebra seftoni Chace (50, 57) . Five specimens.

Thaisinae
Acanthina lugubris (Sowerby) (W, 50, 54, 57) Common.

Epitonhdae

Epitonium sp. cf. E. apiculatum Dall (50). Three specimens.

Epitonium bellastriatum (Carpenter) (54) . One specimen.

Epitonium calif ornicum Dall (W) .

Epitonium sp. cf. E. columbianum Dall (50). One specimen.

Epitonium sawinae Dall (50) . Three specimens.

Turbonilla (Pyrgiscus) halidoma Dall & Bartsch (50) .

Chace — Marine Mollusks of Guadalupe Island 325

Janthinidae
Janthina globosa Blainville (50) . Two small ones.

Eulimidae
Eulima calif ornica (Bartsch) (50). Three specimens.

Pyramellidae

Odostomia (Miralda) aepynota Dall & Bartsch (W, S) .

Odostomia (Menestho) amilda Dall & Bartsch (W) .

Odostomia (Menestho) callipyrga Dall & Bartsch (55). One specimen.

Odostomia (Chrysallida) clementina Dall & Bartsch (W) .

Odostomia (Chrysallida) deceptrix Dall & Bartsch (S).

Odostomia (Iolina) eucosmia Dall & Bartsch (S) .

Odostomia (Ividella) navisa Dall & Bartsch (S, W).

Odostomia (Chrysallida) pulcia Dall & Bartsch (W) .

Odostomia (Ivara) turricula Dall SC Bartsch (S) .

Odostomia (Chrysallida) virginalis Dall & Bartsch (S, W).

Cypraeidae

Cypraea spadicea Swainson (50) . One specimen.

Erato columbella Menke (W, 50). Eight specimens.

Trivia calif orniana (Gray) (54). Four specimens.

Trivia solandri (Gray SC Sowerby) (W, 57) . One specimen.

Ranellidae

Bursa calcipicta Dall (W). Judging from photographs of the type specimen,

this species is very close to the following species.
Bursa calif ornica (Hinds) (S, 50, 54, 57) . Many specimens.

Triphoridae

Triphora sp. cf. T. chamberlini Baker (50) . One specimen.
Triphora pedroana Bartsch (S, W) .

Cerithiopsidae

Cerithiopsis guadalupensis Strong (S) .
Cerithiopsis oxys Bartsch (S, W) . ,
Metaxia diadema Bartsch (S, W) .
Seila montereyensis Bartsch (S, W) .

Cerithiidae

Alabina diomede Bartsch (50). One specimen.

Alabina jordani Strong (S, W) .

Bittium interfossum (Carpenter) (S) .

Diastoma slevini Strong (S, 50, 54, 57) . Common in dredgings.

326 San Diego Society of Natural History

Caecidae

Caecum californicum Dall (S).
Fartulum hemphdli Bartsch (S, W) .

Vermetidae

A letes squamigerus Carpenter (W).

Bivonia compacta Carpenter (57) . Several on Haliotis shells.
Spiroglyphus lituellus (Morch) (W, 50, 57). Six on Haliotis shells.
Vermicularis eburnea (Reeve) (50) . A few small specimens.

TURRITELLIDAE

Turritella orthosymmetra Berry (50, 54). Several specimens, mostly small.

LlTTORINIDAE

Littorina planaxis Philippi (S, W, 50, 54, 57). Common.

FOSSARIDAE

Iselica jenestrata (Carpenter) (S).

LlTIOPIDAE

Alaba jeannettae Bartsch (S) .

Barleeiidae
Barleeia calif ornica Bartsch (S, W) .

RlSSOIDAE

Alvania aequisculpta Keep (S) .

Alvania cosmia Bartsch (S, W) .

Alvania oldroydae Bartsch (S, W).

Alvania purpurea Dall (S, W, 50). A few specimens in dredgings.

Amphithalamus inclusus Carpenter (S).

Amphithalamus tenuis Bartsch (S).

Nodulus kelseyi Bartsch (S) .

Rissoinidae (as used by Dall)

Rissoina calif ornica Bartsch (S, W) .

Rissoina cleo Bartsch (S, W) .

Rissoina guadalupensis Strong (S, W) .

Rissoina lowei Strong (S, W, 57) . Several specimens.

Rissoina willetti Strong (S, W) .

RlSSOELLIDAE

Rissoella bakeri Strong (S).

Chace — Marine Mollusks of Guadalupe Island 327

Hipponicidae

Hipponix antiquatus (Linnaeus) (S, W, 50, 57). Two specimens.
Hipponix tumens Carpenter (S, W, 50, 57) . Three specimens.

Calyptraeidae

Crepidula (Crepipatella) lingulata Gould (S, W, 50, 54, 57). Common on

Astraea undosa Wood.
Crepidula perforans Valenciennes (57) . One specimen.

Vitrinellidae
Macromphalina occidentalis Bartsch (50) . One specimen.

Naticidae
Lamellaria stearnsi orbiculata Dall (57). One specimen.

Acmaeidae

Acmaea digitalis Eschscholtz (S, W, 50, 54, 57). Several specimens.

Acmaea limatula Carpenter (W, 50, 57). A few specimens.

Acmaea mesoleuca var. vespertina Reeve (57). A few specimens.

Acmaea paleacea Gould (S, W).

Acmaea pelta Eschscholtz (S, W) .

Acmaea scabra (Gould) (50). Several specimens.

Lottia gigantea Gray (S, W, 50, 54, 57). Many specimens.

Phasianellidae

Tricolia pulloides (Carpenter) (S, W, 50) .

Tricolia (Eulithidium) rubrilineata (Strong) (S, W).

Turbinidae

Astraea guadalupeana Berry (50). One specimen, with A. inequalis.
Astraea inequalis (Martyn) [= A. gibberosa (Dillwyn)] (57). The two

specimens are close to the original figure of Trochus inequalis (Martyn

1784, pi. 31), and they match the original figures of A. diademata

(Valenciennes 1846, pi. 3, figs. 2-2b).
Astraea lithophora (Dall) (W).

Astraea sp. cf. A. petrothauma Berry (50). One specimen.
Astraea undosa (Wood) (S, 50, 54, 57) . Many specimens; apparently more

variable than on the mainland.
Homolopoma carpenteri (Pilsbry) (W, 57). A few specimens.
Homolopoma paucicostatum (Dall) (S, W) .
Homolopoma luridum (Dall) (55). Two specimens.

328 San Diego Society of Natural History

Liotiidae

Liotia acuticostata Carpenter (S, W) .

Liotia calif ornica Dall (S, 50) . Five specimens.

Liotia fenestrata Carpenter (S) .

Liotia beimi Strong & Hertlein (50, 57) . Several specimens.

Trochidae

Calliostoma splendens Carpenter (S, 50, 57) . Two specimens.

Calliostoma sp. (W) . An unidentified species also in 1957.

Margarites acuticostata Carpenter (W, 50, 54, 57) . Common in dredgings.

Margarites parcipicta (Carpenter) (S, W) .

Norrisia norrisii (Sowerby) (S, W, 50, 54, 57) . Many specimens.

Tegula gallina (Forbes) (S, W).

Tegula gallina multifilosa (Stearns) (50, 57). Several specimens.

Tegula regina (Stearns) (S, 50, 54, 57). Several specimens.

Vitrinellidae

Circulus rossellinus Dall (S) .

Teinostoma invallatum (Carpenter) (S, W) .

Teinostoma supravallatum (Carpenter) (S, W) .

Haliotidae

Haliotis californiensis Swainson (S, 54, 57) . Many specimens.
Haliotis corrugata Gray (S, 55) . Only four or five specimens.
Haliotis cracherodii Leach (S, W) .

Haliotis sp. cf. H. julgens Philippi (S, 54, 57). Not typical of H. julgens;
may be a new species.

FlSSURELLIDAE

Fissurella volcano Reeve (W, 50, 54) . Only a few specimens.
Megathura crenulata (Sowerby) (50, 54). Three specimens; evidently not
common.

Pelecypoda

Arcidae
Area bailyi (Bartsch) (S).

Glycymeridae
Glycymeris guadalupensis Strong (S, W, 50, 54, 57) . Common in dredgings.

Chace — Marine Mollusks of Guadalupe Island 329

Philobryidae
Philobrya setosa (Carpenter) (S, W) .

Mytilidae

Crenella Columbiana Dall (57) . One specimen.
Crenella divaricata (d'Orbigny) (S, W) .
Crenella megas Dall (50) . Fifteen valves.
Lit h o pha ga plumula (Hanley) (54). One specimen.
Modiolus pallidulus Dall (50) . Two specimens.

Pectinidae

Hinnites gigantea (Gray) (W) .

Pecten diegensis Dall (50, 54) . Several valves.

Pecten lowei Hertlein (50) . One valve.

Pecten pernomus Hertlein (50, 54, 57) . Many good pairs.

Limidae
Lima subauriculata (Montague) (50,54). Many valves.

Thraciidae

Thracia diegensis Dall (W) .

Thracia squamosa Carpenter (50, 54, 57) . Many small valves.

Lyonsiidae
Lyonsia calif ornica nesiotes Dall (50, 54, 57) . Several live specimens.

CUSPIDARIIDAE

Cuspidaria pectinata (Carpenter) (50) . Two valves.

Leiomya scaber (Carpenter) (50, 54) . Three pairs and several valves.

ASTARTIDAE

Bernardina baker i Dall (W) .

Carditidae

Glans carpenteri (Lamy) (W) .
Milneria kelseyi Dall (S) .
Milneria minima (Dall) (W) .

Chamidae
Chama buddiana C. B. Adams (50) . Two upper valves.
Chama pellucida Browerip (S, W) .

Thyasiridae
Thyasira barbarensis (Dall) (54). One valve.

330 San Diego Society of Natural History

Ungulinidae
Diplodonta subquadrata Carpenter (50). Eight valves.

LUCINIDAE

Lucina approximate! (Dall) (50,54). Six specimens plus some valves.
Lucina calif ornica Conrad (S, W, 50, 54) . A few valves.

Leptonidae
Kellia laperousii Deshayes (W, 50) . One valve.

Cardiidae

Cardium biangulatum Broderip & Sowerby (54). Several pairs and valves.
Protocardia centifilosa (Carpenter) (50). Two valves.

Veneridae

Antigona fordi Yates (W, 54). Several pairs and valves.
Psephidea cymata Dall (S, W, 50, 54) . Several pairs and valves.
Psephidea salmonea (Carpenter) (54). Three pairs and nine valves.
Transenella puella (Carpenter) (S, W, 50, 54, 57) . Many live specimens and
valves.

Tellinidae
Tellina pacijica Dall (W, 54, 57) . Two pairs.

Semelidae
Semele incongrua Carpenter (54). One specimen.

CORBULIDAE

Grippina calif ornica Dall (S) .

SCAPHOPODA

Dentalhdae

Cadulus fusiformis Pilsbry & Sharp (S) .
Dentalium berryi Smith & Gordon (50) . One specimen.
Dentalium splendidulum Sowerby (50). One specimen in dredgings.
Siphonodentalium quadrifissatum Pilsbry & Sharp (S, W, 50, 57). Many
specimens.

Polyplacophora
Lepidopleuridae
Lepidopleurus rugatus Pilsbry (57). One specimen.

Lepidochitonidae

Lepidochitona sp. (55). One very small specimen.
Nuttallina calif ornica Reeve (57). Three specimens.

Chace — Marine Mollusks of Guadalupe Island 331

ischnochitonidae

Ischnochiton biarcuatus Dall (57) . One specimen.
Ischnochiton mertensi (Middendorff) (57). Two specimens.

ACANTHOCHITONIDAE

Acanthochitona arragonites Carpenter (55). Two specimens.

Pteropoda
Cavolinidae

Carolina inflexa (Lesueur) (50) . Seven specimens.
Carolina occidentalis Dall (50). One specimen.
Clio pyramidata Rang (50). Four fragments.
Curierina columella Rang (50) . One specimen.

Brachiopoda
Terebratu lidae

Terebratulina unguicula Carpenter (50). One valve.

Terebratellidae

Platidia sp. cf. P. anomioides radiata Dall (57). Three specimens and five
valves.

A NEW SPECIES OF GASTROPOD

Ocenebra seftoni sp. nov.

Shell small, white with a light tan-colored band on the shoulder and
another below the periphery and with a few darker spots on the spiral cords and
between the axial ribs on the base. Nuclear whorls \y2, squarish, a prominent
keel at the top of the first whorl and a second keel developing below it on the
last half of the nuclear. Later whorls 5, slopingly shouldered and distinctly
angled at the periphery. Two spiral cords on the first whorl, increasing to 4
on the penultimate and to 9 on the body whorl. Axial ribs 9, rather strong,
reaching from suture to suture and down over the base. Whole surface with
fine axial lamellae, which are strongest where the cords pass over the axial
ribs. Aperture oval, the outer lip thickened, especially where it joins the
previous whorl; three denticles within. Callus rather heavy on the body whorl
and on the slightly twisted columella. Siphonal canal about half as long as the
aperture and well covered, the small umbilical chink nearly covered by the
end of the columellar callus. Measurements of the holotype: height 12 mm.;
diameter 6.2 mm.; height of the aperture 3.4 mm. Color description taken
from one of the paratypes, which is brighter colored than the type.

332 San Diego Society of Natural History

The holotype is number 12955 in the conch ological collection of the San
Diego Natural History Museum. It was dredged in about 40 fathoms by Dr.
Carl L. Hubbs and party, February 2, 1950, at Melpomene Cove, Guadalupe
Island, Baja California, Mexico. Paratypes are at the California Academy of
Sciences and at the Museum of Comparative Zoology, Harvard University.
The species is named for Mr. Joseph Sefton, then owner of the Research Ship
Orca, from which the type was collected.

The angular nuclear whorls indicate that the new species is closely re-
lated to Ocenebra crispatissima Berry, from Catalina Island, and to members
of the O. barbarensis group. It is especially similar to O. crispatissima but
has fewer spiral chords on the body whorl and a shorter siphonal canal.

LITERATURE CITED
Johnson, Clifton W.

1953 Notes on the geology of Guadalupe Island, Mexico. Am. Jour.
Sci. 251: 231-236.

Keen, A. Myra

1937 An abridged check list and bibliography of west North American
marine Mollusca. 1-88. Stanford University Press.

Martyn, Thomas

1784 The universal conchologist . . . vol. 1 [Numbering of plates
varies.]

PlLSBRY, H. A.

1927 Expedition to Guadalupe Island, Mexico, in 1922. Land and
freshwater mollusks. Proc. Calif. Acad. ser. 4. 16: 159-203,
pis. 6-12.

Scharff, Robert Francis

1911 Distribution and origin of life in America, i-xvi, 1-497. Con-
stable & Co., Ltd-, London.

Strong, A. M.

1954 The marine molluscan fauna of Guadalupe Island, Mexico. Min.
Conch. Club So. Calif. 142: 6-10.

Strong, A. M., and G. D Hanna

1930 Marine Mollusca of Guadalupe Island, Mexico. Proc. Calif.
Acad. ser. 4. 19: 1-6.

Valenciennes, Achille

1846 Voyage autour du monde sur ... la Venus, pendant . . . 1836-
1839. Adas.

A NEW MOLLUSK FROM SAN FELIPE,
BAJA CALIFORNIA

BY

E. P. Chace

As it has become easier to collect in the northern part of the Gulf of
California, molluscan material in collections has increased. This additional
material has shown the intergradation of some forms originally described from
scanty material as distinct. On the other hand, however, it has emphasized
the distinctness of some forms that conservative students previously were
unwilling to name. Among these latter forms is the Nassarhis described
below, of which previously only one specimen was known.

Nassarius howardae sp. nov.

Shell of medium size for the genus, white, the thin periostracum light
tan near the aperture but darker above, high spired, the apex acute, the whorls
fairly convex, the suture impressed well below periphery. Nuclear whorls
21/2, smooth, white. Post-nuclear whorls 6. Axial ribs on early whorls 8,
rounded, increasing to 25 on penultimate whorl and more on body whorl but
becoming much weaker on base so that spirals are almost smooth. Spiral
cords on early whorls 3, fairly strong, straplike, separated by channels about
as wide, increasing to 7 or 8 on penultimate whorl and more on body whorl,
forming low nodes where they pass over axial ribs. Aperture less than half
length of shell; outer lip with varixlike thickening and 7 or 8 small elongate
denticles within; inner lip with very thin callus on body whorl; columella
nearly straight, with distinct channel just above its termination. Canal open,
moderately deep. Some specimens with a second varix slightly more than Y2
whorl back from aperture. Measurements of holotype: height 28 mm.; di-
ameter 13}/2 mm.; height of aperture 11 mm.

The holotype is number 12954 in the conchological collection of the San
Diego Natural History Museum. It was collected in December 1956 by Mrs.
E. P. Chace and Mrs. Faye Howard at Almejas Beach, about 5 miles north
of San Felipe, Baja California, Mexico. Paratypes are in the collections of
Dr. S. Stillman Berry (Redlands, California), the California Academy of
Sciences, the Museum of Comparative Zoology at Harvard University, and
the Philadelphia Academy of Natural Sciences. Another specimen of this
species, number 21879 in the collection of the San Diego Natural History
Museum, was collected at San Felipe about 1935 by Mr. J. M. Snyder.

334

San Diego Society of Natural History

Nassarius howardae somewhat resembles N. perpinguis Hinds, but the
shell is heavier and more porcellaneous and the spirals are broader. Nassarius
rhinetes Berry differs in having only 5 spirals on the penultimate whorl, and
its sculpture is much more open.

FlG. 1- Nassarius howardae sp. nov. Holotype (right) and paratype.

TRANSACTIONS

OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY
Volume XII, No. 21, pp. 335-346, figs. 1-3 May 27, 1959

PLEISTOCENE MOLLUSKS FROMTECOLOTE CREEK,
SAN DIEGO, CALIFORNIA

BY

William K. Emerson

American Museum of Natural History
AND

Emery P. Chace

San Diego Natural History Museum

Contents

Introduction 335

Locality Description 336

Composition and Inferred Significance of the Fauna 336

Literature Cited 343

Introduction

This paper records an ecologically significant assemblage of late-Pleistocene
invertebrates from a marine terrace deposit in Tecolote Creek, San Diego
County, California. Although the occurrence of fossils of this age has long been
known in the San Diego area, the assemblage from this locality has not pre-
viously been described. Several locally extinct southern species of mollusks
are recorded here for the first time in the Pleistocene fauna of this region.

We are indebted to George P. Kanakoff of the Los Angeles County
Museum for calling our attention to this exposure and for permitting us to
list the species which he collected from it. James R. Emerson kindly assisted
in collecting. Dr. Leo G. Hertlein of the California Academy of Sciences
collaborated in the identification of some of the fossils.

The collection is deposited in the San Diego Natural History Museum.

336 San Diego Society of Natural History

Locality Description

The fossils occur in a bluff about 20 feet in elevation bordering Linda
Vista Mesa along the southeast side of the mouth of Tecolote Creek, on the
north side of Mission Valley (see figs. 1, 2). The locality is about one-fifth
of a mile northeast of Morena Boulevard along a private road to a Hazard
Construction Company quarry.

The terrace platform angularly truncates fine-grained bluish-gray sand
of the San Diego formation (mid-Pliocene) . The terrace deposits are com-
posed of a basal conglomerate, the fossiliferous sand, and a non-fossiliferous
cover of sand and soil. The fossiliferous sediment is a poorly sorted, coarse,
and well consolidated sand, which weathers dark brown. This sandstone
locally overlies the conglomerate of pebbles and small boulders or, in the
absence of the conglomerate, lies directly on the unevenly eroded surface
of the Pliocene sediments. The contact is only a few feet above the adjacent
valley floor (fig. 3).

Fragments of fossils occur locally in the marine sediments, but complete
specimens are largely confined to thin beds at intervals above the level of
the conglomerate. Oysters, however, are found attached to boulders and
pebbles in the upper part of the conglomerate. Paired valves of Tagelus
californianus commonly appear in the upper part of the sandstone, and occa-
sional paired valves of Chione gnidia and Chione californiensis are present in
the basal part of this bed.

The apparently non-fossiliferous terrace cover is 1 to 10 feet thick and
weathers reddish-brown. It is presumably alluvial in origin and therefore is
late Pleistocene (?) and Recent in age.

Composition and Inferred Significance of the Fauna

The writers collected 61 species of mollusks from three stations in the
bluff. Nine additional molluscan species were obtained from the exposure by
Mr. George P. Kanakoff. The collected fauna therefore totals 70 species of
mollusks, including 34 pelecypods, 1 chiton, and 35 gastropods. The fauna
is enumerated in table 1. Names of species reported by Mr. Kanakoff but
not seen by the writers are enclosed in brackets.

The composition of the fauna suggests a semi-protected shallow-water
assemblage of a bay-lagoon environment. The large number of rock and sand
inhabitants and rocky rubble dwellers require a hard substrate, but the presence
of some tidal-flat inhabitants, such as Melampus, Cerithidea, and Nassarius
tegida, indicates adjacent mudflats. Such substrates exist in San Diego and
Mission Bays at the present time.

The habitat requirements are corroborated by the composition and sequence
of the terrace sediments. The basal conglomerate is overlain by local beds
of coarse sand containing fragmental and minute shells with occasional complete
larger shells. This in turn is overlain by thin zones containing paired specimens
of the bay razor clam, Tagelus californianus, in the finer grained sand near
the top of the deposit.

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Emerson & Chace — Tecolote Creek Mollusks

337

MILES

MEXICO

Fig. 1. The San Diego area, showing late Pleistocene fossiliferous deposits.

The broken line shows the present coastline; the solid line approximately delineates
the present 100-foot contour (modified after Stephens 1929). Numbers indicate the late
Pleistocene fossiliferous localities mentioned in the text: 1. Pacific Beach (Arnold 1903),
2. Railroad Cut (Stephens 1929), 3. Bay Point [Crown Point] (Stephens 1929),
4. Tecolote Creek, 5. West side of Point Loma (Webb 1937), 6. East side of Point
Loma (Stephens 1929), 7. Spanish Bight (Hertlein and Grant 1944), 8. Indian Point
[foot of 26th Street] (Arnold 1903), 9. Border locality (Emerson and Addicott 1953).

338 San Diego Society of Natural History

Table 1. Mollusks from Tecolote Creek

Names in brackets are for species reported by George P. Kanakoff but not seen by
the writers. Occurrence in other Late Pleistocene deposits of the San Diego area is shown,
based on the following reports: (1) Pacific Beach, Arnold (1903); (2) Point Loma,
Webb (1937); (3) Spanish Bight, Hertlein and Grant (1944); (4) Indian Point,
Arnold (1903); and (5) Border locality, Emerson and Addicott (1953).

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Cardium cf. C. biangulatum Broderip

and Sowerby (juvenile)
[Cardium procerum (Sowerby)]
[Cardium quadragenarium (Conrad)]

Chione calif orniensis (Broderip)
[Chione cortezi (Carpenter) ]

Chione gnidia (Broderip and Sowerby)

Corbula luteola Carpenter
[Crassinella branneri (Arnold)]
[Crassinella nuculiformis Berry]

Cryptomya California (Conrad)

Diplodonta sericata (Reeve)

Donax californica Conrad
[Glycymeris subobsoleta (Carpenter) ]

Lucina approximata (Dall)

Lucina nuttalli Conrad

Macoma acolasta Dall

Macoma pacts Pilsbry and Lowe

Macoma yoldiformis Carpenter

Mactra cf. M. californica Conrad

Nucula exigua Sowerby

Ostrea angelica Rochebrune

Ostrea cf. O. lurida Carpenter

Pecten circularis Sowerby

Pecten latiauritus Conrad
[Petricola parallela Pilsbry and Lowe]

Protothaca staminea (Conrad)

Semele quentinensis Dall

Septifer bifurcatus (Conrad)

Siliqua lucida (Conrad)

Tagelus californianus (Conrad)

Tellina cf. T. idae Dall (frag.)

Tellina meropsis Dall
\7jrfaea pihbryi Lowe]

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Emerson & Chace — Tecolote Creek Mollusks

339

Table 1 — Continued

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Lepidochitona keepiana Berry

Gastropoda
Actnaea insessa (Hinds)
Acmaea limatula Carpenter
Acteocina ci. A. culcitella (Gould)
Alabina ci. A. io Bartsch
Alabina tenuisculpta (Carpenter)
Anachis coronata hannana Hertlein and Strong
Bittium interfossa (Carpenter)
Bulla gouldiana Pilsbry
Caecum californicum Dall
Cerithidea californica (Haldeman)
Cerithiopsis carpenteri Bartsch
Crucibulutn spinosum (Sowerby)
Cylichnella attonsa (Carpenter)
Diodora inaequalis (Sowerby)
Eupleura muriciformis Broderip
Littorina scutulata Gould
Lucapinella callomarginata (Dall)
"Mangelia" barbarensis Oldroyd
Melampus olivaceus Carpenter
Micranellum crebricinctum (Carpenter)
Morula lugubris (C. B. Adams)
Nassarius tegula (Reeve)
Ocenebra ci. O. barbarensis (Gabb)
Ocenebra foveolata (Hinds)
Odostomia fetella Dall and Bartsch
Olivella baetica Carpenter
Pedipes liratus Binney
Pedipes unisulcatus Cooper
["Phyllonotus" radix nigritus (Philippi)]
Pyramidella ci. P. ad am si Carpenter
Retusa bar pa (Dall)
Syncera translucens (Carpenter)
Tegula ligulata (Menke)
Triphora pedroana Bartsch
Turbonilla aepynota Dall and Bartsch

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340 San Diego Society of Natural History

With the exception of Macoma acolasta (Dall 1921a), the assemblage is
composed entirely of extant species. The fauna contains, however, a significant
warm-water element comprising species which now are locally extinct (see
table 2) or near the northern endpoint of their range (see table 3). The
presence of this "tropical" element required water temperatures warmer than
now exist in the San Diego area and thus suggests that a northern shift of
marine isotherms had occurred prior to the deposition of these terrace deposits.
Similar warm-water faunas are known from late Pleistocene terrace deposits
of the Los Angeles basin (Woodring et al. 1946) .

Constituents of the Tecolote Creek fauna known to occur in other
assemblages of late Pleistocene age in the San Diego area are shown in table 1.
Of particular ecologic significance are Eupleura muriciformis, Cardium pro-
cerum, Crassinella branneri, Diplodonta sericata, and Chione gnidia, which
are warm- water indicators. They are known from Tecolote Creek and Spanish
Bight, and the first four occur at Indian Point. None of them, however, has
been found at Pacific Beach, and only one, Crassinella branneri, is recorded
from the Border locality. Also, Stephens (1929) listed Chione gnidia and
Diplodonta sericata from the Railroad Cut.

In the San Diego region, warm-water elements are thus known from the
lowest exposed terrace deposits at Spanish Bight (Hertlein and Grant 1944),
at Railroad Cut (Stephens 1929), at Indian Point (Arnold 1903), and on
the protected side of Point Loma (Stephens 1929) . Mixed associations of
warm-water and cool-water indicators are recorded from the Border locality.
Cold-water assemblages are known from the exposed western side of Point
Loma (Webb 1937; Berry 1922) and from Pacific Beach (Arnold 1903).

When the distribution of the faunas is viewed in terms of Pleistocene
topography, a close correlation is evident between the inferred hydroclimate of
the embayment and the composition of the fauna (fig. 1 ) . Associations domi-
nated by warm-water elements lived in semi-protected back-bay habitats;
associations dominated by cool-water elements lived in open-coast habitats; and
mixed associations lived in intermediate habitats.

This distributional pattern is apparently explained by the presence of
near-shore highlands which sheltered parts of the embayment from northwest
weather. Soledad Mountain protected back-bay sites in the Mission Bay and
Mission Valley areas, including warm-water assemblages at the Railroad Cut
and Tecolote Creek. Pleistocene Point Loma island protected the Spanish
Bight and especially the back-bay areas, such as Indian Point. Exposed localities,
Pacific Beach and the windward side of Point Loma island, supported pre-
dominantly cool-water assemblages. The partially protected Border locality,
on the other hand, apparently supported a mixed assemblage with cool-water
elements predominating. Although some mixing of faunal elements may have
resulted from transportation by currents and from redeposition of sediments,
the totality of evidence suggests that the faunal composition reflects local
habitat conditions (Emerson 1956) .

Emerson & Chace — Tecolote Creek Mollusks

341

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342

San Diego Society of Natural History

Table 3. Certain locally extant "southern" species in the Tecolote Creek fauna

Present Range

Gastropoda
Morula lugubris

Pelecypoda
Cardium biangulaturn

Diplodonta sericata

Semele quentinensis
Tellina meropsis

Off Catalina Island and Redondo Beach, California
(Burch 1940) to Panama (Adams 1852)

Off Catalina Island and Redondo Beach, California
(Burch 1944b) to Guayaquil, Ecuador (Hert-
lein and Strong 1947)

? Monterey Bay, California (Smith and Gordon
1948; Burch 1944a) ; San Ignacio Lagoon, Baja
California, to Guayaquil, Ecuador (Hertlein
and Strong 1947)

Point Mugu, California, to Costa Rica (Hertlein
and Strong 1949)

? Off Monterey, California (Burch 1944a); San
Diego, California (Dall 1921b) to Panama
(Burch 1944b)

Several prominent terraces have been noted in this region. Ellis (1919)
mapped the major terraces in coastal southwestern San Diego County and
described five terraces in the San Diego Bay area, namely: Tijuana terrace
at an elevation of 20 to 50 feet; Nestor terrace at 25 to 100 feet; Chula Vista
terrace at 100 to 130 feet; Avondale terrace at 200 to 250 feet; and Otay
terrace at 430 to 525 feet. All but the lowest, the Tijuana terrace, were con-
sidered to be of marine origin. In addition to these, Hertlein and Grant
(1944) mentioned the presence of other, less prominent, poorly developed
and preserved terraces in the area. Hanna (1926) proposed the name La Jolla
terrace for the lowest prominent terrace exposed in the La Jolla region and
continuing southward to Pacific Beach and along the west side of Point Loma
(Webb 1937). Gale (1931) believed the La Jolla terrace to be a northern
extension of the Nestor terrace. Carter (1957) considered the lowest marine
terrace referable to the Tijuana terrace and recognized remnants of it in
several areas of coastal San Diego County. Carter, however, did not demon-
strate the presence of fossiliferous deposits on the platform of his Tijuana
terrace.

Emerson & Chace — Tecolote Creek Mollusks 343

Owing to the remnantal preservation of the lowest exposed major marine
terraces of the area, it is not possible at the present time to establish precise
correlations between the various isolated outcrops. These deposits in the San
Diego area have been designated in part the Bay Point formation, and
correlated with the Palos Verdes formation of the Los Angeles region (Hertlein
and Grant 1939) .

On the basis of the available faunistic and physiographic evidence, it is
assumed that the fossiliferous sediments described from Tecolote Creek were
deposited on the Nestor terrace platform and are referable to the Bay Point
formation. Carbon- 14 dates are known for similar terrace features elsewhere
in California. Deposits from the lowest exposed terrace of the Palos Verdes
Hills of San Pedro (Kulp et al. 1952) , Santa Rosa Island (Orr 1956) , and
Santa Cruz (Bradley 1956) all showed ages greater than 30,000 years. The
modern composition of the faunas, the limited available carbon- 14 data, and
the known physiographic relationships of the terrace deposits, suggest the
age of these faunas to be later than early Pleistocene and older than Wisconsin
time (Addicott and Emerson 1959) .

Literature Cited

Adams, C. B.

1852 Catalogue of shells collected at Panama with notes on synonymy,
station and habitat. Ann. Lye. NY. 5: 222-298 (June),
297-549 (July).

Addicott, Warren O., and William K. Emerson

1959 Late Pleistocene invertebrates from Punta Cabras, Baja California,
Mexico. Am. Mus. Novitates 1925: 1-33, figs. 1-8.

Arnold, Ralph

1903 The paleontology and stratigraphy of the marine Pliocene and
Pleistocene of San Pedro, California. Mem. Calif. Acad. 3:
1-420, pis. 1-37.

Berry, S. Stillman

1922 Fossil chitons of western North America. Proc. Calif. Acad,
ser. 4. 11: 399-526, figs. 1-11, pis. 1-16.

Bradley, William C.

1956 Carbon- 14 date for a marine terrace at Santa Cruz, California.
Bull. Geol. Soc. Am. 67: 675-677, pi. 1.

Burch, John Q. (editor)

1944a Distributional list of the west American marine mollusks from
San Diego, California to the Polar Sea. Min. Conch. Club So.
Calif. 39: 1-29.
1944b Ibid. 41: 1-43.
1945 Ibid. 44: 1-44.

344 San Diego Society of Natural History

Burch, Tom

1940 Addition to the molluscan fauna of California. Nautilus 54:
46, 47, pi. 2, figs. 5-7.

Carter, George F.

1957 Pleistocene man at San Diego. 400 pp., 96 text figs., 6 tables.
Johns Hopkins Press, Baltimore.

Dall, W. H.

1921a New shells from the Pliocene or early Pleistocene of San Quentin
Bay, Lower California. West Am. Sci. 19: 21-23.

1921b Summary of the marine shellbearing mollusks of the northwest
coast of America, from San Diego, California, to the Polar
Sea . . . Bull. U.S. Natl. Mus. 112: 1-217, pis. 1-22.

Ellis, Arthur J.

1919 Physiography. In A. J. Ellis and C. H. Lee, Geology and
ground waters of the western part of San Diego County, Cali-
fornia. U. S. Geol. Surv. Water-Sup. Pap. 446: 20-50, pis. 6, 7.

Emerson, William K.

1956 Pleistocene invertebrates from Punta China, Baja California,
Mexico, with remarks on the composition of the Pacific Coast
Quaternary faunas. Bull. Am. Mus. Nat. Hist. Ill: 313-342,
1 fig., pis. 22, 23.

Emerson, William K., and Warren O. Addicott

1953 A Pleistocene invertebrate fauna from the southwest corner of
San Diego County, California. Trans. San Diego Soc. Nat.
Hist. 11: 429-444, 1 map.

Gale, Hoyt Rodney

1931 Summary of the stratigraphy. In U.S. Grant, IV, and H. R.
Gale, Catalogue of the marine Pliocene and Pleistocene Mollusca
of California and adjacent regions. Mem. San Diego Soc. Nat.
Hist. 1 : 19-78.

Hanna, Marcus A.

1926 Geology of the La Jolla Quadrangle, California. Univ. Calif.
Publ. Bull. Dept. Geol. Sci. 16: 187-246, pis. 17-23, 1 map.

Hertlein, Leo George, and U. S. Grant, IV

1939 Geology and oil possibilities of southwestern San Diego County.

Calif. Jour. Mines Geol. 35: 57-78, figs. 1-8.
1944 The geology and paleontology of the marine Pliocene of San

Diego, California. Part 1, Geology. Mem. San Diego Soc. Nat.

Hist. 2: 1-72 figs. 1-6, pis. 1-18, 1 map.

Emerson & Chace — Tecolote Creek Mollusks 345

Hertlein, Leo George, and A. M. Strong

1940-1951 Eastern Pacific expeditions of the New York Zoological
Society. Mollusks from the west coast of Mexico and Central
America. Zoologica, New York, 25: 369-430, pis. 1-2 (Decem-
ber 31, 1940); 28: 149-168, pi. 1 (December 6, 1943); 31:
53-76, pi. 1 (August 20, 1946); 31: 93-120, pi. 1 (December 5,
1946); 31: 129-150, pi. 1 (February 21, 1947); 33: 163-198,
pis. 1, 2 (December 31, 1948); 34: 63-97, pi. 1 (August 10,
1949); 34: 239-258, pi. 1 (December 30, 1949); 35: 217-252,
pis. 1, 2 (December 30, 1950); 36: 67-120, pis. 1-11 (August
20, 1951).

1955 Marine mollusks collected during the "Askoy" expedition to
Panama, Colombia, and Ecuador in 1941. Bull. Am. Mus. Nat.
Hist. 107: 159-318, pis. 1-3.

Jordan, Eric Knight

1924 Quaternary and Recent molluscan faunas of the west coast of
Lower California. Bull. So. Calif. Acad. 23: 146-156.

Keen, A. Myra

1958 New mollusks from tropical west America. Bull. Am. Paleont.
38: 239-255, pis. 30, 31.
Kulp, J. Laurence, Lansing E. Tryon, Walter R. Eckelman, and
William A. Snell

1952 Lamont natural radiocarbon measurements, II. Science 116:
409-414.

Lowe, Herbert N.

1935 New marine Mollusca from west Mexico, together with a list
of shells collected at Punta Penasco, Sonora, Mexico. Trans.
San Diego Soc. Nat. Hist. 8: 15-34, pis. 1-4.

Orr, Phil C.

1956 Radiocarbon dates for Santa Rosa Island. Santa Barbara Mus.
Nat. Hist. Bull. 2: 1-10.

Smith, Allyn G., and Mackenzie Gordon, Jr.

1948 The marine mollusks and brachiopods of Monterey Bay, Cali-
fornia, and vicinity. Proc. Calif. Acad. ser. 4, 26: 147-245,
figs. 1-4, pis. 3, 4.
Stephens, Frank

1929 Notes on the marine Pleistocene deposits of San Diego County,
California. Trans. San Diego Soc. Nat. Hist. 5: 245-256, 1 fig.

Webb, Robert W.

1937 Paleontology of the Pleistocene of Point Loma, San Diego
County, California. Trans. San Diego Soc. Nat. Hist. 8:
337-348,
Woodring, W. P., M. N. Bramlette, and W. S. W. Kew

1946 Geology and Paleontology of Palos Verdes Hills, California.
U. S. Geol. Surv. Prof. Pap. 207: 1-145, 14 figs., 37 pis.

346

San Diego Society of Natural History

yrm

Jim&>:

S.V-. ti #***

i

*. .•"••^.Vr * * •

> A

* : A,-

Fig. 2. Bluff on the southeast side of Tecolote Creek.
Fossils were collected in the Pleistocene sands exposed above the talus slope.

Fig. 3. Bluff on the southeast side of Tecolote Creek.

Here may be seen the contact of the basal Pliocene sand and the terrace sediments
composed of sand, conglomerate, and non-marine cover.

TRANSACTIONS

OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY
Volume XII, No. 22, pp. 347-406, fig. 1

TYPE LOCALITIES OF VASCULAR PLANTS
IN SAN DIEGO COUNTY, CALIFORNIA

BY

Ethel Bailey Higgins

Associate Curator of Botany
San Diego Museum of Natural History

SAN DIEGO, CALIFORNIA

Printed for the Society

July 22, 1959

Past and present boundaries of San Diego County.

The area of San Diego County has been reduced in the years shown, with the formation
of Inyo, San Bernardino, Riverside, and Imperial Counties. (Redrawn by Wesley Farmer
from a map originally prepared by Bertram B. Moore.)

TYPE LOCALITIES OF VASCULAR PLANTS
IN SAN DIEGO COUNTY, CALIFORNIA

BY

Ethel Bailey Higgins

CONTENTS

Introduction 349

Systematic List of Plants 350

Notes Concerning the Collectors 388

Old Place Names 400

Acknowledgements 400

Literature Cited 401

INTRODUCTION

When a new species of plant is discovered and named, one specimen of
it is designated as the type specimen or type. Later interpretation of the name
must if possible be based on this specimen. The place where the type specimen
is collected is known as the type locality. This locality is important because
any specimen of the species collected there has a certain authenticity in the
interpretation of the species. If the type specimen is lost or unavailable, or if it
is incomplete or abnormal, such another specimen may be especially valuable.

This paper lists type localities of vascular plants within the present boun-
daries of San Diego County. So far as possible, all proposed species and
varieties from this area are included, regardless of whether they are considered
valid. For want of definite data, however, it is sometimes difficult to know
whether a type locality is within the County. In early times, lack of accurate
maps and lack of place names often made precise location difficult even if the
need for it was felt. In doubtful cases, the distribution of the species and the
itinerary of the collector, so far as known, have been taken into account.

As shown in fig. 1, San Diego County has several times been reduced in
size. As a result, some type localities originally in San Diego County now are
in Riverside County or Imperial County. Yet in the botanical literature, some
of these may still be given as in San Diego County.

Many plants of San Diego County were named before the type concept
was established. If the author cited only one collection, this automatically is
the type; if he cited more than one, some later student is permitted to select
one of these as the type. The present paper does not select types; but it records
such selections as are known to have been made, even though the basis for
selection may be questionable. When no selection is known to have been made,
it is possible only to record the several localities originally cited.

350 San Diego Society of Natural History

SYSTEMATIC LIST OF PLANTS

POLYPODIACEAE

Asplenium vespertinum Maxon, Bull. Torrey Club 27: 197. 1900.
"San Miguel Mountain", Kimball, about February 1, 1900.

Cheilanthes Clevelandii D. C. Eaton, Bull. Torrey Club 6: 33. 1875.

"Growing on a mountain about forty miles from San Diego, California,
at an elevation of about 2,500 feet", Cleveland.

Notholaena californica D. C. Eaton, Bull. Torrey Club 10: 27. 1883.

"San Diego Co.", Burbeck, 1878. According to Tryon (1956, p. 73) this
is the type, and the locality is Spring Valley.

Notholaena Candida var. accessita Jepson, Man. Fl. PI. Calif. 27. 1925.
"Upper Vallecito", Jepson 8031.

Notholaena Newberryi D. C. Eaton, Bull. Torrey Club 4: 12. 1873.

"San Diego", Newberry, November 8, 1851. Although another collection
also was cited, this is the type according to Tryon (1956, p. 43).

Pityrogramma triangularis var. viscosa (D. C. Eaton) Weatherby.
Gymnogramme triangularis var. viscosa D. C. Eaton, Ferns N. Am. 2: 16.
1880.

"San Diego", Cleveland.

OPHIOGLOSSACEAE

Ophioglossum californicum Prantl, Ber. Deuts. Bot. Ges. 1: 351. 1883.

"Californien." According to Clausen (1938, p. 160) the type was collected
near San Diego by Cleveland in April 1882.

ISOETACEAE

Isoetes Orcuttii A. A. Eaton, Fern Bull. 8: 13. 1900.

"San Diego . . . low depressions on top of the mesas", Orcutt, June 7,
1884.

SELAGINELLACEAE

Selaginella cinerascens A. A. Eaton, Fern Bull. 7: 33. 1899.
"On bare ground, National City", Kimball.

PINACEAE

Cupressus Forbesii Jepson, Madrono 1: 75. 1922.

"Cedar Canyon, between El Nido and Dulzura", Forbes, December 30,
1907.

Higgins — Type Localities in San Diego County 351

Cupressus Stephensonii C. B. Wolf, Aliso 1: 125. 1948.

"Upper limits of King Creek, a tributary to the South Fork of the San
Diego River, Cuyamaca Mountains ... at about 4,000 ft. elevation on a dry
ridge near the creek", Wolf 9467, December 1, 1938.

Pinus cembroides var. Parryana (Engelmann) Voss.
Pinus Parryana Engelmann, Am. Jour. Sci. ser. 2. 34: 332. 1862.
"On the mountains east of San Diego", Parry, 1850.

Pinus Torreyana Parry, Bot. Mex. Bound. 210. 1859.

"Bluffs near the mouth of Soledad creek, 10 miles north of San Diego",
Parry.

Pseudotsuga macrocarpa (Vasey) Mayr.
Abies macrocarpa Vasey, Gard. Month. Hort. 18:21. 1876.
"Mountains near San Felipe", Newberry, November 1857.

GNETACEAE

Ephedra calif ornica S. Watson, Proc. Am. Acad. 14: 300. 1879.

"Promontory near San Diego and Jamul Valley", Palmer 364 & 365, 1875.
According to Abrams (1923, p. 78) the type locality is Point Loma.

NAIADACEAE

Potamogeton pauciflorus var. calif ornicus Morong, Bot. Gaz. 10: 254.

1885.

"San Diego County", S. B. and W. F. Parish. Many Parish collections
labeled as from San Diego County are from what is now Riverside County.

GRAMINEAE

Agropyron Parishii var. laeve Scribner & Smith, U. S. Dep. Agr. Div.
Agrost. Bull. 4: 28. 1897.
"Fowleys, Cuiamaca Mountains", Palmer 414, 1875.

Agrostis diegoensis Vasey, Bull. Torrey Club 13: 55. 1886.

"San Diego", Orcutt 1058. According to Orcutt (1908, col. 43) the
locality is Chollas Valley, San Diego.

Agrostis Kennedyana Beetle, Bull. Torrey Club 72: 547. 1945.
"San Diego", Grant 896, April 1902.

Bromus grandis (Shear) Hitchcock.

Bromus Orcuttianus var. grandis Shear, U. S. Dep. Agr. Div. Agrost. Bull.

23: 43. 1900.

"La Make, San Diego", Orcutt 472. According to Orcutt (1909, col. 521)
the locality is La Monte, near El Cajon.

352 San Diego Society of Natural History

Bromus Orcuttianus Vasey, Bot. Gaz. 10: 223. 1885.

"Mountains near San Diego", Orcutt, 1884. According to Orcutt (1908,
col. 45) the locality is Smith Mountain and the collector was H. C. Orcutt
[father of C. R. Orcutt].

Calamagrostis densa Vasey, Bot. Gaz. 16: 147. 1891.
"Near Julian", Orcutt.

Calamagrostis koelerioides Vasey, Bot. Gaz. 16: 147. 1891.
"Near Julian", Orcutt.

Danthonia californica var. unispicata Thurber in S. Watson, Bot. Calif.
2: 294. 1880.

"From San Diego to San Francisco", Bolander, Parry, Lemmon.

Deschampsia gracilis Vasey, Bot. Gaz. 10: 224. 1885.
"Mesas about San Diego", Orcutt.

Elymus Orcuttianus Vasey, Bot. Gaz. 10: 258. 1885.
"Near San Diego", Orcutt.

Eragrostis Orcuttiana Vasey, Contr. U. S. Nat. Herb. 1: 269. 1893.
"San Diego", Orcutt 1313, 1885.

Koeleria nitida var. californica subvar. transiens Domin, Bibl. Bot. 65: 234.
1907.

"San Diego", Brandegee, 1903.

Koeleria nitida var. californica subvar. vestita Domin, Bibl. Bot. 65: 234.
1907.

"Cusamacca Mts.", Palmer 405, 1875.

Koeleria pseudocristata var. californica Domin, Magyar Bot. Lapok 3 : 264.
1904.

"Hills, San Diego", Pringle, 1882.

Leptochloa imbricata Thurber in S. Watson, Bot. Calif. 2: 293. 1880.

"Larken's Station, San Diego County", Palmer 404, August 1875; also
Fort Yuma and through the Gila Valley to the Rio Grande.

Phalaris intermedia var. angustata Beal, Grasses N. Am. 2: 182. 1896.

"California", Pringle, 1882. According to Hitchcock (1935, p. 925) the
locality is San Diego.

Poa capillaris Scribner, U. S. Dep. Agr. Div. Agrost. Bull. 11: 51. 1898.
"Potrero", April 9, 1892. "Name of collector not given."

Poa Orcuttiana Vasey, West. Am. Sci. 3: 165. 1887.

"Near San Diego", Orcutt 1070, 1884; also Santa Barbara County. Ac-
cording to Hitchcock (1935, p. 944), the San Diego specimen is the type.

Higgins — Type Localities in San Diego County 353

Sitanion minus J. G. Smith, U. S. Dep. Agr. Div. Agrost. Bull. 18: 12.

1899.

"Jacumba Hot Springs, near monument 233, altitude 900 m.", Schoene-
feldt 3277, May 24, 1894.

Sporobolus altissimus Vasey, Proc. Calif. Acad. ser. 2. 2: 212. 1889.
"San Diego", Palmer, 1888.

Stipa coronata Thurber in S. Watson, Bot. Calif. 2:287. 1880.

"Hills near Julian City", Bolander; also near San Bernardino. Probably
a duplicate type according to Hitchcock (1925, p. 226) is a specimen in the
U. S. National Herbarium collected "in a canyon around springs on hillsides
near Julian City, San Diego Co., Apr. 1872."

Stipa diegoensis Swallen, Jour. Wash. Acad. 30: 212. 1940.

"Along vernal stream in chaparral, Proctor Valley near Jamul", Gander
5778, May 23, 1938.

CYPERACEAE

Carex monticola Dewey in Torrey, Bot. Mex. Bound. 229. 1859.
"Mountains east of San Diego", Parry.

Carex praegracilis Boott, Bot. Gaz. 9: 87. 1884.
"San Diego", Scott, 1880.

Carex spissa Bailey, Proc. Am. Acad. 22: 70. 1886.

"San Diego Co.", Pringle, 1882; also San Juan Capistrano, Arizona, Baja
California.

PALMAE

Washingtonia filifera (Linden) Wendland.

Pritchardia filifera Linden ex Andre, III. Hort. 24: 32, 105. 1877.

"Des bords du Colorado, dans l'Arizona", Roezl, 1869. According to
Parish (1909b, p. 462) the seeds from which the original plants were grown
probably were collected by Roezl from trees cultivated in San Diego.

LILIACEAE

Allium Bullardiae A. Davidson, Bull. So. Calif. Acad. 22: 72. 1923.

"Near San Julian, San Diego Co.", Bullard, April 1923 (Davidson 3575).

Allium praecox T. S. Brandegee, Zoe 5: 228. 1906.

"Common about San Diego, on northern slopes of canons, especially near
the coast." Brandegee mentioned several other localities, but according to
Abrams (1923, p. 396) this is the type locality.

354 San Diego Society of Natural History

Allium tenellum A. Davidson, Bull. So. Calif. Acad. 21: 39. 1922.
"Julian", Bullard, May 1922 (Davidson 3524).

Bloomeria Clevelandii S. Watson, Proc. Am. Acad. 20: 376. 1885.
"Mesas near San Diego", Cleveland, 1874.

Bloomeria crocea (Torrey) Coville.

Allium croceum Torrey, Bot. Mex. Bound. 218. 1859.

"Summit of the mountains east of San Diego", Parry.

Brodiaea Orcuttii (Greene) Baker.

Hookera Orcuttii Greene, Bull. Calif. Acad. 2: 138. 1887.

"San Diego county, near the city of that name, and also thirty miles to
the northward", Orcutt, 1884.

Calochortus concolor Purdy, Proc. Calif. Acad. ser. 3. 2: 135. 1901.
"Laguna, on the edge of the desert, San Diego County", Cleveland.

Calochortus Davidsonianus Abrams, III. Fl. Pac. States 1: 441. 1923.

"Grassy slopes between the Onofre Mountains and the sea", Abrams 3275,
1903.

Calochortus Dunnii Purdy, Proc. Calif. Acad. ser. 3. 2: 147. 1901.
"Near Julian", Dunn. Described from specimens cultivated by Purdy.

Calochortus Weedii Wood, Proc. Acad. Phila. 1868: 169. 1868.
"San Diego", Weed.

Chlorogalum parviflorum S. Watson, Proc. Am. Acad. 14: 243. 1879.
"Cajon Valley", Cleveland, 1877.

Hookera multipedunculata Abrams, Bull. Torrey Club 32: 537. 1905.
"In heavy soil near Cuyamaca Lake", Abrams 3897, June 25, 1903.

Lilium Fairchildii M. E. Jones, Contr. West. Bot. 16: 39. 1930.

From the garden of Mr. Rausch, Palomar Mountain, Jones, July 1929.
The plants were originally collected locally in pine woods in deep shade. The
species was named for Dr. J. H. Fairchild of Claremont.

Nolina interrata Gentry, Madrono 8: 181. 1946.

"Slope west of Dehesa School", Gentry 7330, August 5, 1945.

Yucca schidigera Roezl ex Ortgies, Gartenflora 20: 110. 1871.
Near San Diego, Roezl, 1869.

Yucca Whipplei Torrey, Bot. Mex. Bound. 222. 1859.
"On rocks near San Pasqual", Schott.

Higgins — Type Localities in San Diego County 355

Zygadenus diegoensis A. Davidson, Bull. So. Calif. Acad. 23: 105. 1924.

"Palomar Mts.", Fultz {Davidson 3392). Described from specimens culti-
vated by R. Kessler.

AMARYLLIDACEAE

Agave consociata Trelease, Missouri Bot. Gard. Rep. 22: 53. 1912.
"San Felipe", Parish 413, June 1882.

Agave deserti Engelmann, Trans. Acad. St. Louis 3: 310. 1875.

"Eastern base of the Southern California mountains and in the adjoining
deserts", G. N. Hitchcock, 1875; Palmer, 1875. According to Trelease (1912,
p. 53) these specimens are from east of San Felipe.

Agave Shawii Engelmann, Trans. Acad. St. Louis 3: 314. 1875.

"On the arid hills which overlook the sandy strand of the Pacific in the
southwest corner of California, where the boundary is marked by the initial
monument", Parry, 1850; Hitchcock and Parker, 1875; Palmer, 1875.

ORCHIDACEAE

Piperia Cooperi (S. Watson) Rydberg.

Habenaria Cooperi S. Watson, Proc. Am. Acad. 12: 276. 1877.

"On clay hills near San Diego", Cooper.

Piperia leptopetala Rydberg, Bull. Torrey Club 28: 637. 1901.
"Mountains east of San Diego", Parry, 1850.

FAGACEAE

Quercus agrifolia var. oxyadenia (Torrey) J. T. Howell.
Q. oxyadenia Torrey, Sitgreaves Rep. 172. 1853.

"Santo Isabelle", Woodhouse, December 1851.

Quercus XGanderi C. B. Wolf,, Proc. Calif. Acad. ser. 4. 25: 178. 1944.

"About 300 yards north of the entrance to the Volcan Indian School on
California State Highway No. 79 between Santa Ysabel and Lake Henshaw,
at 3100 feet elevation, in decomposed granite soil, near an occasional stream
just west of the highway", Wolf and Everett 9543, October 20, 1939.

Quercus Palmeri Engelmann, Trans. Acad. St. Louis 3: 393. 1877.

"In the mountains 80 miles east of San Diego", Palmer. According to
Abrams (1910, p. 346) it was first discovered "about five miles west of Jacumba
Hot Spring."

356 San Diego Society of Natural History

POLYGONACEAE

Chorizanthe discolor Nuttall, Jour. Acad. Phila. ser. 2. 1:167. 1848.

"St. Diego", Nuttall, 1836. Although no collector was given for this
species, the next-listed species was collected "With the preceding . . . (Nuttall) ".

Chorizanthe fimbriata Nuttall, Jour. Acad. Phila. ser. 2. 1 : 168. 1848.
"St. Diego", Nuttall, 1836.

Chorizanthe fimbriata var. laciniata (Torrey) Jepson.
C. laciniata Torrey, Pacif. Railr. Rep. 7 (3) : 19. 1857.

"San Felipe", Antisell, May 1855.

Chorizanthe leptotheca Goodman, Ann. Missouri Bot. Gard. 21: 61. 1934.
"Dry hills between Ramona and Ballena", Abrams 3777 , June 19, 1903.

Chorizanthe Orcuttiana Parry, Proc. Davenp. Acad. 4: 54. 1884.
"Exposed sandy soil on Pt. Loma", Orcutt, March 1884.

Chorizanthe procumbens Nuttall, Jour. Acad. Phila. ser. 2. 1: 167. 1848.
"St. Diego", Nuttall, 1836.

Chorizanthe procumbens var. albiflora Goodman, Ann. Missouri Bot.
Gard. 21: 87. 1934.

"Dry slope 2 miles east of Pala", Munz 10372, April 30, 1926.

Chorizanthe Thurberi (A. Gray) S. Watson.

Centrostegia Thurberi A. Gray ex Bentham in DeCandolle, Prod. 14: 27.
1857.

"In collibus arenosis ad San Felipe", Thurber, May 1852.

Chorizanthe uncinata Nuttall, Jour. Acad. Phila. ser. 2. 1 : 167. 1848.
"St. Diego", Nuttall, 1836.

Eriogonum cernuum subsp. viscosum Stokes, Gen. Eriog. 42. 1936.
"Near road, Campo to Buckman Springs", Stokes 3102.

Eriogonum fasciculatum var. maritimum Parish, Muhlenbergia 3: 59.
1907.
"Oceanside", Parish 4445, June 1897.

Eriogonum fasciculatum var. oleifolium Gandoger, Bull. Soc. Bot. Belg.
42: 189. 1906.

"San Diego", Schumo, March 1897. Professor Robert Dovin reports that
the type specimen is from the herbarium of Uselma C. Smith, Philadelphia.
He adds that the collector's name, given by Gandoger as "Schamo", can as
well be read "Schumo".

Higgins — Type Localities in San Diego County 357

Eriogonum Thurberi Torrey, Bot. Mex. Bound. 176. 1859.
"Sandy ravines, San Pasqual", Thurber, May 1852.

Eriogonum trichopes Torrey in Emory, Notes Mil. Rec. 151. 1848.
"Eastern slope of the Cordilleras of California", Emory, 1846.

Eriogonum trichopes subsp. cordatum (Torrey) Stokes.
E. cordatum Torrey in DeCandolle, Prod. 14: 21. 1857.
"California", Fremont; "prope San Felippe", Parry.

Eriogonum verticillatum Nuttall, Jour. Acad. Phila. ser. 2. 1:165. 1848.

"Near St. Diego." Since the paper is primarily on Gambel's collections
and since no collector is given for this species, Gambel's name is implied. It
may be, however, that Nuttall's name was omitted accidentally. It is not certain
that Gambel visited San Diego.

Eriogonum viminium subsp. molestum var. glabrum Stokes, Gen. Eriog.
51. 1936.

"Boulevard to Buckman Springs", Stokes 67, June 28, 1933.

Nemacaulis denudata Nuttall, Jour. Acad. Phila. ser. 2. 1: 168. 1848.

"St. Diego, Upper California, in sandy places near to the sea shore",
Nuttall, 1836.

Nemacaulis foliosa Nuttall, Jour. Acad. Phila. ser. 2. 1: 168. 1848.
"With the above [N. denudataj ', Nuttall, 1836.

CHENOPODIACEAE

Aphanisma blitoides Nuttall ex Moquin-Tandon in DeCandolle, Prod. 13
(2) : 54. 1849.
"Prope San-Diego", Nuttall, 1836.

Atriplex pacifica A. Nelson.

Obione microcarpa Bentham, Bot. Voy. Sulphur 48. 1844.

"San Diego", Hinds, October 1839.

Atriplex Watsonii A. Nelson.

A. decumbens S. Watson, Proc. Am. Acad. 12: 275. 1877.
"Near San Diego", Palmer 331, 1876.

Obione tetraptera Bentham, Bot. Voy. Sulphur 48. 1844.

"From the coast of California, probably San Diego", Hinds, October 1839.

Salicornia depressa Standley, N Am. Fl. 21: 85. 1916.
"San Diego", K. Brandegee, 1899.

Suaeda californica var. pubescens Jepson, Fl. Calif. 1: 447. 1914.
"Del Mar", Jepson.

358 San Diego Society of Natural History

AMARANTHACEAE

Amaranthus Palmeri S. Watson, Proc. Am. Acad. 12: 274. 1877.
"Larkin's Station", Palmer 323, August 1875.

NYCTAGINACEAE

Abronia umbellata var. platyphylla (Standley) Munz.

Abronia platyphylla Standley, Contr. U. S. Nat. Herb. 12: 314. 1909.

"Del Mar", T. S. Brandegee, May 12, 1894.

Mirabilis calif ornica Gray in Torrey, Bot. Mex. Bound. 169. 1859.
"Dry hills, San Diego", Parry, Thurber.

Mirabilis Froebelii (Behr) Greene.

Oxybaphus Froebelii Behr, Proc. Calif. Acad. 1: 72. 1855.
"Prope Warner's Ranch", Froebel.

PORTULACACEAE

Calandrinia heterophylla Rydberg, N. Am. FI. 21: 293. 1932.

"Old Mission Dam, San Diego County", Chandler 5152, April 10, 1904.

Calandrinia maritima Nuttall in Torrey & Gray, FI. N. Am. 1 : 197. 1838.
"St. Diego, California, on the sea coast", Nuttall, May 1836.

Calandrinia muricata Rydberg, N. Am. FI. 21: 293. 1932.
"San Diego", T. S. Brandegee, April 1906.

Calyptridium monandrum Nuttall in Torrey & Gray, FI. N. Am. 1: 198.
1838.

"St. Diego", Nuttall, 1836.

CARYOPHYLLACEAE

Loeflingia squarrosa Nuttall in Torrey & Gray, Fl. N. Am. 1: 174. 1838.
"Sandy plains, St. Diego", Nuttall, 1836.

Polycarpon depressum Nuttall in Torrey & Gray, Fl. N. Am. 1 : 174. 1838.
'On bare sand-hills, near St. Diego", Nuttall, 1836.

Silene multinervia S. Watson, Proc. Am. Acad. 25: 126. 1890.

"Jamuel, San Diego County", Orcutt, April 1885; also Santa Cruz Island.
According to Abrams (1944, p. 160) the type locality is Jamul.

Silene Palmeri S. Watson, Proc. Am. Acad. 11: 124. 1876.
"Cuyamaca Mountains", Palmer, August 1875.

Higgins — Type Localities in San Diego County 359

Silene verecunda var. platyota (S. Watson) Jepson.
S. platyota S. Watson, Proc. Am. Acad. 17: 366. 1882.

"Cuyamaca Mountains", Palmer, 1875; also San Bernardino and San
Jacinto Mountains. According to Abrams (1944, p. 167) the type locality
is Cuyamaca Mountains.

Spergularia Clevelandii (Greene) Robinson.
Tissa Clevelandii Greene, Fl. Fran. 127. 1891.

"Formerly abundant on rather sandy uplands about San Diego"; also San
Francisco.

RANUNCULACEAE

Aquilegia hypolasia Greene, Leafl. Bot. Obs. 2: 141. 1911.

"Between Campbell's and Cameron's ranches", M earns 3657, June 21,
1894; also between Pine Valley and Laguna. According to Munz (1946, p.
106) the type is M earns 3657.

Clematis lasiantha Nuttall in Torrey & Gray, Fl. N. Am. 1 : 9. 1838.
"With the preceding [C. paucijlora~}" ', Nuttall, 1836.

Clematis pauciflora Nuttall in Torrey & Gray, Fl. N. Am. 1: 9, 657. 1838,
1840.
"Near the sea-coast of St. Diego". Nuttall, 1836.

Delphinium coccineum Torrey, Pacif. Railr. Rep. 4: 62. 1857.
"Mountains east of San Diego", Parry, 1850.

Delphinium collinum Ewan, Bull. Torrey Club 63: 338. 1936.

"Ridge betw. Campbells Ranch and Mason Valley". Epling and Robison,
April 4, 1932.

Delphinium cuyamacae Abrams, Bull. Torrey Club 32: 538. 1905.

"Grassy slopes bordering Cuyamaca Lake, altitude 1550 meters", Abrams
3888, June 26, 1902.

Delphinium Parryi var. subglobosum (Wiggins) Munz.
D. subglobosum Wiggins, Contr. Dudley Herb. 1: 99. 1929.

"Open grassy slope near Banner", Wiggins 2003, March 20, 1926.

Myosurus minimus var. apus Greene, Bull. Calif. Acad. 1: 277. 1885.
"Table-lands back of San Diego", Orcutt, April 10, 1884.

Thalictrum coreospermum Greene, Repert. Nov. Sp. 7: 253. 1909.
"Cuyamaca Mountains", Orcutt, July 1889.

Thalictrum magarum Greene, Muhlenbergia 5: 130. 1909.
"Witch Creek", Alderson, April 1894.

360 San Diego Society of Natural History

BERBERIDACEAE

Berberis Higginsae Munz, Aliso 4: 91. 1958.

"Dry rocky point at Boulevard", Johnson and Stark 1351, July 12, 1929.

PAPAVERACEAE

Dendromecon saligna Greene, Pittonia 5: 300. 1905.
"Encenitas", T. S. Brandegee.

Eschscholtzia Clevelandii Greene, Pittonia 5: 248. 1905.

"At and near San Diego", and southward into Baja California.

Eschscholtzia floribunda Greene, Pittonia 5: 247. 1905.
"Near Santa Ysabel", Henshaw, May 15, 1893.

Eschscholtzia physodes Greene, Pittonia 5: 259. 1905.
"Witch Creek", Alder son, April 1894.

Eschscholtzia sanctarum Greene, Pittonia 5: 243. 1905.
"Grand Mesa", Greene?, May 1894.

Meconella kakoethes Fedde, Repert. Nov. Sp. 3: 275. 1907.
"San Diego", Orcutt, 1895.

Platystemon californicus var. nutans T. S. Brandegee, Zoe 5: 177. 1903.
"About San Diego and on many of the islands off the coast."

Platystemon obtectus Greene, Pittonia 5: 186. 1903.
"Witch Creek", Alder son, May 24, 1894.

Platystemon verecundus var. glabrifructifer Fedde, Pflanzenr. 4 (104):
131. 1909.

"San Diego", Greene; Palmer 8a, 1875.

CAPPARIDACEAE

Isomeris arborea Nuttall in Torrey & Gray, FI. N. Am. 1: 124. 1838.
"St. Diego", Nuttall, 1836.

Wislizenia divaricata Greene, Proc. Biol. Soc. Wash. 19: 130. 1906.

"Bonego Springs . . . southern part of the Colorado Desert in San Diego
County", Orcutt, June 23, 1888. Clearly Borrego Springs is meant.

CRUCIFERAE

Agianthus Jacobaeus Greene, Leafl. Bot. Obs. 1: 229. 1906.
"Cuyamaca Mountains", Orcutt 1507, July 1889.

Higgins — Type Localities in San Diego County 361

Arabis pulchra var. glabrescens Wiggins, Contr. Dudley Herb. 1 : 100. 1929.
"Gravelly slope between Julian and Banner", Wiggins 2015, March 20, 1926.

Arabis pulchra var. viridis Jepson, Fl. Calif. 2: 70. 1936.
"Summit of Mountain Springs grade", Jepson 11810.

Arabis sparsiflora var. californica Rollins, Rhodora 43: 402. 1941.
"Near Campo", Abrams 3563, May 24, 1903.

Caulanthus heterophyllus (Nuttall) Payson.

Streptantbus heterophyllus Nuttall in Torrey & Gray, FI. N. Am. 1 : 77. 1838.

"Bushy hills, near St. Diego", Nuttall, 1836.
Caulanthus stenocarpus Payson, Ann. Missouri Bot. Gard. 9: 300. 1923.
"Dry hillsides near Bernardo", Abrams 3364, May 1, 1903.

Lepidium acutidens var. microcarpum Thellung, Mitt. Bot. Mus. Univ.
Zurich 28: 271. 1906.
"San Diego", Jones 3061, 1882; also Pringle, 1882.

Lepidium f lavum var. felipense C. L. Hitchcock, Madrono 3 : 299. 1936.
"San Felipe", Pur pus, 1898.

Lepidium Robinsonii Thellung, Mitt. Bot. Mus. Univ. Zurich 28: 255.

1906.

"San Diego", Jones 3050, 1882; also Orcutt 1039. According to C. L.
Hitchcock (1936, p. 285) the type apparently is Jones 3050.

Streptanthus campestris S. Watson, Proc. Am. Acad. 25: 125. 1890.
"Campo", Vasey and Parish, 1880.

Thysanocarpus laciniatus var. rigidus Munz, Bull. So. Calif. Acad. 31: 62.
1932.

"Laguna Camp, Laguna Mts.", Munz 9701, May 16, 1925.

CRASSULACEAE

Dudleya Abramsii Rose, Bull. N. Y. Bot. Gard. 3: 14. 1903.

"Wet crevices of rocks west of Jacumba", Abrams 3707, June 1, 1903.

Dudleya aloides Rose, Bull. N. Y. Bot. Gard. 3: 15. 1903.
"San Felipe", Orcutt, April 1903.

Dudleya delicata Rose, Bull. N. Y. Bot. Gard. 3: 24. 1903.
"Spencer Valley", Abrams 3791, June 22, 1903.

Dudleya lagunensis (Munz) E. Walther.

Echeveria lagunensis Munz, Bull. So. Calif. Acad. 31: 64. 1932.

"Dry stony slopes, Campbell Ranch, Vallecito Valley", Munz and C. L.
Hitchcock 12612, April 3, 1932.

362 San Diego Society of Natural History

Dudleya lanceolata (Nuttall) Britton & Rose.

Echeveria lanceolata Nuttall in Torrey & Gray, Fl. N. Am. 1: 561. 1840.

"St. Diego", Nuttall, 1836.

Dudleya pulverulenta (Nuttall) Britton & Rose.

Echeveria pulverulenta Nuttall in Torrey & Gray, Fl. N. Am. 1 : 560. 1840.

"St. Diego", Nuttall, 1836.

Hasseanthus Blochmaniae subsp. brevifolius Moran, Des. PL Life 22: 80.

1950.

"Open area in chaparral near edge of mesa, Torrey Pines Park", Moran
3206, April 9, 1949.

Sedum variegatum S. Watson, Proc. Am. Acad. 11: 137. 1876.
"San Diego", Cleveland, May 1875.

Stylophyllum edule (Nuttall) Britton & Rose.
Sedum edule Nuttall in Torrey & Gray, Fl. N. Am. 1: 560. 1840.
"Edges of rocks and ravines, St. Diego", Nuttall, 1836.

Stylophyllum Orcuttii Rose, Bull. N. Y. Bot. Gard. 3: 36. 1903.
"Initial Mexican boundary monument", T. S. Brandegee.

Stylophyllum Parishii Britton, Bull. N. Y. Bot. Gard. 3: 37. 1903.
"Pala", S. B. and W. F. Parish 444, June 1880.

Tillaea erecta var. eremica Jepson, Man. Fl. PL Calif. 450. 1925.
"Vallecito", Jepson 8636.

SAXIFRAGACEAE

Heuchera brevistaminea Wiggins, Contr. Dudley Herb. 1: 100. 1929.

"Shallow soil on a rocky canyon slope a half-mile below Vallecitos View,
Laguna Mountains", Wiggins 2831, September 4, 1927.

Jepsonia Parryi (Torrey) Small.

Saxifraga Parryi Torrey, Bot. Mex. Bound. 69. 1859.

"Dry hills near San Diego and San Luis Rey", Parry.

Lithophragma trifida Eastwood ex Rydberg, N. Am. Fl. 22: 89. 1905.
"Near Sweetwater Dam", Eastwood.

Ribes canthariformis Wiggins, Contr. Dudley Herb. 1: 101. 1929.

"In chaparral on a northeast slope near Moreno Dam", Wiggins 2399,
April 14, 1927. The type plant has been destroyed during road construction,
but others are still to be found in the immediate vicinity.

Ribes indecorum Eastwood, Proc. Calif. Acad. ser. 3. 2: 243. 1902.
"Cajon Heights, near San Diego", Eastwood, March 14, 1891.

Higgins — Type Localities in San Diego County 363

ROSACEAE

Adenostoma fasciculatum var. obtusifolium S. Watson, Bot. Calif. 1: 184.

1876.
A. brevifolium Nuttall ex Rydberg, N. Am. Fl. 22: 396. 1913.

"Near San Diego", Nuttall, 1836.

Adenostoma laxum Gandoger, Bull. Soc. Bot. Fr. 59: 707. 1912.
"San Diego", Brandegee; also Mendocino.

Adenostoma sparsifolium Torrey in Emory, Notes Mil. Rec. 140. 1848.

"Cordilleras of California", Emory, December 2, 1846. According to
Abrams (1910, p. 378) the locality is near Warners Ranch.

Amelanchier alnifolia var. cuyamacensis Munz, Bull. So. Calif. Acad.
31: 65. 1932.

"Cuyamaca Lake", Munz 8099.

Cercocarpus minutiflorus Abrams, Bull. Torrey Club 37: 149. 1910.

"Dry chaparral-covered hills, near San Dieguito (Bernardo)", Abrams
3376, May 4, 1903.

Potentilla Bolanderi var. Clevelandii (Greene) Munz & Johnston.
P. Clevelandii Greene, Pittonia 1 : 102. 1887.

"At Laguna, in the mountains back of San Diego", Cleveland, July 1885.

Potentilla Parishii Rydberg, N Am. Fl. 22: 313. 1908.
"Descanso", S. B. Parish 4523, 1897.

Potentilla truncata (Rydberg) Munz & Johnston.
Horkelia truncata Rydberg, N. Am. Fl. 22: 274. 1908.

"Type collected in Brandegee's garden at San Diego, cultivated plant
brought from Ramona", Chandler 5321.

Prunus fremontii S. Watson, Bot. Calif. 2: 442. 1880.

"Oriflamme Canon", Cleveland; also San Bernardino Mountains. Accord-
ing to Jepson (1936, p. 229) and Abrams (1944, p. 466) the type locality is
Oriflamme Canyon.

Prunus fremontii var. pilulata Jepson, Man. Fl. PI. Calif. 507. 1925.
"Wagon Wash near Sentenac Canon", Jepson 8769.

Rosa Aldersonii Greene, Pittonia 5: 110. 1903.
"Witch Creek", Alderson, June 1894.

Rubus Ganderi Bailey, Gent. Herb. 5: 893. 1945.

"North Peak, Cuyamaca Mountains", Gander A208.

364 San Diego Society of Natural History

Rubus parviflorus forma villosus Fassett, Ann. Missouri Bot. Gard. 28: 324.

1941.

"Near head of Nigger Grade, Palomar Mt.", Gander 6239, August 4,
1938. Specimens of this collection were referred by Fassett to four formae
belonging to two varieties.

LEGUMINOSAE

Amorpha occidentalis Abrams, Bull. N. Y. Bot. Gard. 6: 394. 1910.

"San Diego River, near the Old San Diego Mission", Abrams 3425, May
6, 1903.

Astragalus agninus Jepson, Man. Fl. PI. Calif. 577. 1925.
"Borrego Sprs.", Jepson 8883.

Astragalus coccineus (Parry) T. S. Brandegee.

A. Purshii var. coccineus Parry, West. Am. Sci. 7: 10. 1890.

"Western borders of the Colorado Desert", Orcutt 1514. According to
Jepson (1936, p. 359) the type locality is Mountain Springs.

Astragalus Douglasii var. perstrictus (Rydberg) Munz & McBurney.
Phaca perstricta Rydberg, N. Am. Fl. 24: 344. 1929.

"In a hill valley between Campo and Jocumba", Abrams 3636, May 28,
1903.

Astragalus lentiginosus var. borreganus M. E. Jones, Contr. West. Bot.
8: 3. 1898.
"Borregos Springs", Orcutt.

Astragalus leucopsis var. lonchus M. E. Jones, Rev. N. Am. Astrag. 119.
1923.
"San Diego", Jones 3083.

Astragalus oocarpus A. Gray, Proc. Am. Acad. 6: 213. 1864.
"Mountains east of San Diego", Parry.

Astragalus pomonensis M. E. Jones, Contr. West. Bot. 10: 59. 1902.
"On the tableland of Fallbrook", Jones, 1882.

Astragalus Vaseyi S. Watson, Proc. Am. Acad. 17: 370. 1882.
"Mountain Springs". Vasey, 1880.

Cercis nephrophylla Greene, Repert. Sp. Nov. 11: 111. 1912.

"Some uncertain mountain district in southwestern California, in San Diego
County", Palmer; Vasey, 1875.

Hosackia argophylla A. Gray, Mem. Am. Acad. ser. 2. 5: 316. 1854.
"San Isabel, California, on rocks", Thurber, May 1852.

Higgins — Type Localities in San Diego County 365

Hosackia micrantha Nuttall in Torrey & Gray, Fl. N. Am. 1: 324. 1838.
"Near Monterey", Nuttall, 1836. According to Abrams (1944, p. 551)
this is an error, the specimen probably having been collected at San Diego.

Hosackia prostrata Nuttall in Torrey & Gray, Fl. N. Am. 1: 325. 1838.
"Plains near the sea; St. Diego, and St. Barbara", Nuttall, 1836.

Krameria parvifolia var. imparata Macbride, Contr. Gray Herb. 56: 52.
1918.
"Mountain Springs", Spencer 763.

Lathyrus splendens Kellogg, Proc. Calif. Acad. 7: 90. 1876.
"Southern California", Hutchings.

Lathyrus strictus Nuttall in Torrey & Gray, Fl. N. Am. 1: 276. 1838.
"Bushy places around St. Diego", Nuttall, April 1836.

Lotus confinis Greene, Erythea 1: 258. 1893.

"In the mountains of San Diego Co., Calif., near the United States and
Mexican boundary", Alder son, 1893.

Lotus Spencerae Macbride, Contr. Gray Herb. 53: 13. 1918.

"Stony slopes, Mountain Springs", Spencer 561, March 18, 1917.

Lupinus albifrons var. medius Jepson, Fl. Calif. 2: 252. 1936.
"Mountain Springs grade (at summit)", Jepson 11815.

Lupinus Brittonii Abrams, Bull. N. Y. Bot. Gard. 6: 391. 1910.
"Cottonwood Valley", Abrams 3904, June 5, 1903.

Lupinus Chamissonis var. longifolius S. Watson, Bot. Calif. 1: 117. 1876.

"San Diego", Cleveland. According to Jepson (1936, p. 253) this is the
type locality.

Lupinus densiflorus var. austrocollium C. P. Smith, Bull. Torrey Club
45: 200. 1918.

"Near St. Mary's hospital, San Diego", Abrams 3465, May 12, 1903.
Apparently St. Joseph's Hospital (now Mercy Hospital) was meant.

Lupinus micranthus var. microphyllus S. Watson, Proc. Am. Acad. 8: 535.
1873.

"San Diego", Nuttall, 1836.

Lupinus sparsiflorus var. inopinatus C. P. Smith, Bull. Torrey Club
47: 499. 1920.
"San Diego", T. S. Brandegee, AprH 1903.

Lupinus truncatus Nuttall ex Hooker & Arnott, Bot. Beechey Voy. 336.
1840.

"San Diego", Nuttall, 1836.

366 San Diego Society of Natural History

Phaca Deanei Rydberg, N. Am. Fl. 24: 355. 1929.
"Sweetwater Valley", Deane, May 13, 1883.

Phaca pseudoocarpa Rydberg, N. Am. Fl. 24: 343. 1929.
"Cuiamaca Mountains", Palmer 68, 1875.

Pickeringia montana var. tomentosa (Abrams) Johnston.

Xylothermia montana subsp. tomentosa Abrams, Bull. Torrey Club 34: 263.

1907.

"Near El Nido", Abrams 3530, May 20, 1903.

Psoralea rigida Parish, Bull. Torrey Club 19: 91. 1892.
"Dry hills, Oak Grove", S. B. Parish 643, June 1882.

Thermopsis macrophylla var. senota Jepson, Fl. Calif. 2: 245. 1936.
"Spencer Valley", Alderson.

Trifolium anodon Greene, Pittonia 5: 107. 1903.
"San Diego", T. S. Brandegee.

Trifolium decodon Greene, Pittonia 5: 108. 1903.
"San Diego", T. S. Brandegee, May 20, 1903.

OXALIDACEAE

Oxaiis californica (Abrams) R. Knuth.

Xanthoxalis californica Abrams, Bull. Torrey Club 34: 264. 1907.

"Onofre Mountains", Abrams 3274, April 19, 1903.

RUTACEAE

Cneoridium dumosum (Nuttall) Hooker f.

Pitavia dumosa Nuttall in Torrey 8C Gray, Fl. N. Am. 1: 215. 1838.

"St. Diego", Nuttall, 1836.

EUPHORBIACEAE

Croton californicus var. tenuis (S. Watson) Ferguson.
Croton tenuis S. Watson, Proc. Am. Acad. 14: 297. 1879.

"Potrero", Cleveland 836, 1876.

Euphorbia cinerascens var. appendiculata Engelmann, Bot. Mex. Bound.

186. 1859.

"San Felipe", Thurber 628, May 1852. Although two other collections
also were cited, this one is the type according to Wheeler (1941, p. 188).

Euphorbia hirtula Engelmann ex S. Watson, Bot. Calif. 2: 74. 1880.

"Talley's in the Cuyamaca Mountains", Palmer, July 1875. This is the
type locality according to Abrams (1951, p. 39).

Higgins — Type Localities in San Diego County 367

Euphorbia misera Bentham, Bot. Voy. Sulphur 51. 1844.
"San Diego", Hinds, 1839; also San Quentin.

Euphorbia Palmeri Engelmann ex S. Watson, Bot. Calif. 2: 75. 1880.
"Talley's Ranch in the Cuyamaca Mountains", Palmer 450, July 1875.

Euphorbia patellifera J. T. Howell, Leafl. West. Bot. 1: 53. 1933.

"Near Palm Wash", Howell 3488. According to a letter from Mr. Howell,
the locality is 5 miles west of highway U. S. 99, in the first wash north of Palm
Wash, 35 highway miles south of Indio.

Stillingia linearifolia S. Watson, Proc. Am. Acad. 14: 297. 1879.

"Near Boundary Monument, San Diego", Palmer 449, September 1875;
also San Bernardino.

CALLITRICHACEAE

Callitriche marginata var. longipedunculata (Morong) Jepson.
C. longipedunculata Morong, Bull. Torrey Club 18: 236. 1891.
"On mesas, San Diego", Orcutt, 1884.

BUXACEAE

Simmondsia californica Nuttall, Lond. Jour. Bot. 3: 401. 1844.

"Covering the sides of barren hills, in argillaceous soils, near the sea, in
the vicinity of St. Diego", Nuttall, 1836.

LIMNANTHACEAE

Limnanthes versicolor var. Parishii Jepson, FI. Calif. 2: 412. 1936.
"Stonewall Mine, Cuyamaca Mts.", S. B. Parish 4416.

ANACARDIACEAE

Rhus integrifolia (Nuttall) Bentham & Hooker.

Styphonia integrifolia Nuttall in Torrey & Gray, Fl. N. Am. 1 : 220. 1838.

"On the margins of cliffs, &c. near the sea, around St. Diego &C St. Bar-
bara; common", Nuttall, 1836.

Schmaltzia cruciata Greene, Leafl. Bot. Obs. 1: 139. 1905.

"Hot Springs in the northern part of San Diego Co.", Palmer, 1875.
According to McVaugh (1956, p. 225) this apparently is Warners Hot
Springs.

Styphonia serrata Nuttall in Torrey & Gray, Fl. N. Am. 1 : 220. 1838.
"With the preceding [S. integrifoliay, Nuttall, 1836.

368 San Diego Society of Natural History

Toxicodendron comarophyllum Greene, Leafl. Bot. Obs. 1: 120. 1905.
"Tighe's, near San Diego", Palmer, July 1875.

RHAMNACEAE

Adolphia calif ornica S. Watson, Proc. Am. Acad. 11: 126. 1876.

"At Solidad and Chollas Valley, near San Diego, and near Monterey",
Parry, Cleveland, Palmer. As pointed out by Abrams (1910, p. 415) the last-
named station is unquestionably an error.

Ceanothus austromontanus Abrams, Bull. N. Y. Bot. Gard. 6: 412. 1910.
"Coniferous forests, between Julian and Cuiamaca", Abrams 3966, July 1,
1903.

Ceanothus cyaneus Eastwood, Proc. Calif. Acad. ser. 4. 16: 361. 1927.
"Lakeside", Philbrook, April 1920.

Ceanothus divaricatus var. laetiflorus Jepson, Man. Fl. PI. Calif. 620.
1925.

"Palomar Mt. (Pala Mission)". Jepson 8494.

Ceanothus Orcuttii Parry, Proc. Davenp. Acad. 5: 194. 1889.

"High mountains east of San Diego", Orcutt, May and July 1889. Ac-
cording to Orcutt (1909, col. 548) the locality is Cuyamaca.

Ceanothus Xotayensis McMinn, Ceanothus 273. 1942.
"Otay Mountain", Jensen 401, January 20, 1935.

Ceanothus Palmeri Trelease, Proc. Calif. Acad. ser. 2. 1: 109. 1888.

"Mountains of Southern California", Palmer 42, 1875. According to
Abrams (1951, p. 71) the type locality is Cuyamaca Mountains.

Ceanothus tomentosus var. olivaceus Jepson, Man. Fl. PI. Calif. 621.
1925.

"Clevenger Canon, Ramona", Jepson 8509.

Ceanothus verrucosus Nuttall in Torrey & Gray, Fl. N. Am. 1: 267. 1838.
"Low hills near the coast, St. Diego", Nuttall, 1836.

Condalia Parryi (Torrey) Weberbauer.
Zizyphus Parryi Torrey, Bot. Mex. Bound. 46. 1859.

"Gravelly ravines near San Felipe", Parry, June 1850.

Rhamnus crocea var. pilosa Trelease ex Curran, Proc. Calif. Acad. ser. 2.
1:251. 1888.

"Santa Maria Valley, in the mountains back of San Diego." According
to Wolf (1938, p. 39) the type specimen is labeled "Nuevo, M. K. Curran,
July, 1885". Nuevo is an old name for Ramona, which is in the Santa Maria
Valley.

Higgins— Type Localities in San Diego County 369

VITACEAE

Vitis Girdiana Munson, Proc. Soc. Prom. Agr. Sci. 8: 59. 1887.
"San Diego Co.", Gird; also Los Angeles County.

MALVACEAE

Malvastrum densiflorum var. viscidum (Abrams) Estes.
M. viscidum Abrams, Bull. Torrey Club 34: 264. 1907.

"Dry hillsides near El Nido", Abrams 3528, May 19, 1903.

Malvastrum fasciculatum (Nuttall) Greene.

Malva jasciculata Nuttall in Torrey & Gray, Fl. N. Am. 1 : 225. 1838.

"St. Barbara". Nuttall, 1836. According to Abrams (1910, p. 417) "Nut-
tail's specimens are like the form occurring at San Diego, where he probably
obtained his specimens, as the species has not since been found at Santa
Barbara.

Saviniona suspensa Greene, Leafl. Bot. Obs. 2: 162. 1911.
"San Diego", Vasey, 1889.

Sphaeralcea purpurea Parish ex Jepson, Man. Fl. Pi. Calif. 635. 1925.
"Mountain Spr."; also Coyote Well.

STERCULIACEAE

Fremontia calif ornica var. diegensis M. Harvey, Madrono 7: 108. 1942.
"Bottom of Vallecito Canyon, Laguna Mountains", Munz 9716, May 17,
1925.

CISTACEAE

Helianthemum scoparium var. Aldersonii (Greene) Munz.
H. Aldersonii Greene, Erythea 1: 259. 1893.

"Mountains of the southern borders of San Diego Co., Calif., among
rocks in hard and sterile granitic soil", Alderson, June 1893.

CACTACEAE

Bergerocactus Emoryi (Engelmann) Britton & Rose.

Cereus Emoryi Engelmann, Am. Jour. Sci. ser. 2. 14: 338. 1852.

"On dry hills near the sea shore, about the boundary line", Parry, 1850.

Cereus? californicus (Nuttall) Torrey & Gray, Fl. N Am. 1 : 555. 1840.

"Arid hills and denuded tracts near St. Diego, California, common"
Nuttall, 1836.

370 San Diego Society of Natural History

Echinocactus limitus Engelmann in Coulter, Contr. U. S. Nat. Herb. 3 : 374.
1896.
"Boundary line south of San Diego", G. M. Hitchcock, 1876.

Echinocactus viridescens var. cylindraceus Engelmann, Am. Jour. Sci.
ser. 2. 14: 338. 1852.
"Near San Felipe", Parry.

Echinocereus Engelmannii (Parry) Riimpler.

Cereus Engelmannii Parry ex Engelmann, Am. Jour. Sci. ser. 2. 14: 338. 1852.
"Mountains about San Felipe, on the eastern declivity of the Cordilleras",
Parry.

Ferocactus viridescens (Nuttall) Britton & Rose.

Echinocactus? viridescens (Nuttall) Torrey & Gray, Fl. N. Am. 1: 554. 1840.
"Arid hills &c. near St. Diego", Nuttall, 1836.

Mammillaria incerta Parish in Jepson, Fl. Calif. 2: 549. 1936.

"Vallecito", Parish 450.
Mammillaria tetrancistra Engelmann, Am. Jour. Sci. ser. 2. 14: 337. 1852.

"From San Diego to the junction of the Gila with the Colorado", Parry.

Opuntia acanthocarpa subsp. Ganderi Wolf, Occ. Pap. Rancho Santa Ana
Bot. Gard. 2: 75. 1938.
"3 mi. below the old Vallecito Stage Station". Wolf 9424, June 12, 1938.

Opuntia demissa Griffiths, Missouri Bot. Gard. Rep. 22: 29. 1912.
"East of San Diego", Griffiths 9647, April 2, 1909.

Opuntia echinocarpa var. Parkeri (Engelmann) Coulter, Contr. U. S. Nat.

Herb. 3: 446. 1896.

"San Diego County, California, east side of mountains facing desert", /. C.
Parker, September 1879. The collector's name was originally given as "C. F.
Parker", perhaps through confusion with Charles F. Parker (1820-1883), con-
cerning whom see Ewan (1950, p. 277). According to Dr. Lyman Benson,
however, a slip with the type specimen reads "Campo, on the eastern slope
toward the desert, J. C. Parker, Sept. 1879".

Opuntia Fosbergii Wolf.

O. Bigelovii var. Hoffmannii Fosberg in Pierce & Fosberg, Bull. So. Calif.

Acad. 32: 121. 1933.

"Alluvial fan at the mouth of Cane Brakes Canyon, at the eastern base
of the Laguna Mountains", Fosberg 8602.

Opuntia Parryi Engelmann, Am. Jour. Sci. ser. 2. 14: 339. 1852.
"Eastern slope of the California mountains, near San Felipe", Parry.

Opuntia prolifera Engelmann, Am. Jour. Sci. ser. 2. 14: 338. 1852.
"San Diego, on arid hills and in dry creek beds", Parry.

Higgins — Type Localities in San Diego County 371

Opuntia serpentina Engelmann, Am. Jour. Sri. ser. 2. 14: 338. 1852.
"Dry hillsides, San Diego", Parry.

ONAGRACEAE

Clarkia delicata (Abrams) Nelson & Macbride.
Godetia delicata Abrams, Bull. Torrey dub 32: 539. 1905.

"Frequent on shady slopes between Potrero and Campo", Abrams 3710,
June 3, 1903.

Epilobium Palmeri Leveille, Repert. Sp. Nov. 5: 98. 1908.
"Cuimaca Mts.", Palmer 5368, June 27, 1875.

Gayophytum lasiospermum Greene, Pittonia 2 : 164. 1891.
"Near Julian", Dunn, August 1888.

Godetia epilobioides (Nuttall) S. Watson.

Oenothera epilobioides Nuttall in Torrey & Gray, Fl. N. Am. 1: 511. 1840.
"St. Diego", Nuttall, May 1836.

Oenothera bistorta Torrey & Gray, FI. N. Am. 1: 508. 1840.
"St. Diego", Nuttall, 1836.

Oenothera leptocarpa Greene.

Eulobus calif ornicus Nuttall in Torrey & Gray, FI. N. Am. 1: 515. 1840.

"Bushy plains near St. Diego", Nuttall, April 1836.

Oenothera spiralis var. linearis Jepson, Man. FI. PI. Calif. 684. 1925.
"Sunnyside", Hall 3908.

Oenothera trichocalyx var. cineracea Jepson, Man. Fl. PI. Calif. 681. 1925.
"Borrego Spr.", T. S. Brandegee.

UMBELLIFERAE

Apiastrum angustifolium Nuttall in Torrey & Gray, Fl. N. Am. 1: 644.
1840.

"St. Diego", Nuttall, April 1836.

Apiastrum angustifolium var. tenellum Nuttall in Torrey & Gray, FI. N.
Am. 1: 644. 1840.

Apparently with the preceding.

Apiastrum latifolium Nuttall in Torrey 8C Gray, Fl. N. Am. 1: 644. 1840.
"With the preceding [A. angustifoliumy, Nuttall, April 1836.

Eryngium Parishii Coulter & Rose, Contr. U. S. Nat. Herb. 7: 57. 1900.
"Oceanside", S. B. Parish 4436, 1897.

372 San Diego Society of Natural History

Euryptera lucida Nuttall in Torrey & Gray, Fl. N. Am. 1 : 629. 1840.
"Woods of St. Diego", Nuttall, April 1836.

Peucedanum Pringlei Coulter 8C Rose, Bot. Gaz. 13: 209. 1888.
"San Diego county". Pringle, Parry, Vasey.

Sanicula arguta Greene ex Coulter & Rose, Contr. U. S. Nat. Herb. 7: 36.
1900.

"Hills near San Diego", Pringle, 1882.

Velaea arguta (Nuttall) Coulter & Rose.

Deweya arguta (Nuttall) Torrey & Gray, FI. N. Am. 1: 641. 1840.

"Woods of St. Diego", Nuttall, April 1836.

Velaea arguta var. ternata Coulter & Rose, Bot. Gaz. 14: 282. 1889.
"Cuyamaca Mountains", Orcutt, July 1889.

GARRYACEAE

Garrya Veatchii var. Palmeri (S. Watson) Eastwood.

G. flavescens var. Palmeri S. Watson, Bot. Calif. 1 : 276. 1876.

"Milquatay, 60 miles from San Diego, on the Fort Yuma road", Palmer,
1875.

LENNOACEAE

Pholisma arenarium Nuttall ex Hooker, Hook. Ic. 7: pi. 626. 1844.

"Monterey and San Diego", Nuttall, 1836. As pointed out by Jepson
(1939, p. 17) this plant has not since been found at Monterey, and the citation
probably is an error.

ERICACEAE

Arctostaphylos Clevelandii A. Gray, Proc. Am. Acad. 12: 61. 1876.
"Potrero", Cleveland, September 20, 1876.

Arctostaphylos drupacea (Parry) Macbride.
A. Pringlei var. drupacea Parry, Bull. Calif. Acad. 2: 495. 1887.
"Mountains east of San Diego", Orcutt 543, September 1886.

Arctostaphylos glandulosa var. adamsii Munz, Aliso 4: 95. 1958.

"Dry granitic slopes and benches, north end of Laguna Mts., on highway
to Julian, at 5000 feet", Munz and Balls 17958, August 20, 1952.

Arctostaphylos glandulosa var. australis Adams, Jour. Elisha Mitchell
Soc. 56: 51. 1940.

"West slopes of Otay Mountain", Adams 974a.

Higgins — Type Localities in San Diego County 373

Arctostaphylos glandulosa var. crassifolia Jepson, Madrono 1: 86. 1922.
"Del Mar", Jepson 1606a.

Arctostaphylos otayensis Wieslander & Schreiber, Madrono 5: 43. 1939.
"Otay Mountain", Jensen, Wieslander.

Comarostaphylis diversifolia (Parry) Greene.

Arctostaphylos arguta var. diversifolia Parry, Proc. Davenp. Acad. 4: 35. 1884.
"Jamul Valley", Sanjord. Other localities were mentioned, but according
to Abrams (1910, p. 431) this is the type locality.

Xylococcus bicolor Nuttall, Trans. Am. Philos. Soc. ser. 2. 8:259. 1843.

"Monterey", Nuttall, 1836. As pointed out by Abrams (1910, p. 431)
the species is not known in the Monterey region, and NuttalPs specimen doubt-
less came from San Diego.

PRIMULACEAE

Dodecatheon Clevelandii Greene, Pittonia 1: 213. 1888.

"Dry hills and mesas in the southern part of California, about San Diego
and San Bernardino."

PLUMBAGINACEAE

Limonium mexicanum Blake, Rhodora 18: 59. 1916.
"San Diego", Palmer 216, 1876.

GENTIANACEAE

Swertia Parryi (Torrey) Kuntze.

Frasera Parryi Torrey, Pacif. Railr. Rep. 4: 126. 1857.

"Mountains east of San Diego", Parry.

ASCLEPIADACEAE

Asclepias californica Greene.

Acerates tomentosa Torrey, Bot. Mex. Bound. 160. 1859.

"Mountains east of San Diego", Parry, June 1850; "San Isabel", Thurber,
1852.

Gonolobus californicus Jepson, Man. Fl. PL Calif. 771. 1925.
"Ironwood Well [Yaqui Well]", T. S. Brandegee.

CONVOLVULACEAE

Convolvulus aridus subsp. longilobus Abrams, Contr. Dudley Herb. 3: 358.
1946.
"San Diego", Stokes, June 1895.

374 San Diego Society of Natural History

Convolvulus aridus subsp. tenuifolius Abrams, Contr. Dudley Herb.
3: 359. 1946.
"Hills near Bernardo (San Dieguito)", Abrams 3363, May 1, 1903.

Dichondra occidentalis House, Muhlenbergia 1 : 130. 1906.
"San Diego", Orcutt, January 7, 1884.

POLEMONIACEAE

Eriastrum densifolium subsp. austromontanum (Craig) Mason.
Gilia densijolia var. austromontana Craig, Bull. Torrey Club 61: 391. 1934.
"Dry slope near Nellie, Palomar Mts.", Munz 8341, 1924.

Fenzlia concinna Nuttall, Proc. Acad. Phila. 4: 12. 1848.
"Near Santa Diego", Nuttall, May 1836.

Gilia caruifolia Abrams, Bull. Torrey Club 32: 540. 1905.

"Cuyamaca Mountains, between Cuyamaca Lake and Oriflamme Canon",
Abrams 3940, June 28, 1903.

Gilia floribunda A. Gray, Proc. Am. Acad. 8: 267. 1870.

"California, probably on S. E. borders", Coulter 454. Other specimens
also were cited, but according to Mason (in Abrams 1951, p. 431) this is the
type. It is likely that Coulter collected this specimen in San Diego County on
his trip to the Colorado River in 1832.

Gilia inconspicua var. diegensis Munz, Man. So. Calif. Bot. 599. 1935.
"Under pines, at 5000 ft., Laguna Mts.", Munz 9694.

Gilia lutea var. longistylis Munz, Man. So. Calif. Bot. 599. 1935.
"San Diego", Baker 1608.

Gilia truncata Davidson, Bull. So. Calif. Acad. 22: 72. 1923.

"Near Jacumba", Kessler, April 1923 (Davidson 3572). According to
a letter from Theodore Payne, this specimen was collected by Kessler.

Linanthus androsaceus subsp. luteolus (Greene) Mason.
L. luteolus Greene, Erythea 3: 121. 1895.

"Cuyamaca Mountains", Vasey, June 1880. Another specimen also was
cited, but according to Mason (in Abrams 1951, p. 430) this is the type.

Navarretia densifolia var. jacumbana Brand, Annu. Conserv. & Jard. Bot.
Geneve 15-16: 340. 1913.
"Desert slopes, Jacumba", Abrams 3640, May 29, 1903.

Navarretia hamata subsp. foliacea (Greene) Mason.
N. foliacea Greene, Pittonia 1: 138. 1887.

"San Diego", Orcutt; also Potrero. According to Mason (in Abrams
1951, p. 451) the type locality is San Diego.

Higgins — Type Localities in San Diego County 375

Navarretia macrantha Brand, Pflanzenreich 4(250) : 154. 1907.

"Hugel zwischen Foster und Ramona", Abrams 3765; also San Bernardino
and San Luis Obispo Counties.

HYDROPHYLLACEAE

Eriodictyon crassifolium Bentham, Bot. Voy. Sulphur 35. 1844.
"San Diego", Barclay, October 1839.

Eriodictyon trichocalyx var. lanatum (Brand) Jepson.

E. calif ornicum subsp. australis var. lanatum Brand, Pflanzenreich 4(251) : 142.

1913.

"Zwischen Campo und Jacumba", Abrams 3632.

Eutoca speciosa Nuttall, Jour. Acad. Phila. ser. 2. 1:158. 1848.
"Near St. Diego", Nuttall, 1836.

Nemophila rotata Eastwood, Bull. Torrey Club 28: 159. 1901.

"Near San Diego", Eastwood, February 28, 1891. Of the two collections
cited by Eastwood, this one, according to Hoover (in Jepson 1943, p. 230),
should be considered the type.

Phacelia Aldersonii Greene, Pittonia 5 : 22. 1902.
"Witch Creek", Alder son, 1893.

Phacelia californica var. patula (Brand) Jepson.

P. magellanica forma patula Brand, Univ. Calif. Publ. Bot. 4: 219. 1912.

"Stonewall mine, 4,600 ft. alt. in the Cuyamaca Mts.", S. B. Parish 4423.

Phacelia eremica Jepson, Man. FI. PI. Calif. 823. 1925.
"Rocky canon walls, . . . Collins Valley", Jepson 8852.

Phacelia leucantha Lemmon ex Greene, Pittonia 1: 175. 1888.
"Del Mar", Mr. and Mrs. Lemmon, April 1888.

Phacelia Parryi Torrey, Bot. Mex. Bound. 144. 1859.
"Mountains east of San Diego", Parry, June 1850.

Phacelia polystachya Greene, Pittonia 5 : 19. 1902.
"Witch Creek", Alder son, 1893.

Phacelia whitlavia var. Jonesii forma gracillima Brand, Pflanzenreich
4(251): 71. 1913.
"Fallbrook", Jones 3099.

Pholistoma racemosum (Nuttall) Constance.

Nemophila racemosa Nuttall ex A. Gray, Proc. Am. Acad. 10: 315. 1875.

"San Diego", Nuttall, 1836.

376 San Diego Society of Natural History

BORAGINACEAE*

Allocarya echinacea Piper, Contr. U. S. Nat. Herb. 22: 88. 1920.

"University Heights, San Diego", T. S. Brandegee, April 12, 1902
(Baker 825).

Allocarya inornata Piper, Contr. U. S. Nat. Herb. 22: 106. 1920.
"Ramona", T. S. Brandegee, May 23, 1903 (Baker 3380) .

Amsinckia albicarpa Suksdorf, Werdenda 1 : 90. 1931.
"Colorado Desert", Spencer 251a, May 30, 1917.

Amsinckia angustata Suksdorf, Werdenda 1: 83. 1931.

"Cottonwood Valley", Orcutt 3263, April 5, 1889. Although several other
localities were given, this one, cited first, is the type locality according to Jepson
(1943, p. 325).

Amsinckia caduca Suksdorf, Werdenda 1: 79. 1931.
"San Diego", T. S. Brandegee, March 10, 1903.

Amsinckia curvata Suksdorf, Werdenda 1: 83. 1931.

"Warrens Campo", Eastwood 9419, April 22, 1920. Several other locali-
ties also were cited, but according to Jepson (1943, p. 325) this one, cited first,
is the type locality.

Amsinckia decumbens Suksdorf, Werdenda 1: 86. 1931.

"Grassy places in park, San Diego", Spencer 169, February 22, 1916.

Amsinckia deltoidea Suksdorf, Werdenda 1: 110. 1931.

"Between Jacumba and Mountain Springs", Eastwood 9520, April 24,
1920. Although other localities were cited, this one is the type locality according
to Jepson (1943, p. 322).

Amsinckia diversifolia Suksdorf, Werdenda 1: 79. 1931.

"La Jolla", F. E. and E. S. Clements, March 1, 1914; "San Diego",
Jepson 6666.

Amsinckia Jonesii Suksdorf, Werdenda 1: 69. 1931.
"Fall Brook", Jones 3112, March 25, 1882.

Amsinckia laxa Suksdorf, Werdenda 1: 87. 1931.

"Lakeside", K. Brandegee, June 1906; also Temescal Canyon, Riverside
County.

Amsinckia littoral is Greene ex Suksdorf, Werdenda 1: 99. 1931.
"Peninsula of San Diego", Greene, April 10, 1885.

* Suksdorf' s many segregates in Amsinckia have generally been reduced to synonymy. See
Jepson 1943, p. 318, and Abrams 1951, p. 607.

Higgins — Type Localities in San Diego County 377

Amsinckia Palmer i Suksdorf, Werdenda 1: 65. 1931.
"Southern part of San Diego County", Palmer, 1875.

Amsinckia Spencer ae Suksdorf, Werdenda 1: 86. 1931.
"Colorado Desert", Spencer 251a in part, May 30, 1917.

Cryptantha Clevelandii Greene, Pittonia 1: 117. 1887.

"Shaded places along streamlets in the hills back of San Diego", Cleveland
and Greene, April 1885.

Cryptantha Clevelandii var. florosa Johnston, Contr. Gray Herb. ser. 2.
74: 95. 1925.
"Roadside, Linda Vista", Macbride and Payson 797.

Cryptantha costata Brandegee, Bot. Gaz. 27: 453. 1899.

"Borregos Springs, Colorado desert", T. S. Brandegee, April 18, 1895.

Cryptantha Ganderi Johnston, Jour. Arnold Arb. 20: 386. 1939.

"Near school at Borego Valley, Larrea-Franseria association, 500 ft. alt.",
Gander 5328, April 15, 1938.

Cryptantha micrantha var. lepida (A. Gray) Johnston.

Eritrichium micranthum var. lepidum A. Gray, Syn. Fl. N. Am. 2: 193. 1878.

"San Diego Co.", Cleveland, 1876. Probably from the Laguna or Cuya-
maca Mountains according to Abrams (1951, p. 578).

Pectocarya setosa var. aptera Johnston, Contr. Gray Herb. ser. 2. 70: 38.
1924.
"Dry canyon floor near Campo", Abrams 3571, 1903.

Plagiobothrys allocaryoides Brand, Repert. Sp. Nov. 20: 47. 1924.
"San Diego", Jones 3072.

Plagiobothrys californicus var. gracilis Johnston, Contr. Gray Herb. ser. 2.
68: 73. 1923.

"San Diego", T. S. Brandegee 1658.

VERBENACEAE

Verbena Abramsii Moldenke, Am. Midi. Nat. 24: 750. 1940.

"Hot Springs, in the southern part of San Diego Co.", Palmer 309, 1875.

LABIATAE

Acanthomintha ilicifolia A. Gray.

Calamintha ilicifolia A. Gray, Proc. Am. Acad. 8: 368. 1872.

"California, probably Lower California", Rich. According to Abrams
(1951, p. 635) the type locality is San Diego.

378 San Diego Society of Natural History

Lepechinia Ganderi Epling, Brittonia 6: 363. 1948.

"Otay Mountain, 3000 feet", Epling, Robison, and Gander.

Monardella lanata Abrams, Muhlenbergia 8: 39. 1912.

"Descanso grade, near the top, Cuiamaca mountains", K. Brandegee, July
19, 1906.

Monardella lanceolata var. microcephala A. Gray, Syn. FI. N. Am. ed. 2.
2: 459. 1886.

"Potrero", Orcutt.

Monardella linoides A. Gray, Proc. Am. Acad. 11: 101. 1876.
"Mountains east of San Diego", Palmer, 1875.

Monardella linoides var. viminea (Greene) Munz.
M. viminea Greene, Pittonia 5: 85. 1902.

"Mountains of San Diego Co.", Vasey, 1880.

Monardella macrantha A. Gray, Proc. Am. Acad. 11: 100. 1876.

"Cuiamaca Mountains and near Julian City", Cleveland, Palmer. Accord-
ing to Jepson (1943, p. 433) the type is Palmer 295 from the Cuyamaca
Mountains.

Monardella macrantha var. Hallii Abrams, Muhlenbergia 8: 29. 1912.
"Palomar mountain", Hall 1936, May 1901.

Monardella nana A. Gray, Proc. Am. Acad. 11: 101. 1876.

"Mountains behind San Diego", Cleveland. Near Talley's according to
Abrams (1951, p. 650).

Monardella nana subsp. leptosiphon Abrams.

Monardella villosa var. leptosiphon Torrey, Bot. Mex. Bound. 129. 1859.

"San Felipe", Parry, June 1850. According to Abrams (1951, p. 650) the
type probably is from the mountains west of San Felipe canyon along the old
San Diego to Fort Yuma road.

Monardella sanguinea Greene, Pittonia 5 : 86. 1902.
"Near Julian", Dunn, 1881 and 1888.

Pogogyne Abramsii J. T. Howell, Proc. Calif. Acad. ser. 4. 20: 119. 1931.
"Mesa north of San Diego", Abrams 3446.

Pogogyne nudiuscula A. Gray, Bot. Calif. 1 : 597. 1876.
"Near San Diego", Cleveland.

Salvia Clevelandii (A. Gray) Greene.

Audibertia Clevelandii A. Gray, Proc. Am. Acad. 10: 76. 1874.

"Mountains behind San Diego, California, at the elevation of about 2,200
feet", Cleveland. Near Potrero according to Epling (1938, p. 119).

Higgins — Type Localities in San Diego County 379

Salvia eremostachya Jepson, Man. Fl. PI. Calif. 870. 1925.
"Indian Canon, Collins Valley", Jepson 8847.

Salvia Palmeri (A. Gray) Greene.

Audibertia Palmeri A. Gray, Bot. Calif. 1: 601. 1876.

"Near Tighes Ranch in the mountains northeast of San Diego", Palmer,
July 1875.

Salvia Vaseyi (Porter) Parish.
Audibertia Vaseyi Porter, Bot. Gaz. 6: 207. 1881.
"Mountain Springs", Vasey 500, June 1880.

Satureja Chandleri (T. S. Brandegee) Jepson.
Calamintha Chandleri T. S. Brandegee, Zoe 5: 195. 1905.
"Mount San Miguel", Chandler, May 21, 1904.

Scutellaria Bolanderi subsp. austromontana Epling, Madrono 5: 58. 1939.
"Ad rivulum Carrizo dictum prope Lake Henshaw", Gander 2739, July 10,
1936.

Scutellaria linearifolia Eastwood, Bull. Torrey Club 30: 493. 1903.
"San Diego", Fisher 586, June 1876.

Trichostema Parishii Vasey, Bot. Gaz. 6: 173. 1881.
"San Diego County", Parish and Vasey.

SOLANACEAE

Lycium californicum Nuttall ex A. Gray, Bot. Calif. 1: 542. 1876.

"Near San Diego, on clay-hill slopes", Nuttall, 1836; also Cooper, Cleve-
land.

Nicotiana Clevelandii A. Gray, Syn. Fl. N. Am. 2: 242. 1878.

"In dry bed of streams, Chollas Valley near San Diego", Cleveland; also
Palmer 267, 1875.

SCROPHULARIACEAE

Antirrhinum Coulterianum subsp. Orcuttianum (A. Gray) Pennell.
A. Orcuttianum A. Gray, Bot. Gaz. 9: 54. 1884.

"Near San Diego, and also in adjacent parts of Lower California", Orcutt.

Antirrhinum Nuttallianum Bentham in DeCandolle, Prod. 10: 592. 1848.
"S. Diego", Nuttall, 1836.

Castilleja oblongifolia A. Gray, Syn. Fl. N. Am. 2: 296. 1878.
"Southern borders of San Diego Co.", Palmer.

380 San Diego Society of Natural History

Collinsia concolor Greene, Erythea 3: 49. 1895.

"Presumably from the southern part of San Diego Co.", 'R. D. Anlerson.
The collector's name doubtless is a misprint for that of R. D. Alderson, many
of whose collections from San Diego County were described by Greene.

Cordylanthus filifolius Nuttall ex Bentham in DeCandolle, Prod. 10: 597.
1846.
"San Diego", Nuttall, 1836.

Cordylanthus maritimus Nuttall ex Bentham in DeCandolle, Prod. 10: 598.
1846.
"San Diego", Nuttall, 1836.

Mimulus aridus (Abrams) Grant.

Diplacus aridus Abrams, Bull. Torrey Club 32: 540. 1905.

"Dry rocky ridges at Jacumba, near the boundary monument", Abrams
3656, May 31, 1903.

Mimulus Clevelandii Brandegee, Gard. & For. 8: 134. 1895.

"South side of Cuyamaca Peak . . . not far from the signal station on its
summit", T. S. Brandegee, July 7, 1894.

Mimulus diffusus Grant, Ann. Missouri Bot. Gard. 11: 254. 1924.
"Palomar", Jepscn and Hall 1959, May 29, 1901.

Mimulus Grantianus Eastwood, Proc. Calif. Acad. ser. 4. 20: 153. 1931.
"Campo ... in sandy soil amid the brush", Eastwood 9442, April 23, 1920.

Mimulus latidens (A. Gray) Greene.

M. inconspicuus var. latidens A. Gray, Syn. Fl. N. Am. ed. 2. 2: 450. 1886.

"Flanks of Monte Diable", Brewer, Greene; "Chollas Valley", Orcutt
1179, June 20, 1884. According to Grant (1925, p. 202) the Brewer specimen
is the type; but according to Abrams (1951, p. 704) the Orcutt specimen is
the type.

Mimulus paniculatus Greene, Leafl. Bot. Obs. 1 : 190. 1906.
"Witch Creek", Alderson, May 1894.

Mimulus puniceus (Nuttall) Steudel.

Diplacus puniceus Nuttall, Bot. Mag. 65: pi. 3655. 1838.

"Sandy loam by the borders of small winter streams . . . near St. Diego",
Nuttall, 1836.

Orthocarpus falcatus Eastwood, Bull. Torrey Club 32: 212. 1905.

"Smith or Palomar Mountain, at an elevation of 1500-1800 meters", S. B.
Parish, June 1-4, 1891.

Higgins — Type Localities in San Diego County 381

Penstemon Bridgesii Xheterophyllus Hall, Univ. Calif. Publ. Bot. 6: 169.
1915.

"Vicinity of Nellie, a stage station on Palomar Mountain", Valentien,
July 15, 1910.

Penstemon spectabilis Thurber ex A. Gray, Pacif. Railr. Rep. 4: 119. 1856.
"San Pasqual", Thurber, according to Keck (Abrams 1951, p. 756).

Penstemon ternatus Torrey ex A. Gray, Bot. Mex. Bound. 115. 1859.
"Mountains east of San Diego", Parry.

PLANTAGINACEAE

Plantago ob versa Morris, Bull. Torrey Club 28: 121. 1901.
"Del Mar", Angier 21, May 1894.

RUBIACEAE

Galium angustifolium Nuttall ex Torrey & Gray, FI. N. Am. 2: 22. 1841.
"St. Diego", Nuttall, 1836.

Galium angustifolium var. siccatum (Wright) Hilend & Howell.
G. siccatum Wright, Zoe 5: 54. 1900.

"Del Mar", T. S. Brandegee; also other localities. According to Hilend and
Howell (1935, p. 155) Del Mar is the type locality.

Galium Nuttallii A. Gray.

G. suffruticosum Nuttall in Torrey & Gray, Fl. N. Am. 2: 21. 1841.

"St. Diego", Nuttall, 1836.

CAPRIFOLIACEAE

Lonicera subspicata var. denudata Render, Missouri Bot. Gard. Rep.
14: 176. 1903.

"San Diego", Thurber 558, May 1852; Cleveland, 1874; Palmer 120, 1875;
Orcutt, June 20, 1884; also Santo Tomas hills, Baja California, Mexico. Accord-
ing to McMinn (1951, p. 543) Thurber's specimen is the type.

VALERIANACEAE

Aligera patelliformis Suksdorf, West Am. Sci. 12: 53. 1901.

"Auf feuchten oder nassen Platzen, Stonewall Mine, Cuyamaca-Gebirg,
Meereshbhe 4600 F.", Parish 4539, June 1897.

382 San Diego Society of Natural History

CUCURBITACEAE

Cucurbita palmata S. Watson, Proc. Am. Acad. 11: 137. 1876.

"Cajon Valley", Cleveland; "Larkin's Station near the Jacumba Moun-
tains", Palmer, August 1875.

CAMPANULACEAE

Githopsis filicaulis Ewan, Rhodora 41: 312. 1939.

"Mission Canyon, San Diego", Orcutt, May 8, 1884.

Githopsis specularioides subsp. Candida Ewan, Rhodora 41: 308. 1939.

"Gravelly burn in chaparral, 6 mi. n. of Santa Ysabel", Munz 9806, May
19, 1925.

LOBELIACEAE

Downingia concolor var. brevior McVaugh, Mem. Torrey Club 19: 20.
1941.

"Cuyamaca Lake", Abrams 3851.

Downingia pulchella var. arcana Jepson, Madrono 1 : 100. 1922.
"La Mesa", Jepson 6678.

Nemacladus glanduliferus Jepson, Man. Fl. PL Calif. 975. 1925.
"Wagon Wash, near Sentenac Canon", Jepson 8766.

Nemacladus ramosissimus Nuttall, Trans. Am. Philos. Soc. ser. 2. 8: 254.
1842.
"In sandy soils, near St. Diego", Nuttall, 1836.

COMPOSITAE

Achillea californica Pollard, Bull. Torrey Club 26: 369. 1899.
"Sea coast at Santa Ysabel", Hensbaw, May 1893.

Ambrosia pumila (Nuttall) A. Gray.

Franseria pumila Nuttall, Trans. Am. Philos. Soc. ser. 2. 7: 344. 1841.

"Near St. Diego", Nuttall, 1836.

Aromia tenuifolia Nuttall, Trans. Am. Philos. Soc. ser. 2. 7: 396. 1841.
"Near the coast of St. Diego", Nuttall, May 1836.

Artemisia Palmeri A. Gray, Proc. Am. Acad. 11: 79. 1876.
"Jamuel Valley", Palmer, June 1875.

Aster Orcuttii Vasey & Rose, Bot. Gaz. 16: 113. 1891.
"Cariso Creek Wash", Orcutt.

Higgins — Type Localities in San Diego County 383

Baccharis sorothroides A. Gray, Proc. Am. Acad. 17: 211. 1882.

"Southern borders of California, San Diego Co., near the old Mission
station, the boundary monument, &c", Hayes, Palmer.

Baeria aristata (Nuttall) Coville.

Ptilomeris aristata Nuttall, Trans. Am. Philos. Soc. ser. 2. 7: 382. 1841.
"Near St. Diego", Nuttall, April 1836.

Baeria chrysostoma var. gracilis forma crassa Hall, Univ. Calif. Publ.
Bot. 3: 172. 1907.

"Ocean Beach", K. Brandegee, May 1906.

Baeria Clevelandii A. Gray, Proc. Am. Acad. 19: 22. 1883.
"Near San Diego", Cleveland, 1874.

Bidens californica Nuttall, Trans. Am. Philos. Soc. ser. 2. 7: 368. 1841.
"St. Diego and St. Barbara", Nuttall, 1836; also Chile.

Brickellia arguta var. odontolepis Robinson, Mem. Gray Herb. 1: 103.
1917.

"In desertis Colorado nominatis", Orcutt, 1889.

Brickellia frutescens A. Gray, Proc. Am. Acad. 17: 207. 1882.
"Mountain Springs", Vasey, 1880.

Calais Parryi A. Gray, Pacif. Railr. Rep. 4: 112. 1857.
"Near San Diego", Parry.

Calycoseris Wrightii var. calif ornica Brandegee, Zoe 5 : 155. 1903.
"In sand, near San Felipe", T. S. Brandegee.

Chaenactis glabriuscula var. tenuifolia (Nuttall) Hall.
C. tenuifolia Nuttall, Trans. Am. Philos. Soc. ser. 2. 7: 375. 1841.
"St. Diego", Nuttall, May 1836.

Chaenactis latifolia Stockwell, Contr. Dudley Herb. 3: 128. 1940.

"Jacumba Springs", Eggleston 19747.

Chaenactis Peirsonii Jepson, Madrono 1: 259. 1929.

"Silent Canon, in the desert foothills, southeast end of the Santa Rosa
Mountains", Jepson 11708, April 14, 1927.

Chaenactis tenuifolia var. Orcuttiana Greene, West Am. Sci. 3: 157.
1887.

"Along the beaches about San Diego", Orcutt, Cleveland, Parry, and
others.

Coreopsis californica (Nuttall) H. Sharsmith.

Leptosyne californica Nuttall, Trans. Am. Philos. Soc. ser. 2. 7: 363. 1841.
"Near St. Diego", Nuttall, May 1836.

384 San Diego Society of Natural History

Coreopsis maritima (Nuttall) Hooker f.

Tuckermannia maritima Nuttall, Trans. Am. Philos. Soc. ser. 2. 7: 363. 1841.

"On shelving rocks, near the sea, at St. Diego", Nuttall, 1836.

Corethrogyne brevicula Greene, Leafl. Bot. Obs. 2: 26. 1910.
"Mountains of San Diego Co.", Orcutt, October 1899.

Corethrogyne filaginifolia var. linifolia Hall, Univ. Calif. Publ. Bot. 3: 71.

1907.

"About one kilometer south of Del Mar ... in hard siliceous soil on an
exposed windswept bluff overlooking the sea", K. Brandegee, August 5, 1906.

Corethrogyne filaginifolia var. pacifica Hall, Univ. Calif. Publ. Bot.

3: 73. 1907.

"Pacific Beach . . . just back from the beach ... in rich, loose soil along
a railroad embankment", Pur pus, summer of 1899.

Corethrogyne incana Nuttall, Trans. Am. Philos. Soc. ser. 2. 7: 290. 1841.
"Near St. Diego", Nuttall, May 1836.

Corethrogyne racemosa Greene, Leafl. Bot. Obs. 2: 26. 1910.
"Mountains of San Diego Co.", Orcutt, 1899.

Dysodia porophylloides A. Gray, Mem. Am. Acad. ser. 2. 5: 322. 1855.
"Sandy hills, near San Felipe", Thurber, May 1852.

Encelia farinosa A. Gray ex Torrey in Emory, Notes Mil. Rec. 143. 1848.

Although the locality of collection was not definitely stated, this species
was mentioned in the narrative (p. 103) under the date of November 28 and
29, 1846, when the party was in the vicinity of "Cariso (cane) creek". This
locality was therefore taken by Abrams (1910, p. 480) as the type locality.

Eriophyllum Wallacei var. rubellum (A. Gray) Jepson.
Bahia rubella A. Gray, Bot. Mex. Bound. 95. 1859.

"In a dry valley, near San Felipe", Parry, June 1850.

Franseria Palmeri Rydberg, N. Am. FI. 33: 25. 1922.
"San Diego", Palmer, 1875.

Geraea viscida (A. Gray) Blake.
Encelia viscida A. Gray, Proc. Am. Acad. 11: 78. 1876.
"Near Larkens' Station", Palmer, August 1875.

Gnaphalium bicolor Bioletti, Erythea 1: 16. 1893.

"Common in low thickets among the coast hills about San Diego", Greene?

Gnaphalium microcephalum Nuttall, Trans. Am. Philos. Soc. ser. 2. 7: 404.
1841.

"St. Diego", Nuttall, May 1836.

Higgins — Type Localities in San Diego County 385

Grindelia Hallii Steyermark, Ann. Missouri Bot. Gard. 21: 229. 1934.
"Open meadows about Cuyamaca Lake", Abrams 3957, June 30, 1903.

Haplopappus junceus Greene, Bull. Calif. Acad. 1: 190. 1885.

"San Diego County", Cleveland, Curran; also Baja California, Mexico.

Haplopappus propinquus Blake.
Bigelovia brachylepis A. Gray, Bot. Calif. 1 : 614. 1876.
"Larkens' Station", Palmer, August 1875.

Haplopappus venetus var. decumbens (Greene) Munz.
Isocoma decumbens Greene, Leafl. Bot. Obs. 1: 172. 1906.

Greene gave no locality, but he cited only one specimen, C. F. Baker's
3405, collected by T. S. Brandegee. According to Hall (1928, p. 227) this
came from clay depressions on the mesas near San Diego.

Haplopappus venetus var. oxyphyllus (Greene) Munz.
Isocoma oxyphylla Greene, Leafl. Bot. Obs. 1: 171. 1906.
"Jamul Valley", Palmer, June 1875.

Helianthus gracilentus A. Gray, Proc. Am. Acad. 11: 77. 1876.
"In the mountains 45 miles north-east of San Diego", Palmer.

Hemizonia conjugens Keck, Aliso 4: 109. 1958.

"River-bottom land near Otay", Abrams 3521, May 16, 1903.

Hemizonia fasciculata var. ramosissima (Bentham) A. Gray.
H. ramosissima Bentham, Bot. Voy. Sulphur 30. 1844.

"San Diego", Hinds, October 1839.

Hemizonia floribunda A. Gray, Proc. Am. Acad. 11: 79. 1876.

"California, near the southern boundary, on the Fort Yuma Road, 80
miles east of San Diego", Palmer.

Hemizonia tenella (Nuttall) A. Gray.

Osmadenia tenella Nuttall, Trans. Am. Philos. Soc. ser. 2. 7: 392. 1841.
"St. Diego", Nuttall, May 1836.

Holocarpha virgata subsp. elongata Keck, Aliso 4: 111. 1958.
"San Diego", Keck 1932, October 2, 1932.

Hulsea calif ornica Torrey & Gray, Bot. Mex. Bound. 98. 1859.

"Mountains east of San Diego ... in bushy places", Parry, June 1850.

Hymenopappus wrightii var. viscidulus Jepson, Man. FI. PI. Calif. 1128.
1925.

"Cuyamaca Mts.", T. S. Brandegee.

386 San Diego Society of Natural History

Isocoma leucanthemifolia Greene, Leafl. Bot. Obs. 1: 171. 1906.

"Warren, in the Mountains of San Diego Co.", Orcutt, October 21, 1889.
According to Hall (1928, p. 227) the type is from Warner's Ranch.

Iva Hayesiana A. Gray, Proc. Am. Acad. 11: 78. 1876.

"Jamuel Valley", Palmer, June 1875. Gray also cited a specimen collected
by Hayes at Warners Pass in 1858, but he said that it was indeterminable, the
heads having all fallen from their peduncles.

Machaeranthera hiemalis A. Nelson, Am. Jour. Bot. 21: 580. 1934.
"Devil's Canyon, near Jacumba", Nelson 11190, March 14, 1930.

Machaeranthera lagunensis Keck, Brittonia 9: 238. 1957.

"From 2 miles south of the main recreation area in the Laguna Moun-
tains ... at 5200 ft. elevation ... on dry slopes under pines", Munz and
Balls 17948, August 20, 1952.

Madraglossa carnosa Nuttall, Trans. Am. Philos. Soc. ser. 2. 7: 393. 1841.
"St. Diego . . . growing in the sands of the sea-coast", Nuttall, 1836.

Malacothrix Clevelandii A. Gray, Bot. Calif. 1: 433. 1876.
"Near San Diego", Cleveland; also Baja California.

Microseris breviseta Greene, Pittonia 5 : 8. 1902.
"San Diego", Greene, April 1885.

Microseris heterocarpa (Nuttall) K. Chambers.

Uropappus heterocarpus Nuttall, Trans. Am. Philos. Soc. ser. 2. 7: 425. 1841.
"St. Diego", Nuttall, 1836.

Microseris Parishii Greene, Bull. Calif. Acad. 2: 46. 1886.

"Near San Luis Rey", S. B. and W. F. Parish 955, April 1881; "near
San Diego". Greene, 1885; also near Tulare. According to Chambers (1955,
p. 291) the Parish specimen is the type.

Microseris platycarpha A. Gray.

Calais platycarpha A. Gray, Pacif. Railr. Rep. 4: 113. 1857.

"San Luis Rey", Parry, 1850.

Pentachaeta aurea Nuttall, Trans. Am. Philos. Soc. ser. 2. 7: 336. 1841.

"In dry plains near the sea, in the vicinity of St. Diego", Nuttall, April
1836.

Perityle Emoryi Torrey in Emory, Notes Mil. Rec. 142. 1848.
"Cordilleras of California", Emory, 1846.

Porophyllum caesium Greene, Leafl. Bot. Obs. 2: 155. 1911.
"Cajon Hills", Dunn, May 1, 1891.

Higgins — Type Localities in San Diego County 387

Porophyllum Vaseyi Greene, Leafl. Bot. Obs. 2: 154. 1911.
"Mountain Springs", Vasey, 1880.

Ptilomeris anthemoides Nuttall, Trans. Am. Philos. Soc. ser. 2. 7: 382.
1841.

"Near St. Diego", Nuttall, 1836.

Ptilomeris coronaria Nuttall, Trans. Am. Philos. Soc. ser. 2. 7: 382. 1841.
"Near St. Diego", Nuttall, 1836.

Ptilomeris mutica Nuttall, Trans. Am. Philos. Soc. ser. 2. 7: 382. 1841.
"Near St. Diego", Nuttall, 1836.

Rafinesquia californica Nuttall, Trans. Am. Philos. Soc. ser. 2. 7: 429.
1841.

"Near the sea-coast, in the vicinity of St. Diego", Nuttall, 1836.

Solidago confinis A. Gray, Proc. Am. Acad. 17: 191. 1882.
"Southern borders of California", Palmer, Cleveland, Parish.

Sonchus fallax? var. californicus Nuttall, Trans. Am. Philos. Soc. ser. 2.
7: 438. 1841.

"Around St. Diego", Nuttall, 1836.

Sonchus tenuifolius Nuttall, Trans. Am. Philos. Soc. ser. 2. 7: 438. 1841.
"Round St. Diego, in shady ravines, among rocks", Nuttall, 1836.

Stephanomeria exigua var. deanei Macbride, Contr. Gray Herb. 53: 22.
1918.

"Sweetwater Valley". Deane, July 23, 1888.

Viguiera deltoidea var. Parishii (Greene) Vasey & Rose.
V. Parishii Greene, Bull. Torrey Club 9: 15. 1882.

"San Luis Rey", S. B. and W. F. Parish 963, April 1881.

Viguiera laciniata A. Gray, Bot. Mex. Bound. 89. 1859.

"Rancho Gamacha [ = Jamacha]", Schott, September 1855.

Wyethia coriacea A. Gray, Proc. Am. Acad. 11: 77. 1876.

"On the Mesa Grande, 70 miles north-east of San Diego", Palmer, July
1875.

Wyethia ovata Torrey & Gray in Emory, Notes Mil. Rec. 143. 1848.

"Western side of the Cordilleras of California", Emory, December 4,
1846. According to Abrams (1905, p. 541) the specimens evidently were col-
lected between Warners Ranch and Santa Isabel.

388 San Diego Society of Natural History

NOTES CONCERNING THE COLLECTORS

An attempt is made to identify all those mentioned in the Systematic List
as collectors of type specimens in San Diego County. Recent and well-known
collectors are briefly noted, with references to published biographical material
when known. Information about most current and recent professional botanists
also may be found in American Men of Science (Cattell 1955 and earlier
editions) , though it has not seemed necessary to repeat this reference for each
individual. A special effort has been made to learn about the earlier and more
obscure collectors and their botanical work in the County.

Abrams, LeRoy (1874-1956)

Dr. Abrams was professor of biology at Stanford University and an
authority on the flora of the western United States. In the spring and summer
of 1903 he collected extensively in San Diego County in connection with his
doctoral problem (Abrams 1910). For a portrait and brief tribute, see Baci-
galupi and Mason (1954), and for a sketch of his life and work, see Wiggins
(1957).

Adams, Joseph Edison (1903- )

Dr. Adams is professor of botany at the University of North Carolina.
For a doctoral thesis at the University of California, under Professor Jepson,
he wrote a revision of Arctostaphylos.

Alderson, Rufus Davis (1858-1932)

Mr. Alderson was a newspaperman and teacher who came to California
in 1884 or soon after and returned to his native West Virginia in 1897.
Between 1889 and 1897 he taught school in San Diego County, his posts
including Potrero, Descanso, El Nido, Warner, Glencoe, Santa Ysabel, Spring
Hill, and Witch Creek. From 1893 to 1896 he collected some thousands of
botanical specimens in various parts of the County but especially about Witch
Creek. He sent many duplicates for identification to E. L. Greene and to S. B.
Parish. Other duplicates he sold or exchanged.

Angier, Belle Sumner (Mrs. Walter L. Burn) (1870-1948)

Mrs. Burn's interest was primarily horticultural, but she collected botanical
specimens for her cousin Professor C S. Sargent. At his instance and under
his direction, she propagated seedlings of the Torrey Pine which were distributed
by him. She wrote for the papers, urging the preservation of the Torrey Pine,
and thus helped initiate the interest leading to the ultimate establishment of the
Torrey Pines Preserve.

Antisell, Thomas (1817-1893)

Dr. Antisell was geologist on the Pacific Railroad Survey under Lt. J. G.
Parke from 1854 to 1856. The party reached San Diego from central Cali-
fornia in April 1855, and in May they started for Arizona. For a portrait and
biographical sketch, see Merrill (1906).

Higgins — Type Localities in San Diego County 389

Baker, Charles [or Carl] Fuller (1872-1927)

Professor Baker was an untiring and intensive collector, both botanical and
entomological. He was at Pomona College for several years (1903-1904,
1908-1912) as assistant professor and professor. There he was especially active
in entomology, though his botanical collections also were very rich. From
Pomona he went to the Philippines, where he remained until his death. For
an account of his life, see Essig (1927).

Balls, Edward Kent (1892- )

Mr. Balls is horticulturist at the Rancho Santa Ana Botanic Garden and
a botanical collector of worldwide experience.

Barclay, George

Mr. Barclay was a botanical collector sent on the voyage of H. M. S.
Sulphur to obtain plants and seeds for "his Majesty's garden at Kew". See
under Hinds.

Bestor, Norman

Mr. Bestor was botanical collector on the expedition of the Army of the
West in 1846. See under Emory. As remarked by Ewan (1950, p. 163), Lt.
Emory has always received credit for the collections.

Bolander, Henry Nicholas] (1831-1897)

Dr. Bolander was state botanist of California, succeeding William Brewer
in 1864. In 1871 he became state superintendent of schools, and in this
capacity he was able to visit every section of the state. In 1873 he visited
San Diego, the Cuyamaca Mountains, and the desert at San Felipe. A tribute
was paid him by Jepson ( 1898) .

Brandegee, [Mary] Katharine [Layne] [Curran] (1844-1920)
Brandegee, T[ownshend] S[tith] (1843-1925)

During his earlier years while employed as a civil engineer, Mr. Brandegee
had a lively interest in botany, which later crystallized as a life-time career.
Mrs. Brandegee was curator of botany in the California Academy of Sciences
from 1883 until 1894. In 1894 the Brandegees moved to San Diego, where
they lived until 1906. In this period they collected widely throughout the
County and beyond. In 1906 they moved to Berkeley, where they staid until
their deaths. An account of the^ lives of these two botanists with complete
bibliography was given by Setchell (1926).

Bullard, Frances E. ( ? -1932)

Mrs. Bullard (nee Schmidt) was the wife of Dr. J. H. Bullard, a friend
of Dr. Anstruther Davidson, Los Angeles botanist. On trips with her husband,
Mrs. Bullard collected for Dr. Davidson, who described her findings in the
Bulletin of the Southern California Academy of Sciences. An ardent horticul-
turist, she was the originator of many hybrid Watsonias, put on the market
for her by Theodore Payne. For notes concerning her, see Payne (1954).

390 San Diego Society of Natural History

Burbeck, Anna Leora (1861-1935)

Miss Burbeck, a native of Sherborn, Massachusetts, came to National City
in 1871. Like her friend Miss Kimball, q.v., she collected and pressed native
ferns. She also made some 150 water-color paintings of wild flowers. In 1887
she married Albert Carleton Copeland.

Chandler, H[arley] P[ierce] (1875-1918)

Mr. Chandler was a teacher with a strong avocational interest in botany.
From 1904 to 1905 he was principal of Russ High School in San Diego, and
while here he collected extensively in the County. For a biographical sketch,
with portrait, see Jepson (1929b).

Clements, Frederick E[dward] (1874-1945)
Clements, Edith [Gertrude] S[chwartz] (1877- )

Dr. Frederick Clements, the well-known ecologist, was Professor of Botany
at the University of Nebraska and the University of Minnesota. From 1917
until his retirement in 1941 he was research associate in ecology with the
Carnegie Institution of Washington. For accounts of his life, with portraits
and bibliography, see Shantz (1945) and Pool (1954). Dr. Edith Clements
was her husband's assistant and botanical illustrator. She now lives in La Jolla.

Cleveland, Daniel (1838-1929)

Daniel Cleveland came to San Diego in 1869 and remained for the rest
of his life. Actively engaged in the practice of law, he made the study of botany
his lifelong avocation. He collected widely in the County; and his collections,
sent to Asa Grey, Sereno Watson, and Edward Lee Greene, became the basis
for several new species. He was a charter member of the San Diego Society of
Natural History and its president from 1888 to 1890 and in 1893 and 1894.
The Society inherited the Cleveland collection, an herbarium of some 3000 speci-
mens. For a biographical sketch, with portrait, see Jepson (1929a). Further
details of his life may be found in the San Diego Union, January 4, 1929.

Cooper, James Graham (1830-1902)

Dr. Cooper, after whom the Cooper Ornithological Club was named, is
best known for his pioneer work in western ornithology and mammalogy. How-
ever, he was very active in other branches of natural history, including botany.
He wrote on the vegetation of the Cuyamaca Mountains (1874) . For details of
his life, see Emerson (1898, 1902), Taylor (1919), and Grinnell (1930).

Coulter, Thomas (1793-1843)

Dr. Coulter, an Irishman, was a keen botanist and a prodigious collector.
In 1832 he travelled overland from Monterey to the Colorado River and back,
passing Pala, Warners Pass, and Carrizo, and also making a side trip to San
Diego. He was thus the first botanist to cross the Colorado Desert. His types
of several desert plants must have been collected on this trip, but it is not
known whether they are from San Diego County or from farther east. For
notes on his life, and on this trip in particular, see Coville (1895) and Mc
Kelvey (1955).

Curran, [Mary] Katharine see Brandegee, Katharine

Higgins — Type Localities in San Diego County 391

Deane, George Clement (1854-1930)

Mr. Deane, a native of Cambridge, Massachusetts, came to California
with a partner in the late 1870's and started a vineyard at Bonita, in the
Sweetwater Valley. Most of his later life was spent in Cambridge, but he
returned twice to California. Though primarily interested in birds, he collected
many botanical specimens for his brother Walter Deane, a New England
botanist. For a biographical note, see Allen (1930).

Dunn, G[eorge] Washington] (1814-1905)

Mr. Dunn was a professional collector of natural history materials, includ-
ing plants. He is said to have lived in San Diego for many years. A record
of his activities, with a portrait, was given by Jepson (1934d).

Eastwood, Alice (1859-1953)

Miss Eastwood served as curator of botany in the California Academy of
Sciences for 57 years, from 1892 to 1949, when at the age of 90 she became
curator emeritus. She collected widely, her trips including six to San Diego
County (Wilson 1953). Several of her collections became the types of new
species. A biographical sketch, with bibliography and portrait, appeared in a
volume celebrating her fiftieth year with the California Academy of Sciences
(MacFarland et al. 1949) . A full biography is by Wilson (1955) and a sketch
by Dakin (1954). A tribute to her is from the heart of John Thomas Howell
(1954).

Eggelston, W[illard] W[ebster] (1863-1935)

Mr. Eggelston was botanist with the U. S. Forest Service and Bureau of
Plant Industry from 1910 until his retirement in 1933.

Emory, W[illiam] H[emsley] (1811-1887)

In 1846 Lt. Emory, as topographical engineer, accompanied the Army of
the West from Fort Leavenworth to San Diego on a march culminating in the
Battle of San Pasqual. Entering what is now San Diego County late in Novem-
ber, they passed Carrizo, Warners Ranch, Santa Ysabel, and the Soledad River,
reaching San Diego December 12th. Emory (1848) recounted the details of
this memorable expedition, and to his report was appended Dr. Torrey's catalog
of the plants collected. See under Bestor. For further information, see Mc-
Kelvey (1955). Later Emory was chief astronomer, then commissioner, of the
United States and Mexican Boundary Survey. For an account of his life, see
Carey (1931).

Epling, Carl C[lawson] (1894- )

Dr. Epling is professor of botany at the University of California at Los
Angeles. His specialty is the Labiatae, upon which he has published widely.

Everett, Percy C[harles] (1902- )

Mr. Everett is superintendent of the Rancho Santa Ana Botanic Garden.

Fisher, William J.

In the San Francisco directory from 1866 to 1880, Mr. Fisher was listed
variously as "bookkeeper", "curiosities", and "master mariner", but mosdy as
"naturalist". In 1876 he was librarian of the California Academy of Sciences.

392 San Diego Society of Natural History

Forbes, Charles N[oyes] (1883-1920)

After high school in National City, San Diego County, Mr. Forbes grad-
uated from the University of California. He became assistant botanist, then
curator of botany, at the Bishop Museum, where he remained until his death.
For a biographical sketch, see Gregory (1921, p. 9).

Fosberg, F[rancis] Raymond (1908- )

Dr. Fosberg is botanist for the U. S. Geological Survey. As an under-
graduate at Pomona College and while assistant botanist at the Los Angeles
County Museum, he collected extensively in southern California.

Froebel, Julius (1805-1893)

A political refugee from Germany because of his participation in the Revo-
lution of 1848, Mr. Froebel came to this country and settled near San Francisco
in 1854. He became a member of the California Academy of Sciences. For
biographical notes, see Ewan (1955, p. 15, 49).

Fultz, Francis M[arion] (1857-1948)

Mr. Fultz was a writer, lecturer, and educator, in his later years connected
with the city schools in Santa Barbara and Los Angeles. During this period he
published several books on plant life, the best known of which is "The Elfin
Forest" (Fultz 1920).

Gambel, William (1821-1849)

Mr. Gambel was a friend and protege of Thomas Nuttall, who sent him to
California to collect natural history specimens. Primarily an ornithologist, he
was according to Palmer (1928) the first to spend any length of time in Cali-
fornia. His botanical collections, though apparently small, are important. He
came overland late in 1841. Details of his itineraries in California and his date
of departure are unknown. McKelvey (1955, p. 737) suggested that the San
Diego specimens attributed to him possibly were collected by Nuttall. For
notes on his travels, see McKelvey (1955).

Gander, Frank F[orest] (1899- )

Mr. Gander was curator of botany in the Natural History Museum, San
Diego, from 1934 to 1942. During part of this time he was also supervisor
of nature study in the County schools; and while travelling to visit the schools,
he collected intensively throughout the County. He now maintains a nursery
of native California plants near Escondido.

Gentry, Howard Scott (1903- )

Dr. Gentry is botanist with the Section of Plant Introduction of the U. S.
Department of Agriculture. He has collected widely in Mexico. He has a
special interest in Agave and has many plants in cultivation at his ranch home
near Temecula.

Gird, H[enry] H[arrison] (?1827-1913)

Mr. Gird bought a large ranch near Bonsall in 1876 and moved there in
1880. He had an orchard with many kinds of fruit. For further information
about him, see the San Diego Union, September 9, 1939.

Higgins — Type Localities in San Diego County 393

Grant, George B[ernard] (1849-1917)

Mr. Grant was an amateur botanist who lived for some years in Pasadena
and collected widely in southern California. His herbarium was given to S. B.
Parish and with the Parish herbarium went to Stanford University. For a
biographical note, see Ewan (1950, p. 218).

Greene, Edward Lee (1843-1915)

Professor Greene taught at the University of California from 1885 until
1905, when he moved to Washington, D. C. For details of his life and work,
see Nieuwland (1915), Bartlett (1916), Jepson (1931), and Ewan (1950).
For his bibliography see Kistler (1936).

Griffiths, David (1867-1935)

Dr. Griffiths was an agriculturist and horticulturist with the U. S. Depart-
ment of Agriculture. He was especially interested in cacti and wrote several
articles about them.

Hall, Harvey M[onroe] (1874-1932)

Dr. Hall taught botany at the University of California from 1903 until
1919. For the rest of his life he was investigator in experimental taxonomy
for the Carnegie Institution of Washington, still working in California. He
was a meticulous taxonomist of broad vision, concerned especially with the
Compositae. For a biography and bibliography, see Babcock (1934). A story
of his life with many interesting details was given by Jaeger (1953).

Hayes, Sutton ( ? -1863)

Dr. Hayes was assistant surgeon on the El Paso and Fort Yuma Wagon
Road Expedition of 1857 to 1859. Iva Hayesiana was named "in memory of
the estimable discoverer". He later went to Panama for his health and collected
extensively there until his death. For a biographical note, see Seemann (1863).

Henshaw, Henry W[etherbee] (1850-1930)

Mr. Henshaw, sometime chief of the U. S. Biological Survey, was pri-
marily an ornithologist; but he also collected in a limited way in other fields,
"even plants". He was in southern California in 1892 to 1894, when he visited
Frank Stephens at Witch Creek. For a sketch of his life, see Nelson (1932) .

Hinds, Richard Brinsley (?1812-1847)

Dr. Hinds was surgeon-naturalist on the voyage of H. M. S. Sulphur
around the world in 1835 to 1842. They visited California in 1837 and 1839,
stopping in San Diego in October of 1839. Collections were made by Hinds
and by Mr. Barclay, a collector from Kew. The botany of the expedition was
reported by Bentham (1844-46), with an introduction by Hinds. For an
account of the expedition in western North America, see McKelvey (1955).

Hitchcock, C[harles] Leo (1902- )

Dr. Hitchcock is professor of botany at the University of Washington,
Seattle.

394 San Diego Society of Natural History

Hitchcock, George N. (1843-1907)

Mr. Hitchcock, born in Boston, came to San Diego in 1869. He was a
lawyer but was interested in plants. Together with J. C. Parker, a photographer,
he followed Parry's notes and collected Agave Shawii at the border station.
They sent specimens and photographs to Dr. Engelmann at St. Louis. For
a biographical note, with portrait, see Black (1913).

Howell, John Thomas (1903- )

Mr. Howell is curator of botany at the California Academy of Sciences
and a keen student of the California flora.

Hutchings, James M[ason] (1818-1902)

Mr. Hutchings was known for his connection with the Yosemite Valley
as guide and hotelkeeper, guardian of the state park, and author and lecturer.
For details of his career, see Russell (1947).

Jensen, H[erbert] A. (1906- )

Mr. Jensen, who was with the U. S. Forest Service from 1930 to 1948,
worked with A. E. Wieslander on the Vegetation Type Map project. He is
now a member of the forestry consulting firm of Hammon, Jensen and Wallen,
in Oakland, California.

Jepson, Willis Linn (1867-1946)

Dr. Jepson's whole life at the University of California was devoted to the
study of California botany. For 60 years, from his student days until his death
as Professor Emeritus, it was his only thought. He wrote voluminously on the
subject, and to his writings we are much indebted. He collected extensively
throughout California but did comparatively little in San Diego County. For
a biographical sketch, see Constance (1947); and for an estimate of his place
in California botany, see Keck (1948) .

Johnson, Ernest R[alph] (1890- )

Mr. Johnson was superintendent of the Rancho Santa Ana and was the
first superintendent of the Rancho Santa Ana Botanic Garden.

Jones, Marcus E[ugene] (1852-1934)

Mr. Jones was an amateur botanist who collected widely in western North
America but especially in the Great Basin. He collected comparatively little
in San Diego County. Jones (c. 1930) wrote an account of his own botanical
activities. Jepson (1934c) gave a pithy commentary on his botanical work. An
interesting and more personal account was given by Ewan (1950) and another
by Jaeger (1952).

Keck, David D[aniels] (1903- )

Dr. Keck is head curator of the New York Botanical Garden. He collected
in California while a student and while on the staff of the Stanford laboratory
of the Carnegie Institution of Washington.

Kessler, Robert

Mr. Kessler is an amateur horticulturist, formerly of Los Angeles and now
of Whittier. Apparently at one time he was interested in growing native plants,
and several of these were described as new by Dr. Anstruther Davidson.

Higgins — Type Localities in San Diego County 395

Kimball, Laura Frances (1856-1942)

Miss Kimball was a resident of National City, San Diego County, from
the time of her arrival in California in 1869. An amateur botanist, she was
especially interested in ferns.

Lemmon, John Gill (1832-1908)

Mr. Lemmon was a botanist who, together with his wife, Sara Allen Plum-
mer Lemmon, collected widely in California, Nevada, and Arizona. He was
botanist to the state forestry board in California during its brief existence from
1888 to 1892. Through his writings he did much to popularize botany and
forestry. For a sketch of his life, see Jepson (1933).

Macbride, J[ames] Francis (1892- )

Mr. Macbride is curator of Peruvian botany for the Chicago Natural
History Museum. He lives in California and may be reached through the
Dudley Herbarium, Stanford University.

Mearns, Edgar A[lexander] (1856-1916)

Dr. Mearns, a surgeon with the U. S. Army, was a prodigious collector
of zoological and botanical specimens. From 1891 to 1894, as medical officer
for the United States and Mexican Boundary Survey, he was able to explore
the boundary from El Paso to the Pacific coast, making large and important
collections. For a sketch of his life, see Miller (1933).

Moran, Reid [Venable] (1916- )

Dr. Moran is curator of botany at the Natural History Museum, San
Diego. His special interest is the Crassulaceae.

Munz, Phillip Alexander] (1892- )

Dr. Munz is director of the Rancho Santa Ana Botanic Garden. He is
an authority on the flora of California and a special student of the Onagraceae.

Nelson, Aven (1859-1952)

Dr. Nelson was professor of botany at the University of Wyoming. For
a biographical sketch, see Ewan (1950, p. 271). His portrait appears as the
frontispiece to volume seven of Madrono.

Newberry, John S[trong] (1822-1892)

Dr. Newberry was physician and naturalist on Lt. Ives' expedition to
explore the Colorado River in 1857 and 1858 (Ives 1861). In November of
1857 he crossed the desert from San Diego to Yuma by way of Santa Ysabel
and Warners Ranch. For details of his life, see Merrill (1906, 1934).

Nuttall, Thomas (1786-1859)

Of English birth, Mr. Nuttall came in 1807 or 1808 to Philadelphia,
where he became affiliated with the Academy of Natural Sciences. In 1834
he journeyed overland with the Wyeth Expedition to Oregon. Thence he
sailed by way of the Hawaiian Islands to Monterey and down the coast to San
Dego, finally taking passage for Boston on the Alert, whose voyage was
described by Richard Henry Dana in his well-known Two years before the

396 San Diego Society of Natural History

mast. Dana told of his surprise at finding in San Diego "Professor N ,

of Cambridge, [whom he had] last seen quietly seated in the chair of Botany
and Ornithology, in Harvard University". In San Diego from April 15th to
May 8th, 1836, Nuttall collected many botanical specimens, seventy-one of
which became types. NuttalPs portrait appears as frontispiece to volume two
of Madrono. For accounts of his western travels, see Jepson (1934b) and
McKelvey (1955).

Orcutt, Charles Russell (1864-1929)

Mr. Orcutt was a commercial collector of natural history materials who
made his home in San Diego. He was an indefatigable collector and in the
course of his work made many discoveries. Also he was the editor and publisher
of the West American Scientist. An account of his life was given by Jepson
(1929c).

Palmer, Edward (?1831-1911)

Dr. Palmer was a professional collector whose work was mostly in Mexico,
both on the mainland and in Baja California. Over 200 species are said to
have been named for him as collector. Although he visited San Diego County
several times, his most important collections here, including several types, were
in 1875. For details of his life and collections, see McVaugh (1956).

Parish, Samuel Bonsall (1838-1928)

After the Civil War, Mr. Parish and his brother William started west,
eventually settling in 1872 in the San Bernardino Valley, where they purchased
a ranch. He was an enthusiastic botanical collector for more than forty years,
collecting throughout southern California and being the first to explore some
areas botanically. A voluminous but careful writer, he added much to the
literature of southern California botany. His herbarium was purchased by
Stanford University. Jepson (1932) published a record of his botanical activi-
ties with a portrait and a full bibliography.

Parish, William Fletcher

Mr. Parish to some extent shared the botanical interests of his brother
Samuel and sometimes collected with him.

Parker, J. C.

Mr. Parker, a native of Cincinnati, came to San Diego in 1873 and engaged
in business as a photographer. He made not only portraits but landscape
photographs as well. See under Hitchcock, George N. For a biographical
note, see Anonymous (1890).

Parry, C[harles] C[hristopher] (1823-1890)

Dr. Parry was a physician who served as botanist and geologist with the
United States and Mexican Boundary Survey. He arrived in San Diego by
sea in July 1849 and collected in southern California until March 1851. In
1875 and later he returned to southern California and collected widely. Preston
(1893) published a biographical sketch, to which was appended a bibliography
compiled by Mrs. Parry. Tributes to his work were given by Parish (1909a)
and Jepson (1934a).

Higgins — Type Localities in San Diego County 397

Payson, Edwin Blake (1893-1927)

Dr. Payson was at the University of Wyoming, both as an undergraduate
and later as a professor of botany. For a biographical sketch, see Ewan (1950,
p. 279).

Philbrook, Myrtle (1877- )

Miss Philbrook collected Ceanothus cyaneus on her brother's ranch at
Lakeside, and she and her brother submitted it to Miss Kate Sessions. Recog-
nizing it as a new species, Miss Sessions sent it to Miss Alice Eastwood, who
named it. Miss Philbrook lives in San Diego.

Pringle, Cyrus Guernsey (1838-1911)

Mr. Pringle was a professional botanical collector best known for his
voluminous Mexican collections, distributed to herbaria all over the world.
He visited San Diego in 1882 and made a brief trip into Baja California. For
a biographical sketch, see Evans (1935); and for a full biography, see Davis
(1936).

Purpus, Carl A[lbert] (?185 1-1941)

Dr. Purpus was a botanical collector, whose Mexican collections over many
years were identified and distributed by his friend T. S. Brandegee. Between
1898 and 1912 he gave his address as First and Redwood Streets, San Diego:
this was the Brandegees' address from 1894 until 1906, when they moved to
Berkeley. From 1913 until the end of his life, his home was Hacienda Mirador,
Zacuapam Huatusco, Veracruz. For biographical notes, see Langman (1949)
and Ewan (1950, p. 285).

Rich, William

Major Rich made natural history collections on the United States and
Mexican Boundary Survey.

Robison, William C[ondit] (1914- )

As an undergraduate student at the University of California at Los
Angeles, Mr. Robison accompanied Dr. Carl Epling on several field trips
through San Diego County. He is now a geographer with the Quartermaster
Research and Engineering Center, Natick, Massachusetts.

Roezl, Benedict [or Benito] (?1823-1885)

Mr. Roezl was a horticulturist who collected widely in the Americas, intro-
ducing many kinds of plants into European cultivation and in great quantity.
He had in fact the reputation of being a greedy and ruthless collector. He
visited San Diego in 1869. For an autobiographical account with brief intro-
ductory notes, see Masters and Moore (1874, 1885); and for notes on his visit
to San Diego, see McKelvey (1938, p. 95).

Sanford, Oliver N[ason] (1847-1928)

Mr. Sanford was a civil engineer with a broad interest in natural history,
especially entomology and botany. A native of Boston, he lived in San Diego
about from 1872 to 1899, when he moved to San Francisco. He was a charter
member of the San Diego Society of Natural History.

398 San Diego Society of Natural History

schoenefeldt, ludwig

Mr. Schoenefeldt was a hospital steward in the U. S. Army, who served
in the biological section of the United States and Mexican Boundary Survey.
According to Mearns (1907, p. 6), he collected plants assiduously from Fort
Yuma to the Pacific Ocean and on San Clemente Island, from April 6 to
September 9, 1894.

Schott, Arthur [Carl Victor] (1814-1875)

A native of Germany, Mr. Schott came to the United States in 1850. As
assistant on the United States and Mexican Boundary Survey, he studied the
geology and made large botanical collections. He was with the party of Lt.
Michler, which came to San Diego by sea and on November 16, 1854, departed
for Fort Yuma. For a biographical note, see Standi ey (1920, p. 92).

Schumo, Mrs. Silas L.

According to records of the Academy of Natural Sciences of Philadelphia,
Mr. Schumo donated shell collections, from the late 1890's to 1903. Mrs.
Schumo gave at least one botanical specimen from California, in 1899. Probably
she collected the specimen of Eriogonum. No further information is available
concerning either Mr. or Mrs. Schumo.

Scott, Miss

With the type specimen of Carex prat ■gracilis, according to Dr. Reed
Rollins, there is a long note, presumably in Miss Scott's hand, comparing this
with another species and suggesting that it was new. From this note it appears
that Miss Scott must have had some knowledge of botany. No further informa-
tion about her is available.

Spencer, Mary [Evelyn] F[isk] (1841-1940)

Mrs. Spencer was an amateur botanist who collected extensively in Europe
and, in her later years, in the desert areas of San Diego County. Her herbarium
is at Oberlin College. For a report on her life and work, see Grover (1941).

Stark, B[yron] D[avid] (1894- )

Mr. Stark was a nurseryman for the Rancho Santa Ana Botanic Garden.
He is now retired and lives in Whittier.

Stokes, Susan G[abriella] (1868-1954)

Miss Stokes moved with her family to San Diego in 1886; and she taught
high school here from 1914 to 1936. From college days to the end of her life,
her main interest was the genus Eriogonum. For a sketch of her life and an
appreciation of her work, see Howell (1955).

Thurber, George (1821-1890)

From his early employment in pharmacy, Dr. Thurber became interested
in drug plants and thus took up the study of botany. As botanist, quarter-
master, and commissary, with the United States and Mexican Boundary Survey,
he came overland from Texas to San Diego, reaching what is now eastern San
Diego County in February 1852; and he began the return trip in May 1852.
For his own notes on this trip, see Gray (1855); and for a biographical sketch,
see Woodward (1936).

Higgins — Type Localities in San Diego County 399

Valentien, Anna M[arie Bookprinter] (1862-1947)

Mrs. Valentien was a noted painter and sculptress, who taught arts and
crafts in San Diego for many years. Her husband, Albert Robert Valentien
(1862-1925), after retiring as head decorator for Rookwood Pottery, Cin-
cinnati, began painting the wildflowers of California. The Valentien paintings
in the San Diego Museum of Natural History include some 1200 subjects in
watercolor. Mrs. Valentien collected specimens for her husband to paint; and
these were later sent to Dr. H. M. Hall for identification.

Vasey, George (1822-1893)

Always interested in botany, Dr. Vasey had his first real opportunity as
a botanist when he joined Major John Powell in an expedition to Colorado in
1868 (Darrah 1951). From that time botany was his sole interest. He was
botanist in the U. S. Department of Agriculture from 1872 until his death.
His special interest was the grasses. For a biographical sketch, see Maxon
(1936).

Weed

The original collector of Calochortus Weedii was given simply as "Weed".
His identity remains a mystery. The suggestion is made that he might possibly
be Charles Leander Weed, a photographer for many years resident in San
Francisco and known as the first photographer in the Yosemite (Russell 1947).
Charles Weed's name is absent from the city directory of San Francisco for
several years, including the time preceding the description of the Calochortus.
But there is no positive evidence that he was in San Diego at this time or that
he was the collector.

WlESLANDER, A[LBERT] E[VERETT] (1890- )

Mr. Wieslander is a forester with the California Forest and Range Experi-
ment Station. He collected in San Diego County in the survey of vegetation
for the Vegetation Type Map project (Wieslander 1935).

Wiggins, Ira L[oren] (1899- )

Dr. Wiggins is professor of biology and director of the Natural History
Museum at Stanford University and an authority on the flora of the Sonoran
Desert. From 1921 until 1928 he collected widely in San Diego County in
preparation for the flora of the County that was his doctoral dissertation
(Wiggins 1929).

Wolf, Carl B[randt] (1905- )

Dr. Wolf collected extensively throughout California while he was botanist
at the Rancho Santa Ana Botanic Garden, from 1930 to 1945.

Woodhouse, Samuel Washington] (1821-1904)

Dr. Woodhouse was physician and naturalist with Lt. Sitgreaves' expedi-
tion down the Zuni and Colorado Rivers in 1851. From Fort Yuma they
crossed the Colorado Desert to San Diego in December 1851.

400 San Diego Society of Natural History

OLD PLACE NAMES

A few place names appearing in the list are no longer used and will need
explanation. Daniel Cleveland's notes concerning three of these names were
published by Mason (1937). Further details have been furnished by Mr.
Bertram B. Moore of San Diego, formerly Assistant County Surveyor.

Cordilleras of California. In the report of Emory (1848) this expression
refers to the mountains along the route from San Felipe to Santa Ysabel
by way of Warners Ranch.

Larkin's (or Larken's) Station. According to Cleveland this is an old
name for Jacumba. Mr. Moore agrees, adding that it is shown as "Larkins"
on the San Diego County map of 1889 by J. D. Schuyler. According to
McVaugh (1956, p. 242), "Larkin" appears in township 17-18 south,
range 7 east, on the Immigration Association's map of California, ca. 1885.
McVaugh added that it was on the site of Buckman's Springs, but that
is in township 16 south, range 5 east. Perhaps he meant Bankhead Springs,
which is about 4 miles northwest of Jacumba.

Milquatay, according to Mr. Moore, was an Indian encampment in the
Campo Valley; it is shown on the San Diego County map of 1872 by
M. G. Wheeler, County Surveyor.

Stonewall Mine was about a mile north of Stonewall Peak in the Cuyamaca
Mountains.

Talley's (or Tally's) Ranch according to Cleveland was in a valley at the
base of South Cuyamaca Peak, about 8 miles south of Julian. But accord-
ing to Palmer's notes, as quoted by McVaugh (1956, p. 326), it was about
5 miles south of Julian, in the Stockton Valley. And according to Mr.
Moore it is shown about 4 miles south of Julian, in section 21, township
13 south, range 4 east, on the township plat of 1876 and on the San Diego
County map of 1889 by J. D. Schuyler.

Tighe's (or Tigh's) Ranch was about 7 miles east of Ramona, on the old
road from San Diego to Julian. On the township plat of 1875, according
to Mr. Moore, it is shown in section 16, township 13 south, range 2 east.
According to Cleveland it was also given on some botanical labels as
"Luckett" after an earlier owner.

ACKNOWLEDGEMENTS

I wish to express my appreciation and thanks to those who have helped in
the preparation of this paper: to Dr. Herbert L. Mason and Dr. Phillip A.
Munz, who have made available their card files of type localities; to Professor
Joseph Ewan and the many others, too many to name, who have given informa-
tion for the biographical sketches; to Mr. Bertram B. Moore, who has furnished
careful notes concerning early place names; to Mr. Wesley F. Farmer, who has
redrawn the map used as fig. 1; to Mrs. Mildred Meeder, Librarian of the
Natural History Museum, who with unlimited patience has helped me with
references in the library; and to Dr. Reid Moran, whose careful editorial work
has been unstinted.

Higgins — Type Localities in San Diego County 401

LITERATURE CITED

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Allen, Glover M.

1930 George Clement Deane. Auk 47: 456, 457.

Anonymous

1890 An illustrated history of southern California, i-viii, 9-898. Lewis
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Babcock, Ernest Brown

1934 Harvey Monroe Hall. Univ. Calif. Publ. Bot. 17: 355-368, portrait.

B[acigalupi], R. C, and H. L. M[ason]

1954 LeRoy Abrams, a tribute from the California Botanical Society.
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Bartlett, Henry Harris

1916 The botanical work of Edward Lee Greene. Torreya 16: 151-175,
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Bentham, George

1844-46 The botany of the voyage of H. M. S. Sulphur, under the com-
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Black, Samuel F.

1913 San Diego County, California; a record of setdement, organization,
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Carey, Charles F.

1931 Emory, William Hemsley. Diet. Am. Biog. 6: 153, 154.

Cattell, Jaques, Ed.

1955 American men of science: a biographical dictionary, ed. 9. II, Bio-
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Chambers, Kenton L.

1955 A biosystematic study of the annual species of Microseris. Contr.
Dudley Herb. 4: 207-312, figs. 1-22.

402 San Diego Society of Natural History

Clausen, Robert T.

1938 A monograph of the Ophioglossaceae. Mem. Torrey Club 19 (2) :
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Constance, Lincoln

1947 Willis Linn Jepson. Science 105: 614.

Cooper, J. G.

1874 The botany of the Cuyamaca Mountains. Am. Nat. 8: 90-98.

Coville, Frederick V.

1895 The botanical explorations of Thomas Coulter in Mexico and Cali-
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Dakin, Susanna Bryant

1954 The perennial adventure: a tribute to Alice Eastwood. 1-48. Cali-
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Dana, Richard Henry

1840 Two years before the mast.

Darrah, William Culp

1951 Powell of the Colorado, i-ix, 1-426, figs. 1-17. Princeton Univer-
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Davis, Helen Burns

1936 Life and work of Cyrus Guernsey Pringle. 1-756, map. 2 pis. Uni-
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Emerson, W. Otto

1898 Dr. James G. Cooper. Bull. Cooper Club 1: 1-5, portrait.
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1848 Notes of a military reconnaissance, from Fort Leavenworth, in
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1938 The Californian Salvias. A review of Salvia, section Audibertia.
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1935 Pringle, Cyrus Guernsey. Diet. Am. Biog. 15: 236, 237.

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1950 Rocky Mountain naturalists, i-xvi, 1-358, 9 pis. University of
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1955 San Francisco as a mecca for nineteenth century naturalists. In A
century of progress in the natural sciences. 1-63, illus. California
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Higgins — Type Localities in San Diego County 403

Fultz, Francis M.

1920 The elfin forest of California. 1-267, figs. 1-23. Times-Mirror
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Grant, Adele Lewis

1925 A monograph of the genus Mimulus. Ann. Missouri Bot. Gard.
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Gray, Asa

1855 Plantas novae Thurberianae. Mem. Am. Acad. ser. 2. 5: 297-328.

Gregory, Herbert E.

1921 Report of the director for 1920. Occ. Pap. Bishop Mus. 8: 1-28.

Grinnell, Joseph

1930 Cooper, James Graham. Diet. Am. Biog. 4: 406, 407.

Grover, Frederick

1941 Mary Fisk Spencer. Madrono 6: 82-84, pi. 6.

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1928 The genus Haplopappus: a phylogenetic study in the Compositae.
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Hilend, Martha, and John Thomas Howell

1935 The genus Galium in southern California. Leafl. West. Bot. 1 :
145-168.

Hitchcock, A. S.

1925 The North American species of Stipa. Contr. U. S. Nat. Herb.
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1935 Manual of the grasses of the United States. U. S. Dep. Agr.
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Hitchcock, C. Leo

1936 The genus Lepidium in the United States. Madrono 3: 265-320,
fig. 1, pis. 14-17.

Howell, John Thomas

1954 "I remember when I think . . ." Leafl. West. Bot. 7: 153-164.

1955 Susan G. Stokes, the Eriogonum lady. Leafl. West. Bot. 7: 225-
227.

Ives, Joseph C.

1861 Report upon the Colorado River of the West, explored in 1857 and
1858 by Lieutenant Joseph C. Ives, Corps of Topographical Engi-
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Jaeger, Edmund

1952 Marcus Jones. Calico Print 8 (7) : 8, 9, 25, portrait.

1953 Buckboard botanist. Calico Print 9 (4) : 14, 15, 22, 23, portrait.
Jepson, Willis Linn

1898 Dr. Henry N. Bolander, botanical explorer. Erythea 6: 100-107,

portrait.
1929a Daniel Cleveland. Madrono 1: 267, 268, portrait.

404 San Diego Society of Natural History

1929b Harley Pierce Chandler. Madrono 1: 269, 270, portrait.
1929c Charles Russell Orcutt, natural history collector. Madrono 1: 273,
274.

1931 Greene, Edward Lee. Diet. Am. Biog. 7: 564, 565.

1932 Samuel Bonsall Parish. Univ. Calif. Publ. Bot. 16: 427-444, pi. 32.

1933 Lemmon, John Gill. Diet. Am. Biog. 11: 162, 163.
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1934c Marcus Eugene Jones. Madrono 2: 152-154.

1934d George W. Dunn. Madrono 2: 156, 157, portrait.
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1943 Ibid. 3 (2): 129-464, figs. 366-451.

Jones, Marcus E.

c. 1930 Botanical exploration of Marcus E. Jones, 1875 to 1919. Typescript
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where.

Keck, David D.

1948 The place of Willis Linn Jepson in California botany. Madrono
9: 223-228.

Kistler, Ellen D.

1936 Bibliography of the botanical writings of Edward Lee Greene.
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Langman, Ida K.

1949 Dos figuras casi olvidadas en la historia de la botanica Mexicana.
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Macfarland, F. M., R. C. Miller, and John Thomas Howell

1949 Biographical sketch of Alice Eastwood. Proc. Calif. Acad. ser. 4.
25: ix-xiv, portrait.

Macfarland, F. M., and Veronica J. Sexton

1949 Bibliography of the writings of Alice Eastwood. Proc. Calif. Acad,
ser. 4. 25: xv-xxiv.

M[ason], H. L.

1937 Early California place names used by Daniel Cleveland. Madrono
4: 67.

Masters, M. T., and T. Moore

1874 Benedict Roezl. Gard. Chron. ser. 2. 2: 73, portrait.
1885 Benedict Roezl. Gard. Chron. ser. 2. 24: 521, portrait.

Maxon, William R.

1936 Vasey, George. Diet. Am. Biog. 19: 229, 230.

McKelvey, Susan Delano

1938 Yuccas of the southwestern United States. 1: 1-150, maps 1-8,
pis. 1-80.

1955 Botanical exploration of the Trans-Mississippi West, 1790-1850.
i-xl, 1-1144, 11 maps. Arnold Arboretum of Harvard University.

Higgins — Type Localities in San Diego County 405

McMinn, Howard E.

1951 An illustrated manual of California shrubs, i-xi, 1-663, figs. 1-775.
University of California Press.

McVaugh, Rogers

1956 Edward Palmer, plant explorer of the American West, i-xvii, 1-430,
2 maps, 15 pis. University of Oklahoma Press.

Mearns, Edgar Alexander

1907 Mammals of the Mexican boundary of the United States, Part. 1.
U. S. Nat. Mus. Bull. 56: i-xv, 1-530, figs. 1-126, pis. 1-13.

Merrill, George P.

1906 Contributions to the history of American geology. Rep. U. S. Nat.

Mus. 1904: 189-734, figs. 1-141, pis. 1-37.
1934 Newberry, John Strong. Diet. Am. Biog. 13: 445, 446.

Miller, Gerret S.

1933 Mearns, Edgar Alexander. Diet. Am. Biog. 12: 482, 483.

Munz, Philip A.

1946 Aquilegia: the cultivated and wild columbines. Gent. Herb. 7:
1-150, figs. 1-38.

Nelson, Edward William

1932 Henry Wetherbee Henshaw — Naturalist. Auk 49: 399-427, pis.
15-18.

NlEUWLAND, J. A.

1915 In memoriam. Am. Midi. Nat. 4: 228a, 228b, portrait.

Orcutt, Charles Russell

1908 American plants. 1 : [columns] 1-384. San Diego.

1909 Ibid. 2: [columns] 385-788.

Palmer, T. S.

1928 Notes on persons whose names appear in the nomenclature of Cali-
fornia birds. Condor 30: 261-307, figs. 78-82.

Parish, S. B.

1909a Parry and southern California botany. Plant World 12: 158-162.
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Payne, Theodore

1954 History of the hybrid Watsonia. Gold. Gard. 18 (10) : 4-6.

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1954 Frederick Edward Clements. Ecology 35: 108-112, portrait.
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1893 Biographical sketch of Dr. C. C Parry. Proc. Davenp. Acad.
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Russell, Carl Parcher

1947 One hundred years in Yosemite. i-xiii, 1-226, 52 illus. University
of California Press.

406 San Diego Society of Natural History

Seemann, Berthold

1863 Botanical news. Jour. Bot. 1 : 253-256.

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1926 Townshend Stith Brandegee and Mary Katharine (Layne) (Cur-
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Shantz, H. L.

1945 Frederick Edward Clements. Ecology 26: 317-319, portrait.

Standley, Paul C.

1920 Trees and shrubs of Mexico, Part 1. Contr. U. S. Nat. Herb. 23:
i-xviii, 1-169.

Taylor, Walter P.

1919 Notes on mammals collected principally in Washington and Cali-
fornia between the years 1853 and 1874 by Dr. James Graham
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Trelease, William

1912 The Agaves of Lower California. Missouri Bot. Gard. Rep. 1911:
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Tryon, Rolla

1956 A revision of the American species of Notholaena. Contr. Gray
Herb. 179: 1-106, maps 1-67, figs. 1-58.

Wheeler, Louis Cutter

1941 Euphorbia subgenus Chamaesyce in Canada and the United States
exclusive of southern Florida. Rhodora 43: 97-154, 168-205, 223-
286, pis. 654-668.

Wieslander, A. E.

1935 A vegetation type map of California. Madrono 3: 140-144.

Wiggins, Ira L.

1929 Flora of San Diego County, California; a phytogeographic and
taxonomic study. Unpublished thesis, Stanford University.

1957 LeRoy Abrams. Taxon 6: 61-63.

Wilson, Carol Green

1953 A partial gazeteer and chronology of Alice Eastwood's botanical

explorations. Leafl. West. Bot. 7: 65-68.
1955 Alice Eastwood's wonderland; the adventures of a botanist, i-iv,
1-222, illus. California Academy of Sciences, San Francisco.

Wolf, Carl B.

1938 The North American species of Rhamnus. Monogr. Rancho Santa
Ana Bot. Gard. 1: 1-136, figs. 1-62.

Woodward, Carl R.

1936 Thurber, George. Diet. Am. Biog. 18: 514, 515.

TRANSACTIONS

OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY
Volume XII, No. 23, pp. 407, 408 February 1, 1960

UNii

A NEW RACE OF POCKET GOPHER (THOMOMYS)

FROM SAN FERNANDO MISSION,

BAJA CALIFORNIA, MEXICO

BY

Laurence M. Huey

Curator of Birds and Mammals, San Diego Society of Natural History

The study of a small series of Pocket Gophers (Thomomys) from San
Fernando Mission, Baja California, Mexico, has revealed characters which were
not apparent in the limited number of specimens available to the writer when
he named Thomomys bottae abbotti (Huey 1928) or when he reviewed the
Pocket Gophers of Baja California (Huey 1945). The species is now known
as Thomomys umbrinus (Hall and Kelson, p. 413, 1959) .

The new race may be known as

Thomomys umbrinus brazierhowelli1 subsp. nov.
San Fernando Pocket Gopher

Type. — Adult male; from San Fernando Mission, Baja California, Mexico,
Iat. 30°; collected by A. Brazier Howell and skinned by Laurence M. Huey,
February 26, 1958; no. 18725, collection of the San Diego Society of Natural
History.

Characters. — Thomomys umbrinus brazierhowelli is a dark, dull tawny
colored gopher with a darker and, in some specimens blackish, median dorsal
stripe. It has a blackish colored face with black pouch linings. The dull tawny
color extends over the sides and underparts, while the feet and tip of the tail
are white.

Compared with Thomomys umbrinus abbotti, the race that lives along the
Rosario River to the northwest, T. u. brazierhowelli is much darker, with the
blackish dorsal color and black face and cheek pouches outstanding. The skull
of T. u. brazierhowelli differs from that of T. u. abbotti in being more angular
and rugose, with wider, square arching and heavier-boned zygomatic arches. The
bullae are more rounded and inflated, lacking the anterior flattened surfaces
found in T. u. abbotti.

Named in honor of A. Brazier Howell, a lifelong friend, charter member and past president
of the American Society of Mammalogists.

408 San Diego Society of Natural History

Compared with Thomomys umbrinus catavinensis (Huey 1931) and
Thomomys umbrinus ruricola (Huey 1949) , both of which occur to the south
and are brownish gray in color, T. u. brazierhowelli contrasts sharply: its buff
tawny color sets it apart readily. It is larger than either; and its skull is larger,
more angular, and more heavily boned.

Measurements of Type. — Total length, 235; tail, 76; hind foot, 33;
ear, 5. Skull: greatest length, 40.2; spread of maxillary arches, 25.9; length of
nasals, 14.3; interorbital constriction, 7.0; alveolar length of upper molar series,
8.2.

Range. — Known only from the type locality.

Specimens Examined.— Thomomys umbrinus abbotti, 17 from 1 mi. e.
of El Rosario, Baja California (type locality) . Thomomys umbrinus catavinensis,

12 from Catavina, Baja California (type locality) ; 2 from Rancho Ramona,

13 mi. sw. of San Agustin, Baja California. Thomomys umbrinus ruricola, 1
(the type) from 4 mi. n. of Santa Catarina Landing, Baja California. Thomomys
umbrinus brazierhowelli, 14 from San Fernando Mission, Baja California (type
locality) .

LITERATURE CITED

Hall, E. Raymond, and Keith R. Kelson

1959 The mammals of North America. The Ronald Press Co., New York,
March 31.

Huey, Laurence M.

1928 A new silky pocket mouse and a new pocket gopher from Lower Cali-
fornia, Mexico. Trans. San Diego Soc. Nat. Hist. 5 : 87-90. January
18.

1931 A new species and a new subspecies of pocket gopher. Trans. San
Diego Soc. Nat. Hist. 7: 43-46. December 19.

1945 The pocket gophers of Baja California, Mexico, with descriptions
of nine new forms. Trans. San Diego Soc. Nat. Hist. 10: 245-268,
map. August 31.

1949 Three new races of pocket gophers (Thomomys) from Baja Cali-
fornia, Mexico. Trans. San Diego Soc. Nat. Hist. 1 1 : 53-56. January
31.

01

TRANSACTIONS

OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY
Volume XII, No. 24, pp. 409-412 February 1, 1960

TWO NEW RACES OF PEROGNATHUS SPINATUS
FROM BAJA CALIFORNIA, MEXICO

BY

Laurence M. Huey

Curator of Birds and Mammals, San Diego Society of Natural History

The study of a series of Perognathus spinatus from the rocky region about
the old Mission of San Fernando, together with specimens collected years ago
from localities on the western slope of the Sierra San Pedro Martir and the
southern parts of the peninsula, reveals the presence of two undescribed races,
the descriptions of which follow:

Perognathus spinatus oribates1 subsp. nov.
San Fernando Spiny Pocket Mouse

Type. — Adult male; from San Fernando Mission, lat. 30° N., Baja Cali-
fornia, Mexico; collected by Laurence M. Huey, February 27, 1958; no. 18742,
collection of the San Diego Society of Natural History.

Characters and Comparisons. — Perognathus spinatus oribates is recog-
nizable by its very spiny and grizzled blackish dorsal pelage, darker than in other
races, and its small body, having a proportionately long, well tufted, bicolored
tail. As in all other races of P. spinatus, the underparts and feet are white, with
little or no blending of color on the sides.

Compared with Perognathus spinatus rufescens, the desert race found
living on the light-colored granite slopes of the mountains in northeastern Baja
California, P. s. oribates is much darker in dorsal pelage color, with a skull
slightly heavier-boned in most dimensions. Compared with Perognathus spinatm
prietae, the race that occupies the north central midpeninsular region, P. s.
oribates is slightly darker and more grizzled, with a proportionately longer
tail. The skull of P. s. oribates, when compared with the skulls of both these
races, is rounder as seen either from the side or from the rear and has heavier
maxillary arches and nasals; the bullae also are more rounded and inflated.

From the Greek, oribates, mountain ranging.

410 San Diego Society of Natural History

The newly described race is more closely allied in pelage color to Perognathus
spinatus peninsulae, found at the extreme southern part of the peninsula; how-
ever it is set apart from the southern relative by its smaller cranial and body size.

Measurements of Type. — Total length, 192; tail, 112; hind foot, 22;
ear, 5. Skull: greatest length, 24.9; width across bullae, 12.8; interorbital con-
striction, 6.5; nasals, 9.1; toothrow. 3.5.

Range and Remarks. — Perognathus spinatus oribates is found living on
rocky terrain from the western foothills bordering Llano de San Agustin, at
Iat. 30° N., northward along the arid foothills of the western slopes of the Sierra
San Pedro Martir to Las Cabras, inland and east of San Telmo, near lat. 31° N.

This is the westernmost race of the Perognathus spinatus group, which is an
example of a desert species whose range rounds the southern end of the Sierra
San Pedro Martir to terminate on the western slope in the San Quintin area.
Two species of pocket mice, Perognathus baileyi and Perognathus arenarius, a
kangaroo rat, Dipodomys merriami, and sandpaper plant, Petalonyx linearis, are
other examples of this distribution.

Perognathus spinatus broccus2 subsp. nov.
San Ignacio Spiny Pocket Mouse

Type. — Adult male; from San Ignacio, Iat. 27° 17' N., Baja California,
Mexico; collected by Laurence M. Huey, March 18, 1928; no. 6891, collection
of the San Diego Society of Natural History.

Characters and Comparisons. — This race, Perognathus spinatus broccus,
differs from its nearest relative to the north, P. s. prietae, in its larger size and
proportionately longer tail. The skull is flatter and less rounded. The most
pronounced character, for which the race is named, is a well developed pointed
projection on the underside of the zygomatic arch where the jugal joins the
maxilla.

Compared with the southern race, Perognathus spinatus peninsulae, P. s.
broccus is smaller, with smaller ears and smaller skull. The pointed projection
on the zygomatic arch, however, is larger and more prominently developed.

Measurements of Type. — Total length, 204; tail, 118; hind foot, 22;
ear, 5. Skull: greatest length, 25.6; width across bullae, 13.1; interorbital con-
striction, 6.6; nasals, 9.8; toothrow, 3.4.

Range. — Perognathus spinatus broccus occurs, so far as known, only over
the lava-covered slopes of the Sierra de la Giganta.

Remarks. — The accompanying table gives the average measurements of
the male specimens used in the preparation of this paper. A like tabulation
was made of female specimens, but it gave only an indication of relatively
smaller sizes for that sex.

From the Latin, broccus, projecting.

r l.

II

Perognathus spinatus

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412 San Diego Society of Natural History

SPECIMENS EXAMINED

Perognathus spinatus spinatus: 25. (CALIFORNIA: Morongo Valley, San
Bernardino County, 2; Frink, Imperial County, 6; 3 mi. n. of Bard, Imperial
County, 7; 1 mi. n. of Potholes, Imperial County, 9; Baja California: San
Felipe, 1.)

Perognathus spinatus rufescens: 13. (California: Palm Springs, River-
side County, 1; mouth of Palm Canyon, Borrego Valley, San Diego County,
2; San Felipe Canyon, San Diego County, 3; La Puerta Valley, San Diego
County, 7.)

Perognathus spinatus oribates: 17. (Baja California: Rancho Ramona,
8 mi. n. of Santa Catarina, 4; San Fernando Mission, 12; Las Cabras, 1.)

Perognathus spinatus prietae: 33. (Baja California: Catavina, 1 (not
typical); 25 mi. n. of Punta Prieta, 12; San Borjas Mission, 7; 12 mi. e. of El
Arco, 3; Santa Gertrudis Mission, 6; Barril, 4.)

Perognathus spinatus broccus: 26. (Baja California Sur: San Ignacio,
11; Llano de San Bruno, 1; 12 mi. s. of Mulege, Concepcion Bay, 2; Canipole,
1; Comondu, 11.)

Perognathus spinatus peninsulae: 30. (Baja California Sur: La Paz
Mesa, 19 mi. n. of La Paz, 2; 7 mi. nw. of San Bartolo, 11; Miraflores, 3; 7 mi.
s. of Miraflores, 1; Los Barriles, 1; San Jose del Cabo, 8; 9 mi. ne. of Cape San
Lucas, 2; Cape San Lucas, 2.)

TRANSACTIONS

OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY
Volume XII, No. 25, pp. 413420

FEE
i

COMMENTS ON THE POCKET MOUSE,

PEROGNATHUS FALLAX,

WITH DESCRIPTIONS OF TWO NEW RACES

FROM BAJA CALIFORNIA, MEXICO

BY

Laurence M. Huey

Curator of Birds and Mammals, San Diego Society of Natural History

SAN DIEGO, CALIFORNIA

Printed for the Society

February 1, 1960

TLB ZC

COMMENTS ON THE POCKET MOUSE,

PEROGNATHUS FALLAX,

WITH DESCRIPTIONS OF TWO NEW RACES

FROM BAJA CALIFORNIA, MEXICO

BY

Laurence M. Huey

Curator of Birds and Mammals, San Diego Society of Natural History

During the past thirty years there have accumulated in the collection of
the San Diego Society of Natural History representative specimens of the
Pocket Mouse, Perognathus fallax, from a number of localities ranging from
southern California to central Baja California, Mexico.

The present known range of Perognathus fallax is from latitude 34° 30' N
to latitude 27° 30' N., mainly west of the desert, but it includes both slopes of
the backbone mountains of southern California. However, so far as is now
known no specimens have been collected or records made from the eastern
slopes of the Sierra Juarez or the Sierra San Pedro Martir in northern Baja
California nor from above the 5000 foot level along the western slopes of the
mountains. At the southern extent of the present known range, the species is
found in isolated localities along the Pacific coast and at a few localities inland
where vegetative conditions which furnish food, exist.

As with other mammalian species that occupy both coastal and desert or
near-desert areas, there is shown in the assemblage of specimens of Perognathus
fallax the dark to light or pallid colored pelages. There are also variations
shown in specimens taken from localities along the coastal section of the range
from north to south. This coastal section, too, runs the gamut from a cool
humid to a hot arid climate, though the effect on the species does not parallel
that of the west-to-east climatic variation. This geographical and climatological
influence has developed not only divergent pelage color but size differences
as well. Hence in areas such as the coastal Viscaino Desert region of Baja
California, with its xerothermic plant life, the species has been dwarfed, while
inland, the populations show some characters, particularly those of pelage color
and size, of the more northern populations.

It has been of great interest to find within the ranges of other mammalian
species, whose populations are to he found living down the length of this long
peninsula, or in fact from the cooler humid northern coastal section southward
into the arid central parts, characters that parallel those shown by the Perognathus
fallax group, the subject of the present study. Dwarfing, light and dark pelage,
or brightening of color patterns are amongst the more prominent.

416 San Diego Society of Natural History

Notable changes are likewise to be found in the vegetation both on land
and in the sea. One of the most important factors to consider in the study of
this species of Perognathus is the effect of climatic conditions within its range.
For instance, rainfall and the resulting plant growth which furnishes food upon
which these rodents live, in this coastal area differ considerably from north
to south. At San Diego, California, which is not the northernmost section of
the species' range but which may be taken as a central part of the nominate race,
P. fallax, the average mean rainfall is 9.2 inches annually. Seventy miles south-
ward at Ensenada, Baja California, the average rainfall decreases slightly, though
the flora is similar to that of San Diego, and P. fallax is still to be found in typical
character.

A range of mountains of about 3000 feet elevation lying in an east-west
direction south of Ensenada, reaches the sea. The more arid south face of these
hills marks a sharply changing flora. Below this barrier the annual rainfall average
is much less, only 5.2 inches at San Quintin. However, fog is more prevalent.
In this region, where less rain falls, the vegetation is more scrubby and of
a very different type, with cacti and a dense, very thorny species of wild rose
predominating. Here a darker, robust race of P. fallax is found.

For the southernmost coastal section of the known range of the P. fallax
group, near Punta Eugenio, Baja California, where the race P. f. inopinus is
found, rainfall data are not available; but the flora of the region proves to be
a good index of the annual precipitation : it is very sparse, dwarfed, and thorny.
This is a land of seasonal fogs, with erratic and irregular rainstorms, which,
when occurring, are often short and violent. The influence of the fog is mainly
coastal and epiphytic lichens are found on most of the shrubs. This lack of
regular rainfall reduces annual plants and grasses to a low measure, and the seed-
and grass-eating mammalian fauna is therefore sparse.

The moisture-laden air is gradually dissipated inland with the rise in eleva-
tion of the terrain until the backbone of the peninsula is reached, which so far
as is now known is the range limit of the species P. fallax. Along the greater
part of the length of the peninsula of Baja California, eastward from the moun-
tainous backbone the terrain falls rapidly to the shores of the Gulf of California
and offers no habitat suitable to P. fallax.

In the spring of 1949 the writer visited the Gulf coast at latitude 29° 30' N.
At that time the only rain for over six years had fallen in the previous two
months. Dried and bleached shrubs were everywhere, and the few cattle owned
by a local ranchero had starved to death. It is difficult to express in words the
sharp changes that take place in such short distances in this arid land. Yet the
profound influence upon both plant and mammalian life is quite apparent to
the naturalist.

The types of the three named races were all taken on the perimeter of the
known range of the species.

Perognathus fallax 417

Perognathus fallax fallax Merriam
San Diego Short-eared Pocket Mouse

Perognathus fallax Merriam, N. Am. Fauna. 1: 19, 20. October 25, 1889.

Type. — Adult male; from Reeche Canyon, 3 miles southeast of Colton,

San Bernardino County, California; collected by Frank Stephens, April 21,

15889
1887; no. — — — -, collection of the U.S. National Museum.
22684'

Range. — Southwestern California and northwestern Baja California, from
the vicinity of Riverside and San Bernardino, almost exclusively on the Pacific
drainage to Ensenada, Baja California, inland to San Matias Pass, where examples
show characters strongly influenced by the desert regions, thence north along
the lower slopes of the Sierra Juarez to Jacumba, just north of the International
Boundary. The population of this latter locality is not typical and, oddly enough,
lives within 3 airline miles of the type locality of P. f. pallidum, however in a
very different plant association and at an elevation 1200 feet higher.

Remarks. — It is unfortunate that the type of the nominate race should
have been selected from a population living in the semi-arid San Bernardino
Valley. A more western coastal locality would have offered a far better example
of this race.

Specimens Examined. — California: San Bernardino County — Lytle
Creek, 1; San Bernardino, 1; Herron's Ranch, Reeche Canyon, 4 (topotypes).
Riverside County — Riverside, 1; Box Spring, 1; Perris, 5; Winchester, 1; San
Jacinto Lake (now drained, specimen collected by A. W. Anthony, June 28,
1895), 1; Aguanga, 2. San Diego County — Ballena, 1; Dulzura, 12; Dehesa,
2; Lake Hodges, 8; El Monte, 3; Mission Gorge, 8; Vista, 1; 2 mi. s. of Del
Mar, 6; 3 mi. s. of Del Mar, 3; 2 mi. se. of Bonita, 5; Ocean Beach, 1; San
Diego, 3; mouth of Tia Juana River, 2. Baja California: El Valle de las
Palmas, 3; n. side of Descanso Bay, 1; Ensenada, 1; 6 mi. e. of Ensenada, 1;
Boundary s. of Jacumba, California, 8; Sangre de Cristo, 19; El Valle de la
Trinidad (not typical), 18; summit of San Matias Pass (not typical), 13.
Total 143.

Perognathus fallax pallidus Meatus
Pallid Short-eared Pocket Mouse

Perognathus fallax pallidus Mearhs, Proc. Biol. Soc. Wash. 14: 135, 136.
August 9, 1901.

Type. — Adult female; from Mountain Springs, about 2500 feet elevation,
east slope of the Coast Range near the Mexican Boundary, Imperial County
(formerly San Diego County), California; collected by Dr. Edgar A. Mearns,
May 16, 1894; no. 61007, collection of the U.S. National Museum.

Range. — Southern margin of the Mohave Desert along the north slopes
of the San Bernardino Mountains and western rim of the Colorado Desert
south to the Mexican Boundary.

418 San Diego Society of Natural History

Remarks. — Perognathus fallax pallidus may be expected southward a
short distance below the Boundary; however, extensive collecting produced no
specimens at two very likely places on the eastern desert slopes of the Sierra
Juarez: GaskelPs Tanks, about 25 miles, and Las Palmitas, about 60 miles,
from the type locality.

The apparent end of the range of P. f. pallidus has given cause to investi-
gate the ranges of other vertebrates living near this area. Some interesting
problems both ornithological and mammalogical were found, though with this
casual study, the solution of range limitations was not determined. Much work
remains to be accomplished in this region before definite statements can be made.

Specimens Examined. — California: Riverside County — Cabezon, 1.
San Diego County — San Felipe Narrows, 2; San Felipe Valley, 3; San
Felipe Canyon, 2; Mason Valley ( = La Puerta Valley) , 17; Granite Mountain,
2. Imperial County — Mountain Springs, 26 (type locality). Total 53.

Perognathus fallax majusculus subsp. nov.
San Quintin Short-eared Pocket Mouse

Type. — Adult male; from San Quintin, Baja California, Mexico; collected
by Laurence M. Huey, June 29, 1947; no. 15952, collection of the San Diego
Society of Natural History.

Characters. — This race is darker in dorsal coloration than any other
member of the species, outstandingly so when a series of specimens is viewed.
It has a very robust body with a relatively shorter tail that is brightly bicolored
and well tufted. An unusually large number of spine-like white bristle hairs are
present over the posterior part of the body when in fresh pelage.

The chief differences of this race, however, are cranial. The skull of
P. f. majusculus as compared with skulls of the other races is more heavily boned,
with larger more inflated bullae, and is more arched dorsally when viewed in
profile. There are also minor differences in dentition.

Compared with specimens from the more southern interior localities, P. f.
majusculus is darker in dorsal pelage color and larger in body size. It differs
in the same respects from the southern coastal race, P. f. inopinus, which is
the smallest race of the species.

Measurements of Type. — Total length, 191; tail, 105; hind foot, 24;
ear, 6. Skull: greatest length, 26.7; mastoid breadth, 14.7; interorbital con-
striction, 6.9; nasals, 9.8; tooth row, 3.6.

Range. — San Quintin district, from south of Santo Tomas to El Rosario.
Most abundant on the coastal mesas up to 1200 feet altitude.

Specimens Examined.— Baja California: 3 mi. s. of San Telmo, 4
Las Cabras, 7; Santo Domingo (Iat. 30° 47' N), 17; 1 mi. s. of San Ramon, 6
s. of San Ramon, 2; north end of San Quintin, 3; 10 mi. se. of San Quintin, 14
Santa Maria, near San Quintin, 1; 5 mi. e. of San Quintin, 2; 1 mi. e. of El
Rosario, 2; 4 mi. e. of EI Rosario, 1; 10 mi. e. of El Rosario, 2; mouth of Canyon
San Juan de Dios, 3; Aguaita, 4 (not typical). Total 93.

Perognathus fallax 419

Perognathus fallax inopinus Nelson and Goldman
Turtle Bay Short-eared Pocket Mouse

Perognathus fallax inopinus Nelson and Goldman, Proc. Biol. Soc. Wash. 42:

110. March 25, 1929.

Type. — Adult male; from Turtle Bay (also known as San Bartolome Bay) ,

Baja California Sur, Mexico; collected by Alfred W. Anthony, August 1, 1896;

no. 81059, collection of the U.S. National Museum (Biological Survey Coll.).

Range. — Along the immediate coastal shelf from Turtle Bay (27° 41' N)
to Santa Catarina Landing (29° 50' N) .

Remarks. — The southern race, Perognathus fallax inopinus, differs con-
siderably from its northern relatives both in lighter coloration of pelage and in
smaller body size. Without question these characters are the results of environ-
mental conditions under which P. f. inopinus lives.

Specimens Examined. — Baja California: Santa Rosalia Bay (28° 40'
N) , 4; Santa Catarina Landing, 2. Total 6.

Perognathus fallax xerotrophicus subsp. nov.
Arid Plains Short-eared Pocket Mouse

Type. — Adult male; from 2 mi. northwest of Chapala, Baja California,
Mexico; collected by Laurence M. Huey, October 15, 1930; no. 8310, collection
of the San Diego Society of Natural History.

Characters. — Dorsally this race, when viewed in series, is slightly paler
than P. f. ma jus cuius but is decidedly darker than the coastal form, P. f. inopinus,
and than P. f. pallidus, the race that inhabits the California desert.

In body size, P. f. xerotrophicus is similar to the nominate race, P. f. fallax,
and to the desert form, P. f. pallidus; it is smaller than P. f. jnajusculus and
larger than P. f. inopinus. It differs from them all by having a longer tail.

Several well-marked cranial characters differentiate P. f. xerotrophicus from
its nearest comparatives. From P. f. majusculus, it differs in having a lighter-
boned skull, less inflated bullae, and a smaller braincase, with a flatter less
curved profile and more slender rostrum. Compared with P. f. inopinus, the
skull is heavier boned and larger in size.

Measurements of Type. — Total length, 210; tail, 127; hind foot, 24;
ear, 6. Skull: greatest length, 26.8; mastoid breadth, 14.0; interorbital con-
striction, 6.2; nasals, 10.4; toothrow, 3.7.

Range. — Found in an irregular pattern all inland, from San Fernando
Mission south to Punta Prieta and San Andreas.

Specimens Examined. — Baja California: San Fernando Mission, 26;
5 mi. se. of San Fernando, 1; San Agustin, 21; Rancho Ramona, 7 mi. n. of
Santa Catarina, 15; Onyx, 4; 3 mi. s. of El Marmol, 2; 13 mi. nw. of Chapala,
4; 2 mi. nw. of Chapala (type locality), 5; 25 mi. n. of Punta Prieta, 1; San
Andreas (not typical), 9. Total 88.

«"»••: • . Ll'UI

JAN -1196?

TRANSACTIONS 1^ Jg»

OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY
Volume XII, No. 26, pp. 421-440, figs. 1-9

INHERENT AND APPLIED CAMOUFLAGE

IN THE

SUBFAMILY GEOMETRINAE (LEPIDOPTERA),

INCLUDING THREE NEW LIFE HISTORY STUDIES

BY
John Adams Comstock

San Diego Society of Natural History

SAN DIEGO, CALIFORNIA

Printed for the Society

December 1, 1960

&

■-- i

» ► I . w

•

-•v F*

Fig. 1. Mature larva of Nemoria delicataria Dyar on Eriogonum fasciculatum,
approximately X 5.

INHERENT AND APPLIED CAMOUFLAGE

IN THE

SUBFAMILY GEOMETRINAE (LEPIDOPTERA),
INCLUDING THREE NEW LIFE HISTORY STUDIES

BY

John Adams Comstock

Illustrations by the Author

Contents

Additional Notes on the Life History of Nemoria delicataria Dyar 424

The Life History of Dicborda illustraria Hulst 426

The Early Stages of Merochlora fasiolarta Guenee 431

Host Plants of Geometrinae 435

Bibliography 437

Entomologists have long been fascinated by the remarkable adaptations of
certain insects in protective coloration and form. Among the North American
moths of the subfamily Geometrinae (Hemitheinae), this type of protection is
developed to an unusual degree. The mature insects of this group are pre-
dominantly green of various shades, and their wings are more or less leaflike
so that they blend with their verdant habitat. In addition, the larvae usually re-
semble, in form and color, the particular portion of the specific plant on which
they feed. This frequently is true for nearly the entire cycle of larval growth,
during which they molt several times and not infrequently improve their camou-
flage with each change of skin. Furthermore, the posture of the larva enhances
its imitative protection. In many species the larva attaches to a branch in such
a stance as to simulate exactly a small twig. Some larvae, when in motion, have
a nervous twitching of the body and a jerky progressive gait, which resembles
the quivering motion of small elements of the plant under the influence of a
gentle breeze.

Each species has its individual inherent mechanism of survival. There is a
small part of the group, however, in which the individual seems to exercise a
choice in its camouflage. The larva bites off small fragments of its food plant
and attaches them to short spines on its back with silk as the cementing substance.
Thus it is at all times completely covered with parts of the plant on which it
feeds. Each time the larva molts, this covering has to be shed and immediately
replaced with new fragments. When the larva finally spins a cocoon, it transfers
the camouflage to the outside of the covering layer of silk.

I have referred to this type of activity as "applied camouflage", but it must
be understood that no reasoning faculty is involved, even though it simulates a
rational cause-and-effect reaction. How this habit originated is difficult to ex-
plain, but it persists because of its highly protective survival value.

Many authors have published information on the life histories of members
of this subfamily, although there are still many concerning which nothing is

Uiiiiki

JAN-1

HAiUi
IfNIVERS

424

San Diego Society of Natural History

known. Three new life histories are recorded here. In addition, I include a list
of the known host plants and a bibliography of the published papers covering
nineteen species.

Additional Notes on the Life History
of Nemoria delicataria Dyar

Since the publication in 1945 of my paper on certain portions of the life
history of Nemoria delicataria, additional material has come to hand allowing
amplification of this record. A mature larva was collected on Eriogonum fascicu-
latum var. foliolosum (Nuttall) Stokes, in Del Mar, California, late in August
1958. Notes and drawings were made from this and from the resultant pupa.
On August 14, 1959, while I was collecting at black light, a gravid female came
in, and obligingly laid a quantity of eggs, which made possible the supplemental
information and illustration included here. The eggs were laid on August 15, and
hatched on August 21, 1959. The young larvae were immediately placed on
tender leaves and also flowers of Toyon (Photinia arbutifolia Lindley), which
they avidly accepted.

Fig. 2. Egg of Nemoria delicataria Dyar, in two views, approximately X 60.

Egg (fig. 2). — The egg is a flat oval, 0.75 mm. long, 0.55 mm. wide,
and 0.33 mm. high. When first laid, it is a deep pinkish orange, with a narrow
white line circling the edge of the side wall where it joins the top, and with a
similar circlet around the base. The surface appears granular but on higher
magnification is seen to be covered with a fine grill or network surrounding
hexagonal cells. This network is more clearly defined on the superior surface
of the egg than it is on the side walls.

Comstock — Camouflage in the Geometrinae 425

Larva, first instar. — The length is 1.5 mm. and the head width approxi-
mately 0.25 mm. The head is bright orange and the ocelli black. The body
ground color is bright yellow. A very narrow longitudinal middorsal discon-
tinuous black stripe is present, on each side of which is a wide yellow band.
Lateral thereto is a series of dark spots, one to a segment, on each side. Latero-
inferior thereto is a broad yellow area. Another series of large raised black spots
runs stigmatally. There are numerous small warty tubercles ranged along the
body, a pair of which in the cervical area and another pair in the caudal area
are larger than the others. A large pyramidal process occurs middorsally on
the cauda.

The young larva moves in a series of nervous jerks.

From this point on, no separate record was kept of the larval instars, until
the mature stage, but it was noticeable that most or all of the characters of
maturity were present.

Another noteworthy fart is that the larvae raised on Toyon were uniformly
much darker and more heavily spotted with brown than was the single example
reared on Eriogonum. The latter, having much more pink and yellow was better
adapted to matching the colors of the stems and flowers of its food plant.
Conversely, the darker mottled brown form reared on Toyon was well camou-
flaged when resting on the twigs or stems of its host plant. Since the dark
Toyon-feeding mature larva was described in my earlier (1945) paper I will
include here the description of the pinker color phase.

Mature Larva (fig. 1). — The length is 18 mm. and the head width
approximately 1.4 mm. The head is mottled and speckled with various shades of
brown. The body color is cream-yellow, which is heavily overlaid with spots and
blotches of pinkish brown. Middorsally there is a longitudinal line of broken
brown dashes. The body surface has a frosted appearance, owing to a covering
of very minute raised granules. In the middorsal area each typical segment bears
a single spur, bifurcated at the tip, at the juncture of the forward third of the
segment. Caudal to this on each segment there is a pair of short spurs. Dorso-
laterally, from the 4th to the 7th segments, a prominent spur projects diagonally
forward and laterally, one on the side of each segment. Each is heavily shaded
with pinkish brown. The caudal segment has a pair of long spurs which incline
caudally. Numerous small nodules occur on various portions of the body.

Pupa. — The illustration of the pupa in my paper of 1945 was a photo-
graph in which some of the structural details were blurred. The description also
lacked certain important details. I have therefore prepared a drawing showing
the pupa in three aspects (fig. 3) and add the following description:

The length is 11.5 mm. and the greatest width through the area of the
patagia, 3 mm. The pupa is fusiform, the cephalic end well rounded and the
cauda tapering to a small rounded knob, which is topped by a cremasteric
rosette of short recurved red-brown hooklets. The ground color is yellow-tan,
but this is heavily obscured by a speckled mottling of red-brown. The maxillae
terminate about 0.5 mm. beyond the wing margins, and the antennae reach
0.2 mm. beyond that. The eyes are not prominent. The wings are streaked with

426

San Diego Society of Natural History

light lines on the nervules, interspersed with minute blackish-brown raised dots.
The entire surface of the pupa is finely granular.

An imago emerged October 2, 1958, from a pupa that formed September
8, 1958.

n t

Fig. 3. Pupa of Nemoria delicataria Dyar, in three views, approximately X 6.

The Life History of Dichorda illustraria Hulst

A gravid female of this beautiful green geometrid moth came to black light
on the evening of August 15, 1959, in Del Mar, California. On the following
day numerous eggs were deposited in the rearing jar. These hatched on August
21, 1959, and were immediately transferred to succulent leaves of the known
food plant, Rhus laurina Nuttall.

Egg (fig. 4) . — The egg is a flattened oval 0.8 mm. long, 0.6 mm. wide,
0.2 mm. thick at the more acutely rounded end, and 0.3 mm. thick at the op-
posite end. The color is a light salmon, except for a narrow white stripe en-
circling the upper margin and a similar stripe around the basal margin. These
white marginal stripes are finely studded, but this bead-like arrangement is
discernible only with higher magnification. The side walls are slightly convex
and are granular in texture. The top surface is faintly pitted.

Larva, first instar. — The length when extended is 1.75 mm. and the
head width approximately 0.5 mm. The head is orange-yellow. There are two
pairs of horny protrusions over the crown, and a few short colorless setae are

Comstock — Camouflage in the Geometrinae

427

Fig. 4. Egg of Dichorda lllustraria Hulst, in two views, approximately X 60.

present on the face. The body is predominantly mottled brownish black. A
series of yellow papillae runs longitudinally along the dorso-lateral margin.
These are placed on raised yellow bases. The legs are black, except for the hyaline
terminal segments. The single pair of prolegs is black. The anal prolegs have a
line of raised yellow warty processes on their lateral surfaces. The venter is black.

On September 3, 1959, larvae approximately 5 mm. long, probably in their
second instar, were noted as follows : The head width is approximately 0.75 mm.
The color is mottled dark brown, tending to blackish brown on the front. The
texture is granular. The ocelli and all other parts of the head are nearly uni-
colorous and are difficult to distinguish. The head is narrower than the first
cervical segment. A pair of pointed papillae rising from the first segment extend
forward over the crown of the head. The first three body segments expand pro-
gressively dorso-laterally. From the 4th to the 8th segments this lateral expansion
becomes a series of quadrate flanges suggesting gables, extending outwardly over
the stigmatal area. The 9th to 11th segments lack these "gables". The 11th
segment bears a pair of processes somewhat similar to those arching over the
crown of the head. The body ground color is a soft tan, shaded with areas of
brown and mauve. There is a very narrow middorsal black stripe. The ventral
surface is mottled dark brown. The legs are dark brown and the prolegs a
lighter brown, tinged with pink.

The larva rests in an arched posture. When disturbed, it starts a quivering
motion.

On September 10, 1959, the larvae were approximately 12 mm. in length
when fully extended. This probably represented their third instar. The head

428 San Diego Society of Natural History

width is 1 mm. The color of the face is red-brown. The horns arching over the
crown from the first segment are much reduced in size. The body color and
shape are much as in the previous instar. The most noticeable change is in the
darkening of all the edges and ventral surfaces of all the flanges or "gables".
The two horns on the 11th segment are relatively longer, stouter, and more
blunted at the tips.

From this point on there is little change in the larva. The principal difference
is in the progressive shortening of the "horns" on the first segment and a progres-
sive darkening of certain segments and areas. The three cervical segments
particularly take on a dark mottling that is in contrast to the remaining seg-
ments. A drawing was made of the larva in penultimate instar which is shown
in fig. 5.

Finally, on September 21, 1959, the following notes were made of the
mature larva. The length, extended, is 21 mm. and the head width, 1.6 mm. The
head is unchanged, so far as color and general appearance are concerned. The
horns on the first cervical segment are reduced to round knobs at the anterior
corners of the segment. The anterior five segments are slightly more darkened
and mottled than previously. The middorsal black stripe is more prominent and
clearly defined. On the ventral surface, the midventral longitudinal band is wide,
and complete from the 1st to the 9th segments. It is dark brown and is margined
by a narrow creamy brown line. The dark brown mottling on the first five seg-
ments of the ventral surface is in strong contrast to the remaining segments. The
crochets on the pair of prolegs are separated into two bundles on each foot, the
anterior group smaller than the posterior group. The crochets are brown on a
light brown base.

The larva began spinning a fragile cocoon on September 21, 1959, and two
days later formed a chrysalis. Pupation occurs on a small branch or twig of the
food plant. Freshly cut pieces of leaves are used to cover the outside of the
pupal chamber. The internal network is fragile and irregular, and the pupa is
clearly discernible within it.

Pupa (fig. 6) . — The length is 13 mm. and the greatest width across the
shoulders, 4 mm. The pupa is subfusiform, the cephalic end cupped inwardly as
a shallow V. The entire surface is rough and heavily streaked and spotted, mak-
ing it difficult to distinguish the segmental junctures. The ground color is light
brown or tan, but this is so heavily obscured with black and dark brown spots
as to appear nearly black, except for portions of the abdominal surface. The
antennae extend to the wing margins, as do also the maxillae. The metathoracic
leg extends out under the tip of the maxilla for a distance of about two-thirds of
a segment. The eyes are relatively small and flat and are discernible only on
lateral view. The wing cases are darker than the thorax and abdomen, and the
nervules are raised and topped by lines of black nodules. There is a middorsal
longitudinal dark line on the abdomen, discontinuous at each segmental juncture,
and expanding at the caudal margin of each segment. In the center of this dark
line there is on each segment a white "tear-drop". Lateral to this middorsal dark
line is a longitudinal series of minute black papillae, paired on each segment and

Comstock — Camouflage in the Geometrinae

429

X

X

O

CL,

a.
n

• 5

-e

on

c
o

u

E

3
C

CL

O

>

430

San Diego Society of Natural History

W^c^ fk

'-»• > •

c

•Jfc-

I

m

5,

' i?

Fig. 6. Pupa of Dichorda illustraria Hulst, in three views, approximately X 8.

each one topped by a microscopic seta. The spiracles are raised and relatively
prominent. The cremaster ends in a small subquadrate element, the rounded end
of which is topped by eight minute recurved hooklets.

The first imago emerged October 11, 1959. Preserved examples of the larva
and pupa will be deposited in the Research Department of Yale University.

Comstock — Camouflage in the Geometrinae 431

The Early Stages of Merochlora fasiolaria Guenee

Eggs of this species were obtained from a gravid female taken at black
light in Del Mar, California, August 14, 1959. They were laid on August 15,
and hatched August 25, 1959. The newly emerged larvae were immediately
placed on Artemisia californica Lessing, their recorded food plant.

Fig. 7. Egg of Merochlora fasiolaria Guenee, in two views, approximately X 60.

Egg (fig. 7) . — The egg is a flattened oval, with the sides convex and the
top slightly recessed. It is 0.75 mm. long, 0.50 mm. wide, and 0.22 mm. thick.
The color is a delicate yellow-green. The white flange circling around the junc-
tion of the top and side is well defined, as is also the equivalent white circlet
around the base. The surface of the egg is granular, with apparently no grill-
work or reticulation such as occurs in the eggs of closely related species.

Larva, first instar. — The length is 1.8 mm. and the head width ap-
proximately 0.25 mm. The head is deep yellow and is crossed by two cream-
colored bars, each placed centrally and running vertically across the cheek. These
bars continue as stripes onto the. cervical segments. There are several minute
black dots scattered over the head. The body ground color is yellow but is
heavily obscured by a brownish mottling on the 4th to 9th segments. There are
two longitudinal rows of spiculiferous processes or nodules, each row placed
dorso-laterally. Each nodule has somewhat the appearance of a minute compact
bush, and this effect is heightened by the larval habit of covering the top of
each nodule with granules scraped from the surface of stems of the food plant.
The legs and prolegs are yellow.

The larva is sluggish in habit; when moving it advances with a quivering,
jerky, and uncertain gait.

432 San Diego Society of Natural History

There was great disparity in the rate of growth of the larvae. On October
9, 1959, only eight examples had survived. Six of these were apparently in their
second instar, and two were nearly mature.

Second instar. — The length is 3 mm. and the head width approximately
0.35 mm. The ground color of the face is light brown, on which the two vertical
yellow bands previously noted are present. The body color and form are much
as in the prior instar, but there is more tendency of the larva to cover the pro-
truding dorso-lateral knobs with plant fragments. A narrow middorsal longi-
tudinal brown stripe is evident, running the entire length of the body, and a
wider stigmatal or substigmatal band is present. The legs are brown and the
prolegs mottled yellow-brown. The venter is yellow.

On October 12, 1959, one of the smaller larvae, apparently in its third
instar or possibly its fourth, was recorded as follows: The length is approxi-
mately 8 mm. and the head width 1 mm. The face is mottled brown and tan.
The first cervical segment is characterized by a transverse row of yellow nodules.
The body is considerably obscured by pieces of stems and parts of blossoms
which the larva has attached to the series of nodules along its back. These
nodules have expanded into triangulate processes with spiculate tops. The mid-
dorsal brown stripe is more prominent and is edged laterally by yellow lines.
Spiracularly, the brownish-yellow band is bordered above by a narrow white
stripe and below by white dashes running diagonally on each segment. Inferior
thereto is a narrow longitudinal dark brown band which borders on the yellow
venter. In the center of the wide yellow ventral band is a narrow brown stripe.
The legs and prolegs are mottled brown. The entire surface of the larva is
covered with minute round white dots, which give it a frosted appearance.

The larva rests in an arched posture, with the head curled under the abdo-
men. When it moves, which it does infrequently, it still exhibits the quivering
nervous habit recorded for prior instars.

The self -acquired camouflage, together with the mimetic form and colora-
tion of the larva, makes it almost impossible to see specimens at rest on the food
plant. It is suspected that the high rate of mortality was due to the accidental
discarding of the nearly invisible larvae on dried plants when the food plant
was changed.

On October 16, 1959, the two largest individuals were apparently mature.
An illustration was made of the largest (fig. 8) and the following notes were
recorded: The length of the larva is 16 mm. and the head width, 1.25 mm. The
head is speckled with dark brown and tan dots, the dark elements forming a
wide band running vertically down the center of the face, bordered laterally with
a tan band, and this giving place to a dark patch that covers the side of the
head. On the body, the middorsal brown band is less conspicuous than it was
in the prior instar, and the yellow band bordering it is wider. The second and
third segments are progressively expanded laterally as flattened lobes. These are
in line with the large triangular plates that occur laterally on the succeeding
segments, but these plates extend upward and outward, and each is topped by
a spur. On each of these spurs the larva attaches one or more fragments of the

Comstock — Camouflage in the Geometrinae

433

,<"\

£Off

^

2- h &-A .

Fig. 8. Mature larva of Merochlora fasiolaria Guenee on Artemisia calif ornica,
approximately X 7y2-

434

San Diego Society of Natural History

V,

< !

it

•>

Fig. 9. Pupa of Merochlora fasiolaria Guenee, in three views, approximately X 8.

food plant which it has bitten off. The triangular plates are margined with light
yellow bands which are bordered by a fine brown stripe. Inferior to this is a
triangulate area of speckled tan, which is crossed by a white diagonal bar that
is pointed at one end. There are five of these triangular plates on each side and,
in addition, a single middorsal one immediately superior to the suranal triangle.

The larva apparently shows some instinctive ability of selection in choosing
the elements with which it camouflages itself. If it places a bit of blossom on
one spur, it usually selects the same type of blossom for the opposing spur of
the same segment. The caudal and thoracic spurs usually seem to carry blossom
fragments, and those between are adorned predominantly with cut ends of
elongate leaves. In other particulars the larva of this instar resembles the prior
phase.

On October 22, 1959, the largest larva began spinning a loosely woven
cocoon on a stem of its food plant. A part of this procedure was to remove its
camouflage and attach it to the outer surface of its silk covering. Thus it at-
tained protection for the pupa equal to that maintained in its prior changes.

The second larva began spinning on October 26. At this time the first
pupa was removed from its cocoon, illustrated, and recorded.

Pupa (fig. 9) . — The length is 8 mm. and the greatest width through the
center, 2.5 mm. The pupa is subfusiform, the cephalic end unevenly rounded.
The maxillae extend beyond the wing cases, but a portion is obscured by over-
lapping of the prothoracic legs. The antennae also extend beyond the wing

Comstock — Camouflage in the Geometrinae 435

margins but not so far as do the maxillae. The cremaster is topped by four spurs,
with recurved tips that curl laterally. There are also two pairs of shorter hooklets
anterior to the caudal group, the tips of which curl medially. On the ventral sur-
face of the cauda there is a dark triangular plate shaped like an arrow point.
This bears a few short straight spicules. The spiracles are dark brown, and
slightly protruding. The ground color of the thorax, head, and wings is brownish
tan, with numerous clusters and lines of dark brown spots. The nervules of the
wing cases are accented by these spots, and the central shafts of the antennae are
heavily covered and cross-hatched with them. On the dorsal surface there is a
middorsal longitudinal line of the same character of spots. The ground color of
the abdominal segments is a light tan, slightly tinged with pink.

Host Plants of Geometrinae

McDonnough in 1938 listed 17 genera, including 80 species, for the
Geometrinae. Ten new species have been described since (9 recorded by the late
John L. Sperry), bringing the total to 90 species, besides several subspecies. Ap-
parently there is no published information on the early stages of 65 of these
species. For the remaining 25, a few have been adequately recorded, but others
require additional study and illustration. There follows a list of the species for
which the host plants are known.

Cheteoscelis bistriaria Packard
Solidago sp.

Chlorissa pistaciaria Guenee
Quercus sp.

Chlorissa subcroceata Walker
Quercus sp.

Chlorochlamys chloroleucaria Guenee

Buds and flowers of Achillea millefolium Linnaeus, Apocynum androsaemi-
jolium Linnaeus, Aster sp., Eriogonum jascicidatum Bentham, Eupatorium per-
joliatum Linnaeus, Helenium autumnale Linnaeus, Helianthus sp., Leucanthe-
mum sp., Rubus sp., Rudbeckia hirta Linnaeus, and 'Z.innia sp. (cultivated).

Chlorochlamys zelleraria Packard

Buds and flowers of Chrysanthemum sp.; Eriogonum fasciculatum Bentham.

DlCHORDA ILLUSTRARIA Hulst
Rhus laurina Nuttall.

Dichorda iridaria Guenee
Rhus sp.

Eueana niveociliaria Herrick-Schaffer
Condalia sp.

436 San Diego Society of Natural History

Merochlora faseolaria Guenee
Artemisia californica Lessing.

Mesothea viridipennata Hulst

Populus sp.; Primus sp.; Ribes sp.

Nemoria bistriaria Hubner
Quercus sp.

Nemoria brunnearia Packard

Juglans nigra Linnaeus; Prunus sp.

Nemoria darwiniata Dyar
Salix sp.

Nemoria delicataria Dyar

Eriogonum fasciadatum Bentham; Photinia arbutijolia Lindley.

Nemoria mimosaria Guenee

Abies sp.; Quercus sp.; Rubus idaeus v. aculeatissimus Meyrick; Tsuga sp.

Nemoria pulcherrima Barnes & McDunnough
Quercus sp.

Nemoria punctularia Barnes & McDunnough
Ceanothus spinosus Nuttall; Quercus sp.

Nemoria rubrifrontaria Packard

Ceanothus sp.; Comptonia sp., probably Comptonia peregrina var. aspleni-
folius (Linnaeus) Fernald; Eupatorium sp.

Racheospila herbaria v. hulstiana Dyar*
Flower heads of Lantana camara Linnaeus.

Racheospila rubrolinearia Packard*
Myrica sp.; Quercus sp.

Racheospila rubromarginaria Packard*
Quercus sp.

Synchlora aerata Fabricius

Ageratum sp.; Aster sp.; Rubus sp.

Synchlora denticularia Walker

Rudbeckia sp.; flowers of Solidago sp. and Verbena sp.; Verbesina sp.

Synchlora liquoraria Guenee

Artemisia californica Lessing; buds and flowers of Chilopsis linearis De-

Candolle; Coreopsis sp.; Eriogonum fasciadatum Bentham.

Synchlora rubrifrontaria Packard
Eupatorium sp.

* William T. M. Forbes (1948) placed these three species of Racheospila in the genus
Nemoria.

Comstock — Camouflage in the Geometrinae 437

Bibliography

The references to life history studies in the literature evidence the need of
much additional research in this important group of insects. Very few of the
published records of metamorphoses are complete, and there is a regrettable
scarcity of good illustrations. The following bibliography includes only those
references that carry information on life histories, food plants, or parasites.

Anonymous

1890 Some of the bred parasitic Hymenoptera in the national collection.
Insect Life 2: 348-353. [Food plants and parasite of Synchlora rubri-
jrontaria, Chlorochlamys zelleraria, and Synchlora aerata (as Aplodes
rubtvora) , p. 352.]

Bird, Ralph D.

1927 Notes on insects bred from native and cultivated fruit trees and shrubs
of southern Manitoba. Canad. Ent. 59: 124-128. [Food plant of
Nemoria mimosarta,p. 125.]

Comstock, John A.

1945 Notes on the early stages of Nemoria delicataria Dyar. Bull. So. Calif.
Acad. 44: 20, 21, pis. 9, 10. [Larva, pupa, food plant.]

1960 This publication. [Egg, larva, pupa, and food plant of Nemoria
delicataria, Dicborda illiistraria, and Merochlora jaseolaria.~\

Comstock, John A., and Charles M. Dammers

1934 Additional notes on the early stages of California Lepidoptera. Bull.
So. Calif. Acad. 33: 25-34, pis. 3-13. [Larva, pupa, and food plant of
Chlorochlamys chloroleucaria, pp. 29, 30, pi. 8.]

1937 Notes on the early stages of three California moths. Bull. So. Calif.
Acad. 36: 68-78, pis. 29-40. [Larva, pupa, and food plant of Synch-
lora liquoraria, pp. 71-74, pis. 34-35a; larva, pupa, and food plant of
Nemoria pulchemma (as N. naidana) , pp. 74-78, pis. 36-40.]

Comstock, John A., and Christopher Henne

1940 Notes on the early stages of Nemoria pistaciaria Pack. Bull. So. Calif.
Acad. 39: 78-80, pis. 9-11. [Egg, larva, pupa, and food plant of
Nemoria punctularia (as N. pistaciaria) .]

Dethier, Vincent G.

1942 Notes on the life histories of five common Geometridae. Canad. Ent.
74: 225-234, pis. 14-17. [Larva and pupa of Chlorochlamys chloro-
leucaria, pp. 231-233, pis. 15, 17; larva of Nemoria rubrofrontaria, p.
233, pis. 15, 16.]

438 San Diego Society of Natural History

Dyar, Harrison G.

1894 Descriptions of certain geometric! larvae. Ent. News 5: 60-64. [Larva,
pupa, and cocoon of Synchlora dentictdaria (as S. excurvaria) , p. 62.]

1899a Life histories of North American Geometridae. — I. Psyche 8: 310,
311. [Egg, larva, and food plant of Nemona mimosaria.~]

1899b Life histories of North American Geometridae. — II. Psyche 8: 386,
387. [Egg and larva of Chlorissa subcroceata.~\

1900a Life histories of North American Geometridae. — XIII. Psyche 9:
93, 94. [Egg and larva of Synchlora aerata (as S. glaucaria) .]

1900b Life histories of North American Geometridae. — XV. Psyche 9:
118, 119. [Egg, larva, pupa and cocoon of Eueana niveociliaria (as E.
saltusaria) .]

1901a Notes on the winter Lepidoptera of Lake Worth, Florida. Proc. Ent.
Soc. Wash. 4: 446-485. [Brief note on Eueana niveociliaria (as
Racheospila saltusaria), p. 456; larva and food plant of Racheospila
herbaria v. hulstiana, p. 457.}

1901b Life histories of North American Geometridae. — XXVIII. Psyche
9: 287, 288. [Egg, larva, cocoon, and food plant of Mesothea viridi-
pennata.~\

1904 Life histories of North American Geometridae. — LVII. Psyche 11:
121. [Egg, larva, and food plant of Nemoria darwiniata.~\

Forbes, William T. M.

1948 Lepidoptera of New York and neighboring states. Part II. Cornell Univ.
Agr. Exp. Sta. Mem. 274: 1-263, figs. 1-255. [Larva and food plant
of Nemoria mimosaria, p. 113; larva and food plant of N. rubrijrontaria
p. 113; larva and food plant of Racheospila rubroltnearia, p. 113; food
plant of R. rubromarginar'ia, p. 114; larva of R. lixiaria, p. 114; larva
and food plant of Nemoria brunnearia, p. 114; food plant of Dichorda
iridaria, p. 114; larva and food plant of Synchlora aerata, p. 115; larva
and food plant of S. denticularia, p. 115; larva and food plant of
Chlorochlamys chloroleucaria, p. 116; larva and food plant of Chlorissa
pistaaaria, p. 116.]

French, C. H.

1877 Trans. Dep. Agr. Illinois 15: 238. [Larva of Synchlora aerata.']

GOODELL, L. W.

1878 Notes on the early stages of some moths. Canad. Ent. 10: 66, 67.
[Larva of Nemoria brunnearia, p. 66.]

1880 On the early stages of four geometrid moths. Canad. Ent. 12: 235,
236. [Larva, pupa, and food plant of Chlorochlamys chloroleucaria.~]

Comstock — Camouflage in the Geometrinae 439

Hulst, George D.

1879 Notes on Nemoria chloroleucaria, Guen. Bull. Brooklyn Ent. Soc. 2:
78. [Larva and pupa of Chlorochlamys chloroleucaria.]

1888 Larva of Chlorosea bistriaria, Pack. Ent. Am. 3: 193, 194. [Larva,
pupa, and food plant of Cheteoscelis bistriaria.]

Kellicott, D. S.

1885 Eumacaria brunnearia, Packard. Canad. Ent. 17: 32, 33. [Larva, pupa,
and food plant of Nemoria brunnearia.]

McFarland, Noel

1959 Letter to the author. [Food plants of Nemoria punctularia, Synchlora
liquoraria, Dichorda illustraria, and Chlorochlamys z^Ueraria.]

Packard, Alpheus S.

1876 A monograph of the geometrid moths or Phalaenidae of the United
States. Rep. U. S. Geol. Surv. Terr. 10: 1-607, pis. 1-13. [Larva of
Synchlora aerata (as S. rubivoraria) , pp. 382, 383; larva and pupa of
Nemoria rubrifrontaria, p. 387; larva and pupa of N. mimosaria, p.
389 (quotes Walsh) ; larva of Synchlora denticidaria, pi. 13 (after
Abbot's manuscript drawing) .]

1881 Insects injurious to forest and shade trees. U. S. Ent. Comm. Bull. 7:
1-275, figs. 1-100. [Larva of Nemoria mimosaria, p. 49. Also in 1890
edition, p. 189.]

1884 Life-histories of some geometrid moths. Am. Nat. 18: 933-936. [Larva
and pupa of Nemoria mimosaria, pp. 933, 934.]

Riley, C. Valentine

1869 The raspberry geometer, Aplodes rubivora n. sp. First Missouri Rep.
139. [Larva and food plant of Synchlora aerata."]

Saunders, William

1873 Insects injurious to the raspberry. Rep. Ent. Soc. Ontario, 16. [Larva
of Synchlora aerata.]

1883 Insects injurious to fruit. 1-436, figs. 1-440. London and Philadelphia.
[Larva and pupa of Synchlora aerata, p. 316.]

Walsh, Benjamin D.

1864 On certain remarkable or exceptional larvae, Coleopterous, Lepidop-
terous and Dipterous . . . Proc. Bost. Soc. Nat. Hist. 9: 286-318.
[Larva, pupa, and food plant of Nemoria mimosaria (as Hippar-
chiscus venustus) , pp. 300-302.]

|JAN-31S

HARVARD
UNIVERSITY

TRANSACTIONS

OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY
Vol. XII, No. 27, pp. 441-448, figs. 1-4 November 15, 1960

DIFFERENTIATION OF THE SOUTHWESTERN
TORTOISES (GENUS GOPHERUS),
WITH NOTES ON THEIR HABITS

BY

Chapman Grant

Contents

Eggs 442

Growth of the Shell 443

Proportions of Adults 443

Color Perception : 444

Death by Direct Insolation 447

Behavior in Captivity 447

Care in Captivity 448

Literature Cited 448

The true land tortoises of the southern United States and northern Mexico
comprize three species of the genus Gopherus. The gopher tortoise, G. poly-
phemiis, of the southeastern states, is said to range into the Carolinas and west-
ward towards the Mississippi. It digs burrows, lays spherical eggs, is less than half
as high as long, and lacks a lateral flare of the posterior part of the carapace. The
desert tortoise, G. agassizi, occurs in southeastern California, western Arizona,
and the southern tips of Nevada and Utah, and in Sonora, Mexico. Like the
gopher tortoise, it lays spherical eggs, digs burrows, and is less than half as high
as long; but the carapace flares posteriorly, being widest at about the fifth neural
scute. It is apparently confined to regions where the creosote bush (Larrea
divaricata) abounds. Occupying territory between these two species, but not
coming in contact with either unless with G. agassizi in Mexico, is the smaller
Texas tortoise, G. berlandieri, found south and southwest of San Antonio
and into Nuevo Leon and Tamaulipas, Mexico. It is apparently confined to
districts where prickly pear (Opuntia sp.) is found. It lays elongated eggs,
possibly does not burrow, has a posterior flare like that of G. agassizi, is over
half as high as long, and has a proportionately larger head; and the nuchal scute
is reduced or wanting. There is frequently a yellow stripe across the snout behind
the nostrils and a yellow spot behind each orbit. Other distinctions are discussed
herein. Little is known of the very young.

Twenty-one specimens of G. berlandieri were collected April 10 to 15,
1960, near Hebbronville, Jim Hogg County, Texas. Various sizes and both

442 San Diego Society of Natural History

sexes were taken in order to experiment on color perception and to determine
the size and shape of eggs and the proportions and gross anatomy of adults and
thus determine secondary differences between this and the other two species. In
this paper G. berlandieri is compared to G. agassizi.

The temperature at Hebbronville was in the 90's F. by day, and the tortoises
were beginning to emerge from hibernation. It is said in Hebbronville that the
tortoises hibernate by squirming into the soil, leaving the top of the carapace
exposed. A letter from Mr. V. W. Lehman, Wildlife Manager, King Ranch,
July 29, 1960, states, "We know that it utilizes burrows for both shelter and
hibernation, but there are so many coyote, skunk, armadillo, opossum, and badger
dens in the Rio Grande Plain that there is small purpose for the tortoise to make
additional excavations." Mr. Lehman's observations may explain why this
tortoise has not developed the shovel-like forelegs of G. agassizi. I cannot
agree that opossums dig.

It is generally agreed that members of this genus cannot swim. A recently
published statement that G. polyphemus enters the water voluntarily and swims
purposefully is unconfirmed. The other two species seem helpless in water.

In Hebbronville there is a rumor that the tortoise destroys quail eggs, but
Mr. Lehman writes, "I can state with certainty that the tortoise is not a predator
on quail eggs as is often supposed." In the Texas Field Trials for hunting dogs,
the score of the dogs is not reduced for "pointing" tortoises, which possibly
smell like quail. The local name for the tortoise is "sand terrapin".

Eggs

According to the literature, the eggs of G. berlandieri are elongate, but no
measurements or ratios appear. Eggs of G. agassizi and G. polyphemus are
nearly spherical. A captive G. berlandieri laid three eggs June 8, 1960, buried
almost vertically about an inch below the surface and tightly packed. The order
of deposition of these is not known. Another egg was found on the surface
June 19, another July 9, and another July 13. All eggs were hard and all were
nearly circular in cross section except No. 5, which was noticeably flattened,
with transverse diameters of 32 and 34 mm. The egg was squeezed and appeared
to be hard and firm. Yet this egg when remeasured July 15 was found to have
become nearly circular in section. The eggs are white and smooth with a fine
granular surface detectable by scratching with the fingernail. The yolk is visible
and settles when the egg is tilted. These eggs were sterile, indicating that copula-
tion had not occurred prior to hibernation or as late as early April.

Dimensions in mm. Width-length ratio Date laid

1960

Ime

:nsions in mm.

Width-length

ratio

Date

(using greatest

width)

1.

49 x 32.5

.66

June 8,

2.

44 x 31

.70

3.

43 x 31

.72

4.

50 x 34

.68

June 19

5.

40 x 32 x
(40 x 33 on

34
July 15)

.82

July 9

6.

43 x 34

.79

July 13

Grant — Southwestern Tortoises 443

The average axial difference is about 12 mm. The longest egg is about 29
percent as long as the carapace of the female.

Grant (1936) gave the dimensions of three G. agassizi eggs:

Dimensions in mm. Width-length ratio

(using greatest width)

1. 42 x 36.5 .87

2. 42 x 37.5 x 35.7 .89

3. 42 x 36 .85

The average axial difference is about 6 mm. or about half that of G. ber-
landieri, which is about 12 mm.

The maximum clearance between pygal and anals is 19 mm. in the specimen
of G. berlandieri which laid the six eggs. Even without any fleshy parts of the
cloacal region being extended into this constriction during deposition, the space
through which the eggs must pass is 15 mm. less than the diameter of the widest
eggs. Therefore the eggs of this species must be leathery when laid as is the
case in G. agassizi-

Una Nichols, who has kept G. agassizi in captivity in San Diego, Cali-
fornia, for thirty years, stated to me that on six occasions she had seen eggs laid
and had immediately handled them. The eggs when first laid are flattened to
about a third of their final diameter, but they become hard and nearly spherical
in a few minutes.

Miller (1932, p. 190) wrote of the desert tortoise: "The egg which was
dropped into my hand felt quite warm . . . five degrees [C] above the [tem-
perature of] the surrounding air . . . At the moment of deposition the egg is
decidedly moist so that the shell appears quite translucent although perfectly
hard. It is not like some lizard eggs (Gecko), soft at deposition and dependent
upon exposure to the air for hardening."

Growth of the Shell

The smaller the openings between the carapace and plastron, the greater
the protection from predators. As the tortoise grows, these openings must
enlarge to allow for the growth of legs and head, and it might be expected that
they would enlarge proportionately. Actually in the desert tortoise, the head
clearance, or anterior opening, increases more than proportionally to the length
of the carapace. The posterior, or anal clearance, which is used only for excretion,
mating, and egg laying, apparently does not enlarge after maturity, remaining
smaller than the width of the eggs.

Proportions of Adults

The proportions of an adult male of G. berlandieri (carapace 176 mm.
long) and those of an adult male of G. agassizi (carapace 220 mm. long) show
differences that are not so pronounced in the young.

G. berlandieri

G

. agassizi

.26

.23

.17

.16

.65

.68

.28

.38

.72

.64

.82

1.00

.42

.90

444 San Diego Society of Natural History

Head length1 to carapace length2
Head width'5 to carapace length
Head width to head length
Anterior head width1 to head length
Forefoot length to forefoot width'
Hind foot length to hind foot width'5
Foreleg width7 to head length

1 From the end of the fleshy snout to the dissected end of the supra-occipital.

- Between the rear of the pygal and (in G. berlandieri) the most anterior line joining the
ends of the first or second pair of marginals, or (in G. agassizi) the anterior end of the
nuchal.

3 At the widest point, just posterior to the orbits.

4 Just anterior to the orbits. The greater constriction in G. berlandieri gives it a Chelydra-
like appearance.

5 Length, from the base of the center claw to the center of the crease that denotes the end
of the foot. Width, between the outside bases of the outer claws.

6 Length, from the base of the second claw, excluding adjoining spines, to the heel, excluding
spines. Width, between the outer edges of the bases of the outer claws.

7 From edge to edge at the widest point, excluding the spines on the outer edge.

Color Perception

Walls (1942) discussed color discrimination in turtles in general, not
including Gopherus, stating that they are red-green perceptive and, curiously,
also violet perceptive, although violet is at the opposite end of the spectrum.
He also intimated that they may be blue-yellow colorblind. My experiments
with captive Testudo in Puerto Rico and Terrctpene in Indiana and Kansas
showed that there is a marked reaction to any red object.

Dr. Meyer Wiener, of Coronado, California, wrote in a letter of July 17,
1960, "You are perfectly right in assuming that the eyes of turtles are adjusted
to diurnal living. The retina consists almost entirely of cones."

My experiments with G. berlandieri cover the combination of color percep-
tion, olfaction, and taste, which seem to be used in that order. Red appears
to be the most attractive color and a red object is immediately investigated,
smelled, tasted, and eaten if suitable. Blue, yellow, and white objects appear
not to register except as objects and are not sampled unless accompanied by a
pleasing smell. Thus peeled bananas in natural color or dyed blue, yellow, or
green, are eaten. Flowers and leaves of red hibiscus (Hibiscus rosa-sinensis)
are readily eaten, but white hibiscus flowers are ignored unless dyed red, when
they are immediately perceived and eaten. If dyed blue or green they are eaten
only casually with the green leaves. This may indicate that hibiscus flowers
are chosen by color and that the taste is agreeable. I cannot detect any odor
from this flower.

Grant — Southwestern Tortoises

445

Fig. 1. Head and intromittent organ of G. berlandieri (left) and of G. agassizi.

Note the near equality of head size of two mature males of substantially different body
size (carapace of G. berlandieri 176 mm. long; that of G. agassizi 220 mm. long). The
single intromittent organs differ principally in pigmentation.

FlG. 2. Eggs of Gopherus berlandieri, laid in captivity, June and July 1960.

446

San Diego Society of Natural History

Grant — Southwestern Tortoises 447

From experiments with available plants and vegetables, it appears that red
hibiscus flowers are the first choice; next are the green pods of black-eyed peas
or string beans, next the fruit of Opuntia when red-ripe, next the pads of spine-
less Opuntia or the tender pads of spiny Opuntia. Sparingly eaten are red or
pink carina blossoms, purple iris blossoms, and apples. The apples are more
readily eaten if dyed red, but will finally be consumed even after they have dried
to a brown morsel. Apparently the smell of dried apple is more pleasing than
that of fresh apple. Apples must be quartered, for the tortoise cannot bite or
hold a whole one. Slices of fresh onion dyed red are smelled, tasted, and rejected:
apparently the smell is not offensive, but the taste is. Lettuce and green cabbage
are rejected, but red cabbage or cabbage or lettuce dyed red are sparingly eaten.
They struggle to eat the sweet coarsely fibrous covering of the seeds of the queen
palm (Arecastrum romanzoffianum) . Watermelon, beets, red roses, fuchsias,
red plums, red beef, white bread dyed red, cantaloupe unless dyed red, all were
ignored. When the red is eaten off a red-dyed cantaloupe or banana, they become
less attractive. Brushing food that is not particularly liked with vanilla extract
had no effect. Frequently when an animal is not eating normally, it can be
induced to eat its usual foods if these are flavored with vanilla.

Experiments were made in the dark with an ordinary electric bulb. The
tortoises moved about slowly a couple of feet beyond the lighted area and then
returned. With a red bulb the reactions were the same except that white straws
etc., appearing bright red, were closely inspected, probably as potential food.

Death by Direct Insolation

On July 20, 1960, a healthy male of G. berlandieri was exposed to direct
sun in a carton 12 inches deep. It hurried around inside the carton with great
agitation for ten minutes and then stopped with its head in a two-inch strip of
shade. The temperature in the shade adjacent to the carton was 103 degrees F.
Twenty minutes later it was dead and rigid, and it may have died immediately
after coming to a halt. The remaining tortoises, in the shade at 103 degrees F.,
were not affected.

Behavior in Captivity

The Texas tortoise seems more sensitive to the sun than is G. agassizi, as
shown by its endeavor to get into shade. Its activity is limited to short periods
in the morning and afternoon during optimum temperature and light conditions.
Doubtless these periods would be longer in the field if the animal were hungry,
but captives do not eat daily, sometimes skipping two days. It is more sensitive
to darkness than are chickens and immobilizes while there is still light enough
for a person to read a newspaper. The spot or corner selected to spend the night
is usually the same for the various individuals. If moved from the selected spot
after dark, some will make a blundering effort to move, whereas others will
spend the remainder of the night where placed.

Digging seems to be of two kinds. They can squirm down into the soil,
aided by the recurved marginals which act like little plows, lifting the dirt

448 San Diego Society of Natural History

upwards; or they burrow, flicking the dirt out with the forefeet and pushing
it out with the hind feet.

Apparently undesired grass and twigs are eaten because the tortoise is
unable to discard them because of its clumsiness.

While climbing in attempting to escape or find shelter, the Texas tortoise
may frequently fall wrong-side-up. Because of the domed carapace, it is usually
somewhat tilted to one side. The first act is usually to urinate and then lie
quietly for some seconds. Then the snout is pressed against the ground; the
upper foreleg is flailed to overcome inertia so as to rotate the body towards the
snout. The lower foreleg can then be moved next to the snout, and the lower
hind leg is used to pry the body over.

No attempts to mate were seen although the temperature was in the 90's F.
for a week in July and then up to 104 the following week and in the 90's the last
week of July. They were penned outdoors with shelter and water but were not
seen to drink.

Care in Captivity

Many captive desert tortoises have died because of unfavorable conditions.
They will not eat except in broad daylight and at an optimum temperature of
from about 80 to 90 degrees F. Food that can be bitten off of larger pieces is
preferable to chopped food. Lettuce and apples are not a whole food, whereas
green string beans, the pads of Opuntia ficus-indica, and the fruit of any Opuntia,
are whole foods and readily eaten. Some other fruits and vegetables are eaten
if dyed red with an odorless tasteless vegetable dye, which may be applied with
a pastry brush. Captive tortoises must have shade, and they must be supplied
with water unless Opuntia or other succulent food is eaten. They must also be
allowed to burrow or furnished with low shelter.

Literature Cited

Grant, Chapman

1936 The southwestern desert tortoise, Gopherus agassizii. Zoologica
21: 225-229.

Miller, Loye

1932 Notes on the desert tortoise {Testudo agassizii)- Trans. San Diego
Soc. Nat. Hist. 7: 187-206, pis. 10, 11.

Walls, Gordon Lynn

1942 The vertebrate eye and its adaptive radiation. Cranbrook Inst. Sci.
Bull. 19: i-xvi, 1-785, figs. 1-197, pi. 1, frontisp.

LIBRA

AUG 2 2
HARVA

TRANSACTIONS L '

OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY
Volume XII, No. 28, pp. 449-476, figs. 1-3

MARINE MOLLUSKS FROM LOS ANGELES BAY,
GULF OF CALIFORNIA

BY

James H. McLean

Hopkins Marine Station, Pacific Grove, California

SAN DIEGO, CALIFORNIA

Printed for the Society

August 15, 1961

MARINE MOLLUSKS FROM LOS ANGELES BAY,
GULF OF CALIFORNIA

BY

James H. McLean
Contents

Introduction 451

Previous Collecting in the Los Angeles Bay Area 452

Oceanographic Considerations 452

Description of the Collecting Areas 453

Faunal Relationships 455

Systematic Account 457

Acknowledgments 474

Literature Cited — ~ 474

Introduction

Los Angeles Bay, Baja California, is on the west shore of the Gulf of
California at 29 degrees north latitude and thus about 275 miles south of the
California boundary (fig. 1). Although several small collections of mollusks
have been reported from this part of the Gulf, no extensive list is published.
Now two large collections and three small ones make possible a preliminary
check list. Though doubtless incomplete, this list probably includes at least all
the common species. It is offered as a working list for use at the new Vermilion
Sea Field Station, of the San Diego Museum of Natural History, at Los Angeles
Bay. Furthermore, it includes several interesting range extensions.

Mrs. Faye Howard of Pacific Palisades, California, collected mollusks at
Los Angeles Bay on September 9 and 10, 1957, and from February 22 to April
2, 1960. Her collection of 362 species is the main basis for the list. Also, from
June 9 to 12, 1960, I collected 156 species of mollusks. Several people have
contributed identifications: Dr. A. Myra Keen, most of the small pelecypods;
Dr. S. Stillman Berry, Caecidae, chitons; Dr. William K. Emerson, scaphopods;
Dr. Leo G. Hertlein, brachiopods and some gastropods; and Mr. Allyn G.
Smith, additional chitons. A small collection by Miss Josephine Scripps and one
by Mr. Wesley Farmer, both identified by Mr. Emery P. Chace, also are included
in the report, and likewise a small collection by Mr. Robert Martin, identified
by Dr. S. Stillman Berry.

452 San Diego Society of Natural History

Previous Collecting in the Los Angeles Bay Area

Nine records of molluscan collecting in the vicinity of Los Angeles Bay
appear in the literature. (1) The "Albatross" voyage of 1911 reported by
Townsend (1916) collected mollusks at Angel de la Guarda Island. (2)
Oldroyd (1918) described Acanthina angelica and recorded 46 other species of
mollusks from Angel de la Guarda Island. (3) The 1921 expedition of the
California Academy of Sciences to the Gulf of California dredged in Los
Angeles Bay. The type of Haminoea angelensis was taken then. A general ac-
count of this expedition was given by Slevin (1923). (4) The Second Bingham
Oceanographic Expedition to the Gulf of California dredged in Los Angeles
Bay in 1926. Boone (1928) listed 16 species of mollusks found at that time.
(5) Lowe (1935) described two species from Los Angeles Bay, Calliostoma
angelenum and Clavus pembertoni, which he collected there in 1933. (6) The
Hancock Pacific Expeditions of 1936 and 1937 both visited Los Angeles Bay.
Fraser (1943) listed five dredging stations and three shore stations, but the
mollusks have not been reported. (7) Steinbeck and Ricketts visited Los Angeles
Bay on their collecting cruise of 1940. They concentrated on shore collecting of
invertebrates, and mentioned 28 common species of mollusks from Los Angeles
Bay in their Sea of Cortez (1941). (8) Emerson and Puffer (1957) reviewed
collections made on the E. W '. Scnpps cruise of 1940 and reported 26 species
from Angel de la Guarda Island, 12 species dredged in the Ballenas Trench
west of the island, and 66 species from Las Animas Bay, to the southeast of
Los Angeles Bay. (9) The 1957 Puritan-American Museum of Natural History
Expedition to Western Mexico collected at the south and north ends of Angel
de la Guarda Island. A general account of this expedition was given by Emerson
(1958).

Oceanographic Considerations

Los Angeles Bay has an area of about 25 square miles. It is protected at
the entrance by several small islands (fig. 1). Angel de la Guarda Island, the
second largest island in the Gulf, lies east of the Bay, approximately 20 miles off
shore. The Gulf sea-floor outside the Bay slopes gradually to a depth of over
600 fathoms in the Ballenas Trench, then rises abruptly to Angel de la Guarda
Island, which reaches an elevation of 4315 feet (Anderson 1950). The 600-
fathom basin of the Ballenas Trench is isolated from other areas of similar depth
by bottom of less than 200 fathoms.

Unusual hydrographic conditions are found in the Los Angeles Bay area.
Roden and Groves ( 1959, p. 1 1) reported:

One of the outstanding features of the Gulf of California is the large annual
range of sea surface temperatures; at Puerto Penasco in the north the range is about
16°C; at Cabo San Lucas in the south it is about 9°C . . . An interesting feature
is an area of low temperatures around Isla Angel de la Guarda which is found
throughout the year. Along the coast, temperatures are somewhat dependent upon
upwelling; in winter low temperatures are generally found along the east coast and
in summer along the west coast.

McLean — Mollusks of Los Angeles Bay 453

Unfortunately, no year-around temperature data are available, but up-
welling is known to occur in the area of Los Angeles Bay and Angel de la
Guarda Island. During June the water was so cold that I needed a rubber suit
for skin-diving.

The tidal range in the vicinity of Los Angeles Bay is about three meters.
The tides are characterized by diurnal inequality, i.e. successive high tides are
markedly unequal and likewise successive low tides. Tidal currents of six knots
have been reported in the Ballenas Strait (Roden and Groves 1959).

During my stay at Los Angeles Bay in early June, a "red tide" plankton
bloom was present in front of the village but not around the small islands. The
organism was not a dinoflagellate such as causes red tides in southern California
but had the appearance of an unbranched red filament, just visible without
magnification. Dr. E. Yale Dawson, the algal specialist to whom I wrote, sug-
gested that it was "probably the cyanophyte that gave rise to the name of
'Vermilion Sea' for the Gulf of California in the days of Coronado." Where
it occurred in high concentrations, the water was rust-red and slimy to the touch.
This condition had prevailed in the bay during most of May. My attempts to
preserve specimens in alcohol were unsuccessful because the filaments disinte-
grated. Unlike the dinoflagellate red tides, this planktonic bloom did not seem
to have any detrimental effects on fish or invertebrates. The field station will
provide an opportunity to investigate this phenomenon when it occurs again.

Description of the Collecting Areas

The rocky area in front of the field station (fig. 3) was collected in-
tensively. The most abundant of the mollusks found there are listed in table 1.
Inasmuch as wave action and the effect of tidal currents are slight, the substrate
surrounding the rocks is muddy, and the assemblage therefore includes some
forms typical of a restricted bay environment as well as others more typical of
the open coast.

Steinbeck and Ricketts (1941, pp. 220, 221) mentioned some of the in-
vertebrates they collected at this spot and gave additional records in the appendix
to their book. My identifications of invertebrates other than mollusca are based
upon their photographs, records, and descriptions.

At highest tides the stone wall in front of the station is covered by water,
but the rocks of this wall are barren. The lowermost stones of the wall, and the
boulders below, support the large barnacle Tetraclita squamosa stalactifera
(Lamarck) and the smaller barnacle Chthamalus anisopoma Pilsbry, among
which oysters and limpets are found. Three species of mollusks occur in great
numbers under rocks. A large rock I overturned had under it about 60 individuals
of the pulmonale Onchidella binneyi, 30 of the chiton Acanthochitona exquisita,
and 25 of the gastropod Acanthina angelica.

A black holothurian is conspicuous under rocks exposed by the higher of
the low tides. The lowest tides expose a much greater profusion of life. Algae
are noticeable only at the lowest levels, especially a brown alga with fan-shaped
blades, Padina durvillaei Bory, and a green alga, Codium cuneatum Setchell and
Gardner. Most noticeable at the lowest tides is a large red sponge, colonies of

454

San Diego Society of Natural History

under sides of rocks

among barnacles

>> »

under rocks

attached among barnacles

on sand near rocks

on

rocks

Table 1. Mollusks abundant on rocks in front of the field station
Exposed at mid-tide :
Chiton

Acanthochitona exquisita
Gastropods

Acmaea mi tell a fayae

A. turveri

Acanthina angelica

Onchidella binneyi
Pelecypods

Hormomya adamsiana

Ostrea concbaphila

O. palmida
Exposed at both low tides :
Gastropods

Cerithium sculptum

Mitrella ocellata

Acmaea strongiana

Nomaeopelta dalliana

Tegula mariana
Pelecypods

Ar cops is solid a

Cardita affinis

Isognomon chemnitzianus
Exposed only at the lower low tides
Gastropods

Crassispira nympbia

Fusinus cinereus

Morula ferruginosa

Pyrene fuscata

Turbo fluctuosus

Strombina maculosa

Crucibidum spinosum

Cypraea annettae

Diodora alta

D. digue ti

D. inaequalis

Marginella calijornica

Mitra solitaria

Serpulorbis margaritaceus

attached to under-surfaces of rocks

exposed on, or under, rocks

always under rocks

attached to sides of rocks

which reach a foot in diameter. Under rocks will be found a black sea urchin,
Arbacia incisa A. Agassiz, a many-rayed star, Heliaster kubijiiji Xantus, and
numerous brittle stars, sponges, and tunicates. Octopus bimaculatus may be en-
countered among rocks underwater. The depth of the water increases gradually,
but I could not explore the subtidal area because of the red tide.

McLean — Mollusks of Los Angeles Bay 455

Mrs. Howard collected on the east shore of the bay, where she reported
that the intertidal zone is steeper than on the west side and surf and tidal action
are stronger. The molluscan assemblage is typical of a more exposed coast.
Tegula rugosa, an abundant north Gulf gastropod, occurs there in large numbers
though not in the more protected environment of the bay.

Just north of the station and in front of the village, the lowest tides expose
a flat of muddy sand below the beach. This flat increases in width toward the
sandspit to the north. The dominant animals of the flat are the sand dollars
Encope grandis L. Agassiz and Encope californica Verrill. They are found
buried just under the surface of sand or exposed in such numbers that they may
cover entire areas. Two or three commensal crabs, Dissodactylus sp., were usually
found with each sand dollar. Such a high concentration of sand dollars may
discourage burrowing pelecypods. The gastropods Nassarius tiarula and, in
lesser numbers, N. moestus occur on the sand surface by the thousands. The
large gastropod Terebra variegata moves just below the surface of the sand.
On the sandspit, cobbles and small rocks support some rock-living forms.

The small islands are separated from shore by water 10 to 25 fathoms
deep (fig. 1 ) . The intertidal zone on the islands is steep and, except for a few
gravel beaches, is composed entirely of small boulders (fig. 2). At low tide
level, the alga Padina durvillaei Bory grows more luxuriantly than on the rocks
in front of the station. Around the islands, a Sargassum with dense branches
is anchored to the subtidal rocks within several yards of shore, forming the
equivalent of a kelp bed.

Subtidally, the long-spined urchin Centrostephanus coronatus (Verrill)
practically fills the rock crevices, and the diver must avoid its sharp spines. The
coral Pontes californica Verrill (see Squires 1959) forms extensive green en-
crustations and pinnacles on the rocks, and encrusting red coralline algae cover
large areas of rock surfaces. A specimen of the large pecten Lyropecten sub-
nodosus, loosely attached by byssal threads, was encountered. The jingle oyster
Pododesmus pernoides thrives on the upper surfaces of boulders. Under surfaces
of rocks yielded many specimens of the limpet Acmaea semirubida. SCUBA
diving at greater depths should prove rewarding.

Faunal Relationships

This check list records 405 species of mollusks, of which 135 are pelecypods,
248 are gastropods, 3 are scaphopods, and 19 are chitons. Of this number, 16
are doubtfully identified ("cf.") and 34 are identified only by genus. I hope to
describe some of the new species in a later paper.

Twenty- four species collected at Los Angeles Bay appear to be major north-
ward extensions of known range, not to my knowledge having been reported
from the Gulf of California or from north of Mazatlan (range data from Keen
1958). These are: Pseudochama corrugata, P. panamensis, Mactra fonsecana,
Apolymetis cognata, Tagelus bourgeoisae, Arene hindsiana, Balcis gibba, Opalia
diadema, Cyclostremiscus panamensis , Solariorbis elegans, S. narinensis, Teinos-
toma cf. T. herbertianum, Rissoina io, Heliacus mazatlanicus , Serpulorbis mar-

456 San Diego Society of Natural History

garitaceus, Eupleura nitida, Mitrella delicata, Nassarina pammicra, Mitra ery-
tbrogramma, Crassispira atramentosa, Mangelia phyrne, Turricula dolenta, T.
howelli, and Placiphorella blainvillei. The northernmost locality previously
known for each of these is mentioned in the checklist. Four species collected at
Los Angeles Bay appear to be southward extensions of known range, occurring
in the California region but not to my knowledge reported before from the Gulf.
These are: Macoma indentata, Balcis rutila, Leptochiton cancellatus, and Lepi-
dochitona keepiana.

The molluscan fauna is Panamic in its affinities. The northern Gulf, how-
ever, supports some members of the Californian province. The following species
with ranges centered in the Californian province occur at Los Angeles Bay
(range data from Keen 1937) : Aequipecten latiaurata, Transenella tantilla,
Cooperella subdiaphana, Macoma indentata, Cryptomya californica, Corbula
luteola, Tegula ligidata, Balcis rutila, Metaxia diadema, Polinices reclusianus,
Trivia californiana, Cypraeolina pyriformis, Mangelia barbarensis, Melampus
olivaceus, Wdliamia peltoides, Leptochiton cancellatus, and Lepidochitona
keepiana. Some of these species may be isolated in the north end of the Gulf,
while others may eventually be shown to occur along the entire Baja California
peninsula and throughout the Gulf. None are to be expected south of Mazatlan.
Many other species, as for example, Chione calif orniensis, are found in southern
California and the Gulf but range well into the Panamic province (see Hertlein
and Emerson 1956) .

More striking than the presence of these northern elements is the apparent
absence of a considerable number of species common on the eastern shore of the
Gulf. The following larger gastropods, not known at Los Angeles Bay, I have
collected at either Guaymas or Puerto Penasco, or both: Astrea unguis, Tumtella
gonostoma, Cypraea arabicula, Thais biserialis, T. speciosa, T. triangularis,
Purpura patula pansa, Anachis varia, Parametaria dupontii, Pyrene haemostoma,
P. major, Cantharus sanguinolentus, Conus brunneus, C. nux, C. perplexus, C.
princeps, C. purpurascens, and C. virgatus. Many of these species occur on the
west side of the Gulf to the south, for example at San Marcos Island (Emerson
and Puffer 1957). Some of them occur at least sporadically to the north: Mrs.
Howard tells me that Anachis varia and Conus perplexus occur at San Felipe,
Parametaria dupontii at Puertocitos, Conus nux and C. princeps at San Luis
Gonzaga Bay. Additional collecting is needed to show whether all these species
actually are absent from the Los Angeles Bay area and, if so, whether they are
absent also along most of the northwestern Gulf coast. Adequate temperature
data are not available to demonstrate that the absence of these species at Los
Angeles Bay can be accounted for by year-around cool temperatures maintained
by upwelling, particularly in the summer. Their exclusion could result from
failure of the area to reach temperatures necessary for reproduction.

Less well known is a faunal element that appears to be restricted to the
extreme northern and northwestern shores of the Gulf. Species in this category
that are common at Los Angeles Bay but rare or absent at Guaymas on the
eastern shore include Acmaea strongiana, Nomaeopelta dalliana, Acanthina an-
gelica, Crassispira nymphia, and Acanthochitona exquisita.

McLean — Mollusks of Los Angeles Bay 457

Systematic Account

The nomenclature and the order in the checklist follow Keen (1958) except
that the arrangement of the chitons follows Smith (1960). Most of the species
are illustrated in Keen's book; and illustrations of the minute-shelled species,
which she omitted, may be located through her bibliography. The dates of de-
scription of species are given only for those species that she did not include.

The collecting stations referred to in the list (west shore rocks, sand flat,
sandspit, east shore rocks, estero, islands) are shown on the map (fig. 1). The
collector's initial is given in parentheses: B for a collection made by Mr. Robert
Martin in April 1957, now in the collection of Dr. S. Stillman Berry; F for a
collection made by Mr. Wesley Farmer in April 1960; H for Mrs. Howard's
collection made September 9 and 10, 1957, and February 22 to April 2, 1960;
M for the writer's collection, June 9 to 12, 1960; S for the collection made by
Miss Josephine Scripps in March 1960. The Farmer (F) and Scripps (S)
collections are in the San Diego Museum of Natural History. Los Angeles Bay
records of Boone (1928) and Steinbeck and Ricketts (1941) have been included
for the few species of their lists not otherwise recorded.

Twenty-three of Mrs. Howard's records cited as "on Spondylus" were taken
from 18 shells of Spondylus princeps, presumably from relatively deep water,
which she obtained from a local fisherman. A large number of species are re-
ported only "from drift". These minute shells, deposited on the far end of the
sandspit by the outgoing tide, were laboriously sorted by Mrs. Howard. The
species referred to as beach and drift shells were not found living by us. Many
of these live in deeper water according to the bathymetric ranges given by Keen
(1958). Little attempt has been made to indicate relative abundance, except for
the mention of common and scarce species. All live-collected specimens were
taken at low tide unless indicated otherwise.

Class Pelecypoda

NUCULIDAE

Nucula cf. N. declivis Hinds. Worn valves from drift (H) .
Nucula cf. N. Unki Dall. A fragment from drift (H) .

Arcidae
Area (Area) mutabilis (Sowerby) . East shore under rocks (H) .
Area (Area) pacifica (Sowerby) . East shore under rocks (H) .
Barbatia (Barbatia) lurida (Sowerby) . Attached to under surfaces of large rocks

(M).
Barbatia (A car) gradata (Broderip and Sowerby) . Under rocks (H) .
Barbatia (Cueullaearca) reeveana (Orbigny) . West shore under rocks (H) .
Barbatia (Fugleria) illota (Sowerby). Under large rocks (H,F).
Areopsis solida (Sowerby) . Abundant on undersurfaces of rocks (H,M,F) .
Anadara (Larkinia) multieostata (Sowerby) . Half buried in sand (H,M) .
Anadara (Scapharca) cepoides (Reeve). Just under surface of sand on sandspit

(H).

458 San Diego Society of Natural History

Glycymeridae

Glycymeris (Glycymeris) bicolor (Reeve). Fresh valves on sandflat (H).
Glycymeris (Glycymeris) gigantea (Reeve). Living on side in sand near rocks

(H,M).
Glycymeris (Glycymeris) maculata (Broderip) . Just below surface of coarse sand

(HM).
Glycymeris (Glycymeris) multicostata (Sowerby). Just below surface of sand

(H,M).

Mytilidae

Brachiodontes multiformis (Carpenter) . On rocks among barnacles (H,M) .

Hormomya adamsiana (Dunker) . Abundant at mid-tide level among oysters and
large barnacles (H,M).

Crenella divaricata (Orbigny) . Valves from drift (H) .

Litbophaga (Microforceps) aristata (Dillwyn). Boring in vermetids and nest-
ling among barnacles (H,M,B) .

Lithophaga (Labis) attenuata rogersi Berry. Living in worm tubes (H) .

Litbophaga (Leiosolenus) spatiosa (Carpenter). Valves from drift (H).

Gregariella cf. G. chenui (Recluz) . On Spondylus (H) .

Modiolus capax (Conrad) . Exposed among rocks and cobbles (H,M,F,S,B) .

Pteriidae

Pteria sterna (Gould) . Fresh valves on outer side of sandspit; living attached to
gorgonians (H,F) .

PlNNIDAE

Pinna rugosa (Sowerby) . Anchored in sand among rocks on outer side of sand-
spit and on east shore (H,M,F) .

Atrina tuberculosa (Sowerby). In sand near rocks, especially on east shore
(H,M,F).

ISOGNOMONIDAE

Isognomon (Melina) chemnitzianus (Orbigny). Common attached to under-

surfaces of rocks, low to mid-tide levels (H,M) .
Isognomon (Melina) janus Carpenter. Under rocks; less common (H,M,F) .

OSTREIDAE

Ostrea angelica Rochebrune. East shore, on cobbles (H) .

Ostrea columbiensis Hanley. Valves on beach (H) .

Ostrea conchaphila Carpenter. Common attached to rock among small barnacles

at mid-tide level (H,M) .
Ostrea fisberi Dall. Valves from islands (H,M) .

Ostrea palmula Carpenter. Common in bay, exposed on rocks at mid- tide level
(H,M,B).

McLean — Mollusks of Los Angeles Bay 459

Pectinidae

Pecten vogdesi Arnold. Valves common on beach (H,M) .

Aequipecten (Plagioctenium) circularis (Sowerby). Valves on beach (H,M).

Aequipecten (Leptopecten) latiauratus (Conrad, 1837). One specimen attached

to Spondylus (H).
Cyclopecten pernomus (Hertlein) . Valves in drift (H) .
Lyropecten (Lyropecten) subnodosus (Sowerby) . Valves on beach; living sub-

tidally near islands (H,M,F,S).

Limidae

Lima (Lima) tetrica Gould. One juvenile from islands (M) .
Lima (Limaria) hemphilli Hertlein and Strong. Valves (H) .
Lima (Promantellum) pacifica Orbigny. Valves (M) .

Spondylidae

Spondylus princeps Broderip. Subtidally from islands (H).

Spondylus calcijer Carpenter. Large rocks on islands at low tide level (H) .

Plicatula spondylopsis Rochebrune. Attached to rocks, west shore (M).

Anomiidae

Anomia adamas Gray. Plentiful on cobbles outside sandspit (H) .
Anomia peruviana Orbigny. On cobbles outside sandspit (H,M,F,S).
Pododesmus cepio (Gray) . Common on rocks and pelecypod valves (H,M) .
Pododesmus (Tedinia) pernoides (Gray). Large specimens on subtidal boulders

off islands. (M) .
Placunanomia cumingii Broderip. Valves from sandspit (H) .

Crassatellidae

Crassatella (Hybolophus) digueti Lamy. Valves, east shore (H) .
Crassinella pacifica (C. B. Adams) . Valves, with Spondylus (H) .

Carditidae

Cardita afjinis Sowerby. Under rocks (H) .

Cardita affinis var. calif ornica Deshayes. Abundant under rocks in bay (H,M,F) .

Lucinidae

Lucina (Bellucina) cancellaris Philippi. Valves from drift (H) .
Lucina (Callucina) lampra (DAY) . Valves from drift (H) .
Lucina (Parvilucina) approximata (Dall) . Valves from drift (H) .
Anadontia edentuloides (Verrill). Valves (H.M).
Codakia distinguenda (Tryon) . Valves (H,M).
Ctena mexicana (Dall) . One valve from drift (H) .
Divalinga eburnea (Reeve). Valves (H,M).

460 San Diego Society of Natural History

DlPLODONTIDAE

Diplodonta (Diplodonta) inezensis (Hertlein and Strong). Valves (H).
Diplodonta (Diplondonta) orbellus (Gould) . Nestling among rocks (H,M) .
Diplodonta (Diplodonta) subquadrata (Carpenter). Nestling on east shore

among rocks (H) .
Diplodonta (Felaniella) sericata (Reeve) . Valves (H,M,F) .

Erycinidae

Aligena cokeri Dall. Valves from drift (H) .

Aligena nucea Dall. Valves from drift (H) .

Basterotia peninsularis Jordan. Valves from drift (H) .

Galeomma mexicanum Berry, 1959. Valves from drift (H) .

Kellia sub orbicularis (Montagu) . Juveniles nestling en Spondylus (H) .

Mysella cf. M. compressa (Dall) . Valves from drift (H) .

Solecardia eburnea Conrad. Valves (H) .

Chamidae
Chama buddiana C. B. Adams. On rocks north of sandspit (H,F) .
Chama jrondosa mexicana Carpenter. East shore (H). Known range extended

north from La Paz.
Chama sordida Broderip. A large specimen on Spondylus (H) .
Chama squamuligera Pilsbry and Lowe. Juveniles on Spondylus; adults from

east shore (H).
Pseudochama corrugata (Broderip) . On rocks, east shore (H) . Known range

extended north from southern Mexico.
Pseudochama panamensis (Reeve). On cobbles, east shore (H). Known range

extended north from Panama.
Pseudochama saavedrai Hertlein and Strong. Common on rocks of west shore

(M).

Cardiidae

Trachycardium (Trachycardium) consors (Sowerby). Just under sand on sand-
spit (H,M).

Trachycardium (Mexicardia) panamense (Sowerby). Just under sand inside
sandspit (H).

Papyridea aspersa (Sowerby) . Valves common (H,M).

Trigonicardia (Trigonicardia) granulifera (Broderip and Sowerby). Valves
with Spondylus (H) .

Trigonicardia (Americardia) biangulata (Broderip and Sowerby). Just under
sand on sandflat (H) .

Laevicardium clarionense (Hertlein and Strong) . Just under sand on sandflat

(H).

Laevicardium elatum (Sowerby) . Fresh valves on sandflat (H) .
Laevicardium elenense (Sowerby). Just under sand on sandflat (H,M).
Laevicardium elenense var. apicinum (Carpenter). Just under sand on sandflat

(H)\

Nemocardium (Micro car dium) pazianum (Dall). Valves with Spondylus (H).

McLean — Mollusks of Los Angeles Bay 461

Veneridae

Periglypta multicostata (Sowerby) . Valves from midden (H) .

Gouldia calif ornica Dall. Valves in drift and on Spondylus (H). Known range

extended north from La Paz.
Transennella tantilla (Gould) . Common in drift (H) .
Pitar (Pitar) newcombianus (Gabb). Just under surface of sand (H).
Megapitaria squalida (Sowerby). Just under surface of sand on sandflat (H).
Dosinid ponderosa (Gray). Living below surface on sandflat (H,M).
Chione (Chione) californiensis (Broderip). Among rocks (H,M).
Chione (Chione) undatella (Sowerby). Among rocks (H,M).
Chione (Chionista) cortezi (Carpenter). Valves (H).
Chione (Timoclea) picta Willet. Valves (H) .
Anomalocardia subimbricata (Sowerby). Valves (S). Known range extended

north from La Paz.
Anomalocardia tumens (Verrill). Valves common on islands (H,M).
Protothaca grata (Say) . Abundant on sand, near rocks (H,M,F,S) .
Protothaca (Leukoma) asperrima (Sowerby). Valves (H,M).

Petricolidae
Petricola (Rupellaria) robusta Sowerby. Valves (H).

COOPERELLIDAE

Cooperella subdiaphana (Carpenter) . Valves from drift (H) .

Mactridae

Mactra (Mactrotoma) dolabrijormis (Conrad) . One valve (H).

Mactra (Micromactra) fonsecana Hertlein and Strong. Valves (H). Known

range extended north from Nicaragua.
Anatina (Raeta) undulata (Gould) . Valves (H,M).
Mulinia coloradoensis Dall. Valves (H) .

Tellinidae

Tellina (Merisca) reclusa Dall. Valves (H) .

Tellina sp. Abundant in drift (H). A small tellin to be described by Dr. Myra
Keen.

Apolymetis cognata (Pilsbry and Vanatta) . One valve (H). Known range ex-
tended north from Nicaragua.

Macoma indentata Carpenter, 1866. Valves common on sandflat (H,M) . Known
range extended south into the Gulf.

Strigilla cicercula (Philippi). Valves (S).

Strigilla costulifera (Mbrch) . Valves (H).

462 San Diego Society of Natural History

Sanguinolariidae

Gari (Psammocola) regularis (Carpenter). Valves on sandflat (H).

Heterodonax bimacidatus (Linnaeus). Abundant in gravel, especially on is-
lands, low to mid-tide levels (H,M,S) .

Tagelus (Tagelus) calif ornianus (Conrad) . Valves on sandflat (H,M,F) .

Tagelus (Mesopleura) bourgeoisae Hertlein. Valves (H) . Known range ex-
tended north from Salina Cruz, Mexico.

Tagelus (Mesopleura) peruvianus Pilsbry and Olsson. Valves (H) .

Semelidae
Semele bicolor (C. B. Adams) . Valves (H) .
Semele corrugata calif ornica (Reeve) . Valves from islands (M).
Semele flavescens (Gould) . Valves (H,M) .
Semele guaymasensis Pilsbry and Lowe. Valves (H) .
Semele jaramija Pilsbry and Olsson. One valve from drift (H) . Second record

for this species.
Abra tepocana Dall. Valves with Spondylus (H) .
Cumin gia lamellosa Sowerby. Valves from drift (H) .

Myacidae
Cryptomya calif ornica (Conrad) . Valves (H).

CORBULIDAE

Corbula (Caryocorbula) luteola Carpenter. Under rocks (H,M) .
Corbula (Tenuicorbida) fragilis Hinds. With Spondylus (H) .

Gastrochaenidae
Gastrocbaena ovata Sowerby. Valves from drift (H) .

HlATELLIDAE

Hiatella arctica (Linnaeus) . Nestling under rocks (H,M) .

Lyonsiidae
Lyonsia (Lyonsia) gouldii Dall. Just under surface of sand (H) .
Lyonsia (Phlycticoncha) lucasana Bartsch and Rehder. Valves (H) .

Thraciidae
Thracia squamosa Carpenter. One valve (H) .
Cythodonta lucasana Dall. One valve (H) .

Class Scaphopoda

Dentaliidae
Dentalium (Graptacme) sp. Fragments from drift (H) .
Dentalium (Tesseracme) quadrangulare Sowerby. Specimens from drift (H).

SlPHONODENTALIIDAE

Cadulus sp. Fragments from drift (H) .

McLean — Mollusks of Los Angeles Bay 463

Class Gastropoda

ACMAEIDAE

Acmaea atrata Carpenter. One specimen from drift (H) .

Acmaea mitella jayae Hertlein. Among small barnacles at mid-tide level (H,M) .

Acmaea pediculus (Philippi) . Uncommon, immediate subtidal level on islands

(M).

Acmaea semirubida Dall. Abundant on under side of subtidal rocks off islands
(H,M).

Acmaea strongiana Hertlein. Abundant on and under rocks at mid-tide level
(H,M).

Acmaea turveri Hertlein and Strong. Among small barnacles at mid-tide level
(H,M).

Nomaeopelta dalliana (Pilsbry). Bay and island rocks, buried in sand during
the day (H,M,F,S,B).

Nomaeopelta mesoleuca (Menke). East shore rocks (H). This may be separ-
able from the southern form.

Nomaeopelta stanfordiana (Berry). Mid-tide level, east shore rocks (H).

FlSSURELLIDAE

Fissurella rugosa Sowerby. Specimens from drift (H,M) .

Diodora alta (C B. Adams) . Common under rocks in bay (H,M) .

Diodora constantiae Kanakoff, 1953. Shells in drift (H) .

Diodora inaequalis (Sowerby). Under rocks in bay (H,M). The aperture is

elongate and shows black mantle tissue and an epipodium mottled with

brown.
Diodora digueti (Mabille). Under rocks in bay (H,M). The aperture is oval

and shows light-colored tissue. The epipodium is gray with small brown

spots.
Hemitoma (Montfortia) hermosa Lowe. A juvenile specimen from drift (H).

Trochidae

Calliostoma angelenum Lowe. Uncommon at low-tide level and subtidally

(H,M).
Calliostoma eximium (Reeve) . Partially buried on sandflat (H) .
Calliostoma marsballi Lowe. Under rocks outside sandspit (H) .
Tegula (Agathistoma) ligulata (Menke). Small specimens under rocks

(H,M,S) .
Tegula (Agathistoma) mariana Dall. Common under rocks (H,M,F,S) .
Tegula (Omphalius) rugosa (A. Adams). Abundant on rocks of east shore

(H,M).
Solariella triplostephanus Dall. One small specimen from drift (H).

464 San Diego Society of Natural History

Turbinidae

Turbo fluctuosus Wood. Common under and on rocks (H,M) .
Turbo saxosus Wood. On rocks of east shore (H,S) .

LlOTIIDAE

Arene hindsiana Pilsbry and Lowe. Two specimens under subtidal rocks on
islands (M) . Known range extended north from Mazatlan.

Phasianellidae
Tricolia ( Eulithidium ) substriata (Carpenter, 1864). One specimen from drift

(M).

Tricolia cf. T. (Eulithidium) typica (Dall, 1908). Specimens from drift (H).

Neritidae
Nerita (Ritena) scabricostata Lamarck. Four specimens; apparently scarce (H).
Nerita (Theliostyla) funiculata Menke. Abundant on rocks at mid-tide level

(H,M).
Neritina luteofasciata Miller. On sandflat and in estero (H,M) .

Phenacolepadidae
Pkenacolepas malonei Vanatta. Specimens from drift (H) .
Phenacolepas osculans (C. B. Adams) . Under large rocks north of sandspit (H) .

EULIMIDAE

Balcis gibba (de Folin, 1867). Two specimens on Spondylus (H). Known

range extended north from Panama.
Balcis mexicana (Bartsch, 1917) . Two specimens under rocks of west shore (M) .
Balcis cf. B. rutila (Carpenter, 1856). Four live specimens from sandspit (H).

Close to the Californian B. rutila.
Balcis, 3 spp. indet. (H;H;H).

Eulima lapazana (Bartsch, 1917). One juvenile on Spondylus (H).
Niso hipolitensis Bartsch, 1917. One specimen from drift (H) .

Epitoniidae

Epitonium (Nitidiscala) apiculatum Dall. Hermit crab shells (H,M).

Epitonium (Nitidiscala) hexagonum (Sowerby). Specimens from drift (H).

Epitonium (Nitidiscala) oerstedianum Hertlein and Strong. Two specimens
from drift (H) .

Opalia (Dentiscala) crenimarginata (Dall). East shore (H).

Opalia (Dentiscala) diadema (Sowerby). East shore (H). Known range ex-
tended north from Mazatlan.

LlTTORINIDAE

Littonna dubiosa C. B. Adams. Common in drift (H,M). Specimens with
varying numbers of keels may also be referable to this species.

McLean — Mollusks of Los Angeles Bay 465

VlTRINELLIDAE

Cyclostremiscus panamensis (C. B. Adams, 1852). Four specimens from drift

(H) . Known range extended north from Mazatlan.
Cyclostremiscus sp. Two specimens from drift (H) .
Macromphalina sp. Three specimens from Spondylus (H) .

Solariorbis elegans Pilsbry and Olsson, 1952. Nine specimens (H). Known

range extended north from Ecuador.
Solariorbis narinensis Pilsbry and Olsson, 1952. One immature shell (H).

Known range extended north from Colombia.
Teinostoma amplectans Carpenter, 1857. Common in drift (H) .
Teinostoma cf. T. herbertianum Hertlein and Strong, 1951. One specimen (H).

Slightly larger than this species described from Costa Rica.
Vitrinella sp. One worn specimen (H) .

Rissoidae

Alvania gallegosi Baker, Hanna, and Strong, 1930. One specimen from drift
(H).

Alvania lirata (Carpenter, 1856) . Abundant in drift (H) .

Alvania monserratensis Baker, Hanna, and Strong, 1930. Common in drift (H).

Alvania sp. One specimen from islands (M) .

Barleeia alderi (Carpenter, 1856) . Common in drift (H,M) .

Barleeia subtenuis (Carpenter, 1856) . Common under rocks (H,M) .

?Diala sp. Five specimens from drift (H) .

Rissoina burragei Bartsch, 1915. Abundant in drift; living at low-tide level
(H,M).

Rissoina io Bartsch, 1915. One perfect specimen from drift (H). Known range

extended north from the Galapagos Islands.
Rissoina, 2 spp. indet. (H;H).

ASSIMINEIDAE
Assiminea compacta (Carpenter, 1863). Specimens from drift (H).

Truncatellidae
Truncatella bairdiana C. B. Adams, 1852. Four specimens from drift (H).

TURRITELLIDAE

Turritella anactor Berry. Shells on beach (H) .

Turrit ella clarionense Hertlein and Strong. One shell on beach (H) .

Vermicularia pellucida (Broderip and Sowerby). Abundant, nestled among
rocks (H,M).

466 San Diego Society of Natural History

Architectonicidae

At chitectonica nobilis Roding. Shells on beach (H) .

Heliacus bicanaliculatus (Valenciennes) . On rocks of east shore (H,M) .
Heliacus mazatlanicus Pilsbry and Lowe. Specimens from drift (H) . Known
range extended north from Mazatlan.

Caecadae

Caecum "firmatum" of authors, not of C. B. Adams. Abundant in drift (H) .
Has no satisfactory name according to Dr. Berry.

Caecum sp. Six juveniles from drift (H) .

Elephantanellum heptagonum (Carpenter, 1857). Common in drift (H).

?Elephantulum mirificum (de Folin, 1867). One specimen from drift (H).

Elephantulum ? sp. One juvenile from drift (H) .

MODULIDAE

Modulus cerodes (A. Adams) . Subtidally from islands (H,M) .
Modulus disculus (Philippi) . One hermit crab shell (H) .

Vermetidae

Vermetus (Thylaeodus) indentatus (Carpenter) . Common on rocks (H,M) .
Serpidorbis (Cladopoda) mar garitaceus (Chenu) . On rocks of west shore (M) .

Cerithiidae

Cerithium maculosum Kiener. Shells on beach (H) .

Cerithium (Liocerithium) sculptum Sowerby. Common on sand near rocks

(H,M,B,S).
Cerithium stercusmuscarum Valenciennes. On sandspit, not common (H).
Ceritbiopsis (Cerithiopsida) kinoi Baker, Hanna, and Strong, 1938. Specimens

from drift (H).
Ceritbiopsis, 3 spp. indet. (H;H;M).

Metaxia diadema Bartsch, 1907. One specimen under rocks of west shore (M) .
Seila assimilata (C B. Adams, 1852). Common under rocks and associated with

the large red sponge (H,M) .
Alaba interruptilineata Pilsbry and Lowe, 1932. Common in drift (H).
Alaba jeannettae Bartsch, 1910. Specimens from drift (H,M) .
Alaba supralirata Carpenter, 1856. Specimens from drift (H) .
Triphora cL T. evermanni Baker, 1926. Worn specimens from drift (H).
Triphora, 2 spp. indet. (H;M) .
Alabina diomedeae Bartsch, 1911. Very abundant in drift (H). One or more of

at least five separable forms may be referred to this species.
Alabina sp. One specimen from drift (H) .

McLean — Mollusks of Los Angeles Bay 467

potamididae

Cerithidea albonodosa Gould and Carpenter. At high-tide level on sandflat

(H,F).
Cerithidea mazatlanica Carpenter. On sandflat (H,M,F) .

HlPPONICIDAE

Hipponix serratus Carpenter. Shells on beach (H,M) .

FOSSARIDAE

Fossarus (s.l.) sp. Under rocks of west shore and in drift (H,M) .

Calyptraeidae
Cheilea cepacea (Broderip) . One specimen on beach (H) .
Crepidula aculeata (Gmelin) . On Spondylus (H) .
Crepidula arenata (Broderip) . On shells (H,S) •
Crepidula excavata (Broderip) . On shells (H,M,F) .
Crepidula incurva (Broderip) . On shells (M) .
Crepidula onyx Sowerby. On shells (H,M,S) .
Crepidula perforans (Valenciennes) . In shell apertures (H,M) .
Crepidula uncata Menke. Shells from drift (H) .

Crucibulum scutellatum (Wood) . On or under rocks in bay (H,M,F) .
Crucibidum spinosum (Sowerby) . Small specimens common under rocks
(H,M,F,S,B).

Naticidae

Natica (Natica) chemnitzu Pfeiffer. On sandflat and in estero (H) .
Natica (Stigmaulax) broderipiana Recluz. One shell on beach (S) .
Natica (Stigmaulax) elenae Recluz. Shell on beach of island (H) .
Polinices (Polinices) bifasciatus (Gray) . On sand inside sandspit (H) .
Polinices (Polinices) uber (Valenciennes) . On sandspit (H) .
Polinices (Neverita) reclusianus (Deshayes). On sandflat (H,M,F).

Cypraeidae
Cypraea (Zonaria) annettae Dall. Under rocks in bay (H,M,F,S)-

Eratoidae
Erato (Hespererato) columbella Menke. Shells from drift (H) .
Trivia (Pusula) calijorniana (Gray) . Under rocks in bay (H) .
Trivia (Pusula) elsiae Howard and Sphon, 1960. One shell on beach at east

shore (H) .
Trivia (Pusula) solandri (Sowerby) . Under rocks (H) .
Jenneria pustulata (Solander) . Shells on beach (H) .

Ovulidae
Neosimnia quaylei (Lowe) . On gorgonians in shallow water outside sandspit
(H) . Red- and white-shelled forms.

468 San Diego Society of Natural History

Strombidae
S trombus galeatus Swainson. Shells on beach (H) .
Strombus gracilior Sowerby. On sand near the sandspit (H,M,S) •
Strombus granulatus Swainson. On sand near the sandspit (H) .

Cassididae
Cassis (Semicassis) centiquadrata (Valenciennes). Shells on beach (H).

FlCIDAE

Ficus ventricosa (Sowerby) . Shells on beach (H) .

Cymatiidae
Cymatium (Tumtriton) gibbosum (Broderip) . On rocks in bay (H) .

MURICIDAE

Murex elenensis Dall. Shells on beach (H,M,S) .

Murex recurvirostris Broderip. Shells on beach (H) .

Hexaplex brassica (Lamarck) . Reported by Boone (1928) .

Hexaplex erythrostomus (Swainson). On sand near rocks (H,M,S,B).

Muricanihus nigritus (Philippi). One shell on beach (H). Apparently scarce.

Muricanthus princeps (Broderip). Immature specimen from islands (H).

Pterynotus erinaceoides (Valenciennes) . On rocks in bay (H) .

Ocenebra parva (E. A. Smith) . One shell, from islands (M) .

Eupleura muriciformis (Broderip). On rocks north of sandspit (H).

Eupleura nitida (Broderip). One shell on beach (H). Known range extended

north from Mazatlan.
Muricopsis armatus (A. Adams) . Shells on beach (H) .
Typhis (Typbinellus) quadratus Hinds. On base of gorgonian (H) .
Coralliophila (Coralliophila) hindsii (Carpenter). One shell on beach (M).
Acanthina (Acanthina) angelica I. Oldroyd. Abundant on rocks at mid-tide

level (H,M,F,S).
Acanthina (Acanthina) tyrianthina Berry. On rocks of east shore (H).
Acanthina (Neorapana) tuberculata (Sowerby). On rocks (H,M,F,S,B).
Morula ferruginosa (Reeve). Abundant on rocks in bay and on islands (H,M,

F,B).

COLUMBELLIDAE

Anachis coronata (Sowerby) . On rocks, not common (H,M) .
Anachis (Glyptanachis) hilli Pilsbry and Lowe. Scarce, under rocks (H).
Decipifus lyrta (Baker, Hanna, and Strong). One shell from drift (H).
Decipifus gracilis McLean, 1959. Under rocks in bay (H) . Second record;

described from Guaymas.
Mitrella delicata (Reeve). Shells in drift (H). Known range extended north

from Manzanillo, Mexico.

McLean — Mollusks of Los Angeles Bay 469

Mitrella densilineata (Carpenter). Specimens from drift (H). Known range ex-
tended north from Cape San Lucas.

Mitrella dorma Baker, Hanna, and Strong. Under rocks in bay (H,M,F).

Mitrella grant i Lowe. Drift specimen (H) . Known range extended south from
San Felipe.

Mitrella ocellata (Gmelin) . Common on sand near rocks (H,M) .

Nassarina (Z.anassarina) pammicra Pilsbry and Lowe. Under rocks in bay (H).
Known range extended north from Nicaragua.

?Nassarina, 3 spp. Three separable shells from drift (H;M;M). A genus close
to Nassarina but lacking axial ribs.

Pyrene juscata (Sowerby) . Small specimens common in bay; large specimens on
islands (H,M,F,S,B).

Strombina maculosa (Sowerby). Common on sand near rocks (H,M,S,B)-

BUCCINIDAE

Hanetia macrospira Berry. On rocks near sand (H,M) .

Melongenidae
Melongena patula (Broderip and Sowerby). Reported by Boone (1928).

Nassariidae
Nassarius angulicostis (Pilsbry and Lowe). Shells from drift (H).
Nassarius versicolor (C. B. Adams) . Shells on beach (M,S) .
Nassarius ( Arcularia) iodes (Dall). Inside sandspit (H).
Nassarius (Arcularia) moestus (Hinds). Common on sandflat (M).
Nassarius (Arcularia) tiarula (Kiener). Abundant on sandflat (M,F,S,B).

Fusinidae
Fusinus (Fusinus) dupetitthouarsi (Kiener) . Shells on beach (H) .
Fusinus ambustus (Gould) . Bay rocks (H) .
Fusinus (Aptixis) cinereus (Reeve). Common on and under rocks (H,M,B)-

Olividae
Oliva (Oliva) spicata (Roding) . On sandflat (H,M) .
Olivella (Olivella) dama (Wood) . On sand near rocks (H,M,F,S) .
Olivella (Olivella) fletcherae Berry.. Specimens from drift (H) .

Mitridae
Mitra (Mitromica) solitaria C. B. Adams. Common under rocks in bay (H,M).
Mitra (Strigatella) dolorosa Dall. East shore rocks and subtidally off islands

(HJM).

Mitra (Strigatella) mexicana Dall. One specimen on beach (H). The second

record of this species.
Mitra (Tiara) erythrogramma Tomlin. One hermit crab shell (S) . Known

range extended north from Nicaragua.

470 San Diego Society of Natural History

Harpidae
liar pa crenata Swainson. Recorded by Boone ( 1928) .

VOLUTIDAE
Lyria (Enaeta) cumingii (Broderip). On sand near rocks (H,M,S).

Marginellidae

Cypraeolina pyrijormis (Carpenter, 1865). Common under rocks and in drift

(H,M).
Cystiscus politulus (Carpenter, 1857) . Abundant in drift (H) .
Marginella calif ornica Tomlin. Common under rocks in bay (H,M,B) .
Marginella cf. M. jewetii Carpenter, 1865. A worn juvenile from drift (H) .

Cancellariidae

Cancellaria (Cancellaria) obesa Sowerby. Shells on beach (H) .

Cancellaria (Euclia) cassidiformis Sowerby. Seven specimens moving under sand

on sandflat in March (H) . Stalked egg capsules (identified by Dr. Gunnar

Thorson) found on sandflat in June (M).

Turridae
Daphnella cf. D. bartschi Dall. One specimen, under rocks on west shore (M) .
Knejastia juniculata (Kiener) . One worn shell on beach (H) .
Knefastia walkeri Berry. One specimen on beach (H) .
Clavus (Cymatosyrinx) sp. One worn juvenile from drift (H) .
Clavus (Elaeocyma) aeginus (Dall). Hermit crab shells on sandflat (M).
Clavus (Imaclava) pembertoni Lowe. Shell on beach (H) .
Crassispira atramentosa (E. A. Smith). One specimen from islands (M).

Known range extended north from Panama.
Crassispira nymphia Pilsbry and Lowe. Common under rocks (H,M,F,S) .
Crassispira pluto Pilsbry and Lowe. Under rocks (H,M) .
Mangelia (Mangelia) barbarensis I. Oldroyd, 1924. Specimens from drift (H) .

Known range extended south into the Gulf.
Mangelia (Agathotoma) pbryne (Dall) . Under rocks on west shore (H,M) .

Known range extended north from Panama.
Mangelia (Agathotoma) sp. Three specimens from drift (H) .
Mangelia cf. M. (Agathotoma) subdiaphana Carpenter. Specimens from drift

(H).
Clathurella (Lioglyphostoma) sp. One perfect specimen (H) . Probably new.
Clathurella (Nannodiella) trichoides (Dall, 1919). One worn specimen with

Spondylus (H).
Notocytharella niobe (Dall). Two specimens from drift (H). Known range

extended north from Cape San Lucas.
Tenaturris nereis (Pilsbry and Lowe). Specimens from drift (H). This species

is now known in collections from other localities in the northern Gulf.

McLean — Mollusks of Los Angeles Bay 471

Tcnaturris phaethusa (Dall, 1919) . Specimens from drift (H,M) .

Tenaturris enryclea (Dall, 1919) . Six specimens from drift (H) .

Turricula (Fusiturricula) dolenta (Dall) . One specimen on beach (H) . Known

range extended north from Panama.
Turricula (Fusiturricula) howelli (Hertlein and Strong) . Two specimens on

beach (H) . Known range extended north from Costa Rica.
?Turricula sp. One worn specimen on beach (H) .
Pleurohria picta (Reeve) . One specimen on beach (H) .
Tiariturris spectabilis Berry. Three specimens on beach (H) .

CONIDAE

Conus tornatus Sowerby. One specimen on beach (H) .

Conus ximines Gray. On sandspit near cobbles (H) .

Conus cf. C. dispar Sowerby. Worn specimens on beach (H) .

Conus recurvus Broderip. Specimens on beach (H) .

Conus regularis Sowerby. Worn shells on sandspit (H) .

Conus cf. C. scalaris Valenciennes. Worn specimens on beach of islands (H,M) .

Terebridae

Terebra (Microtry petes) sp. One specimen with Spondylus (H). Probably
new.

Terebra (Strioterebrum) variegata Gray. Just under surface of sand on sand-
flat (H,M,F,S).

BULLIDAE
Bulla gouldiana Pilsbry. Outer side of sandspit (H,M,F,S) .

Akeridae .

Haminoea angelensis Baker and Hanna. Inner side of sandspit (H).
Haminoea strongi Baker and Hanna. Inner side of sandspit (H,M).

SCAPHANDRIDAE

Atys casta Carpenter. Specimens from drift (H). Known range extended north
from Cape San Lucas.

ACTEONIDAE

Acteon trask'i Stearns. Specimens from drift (H) .

ACTEOCINIDAE

Acteocina carinata (Carpenter) . Common in drift (H) .
Acteocina injrequens (C. B. Adams) . Common in drift (H,M) .
Sulcoretusa paziana (Dall, 1919) . One specimen from drift (H) .

Pyramidellidae
Pyramidella (Longchaeus) adamsi Carpenter. Five specimens from drift (H) .
Odostomia (Cbrysallida) viscainoana Baker, Hanna, and Strong, 1928. Under
rocks in bay; common in drift (H,M) .

472 San Diego Society of Natural History

Odostomia (Chrysallida) audax Baker, Hanna, and Strong, 1928. One speci-
men from drift (H). Known range extended north from Cape San Lucas.

Odostomia (Chrysallida) sp. One fresh specimen from Spondylus (H).

Odostomia (Pyrgulina) herrerae Baker, Hanna, and Strong, 1928. Specimens
from drift (H,M).

Turbonilla (Chemnitzia) muricata (Carpenter, 1856). Two specimens from
Spondylus (H) . Carpenter's type was likewise on Spondylus, upon which
this Turbonilla may be parasitic.

Turbonilla (Strioturbonilla) buttoni Dall and Bartsch, 1909. One specimen
from drift (H) .

Turbonilla, 3 spp. indet (H;H;M) .

Iselica fenestrata (Carpenter, 1864) . Three adults and one juvenile on Spondylus
(H).

Placement of the genus Iselica in the Pyramidellidae has recently been sug-
gested by Dr. S. Stillman Berry following Theile (1929, p. 233) . The treatment
is supported by study of Iselica fenestrata, which has a partially heterostrophic
nucleus, a single columellar fold, and an operculum similar in shape and mor-
phology to that of Odostomia (Evalea) sp., and which shows no indication of
a radula in softened specimens. The present record of the species on Spondylus
is also suggestive of the parasitism characteristic of pyramidellids.

Iselica cf. /. ovoidea (Gould, 1852). Specimens from drift (H).

Cavolinidae
Carolina longirostris (Deshayes in Lamarck, 1836, ex Leseur MS). One speci-
men from drift (H) .

Ellobiidae
Melampus olivaceus Carpenter. Common in beach drift at high-tide level (H,M) .

SlPHONARIIDAE

Siphonaria maura Sowerby. Reported by Steinbeck and Ricketts (1941) [as
S. aequilirata Reeve] to be common on rocks near sand. Not seen by us.

Williamia peltoides (Carpenter) . Common on undersurfaces of subtidal rocks
off islands (M).

Trimusculidae

Trimusculus reticularis (Sowerby). Specimens from drift (H).
Onchidella binneyi (Stearns). Abundant under rocks at mid-tide level, exposed
at night (H,M,F) .

Class Amphineura

Lepidopleuridae

Lepidopleurus (Leptochiton) cancellatus Sowerby, 1839. Two specimens (H).
Apparently not differing from this northern species, according to Mr. Smith,
and thus extending the known range from southern California into the Gulf.

McLean — Mollusks of Los Angeles Bay 473

ischnochitonidae
Ischnochiton (Radsiella) tridentatus Pilsbry. Four specimens (M).
Lepidozona clathrata (Reeve) . Common under rocks (H,M) .
Lepidozona subtilis Berry. Abundant under rocks (H,M) .
Stenoplax ci. S. aetbona (Dall). One specimen (M). Needs to be compared

with the type of this species according to Dr. Berry.
Stenoplax I imaci for mis (Sowerby). Under rocks (H,M,B) .
Stenoplax mariposa (Dall). Abundant under rocks (H,M,F). Many color

phases.
Stenoplax conspicua sonorana Berry. Under rocks (H,M,B) .
Lepidoehitona k^epiana Berry, 1948. One specimen (H) . Known range extended

from San Ignacio Lagoon into the Gulf according to Mr. Smith.
Callistochiton gabbi Pilsbry. Under rocks (H,M) .
Callistochiton infortunatus Pilsbry. Under rocks (H,M).
Nuttalina crossota Berry. Four specimens under rocks in bay (H) .

Chaetopleuridae
Chaetopleura lurida (Sowerby). Recorded by Steinbeck and Ricketts (1941).
Chaetopleura mixta (Dall). Described from Los Angeles Bay. Not seen by us.
Chaetopleura sp. A juvenile (M) .

Mopaliidae

Placiphorella blainvillei (Broderip). One specimen found on a large Pinna
rugosa (H) . A new record, extending the known range north from Pan-
ama according to Mr. Smith.

Chitonidae
Chiton virgulatus Sowerby. Common under rocks (H,M) .

ACANTHOCHITONIDAE

Acanthochitona cf. A. angelica Dall. Three specimens (H). Needs to be com-
pared with the type of this species according to Mr. Smith.

Acanthochitona exquisita (Pilsbry). Abundant under rocks at mid-tide level
(H,M,F).

Phylum BRACHIOPODA
Class Testicardines

DlSCINIDAE

Discinisca cumingii (Broderip) . Many specimens on Spondylus (H) .
Discinisca strigata (Broderip) . Equally abundant on Spondylus (H) .

Terebratellidae
Argyrotheca lowei Hertlein and Grant. Four specimens on Spondylus (H).

474 San Diego Society of Natural History

Acknowledgments

While many people have given help in connection with this paper, I must
give special recognition to Mrs. Faye Howard, whose collection forms the main
basis of the list. She has made her collection available and has given me infor-
mation about collecting conditions. Appreciation is due to Dr. A. Myra Keen
of Stanford University who has been generous with her time and advice. Thanks
are also due to Messrs. S. Stillman Berry, Emery P. Chace, Leo G. Hertlein, and
Allyn G. Smith for help with identifications, and to Messrs. Joshua L. Baily, Jr.,
William K. Emerson, and Reid Moran for suggestions and assistance in the
preparation of this paper. I am grateful for the use of the facilities of the Ver-
milion Sea Field Station, which has been equipped with funds from the National
Science Foundation.

Literature Cited

Anderson, C. A.

1950 1940 E. W. Scripps cruise to the Gulf of California. Part I. Geo-
logy of islands and neighboring land areas. Geol. Soc. Am. Mem.
43:' 1-53.

Boone, Lee

1928 Scientific results of the second oceanographic expedition of the
"Pawnee," 1926. Mollusks from the Gulf of California and the
Perlas Islands. Bull. Bingham Oceanogr. Coll. 2: 1-17.

Emerson, W. K.

1958 Results of the Puritan- American Museum of Natural History
expedition to western Mexico. 1. General account. Am. Mus.
Novitates, 1894: 1-25.

Emerson, W. K., and E. L. Puffer

1957 Recent mollusks of the 1940 "E. W. Scripps" cruise to the Gulf
of California. Am. Mus. Novitates, 1825: 1-57.

Fraser, C. M.

1943 General account of the scientific work of the Velero III in the
eastern Pacific, 1931-41. Part III. A ten-year list of the Velero III
collecting stations. Allan Hancock Pac. Exp. 1: 259-431.

Hertlein, L. G, and W. K. Emerson

1956 Marine Pleistocene invertebrates from near Puerto Penasco, Sonora,
Mexico. Trans. San Diego Soc. Nat. Hist. 12: 154-176.

McLean — Mollusks of Los Angeles Bay 475

Keen, A. M.

1937 An abridged check list and bibliography of West North Ameri-
can marine Mollusca. 1-87, figs. 1-2. Stanford University Press.

1958 Sea shells of tropical West America; marine mollusks from Lower
California to Colombia, i-xi, 1-624, illus. Stanford University Press.

Lowe, H. N.

1935 New marine mollusca from west Mexico, with a list of shells col-
lected at Punta Penasco, Sonora, Mexico. Trans. San Diego Soc.
Nat. Hist. 8: 15-34.

Oldroyd, I. S.

1918 List of shells from Angel and Tiburon Islands, Gulf of California,
with a description of a new species. Nautilus 32: 26-27.

Roden, G. I., and G. W. Groves

1959 Recent oceanographic investigations in the Gulf of California. Jour.
Mar. Res. 18: 10-35.

Shepard, F. P.

1950 1940 E. W. Scripps cruise to the Gulf of California. Part III.
Submarine topography of the Gulf of California. Geol. Soc. Am.
Mem. 43: 1-32.

Slevin, J. R.

1923 Expedition of the California Academy of Sciences to the Gulf of
California in 1921. General account. Proc. Calif. Acad. Sci., ser.
4. 12: 55-72.

Smith, A. G.

1960 Amphineura. In Treatise on invertebrate paleontology, Geol. Soc.
Am. Part I, Mollusca 1:41-76.

Squires, D. F.

1959 Results of the Puritan- American Museum of Natural History
Expedition to western Mexico. 7. Corals and coral reefs in the Gulf
of California. Bull. Am. Mus. Nat. Hist. 118:367-431.

Steinbeck, John, and E. F. Ricketts

1941 Sea of Cortez, i-x, 1-598, pis. 1-40. Viking Press, New York.

Thiele, Johannes

1929 Handbuch der systematischen Weichtierkunde. Teil I, 1-376.
Gustav Fischer, Jena.

TOWNSEND, C. H.

1916 Voyage of the "Albatross" to the Gulf of California in 1911. Bull.
Am. Mus. Nat. Hist. 35: 399-476.

476

San Diego Society of Natural History

Fig. 2. A cove on one of the small islands near Los Angeles Bay, showing the
steep intertidal zone composed of small boulders.

, /C»

Fig. 3. The Vermilion Sea Field Station from low water line, showing the
intertidal zone.

] 3 -.

TRANSACTIONS

OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY
Volume XII, No. 29, pp. 477-480 August 30, 1962

TWO NEW SPECIES OF BROAD-FACED,

FIVE-TOED KANGAROO RATS

(GENUS DIPODOMYS)

BY

Laurence M. Huey

Curator Emeritus of Birds and Mammals, San Diego Society of Natural History
The discovery of an isolated population of broad-faced kangaroo rats on
the higher mesas of the Sierra San Borja in Baja California and another along
the San Luis Rey River valley in northern San Diego County, California, is of
outstanding interest. Thus two species are added to the mammal relicts whose
ancestors must have once occupied much larger areas, but which are known only
from a broken chain of small living populations in small restricted localities.
Three such remnant mammalian populations in northern and central Baja Cali-
fornia are (1) a chickaree, Tamiascinrus douglasii mearnsi, on the forested sum-
mit of the Sierra San Pedro Martir; (2) a large broad-faced kangaroo rat,
Dipodomys gravipes, on the low coastal mesas and plains about San Quintin; and
(3) a striped chipmunk, Eutamias merriami meridionalis, known only from a
very limited region of steep canyons and cactus-covered areas at 1000 to 1200
feet elevation about San Pablo, 25 miles north of San Ignacio. Two such species
living in southern California are a broad-faced kangaroo rat, Dipodomys stephensi,
in the arid Sonoran Zone in the San Jacinto Valley region of Riverside and
southwestern San Bernardino Counties, and a large silky pocket mouse, Perog-
nathus alticolus, in the pine-bracken softsoiled area at 5000 feet altitude in the
western San Bernardino Mountains. Others could be named farther to the north,
but these will suffice.

In March 1960, Mr. Glen Ives, then a staff member of the Natural History
Museum, made a small collection of mammals at the San Juan Mine in central
Baja California, Mexico. This collection included a new species of kangaroo rat
{Dipodomys) . This discovery led to a renewed interest in a specimen collected
by the writer in 1920 near Bonsall, San Diego County, California, and called
"an anomaly" by Grinnell (1922, p. 66). Two more specimens collected near
Bonsall in the summer of 1961 show this "anomaly" to be a well defined new
species. Both species are described below.

Dipodomys antiquarius sp. nov.
San Borja Kangaroo Rat

Type. — Adult female; from the San Juan Mine, Sierra San Borja, Baja
California, Mexico, alt. 4000 feet, lat. 28°41'N, long. 113°37/W; collected by
Glen Ives, March 30, 1960; No. 18901, collection of the San Diego Society
of Natural History.

478 San Diego Society of Natural History

Description. — Dipodomys antiquarius is a middle-sized five-toed member
of the broad-faced heermanni group. The general dorsal coloration is light buff.
The cheeks and area surrounding the eyes are lighter colored, almost white, this
light area continuing to the ears and in some specimens extending around the
ears to form conspicuous white areas behind them. The sides are much lighter
colored than the back, the white of the under parts intermingling well above the
break-line between the belly and the sides. The hip stripe is broad and white. The
ears are blackish inside and behind, with buff-colored folded edges and scarcely
any white on the tip of the pinna. The tail is moderately long, with white side
stripes about as wide as the upper and lower black stripes, the white stripes ex-
tending well into the fairly large tail tuft. The arietiform facial marking is nar-
row, almost obscure, near the nose but is broad and black at the base of the longer
whiskers. The eyelids are black and prominent in the light-colored facial area.
The plantar stripes on the hind feet are well marked and black.

The skull of D. antiquarius has a slender rostrum with broad-angled, wide
zygomatic arches. The jugals are straight. The mastoid bullae are well inflated
but not to the point of crowding the interparietal into a deep sulcus. The audi-
tory bullae are large and bulging, giving the skull its greatest breadth.

Measurements of Type. — Total length, 274; tail, 155; hind foot, 40;
ear (dried skin), 10; greatest length of skull, 39.1; width across bullae, 26.1;
spread of maxillary arches, 21.3; greatest length of nasals, 13.7; width of maxil-
lary arch at middle, 5.1.

Comparisons. — The nearest relative so far known is Dipodomys stephensi,
found only in the San Jacinto Valley region of Riverside and extreme south-
western San Bernardino Counties, California. Dipodomys antiquarius is about
equal in size but is lighter colored dorsally and has whiter and lighter colored
facial markings. The white tail stripes are as wide as the ventral and dorsal black
stripes rather than narrower, as in D. stephensi, and they extend farther into the
tip of the tail, making a much lighter colored tail tuft.

The greatest differences, however, are cranial. Dipodomys antiquarius has
a more slender rostrum and slightly less angled zygomatic arches than has D.
stephensi. The auditory bullae are larger and more swollen over-all, and the
mastoid bullae are much more inflated. The pterygoid processes are heavier and
not parallel like those of D. stephensi.

To Dipodomys gravipes, the only other broad-faced kangaroo rat found in
Baja California, D. antiquarius shows but little resemblance except in the wide-
spreading maxillary arches. In all respects, D. antiquarius is less robust within
and without.

Range.— So far as known, the region, about 4000 feet altitude, at the San
Juan Mine, in the Sierra San Borja.

Remarks. — The isolated species in restricted spots southward to the mid-
peninsula region, probably are remnants of wider spread former populations. The
Pliocene submersion of the central peninsula and the volcanic and orogenic
periods doubtless broke up the old populations and left many areas that were not
reinhabitable by the small, less vigorous remnants. Possibly also, more vigorous

Huey — Two New Kangaroo Rats

and better adapted species helped confine these relicts to circumscribed ranges.
There seems but little doubt that the wide-faced character shown in several well-
marked species of Dipodomys is very old.

Dipodomys cascus1 sp. nov.
Bonsall Relict Kangaroo Rat

Type. — Adult male; from 1 mile east of Bonsall, San Diego County,
California; altitude 350 feet; collected by Laurence M. Huey, August 17, 1961;
No. 18961, collection of the San Diego Society of Natural History.

Description. — Dipodomys cascus is a medium-sized, five-toed, broad-faced
species and is also a member of the heermanni group. The dorsal color is light
wood-brown with slightly paler cheeks, face, and sides. The lighter color on the
sides does not extend as high toward the back as on some other kangaroo rats
and is in sharp contrast with the white underparts at the break-line. The hip
stripes are broad and white. The ears are small and are black inside and on the
folded edges, with a conspicuous white tip on the end of each pinna. Above each
eye is a prominent roundish white spot, 2-3 mm. in diameter. The eyelids are
black and appear as rings around the eyes. The arietiform facial marking is
black, broad, and heavy, extending from each whisker patch across the nose. The
bicolored tail, which is noticeably heavy-boned in the flesh, is of medium length.
The white side stripes are narrower than the black upper and lower stripes and
extend well into the scantily tufted tail tip.

The skull of Dipodomys cascus has several outstanding characters. The
mastoid bullae are only moderately inflated in breadth but are clearly bulbous
and squarish when viewed posteriorly. The zygomatic arches are extremely wide-
spread, closely rivaling the mastoid bullae in forming the widest part of the
skull. This character gives a rounder appearance when viewed dorsally, rather
than the triangular shape shown in the skulls of many other species of Dipo-
domys. The maxillary arches are exceptionally wide and the jugals are not parallel
but are bowed outward at the middle.

Measurements of Types. — Holotype: total length, 272; tail, 158; hind
foot, 39; ear, 10; greatest length of skull, 37.4; width across bullae, 24.8; spread
of maxillary arches, 23.7; greatest length of nasals, 13.4; width of maxillary arch
at middle, 6.1; weight, 68.5 gr. Female paratype (No. 18962, coll. San Diego
Society of Natural History) : total length, 270; tail, 156; hind foot, 40; ear, 11;
greatest length of skull, 39.3; width across bullae, 24.8; spread of maxillary
arches, 23.0; greatest length of nazals, 13.4; width of maxillary arch at middle,
6.1; weight, 64.0 gr.

Comparisons. — Dipodomys cascus resembles Dipodomys agilis simulans
in general outward appearance. They live on common ground and must be
closely examined to notice the slightly lighter dorsal color and smaller ears of
D. cascus. However, when the skulls are compared, the two species are far apart
and easily determined.

The nearest wide-faced relative is Dipodomys stephensi, found at the much

1 =01d (Latin), in reference to its ancient lineage.

480 San Diego Society of Natural History

higher elevation of the San Jacinto Valley region north of the Palomar Mountain
chain. Compared with D. stephensi, D. cascus is darker in over-all dorsal colora-
tion and has a slightly flatter and wider skull, with wider spreading zygomatic
arches and bowed jugals. Viewed posteriorly, the bullae of D. cascus are notice-
ably deeper and more squarish than those of D. stephensi. Compared with Dipo-
domys antiquarius , the wide-faced species from central Baja California, D. cascus
is similar in size but decidedly darker in dorsal coloration. The skull of D. cascus
differs from that of D. antiquarius in having less inflated bullae, more sharply
angled and wider zygomatic arches, and bowed jugals.

Range. — Known only from a small area at 300 to 400 feet elevation in the
San Luis Rey River valley east of Bonsall, San Diego County, California. The
original specimen was from 5 miles northeast of Bonsall.

Remarks. — The character that most impressed Grinnell in the one specimen
he saw of D. cascus was the solid black tail. He also noted that the spread of the
maxillary arches was 20.5 mm as compared to a mean of 22.5 and a range of 21
to 23.8 for D. stephensi but that the specimen was immature. These two
characters caused him to call the specimen an anomaly: otherwise, he found it
essentially similar to D. stephensi. Actually, the tail was solid black only because
the preparator had accidentally spilled eternal ink on it; and the spread of the
maxillary arches in both new specimens is greater than the mean for D. stephensi.
Thus the two seemingly anomalous characters noted by Grinnell do not hold.
However, the study of the two new specimens shows other differences, as detailed
above, overlooked by Grinnell.

That this struggling relict population of kangaroo rats is still to be found in
the tiny known range, where it has had to contend with poisoning campaigns and
human agricultural invasion, is almost a miracle. Its future seems extremely
doubtful, and before many years this species probably will pass into extinction.

It is also interesting to note that Dipodomys cascus represents another link
in the chain of broad-faced kangaroo rats whose ancestors appear to have lived
in past ages before the great Pliocene disturbances. Careful collecting by "seeing
eyed" mammalogists probably will find other such restricted populations belong-
ing to this broad-faced chain along the foothills in northern Baja California.

SPECIMENS EXAMINED

Dipodomys antiquarius. Baja California: San Juan Mine, Sierra San Borja, 8.

Dipodomys cascus. California: San Diego County — 1 mi. e. of Bonsall, 2.

Dipodomys gravipes. Baja California: 3 mi. s. of San Telmo, 2; Santo
Domingo (type locality), 19; San Quintin Plain, 14; Santa Maria, 1; near San
Quintin, 2; San Quintin, 1; Agua Chiquita Canyon, 1. Total, 40.

Dipodomys stephensi. California: Riverside County — San Jacinto Valley,
1; San Jacinto Lake, 1; Ethanac, 1; Perris, 1; 4 mi. s. of Perris, 25. Total, 29.

LITERATURE CITED

Grinnell, Joseph

1922 A geographical study of the kangaroo rats of California. Univ. Calif.
Publ. Zool. 24: 1-124, figs. A-X, pis. 1-7. June 17.

INDEX

Transactions of the San Diego Society of Natural History,

Volume 12

New names are in boldface type. Page numbers for principal entries also are in
boldface, those for illustrations in italic. The index was prepared by Mrs.
Ann Sprigg.

Abies macrocarpa, 351
Abra tepocana, 462
Abrams, LeRoy, 388
Abronia platyphylla, 358

umbellata platyphylla, 358
Acanthina angelica, 166, 169, 453, 454,
456, 468

lugubris, 179, 198, 298, 303

muricata tuberculata, 166

paucilirata, 198, 298

spirata, 198, 298

tuberculata, 468

tyrianthina, 468
Acanthochitona angelica, 473

arragonites, 331

avicula, 201

exquisita, 453, 456, 473
Acanthomintha ilicifolia, 377
Acerates tomentosa, 373
Achillea californica, 382

millefolium, 435
Acmaea asmi, 300

atrata, 463

dalliana, 169

digitalis, 300, 327

insessa, 201, 300, 339

limatula, 300, 327, 339

mesoleuca, 166

mesoleuca vespertina, 327

mitella fayae, 454, 463

mitra, 300

paleacea, 300, 327,

pediculus, 463

pelta, 300, 327,

pelta navicelloides, 300

persona, 300

persona strigatella, 295, 300

scabra, 300, 327

semirubida, 455, 463

scutum, 300

strongiana, 454, 456, 463

turveri, 454, 463
Acteocina angustior, 323

carinata, 471

culc.tella, 196, 297, 339

inculta, 196

infrequens, 186, 196, 471

magdalenensis, 187, 196, 323

planata, 323
Acteon punctocaelata, 196, 323

traskii, 196, 471
Actocharis, 109

Actosus nigriventris, 135, 136, 151
Adams, Joseph Edison, 388
Adenostoma brevifolium, 363

fasciculatum obtusifolium, 363

laxum, 363

sparsifolium, 363
Adolphia californica, 368
Aequipecten circularis, 459

latiauratus, 456, 459
Aesopus arestus, 324

eurytoides, 324

myrmecoon, 298

sanctus, 324
Agave-boring butterfly, 231-61, 263-75
Agave consociata, 355

deserti, 270, 355

goldmaniana, 238

orcuttiana, 238

schottii, 268, 273

sebastiana, 238

shawii, 234, 236, 237, 238, 268, 269,
355
Agianthus jacobaeus, 360
Agkistrodon contortrix, 33, 35, 39

piscivorus, 35, 39
Agropyron parishii laeve, 351
Agrostis diegoensis, 351

kennedyana, 351
Alaba catalinensis, 200

interruptilineata, 466

jeannettae, 200, 326, 466

supralirata, 466
Alabina diomede, 325, 466

io, 339

jordani, 322, 325

monicensis, 186, 199

sp., 466

tenuisculpta, 339

tenuisculpta diegensis, 199

tenuisculpta phalacra, 187, 199
Alderson, Rufus Davis, 388
Aleochara, 212
Aleocharinae, 107, 215-20

482

San Diego Society of Natural History

Aletes, 166

squamigerus, 200, 299, 326
Aligena cerritensis, 195

cokeri, 460

nucea, 460
Aligera patelliformis, 381
Allium bullardiae, 353

croceum, 354

praecox, 353

tenellum, 354
Allocarya echinacea, 376

inornata, 376
Alvania acutelirata, 200, 299

aequisculpta, 326

cosmia, 326

gallegosi, 465

lirata, 465

monserratensis, 465

oldroydae, 326

purpurea, 299, 326

sp., 465
Amaranthus palmeri, 358
Amblopusa, 109, 111, 112, 113, 114, 115,
127-129, 139, 212

borealis, 127, 128-29, 150

brevipes, 111, 125, 127, 128, 129, 150
Amblyderus, 212
Ambrosia pumila, 382
Amelanchier alnifolia cuyamacensis, 363
Amiantis callosa, 195, 296, 301
Amorpha occidentalis, 364
Amphissa undata, 298

versicolor, 198, 298
Amphithalamus inclusus, 326

tenuis, 295, 299, 326
Amsinclcia albicarpa, 376

angustata, 376

caduca, 376

curvata, 376

decumbens, 376

deltoidea, 376

diversifolia, 376

jonesii, 376

laxa, 376

littoralis, 376

palmeri, 377

spencerae, 377
Anachis coronata, 468

coronata hannana, 339, 342

hilli, 468

lineolata, 186, 198
minuta, 186

subturrita, 324
varia, 456
Anadara cepoides, 457

multicostata, 457
Anadontia edentuloides, 459
Anatina undulata, 461
Anatrechus, 213n
Angier, Belle Sumner, 388

Anomia adamas, 459

peruviana, 165, 178, 195, 338, 459
Anomalina schmitti, 188, 189
Anomalocardia subimbricata, 461

tumens, 461
Anopsisus microphthlalmus, 135
Antarctophytosus, 109, 111, 115, 139-40,
216

atriceps, 139, 140, 151

darwini, 140
Anthicus, 212
Antigona fordi, 330
Antiplanes perversa, 197
Antirrhinum coulterianum orcuttianum, 379

nuttallianum, 379

orcuttianum, 379
Antisell, Thomas, 388
Antropora tincta, 167
Aphanisma blitoides, 357
Apiastrum angustifolium, 371

angustifolium tenellum, 371

latifolium, 371
Apocynum androsaemifolium, 435
Apolymetis biangulata, 178, 195

cognata, 165, 455, 461
Aquilegia hypolasia, 359
Arabis pulchra glabrescens, 361

pulchra viridis, 361

sparsiflora californica, 361
Arbacia incisa, 454
Area bailyi, 328

gradata, 165, 178

mutabilis, 457

paciflca, 457

reeveana, 178

solida, 165, 178
Archaeology

fish remains from aboriginal sites in So-
nora, 279-86

middens in northwest Baja California, the
mollusks identified, 294
Architectonica nobilis, 466
Arcopsis solida, 454, 457
Arctostaphylos arguta diversifolia, 373

clevelandii, 372

drupacea, 372

glandulosa adamsii, 372

glandulosa australis, 372

crassifolia, 373

otayensis, 373

pringlei drupacea, 372
Arena, 109

Arene hindsiana, 455, 464
Argyrotheca lowei, 473
Arizona

Dipodomys deserti arizonae, 99-100

Thomomys bottae cedrinus, 100-101
Aromia tenuifolia, 382
Artemisia palmeri, 382

californica, 43 1

Index

483

Arthropods. See Crustacea and various

orders of insects
Asclepias californica, 373

subulata, 239
Asplenium vespertinum, 350
Assiminea compacta, 465

translucens, 299
Aster orcuttii, 382
Astraea guadalupeana, 322, 327

petrothauma, 327

inequalis, 327

lithophora, 327

undosa, 179, 327

unguis, 456
Astragalus agninus, 364

coccineus, 364

douglasii perstrictus, 364

lentiginosus borreganus, 364

leucopsis lonchus, 364

oocarpus, 364

pomonensis, 364

purshii coccineus, 364

vaseyi, 364
Atrina tuberculosa, 458
Atriplex canescens, 239

decumbens, 357

pacifka, 357

watsonii, 357
Atys casta, 471
Audibertia clevelandii, 378

palmeri, 379

vaseyi, 379
Austromalota, 139
Baccharis sorothroides, 383
Baeostethus, 109, 111, 113, 115, 137-39

chiltoni, 137-39
Baeria aristata, 383

chrysostoma gracilis crassa, 383

clevelandii, 383
Bahia rubella, 384
Baja California

Crotalus enyo furvus, 49-64

geology of northwest coast, 293, 294

insular snakes, Gulf of California, 67-97

intertidal Coleoptera, describing larvae,
207-229

marine mollusks and brachiopods of
Guadalupe Island, 319-32

marine mollusks and brachiopods of Los
Angeles Bay, 449-76

Megathymus comstocki, 231-261,
263-75

mollusks of middens, 294

names of islands, 70, 95

Nassarius howardae, 333, 334

Neotoma lepida aridicola, 287-88

new species of Dipodomys, 477-80

Perognathus fallax tnajusculus &
xerotrophicus, 413-20

Perognathus formosus infolatus, 1-2

Perognathus spinatus oribates & broc-
cus, 409-12

Pleistocene mollusks of northwest coast,
289-308

Pliocene and Pleistocene faunas of
Turtle Bay, 177-79

reptiles, 67-97

Thomomys umbrinus brazierhowelli,
407
Baker, Charles (or Carl) Fuller, 389
Balanus, 167, 294

amphritrite, 167
Balcts gibba, 455, 464

mexicana, 464

micans, 186, 198

monicensis, 186, 198

oldroydi, 186, 198

rutila, 186, 198, 298, 456, 464

sp., 464

thersites, 198, 298, 301
Balls, Edward Kent, 389
Barbarofusus barbarensis, 197
Barbatia bailyi, 296

gradata, 457

illota, 457

lurida, 457

reeveana, 457
Barclay, George, 389
Barleeia alderi, 465

californica, 326

haliotiphila, 299

oldroydi, 299, 303

subtenuis, 465
Basterotia peninsularis, 460
Belonuchus, 311
Bembidion, 212
Berberis higginsae, 360
Bergerocactus emoryi, 369
Bernardina bakeri, 329
Bestor, Norman, 389
Bidens californica, 383
Bigelovia brachylepis, 385
Bittium attenuatum, 199

mterfossum, 299, 325, 339

purpureum, 299

rugatum, 199

rugatum giganteum, 199

rugatum larum, 199
Bivetopsia bullata, 191, 193. 197, 204,

205
Bivonia compacta, 326
Bledius, 212

Bloomeria clevelandii, 354
Bolander, Henry Nicholas, 389
Bolitocharini, 107
Bolivina interjuncta, 188
Boreotrophon stuarti, 198
Bornia retifera, 191, 195
Botanical collectors in San Diego County,
388-99

484

San Diego Society of Natural History

Brachidontes multiformis, 165, 458
Brachiopods

Guadalupe Island, 322, 331
Los Angeles Bay, Baja California, 473
Brandegee, Mary Katherine Layne, 389
Brandegee, Townshend Stith, 389
Brattstrom, Bayard H.

The fossil Pit-vipers (Reptilia: Crotali-
dae) of North America, 31-46
Brickellia arguta odontolepis, 383

frutescens, 383
Brodiaea orcuttii, 354
Bromus grandis, 351
orcuttianus, 352
orcuttianus grandis, 351
Bryobiota, 109, 112, 114, 115, 129-31,
140, 212
bicolor, 129, 130-31, 150
Bryonomus, 212

Bryothinusa, 109, 111, 112, 114, 115,
131-32, 212
catalinae, 131, 132, / 50
Bryozoa

Pleistocene of Sonora, 167
Bulla gouldiana, 339, 471

punctulata, 197
Bullard, Frances E., 389
Burbeck, Anna Leora, 390
Burn, Mrs. Walter L., 388
Bursa calcipicta, 325

californica, 179, 199, 299, 301, 325
Butterfly. See Lepidoptera
Bythinopsis, 10
Cadulus fusiformis, 196, 330

sp., 462
Caecum californicum, 200, 299, 326, 339
firmatum, 466
licalum, 295, 299
sp., 466
Cafius, 212
decipiens, 1 3 1
opacus, 1 3 1
Calais parryi, 383

platycarpha, 386
Calamagrostis densa, 352

koelerioides, 352
Calamintha chandleri, 379

ilicifolia, 377
Calamus taurinus, 281, 282
Calandrinia heterophylla, 358
maritima, 358
muricata, 358
Caleschara mexicana, 167
California, Gulf of, See Gulf of California
California. See also San Diego County

intertidal Coleoptera, describing larvae,

207-229
Pleistocene mollusks of Potrero Canyon,
181-205

Calliostoma angelenum, 463
annulatum, 201
doliarium, 201,300, 303
eximium, 463
gemmulatum, 201, 300
gemmuloides, 169
gloriosum, 193, 201, 204, 205
ligatum, 300, 303
marshalli, 463
sp., 328

splendens, 201, 328
supragranosum, 201
tricolor, 201
Callistochiton crassicostatus, 201, 301
infortunatus, 473
gabbi, 473

palmulatus mirabilis, 201, 301, 303
Callitriche longipedunculata, 367

marginata longipedunculata, 367
Calochortus concolor, 354
davidsonianus, 354
dunnii, 354
weedii, 354
Calycoseris wrightii californica, 383
Calyptraea contorta, 200
Calyptridium monandrum, 358
Cameronium, 108, 111, 115, 137
flavipenne, 137
obockianum, 137
Camouflage, inherent 8C applied, in Geom-

etridae, 421-40
Cancellaria cassidiformis, 179, 470
cooperi, 297, 303
obesa, 470
Cantharus lugubris, 324

sanguinolentus, 456
Carabidae

California intertidal, describing larvae,
207-229
Cardita affinis, 459

affinis californica, 459
Cardium biangulatum, 330, 338, 342
elatum, 162, 165, 169
procerum, 165, 338, 341
quadragenarium, 338
Carex monticola, 353
praegracilis, 353
spissa, 353
Carpel imus, 212
Cassidulina californica, 188

translucens, 188
Cassis centiquadrata, 468
Castilleja oblongifolia, 379
Catorama, 212
Caulanthus heterophyllus, 361

stenocarpus, 361
Cavolina inflexa, 331
longirostris, 472
occidentalis, 331
telemus tricuspida, 185, 196

Index

485

Ceanothus austromontanus, 368

cyaneus, 368

divaricatus laetiflorus, 368

orcuttii, 368

otayensis, 368

palmeri, 368

tomentosus olivaceus, 368

verrucosus, 368
Centrifuga leeana, 191, 198
Centrostegia thurberi, 356
Centrostephanus coronarus, 455
Ceratostoma nuttalli, 298, 324
Cercis nephrophylla, 364
Cercocarpus minutiflorus, 363
Cercyon, 212
Cereus? calif ornicus, 369

emoryi, 369

engelmannii, 370

pringlei, 57

schottii, 57
Cerithidea albonodosa, 467

californica, 339

mazadanica, 467
Cerithiopsis antefilosa, 200, 299

antemunda, 299

arnoldi fossilis, 200

carpenteri, 295, 299, 339

cosmia, 200

diegensis, 299

guadalupensis, 325

kinoi, 466

oxys, 325

pedroana, 200

pedroana fatua, 200

sp., 299, 466
Cerithium maculosum, 466

sculptum, 454, 466

stercusmuscarum, 466
Chace, Emery P.

Additional notes on the Pliocene and
Pleistocene fauna of the Turtle Bay
area, Baja California, Mexico, 177-79

A new mollusk from San Felipe, Baja
California, 333-34

The marine molluscan fauna of Guada-
lupe Island, Mexico, 319-32

Pleistocene mollusks from Tecolote Creek,
San Diego, California, 335-46^
Chaenactis glabriuscula tenuifolia, 383

latifolia, 383

peirsonii, 383

tenuifolia, 383

tenuifolia orcuttiana, 383
Chaetopleura lurida, 473

mixta, 473

sp., 473
Chama buddiana, 329, 460

frondosa mexicana, 165, 460

pellucida, 195, 296, 329

sordida, 460

squamuligera, 460
Chandler, Harley Pierce, 390
Cheilanthes clevelandii, 350
Cheilea cepacea, 467
Cheteoscelis bistriaria, 435
Chilomeniscus, 70

cinctus, 71, 73

punctatissimus, 71, 73-74

savagei, 71-73, 71
Chilopsis linearis, 436

Chione californiensis, 157n, 165, 169, 178,
336,338,461

cancellata, 157

cortezi, 338, 461

fluctifraga, 169

gnidia, 165, 178, 190, 195, 336, 338, 341

picta, 190, 195,296,303,461

undatella, 195, 461

undatella simillima, 296
Chiton virgulatus, 473
Chlorissa pistaciaria, 435

subcroceata, 435
Chlorochlamys chloroleucaria, 435

zelleraria, 435
Chlorogalum parviflorum, 354
Chorizanthe discolor, 356

fimbriata, 356

fimbriata laciniata, 356

laciniata, 356

leptotheca, 356

orcuttiana, 356

procumbens, 356

procumbens albiflora, 356

thurberi, 356

uncinata, 356
Chthamalus anisopoma, 453
Cincindela, 212
Circulus rossellinus, 328
Clarkia delicata, 371
Clathurella sp., 470

trichoides, 470
Clavus aeginus, 470

empyrosia, 297

hemphilli, 197, 297

pembertoni, 470

sp., 470
Clematis lasiantha, 359

pauciflora, 359
Clements, Edith Gertrude Schwartz, 390
Clements, Frederick Edward, 390
Cleveland, Daniel, 390
Cliff, Frank S.

Snakes of the islands in the Gulf of Cali-
fornia, Mexico, 67-97
Clio pyramidata, 331
Cneoridium dumosum, 366
Codakia distinguenda, 165, 459
Codium cuneatum, 453
Coleophysis carinata, 196

harpa, 197

486

San Diego Society of Natural History

Collinsia concolor, 380
Coleoptera

California intertidal, describing larvae,
207-229
distribution in intertidal zone, 212

Hesperus, revision of U.S., 311-18

Pselaptrichus, revision, 3-28

Phytosi, review of genera and revision of
west American species, 103-51
Comarostaphylis diversifolia, 373
Comstock, John Adams

Inherent and applied camouflage in the
subfamily Geometrinae (Lepidop-
tera), including three new life history
studies, 421-40

Notes on the metamorphosis of an Agave-
boring butterfly from Baja Califor-
nia, Mexico, 263-275
Comptonia peregrina asplenifolius, 436
Condalia parryi, 368
Coniophis, 33

Conopeum commensale, 167
Conus brunneus, 456

californicus, 179, 197, 297, 301, 323

dispar, 471

fergusoni, 179

nux, 456

perplexus, 179, 456

princeps, 456

purpurascens, 179, 456

recurvus, 471

regularis, 166, 471

seal arts, 471

tornatus, 471

virgatus, 456

ximines, 471
Convolvulus aridus longilobus, 373

aridus tenuifolius, 374
Cooper, James Graham, 390
Cooperella subdiaphana, 195, 456, 461
Coralliophila hindsii, 468
Corbula fragilis, 462

luteola, 196, 297, 338, 456, 462

nasuta, 165
Cordilleras of California, 400
Cordylanthus filifolius, 380

maritimus, 380
Coreopsis californica, 383

maritima, 384
Corethrogyne brevicula, 384

filaginifolia linifolia, 384

filaginifolia pacifica, 384

incana, 384

racemosa, 384
Corvina, orangemouth, 281
Coulter, Thomas, 390
Crassatella digueti, 459
Crassinella branneri, 195, 338, 341
nuculiformis, 195, 338, 341
pacifica, 459

varians, 178
Crassispira atramentosa, 456, 470
nymphia, 454, 456, 470
pluto, 470
Crenella Columbiana, 329
divaricata, 329, 458

megas, 329
Crepidula, 166

aculeata, 467

adunca, 200, 299

arenata, 467

coei, 299

excavata, 200, 467

incurva, 467

lingulata, 327

nummaria, 200, 300

onyx, 179, 200, 300, 467

perforans, 327, 467

uncata, 467
Crepipatella lingulata, 200, 300
Crotalidae

North American fossil, 31-46

North American Pleistocene, 39

North American Pliocene, 39

oldest known, 33
Crotalus, 69

scale terminology, 49-52

adamanteus adamanteus, 35-36, 37, 45

adamanteus pleistofloridensis, 35-36,
37,39,45,46

atrox, 36, 39, 80

catalinensis, 80-82, 97

enyo, 49-64, 82

enyo cerralvensis, 82-84, 96

enyo enyo, 54-62, 84

enyo furvus, 52-63

giganteus, 36-37, 39, 45, 46

horridus, 37-38, 39

lepidus, 38, 39

mitchelli, 38, 39

mitchelli mitchelli, 84

mitchelli muertensis, 85

mitchelli pyrrhus, 84, 85

molossus estebanensis, 85

molossus molusses, 86

potterensis, 38, 39

ruber lucasensis, 80, 87

ruber ruber, 56-57, 80, 87

scutulatus, 38, 39

tortugensis, 80, 87-88

viridis, 33,38, 44
Croton californicus tenuis, 366

tenuis, 366
Crucibulum imbricatum, 179

scutellatum, 467

spinosum, 169, 179, 200, 339, 454, 467
Crustacea

Pleistocene, of Sonora, 167

Pleistocene, of Turtle Bay, 179

Index

487

Cryptantha clevelandii, 377

clevelandii florosa, 377

costata, 377

ganderi, 377

micrantha lepida, 377
Cryptochiton stelleri, 301, 303
Cryptomya californica, 196, 297, 338. 456,

462
Ctena mexicana, 459
Cucurbita palmata, 382
Cumingia californica, 297

lamellosa, 462
Cupressus forbesii, 350

stephensonii, 351
Curran, Mary Katharine, 390
Cuspidaria pectinata, 329
Cuvierina columella, 331
Cyclopecten pernomus, 459
Cyclostremiscus panamensis, 455, 465

sp., 465
Cylichna attonsa, 197, 323
Cylichnella attonsa, 339
Cymatium gibbosum, 468
Cynoscion macdonaldi, 281, 282, 286

xanthulus, 281, 282, 286
Cypraea annettae, 166, 179, 454, 467

arabicula, 179, 456

spadicea, 325
Cypraeolina pyriformis, 297, 324, 456. 470
Cystiscus jewettii, 297, 324

minor, 324

politulus, 324, 470

regularis, 197, 297

subtrigona, 197, 297
Cythodonta lucasana, 462
Danthonia californica unispicata, 352
Daphnella bartschi, 470
Deane, George Clement, 391
Decipifus gracilis, 468

lyrta, 468
Delphinium coccineum, 359

collinum, 359

cuyamacae, 359

parryi subglobosum, 359

subglobosum, 359
Dendraster ashleyi, 178

gibbsi humilis, 178
Dendromecon saligna, 360
Dentalium berryi, 330

neohexagonum, 196, 297

quadrangulare, 462

sp., 462

splendidulum, 330
Deschampsia gracilis, 352
Deweya arguta, 372
Diacria trispinosa, 193, 196, 204, 205
Diala sp., 465
Diastoma slevini, 322, 325
Diaulota, 109, 111, 112, 113, 115, 119-
26, 139,212,214,216,217-20

key to larvae, 217

brevipes, 125

densissima, 114, 119, 120-21, 122, 124,
129,24*
larva, 217-218, 219, 227

fulviventris, 120, 121-22, 124, 148-49,
214
larva, 217, 218-19, 227

harteri, 120, 123-124, 149
larva, 219-20, 228

insolata, 120, 121

megacephala, 120, 124-25, 149
larva, 217, 219-20, 228

vandykei, 120, 125-26, 129, 149
larva, 217, 219, 228
Dichondra occidental is, 374
Dichorda illustraria, 426-30

iridaria, 435
Diglotta, 115, 212
Diodora alta, 454, 463

aspera, 201, 301

constantiae, 463

digueti, 454, 463

inaequalis, 166, 339, 454, 463
Diplacus aridus, 380

puniceus, 380
Diplodonta inezensis, 460

orbella, 165, 178, 195, 296, 460

sericata, 338, 342, 460

subquadrata, 330, 460
Dipodomys agilis simulans, 479

antiquarius, 477-80

cascus, 479-80

deserti arizonae, 99-100

deserti deserti, 99, 100

deserti sonoriensis, 100

gravipes, 477-80

stephensi, 477-80
Discinisca cumingii, 473

strigata, 473
Dissodactylus sp., 455
Divalinga eburnea, 459
Dodecatheon clevelandii, 373
Donax californica, 338

gouldii, 196, 297, 301

gracilis, 165
Dosinia ponderosa, 165, 190, 195, 461
Downingia concolor brevior, 382

pulchella arcana, 382
Drake, Robert J.

Type material of Eucalodium orcutti Dall
(Gastropoda: Pulmonata) f rom Oax-
aca, Mexico, 309-10
Dudleya abramsii, 361

aloides, 361

delicata, 361

lagunensis, 361

lanceolata, 362

pulverulenta, 362

488

San Diego Society of Natural History

Dunn, George Washington, 391
Dyschirius, 212
Dysodia porophylloides, 384
Eastwood, Alice, 391
Echeveria lagunensis, 361
lanceolata, 362
pulverulenta, 362
Echinocactus limitus, 370
Echinocactus? viridescens, 370
Echinocactus viridescens cylindraceus, 370
Echinocereus engelmannii, 370
Echinoderms

Pleistocene, of Sonora, 166
Pliocene, of Turtle Bay, 178
Eggleston, Willard Webster, 391
Elephantanellum heptagonum, 466
Elephantulum mirifucum, 466

sp., 466
Elymus orcuttianus, 352
Emerson, William K.

Marine Pleistocene invertebrates from near
Puerto Penasco, Sonora, Mexico,
153-75
Emerson, William K., and Emery P. Chace
Pleistocene mollusks from Tecolote Creek,
San Diego, California, 335-46
Emory, William Hemsley, 391
Emphyastes, 212
Emplenota, 212
Encelia farinosa, 384

viscida, 384
Encope, 166

grandis, 455
Endeodes, 212, 214, 220-222
collaris

larva, 220-221, 229
insularis
larva, 222
pupa, 222, 229
rugiceps, 221-22
Engina strongi, 193, 197, 204, 205
Ensis calif ornicus, 196
Epantius, 212
Ephedra californica, 351
Epilobium palmeri, 371
Epilucina californica, 195, 296
Epinephelus analogus, 280
Epitonium apiculatum, 324, 464
bellastriatum, 198, 324
californicum, 324
clarki, 186, 198
columbianum, 324
cooperi, 198
hexagonum, 464
indianorum, 298
insculptum, 198
oerstedianum, 464
sawinae, 191, 198,324
tinctum, 198, 298
Epling, Carl Clawson, 391

Eragrostis orcuttiana, 352

Erato columbella, 199, 299, 325, 467

vitellina, 299
Eriastrum densifolium austromontanum, 374
Eriodictyon californicum australis lanatum,
375

crassifolium, 375

trichocalyx lanatum, 375
Eriogonum cernuum viscosum, 356

cordatum, 357

fasciculatum, 435, 436

fasciculatum foliolosum, 424

fasciculatum maritimum, 356

fasciculatum oleifolium, 356

trichopes, 357

trichopes cordatum, 357

thurberi, 357

verticillatum, 357

viminium molestum glabrum, 357
Eriophyllum wallacei rubellum, 384
Eriphia squamata, 167, 170
Eritrichium micranthum lepidum, 377
Eryngium parishii, 371
Eschscholtzia clevelandii, 360

floribunda, 360

physodes, 360

sanctarum, 360
Eucalodium orcutti, 309, 310
Eueana niveociliaria, 435
Eulima californica, 325

lapazana, 464
Eulobus californicus, 371
Eupatorium perfoliatum, 435
Euphorbia cinerascens appendiculata, 366

hirtula, 366

misera, 239, 367

palmeri, 367

patellifera, 367
Euphytosus schenklingi, 135
Eupleura muriciformis, 339, 341, 468

nitida, 456, 468

triquetra, 179
Euryptera lucida, 372
Euspira lewisii, 200, 300, 301
Eutamias merriami meridionalis, 477
Eutoca speciosa, 375
Everett, Percy Charles, 391
Fartulum hemphilli, 200, 299, 326

occidentale, 200, 299

orcutti, 299
Fenzlia concinna, 374
Ferocactus fordii, 239

viridescens, 370
Ficus ventricosa, 468
Fish remains from aboriginal sites in Sonora,

279-86
Fisher, William J., 391
Fissurella rugosa, 463

volcano, 294, 301,328

Index

489

Follett, W. I.

Fish remains from aboriginal sites in the
Punta Peiiasco region of Sonora,
Mexico, 279-286
Forbes, Charles Noyes, 392
Forreria belcheri, 198
Fosberg, Francis Raymond, 392
Fossarus sp., 467
Franseria chenopodiifolia, 239

palmeri, 384

pumila, 382
Frasera parryi, 373
Fremontia californica diegensis, 369
Froebel, Julius, 392
Fultz, Francis Marion, 392
Fusinus ambustus, 469

cinereus, 454, 469

dupetitthouarsi, 469

kobelti, 324

luteopictus, 197, 298
Galeomma mexicanum, 460
Galium angustifolium, 381

angustifolium siccatum, 381

nuttallii, 381

siccatum, 381

suffruticosum, 381
Gambel, William, 392
Gander, Frank Forest, 392
Gari regularis, 462
Garrya flavescens palmeri, 372

veatchii palmeri, 372
Gastrochaena ovata, 462
Gastropoda

Eucalodium orcutti from Oaxaca. 309,
310
Gayophytum lasiospermum, 371
Gentry, Howard Scott, 392
Geology

northwest Baja California, 293-94

Potrero Canyon, California, 185, 188

Puerto Peiiasco area, Sonora, 158-64
Geometridae

camouflage; life history studies, 421-40
Geraea viscida, 384
Gilia caruifolia, 374

densifolia austromontana, 374

floribunda, 374

inconspicua diegensis, 374

lutea longistylis, 374

truncata, 374
Gird, Henry Harrison, 392
Girella simplicidens, 282, 283
Githopsis filicaulis, 382

specularioides Candida, 382
Glans afnnis californica, 165

carpenteri, 329

subquadrata, 296
Glycymeris bicolor, 458

gigantea, 162, 458

guadalupensis, 322, 328

maculata, 165, 458

multicostata, 165, 458

profunda, 296, 303

subobsoleta, 296, 338
Gnaphalium bicolor, 384

microcephalum, 384
Godetia delicata, 371

epilobioides, 371
Gonolobus californicus, 373
Gopherus, 441-48

agassizi, 441-48

berlandieri, 441-48

polyphemus, 441-42
Gouldia californica, 461
Grant, Chapman

Differentiation of the southwestern tor-
toises (genus Gopherus), with notes
on their habits, 441-48
Grant, George Bernard, 393
Greene, Edward Lee, 393
Gregariella chenui, 458
Griffiths, David, 393
Grindelia hallii, 385
Grippina californica, 330
Grouper, Gulf, 280, 283
Guadalupe Island

marine mollusks and brachiopods, 319-32

Pleistocene fossils, 321
Gulf of California

insular snakes, 67-97

marine mollusks of Los Angeles Bay, 449-
76

names of islands, 70, 95

north Gulf fauna, relationships, 167-71
Gymnogramme triangularis viscosa, 350
Habenaria cooperi, 355
Hadrotes, 212
Haliotis sp., 301

californiensis, 322, 328

corrugata, 328

cracherodii, 294, 300, 328

fulgens, 328

rufescens, 301
Halistylus pupoideus, 300
Hall, Harvey Monroe, 393
Haminoea angelensis, 323, 471

strongi, 471
Hanetia elegans, 187, 191, 193, 197, 204,
205

macrospira, 469
Haplopappus junceus, 385

palmeri, 239

propinquus, 385

squarrosus, 239

venetus decumbens, 385

venetus oxyphyllus, 385
Harbison, Charles F.

A new species of Megathymus from Baja
California, Mexico, 231-261

490

San Diego Society of Natural History

Harfordia monksae, 197, 298

Harpa crenata, 470

Hasseanthus blochmaniae brevifolius, 362

Hayes, Sutton, 393

Helagras, 33

Helenium autumnale, 435

Heliacus bicanaliculatus, 466

mazatlanicus, 455, 466
Helianthemum aldersonii, 369

scoparium aldersonii, 369
Helianthus gracilentus, 385
Heliaster kubiniji, 454
Hemitonia hermosa, 463
Hemizonia conjugens, 385
fasciculata ramosissima, 385
floribunda, 385
ramosissima, 385
tenella, 385
Henshaw, Henry Wetherbee, 393
Here excavata, 195, 296
Herdein, Leo George, and William K.
Emerson
Marine Pleistocene invertebrates from near
Puerto Pefiasco, Sonora, Mexico,
153-175
Hesperus

revision of U. S., 311-18
apicialis, 313,314, 315, 316, 318
arizonicus, 3 12, 3 16, 318
baltimorensis, 314, 316, 318
stehri, 312,313,316, 318
Heterodonax bimaculatus, 462
Heuchera brevistaminea, 362
Hexaplex brassica, 468

erythrostomus, 468
Hiatella arctica, 196, 297, 462
Higgins, Ethel Bailey

Type localities of vascular plants in San
Diego County, California, 347-406
Hinds, Richard Brinsley, 393
Hinnites giganteus, 296, 329
Hipponix antiquatus, 179, 200, 299, 327
serratus, 467
tumens, 200, 299, 327
Hippopodinella adpressa, 167
Hitchcock, Charles Leo, 393
Hitchcock, George N., 394
Holocarpha virgata elongata, 385
Homalopoma carpenteri, 201, 300, 327
luridum, 327
paucicostatum, 327
Hookera multipedunculata, 354

orcuttii, 354
Horkelia truncata, 363
Hormomya adamsiana, 165, 454, 458
Hosackia argophylla, 364
micrantha, 365
prostrata, 365
Howell, John Thomas, 394

Huey, Laurence M.

A new race of Dipodomys and a new
race of Thomomys from Arizona,
99-101
A new form of Perognathus formosus

from Baja California, Mexico, 1-2
A new race of pocket gopher (Thomo-
mys) from San Fernando Mission,
Baja California, Mexico, 407-8
A new race of wood rat (Neotoma) from
the Gulf side of central Baja Cali-
fornia, Mexico, 287-88
Comments on the pocket mouse, Perog-
nathus fallax, with descriptions of
two new races from Baja California,
Mexico, 413-20
Two new races of Perognathus spinatus
from Baja California, Mexico, 409-
12
Two new species of broad-faced, five-toed
kangaroo rats (genus Dipodomys),
477-80
Hulsea californica, 385
Hutchings, James Mason, 394
Hyalina californica, 197, 297, 324
Hymenoclea monogyra, 239
Hymenopappus wrightii viscidulus, 385
Hypsiglena, 69

torquata gularis, 74
torquata ochrorhyncha, 74
torquata tortugaensis, 74
torquata venusta, 74-75
Insects. See the various orders
Irus lamellifer, 296, 303
Ischnochiton acrior, 169, 191, 201
biarcuatus, 331
conspicuus, 201
mertensi, 331
pectinulatus, 201
sanctaemonicae, 201
tridentatus, 473
Iselica fenestrata, 326, 472

ovoidea, 472
Isocoma decumbens, 385
leucanthemifolia, 386
oxyphylla, 385
Isoetes orcuttii, 350
Isognomon chemnitzianus, 165, 454, 458

janus, 458
Isomeris arborea, 360
Iva hayesiana, 386
Janthina globosa, 325
Jaton festivus, 198, 298
Jenneria pustulata, 467
Jensen, Herbert A., 394
Jepson, Willis Linn, 394
Jepsonia parryi, 362
Johnson, Ernest Ralph, 394
Jones, Marcus Eugene, 394
Juglans nigra, 436

Index

491

Kangaroo rat

Arizona desert, 99

Bonsall relict, 479

San Borjas, 477
Keck, David Daniels, 394
Kelletia kelletu, 197, 298
Kellia laperousii, 195, 296, 330

suborbicularis, 460
Kessler, Robert, 394
Kimball, Laura Frances, 395
Knefastia funiculata, 470

walkeri, 470
Koeleria nitida californica transiens, 352

nitida californica vestita, 352

pseudocristata californica, 352
Krameria parvifolia imparata, 365
Kurtzia arteaga roperi, 197

beta, 323
Lacuna carinata, 200, 299, 303

carinata aurantiaca, 200

unifasciata, 200, 299
Laevicardium clarionense, 460

elatum, 195, 460

elenense, 460

elenense apicinum, 460
Lamellaria stearnsi orbiculata, 327
Lampropeltis, 70

catalinensis, 75

getulus, 37

nitida, 75
Lantana camara, 436
Larkin's (Larken's) Station, 400
Larvae

Geometrinae, 424-39

intertidal Coleoptera, 207-229

Megathymus comstocki, 267-75
Lasaea cistula, 296

subviridus, 295, 296
Lathyrus splendens, 365

strictus, 365
Leiomya scaber, 329
Lemmon, John Gill, 395
Lepechinia ganderi, 378
Lepidium acutidens microcarpum, 361

flavum felipense, 361

robinsonii, 361
Lepidochitona keepiana, 339, 456, 473

sp., 330
Lepidopleurus cancellatus, 472

rugatus, 330
Lepidoptera

Geometridae: camouflage; life history
studies, 421-40

Megathymus comstocki, 231-261
metamorphosis, 263-75
Lepidozona clathrata, 473

pectinulatus, 301

subtilis, 473
Leptochiton cancellatus, 456

clarki, 201

Leptochloa imbricata, 352
Leptopecten latiauratus, 194, 296, 301

latiauratus monotimeris, 194, 296

sp., 296
Leptosyne californica, 383
Leptotyphlops, 69
Lichanura roseofusca gracia, 70
Lilium fairchildii, 354
Lima hemphilli, 195, 459

pacifica, 459

subauriculata, 329

tetrica, 459
Limnanthes versicolor parishii, 367
Limonium mexicanum, 373
Linanthus androsaceus luteolus, 374

luteolus, 374
Liotia acuticostata, 328

californica, 328

fenestrata, 328

heimi, 328
Liparocephali, 109

Liparocephalus, 109, 111, 112, 113, 115,
116-19, 120,139,212,216-17
key to North American species, 117

brevipennis, 114, 116, 117-18, 119, 148,
216

cordicollis, 117, 118-19, 129, 148, 214,
216
Lithophaga, 165

attenuata rogersi, 458

aristata, 458

plumula, 195, 329

spatiosa, 458
Lithophragma trifida, 326
Littorina dubiosa, 464

planaxis, 200, 299, 301,326

scutulata, 200, 294, 299, 301, 339
Loeflingia squarrosa, 358
Lonicera subspicata denudata, 381
Lophocereus schottii, 239
Los Angeles Bay

marine mollusks, 449-76
Lottia gigantea, 179, 294, 300, 301, 327
Lotus confinis, 365

spencerae, 365
Lowe, Charles H., and Kenneth S. Norris

Analysis of the herpetofauna of Baja
California, Mexico. I. Introduction,
49. II. Analysis of subspecific differ-
entiation in Crotalus enyo, the Baja
California Rattlesnake, 49-64
Lucapinella callomarginata, 301, 339
Lucina approximata, 330, 338, 459

californica, 330

cancellaris, 459

lampra, 165, 459

nuttalli, 178, 338
Lucinisca nuttalli, 195, 296
Lupinus albifrons medius, 365

brittonii, 365

492

San Diego Society of Natural History

chamissonis longifolius, 365

densiflorus austrocollium, 365

micranthus microphyllus, 365

sparsiflorus inopinatus, 365

truncatus, 365
Lycium californicum, 239, 379
Lyonsia californica nesiotes, 329

gouldii, 462

lucasana, 462
Lyria cumingii, 470
Lyropecten subnodosus, 455, 459
Macbride, James Francis, 395
Machaeranthera hiemalis, 386

lagunensis, 386
Machaerocereus gummosus, 239
Macoma acolasta, 338, 341

indentata, 195,456,461

nasuta, 196, 296, 301

pads, 338, 341

secta, 196,297,301

yoldiformis, 196, 338
Macromaphalina occidentalis, 327

sp., 465
Macron aethiops, 179

lividus, 298
Mactra californica, 196, 338

dolabriformis, 461

fonsecana, 455, 461
Madraglossa carnosa, 386
Malachiidae

California intertidal, describing larvae,
207-229
Malacothrix clevelandii, 386
Malva fasciculata, 369
Malvastrum densiflorum viscidum, 369

fasciculatum, 369

viscidum, 369
Mammals

Dipodomys deserti arizonae, 99-100

Dipodomys antiquarius & cascus,
477-80

Neotoma lepida aridicola, 287-88

Perognathus fallax majusculus & xero-
trophicus, 413-20

Perognathus formosus infolatus from
Baja California, 1-2

Perognathus spinatus oribates & broc-
cus, 409-12

Thomomys bottae cedrinus, 100-101

Thomomys umbrinus brazierhowelli,
407
Mammillaria brandegeei, 239

incerta, 370

tetrancistra, 370
Mangelia barbarensis, 323, 339, 456, 470

cetolaca, 197

interlirata, 297, 323

phryne, 456, 470

sp., 470

subdiaphana, 470

variegata, 197, 297
Margarites acuticostata, 328

parcipicta, 328
Marginella californica, 169, 454, 470

jewetii, 470
Marsh, Gordon A., and Robert O. Schuster

A preliminary revision of the genus Psel-
aptrichus (Coleoptera: Pselaphidae),
3-28
Masticophis aurigulus, 76

barbouri, 75

bilineatus, 76-77

bilineatus lineolatus, 76

flagellum piceus, 77-78, 79

lateralis, 76
Maxwellia gemma, 198, 298, 324

santarosana, 298
McLean, James H.

Marine mollusks from Los Angeles Bay,
Gulf of California, 449-76
Mearns, Edgar Alexander, 395
Meconella kakoethes, 360
Megapitaria squalida, 165, 178, 461
Megasurcula carpenteriana, 186, 197, 297

carpenteriana tryoniana, 197, 297
Megatebennus bimaculatus, 201, 301
Megathura crenulata, 328
Megathymus comstocki 231-58, 259-61,
263-75
metamorphosis, 267-275
parasites, 273

evansi, 267, 268

mariae, 254-55

polingi, 255-56, 268, 269, 273

stephensi, 235, 253-54, 267-70
larva, 272

yuccae-martini, 269

yucca-navajo, 235
Melampus olivaceus, 339, 456, 472
Melongena patula, 469
Merochlora fasiolaria, 431-435
Mesothea viridipennata, 436
Metaxia diadema, 200, 325, 456, 466
Mexico. See also Baja California, Sonora

Eucalodium orcutti from Oaxaca, 309,
310
Micranellum crebricinctum, 200, 339
Micrarionta levis, 177
Microseris breviseta, 386

heterocarpa, 386

parishii, 386

platycarpha, 386
Micruroides euryxanthus, 80
Middens in northwest Baja California, the

mollusks identified, 294
Milneria kelseyi, 296, 329

minima, 329
Milquatay, 400
Mimulus aridus, 380

clevelandii, 380

Index

493

diffusus, 380

grantianus, 380

inconspicuus latidens, 380

latidens, 380

paniculatus, 380

puniceus, 380
Mirabilis californica, 358

froebelii, 358
Mitra catalinae, 297, 324

dolorosa, 469

erythrogramma, 456, 469

fultoni, 197, 324

idae, 197, 298, 303

mexicana, 469

solitaria, 454, 469
Mitrella carinata, 198, 298, 324

carinata gausapata, 198, 298

delicata, 456, 468

densilineata, 469

dorm a, 469

gouldi, 198

grand, 469

ocellata, 454, 469

tuberosa, 198
Mitromorpha aspera, 297, 303

crassaspera, 323

filosa, 297, 323

interfossa, 191, 197
Modiolus capax, 165, 169, 458

fornicatus, 195, 296

pallidulus, 329
Modulus cerodes, 466

disculus, 466
Mojarra garabata, 281
Mojarron chino, 281
Mollusks

Eucalodium orcutti from Oaxaca, 309,
310

Guadalupe Island marine, 319-32

Los Angeles Bay, Baja California, marine,
449-76

of middens in northwest Baja California,
294

Pleistocene of northwest Baja California,
289-308
Recent ranges, 302-3

Pleistocene of Potrero Canyon, California,
181-205
faunal affinities, 192-93
north-ranging Recent species, 191
south-ranging Recent species, 190-91

Pleistocene of Sonora, 153-75

Pleistocene of Tecolote Creek, San Diego,
335-46

Pliocene and Pleistocene of Turde Bay,
178-79
Moniliopsis incisa, 197

incisa fancherae, 197

incisa ophioderma, 197

Monardella lanata, 378

lanceolata microcephala, 378

linoides, 378

linoides viminea, 378

macrantha, 378

macrantha hallii, 378

nana, 378

nana leptosiphon, 378

sanguinea, 378

villosa leptosiphon, 378

viminea, 378
Moore, Ian

A revision of the Pacific coast Phytosi,
with a review of the foreign genera
(Coleoptera: Staphylinidae) 103-52

Notes on some intertidal Coleoptera, with
descriptions of the early stages (Cara-
bidae Staphylinidae, Malachiidae) ,
207-29

The North American species of Hesperus
Fauvel, with descriptions of two new
species (Coleoptera: Staphylinidae),
311-18
Mopalia acuta, 201

ciliata, 301

hindsi, 301

muscosa, 301

sp., 301
Moran, Reid Venable, 395
Morula ferruginosa, 454, 468

lugubris, 339, 342
Motschulskium, 212

sinuaticolle, 13 1
Mugil cephalus, 281, 282, 283, 286
Mulinia coloradoensis, 461

pallida modesta, 190, 196
Mullet, striped, 281
Munz, Phillip Alexander, 395
Murex elenensis, 468

erythrostomus, 166

radix nigritus, 166

recurvirostris, 468
Muricanthus nigritus, 468

princeps, 468
Muricopsis armatus, 468
Mustelus lunulatus, 280, 283
Mycteroperca jordani, 280, 283
Myosurus minimus apus, 359
Myrtillocactus cochal, 239
Mysella aleutica, 195

compressa, 460
Mytilus californianus, 294, 296, 301
Nassa cerritensis, 191, 197

delosi, 197, 298, 303

fossata, 197, 294, 298

mendica, 198, 298

mendica cooperi, 198, 298

perpinguis, 198, 298, 301

tegula, 198

494

San Diego Society of Natural History

Nassarina pammicra, 456, 469

sp., 469
Nassarius angulicostis, 469

howardae, 333-534

insculptus, 324

iodes, 469

leucops, 166

moestus, 455, 469

perpinguis, 334

rhinetes, 334

tegula, 179, 339
Nassarius tiarula, 455, 469

versicolor, 469
Natica broderipiana, 467

chemnitzii, 467

elenae, 467
Navarretia densifolia jacumbana, 374

foliacea, 374

hamata foliacea, 374

macrantha, 375
Nelson, Aven, 395
Nemacaulis denudata, 357

foliosa, 357
Nemacladus glanduliferus, 382

ramosissimus, 382
Nemocardium pazianum, 460
Nemophila racemosa, 375

rotata, 375
Nemoria bistriaria, 436

brunnearia, 436

darwiniata, 436

delicataria, 422, 424-26

mimosaria, 436

pulcherrima, 436

punctularia, 436

rubrifrontaria, 436
Neosimnia quaylei, 467
Neotoma lepida arenacea, 288

lepida aridicola, 287-88

lepida egressa, 288

lepida felipensis, 287, 288

lepida gilva, 288

lepida intermedia, 288

lepida molagrandis, 287, 288

lepida notia, 288

lepida pretiosa, 288

lepida ravida, 287, 288
Nerita funiculata, 464

scabricosta, 166, 464
Neritina luteofasciata, 454
Neurodromicus, 33
Neverita reclusiana, 200

reclusiana alta, 200, 300

reclusiana imperforata, 200

sp., 300
Newberry, John Strong, 395
Nicotiana clevelandii, 379
Niso hipolitensis, 464
Nodulus kelseyi, 295, 299, 326
Nolina interrata, 354

Nomaeopelta dalliana, 454, 456, 463

mesoleuca, 463

stanfordiana, 463
Norris, Kenneth S. See Lowe, Charles H.,

and Kenneth S. Norris
Norrisia norrisii, 201, 300, 328
Notholaena californica, 350

Candida accessita, 350

newberryi, 350
Notocytharella niobe, 470
Nucella emarginata ostrina, 198, 298, 301
Nucula declivis, 457

exigua, 338, 341

linki, 457

suprastriata, 194
Nuculana taphria, 194, 296, 301
Nuttall, Thomas, 395
Nuttallina californica, 301, 330

crossota, 473
Obione microcarpa, 357

tetraptera, 357
Ocenebra barbarensis, 191, 198, 332, 339

circumtexta, 198

crispatissima, 332

foveolata, 198, 298, 339

gracillima, 298, 324

interfossa, 198, 298

interfossa beta, 298, 303

parva, 468

poulsoni, 198,298,301,324

seftoni, 320, 324, 331-32
Ochthebius, 212
Ocinebra. See Ocenebra
Octopus bimaculatus, 454
Odostomia aepynota, 325

amilda, 325

audax, 472

California, 199

callipyrga, 325

clementina, 325

deceptrix, 325

domlla, 199, 298

eucosmia, 325

eugena, 199

farallonensis, 187, 199

fetella, 339

gravida, 187, 199

helena, 199

herrerae, 472

io, 298

minutissima, 199

navisa, 325

nemo, 199

phanea, 298, 303

phanella, 199, 298

pulcia, 325

sp., 299, 472

trachis, 295, 299

turricula, 325

virginalis, 295, 299, 325

Index

495

viscainoana, 471
Oenothera bistorta, 371

epilobioides, 371

leptocarpa, 371

spiralis linearis, 371

trichocalyx cineracea, 371
Oliva davisae, 179

incrassata, 166, 179

spicata, 166, 469
Olivella baetica, 197, 297, 339

biplicata, 179, 197, 294, 297, 301

dama, 166, 469

fletcherae, 469

pedroana, 197, 297
Onchidella binneyi, 453, 454, 472
Opaleye, Gulf, 282
Opalia chacei, 298, 303

crenimarginata, 464

diadema, 455, 464

insculpta, 298
Ophiodermella incisa fancherae, 297

ophioderma, 297
Ophioglossum californicum, 350
Opuntia, 239

acanthocarpa ganderi, 370

bigelovii hofrmannii, 370

demissa, 370

echinocarpa parkeri, 370

fosbergii, 370

parryi, 370

prolifera, 370

serpentina, 371
Orcutt, Charles Russell, 309-10, 396
Orthocarpus falcatus, 380
Osmadenia tenella, 385
Ostrea angelica, 165, 338, 341, 458

columbiensis, 458

conchaphila, 165, 454, 458

cumingiana, 178

fisheri, 458

lurida, 194, 296, 338

megodon, 178, 296, 303

palmula, 165,454,458

vespertina, 178
Oxalis californica, 366
Oxybaphus froebelii, 358
Pachygrapsus crassipes, 169
Padina durvillaei, 453, 455
Paleoecology

Pleistocene, northwest Baja California,
301-2

Pleistocene, northwest Sonora, 170

Pleistocene, Potrero Canyon, California,
189-92
Palmer, Edward, 396
Pandora punctata, 195
Panope generosa, 196
Papyridea aspersa, 460
Paralabrax maculatofasciatus, 280
Parametaria dupontii, 324, 456

Paraphytosus, 111, 135, 139
Parish, William Fletcher, 396
Parish, Samuel Bonsall, 396
Parker, J. C, 396
Parry, Charles Christopher, 396
Parvilucina tenuisculpta, 195
Payson, Edwin Blake, 397
Pecten bellus hemphillii, 178

callidus, 178

cerrosensis, 178

circularis, 165, 178, 338

circularis circularis, 341

cristobalensis, 178

diegensis, 296, 329

hakei, 178

latiauritus, 338

lowei, 329

pernomus, 329

vogdesi, 165, 459
Pectocarya setosa aptera, 377
Pectunculus giganteus, 162
Pedipes liratus, 339

unisulcatus, 339
Penstemon bridgesii X heterophyllus, 381

spectabilis, 381

ternatus, 381
Pentachaeta aurea, 386
Periglypta multicostata, 461
Periploma planiusculum, 195
Perityle emoryi, 386
Perognathus alticolus, 477

fallax fallax, 417

fallax inopinus, 419

fallax majusculus, 418-19

fallax pallidus, 417-18

fallax xerotrophicus, 419

formosus cinerascens, 2

formosus formosus, 2

formosus infolatus, 1-2

formosus mesembrinus, 2

formosus mohavensis, 2

spinatus broccus, 410-12

spinatus oribates, 409-12

spinatus peninsulae, 410-12

spinatus prietae, 409-12

spinatus rufescens, 409-12

spinatus spinatus, 411-12
Petaloconchus complicates, 299
Petricola californiensis, 296

carditoides, 296

parallela, 338, 341

robusta, 461
Peucedanum pringlei, 372
Phaca deanei, 366

perstricta, 364

pseudoocarpa, 366
Phacelia aldersonii, 375

californica patula, 375

eremica, 375

leucantha, 375

496

San Diego Society of Natural History

magellanica patula, 375
parryi, 375
polystachya, 375
whidavia jonesii gracillima, 375
Phalaris intermedia angustata, 352
Phaleria, 212
Phenacolepas malonei, 464

osculans, 464
Philbertia crystallina, 323
Philbrook, Myrtle, 397
Philobrya setosa, 329
Philonthus, 3 1 1
apicalis, 315
apicialis, 315
baltimorensis, 3 14
haematurus, 315
Pholad, 178
Pholadidea penita, 297
Pholisma arenarium, 372
Pholistoma racemosum, 375
Photinia arbutifolia, 424
Phycocoetes, 212
testaceus, 131
Phyconumus, 212

Phyllonotus radix nigritus, 339, 341
Phyllorhynchus arenicola, 78
Phytosi

characters, 109
ecology, 110

generic characters, 110-15
key to genera, 115
key to larvae, 216

review of genera and revision of west
American species, 103-51
Phytosus, 108, 109, 112, 114, 115, 135-36,
216
(Actosus), 112, 115
(Anopsisus), 115
balticus, 136, 151
bicolor, 130
dimidiatus, 136
fenyesi, 136
littoralis, 136
maritimus, 134
nigriventris, 115, 136, 216
opacus, 108n

spinifer, 115, 135, U6, 151
Pickeringia montana tomentosa, 366
Pinna rugosa, 458
Pinus cembroides parryana, 351
parryana, 351
torreyana, 351
Piperia cooperi, 355

Ieptopetala, 355
Pitar newcombianus, 461
Pitavia dumosa, 366
Pituophis, 70

Pit-vipers, North American fossil, 31-46
Pityrogramma triangularis viscosa, 350
Placiphorella blainvillei, 456, 473

Placunanomia cumingii, 459

Placusa, 109n

Plagiobothrys allocaryoides, 377

calif ornicus gracillis, 377
Plantago obversa, 381
Platidia anomioides radiata, 331
Platyodon cancellata, 196
Platystemon californicus nutans, 360
obtectus, 360

verecundus glabrifructifer, 360
Pleistocene

fauna of Turtle Bay, 177-79
fossils on Guadalupe Island, 321
marine climates in northwest Baja Califor-
nia, 302
marine climate in Sonora, 171
marine invertebrates from Sonora, 153-75
mollusks of northwest Baja California,

289-308
mollusks of Potrero Canyon, California,

181-205
mollusks of Tecolote Creek, San Diego,
335-46
Pleuroliria picta, 471
Plicatula spondylopsis, 459
Pliocene

fauna of Turtle Bay, 177-79
Poa capillaris, 352
orcuttiana, 352
Pocket gopher

Chemehuevis Mountain, 100-101
San Fernando, 407-08
Pocket mouse, San Fernando spiny, 409-12
San Francisquito, 1
San Ignacio spiny, 410-12
short-eared, 413-20
Pododesmus cepio, 459
macroschisma, 195, 296
pernoides, 455, 459
Pogogyne abramsii, 378

nudiuscula, 378
Polinices bifasciatus, 467
draconis, 187, 200
reclusianus, 166, 169, 179, 456, 467
uber, 166, 467
Polycarpon depressum, 358
Polypea, 109, 132
Pomaulax undosus, 201, 300
Pontamalota, 212

opaca, 108n
Porgy, Pacific, 281
Porites californica, 455
Porophyllum caesium, 386

vaseyi, 387
Potamogeton pauciflorus californicus, 351
Potentilla bolanderi clevelandii, 363
clevelandii, 363
parishii, 363
truncata, 363

Index

497

Potrero Canyon, Pleistocene mollusks, 181-

205
Pringle, Cyrus Guernsey, 397
Pritchardia filifera, 353
Protocardia centifilosa, 330
Protothaca asp>errima, 461

grata, 165, 178, 461

staminea, 294, 296, 338

tenerrima, 195
Prunus fremontii, 363

fremontii pilulata, 363
Psammobia edentula, 196
Pselaphidae

Pselaptrichus, revision, 3-28
Pselaptrichus, 3-28

carinatus, 16-18, 26, 27, 28

cavatus, 21-23, 24, 27, 28

curiosus, 8, 16, 25

gibbosus, 20, 21, 23, 24, 27, 28

oculatus, 18-20, 26, 27, 28

ornatus, 20-21, 24, 26, 27, 28

rectus, 13-15, 26

rothi, 10, 16, 19,20,21,24

spinosus, 15-16, 26, 27, 28

tuberculipalpus, 6, 10, 12-13, 14, 15, 16,
18, 26, 28
Psephidea cymata, 330

salmonea, 330
Pseudochama corrugata, 455, 460

exogyra, 178, 195, 296

panamensis, 455, 460

saavedrai, 460
Pseudomelatoma penicillata semiinflata, 197,

295, 297
Pseudorotella bibbiana, 300

invallata, 300

supravallata, 300
Pseudotsuga macrocarpa, 351
Psoralea rigida, 366
Pteria sterna, 458
Pteropurpura trialatus, 198
Pterynotus erinaceoides, 468

petri, 198
Ptilomeris anthemoides, 387

aristata, 383

coronaria, 387

mutica, 387
Puerto Penasco, Pleistocene marine inverte-
brates, 153-75
Pulmonata

Eucalodium orcutti from Oaxaca, 309,
310
Punta Pefiasco, fish remains from aboriginal

sites, 279-86
Pupillaria optabilis, 201

parcipictus, 300
Purpura f estiva, 179

patula pansa, 456

Purpus, Carl Albert, 397
Pusula californianus, 199, 299

solandri, 199
Pyramidella adamsi, 339, 471
Pyrazus humboldti, 166
Pyrene fuscata, 454, 469

haemostoma, 456

major, 456
Quercus agrifolia oxyadenia, 355

ganderi, 355

oxyadenia, 355

palmeri, 355
Racheospila herbaria hulstiana, 436

rubrolinearia, 436

rubromarginaria, 436
Rafinesquia californica, 387
Rattlesnake

dusky, 52-63
Reptiles

Crotalus enyo furvus, 49-64

Gopherus, 441-48

insular snakes, Gulf of California, 67-97

Pit-vipers, North American fossil, 31-46
Retusa harpa, 323, 339
Rhamnus crocea pilosa, 368
Rhus integrifolia, 367

laurina, 329, 426
Ribes canthariformis, 362

indecorum, 362
Rich, William, 397
Rissoella bakeri, 322, 326
Rissoina burragei, 465

californica, 326

cleo, 295, 299, 326

guadalupensis, 322, 326

io, 455, 465

lowei, 322, 326

pleistocena, 200

sp., 465

willetti, 322, 326
Robison, William Condit, 397
Roezl, Benedict (or Benito), 397
Rosa aldersonii, 363
Rubus ganderi, 363

idaeus aculeatissimus, 436

parviflorus villosis, 364
Rudbeckia hirta, 435
Salicornia depressa, 357
Salvadora hexalepis deserticola, 79

hexalepis hexalepis, 78-79

hexalepis klauberi, 79
Salvia clevelandii, 378

eremostachya, 379

palmeri, 379

vaseyi, 379
San Diego County

new species of Dipodomys, 477-80

old place names, 400

past and present boundaries, 348

plant collectors, 388-99

498

San Diego Society of Natural History

Pleistocene mollusks of Tecolote Creek,
335-46

type localities of vascular plants, 347-406
San Felipe

Nassarius howardae, 333, 334
Sanford, Oliver Nason, 397
Sanguinolaria nuttalli, 297
Sanicula arguta, 372
Saprinus, 212
Satureja chandleri, 379
Saviniona suspensa, 369
Saxidomus nuttalli, 195, 296
Saxifraga parryi, 362
Schizothaerus nuttallii, 196, 297
Schmaltzia cruciata, 367
Schoenefeldt, Ludwig, 398
Schott, Arthur Carl Victor, 398
Schumo, Mrs. Silas L., 398
Schuster, Robert O., and Gordon A. Marsh

A preliminary revision of the genus Psel-
aptrichus (Coleoptera: Pselaphidae) ,
3-28
Scoliodon longurio, 280, 283
Scott, Miss, 398
Scutellaria bolanderi austromontana, 379

linearifolia, 379
Sedum edule, 362

variegatum, 362
Seila assimilata, 466

montereyensis, 200, 299, 325
Selaginella cinerascens, 350
Semele bicolor, 462

corrugata californica, 462

decisa, 196

flavescens, 165, 178, 462

guaymasensis, 165, 462

incongrua, 330

jaramija, 462

quentinensis, 338, 342

rubropicta, 196

rupicola, 297
Septifer bifurcatus, 294, 296, 338
Serpulorbis margaritaceus, 454, 455, 466
Shark, Pacific sharpnose, 280
Silene multinervia, 358

palmeri, 358

platyota, 359

verecunda platyota, 359
Siliqua lucida, 196, 297, 338

patula, 297, 303
Simmondsia californica, 367
Sinum debile, 200

scopulosum, 201
Siphonaria maura, 472
Siphonodentalium quadrifissatum, 330
Sitanion minus, 353
Skenea californica, 201
Smoothhound, sicklefin, 280
Snakes. See Reptiles
Solariella triplostephanus, 463

Solariorbis elegans, 455, 465

narinensis, 455, 465
Solecardia eburnea, 460
Solen sicarius, 196
Solenosteira pallida, 166
Solidago conflnis, 387
Sonchus fallax? californicus, 387

tenuifolius, 387
Sonora, 70

mosaueri, 79
Sonora, Mexico

fish remains from aboriginal sites, 279-86

Pleistocene marine invertebrates, 153-75
Spencer, Mary Evelyn Fisk, 398
Spergularia clevelandii, 359
Sphaeralcea purpurea, 369
Spiroglyphus lituellus, 200, 299, 326
Spisula catilliformis, 196

dolabriformis, 196, 297

hemphilli, 196, 297, 301
Spondylus calcifer, 459

princeps, 457, 459
Sporobolus altissimus, 353
Staphylinidae

California intertidal, describing larvae,
207-29

Phytosi, review of genera and revision of
west American species, 103-51

revision of U. S. Hesperus, 311-18
Staphylinus apicialis, 311, 315

baltimorensis, 311, 314

rufipennis, 3 1 1
Stark, Byron David, 398
Stenoplax aethona, 473

acrior, 301

conspicuus, 301

conspicua sonorana, 473

heathiana, 169n

limaciformis, 473

magdalenensis, 169n

mariposa, 473
Stephanomeria exigua deanei, 387
Stillingia linearifolia, 367
Stipa coronata, 353

diegoensis, 353
Stokes, Susan Gabriella, 398
Stonewall Mine, 400
Stramonita biserialis, 198, 298
Streptanthus campestris, 361

heterophyllus, 361
Strigilla cicercula, 461

costulifera, 461
Strombiformis raymondi, 185, 199
Strombina maculosa, 454, 469
Strombus galeatus, 166, 468

gracillior, 468

granulatus, 179, 468
Strongylocentrotus purpuratus, 294
Stylophyllum edule, 362

orcuttii, 362

Index

499

parishii, 362
Styphonia integrifolia, 367

serrata, 367
Suaeda californica pubescens, 357
Sulcoretusa paziana, 471

xystrum, 187, 196, 204, 205
Swertia parryi, 373
Syncera translucens, 339
Synchlora aetata, 436

denticularia, 436

liquoraria, 436

rubrifrontaria, 436
Tachys, 212
Tagelus bourgeoisae, 455, 462

californianus, 165, 169, 336, 338, 462

peruvianus, 462

subteres, 196
Talley's (Tally's) Ranch, 400
Tamiascurus douglasii mearnsi, 477
Tarphiota, 212
Tecolote Creek

Pleistocene mollusks, 335-46
Tegula aureotinaa, 179, 201, 300

brunnea, 300, 303

brunnea fluctuata, 300, 303

funebralis, 294, 300

gallina, 179,201,300,328

gallina multifilosa, 300, 328

ligulata, 201, 300, 339, 456, 463

marcida, 300

marcida ssp., 300

mariana, 166, 454, 463

regina, 328

rubroflammulata, 166

rugosa, 166, 455, 463
Teinostoma amplectans, 465

herbertianum, 455, 465

invallatum, 328

supravallatum, 328
Tellina bodegensis, 296

buttoni, 195

idae, 195, 338

meropsis, 338, 342

pacifica, 330

reclusa, 461

sp., 461
Tenaturris euryclea, 471

nereis, 470

phaethusa, 471
Terebra armillata, 179

pedroana, 197, 297

sp., 471

variegata, 455, 471
Terebratulina unguicula, 331
Tetraclita, 294

squamosa albescens, 179

squamosa stalactifera, 453

squamosa stalactifera confinis, 167
Thais biserialis, 179, 456

haemastoma biserialis, 166

speciosa, 456

triangularis, 456
Thalassotrechus, 212, 213-15

barbarae, 213-15

larva, 213,214, 215, 226

barbarae barbarae, 213-14

barbarae nigripennis, 213-14

nigripennis, 213
Thalictrum coreospermum, 359

magarum, 359
Thermopsis macrophylla senota, 366
Thinopinus, 212

Thinusa, 109, 112, 113, 114, 115, 132-35,
137,212
key to species, 134

divergens, 135

fletcheri, 133, 134, 135

maritima, 132, 133, 134-35, 151

nigra, 135

obscura, 134

robustula, 135
Thomomys bottae, 99-101

bottae cedrinus, 100-101

bottae desertorum, 101

bottae desitus, 101

bottae hualpaiensis, 101

umbrinus abbotti, 407-08

umbrinus brazierhowelli, 407-08

umbrinus catavinensis, 408

umbrinus ruricola, 408
Thracia diegensis, 329

squamosa, 329, 462
Thurber, George, 398
Thyasira barbarensis, 329
Thysanocarpus laciniatus rigidus, 361
Tiariturris spectabilis, 471
Tiburon mamon, 280
Tighe's (Tigh's) Ranch, 400
Tillaea erecta eremica, 362
Tissa clevelandii, 359
Tivela stultorum, 296, 301
Tonicella lineata, 301
Tortoise, southwestern, 441-48
Totoaba, 281
Tortuava, 281

Toxicodendron comarophyllum, 368
Trachycardium consors, 460

panamense, 460

procerum, 178, 190, 195

quadrigenarium, 195, 296
Transennella puella, 330

tantilla, 195, 296, 456, 461
Trechus barbarae, 213
Trichostema parishii, 379
Tricolia compta, 201, 300

pulloides, 201, 300,327

rubrilineata, 327

substriata, 464

typica, 464

variegata, 191, 201

500

San Diego Society of Natural History

Trifolium anodon, 366

decodon, 366
Triggerfish, fine-scale, 282
Trigoniocardia biangulata, 195, 460

granulifera, 460
Trimusculus reticulatus, 297, 472
Triphora chamberlini, 325

evermanni, 466

hemphilli, 295, 299

pedroana, 200, 299, 325,339

sp., 466
Trivia californiana, 325, 456, 467

elsiae, 467

solandri, 325, 467
Truncatella bairdiana, 465

californica, 179

stimpsoni, 299
Tuckermannia maritima, 384
Turbo fluctuosus, 166, 179, 454, 464

saxosus, 464
Turbonilla aepynota, 339

almo, 199, 298

aresta, 298

asser, 199

buttoni, 298, 472

canfieldi, 199

carpenteri, 199

halidoma, 324

keepi, 199

lammata, 199

lowei, 199

lowei pedroana, 199

muricata, 472

pentalopha, 199

regina, 199

sp., 298, 472

stylina, 199

tenuicula, 199, 298

torquata, 199

tridentata, 199

weldi, 199
Turcica caffea, 201
Turricula dolenta, 456, 471

howelli, 456, 471

maculosa, 179

sp., 471
Turritella anactor, 465

clarionense, 465

cooperi, 200, 299

gonostoma, 166, 456

orthosymmetra, 326
Turtle Bay, Pliocene and Pleistocene faunas,

177-79
Tylodina fungina, 323
Type localities

vascular plants of San Diego County,
347-406
Typhis quadratus, 468

Umbraculum ovale, 323

Uropappus heterocarpus, 386

Uvigerina peregrina, 188

Valentien, Anna Marie Bookprinter, 399

Valentine, James W.

Late Pleistocene faunas from the north-
western coast of Baja California,
Mexico, 289-308

Upper Pleistocene Mollusca from Potrero
Canyon, Pacific Palisades, Califor-
nia, 181-205
Vascular plants

type localities in San Diego County, 347-
406
Vasey, George, 399
Velaea arguta, 372

arguta ternata, 372
Velutina laevigata, 300, 303
Verbena abramsii, 377
Vermetus indentatus, 466
Vermicularia eburnea, 326

pellucida, 465
Vermilion Sea Field Station, 45 1, 474, 476
Verrunculus polylepis, 282, 283, 286
Vertebrates. See common names of classes
Vestitrichus, 8
Viguiera deltoidea parishii, 387

laciniata, 387

parishii, 387
Vitis girdiana, 369
Vitrinella, 201

oldroydi, 300

sp., 465
Vitularia salebrosa, 179
Volvulella cylindrica, 197
Washingtonia filifera, 353
Weed, 399

Wieslander, Albert Everett, 399
Wiggins, Ira Loren, 399
Williamia peltoides, 197, 323, 456, 472
Wislizenia divaricata, 360
Wolf, Carl Brandt, 399
Woodhouse, Samuel Washington, 399
Wood rat

San Francisquito white-footed, 287
Wyethia coriacea, 387

ovata, 387
Xanthoxalis californica, 366
Xylococcus bicolor, 373
Xylothermia montana tomentosa, 366
Yolida cooperi, 194
Yucca schidigera, 354

whipplei, 354
Zirfaea pilsbryi, 196, 297, 338
Zizyphus parryi, 368
Zonaria spadiciea, 299
Zygadenus diegoensis, 355

/

)

Date Due

■ APR 3 o 20M-N
JAN 0 5 2QD4

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