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PROCEEDINGS

OF THE

California Academy of Sciences

FOURTH SERIES

VOL. XXXII

SAN FRANCISCO
PUBLISHED BY THE ACADEMY
1962-1968

COMMITTEE ON PUBLICATIONS
George E. Lindsay, Chairman

Dr. Epwarp L. KeEsseEt, Editor Dr. LEo G. HERTLEIN

No.

CONTENTS OF VOLUME XXXII

MILLER, Rospert C. G Dallas Hanna (frontispiece; 2 fig-
tires). ye ublisheds Ajraly 24) 9625 bee Be ee

DuruHAM, J. Wyatt. Corals from the Galapagos and Cocos
Islands (9 figures). Published June 29, 1962

KESSEL, EpwArp L., and Berta B. KesseLt. Mating be-
havior and activity-rest periodicity in Protoclythia cali-
fornica (Diptera: Platypezidae) (3 figures). Published
Aft Opel (yee eee ee RE ees oe AE i SE

EXLINE, HARRIET. Two gnaphosid spiders from Arkansas
cietieures). Published June. 29, 1962 22 = St

FRIZZELL, Don L., and C. Kurt LaAmper. Distinctive
“Congrid Type” fish otoliths from the lower Tertiary of
the Gulf Coast (Pisces: Anguilliformes) (13 figures).
Published June 29, 1962 ___ as eae. Ss Sipe i as

SWAN, LAWRENCE W., and ALAN E. Leviton. The Her-
petology of Nepal: A history, check list, and zoogeograph-
ical analysis of the Herpetofauna (4 figures). Published
PLU 2) Hie 1 OG Vee as as epee SORE RE ARI aes to cee

LreEcH, HucH B. Centrodera spurca (LeConte) and two
new species resembling it, with biological and other notes
(Coleoptera: Cerambycidae) (28 figures; 3 tables). Pub-
ishecam aye 20m LOGS) a2 ta a ae cee eee

HERTLEIN, LEo GEorGE. Contribution to the biogeography
of Cocos Island, including a bibliography (4 figures). Pub-
LUSH lu Fe pe ecg IS 0 geht ae cae ie a ae

BuTLer, C. P. The Goose Lake fragments (18 figures; 1
table); ePublished May 20; 1963. 2

FouteT, W. I., and LILLIAN J. DEmpsTER. Relationships
of the percoid fish Pentaceros richardsoni Smith, with de-
scription of a specimen from the coast of California (4
fagumes; 2 tables): Published’ May 20, 1963 ....-. =.

41-56

57-78

79-85

87-101

103-147

149-218

219-289

291-313

No.

Index to Volume XXXII

LS:

lO:

Bulli

- 18:

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7AO

Errata

CHESTERMAN, CHARLES W. Contributions to the petro-
graphy of the Galapagos, Cocos, Malpelo, Cedros, San
Benito, Tres Marias, and White Friars Islands (7 fig-
ures). Published*September 10, 1963 2 ===

HERALD, Ear S. Studies on the Atlantic American pipe-
fishes with descriptions of new species (2 tables). Pub-
lished July 28,1963. = Eee

Orr, Ropert T., and THomas C. PouLTEer. The pinniped
population of Ano Nuevo Island, California (12 figures;
3 tables). Published October’ 8; 1965 = = aaa

HAMILTON, WILLIAM J., III, and MARIAN E. HAMILTON.
Breeding characteristics of yellow-billed cuckoos in Arizona
(2 figures). Published October’ 8,19}.

SMITH, ALLYN G. The larval development of chitons
(Amphineura) (11 figures). Published October 24, 1966

GREEN, JOHN WAGENER. Revision of the Nearctic species
of Sais (Cantharidae: Coleoptera) (65 figures). Pub-
lished October 24,. 1966 2.0. 0 eS ee

CHIvERS, Dustin D. Observations on Pleurobranchaea
californica’ MacFarland, 1966 (Opisthobranchia, Nota-
spidea) (4 figures). Published November 22, 1967 —_

CHURCH, CLIFFORD C. Lower Cretaceous Foraminifera of
the Orchard Peak-Devils Den Area, California (8 plates).
Published, February 23, 1963 2

HoweELL, JOHN THoMmAS, and DuNCAN M. Porter. The
plant genus Polygala in the Galapagos Islands. Pub-
lished) «ume. 27, 1968) 1.4 oR eo ee

McC intTock, ELIzABETH, and WALTER KNIGHT. Flora of
the San Bruno Mountains, San Mateo County, California
(14 figures; 5 plates). Published November 29, 1968 __

339—sa2

363-375

377-404

405-432.

433-446

447-513 |

515-521

523-580

581-586

587-677»
679-722

722

This volume is dedicated to

G DALLAS HANNA
on the occasion of his seventy-fifth birthday

by the Academy’s Staff and Board of Trustees, with the
utmost respect and appreciation for his long service to the
Academy and his unfailing devotion to its principles and
purposes, and for his outstanding contributions to science.

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PROCEEDINGS
OF THE

CALIFORNIA ACADEMY OF SCIENCES

Fourth Series

G Dallas Hanna Anniversary Volume

Vol. XXXII, No. 1, pp. 1-40; frontispiece; 2 figs. April 24, 1962

G DALLAS HANNA
by
Robert C. Miller

California Academy of Sciences

Some men find adventure in travel, exploration, and discovery—visiting
the far and difficult places of the earth. Others find it in the library and
the laboratory, in study and thought and experiment—adventure of the
mind. To a chosen few it is given to know both kinds of adventure, and
to know them in continuing and brimming measure. One of those few is
the subject of this biographical sketch.

G Dallas Hanna was born in Carlisle, Arkansas, on April 24, 1887. He
oraduated from the University of Kansas in 1911, where he distinguished
himself by his proficiency in zoology, paleontology, and chemistry. At in-
tervals thereafter—in a detour to Alaska presently to be described—he took
an A.M. degree at the University of Kansas in 1913 and a Ph.D. at George
Washington University in 1918.

His first name is G, just lke that, no period. This causes no end of
confusion among editors and bibliographers. Acquaintances of fifty years’
standing have been known to call him Dallas, but most of his friends and

[5]

6 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 47H SER.

colleagues skirt the question of which name to use by comfortably addressing
him as “Doe.”

The newly fledged graduate of the University of Kansas promptly
obtained a job with the United States Bureau of Fisheries, and was just
as promptly assigned to the Bristol Bay area of Alaska, where he arrived
in the spring of 1911 at a point on the map labelled Nushagak. His mis-
sion seems to have been to find out everything he could about the fisheries,
aquatie biology, and miscellaneous wildlife resources of the area, so that the
Bureau of Fisheries could plan administrative and conservation measures.

This is pretty remote country even today, and one ean imagine the prob-
lems of transportation and communieation fifty vears ago. But the young
man from the middle west took to the new life as if he had been born to it.
The basic means of transportation being by boat in summer and by dog sled
in winter, he quickly became an expert seaman and an expert dog-sled
driver.

In the winter of 1912-13 he made an historie trip of a thousand miles
by dog sled, from the head of Bristol Bay to Idadorod and return. On the
first leg of the trip, from Bristol Bay to Bethel on the Kuskokwim River,
he had a companion driving another sled. But on the last and most arduous
leg of the trip, up the Kuskokwim River and over a 5000-foot pass in the
Kuskokwim Range in the dead of winter, he went it alone with no com-
panions except his seven sled dogs—animals half-wild and savage, but strong
and persevering under the control of a compelling man.

On this trip he collected 800 birds and mammals for the United States
National Museum, including ptarmigan, fox, mink, otter, shrews, and snow-
shoe rabbits. All that science needed was the skin and skulls. The meat he
apportioned between himself and his sled dogs. The purpose of this trip
was to communicate with Washington and get instructions for the following
summer. Idadorod, then a thriving metropolis of 400 people—mostly en-
gaged in placer mining in frozen ground—was the nearest telegraph station!

His new assignment was to the Pribilof Islands—more remote and wet
and cold and fog-bound even than Bristol Bay. Here he kept count of the
Alaskan fur seal herd—then as now governed by a treaty with the Japanese
geovernment—and formed lasting friendships with the local human inhabi-
tants, both Aleuts and the few resident whites, through his interest in them
and his ingenuity in suggesting new solutions to the particular problems of
an inhospitable environment.

In addition to his duties as custodian and census-taker of the fur seals
on their breeding grounds, Dr. Hanna studied the general natural history
of the Pribilofs, and published papers on both the birds and the mammals.
He also interested himself in the geology and paleontology of the islands,
and took up the study of fossil diatoms—a specialty that was to play an
important role in his subsequent career.

Vou. XXXIT] MILLER: G DALLAS HANNA Hi

In 1919 he accepted appointment as Curator of the Department of Pa-
leontology (now Geology) in the California Academy of Sciences, at the
invitation of the Academy’s then director, Barton Warren Evermann, who—
in his previous capacity of chief of the Alaska Division of the U. S. Bureau
of Fisheries—had acquired first-hand knowledge of Dr. Hanna’s brilliant
mind and versatile abilities.

Among the first problems he took up in his new position were studies
of diatoms, radiolarians, and silicoflagellates, all of which are unicellular
organisms—most of them microscopic—with a siliceous skeleton. Being al-
ways of a practical turn of mind, Dr. Hanna was aware that these organisms
could be useful in the identification of geological strata in the search for oil.
His services came into demand by the oil companies, and he began doing
part-time work for Pacific Oil Company and Associated Oil Company.

He made arrangements with these two companies to do experimental
work with oil-well sediments to determine whether microfossils could reliably
be used for the correlation of sediments. This project was approved and
work began at the Academy on May 16, 1923. To assist him, Dr. Hanna
employed Roy T. Hazzard from the University of California, and subse-
quently H. C. Driver. By December the work had progressed so favorably
that Dr. Hanna suggested that a laboratory be established in the Associated
Oil Building. This plan was approved in February, 1924, and on April 3,
1924, the laboratory was completed and ready for work in Room 637, at 79
New Montgomery Street, San Francisco. This is believed to be the first lab-
oratory for micropaleontology established by an oil company on the Pacific
Coast. Other oil companies soon followed suit; but there is no question that
Dr. Hanna pioneered this type of work, and that he added impetus to it
through his scientific investigations and through his training of younger men
who later became heads of such laboratories. He also pioneered in the sys-
tematic study of the foraminiferal content of oil-well cores. In summary, he
laid the foundation for a science that has been universally practised since,
and used to great advantage by all major oil companies.

Of his numerous published papers on geology and paleontology, probably
the one which has had the greatest influence in the search for oil in Cali-
fornia is one of the shortest. It bears the title, ““A Geologie Section in the
Center of the San Joaquin Valley, California.”” (See Bibliography, no.
208.) Mr. J. A. Taff was joint author. This paper described the geologic
formations penetrated by three wells drilled for oil in the center of the San
Joaquin Valley in Fresno County. The drill holes showed for the first time
that formations capable of producing oil were easily within the reach of the
drill on the valley floor and that they were thinning rapidly to the east. The
publication of this information resulted in extensive drilling throughout the
valley and resulted in the discovery of many oil and gas fields; and the
drilling is still going on.

8 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER.

Not content with the scientific aspects of drilling for oil, he interested
himself also in the purely mechanical aspects. He produced the first and
assisted on subsequently improved designs of the Reiber-Hanna well sur-
veying instrument used to indicate the orientation of well cores. Patent no.
1,665,058, issued to him on April 13, 1928, and assigned to the Associated
Oil Company, covered his first instrument developed as a “Means for obtain-
ing data in earth bores.” (This story is covered in an article by A. T.
Parsons, engineer of Frank Reiber, Inc., in the Associated Oil Company’s
magazine, The Record, for December, 1930.) Although such tools are com-
monplace now, twenty-five or thirty years ago they were just being put into
operation.

Figure 1, Packing into Burl’s Creek, southeast Alaska, 1938.

VoL. XXXIT] MILLER: G DALLAS HANNA 9

Dr. Hanna’s contributions to the oil industry were recognized in part by
his election as vice-president of the Society of Economic Paleontologists and
Mineralogists in 1928-29, and his election as president of that organization
in 1932-33. An equally convincing evidence of the respect in which he is
held among his peers is the fact that in 1937 and again in 1938 he was chosen
as leader and chief geologist of a joint expedition, sponsored by Tidewater
Associated, Standard of California, and Union Oil companies, to conduct
detailed geological work in the Cold Bay and Yakataga prospective oil dis-
tricts of Alaska.

As already mentioned, Dr. Hanna’s interest in microfossils, especially in
diatoms, antedated his work in petroleum geology. It began while he was
stationed on the Pribilof Islands. In 1916, when working on a fossil diatom
deposit on St. Paul Island, he designed a new type of “mechanical finger”
clamped to his microscope for greater ease in handling and mounting indi-
vidual diatom specimens for study. This is still the best apparatus available
for such extremely delicate work and is in constant use today. His work on
the diatom flora of the western part of the United States has been extensive.

His most important work on diatoms has never been published although
it is still being used in reference work; it consists of two sizable manuscripts
on file in the California Academy of Sciences. The first of these is a collec-
tion of all references to diatoms, living and fossil, which had been published
up to 1929, covering the mainland area from the Bering Sea to Panama. It
includes records of fossil diatoms occurring as far east as Utah and the east-
ern boundary of British Columbia. This manuscript, dated January 1, 1930,
runs 519 pages of typeseript and is labelled ““An Index to West American
Diatoms.” The second unpublished work is entitled “An Index to Atlas der
Diatomaceen-Kunde, by Adolf Schmidt.” The Schmidt Atlas began in 1874
and has been continued by several subsequent German contributors. This
index is an authoritative and useful contribution to the literature on diatoms,
being more complete than the prior index of this great work. Dr. Hanna’s
index is dated 1938 and includes 470 pages of typescript. As a result of Dr.
Hanna’s interest in diatoms, the California Academy of Sciences has one of
the best reference libraries on the subject in existence. He is recognized
internationally as well as nationally as one of the leading experts on the
subject. And even now he is imparting his knowledge and experience to a
graduate student under a National Science Foundation grant.

In the course of his study of diatoms, he found it necessary to mount the
specimens in a medium of higher index of refraction than any yet available.
Canada balsam, the medium most commonly used, did not provide sufficient
resolution; he couldn’t see the details. The challenge to find something better
brought him to a study of other natural resins, and of synthetic resins. Call-
ing on his early proficiency in chemistry, he presently came up with a syn-
thetic resin made from naphthalene and formaldehyde. To this excellent

10 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 47TH SER.

resin, which had a much higher index of refraction, he gave the name “Hy-
rax.”’ It is now made and sold commercially by biological supply houses.
Later he developed an even better mounting medium which he has ealled
“Pleurax.”’ With an improved mounting medium, plus a method of coating
specimens under high vacuum with a film of realgar (an arsenie sulphide),
Dr. Hanna proved the possibility of photographing diatoms under magnifi-
cations up to several thousand times to show their minute structure in
pertect focus. This was a significant breakthrough toward a more complete
knowledge of diatom structure.

In recognition of these contributions to microscopy, Dr. Hanna was made
a Fellow of the British Royal Microscopical Society. And because of his
extensive knowledge of microfossils and micro-techniques, he accepted an
appointment in 1928 to serve on the Committee on Micropaleontology in the

Figure 2. At work on diatoms.

Vou. XXXII] MILLER: G DALLAS HANNA alg

Division of Geology and Geography of the National Research Council and
also contributed to the work of a companion Committee on Paleobotany.

His interest in improving visibility through the microscope led him into
the further study of optics, which he pursued with characteristic thorough-
ness. This had at least two important consequences.

At the beginning of World War IT the United States was extremely short
on optical equipment, and in facilities and know-how for manufacture of
such equipment. We had depended largely on Germany and Japan for
optical supplies, and we were at war with both of them. There was really
a critical shortage of such ordinary things as prisms for binoculars and other
instruments, not to mention the optical parts for gun sights, range finders,
and related equipment.

Dr. Hanna’s initial contribution to the war effort involved the manufac-
ture of Amici or “roof” prisms of which the U. S. Army was in desperate
need. These were thumb-sized prisms used in range finders, which bent the
hight beam 90 degrees and erected the image at the same time. The 90-degree
“roof” angle had to be correct to provide accuracy of one foot in an eight-
mile distance. There was no known mechanical method of reaching such
accuracy—it had to be achieved by hand. Working with a small group of
amateur telescope makers and others, he perfected each of the 76 separate
processes required to make these prisms, trained the group of people working
with him, and delivered 200. All passed the rigorous tests with a 100 per
cent acceptance by the Army.

While engaged in this work, Dr. Hanna learned one day that a submarine
at Mare Island had been out of commission for several weeks on account of
a broken head-prism in its periscope. He asked whether the parts of the
broken prism were available. They were. Using these to determine size and
angles, he turned out a new head prism over a week-end and the submarine
was returned to service. This resulted almost immediately in a contract with
the U. S. Navy which kept the Academy exceedingly busy till the end of the
war. Twenty-five thousand instruments were repaired or serviced—hinocu-
lars, gun sights, navigation instruments—and eleven thousand optical parts
were manufactured or repaired in the Academy’s shop by a corps of some 50
instrument repair people and a few volunteer glass grinders and polishers,
all under Dr. Hanna’s direction.

When “black glass” for bearing-circle mirrors used in navigation became
unavailable, he ingeniously substituted obsidian, a natural black glass of
voleanic origin available in quantity at several locations in California, and
found it just as good or better. Under the Navy contract Dr. Hanna and his
eroup handled nearly all optical repair work for the Pacific Fleet and for
some 7000 vessels of the Armed Guard (Liberty and Victory ships). When
the job was over, the group could boast with pride that in spite of emer-
gencies and close deadlines it ‘never missed the sailing date of a ship.”

12 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER.

After this work terminated at the close of the war, Dr. Hanna and his as-
sociates received a “well done” from the Navy in the form of a certificate
awarded to the Academy “in recognition of exceptional accomplishment in
behalf of the United States Navy and of meritorious contribution to the
national war effort.”

At the end of the war the Academy found itself with a well-equipped
instrument shop and a group of highly skilled optical instrument workers.
The Board of Trustees had in the meantime become interested in building
a planetarium, and had a sum of money available for that purpose. But
they were unable to obtain a star projector. All of these instruments had
been built in the Zeiss optical works in Germany, and at the end of the war
the Zeiss plant had been dismantled and shipped to Russia. In this dilemma
Dr. Hanna suggested, “Why not build the star projector ourselves in our
own shop?” The Trustees, after some proper hesitation at so brash a pro-
posal, agreed—and thus the Morrison Planetarium came into being. Now
in operation for nearly a decade, it is unique in design with many new
features and is universally conceded to be one of the best in the world.

One of Dr. Hanna’s most unique and characteristic contributions to this
project was the method he devised for making the star plates. Grains of
ecarborundum powder, sorted for relative size to correspond to stars of dif-
ferent magnitudes, were placed in proper position on a flat glass surface by
means of a device similar to the “mechanical finger” for mounting diatoms
mentioned above. Then the glass was aluminized by an evaporation process
in a vacuum chamber, making it opaque. Finally the carborundum particles
were brushed off, leaving apertures through which light could be projected,
forming the star-images. All previous “artificial stars” had been projected
through small round holes in a metal plate so that they appeared on the
planetarium dome as circular spots or dises. The stars achieved by the ear-
borundum process were of irregular shape, producing a more realistic night
sky.

Dr. Hanna is remarkably ingenious in designing and buildine special
equipment for researech—his own or others. Seores, if not actually hundreds,
of scientists are indebted to him for advice on how to build or modify equip-
ment to meet their particular needs; and the Academy has had numerous
contracts for “research and development” from state, Federal and private
agencies that wished to utilize Dr. Hanna’s special inventiveness in design-
ing equipment.

A eurrent example of this is his present work on a new and inexpensive
method of illustrating secientifie papers in full color which has lone been a
research scientist’s unfulfilled dream. Learning of research work along this
line by the Eastman Kodak Company, Dr. Hanna went to Rochester, New
York, in December of 1954, to study their process and equipment. With the
information obtained at Rochester, he built the necessary equipment in the

VoL. XXXIT] MILLER: G DALLAS HANNA 3

Academy's instrument shop and then began a slow but gradually winning
battle over many obstacles, both mechanical and optical, that no one lacking
Dr. Hanna’s persistence and his insistence on perfection could have won.
This has been a teamwork job, first with Charles EK. Crompton and now with
Maurice C. Giles on the photographic side, and with himself and his wife,
Margaret M. Hanna, on the multilith reproduction side.

It was not until 1960 that Dr. Hanna felt that the new three-color offset
process was sufficiently advanced to provide the first published color illustra-
tion, which appeared in the Academy’s Annual Report for that year. Since
then other color plates, produced by him and his associates in the Academy’s
photographie laboratory and instrument shop, have been included in several
recent numbers of the Academy’s Proceedings and Occasional Papers as well
as in a number of publications of other scientific societies. Each one has been
better than the one preceding, but he is still working toward still greater
perfection of the process. The beautiful color frontispiece of the yellow
arctic poppy, appearing in A Flora of the Alaskan Arctic Slope, by Iva L.
Wiggins and John Hunter Thomas, just published by the Arctic Institute of
North America, is an excellent example of this color work.

At this writing, comparable printed color reproductions based on the
Eastman offset process are produced nowhere else in the country. The high
quality of the color reproductions achieved by the Hanna color team, to-
gether with the ability to show extremely fine detail, makes this work a
unique contribution to scientific illustration. And one great advantage is
that the cost is of a magnitude no greater than for comparable black-and-
white reproductions.

The Academy’s work on scientific illustrations in color is already fairly
well-known. As one evidence of this, the University of Costa Riea sent the
superintendent of its University press to the Academy in the fall of 1960
to study our equipment and methods.

Dr. Hanna is well-known among his friends and associates as an excellent
field collector and photographer. He has made many important additions
of bird skins, bird eges, fossil vertebrates and invertebrates, living land,
fresh-water and marine mollusks, mineral specimens, plants, and other
things to the research collections of many museums, especially the U. S.
National Museum in Washington, D.C., and the California Academy of
Sciences. Size is no criterion for him as items he has collected range from
huge fossil whale skulls to the tiniest of diatoms and radiolaria. The excel-
lence and extent of the fossil invertebrates and recent shells in the research
collections of the Academy’s Department of Geology, consisting of nearly
38,000 locality numbers, together with a special type collection of more than
12,350 specimens, have developed over the years under his supervision as the
department’s Curator.

14 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER.

As a research scientist in the field, Dr. Hanna has few if any equals. He
takes advantage of opportunities to get into little-known country on colleet-
ing trips, short or long, provided there is some worth-while scientifie objeec-
tive to be attained. He is “tops” as a camper, camp cook, and an interesting
companion around a campfire—and this without regard to weather or other
problems. He has represented the Academy on several major expeditions,
including one to Guadalupe Island, Mexico, in 1922, and another to the
Revillagigedo Islands, Mexico, in 1925. He was on the Orca expedition to
the Gulf of California in 1953. In 1949 he designed the specially massive
dredging and trawling equipment used on the U. S. Navy net-tender Mul-
berry for an Academy-sponsored geological exploration of the sea bottom off
central California under the auspices of the Office of Naval Research. Dr.
Hanna directed operations aboard the Mulberry for the 24 days she was at
sea on this project, which resulted in much new knowledge of the underwater
geology of the area in depths down to 2000 fathoms. In the following year
he spent several weeks on the California Division of Fish and Game fisheries
research vessel NV. B. Scofield, off the coast of northern California collecting
marine invertebrates and rocks from considerable depths for a further study
of the underwater geology. These are but a few of his longer periods in the
field. His collecting trips of shorter duration would make a long list, much
too extensive to be covered here.

One of the ecapstones of Dr. Hanna’s recognition as a scientist came in
1959 when his alma mater conferred on him the “Erasmus Haworth Dis-
tinguished Alumni Honors in Geology,” which was awarded by the Univer-
sity of Kansas and presented to him at the annual meeting of the American
Association of Petroleum Geologists held that year in Dallas, Texas.

This account thus far has emphasized Dr. Hanna’s practical aeeomplish-
ments. We will now attempt to show that he is just as impractical as any-
body. Witness his papers on “New and Interesting Records of Pribilof
Island Birds,” “Mammals of the St. Matthew Islands, Bering Sea,” “Random
Notes on Alaska Snow Buntings,” “Extremely Diversified Habitat of a
Marine Gastropod,” “Fossil Diatoms Dredged from Bering Sea.” Dr. Hanna
is first and last a scientist, who reports carefully and accurately any observa-
tions that come within his field of competence, which, as we have already
shown, is a very wide field. His voluminous field notes, meticulously written,
often after a long, hard day in the field, cover many years of collecting work
and observation. They contain much of interest, both scientifie and his-
torical. If the things he finds out happen to have some practical use, he has
no objection; but his primary purpose is to discover the facts for themselves.

His love of the far north is a part of his character, and he returns to
Alaska at every opportunity. In the summer of 1955 he served as Director
of the Arctic Research Laboratory at Point Barrow, and the following sum-
mer he returned and spent an entire year (1956-57) at Point Barrow as ¢

VoL. XXXIT] MILLER: G DALLAS HANNA 15

special investigator, accompanied by his wife, Margaret, who served in her
special capacity as a geological artist and illustrator. His most recent trip
was with Academy Trustee Kenneth K. Bechtel in the summer of 1960, when
he revisited St. Paul Island in the Pribilofs where he had been stationed
nearly half a century ago.

To close this altogether inadequate account of G Dallas Hanna—the
scientist and the man—we can do no better than to quote what a long-time
friend and close associate has written:

“The unique qualities of Dr. Hanna’s mind and personality, which have
made him stand out in the fields of biology, geology, and paleontology, are
his keen, critical powers of observation; his clear, analytical mind; his
patience and persistence in the face of obstacles; his retentive memory and
his ability to associate related facts; his ability to inspire those around him
to become interested in what he is doing; the power to inspire in others con-
fidence and belief in their own worth and ability; the personality to make
friends of all kinds of people from all walks of life; in experimentation, the
attitude of patience to try many things and to discard the failures without
regret; the mental attitude to explore all sides of a question and the courage
to challenge accepted ideas and viewpoints; the honesty always to give full
eredit to his co-workers for their contribution in a joint project; the innate
modesty to minimize or play down his own part in a joint accomplishment;
and in speaking of others, to praise their good qualities and minimize the
less admirable ones—to speak well of others or not at all.”

Author’s Note: In preparing this biographical sketch I have received valued as-
sistance and information from a number of Dr. Hanna’s close associates over the
years. These include his brother, Marcus A. Hanna, Clifford C. Church, Leo G.
Hertlein, Allyn G. Smith, and the late G. Clark Gester and William S. W. Kew.

—R. C. M.

BIBLIOGRAPHY OF G DALLAS HANNA
1909-1961

1909

1. The Mollusea of Douglas County, Kansas—Gasteropoda. Nautilus, vol. 23, no. 6,
pp. 81-82. November—December.

2. [Plates 5 and 7 from drawings by Hanna, and some retouching of photographs
on other plates in] Carboniferous air-breathing vertebrates of the United States
National Museum, by Roy L. Moodie. Proceedings of the United States National
Museum, vol. 37, no. 1696, pp. 11-28, pls. 4-10. September 23.

1910
Three shells not hitherto reported from the District of Columbia. Nautilus, vol.
24, no. 2, p. 24. June.

4. [Figure 3 drawn by Hanna inj The alimentary canal of a Carboniferous sala-

10.

12.

13.

14.

15.

16.

CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER.

mander, by Roy L. Moodie. American Naturalist, vol. 44, no. 522, pp. 367-375,
figs. 1-4. June.

[Figures on all six plates drawn by Hanna except pl. 49, fig. 1 in] The
Temnospondylous Amphibia and a new species of Hryops from the Permian of
Oklahoma, by Roy L. Moodie. Kansas University Science Bulletin, vol. 5, no. 3,
pp. 235-253, pls. 49-54. March.

1907.
[Four drawings by Hanna in] A new labyrinthodont from the Kansas coal
measures, by Roy L. Moodie. Proceedings of the United States National Mu-
seum, vol. 40, no. 1796, pp. 489-495. January 30.
[Drawings by Hanna in] Two amphibians, one of them new, from the Carbon-
iferous of Illinois, by Roy L. Moodie. Proceedings of the United States National
Museum, vol. 40, no. 1828, pp. 429-433, figs. 1-2. May 8.
The American species of Sphyradium with an inquiry as to their generic rela-
tionships. Proceedings of the United States National Museum, vol. 41, no. 1865,
pp. 371-376, figs. 1-4. October 14.

1912
[Five drawings on three plates by Hanna in] On a comparison of three
skulls: Castoroides ohioensis, Castoroides kansensis, and Castor, by H. T. Mar-
tin. Kansas University Science Bulletin, vol. 6, no. 6, pp. 389-396, pls. 24-27.
January.
[Plate 2 drawn by Hanna in] The skull structure of Diplocaulus magnicornis
Cope and the amphibian order Diplocaulis, by Roy L. Moodie. Journal of Mor-
phology, vol. 23, no. 1, pp. 31—48, pls. 1-2. March 20.
[Plates all from retouched photographs and drawings by Hanna; three plates
are restorations; in] The Pennsylvanic Amphibia of the Mazon Creek, Illinois,
shales, by Roy L. Moodie. Kansas University Science Bulletin, vol. 6, no. 2, pp.
323-359, pls. 1-14. January.

1913
[Senior author with Edward C. Johnston of] A Pleistocene molluscan fauna
from Phillips County, Kansas. Kansas University Science Bulletin, vol. 7, no. 3,
pp. 111-121, pl. 18. January.

1914
[Note on] Interesting mammals on the Pribilof Islands. Proceedings of the
Biological Society of Washington, vol. 27, p. 218. October 31.

1915
[Three text figures drawn by Hanna in] Some methods of studying fossil
Amphibia embedded in coal, by Roy L. Moodie. Kansas University Science
Bulletin, vol. 9, no. 16, pp. 187-193, figs. 1-3. September.

1916
[Figures 6, 7, 9, 11, 14a, 15, 15a, 16-19, 21-27, 29, 35, 38-40, 42 and plates 4, 5,
and 16 drawn by Hanna in] The Coal Measures Amphibia of North America, by
Roy L. Moodie. Carnegie Institution of Washington, publ. no. 238, pp. i-x, 1-122,
pls. 1—26, figs. 1-43. September 28.
Records of birds new to the Pribilof Islands including two new to North
America. Auk, vol. 33, no. 4, pp. 400-403. October.

VoL

18.

IGE

20.

Zils

22.

23.

24.

25.

26.

27.

28.

29.

30.

. XXXIT] MILLER: G DALLAS HANNA 17

A new mollusk of the genus Pisidium from Alaska, with field notes by G Dallas
Hanna, by Victor Sterki. Proceedings of the United States National Museum,
vol. 51, no. 2160, pp. 475—477, figs. 1-2. December 16.

1917
Fur-seal census, Jn Alaska Fisheries and Fur Industries in 1915. Appendix 3
to Report of the United States Commissioner of Fisheries for the fiscal year
ended June 30, 1915, Bureau of Fisheries, doc. no. 834, pp. 83-96. January 6.

Branded seals. Jn Alaska Fisheries and Fur Industries in 1915. Appendix 3 to
Report of the United States Commissioner of Fisheries for the fiscal year ended
June 30. Bureau of Fisheries, doc. no. 834, pp. 97-103. January 6. [Much other
matter on the Pribilof Islands in this report prepared by Hanna. ]

The summer birds of the St. Matthew Island Bird Reservation. Auk, vol. 34, no.
4, pp. 403-410. October.

Fur-seal census, 7n Alaska Fisheries and Fur Industries in 1916. Appendix
2 to the Report of the United States Commissioner of Fisheries for the fiscal
year 1916, doc. no. 838, pp. 91-102. August 14.

1918
Branded seals. 7n Alaska Fisheries and Fur Industries in 1917. Appendix 2 to
the Report of the United States Commissioner of Fisheries for the fiscal year
1917, doc. no. 847, pp. 86-87. December 28.

Fur-seal census. Jn Alaska Fisheries and Fur Industries in 1917. Appendix 2 to
the Report of the United States Commissioner of Fisheries for the fiscal year
1917, doc. no. 847, pp. 97-123. December 28. [Much other matter on the Pribilof
Islands in this report prepared by Hanna. ]

i)
Additions to the avifauna of the Pribilof Islands, Alaska, including species new
to North America. Journal of the Washington Academy of Sciences, vol. 9, no.
6, pp. 176-177. March.
The introduction of Acanthinula harpa and Circinaria vancouverensis into St.
Paul Island, Alaska. Nautilus, vol. 32, no. 4, p. 148. April.

[Photographs on pages 264, 274-276, and 280 taken by Hanna and published in]
The northern fur-seal problem as a type of many problems of marine zoology,
by Dr. Barton Warren Evermann. Scientific Monthly, vol. 9, no. 3, pp. 263-282.
September.

Geological notes on the Pribilof Islands, Alaska, with an account of the fossil
diatoms. American Journal of Science, ser. 4, vol. 48, art. 18, pp. 216—224.
September.

Birds. Jn Alaska Fisheries and Fur Industries in 1918. Appendix 7 to the Re-
port of the United States Commissioner of Fisheries for the fiscal year 1918,
doc. no. 872, pp. 105-107. December 8.

Fur-seal census, Pribilof Islands, 1918. 7n Alaska Fisheries and Fur Industries
in 1918. Appendix 7 to the Report of the United States Commissioner of Fish-
eries for the fiscal year 1918, doc. no. 872, pp. 116-128, pls. 6-10. December 8.

1920
A comparative anatomical discussion of the five species of Cerions involved in
the breeding experiments. 7n Experiments in the breeding of Cerions, by Paul
Bartsch. Carnegie Institution of Washington, vol. 14, no. 282, pp. 7-13, pls. 1-6.

Sl.

40.

41.

42.

43.

44,
45.

46.

47.

49.

50.

CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER.

[Announcement of the appointment of G Dallas Hanna as Curator of Inverte-
brate Paleontology, California Academy of Sciences. Nautilus, vol. 33, no. 3, p.
105. January. |

Additions to the avifauna of the Pribilof Islands, Alaska, including four species
new to North America. Auk, vol. 37, no. 2, pp. 248-254. April.

Mammals of the St. Matthew Islands, Bering Sea. Journal of Mammalogy, vol.
1, no. 3, pp. 118-122. May.

Miocene land shells from Oregon. Kansas University Science Bulletin, vol. 13,
no. 1, pp. 3-12, 1 pl. May.

Pleistocene mollusks from Wallace County, Kansas. Kansas University Science
Bulletin, vol. 13, no. 2, pp. 17-19. May.

[Annual report of the] Department of Invertebrate Paleontology. In Report of
the Director of the Museum for the year 1919. Proceedings of the California
Academy of Sciences, ser. 4, vol. 9, no. 15, pp. 387-889. August 4.

Fossil mollusks from the John Day Basin in Oregon. University of Oregon Pub-
lication, vol. 1, no. 6, pp. 1-8, pl. 1. August.

New and interesting records of Pribilof Island birds. Condor, vol. 22, no. 5, pp.
173-175. September—October.

Fur-seal census, Pribilof Islands, 1919. Zn Alaska Fisheries and Fur Industries
in 1919. Appendix 9 to the Report of the United States Commissioner of
Fisheries for the fiscal year 1919, doc. no. 891, pp. 106-117, pls. 3-4. Decem-
ber 11.

Birds of the Alaska fur-seal islands. Gull, vol. 2, no. 12, pp. 1-2. December.

1ePAl
[Note on] Results of fur seal protection. Gull, vol. 3, no. 1, p. 2. January.

Modiolus demissus Dillwin, in San Francisco Bay. Nautilus, vol. 34, no. 3, pp.
91-92. January.

[Photographs on pages 30, 44, and 59, and note on page 67 by Hanna in] The
Story of Matka, by David Starr Jordan. World Book Company, Animal Life
Series, no. 1.

The Pribilof sandpiper. Condor, vol. 238, no. 2, pp. 50—57, fig. 18. March—April.

[Review of] Lake Maxinkuckee, a physical and biological survey, by Barton
Warren Evermann and Howard Walton Clark. Nautilus, vol. 34, no. 4, p. 142.
April.

[Three photographs on plate 2 by Hanna in] A new aberrant form of Vanessa
virginiensis Drury (Huntera Fabr.), by Chas. L. Fox. Psyche, vol. 28, no. 2,
pp. 45-46, pl. 2. April.

[Coauthor with Joseph Mailliard of] New bird records for North America with
notes on the Pribilof Island list. Condor, vol. 23, no. 3, pp. 93-95. May—June.

[Annual report of the] Department of Invertebrate Paleontology. Zn Report of
the Director of the Museum for the year 1920. Proceedings of the California
Academy of Sciences, ser. 4, vol. 10, no. 12, pp. 143-146. July 2.

A fossil ammonite exhibited. Golden Gate Pathfinder, vol. 2, no. 15, p. 15.
July 15.

Galls and gall insects in Academy of Sciences. Golden Gate Pathfinder, vol. 2,
no. 16, p. 12. July 24.

VoL. XXXIT] MILLER: G DALLAS HANNA 19

51.

52.

53.

D4.

55.

56.

57.

58.

59.

60.

61.

62.

63.

64.

65.

66.

67.

68.

69.

73.

Flower exhibit in the Academy of Sciences. Golden Gate Pathfinder, vol. 2, no.
U7, 10s 16. afl oil.

Pictures of California lilies. Golden Gate Pathfinder, vol. 2, no. 18, p. 15. Aug-
ust 7.

[Notes on] A lecture by a plant expert, by Dr. Albert Mann (on diatoms).
Golden Gate Pathfinder, vol. 2, no. 19, p. 15. August 14.

Steller sea lion group, Academy of Sciences. Golden Gate Pathfinder, vol. 2,
MOO pe lO. fies eIuly, 155

California sea lions. Golden Gate Pathfinder, vol. 2, no. 20, p. 16, 1 fig. Aug-
ust 21.

Grizzly bear shot by Dr. Pope with bow and arrow—now being installed in
Academy of Sciences Museum. Golden Gate Pathfinder, vol. 2, no. 21, p. 15, 1 fig.
August 28.

Hair seals in California Academy of Sciences. Golden Gate Pathfinder, vol. 2,
no. 21, p. 16, 1 fig. August 28.

Alaska fur seals. Hamlyn’s Menagerie Magazine, ca. August.

The mule deer group, Academy of Sciences. Golden Gate Pathfinder, vol. 2, no.
22, p. 16, 1 fig. September 4.

Desert bird group, Academy of Sciences. Golden Gate Pathfinder, vol. 2, no. 23,
p. 16, 1 fig. September 11.

The fur-seal group, Academy of Sciences. Golden Gate Pathfinder, vol. 2, no. 24,
p. 16, 1 fig. September 18.

The white pelican group, Academy of Sciences. Golden Gate Pathfinder, vol. 2,
no. 25, p. 15, 1 fig. September 25.

Genital organs of hermaphroditic fur seals. American Naturalist, vol. 55, no.
640, pp. 473-475. September—October.

Frederick Morton Chamberlain [Obituary]. Science, now ser., vol. 54, no. 1397,
pp. 323-324. October 7.

[Sunday afternoon lectures at the Academy of Sciences]. Golden Gate Path-
finder, vol. 2, no. 26, p. 3. October 2.

The antelope group, Academy of Sciences. Golden Gate Pathfinder, vol. 2, no.
26, p. 16, 1 fig. October 2.

[Sunday afternoon lecture at the Academy of Sciences]. Golden Gate Path-
finder, vol. 2, no. 27, p. 13. October 9.

The coyote group, Academy of Sciences. Golden Gate Pathfinder, vol. 2, no. 27,
p. 16, 1 fig. October 9.

Farallone Islands’ bird rookery in the Academy of Sciences. Golden Gate Path-
finder, vol. 2, no. 28, p. 16, 1 fig. October 16.

Progress on Steinhart Aquarium. Golden Gate Pathfinder, vol. 2, no. 29, p. 10.
October 23.

The California condor group, Academy of Sciences. Golden Gate Pathfinder,
vol. 2, no. 29, p. 16, 1 fig. October 23.

The mountain lion group, California Academy of Sciences. Golden Gate Path-
finder, vol. 2, no. 30, p. 16, 1 fig. October 29.

Epiphragmophora fidelis (Gray) near San Francisco Bay? Nautilus, vol. 35, no.
2, pp. 34-35. October.

85.

86.

93.

94.

CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER.

Frederick Morton Chamberlain [Obituary]. Nautilus, vol. 35, no. 2, p. 60.
October.

[Introduction to] Insects of the Pribilof Islands, Alaska, by E. C. Van Dyke,
et al. Proceedings of the California Academy of Sciences, ser. 4, vol. 11, no. 14,
pp. 153-195, figs. 1-4a. November 2.

The wild birds of Golden Gate Park. Golden Gate Pathfinder, vol. 2, no. 31, p.
16. November 6.

The tree ferns of Golden Gate Park. Golden Gate Pathfinder, vol. 2, no. 32, p.
16. November 13.

[Announcement of] Appointment of Dr. B. W. Evermann, Director, and Mr.
Alvin Seale, Superintendant, of new Steinhart Aquarium. Science, new ser.,
vol. 54, no. 1403, p. 489. November 18.

The Roosevelt elk group. Academy of Sciences. Golden Gate Pathfinder, vol. 2,
no. 33, p. 16, 1 fig. November 20.

The Academy of Sciences expedition to South America. Golden Gate Path-
finder, vol. 2, no. 34, p. 5. November 27.

The San Joaquin Valley water fowl group, Academy of Sciences. Golden Gate
Pathfinder, vol. 2, no. 34, p. 16, 1 fig. November 20.

Exhibit of painted flowers in California Academy of Sciences. Golden Gate
Pathfinder, vol. 2, no. 36, p. 10. December 11.

The robins of the Park. Golden Gate Pathfinder, vol. 2, no. 36, p. 16. Novem-
ber 20.

Fur-seal census, Pribilof Islands, 1920. Jn Alaska Fisheries and Fur Industries
in 1920. Appendix 6 to the Report of the United States Commissioner of Fish-
eries for the fiscal year 1921, doc. no. 909, pp. 104-121, fig. 3. December 12.
Natural-history records of Pribilof Islands. Jn Alaska Fisheries and Fur In-
dustries in 1920. Appendix 6 to the Report of the United States Commissioner
of Fisheries for the fiscal year 1921, doc. no. 909, pp. 122-127. December 12.
The sparrows of Golden Gate Park. Golden Gate Pathfinder, vol. 2, no. 38, p.
16. December 25.

Alaska fur seals. Year Book of the Amateur Menagerie Club of London for 1921.

1922
Notes on the anatomy of Strobilops labyrinthica (Say). Nautilus, vol. 35, no.
3, pp. 91-93, figs. 10-17. January.
[Review of] The gray garden slug with notes on allied forms, by A. L. Lovett
and A. B. Black. Nautilus, vol. 35, no. 3, p. 97. January.
Frederick Morton Chamberlain [Obituary]. Indiana University Alumni Quar-
terly, vol. 9, no. 1, pp. 94-95. January.
A superior eye splice. Popular Mechanics, p. 288. February.
[Senior author with Barton W. Evermann of] An exhibit of fur-bearing ani-
mals planned by the California Academy of Sciences. Golden Gate Pathfinder,
vol. 3, no. 10, p. 15. March 5.
[Senior author with William M. Grant of] Genera of diatoms characteristic of
marine and fresh water. Mining in California, California State Mining Bureau,
vol. 18, no. 2, pp. 59-76, pls. 1—5, figs. 1-27. February.

The 1921 expedition of the California Academy of Sciences to the Gulf of Cali-
fornia. Science, new ser., vol. 55, no. 1421, pp. 305-307. March 24.

Vol

95.

96.

Silke
98.
99.

100.

101.

102.

103.

104.

105.

106.

107.

108.

109.
110.
itil.

112.
113.

114.

119.

120.

. XXXII] MILLER: G DALLAS HANNA 21

Pictures of some of California’s wild flowers exhibited. Golden Gate Pathfinder,
vol. 3, no. 14, p. 16. April 2.

[Note on] Gift of the Johnston Collection of land and fresh-water shells to the
California Academy of Sciences. Nautilus, vol. 35, no. 4, pp. 133-134. April.
Some other parks. Golden Gate Pathfinder, vol. 3, no. 17, p. 16. April 23.

San Francisco needs a zoo. Golden Gate Pathfinder, vol. 3, no. 18, p. 16. May 7.
What becomes of the fur seals. Science, new ser., vol. 55, no. 1428, pp. 505-507.
May 12.

The new robin group in the California Academy of Sciences. Golden Gate Path-
finder, vol. 3, no. 20, p. 16. May 21.

Wild flowers exhibited at the California Academy of Sciences. Golden Gate
Pathfinder, vol. 3, no. 21, p. 16. May 28.

Nature’s curious mistake; mussels of ship bottom capture small fish. Popular
Mechanics, p. 728. May.

The ivory carvers of Bering Sea. Golden Gate Pathfinder, vol. 3, no. 20, pp.
761, 763, 4 illustrations. May.

[Brief note on Columella alticola (Ingersoll) in] Land snails from the Ca-
nadian Rockies, by S. Stillman Berry. Canada Department of Mines, Victoria
Memorial Museum, Bulletin no. 36, Biological Series no. 8, p. 15. May 9.

The Aleutian rosy finch. Condor, vol. 24, no. 3, pp. 88-91, fig. 32. May—June.
Bird drives in the Yukon Delta. Condor, vol. 24, no. 3, p. 100. May—June.

The quails of the Park. Golden Gate Pathfinder, vol. 3, no. 22, p. 16. June 4.
Manzanita, the little apple of California. Golden Gate Pathfinder, vol. 3, no. 24,
p. 16. June 18.

Relics of appreciation. Golden Gate Pathfinder, vol. 3, no. 25, p. 16. June 25.
The Pismo clam. Golden Gate Pathfinder, vol. 3, no. 27, pp. 6-7. July 9.

New exhibit of American game animals. Golden Gate Pathfinder, vol. 3, no. 29,
pp. 6-7. July 23.

The status of Helix oregonensis Lea. Nautilus, vol. 36, no. 1, pp. 12-14. July.
[Notes on diving of cormorants and death of old squaws and harlequin ducks
in] Report of E. H. Forbush, Director, Division of Ornithology, Department of
Agriculture, Commonwealth of Massachusetts, Bull. 8, pp. 29, 32, 41.
Commission of scientists returns to San Diego after investigations of rare
marine animals. The San Diego Union, August 17.

[Annual report of the] Department of Invertebrate Paleontology. Jn Report of
the Director of the Museum for the year 1921. Proceedings of the California
Academy of Sciences, ser. 4, vol. 11, no. 21, pp. 636-638. August 21.
Guadalupe Island. Golden Gate Pathfinder, vol. 3, no. 34, p. 2. August 27.
Fossil fresh-water mollusks from Oregon contained in the Condon Museum of
the University of Oregon. University of Oregon Publication, vol. 1, no. 12, pp.
1-22, pls. 1-4. August 29.

The reindeer herds of the Pribilof Islands. Scientific Monthly, vol. 15, no. 2,
pp. 181-186, 5 illustrations. August.

The protection of wild life. Golden Gate Pathfinder, vol. 3, no. 36, pp. 2, 15.
September 10.

Why not protect the fur-seal herds of the Southern Hemisphere? Australian
Zoologist, vol. 3, pt. 1, pp. 11-13. September 15.

121.

129.

130.
131.

132.

133.

137.

138.

139.

CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER.

The mountain lion group, California Academy of Sciences. Golden Gate Path-
finder, vol. 3, no. 37, pp. 2, 4. September 17.

A recent scientific expedition to the islands off the west coast of Lower Cali-
fornia. Golden Gate Pathfinder, vol. 3, no. 38, p. 2. September 24. (Also pub-
lished in The Catalina Islander, vol. 9, no. 37, pp. 6-7. September 22.)

[Sunday afternoon lectures at the California Academy of Sciences.] Golden
Gate Pathfinder, vol. 3, no. 39, pp. 2, 6. October 1.

Elephant seals. Golden Gate Pathfinder, vol. 3, no. 40, p. 2. October 8.
Attu Island basketry. Golden Gate Pathfinder, vol. 3, no. 42, p. 2. October 22.

Extremely diversified habitat of a marine gastropod. Ecology, vol. 3, no. 4, p.
347. October.

A recent scientific expedition to the islands off the west coast of Lower Cali-
fornia. Science, new ser., vol. 56, no. 1453, pp. 503-504. November 3.

[Unsigned notice of] Dr. Evermann’s attendance at the Pan-Pacific Commercial
Conference at Honolulu, October 25—-November 8, 1922. Science, new ser., vol.
56, no. 1453, p. 508.

1923
[Plates 52-57 from photographs supplied by Hanna in] A fauna from the middle
Eocene shales near Vacaville, California, by Dorothy Bryant Kemper Palmer.
University of California Publications, Bulletin of the Department of Geological
Sciences, vol. 14, no. 8, pp. 289-318, pls. 52-57. November 3.
What is the future of the fur seal? Fur Trade Review, pp. 357-359. January.

Upper Miocene lacustrine mollusks from Sonoma County, California. Proceed-
ings of the California Academy of Sciences, ser. 4, vol. 12, no. 3, pp. 31-41, pls.
1-3. January 2.

[Senior author with Emmet Rixford of] Notes on some land snails of the Sierra
Nevada mountains, with description of a new species. Proceedings of the Cali-
fornia Academy of Sciences, ser. 4, vol. 12, no. 4, pp. 43-50, pl. 4. January 2.

A new species of Carychium from Vancouver Island, British Columbia. Pro-
ceedings of the California Academy of Sciences, ser. 4, vol. 12, no. 5, pp. 51-53,
fig. 1. January 2.

Random notes on Alaska snow buntings. Condor, vol. 25, no. 2, pp. 60—65, figs.
22-23. March—April.

[Review of] Fauna from the Eocene of Washington, by Charles E. Weaver and
Katharine Van Winkle Palmer. Nautilus, vol. 36, no. 4, pp. 141-142. April.

[Photograph of Vostoshni Rookery, St. Paul Island, by Hanna in] Conservation
of marine life of the Pacific, by Barton W. Evermann. Mid-Pacific Magazine,
vol. 25, no. 4, p. 304. April. (Also published in Catalina Islander, vol. 10, no. 16,
pp. 1-2. May 2.)

[Review of] Fauna from the Eocene of Washington, by Charles E. Weaver and
Katharine Van Winkle Palmer. Journal of Geology, vol. 31, no. 3, pp. 261-263.
April—May.

[Comments, field notes, photographs, and other material contributed by Hanna
in] A biological survey of the Pribilof Islands, Alaska, by Edward A. Preble,
W. L. McAttee, et al. United States Department of Agriculture, Bureau of Bio-
logical Survey, North American Fauna no. 46, pp. 1-257, pls. 1-15. June 20.

[Data and photograph furnished by Hanna for article] Uncle Sam’s seal farm,
by Louise Meilike. Oakland (California) Tribune Magazine, p. 11. May 20.

VOL

140.

141.
142.

143.

144.

145.
146.

147.

148.

149.

150.
151.
152.
153.
154.

155.

156.

157.

. XXXII] MILLER: G DALLAS HANNA 23

[Map of Gulf of California showing expedition route furnished by Hanna for]
Expedition of the California Academy of Sciences to the Gulf of California in
1921—General Account, by Joseph R. Slevin. Proceedings of the California
Academy of Sciences, ser. 4, vol. 12, no. 6, map opposite p. 72. June 2.

[Senior author with A. W. Anthony of] A cruise among desert islands. National
Geographic Magazine, vol. 44, no. 1, pp. 70-99, 33 illustrations. July.

Note on Lymnaea hemphilliana (Baker). Nautilus, vol. 37, no. 1, pp. 23-25.
July.

Results of preliminary examination of seven samples of sediments from near
Lomita. Bulletin of the Southern California Academy of Sciences, vol. 22. pt.
2, p. 64. July.

Pleistocene fresh-water mollusks from north central Texas. Nautilus, vol. 37,
no. 1, pp. 25-26. July.

[Annual report of the] Department of Invertebrate Paleontology. In Report of
the Director of the Museum for the year 1922. Proceedings of the California
Academy of Sciences, ser. 4, vol. 11, no. 23, pp. 680-682. August 22.

Some Eocene foraminifera near Vacaville, California. University of California
Publications, Bulletin of the Department of Geological Sciences, vol. 14, no. 9,
pp. 319-328, pls. 58-59. October 31.

Rare mammals of the Pribilof Islands, Alaska. Journal of Mammalogy, vol. 4,
no. 4, pp. 209-215, pl. 23. November.

[Review of] North American later Tertiary and Quaternary Bryozoa, by Ferdi-
nand Canu and Ray S. Bassler. Journal of Geology, vol. 31, no. 8, pp. 686-687.
November—December.

Expedition of the California Academy of Sciences to the Gulf of California in
1921—Land and freshwater mollusks. Proceedings of the California Academy
of Sciences, ser. 4, vol. 12, no. 26, pp. 483-527, pls. 7-11. December 31.

1924

Succinea avara Say, from the tar pits of California. Nautilus, vol. 37, no. 3, p.
106. January.

[Review of] Mollusca of the southwestern states, by Henry A. Pilsbry and
James H. Ferriss. Nautilus, vol. 37, no. 3, pp. 107-108.

[Note on] Sperm whales at St. George Island, Bering Sea. Journal of Mammal-
ogy, vol. 5, no. 1, p. 64. February.

Temperature records of Alaska fur seals. American Journal of Physiology, vol.
68, no. 1, pp. 52-53. March.

Fresh-water mollusks of Eagle Lake, California. Proceedings of the California
Academy of Sciences, ser. 4, vol. 13, no. 7, pp. 131-136, pl. 1. March 18.

[Senior author with E. G. Gaylord of] Description of a new genus and species
of fresh-water gastropod mollusk (Scalez petrolia) from the Etchegoin Pliocene
of California. Proceedings of the California Academy of Sciences, ser. 4, vol. 13,
no. 9, pp. 147-149, figs. 1-2. March 18.

Rectifications of nomenclature. Proceedings of the California Academy of Sci-
ences, ser. 4, vol. 13, no. 10, pp. 151-186. March 18.

Smaller foraminifera for stratigraphy. Bulletin of the American Association
of Petroleum Geologists, vol. 8, no. 2, pp. 246-250. March—April.

158.

159.

160.

163.

164.

165.

166.

167.

168.

169.

CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER.

[Senior author with Mary E. McLellan of] A new species of whale from the
type locality of the Monterey Group. Proceedings of the California Academy
of Sciences, ser. 4, vol. 13, no. 14, pp. 237-241, pls. 5-9. June 14.

Insects in the California tar traps. Science, new ser., vol. 59, no. 1538, p. 555.
June 20.

[Notice of the] Resignation of A. H. Proctor. United States Department of
Commerce, Fisheries Service Bulletin, no. 111, pp. 2-3. August 1.

A little about diatoms, with photographs by W. M. Grant. The Record (Associ-
ated Oil Company, San Francisco), vol. 5, no. 9, pp. 6, 10-11, 10 figs. September.
(Also published in California Engineer, University of California, Berkeley, vol.
3, no. 4, pp. 107-108, 8 figs. December. )

[Senior author with H. L. Driver of] The study of subsurface formations in
California oil field development. Summary of Operations, California Oil Fields,
California State Mining Bureau, vol. 10, no. 3, pp. 5—26, figs. 1-10. September.

[Senior author with Marcus A. Hanna of] Foraminifera from the Eocene of
Cowlitz River, Lewis County, Washington. University of Washington Publi-
cations in Geology, vol. 1, no. 4, pp. 57—63, pl. 13. October.

[Note on] Gigantic amphibian from Kansas Coal Measures. Pan-American
Geologist, vol. 42, no. 3, pp. 235-236. October 10.

[Annual report of the] Department of Invertebrate Paleontology. Zn Report of
the Director of the Museum for the year 1923. Proceedings of the California
Academy of Sciences, ser. 4, vol. 12, no. 33, pp. 1264-1265. October 10.

1925
Miocene marine vertebrates in Kern County, California. Science, new ser., vol.
61, no. 1568, pp. 71-72. January 16.

[Coauthor with Frank M. Anderson of] Fauna and stratigraphic relations of
the Tejon Eocene at the type locality in Kern County, California. Occasional
Papers of the California Academy of Sciences, vol. 11, pp. 1-249, pls. 1-16.
March 18.

[ Discussion of] Diatom theory of origin of petroleum in California, by Jun-ichi
Takahashi [Sendai, Japan]. Bulletin of the Geological Society of America, vol.
36, no. 1, p. 207. March 30.

[Coauthor with E. G. Gaylord of] Correlation of organic shales in the southern
end of the San Joaquin Valley, California. Bulletin of the American Associa-
tion of Petroleum Geologists, vol. 9, no. 2, pp. 228-234, pls. 4-5. Marech—April.

Expedition of the California Academy of Sciences to the Revillagigedo Islands.
Science, new ser., vol. 61, no. 1579, pp. 359-360. April 3.

Naturalists to explore strange islands off Mexico. [Article in] San Francisco
Examiner (Sunday), p. K7, 4 illustrations. April 12.

Some land shells from the Aleutian Islands, Alaska. Nautilus, vol. 38, no. 4, pp.
122-125. April.

[Senior author with H. Walton Clark of] Lymnaea auricularia (Linn.) in Cali-
fornia. Nautilus, vol. 38, no. 4, pp. 125-127. April.

[Report of the] Department of Invertebrate Paleontology. In Report of the
Director for the year 1924. Proceedings of the California Academy of Sciences,
ser. 4, vol. 13, no. 28, pp. 476-478. May 29.

VOL.

175.

183.

184.

185.

186.

187.

188.

189.

190.

isl

192.

193.

XXXII] MILLER: G DALLAS HANNA 25

[Coauthor with Barton W. Evermann of] The Steller sea lion rookery on Afio
Nuevo Island, California, in 1924. Journal of Mammalogy, vol. 6, no. 2, pp. 96—
99, pls. 8-10. May.

Scientists return with rarities from islands off Mexico. [Article in] San Fran-
cisco Examiner (Sunday), p. K3, 10 illustrations. June 28.

[Senior author with EK. G. Gaylord of] Organic shales in southern San Joaquin
Valley. Petroleum World (Los Angeles), vol. 10, pp. 66, 68. June.

[Senior author with Merle C. Israelsky of] Contribution to the paleontology of
Peru. Proceedings of the California Academy of Sciences, ser. 4, vol. 14, no. 2,
pp. 37-75, pls. 7-8. July 21.

[Photograph supplied by Hanna of fur-seal census being taken on the Pribilof
Islands]. Popular Mechanics, vol. 44, no. 1, p. 87. July.

Zalophancylus, a fish vertebra, not a mollusk. Nautilus, vol. 39, no. 1, pp. 18-19.
July.

The extraction of fossils from refractory rocks. Journal of Geology, vol. 33, no.
5, pp. 545-547. July-August.

Expedition to Guadalupe Island, Mexico, in 1922—General Report. Proceedings
of the California Academy of Sciences, ser. 4, vol. 14, no. 12, pp. 217—275, pls.
15-19, figs. 1-2. September 5.

Tertic expanse of North American continent in Pacific Ocean. Pan-American
Geologist, vol. 44, no. 2, p. 147. September.

The age and correlation of the Kreyenhagen shale in California. Bulletin of the
American Association of Petroleum Geologists, vol. 9, no. 6, pp. 990-999.
September.

[Senior author with H. Walton Clark of] Additional records for Lymnaea
auricularia. Nautilus, vol. 39, no. 2, p. 71. October.

Was there a Pacific continent? Science, new ser., vol. 57, no. 1613, pp. 491—492.
November 27.

Data on the age and correlation of the Kreyenhagen shale in Fresno County,
California. Petroleum Age, vol. 10, pp. 60, 78-80. December.

1926
Paleontology of Coyote Mountain, Imperial County, California. Proceedings of

the California Academy of Sciences, ser. 4, vol. 14, no. 18, pp. 427-503, pls. 20-29,
fig. 1. March 23.

Expedition to the Revillagigedo Islands, Mexico, in 1925—General report. Pro-
ceedings of the California Academy of Sciences, ser. 4, vol. 15, no. 1, pp. 1-113,
pls. 1-10, figs. 1-7. March 30.

Microscopical research in California petroleum fields. The Oil and Gas Journal
(Tulsa, Oklahoma), vol. 24, no. 45, p. 96. April 1.

[Senior author with Wm. M. Grant of] Expedition to the Revillagigedo Islands,
Mexico, in 1925, (11)—-Miocene marine diatoms from Maria Madre Island, Mex-
ico. Proceedings of the California Academy of Sciences, ser. 4, vol. 15, no. 2, pp.
115-193, pls. 11-21, fig. 1. April 16.

[Annual report of the] Department of Invertebrate Paleontology. Jn Report of
the Director for the year 1925. Proceedings of the California Academy of Sci-
ences, ser. 4, vol. 14, no. 20, pp. 546-547. April 28.

Further notes on Scalez petrolia. Nautilus, vol. 40, no. 1, pp. 14-16. July.

203.

204.

205.

207.

208.

209.

210.

211.

CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER.

[Coauthor with J. A. Taff of] Notes on the age and correlation of the Moreno
shale. Bulletin of the American Association of Petroleum Geologists, vol. 10,
no. 8, pp. 812-814. August.

Concerning Shark-Tooth Hill. Standard Oil Bulletin, vol. 14, no. 4, pp. 12, 16, 2
figs. August.

Grizzly bear specimen at San Francisco Academy. [Article in] San Francisco
Examiner (Sunday), p. N5. November 21.

Disintegrating shales by mechanical attrition. Micropaleontology Bulletin, vol.
1, no. 3, pp. 1-2. December 15.

[Abstract of] Untersuchungen itiber den Bau der Diatomeen, by Friedrich
Hustedt. Biological Abstracts, vol. 1, no. 1, p. 143. December.

Abstract of no. 189. Biological Abstracts, vol. 1, no. 1, p. 162. December.

US PAT
“AFS”, a new resin of high refractive index for mounting microscopic objects.
Science, new ser., vol. 65, no. 1672, pp. 41-42. January 14.

The photography of small objects. Transactions of the American Microscopical
Society, vol. 46, no. 1, pp. 15-25. January.

[Notes on microscopic fossils in] The marine Miocene deposits of North Colum-
bia, by Frank M. Anderson. Proceedings of the California Academy of Sciences,
ser. 4, vol. 16, no. 3, p. 88. January 31.

[Photographs of Arizona Cretaceous fossils supplied by Hanna for] Fossils
from the Mancos-Dakota-Tununk formation in the vicinity of Steamboat, Ari-
zona, by Albert B. Reagan. Proceedings of the Indiana Academy of Sciences,
1926, vol. 36, pp. 119-127, figs. 1-26. 1927.

[Annual report of the] Department of Invertebrate Paleontology. Jn Report of
the Director for the year 1926. Proceedings of the California Academy of Sci-
ences, ser. 4, vol. 15, no. 18, p. 538. March 31.

[Coauthor with Fred Baker of] Expedition of the California Academy of Sci-
ences to the Gulf of California in 1921—Marine Mollusca of the order Opistho-
branchiata. Proceedings of the California Academy of Sciences, ser. 4, vol. 16,
no. 5, pp. 123-135, pl. 4. April 22.

[Senior author with Leo George Hertlein of] Expedition of the California
Academy of Sciences to the Gulf of California in 1921—-Geology and Paleon-
tology. Proceedings of the California Academy of Sciences, ser. 4, vol. 15, no. 6,
pp: 1387—157, pl. 5. April 22:

[Coauthor with Joseph A. Cushman of] Foraminifera from the Eocene near
Coalinga, California. Proceedings of the California Academy of Sciences, ser. 4,
vol. 16, no. 8, pp. 205-229, pls. 13-14. April 22.

[Coauthor with J. A. Taff of] A geologic section in the center of the San
Joaquin Valley. Proceedings of the California Academy of Sciences, ser. 4, vol.
16, no. 16, pp. 509-515. April 27.

Exact localities of certain California land shells. Nautilus, vol. 40, no. 4, pp.
124-125. April.

A note on the geology of St. Matthew Island, Bering Sea. American Journal
of Science, ser. 5, vol. 13, no. 77, p. 450. May.

Synthetic resin as a mounting medium. Science, new ser., vol. 65, no. 1693, pp.
575-576. June 10.

VOL

212.

213.

214.

215.

216.

217.

218.

225.

226.

227.

228.

229.

230.

231.

. XXXIT] MILLER: G DALLAS HANNA 27

[Abstract of] The fossil diatom deposit at Spokane, by Albert Mann. Biological
Abstracts, vol. 1, no. 4, p. 539. June.

[Photograph of Aulacodiscus Kittoni Arnott supplied by Hanna for] Diatom
“epidemics” at Copalis Beach, Washington, by L. F. Becking, C. F. Tolman, ef al.
Economic Geology, vol. 22, no. 4, p. 360, fig. 2. June—July.

Helminthoglypta traskii (Newcomb) at “Fort Tejon,” Kern County, California.
Nautilus, vol. 41, no. 1, pp. 32-34. July.

[Review of] New specific names for Austral Mollusca, by H. J. Finlay. Nautilus,
vol. 41, no. 1, pp. 34-36. July.
Geology of west Mexican islands. Pan-American Geologist, vol. 48, no. 1, pp.
1-24, pls. 1-8, figs. 1-3. August.

[Abstract of] Diatomeas fossiles del prebelgranese de Mirmar (Prov. de Buenos
Aires), por Joaquin Frenguelli. Biological Abstracts, vol. 1, no. 5, p. 717.
August.

The lowest known Tertiary diatoms in California. Journal of Paleontology,
vol. 1, no. 2, pp. 103-127, pls. 17-21. August.

Cretaceous diatoms from California. Occasional Papers of the California Acad-
emy of Sciences, no. 13, pp. 1-48, pls. 1-5. September 17.

[Senior author with Leo George Hertlein of] Notes on Ostrea californica Mar-
cou. Nautilus, vol. 41, no. 2, pp. 45-47. October.

Abstract of no. 199. Biological Abstracts, vol. 1, no. 6, pp. 774-775. October.
[Abstract of] On the effects of B-radiation on Colpidium colpoda as seen in
stained specimens, by J. C. Mottram. Biological Abstracts, vol. 1, no. 6, p. 819.
October.

Abstract of no. 191. Biological Abstracts, vol. 1, no. 6, p. 953. October.
[Abstract of] Uber die Diatomeen-Assoziationen des Dorfes Szamosfalva bei
Kolozsvar, by B. v. Cholnoky. Biological Abstracts, vol. 1, no. 6, pp. 1005-1006.
October.

[Coauthor with Joaquin Frenguelli of abstract of] Diatomee della valle d’Aosta,
by Piero Bevra Giaj. Biological Abstracts, vol. 1, no. 6, p. 1181. October.
[Coauthor with Joaquin Frenguelli of abstract of] Diatomee raccolte nell ’Orto
Botanico della Universita di Genova, by Piero Bevra Giaj. Biological Abstracts,
vol. 1, no. 6, p. 1181. October.

[Coauthor with Joaquin Frenguelli of abstract of] Ein Beitrag zur Kenntnis
der Diatomeenflora von BOhmen—lI, by E. Sprenger. Biological Abstracts, vol.
1, no. 6, p. 1182. October.

[Abstract of] List of Quaternary and Tertiary Diatomaceae from deposits of
southern Canada, by C. S. Boyer. Biological Abstracts, vol. 1, nos. 7-8, p. 1198.
November—December.

[Abstract of] Biogenesis of hydrocarbons by diatoms, by C. F. Tolman. Bio-
logical Abstracts, vol. 1, nos. 7-8, p. 1200. November—December.

[Senior author with C. C. Church of] A collection of Recent foraminifera taken
off San Francisco Bay, California. Journal of Paleontology, vol. 1, no. 3, pp.
195-202. December.

1928

Silicoflagellata from the Cretaceous of California. Journal of Paleontology, vol.
1, no. 4, pp. 259-2638, pl. 41. January.

239.

240.

250.

CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER.

[Abstract of] Preliminary statement regarding the diatom “epidemics” at
Copalis Beach, Washington, and an analysis of diatom oil, by L. B. Becking,
et al. Biological Abstracts, vol. 2, nos. 1-2, p. 27. January—February.

[Abstract of] Sur la Schmardaella lutzi Michaelsen, by A. Lutz. Biological Ab-
stracts, vol. 2, nos. 1-2, p. 335. January—February.

Abstract of no. 186. Biological Abstracts, vol. 2, nos. 1-2, p. 342. January—
February.

[Abstract of] Diatoms from the dinner table, by Manfred Voigt. Biological
Abstracts, vol. 2, nos. 3-5, p. 632. March—May.

Abstract of no. 228. Biological Abstracts, vol. 2, nos. 3-5, p. 704. March—May.
Means for obtaining data in earth bores. United States Patent Office, Patent
No. 1,665,058, pp. 1-5, 1 pl. Issued April 3.

The riddle of the origin of oil. The Record (Associated Oil Co., San Francisco),
vol. 9, no. 4, pp. 6-7, 3 figs. (one of the author). April.

[Annual report of the] Department of Paleontology. Jn Report of the Director
for the year 1927. Proceedings of the California Academy of Sciences, ser. 4,
vol. 16, no. 24, pp. 745-747. May 22.

An early reference to the theory that diatoms are the source of bituminous
substances. Bulletin of the American Association of Petroleum Geologists, vol.
12, no. 5, pp. 555-556. May.

[Senior author with C. C. Church of] Freezing and thawing to disintegrate
shales. Journal of Paleontology, vol. 2, no. 2, p. 131. June.

[Coauthor with Fred Baker and A. M. Strong of] Some Pyramidellidae from
the Gulf of California. Proceedings of the California Academy of Sciences, ser.
4, vol. 17, no. 7, pp. 205-246, pls. 11-12. June 29.

[Abstract of] Synopsis of North American Diatomaceae, Part I, by Charles S.
Boyer. Biological Abstracts, vol. 2, nos. 6-8, p. 1197. June-August.

[Abstract of] Study of algae distribution, by Samuel Eddy. Biological Ab-
stracts, vol. 2, nos. 6-8, p. 1198. June-August.

[Abstract of] Beitrage zur Kenntnis der Diatomeen, by K. Gemeinhardt. Bio-
logical Abstracts, vol. 2, nos. 6-8, p. 1199. June-August.

[Abstract of] Beitrage zur Kenntnis Osterreichischer Diatomeen und ihrer
Verbreitung, by Rudolph Handmann. Biological Abstracts, vol. 2, nos. 6-8, p.
1200. June-August.

[Abstract of] Origin of California petroleum, by F. M. Anderson. Biological
Abstracts, vol. 2, nos. 6-8, p. 1256. June-August.

The Monterey Shale of California at its type locality with a summary of its
fauna and flora. Bulletin of the American Association of Petroleum Geologists,
vol. 12, no. 10, pp. 969-983, pls. 7-10. October.

[Coauthor with F. M. Anderson of] Solen novacularis, a name for an Eocene
fossil from California. Nautilus, vol. 42, no. 2, pp. 65-66. October.

The age of diatom-bearing shales at Malaga Cove, Los Angeles County, Cali-
fornia. Bulletin of the American Association of Petroleum Geologists, vol. 12,
no. 11, pp. 1109-1111. November.

1929
Mammoth tusks found near Oroville, California. Department of Natural Re-
sources, [California] Division of Mines and Mining, Report 25 of the State
Mineralogist, vol. 25, no. 1, pp. 88-90, 2 figs. January.

VoL

252.

253.

254.

255.

256.

257.

258.

259.

260.

261.

262.

263.

264.

265.

266.

267.

268.

269.

. XXXIT] MILLER: G DALLAS HANNA 29

[Abstract of] Uber Einschlussmittel fiir Diatomeen, by R. W. Kolbe. Biological
Abstracts, vol. 3, nos. 1-3, p. 262. January—March.

[Abstract of] Continuous and discontinuous variations in nature, by Clarence
Tierney. Biological Abstracts, vol. 3, nos. 1-3, p. 264. January—March.

[Abstract of] La composition chimique des substances végétales et des houilles.
Le réle des substances végétales dans la formation de la houille, by Andre
Duparque. Biological Abstracts, vol. 3, nos. 1-3, p. 289. January-March.

[Abstract of] La structure microscopique des lignites. Comparison avec la
structure miscroscopique de la houille, by Andre Duparque. Biological Ab-
stracts, vol. 3, nos. 1-3, pp. 289-290. . January—March.

[Abstract of] Le role des tissues lignifiés dans la formation de la houille, by
Andre Duparque. Biological Abstracts, vol. 3, nos. 1-3, p. 290. January—March.

[Abstract of] Remarques sur la nature des quatre constituants macroscopiques
de la houille, by Andre Duparque. Biological Abstracts, vol. 3, nos. 1-3, p. 290.
January—March.

[Two photographs on plate 1 and quoted notes from Hanna in] Life history of
the sea lions on the California coast, by John Rowley. Journal of Mammalogy,
vol. 10, no. 1, pp. 1-36, pls. 1-3. February.
[Senior author with William M. Grant of] Brackish-water Pliocene diatoms
from the Etchegoin formation of central California. Journal of Paleontology,
vol. 3, no. 1, pp. 87-100, pls. 11-14. March.

[Notes on microscopic fossils by Hanna in] Marine Miocene and related de-
posits of north Columbia, by Frank M. Anderson. Proceedings of the California
Academy of Sciences, ser. 4, vol. 18, no. 4, p. 89. March 29.

A new species of land snail from Kern County, California. Proceedings of the
California Academy of Sciences, ser. 4, vol. 18, no. 6, pp. 217-218, pl. 24, figs.
7-9. April 5.

[Senior author with Leo George Hertlein of] A new species of land snail from
Coahuila, Mexico. Proceedings of the California Academy of Sciences, ser. 4,
vol. 18, no. 7, pp. 219-220, pl. 24, figs. 5-6. April 5.

[Report on work relating to diatom collections] Jn Annual Report for 1928-29,
National Research Council, Division of Geology and Geography, Committee on
Paleobotany, Appendix J, p. 5. Washington, D.C., April 27.

[Abstract of] Zur Okologie, Morphologie und Systematik der Brackwasser-
Diatomeen, by R. W. Kolbe. Biological Abstracts, vol. 3, nos. 4-6, p. 879. April—
June.

[Abstract of] El gigantismo en los protozoarios, by Alicia E. Reyes. Biological
Abstracts, vol. 3, nos. 4-6, p. 951. April—June.

[Annual report of the] Department of Paleontology. In Report of the Director
of the Museum for the year 1928. Proceedings of the California Academy of
Sciences, ser. 4, vol. 17, no. 12, pp. 331-333. May 22.

Another synthetic resin useful in microscopy. Science, new ser., vol. 70, no.
1801, pp. 16-17. July 5.

[Abstract of] Baltiska tidsbestamningar och baltisk tidsindelning vid Kalmar-
sund, by H. Thomasson. Biological Abstracts, vol. 3, nos. 7-8, p. 1483. July—
August.

[Abstract of] Observaciones sobre el macronticleo de Chilodon wncinatus Ehrb.,
by E. Fernandez Galiano. Biological Abstracts, vol. 3, nos. 9-10, p. 2074. Sep-
tember—November.

270.

274.

279.

CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER.

Fossil diatoms dredged from Bering Sea. Transactions of the San Diego So-
ciety of Natural History, vol. 5, no. 20, pp. 287-296, pl. 34. December 31. (Ab-
stract in Pan-American Geologist, vol. 52, no. 2, p. 160. September.)

1930
Pyrgulopsis nevadensis (Stearns) in Oregon. Nautilus, vol. 43, no. 3, pp. 103-
104. January.

[Seven figures of diatoms from photographs by Hanna in] Kettleman Hills Oil
Field—Part II, by J. A. Taff. Record (Associated Oil Company, San Francisco),
vol. 11, no. 2, p. 9, 7 figs. February.

[Annual report of the] Department of Paleontology. Jn Report of the Director
of the Museum and of the Aquarium for the year 1929. Proceedings of the Cali-
fornia Academy of Sciences, ser. 4, vol. 18, no. 18, pp. 558-560. April 8.

[Abstract of] Distinctions among certain genera of larger Foraminifera for the
field geologist of the East Indies, by I. M. van der Vlerk and R. E. Dickerson.
Biological Abstracts, vol. 4, no. 4, p. 1257. April.

[ Abstract of] Landshells of the Galapagos Islands, by William Healey Dall and
Washington Henry Ochsner. Biological Abstracts, vol. 4, no. 4, p. 1267. April.

[Report on diatom work] Jn Annual Report for 1929-30, National Research
Council, Division of Geology and Geography, Committee on Paleobotany, Ap-
pendix P, pp. 5, 7-8, 9. Washington, D.C., May 3.

[Abstract of] Fresh-water diatoms from Ancoy, south China, by B. W. Skvort-
zow. Biological Abstracts, vol. 4, no. 5, pp. 1524-1525. May.

[Coauthor with A. M. Strong of] Marine Mollusca of Guadalupe Island, Mexico.
Proceedings of the California Academy of Sciences, ser. 4, vol. 19, no. 1, pp. 1-6.
June 4.

[Coauthor with A. M. Strong of] Marine Mollusca of the Revillagigedo Islands,
Mexico. Proceedings of the California Academy of Sciences, ser. 4, vol. 19, no. 2,
pp. 7-12. June 4.

[Coauthor with A. M. Strong of] Marine Mollusca of the Tres Marias Islands,
Mexico. Proceedings of the California Academy of Sciences, ser. 4, vol. 19, no. 3,
pp. 13-22. June 4.

A review of the genus Rouvia. Journal of Paleontology, vol. 4, no. 2, pp. 179-
188, pl. 14, figs. 1-8. June.

Observations on Lithodesmium cornigerum Brun. Journal of Paleontology, vol.
4, no. 2, pp. 189-191, pl. 14, figs. 9-10. June.

The growth of Omphalotheca. Journal of Paleontology, vol. 4, no. 2, p. 192, pl.
14, fig. 11. June.

[Review of] Premiéres observations microscopiques sur le schiste bitumineux
du volgien inférieur, by M. D. Zalessky. Journal of Paleontology, vol. 4, no. 2,
pp. 202-203. June.

Porosity of diatomite. Engineering and Mining Journal, vol. 130, no. 1, pp. 7-8,
figs. 1-7. July 10.

[Coauthor with Fred Baker and A. M. Strong of] Some Rissoid Mollusca from
the Gulf of California. Proceedings of the California Academy of Sciences, ser.
4, vol. 19, no. 4, pp. 28-56, pl. 1, figs. 1-4. July 15.

[Coauthor with Fred Baker and A. M. Strong of] Some Mollusca of the family
Hpitoniidae from the Gulf of California. Proceedings of the California Academy
of Sciences, ser. 4, vol. 19, no. 5, pp. 41-56, pls. 2-3. July 15.

VoL. XXXIT] MILLER: G DALLAS HANNA 31

288.

289.

290.

291.

292.

293.

294.

295.

296.

297.

298.

299.

300.

301,
302.

303.

3804.

305.

306.

307.

Geology of Sharktooth Hill, Kern County, California. Proceedings of the Cali-
fornia Academy of Sciences, ser. 4, vol. 19, no. 7, pp. 65—83, figs. 1-3. July 15.
[Senior author with J. L. Nicholson of] Rediscovery of Polygyra roperi Pilsbry.
Nautilus, vol. 44, no. 1, pp. 17-18. July.

Silicoflagellates from the Cantua shale. Pan-American Geologist, vol. 54, no. 1,
pp. 79-80. August.

[Review of] Diatoms from the Cantua shale. Pan-American Geologist, vol. 54,
no. 1, p. 80. August.

The dates of publication of Tempére and Peragallo’s ““Diatomées du Monde
Entier, edition 2.” Journal of Paleontology, vol. 4, no. 3, pp. 296-297.
September.

Hyrax, a new mounting medium for diatoms. Journal of the Royal Microscopi-
cal Society, ser. 3, vol. 50, pt. 4, no. 20, pp. 424-426. December.

Remains of Holothuroidea from the Carboniferous of Kansas. Journal of
Paleontology, vol. 4, no. 4, pp. 413-414, pl. 40, figs. 1-7. December.

A new genus of Silicoflagellata from the Miocene of Lower California. Journal
of Paleontology, vol. 4, no. 4, pp. 415-416, pl. 40, figs. 8-18. December.

1931
[Abstract of] West American Mollusca of the genus Phasianella, by A. M.
Strong. Biological Abstracts, vol. 5, no. 2, p. 645. February.
[Abstract of] Notes on Lower Tertiary deposits of Columbia and their mollus-
can and foraminiferal fauna, by F. M. Anderson. Biological Abstracts, vol. 5,
no. 2, p. 637. February.
[Abstract of] Silicoflagellates from the Lillis shale. Bulletin of the Geological
Society of America, vol. 42, no. 1, p. 306. March 31.
[Abstract of] Diatoms from the Lillis shale. Bulletin of the Geological Society
of America, vol. 42, no. 1, pp. 306-307. March 31.
Minutes of the meeting of the Pacific coast branch of the Paleontological So-
ciety. Bulletin of the Geological Society of America, vol. 42, no. 1, pp. 365-371.
March 31.
Illustrating fossils. Journal of Paleontology, vol. 5, no. 1, pp. 49-68, pl. 8. March.
Diatoms and silicoflagellates of the Kreyenhagen shale. California Division of
Mines, Report 27 of the State Minerologist, vol. 27, no. 2, pp. 187-201, pls. A-E.
April.
[Senior author with C. C. Church of] David Dwight Baldwin, 1831-1912. In
Report of the Director of the Museum and of the Aquarium for the year 1930.
Proceedings of the California Academy of Sciences, ser. 4, vol. 19, no. 14, pp.
431-436. May 29.
[Annual report of the] Department of Paleontology. Jn Report of the Director
of the Museum and of the Aquarium for the year 1930. Proceedings of the Cali-
fornia Academy of Sciences, ser. 4, vol. 19, no. 14, pp. 447-448. May 29.
[Senior author with Allyn G. Smith of] Notes on Acmaea funiculata (Carpen-
ter). Nautilus, vol. 45, no. 1, pp. 21—25, pl. 2. July.
[Review of] Some notes on liver-fluke investigation, by J. N. Shaw. Nautilus,
vol. 45, no. 1, pp. 34-35. July.
[Review of two papers on] Salmon poisoning, by B. T. Simms, et al. Nautilus,
vols 45; nol 1, p. 35. July.

310.

311.

312.

313.

316.

321.

322.

323.

324.

CALIFORNIA ACADEMY OF SCIENCES [ Proc, 4TH SER.

Lichtenstein on California birds. Condor, vol. 33, no. 5, pp. 211-213. September—
October.

[Senior author with W. M. Grant of] Diatoms of Pyramid Lake, Nevada. Trans-
actions of the American Microscopical Society, vol. 50, no. 4, 281-297, pls. 25-27.
October.

The adjustment of low power binocular microscopes. Micropaleontology Bul-
letin, vol. 3, no. 1, pp. 1-3, figs. 1-2. December 15.

1932
[Four photographs of micro-fossils by Hanna in] It’s fun to hunt fossils, by J.
Stephens Sullivan. Sunset Magazine, vol. 68, no. 1. January.

The diatoms of Sharktooth Hill, Kern County, California. Proceedings of the
California Academy of Sciences, ser. 4, vol. 20, no. 6, pp. 161—263, pls. 2-18.
January 8.

Minutes of the meeting of the Pacific coast branch of the Paleontological So-
ciety. Bulletin of the Geological Society of America, vol. 43, no. 1, pp. 287-292.
March.

[Abstract of paper delivered at annual meeting of Pacific coast branch of
Paleontological Society] Optical properties of Hyrax. Bulletin of the Geological
Society of America, vol. 43, no. 1, p. 291. March.

[Abstract of paper delivered at annual meeting of Pacific coast branch of Pale-
ontological Society] Desmostylus tooth dredged in Monterey Bay. Bulletin of
the Geological Society of America, vol. 43, no. 1, p. 291. March. (Also in Pan-
American Geologist, vol. 56, no. 1, p. 71. August, 1931.)

[Six photographs by Hanna in] Observations on the head markings of the
golden-crowned sparrow, by Joseph Mailliard. Condor, vol. 34, no. 2, pp. 66—70,
fig. 9. March—April.

[Report on diatom work] /n Annual Report for 1931-32, National Research
Council, Division of Geology and Geography, Committee on Paleobotany, Ap-
pendix P, pp. 5, 6, 9. Washington, D.C. April 23.

A protest against the use of ‘“Navicula”’ for a group of Mollusca. Nautilus, vol.
45, no. 4, pp. 118-120. April.

[Notes on diatoms and seven figures by Hanna in] Earth History, by Luther C.
Snyder. The Century Company. August.

Barton Warren Evermann, 1853-1932. Science, new ser., vol. 76, no. 1971, pp.
317-318. October 7. (Also published in Copeia, 1932, no. 4, pp. 160-162, portrait.
December 31.)

Pliocene diatoms of Wallace County, Kansas. University of Kansas Science
Bulletin, vol. 20, pt. 2, no. 21, pp. 369-394, pls. 31-34. May 15.

The Templeton Crocker Expedition of the California Academy of Sciences.
Science, new ser., vol. 76, no. 1974, pp. 375-377. October 28.

[Senior author with A. G. Smith of] Location of the collection of shells as-
sembled by Rev. J. Rowell. Nautilus, vol. 46, no. 2, p. 70. October.

[Photographs by Hanna of fossils on plates 2 and 3 in] Geology of Santa Cruz
Island, Santa Barbara County, California, by Carl St. J. Bremner. Santa Bar-
bara Museum of Natural History, Occasional Papers, no. 1, pp. 1-33, pls. 1-4.
November 1.

VOL.

325.

326.

327.

328.

329.

330.

331.

332.

333.

334,

335.

336.

337.

338.

339.

XXXIT] MILLER: G DALLAS HANNA

we
Co

193:
[Senior author with Allyn G. Smith of] Two new species of Monadenia from
northern California. Nautilus, vol. 46, no. 3, pp. 79-86, pl. 5 and pl. 6, figs. 8—9.
January.
The name “Lillis formation” in California geology. Bulletin of the American
Assocation of Petroleum Geologists, vol. 17, no. 1, pp. 81-84. January.

Minutes of the meeting of the Pacific coast branch of the Paleontological So-
ciety, Stanford University, California, Saturday, April 9, 1932. Bulletin of the
Geological Society of America, vol. 44, pt. 1, pp. 216-220. February 28.

Utilization of diatoms in western North America. Jn Annual Report for 1932-
33, National Research Council, Division of Geology and Geography, Committee
on Micropaleontology, Exhibit C to Appendix C, pp. 1-2. Washington D.C.
April 22.

[Report on diatom work and on card index of 8000 references on diatom litera-
ture published since the appearance of the 1892 catalogue, together with an
index of the genera of diatoms, both available to students.] Jn Annual Report
for 1932-33, National Research Council, Division of Geology and Geography,
Committee on Paleobotany, Appendix D, pp. 3, 4, 5, 8, 13, 14. Washington, D.C.
April 22.

A new outbreak of Helix pisana in California. Nautilus, vol. 46, no. 4, pp. 139—
140. April.

[Senior author with Achille Forti of] Contribuzioni diatomologiche—XIV—
Schulziella. Atti del Reale Instituto Veneto di Scienze, Lettere ed Arti, tom. 92,
pt. 2, pp. 1277-1282, 1 pl. August 28.

Fresh-water diatoms from Oregon. Jn The Trout Creek flora of southeastern
Oregon, by Harry D. MacGinitie. Carnegie Institution of Washington, publ. no.
416, pp. 43-45. October 26.

The dates of publication of ‘Synopsis des Diatomées de Belgique,” by Henri
Van Heurck. Annals and Magazine of Natural History, ser. 10, vol. 12, no. 70,
pp. 442-443. October.

[Review of] An index to the genera and species of the Diatomaceae and their
synonyms, 1816-1932, Part 1, by Frederick Wm. Mills. Transactions of the
American Microscopical Society, vol. 52, no. 4, p. 363. October.

[Abstract of] Diatomeen des Faulschlammes von Kostenthal in Oberschlesien,
by V. Torka. Biological Abstracts, vol. 7, no. 9, p. 2231. November.

[Coauthor with A. M. Strong and L. G. Hertlein of] The Templeton Crocker
Expedition of the California Academy of Sciences, 1932, No. 10—Marine Mol-
lusca from Acapulco, Mexico, with notes on other species. Proceedings of the
California Academy of Sciences, ser. 4, vol. 21, no. 10, pp. 117-130, pls. 5-6.
December 21.

[Note on] Diatomite from Turtle Bay. Jn Additions to the Pliocene fauna of
Turtle Bay, Lower California, with a note on the Miocene diatomite, by L. G.
Hertlein. Journal of Paleontology, vol. 7, no. 4, pp. 440-441. December.

[Note on] Preservation of type specimens. Jn letter quoted by L. W. Stephen-
son. Journal of Paleontology, vol. 7, no. 4, pp. 442-443. December.
[Arrangement and photography of invertebrate fossils in figure 58 (with C. C.
Church) in] Geology of California, by Ralph D. Reed. American Association of
Petroleum Geologists, fig. 58 (opposite p. 288). Tulsa, Oklahoma, 1933. (Re-
printed 1936, 1951).

341.

343.

344.

355.

CALIFORNIA ACADEMY OF SCIENCES [ Proc. 47H SER.

Diatoms of the Florida peat deposits. 23rd—24th Annual Report of the Florida
State Geological Survey, pp. 65-119, pls. 1-11.

1934

[Abstract of] An index to the genera and species of the Diatomaceae and their
synonyms, 1816-1932, Parts 1 and 2, by Frederick Wm. Mills. Biological Ab-
stracts, vol. 8, no. 2, p. 485. February.

Use of fossil diatoms for geological correlation. Jn Annual Report for 1933-34,
National Research Council, Division of Geology and Geography, Committee on
Micropaleontology, Exhibit E to Appendix A, pp. 18-19. Washington, D.C.,
April 28.

[Work on fossil diatoms] In Annual Report for 1933-34, National Research
Council, Division of Geology and Geography, Committee on Paleobotany, Appen-
dix, pp. 6, 8, 15. Washington, D.C., April 28.

[Review of] An index to the genera and species of the Diatomaceae and their
synonyms, Parts 2-9, by Frederick Wm. Mills. Transactions of the American
Microscopical Society, vol. 53, no. 2, p. 200. April.

Carl Ewald Grunsky [Obituary]. Science, new ser., vol. 79, no. 2060, p. 556.
June 22.

Additional notes on diatoms from the Cretaceous of California. Journal of
Paleontology, vol. 8, no. 3, pp. 352-355, pl. 48. September.

Land shells from the upper Eocene Sespe deposits, California. Journal of the
Washington Academy of Sciences, vol. 24, no. 12, pp. 539-541, figs. 1—3a.
December 15.

1935

[Senior author with W. M. Grant of] Lunar rings. Science, new ser., vol. 81,
no. 2088, pp. 18-19. January 4.

Senior author with Leo G. Hertlein of] Longevity of Mitra in captivity. Nauti-
lus, vol. 48, no. 3, pp. 90-91. January.

[Report on work in micropaleontology in California during the year 1934]. In
Annual Report for 1934-35, National Research Council, Division of Geology and
Geography, Committee on Micropaleontology, Appendix C, pp. 5-6. Washington,
IDACE, ANoreatll 7,

[Report of work on diatoms] Jn Annual Report for 1934-35, National Research
Council, Division of Geology and Geography, Committee on Paleobotany, Ap-
pendix H, pp. 7, 18. Washington, D.C., April 27.

[Coauthor with J. A. Taff and C. M. Cross of] Chico Cretacic at type locality
[Abstract]. Pan-American Geologist, vol. 64, p. 72. August.

[Abstract of] Interesting whale jaw from Kern County, California. Pan-Ameri-
can Geologist, vol. 64, pp. 79-80. August. (Also reviewed in Proceedings of the
Geological Society of America, 1935, p. 419. June, 1936.)

[Review of] An index to the genera and species of Diatomaceae and their
synonyms, by Frederick W. Mills. Science, new ser., vol. 82, no. 2114, p. 14.
July 5.

[Coauthor with Frank M. Anderson of] Cretaceous geology of Lower California.

Proceedings of the California Academy of Sciences, ser. 4, vol. 23, no. 1, pp.
1—34, pls. 1-11, figs. 1-2. December 23.

VoL. XXXIT] MILLER: G DALLAS HANNA 35

356.

356a.

357.

358.

359.

361.

363.

364.
365.

366.

367.

368.

1936
[Coauthor with M. Vonsen of] Borax Lake, California. California Journal of
Mines and Geology, Report of the State Mineralogist, pp. 99-108, figs. 1-5.
January.

[Report of work on diatoms] /n Annual Report for 1955-36, National Research
Council, Division of Geology and Geography, Committee on Paleobotany, Ap-
pendix D, p. 16. Washington, D.C., May 2.

Notes on fossils from the Tampico region. In Geology of the Tampico region,
Mexico, by John M. Muir. American Association of Petroleum Geologists, pp.
122, 124,135. June.

Review with J. A. Taff and C. C. Church of no. 352. Proceedings of the Geologi-
cal Society of America, 1935, pp. 348-349. June, 1936.

[Photograph of Clestes cuneatus Cope, a ganoid fish from the o6litic limestones,
Ephraim, Utah, supplied by Hanna for] A study of Utah fossil fishes with de-
seription of a new genus and species, by Vasco M. Tanner. Proceedings of the
Utah Academy of Science, Arts and Letters, vol. 13, pp. 88-89, pl. 2. July.

A new land shell from the Eocene of California. Journal of Paleontology, vol.
10, no. 5, pp. 416-417, fig. 1. July.

[Photograph of Steinhart Aquarium supplied by Hanna for] Das Aquarium,
vol. 10, p. 125. Berlin, July.

Permanent preservation of small zoological specimens. Science, new ser., vol.
84, no. 2186, pp. 463-464, fig. 1. November 20. [Describes a method of perma-
nent sealing in glass tubes, with alcohol or formalin. ]

Notes on the localities of fossil diatoms in California. Bulletin de la Société
Francaise de Microscopie, vol. 5, no. 3, pp. 109-111.

1937
Abstract of no. 360. Biological Abstracts, vol. 11, no. 4, p. 1050. April.

[Report of work on diatoms] /n Annual Report for 1936-37, National Research
Council, Division of Geology and Geography, Committee on Paleobotany, Ap-
pendix C, pp. 12, 13, 19, 19-20. Washington, D.C., May 1.

Report on diatoms. Jn Annual Report for 1936-37, National Research Council,
Division of Geology and Geography, Committee on Micropaleontology, Exhibit
A to Appendix E, pp. 10-11. Washington, D.C., May 1.

[Senior author with Allyn G. Smith of] Some notes on an old race of California
land snail with descriptions of three new forms. Nautilus, vol. 51, no. 1, pp.
10-17, pl. 1. July.

[Senior author with A. G. Smith of] The mollusks of Lake Tahoe. California
Fish and Game, vol. 23, no. 3, pp. 244-245. July. (Also published in Nautilus,
vol. 52, no. 1, pp. 34-36. July, 1938.)

[Abstract of] Resti silicei de microorganismi dei travertini della Somalia, by G.
Frenguelli. Biological Abstracts, vol. 11, no. 7, pp. 1814-1815. August—October.
[Senior author with C. C. Church of] Notes on Marginulina vacavillensis
(Hanna). Journal of Paleontology, vol. 11, no. 6, pp. 530-531. September.

An illuminator for opaque objects. Journal of the Royal Microscopical Society,
vol. 57, pp. 11-14, figs. 1-2.

Invertebrates. Jn Methods of Paleontology, by Charles L. Camp and G Dallas
Hanna, part 2, pp. 77-149, figs. 48-58. University of California Press, Berkeley,
1937.

373.

374.

376.

378.

385.

386.

387.

388.

CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER.

1938

[Senior author with Leo George Hertlein of] New Tertiary mollusks from
western North America. Journal of Paleontology, vol. 12, no. 1, pp. 106-110,
pl. 21. January.

[1930-31 index to] Proceedings of the California Academy of Sciences, ser. 4,
vol. 19, pp. 483-496. May 16.

[Coauthor with Fred Baker of] Some Mollusca of the families Cerithiopsidae,
Cerithiidae and Cyclostrematidae from the Gulf of California and adjacent
waters. Proceedings of the California Academy of Sciences, ser. 4, vol. 23, no.
15, pp. 217-244, pls. 17-23. May 24.

[Coauthor with Fred Baker of] Columbellidae from western Mexico. Proceed-
ings of the California Academy of Sciences, ser. 4, vol. 23, no. 16, pp. 245-254,
pl. 24. May 24.

The type locality of “Helix Tudiculata var. Tularensis Hemphill”. Nautilus,
vol. 52, no. 1, pp. 7-9. July.

1939

[Senior author with Leo George Hertlein of] Campanile greenellum, a new
species from the early Eocene of California. Journal of Paleontology, vol. 13,
no. 1, pp. 100-102, figs. 1-2. January.

[Report of work in micropaleontology] Jn Annual Report for 1938-39, National
Research Council, Division of Geology and Geography, Committee on Micropale-
ontology, Appendix D, p. 5. Washington, D.C., April 29.

The Emmet Rixford Collection. Nautilus, vol. 52, no. 4, p. 139. April.

Exotic Mollusea in California. Bulletin of the Department of Agriculture, State
of California, vol. 28, no. 5, pp. 298-321, pls. A-D, figs. 1-2. May.

Extension of range of Tethys californicus Cooper in California. Nautilus, vol.
Mais io, Il, 10, ah Alwihy.

Say’s Conchology. Nautilus, vol. 53, no. 1, pp. 34-35. July.

[Senior author with W. M. Grant of] Preliminary note on a technique for
mounting diatoms in realgar and other substances. Journal of the Royal Micro-
scopical Society, ser. 3, vol. 59, pt. 3, pp. 174-176. September.

[Senior author with Allyn G. Smith of] Notes on some forms of Oreoheliz stri-
gosa (Gould). Proceedings of the California Academy of Sciences, ser. 4, vol.
23, no. 25, pp. 381-392, pls. 33-36. December 29.

1940

The anatomy of Succinea chrysis Westerlund (Mollusca, Gastropoda). Journal
of Morphology, vol. 66, no. 1, pp. 115-129, pls. A-B. January 2. [This is based
on Dr. Hanna’s dissertation submitted in partial fulfillment of the requirements
of George Washington University for the degree of Doctor of Philosophy, which
was actually conferred in 1918. ]

[Note on high vacuum technique for mounting diatoms] /n Annual Report for
1939-40, National Research Council, Division of Geology and Geography, Com-
mittee on Micropaleontology, Appendix B, p. 7. Washington, D.C., April 27.

[Report of work on diatoms] Jn Annual Report for 1939-40, National Research

Council, Division of Geology and Geography, Committee on Paleobotany, Ap-
pendix C, p. 22. Washington, D.C., April 27.

VOL

389.

390.

391.

392.

393.

394.

395.

396.

397.

398.

399.

400.

401.

402.

. XXXIT] MILLER: G DALLAS HANNA 37

[Coauthor with J. A. Taff and C. M. Cross of] Type locality of the Cretaceous
Chico formation. Bulletin of the Geological Society of America, vol. 51, pp.
1311-1327, pls. 1-2, fig. 1. September 1.

[Senior author with W. M. Grant of] Apparatus for mounting diatoms in real-
gar and other substances. Journal of the Royal Microscopical Society, ser. 3,
vol. 60, pp. 152-160, pl. 1, figs. 1-4. September.

[Eight photographs of 1940 operations of the California Division of Fish and
game vessel N. B. Scofield supplied by Hanna for] Fishery research, by G. H.
Clark. California Conservationist, vol. 5, no. 10, p. 5 and back cover. October.

1941

Flying around Mount St. Elias. Women’s City Club Magazine (San Francisco),
vol. 14, no. 12, pp. 14-15, 2 illustrations. January.

[Report on work in micropaleontology] Jn Annual Report for 1940-41, National
Research Council, Division of Geology and Geography, Committee on Micro-
paleontology, Appendix I, p. 7. Washington, D.C., May 3.

[Senior author with Leo George Hertlein of] Characteristic fossils of Califor-
nia. California Division of Mines, Bull. 118, pt. 2, pp. 165-182, with 8 pages of
figures. Pre-print, August; available September 11.

1945

[Review of micropaleontological work in California] Jn Annual Report for
1942-43, National Research Council, Division of Geology and Geography, Com-
mittee on Micropaleontology, Appendix O, pp. 3-4. Washington, D.C., May 1.

1944

[Report on work in micropaleontology] Jn Annual Report for 1943-44, Na-
tional Research Council, Division of Geology and Geography, Committee on
Micropaleontology, Appendix I, p. 5. Washington, D.C., April 29.

1946

[Introduction to] Notes on the geology and general paleontology of the Moreno
shale. In Diatoms of the Moreno shale [California], by John A. Long, ef al.
Journal of Paleontology, vol. 20, no. 2, pp. 82-94, fig. 1. March.

1947

Alaska is a peaceful place. Let’s Get Associated (Tidewater Associated Oil Com-
pany, San Francisco), vol. 6, no. 11, pp. 12-13, 3 illustrations. November.

1948
Comments regarding the McArthur inverted microscope. Journal of the Royal
Microscopical Society, ser. 3, vol. 69, pt. 2, pp. 86-87. May.
Indiscriminate scattering of exotic mollusks. Nautilus, vol. 62, no. 1, pp. 29-31.
July.
[Notes on shellfish poisoning]. Jn the marine mollusks and brachiopods of
Monterey Bay, California, and vicinity, by Allyn G. Smith and Mackenzie
Gordon, Jr. Proceedings of the California Academy of Sciences, ser. 4, vol. 26,
no. 8, pp. 163-164. December 15.
Animals and oil traps. Wasmann Collector, vol. 7, no. 4, pp. 138-139. December.

403.

404.

405.

406.

407.

408.
409.
410.

Ane.

412.

413.

417.

418.

CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER.

1949

[Senior author with A. M. Strong of] West American mollusks of the genus
Conus. Proceedings of the California Academy of Sciences, ser. 4, vol. 26, no. 9,
pp. 247-322, pls. 5-10, figs. 1-4. January 28.

A synthetic resin which has unusual properties. Journal of the Royal Micro-
scopical Society, vol. 69, pp. 25-28. Received March 15.

[Senior author with Leo G. Hertlein of] Two new species of gastropods from
the middle Eocene of California. Journal of Paleontology, vol. 23, no. 4, pp.
392-394, pl. 77, figs. 1-2. July.

1950
Some shells of Clear Lake, northern Modoc County, California. Nautilus, vol.
63, no. 3, pp. 97-99. January.

1951
A new west American nudibranch mollusk. Nautilus, vol. 65, no. 1, pp. 1-3, pl.
2, figs. 1-5. Juliy.
Harold Heath, 1868-1951. Nautilus, vol. 65, no. 1, pp. 28-30. July.
Frank Mace MacFarland, 1869-1951. Nautilus, vol. 65, no. 1, pp. 30-32. July.

Alaska fur seals, Callorhinus ursinus, observed off San Francisco Bay, Califor-
nia. Journal of Mammalogy, vol. 32, no. 3, pp. 364-365. August.

[Figures 1 and 5 from photography by Hanna in] Obsidian. California Division
ot Mines, Mineral Information Service, vol. 4, no. 10, pp. 1—7, figs. 1-5. October 1.
Diatom deposits. Jn Geologic guidebook of the San Francisco Bay counties.
California Division of Mines, Bull. 154, pp. 281-290, figs. 1-3 (35 microphoto-
graphs of diatoms). December.

Geology of the Farallon Islands. In Geologic guidebook of the San Francisco
Bay counties. California Division of Mines, Bull. 154, pp. 301-310, figs. 1-10.
December.

1952
[ Notes on the use of obsidian for optical purposes.] Jn The Amateur Astrono-
mer, by Albert G. Ingalls. Scientific American, vol. 186, no. 2, pp. 84-87.
February.
[Senior author with Olaf P. Jenkins of] The Arvin-Tehachapi earthquake.
Pacific Discovery, vol. 5, no. 5, pp. 28-29, 2 photographs. May—June.
Geology of the continental slope off central California. Proceedings of the Cali-
fornia Academy of Sciences, ser. 4, vol. 27, no. 9, pp. 325-374, pls. 7-14, fig. 1.
July 11.
[Coauthor with A. G. Smith of] A rare species of chiton from the Pioneer Sea-
mount off central California. Proceedings of the California Academy of Sci-
ences, ser. 4, vol. 27, no. 14, pp. 389-392, pl. 20, figs. 6, 10-11. July 11.

1953
Glacier flight, an aerial reconnaissance of the St. Elias Range. Pacific Dis-
covery, vol. 6, no. 3, pp. 18—21, 4 photographs. May—June.
[Fisherman’s luck. Academy News Letter, no. 163, p. 3. July.]

VOL.

420.

421.

422.

423.

424.

425.
426.

427.

428.

429.

433.

434.

435.

XXXIT] MILLER: G@ DALLAS HANNA 39

The overhaul and adjustment of binoculars. Jn Amateur Telescope Making
(Book 3) (A. G. Ingalls, ed.), pp. 218-252, 10 figs. Scientific American, Inc.
Received December 10.

A reflecting collimator for precise measurement of prism angles Jn Amateur
Telescope Making (Book 3) (A. G. Ingalls, ed.), pp. 253-263, 3 figs. Received
December 10.

Making rhomboid prisms. Jn Amateur Telescope Making (Book 3) (A. G.
Ingalls, ed.), pp. 264—268, 2 figs. Received December 10.

Separation of abrasives on a laboratory scale. Jn Amateur Telescope Making
(Book 3) (A. G. Ingalls, ed.), pp. 269-276, 2 figs. Scientific American, Inc. Re-
ceived December 10.

1954
[Senior author with A. G. Smith of] Rediscovery of two California land snails.
Nautilus, vol. 67, no. 3, pp. 69-76, pl. 8, figs. 5-7, text fig. 1. January.

1955
Preparation of Nudibranchiata. Nautilus, vol. 68, no. 3, pp. 105-106. January.
[Reprint of no. 425 in] How to collect shells, p. 72. American Malacological
Union. March 31.
[Photograph of Dr. Hanna. Jn Named director of Arctic project. Academy
News Letter, no. 183, pp. 3-4, photograph on p. 1. March. ]
[Abstract of] Suggestions regarding additional submarine geological work in
the Arctic. Bulletin of the Geological Society of America, vol. 66, no. 12, pt. 2,
pp. 1702-1703. December. (Also abstracted in Proceedings of the 6th and 7th
Alaskan Scientific Conterence, 1955-56, pp. 78-79. 1959.)

1956
[Coauthor with Robert P. Dempster of] Preserving Artemia eggs in high
vacuum. Aquarium Journal (San Francisco), vol. 27, no. 3, 2 pp., 1 illustration.
March.
Land and fresh-water mollusks of the Arctic slope, Alaska. Nautilus, vol. 70,
no. 1, pp. 4-10, pl. 1. July.
Distribution of west American deposits of fossil diatoms. Bios, vol. 27, no. 4,
p. 227-231. December.
[Diatom identifications in] Occurrence of diatomaceous earth near Kenai,
Alaska, by George Plafker. United States Geological Survey, Bulletin 1039—B,
p. 28.

[Annotated bibliography of] Silicoflagellata. In Treatise on Marine Ecology
and Paleoecology, Harry S. Ladd, ed. Geological Society of America, Memoir
67, vol. 2, pp. 745-746. March 25.

Collecting shells in the Arctic. Minutes of the Conchological Club of Southern
California (John Q. Burch, ed.), no. 168, pp. 2-8. June.

1958

[Senior author with Leo George Hertlein of] Marine shells of Middleton Island,
Alaska. Nautilus, vol. 72, no. 3, pp. 78-84, pl. 10. January.

40 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 47H SER.

436. Peach collecting in the Arctic. Veliger (Northern California Malacozoological
Club, Rudolf Stohler, ed.), vol. 1, no. 4, pp. 25-26. April 1.

1961

437. [Senior author with Leo George Hertlein of] Large terebras (Mollusca) from
the eastern Pacific. Proceedings of the California Academy of Sciences, ser. 4,
vol. 30, no. 3, pp. 67-80, pls. 6-7 (in color). August 31.

438. [Preservation of] Nudibranchiata. Jn How to collect shells, 2nd ed., p. 54.
American Malacological Union.

Epiror’s Note: Dr. Hanna’s published writings and other scientific contributions

have been compiled by members of the Academy’s staff from several sources, includ-
ing a current file maintained in the Department of Geology. It has been made as in-
clusive as possible from available material.

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

FOURTH SERIES

G Dallas Hanna Anniversary Volume

Vol. XXXII, No. 2, pp. 41-56; 9 figs. June 29, 1962

SCIENTIFIC RESULTS OF THE GALAPAGOS-EXPEDITION
1953-54 OF THE INTERNATIONAL INSTITUTE FOR SUBMARINE

RESEARCH, VADUZ (LIECHTENSTEIN), LEADER DR. HANS HASS

CORALS FROM THE
GALAPAGOS AND COCOS ISLANDS

J. Wyatt Durham

University of California, Berkeley, California

During 1953-1954 the Galapagos Expedition of the International Insti-
tute for Submarine Research, Vaduz (Liechtenstein), leader Dr. Hans Hass,
on board the Xarifa, visited both Cocos Island and the Galapagos Islands in
the tropical Eastern Pacific and made various collections. Dr. Georg Scheer,
of the Hessisches Landesmuseum, Darmstadt, Germany, a member of the
expedition, collected numerous corals by skin diving. Through his courtesy
and patience I have had the opportunity of examining the corals and pre-
paring this report. The major part of the collection is housed in the Hes-
sisches Landesmuseum. Thanks to Dr. Scheer duplicate specimens of many

pean

42 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4711 SER.

of the species have been deposited in the Museum of Paleontology, Univer-
sity of California, Berkeley, California, under accession no. 2070. The
material at Berkeley includes representatives of Balanophylha scheeri, new
species; Pavona clivosa Verrill; Pavona gigantea Verrill; Pavona varians
Verrill; Pavona (Polyastra) ponderosa (Gardiner) ; Pocillopora damacornis
(Linnaeus); Pocillopora damicornis cespitosa Dana; Pocillopora elegans
Dana; Pocillopora meandrina Dana; Pocillopora verrucosa (Ellis and
Solander) ; Porites excavata Verrill; Porites lobata Dana (?); Psammocora
profundacella Gardiner; Psammocora (Stephanaria) stellata (Verrill) ; and
Tubastrea tenwilamellosa (Milne Edwards and Haime).

In identifying this collection of corals a broader species concept, in part
following Squires (1959), than used previously (Durham, 1947; Durham and
Barnard, 1952), has been employed in the potentially reef-building and
taxonomically troublesome genera Pavona, Pocillopora, and Porites. As a
result some of the previously employed names are hidden in the synonymies
and the list of species in these genera has a marked Indo-Pacific aspect.

Durham and Barnard (1952) have summarized most of the available
data on stony corals from Cocos Island and the Galapagos Islands, as well
as other Eastern Pacific areas. Vaughan (1906) described 3 species from the
Galapagos Islands that were not included in the paper by Durham and
Barnard. Since then Squires (1959) has presented an extensive discussion
of the corals of the Gulf of California, Durham and Allison (1960, pp. 70-
76) have tabulated available distributional data on living and fossil corals
of the Eastern Pacifie at the generic level, and Hertlein and Emerson (1957,
pp. 7-8) listed five corals previously unrecorded from Clipperton Island.

Cocos Island is situated a little over 500 kms. southwest of Costa Rica
and about 700 kms. northeast of the Galapagos Islands. During most of the
year it seems to be in the path of the Equatorial Counter Current. Durham
and Barnard (1952) recorded 7 species (table 1) from this area.

The Galapagos Islands are about 1000 miles off the Eeuadorian Coast and
are within the path of the westward flowing Peru current although a counter
oyral from the Equatorial Counter current is directed toward this area for
at least part of the year. Durham and Barnard (1952) recorded 21 species
and Vaughan (1906), had described an additional 3 species, making a total
of 24 species (table 1) previously known from these islands.

The corals collected by Dr. Scheer include representatives of at least 20
species and varieties, of which 4 have not been recorded from the Eastern
Pacific previously. Three of these species (Pavona (Polyastra) ponderosa
Gardiner, Porites lobata Dana(?), Psammocora profundacella Gardiner) are
members of the Indo-Pacific coral reef fauna. All three, however, are mem-
bers of genera that were previously known to occur in the Eastern Pacifie
and that are widespread throughout the warmer parts of the Indian and

-acifie Oceans.

Vou. XXXII] DURHAM: GALAPAGOS-COCOS ISLAND CORALS 43

Uf 6'
XARIFA EXPEDITION 1953-54

COCOS ISLAND cS
LOCALITIES ee G Ulloa Is

FROM DATA BY DRGEORG SCHEER
COLLECTING LOCALITY

ES
® Wenman GALAPAGOS ISLANDS XS

Abingdon
® Bindloe

North 90°40'W
Seymour |
Mosquera

a, Eb

North Seymour Guy Fawkes
South Seymour
Conway Bay f SN

XARIFA EXPEDITION 1953-54
GALAPAGOS ISLANDS LOCALITIES

FROM DATA BY DR.GEORG SCHEER
—o) COLLECTING LOCALITY

Figure 1. Collecting localities, Cocos and Galapagos islands.

44 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4rH SER.

TABLE 1. Corals previously reported from Galapagos Islands and Cocos Island
(Vaughan, 1906; Durham and Barnard, 1952).

Galapagos Cocos
Island Island

Astrangia equatorialis Durham and Barnard
Astrangia gardnerensis Durham and Barnard
Astrangia hondaensis Durham and Barnard (?)
Balanophyllia galapagensis Vaughan
Balanophyllia osburni Durham and Barnard
Caryophyllia diomedae Von Marenzeller
Cladocora debilis Milne Edwards and Haime
Cycloseris mexicana Durham
Desmophyllum galapagense Vaughan
Endopachys vaughani Durham
Flabellum daphnense Durham and Barnard
Kionotrochus (2?) avis Durham and Barnard
Kionotrochus (2?) hoodensis Durham and Barnard
Leptoseris digitata Vaughan (?) x
Lophosmilia wellsi Durham and Barnard
Madracis asperula Milne Edwards and Haime
Madracis sp.
Madrepora galapagensis Vaughan
*Pavona clivosa Verrill
Pavona cf. explanulata (Lamarck) Ki
Pocillopora damicornis cespitosa Dana x
Pocillopora elegans Dana x
Psammocora (Stephanaria) stellata (Verrill) x X

bd bd bd bedi bd od Pd! od Ol OM OP OSS
mow

a a a a

Sphenotrochus hancocki Durham and Barnard
Thecopsammia pourtalesi Durham and Barnard
Tubastrea tenuilamellosa (Milne Edwards and Haime)

*Nomenclature corrected to agree with Squires (1959) 24

=~]

An important result of this expedition has been the addition of distribu-
tional data (see figure 1 for collecting localities) to the scanty store of infor-
mation on corals of the tropical Eastern Pacific. The present collection
(table 2) adds 11 taxa to the 7 previously recorded from Cocos Island and
7 to the 24 previously known from the Galapagos. The addition of 10 species
to the coral fauna of these islands emphasizes how little systematic coral
collecting has been done in the Eastern Pacific (see: Durham and Barnard,
1952, pp. 3-4; Squires, 1959, pp. 372-873).

The recorded fauna of the Galapagos Islands (table 3) now totals 30
species, of which only 13 belong to the hermatypic group. Highteen species,
including 14 hermatypie types are now known from Cocos Island. Thirty-
eight species, of which 20 are hermatypic, are now recorded from these two
island areas. It seems probable that all of the hermatypie species may
eventually be found in both areas. Fourteen of the 20 hermatypie species

VoL. XXXII]. DURHAM: GALAPAGOS-COCOS ISLAND CORALS 45

TABLE 2. Corals collected by the Galadpagos-Expedition 1953-5 }.

; = Cocos
Galapagos Islands

pag Island
Localities

10 << S33 OS oN OO SOS s- cr

rererereoaandaand & ©& oO co

Sool morn

Astrangia dentata Verrill x
Astrangia equatorialis Durham and Barnard
Astrangia gardnerensis Durham and Barnard Xe
Balanophyllia osburni Durham and Barnard K
*Balanophyllia scheeri, new species x
Cycloseris elegans (Verrill) x
Pavona clivosa Verrill x x
Pavona gigantea Verrill XS x
Pavona varians Verrill x
*Pavona (Polyastra) ponderosa Gardiner
Pocillopora damicornis (Linnaeus) x x xk xX
Pocillopora damicornis cespitosa Dana
Pocillopora elegans Dana 5 ?
Pocillopora meandrina Dana x
Pocillopora verrucosa (Ellis and Solander) x x x
Porites excavata Verrill x
*Porites lobata Dana(?) Xk x
*Psammocora profundacella Gardiner x x
Psammocora (Stephanaria) stellata (Verrill)

Tubastrea tenuilamellosa (Milne Edwards
and Haime)

*Not previously reported in Eastern Pacific.

“

“HM
>)
os
*
ta

*
MM
al
Pal
rac]

are also known from the coast of the Americas. In addition, the hermatypic
species Agaricia sp., Leptoseris panamensis Durham and Barnard, Porites
panamensis Verrill, Psammocora brighami (Vaughan), and Solenastrea
ecuadoriana Durham and Barnard have been recorded (Durham and Bar-
nard, 1952) from the adjacent mainland coast. It seems probable that most
or all of these species may eventually be found in the offshore island areas.
Despite the increased distributional data afforded by the present collection
it is notable that no additional genera of hermatypic corals were added to
the known fauna. Genera such as Acropora, Montipora, Favia, Cyphastrea,
and Plesiastrea continue to be unrecorded on the American side of the East
Pacific Barrier (see Durham and Allison, 1960, pp. 69, 70-76). Likewise the
disappearance from the region of many of the hermatypic genera recorded
as fossils (Durham and Allison, 1960, pp. 71-73) from the Pacifie shores of
the Americas is still enigmatic. For example Durham and Allison list 30

46 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4711 SER.

extant genera of hermatypic corals that are known to occur in the fossil
record of this region. Although many of these genera are still present in the
Indo-Pacific region, only 10 are now living in the Eastern Pacific.

TaBLe 3. Summary of Corals recorded from Galapagos Islands and Cocos Island.

3
oD! zp
eS lutlas |e
Soe | Sa | ecules
aim | oF Cie) a 8
Oi or ote) me AY
Astrangia dentata Verrill xX x
Astrangia equatorialis Durham and Barnard x
Astrangia gardnerensis Durham and Barnard x
Astrangia hondaensis Durham and Barnard (?) Rs Si
Balanophyllia galapagensis Vaughan Xe
Balanophyllia osburni Durham and Barnard Oe
Balanophyllia scheeri, new species x
Caryophyllia diomedae Von Marenzeller x x
Cladocora debilis Milne Edwards and Haime x x
Cycloseris elegans (Verrill) os x
Cycloseris mexicana Durham x x x
Desmophyllum galapagense Vaughan x
Endopachys vaughani Durham x x x
Flabellum daphnense Durham and FParnard x
Kionotrochus (?) avis Durham and Barnard xe
Kionotrochus (2?) hoodensis Durham and Barnard x
Leptoseris digitata Vaughan (?) x x x
Lophosmilia wellsi Durham and Parnard x
Madracis asperula Milne Edwards and Haime xe
Madracis sp. Xe x
Madrepora galapagensis Vaughan exe
Pavone clivosa Verrill x ag
Pavona gigantea Verrill x x
Pavona varians Verrill x xX x
Pavona ct. explanulata (Lamarck) x x
Pavona (Polyastra) ponderosa (Gardiner ) x Xe
Pocillopora damicornis (Linnaeus ) x x x x
Pocillopora damicornis cespitosa Dana x x x
Pocillopora elegans Dana x x x x
Pocillopora meandrina Dana x xx x
Pocillopora verrucosa (Eliis and Solander ) Xs x x x
Porites excavata Verrill x x
Porites lobata Dana (?) x *
Psammocora profundacella Gardiner x x x
Psammocora (Stephanaria) stellata (Vervill) K x x x
Sphenotrochius hancocki Durham and Barnard x x
Theocopsammia pourtalesi Durham and Earnard x
Tubastrea tenuilamellosa (Milne Edwards and Haime) x x aN

Total 30 18 19 12

VoL, XXXIT]

DURHAM: GALAPAGOS-COCOS ISLAND CORALS 47

DESCRIPTION OF COLLECTING LOCALITIES (see figure 1) oF THE
GALAPAGOS-EXPEDITION 1953-54 (‘“XARtFA—E XPEDITION” )

GALAPAGOS ISLANDS

Loeality

Locality

Loeality

Locality

Loeality

Locality

Loeality

74.

80.
Ou:

90.

92.

98.
100.

Cocos ISLAND

Locality

Loeality

Locality

104.
107.

114.

Osborn Island, near Hood Island. About 30 m. offshore
on northwest coast. Depth 3 m.

From depth of 2 m. along steep shore of northwest coast
of Osborn Island.

From west shore of Xarifa Island, near Hood Island.

About 30 m. from beach, south side of southwest point of
rardner Island. Depth between 5 and 6 meters.

From southeast side of Gardner Island.

Guy Fawkes Islands, near Indefatigable Island. From
between the two islets, depth 5 m., on steep rocks.

North shore of North Seymour Island, from depth of 6 m.
on steep shore.

Mosquera Island, between North Seymour and South
Seymour Island. Along west side, about 50 m. offshore.
Depth 5 m.

East side of Darwin Bay, Tower Island, steep shore.

Near small inlet on north shore of Darwin Bay, Tower
Island.

From northeast side near Ulloa Island, depth 5 m.

From shore of Wrack Islet, Wafer Bay, northwest side of
Cocos Island. Depth 8 m.

From depth of 7 to 10 meters, Wafer Bay, northwest side
of Cocos Island.

SYSTEMATICS
Family PocILLOPoRIDAE Gray, 1842

Genus Pocillopora Lamarck, 1816

Type species Pocillopora acuta Lamarck.

Pocillopora damicornis (Linnaeus).

Millepora damicornis LINNAEUS, 1758, Systema Natura, Ed. 10, p. 791.
Pocillopora lacera VrERRILL, 1869, Proc. Essex Inst., Vol. 6, p. 100; DurRHAm and
BarNArD, 1952, Allan Hancock Pac. Exped., vol. 16, no. 1, pp. 20-21, 26, pl. 1, fig. 4.

48 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Pocillopora damicornis (Linnaeus), WELLS, 1954, U.S. Geol. Surv. Prof. Pap. 260,
p. 412, pl. 99, fig. 2; Squires, 1959, Bull. Amer. Mus. Nat. Hist., vol. 118, pp. 408—
409, pl. 34, fig. 2.

OccuRRENCE. Xarifa localities 74, 79(?), 107, 114. Cocos and Galapagos
islands. Recorded from Panama to Indian Ocean. The illustrated specimen
(fig. 9) is referred to this species with some doubt because of its extreme
“roughness.”

Pocillopora damicornis var. cespitosa Dana.

Pocillopora cespitosa DANA, U.S. Expl. Exped., Zooph., p. 525, pl. 49, figs. 5, 5a.

Pocillopora damicornis var. cespitosa Dana, HOFFMEISTER, 1925, Carnegie Inst.,
Wash., Publ. Dept. Mar. Biol., vol. 22, pp. 16-17; DuRHAm and BARNARD, 1952,
Allan Hancock Pac. Exped., vol. 16, no. 1, p. 20, pl. 1, figs. 3a—3c.

OccuRRENCE. Xarifa localities 74, 89(?). Galapagos Islands. Recorded
from Panama and widely through the Indo-Pacific.

Pocillopora elegans Dana.

Pocillopora elegans DANA, 1846, U.S. Explor. Exped., Zooph., p. 532, pl. 51, figs. 1, 1a;
WELLS, 1954, U.S. Geol. Surv., Prof. Pap. 260, p. 413, pl. 95, fig. 2; Squires, 1959,
Bull. Amer. Mus. Nat. Hist., vol. 118, pp. 409-410, pl. 34, fig. 5.

Pocillopora capitata VERRILL, 1864, Bull. Mus. Comp. Zool. Harvard Coll., vol. 1, p. 60.

Pocillopora capitata var. robusta VERRILL, 1870, Trans. Conn. Acad. Arts Sci., vol. 1,
pp. 521-522.

Pocillopora robusta Verrill, DURHAM and BARNARD, 1952, Allan Hancock Pac. Exped.,
vol. 16, no. 1, pp. 26—28, pl. 1, figs. 5a—5b, 6.

OccuRRENCE. Xarifa localities 74, 76(?), 107, 114. Cocos and Galapagos
islands. Reeorded from the Gulf of California to the Indo-Pacifie.

Pocillopora meandrina Dana.

Pocillopora meandrina DANA, 1846, U.S. Explor. Exped., Zooph., p. 529, pl. 50, figs.
3, 8a; VAUGHAN, 1918, Carnegie Inst., Wash., Pap. Dept. Mar. Biol., vol. 9, p. 78;
Squires, 1959, Bull. Amer. Mus. Nat. Hist., vol. 118, pp. 410-411, pl. 34, fig. 4.

OccURRENCE. Xarifa locality 114, Cocos Island. Recorded from Gulf of
California to the Indo-Pacific.

Pocillopora verrucosa (Ellis and Solander).

Madrepora verrucosa ELLIs and SoLANDER, 1786, Nat. Hist. Zooph, p. 172.

Pocillopora capitata var. porosa VERRILL, 1869, Proc. Essex Inst., vol. 6, p. 99.

Pocillopora porosa Vervill, DurtAM and BARNARD, 1952, Allan Hancock Pac. Exped.,
VOL Gammon espe 2G:

Pocillopora verrucosa (Ellis and Solander), WrELLS, 1954, U.S. Geol. Surv. Prof. Pap.
260, p. 413, pl. 98, figs. 5-6; DuriHAmM, in Hertlein and Emerson, 1957, Amer. Mus.

Novitates, no. 1859, p. 7; Squrres, 1959, Bull. Amer. Mus. Nat. Hist., vol. 118,
p. 410.

VoL. XXXII] DURHAM: GALAPAGOS-COCOS ISLAND CORALS 49

OccuRRENCE. Xarifa localities 90, 100, 107. Cocos and Galapagos islands.
Recorded from Gulf of California, Clipperton Island, and westward to
Indian Ocean.

Family THAMNASTERIDAE Vaughan and Wells, 1943
Genus Psammocora Dana, 1846
Type species Pavona obtusangula Lamarck.

Psammocora (Psammocora) profundacella Gardiner.
(Figure 6.)

Psammocora profundacella GARDINER, 1898, Proc. Zool. Soc. London for 1898, p. 537,
pl. 45, fig. 3; VAUGHAN, 1918, Carnegie Inst. Wash., Pap. Dept. Biol., vol. 9, p. 78.

Hypvortyre, Univ. Calif. Mus. Paleo., no. 30725, Xarifa locality 114.

OccuRENCE. Xarifa localities 92 and 114. Cocos and Galapagos islands.
Previously recorded from Funafuti and Fanning Island.

Subgenus Stephanaria Verrill, 1867.
Type species Stephanocora stellata Verrill.

Psammocora (Stephanaria) stellata (Verrill).

Stephanocora stellata VERRILL, 1866, Proc. Boston Soc. Nat. Hist., vol. 10, p. 330.

Stephanaria stellata VERRILL, 1870, Trans. Conn. Acad. Arts Sci., vol. 1, pp. 545-546,
pl. 9, figs. 4, 4a.

Psammocora (Stephanaria) stellata (Verrill), DurHAM and Barnarp, 1952, Allan
Hancock Pac. Exped., vol. 16, no. 1, pp. 29-30, pl. 2, fig. 8.

OccurRENCE. Xarifa locality 74, Galapagos Islands. Previously recorded
from the Galapagos Islands, Cocos Island, Hawaiian Islands, and Panama to
the Gulf of California.

Family AGARICHDAE Gray, 1847
fenus Pavona Lamarck, 1801
Type species Pavona cristata Lamarck.

Pavona (Pavona) clivosa Verrill.

Pavonia clivosa VERRILL, 1869, Proc. Boston Soc. Nat. Hist., vol. 12, pp. 395-396.

Pavona (Pavona) clivosa Verrill, DURHAM and BARNARD, 1952, Allan Hancock Pac.
Exped., vol. 16, no. 1, pp. 37, 42, pl. 2, fig. 12.

Pavona (Pseudocolumnastrea) galapagensis DURHAM and BARNARD, 1952, Allan Han-
cock Pac. Exped., vol. 16, no. 1, pp. 44—45, pl. 3, figs. 16a—16b.

Pavona clivosa Verrill, SqutRES 1959, Bull. Mus. Nat. Hist., vol. 118, pp. 412-413, pl.
SVs likey ae

50 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4T1r SER.

OccuRRENCE. Xarifa localities 76 and 92, Galapagos Islands. Previously
reported from Eeuador, the Galapagos Islands, Panama and the Gulf of
California.

Pavona (Pavona) gigantea Verrill.

Pavonia gigantea VERRILL, 1869, Proc. Boston Soc. Nat. Hist., vol. 12, pp. 394-395.

Pavona (Pavona) gigantea Verrill, DuRtmAM and BARNARD, 1952, Allan Hancock Pac.
Exped: vol: W6; no: 15 p: 43, plo 3, tie. 14.

Pavona gigantea Verrill, Squrres, 1959, Bull. Amer. Mus. Nat. Hist., vol. 118, pp. 413—
414, pl. 32, fig. 2.

OccuRRENCE. Xarifa localities 79 and 92, Galapagos Islands. Previously
recorded from Clipperton Island, and Panama to Gulf of California.

Pavona (Pavona) varians Verrill.
(Figure 3.)

Pavonia varians VERRILL, 1864, Bull. Mus. Comp. Zool., vol. 1, no. 3, p. 55.

Pavona varians Verrill, VAUGHAN, 1907, U.S. Nat. Mus. Bull. 59, p. 135, pl. 38, figs. 1,
la; WELLS, 1954, U.S. Geol. Surv. Prof. Pap. 260, p. 442, pl. 152, figs. 3—4.

Pavona (Pavona) cf. varians Verrill, DURHAM and BARNARD, 1952, Allan Hancock
Pac. Exped., vol. 16, no. 1, p. 43, pl. 3, fig. 15.

Hypotype, Univ. Calif. Mus. Paleo., no. 30721, Xarifa locality 114.
OccURRENCE. Xarifa locality 114, Cocos Island. A widespread species,

ranging from the Red Sea and Great Barrier Reef to the Hawaiian Islands
and Colombia.

Subgenus Polyastra Ehrenberg, 1854

Type species Polyastra venosa Ehrenberg.

Pavona (Polyastrea) ponderosa (Gardiner).

(Figure 5.)

Agaricia ponderosa GARDINER, 1905, Fauna and Geogr. Maldive and Laccadive Arch.,
vol. 2, supl. 1, p. 937, pl. 89, figs. 1-2; VaAuGHAN, 1918, Carnegie Inst. Wash., Pap.
Dept. Mar. Biol., vol. 9, p. 140; YaBe, SuciyAMA, and Eeucut, 1936, Sci., Repts.
Tohoku Imp. Univ., Ser. 2, (Geol.), Spec. vol. 1, p. 55, pl. 27, fig. 5, pl. 38, fig. 1,
pl s2) figs 1

Hyportyrpr, Univ. Calif. Mus. Paleo., no. 30724, Xarifa locality 114.

OccurRENCE. Xarifa locality 114, Cocos Island. This species is known
from the Maldives to the Bonin Islands.

Vou. XXXII] DURHAM: GALAPAGOS-COCOS ISLAND CORALS 51
Family Poriripar Gray, 1842
Genus Porites Link, 1807

Type species Jadrepora porites Pallas.

Porites californica Verrill.

Porites californica VERRILL, 1870, Trans. Conn. Acad. Arts Sci., vol. 1, p. 504;
DuRHAM and EARNARD, 1952, Allan Hancock Pac. Exped., vol. 16, no. 1, p. 46, pl.
3, figs. 17a—17b; Squires, 1959 (in part), Eull. Amer. Mus. Nat. Hist., vol. 118,
pp. 420-422, pl. 32, figs. 3-6, pl. 33, figs. 3-4 (only).

Porites porosa VERRILL, 1870, Trans. Conn. Acad. Arts Sci., vol. 1, p. 504.

Porites nodulosa VerRRILL, 1870, Trans. Conn. Acad. Arts Sci., vol. 1, pp. 505-506;
DuriwAM and BarnNaArpD, 1952, Allan Hancock Pac. Exped., vol. 16, no. 1, pp. 46—
ANT vols Byecinifess DUS

OccuRRENCE. Xarifa locality 107, Cocos Island. Previously recorded from
Panama to Magdalena Bay and to the head of the Gulf of California.

Porites excavata Verrill.

Porites excavata VERRILL, 1870, Trans. Conn. Acad. Arts and Sci., vol. 1, pp. 504-505;
Squires, 1959, Bull. Amer. Mus. Nat. Hist., vol. 118, pl. 33, figs. 1-2.

OccuRRENCE. NXarifa locality 107, Cocos Island. Recorded previously
from Panama. The ealices on one specimen are smaller than those on the
type as figured by Squires. Squires suggests that his P. baueri, described
from the Tres Marias Islands, is closely related.

Porites lobata Dana (7?)
(Figure 8.)

Porites lobata DANA, 1846, U.S. Expl. Exped., Zooph., p. 562, pl. 55, fig. 1; VAUGHAN,
1907, U.S. Nat. Mus. Bull. 59, pp. 196-198, pl. 81, figs. 1-la, 1b; Vaueuan, 1918,
Carnegie Inst. Wash., Pap. Dept. Mar. Biol., vol. 9, p. 192.

Iypotyrr, Univ. Calif. Mus. Paleo., no 30726, Xarifa locality 98.

OccurRENCE. Xarifa localities 76, 79, and 98, Galapagos Islands. The
specimens have deeper calices and the top of the wall more fused than in
typical P. lobata. They also resemble P. paschalensis Vaughan, described
from Faster Island, but the calices average smaller in size and the columella
is more compressed. P. lobata is widespread in the Central Pacific.

Family FUNGHDAE
Genus Cycloseris Milne Edwards and Haime, 1849.

Type species Fungia cyclolites Lamarck.

52 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SEr.

Cycloseris elegans (Verrill).

Fungia elegans VERRILL, 1870, Am. Jour. Sci., ser. 2, vol. 49, p. 100.
Cycloseris elegans (Verrill), DURHAM and BARNARD, 1952, Allan Hancock Pace.

Exped., vol. 16, no. 1, p. 52, pl. 4, figs. 20a—20b; Squtres, 1959, Bull. Amer. Mus.
Nat. Hist., vol. 118, p. 414.

OccuRRENCE. Xarifa locality 80, Galapagos Islands. Previously recorded
from La Paz to Panama.

Family RuizancuparE d’Orbigny
Genus Astrangia Milne Edwards and Haime, 1848
Type species Astrangia michelinii Milne Edwards and Haime.
Astrangia dentata Verrill.

Astrangia dentata VERRILL, 1866, Proc. Boston Soc. Nat. Hist., vol. 10, p. 332; DuRHAM,
1947, Geol. Soc. Amer. Mem. 20, p. 28, pl. 5, fig. 6.

OCCURRENCE. Xarifa locality 107, Cocos Island. Previously reported from
La Paz to Panama.

Astrangia equatorialis Durham and Barnard.

Astrangia equatorialis DURHAM and BARNARD, 1952, Allan Hancock Pac. Exped., vol.
16, no. 1, p. 69, pl. 6, figs. 29a—29b.

OccuRRENCE. Xarifa locality 98, Galapagos Islands. Previously reported
from the Galapagos Islands.

Astrangia gardnerensis Durham and Barnard.

Astrangia gardnerensis DURHAM and BARNARD, 1952, Allan Hancock Pac. Exped.,
Vol. 165 nos de%ps 70) plaid, fies 277:

OccurRENCE. Xarifa locality 75, Galapagos Islands. Originally described
from the same bay.

Family DENDROPHYLLUDAE Gray, 1847
yenus Balanophyllia Searles Wood
Type species Balanophyllia calyculus Searles Wood.

Balanophyllia osburni Durham and Barnard.
(Figure 2a, d.)

Balanophyllia osburni DurwAM and BARNARD, 1952, Allan Hancock Pac. Exped., vol.
16, no. 1, pp. 100-101, pl. 15, figs. 63a—63d.

OccuURRENCE. Xarifa locality 74, Galapagos Islands. Described from the
Galapagos.

VoL. XXXII] DURHAM: GALAPAGOS-COCOS ISLAND CORALS D3

Balanophyllia scheeri Durham, new species.
(Figures 2b, 2c, 4, 7.)

Corallum elongate, tall; attachment base broken but apparently broad;
cealice round to slightly oval; calicular fossa deep; columella oval in outline,
rising slightly above base of fossa, parietal, moderately compact; septa ap-
parently in five incomplete cycles (fig. 2e¢) ; first cycle septa broadly rounded
at top (fig. 2b) inner edge smooth, nearly vertical, fused to columella at
depth; second cycle septa narrow at top, usually smooth, descending obliquely
to columella; third cycle septa narrow, inner edge dentate; fourth cycle septa
more prominent than third cycle, dentate, fused to one another about mid-
way to columella and in turn fused to columella at higher point than first
and second cycle septa; fifth cycle septa narrow, fused to inelude fourth
cycle, developed only in outer part of each system; septa considerably
thinner than adjacent interspaces, laterally finely granulate; wall porous,
with epitheca of varying thickness for one-half to two-thirds height of coral-
lum; costae corresponding to all septae, covered by a row of fine granules;
first cycle septa slightly more prominent than others.

Figure 2. a, d. Balanophyllia osburni Durham and Barnard. After hypotype in
Hessisches Landesmuseum, no. X 1:74-7. a. Lateral profile of 4 cycles of septa; d.
Plan view of part of calice. b, c. Balanophyllia scheeri, n. sp. After holotype, Univ.
Calif. Mus. Paleo. no. 30722. b. Lateral profile of 4 cycles of septa; c. Plan view of
part of calice.

54 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER.

DIMENSIONS OF HOLOTYPE. Height (incomplete) 22.4 mm., maximum dia-
meter 9.0 mm., depth of fossa to top of columella 3.6 mm.

HowuotyveeE: Univ. Calif. Mus. Paleo., no. 30722. Paratypes: Univ. Calif.
Mus. Paleo., no. 30723 and Hessisches Landesmuseum nos. XI:114—9e,
XT :114—9d; all from Xarifa locality 114.

OccuRRENCE. Xarifa locality 114, Wafer Bay, Cocos Island. This species
resembles single corallites of Dendrophylla fistula (Aleock) but differs by
the presence of the fifth cycle septa, and the slightly prominent first cycle
costae. Balanophyllia galapagensis Vaughan has a shallower fossa, lacks the
fifth evele septa, and has a heavier epitheca.

Genus Tubastrea Lesson, 1854.

Type species Tubastrea coccinea Lesson.

Tubastrea tenuilamellosa (Milne Edwards and Haime).

Coenopsammia tenuilamellosa MILNE Epwarpbs and Harms, 1848, Ann. Sci. Nat., ser.
Bevo aK, Toe TUL Tol, Al, anes, IGE

Astropsammia pedersenii VERRILL, 1869, Proc. Boston Soc. Nat. Hist., vol. 12, p. 392.

Tubastrea tenuilamellosa (Milne Edwards and Haime), DurmAM and BARNARD, 1952,
Allan Hancock Pac. Exped., vol. 16, no. 1, pp. 105-106, pl. 12, fig. 50d.

Tubastrea aurea Squires, 1959, Bull. Am. Mus. Nat. Hist., vol. 118, pp. 427-428 (pro
parte, non Quoy and Gaimard, 1833).

OccuRRENCE. Xarifa localities 74, 89, 104, 107, and 114. Galapagos and
Cocos islands. Previously recorded from San Marcos Island, Gulf of Cali-
fornia to the Galapagos. One specimen from locality 114 closely resembles
Verrill’s Astropsammia pederseni.

FicurE 3. Pavona varians Verrill (xX 0.83). Hypotype, Univ. Calif. Mus. Paleo.,
no. 30721, Xarifa locality 114, Cocos Island.

Ficures 4, 7. Balanophyllia scheeri, new species (xX 1.66). 2, holotype, Univ.
Calif. Mus. Paleo., no 30722; 5, paratype, Univ. Calif. Mus. Paleo., no. 30723. Both
from Xarifa locality 114, Cocos Island.

FIGURE 5. Pavona ponderosa (Gardiner) (x 0.83). Hypotype, Univ. Calif. Mus.
Paleo., no 30724, Xarifa locality 114, Cocos Island.

Figure 6. Psammocora profundacella Gardiner (x 0.83). Hypotype, Univ. Calif.
Mus. Paleo., no. 30725, Xarifa locality 114, Cocos Island.

Ficure 8. Porites lobata Dana (?) (x 0.83). Hypotype, Univ. Calif. Mus. Paleo.,
no. 30726, Xarifa locality 98, Darwin Bay, Tower Island, Galapagos Island.

Ficure 9. Pocillopora damicornis (Linnaeus) (?) (xX 0.83). Hypotype, Univ.
Calif. Mus. Paleo., no. 30727, Xarifa locality 79, Gardner Island, Galapagos Islands.

GALAPAGOS-COCOS ISLAND CORALS

DURHAM:

XXXII]

VOL.

56 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.
LITERATURE CITED

DuRHAM, J. W.
1947. Corals from the Gulf of California and the North Pacific Coast of America.

Geological Society of America, mem. 20, pp. 1-68, pls. 1-14.

DuRHAM, J. W., and E. C. ALLISON
1960. The geologic history of Baja California and its marine faunas. In sym-

posium: The Biogeography ot Baja California and adjacent seas. Sys-
tematic Zoology, vol. 9, pp. 47-91.

DurRHAM, J. W. and J. L. BARNARD
1952. Stony corals of the Eastern Pacific collected by the Velero III and Velero

IV. Allan Hancock Pacific Expeditions, vol. 16, no. 1. pp. 1-110, pls. 1-16.

HERTLEIN, L. G., and W. K. EMreRsSon
1957. Additional notes on the invertebrate fauna of Clipperton Island. American

Museum Novitates, no. 1859, pp. 1-9.

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.
Bulletin of the American Museum of Natural History, vol. 118, Art. 7,
pp. 367-432, pls. 28-34.

VAUGHAN, T. W.
1906. Reports of the scientific results... “‘Albatross”,... Part VI, Madreporaria.

Bulletin of the Museum of Comparative Zoology, Harvard College, vol.
50, pp. 61-72, pls. 1-10.

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

FOURTH SERIES

G Dallas Hanna Anniversary Volume

Vol. XXXII, No. 3, pp. 57-78; 3 figs. June 29, 1962

MATING BEHAVIOR AND ACTIVITY-REST
PERIODICITY IN PROTOCLYTHIA CALIFORNICA
(DIPTERA: PLATYPEZIDAE)'

Edward L. Kessel

University of San Francisco
and

California Academy of Sciences

and

Berta B. Kessel

University of San Francisco

INTRODUCTION

The first direct observation dealing with the mating behavior of flat-
footed flies was made by Snow (1894). This involved a single mating pair of
the species Calotarsa calceata (Snow). The flies were captured with a lucky
stroke of the net as they passed within an inch or two of the observer’s eye
and therefore too close to be brought into focus. They did not remain in
copula in the net, so Snow was unable to contribute any notes on the mating

1. Based on studies aided by National Science Foundation grant G19006.

[57 J

APR 1 8 1968
WOODS HOLE, MASS.

58 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4r1 SER.

behavior of these flies except to mention that at the time of the capture there
were a dozen or so males of this same species engaged in zigzag flight overhead.

Except for oceasional observations (e.g., Verrall, 1901; Kessel, 1948) that
males of other platypezid species engage in aerial dances, knowledge of the
mating habits of flat-footed flies did not progress for another sixty-seven
vears. In the meantime we had observed the mating behavior of certain
balloon flies, species of the genera Empimorpha, Hilara, and Empis (Kessel
and Karabinos, 1947; Kessel and Kessel, 1951; Kessel, 1955 and 1959), which
belong to the Empididae, a family that most certainly is closely related to
Platypezidae. The males of these empidids dance in swarms similar to those
of platvpezids, and in the ease of these balloon flies we were able to observe
the female enter the swarm of males, embrace with one of them, and then
settle with him on a nearby shrub until the mating activities were completed.
The observation of platypezid and empidid males dancing in similar swarms
led to the supposition that the females of flat-footed flies, like those of balloon
flies, enter the swarms of males to find their mates. But no evidence of such
behavior was forthcoming until recently when we were able to observe the
mating activities of Platypezina pacifica Kessel.

MatING BEHAVIOR OF PROTOCLYTHIAS

Our observations on Platypezina pacifica had barely been published
(IKXessel and Kessel, 1961) when we discovered flies of another platypezid
genus engaged in mating activities. These belonged to the species Proto-
clythia californica Kessel, and our observations were made at Steckel Park,
Ventura County, California.

We arrived at Steckel Park early on the afternoon of December 19, 1961.
The first hour of searching yielded no platypezids, but at 1:55 o’clock we
came upon a mating pair of the species mentioned. They were resting on a
horizontal leaf of a Solanwm plant which was growing in a protected area
near the river and in a spot surrounded by willow trees and bushes.

As on the occasion of our discovery of the mating pairs of Platypezina
pacifica, we were startled by so rare a sight. Again our first reaction was to
look up for the expected swarm of dancing males, and again there they were
directly overhead. The general picture was the same, even to the chimney-
like opening in the trees where the males engaged in their erratic zigzag
flight. Once more we found the mating flies sitting with their heads directed
away from each other, the larger male with his wings extended over those of
the female. As before, the flies remained very quiet until they were dis-
turbed, and then only the female responded, moving forward and towing the
male behind her. When the insects were taken in the net they remained
attached and continued in this union until they were transferred to the
cyanide jar. They are mounted on the same pin, but on separate points, and
are deposited in the collection of the California Academy of Sciences.

Vou. XXXII] KESSEL AND KESSEL: PROTOCLYTHIA BEHAVIOR 59

Figurel. The mating “chimney”

redwoods.

~ ra ee
eae,

Figure 2. The mating “chimney” of Protoclythia californica surrounded by

willows.

60 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47H SER.

Although the general mating pictures for Platypezina pacifica and Pro-
toclythia californica proved to be basically similar, we observed obvious
differences pertaining to the trees outlining the “chimney,” that vertically
open shaft in which the males danced (figs. 1 and 2).? In the case of Platy-
pezina pacifica the trees were coast redwoods (Sequow sempervirens) while
for Protoclythia californica they were willows (Salix sp.). A second dif-
ference involved the height of the trees, the redwoods of the Platypezina
“chimney” being more than 200 feet tall, while the willows which lined the
swarming opening of the protoclythias were only 20 to 25 feet high. It is of
interest to note in this connection that we have always found platypezinas
associated with areas where coniferous trees are growing, while we have
collected many specimens of Protoclythia californica far removed from such
forests.

Data are given in table 1 for the Steckel Park collection spot at the time
(1:55 P.M.) the mating flies were taken. These include altitude of the sun,
temperature, relative humidity, hght intensity, barometric pressure, and
wind velocity.

In order to make certain that the flies which were dancing in the “chimney”
actually were males of Protoclythia californica, a net was passed through
the swarm. The three specimens that were taken by this stroke were all males
of this species. No attempts were made to capture more of the insects at that
time and the remainder of the afternoon was spent in observing their actions.

Unlike the swarm observed by Snow (1894) for Calotarsa calceata and
those which we have seen for Calotarsa insignis Aldrich and Platypezina
pacifica (Kessel and Kessel, 1961), all of which were at a height of fifteen
feet or more, the aggregation of dancing males in the case of Protoclythia
californica was much lower, the altitude of flight frequently being as low as
five feet. This made it possible to observe the behavior of individual insects
in the swarm in more detail than had ever been possible in other species.

Reference has been made to the parallels which we have observed between
the epigamic behavior of balloon flies and that which we have seen exhibited
in platypezids. In our publications dealing with balloon flies we have more
than once made the statement that we have observed the female enter a
swarm of males, embrace one of them, and then settle with him on a nearby
shrub. Also, in our last paper (Kessel and Kessel, 1961) we recorded our
observation of a mating pair of Platypezina pacifica descending from the
swarm of males and added the comment: “It is evident that the female, like
those of balloon flies, had selected her mate from the dancing swarm, and the
pair were descending to the groundeover plants to complete their nuptial
activities.” In these instances we have attributed the initiative in selecting
a mate to the female, but from our closer observation of swarming proto-

2. We wish to thank Mr. Maurice Giles for preparing the three black-and-white prints from our koda-
chromes for use as illustrations in this paper.

VoL. XXXII] KESSEL AND KESSEL: PROTOCLYTHIA BEHAVIOR 61

Table 1

Generalized data for various times of the day at the “chimney” location in Steckel
Park, Ventura County, California, on December 19 and 20, 1961. Pacific Standard
Time is given to the nearest 5 minutes and no correction is made for the east-of-
meridian location. Temperature is given to the nearest degree Fahrenheit and rela-
tive humidity to the nearest per cent. Light intensity with the meter directed up the
“chimney” is recorded to the nearest 50 foot-candles and with it pointed toward the
ground under the “chimney” the light intensity is given to the nearest 5 foot-candles.
The barometric pressure on both days was 29.6 inches and the wind velocity was

never higher than 5 miles per hour.

Time Altitude Relative Light up Light down

12SSe40s of sun Temperature humidity “chimney” “chimney”
9:05 A.M. 20°10’ 60° 78% 300 20
es 21°43" 63° 15% 300 25
9:40 rays) > Sy 64° T4A% 400 30
10:30 29°40’ 64° 72% 500 30
11:00 31°20’ 64° 67% 500 30
ills 40) 32°18’ 64° 67% 500 30
12:00 M. 33°07’ 64° 64% 450 30
12:55 P.M. alls 66° 64% 400 30
1:05 ad ats 66° 64% 300 25
aus 29°55! 65° 65% 300 25
1:30 28°50’ 65° 67% 300 25
IS ys 26°48’ 64° 68% 300 20
2255 19°40’ 64° 72% 300 20
SEO ie Abe 64° 76% 250 10

elythias we are now convinced that the male is not passive in the mate-
selecting process.

It is evident that in both balloon flies and platypezids mating takes place
only in the swarms where the males are waiting for the females to come to
them. We have often observed males and females belonging to the same
species sitting in close proximity on the same leaf but in only one ease did
the sexes not ignore one another under such circumstances. The exception
occurred when we were collecting in the Botanical Gardens of the University
of California at Los Angeles and involved Clythia (== Platypeza) agarici
(Willard). A male that was sitting close to a female suddenly jumped on her
back but she promptly threw him off. It appears that the female must be
stimulated by the recognition of her own special species’ pattern of epigamic
behavior before she will accept a mate, and that she encounters this pattern
only in the swarm of dancing males. She doubtless recognizes them as be-
longing to her own species by some such specific symbol as the flashes of their
balloons (Empis, Empimorpha, Hilara), the scintillations of the flags on
their feet (Calotarsa insignis), or other decorations and characteristics of
their extended posterior tarsi (Calotarsa calceata, Platypezina pacifica, Pro-
toclythia californica). Once she has recognized the swarm as consisting of

62 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4r11 SEr.

males of her species she flies in among them. On a leaf she may not recognize
a potential mate, at least she repulses any advanees that he may make, but in
the proper swarm she is receptive and perhaps even eager. In the ease of the
balloon fly Empimorpha geneatis Melander we saw the female enter the
swarm of males and there embrace with one of them. We presumed at the
time that she had taken the initiative and selected a mate, but the joining of
the flies happened so quickly once she had entered the swarm that it was
impossible to be sure which sex had made the advances.

Because of the lower swarming flight of Protoclythia californica we were
near enough to the insects to ascertain that all of the dancing flies were
males. It was also easily seen that with even the slightest indication of a
breeze they would all face into it, “standing” still in the air or rising and
falling with only slow progress forward. When a male had finally reached
the periphery of the “chimney” area, he would swine back more or less to
the opposite margin of the swarm and once more face into the breeze. When
there was no evident air movement, the flies would usually merely turn about
and progress slowly through the swarm to the opposite side of the “chimney.”
It was not possible for us to determine if there was a sheht updraft of
warmer air in the vertically open shaft. There is opportunity to speculate
on the possible relationship between air movements or temperature gradients
and the selection of these “chimneys” for nuptial flights.

In watehing the swarm of protoelythias we noticed that when a male came
closer than about four inches to another fly, the two insects would invariably
dart toward each other, only to veer away before contact was made. It
seemed as though the males were inspecting one another, each to determine if
the other fly was a female. The males were very alert in these actions and
any female entering the swarm would have been grabbed by the first male
she approached in her line of flight.

The compound eyes of male platypezids are both larger and more complex
than those of the female. They are so large, in fact, that they are contiguous
alone the midline in front. As for the complexity of the eyes themselves,
each is divided by a shallow groove into an upper portion possessing larger
facets and a lower part having smaller facets. By contrast, the female has
her compound eyes widely separated and all of her facets are of uniform size,
about equivalent to those of the lower area of the male’s eve. There has been
considerable speculation as to the significance of these differences. Presum-
ably the more complex eyes of the male give him wider and sharper vision
than the female enjoys. It seemed illogical, therefore, to suppose that this
more elaborate optical equipment is without significance in the matine be-
havior of these flies which would certainly be the case if the male did nothing
but dance with his fellows and wait passively for a female to select him. We
now know that in the swarm the male darts aggressively at every fly that

Vot. XXXII] KESSEL AND KESSEL: PROTOCLY7IIIA BEHAVIOR 63

comes within range. The first male to recognize the female and to get to her
will take her as his mate.

We have presumed that the events leading up to pairing ave similar in
the balloon flies and the platypezids, and this in spite of the different copula-
tion positions which the members of their respective families, Empididae and
Platypezidae, typically assume. In the eases of the balloon flies which we
have observed the male sits astride the female with his head in the same
direction as hers, whereas in the platypezids the male and the female are
directed away from each other and with only the tips of their abdomens in
contact. These positions seem to be correlated with the nature of the male
genitalia which in empidids are characteristically directed upward and for-
ward and in typical platypezids are pointed downward and forward. These
would seem to be major distinctions until one considers the fact that the
male platypezid, as he approaches the completion of metamorphosis, has
genitalia positioned like those of empidids. Only later, but while the fly is
still in the puparium, do the genitalia begin to rotate to assume their final
direction. By the time of emergence, the rotation is nearing completion of a
180° movement, and by the time the male is sexually mature the hypopygium
is directed downward as well as forward. It is likely that this difference
achieved by the 180° rotation of the genitalia in platypezids requires diver-
gent copulatory positions in the two families. But in no way do the different
positions and structures here described indicate that the two families are not
closely related as we have believed.

Returning to the consideration of the swarm of Protoclythia californica
which we observed at Steckel Park, some of the dancing males continued
their activities as late as 3:10 P.M. when the san disappeared behind the hills
(see table 1 for data). However, their numbers had been gradually diminish-
ing before that time. Almost as the sun vanished, the last males disappeared
from the “chimney.” No platypezids had been seen on the bushes since 2:55
o’eloek.

Early next morning we returned to the swarming site to ascertain when
and under what conditions the males would resume their dancing, but in
spite of the fact that the day was clear and sunny like the one before, no flies
appeared in the “chimney” until early afternoon. Nevertheless, at 9:05 A.M.
(see table 1 for data) we did begin to see protoelythias of both sexes moving
about on the leaves of bushes under the “chimney” area, stopping here and
there to feed. For approximately the next two hours the flies continued their
morning exercise and feeding period. Then came the rest period, beginning
as abruptly as had the morning activity period which it replaced.

ACTIVITY-REST PERIODICITY

Our many years of platypezid collecting during all of the seasons, from
Alaska to Mexico and from the Atlantic to the Pacific, had led us to conclude

64 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4711 SER.

that the more sun-loving flat-footed flies interrupt their daily aetivities to
take a siesta during the middle of the day. So definite was this feeling that
we often found ourselves planning our collecting stops so as to avoid the
hour or so before and after noon. When we made a point of looking into the
matter we found that our impression agreed with the facts. While those
species of Platypezidae which frequent environments of deep shade do not
engage in well defined rest periods, all of those forms which are to be found
typically in habitats of filtered sunshine seem to resort to a mid-day period
of inactivity. Representative of the shade-dwelling forms are Platypezina
pacifica, Platypezina diversa (Johnson), Calotarsa insignis, Agathomyta lu-
cifuga Kessel, Agathomyia nemophila Kessel, and Agathomyva sylvania Kes-
sel. Examples of the sun-loving species are Protoclythia californica, Clythia
agarict (Willard), Clythia polyport (Willard), Clythia dymka Kessel,
Clythia cinerea (Snow), Clythia hunteri Kessel, and Clythia coraxa Kessel.
The species of Metaclythia would doubtless fall into this second eategory.
Our experience indicates that the species of Callomyia are intermediate in
their sun-shade inclinations.

Reference has been made to our observation that protoelythias indulge in
a morning activity period during which they feed and run about on the
leaves. It has also been our observation that all of the sun-loving species
have such a time of activity and that they exhibit a corresponding period in
the afternoon. Their mid-day rest occupies the time between these two.
Everywhere we have been impressed by the rather sudden onset and abrupt
termination of these periods. We were also impressed by the fact that the
activity periods of C. cinerea in Alaska are longer than those exhibited by
this species in the San Francisco Bay area of California. Similarly, for C.
corara the activity periods are longer in the San Francisco region than are
those which we found for this species in San Diego County, California, some
375 miles to the south. These longer activity periods would seem to be re-
lated in some way to the longer days one encounters in our hemisphere as he
travels north during the time between the vernal and autumnal equinoxes.

We presumed that such physical factors of the environment as tempera-
ture, relative humidity, and light intensity were the stimuli which regulated
the activity and rest periods of these flies, but having provided ourselves
with thermometer, hygrometer, and heht meter we were quickly convineed
that the data recorded by these instruments did not reveal the whole story.
Too often there had been no changes or only minor ones in the factors just
mentioned when there would be a sudden termination to the mid-day rest
period. For instance, where at most only an oceasional platypezid had been
seen during the previous two hours or so, we were suddenly confronted with
the problem of which fly to catch.

Considering this matter, we began to think about the discoveries which
have been made in recent years in connection with the cireadian phenomena

VoL. XXXII] KESSEL AND KESSEL: PROTOCLYTHIA BEHAVIOR 65

of insects and other organisms (von Frisch, 1950; von Frisch and Lindauer,
1954; Lindauer, 1954, 1960; Lees, 1960; Pittendrigh, 1954, 1960; Bruce and
Pittendrigh, 1957; Harker, 1960; Birukow, 1960; Breamer, 1960; and Renner,
1957, 1958, 1959, 1960).2 Could it be that we were dealing with a biological
time clock, some internal mechanism which functions with a twenty-four hour
periodicity and prompts our flies to become active or inactive, depending
upon the rhythm of their particular species? If such an endogenous mech-
anism does exist, does it act alone, or is it correlated with and triggered
by certain periodically recurring exogenous factors?

Renner (1958, 1959) has shown that the time-sense of honeybees is based,
under normal circumstances, on both endogenous and exogenous elements.
He was able to show that while the insects were able to orient themselves in
time by means of an endogenous mechanism alone under laboratory condi-
tions, they used both internal and external factors when they were in their
natural surroundings. It seemed reasonable, therefore, to suspect that our
flies, which like the bees are among the highest of insects, may possess a
somewhat similar internal clock, a mechanism which is synehronized with
one or more recurrent environmental factors.

Aschoff (1960) has applied the term “Zeitgeber” to the sum-total of the
recurring components of the environment which serve to synehronize an
organism’s cireadian rhythm with its surroundings. Adapting this word to
our English terminology, it should be pointed out that the zeitgeber must be
periodically operative. An exogenous timer of this type might include alter-
nating factors such as light and darkness, continuously changing factors
such as the daily course of temperature, or even short signals which occur
only once or twice in a twenty-four hour period. However simple or complex
the zeitgeber may be, its function is to determine phase, to synchronize any
multiplicity of circadian clocks that may be present within an organism, and
to synehronize and keep in phase all of the individuals of the species which
are in the environment under consideration. As Aschoff has also pointed out,
this last function of the zeitgeber implies that all of the individuals of the
species have the same sensitivity to the timer, because if this were not the
ease the organisms would not be in phase and one would not find what we
eall a species’ pattern.

We may suppose that the rest and activity periods of platypezids indi-
eate a circadian periodicity which is triggered by a zeitgeber but, as we have
already suggested, the exogenous timer in this case must involve an environ-
mental component other than temperature, relative humidity, and light in-
tensity. Some or all of these may contribute to the zeitgeber, but we are
convinced that some other factor must be the recognizable exogenous element.

In considering what the missing component might be, we thought about

3. We are indebted to Dr. William J. Hamilton of the California Academy of Sciences for referring us to
some of this literature and also for lending us certain reprints from his personal files.

66 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

our observation that those species of platypezids which inhabit deep woods
lack a mid-day rest period. We also recalled the longer activity periods of
sun-loving platypezids as one travels northward to Alaska during the longer
days of summer. We wondered, therefore, if the altitude of the sun could be
the critical element of the zeitgeber which had been missing from our data.
Perhaps the time-sense of our flies which lets them know when to begin and
when to terminate their morning and afternoon activity periods is dependent
upon a cireadian periodicity which is triggered by a “built-in” sextant. In
order to test this hypothesis of a sun-oriented rhythm we decided to add a
sextant to our equipment. Dr. G Dallas Hanna of the California Academy of
Sciences advised us to use a bubble sextant Inasmuch as we would usually be
working where it would be impossible to observe the horizon. We are in-
debted to Dr. Hanna for the loan of such an instrument, the one we have
been using to obtain our sun-altitude readings.

The rest period for Protoclythia californica at Steckel Park on December
20, 1961, began at approximately 11:00 A.M. At that time a male alighted
on the Solanum plant under the “chimney” where the copulating pair had
been captured the day before. He walked about for a second or two and then
settled down near the edge of the leaf with his head directed toward its tip
(fig. 3). This was a characteristic stance and position on the leaf for a
platypezid during the mid-day rest period as we had observed it in several

Figure 3. The observed male of Protoclythia californica may be located sitting
along the margin of the large leaf where he was difficult to see and maintained his
position for the entire mid-day rest period.

Vou. XXXII] KESSEL AND KESSEL: PROTOCLYTHIA BEHAVIOR 67

species of Clythia. These insects always seem to rest at or very near the edge
of the leaf and with the head pointed toward its tip. Often they take up a
position at the very tip of the leaf. The protective value of these marginal
sites is evident when one considers the hazards to which the leaf margin is
subject and the leaf sears which result therefrom. The fact that the fly sits
with his body parallel to the leaf veins also adds to his protection. While at
such times the insects show no interest in feeding and are not easily dis-
turbed, collecting them during this rest period is not ordinarily profitable
because they are so easily overlooked.

It was evident almost immediately that the male before us had settled
down for his mid-day rest. At noon he had shown absolutely no movement
and no other platypezids had been observed. This inactivity continued until
12:55 P.M. when the first signs that the rest period was ending became evi-
dent. While the male under observation still sat undisturbed, two other
males were noticed walking about on other plants in the area. A third male
was observed dancing alone in the “chimney.” Data for the several times
mentioned in this account are given in table 1.

At 1:05 P.M. the male under special observation shifted himself and
walked around to the opposite side of the leaf, this time taking up a position
with his head directed toward the petiole. Shortly thereafter he flew away
but we were unable to observe whether or not he entered the “chimney.” By
this time there were several flies dancing there and by 1:15 o'clock their
number had increased to more than a dozen.

Our observations at Steckel Park indicate therefore that the mid-day rest
period for Protoclythia californica on December 20, 1961, began at approxi-
mately 11:00 A.M. and terminated shortly after 1:00 P.M. Because of the
symmetrical shape of the sun’s altitude curve, each altitude achieved during
the day, except that of noon, oceurs twice that day, once in the morning and
once in the afternoon. Therefore, the altitude of the sun at 11:00 A.M. and
at 1:00 P.M. would be the same on any one day, providing one is dealing
with solar time. Our data given in table 1 are only approximate as noted in
the explanation. These generalized data, plus the fact that for the most part
we are dealing with a single set of observations made in one place under one
set of conditions, justify only generalized conclusions. We present these
here in the nature of a preliminary report.

The sextant reading taken at 11:00 o’clock when the male under special
observation began his siesta was 31°20’. At the time he flew away, the sun
had declined to an altitude of 30°35’. Because the male in question did not
“wake up” until after some of the other protoclythias in the vicinity had be-
come active, we must conclude that not all of a species’ individuals in an area
begin and end their rest period at the same instant. It may seem to the col-
lector in the field that they do, but it is evident that we must allow a little
individual variation even in response to a biological clock such as we are

68 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

dealing with here. Allowing this at the beginning and again at the end of the
rest period, we approximate the angle of 30 degrees as the altitude of the
sun which indicates the onset and the termination of the siesta time, as well
as the conclusion of the morning and the beginning of the afternoon activity
periods.

It has been stated that the first protoclythias observed in the morning ap-
peared at 9:05 o’clock at which time the angle of the sun was 20°10’. It has
also been noted that the last flies seen in the afternoon disappeared at 3:10
o’clock when the angle of the sun was 17°45’. Until then some of the males had
been dancing in the “chimney” although their numbers had been diminishing
for some minutes. The last flies to be seen walking or running about on the
leaves had been observed 15 minutes earlier. Because no swarming of the
males occurred during the morning and this phenomenon has been shown
to be a part of the mating behavior, we may regard it as something special
which is not entirely regulated by the same factors which control the regular
activity periods. While it is true that some of the males remained in the
swarm until the sun disappeared behind the hills, the last flies seen walking
about and feeding on the leaves had been observed at 2:55 P.M. when the
angle of the sun’s altitude was 19°40’. This figure agrees well with the 20°10’
of the sextant reading when the insects began their activity-feeding period
in the morning. Our data, therefore, lead us to the generalization that the
times of activity for Protoclythia californica oceupy the periods when the
altitude of the sun is between the angles of 20 and 30 degrees.

Owing to the lower maximum altitude of the sun and its less abrupt
ascendancy and descendaney during the longer summer days in more north-
ern latitudes, there is a longer period each morning and afternoon in such
regions during which the angle of the sun is between 20 and 30 degrees. On
this basis one would expect that the activity periods of the sun-loving species
of platypezids would be correspondingly extended so as to both begin earlier
and last later. This result would be in accord with our observations that the
activity periods of Clythia cinerea in Alaska are longer than those of the
same species in California at the same time of the summer. This extension
of the activity periods obviously reduces the length of the rest periods. In
fact we may suppose that for those platypezids which respond to the same
sun angles in the same ways as do protoclythias, the mid-day rest period
would not only be abbreviated in Alaska, but actually eliminated wherever
the altitude of the sun does not pass above the 30-degree mark.

Examination of table 1 does not suggest that the temperature factor is
directly involved with the onset and termination of the activity periods. The
same may be said for the data given in the two light-intensity columns. On
the other hand the figures on relative humidity reveal that there was an
abrupt 5 per cent drop in the value of this factor between 10:30 and 11:00
A.M., just prior to the beginning of the mid-day rest period. However, the

VoL. XXXII] KESSEL AND KESSEL: PROTOCLYTHIA BEHAVIOR 69

data show no immediate return at the conclusion of the rest period to the
higher relative-humidity reading of 72 per cent obtained at 10:30 o’clock.
This figure was reached by 2:55 P.M. but that was when the afternoon
activity period was ending, not beginning. We may conclude, therefore, that
the data support our general field impressions. Relative humidity, like tem-
perature and light intensity, seems to play no direct role in the regulation
of the activity and rest periods of platypezids. Possibly, acting together with
temperature and barometric pressure to determine the saturation deficit, it
may participate in the zeitgeber. There is little doubt, however, that the
data here reported point to the angle of the sun’s altitude as the time signal
or chief exogenous regulator of the circadian activity-rest periodicity in
platypezids.

Granting that these flies do possess such a cireadian periodicity, one may
well ask what the value of it could be to the species. By its very nature, such
a rhythm involves innate factors, so we must conclude that it is a physiologi-
cal phenomenon with a genetic basis. The species’ pattern of restricted
feeding-activity periods, with rest intervals between them, is likely to possess
a survival value. In considering what this advantage to the species might be,
it is logical to determine first what are the chief natural enemies of the flies.
There seems to be no doubt that there are birds, those species which frequent
the marginal woods which also constitute the filtered-sun habitats of the
platypezids here being considered. If the birds, too, have periods of greater
and less activity, then it is advantageous to the flies to avoid those intervals
when the birds are more active. When platypezids are running about, they
are among the most obvious of insects considering their small-to-medium size.
So unique is their manner of movement, running to and fro in a series of
darts and stops, that it is often easy to spot one of them from a distance of
many feet. While this attention-attracting behavior constitutes a welcome
advantage to the entomologist who is searching for these relatively rare in-
sects, it would seem to constitute a real disadvantage to the flies in the matter
of their bird predators. But this disadvantage would be overcome, in part at
least, if the flies would alternate activity periods with the insectivorous and
omnivorous birds which occupy the same habitat.

On our extensive collecting trips, during which we eamped out most of
the time, we have gained the impression that in general birds are early
risers and, after the night’s fast, spend the first several hours of the morning
in search of food. When there is an abundance of food, many of the birds
cease their hunting in the middle of the morning and retire for a rest period.
Contrasting platypezids with birds, we have seen that these flies are late
risers and do not make their appearance in numbers until the middle of the
morning, or about the time that the number of feeding birds is noticeably
diminished. It is apparent, therefore, that while the flies are feeding, many
of the birds are likely to be resting. Late in the morning and continuing

70 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER.

until the early afternoon comes the platypezids’ mid-day rest period during
which time there may be another surge of active birds. Once again, in the
early afternoon, fewer birds are in evidence and the flies come out for an
activity period. General feeding time for the birds comes once more in the
late afternoon, but by this time the platypezids have disappeared for the day.

In order to see if there might be some credibility to this hypothesis, we
decided to keep a record of the birds which visited the feeding station at our
home in Novato, California, on a particular day which happened to be Feb-
ruary 17, 1962. The feeding platform is located in ideal platypezid territory
along the margin of virgin woods where we have often observed flat-footed
flies of several species. The trees making up the woods are mostly oaks and
California laurels, and underneath them is an understory of low bushes. All
birds visible in the immediate area at the time of a count were included, re-
oardless of whether they were on the platform, or on the ground, or in the
bushes. For the period between 8:00 and 10:00 A.M. and after 3:00 P.M.,
the counts were made every 15 minutes. From 10:00 A.M. to 3:00 P.M. they
were made every 5 minutes.

The results of the count (table 2) fit in well with the hypothesis which we
have proposed. The expected morning surge was very evident with its aver-
age of 17.4 birds per count. Then as the morning activity period for the
flies began at 9:00 o’clock, the number of birds dropped off abruptly. For
the entire fly-activity period ending at 11:00 A.M. the birds figures remained
rather low with an average of 2.44 per count. During the mid-day rest
period for the flies, from 11:00 until 1:00 o’clock, there was a substantial
increase of some 50 per cent in the average number of feeding birds present
to give a figure of 3.74 per count. Between 11:30 and 11:55 there was a
noticeable rise in the counts, with a maximum number of 12 birds represent-
ing 4 species (junco, brown towhee, titmouse, and chickadee). These figures
indicate that it would be worthwhile for the platypezids to abstain from en-
eaging in their attention-attracting movements during this period of greater
activity for the birds. For the afternoon activity periods of the flies, from
1:00 to 3:00 o’clock, the average number of birds per count dropped to 2.56.
During this interval the birds which we saw appeared to be more lethargic
and one even seemed to be asleep on the feeding platform for several minutes.
After 3:00 o’clock, when the second fly-activity period of the day was ended,
there was no evident increase in bird numbers until about 4:15 P.M. At
4:45 o'clock when the last count was made, there had been an increase to 10,
perhaps representing the beginning of the expected surge in numbers of the
evening feeding period of the birds. Although the average number of birds
per count had increased to 3.12 between 3:00 and 4:45, this increase was so
slow in coming that as far as danger from birds is concerned these figures
cive little reason for the flies to go into hiding as early as 3:00 P.M. Of
course, if the activity periods of these insects are determined on the lower

Vou. XXXII] KESSEL AND KESSEL: PROTOCLYTHIA BEHAVIOR Tal

Table 2
Number of birds observed at different times of the day at a feeding station located
at Novato, California, and recorded on February 17, 1962, together with the average
number of birds per count for the several periods which are indicated.

Tea Sid he Number Period and Sai Number Period and
average average

8:00 A.M. 12 12:35 3

Seeley 15 Morning bird- 12:40 oY Mid-day rest
8:30 22 activity 12:45 4 period for
8:45 18 period. 12:50 4 flies (cont. )
9:00 20 Average 17.4 12:55 4

9:15 4 1:00 0

9:30 3 ILS(05) 3

9:45 1 1:10 6

10:00 7 Morning fly- 12315) 3

10:05 0 activity 20 1

10:10 2 period. ISPs) 2

10:15 4 Average 2.44. 1:30 4

10:20 2 Decrease in 135 2
10:25 5 bird activities. 1:40 2 Afternoon fly-
10:30 2 1:45 2 activity
10:35 3 1:50 2 period.

10:40 2 155 3 Average 2.56.
10:45 il 2:00 3 Decrease in
10:50 i ZEOS 3 bird activities.
1OESS 2 2:10 2

11:00 0 2:15 3

“Sonos hice ane ener ae a Bee ne eo ee 2:20 2

11:05 5 PAR DAS: 3

Tabeak(t) 4 220 3

ilibga ls i Pa ay5 2

11:20 1 2:40 1
11325 0 Mid-day rest 2:45 5

11:30 4 period for 2:50 3

ULERY 12 flies. 2:55 3

11:40 8 Average 3.74. 3:00 1

11:45 4 WTAGTESHSS) WN Scene coc temo nec teeeten sedan cones eceeeceearSna sa vieceanct fot coeatecs uote
11:50 6 bird activities. BIS 155 il

WAL 55s 6 3:30 2

12:00 M 7 Betsy 0 Evening bird-
127105 P.M. 2 4:00 2 activity

12:10 0 4:15 5 period.

ne ppltey 4 4:30 4 Average 3.12.
12:20 2 4:45 10

12:25 Fn eenenee cence ncen ce cenencececetnensecacoentasaceecenansesececeositenencoseeenestics
12:30 3

level by a particular angle of the sun’s altitude as we have postulated, then
this could not be decreased for the afternoon period without likewise de-

72 CALIFORNIA ACADEMY OF SCIENCES [Proc. 471 SER.

creasing it for the morning period. And this would put the flies into greater
jeopardy by forcing their early activities into conflict with the morning
surge of feeding birds.

It is recognized, of course, that to be wholly significant the counts of
birds observed feeding at different times during the day should have been
made in the same place and same time of year as the fly study with which
they are compared, not 300 miles or so more to the north and 2 months later
as they were. But because it was impossible to make the counts at Steckel
Park on December 20, these less-desirable data from Novato and recorded on
February 17 are substituted.

Our comments concerning the attention-attracting movements of platy-
pezid flies as they run about on leaves in their unique jerky tempo, deserve
additional comment. One cannot help but wonder why such a jeopardy-
inviting activity has been maintained by these flies. Its disadvantages are so
obvious when we consider the matter of bird enemies. Does this habit seem-
ingly have any conpensating advantages?

Howlett (1921) is of the opinion that it does have such advantages. While
he agrees that the unique movements of flat-footed flies are attention arrest-
ing, and therefore can hardly make these insects less conspicuous to such
important enemies as birds, nevertheless he believes that they have a pro-
tective value in that they make the capture of the fly more difficult than if it
were sitting still or “standing” in the air. Howlett also cites the cases of
several arthropods which supposedly obtain protection by means of their
obvious tetanie actions. He refers to several of these which, when disturbed,
eause the body to vibrate so rapidly that the observer sees only a blurred
area. He thinks that it is likely that in platypezids “the rapidly alternating
translatory motion of the whole insect may be another means of achieving a
similar result.”

While Howlett attempts to make a convincing ease of these points, our
observations indicate that he is wrong in his major premise that the darting-
stopping movements of platypezids are executed only when the flies are dis-
turbed by the movement of some object in the vicinity. Over and over again
we have eautiously approached a likely looking “platypezid bush” from
behind good cover and from our vantage point observed several flies vari-
ously engaged. While at the moment some were quietly feeding, the others
were moving about in their to-and-fro fashion searching for food. A few
minutes later and without cause for alarm, the feeders might be running and
the runners quieted down. We can hardly agree, therefore, with Howlett’s
contention that the movements are wholly defensive maneuvers.

Platypezids are often extremely quick in movement. This is particularly
true of their departure flights, that is, when they leave the immediate vici-
nity, not just move to an adjacent leaf. They may have been “floating” in
the air in zigzag fashion, “standing” practically motionless in the air, or

Vou. XXXII] KESSEL AND KESSEL: PROTOCLYTHIA BEHAVIOR 73

running or sitting on leaves. In any case their departure is instantaneous,
without any discernible preparatory movements or attitudes. It is also
instantaneous in the actual departure which is so rapid that the eye can very
seldom follow it. It can be truthfully said in almost every instance that “if
you saw it leave it was not a platypezid.” Most frequently the fly leaves
completely, that is, it does not just fly a few feet or yards and then settle
down or return.

Very few, if any, of the dipterans that we have encountered have this
habit of complete and rapid departure, or at least utilize it at all frequently.
In the platypezids it is used on many occasions: when they are disturbed by
a sudden breeze, another insect, a bird, or a falling twig or leaf, sometimes
even by abrupt exposure to bright sun or heavy shadow. It is also used when
a person causes a disturbance by close approach, considerable movement, or
shaking the foliage. Significantly, however, the same method is followed
when the most critical observation can discern no disturbing condition; this
is just the usual manner in which platypezids depart from any particular
spot.

A point of special interest in connection with the departure flight is the
fact that it can take place in the midst of any of the fly’s normal activities.
As already mentioned, flies leave from a place in the air or from a leaf.
Furthermore, when from a leaf this flight may occur in an entirely typical
manner whether the fly has been busily feeding at one spot, running rapidly
over a leaf, or remaining very quiet during a rest period. In any of these
cases the fly is there and the next instant it is not. It should be pointed out,
however, that quite logically these flies spontaneously take flight less often
and are disturbed least by external factors when they are quietly feeding or
in a rest period, or when it is relatively dark and cold. Departure flights are
more frequent and most easily caused by disturbance when the flies are
running on leaves, or “standing” in the air, or when the light intensity and
temperature are high.

In considering the subject of external disturbance of platypezid activity,
aside from the relation of actual departure flights, it can be said that any
one of many nonextreme factors may bring about a change of activity. This
is most easily seen when a fly is busy feeding in one spot and, when disturbed
only slightly, begins to run in typical jerky fashion over the leaf. It should
be remembered in this connection that the fly’s feeding activity on the leaf
consists normally of two phases, one of more or less rapid running and at the
same time testing for the presence of food and the momentary stopping to
actually take food when it is found in tiny amounts, the other phase involv-
ing feeding at one spot of greater abundance. When the flies are undisturbed
these two phases alternate with no regular pattern whatsoever. The two
phases together make up the general pattern and shift according to the cir-
cumstances. One of the conditions which seem capable of serving as the

74 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47m Ser.

stimulus for a temporary shift to the running phase, yet without breaking
the pattern, is some slight external disturbance such as the casual yet obvious
approach of a person. Howlett’s observations, limited to a study of two
platypezids, led him to interpret this shift to the running phase as a special
defensive maneuver.

Perhaps the running behavior of flat-footed flies does have some such
protective value as Howlett has suggested. Our opinion, however, is that this
is of minor importance, if indeed it has any such significance. Instead, we
are convinced that the advantage to the species of this display of so much
haste and such an expenditure of energy is correlated with the search for
food. Platypezid flies are not predaceous, so they need not run rapidly to
eatech prey. But they do glean their nourishment from inconspicuous accumu-
lations on such flat surfaces as leaves, and because of the limited quantity
of this food often available to them they have survived by the utilization of
an innate capacity to scrounge for their food. Such scrounging becomes most
efficient when the movements are rapid and to the point of discovery.

That platypezids utilize their characteristic jerky movements when they
are completely undisturbed and their only motivation is the search for food
was made very clear to us by the following laboratory observations. Larvae
of both Calotarsa insignis and Clythia agarici had been collected in large
numbers and placed in rearing cages. When it came time for the insects to
emerge as adults, so many of them appeared that after we had mounted and
preserved all that we could use, a hundred or more flies of each species were
left over. So we decided to release them in the laboratory and to observe
their behavior.

The room was a small one measuring about 7 by 14 feet. There were no
windows and the only natural light was from a small skylight at one end of
the ceiling. Artificial illumination was provided by two overhead fixtures
and a desk lamp. Many of the flies had emerged as long as two days previ-
ously and none of them had received nourishment. When these hungry flies
were released they dispersed and within a few minutes they seemed to be
everywhere, on the floor, bookease, collection ease, filing cabinet, and desk.
And everywhere they exhibited the same to-and-fro, stop-and-go search for
something to eat. As we had expected, few of them went to the skylight;
instead, most of them went to the lower levels of the room, particularly the
desk and the floor. It is of interest to note that this preference for the lower
levels agrees with their lower-story habitats in woods.

Although all of the flies seemed to us to ignore our presence and to be
concerned only with their search for food, one might argue that the reverse
was true and that in conformity with Howlett’s theory they were exhibiting
their incessant erratic movements because they were in artificial surround-
ings and very much disturbed. But this possibility was eliminated very soon
when some of the flies on the desk happened upon the cleared area where

VoL. XXXII] KESSEL AND KESSEL: PROTOCLYTHIA BEHAVIOR 75

we had just eaten our sandwich lunch. All of the crumbs had been brushed
up and nothing of food value remained as far as we could see. But to the
flies there were spots of nourishment present and they stopped to feed on
them. The insects were so intent on their feeding that they were undisturbed
when we placed an open-base-type binocular microscope over them and
brought them into focus. By this means we were able to see them feeding in
house-fly fashion on the film spots of food which they had discovered on the
desk. But finishing in one spot they were always in a hurry to find another,
and in so doing fell into their familiar pattern of movement.

We are therefore convinced that these peculiar motions indulged in by
platypezids are not defensive measures such as to make them more difficult
to capture or to scrutinize, but rather are only a part of their search-for-food
pattern. If by such movements the flies also run the danger of attracting the
attention of birds, that is a hazard which the species must endure. If this
hazard has been reduced by the expedient of alternating feeding periods with
birds, this constitutes a survival factor worth perpetuating. But the busi-
ness of getting food must go on.

SUMMARY

The mating behavior of Protoclythia californica here described involves
only the second oceasion on which mating pairs of flat-footed flies have been
studied; the first observations dealing with Platypezina pacifica having al-
ready been reported by us. Owine to the lower height of the willow trees
which outlined the swarming “chimney” of P. californica as compared with
the very tall redwoods of the platypezina “chimney,” the level of the swarm
was much lower in the ease of the protoclythias. This made it possible to
study them at very close range and thereby gain a clearer picture of the
epigamie behavior of these flies. It is evident that the matine behavior of
these two species is essentially the same and presumably it is representative
not only of the two genera to which they belong but also of the family
Platypezidae.

The males gather in swarms to dance in a “chimney” outlined by trees.
Here they wait for individual females to approach the swarm. While the
females have always been observed to ignore or repulse the advances of males
in other situations, they are positively erotropic once they recognize that the
swarm represents their own species. The receptive female enters the swarm
of males and is grabbed by the first one she approaches in her line of flight.
Copulation is initiated in the swarm and union is accomplished with the
heads of the flies pointing in opposite directions. The female assumes the
initiative in flight, the male being towed backward as the pair descend to
complete their nuptial activities on a lower-story shrub below the “chimney.”
The female continues to haul the male around as she walks over the leaves in
search of a suitable resting place. The peculiar mating position assumed by

76 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

these flies seems to be correlated with the nature of the male genitalia which
are rotated 180 degrees so as to be directed downward and forward.

Long experience in collecting flat-footed flies convinced us that the more
sun-loving species which inhabit the areas along the margins of woods have
well defined morning and afternoon periods of activity, with a rest period
occupying the mid-day interval between them. We were impressed by the
rather sudden onset and abrupt termination of these periods and also by our
observation that the activity periods for a particular species are longer in
the more northern latitudes.

It is our opinion that the activity-rest periodicity of these flies is regu-
lated by a cireadian rhythm whieh functions as a biological clock on a twenty-
four hour basis. Searching for the exogenous factors which may trigger this
presumed endogenous mechanism, we concluded that temperature, relative
humidity, and hght intensity do not qualify for this role. Only the altitude
of the sun seems to be identifiable as an exogenous timing element.

Observations made at Steckel Park, Ventura County, California, on De-
cember 20, 1961, indicate that the activity times of this species coincide with
the periods when the sun’s altitude is between the angles of 20 and 30 de-
orees. On the day and at the place stated, the morning activity period lasted
approximately from 9:00 to 11:00 o’clock, the rest period from 11:00 until
1:00 o’clock, and the afternoon aetivity period from 1:00 to 3:00 P.M.

In consideration of what value this periodicity may possess for the flies,
we propose that it is advantageous for them to alternate activity periods
with birds, which are their chief predators. Our general impression, gained
during mueh field work and supported by limited data, is that birds tend to
be early risers and spend the first hours of the day in search of food, after
which they may retire for a rest period. Platypezids, by contrast, are late
risers and do not make their appearance in numbers until the middle of the
morning, so while the flies are feeding many of the birds are resting. When
the flies take their mid-day siesta, there may be another increase of feeding
birds. As the birds once more become less active in the afternoon, the flies
appear again for an activity period. General feeding time for the birds is
evident again in the late afternoon, but by this time the platypezids have
retired for the day.

Platypezids characteristically attract attention by their to-and-fro stop-
and-go0 movements as they run about on leaves. It was proposed long ago
that these darting-stopping movements are executed only when the flies are
disturbed and that they have the protective value of making capture more
difficuit. We cannot agree with this viewpoint. Instead, we hold that the
jerky running of these flies is merely a phase of their feeding activity, a part
of their search-for-food pattern. Adult platypezids glean their nourishment
from inconspicuous accumulations on such flat surfaces as leaves and, be-
cause of the limited supply, they must often serounge for their food. Such

Vou. XXXII] KESSEL AND KESSEL: PROTOCLYTHIA BEHAVIOR 77

serounging becomes most productive when the movements of the search are
rapid and the testing stops are brief.

LITERATURE CITED

ASCHOFE, J.

1960. Exogenous and endogenous components in circadian rhythms. Cold Spring

Harbor Symposia on Quantitative Biology, vol. 25, pp. 11-28.
BIrRUKOw, G.

1960. Innate types of chronometry in insect orientation. Cold Springs Harbor

Symposia on Quantitative Biology, vol. 25, pp. 403-412.
BREAMER, W.

1960. A critical review of the sun-azimuth hypothesis. Cold Spring Harbor

Symposia on Quantitative Biology, vol. 25, pp. 412-427.
Bruce, V. G., and C. S. PITTENDRIGH
1957. Endogenous rhythms in insects and microorganisms. American Natural-
ist, vol. 91, pp. 179-195.
FRiscH, K. von
1950. Die Sonne als Kompass in Leben der Bienen. Experimentia, vol. 5, p. 142.
Friscu, K. von, and M. LInDAUER

1954. Himmel und Erde in Konkurrenz bie der Orientierung der Bienen. Die

Naturwissenschaften, vol. 41, pp. 245-153.
HARKER, JANET EH.

1960. Endocrine and nervous factors in insect circadian rhythms. Cold Spring

Harbor Symposia on Quantitative Biology, vol. 25, pp. 279-287.
How tert, F. M.

1921. Protective movements and range of vision in platypezid flies. Report of
the Proceedings of the Fourth Entomological Meeting Held at Pusa on
the 7th to 12th February, 1921. Calcutta. Superintendent Government
Printing, India. Pp. 279-286.

KESSEL, HE. L.

1948. A review of the genus Platypezina Wahlgren, announcement of its presence
in the New World, and the description of a new species (Diptera:
Clythiidae). Wasmann Collector, vol. 7, pp. 47-64.

1955. The mating habits of balloon flies (Diptera: Empididae). Systematic
Zoology, vol. 4, pp. 97-104.

1959. Introducing Hilara wheeleri Melander as a balloon maker, and notes on
other North American balloon flies (Diptera: Empididae). Wasmann
Journal of Biology, vol. 17, pp. 221-230.

Kesser, E. L. and J. V. KARABINOS

1947. Empimorpha geneatis Melander, a balloon fly from California, with a
chemical examination of its balloons (Diptera: Empididae). Pan-
Pacific Entomologist, vol. 23, pp. 181-192.

KEsseEL, KE. L. and Berta B. KESSEL

1951. A new species of balloon-bearing Hmpis and an account of its mating
activities (Diptera: Empididae). Wasmann Journal of Biology, vol. 9,
pp. 137-146.

1961. Observations on the mating behavior of Platypezina pacifica Kessel (Dip-
tera: Platypezidae). Wasmann Journal of Biology, vol. 19, pp. 295-299.

78 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4rH SER.

Lees, A. D.
1960. Aspects of animal photoperiodism. Cold Spring Harbor Symposia on
Quantitative Biology, vol. 25, pp. 261-268.

LINDAUER, M.

1954. Dauertanz im Bienenstock und ihre Beziehung zur Sonnenbahn. Die
Naturwissenschaften, vol. 41, pp. 506-507.

1960. Time-compensated sun orientation in bees. Cold Spring Harbor Symposia
on Quantitative Biology, vol. 25, pp. 371-377.

PITTENDRIGH, C. S.

1958. On temperature independence in the clock-system controlling emergence
time in Drosophila. Proceedings of the National Academy of Sciences,
vol. 40, pp. 1018-1029.

1960. Circadian rhythms and the circadian organization of living systems. Cold
Spring Harbor Symposia on Quantitative Biology, vol. 25, pp. 159-184.

RENNER, M.

1957. Neue Versuche iiber den Zeitsinn der Honigbiene. Zeitschrift fiir verglei-
chende Physiologie, vol. 40, pp. 85-118.

1958. Der Zeitsinn der Arthropoden. Ergebnisse der Biologie, vol. 20, pp. 127—
158.

1959. tiber ein weiteres Versetzungs experiment zur Analyse des Zeitsinnes und
der Sonnenorientierung der Honigbiene. Zeitschrift fiir vergleichende
Physiologie, vol. 42, pp. 449-483.

1960. The contribution of the honeybee ito the study of time sense and astro-
nomical orientation. Cold Spring Harbor Symposia on Quantitative
Biology, vol. 25, pp. 361-367.

Snow, W. A.
1894. American Platypezidae. Kansas University Quarterly, vol. 3, pp. 143-152.
VERRALL, G. H.

1901. Platypezidae, Pipunculidae, and Syrphidae of Great Britain. British Flies,

vol. 8, pp. 1-691.

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

FOURTH SERIES

G Dallas Hanna Anniversary Volume

Vol. XXXII, No. 4, pp. 79-85; 12 figs. June 29, 1962

TWO GNAPHOSID SPIDERS FROM ARKANSAS'
by

Harriet Exline~

Research Associate

California Academy of Sciences

Rachodrassus echinus Chamberlin (1922; Kentucky) was proposed with
a brief deseription and without illustration; Drassylus covensis is a new
species. Both were collected by Otis Hite in pitfall traps in the forest litter
of Cove Creek Valley, Arkansas, during April and May, 1961. In so far as
has been determined, genitalia of spiders of the family Gnaphosidae are de-
seribed here for the first time.

I am indebted to Dr. Willis J. Gertsch, American Museum of Natural
History, for checking identifications, and to Dr. Herbert W. Levi, Museum
of Comparative Zoology, Harvard University, for sketches of the palpus and
epigynum of the type and allotype of Rachodrassus echinus Chamberlin and
the epigynum of the type of “R. chera Chamberlin.”

1. Work supported by National Science Foundation grant G17564, under direction of Dr. Willard H. Whit-

comb, Department of Entomology, Agricultural Experiment Station, University of Arkansas.
2. Mrs. Don L. Frizzell, Rolla, Missouri.

APR 1 § 1903
WOODS HOLE, MASS.

80 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Family GNAPHOSIDAE

Rachodrassus echinus Chamberlin.
(Figures 7-12.)

Rachodrassus echinus CHAMBERLIN, 1922, Biol. Soc. Wash., Proc., vol. 35, pp. 146, 160.
Male holotype and female allotype from near Mammoth Cave, Kentucky, in
Museum of Comparative Zoology. Type species of Rachodrassus. The specific
name is a Latin noun in apposition, meaning “sea-urchin.”

Rachodiassus chera CHAMBERLIN, 1922, Biol. Soc. Wash., Proc., vol. 35, p. 161. Fe-
male holotype from near Mammoth Cave, Kentucky, in Museum of Comparative
Zoology.

Mane. Carapace yellow covered with gray hair-like setae that lie prone
and face mid-line between thoracic groove and ocular area, and a few erect
setae. Legs, palpi, mouthparts and sternum yellow, thickly covered with
prone and erect setae. Abdomen dull yellow with an irregular gray pattern,
venter grayish yellow, spinnerets clear yellow.

Carapace with steeply inclined sides, equally high from behind thoracic
eroove to near eye region, which is lower; evenly rounded on sides, but nar-
rowed anteriorly. Chelicerae short, somewhat geniculate; anterior margin of
fang groove with three teeth, posterior margin with two teeth, all well sep-
arated and sharp. Clypeus narrow, less than a radius of anterior lateral eye.
Anterior eyes in proecurved row, anterior medians smaller than other eyes,
separated by a radius, nearer anterior laterals. Posterior eyes in a straight
row; median eyes round, slightly smaller than laterals, and nearer each other
than laterals. Median ocular area a little wider behind than long. Legs
moderately robust with many long spines; relative length 4,1,2,5. Fourth

Figure 1. Drassyllus covensis Exline, new species, male palpus, ectal view.
Figure 2. Drassyllus covensis Exline, new species, male palpus, ventral view.
Figure 3. Drassyllus covensis Exline, new species, epigynum of female.

Figure 4. Drassyllus covensis Exline, new species, female genital plate, dorsal view.

Figure 5. Drassyllus covensis Exline, new species, bulb of male palpus expanded,

dorsal view: a) median apophysis showing the two segments, c) con-

ductor, e) embolus, f) fulcrum or terminal apophysis.

Figure 6. Drassyllus covensis Exline, new species, abdomen of male.

Figure Rachodrassus echinus Chamberlin, abdomen of male.

Figure 8. Rachodrassus echinus Chamberlin, female epigynum.

Figure 9. Rachodrassus echinus Chamberlin, genital plate of female, dorsal view.

Figure 10. Rachodrassus echinus Chamberlin, male palpus, ectal view.

Figure 11. Rachodrassus echinus Chamberlin, male palpus, ventral view.

Figure 12. Rachodrassus echinus Chamberlin, tip of bulb of male palpus, lateral
view: a) median apophysis, aa) accessory apophysis of tegulum, ¢c)
conductor, e) embolus, f) fulcrum or terminal apophysis.

a
a)
Q
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os
Q
NN
D
So
ce
Q
=
zi
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3
=
—
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EXLINE

VOL. XXXII]

82 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER.

tibiae with two mid-dorsal spines, third tibiae with one; all tibiae with three
pairs of ventral spines.

Abdomen slender, without scutum. Anterior spinnerets very long, stout,
separated by their radius. Posterior spinnerets shorter, slender, with very
short terminal segment.

Palpal tibia longer than patella, with a short, robust, ventrally pointed
apophysis. Tarsus long, narrow, with bulb thick. Bulb with terminal ful-
erum (terminal apophysis) bent back in a double curve at tip; embolus
heavily sclerotized, broad, with three distal teeth, the median one bearing
duet; conductor membranous; median apophysis large, with a broad basal
tooth and distal hook. A dorsal projection of the tegulum has a large, mem-
branous folded tip. When expanded artificially, at least, the fulerum en-
velops and grasps the embolus.

Total length, 6.5 mm., 5.5 mm. without chelicerae and spinnerets. Cara-
pace 2.3 mm. long, 2.0 mm. wide. First leg: femur, 2.1 mm.; patella-tibia,
3.0 mm.; metatarsus, 1.6 mm., tarsus, 1.2 mm. Second leg: femur, 2.0 mm.;
patella-tibia, 2.6 mm.; metatarsus, 1.4 mm.; tarsus, 1.1 mm. Third leg:
femur, 1.7 mm.; patella-tibia, 2.2 mm.; metatarsus, 1.5 mm.; tarsus, 1.0 mm.
Fourth leg: femur, 2.3 mm.; patella-tibia, 2.9 mm.; metatarsus, 2.6 mm.;
tarsus, 1.2 mm. The eight males range from 5.5 mm., in total length, to
6.5 mm.

Fema.e. Color, pattern, and structure similar to male. Chelicerae more
eeniculate; posterior eye row a little recurved, and eyes farther apart; ocular
area much wider behind than in front; first tibiae with only two pairs of
ventral spines, second tibiae with two pairs and one distal; anterior spin-
nerets separated by a diameter.

Epigynum swollen with a long, wide anterior fold, with deep grooves
bordering it laterally. A pair of conspicuous horizontal ridges posterior to
fold. Grooves anterior to ridges are confluent with grooves on each side of
anterior fold. A pair of large, contiguous, heavily sclerotized, oval seminal
receptacles lies in the posterior half. Three small, deep, sclerotized depres-
sions occur, one over each seminal receptacle and one between them. If the
genital plate is removed, dissected, and cleared, a slender, irregular tube can
be seen on each side, opening at the anterior end of the groove bordering the
anterior fold. At the level of the lateral ridges, this enters a large sclerotized
oval body, and passes posteriorly to the seminal receptacle, where it becomes
affixed as a wide, sclerotized anterior part. The structure and function of the
parts are not entirely understood. The tube is only wide enough for the duct
of the embolus to enter. There seem to be no openings anterior to the lateral
ridges into the sclerotized bodies. The lateral ridges may act as a lock for
the combined radix and embolus, the duct of the latter becoming free and
following the grooves to the small anterior opening.

Total length, 6.8 mm.; 6.1 mm. without chelicerae and spinnerets. Cara-

VoL. XXXIT] EXLINE: TWO GNAPHOSID SPIDERS 83

lord

pace, 2.6 mm. long, 2.1 mm. wide. First lez: femur, 1.7 mm.; patella-tibia,
2.6 mm.; metatarsus, 1.1 mm.; tarsus, 0.8 mm. Second leg: femur, 1.7 mm.;
patella-tibia, 2.6 mm.; metatarsus, 1.1 mm.; tarsus, 0.8 mm. Third leg: femur,
1.6 mm.; patella-tibia, 2.2 mm.; metatarsus, 1.4 mm.; tarsus, 0.8 mm. Fourth
leg: femur, 2.3 mm.; patella-tibia, 3.0 mm.; metatarsus, 2.4 mm.; tarsus, 1.2
mm. The five females range from 5.6 mm. to 6.8 mm. in length.

Rachodrassus chera Chamberlin is synonymized with R. echinus as a
paucity of spines was the basis for separation of R. chera, and the spination
of the ventral tibiae proves to be quite variable. The epigyna of the two
nominal species are very similar, although the grooves of that of R. chera
are darker. One female taken in Arkansas also has darkened grooves, filled
with chitin or a resinous material. Rachodrassus flavus Chamberlin and
Woodbury (1929, p. 134, pl. 2, figs. 1, 2; erroneously given as plate 1), from
Utah, has the anterior eye row straight; the male palpus seemingly is simpler
than that of R. echinus, with a flat tibial apophysis rounded at the tip; and
the female epigynum has a quite different appearance.

Drassyllus covensis Exline, new species.
(Figures 1-6.)

Holotype male and allotype female from Cove Creek, Washington County,
Arkansas, collected by Otis Hite, May 13, 1961, in collection of California
Academy of Sciences. The specific name is a Latinized adjective based on the
type locality.

Mate. Carapace, mouthparts, sternum, and some leg segments pale
orange with light gray infusions. Distal part of femora, patellae and tibiae
mostly dark gray. Abdomen grayish white, nearly covered with a blackish
eray pattern; basal scutum orange overcast with blackish gray. Venter pale
with a pair of large grayish black patches uniting in front of spinnerets.

Carapace low, evenly arched, highest at posterior end of thoracic groove,
wide with sides evenly curved, somewhat narrowed anteriorly; sparsely
covered with setae that incline toward thoracie groove. Clypeus vertical,
height about equal to diameter of anterior median eyes. Posterior eye row
procurved; posterior median eyes oblique, very slightly larger than posterior
laterals, all separated by less than a radius. Shape and spacing of eyes some-
what variable. Anterior median eyes smaller than others, separated by nearly
a diameter. Median ocular area a little longer than wide, slightly wider
behind.

Chelicerae geniculate, slender, with elongate boss. Anterior margin of
fang furrow armed with 3-5 small dark teeth; posterior margin with three
well spaced, pale denticles.

Legs with metatarsi and tarsi slender, tibiae IV slightly enlarged. Tibiae
IIT and IV without median dorsal spines. Two, a basal and median, ventral

84 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4ru Ser.

spines on tibiae I and II; three pairs of ventral spines on tibiae IIT and IV.
Spines on anterior tibiae are not present on all males. Legs 4,1,2,5 in rela-
tive length.

Abdomen with triangular seutum on basal half. Anterior spinnerets
separated by their diameter. Posterior spinnerets equal to anteriors in length,
much more slender, with a short terminal segment.

Palp with patella longer than tibia without apophysis. Tibial apophysis
extending a third eymbial length, broad at base, tapering to a point, bent
dorsally near tip. In ventral and lateral views of the unexpanded bulb, the
conspicuous structures are parts of the two-jointed, protruding median
apophysis. The long, slender, curved embolus lies against the eymbium. The
terminal apophysis or fulerum is short, blunt at tip with notched margin,
only the tip showing in ventral view. The conductor, invisible in the unex-
panded bulb, is a thin plate with only the edge sclerotized, lying between the
median apophysis and embolus.

Total length, 4.6 mm. including chelicerae and spinnerets. Carapace, 1.9
mm. long, 1.5 mm. wide. First leg: femur, 1.5 mm.; patella-tibia, 2.0 mm.;
metatarsus, 1.0 mm.; tarsus, 0.8 mm. Second leg: femur, 1.3 mm.; patella-
tibia, 1.6 mm.; metatarsus, 0.7 mm; tarsus, 0.7 mm. Third leg: femur, 1.0
mm.; patella-tibia, 1.3 mm.; metatarsus, 0.7 mm.; tarsus, 0.6 mm. Fourth leg:
femur, 1.5 mm.; patella-tibia, 2.1 mm.; metatarsus, 1.5 mm.; tarsus, 0.8 mm.
Nine males range from 4.6 mm. to 5.3 mm. in length.

FremMaLe. Pattern and colors not so distinet as in male. Structure es-
sentially the same: differs by having clypeus slanting and not quite so high;
posterior median eyes about a radius apart, nearer lateral eyes; first tibiae
without ventral spines, second tibiae with one spine. Epigynum large, flat,
highly sclerotized. Openings lateral about midway, seemingly leading into
sclerotized bursae. Tube from each bursa forms an anterior lateral loop,
bordering bursa medially and forming a_ postero-lateral loop leading to
seminal receptacles on posterior edge of epigynum. Seminal receptacles con-
tiguous, with a dorsal thickening from which the fertilization tubes origi-
nate. Total length, 6.8 mm. Carapace, 1.9 mm. long, 1.4 mm. wide. First
lee: femur 1.4 mm.; patella-tibia, 1.9 mm.; metatarsus, 0.9 mm.; tarsus, 0.7
mm. Second leg: femur, 1.38 mm.; patella-tibia, 1.6 mm.; metatarsus, 0.7
mm.; tarsus, 0.6 mm. Third leg: femur, 0.9 mm.; patella-tibia, 1.3 mm.;
metatarsus, 0.7 mm.; tarsus, 0.6 mm. Fourth leg: femur, 1.5 mm.; patella-
tibia, 2.1 mm.; metatarsus, 1.5 mm.; tarsus, 0.7 mm. A second female meas-
ures 5.0 mm. length.

Drassyllus covensis is closely related to D. dentelifer Chamberlin (1936,
p. 15, figs. 1, 2) from Florida. In the latter the tibial apophysis is not so
long, the position of the embolus is more ventral, the median apophysis is
shorter, the abdomen is darker and without pattern, the lower margin of the

VoL. XXXII] EXLINE: TWO GNAPHOSID SPIDERS 85

chelicera has only two teeth, the posterior tibiae bear a median dorsal spine,
and the posterior median eyes are very large.

LITERATURE CITED

CHAMBERLIN, R. V.
1922. The North American Spiders of the Family Gnaphosidae. Biological So-
ciety of Washington, Proceedings, vol. 35, pp. 145-172.
1936. Further records and descriptions of North American Gnaphosidae. Amer-
ican Museum Novitates, no. 853, pp. 1-25, 47 text figs.
CHAMBERLIN, R. V., and A. M. WoopBury
1929. Notes on the Spiders of Washington County, Utah. Biological Society of
Washington, Proceedings, vol. 42, pp. 131-142, pls. 1-2 (explanations
transposed).

an Wer 3 “al

Sasek" yi“

TV THLE

wey mp

ae}
=i A op ~ 2 Oe hy Pe aie
t 5 ra * AW.

i. ; bu tr~iye |

a aa
Lp mis

enh el

a»

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

FOURTH SERIES

G Dallas Hanna Anniversary Volume

Vol. XXXII, No. 5, pp. 87-101; 13 figs. June 29, 1962

DISTINCTIVE *‘CONGRID TYPE” FISH OTOLITHS
FROM THE LOWER TERTIARY OF THE GULF COAST
(PISCES: ANGUILLIFORMES)'

Don L. Frizzell and C Kurt Lamber

Missouri School of Mines and Metallurgy, Rolla, Missouri

INTRODUCTION

“Congrid type” sagittae characterize some Recent, presumably anguilli-
form, fishes (Frost, 1926) and range as fossils at least from the lower Eocene.
They show genetic unity in the groups containing them, although relation-
ships even at the ordinal level have not been satisfactorily established.

Two nominal species in the Gulf Coast Tertiary are known from otoliths
of this type: “Congermuraena” sector (Koken) (1888, pp. 292-293, pl. 17,
figs. 14-16; as Otolithus (Platessae)) and “Conger” brevior (Koken) (1888,
pp. 293-294, pl. 18, fig. 7; as O. (Congeris)). “Congermuraena” sector, de-
seribed as ranging from Claiborne Eocene to Vicksburg Oligocene, includes
a complex of species that will be treated in a later paper. “Conger” brevior,
described from the Jackson Eocene of Jackson, Mississippi, is redescribed
from suites of well preserved specimens from Hocene and Oligocene strata of
Mississippi and Alabama.

1. Work supported by National Science Foundation Research grant G—-10221, 1959-1961.

[ 87 }

Marine Biological Laboratory
BIBRA NR YT

APR 1 8 1963

aankow ys

iAaminhARr iF

88 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 47H SER.

Five new species are proposed here, from Hocene-Oligocene beds of the
Julf Coast, in order to make names available for these fossils. All are re-
ferred to the very broadly based form-genus “Conger,” although only one
(C.?2 vetustus) shows a close resemblance to otoliths of the Recent genus
Conger Schaeffer. At least three genera are represented by the described
species, but comparative material now available does not justify an attempt
at generic allocation.

ACKNOWLEDGMENTS

Assistance in various phases of research on fossil fish otoliths has been
acknowledged in an earlier paper (Frizzell and Lamber, 1961). To that list
we would add: Mr. Emmett Adams, Jackson, Mississippi, who accompanied
Frizzell to the Moody’s Braneh outerop in Jackson; Mr. C. O. Ketler, Hi-
wannee, Mississippi, for loeality information; Miss Winnie MecGlamery,
formerly of the Alabama Geological Survey, for data on collecting localities;
and Messrs. J. G. MeVay and J. A. McClinton, Lone Star Cement Corpora-
tion, St. Stephens, Alabama, who made it possible to collect at the St.
Stephens quarry.

MorPHOoLoGY OF CONGRID TYPE SAGITTA

The congrid type sagitta (figure A) has a typically ovate outline, highest
toward the anterior and lowest at the posterior end, with dorsal dome (D)
developed in some forms. The suleus is not divided, although caudal and
ostial regions (CR and OR) are recognized. An anterior border (AB) sepa-
rates the ostial region from the anterior margin, and a characteristic ostial
channel (OC) opens onto the dorsal margin or anterodorsal slope.

COLLECTING LOCALITIES

The otoliths described here were found in samples from the following
localities :

OLIGOCENE, VicksBuRG GROUP.

Byram marl. Old Byram, Hinds County, Mississippi (type locality) ;
bank of Pearl River below suspension bridge; irregular beds of shell drift
within sandy shell marl of formation. Collectors: D. lL. Frizzell, August 31,
1957; D. L. and H. E. Frizzell, August 21, 1959; C K. Lamber, November 26,
1959. (Stations F—57-5; F—-59-1C; CKL-59-8, 9.)

Vicksburg, Warren County, Mississippi; small road cut on gravel road
adjacent to north boundary of National Cemetery, ca. 30 yards east of U.S.
Highway 61; highly glauconitie, indurated shell marl containing stringers of
non-indurated broken shell material. Collector: C K. Lamber, November 27,
1959. (Station CKL-—59-16.)

VoL. XXXIT] FRIZZELL AND LAMBER: FISH OTOLITHS 89

Glendon limestone. Quarry of Marquette Cement Company, about one
mile southwest of Brandon, Rankin County, Mississippi; glauconitic shell
marl above basal limestone stratum of formation; various places within
quarry limits. Collectors: D. L. Frizzell, November 26, 1959; © K. Lamber,
November 26, 1959. (Stations F—-59-13, 15; CKL-—59-3, 4, 7.)

Mint Spring marl. National Cemetery, Vicksburg, Warren County, Mis-
sissippi (type locality); falls of Mint Spring Bayou at south boundary of
cemetery; fossiliferous shell detritus about 114 feet above basal contact (with
Forest Hills formation). Collectors: D. L. Frizzell, November 23, 1959; C. K.
Lamber, November 27, 1959. (Stations F-59-6, 7, 8; CKL-—59-12, 13.)

2 ato,

Red Bluff clay. Hiwannee (formerly Red Bluff), Wayne County, Mis-
sissipp1 (type locality) ; bank of Chickasawhay River; shell marl pockets in
plastic green clay near low water level. Collectors: D. L. Frizzell and A. R.
Troell, Jr., August 4, 1960. (Station F—60-11.)

Red Bluff equivalent. Quarry of Lone Star Cement Corporation, ca. 2
miles northeast of town of St. Stephens, Washineton County, Alabama;
ereenish to white glauconitie marl, forming lowest caleareous unit in quarry;
sample taken 6-10 inches below over-lying 4-5 foot very light gray indurated
limestone. Collector: D. L. Frizzell, September 3, 1957. (Station F—57-8.)

EOCENE, JACKSON GROUP.

Danville Landing shale. Duty, Catahoula Parish, Louisiana; bank of
Ouachita River at Duty ferry landing, east of town and off Louisiana High-
way 124; lower shell marl bed of formation exposed at low water level. Col-
lectors: D. L. Frizzell, C K. Lamber, and W. C. Horton, November 24, 1960.
(Stations F-60—-A2; CKL-60-2.)

Moody’s Branch marl. Riverside Park, Jackson, Mississippi (reassigned
type locality); basal greensand of formation, with abundant disseminated
fossil fragments, just above the blue clay of underlying Cockfield formation.
Collectors: E. Adams and D. l. Frizzell, November 24, 1959. (Stations F—
59-9, 10, 11.)

Montgomery Landing, Montgomery, Grant Parish, Louisiana; east bank
of Red River, 500 to 1,000 yards downstream from ferry landing (locally
known as Creole or “Creola”’ Bluff) ; light, extremely shelly marl. Collectors:
D. L. Frizzell, C K. Lamber, and W. C. Horton, November 23-24, 1960.
(Stations F-60—A1; CKL-60-1.)

Yazoo County, Mississippi; bank of Techeva Creek (also “Tesheva” on
U.S.G.S. maps) at bridge on Mississippi Highway 483, just north of town of
Midway, which is northward from Benton; blue to blue-gray marly sand-
stone containing shell debris. Collectors: D. L. Frizzell and A. R. Troell,
Jr., July 31, 1960. (Station F—60-—3. )

90 CALIFORNIA ACADEMY OF SCIENCES [Proc. 471 Ser.
EOCENE, WIxLcox GROUP.

Bashi marl. Meridian, Lauderdale County, Mississippi; bank of drainage
diteh on south side of ““Bypass 80”; lenticular bed of non-indurated shell
debris overlying a bed of conspicuous white sand. Collectors: D. L. Frizzell,
November 28, 1959; D. L. Frizzell and A. R. Troell, Jr., August 2, 1960.
Stations F—59-16; F—60-8.)

SYSTEMATIC DESCRIPTIONS
Order ANGUILLIFORMES
Family CONGRIDAE
(ienus Conger Schaeffer, sensu latissimo

ixcept for Conger? vetustus, that may be congenerie with Recent species
of the genus, the species here described are not believed to belong to the
genus Conger s.s. of the living fauna. “Conger” (in quotes) is a form-genus,
appled only provisionally, that will be revised when more becomes known
of the otoliths of Recent anguilliform fishes.

‘“‘Conger’’ brevior (IXoken).
(Figures 4 a—-b, 10 a-—d.)

Otolithus (Conger) brevior Koken, 1888, Deutsch. Geol. Ges., Zeitschr., Bd. 40, pp.
293-294, pl. 18, fig. 7.

O. (Conger) brevior (KOKEN). PostHumus, 1924, Foss. Cat., no. I, pars 24, p. 8.

DESCRIPTION. Sagitta medium size (maximum leneth observed, 7.1 mm.),
somewhat ovate with pronounced angularity, moderately high (height /length
ratios, 61 to 67 per cent), moderately inflated, greatest height slightly an-
terior to vertical midline. Dorsal margin risine from sharply rounded
anterior margin in low asymmetrical arch, with greatest curvature in pos-
terior portion; posterior marein sharply rounded to join broad asymmetri-
eallvy arched ventral margin; greatest curvature of ventral margin near
position of greatest height. Inner face moderately convex, smooth, with
prominent suleus but lacking area; suleus undivided, separated from anterior
by broad border, extending from about anterior fifth of sagitta to near
posterior third; suleus opens onto dorsal margin through ostial channel;
ostial region shehtly to moderately excavated, elongate, contiguous with
caudal region and sometimes set off from it by faint constriction of sides, its
anterior margin truneate; ostial channel considerably posterior to anterior
margin of ostial region, moderately long, with subparallel sides, slightly
flaring dorsally to open onto dorsal margin near position of greatest height;
caudal region deeply excavated, with subparallel sides, slightly longer than
ostial region on most specimens; erista superior faintly developed or absent;

VoL. XXXIT] FRIZZELL AND LAMBER: FISH OTOLITHS 91

Figure A. Structures of the congrid type sagitta. A-area; AB-anterior border;
CR-caudal region of sulcus; D-dorsal dome; OC—ostial channel; OR -ostial region.

erista inferior moderately marked, region above sulcus convex, lacking area.
Outer face somewhat convex, flattened in central region, sculptured with
weak undulations especially in marginal regions. Dimensions (in mim.) :

Length Height Thickness
2.0 1.6 0.6
3.3 Bal 0.8
4.4 Hee) 1.3
4.7 3.0 1.4
4.9 3.0 E3
3.0 3.6 1.6
5.4 3.0 aba
5.4 3.4 1.3
3.0 a. 1.4
6.5 4.2 Jel,

92 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4r1 SER.

COMPARISONS OF SAGITTAE. “Conger” brevior is very similar to “C.”
sanctus, new species, that occurs with it in the Oligocene but not in the upper
Kocene. “Conger” brevior, however, 1s less evenly ovate or elliptical and has
no area. Distinction of these species must be based on comparison of suites
of specimens, unless they are perfectly preserved.

The species differs from “C” dissimilis, new species, in the less nearly
ovate outline and the projection of the ostial region forward of the ostial
ehannel. The position of the ostial channel also distinguishes “C.” meridies,
new species.

A number of species related to “C” brevior have been described from
Tertiary deposits of other areas, under the names Conger, “Otolithus (Bro-
tulidarum),” “O. (Congeris),” “O. (Congridarum),” “O. (incertae sedis ),”
Heterenchelys, and Uroconger. The relationship of most of these to “C.”
brevior cannot be established without comparison of specimens with suleus
and ostial channel perfectly preserved. For example, Priem compared “O.
(Congeris)” papoimts (1906, pp. 275-276, text figs. 40-45) and later “O.
(Congeris)” duvergiert (1914, pp. 249-250, text fig. 9) with “C.” brevior.
Neither comparison is valid, as Priem’s illustrations are of eroded specimens
and Koken’s type figure is completely inadequate.

TyPE LocALITy. Hocene, Jackson group, Moody’s Branch marl; Jackson,
Mississipp1.

RANGE AND DISTRIBUTION. Oligocene, Vicksburg group: Byram marl, Old
Byram and Vicksburgh, Mississippi; Glendon limestone, Brandon, Missis-
sippl; Mint Spring marl, Vicksburg, Mississippi; Red Bluff clay, Hiwannee,
Mississippi, and equivalent strata at St. Stephens, Alabama. Eocene, Jack-
son group: Moody’s Branch marl, Montgomery, Louisiana, and near Midway,
Mississipp1.

Remarks. The type figure shows an immature specimen that could be-
long to this population or to that of “C.” sanctus, new species. It is specifi-
cally unidentifiable. We are applying the name “C.” brevior to this form, as

Figure 1. “Conger” sanctus Frizzell and Lamber, new species, holotype; Vicks-
burg group, Alabama; length, 9.0 mm. la. Inner face of right sagitta. 1b. Outer face.

Figure 2. “Conger” meridies Frizzell and Lamber, new species, holotype; Wilcox
group, Mississippi; length, 4.3 mm. 2a. Inner face of right sagitta. 2b. Outer face.

Figure 3. “Conger” fornicatus Frizzell and Lamber, new species, holotype; Jack-
son group, Louisiana; length, 3.6 mm. 3a. Inner face of right sagitta. 3b. Outer face.

Figure 4. “Conger” brevior (Koken); Vicksburg group, Mississippi; length, 5.5
mm. 4a. Inner face of left sagitta. 4b. Outer face.

Figure 5. “Conger” dissimilis Frizzell and Lamber, new species, holotype; Jack-
son group, Louisiana; length, 4.5 mm. 5a. Inner face of left sagitta. 5b. Outer face.

Figure 6. Conger? vetustus Frizzell and Lamber, new species, holotype; Jackson
group, Louisiana; length, 4.4 mm. 6a. Inner face of right sagitta. 6b. Outer face.

VoL. XXXII] FRIZZELL AND LAMBER: FISH OTOLITHS 93

“C.” sanctus has not been identified at the type level of the nominal species.
Details of the sulcus, as shown by Koken, are due to erosion of the anterior
boundary of the ostial channel. The horizontal position of the suleus, in his
figure, is believed to be an error of the artist.

94 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4711 SER.

‘‘Conger’’ dissimilis 'rizzell and Lamber, new species.
(FIGURES 5 a—b, 12 a-d.)

DESCRIPTION. Sagitta small to medium size (maximum leneth observed,
6.8 mm.), somewhat ovate in outline, high (height/length ratios, 71 to 80
per cent), moderately inflated, greatest height near anterior third of sagitta.
Dorsal margin rising from rounded anterior margin in undulating high arch
(or small dome) to meet sharply rounded posterior margin; ventral margin
broadly and asymmetrically arched with greatest curvature coincident with

, SS wi : 7a
~ ae ees PEE j SS SS
=e 7b 7c 7d
a
=) —_— ae J —————— Ss z (Ce ; EN eS
a a a
See ae ——_ SS
8a Sb 8c 8d

» SS iS me > ; i “s > Px wen iss
Cz ay) a= ) e Gy, ( (2 ¥
Sar Sk ee Sa es 4
10a 10b 10c 10d
pa sa a
Z > ~ 3 { 5 Page iS 7a ; (GnING
( Eee ®) ee | Ges ( a=
Sais é ; rar i hee ; <a
—< <5, llc
———— a> = \ = 2 = =m
ly a . } es t >
Yigal ee a ee a *~
f yy 7 —_ ) ae ) a )
me an ae |) ae

VoL. XXXII] FRIZZELL AND LAMBER: FISH OTOLITHS 95

position of greatest height. Inner face moderately convex, smooth, with
prominent suleus and sometimes slightly developed area; suleus undivided,
separated from anterior by narrow border, extending from near anterior
eighth of sagitta to near posterior third; suleus opens onto dorsal margin
through ostial channel; ostial region slightly excavated, contiguous with
caudal region and sometimes set off from it by slight constriction of sides,
its anterior margin rounded; ostial channel slightly posterior to anterior
margin of ostial region, short to medium lone, with nearly vertical sides,
moderately flaring dorsally to open onto dorsal margin anterior to position
of greatest height; caudal region moderately to deeply excavated with sub-
parallel sides, of about same length as ostial region; cristae inferior and
superior well marked; area slightly to very slightly impressed, somewhat
triangular, extending upward into dorsal arch or dome. Outer face mod-
erately convex, thickest at center, with poorly developed rugosities and
pustules toward margins. Dimensions of holotype and selected paratypes
(in mm.) :

Figures 7-12. Diagrammatic sketches of inner face.

Figure 7. ‘Conger’ sanctus Frizzell and Lamber, new species; holotype and
paratypes. la. Holotype, reversed; Vicksburg group, Alabama; length, 9.0 mm. 1b.
Reversed; Vicksburg group, Mississippi; length, 5.6 mm. le. Vicksburg group, Mis-
sissippi; length, 6.0 mm. 1d. Vicksburg group, Alabama; length, 4.2 mm.

Figure 8. Conger? vetustus Frizzell and Lamber, new species; holotype and
paratypes. Sa. Holotype, reversed; Jackson group, Mississippi; length, 4.4 mm. 8b.
Reversed; Jackson group, Louisiana; length, 4.2 mm. 8c. Reversed; Jackson group,
Louisiana; length, 3.6 mm. 8d. Reversed; Jackson group, Louisiana; length, 2.8 mm.

Figure 9. “Conger” fornicatus Frizzell and Lamber, new species; holotype and
paratypes; Jackson group, Louisiana. 9a. Holotype, reversed; length, 3.6 mm. 9b.
Reversed; length, 3.0 mm. 9c. Reversed; length, 2.8 mm. 9d. Reversed; length, 3.1
mm.

Figure. 10. “Conger” brevior (Koken); Vicksburg group, Mississippi. 10a. Same
as figure 4; length 5.5 mm. 10b. Length, 5.4 mm. 10c. Length, 5.8 mm. 10d. Re-
versed; length, 4.7 mm.

Figure 11. “Conger” meridies Frizzell and Lamber, new species; holotype and
paratypes. Wilcox group, Mississippi. 1la. Holotype, reversed; length, 4.8 mm. 11D.
Reversed; length, 3.6 mm. llc. Length, 2.7 mm. 11d. Length, 2.5 mm.

Figure 12. “Conger” dissimilis Frizzell and Lamber, new species; holotype and
paratypes; Jackson group, Louisiana. 12a. Holotype; length, 4.5 mm. 12b. Length,
5.38 mm. 12c. Reversed; length, 4.6 mm. 12d. Reversed; length, 4.3 mm.

96 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER.

Length Height Thickness Remarks
2.6 19 0.9
3.8 2.9 iG
3.8 3.0 0.9
4.3 3.4 1.3
4.4 3.0 1.3
4.5 3.4 1.3 Holotype
4.6 3.6 1:5
5.2 4.0 1.5
5.8 4.1 Ey
6.8 5.0 1.5

COMPARISONS OF SAGITTAE. “Conger” dissimilis differs from “C.” brevior
(Koken) and “C.” sanctus, new species, by its higher outline, greater con-
vexity, and in lacking a marked anterior projection of the ostial region
beyond the ostial channel. It is separated from “C.” meridies, new species,
by its greater relative height and angularity in outline. As with “C.” brevior
(see above), comparison with deseribed extra-American forms is unprofit-
able at this time.

Type LocaLity. Eocene, Jackson group, Moody’s Branch marl; Mont-
eomery ferry landing, Montgomery, Louisiana.

RANGE AND DISTRIBUTION. The species is known only from the type locality.

Remarks. The specific name is a Latin adjective indicating the lack of
similarity of this form to the sagittae of associated species.

‘‘Conger’’ fornicatus Frizzell and Lamber, new species.
(FIGuRES 3 a-b, 9 a-d.)

DESCRIPTION. Sagitta small (maximum length observed, 3.6 mm.), some-
what trianeular in outline, very high (height/length ratios, 77 to 88 per
cent), slightly inflated, greatest height at or forward of vertical midline.
Dorsal margin rising from acutely rounded anterior margin in slightly in-
curved arch to form high dorsal dome, then descending in slightly ineurved
arch to join sharply rounded posterior margin; dorsal dome highly developed,
somewhat quadrate, with bevelled top sloping toward anterior margin; ven-
tral margin broadly arched, with greatest curvature slightly forward of
position of greatest height. Inner face almost flat, smooth, with prominently
developed suleus and area; suleus undivided, separated from anterior by
moderately broad border, extending from about anterior sixth to posterior
third of sagitta; suleus opens onto dorsal margin through ostial channel;
ostial region slightly excavated, short, contiguous with caudal region and
usually set off by slight constriction (on well preserved specimens), its an-
terior margin truncate; ostial channel slightly posterior to anterior margin
of ostial region, moderately long; anterior boundary of ostial channel nearly

VOL. XXXII] FRIZZELL AND LAMBER: FISH OTOLITHS 97

straight, bending forward at termination, posterior boundary bending back-
ward and upward; caudal region deeply excavated, with subparallel sides,
longer than ostial region; cristae superior and inferior well developed, with
erista superior following configuration of posterior boundary of ostial chan-
nel; area very prominent, deeply impressed, variable in outline (usually
somewhat triangular), extending high into dorsal dome. Outer face smooth,
slightly to moderately convex, thickest at center and in region of dorsal
dome. Dimensions of holotype and selected paratypes (in mm.) :

Length Height Thickness Remarks
19 1.6 0.5
2.0 1.6 0.5
PT rap 0.6
2.7 2.4 0.7
2.8 2.0 0.7
3.0 2.1 0.8
3.0 2.3 0.8
ot 2.4 0.8
3.6 2.0 1.0
3.6 2.8 0.9 Holotype

COMPARISONS OF SAGITTAE. “Conger” fornicatus is distinguished by its
characteristic outline from all other forms known to us, ineluding those illus-
trated in available literature.

Tyrer LocaLity. Eocene, Jackson group, Danville Landing shale (lower
shell marl bed), Duty ferry landing, Duty, Louisiana.

RANGE AND DISTRIBUTION. The species is known only from the type locality.

ReMaRKsS. The specific name fornicatus is a Latin adjective indicating the
arched appearance of the sagitta produced by the development of the dorsal
dome.

‘‘Conger’’ meridies Frizzell and Lamber, new species.
(Figures 2 a-b, 11 a-d.)

DESCRIPTION. Sagitta medium small to medium size (maximum leneth
observed, 6.7 mm.), somewhat ovate with pronounced angularity, moderately
high (height/length ratios, 59 to 73 per cent), slightly to moderately inflated,
ereatest height at or anterior to vertical midline. Dorsal margin rising from
sharply rounded anterior margin in moderately asymmetrical arch with
ereatest curvature at position of greatest height; posterior margin rounded
with development of rugosities, joining broadly and asymmetrically arched
ventral margin with greatest curvature at position of maximum height. Inner
face smooth, slightly convex, with prominent sulcus but lacking area; sulcus

98 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 47TH SER.

undivided, separated from anterior by very narrow border, extending from
about anterior tenth to near posterior third of sagitta; suleus opens onto
dorsal margin through ostial channel; ostial region moderately excavated,
short, contiguous with caudal region and sometimes set off from it by slight
constriction of sides, its anterior margin rounded; ostial channel slightly
posterior to anterior margin of ostial region, moderately long, directed up-
ward and slightly backward, expanding in central region before constricting
somewhat and opening onto dorsal margin at position of greatest height;
caudal region longer than ostial, deeply excavated, with subparallel sides,
expanding and rounded at posterior end; crista superior well marked, inten-
sifying outline of sulcus; crista inferior shghtly less developed; area absent.
Outer face moderately convex on most specimens, thickest at center, with
undulating surface along margins. Dimensions of holotype and selected
paratypes (in mm.) :

Length Height Thickness Remarks
2.0 oer jE
2.1 1.6 1.0
3.3 2.0 0.9
3.3 2.1 0.9
3.6 2.4 0.9
3.6 2.6 feat
3.8 2.8 all
4.3 2.8 i Holotype
4.5 2.9 0.9
4.8 3.2 1.0

COMPARISON OF SAGITTAE. “Conger” meridies is lower, less convex, and
less angular in outline than “C.” dissimilis, new species. It somewhat re-
sembles “C.” brevior (Koken) and ‘C.” sanctus, new species. Those, how-
ever, are distinguished by an anterior prolongation of the ostial region of
the suleus, forward of the ostial channel, that is much less developed in “C.”
meridies. As with “C.” brevior (see above), similarities to extra-American
forms cannot be determined at this time.

TypPE LocALITY. Hocene, Wilcox group, Bashi marl; Meridian, Mississippi.

RANGE AND DISTRIBUTION. “Conger” meridics is known only from un-
common otoliths at its type locality.

Remarks. The specific name, the Latin word for a geographie meridian,
is in recognition of the type locality of the species: Meridian, Mississippi. It
iS a noun in apposition.

‘““Conger’’ sanctus Frizzell and Lamber, new species.
(FriGuREs 1 a—b, 7 a-d.)

VoL. XXXIT] FRIZZELL AND LAMBER: FISH OTOLITHS 99

DESCRIPTION. Sagitta moderately large (maximum length observed, 9.0
mm.), somewhat ovate in outline, moderately high (height/length ratios, 62
to 72 per cent), moderately inflated, greatest height variable in relation to
position of vertical midline. Dorsal margin rising from sharply rounded
anterior margin in low nearly symmetrical arch; posterior margin mod-
erately rounded to join arched ventral margin. Inner face moderately con-
vex, smooth, with prominent suleus and shallow area; suleus undivided,
separated from anterior by narrow border, extending from about anterior
eighth of sagitta to near posterior third; suleus opens onto dorsal margin
through ostial channel; ostial region slightly excavated, elongate, contiguous
with caudal region and sometimes set off by slight constriction of its sides,
its anterior margin truncate; ostial channel very much posterior to anterior
margin of ostial region, moderately long to long, with subparallel sides,
slightly flaring dorsally to open onto dorsal margin slightly anterior to
position of greatest height; caudal region deeply excavated, as long as ostial
region, with nearly parallel sides; crista superior marked above caudal
region, degenerate along ostial region; erista inferior well marked; area
variable in outline and size (predominantly elliptical), slightly to moderately
impressed. Outer face slightly to moderately convex, flattened in central
region; sculpture lacking or consisting of irregular undulations or bosses
adjacent to margins. Dimensions of holotype and selected paratypes (in
mm.) :

Length Height Thickness Remarks
2.0 1.8 0.8
3.3 2.1 0.8
4.2 Det 0.9
4.8 3.1 Aad
5.6 3.8 12
D.t 3.6 1.3
6.0 4.1 1.4
6.1 4.0 1.5
9.0 5.6 2.3 Holotype

COMPARISONS OF SAGITTAE. “Conger” sanctus is extremely like “C.” brevior
(Koken), but is distinguished by the presence of an area and by the slight
difference in outline. “Conger” meridies, new species, and “C.” dissimilis,
new species, also lack the area and differ in outline, as well as having less
well developed projection of the ostial region anterior to the ostial channel.

As with “C.” brevior (Koken) (see above), similarities to extra-American
forms cannot adequately be determined from the literature. One species
from the European upper Oligocene (“C.” fallax (Koken), 1891, p. 139, pl.
10, fig. 3: as Otolithus (incertae sedis)), however, appears to be extremely
similar to “C.” sanctus, and the two may prove to be identical.

100 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 47TH SER.

Type LocaLity. Oligocene, Vicksburg group, Red Bluff clay equivalent;
St. Stephens Quarry, St. Stephens, Alabama.

RANGE AND DISTRIBUTION. Oligocene, Vicksburg group: Glendon lime-
stone, Brandon, Mississippi; Red Bluff clay, Hiwannee, Mississippi; Red
Bluff clay equivalent, St. Stephens, Alabama.

Remarks. The specific name sanctus (Latin for “sacred”’) refers to the
type locality at St. Stephens Quarry. It is an adjective.

Conger? vetustus I*rizzell and Lamber, new species.
(FIGURES 6 a—b, 8 a-d.)

DescripTION. Sagitta small to medium large (maximum length observed,
4.8 mm.; one broken specimen has an estimated length of 7.3 mm.), sub-
lanceolate, low (height /length ratios, 40 to 50 per cent), moderately inflated,
ereatest height at or slightly anterior to vertical midline. Dorsal margin
rising from acute anterior margin in low arch; posterodorsal slope with one
or more pronounced coarse rugosities (in well preserved specimens) before
meeting acute posterior margin; ventral margin broadly arched. Inner face
moderately convex, smooth, with prominent suleus and area; sulcus un-
divided, separated from anterior by moderately broad border, extending
from about anterior fifth to posterior third; suleus opens onto dorsal margin
through ostial channel; ostial region prominently excavated, short, contiguous
with caudal region and sometimes set off from it by constriction of sides, its
anterior margin rounded; ostial channel slightly posterior to anterior margin
of ostial region, short and very wide; anterior boundary of ostial channel
nearly straight, bent forward at termination, posterior boundary extending
far backward near dorsal margin; caudal region deeply excavated, with sub-
parallel sides, more than twice length of ostial region; crista superior well
marked, bending to follow configuration of posterior boundary of ostial
channel; crista inferior marked, especially below caudal region; area deeply
impressed, predominantly elongate-elliptical. Outer face smooth, moder-
ately convex, thickest along horizontal midline. Dimensions of holotype and
selected paratypes (in mm.) :

Length Height Thickness Remarks
2.0 Jigll 0.6
2.6 £2 0.6
2.8 1.4 0.7
3.0 12 0.6
3.6 135 0.6
4.2 1.8 0.7
4.4 1.8 0.7 ILolotype
4.7 1.9 OF

VoL. XXXIT] FRIZZELL AND LAMBER: FISH OTOLITHS 101

COMPARISONS OF SAGITTAE. Conger? vetustus is unlike other forms yet
encountered in the American lower Tertiary. The sagitta resembles that of
Conger conger (Linnaeus) of the Recent (Chaine, 1938, pp. 234-241, pl. 17).
It differs from the adult sagitta of C. conger in having a shorter, narrower,
better defined, and more sloping suleus, and the anterior end is more sharply
rounded. The sagitta of C.? vetustus, however, is extremely similar to that
of juvenile C. conger as figured by Chaine.

TYPE LOCALITY. Eocene, Jackson group, Moody’s Branch marl; Riverside
Park, Jackson, Mississippi.

RANGE AND DISTRIBUTION. The species is known from the Moody’s Branch
marl of Jackson, Mississippi, and Montgomery, Louisiana. It is rare at both
localities.

Remarks. The similarity of C? vetustus to otoliths of young Conger
conger (see above) may have some phylogenetic significance. It suggests
that C.? vetustus belongs to a lineage that is ancestral to living species of
Conger S.s.

The specific name is a Latin adjective meaning ancient.

LITERATURE CITED

CHAINE, J.
1938. Recherches sur les Otolithes des Poissons. Etude Descriptive et Compara-
tive de la Sagitta des Téléostéens (Suite). Société Linnéenne de Bor-
deaux, Actes, tome 90, pp. 1-258, pls. 1-18.

FRIZZELL, D. L., and C K. LAMBER
1961. New Genera and Species of Myripristid Fishes, in the Gulf Coast Cenozoic,
known from otoliths (Pisces, Beryciformes). University of Missouri,
School of Mines and Metallurgy, Bulletin, Technical Series, no. 100, pp.
1-25, 5 pls., 2 text figs.

Frost, G. A.
1926. A comparative study of the Otoliths of the Neopterygian Fishes (con-
tinued). III. Order Apodes. Annals and Magazine of Natural History,
ser. 9, vol. 17, pp. 99-104, pl. 4.

KOKEN, E.

1888. Neue Untersuchungen an tertidren Fisch-Otolithen. Deutschen geolo-
gischen Gesellschaft, Zeitschrift, Band 40, pp. 274-305, pls. 17-19.

1891. Neue Untersuchungen an tertiiren Fisch-Ctolithen. II. Deutschen geolo-
gischen Gesellschaft, Zeitschrift, Band 43. pp. 77-170, pls. 1-10, text figs.
1-27, 1 range chart.

PRIEM, F.
1906. Sur les Otolithes des Poissons Kocénes du Bassin Parisien. Société Géolo-
gique de France, Bulletin, 4° série, tome 6, pp. 265-280, text figs. 1-51.
1914. Sur des Otolithes de Poissons Fossiles des Terrains Tertiaires Supérieurs
du Sud-Oest de la France. Société Géologique de France, Bulletin, 4°
série, tome 14, pp. 244-278, 74 text figs.

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PROCEEDINGS
OF THE
CALIFORNIA ACADEMY OF SCIENCES

FOURTH SERIES

G Dallas Hanna Anniversary Volume

Vol. XXXII, No. 6, pp. 103-147, 4 figs. June 29, 1962

THE HERPETOLOGY OF NEPAL:
A HISTORY, CHECK LIST,
AND ZOOGEOGRAPHICAL ANALYSIS
OF THE HERPETOFAUNA
by
Lawrence W. Swan'
San Francisco State College
San Francisco, California

and

Alan E. Leviton

California Academy of Sciences

San Francisco, California

INTRODUCTION

Herpetological collecting in Nepal has been sporadic and the literature
describing the Nepalese herpetofauna has appeared irregularly and infre-
quently for more than a century. Prior to 1948 little was known of the fauna
of the country beyond the centrally located capital of Katmandu, but during
the last few years the remote regions have become more readily accessible
and several important collections of animals and plants have been made.

1. Research Associate, Department of Herpetology, California Academy of Sciences.

[ 103 |

Marine Biological Laboratory
LIBRARY

APR 1 8 1968
WOODS HOLE, MASS.

104 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4711 SER.

Motivation for the present paper, in which we attempt to collate all
hitherto published data dealing with Nepalese amphibians and reptiles, was
derived from a joint interest in Himalayan zoogeography on the part of the
authors. Nepal lies at the crossroads of four major faunal units. The
Mediterranean and West Chinese divisions of the Palearctic Region meet
the Indian and Indochinese subregions of the Oriental Region, and an
understanding of the Nepalese fauna is basic to a study of the evolution and
distribution of the high altitude fauna of southeast Asia.

The present report is divided into three parts. The first, a history of
herpetologiecal work in Nepal, is based on a collation of published works
reporting on collections of amphibians and reptiles obtained in that country.
A eheek list of amphibians and reptiles known to occur in Nepal follows;
in this list all literature containing Nepal records are noted in the respective
species synonymies and are the only citations so listed. The third section of
this report represents a zoogeographical analysis of the herpetofauna of
Nepal. In view of the facet that this analysis is based upon an incomplete
knowledge of the country’s fauna, we have prepared a detailed table (table
I), in which all species known to occur in Nepal and adjacent areas, or in
adjacent areas but which have not yet been recorded from Nepal, have been
listed. The table indicates the geographical relationships of the known
fauna, and suggests species which are still likely to be found in the country.

The writers are grateful to Drs. Edward L. Kessel and Leo G. Hertlein,
editors of the Aecademy’s Proceedings, for their patience and their careful
editing of the paper. The authors are solely responsible for the accuracy
of the data presented and for the zoogeographie interpretation.

HISTORY

The earliest collector of Nepalese amphibians and reptiles, to the best
of our knowledge, was Brian Hodgson who occupied the British Residence
in Katmandu during the years 1820 to 1822, and 1824 to 1843. Although
Ilodgson was preceded in Nepal by at least two earlier naturalists, Nathaniel
Wallich, in 1817, and Francis Buchanan-Hamilton, in 1802-1803, both of
whom resided in Katmandu for short periods, there is no evidence that
either obtained any amphibians or reptiles for museum specimens.

During his residence in Nepal, Hodgson distinguished himself as an
authority in many disciplines, among which his researches in ethnology,
Buddhism, ornithology, and mammalogy are preeminent. In addition to his
normal diplomatic duties he found time to publish 127 papers on zoological
subjects, although none dealt with amphibians or reptiles. His prime econtri-
bution to the field of herpetology was through his collection of Nepalese
fauna, the specimens of which, presented to the British Museum in 1858,
included 9,512 specimens of birds, 903 mammals, and 84 reptiles, and a

VoL. XXXII] SWAN AND LEVITON: HERPETOLOGY OF NEPAL 105

number of drawings of which there were 1241 sheets of birds, 557 sheets of
mammals, and 55 sheets of reptiles (Hunter, 1896).

Several years before his final return to England, delayed until 1858,
a few of Hodgson’s specimens found their way to London and were described
by Cantor (1839); they were also included in Gray’s Catalogues of the
British Museum herpetological collection (1844-1850). On his arrival in
England, Hodgson turned over the bulk of his collections of amphibians
and reptiles to Albert Giinther who, in a series of three papers (1858, 1860.
1861), described their contents. In 1864, Giinther monographed the rep-
tilian fauna of “British India,” incorporating the results of his earlier
studies of the Hodgson material in the volume. In addition, he listed a
single specimen of the agamid lizard Calotes versicolor, said to have been
collected in Nepal by Thomas Hardwicke, and a turtle, Chitra indica, sup-
posedly obtained by the botanist Hugh Falconer who, with Hardwicke, was
one of several naturalists to visit Nepal in the early years of British contact
with that country. All other Nepalese records given by Giinther are at-
tributed to Brian Hodgson.

Hodgson’s influence at the Nepalese court enabled him to obtain a
unique concession from the Prime Minister to permit Joseph Hooker ac-
cess to eastern Nepal during the course of the latter’s botanical travels in
Sikkim. Hooker’s collection of amphibians and reptiles are all reported as
being obtained in Sikkim; there is no precise indication in known literature
sourees that he obtained any amphibians or reptiles while he was traveling
in Nepal between November 5, 1848 and December 15, 1848 (Tooker, 1854,
vol. 1, pp. 186-280).

Although Katmandu was visited periodically by European naturalists
(ineluding Hermann Schlagintweit, in 1856) and by Nepalese and Indian
collectors during a sixty year period (1858-1906) following Hodgson’s
work there, there is scarcely any published evidence to indicate that am-
phibians or reptiles were collected. There is, however, one notable exception.
In his lists of amphibians and reptiles in the Indian Museum (1891 and
1892), William Sclater gives Katmandu as the locality for specimens of sev-
eral species stored in that museum’s collections. Unfortunately, there is no
indication who collected the material.

In 1907, Nelson Annandale, George Boulenger, and Frank Wall de-
scribed a substantial collection of amphibians and reptiles obtained in the
vicinity of Katmandu by R. Hodgart. Several new records, including two
species of frogs, five lizards, and five snakes, are given for Nepal. Subse-
quently, in a series of papers published by Wall (1907-1924), a number of
snakes were recorded for the first time from the country. Included among
these is Psammodynastes pulverulentus, the only indication for its oceur-
rence in Nepal being a distribution map published by Wall, in 1910, in
which the locality Butal [= ? Butwal], Nepal, is given as a collecting site.

106 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

In 1913, Boulenger described a small collection of amphibians and
reptiles taken at the extreme eastern frontier of Nepal, immediately adja-
cent to the Darjeeling District of Bengal, collected by N. H. Stevens. Two
new snake records were included.

With publication of the reptile volumes in the Fauna of British India
series (1931, 1935, and 1943), the widely scattered literature on reptiles
of southern Asia was brought together for the first time. The three volumes
on reptiles, written by Maleolm Smith, stand as the single most important
contribution to Asian herpetology published to date. In preparing these
works it was obviously impractical for Smith to give detailed locality,
collector, date, and available ecological information for all the material he
examined. Consequently, specific consideration of the reptiles of any one
region, tor example Nepal, is not possible in these volumes, and resort must
be made to the original literature. Regretfully, a comparable volume dealing
with the amphibians of southern Asia has not been published.

Since 1948, Nepal has been visited by numerous expeditions, and there
are few portions of the country which have not been investigated by persons
interested in the fauna and flora. Nevertheless, the published literature
suggests that only a few expeditions have obtained amphibians and reptiles
in the course of their activities.

In 1949, an expedition under the leadership of H. W. Tilman explored
the Langtang Himal, north of Katmandu. Herpetological collections were
made by Oleg Polunin and were deposited in the British Museum where
Maleolm Smith examined them and prepared a report published in 1951.
An interesting ecological analysis of this Himalayan area was prepared by
Polunin and published as an appendix to Tilman’s “Nepal Himalaya” (1951).

Western Nepal was visited, in 1952, by an expedition sponsored by the
British Museum and the Royal Horticultural Society. Oleg Polunin accom-
panied this group and again secured a number of amphibians and reptiles
which are of special interest because of the altitude records. Malcolm Smith
and James C. Battersby of the British Museum deseribed this collection in
1953. Six new records of amphibians and reptiles in Nepal were reported on
in this paper.

In 1954, the California-Himalaya Expedition to Makalu traversed east-
ern Nepal in the vicinity of the Arun River (a descriptive summary of this
expedition was prepared by Houston and Long [1955] and a discussion of
high-altitude ecology by Swan [1961]). The collections of amphibians and
reptiles, made by Lawrence Swan, and including eight new records and a
new species of frog from Nepal, were deposited in the Natural History
Museum of Stanford University. Alan Leviton, George Myers, and
Lawrence Swan reported on this collection in 1956 and, as an addendum to
the paper, the authors also described a small collection of amphibians from
Katmandu and Pokhara obtained by Alan Taft earler that vear.

VoL, XXXIT] SWAN & LEVITON: HERPETOLOGY OF NEPAL 107

Recently, a few specimens of Natrix piscator were collected by the joint
Harvard-Yale universities expedition to central Nepal. In addition, several
snakes and frogs from Nepal were obtained for the Chicago Natural History
Museum by Dr. Robert L. Fleming in the vicinity of Katmandu. These
collections are reported in this paper for the first time. Collections of am-
phibians and reptiles made by L. W. Swan during 1960 in eastern Nepal
between Katmandu and Darjeeling are not reported in this paper.

CHECKLIST OF NEPALESE AMPHIBIANS AND REPTILES?

Class AMPHIBIA
Order SALIENTIA

Family Bufonidae

Bufo andersoni Boulenger.

Bufo andersoni, LEVITON, MYERS, and Swan, 1956, p. 4 (above Tamur River [1000
Peale

RANGE IN NEPAL. Eastern? (Tamur Valley).

Bufo himalayanus Ciinther.

Bufo himalayanus GUNTHER, 1864, p. 422 (type loc.: Nepal and Sikkim). BOULENGER,
1882, p. 305 (Nepal). Smirn, 1951, p. 727 (Thangjet [5000 ft.]; Syarpagaon
[Langtang Khola, 8000—9000 ft.]; Rasua Garhi District). SmMitH and Bar-
TERSBY, 1953, p. 703 (Jumla [7600 ft.]).

Bufo melanostictus (nec Schneider) GUNTHER, 1860, p. 165 (Nepal); 1851, p. 220
(Nepal).

RANGE IN Nepan. Central (Thangjet*, Syarpagaon, Rasua Garhi); West-
ern (Jumla).

Bufo melanostictus Schneider.

Bufo melanostictus, SCLATER, 1892, p. 27 (Katmandu). BouLeNnaer, 1907, p. 149 (Chit-
long; Soondrijal). Lrviron, Myers, and Swan, 1956, pp. 4 and 14 (near Khand-
bari [7000 ft.]; near Dhankuta [6000 ft.]; near Num [5000 ft.]; near Yetung
[8500 ft.]; Kalimati).

Chicago Natural History Museum 83096 (Patan).
RANGE IN Nepau. Eastern (Dhankuta, Khandbari, Num, Yetung) ; Cen-
tral (Chitlang, Kalimati, Katmandu, Sundarijal, Patan).

2. Many of Giinther’s records (between 1858 and 1864) noted in the following checklist were based
on collections made by Brian Hodgson. Most of the specimens probably came from the vicinity of Katmandu,
but it is known that Hodgson employed collectors to obtain specimens from elsewhere in the country. Unless
otherwise noted, all of Giinther’s records are presumed to have been based on collections from central Nepal,
from in and about the capital city of Katmandu.

3. Nepal has been divided into Western, Central, and Eastern portions based on the major river systems
of the country. See map, pages 120-121, fig. 1.

4. The spelling of locality names has been corrected to follow the Survey of India Eight Mile Map of
Nepal (1934) and the 1 inch to 4 mile map sheets of Nepal (1955).

108 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47H Serr.

Family Microhylidae

Microhyla ornata (Duméril and Bibron).
Microhyla ornata, Lrviron, Myrrs, and Swan, 1956, p. 5 (Khandbari [4000 ft.]).
RANGE IN NEPAL. Hastern (Khandbari).

Family Pelobatidae

Scutiger sikkimmensis (Blyth).

Scutiger sikkimensis, SmMirn, 1951, p. 727 (Langtang Village [11,000 ft.]). Smirn
and Barrerssy, 1953, p. 703 (Khola north of Maharigaon [13,500 ft.]).
RANGE IN NEPAL. Central (Langtang Village); Western (Maharigaon).

Family Ranidae

Rana breviceps Schneider.
Rana breviceps, LEVITON, Myers, and Swan, 1956, p. 5 (Dharan [1000 ft.]).
RANGE IN NEPAL. Eastern (Dharan).

Rana cyanophlyctis Schneider.

Rana cyanophlyctis, ScLaTrr, 1892, p. 2 (Katmandu). BouLencrer, 1907, p. 150
(Soondrijal; Pharping); 1920, p. 12 (Nepal). Lreviron, Myers, and Swan, 1956,
pp. 6, 14 (Tamur River [500 ft.]; near Dhankuta [4500 ft.]; near Num [6000
ft.]; near Khandbari [4500 ft.]; above Arun River [3500 ft.]; Sandarijal; near
Pokhara [Mardi Khola] ).

RANGE IN NepAL. Eastern (Arun River, Dhankuta, Khandbari, Num,
Tamur River); Central (Katmandu, Pharping, Pokhara, Sundarijal).

Rana formosa ((iinther).

Rana formosa, BOULENGER, 1907, p. 151 (Soondrijal). Smiru, 1951, p. 727 (Rasua
Garhi [6000 ft.]).
RANGE IN NEPAL. Central (Rasua Garhi, Sundarijal).

Rana liebigii Giinther.

Rana liebigii GUNTHER, 1860, p. 157, pl. 28, fig. A (Nepal and Sikkim); 1861, p. 220
(Nepal); 1864, p. 407 (Nepal). BouLrencer, 1913, p. 337 (Sandakpho [11,500
ft.]); 1920, p. 80 (Nepal [types]). Smiru, 1951, p. 727 (Rasua Garhi [6000 ft.]).
LEVITON, MYERS, and SwAN, 1956, p. 6 (near Num [5000 ft.]).

RANGE IN NEPAL. Eastern (Num, Sundakphu); Central (Rasua Garhi).

Rana limnocharis Wiegmann.

Rana limnocharis, SCLATER, 1892, p. 6 (Katmandu). BOULENGER, 1907, p. 151 (Soon-
drijal). Lrviron, Myers and Swan, 1956, p. 6 (above Yetung [7000 ft.]; near
Dhankuta [4500 ft.]; Arun River [1000 ft.]; below Yetung [4000 ft.]; near
Num [3500 ft.]; above Dhankuta [6000 ft.]).

RANGE IN NEPAL. Eastern (Arun River, Dhankuta, Num, Yetung);
Central (Katmandu, Sundarijal).

Rana monticola (Anderson).
Leptobatrachium monticola, BOULENGER, 1907, p. 149 (Soondrijal).
RANGE IN NEPAL. Central (Sundarijal).

Vou. XXXII] SWAN AND LEVITON: HERPETOLOGY OF NEPAL 109

Rana polunini Smith.
Rana polunini SmitTu, 1951, p. 727 (Langtang Village [11,000 ft.], Nepal).
RANGE IN NEPAL. Central (Langtang Village).

Rana swani Myers and Leviton.
Rana swani MYeRs and Leviton, 1956, p. 4 (Dharan [1000 ft.], Nepal).

RANGE IN NEPAL. Hastern (Dharan; may also be represented in Arun
Valley [see Leviton, Myers, and Swan, 1956, p. 6]).

Rana tigrina Daudin.

Rana tigrind, GUNTHER, 1860, p. 164 (Nepal); 1861, p. 220 (Nepal). BOoUuLENGER,
1907, p. 151 (Katmandu [4000-5000 ft.]; Soondrijal); 1920, p. 19 (Nepal).
LEVITON, Myers, and Swan, 1956, p. 14 (Kalimati).

Chicago Natural History Museum 83091-83094 (Katmandu).
RANGE IN NEPAL. Central (Kalimati, Katmandu, Sundarijal).

Family Rhacophoridae

Rhacophorus maculatus (Gray).

Polypedates maculatus, GUNTHER, 1861, p. 220 (Nepal).

Rhacophorus maculatus, Leviron, Myrrs, and Swan, 1956, p, 9 (above Num
[5000 ft.]).
RANGE IN NEPAL. Eastern (above Num); Central (without exaet lo-

eality data’).

Rhacophorus maximus Giinther.

Rhacophorus maximus GUNTHER, 1858, p. 83 (Nepal); 1860, p. 165 (Nepal); 1861,
p. 220 (Nepal); 1864, p. 435 (Nepal [5200 ft.]).
RANGE IN NEPAL. Central (without exact locality data’).

Class REPTILIA
Order CROCcODILIA
Family Crocodilidae

Gavialis gangeticus (Gmelin).
Gavialis gangeticus, GUNTHER, 1861, p. 215 (Nepal).
RANGE IN NEPAL. Central (without exact locality data’).

Order TESTUDINATA
Family Emydidae

Kachuga dhongoka (Gray).

Batagur dhongoka, GUNTHER, 1861, p. 214 (Nepal); 1864, p. 42 (Nepal). Titroparp,
1876, p. 22 (Nepal).
RANGE In NEPAL. Central (without exact locality data’).

Kachuga kachuga (Gray).
Batagur lineata (Gray), GinruHer, 1861, p. 214 (Nepal).
Batagur kachuga, THronaLp, 1876, p. 19 (Nepal).

110 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER.

Kachuga kachuga, Smirn, 1931, p. 131 (Nepal).
RANGE IN NEPAL. Central (without exact locality data’).

Family Testudinidae

Testudo elongata Blyth.
Testudo horsfieldi, GUNTHER, 1861, p. 214 (Nepal); 1864, p. 7 (Nepal).

Smith (1931, p. 148) states, “Hodgson obtained a specimen in the Saul
forests of Nepal and has left a fine coloured sketch of it in his collection of
drawings.” It is uncertain whether Smith considered Hodgson’s drawing
(which Giinther described as 7. horsfieldi, supra cit.) to be T. elongata or
whether Smith’s reference is in error.

RANGE IN NEPAL. Central (without exact locality data’).

Family Trionychidae

Chitra indica (Gray).

Chitra indica, GUNTHER, 1861, p. 214 (Nepal); 1864, p. 50 (Nepal). Smirn, 1931,
p. 162 (Nepal).
RANGE IN NEPAL. Central (without exact locality data”).

Trionyx gangeticus Cuvier.

Trionyx gangeticus, GUNTHER, 1861, p. 214 (Nepal); 1864, p. 47 (Nepal). SMITH,
1931, p. 167 (base of Nepal foothills).

Trionyx javanicus GRAY, GUNTHER, 1861, p. 214 (Nepal).

RANGE IN NEPAL. Central (without exact locality data’).

Order SQUAMATA
Suborder Sauria
‘amily Agamidae

Agama tuberculata Gray.

Agama tuberculata, ANNANDALE, 1907, p. 154 (Chitlong). Smiru, 1935, p. 214
(Kashmir to Katmandu); 1951, p. 728 (Chattekhola [Rasua Garhi District],
Rasua Garhi [7000 ft.]). SmirnH and Barrerssy, 1953, p. 703 (Jumla [7600 ft.]).

RANGE IN NEPAL. Central (Chattekhola, Rasua Garhi, Chitlang) ; West-
ern (Jumla).

Calotes versicolor (Daudin).

Calotes versicolor, Gray, 1845, p. 243 (Nepal). GiUnruer, 1860, p. 140; 1864, p. 215
(Nepal). ANNANDALE, 1907, p. 153 (Katmandu). SmiryH, 1935, p. 189 (Nepal) ;
1951, p. 728 (Rasua Garhi District). Smiru and BarTerssy, 1953, p. 703 (Lapha
[Karnali Valley, 4000 ft.]). Lrvrron, Myers, and Swan, 1956, p. 10 (above
Dharan [1500 ft.], Arun Valley [1000 and 2000 ft.], near Khandbari [3000, 4500
and 7000 ft.], below Yetung [4000 ft.]).

RaNcE in Nepau. Eastern (Arun River Valley, Dharan, Khandbari,

Yetung); Central (Katmandu, Rasua Garhi).

VoL. XXXII] SWAN AND LEVITON: HERPETOLOGY OF NEPAL Latet

Japalura major (Jerdon).

Japalura major, SMITH and BATTERSBY, 1953, p. 703 (Barbung Khola, Kakkatgaon
[1200 ft. sic], above Rimi [10,500 ft.], Samala [7500 ft.], Chaudhabise Khola
[10,000 ft.]).

RANGE IN NEPAL. Western (Barbung Khola, Kakkatgaon, Rimi, Samala,
Chaudhabise Khola).

Japalura tricarinata (Blyth).

Acanthosaura tricarinata, ANNANDALE, 1907, p. 153 (Chandragiri [8000 ft.]).
Japalura tricarinata, SmMiTH, 1935, p. 169 (eastern Nepal); 1951, p. 728 (Langtang
Khola [9500 ft.]).

RANGE IN NEPAL. Central (Chandragiri, Langtang Khola).

Family Gekkonidae

Cosymbotus platyurus (Schneider).

Hemidactylus nepalensis ANNANDALE, 1907, p. 151 (Katmandu).
Platyurus platyurus, Smiru, 1935, p. 102 (Nepal).
Cosymbotus platyurus, Leviton, MYERS, and Swan, 1956, p. 9 (Khandbari [4000 ft.]).

RANGE IN NEPAL. Eastern (Khandbari); Central (Katmandu).

Hemidactylus frenatus Schlegel.
Hemidactylus frenatus, Leviron, Myers and Swan, 1956, p. 10 (Dharan [1000 ft.]).
RANGE IN NEPAL. Eastern (Dharan).

Family Scincidae

Leiolopisma himalayanum (Giinther).

Lygosoma himalayanum, ANNANDALE, 1907, p. 154 (Chitlong).

Leiolopisma himalayanum, Smiru, 1935, p. 299 (Nepal). SmirH and BATTERSBY,
1953, p. 703 (Jumla [7600 ft.]).
RANGE IN Nepau. Central (Chitlang); Western (Jumla).

Leiolopisma ladacensis (Giinther).

Leiolopisma ladacense, SMIrH and BaAtTrerssBy, 1953, p. 703 (Balangra Pass, Tibrikot
[11,500 to 12,000 ft.], Phoksumdo Tal [14,000 ft.], Pemringgaon [16,000 ft.],
Kahajeng Khola [18,000 ft.]).

RANGE IN NEPAL. Western (Balangra Pass, Tibrikot, Phoksumdo Tal,

Pemringgaon, Kahajeng Khola).

Leiolopisma sikkimense (Blyth).
Lygosoma sikkimense, ANNANDALE, 1907, p. 154 (Chitlong, Katmandu).
Leiolopisma sikkimense, Smiru, 1935, p. 301 (Chitlong, Katmandu); 1951, p. 728
(Thangjet [5000 ft.]).
RANGE IN NepAu. Central (Chitlang, Katmandu, Tangjet).

Mabuya carinata (Schneider).
Mabuya carinata, Leviron, Myers, and Swan, 1956, p. 11 (Yetung [4500 ft.]).
RANGE IN NEPAL. Eastern (Yetung).

112 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4711 SER.

Mabuya macularia (Blyth).

Tiliqua rufescens (nec Shaw) GRay, 1853, p. 388 (Nepal). GUNTHER, 1860, p. 160
(Nepal); 1861, p. 215 (Nepal).

Euprepes rufescens, GUNTHER, 1863, p. 79 (Nepal).

Mabuia macularia, ANNANDALE, 1907, p. 154 (Terai near Raxaul [Nepal frontier]).
RANGE IN Nepau. Central (Raxaul).

Riopa punctata (Gmelin).
Riopa punctata, Leviton, Myers, and Swan, 1956, p. 11 (Dharan [1000 ft.]).
RANGE IN Nepau. Eastern (Dharan).

amily Varanidae

Varanus flavescens (Cray).
Empagusia jlavescens, GUNTHER, 1860, p. 159 (Nepal); 1861, p. 215 (Nepal).
Varanus flavescens, GUNTHER, 1864, p. 65 (Nepal).

RANGE IN NEpAu. Central (without exact locality data’).

Varanus monitor (Linnaeus).

Varanus heraldicus (Gray), Gray, 1845, p. 8 (Nepal). Gwtnruer, 1860, p. 160
(Nepal); 1861, p. 215 (Nepal).

Varanus dracaena (Gray), GUNTHER, 1864, p. 65 (Nepal).

Varanus monitor, SmirH, 1935, p. 402 (Nepal). Lrevrron, Myers, and Swan, 1956,
p. 11 (sight record; below Khandbari [4000 ft.], and north of Dhankuta).

RANGE IN Nepau. Eastern (below Khandbari, north of Dhankuta; both
sight records); Central (without exact locality data’).

Suborder Serpentes
Family Boidae

Python molurus (Linnaeus).

Python molurus, GUNTHER, 1861, p. 215 (Nepal; known from colored drawing).
WALL, 1907, p. 155 (Bichiakoh [Nepal terai]).

RANGE IN NEPAL. Central (Bichiakoh).

Family Colubridae

Boiga ceylonensis ((iinther).
Dipsadomorphus nuchalis (GUNTHER), WALL, 1924, p. 872 (Chitlong).
Boiga ceylonensis, SMITH, 1943, p. 351 (Chitlong).

RANGE IN Nepau. Central (Chitlang).

Boiga multifasciata (Blyth).

Dipsadomorphus multifasciata, Wart, 1907, p. 157 (Chitlong); 1924, p. 871
(Chitlong).

Boiga multifasciata, Smirn, 1943, p. 357 (Nepal).
RANGE IN NEPAL. Central (Chitlang).

VoL. XXXII] SWAN AND LEVITON: HERPETOLOGY OF NEPAL 113

Boiga trigonata (Schneider).

Chicago Natural History Museum 83087 (Ampipalbhanjan [4000 ft.]) ;
CNHM 83089 (Katmandu [4000 ft.]|).

RANGE IN NEPAL. Central (Ampipalbhanjan, Katmandu).

Coluber fasciolatus Shaw.
Coryphodon fasciolatus, GUNTIIER, 1861, p. 218 (Nepal).
RANGE IN NEPAL. Central (without exact locality data’).

Elaphe cantoris (Cantor).

Spilotes reticularis (Cantor), GUNTHER, 1858, p. 249 (Nepal); 1860, p. 163 (Nepal);
1861, p. 218 (Nepal).
Composoma reticulare, GUNTHER, 1864, p. 245 (Nepal).

RANGE IN NEPAL. Central (without exact locality data’).

Elaphe hodgsonii (Giinther).

Spilotes hodgsonii GUNTHER, 1860, p. 156 (Nepal and Ladak [15,200 ft.]; 1861, p.
218 (Nepal).

Compsosoma hodgsonii, GUNTER, 1864, p. 246 (Nepal). TrmeroBaLp, 1876, p. 166
(Nepal).

Coluber hodgsonti, ScLATER, 1891, p. 31 (Katmandu).

Elaphe hodgsonii, SMiru and Barrerssy, 1953, p. 704 (Tarakot [10,500 ft.]).
RANGE IN NEPAL. Central (without exact locality data?) ; Western (Tara-

kot).

Elaphe radiata (Schlegel).

Elaphe radiata, Leviron, Myers, and Swan, 1956, p. 12 (Arun Valley [1000 ft.]).
Chicago Natural History Museum 83098 (Hitora [4500 ft.]).
RANGE IN NEPAL. Eastern (Arun River Valley); Central (Hitaura).

Liopeltis rappii (Giinther).

Ablabes rappii Giiwruer, 1860, p. 154, pl. 26, fig. B (Nepal and Sikkim [5340 ft.]);
1861, p. 217 (Nepal); 1864, p. 225 (Nepal).

Ablabes owenii, GUNTHER, 1861, p. 217 (Nepal).

Liopeltis rappii, WAL, 1924, p. 865 (Nepal). Smirn, 1943, p. 186 (Nepal).
RANGE IN Nepau. Central (without exact locality data’).

Lycodon aulicus (Linnaeus).

Lycodon aulicus, GUNTER, 1860, p. 164 (Nepal); 1861, p. 219 (Nepal); 1864, p. 316
(Nepal). ScuiaTer, 1891, p. 14 (Katmandu). Watt, 1907, p. 152 (Katmandu
[4500 ft.]). Smiru, 1943, p. 263 (Nepal).

Chicago Natural History Museum 83090 (Katmandu [4000 ft.]).
RANGE IN Nepau. Central (Katmandu).

Natrix himalayana (Giinther).

Tropidonotus himalayanus GUNTHER, 1864, p. 265, pl. 22, fig. H (Nepal and Sikkim).
THEOBALD, 1876, p. 178 (Nepal).
RANGE In Nepau. Central (without exact locality data’).

114 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Natrix parallela (Boulenger).
Tropidonotus parallelus, BOULENGER, 1913, p. 337 (Maikhola Valley [7000-10,000 ft.]).
RANGE IN NeEpAu. Eastern (Mai Khola).

Natrix piscator (Schneider).

Tropidonotus quincunciatus Schlegel, GUNTHER, 1858, p. 63 (Nepal); 1860, p. 162

(Nepal); 1861, p. 217 (Nepal); 1864, p. 260 (Nepal).
Tropidonotus piscator, WALL, 1907, p. 157 (Pharping [5000 ft.]).

Museum of Comparative Zoology, Harvard College 58224 (Patlikhot) ;
MCZ 58235-58237 (Pokhara, Phewatal).

Chicago Natural History Museum 83081-83082 (Ampipalbhanjan [4000
ft.]); CNHM 83097 (Patan).

RANGE IN NEPAL. Central (Ampipalbhanjan, Patan, Pharping, Pat-
likhot, Phewatal, Pokhara).

Natrix platyceps (Blyth).

Tropidonotus chrysargus (nec Boie) GUNTHER, 1858, p. 70 (Nepal); 1860, p. 162
(Nepal); 1861, p. 217 (Nepal). Watt, 1907, p. 156 (Chitlong).

Tropidonotus platyceps, GUNTHER, 1860, p. 162 (Nepal); 1861, p. 217 (Nepal); 1864,
p. 264 (Nepal [4000-7000 ft.]). TuroBaLp, 1876, p. 174 (Nepal). WAttL, 1907,
p. 152 (Pharping [5000 ft.]).

Tropidonotus firthi WALL, 1914, p. 166 (Chitlong).

Rhabdophis firthi (Wall), WALL, 1923, p. 606 (Chitlong).

Natriz platyceps, SM1TH, 1951, p. 728 (Thangjet [5000 ft.]). Smirn and BATTERSBY,
1953, p. 703 (Jumla [7600 ft.], Khanglagaon [8000 ft.], Balangra Pass
A000 N EIDE
RANGE IN NeEpAL. Central (Chitlang, Pharping, Tangjet); Western

(Balangra Pass, Jumla, Khanglagaon).

Natrix stolata (Linnaeus).

Tropidonotus stolatus, GUNTHER, 1858, p. 68 (Nepal); 1860, p. 162 (Nepal); 1861,
p. 217 (Nepal). ScLaATeEr, 1891, p. 39 (Katmandu). WaAtt, 1907, p. 156 (Gowchar,
Pharping [5000 ft.]).

Chicago Natural History Museum 83095 (Katmandu [4400 ft.]).
RANGE IN NEPAL. Central (Gowehar, Katmandu, Pharping).

Oligodon albocintus (Cantor).

Simotes purpurascens (nec Schlegel) GtUinruecr, 1858, p. 245 (Nepal); 1860, p. 161
(Nepal) ; 1861, p. 216 (Nepal).
Simotes punctulatus GUNTHER, 1864, p. 217 (Nepal).

RANGE IN NEPAL. Central (without exact locality data2).

Oligodon arnensis (Shaw).

Simotes russellii (Daudin), GUNTHER 1858, p. 24 (Nepal); 1860, p. 161 (Nepal);
1861, p. 216 (Nepal); 1864, p. 213 (Nepal).

RANGE IN NEPAL. Central (without exact locality data?).

VoL. XXXIT] SWAN & LEVITON: HERPETOLOGY OF NEPAL 115

Oligodon erythrogaster Boulenger.

Oligodon erythrogaster BouLENGER, 1907, p. 217 (Nagarkote [6000 ft.]). Watt,
1923, p. 321 (Nepal). Smirn, 1943, p. 232 (Nagarkote); 1951, p. 728 (Thangjet
[5000 ft.]. Smiru and Barrersspy, 1953, p. 707 (8 miles west of Tibrikot
[8500 ft.]).

RANGE IN NEPAL. Central (Nagarkote, Tangjet); Western (Tibrikot).

Psammodynastes pulverulentus (Boie).
Psammodynastes pulverulentus, WALL, 1910, p. 76 (Butal).
RANGE IN NEPAL. Central (Butal |= ? Butwal]).

Pseudoxenodon macrops (Blyth).
Pseudorenodon macrops, BOULENGER, 1913, p. 338 (Maikhola Valley).
RANGE IN NEPAL. Eastern (Mai Khola).

Ptyas mucosus (Linnaeus).

Coluber dhumna Cantor, 1839, p. 52 (Nepal).

Coryphodon blumenbachii (Merrem), GUNTHER, 1858, p. 111 (Nepal); 1860, p. 163
(Nepal); 1861, p. 218 (Nepal).

Zamensis mucosus, WALL, 1907, p. 157 (Kakani, Gowchar).

Ptyas mucosus, Leviton, MYERS, and Swan, 1956, p. 12 (Tamur River Valley
[500 ft.J).
RANGE IN NEPAL. Eastern (Tamur Valley); Central (Gowehar, Kakani).

Sibynophis collaris (Gray).

Ablabes collaris, GUNTHER, 1858, p. 28 (Nepal); 1860, p. 161 (Nepal); 1861, p. 216
(Nepal); 1864, p. 228 (Nepal).
RANGE IN NEPAL. Central (without exact locality data’).

Sibynophis saggittarius (Cantor).
Sibynophis sagittarius, Smirn and Batrerspy, 1953, p. 704 (Dang Plain [1500 ft.]).
RANGE IN NEPAL. Western (Dang Plain).

Trachischium fuscum (Blyth).

Trachischium fuscum, GUNTH= 8, 1860, p. 161 (Nepal); 1861, p. 215 (Nepal).
Ablabes fuscus, GUNTHER, 1864, p. 225 (Nepal [8500 ft.]).

RANGE IN NEPAL. Central (without exact locality data‘).

Trachischium guentheri Boulenger.
Trachischium guentheri. Scrarer, 1891, p. 11 (Katmandu).
RANGE IN NEPAL. Central (Katmandu).

Trachischium tenuiceps (Blyth).
Ablabes tenuiceps, GUNTHER, 1864, p. 224 (Nepal). THEOBALD, 1876, p. 154 (Nepal).
Trachischium tenuiceps, WALL, 1907, p. 156 (Chandragiri [8000 ft.]). Smirn, 1945,
p. 323 (Nepal).
RANGE IN Nepau. Central (Chandragiri).

116 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4rH SER.

Xenochrophis cerasogaster (Cantor).
Tropidonotus cerasogaster, GUNTHER, 1861, p. 218 (Nepal; based on colored drawing).
RANGE IN NEPAL. Central (without exact locality data’).

Zaocys nigromarginatus (Blyth).

Coryphodon carinatus (part) GUNTHER, 1858, p. 250 (Nepal); 1860, p. 163 (Nepal);
1861, p. 219 (Nepal).

Zaocys nigromarginatus, GUNTHER, 1864, p. 257 (Nepal [7100 ft.]). THroBALD, 1876,
p. 172 (Nepal). Smiru, 1943, p. 165 (Nepal).
RANGE IN NEPAL. Central (without exact locality data’).

Family Elapidae

Calliophis macclellandii (Reinhardt).

Elaps univirgatus GUNTHER, 1858, p. 231 (Nepal); 1860, p. 164 (Nepal).

Callophis macclellandii, GUNTHER, 1861, p. 219 (Nepal); 1864, p. 249 (Nepal). Trro-
BALD, 1876, p. 214 (Nepal). ScLaTER, 1891, p. 56 (Katmandu). Smirn, 1943, p.
423 (Nepal).
RANGE IN Nepau. Central (Katmandu; see footnote “2” regarding

Ginther records, p. 107).

Family Viperidae

Agkistrodon himalayanus (Giinther).

Ancistrodon himalayanus, SMirn and Barrerspy, 1953, p. 704 (8 miles west of Tibri-
kot [8500 ft.], Jumla [9500 ft.], Sialgarhi [9000—10,000 ft.], Turikot [10,000 ft.]).
RANGE IN NEPAL. Western (Jumla, Sialgarhi, Tibrikot, Turikot).

Trimeresurus albolabris Gray.

Lachesis gramineus, (part) Watt, 1907, p. 157 (Katmandu).
Trimeresurus albolabris, SmMirH, 1943, p. 523 (Katmandu); 1951, p. 728 (Thangjet
[5000 ft.], Syarpagaon [9000 ft.]).

RANGE IN NepAL. Central (Katmandu, Syarpagaon, Tangijet).

Trimeresurus monticola Giinther.

Trimeresurus monticola GUNTHER, 1864, p. 388, pl. 24, fig. B (Nepal). Scrarer, 1891
(Katmandu).

Trimeresurus maculatus (nec Gray 1842, but Gray 1853), Giinrmer, 1858, p. 266
(Nepal).

Parias maculata, GUNTHER, 1860, p. 164 (Nepal); 1861, p. 220 (Nepal).

Lachesis monticola, BOULENGER, 1896, p. 548 (Nepaul). WALt.tL, 1907, p. 157 (Chitlong,
Kakani).

RANGE IN NEPAL. Central (Chitlang, Kakani, Katmandu).

Trimeresurus stejnegeri Schmidt.

Chicago Natural History Museum 83081-83082 (Ampipalbhanjan [4000
ft.|) ; CNHM 83088 (25 miles north of Katmandu [4000 ft.] ).
RANGE IN Nepau. Central (Ampipalbhanjan, Katmandu).

Vou. XXXII] SWAN AND LEVITON: HERPETOLOGY OF NEPAL iG

RECORDS OF UNCERTAIN STATUS

AMPHIBIA
Family Pelobatidae

Scutiger sikkimmensis (Blyth).

UNCERTAIN ADDITIONAL RECORD. Sclater (1892, p. 30) records this species
from the Singalehla Range, Sikkim, from an altitude of 12,000 ft. This alti-
tude suggests a collection station on the frontier between Sikkim and Nepal.

Family Ranidae

Rana swani Myers and Leviton.

UNCERTAIN IDENTIFICATION. Uncertain identification of two small frogs
taken from above the Arun River [1500 ft.| has been recorded by Leviton,
Myers, and Swan (1956, p. 9).

REPTILIA
Family Colubridae

Coluber monticolus Cantor.

UNCERTAIN SYNONYMY. Cantor’s species (1839, p. 52) based on a speci-
men from Nepal may be a synonym of Oligodon arnensis (fide Boulenger,
HB845 py 229).

Hurriah sanguiniventer Cantor.

UNCERTAIN SYNONYMY. The identification of Hodgson’s drawing of a
very distinctive snake described by Cantor (1839, p. 52) as H. sanguwini-
venter and based on a specimen from the Valley of Nepal is uncertain.
Smith (1943, p. 257) assigns the snake to the genus Lycodon (see also Giin-
ther, 1864, p. 222).

Lycodon jara (Shaw).

UNCERTAIN NEW RECORD. Smith (1943, p. 260) indicates that this animal
is found in the “Eastern Himalayas as far west as longitude 85°.” This
would place the western limits in the vicinity of Katmandu. We know of
no specific records to substantiate this distribution.

Natrix subminiata (Schlegel).

UNCERTAIN NEW RECORD. Wall (1923, p. 606) indicated that this species
was found from “Nepal to Sikkim.” No specific collection sites were men-
tioned by him. Smith (1943, p. 302) states, “The whole of the Indo-Chinese
subregion as far as Sikkim in the north-west.”

118 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4rit SER.

Simotes octolineatus (Schneider).

UNCERTAIN SYNONYMY. Giinther (1861, p. 216) recorded a specimen iden-
tified as S. octolineatus from Nepal. This certainly is in error for Simotes
[= Oligodon | octoinectus is an Indonesian species. Hither Giinthee’s identi-
fication was in error or there was a mixup in locality data. Perhaps Ginther
had before him a specimen later referred to as O. erythrogaster by Boulen-
ger, a species known from Nepal which bears a superficial similarity to the
Indonesian species in color pattern.

Spilotes melanurus Schlegel.

UNCERTAIN SYNONYMY. Two half grown specimens identified as S. mela-
nurus were recorded from Nepal by Giinther, 1858, p. 97; 1860, p. 163;
1861, p. 218). That species is known from Indonesia; it has not been taken
in the Indo-Burmese region. Either Giinther’s identification was in error
or there has been a mixup in locality data.

Family Elapidae

Naja tripudians Merrem.

UNCERTAIN NEW RECORD. Concerning the distribution of Naja tripudians,
Gunther (1861, p. 219) states, “No record from Nepal” but “Hodgson pre-
sumes it to be there.” Smith (1943, p. 426) indicates that the subspecies
N. n. kaouthw is found “ ... as far west as Nepal;” and in his map (fig.
139, p. 484) both N. n. naja and N. n. kaouthia are shown to occur within
the frontiers of Nepal. We do not know of any specific records of this snake
having been taken in that country though we agree with Hodgson and
feel certain it is there.

Family Viperidae

Trimeresurus viridis (Daudin).

UNCERTAIN SYNONYMY. GiNTHER (1861, p. 220) records a specimen of
T. viridis from Nepal. That nominal species has been placed in the synon-
ymy of T. gramineus, a species common to Peninsular India but which has
not been reported from as far north as Nepal. Giinther’s specimen may
belong to T. stejnegert, a species which is closely similar to 7. gramineus
and which has been reported from Nepal.

THE ZOOGEOGRAPHY OF NEPAL AND ADJACENT AREAS

The Indo-Chinese Subregion of the Oriental Region as defined by Wal-
lace (1876) includes large portions of southeastern Asia and extends as a
distinet western spur into the forested Himalayas. The fauna of Sikkim
clearly exhibits affinities with Burma, Thailand, and southern China,

VoL. XXXII] SWAN AND LEVITON: HERPETOLOGY OF NEPAL 119

whereas that of the western Himalayas appears to be a composite of faunal
elements drawn largely from the Indian Subregion of the Oriental Region
and the adjacent Palearctic Region. The country of Nepal (figure 1)
bridges an intervening zone between the differing eastern and western
Himalayan faunae. It represents an area of diminution, or cessation, of the
Indo-Chinese Subregion.

An analysis of the distribution of Himalayan amphibians, lizards, and
snakes suggests that the typical Indo-Chinese herpetofauna is essentially
intact as far west as Sikkim and the Darjeeling District of Bengal. Fifty-one
species found there (46 per cent of the local fauna) have a continuous dis-
tribution into the mountains; they also extend into Burma and other south-
eastern Asian portions of the Indo-Chinese Subregion (species designated
TE and IW im table I [see appendix] and summarized in figure 2). Thirty-
two additional species (29 per cent of the local fauna) are confined to the
Himalayas (EH, EW, NE in table I). And a Panoriental and Indian com-
plement of 29 species (25 per cent of the local fauna) found in the herpeto-
fauna of the Himalayas east of Nepal extends from the plains of India into
the mountains.

Only 15 typically Indo-Chinese species are known in Nepal, and one ad-
ditional species, Ophisaurus gracilis, should be found there (on the basis of
its presence beyond Nepal in the western Himalayas).

The obvious diminution of the Indo-Chinese fauna to the west of Sikkim
is not very likely an artifact resulting from incomplete information and
insufficient collection data from Nepal. The fauna of the western Himalayas,
from the Nepalese frontier to the vicinity of Simla, is relatively as well
known as is the plains fauna of India. If the distribution of the amphibians,
lizards, and snakes in the areas contiguous with Nepal is analyzed, a fair
approximation of the anticipated Nepalese species may be obtained. Of the
widespread Panoriental and Indian species known to ascend into the moun-
tains to the east and west of Nepal (in table I, species designated OR which
are also recorded in the West Himalayas and Sikkim-Darjeeling), 8 may be
anticipated in the mountains of Nepal. All have been collected there. Simi-
larly, 7 of 9 expected Panoriental-Indian lizards have been collected in
Nepal. However, only 12 of 28 anticipated Panoriental-Indian snakes have
been obtained. If the same analysis is applied to amphibians, lizards, and
snakes that are restricted to the mountains and are found on both sides
of Nepal (i. e., in the western Himalayas and Sikkim-Darjeeling, table I,
EW, IW and MB), the results indicate a collection of 2 out of 3 antici-
pated amphibians and all 9 anticipated snakes. A single lizard, Ophisaurus
gracilis, which is rare in the western Himalayas falls in this category and
has not been obtained in Nepal. In summary then, 10 of 11 anticipated
amphibians, 7 of 10 anticipated lizards, and 21 of 37 anticipated snakes
have been collected in Nepal. This information may be used as an approxi-

120 CALIFORNIA ACADEMY OF SCIENCES [Proc. 411 SER.

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phibians and reptiles have been obtained.

VoL. XXXII] SWAN AND LEVITON: HERPETOLOGY OF NEPAL 121

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VoL. XXXIT] SWAN AND LEVITON: HERPETOLOGY OF NEPAL 123

mate index of the present knowledge of the Nepalese fauna, and sufficient
information is available to permit a meaningful assessment of the zoo-
geography of the country. It is evident then that the dramatic diminution
from 51 Indo-Chinese species in Sikkim-Darjeeling to 15 Indo-Chinese
species in Nepal is scarcely due to ignorance of that fauna. It is significant
that the Indo-Chinese fauna in Nepal is a montane fauna and from the
preceding data it can be seen that collections in the mountains of Nepal
have yielded 11 of 13 anticipated montane species.

The striking diminution from 51 Indo-Chinese species in Sikkim-
Darjeeling to 15 Indo-Chinese species in Nepal is, in all likelihood, associated
with the obvious faunal barrier of the Singalehla Range, on the eastern
frontier of Nepal. However, the occurrence of two Indo-Chinese snakes,
Natrix parallela and Pseudoxenodon macrops, which have been collected
west of this barrier (Mai Khola, southwest of Sundakphu) offers a tenuous
suggestion that a gradual decline in Indo-Chinese species may occur in the
lam district of Nepal or the Tamur Valley. This zone of reduction would
be of limited extent inasmuch as collections made in the Arun Valley
some 50 miles from the eastern frontier of Nepal reveal a marked change
in the herpetofauna. Among 16 species of amphibians and reptiles obtained
in this area only one, Cosymbotus platyurus, may be considered as typically
Indo-Chinese. It is conceivable that outposts of Indo-Chinese reptiles and
amphibians may be localized in isolated forests in the Arun Valley where
aspects of the flora and some insects and birds are characteristic of Sikkim.
But the primary herpetofauna indicates a more easterly termination of
the Indo-Chinese Subregion.

One factor affecting this apparent termination is the greater human
population and more extensive cultivation of the land in eastern Nepal.
Agriculture and associated deforestation is predominant up to 7000 feet and
intact forests are, for the most part, above this level. Along the valley
bottom and near the river itself, fields and villages are less frequent and
lowland forests in various degrees of preservation are usual in this situation.
Deep within the Himalayas where the lowest valley elevations exceed 3000
feet, the valley forests show the greatest similarity to the forests of Sikkim.
It is perhaps in such isolated localities that the Indo-Chinese fauna remains
relatively distinct.

A second factor affecting the reduction of the Indo-Chinese fauna is the
apparent replacement of the Indo-Chinese forms by species typical of the
Indian plains. This is clearly emphasized in the Arun Valley by the abun-
dance of lowland species such as Calotes versicolor and Ptyas mucosus; it is
even more strikingly demonstrated by the recent discovery of Bufo ander-

Figure 2. Comparisons of faunal groups in Nepal and adjacent areas. Species
have been included in eastern, central, or western Nepal if they have been taken in
areas both to the east and to the west.

124 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4vH SER.

soni, Microhyla ornata, and Elaphe radiata, four plains species reported
from the eastern Himalayas for the first time in 1956 (Leviton, Myers,
and Swan).

Additional information concerning the fate of the Indo-Chinese fauna
may be obtained from an analysis of the well-known herpetofauna of the
western Ilimalayas adjacent to the western frontier of Nepal. Whereas
there are 112 species of reptiles and amphibians in the Sikkim-Darjeeling
area (figure 2), the mountains between Simla and Nepal hold 54 species,
of which five are typically Indo-Chinese. The major portion of the west
Himalayan herpetofauna is composed of Panoriental-Indian species (31
species or 57 per cent), but there is a sizeable representation of endemic
Himalayan species primarily of Indo-Chinese affinity (18 species, 33 per
cent). The presence of this latter discrete group obviously distinguishes the
western Himalayan fauna, and it does not seem fitting to assign the western
Himalayas to the Indian Subregion.

Aside from the ubiquitous Panoriental-Indian forms, the Indo-Chinese
species (augmented by the endemic Himalayan species many of which
are closely related to typical Indo-Chinese forms) represent a clear majority
of the mountain forms in the western Himalayas. Although there is a great
reduction in the typical Indo-Chinese fauna in a westward trend from Dar-
jeeling to Simla (51 species to 5 species) (figure 2) there is also a concurrent
reduction in the total fauna (112 species to 54 species). Exeepting a few
additional species characteristic of the Palearctic, no outstanding new as-
semblage of amphibians or reptiles appears in the western Himalayas. The
-anoriental-Indian fraction remains essentially the same (29 species to
31 species); this draws attention to the fact that the western recession of
the fauna is restricted to the inountain forms. It is clear that there is no
eategory to which the western Himalayas may be assigned legitimately
except to the Indo-Chinese Subregion. There is a manifest change in the
herpetofauna of the Indo-Chinese Subregion at the eastern frontier of
Nepal, but unless the 700 miles between Sikkim and Simla is considered a
prolonged transition zone, the Indo-Chinese Subregion apparently extends
through Nepal as far as Simla in the western Himalayas.

The boundary lines which cireumseribe any zoogeographie province are
approximations. Since the time of Alfred Russell Wallace, faunal limits
have been indicated geographically with simple lines. Between contiguous
major faunal zones, small or large transitional areas have been assumed.
In Nepal and the western Himalayas, what may appear outwardly to be a
prolonged transition area of the Indo-Chinese fauna is perhaps something
different. The dissected mountainous region distinguishes cultivated valley
slopes harboring a predominant Panoriental-Indian fauna from the montane
and valley forests where a higher incidence of Indo-Chinese and endemic
species are in evidence. The frequeney and size of these faunal pockets

VoL. XXXII] SWAN AND LEVITON: HERPETOLOGY OF NEPAL 125

apparently decreases westward with a concurrent decrease in precipitation.
But there is no dramatic increase of a new and different fauna; there is
essentially a simple diminution of the mountain and forest forms. Beyond
Simla, where Palearctic species become frequent, the Indo-Chinese species
which still persist may create a minor transition zone of a classical sort
within the boundary of Palearctic Region. In summary, it would seem that a
transition zone should reflect a decrease of one fauna with an accompanying
proportional increase of another fauna as a new province is approached.
When there is an alteration of the fauna involving primarily a reduction of
this fauna, and there is no accompanying significant increase in a new
assemblage of species, the area involved is not in reality a transition zone.
The latter apples to the Indo-Chinese Subregion in Nepal and the western
Himalayas. The region seems to represent an intermediate condition where
a fauna of eastern affinities is in the process of being extinguished while
a fauna of western origin has not yet become prominently developed. On
the map (figure 3), the protracted and weak extension of the Indo-Chinese
Subregion is indicated by localized areas in the mountains which are sepa-
rated by segments of the Indian Subregion. The very minor inroads of the
Palearctic species into the Himalayas proper to the east of Simla have been
superimposed onto the Panoriental-Indian fauna rather than the Indo-
Chinese fauna.

At the present time, 69 species of amphibians and reptiles have been
reported from Nepal. Of these, two frogs, Rana polunini and Rana swana,
are known to be restricted to the country; both were discovered in recent
vears. Rana polunini is closely related to R. blanfordi and may be a high
altitude derivative of the latter widespread Himalayan species. Rana swani
shows affinity with 7'omopterna ranids in southwest India and is probably
a Himalayan representative of a relictual group of frogs which at one time
were more widely distributed. Nepal is also the type locality of a few other
species whose distributions extend beyond its frontiers (Rhacophorus
marimus, Oligodon erythrogaster, and Trimeresurus monticola), and the
types of several species have been recorded together from Nepal and Sikkim
(Hodgson’s and Hooker’s collections, respectively ).

Some approximation of the number of endemic amphibians and reptiles
to be anticipated in Nepal can be obtained by comparing the fauna in Nepal
with that of equal areas to the east and west. In the portion of the Himalayas
extending from Nepal to the gorge of the Brahmaputra in the east, there are
now 14 frogs, 7 lizards, and 9 snakes (total 30 species) which can be con-
sidered as restricted to this part of the Himalayas. Nearly all of these species
are from Sikkim and Darjeeling, inasmuch as Bhutan and the Assam
Himalayas are in a category of ignorance far greater than Nepal. In the
western Himalayas, in an area extending from central Kashmir to the
western frontier of Nepal, 2 frogs, 2 lizards and 2 snakes (total, 6 species)

[Proc. 471 SER,

CALIFORNIA ACADEMY OF SCIENCES

126

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VoL, XXXII] SWAN & LEVITON: HERPETOLOGY OF NEPAL 127

may be considered indigenous. From this it follows that the number of
endemic forms in Nepal is between 6 and 30 species, with the probability
that the number is closer to the former figure.

The large group of typically Himalayan species (table I, EH, EW, WH)
is composed for the most part of species having affinities with the Indo-
Chinese Subregion, for example, T'rachischium guentheri, a member of a
genus with several representatives in the eastern Himalayas and the Khasi
Hills. On the other hand Himalayan species such as Elaphe hodgsoni and
Leiolopisma himalayanum, which range widely in the far western Hima-
layas, Kashmir and Ladak, as well as Nepal, likely emanate from areas to
the west where there are other representatives of the same genera. These
species and a few others with a similar distribution may be considered as
having Palearctic affinities. That portion of the Palearctic Region adjacent
to western India has been delineated by Wallace as the “Mediterranean
Subregion” and species associated with this subregion which enter the
Himalayas are identified by the symbol ME in table I.

In addition to the purely mountain species that can be assigned either
to the Indo-Chinese or Palearctic faunae, there is an aggregation of species
which generally occupy an elevated wet zone in the Himalayas and which
are fundamentally related to species in Western China. Preeminent in this
eroup is the high altitude pelobatid frog Scutiger sikkimmensis. Although
this zone is best delineated in the Himalayas by a single amphibian species,
its identity is supported by many plants (species groups of Rhododendron,
Primula, ete.) and birds (particularly pheasants such as the genera Itha-
ginis, Lophophorus, and Tragopan). In addition, the lizard genus Japalura,
with 5 recognized species in west China (other species also present in the
Khasi Hills, north Burma and north Viet Nam) and 5 species in the Hima-
layas, may represent this biogeographic province in the Himalayas. Scutiger
sikkimmensis, Japalura major, and Japalura tricarinata are among the few
amphibians and reptiles found above 10,000 feet in the outer, wetter
Himalayas. These species have not been segregated into a special category
in table I, but they have been considered as belonging to a distinct subregion
(in figures 2-4).

Other species with a distinct association with the fauna of west China
are the frogs Scutiger alticola, Scutiger mammata®, and Altirana parkert.
These three species have been collected on the Tibetan Plateau immediately
north of Nepal and presumably may be found in those parts of the country
which are extensions of the plateau. They are apparently widespread species
but not continuous in their distribution, for they are restricted to scattered
lakes and streams. Scutiger mammata, or a closely related species, probably

5. Aelurophryne mammata was recently shown to be congeneric with Scutiger sikkimmensis (Myers
and Leviton, 1962).

Figure 3. Zoogeographic subregions of Nepal and adjacent areas.

[Proc. 4v11 SEr.

CALIFORNIA ACADEMY OF SCIENCES

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Vor. XXXII] SWAN AND LEVITON: HERPETOLOGY OF NEPAL 129

extends as far as Kashmir. It appears, therefore, that the West-Chinese
fauna is divided between a wet Himalayan group and a contingent on the
dry plateau of Tibet. The latter group expresses Asian Palearctic rather
than Oriental characteristics and the West-Chinese herpetofauna as a whole
may be considered as essentially Palearetic. This is emphasized by the
distribution of other vertebrate species, including hynobiid salamanders,
which, like Scutiger, extend west across Tibet to Afghanistan. Perhaps the
West-Chinese Province is in reality identical to the western portion of the
Manchurian Subregion of Wallace which was conceived as a vast area em-
bracing most of northern China and Japan with an extension into the Hima-
layas. A tentative appraisal of this subregion suggests that the mountain
fauna of west China is sufficiently distinct to merit the status of a related
but distinct subregion of the Palearetie.

Whereas the amphibian representatives on the plateau adjacent to Nepal
are affiliated with West-Chinese species, the saurian complement (Agama,
Phrynocephalus, Alsophylax, ete.) is entirely related to the fauna of
Afghanistan, Iran, and the Caspian area. This is the eastern limit of the
Mediterranean Subregion as conceived by Wallace, but the invasion of Tibet
by members of this subregion may well extend the Mediterranean Palearctic
to the borders of Sikane and Szechwan. One of the species typical of this
eroup of Mediterranean-plateau species, Leiolopisma ladacense, has been
collected in Nepal near the Tibetan border at 18,000 feet, a record altitude
for any amphibian or reptile.

The Tibetan Plateau herpetofauna directly north of Nepal and in
places actually entering within the frontiers of Nepal is therefore a curious
mixture derived from two Palearctic subregions. The amphibians (and per-
haps the relictual Tibetan snake, Thermophis baileyz) live near water and
have West-Chinese affinities, whereas the lizards have opposite origins and
represent a Mediterranean extension into the area. Similarly there is a
West-Chinese group of frogs and lizards in the Himalayas proper which
are only weakly delimited by altitude from a small group of Himalayan
lizards and snakes with Mediterranean affinities (Agama _ tuberculata,
Elaphe hodgsoni, ete.). This Himalayan-Mediterranean fauna is, however,
a minority conflux of species among the far more numerous Panoriental-
Indian, Indo-Chinese, and Himalayan amphibians and reptiles. This con-
fusion of faunal groups is an outcome of both vertical and horizontal pat-
terns of distribution in the mountains; the interaction between these two
factors as they relate to the zoogeography of Nepal is shown in figures 3
and 4.

Figure 4. Altitudinal distribution of Nepalese amphibians, lizards and snakes
and of the zoogeographic subregions of the Nepal Himalayas. Some species have
been included among the Plateau species that have not been obtained within the
borders of Nepal.

130 CALIFORNIA ACADEMY OF SCLENCES [ Proc. 471 SER.

The altitudinal distribution of the various faunal groups (figure 4) is
characterized by wide overlapping zones rather than discrete altitude-
limited belts. Two altitudinal zones which are occupied exclusively by
discrete faunal groups are confined to elevations between 11,500 feet and
13,500 feet on the southern exposure of the Himalayas and between 17,000
and 18,000 feet on the Tibetan slope (West-Chinese and Mediterranean
subregions, respectively). On the Tibetan slope of the Himalayas and on
the plateau below 17,000 feet, as described above, the Mediterranean and
West-Chinese faunae occupy similar altitudes but can be distinguished
ecologically inasmuch as the lizards have dry habitats whereas the am-
phibians live near streams and lakes. Below 11,500 feet and above 8,500
feet on the southern flank of the Himalayas, the West-Chinese and Medi-
terranean species are joined in the Nepal Himalayas by two Himalayan
species, Rana leibigii and Rana polunini, which appear to range considerably
higher than other species with Indo-Chinese affinities. The elevated distri-
bution of these frogs (and possibly R. blanfordi, which has not yet been
eolleeted in Nepal), suggests that they may be related to the group of ranid
frogs (R. pleuraden, R. phrynoides, and R. boulengert) which are dis-
tributed in the mountains of West-China. If this relationship has any
real basis, the Himalayan frogs may have West-Chinese rather than Indo-
Chinese affinities. At the present time they are viewed as exceptional species
and have not been considered as part of the West-Chinese fauna.

If a few altitudinal records obtained from the western Himalayas are
utilized to reinforce the limited information from Nepal itself, the alti-
tudinal distribution of the Mediterranean fauna in the Himalayas of Nepal
appears more intelligible. Species such as Hlaphe hodgsoni and Agkistrodon
himalayanus appear to range on both the north and south sides of the
Himalayas and may be part of both plateau and Himalayan segments of the
Mediterranean fauna. They ascend to much higher altitudes on the north
slope, both species apparently exceeding 15,000 feet. Toward the southern
flank of the Himalayas these species and other Mediterranean forms such
as Agama tuberculata, Leiolopisma himalayanum, and Natrix platyceps
rarely exceed 10,000 feet. These same species, however, are found as low
as 3000 feet. Some of these species, particularly Agama tuberculata, appear
to occupy rocky, drier areas and to some degree these Mediterranean forms,
despite their wide range, are ecologically distinet from the Indo-Chinese,
West-Chinese, and Panoriental-Indian species which are altitudinally con-
tiguous. It appears therefore, that the Mediterranean species have wide
altitudinal ranges and in western Nepal extend from the southern slopes
continuously through the Himalayas to join the Mediterranean fauna of
the plateau. In eastern Nepal and the eastern Himalayas, the Mediterranean
fauna is divided, as is the West-Chinese fauna, into separate Himalayan and
plateau portions.

VoL. XXXII] SWAN AND LEVITON: HERPETOLOGY OF NEPAL 131

The altitudinal distribution of the Indo-Chinese fauna (including typi-
eally Himalayan species) extends from the lower foothills up to 11,500
feet (if Rana hebign and R. polunini are considered). It is, however, pri-
marily confined between 3000 feet and 7000 feet and at these altitudes it is
rivalled by the more widespread Panoriental-Indian species. The latter
fauna, however, is predominant below 3000 feet.

In many ways the altitudinal distribution of amphibians and reptiles
mirrors the altitudinal distribution of vegetation. The dominant forest
categories (figure 4) fairly coincide with the range of the Indo-Chinese
and Panoriental-Indian amphibians and reptiles which are confined to the
Lower, Middle, and Upper Monsoon forests. However, most of these species
do not exceed an altitude of 6000 feet, which approximates the altitude of
killing frosts in winter. Rhododendron trees of several species extend from
below 6000 feet to tree line, which in eastern Nepal reaches a maximum
altitude of 13,500 feet. They are, however, more prominent in the deciduous
forests (8,500 feet to 10,500 feet) and in the zone of conifer forests (10,500
feet to 13,500 feet). The altitudinal distribution of japalurid lizards re-
sembles the distribution of rhododendron forests; this vegetation mixes
with the Upper Monsoon forests and the deciduous forests. In the latter
zone Japalura tricarinata and J. major are the commonest reptilian species
encountered. The rhododendron-conifer forests coincide strikingly with the
range of Scutiger sikkimmensts. In general, it would seem that the Monsoon
forests harbor Oriental species (Panoriental-Indian and Indo-Chinese),
whereas forests which include rhododendron trees incorporate a West-
Chinese herpetofauna. The Mediterranean species extend over the alti-
tudinal range of nearly all of these forests but they are primarily non-forest
species.

A belt of piedmont forest mixed with swamps and open fields, a foothill
and plains region known as the Terai, separates the Himalayas from the
true plains of India and Nepal. The amphibians and reptiles of this zone
are known primarily from collections obtained in the Terai near the Dar-
jeeling District east of Nepal. With the possible exception of Elachistodon
westermanni and Bungarus lividus there appear to be no reptiles or am-
phibians which are exclusively confined to the Terai zone. The herpetofauna
of this intermediate area seems to be composed entirely of Panoriental-
Indian species although many of the Terai forms do not extend far into the
adjacent plains. Examples of this latter fauna which appear to be confined
to the plains within 100 miles of the Himalayas are: Rhacophorus taeniatus
and R. tuberculatus of Assam, and Boiga forsteni of Bihar and Uttar Prad-
esh. The Terai fauna and the plains fauna of Nepal are very poorly known
and the great majority of collections in this portion of the country were
made by Brian Hodgson and his collectors. Comparing the Nepalese plains
with the well-known fauna of the adjacent Indian plains, it is found that 5

132 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4711 SER.

chelonian species have been collected in Nepal out of 11 species which may
readily occur in Nepal. Similarly, 7 of 15 lizards have been obtained and 11
of 37 plains snakes. Perhaps the best indication of the status of collections
on the plains of Nepal is reflected in the fact that at the present time there
seems to be no published reeord of the occurrence of the common cobra in
Nepal (though Maleolm Smith [1948, p. 434] shows it to be present in his
map). It is obvious that collections on the Nepal plains would rapidly in-
erease the known fauna of the country and, inasmuch as the plains fauna
is richer in adjacent Bengal, it may be assumed that most new records of
amphibians and reptiles will be obtained from the southeastern corner of
Nepal.

Much of the preceding discussion has referred entirely to amphibians,
lizards, and snakes as the Nepalese ecrocodilians and chelonians are primarily
plains species and are poorly represented in collections. Only one tortoise,
Testudo elongata, collected in Nepal can be considered as typical of the
Indo-Chinese Subregion, the remainder being Indian and Panoriental. The
presence of 7. elongata in Nepal and the oceurrence of Geoemyda tricari-
nata and G. trijuga in the Darjeeling District immediately east of Nepal
point to a eurious affinity between the eastern Himalayas and the hills of
Chota Nagpur over 300 miles south of Nepal and separated from the
Ilimalayas by the wide plain of the Ganges. All of these tortoises, together
with two Indo-Chinese lizards (Leiolopisma sikkimensis of the eastern
Himalayas as far as central Nepal and Sphenomorphus maculatum of the
eastern Ilimalavas and southeast Asia), are found in the restricted area
of Chota Nagpur. The five species do not represent a majority of the reptiles
in the area (although they may have some ecological distinction), but as a
eroup of relictual species surrounded by ubiquitous Oriental forms, their
presence and significance may be emphasized by referring this isolated
pocket to the Indo-Chinese Subregion. The species just cited indicate that
a continuous fauna, presumably a forest fauna, likely extended across the
Gangetic plains at one time. The hiatus in the distribution of the Indo-
Chinese fauna calls attention to a recession of the subregion and climatic
changes of the past. The Chota Nagpur link with the Indo-Chinese Sub-
region is based on the presence of identical species in the two areas. If
genera and allied species are taken into consideration, some representatives
of the Indo-Chinese Subregion can be associated with species in Chota Nag-
pur, the western Ghats of southwest India and the Ethiopian Region. This
distribution presumably represents a relictual situation dating from the Ter-
tiary Period, and it would seem that the tortoises and lizards of Chita Nag-
pur, still specifically linked to the Indo-Chinese Subregion, represent the last
segment of this earlier distribution pattern.

It would seem that the fauna of the Gangetic plains is newer and is
occupying a region which has been deforested through the ageney of a

VoL. XXXII] SWAN AND LEVITON: HERPETOLOGY OF NEPAL 133

changing climate which in turn has recently been aided by man. The dis-
tributional patterns of animals of central Asia and of the Himalayas also
show the effects of climatic change, obviously reflecting a trend toward arid
conditions. The narrow altitudinally limited wet and cold zone oceupied by
Scutiger sikkimmensis harbors populations which are totally isolated from
each other by deep tropical valleys, and it is obvious that this West-Chinese
fauna was formerly more widely distributed. On the plateau of Tibet, the
isolated pockets of amphibians and snakes indicate that this portion of
the West-Chinese fauna was at one time also far more extensively dis-
tributed. Superimposed on this West-Chinese fauna are the wide-ranging,
successful, desert lizards related to the Mediterranean Palearctic. The
total pattern clearly suggests the encroachment of an arid environment in
central Asia which has depopulated a preceding West-Chinese fauna and
isolated a fringe of this fauna on the outer slopes of the Himalayas, which
are still wet and cool. The Mediterranean invasion has also involved the
southern slopes of the Himalayas, but, as noted earlier, the several species
which typify this expansion are as yet a very minor percentage of the total
herpetofauna. The climatic modifications on the Gangetic plain are pre-
sumably related to the changes in central Asia, and the plains fauna also
contains a number of species which evidence Mediterranean affinities. It is
significant that the deforested Gangetic plain has been occupied by ubiqui-
tous species from the surrounding areas, leaving few if any relictual
species. However, occupation of this area has not been accomplished by any
substantial number of Indo-Chinese species. The latter, it seems, have re-
treated with the forests and remain remarkably discrete in the Himalayas east
of Nepal. In Nepal itself and in the Himalayas to the west, the plains fauna
has made successful inroads into the Himalayan valleys and the Indo-Chinese
fauna in these areas appears to be declining toward a relictual status.

In summary, the zoogeography of Nepal is a complex of interdigitating
faunal subregions. The Indo-Chinese fauna (composed of species which
range from southeastern Asia into the Himalayas together with the ma-
jority of strictly Himalayan species) becomes greatly reduced on the
eastern frontier of Nepal, but continues westward through Nepal, pri-
marily in isolated forest areas which lie between 3000 and 7000 feet. A
large group of widespread Panoriental-Indian species extend from the
plains and are the dominant species on the plains of Nepal and in the
Valleys and the lower foothills. They are also largely coextensive in their
altitudinal range with the Indo-Chinese fauna. The plains fauna is also
composed of species which do not extend into the mountains. Several species
of lizards and snakes which extend from Kashmir into Nepal from the west
and which are derivatives of the Mediterranean Subregion fauna, have a
wide altitudinal distribution. On the southern slopes of the Himalayas they
are found generally below 10,000 feet; but in western Nepal this fauna links

134 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER.

with Mediterranean species of the Tibetan Plateau and on the frontier of
Nepal these species ascend as high as 18,000 feet. The Mediterranean fauna
in Nepal overlaps the altitudes occupied by the Indo-Chinese and Pan-
oriental-Indian species, but they are to some extent ecologically distinct
from the latter groups. The highest elevations on the southern exposure of
the Himalayas are dominated by a few species with West Chinese affinities.
Similarly, on the plateau immediately north of Nepal (and presumably in
some places within Nepal itself), there are additional West-Chinese species
which are not continuous with the Himalayan forms. The Tibetan Plateau
is therefore dominated by wide-ranging Mediterranean lizards which overlap
a seemingly relictual group of West-Chinese amphibians. It appears that
the West-Chinese and Indo-Chinese faunae are retreating from Nepal and
Tibet, whereas there is a concurrent expansion of Mediterranean species
from the west and Panoriental-Indian species from the south. With Indo-
Chinese, Indian, and Panoriental faunae of the Oriental Region combining
vertically and horizontally in wet and dry areas with two groups of Mediter-
ranean Palearctic species and two groups of West-Chinese Palearctic species,
the zoogeography of Nepal is somewhat bewildering. It has a labyrinthian
quality which rivals the distribution patterns of any area of similar dimen-
sions on the face of the earth, and it is particularly instructive concerning
the manner in which two major zoogeographie regions converge and meet
in a montane faunal barrier which is common to both regions.

VoL. XXXII] SWAN AND LEVITON: HERPETOLOGY OF NEPAL 135

APPENDIX

Table I represents a list of amphibians and reptiles known to occur in
Nepal and the immediately adjacent areas. Information on the Tibetan,
Indian, and Himalayan species and their distribution has been compiled
with the aid of an unpublished work by Swan (1947). The area designations
and the symbols utilized in the table are as follows:

ADJACENT Puatns. The plains of India within approximately 50 miles
of the southern frontier of Nepal which are essentially continuous
with the plains area of Nepal itself.

W. Himatayas. The mountainous area between Simla and the western
frontier of Nepal.

W. NeEpau, C. Nepau, E. NEPA. Divisions of Nepal into western, central,
and eastern portions based upon the drainage systems of the major
rivers (see map).

SIKKIM-DARJEELING. The mountainous area of Sikkim and the Dar-
jeeling District immediately to the east of Nepal.

ADJACENT TiBET. The southern plateau area of Tibet adjacent to Nepal
which is north of the primary erest of the Himalayas.

X. This indicates the known presence of the species.

X1, 2, 3. These indicate the known presence of the species in Nepal and
the number of times it has been obtained by different collectors in
the area.

O. This indicates a real expectation of the species in Nepal based on the
known presence of the species in areas to the east and the west.

P. This indicates the possible presence of the species in Nepal based on
the known presence of the species on the plains of India adjacent
to Nepal.

OR. Species with a distribution typically Indian or Panoriental which
may be found in wide areas of the plains and mountains of India
and may frequently extend into S. E. Asia.

IE. Species with a distribution characteristic of the Indo-Chinese Sub-
region and which extend into the Eastern Himalayas.

IW. Species with a distribution characteristic of the Indo-Chinese Sub-
region and which extend into the Himalayas west of Nepal.

EH. Species with a distribution essentially confined to the Hastern
Himalayas.

EW. Species with a distribution essentially confined to the Himalayas,
including both eastern and western portions.

136 CALIFORNIA ACADEMY OF SCIENCES

[ Proc. 4711 SER.

WH. Species with a distribution essentially confined to the Western

Himalayas.

ME. Species with a distribution which appears to extend from the
Mediterranean Subregion into the Himalayas as far as the Eastern

Himalayas.

TB. Species with a distribution on the Tibetan plateau adjacent to

Nepal.

The data presented in table I are summarized for convenient reference
in table II. The data have been grouped by areas and major subordinal

or ordinal groups of amphibians and reptiles.

A distributional analysis of faunal groups by faunal subregions is

presented in table ITT.

TAPLE I

Distribution of Amphibians and Reptiles of Nepal and Adjacent Areas

Adjacent
Plains

AMPHIBIA
APODA

CoECILUDAE
Ichthyophis glutinosis

URODELA
SALAMANDRIDAE

Lp OLOURLLONAUCTIALCOS (eae eee

ANURA
BREVICIPITIDAE

MACE OMA LO ONT ee x

BUFONIDAE
BU OROMG CTS 01) =a een ee x.
SU OMT CLG ONS = ee ee

SU OMICLONO SULGLIL Sea XG

PELOBATIDAE

Megophrys major Dec ce
SCHULUO Crom (HUELGO LO een een
SGWEUCET MIC TLIN CLG =e

Scutiger sikkimmensis........................

Western

Himalayas

West
Nepal

Central
Nepal
East

Nepal

Sikkim-

Darjeeling

Tibet

Distribution
Category

Vou. XXXII] SWAN AND LEVITON: HERPETOLOGY OF NEPAL

TABLE I (Continued )

Distribution of Amphibians and Reptiles of Nepal and Adjacent Areas

ANURA (Continued)

Adjacent
Plains

RANIDAE

AUEOIRTUG NON AIXE Ae eee

EO TUCUMICLTUTECL TUG. UUU eee ene ee nec ceeeecee

HL ILCINOS S ITU CTUS US tac ers ee nese Aeceee ce
LEONA: OUCMOU OTOH ae ee oe
ROG. WFERCH) 2 ee ee x
DT LOUI (GU CUILO MTVU UGUD Sees eee x
LROIOG. If OU ONO) ee eae
TREC KODE Gs
LEACH OKT DOL Se ee
IRGHG) AHH ODOG | ee
TERCNOD TOTEDLOOC WOH POS eee x
TRS UOC ON ee cee
TEXT) TDI HE SER OG Pee eee
LEROIEINGL © JODUOH OU eee ee
LECH OGE SYORCHA Rasa ee
IRCHNTA OCH OC ee ae Ae ee ea ER xX
TROHOC) EU CO e tli Aa teen ee
(STAC DOSS CHRON KORO So a

DMUnOUS NWUMOLGYONG. 22 ee

RHACOPHORIDAE

PTAOMEWSRONNANAGI.-. 2 ce
UUCULUL SOUL OU Seen ee: Soe
DOGO DMOTRUS FET COU nese es oe nee
Rhacophorus leucomystaa..................
MWACOPNOTUS MaculatUs...-......--------- xX

Fe LAGCOD MOTUS. “NYMLTNWS =. ----------ceee-e-=

REPTILIA

CHELONIA

EMYDIDAE

Geoclemys hamiltoni................-.-------- xX
GeOoeMmyad tiACOTINGtd — 2... ==.
HEDETROEKTR TRON ONT cee
ERORUCUO= GUT Alien xX
CHU Cs ANONG ORG xX

Western

mM eM

Himalayas

West

Se eo ©

Nepal

Central
Nepal
Hast

Nepal

Sikkim-
Darjeeling

AM MMM nM MK OM

hd
1

Tibet

Distribution
Category

Ei neo Ge shki-« 6 6 2a eis
Cet ett aaa eae De

OR
IE
IE

138 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

TABLE I (Continued)
Distribution of Amphibians and Reptiles of Nepal and Adjacent Areas

2 80 =
a Shs ~ ae Bee
Su. a s - By =o
38 2S o©3 Be os eee Eb
R (Continued) ga G8 89 9S. 2 3 3 ee
EPTILIA ontinue = Ra ® is = “4
ace ah EO FZ 04 BZ we seer

Emypipat (Continued )

Kachuga kachuga.........--.----- one Peso x 12 5: a OR

ECHO CS ONUUG eee

~
me)
o)
es)

EG CHAU ot CCGG ees es eee

>
a)
me}
ae)
o)
Be)

TESTUDINIDAE

TES GUC ONCUOTUG CE Cree ane >aL @) x 189)

TRIONYCHIDAE

COUT GAOL oe eee xX P Ol Ae OR
UISUSS CHIU SHA OIETLGLC,L Opeee ene eee x B 12 12 OR
i ONA ERO OMOCELC Se ee xX 12 xP OR
TEFL OFA NTU eee a ee x 12 OR
TALON LUC UG een ee eee xX 12 12 P OR
CROCODILIA

CROCODILIDAE

C7OCOOUUS MD ULUUSU Se x 12 12 Ie OR
COE UTHIOS: GOUT ORO CTS ccccecccce cece ON 12 xe le OR

SQUAMATA-SAURIA
AGAMIDA®

AG OUT TIU UGLY OL10 Os neeeee

A
4
ze
ce]

Agama tuberculata__..__.... tees ne erases xX X1 X2 ME
CULES UCSC x x X1 X4 Xi x OR

Japalura kumaonensis..........-.---.---..-.--

”~
a
q

Japalura major

4
a
a
ss

Japalura tricarinata...........----.----.-------- De (0) x EH
OD OUT OA CLC O CIE Ce ee x EH
Phrynocephalus theobaldi.. =... x TB
SLOT) OMUEUG CIE OMU Cee ee x Ie P 12 OR
ANGUIDAE

Ophisaurus gracilis.............------------------ Ke O O O x IW
GEKKONIDAE

ANUISOPOI ODAC, ONTO OUI) ee eee x TB
Cosymbotus platyurws.................------- ole E Xs Sex IE

Cyrtodactylus fasciolatwss.-- x WH

Vou. XXXII]. SWAN AND LEVITON: HERPETOLOGY OF NEPAL

TABLE I (Continued )

Distribution of Amphibians and Reptiles of Nepal and Adjacent Areas

=
SQUAMATA SAuRIA (Continued) Zs
GEKKONIDAE (Continued)
Cyrtodactylus gubernatoris_............
Cyrtodactylus khasiensis.................-..--
Cyrtodactylus lawderanus...............-.-- x
(GQOONO. ATCT O ee nee x
Henniadachyluus OOWnINng?1......-.-.----..--.--.--
[HECK LLES WiC) =e x
Hemidactylus flaviviridis...................- ox
FENEMVAGCTYVES, JNENQMUWS...-.----.......-------- xX
EMCI ALGLYUWSNOORNOUL. 2 =.
SCINCIDAE
Leiolopisma sikkimense...................
Leiolopisma himalayanum...........-----
IGeLOVOPUSING TAAGGENSE........-----.--..-—--_--
LWACHOCULTD., (COP ULON Ke ene eee XxX
WVORGTUMY COM TUCLGIEUG TG -= 2 xX
Mabuya multifasciata......- ee oes
Ophisops jerdoni__.... Me oes ee eee xX
eh OMI UUO OU IUGCE OE see cee xX
PF tsDC CUO UUTU CU UE Osanna oe a on ae means xX
Sphenomorphus indicum......................
Sphenomorphus maculatum.....-
VARANIDAL
WCACHOUS: UCHIOS GG ae x
W GIRASOLE x
WGRUROS SOOO eee xX
SQUAMATA-SERPENTES
BOIDAE
LEY COMCUS 22 a ee x
EIU OMMmITUOLUTU Se: saree =e eS ae x
COLUBRIDAE
AUT UC TAULL GIN ONUELCUTUGH see
ANGEHUUIG NOSWEO. 2. ---- 2 wee xX

Western
Himalayas

Nepal

West

X1
X1

Central
Nepal

X2
X1

East
Nepal

Darjeeling

Sikkim-

Tibet

139

Distribution

Category

VoL. XXXII] SWAN AND LEVITON: HERPETOLOGY OF NEPAL 141

TABLE I (Continued)
Distribution of Amphibians and Reptiles of Nepal and Adjacent Areas

5 = : ~ ie : Ss}

G4 8s = £4 a eo = 5
SQUAMATA-SERPENTES (Continued ) de os E S 8 B % 5 ca 2 =
COLUBRIDAE (Continued) Stig Hh GE FO aan eS
INCASE TO CTD OHI CH [OHO ioe ee eee ener “al ©) x TE
INVOHETPOGR (XO ROT IU QU Fee eee oe eee al Ke IE
IN GAPOGE. (DOS OOHEOY (eect ene meee ee x x O XAaaO xe OR
DOR POLD FOUN TIO Oa ee nee ee x xaill Xoo >< ME
INOUE OLD. SUQUGT AG aaa re ee x xX O X40 xe OR
INT ORETPUEE SOMDY TON HOOT SD ae oe eee x IE
OMGOCONOUOOCINCIUS= = = ees sal» <6) € IE
CIOIG DO Tim GTEC TUS US essa 2c ae ene axe aXe O xall OR
(QWMGIOGNOMD. (CO RCUC ULI ace eee eee x P P OR
Onigodon. eryLinrogaster........-....----.- ell x @ NE EW
OMGOGOTE GUQUaNG {eT 2... -.- 2 X EH
(ONG OGKOT. TH GUCH GOR een eee ere HE BH
TPO AQIS. GEOK OO NCH PUTIN ape eee ee x IE
PUP EOES TSO ECO ree oe ee ee axe IE
Psammodynastes pulverulentus...... Dal -@ x IB
PSGNIMODNIS CORMANGHUS =. 2-2-2. -222-2------ x d:€ P P 12 OR
IE ROMRTIODIDIVUS. UGA ex aa x P OR
PSCUCOLENOAON MGACTOPS: ..._.-.2--:-------=-- X1 x IE
IPDS. GPU COBSO KS eee a ee eee x x O XZ eal: x OR
STOOD OOS. COUCH AO ee a ee eee x O X1 O x IW
SaDynNoOpnis Sagvtlarwus.._....-.-----22-2-<--=- xe Bx X1 OR
DU CUGTUUS CIUUUTIU RI AES CUM ene ooo as 2a 2. esses 22 =- Ox O X1 O x EW
LECGHUSGHVULNE: Quenvwent...------ xX1 O xe EH
TEROCIES GRIMM LLCUC.— = xX WH
TGWCHISGHIWM LENUIGEDS. =... X2 O >. KH
Xenochrophis cerasogaster..__......-------- x X1 OR
Zaocys nigromarginatus.............-----.--- Xi TO x I}
DASYPELTIDAE
Elachistodon westermanni.................- xX 12 >. EH
ELAPIDAE
TUG OU Sm ONLIUG OU OU CS ans eee wna x 18)
I BOO KOTFUS. (GONTPUNI GT =e ee ee x xX 12 1E P OR
PES URUGOGUUS i (DS GUCHU WS een oa anne Dak 24 OR

IBSPTOGKOPOS HOPMAN «se errno x EH

142 CALIFORNIA ACADEMY OF SCIENCES

TABLE I (Continued)

Distribution of Amphibians and Reptiles of Nepal and Adjacent Areas

_
S|
fed)
oO
=
SQUAMATA-SERPENTES (Continued ) 3
ELAPIpDAE (Continued)
UU G CUS ITVUO C Tee
PES QOVUG CUYGUS a UG OULU tae ae oe xX
Calliophis macclellandi......................--
TN EGRO he (OP) 0 eee eo eee ee ees axe
Onhop aguseGnn Gi x
TyYPHLOPIDAE
MaypNlOPS WOCMTAOTYMUCTIUS as
MAU OM) SIMO 1 CUTIE UTS serene ox
DODDS GOPODO is srcccrcce seacoast
TE DIBUOD SMOLUG OV ED 1S eee eee
EV DUODWS. (NOWTOCTUD xx c2cccaocseecoemeconsensaascee aX

VIPERIDAE

Agkistrodon himalayanus..................--
LTANVETRESILLILS QUO OUCO TUS mre nee
TUVERESIUTUS NEGU IG MURU TAU Sareea ee Fe
TIUUVCTESUTUS UONUCUGOU Oe nee
TEE AUTVCTEC SULTUSHED OD COT Tae
TUNE RESUTUS ES LCIILEG Clee eee

WG DOIRGR PUSS CN a i ao oe cee cee pace eee XG

Plains

Western
Himalayas

ial

oe)

Central
Nepal
East

Nepal

Ss © © ho

[Proc. 4TH SER.

Sikkim-
Darjeeling

AK mM MM OM

Tibet

Distribution
Category

S) Cys! wey
teas

EW
OR
IE
EH
OR

ME
IW
IE
IE
IE
IE
OR

Vou. XXXII] SWAN AND LEVITON: HERPETOLOGY OF NEPAL

TABLE II
Distributional Analysis by Areas:
Totals Based on Summary of Data Presented in Table I

a FS

S, GS =

SH Gi vs S35 oc

oS OF O & a a Ey 5
eee 40 2 EA om fe
Specles™ Collected. 2k ts leis. ie Beds, 8 att) 2 12
Additional species expected (QO)... 10 3 7
Plains species possibly present (P)... 0 0 0
CHELONIA
MECLESmICOMECtCd pee --2-c0-: 2s ie eee ts il 0 5
Additional species expected (OO)... 0
Plains species possibly present (PP)... 5
CROCODILIA
STOSCISS: COSC EL Ieee ee 2 0 0 i 0
Additional species expected (QO)... 0
Plains species possibly present (P) .. % if 2
SAURIA
SPEeclesmeolechedy sy eis 2S es 15 13 5 9
Additional species expected (QO)... 4 2
Plains species possibly present (P) = 5 4 a
SERPENTES
species collected) 202 2....2-.22e ee ets 331 31 5 9 4
Additional species expected (OO)... iM) 7 30

Plains species possibly present (P)... 9 9 14

Sikkim-
. Darjeeling

bo
-~]

143

Tibet

ioe)

[Proc. 4TH SER.

CALIFORNIA ACADEMY OF SCIENCES

144

satoeds 69 eee eae es Ba UO ON 1020) )

soroeds gpT ccc sawedg ny 1070,1 ‘samday pup sumquydwy

foe cout mee “68. TE" 298 = Ss cme ol G 9. &&

I L 0 0 I F 0 0 0 0 0 PR Re ee SS CST) ee rOC NI
G G € g G G 0 0 0 0 0 0 (HIN) WeARTRULLTT “P[—-UvOUR.L TO} po
r L 0 G I f 0 0 0 0 0 T (°° ** CHA) URArleUtyyT W10489 Ah
L 6 f G 0 0 0 0 0 0 € $ °° °° (MM) Uedereunry “M pure “Gy
L eG 6 G t 0 0 0 0 g Or ‘°° * CHM) uedvfeulyy W104sry
P G € g 0) | 0 0 0 0 [ T ‘(AJ) Uedvpeury] “M—esouttpoopuy
ae, or 6 [ee 1 i) I e 0 0 [ g °¢ (Gy) wedepeuryy “G[—-esoutypopuy
Bue Te. tech 9S 2k wl) OC Gi ee G 8 g ocr? (YO) TeyUelO-ueIpu]
[edoNn IV [edoNn ITV jedan lv [eden Iv 1eden IV [edoNn IV

soltjdey ¥ soyuadiag BLINVS BruolayO BI [poop viqrydury

sueiqrydury

poeurqurory

sdnowy ounng fo sishpup jouoynguysig (1 219L “Of $1101)
Tl) S1a¥ 1,

VoL. XXXII] SWAN AND LEVITON: HERPETOLOGY OF NEPAL 145

BIBLIOGRAPHY

ANNANDALE, NELSON
1907. Introductory note and Lacertilia. In: Boulenger, et al. Reports on a col-

lection of Batrachia, reptiles, and fish from Nepal and the Western
Himalayas. Records of the Indian Museum, vol. 1, pp. 149, 151-155.

BOULENGER, GEORGE ALBERT

1882. Catalogue of the Batrachia salientia s. ecaudata in the collection of the
British Museum. Second edition. London. xvi + 503 pp., 30 pls.

1894. Catalogue of the snakes in the British Museum (Natural History).
Volume II, containing the conclusion of the Colubridae aglyphae. Lon-
don, xi + 382 pp., 20 pls.

1896. Catalogue of the snakes in the British Museum (Natural History).
Volume III, containing the Colubridae (Opisthoglyphae and Protero-
glyphae), Amblycephalidae, and Viperidae. London, xvi + 727 pp.,
25 pls.

1907. Batrachia. In: Boulenger, et al. Reports on a collection of Batrachia,
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the Indian Museum, vol. 1, pp. 149-151.

1907. Description of a new snake from Nepal. Records of the Indian Museum,
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1913. A list of the reptiles obtained by N. H. Stevens in Upper Assam and the
Eastern Himalayas. Records of the Indian Museum, vol. 9, pp. 337-338.

1920. A monograph of the south Asian, Papuan, Melanesian, and Australian
frogs of the genus Rana. Records of the Indian Museum, vol. 20, 226 pp.

BOULENGER, GEORGE ALBERT, NELSON ANNANDALE, FRANK WALL, and
CHARLES TATE REGAN

1907. Reports on a collection of Batrachia, reptiles, and fish from Nepal and
the Western Himalayas. Records of the Indian Museum, vol. 1, pp. 149—
158, pl. 6.

CANTOR, THEODORE EDWARD

1839. Spicilegium Serpentium Indicorum. Proceedings of the Zoological So-
ciety of London, 1839, pp. 31-34, 49-55.

GRAY, JOUN EDWARD

1842. Synopsis of the species of rattlesnakes, or family of Crotalidae. Zoologi-
cal Miscellany, pp. 47-51.

1844. Catalogue of the tortoises, crocodiles, and amphisbaenians, in the collec-
tion of the British Museum, London. viii + 80 pp.

1849. Catalogue of the specimens of snakes in the collection of the British
Museum. London. xv + 125 pp.

1850. Catalogue of the specimens of Amphibia in the collection of the British
Museum. Part II. Batrachia gradientia, etc. London. 72 pp., pls. 3—4.

1845. Catalogue of the specimens of lizards in the collection of the British
Museum. London. xxviii + 289 pp.

146 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER.

Gray, JOHN Epwarp (Continued)
1853. Descriptions of some undescribed species of reptiles collected by Dr.
Joseph Hooker in the Khassia Mountains, East Bengal and Sikkim-
Himalaya. Annals and Magazine of Natural History, ser. 2, vol. 12, pp.
386-392.
GUNTHER, ALBERT
1858. Catalogue of the colubrine snakes in the collection of the British Museum.
London. xvi + 281 pp.

1860. Contributions to a knowledge of the reptiles of the Himalaya Mountains.
Proceedings of the Zoological Society of London, 1860, pp. 148-175, pls.
25-28.

1861. List of the cold-blooded Vertebrata collected by B. H. Hodgson, Esq., in
Nepal. Proceedings of the Zoological Society of London, 1861, pp. 213-
PAPA be

1864. Reptiles of British India. London. xxvii + 452 pp., 46 pls.
Hooker, Josepu D.
1854. Himalayan Journals. London. xxix + 408, xii + 487 pp.
Houston, RicHarp C., and WILLIAM LONG
1955. The California Himalayan Expedition to Makalu. Sierra Club Bulletin,
vol. 40, pp. 1-17.
HUNTER, WILLIAM WILSON (SIR)
1896. Life of Brian Houghton Hodgson. London. 390 pp.
LeEviron, ALAN EDWARD, GEORGE SPRAGUE MYERS, and LAWRENCE WESLEY SWAN
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MYERS, GEORGE SPRAGUE, and ALAN EDWARD LEVITON
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ger, Aelurophryne and Oreolalax). Copeia (in press).
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1951. The natural history of the Langtang Valley. In: H. W. Tilman, Nepal
Himalaya. Cambridge University Press, pp. 242-265.
SCLATER, WILLIAM LUTLEY
1891. List of snakes in the Indian Museum. Calcutta. x + 79 pp.
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SMITH, MaLcotm ARTHUR

1931. The fauna of British India, including Ceylon and Burma. Reptilia and
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1935. The fauna of British India, including Ceylon and Burma. Reptilia and
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Indo-Chinese subregion. Reptilia and Amphibia. Vol. 3. Serpentes.
London. xii + 583 pp., 1 map.

Vout. XXXII] SWAN AND LEVITON: HERPETOLOGY OF NEPAL 147

SmitTH, MALtcotm ArtTuur (Continued)

1951. Ona collection of amphibians and reptiles from Nepal. Annals and Maga-
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SmITH, MALCOLM ARTHUR, and JAMES C. BATTERSBY

1953. On a collection of amphibians and reptiles from Nepal. Annals and Maga-
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THEOBALD, WILLIAM
1876. Descriptive catalogue of the reptiles of British India. Calcutta. x + 238 +
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WALL, FRANK

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WALLACE, ALFRED RUSSELL
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asi iii
_ A 6 eee?

= t=

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

FOURTH SERIES

G Dallas Hanna Anniversary Volume

Vol. XXXII, No. 7, pp. 149-218, 28 figs., 3 tables

May 20, 1963

CENTRODERA SPURCA (LECONTE) AND TWO NEW

SPECIES RESEMBLING IT, WITH BIOLOGICAL

AND OTHER NOTES
(COLEOPTERA: CERAMBYCIDAE)

Hugh B. Leech

California Academy of Sciences

CONTENTS

ee SNC AGTU TPS Ay eis tks ea stdin sae Pl A hs et elane 2 nee
PCM OM OMTCTNUS Ms Hee hee as cc 2 he sod bn td wea eo ay woe Shee
Remimecnuroacre o. Wi.-WeConte-... 0. 6. s60 04 26 3.2 a ee eee eee
RR MeMPSeTIE A AM mee Re Sai la cata, PAL ia Wi aeia Sie st Pt-o eee

Key to the species of the Centrodera spurca group ...............-.
OER SI Ured WEOONE 2 6. so. '0 oo alate oe a ele awl anton s Dena ende alee
Centrodera autumnata Leech, new species .........5..-0002.00005
Cemroderasdayzlueecth, new Species’... . 2... fe ule Sales Ja ticle ee ae

The wing venation of some species of Centrodera ................
The eggs of the species of the C. spurca group
neweva Ol CeMUnO@ Crd SPUTCE lees eles Rela he Woe Soleo dee ale
Riese CentinOdera SPUTCO Loss. cds oo Ae RES Ll Bae ee

tinnne unl Cc

MAY 2.9 1982

Marine Biological Laboratory;
DBR AR YX

RASS.

150
151
154
156
158
160
173
178
184
187
188
190

150 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser.

CoNTENTS—Cont.

A partial life history of Centrodera spurca, with notes on other

SPECIES” sxc acess cs ees Ha Sas Same ae hoe nial es ee il
Flight period of the adults of the Centrodera spurca group ........ 196
Habits of the adults of Centrodera spp.

Daytime retreats’ 2..5. 5. 2d: see eon cee ee eee 201

Attraction to light. .\.wacdge deen oelasapeeece teen ae 201

Ability to walk-on vertical panes‘of glass ...9.2: 225502 eee 203

Adults on-flowers. ..o0Ss seac6 ceeo mel ce 2 oaks o ae ee 203

Attraction of some western species to baits .................. 205

Stridulation:« S.. $4 << magewae 2 eae) ahead Occ eee eee 205

Copulation: . <4 42.028 2 seme ose. e odio Snr eee 206
Dubious records: . .....< eh. NES ER AS a 207
Taiterature “Cited «5.5 <<. ¢ .. dsatewccressene oxenew oecdrencone eet tae eee 208

INTRODUCTION

Centrodera spurca (LeConte) is one of the few large Cerambycidae at-
tracted to hght during the relatively cool evenings of the Pacific Coast of
Canada and the United States. Of the species so taken, the only ones likely
to be confused with it are those described as new in this paper, plus Ortho-
leptura valida (LeConte) and O. insignis Fall’ (fig. 1).

A big brownish beetle with antennae fully an inch long in the male
(fig. 2), crawling up the elass or buzzing through an open window after
dusk, is enough to excite most collectors. Having taken the species in
British Columbia I greeted those which landed on my windows in Mill
Valley, California, as old friends, but bottled them just the same. The
first came in mid-May, 1948, and it did not take long to build up an ade-
quate series. Those which followed in June and July were not collected,
but in late August I finally realized that the beetles on the windows were
smaller than usual and had a different facies. Comparison with the earlier
set confirmed that there were two species, and examination of material in
collections showed that both had been identified as C. spurca for many years.
In fact the example used for the drawing in R. Hopping’s paper on the
Lepturini (1937, pl. III, fig. 4) proved to be the new species.

Knowing of my interest in these large centroderas, Willis C. Day eol-
lected some at light along the Scott river in northern California in August,
1949, and they turned out to be a second new species. A larger series of
this from adjacent Oregon was soon submitted by Arthur T. MeClay. From
their known distribution the two new species are allopatric, but C. spurca

1. Swaine and R. Hopping (1928, p. 38) synonymized Ortholeptura Casey under Anoplodera Mulsant,
but Linsley (1942, p. 51) raised it to generic status again.

VoL. XXXIT] LEECH: CENTRODERA SPURCA AND ALLIES 151

Figure 1. Males of Centrodera spurca and Ortholeptura valida, two species
which are attracted to light in many of the same areas. The short legs and unarmed
prothorax distinguish Ortholeptura at a glance.

is sympatrie with both of them. The three are almost enough alike to be
ealled sibling species, but probably were confused because C. spurca is so
distinctive and easily recognized that nobody bothered to examine a series
critically.

ACKNOWLEDGMENTS

It is a pleasure to acknowledge help from the owners of private collec-
tions, and the persons in charge of institutional collections, who have loaned
specimens for study; their names are listed below with the letters used to
designate the collections, many of which are referred to in the text.

In addition, the following persons have generously given their time to
answer questions, compare specimens with types, or verify other data:

152 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SEr.

Figure 2. A male of Centrodera spurca on a leaf of madrono, at night; the
beetles are nocturnal.

kt. P., Allen, J. Balfour-Browne, W. EF. Barr, R. S: Beal, Jn= Pe Darlime=
ton, Jr.. W. C. Day, K. M. Fender, L. G. Gentner, G. A. Hardy, J. N. Knull,
G. Kuschel, F. Lane, J. D. Lattin, the late G. P. Mackenzie, P. Rubtzoff,
J. Sedlacek, the late G. Stace Smith, P. J. Spangler, P. C. Ting. Through
the kindness of P. D. Hurd, Jr., I have been able to use a base map pre-
pared for the California Insect Survey of the Department of Entomology,
The University of California, Berkeley. E. G. Linsley and J. A. Chemsak
have read the manuscript, but the responsibility for any remaining errors
is of course mine. Help by members of my family in field work is equally
appreciated. HE. lL. Kessel kindly took two of the photographs.

AHH A. H. Howden, Ottawa, Ontario.

BM B. Malkin; collection now in the Chicago Natural History
Museum.

CAS California Academy of Sciences, San Francisco; includes speci-

mens from many private collections, such as those of E. C.
Van Dyke, F. E. Blaisdell, E. G. Linsley, J. O. Martin,
J. W. Green, R. Hopping.

CIS California Insect Survey, University of California, Berkeley
(P. D. Hurd, Jr., J. A. Powell, J. A. Chemsak).
CNC Canadian National Collection, Ottawa, Ontario (H. Howden).

DG D. Giuliani, San Anselmo, California.

VoL. XXXIT]J

DR
GHN
ILR
JGE
JNK
JS
LACM

LGG
ML
OHSU
OSDA

OSU
PMV

RBH
RWD
SDNHM
SISC

SRP
UBC

LEECH: CENTRODERA SPURCA AND ALLIES 153

D. Rentz, Novato, California.

G. H. Nelson, Colton, California.

I. LaRivers, Reno, Nevada.

J. G. Edwards, San Jose, California.

J. N. Knull, Ohio State University, Columbus, Ohio.

J. Schuh, Klamath Falls, Oregon.

Los Angeles County Museum, Los Angeles, California (F. S.
Truxal).

L. G. Gentner, Medford, Oregon.

M. Lundgren, Oakland, California.

Ohio State University, Columbus, Ohio (J. N. Knull).

Oregon State Department of Agriculture, Salem, Oregon (K.
Goeden).

Oregon State University, Corvallis, Oregon (J. D. Lattin).

Provincial Museum of Natural History, Victoria, British Co-
lumbia (G. A. Hardy).

k. B. Hutt, Pullman, Washington.

R. W. Dawson, Pullman, Washington.

San Diego Natural History Museum, San Diego, California (G.
Marsh and C. F. Harbison).

San Jose State College, San Jose, California (J. W. Tilden and
J. G. Edwards).

S. R. Piazza, San Jose, California.

University of British Columbia, Vancouver, British Columbia
(G. J. Spencer and G. G. E. Seudder; now ineludes the G.
Stace Smith collection).

University of California, Davis, California (A. T. MecClay; in-
eludes material from his own collection).

University of Idaho, Moscow, Idaho (W. F. Barr and A. Walz).

University of San Francisco, San Francisco, California (E. L.
Kessel).

United States National Museum, Washington, D. C. (P. J.
Spangler; selected specimens).

University of Washington, Seattle, Washington (M. H. Hateh;
the M. H. Hatch and K. M. Fender collections are included
in the University collection).

Washington State University, Pullman, Washineton (M. T.
James).

W. E. Hazeltine, W. R. Bauer and J. S. Buckett collected and
donated specimens.

154 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Genus CENTRODERA JJ. L. LeConte

Centrodera LeContr, 1850. Jour. Acad. Nat. Sci. Phila., 2nd Ser. (N. S.), vol. 1, no.
4, p. 325; LeContr, 1862. Proc. Acad. Nat. Sci. Phila., 1862, pp. 38, 41; La-
CORDAIRE, 1869. Hist. nat. insectes, vol. 8, pp. 426, 432; PRovANCHER, 1877.
Petite faune ent. Canad., vol. 1, pp. 580, 606; LrConTE, 1873. Smithson. Mis-
cell. Coll., no. 265, p. 328; LreConTr and Horn, 1883. Smithson. Miscell. Coll.,
no. 507, p. 313; WickHAM, 1897A. Canad. Ent., vol. 29, no. 4, p. 88; BLATCHLEY,
1910. Coleopt. Indiana, p. 1046, SwaAINE and R. Hoppine, 1928. Nat’l. Mus.
Canada, bull. no. 52, pp. 10, 14; Brapiey, 1930. Manual gen. beetles, p. 235;
CHAGNON, 1936. Le Nat. Canad., vol. 63, nos. 8 and 9, p. 205; R. Hoppine, 1937.
Nat’l. Mus. Canada, bull. no. 85, p. 10; Knut, 1946. Ohio Biol. Survey Bull.
39, pp. 151, 174; JAQquEs, 1951. How to know the beetles, p. 255.

Centrodera subgenus Apatophysis Chevrolat, GressirT, 1951. Longicornia, vol. 2,
p. 48.

Parapachyta CAsry, 1913. Mem. Coleopt., vol. 4, pp. 216-217.

It was J. Thomson (1864, p. 145) who designated Rhamnusium ? de-
coloratum Harris, 1841, as the type-species of Centrodera LeConte. Swaine
and R. Hopping (1928, p. 10) apparently overlooked this, and stated the
type-species to be “(Rhagium) decoloratum Harris,” by monotypy. In fact,
LeConte cited and described two species following his description of Cen-
trodera (p. 325; Centrodera was actually first given in the key on p. 316,
but without included species, and cited as Centroderus by error on p. 312):
1. C. decolorata, with snynonyms “Rhamnusium ? decoloratum! Harris Ins.,
Toxotus rubidus, Dej. Cat., T. rubidus [rudibus in Haldeman’s original
description, by a printer’s error], Hald. 58,” from Niagara and Massa-
chusetts. 2. C. picta, with the reference ‘“T'oxotus pictus Hald 58,” from
Pennsylvania and South Carolina.

It is surprising that Haldeman and LeConte did not include the generic
name Centrodera in their editing of the Melsheimer Catalogue, since in their
part of the Preface (1853, p. vill) they stated that so far as known to
them their additions to the original manuscript “include all the species
published up to January, 1852.”

Casey (1913, p. 216) proposed Parapachyta for “Pachyta spurca Lee.,
a large pallid and coarsely sculptured species of the true Pacifie coast
fauna.” Dr. Paul Spangler has been so kind as to examine Casey’s series
for me, and reports “In the Casey collection are four specimens under the
name Parapachyta spurca LeC. These agree with our C. spurca with the
strongly recurved hair on the base of the elytra and the hind margin of the
dth sternite very distinctly margined. Apparently Casey had LeConte’s
spurca.”

Gressitt (1947, p. 191) said that the Old World genus Apatophysis
Chevrolat, 1860, probably should be considered a subgenus of Centrodera

VoL. XXXII] LEECH: CENTRODERA SPURCA AND ALLIES 155

LeConte, 1850. Four years later (1951, pp. 48-50) he so placed it, without
further comment, and without giving characters to distinguish it from the
nominate subgenus.

According to Gahan (1906, pp. 68, 69) a characteristic of the species of
Apatophysis is that the males have the antennae “serrate,” i.e. segments
5 to 10 “compressed to a sharp edge in front and angulate at the apex.’ In
the two males I have seen, one identified as A. sinica A. Semenov-Tian-
Shanskij, the other as A. serricornis (Gebler), both have antennal segments
6 to 10 distinctly produced at the outer apical angle, and the elytra are
rather densely covered with fine, short hairs. The females are said to be
less pubescent than the males, or even glabrous, and to have the elytra a
little shortened, exposing the last one or two abdominal segments, thus re-
sembling prionines. Figures of both sexes of A. barbara Lueas are given by
Villiers (1946, p. 39, figs. 107, 108).

In the deseribed species of Centrodera the elytral hairs all arise from
the coarse elytral punctures. In the two males of Apatophysis mentioned
above, it was at once apparent that the elytral vestiture arises from all
over the interspaces between the coarse punctures, but few or none actually
from them. Dr. Frederico Lane has been so kind as to check this character
in the species of Apatophysis in the collections of the British Museum (Nat.
Hist.) : A. toxotoides Chevrolat, 1 male (and a female marked ? barbara
Lueas), Sahara; A. caspica Semenow, 2 males from Afghanistan, possibly
syntypes; A. kamarowi Semenow, 1 male, Turkestan; A. modica Gahan, 2
male syntypes, one marked type; A kashmiriana Semenow, 9 males and 3
females, Kashmir; A. montana Gahan, the male type, West Himalayas. He
summarizes his notes (letter of April 15, 1962) as follows: ‘All species of
Apatophysis examined have vestiture on the interspaces, and exceptionally
a few hairs as well in the coarser punctures.” He also suggests that the
palpi may offer characters for the generic separation of Apatophysis and
Centrodera.

On the bases of the differences in the elytral vestiture, the antennae of
the males, the body form and elytra of the females, and the distribution, I
believe that the species of Apatophysis are generically distinet from those
of Centrodera.

Gressitt cited the type-species of Apatophysis as Leptura serricornis
Gebler, 1843; but Thomson (1864, p. 147) designated A. toxotoides Chev-
rolat, 1860. Actually, the generic name was monotypic upon proposal,
since in both Chevrolat’s preliminary (1860A, p. 96) and formal (1860B,
p. 304) descriptions of A. toxotoides, no other species was mentioned in
combination with the name A patophysis.

Van Dyke (1927, pp. 102-103) gave a key to the Pacific Coast species
of Centrodera, but did not include C. spurca (LeConte).

156 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Swaine and R. Hopping (1928, p. 14, footnote 2) remarked that “The
genus Centrodera Lec. is doubtfully distinet from Xylosteus Friv.”; later
(1937, p. 14) R. Hopping wrote “The comparatively small eyes and type
of maculation will not allow it [7.e. the only American species, X. ornatus
LeConte, 1873] to be placed in the genus Centrodera, although C. picta
somewhat approaches this species in maculation.”” Gressitt (1947, p. 191)
removed Y. ornatus from Xylosteus to the genus Leptorhabdium Kraatz,
Isis):

MALE GENITALIA

In describing the curved male genital organ of Centrodera spurca (fig.
12) one faces the problem of deciding which side is morphologically dorsal
and which ventral. Because of the curvature, the organ lies on its side when
at rest in the abdomen, and is rotated during protrusion. This problem has
been referred to by Lindroth and Palmén (1956, p. 72) who state “The
simplest method is no doubt to speak about dorsal side, left side ete. ac-
cording to the position of the organ when extended and in activity. We
propose that, if necessary, the original (morphological) position of the
aedeagus and its details be expressed by the terms euw-dorsal, eu-ventral, eu-
dextral, eu-sinistral.”

Even this is not as simple as it sounds. Guignot (1931, p. 53), discussing
the male genitalia of Dytiscidae, in which the organ rests on its side, wrote
“Au moment de l’éreetion, aedeagus, tout en faisant saille par la fente
eénitale, tourne de 45° sous l’influence de ses muscles rotateurs. C’est dans
cette position simple d’érection (et non pas au moment de |’introduction
dans le vagin de la femelle, car alors le mouvement s’exagére et la pointe du
pénis devient presque antérieure) que sont faites toutes les descriptions, et
elles deviendraient incompréhensibles, si on ne se figurait pas nettement la
position de l’organe. A ce moment la base du pénis est antérieure, la sommet
est postérieur, le bord convexe est dorsal et le bord coneave ventral; un
parameére se trouve a droite et le second a gauche, le tegmen est dorsal et
plus ou moins a droite.”

But F. Balfour-Browne (1940B, pp. 126-128, figs. 2-4; see comparable
figs. in his 1932 book, p. 45, and pp. 23-24, fig. 6, in his 1940A book), also
discussing the Dytiscidae, draws the opposite conclusion: that the convex
side is ventral, the concave dorsal. His argument is based in large part in
conditions in typical hydrophilids, Hydrophilus spp. and Hydrobius
fuscipes (Linnaeus), in which the genital organ is flattened dorso-ventrally
and lies flat in the abdomen. He states (loc. cit., p. 126, with reference to
fig. 1 on p. 127) “An examination of the aedeagus of an Hydrophilid, where
there can be no question as to which is the dorsal side, shows that the open-
ing of the ejaculatory duct, the ‘gonopore,’ is always on the ventral side

VoL. XXXIT] LEECH: CENTRODERA SPURCA AND ALLIES 157

below the apex. In the Hydradephaga | which includes the Dytiscidae], the
gonopore is on the convex side.” On the other hand G. Kuschel, who has
made detailed comparative studies of the genitalia in the Curculionidae, and
investigations in other families of the Phytophagoidea, has found (verbal
communication, 1962), that on the basis of muscular and other attachments
of the genital capsule as it lies at rest in the abdomen, the opening through
which the intromittent organ is everted is always on the (morphologically )
true dorsal side.

The person who has studied the male genitalia of the Coleoptera most
broadly is R. Jeannel. The organs were used extensively in Jeannel and
Paulian’s 1944 elassification of the Order, but it is in Jeannel’s 1955 paper
that the subject is discussed and illustrated most fully. The range of strue-
tural variety shown is truly amazing, and suggests at once that it may not
be possible to give a statement as to which side is dorsal, and have it apply
universally. This is verified in Jeannel’s discussion of the tubular type of
median lobe, which consists of two longitudinal sclerites, one of which sur-
passes the other and forms the “apex,” as in Centrodera spp. He says (p. 22)
“On comprend ainsi que l’apex étant formé tantot par la paire sternale,
tantot par la paire tergale, l’orifice apical sera tergal ou sternal par rapport
a l’apex, selon les eas.” In Centrodera the orifice near the apex of the
aedeagus is comparable to Jeannel’s figure 5 of a trechid, and thus dorsal,
z.e. the convex side of the aedeagus is dorsal. This is in agreement with
figures by Villiers (1946, p. 8), Ehara in his major paper on the male
genitalia of Japanese Cerambycidae (1954), and most others who have il-
lustrated the organs of this family.

There is another basic subject upon which published opinions differ
ereatly, and that is the nomenclature of the parts of the genital capsule.
In the elassical study Sharp and Muir (1912) used the very descriptive
terms median and lateral lobes, the whole being the aedeagus, but pointed
out that the “lateral lobes” are not always lateral, and the term is thus in-
appropriate; paramere and tegmen were suggested. Jeannel (1955) followed
Sharp and Muir. Lindroth and Palmén (1956) and Lindroth (1957) prefer
penis and tegmen, the outer parts of the tegmen being the parameres. Snod-
grass (1957) uses aedeagus for the median lobe of Sharp and Muir, and
parameres for the lateral lobes; I am following his usage. Other views to
be considered are found in papers by Gilbert (1953), Wood (1953) and
Michener (1956).

In Centrodera decolorata the genital armature is elongate and only
slightly curved, so it is able to he on its ventral (concave) surface when re-
tracted into the abdomen, and is merely extruded, then bent downward and
forward, to be in position for copulation. In C. spurca and allies, it is more
strongly curved (figs. 12, 16, 17) and is nearly always found lying on its

158 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser.

left side, 7.e., with its apex pointing to the right side of the abdomen,
though occasionally almost as in C. decolorata. During protrusion for copu-
lation, an armature previously on its side must make a twist of 90 degrees.
In C. nevadica and other small western species examined it lies on its left
side when retracted.

KEY TO THE SPECIES OF THE Centrodera spurca GrRoUP

1. Elytral vestiture (in profile; fig. 3) inconspicuous, the hairs short, strongly
recurved, decumbent. Fifth abdominal sternite with hind edge differentiated,
narrowly thickened, usually clearly margined apically? and narrowly to base

Figure 8. Centrodera spurca. Part of an elytron to show the inconspicuous
vestiture of short, recurved, decumbent hairs.

at sides (fig. 5), often emarginate or broadly subcrenulate, especially in female.
Elytral apices each normally with small tooth at sutural angle. Outer antennal
segments of male smooth, satiny (fig. 6). Larger species, males averaging 22
mm. in length, females 25 mm. An early season species, late March—early
August, commonest in June and July; widespread, southern British Columbia
and northern ldahorto southern’ Calitornia (Giles 4) ee C. spurca
— EHlytral vestiture not very conspicuous, with some recurved hairs, but most of
those in basal half standing out freely at an angle (fig. 4). Fifth abdominal
sternite with at most an unobtrusive fine margin at apex in female. Elytral

2. Not appreciably margined in only four specimens of a total of 1061 studied; these are all females
from Washington and California.

VoL. XXXIT] LEECH: CENTRODERA SPURCA AND ALLIES 159

Figure 4. Centrodera dayi. Profile of an elytron, showing the suberect vesti-
ture.

apices not toothed at sutural angles. Outer antennal segments of male slightly
to obviously bristling (figs. 7, 8). Smaller species, males averaging 17 mm. in
length, females 18 mm. Later appearing species, June—September, commonest
in August; distribution either adjacent to the Cascade mountains from southern
Washington to the Klamath mountains and western foothills of the Sierra
Nevada of California, or along the coast of California only.................................... 2
2. Front tarsus notably short and broad (fig. 9). Hind tarsus comparatively short,
first segment very gradually widened from base to apex, appreciably broadened
apically; second segment noticeably triangular, flattened dorsally (fig. 10).
Male with short and appreciably cuneate elytra (fig. 11), antennae with seg-
ments 6 to 11 and apical half of 5 with exceedingly short, fine, almost decumbent
hairs, the general effect at < 25 fairly smooth (fig. 7). Elytra of female almost
straight-sided (fig. 11). Known from southern Washington to the foothills ad-
jJacent tothe Central Valley of California (figs: V4) 15)-22 C. dayi
— Front tarsus more elongate, narrower (fig. 9). Hind tarsus with first segment
long, parallel-sided, very slightly broadened apically; second segment elongate,
very gradually widening from base to apex, rounded on top (fig. 10). Male with
gradually narrowing, nearly parallel-sided elytra, outer antennal segments at
< 25 bristly (fig. 8). Female with elytra narrowed behind humeri then widened
again (fig.11). Known only from the coast of California (fig. 15)..C. autumnata

160 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Centrodera spurca (LeConte) .

Toxotus spurcus LeCon’s, 1857. Rept. ins. coll. survey, p. 63 (this is a preprint of
the following item); LeConrr, 1860. In: Reports expl. survey . . . Mississippi
._.. Pacific Ocean, vol. 12, part 3, p. 63; LAcoRDATRE, 1869. Gen. coleopt., vol. 8,
p. 439, footnote 1; LreConre, 1870. Ann. Mag. Nat. Hist., ser. 4, no. 35, vol. 6,
p. 402.

Pachyta spurca LeConte, LENG, 1890. Ent. Amer., vol. 6, part 5, pp. 97, 98; FALL,
1901. Occ. Pap. Calif. Acad. Sci., vol. 8, p. 148; Garnerr, 1918. Canad. Ent.,
vol. 50, part 6, p. 212.

Parapachyta spurca LeConte, Casry, 1913. Mem. Coleopt., vol. 4, p. 216; Harpy,
1926A. Rept. Provin. Mus. (1925), p. C 28, pl. IV, fig. 3; Harpy, 1926B. Ceram.
Vane. Isl., p. 5, pl. IV, fig. 3; Essie, 1926. Ins. West. N. Amer., p. 452; Moore,
1937. Oce. Pap. San Diego Soc. Nat. Hist., vol. 2, p. 88.

Pachyta (Parapachyta) spurca LeConte, Doane et al., 1936. For. Ins., pp. 176-177.

Centrodera spurca LeConte, R. Horpine, 1937. Nat’l. Mus. Canad., bull. 85, p. 11
(in part, but not the fig. [4] on pl. III, for which see C. autumnata) ; Harpy,
1942. Proc. Ent. Soc. Brit. Col., vol. 39, p. 10.

Evodinus spurcus J. Leconte, AuRiviriius, 1912. In: Coleopt. Catal., Pars. 39,
p. 188; Saaas, 1936. Ann. Zool. Soc. Zool.-Bot. Fenn. Vanamo, vol. 4, no. 1,
D: 73:

Evodinus (Centrodera) spurcus, SAALAS, 1936. Ann. Zool. Soe. Zool.-Bot. Fenn.
Vanamo, vol. 4, no. 1, p. 82.

Typocerus cervinus WALKER, 1866. In: Lord’s Nat. Vane. Isl. and Brit. Col., vol. 2,
1. Sart

Centrodera spurca was deseribed as from Steilacoom, Washington Terri-
tory; this is in what is now Pierce County, just southwest of Tacoma,
Washington. However, on page 23 of the same paper LeConte recorded it
from Oregon; this is explained by his statement on page 3: “. .. (and
Washington Territory, which is, for purposes of convenience, always in-
cluded when Oregon is referred to in these pages) ...’’ The type is a female,

Figure 5. Centrodera spurca. Fourth and fifth abdominal sternites of a female,
the fifth distinctly margined.

VoL. XXXIT] LEECH: CENTRODERA SPURCA AND ALLIES 161

and there is a female before me which was compared with it in 1949 by
P. J. Darlington, Jr.

The type of Typocerus cervinus Walker is in the British Museum (Nat.
Hist.), and the museum’s Accession Catalogue entry 64-18 shows “British
Columbia Coll. Boundary Commission Collector J. K. Lord Esq.”
LeConte examined it and synonymized it (1870, p. 402) with his Toxotus
spurcus. Frederico Lane was so kind as to compare a female of C. spurca
with Walker’s specimen, which has three labels on the pin: 1. Type,

Brit 64

etolum 18°’
mm. long, 8.5 mm. wide at the humeri; the antennae are approximately
21.25 mm. long, the elytra 19 mm., the fifth abdominal sternite is margined
and slightly emarginate apically, and each elytron has a small spur at the
sutural apex. These characters are all in agreement with those of a medium-
sized female of C. spurca, with which species it was synonymized by LeConte
in 1870 and by Leng in 1890.

It is puzzling that Leng, in his 1890 synopses of the Cerambycidae, put
C. spurca in Pachyta, since a specimen will trace correctly to Centrodera in
his generic key (p. 65), which was taken from LeConte and Horn’s 1883
“Classification,” it in turn having been taken from LeConte’s 1873 “Classifi-
cation.”

3. cervinus [in Walker’s handwriting]. The type is 24.25

Mate. Form elongate, broadest at humeri, elytra gradually narrowing
from base to truncate apex (fig. 11). Length 19 to 26 mm., average 22.2
mm.; width at humeri 5 to 7 mm., average 6.02 mm. Elytra pale yellowish-
brown, with luminous golden sheen from reflection of their shagreened
under surface seen through the almost transparent upper surface (and
resembling the glow seen in some well-worn micaceous schist stones) ;
usually with a rounded black or brownish antemedian dot near side, show-
ing dorsally on each elytron but actually on lower surface; thorax, head,
antennae, legs, and undersurface a little darker, pale reddish-brown, eyes
and tips of mandibles black; pubescence golden yellow. Head densely, ir-
recularly, moderately coarsely punctate dorsally, most coarsely between
eyes where, as on clypeus, surface may be somewhat rugulose; each pune-
ture on mandibles, labrum, elypeus, front, occiput, and undersurface of
head giving rise to a hair; mid-cranial suture an impressed line from base
of clypeus to declevity between eyes; width of vertex between eyes a little
greater than width of an eye. Antennae longer than body, usually sur-
passing apices of elytra by segments 10 and 11, apical (11th) segment
usually constricted at apical five-sevenths, giving the illusion of a twelfth
segment; segment eleven one and one-half times as long as scape and three-
fifths longer than first segment of hind tarsus; scape with moderately long
appressed hairs, and sparse seattered shorter hairs which stand out at an

162 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Figures 6-8. Ninth antennal segments of males of three species of Centrodera.
Fig. 6. C. spurca; the fine hairs are closely appressed, the sensory hairs stand out
clearly. Fig. 7. C. dayi; the fine hairs are not decumbent yet only partially ob-
scure the longer sensory hairs. Fig. 8. ©. autumnata,; the fine hairs are semierect
and almost as long as the sensory hairs, giving a bristly appearance.

VoL. XXXIT] LEECH: CENTRODERA SPURCA AND ALLIES 163

angle of about 45 degrees, segments 2 to 5 with progressively shorter ap-
pressed vestiture, segments 6 to 11 with a dense covering of short appressed
hairs lying parallel to the length of the segments, with occasional short
sensory hairs projecting at about 45 degrees (fig. 6), the effect at about 10
being as of satin, 7.e. holosericeus. Inflated disk of pronotum a little more
coarsely and closely punctured than is head between eyes, each puncture

C. SPURCA C.AUTUMNATA C. DAY!

Figure 9. Basal three segments of the front tarsi of Centrodera spurca, C. au-
tumnata and C. dayi to show relative lengths and widths; vestiture and punctation
omitted.

164 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

C.SPURCA C.AUTUMNATA _ C. DAYI

Figure 10. Basal three segments of the hind tarsi of Centrodera spurca, C.
autumnata and C. dayi; punctation and vestiture omitted.

VoL. XXXIT] LEECH: CENTRODERA SPURCA AND ALLIES 165

with a fine recurved hair, except at sides near base where some of the hairs
are long and projecting; lateral tubercles slightly antemedian. Elytral
width at humeri, to length, about as is 7.3 to 17.7; coarsely closely punctured
at base, except on humeri, and progressively less coarsely and more
shallowly punctured from base toward apex, each puncture giving rise to
a short fine recurved hair; elytral apices truncate, sutural angles usually
with a small blunt spine. Abdominal sternites clothed with fine appressed
golden hairs which do not obscure surface, and sparse longer and less de-
eumbent hairs, especially at apices of segments and along median line.
Fifth visible abdominal sternite distinctly thickened and margined apieally,
margin extending narrowly along sides nearly to base, apex usually broadly
emarginate and a little impressed at middle; pygidium emarginate apically.
Hind femur reaching to slightly beyond apical eighth of elytra, vestiture
of hind margin short, even; first segment of hind tarsus as long as segments
2 and 3 combined, segment 1 gradually widening from base to apex, seg-
ment 2 one and a half times as wide at apex as at base (fig. 10). Male
gemtalia: On the basis of the orientation accepted in the general discus-
sion of the male genitalia, earlier in this paper, the genital armature
hes on its left side in the abdomen. Aedeagus tubular, elongate, curved
(fig. 12A), basal third bilobed in dorsal view (fig. 12D); apical two-
thirds longitudinally divided into dorsal and ventral sclerites which
can gape apart but are actually joined by a phable membrane (figs.
12A, 12B, stippled area), ventral side longer than dorsal, more strongly
sclerotized and slightly hooked apically; sclerotized area of apical two-thirds
of dorsal side narrowing apically, with area along each side of median line
differentiated, 7.e. more strongly sclerotized and darkened, ending in slightly
spreading bifid prominenees (figs. 12A, 12B). Intromittent organ, when
everted, about as long as aedeagus, with two irregularly shaped armatures
(when the endophallus is retracted and folded within the aedeagus the pair
of armatures appear as in fig. 13, but when it is extruded and inflated to
to its full dimensions the two armatures are on opposite sides and separated
by a distance equal to the length of one of them), and a pair of small, nearly
quadrate rasp-surfaced areas. Tegmen elongate, slightly curved, encircling
aedeagus (as in fig. 17) at about mid-point, but independently moveable;
apical third thickened, flattened, developed into two separate parameres,
each with dense reddish-brown hairs on outer edge and apex (fig. 12C)
but having only about 50 thin paler hairs on its inner or ventral surface;
basal ends not coalesced, but fitting into hollow of basal third of ventral
surface of aedeagus, as in fig. 17A.

FEMALE. Length 20 to 30 mm., average 24.9, width at humeri 6 to 8.5
mm., average 7.3. Form stouter than in male, elytra almost paralled-sided
in basal five-sixths. Antennae shorter than in male, barely reaching to

166 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser.

apical sixth of elytra; 11th segment as long as scape, not constricted; an-
tennal vestiture as in male.

VARIATION. There are occasional specimens in which the elytra show
six or seven narrow paler lines; these are on the lower surface and show

Ne ec alac Sey ~ : e
Figure 11. Males (upper row) and females of Centrodera spurca, C. dayi and
C. autumnata, respectively.

VoL. XXXIT] LEECH: CENTRODERA SPURCA AND ALLIES 167

through to give the appearance of light vittae. Most examples seen have
been from the northeastern distribution of the species.

The considerable range in size has been noted above. Some specimens
are more reddish- than yellowish-brown, but this may result from differences
in methods of killing and preparing for mounting. The antemedian lateral

CENTRODERA SPURCA

Figure 12. Male genitalia of Centrodera spurca. A. The tubular aedeagus in
profile; it consists of two longitudinal sclerites, here shown separated by the in-
flated pliable connecting membrane (stippled); the dorsal sclerite is on the left
and ends in a (bifid) prominence. B. Dorsal view of the apical part of fig. A;
the dorsal sclerite with its bifid tip overlies the stippled connecting membrane.
C. Dorsal view of the flattened apical third of the tegmen (—parameres; for their
normal position in relation to the aedeagus see fig. 17A). D. Dorsal view of the
bilobed basal third of the aedeagus, i.e., the upper right part of fig. A.

168 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SEr.

dot on each elytron varies from a maximum diameter of 1 mm. to the tiniest
observable spot, and contrary to R. Hopping’s statement (1937, p. 11) is
completely lacking in about ten per cent of the beetles studied; it varies
also from black to a pale brown. R. Hopping’s reeord (1921, second page) of
a series of Pachyta spurca showing “much variation in maculation” surely
resulted from a lapse of memory as to the genus and species on the part
of F. E. Blaisdell, who recorded the minutes of the meeting.

The apex of the fifth abdominal sternite varies from rather evenly
rounded to broadly emarginate. It may be narrowly or weakly margined,
but is usually strongly so, especially in the female, in which the margin
tends to be broader and less regular than in the male (fig. 5). Of the 1061

CENTRODERA SPURCA C. AUTUMNATA Cc. DAY!

Figure 13. Armature of the intromittent organ of the male genitalia of Cen-
trodera spurca, C. autumnata and C. dayi. For C. spurca the paired armatures are
shown as they lie partially superimposed when the organ is retracted within the
aedeagus; when the organ is everted and inflated during copulation they are on
opposite sides of it and separated by a distance about equal to the length of one of
them.

specimens of C. spurca examined only four, all females, do not show an ap-
preciable margining; they are from Pullman and Rock Island, Washington,
and Hat Creek and Meadow Valley, California. All are easily separated
from the species with non-margined fifth sternite by the short recurved
elytral hairs.

DistRIBUTION. There are not many published records for C. spurca and
its synonym T'ypocerus cervinus Walker; a few of those for Oregon and
California may prove to have been based on the new species described in
this paper, but I believe over 90 per cent truly refer to C. spurca. All
records known to me are listed here in abbreviated form; most of the full
references may be found in the Leng Catalogue and its supplements, though
many are in the terminal bibliography of the present article. I have not

VoL. XXXII] LEECH: CENTRODERA SPURCA AND ALLIES 169

used any of these records in my maps and tables unless I have seen the
actual specimens.

LeConte, 1857, p. 23, “Or.”; p. 63, Steilacoom, Washineton Territory
(reference is to the type specimen; see explanation in first paragraph
following the synonymy of C. spurca).

LeConte, 1860—a repetition of the above.

Walker, 1866, p. 332. British Columbia.

Lacordaire, 1869, p. 489. Orégon.

LeConte, 1869, p. 371. Vancouver’s Island [and/or] British Columbia.

semminger and Harold, 1872, p. 2859. “California. Vancouv. Ins.”

Leng, 1890, p. 98. ‘“Cal., Vane., Nev.”

Fall, 1901, p. 148. Echo Mountain, Los Angeles County, California.

Currie, 1904, p. 28. Kaslo, British Columbia.

Harvey, 1907, p. 4. Victoria, British Columbia.

Wright and Coolidge, 1908, p. 68. Towle, Placer County, California, in
June and July.

Aurivillius, 1912, p. 188. “Californien, Nevada, Vancouver-insel.”’

Casey, 1913, p. 217. “. . . species of the true Pacific Coast fauna.”
Woodworth, 1913, p. 228. California (as Pachyta spurcata [sic!] Le-
Conte).

Gibson, 1917, p. 150. ‘““Swanlake, B. C.”” (This is Swan Lake, a few miles
north of Vernon, British Columbia. )

rarnett, 1918, p. 212. “. .. taken by Fall at Echo Mt., Southern Cali-
fornia. Found by Van Dyke at Santa Moniea.”’

Leng, 1920, p. 271. “Nev.—Vane. So. Cal.”

Baumberger, 1921?. St. Helena, Napa County, California. (Date of pub-
leation uncertain; page not numbered. )

Hardy, 1926A, p. C28. Victoria, Sidney, Shawnigan, Dunean, British
Columbia. British Columbia to California.

Hardy, 1926B, p. 5. (As above.)

Essig, 1926, p. 452. California, Nevada, Oregon, Washington, and Brit-
ish Columbia.

Harvey, 1926, p. 5. (A republication of his 1907 list.)

Canova, 1936, p. 129. Corvallis, Alsea, Junction City, Minam National
Forest 20 miles NW. of Bly; all in Oregon.

Doane et al., 1936, pp. 176-177. “... throughout the Pacific Coast.”

Saalas, 1936, p. 73. ““Nordamerika.”’

Hopping, 1937, p. 11. British Columbia, Idaho and California.
probably oceurs in Oregon and Washington. Nevada is mentioned
in the literature.”

Moore, 1937, p. 88. “. . . San Diego in April and July . . . Warner’s
Spring in July.” (California. )

“ce

170 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Hatch, 1939, p. 29. Eastern and western Washington.

Hardy, 1942, p. 10. 3.5 miles N. of Victoria, British Columbia.

L. E. Ricksecker, then of Santa Rosa, California, offered C. spurca for
sale in his “Price list of Coleoptera of the Pacifie Coast, No. 15. January 1,
1897,” and probably in earlier lists, though I have not seen them. If he
obtained his specimens at his ranch “Sylvania,” near what is now Camp
Meeker, Sonoma County, he probably had representatives of both true C.
spurca and one of the species here described as new.

As known to me from actual specimens, C. spurca occurs across southern
British Columbia except for the mainland west of the coast mountains, from
southern Vancouver Island (Ucluelet-Nanaimo-—Victoria) to the East Koo-
tenays (Creston), with the most northerly record at Salmon Arm, lat.
50° 41’ N., long. 119° 18’ W.; thence through Washington, northern and
mid-western Idaho and adjacent Oregon, Oregon just east of the coast
range to San Diego via the coast of California, and to the Greenhorn moun-
tains of Kern County via the Cascades and Sierra Nevada, with a few
records for western Nevada. The species must occur in northwestern Mon-
tana; a single specimen labeled Salt Lake City, Utah, has been seen. Ex-
cept that there are no records of C. spurca from the British Columbia
mainland west of the coast mountains, its distribution is remarkably like
that of another cerambyeid, Ergates s. spiculatus (LeConte); see the map,
fig. 8, in Linsley’s 1962 paper.

In California, C. spurca is known from the following counties, listed
in north-south sequence, first for the coast and adjacent mountain areas
west of the Central Valley: Del Norte, Humboldt, Trinity, Mendocino,
Sonoma, Lake, Marin, Contra Costa, Alameda, San Mateo, Santa Clara,
Santa Cruz, Monterey, San Luis Obispo, Ventura, Los Angeles, San Ber-
nardino, Riverside, San Diego. For the Cascades and Sierra Nevada, Siski-
vou, Modoe, Shasta, Lassen, Plumas, Tehama, Butte, Nevada, Placer,
Eldorado, Alpine, Calaveras, Tuolumne, Mariposa, Mono, Madera, Fresno,
Tulare, Inyo, and Kern. As is obvious from the map (fig. 14) the species
should be looked for in the Sierra Juarez and Sierra San Pedro MaArtir of
Baja California, México.

In the coastal region of California C. spurca has been taken from about
700 feet above sea level at Carmel, Monterey County, to 5800 feet near the
top of Junipero Serra peak, the highest point in the Santa Lucia moun-
tains of the same county, and some 40 miles southeast of Monterey. In
Mill Valley, Marin County, it occurs down to an elevation of not more than
100 feet above sea level, while in southern California it has been taken at
Lake Arrowhead, San Bernardino County, at 5100 feet. In the southern
Cascades there are records from about 2280 feet at Dunsmuir, Siskiyou
County, to 3200 feet at Hat Creek, Shasta County, 4450 feet at Alturas,

VoL. XXXIT] LEECH: CENTRODERA SPURCA AND ALLIES 171

Modoe County, and about 6700 feet at Summit Lake in Lassen Volcanic
National Park. In the Sierra Nevada the lowest station seems to be Quincy,
Plumas County at 3400 feet, and Twain Harte, Tuolumne County at 3600;

@ CENTRODERA DAYI
OC. SPURCA
© c. DAYI & C. SPURCA

most

Figure 14. The distributions of Centrodera spurca and C. dayi, plotted from the
locality labels of specimens seen during this study; no records from the literature
have been mapped unless represented by specimens. Note that the two species
have been taken at the same places in sixteen cases.

172 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

thence to 7000 feet at Huntington Lake, Fresno County, and to between
8000 and 11,000 feet near Glacier Lodge, Big Pine Creek, Inyo County, on
the east slope of the Sierra Nevada.

Centrodera spurca has not been reported from the Rocky Mountains to
my knowledge; it is not in Mank’s 1934 Glacier Park list. Neither is it in
the Utah list (Knowlton and Wood, 1950), but there is a female specimen
in the Ohio State University collection labeled “Salt Lake City, Utah,
X.16.1952. R. E. Rodock. R. E. Rodock Collection.” October is an amazingly
late date for this species.

O CENTRODERA AUTUMNATA <> G= °-
@ Cc. DAY! IN CALIFORNIA

DRAFT 1955 Waser
us 4

Figure 15. The known distribution of Centrodera autumnata, and the California
records for C. dayi.

VoL. XXXIT] LEECH: CENTRODERA SPURCA AND ALLIES 173

Centrodera autumnata Leech, new species.

Centrodera spurca LeConte, R. Hoppine, 1937. Nat’l. Mus. Canada, bull. no. 85,

p. 11 (in part, including the fig. on pl. III).

A species resembling C. spurca but averaging smaller, with suberect
elytral pubescence, bristly outer antennal segments in the male, non-
margined fifth abdominal sternite; appears later in the season, and is re-
stricted to coastal California.

Ho.Lotyre, male, Minh VALLEY, Marin County, Cauirornia, 16.VIII.49,
at light (Hugh B. Leech). In the California Academy of Sciences (Ento-
mology ).

Length 18 mm., width at humeri 5.2 mm. Form elongate, elytra very
gradually tapering from base toward apex. Color yellowish-brown above
and below; tips of mandibles, eyes, and a small oval sublateral spot show-
ing through from lower surface of each elytron just before middle, black;
elytra pale yvellowish-brown, darkest at base, pronotum and antennae a
little darker than elytra, head reddish-brown, tarsal claws and mandibular
attachments rufous. Head dull between eyes, otherwise shining; punctures
on elypeus irregular in sizes and distribution, those on vertex, occipital
area and submental region more regular and denser, each puncture giving
rise to a hair; width of vertex between eyes slightly narrower than width
of an eye, mid-cranial suture apparent but its course very little depressed.
Antennae longer than elytra by leneth of last two segments, 11th segment
one and a half times as long as scape and a little longer than first segment
of hind tarsus, scape reaching to beyond hind margin of eye; scape, seg-
ments 2 to 4 and basal half of 5 shining, finely punctured, clothed with
closely appressed hairs and widely spaced suberect fine sensory hairs; apical
half of segment 5 and segments 6 to 11 in their entirety dull, the vestiture
dual, consisting of fine decumbent hairs, hard to see, and a dense covering
of suberect stiff hairs like the pile of a rug, two-thirds as long as the sensory
hairs and projecting at the same angle; the outer segments thus have a
bristly appearance as in fig. 8. Pronotum moderately coarsely punctate,
punctures sparser on preapical and prebasal transverse grooves, coarsest at
sides of disk where some punctures are contiguous, integument there sub-
rugose; vestiture sparse, of fine erect hairs, especially at sides basally, and
of shorter decumbent hairs whieh are less obvious; lateral tubercles blunt,
slightly antemedian, prothoracie width at (and including) tubercles 95 per
cent of width at base and about one-third greater than width at apex.
Elytra widest at humeri, gradually tapering to apices, fastest in apical
sixth; width at humeri is to length as 11 is to 26; punctures coarse and
dense basally except on humeri, discal punctures separated from one an-
other by about one-half their own widths, becoming progressively smaller
and shallower toward elytral apices, densest just behind humeri; apices

174 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

vaguely truncate, without tooth at sutural angle. The hairs arising from
elytral punctures short, standing out at an angle, especially in discal region;
elytral surface more opaque than in C. spurca, golden subsurface reflections
less evident. Front, middle, and hind tarsi progressively narrower and more
elongate, first three segments of front tarsus seven-tenths as long as front
tibia; first segment of hind tarsus long, narrow, parallel-sided, very slightly
broadened apically (fig. 10), one-third as long as hind tibia; second seg-
ment of hind tarsus rounded on top, almost half as long as first. Hind
femur reaching slightly bevond apical seventh of elytra, with a series of
sparse long hairs along hind margin, from tip of trochanter nearly to apex,
hairs from half to three-quarters as long as femur is wide and inclined some-
what toward abdomen. Male genitalia (see fig. 16, and the general dis-

C. AUTUMNATA

Figure 16. Male genitalia of Centrodera autumnata. A. Profile of the tubular
aedeagus, the upper (on the left side) and lower sclerites here shown separated
by the pliable connecting membrane, stippled. B. Dorsal view of the apical part of
the aedeagus. C. Dorsal view of the parameres.

VoL. XXXIT) LEECH: CENTRODERA SPURCA AND ALLIES 175

cussion of the organs in the early part of this paper). Essentially as in
C. spurca, but differing as follows. Median line of apical two-thirds of
dorsal side of aedeagus much less strongly or not at all differentiated in
color, ending in a less prominent knob which is not bifid apically, but with
transverse rugae on the depressed neck just before apical bulb. “Floor”
of median area of tegmen deeper than in C. spurca, hairs of parameres
shorter and paler in color, inner face of paramere with only about 25 hairs.
Armatures of intromittent organ as in fig. 13; it is possible that their ap-
parent difference in shape from those of C. spurca may be due in part to
some slight difference in the angle of viewing.

ALLOTYPE, female, Mill Valley, Marin County, California, 13.VIII.1949,
at light (H. B. Leech) [CAS]. Length 17 mm., width at humeri 5 mm.
Similar to the male but more robust (as in fig. 11), elytra not tapering grad-
ually from base to apex but a little wider at apical quarter than at base,
apices rounded. Antennae shorter than body, reaching to apical quarter
of elytra (equivalent to third abdominal sternite), segment 11 seven-tenths
as long as scape; hairs on segments 6 to 11 and on apieal half of 5 appressed,
except for the scattered suberect sensory hairs. Fifth abdominal sternite
lightly maregined at sides near base; intereoxal process of first abdominal
sternite a little blunter apically than in holotype.

PaRATYPES, all from California. Marin County: 55¢'¢, 72 2 topo-
types, taken by members of my family or myself at our former home at 427
Rose Avenue, Mill Valley (elevation 600 feet), nearly all attracted to
lights and collected on the windows of the house, 1948-1956; 1% in May,
50 ¢ in July, 400 o and 52 2 in August, 11d’ v and 22 ? in September
[CAS]. Also the following from other parts of Mill Valley; 1¢ 23.V1.1925,
Pousecovn1925, Io 8. VIIT1925, fo 3.1X.1924 (Ce. P. Van Duzee)
aoe tos IVI 1954, 1, 22 9 17. VITL1950 (H. S. Ross) [CAS];
eo 7.VIL1959, 19 16.VIT.1959, 10, 12 VIIT.1959 (J. Sedlacek) [J. Sed-
lacek]; 19 23.1X.1950 (F. X. Williams) [CAS]; 19 3.1X.1950 (D. Kelley)
(exsieeeico Go i VILNI953, So's, 19 LOVEITII953, to 23:VIIE1993,
Sem ov LIN 19538, 20 og 1.1X%.1953, 26 o 9.1X.1953, ho 9.1X.1954, 1
essa aco o 16.VITI1955, 16° 23.VII1.1955 (all by A. LL. Mathis;
those for 1955 labeled “Light Trap Collecting”) [UCD]; 1¢7.V1II.1961,
Bootjack Camp, south slope of Mt. Tamalpais (H. B. Leech) [CAS]. Also
the following from other places in Marin County; Novato, 1¢, 12 16.VIII.
1954, at light (H. B. Leech) [CAS. Observed in copulation on August 17];
1¢ 6.1X.1948, 10 18.VIII.1954 (KE. L. Kessel) [CAS]; Umdelelannyoni,
Novato, 20 & 5.VII.1952, 2¢ f 26.VII.1952 (E.L. Kessel) [CAS]; Laguni-
tas, 1¢ 30.VII.1921 (F. E. Blaisdell) [CAS]; Lansdale, 1¢ 30.VIII.1914
(R. Hopping collection. The pin carries a pink label marked “PI.” and the
specimen is the one used by George R. Hopping in making his drawing tor

176 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser.

fig. 4, pl. III of R. Hopping’s 1937 paper on the Lepturini) [CAS]; Toll
House, 206, 22 2 5.VIII.1949 (Alice Edwards, J. G. Edwards) [JGE];
Woodaere, 1o' 27.V1II.1955, 36 oh 3.VIII.1955, 10 16.VIII.1955 (HB i:
Mathis) [UCD]; Fairfax, 1¢ 25.VIII.1953 (H. L. Mathis) [UCD]; Straw-
berry Point, 1¢ 6.VIII.1953 (H. L. Mathis) [UCD]; San Anselmo, 1¢
24.V11.1940 [CAS]; Ross, 1% 3.VIII.1955, 1¢ 16.VIII.1955 (H. L. Mathis)
[UCD]. Humsotpt County: Dyerville, 1¢ 18.VIII.1951 (R. W. Dawson)
[RWD]. Sonoma County: Rio Nido, 1¢ 31.VII.1946 (D. Giuliani) [DG];
Guerneville, 1¢ 11.VIII.1948 (D. Giuliani) [DG]; Stewarts Pt., 1¢ VI.
1942, 1¢ VII.43 [CIS]. San Matro Country: La Honda, 1¢° 14.VIII.1925
(U. S. Grant IV) [SDNHM]. Santa Ciara County: Cupertino, 1d
20.V1.1939 (K. S. Hagen) [CIS]; Los Gatos, 1¢° 16.VIII.1954 (Sr. Mary
Baptista) [USF]; Alum Rock Park, 1¢ 17.V11.1949 (S. R. Piazza) [SRP];
San Jose, 1¢ 17.V11.1931, 1¢ 22.VI1.1931 [CIS]. Santa Cruz County:
Big Basin Redwood State Park, 1¢ 1.1X.1953 (P. H. Arnaud, Jr.) [CAS];
Big Basin, 1¢ 10.VIII.1933, 2¢ f& 12.VIII.1933 (W. H. Lange) [UCD],
1¢ 17.V11.1940 (B. Brookman) [UI]; Boulder Creek, 3¢'¢ 6.VIII.1941
(J. W. Tilden) [SJSC], 10 20.V1II.1949 (W. E. Hazeltine) [CAS], 19
21.VII1.1935 (B. E. White) [CAS]; Brookdale Lodge, 1¢ 18.VIII.1940
(K. Frick) [CIS]; Ben Lomond, 400, 22 ? 30.VII.1959 (D. Rentz)
[DR], 1¢ 3.VIII.1959 (C. Wemmer) [CAS], 4¢'¢ 31.VII1L.1962 (C. D.
MacNeill) [CAS], 1 “1918” (Mary Knowles; R. Hopping Collection)
[CAS]; Mount Hermon, 19 16.VIII.1949 (W. E. Hazeltine) [CAS]; Santa
Cruz hills south of Felton, 3¢ & 27.VI1.1961 (G. Follin) [CAS]; Santa
Cruz; 1¢ VUIL37 [CIS],. 16% 2.VI1941, 1o 7.VIL1941, top 10y ieee
20 3h 12.V11.1941, 18 17.V11.1941 (J. W. Tilden) [SJSC]; Highland Dis-
trict, 29 9 .2.1X.1956 (S. M. Fidel) [UCD]; “Santa Cruz(Co72iioeyaiie
1916, 20h VII.1917 (E. R. Leach) [CAS]. Montery County: Junipero
Serra Peak, Santa Lucia Mountains, on peak ca. 5800 ft. elev., at light,
49 8.VIII.1956 (H. B. Leech) [CAS]. Santa BarBara County: Car-
pinteria, 19 1.1X.1935 (B. E. White) [CAS]. San Brernarpino County:
Barton Flats, 2¢ ¢% 20.VIII.1936, 1¢ 21.VIII.19386 [CAS]. Riversipe
County: Idyllwild, 1¢ VIII.1946 (C. Harnage) [UCD]. San Dieco
County: Newton, 1¢ 14.VII.1949 (D. J. & J. N. Knull) [JNK]; Laguna,
1¢ 22.VI1.19384 (C. C. Searl) [SDNHM].

Additional specimens studied but not designated as paratypes are the
following: Marin County; Mill Valley, 1¢ 28.V1.1959 “at light, first of
season” (H. B. Leech), 3¢' ¢ 18.VIII.1950 (H. B. Leech) [CAS]; “Redw.
C. Ft. Hills” [= Redwood Canyon foothills near Muir Woods?], 20 o
18.VIII.1946, 1¢ 21.VII1.1946 (D. Giuliani) [DG]. Santa Cruz County;
Ben Lomond, 19 3.V1.1946, 1¢ 8.VI.1946 (W. Lee) [CAS]. Santa Bar-
BARA COUNTY; Santa Barbara, 1? (F. E. Winters) [CAS].

VoL. XXXIT] LEECH: CENTRODERA SPURCA AND ALLIES 177

In addition to those being returned to their owners, paratypes will be
deposited in the following institutions: the U. S. National Museum, Wash-
ington, D. C.; the Canadian National Collection, Ottawa, Ontario; the Brit-
ish Museum (Nat. Hist.), London; the University of British Columbia,
Vancouver; the University of Washington, Seattle; Oregon State University,
Corvallis.

VARIATION. The paratype males vary in size from a length of 14 mm.
and a width at the humeri of + mm., to a length of 21 mm. and width of
6 mm., with averages of 18.2 mm. and 5 mm. respectively; paratype females
from a length of 16 mm. and width of 4.5 mm. to 22.5 mm. and 6.75 mm.,
with averages of 18.8 mm. and 5.5 mm. There is an appreciable variation
in color, from pale yellowish-brown to distinctly reddish-brown. It is hard
to tell how much of the variation is natural because all living examples I
have seen have been yellowish-brown; I suspect the darker color of some
dried specimens results in part from different methods of killing and pre-
serving.

The elypeus may be somewhat flattened, smooth, and impunctate in front
as is the labrum, or it may be punctate right up to its front edge. The apex
of the seutellum varies from broadly rounded to pointed, and is occasionally
slightly irregular but not emarginate. The elytral apices vary from dis-
tinetly truncate to rounded, or even incised. Only a single specimen has
been seen in which the semi-erect hairs of the basal half of the elytra are
so abraded as to give difficulty in the key. The much finer erect hairs of the
pronotum are often matted down or worn off, except behind the lateral
tubercles; these latter vary from softly rounded protuberances to almost
spinous processes. The intercoxal process of the first abdominal sternite of
some females is as sharply pointed as in the male, in others it is blunter
or even rounded apically. None of these variations is correlated with dis-
tribution.

Remarks. Males of C. autuwmnata resemble those of C. spurca fairly
closely in form but average smaller and are commonest later in the season;
both have antennae surpassing the elytral apices by the length of the last
two segments. Centrodera autumnata is easily recognized by the suberect
elytral pubescence, non-margined fifth abdominal sternite, long narrow hind
tarsi, fuzzy-appearing outer antennal segments (see figs. 8, 10). These
same characters, except the last, will also separate females of C. autumnata
from those of C. spurca.

Males of C. autwmnata may be separated from those of C. dayi by their
more elongate and less triangular elytra, longer antennae with the eleventh
segment longer than the first segment of the hind tarsus, and their narrower
and more elongate tarsi, especially the front tarsi (figs. 9, 11, 18). Females

178 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

of C. autumnata are recognized by their narrower and more elongate hind
tarsi, with the second segment rounded rather than flattened on top.

DisTRIBUTION. Centrodera autumnata is known only from the coastal
area of California, from Dyerville, Humboldt County to San Diego County
(fig. 15). It occurs from virtually sea level at Mill Valley, Marin County,
to 5800 feet in the Santa Lucia Mountains of Monterey County, but is un-
known from the Monterey coast where C. spurca is not rare. Although its
distribution is entirely within the southwestern edge of that of C. spurca
(figs. 14, 15) and they have nine localities from Sonoma County to River-
side County in common*, they must have very different ecological require-
ments. For instance, at Mill Valley both species are remarkably common,
especially C. autumnata; yet just across San Francisco Bay in Berkeley and
the Oakland hills, where C. spurca is fairly common, there are no records for
C. autumnata. A little south of the bay, however, it occurs further east than
the Oakland hills (fig. 15).

The gap between the Humboldt County and Sonoma County localities
is almost certainly because there has been little collecting at light in the
late summer; but similar gaps south of Monterey County are likely to indi-
cate extensive areas of unsuitable habitats. The species should be looked
for in the northern mountains of Baja California, México. The finding of
C. dayi at Rumsey, Yolo County, some 50 miles due east of the Sonoma
County reeords for C. autumnata, suggests that their distributions may
over lap in the Eel River country.

Centrodera dayi Leech, new species.
? Centrodera hirsuta R. Hoprtne (Ms., nomen nudum), 1939. In Hateh, Prelim.
list Coleopt. Wash., p. 29.

A species resembling C. spurca but which has shorter antennae in the
male, shorter front tarsi, suberect elytral vestiture, shorter elytra which are
distinctly cuneate in most males, and coarser elytral punctation especially
in the apieal half. Known from south eentral Washineton to the foothills
of the Sierra Nevada in central California.

Ho.Lotypr, male, Scorr River at KLAMATH RIVER, SISKIYOU CoUNTY,
CALIFORNIA, 1.VIII.1949, at light (W. C. Day). In the California Academy
of Sciences (Entomology ).

Length 19.2 mm., width at humeri 5.7 mm. Form moderately elongate
(as in fig. 11), elytra tapering regularly from humeri to near apices. Color
brown, eyes, tips of mandibles and antemedian lateral spot (showing
through from underside of each elytron), black; head, thorax, legs, basal
four segments of antennae and basal third of elytra reddish-brown, ab-

3. Centrodera autumnata flies later in the season, when fewer collectors are in the field, and the two may
actually have more localities in common than museum records indicate.

VoL. XXXIT] LEECH: CENTRODERA SPURCA AND ALLIES 179

domen, outer antennal segments and apical two-thirds of elytra yellowish-
brown. Head shining; punctures of irregular sizes dorsally, less coarse and
more evenly distributed on submental area, each giving rise to a hair;
width between eyes slightly less than width of an eye; mid-cranial suture
clearly, evenly impressed. Antennae reaching to just beyond elytral apices,
11th segment a little shorter than scape but equal in length to first segment
of hind tarsus, scape reaching to beyond hind margin of eye; scape, seg-
ments 2 to 4 and basal two-thirds of 5 shining, finely punctured, clothed
with appressed hairs, and some widely spaced suberect fine sensory hairs;
apical third of segment 5 and segments 6 to 11 dull, with a vestiture of
nearly decumbent fine hairs from which the sparse suberect sensory hairs
stand out clearly as in fig. 7, the whole having a slightly velvety appearance;
11th segment tapering from apical three-fifths to tip. Pronotum strongly
inflated between preapical and prebasal transverse impressions, closely and

Cc. DAY!

Figure 17. Male genitalia of Centrodera dayi. A. Profile of the tubular aedea-
gus, with the tegmen in its normal position over it, the parameres covering the
apical portion of the dorsal sclerite. B. Apical part of the aedeagus, dorsal view.
C. Parameres, dorsal view.

180 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

rather evenly punctate, punctures about comparable in size to those at
middle of elytra; vestiture of fine mostly appressed hairs, one from each
puncture; blunted lateral tubercles shghtly antemedian, width of prothorax
across tubereles a trifle wider than at base, apex five-sevenths width of base.
Apex of seutellum rounded. Elytra widest at humeri, regularly tapering
to apical seven-ninths where they are a little less than nine-fourteenths
width at base, then more rapidly to apices; width at humeri about six-
fourteenths the elytral length; punctures coarse basally, fairly regularly in
oblique series of four between traces of costae following courses of the
longitudinal tracheae, gradually smaller and shallower from base to apex,
those near apex comparable to discal pronotal punctures but shallower ;
apices subtruneate, without tooth at sutural angle. Each elytral puncture
giving rise to a hair, of which those on diseal area stand out at an angle,
suberect, those at sides and toward apices more nearly decumbent. Front
and middle tarsi much broader than hind tarsi (as in figs. 9, 10), front pair
notably broad; first three segments of front tarsus six-tenths as long as
front tibia; first seement of hind tarsus one-third length of hind tibia, very
gradually widening from base to apex, appreciably broadened apically, see-
ond segment flattened dorsally, a little less than two-fifths length of first.
Hind femur with a row of sparse hairs along hind margin, from tip of
trochanter nearly to apex, hairs less than half as long as femur is wide.
Male genitalva as in figs. 18, 17; for general description see C. spurca.

ALLOTYPE, female, same data as for holotype but collected on August 7;
in California Academy of Sciences (Entomology). Length 17.5 mm., width
5.7 mm. Generally similar to male but more robust (as in fig. 11). Head
and thorax tinged with piceous, elytra more uniformly yellowish-brown,
more parallel-sided, less tapering than in male. Antennae reaching to just
beyond apical two-thirds of elytra (7.e. not quite to apex of second ab-
dominal sternite); segment 11 is to scape as 2 is to 2.75; vestiture of see-
ments 6 to 11 and apical half of 5 appressed, except for scattered suberect
sensory hairs. Fifth abdominal sternite exceedingly narrowly margined
apieally; elytral apices rounded.

ParATypPes. WASHINGTON: Yaxima County; Yakima, 1¢ 10.VIII.
1931 (A. R. Rolfs. R. Hopping collection) [CAS], 1? 10.VII.1936, eleva-
tion 1025 feet (R. W. Every) [CIS]; Selah, 19 (Rufus Kiser) [UW].
WaLLA WaLua County; Walla Walla, 1¢ 30.VI1I.1952 (M. C. Lane)
JNK], 1¢ 30.VII.1947 (W. C. Cook) [JS]; Kooskooskie, 16° 1.VIII.1932,
1 21.VIII.19382 (M. C. Lane) [USNM]. OREGON: Cotumsi1a County;
St. Helens, 1¢ 21.VII.1936, at light (K. Gray, J. Schuh) [OSU]. Hoop
River County; Mid Col. Expt. Sta., Hood River, 2¢' ¢ 30.VI1.1957, 1¢
3.VITI.1957, 1 gf 23.VIII.1957 (Clive D. Jorgensen) [OSU], 12 3.VII1.1957
(Clive D. Jorgensen) [RBH]; Hood River, 12 18.VII1.1954, at light (Paul

VoL. XXXIT] LEECH: CENTRODERA SPURCA AND ALLIES 181

O. Ritcher) [AHH]. SHermMan County; Pine Grove Dist., Maupin, 10
May-June °o2 [UW]. Yamuitit County; McMinnville, 1¢ 24.VI1I.1944,
19 1.VIII.1949, 19 4.VIIT.1952 (kK. M. & D. M. Fender) [UW]. Marion
County; Salem, 1¢', 19 31.V11.1959, 3¢.¢, 39 9 31.VIII.1959, Blk, Lt.
Trap (Harold Foster) [OSDA]. Benron County; Corvallis, 1 @ 7.VII.1931
(N. P. Larson), 19 8.VIII.1952 (Paul O. Ritcher) [OSU], 19 16.VIII.1941
(K. M. & D. M. Fender) [UW], 12 8.VIII.1925 [USNM, ex Brooklyn
Museum Collection]; Monroe, 1¢ 6.VII.1931 (N. P. Larson) [OSU], 1¢
23.V1I.1931, in moth trap (N. P. Larson) [OSU], 1¢ 30.VII.1931 (Joe
Schuh) [JS]. Lane County; Goshen, 1¢ VIII.1941 (R. Fauts [sic/])
[BM]. Dovueuas County; Sutherlin, 12 30.VI1.1944 (Elwood Mabry)
[GHN]. Jackson County; Medford, 1? 3.VIII.1946 (C. Fitch) [UCD],
5o4,32 2 11.VIIT.1944, 250,19 12.VII1.1944,1¢,39 2 20.VIII.1944,
32 9 29.VIII.1944, all in light trap (C. Fitch) [UCD]; Talent, 1¢, 1¢
24. VII.1938, In ec. moth bait pan (L.G. Gentner) [LGG, UCD], 19 15.VITI.
1940 (L. G. Gentner) [UCD]; Green Springs, 1¢', 1? 27.VIII.1961 (J. S.
Buckett) [UCD]. CALIFORNIA: same data as holotype, 206, 19
[CAS], same data but August 10, 2¢¢% [CAS]. Siskrvou County; Scott
fever, 19°10:V 111-1949, collected at light (W. C. Day) [CAS]... Yoto
County; Rumsey, 19 5.VIII.1955 (EH. A. Kurtz) [UCD]. SHasta County;
Hat Creek, 19 26.VII.1951, 19 25.VII.1952 (G. F. Pronin) [CAS], 1¢
Sev t942, Bhicht at licht [CIS], 16, 19 11.VIIL1956, 2.9 9 12-VIIT.
Hse eee 17. VIM1956, 19 18. VIIL1956, 1° 19. VIIL.1956 (H. Ruekes,. Jr:)
[CIS], 19 28.VII.1957 [CIS]. PLumas County; 4 miles W. of Quincy,
19 26.V1.1949 (W. R. Schreader) [UCD], 192 16.VII.1949 (F. Morishita)
[CIS]; Johnsville, 1¢ 30.VII.1960, 304,192 9.VIII.1961 (J. S. Buckett)
[UCD]. Extporapo County; Georgetown |erroneously stated to be Placer
County on the label] 19 29.VIII.1948 [DG]. TuoLUMNE County; Twain
Harte, 19 9.VIIL1958 (D. C. Rentz) [DR], 1¢ 20.VITI.1960 (M. Lund-
gren) [ML]; near Groveland, 1¢ 27.VII.1954 [CIS].

The following additional specimens were studied, but are too damaged
to be made paratypes: 1 topotypic male, 10.VIII.1949 [CAS]; Mid Col.
Expt. Sta., Hood River, Oregon, 1¢ 21.VIII.1957; Goshen, Lane County,
Oregon, 1¢ VIII.1941 (R. Fauts [sic!] [BM]; Medford, Oregon, 12?
11. VIII.1944, Light trap (C. Fitch) [UCD]; Minam N. F. [Oregon],
1¢@ 95.VIII.1914, (Or. Ex. Sta. No. 1458) [OSU]; Placerville, Eldorado
County, California, 19 29.VIII.1948 [DG].

In addition to those returned to their owners, paratypes will be de-
posited in the Canadian National Collection, Ottawa, and the British
Museum (Natural History), London.

VARIATION. Paratype males vary in length from 14.5 to 19.4 mm., and
in width at the humeri from 4.00 to 5.6 mm., with averages of 17.3 and 5.2

182 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser.

Cc. Cc. Cc.
SPURCA AUTUMNATA DAY |

Figure 18. Last three antennal segments of males and females of Centrodera
spurca, C. autwmnata and C. dayi; vestiture omitted.

VoL, XXXII] LEECH: CENTRODERA SPURCA AND ALLIES 183

mm. respectively; paratype females vary in length from 13.4 to 20.5 mm.,
and in width from 4.7 to 6.5 mm., with averages of 18.1 and 5.5 mm. re-
spectively. The elytral color varies from a pale yellowish-brown to a de-
cidedly reddish-brown. The apex of the secutellum is rounded in all
specimens seen, though it varies a little in width.

The outer antennal segments of the females vary in length more than
in related species. Most specimens have segments 10 and 11 as in fig. 18,
but 11 may be shorter or longer than 10, and in some eases they are long
enough to be confused with the same segments of the male. The fifth ab-
dominal sternite varies from very narrowly and obscurely margined
apically to completely unmargined, in the female; it is unmargined in the
male.

ReMARKS. In a mixed series of the three species, males of C. dayi can
usually be picked out because of their short and somewhat triangular elytra;
females resemble those of C. autumnata but are more heavy shouldered
and straight sided (fig. 11). The sexes are harder to separate in C. dayi
than in the other two species; males have more strongly tapering elytra,
longer antennae, and less smoothly vestitured outer antennal segments;
antennal segments 10 and 11 are usually appreciably longer in the male.

Centrodera dayi may be separated from C. spurca by its short body
form (fig. 11), suberect elytral pubescence, much coarser punctation in the
apical half of the elytra, short broad foretarsal segments (fig. 9), shorter
antennae, and the bristling vestiture of the outer antennal segments in the
male. In males of C. dayi antennal segment 11 is barely or not as long as
the seape; in C. spurca and C. autumnata it is as lone as or longer than the
scape and segment two combined.

The male of C. dayi is distinguished from that of C. autumnata by its
more strongly tapered elytra (fig. 11), much broader fore tarsi (fig. 9), and
short antennae which barely reach beyond the elytral apices. Females differ
from those of C. autumnata by their stouter build, straight sided elytra
(distinctly narrowed behind the humeri then widened again in C. autwm-
nata), and shorter and broader fore tarsi (figs. 9, 11). Both sexes of C.
day? have shorter hairs along the hind margin of the hind femur, starting
at the tip of the trochanter, than does C. autumnata (specimens must be
clean to show this properly), and a more evenly inflated pronotal disk.

DIstTRIBUTION. In general, one may say that C. dayi follows the Cascade
mountains from Washington to California (fig. 14), where it goes due south
at least to Yolo County on the west side of the Sacramento Valley (a single
record for Rumsey, elevation 300 feet; see fig. 15). Via the end of the
Caseades in the Mt. Lassen region it reaches the Sierra Nevada and thence
the western foothills, at elevations of 2000 to 4000 feet, to just west of
Yosemite National Park.

184 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SE.

Its occurrence at Walla Walla, Washington, suggests that it may yet be
found in western Idaho. The several localities in and to the northern border
of the Willamette Valley of Oregon make one expect it west of the Cascade
mountains in Washington, though it is not in the Willapa Bay list of Hatch
and Kineaid. More collecting in northwestern California may show its dis-
tribution to overlap the northeastern edge of that of C. autumnata.

THE WING VENATION OF SOME SPECIES OF Centrodera

Swaine and R. Hopping gave a detailed drawing of the basal half of the
wing of C. spurca (1928, pl. XI), and a photograph of the wing of C. de-
colorata (pl. XIII, fig. 19). Saalas studied and figured the wings of rep-

Figure 19. Wing of Centrodera spurca.

resentative species of the major categories of Cerambycidae. He remarked
(1936, p. 71) that the wing of C. decolorata as figured by Swaine and Hop-
ping does not differ in any significant way from that of species of the genus
Rhagium, and that the short extra branching of Cu, they show? is probably
just an anomaly. On the basis of their figure of part of the wing of C.
spurca, he refers the species without further explanation to the genus
Evodinus (p. 73), noting that CuZ is well developed and Cu, is 3-branched,
as for example in C. decolorata.

4. There is still no universally accepted opinion as to the homologies of the wing venation of the
Coleoptera. The anal veins of Forbes (1923) and of Swaine and Hopping are the cubitals of Saalas. The

wedge-cell of Forbes, significant in Centrodera, is the anal cell of Swaine and Hopping, and the Cubitalzelle of
Saalas.

VoL. XXXIT] LEECH: CENTRODERA SPURCA AND ALLIES 185

An entire wing of C. spurca is shown in fig. 19; the wedge-cell is large,
and 2nd A, goes off just below it (using the venational nomenclature of
Forbes, 1923). In C. decolorata, however, 2d A, goes off from an angle of

cu-3

C. DECOLORATA isth Y)

IstA
2dAy 2dAo 2dAg 3dA9

Figure 20. The wedge-cell region of a wing of Centrodera decolorata. The
venational nomenclature is here according to Forbes, 1923.

Cu,

Cu, c Cup A

Figure 21. The wedge-cell region of a wing of Centrodera decolorata. Vena-
tional nomenclature according to Saalas, 1936.

the wedge-cell itself, and this is constant in the eight examples studied; but
the branching of the first anal in these same wings is so variable (figs. 20,
21, 22, 23) that three branches cannot be cited as typical. Of the wings
examined but not illustrated, one is almost as in fig. 20, two are duplicates
of figs. 21 and 22 respectively, while the fourth has a spur as in Swaine and
Hopping’s illustration (pl. XIII, fig. 19) but the outer branch is bifid as in
my fig. 22. Obviously a 3-branched form cannot be considered typical.

The venation of C. autumnata is like that of C. spurca, but the wedge-
cell is smaller; in C. day? it is a little smaller still. In C. sublineata LeConte

186 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

and (. nevadica LeConte, the wedge-cell is absent and Ist A is only 2-
branched: but in an undescribed species related to the later, in which Ist A
is also 2-branched, there is a very small but clearly defined wedge-cell. In
the few specimens of C. picta I have checked, there is no wedge-cell, Ist A
is 3-branched, but 2d A, is not joined to 2d A. In the only wing of Apato-
physis sp. which I have examined there is no wedge-cell, Ist A is 2-branched,
and there is no trace of a cross connection between Ist A and 2d A. These
facts seem to present only difficulties to the systematist, but ultimately they
may help in finding a key to the puzzle of relationships and generic segre-
gates.

ZZ.

Figure 22. The wedge-cell region of a wing of Centrodera decolorata.

Figure 23. The wedge-cell region of a wing of Centrodera decolorata.

VoL. XXXIT] LEECH: CENTRODERA SPURCA AND ALLIES 187

THE EGGS OF THE SPECIES OF THE Centrodera spurca GRoUP®

Centrodera autumnata. The female mentioned in the paragraphs under
the heading ‘‘Copulation’’ later in this paper was put in Bouin’s solution,
and the abdomen subsequently dissected. It was packed full of large,
white eggs, so stuffed in fact that some projected up into the thoracic
cavity. All 49 eggs were mature.

The ege: length 2.65 to 3.00 mm., width 0.80 to 0.95 mm. Form elon-
gate ovoid, one end more rapidly narrowed than the other, apices rounded;
chorion dull, covered with minute spines which are sparser at apices, not
seattered, but arranged in closely spaced longitudinal lines.

Centrodera dayi. A few eggs, dissected from a dried specimen, were
rendered turgid by being soaked in a detergent solution. They appear
to be inseparable from those of C. autumnata, and have the same spinose
chorion.

Centrodera spurca. The eggs of this species are readily distinguished
from those of C. autumnata and C. dayi by their surface sculpture.

The ege: length 2.65 to 2.90 mm., width 0.70 to 0.80 mm. Fusiform,
nearly paralled-sided, apices rounded or slightly truneated. Chorion dull,
smooth, with faint net-like reticulation of regular, minute hexagonal cells;
no surface irregularities or spines apparent at X112. Color white.

Figure 24. The larva of Centrodera spurca, lateral view.

5. In life the integument of C. spurca and C. autumnata (1 have not seen C. dayi alive) is remarkably
transparent. The underside of the abdomen is so transparent that in a gravid female the individual eggs are
clearly visible, while if the antennae are held against even a moderate light the contents of the basal four
or five segments can be seen.

188 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

THe Larva or Centrodera spurca

The larvae before me (see mention of them under the heading “Life
History”) are from 30 to 34 mm. in length, but only one is fully extended
and it may not be mature. Judging by the variation in length of the
beetles, it is likely that mature larvae vary from 25 to over 40 mm. in length.

Form elongate, cylindrical, virtually without taper except for the last
two abdominal segments (fig. 24); integument shining, slightly wrinkled,
sparsely clothed with slender copper-colored hairs. Head yellowish-brown,
frons in front of the transverse line reddish-brown, becoming black along
frontal margin, mandibles black, their articular areas dark reddish-brown
to black; clypeus pale yellowish-brown, reddish at base, labrum reddish-
brown, pale apically, maxillae and palpi in part reddish-brown. Thorax
and abdomen white, prothorax with narrow yellowish-brown band near
front margin, band widening laterally; spine-bearing tumidity on ninth
abdominal tergum yellow, spine itself brown (figs. 25, 26); spiracles and
legs brown.

Head suborbicular with scattered slender setae, frons flattened and a
little depressed; labrum transversely suborbicular, rounded in front, length

Figure 25. Larva of Centrodera spurca. Abdominal tip in profile, showing the
mammilate tubercle on the tumidity at the apex of tergite 9.

VoL. XXXIT] LEECH: CENTRODERA SPURCA AND ALLIES 189

to width as 2 is to 2.5, whole subcireular margin ciliate, most densely an-
teriorly; clypeus three times as wide as long; mandibles short, thick, blunt,
cutting edge oblique with a flat grinding area adjacent, a slight angle
present at inner end of cutting edge; antennae conical, hardly protruding
beyond sockets, basal membrane large, not retractile. Three indistinct ocelli
present, two close together just below level of antenna, one on a level with
antenna and at a distance about equal to width of basal membrane of an-
tenna. Anterior edge of hypostome broadly curved, ventral mouthparts
extended; apical (third) segment of labial palpus broader and a trifle longer
than that of maxillary palpus, maxillary lobe (lacinia) a little broader
than first segment of maxillary palpus, as long as first and second segments
combined, beset with strong golden setae apically and internally; mentum
quadrate, as broad as a stipes; gula well defined, slightly protuberant, al-
most twice as long as wide. Prothorax widest in front of middle, narrowing
posteriorly; pronotum a little roughened on each side of median line in
pigmented area near front margin, and along hind margin. Abdominal
tergites 1 to 7 with dorsal ampullae finely asperate, with small shining areas,
ampullae marked by two transverse folds and an anterior one marking off
a narrowly fusiform transverse area (the shape is almost exactly as in
Craighead’s 1923 figure for the larva of Anoplodera mtens (Forster), pl.

aipaine # ‘ 2! ee z z

Figure 26. Larva of Centrodera spurca. End of the abdomen in ventral view,
showing the form of the ninth tergite with its single median tubercle.

190 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4rH Serer.

XVII, fig. 5). Tergite 8 transversely wrinkled but without ampullae; ter-
cite 9 irregularly wrinkled, median area before hind margin raised, some-
what triangularly tumid, with a single short mammillate tubercle at apex.

Legs about as long as labrum is wide, femur and tibiotarsus each twice
as long as trochanter, unguiculus a little shorter than tibiotarsus; unguicu-
lus pale in basal half, brown in apical half, with a strong seta arising
laterally at apex of pale area and projecting beyond tip of segment. Eu-
sternum somewhat triangular but with apex (cephalad) broadly rounded;
mesosternum and metasternum each divided into two equal areas by a
median transverse groove, surface finely asperate; abdominal sternites 1 to 7
each similarly divided by transverse groove, surface finely asperate, front
half of each division with transverse lines of low tubercles, 5 on each side
of median line of abdomen; sternites 8 and 9 with transverse rugae or
wrinkles. Spiracles suborbicular.

In Duffy’s key to Lepturinae (1953, p. 67) the larva runs to couplet 2,
and agrees with the first part of the first choice in having a terminal spine
on the 9th abdominal segment, and a divided frons, but does not agree as
to the remaining characters. In Craighead’s key (1928, p. 82) it traces to
numbered couplet 1, but will not run to Centrodera in the second half be-
cause the gula is almost twice as long as wide and the abdomen has only
one caudal spine; neither will it run to couplet 2.

THE Pupa or Centrodera spurca

The following scanty observations are based on the cast skin of the
reared male mentioned in the section ‘Life History.”

Pronotum with marginal line of setae, probably with two linear diseal
groups, and scattered setae between them and marginal line. Metanotum
with a group of about 35 setae on each side of median line; these are the
longest and strongest setae on the pupa. Abdominal tergites with all setae
long, slender, copper-colored as on rest of body. Abdominal tergites 1 to 5
each with two groups of about 14 slender setae on a tumid area on each
side of median line, tergites 6 and 7 with about 12 setae in each group; ter-
gite 8 with an undivided band of them, tergite 9 with 2 discal setae and
humerous apical setae on each side apically. Apex of 9th abdominal tergite
without urogomphi, but with two small seta-bearing tubereles, and a single
median seta just beyond them. Femur with about 10 setae alone outer face
near apex; tarsus with a single seta near apex.

The larvae of C. autumnata and C. dayi are unknown, although the
adults are common in at least parts of their ranges. The fact that there are
no known rearines of either of these medium sized forest insects from trees,
logs, or stumps suggests that the larvae may spend their lives in the soil;

VoL. XXXII] LEECH: CENTRODERA SPURCA AND ALLIES 191

possibly they feed on buried wood, or upon living roots, and almost certainly
they pupate in the soil.

A Partial Lire History or Centrodera spurca, with
NovTEsS ON OTHER SPECIES

Very little has been published on the life histories and habits of the
species of Centrodera. In 1894 Hopkins (1894B, p. 150) cited the larvae
of a species of cerambycid as causing extensive damage to the heartwood
of both lving and dead tulip trees [Liriodendron tulimfera| in West
Virginia®. In 1896 (p. 245) he recorded having found a newly transformed
adult in its pupal cell in the heartwood of a chestnut tree [Castanea sp.|
and stated “This beetle was identified for me through the kindness of Mr.
Howard, of the Division of Entomology, as Centrodera bicolor.” This was
an inadvertent error, as there is no such deseribed species in the genus. His
paper was republished in West Virginia with some changes in the text and
the addition of a plate, in 1897 (pp. 143-152, pl. III), and the above sen-
tence changed to read “This beetle was identified for me through the kind-
ness of Dr. Howard, of the U. S. Division of Entomology, as Centrodera
decolorata, Harr, by Mr. Linell.”

In the next paragraph he recorded another example cut from a tulip
log, and wrote that the species was “one of the most destructive wood-boring
insects that infest the wood of living trees” previously wounded by fire.
In another part of the 1897 report (p. 81) he ealled it the Destructive
Heartwood Borer, while on page 97 he wrote ‘‘Trees of all kinds in all
sections of the State that have been injured by fire or other causes, sufficient
to induce a diseased condition of the wood adjoining the wound, often have
the heartwood literally ruined by the destructive heartwood borer. This
pest extends its destructive depredations to the sound wood, which results
in the final decay of all of the inner portion. This is quite a serious trouble
and causes the loss of a large amount of timber.” Footnote 4: “Centrodera
decolonata [sic!|, Harr.” |Refers to heartwood borer four lines above. |

If Hopkins correctly associated adults and damage by larvae, it is re-
markable that the species has not gained subsequent attention. Craighead
(1923, p. 85) repeated Hopkins’ chestnut and tulip heartwood host records,
but in his 1949 report did not mention the genus or the species.

Craighead (loc. cit.) also recorded the larvae of C. decolorata from old
wet decaying oak logs (Quercus sp.) and a dead chestnut tree in Pennsyl-
vania. He cited pupation from May to July, in “a round cell of frass con-
structed before pupation. An adult was reared in early September.”

Wickham (1897B, p. 170) mentioned that the adults of C. decolorata
occur on beech, and this was repeated by Felt (1906, pp. 428, 456) who

6. Also in an 1894 paver in The Timberman, which reference I have not seen.

192 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

also recorded it as cut from butternut and gave a figure of the beetle.
Blatehley (1910, p. 1048) gave beech and maple, as did Procter (1946,
p. 177). Morris (1916, p. 20) recorded the species on or from a maple
stump. Leng (1928, p. 437) cited larvae in oak, chestnut, and tulip poplar.
Beaulne (1932, p. 199) listed the larvae as injurious to the following host
plants: apple, beech, maple, oak, chestnut; and Chagnon (1936, p. 209) as
oceurring in walnut [Juglans cinerea] and beech [Fagus grandifolia].

Knull (1932, p. 63) reported finding a teneral adult of Centrodera
picta Haldeman in its pupal cell in the decayed part of a living yellow
bireh [Betula lutea] in September; in 1946 (p. 175) he recorded the adults
as frequenting “flowers of mountain maple (Acer spicatum).” Smith (1900,
p. 291; 1910, p. 380) gave “rare on dry hickory;”. Felt (1906, p. 715) re-
peated Smith’s reeord. Leng and Davis (1924, p. 59) also cited hickory and
stated that C. picta had been taken flying to light on Staten Island, New
York.

The larval habits of the North African Apatophysis barbara Lucas were
deseribed by Peyerimhofft (1926, pp. 351-852). He found them tunneling
in a dead specimen of Limoniastrum Guyonianum (Plumbaginaceae), a
desert tree, and caged a section, partially burying it in dampened sand.
The larval stage lasted for at least seven vears! During the intense heat
of the summer they seemed to remain quiescent in their galleries in the
wood, but in winter when the sand was cool they tunneled out and round
about in it, perhaps looking for additional food supplies. Pupation took
place both in the sand and in the wood, and adults emerged in May and
June.

From what is now known, the larvae of C. spurca have some of the
same habits. They commonly feed in rotting stumps and roots (and pos-
sibly on living roots) of several kinds of trees and shrubs; they wander
freely through the soil, and pupate in the wood or in the soil as it suits
them. Nothing is known of the larval stages of C. autwmnata and C. day.

The first statement of a host plant for C. spurca, so far as I know, was
by Garnett (1918, p. 212): “Breeds in Pseudotsuga tarifolia.” I have
been unable to trace the source for his record, which was repeated by Essig
(1926, p. 452), Hardy (1926A, p. C28- 1926B,-p. 5), Doane ei akaGeso
p. 177) and Canova (1936, p. 129). Later, on the basis of material collected
by Mr. Lohbrunner, Hardy (1942, p. 10) was able to correct this.

In my field notebook for 1929 there is the following entry for April 25,
at Salmon Arm, British Columbia: “1 Parapachyta spurca; this was found
in the ground, & had changed very recently, as it was quite soft, & very
white.” Apparently I did not associate it with any rotting or other wood.
There is a female in the collection of the University of British Columbia,
labeled as taken in Victoria, B. C., January 28, 1917, by W. D. [W. Downes] ;

VoL. XXXIT] LEECH: CENTRODERA SPURCA AND ALLIES 193

judging by the date it probably was dug out of the ground. I have seen a
male and a female in the collection of the Provincial Museum of Natural
History at Victoria labeled “Saanich, B. C., 14.11.35. Ed Lohbrunner. Dug
out of ground near oak trees & Rosa nutkana.” Hardy (1942, p. 10) wrote
“Adults have been dug out of the ground in the vicinity of Garry oak
trees among the roots of Rosa nutkana in February. Large larvae were
found in gall-like swellings at the base of the rose bush but as I was un-
successful in rearing them, proof as to their identity is lacking.”

In a letter to me dated October 27, 1948, Mr. Hardy amplified this:
“Although the host tree of C. spurca as I know it, 1s evidently Garry Oak,
I have never been able to rear larvae found in the roots or base of trunk
of these trees through to the adult; they have always died or disappeared
in some way. I believe I mentioned in one of the B. C. proceedings that
possible larvae and certainly the adults have been dug out of the ground
at the base of the oaks in February and March, while I have several times
taken large larvae presumed to be this species from burrows at the base
of dying trees. This same type of larvae |stc!] has also been found in old
roots of the wild rose growing near the oaks, so you will see that uncertainty
is still rampant.”

The statement by R. Hopping (? 1921, second page) that he had found
Pachyta spurca breeding in six species of pine and some six species of shrubs
in the Sierra of California, at altitudes of from 3000 to 4000 feet, is ob-
viously a lapse of memory by the recording secretary, as to the genus and
species concerned.

On May 16, 1954, I was enlarging a small hillside vegetable garden at
427 Rose Avenue, Mill Valley, California, and dug into a bank containing
a stump of serub oak, Quercus Wislizent var. frutescens. The green tree
had been felled some four years earher. A pecular burrow-inhabiting
predaceous carabid larva was found, so a large tin was nearly filled with big
lumps and smaller bits of earth from the site, and the larva put into a
hollow at the top. Tightly lidded, the tin was taken into the house with the
expectation that I would add food and rear the carabid.

Unfortunately I became ill with pneumonia, and the tin was not opened
again until October 11. By then there was no sign of the carabid larva,
but in searching for it I broke up all the lumps of soil, and was amazed to
find a freshly transformed male of C. spurca in one (figs. 27, 28). This
must have been in its pupal cell, presumably as a mature larva, when I
picked up the lump in May. At this time I suspected that the larva might
have been working in the oak stump, then left it to pupate in the soil, as
Craighead (1923, p. 85) recorded for Anthophilax and some other genera.

By happy coincidence, on May 17, 1954, the late Gordon Stace Smith
obtained a teneral female of C. spurca, and a larva in association with it,

Figure 27. Recently transformed male of Centrodera spurca in its pupal cell
in a lump of soil excavated from a bank; see text for details. (Photograph by E. L.
Kessel. )

Figure 28. Close up of the teneral Centrodera spurca in its pupal cell. Note
that the inner surface of the cell is unlined. (Photograph by E. L. Kessel.)

VoL. XXXIT] LEECH: CENTRODERA SPURCA AND ALLIES 195

at Creston, British Columbia. In his letter of June 15 he wrote “Re Centro-
dera spurca: the Amelanchier roots where I took them were quite rotten,
almost crumbling. Actually, the adult occupied only half itself [sze!] in
the root, the other half in a sort of mud cell under the root. The larva was
only an inch or so from the adult, but quite in the root. Now I have several
times dug up larvae I believe the same, but could never before definitely
associate them, though occasionally, adults have been so exposed. I assumed
both were right in the soil, but probably in a fragment of rotten wood.”
There is a female in his collection, now at the University of British Colum-
bia, labeled Creston, B.C., February 14, 1952, and “exeavated from soil.”’

Another likely host may be mentioned. In the summer of 1931 my
father found a larva feeding in the roots of a living Soopalallie (Shepherdia
canadensis; Eleagnaceae) at Salmon Arm, British Columbia. On the basis
of its size, and the known local cerambycid fauna, it had to be Tragosoma
depsarwus (Linnaeus) or Centrodera spurca. We failed to rear it, but since
T. depsarius is known from conifers, I have little doubt that our larva was
that of C. spurca.

The larval skin from the pupal cell of my reared Mill Valley specimen,
with Mr. Stace Smith’s larva, enabled me to identify an apparently mature
larva found by my son Robin on January 24, 1954. It was dead, but in ex-
cellent condition, lying on the bottom of a small pool in Caseade Creek,
Mill Valley. No doubt it had been washed by a freshet from its pupal cell
in the stream bank.

On April 10, 1955, my son Thomas dug up a small pine tree planted as
a seedling five years before; it was at the edge of a group of Coast Red-
woods, Sequoia sempervirens, at our home in Mill Valley. In the process
he uncovered a male of C. spurca and two larvae. One larva was in its pupal
cell, and died there in late July without pupating. The other was free in
the soil, 33 mm. long, and possibly immature; it is shown in fig. 24. All were
in soil interlaced with small roots, including those of redwood, madrofio
(Arbutus Menziesu) and hazel (Corylus californicus), but no oak. We did
not notice anything else in the soil, other than humus, on which the larvae
could have fed. This, coupled with Hardy’s records of oak and wild rose,
and Stace Smith’s of service berry, suggest that they may be general feeders,
or that hke the African A patophysis barbara they may wander through the
soil for some distance from their main food supply.

Since the above was written I have seen a female [USNM] which has
the following label data “San Mateo Co. Cal., larva collected IT.17.1935.

‘

larval host dead Madrone stump. Taken out of pupal cells on X.4.34. P. C.
Ting Collector. Ase No. 81. Parapachyta spurea (Lee.) det. P.C. T.” Mr.
Ting tells me that the file of notes in which the original data were contained
was destroyed in a flood at Reno, Nevada; so it is impossible to tell which of

196 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

the above year dates is correct. At any rate, he appears to have made the
first definite association of larvae and adults.

The finding of two sizes of larvae and an adult at the same site in April,
at Mill Valley’, and an adult in its pupal cell in October, makes it fairly
sure that the life cycle takes at least two years. Some mature larvae must
form their pupal cells, pupate, and tranform into adults, in the fall. How-
ever, the fact that in the San Francisco Bay area of California the species
is on the wing over a period of four months (table I) suggests that some
larvae may not pupate until the spring. Adults of the other two species
(tables II, III) are most numerous later in the year than is C. spurca, and
presumably overwinter as larvae only; Linsley (1961, p. 9) remarks that
“Species which are active in late spring, summer, and fall usually pupate
shortly before emergence, remaining in the cell as adults for only a short
time.”

The pupal cell shown in figs. 27, 28 is completely unlined, its inner sur-
face appearing as if pressed and rubbed or tamped smooth by the larva
when the soil was damp.

Where the females lay their eggs is unknown. I made several attempts
to follow flying specimens at night, with the aid of a strong flashlight, but
that is not easily done on a wooded hillside, and they always disappeared.
On the basis of larval habitat it is likely that the eggs are laid in the soil,
as are those of at least some species of the Old World genus Apatophysis,
which Gressitt considered to be a subgenus of Centrodera. Certainly the
males resemble species of the C. spurca group, but perhaps we should not
expect the egg laying habits to coincide, because as Butovitch has remarked
(1939, p. 225) the species of Apatophysis are desert-inhabiting forms in
which the females show morphological adaptations similar to those of desert
prionids: elongated abdomen, widely separated hind legs, shortened elytra
and very long ovipositor.

There are no published records as to the longevity of the adults of
Centrodera. My only information is based on a female of C. spurca of
unknown age, collected at light in Mill Valley, California, on June 20. She
was kept in a jar containing some damp sand and sawdust, and lived until
July 14; her only source of food was a very weak solution of honey.

FLIGHT PERIODS OF THE ADULTS OF THE Centrodera spurca GROUP

Centrodera spurca is a species of late spring and summer. Ninety-nine
per cent of the 1061 specimens studied were taken during the months of
May—August, inclusive, and 87 per cent of them in June and July. There
are a few exceptional records; E. I. Schlinger took a female at Glendale,

7. In early April, 1959, Dr. E. S. Ross dug up several adults of C. spurca at his property in Mill Valley,
an additional record.

VoL. XXXIT] LEECH: CENTRODERA SPURCA AND ALLIES Ug)

Los Angeles County, California, March 21, 1947 [UCD], and E. A. Dodge
found a female at Santa Cruz, Santa Cruz County on “III.17,” which may
have been March 17, or March, 1917. It is possible that both these specimens
were found in the soil, and are not actual flight appearances, since one
would expect males to be out before females. However, exceptional climatic
conditions may bring out beetles which have pupated in exposed situations.
For instance E. S. Ross mentioned to me on April 6, 1959, that he had
recently dug up several adults of C. spurca in Mill Valley, Marin County,
California, and that a specimen had come to light at his window “at least
two weeks ago,” 7.e. during the last week of March. There had been an
exceptionally warm spell during the second half of March that year.

Table I summarizes the dates of capture of the examples of C. spurca
seen by me, arranged by geographic regions from north to south, with di-
visions east and west where the records permit. The main emergence is
obviously in June and July, both on the coast and in the mountains, but it
is interesting to compare the totals of males and females for each of the two
months. It is surprising how many males are still around in August.

Following are the ‘First of season” records for specimens attracted to
our (indoor) house lights at 427 Rose Avenue, Mill Valley: 1950 (May 11),
1951 (May 8), 1952 (May 6), 1953 (May 3), 1954 (May 3), 1955 (May 11),
1956 (May 14), 1957 (April 28), 1958 (May 18), 1959 (May 5). All these
specimens were males. Of course it is uncertain that our lights attracted
the actual first emergents of the area.

Another point brought out in table I has to do with northern distribu-
tion. Although C. spurca is common enough on southern Vancouver
Island, British Columbia (of 39 specimens, my most northerly records are
in about the same latitude for the inner and outer coasts, Nanaimo and
Ucluelet respectively), and equally common in the southern interior of the
mainland (Seton Lake to Creston), I have no records for the mainland west
of the coast mountains. Neither is the species in Stace Smith’s lists (1929,
1930) of the beetles from Copper Mountain, near Princeton and just east
of the mountains. Yet the Vancouver area and the Lower Fraser Valley
have had resident collectors for well over 60 years. I have not seen enough
material from northwestern Washington to know how far south of British
Columbia this state of affairs continues.

Centrodera autumnata is common during July, August and September,
reaching its peak in August, by which time most C. spurca have disappeared.
I took one male at Mill Valley on June 28, 1959, and have seen a male and
a female from Ben Lomond dated June 3, 1946; this surprisingly early date
is substantiated by a single male taken at light at Mill Valley on May 29,
1958. The first specimen for 1954 is dated July 11, and that for 1957, July

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CALIFORNIA ACADEMY OF SCIENCES

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198

199

LEECH: CENTRODERA SPURCA AND ALLIES

VoL. XXXIT]

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200 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

TaBLE II. Captures of adults of Centrodera autumnata by months, region-
ally.

May June July August Sept.

Centrodera autumnata, EEE

new species ¢ 9|¢ e|¢ e|¢ ele 9

California: Humboldt County,

south to south shore of
Marin County 1 — 2 — 16 1 S4:500i(2

California: San Francisco Co.
south to Monterey County — — 2 1 26 4 24 4 1 2

California: San Luis Obispo
Co. south to San Diego
County See Se SS 2 Se eee

TOTALS 1 —

TaBLE II]. Captures of Centrodera dayi adults by months, regionally.

June July August Sept.
Centrodera dayi, new species i. © | & | ieee | o 2

Washington: west of Cascades SS 88 Se ee Eee

Washington: east of Cascades — — 1 2 38 — — =
Oregon: west of Cascades — — § 4 400° =
Oregon: east of Caseades, including

Medford, Talent 1 —& 861 12 ti
California: Cascade mts., east and

south to Lassen Co.; also to

Yolo Co. on west side of

Central Valley — — = 1. ieee
California: Sierra Nevada. Plumas

Co. south to Tuolumne Co. — | 2% 2 40S

TOTALS ie el Ee 10 30 n —_—- —

VoL. XXXIT] LEECH: CENTRODERA SPURCA AND ALLIES 201

4. These early emergences are probably due to the same causes as in C.
spurca, q.v.

The records are summarized in table II. Note the surprising disparity
in the numbers of females as compared to males in this species, in constrast
with the totals for C. spurca and C. dayi in tables I and III. Of course this
is not a true figure of the proportions of the sexes, but only of relative at-
traction to light, though it may indicate a difference in habitat in C. au-
tumnata females.

As shown in table ITT, C. dayi appears to have a remarkably short flight
period. Of the 94 specimens studied, from southern Washington to central
California, all but two were collected in July and August, with by far the
ereatest number in August. Of course there are fewer collectors in the field
during September than in the summer, but this fact hardly explains the
complete lack of September records.

HABITS OF THE ADULTS oF Centrodera SPP.

DAYTIME RETREATS. The adults of C. spurca, C. autumnata and C. dayi
are nocturnal; although fewer than half of those seen are documented as
having been taken at light, I suspect that at least ninety per cent were col-
lected in this manner. One is occasionally found in plain sight during the
day, near an electric ight which has been on all night, but they normally
hide successfully.

To the best of my knowledge the only person to take specimens of C.
spurca during the day on a number of occasions has been that outstanding
collector, the late Gordon Stace Smith. In a letter to me of September 7,
1948, about finding them at Creston, British Columbia, he stated that the
best way was to beat the overhanging branches of Douglas fir (Pseudotsuga
Menziesu.), but that he had beaten them also from cedar (Thuja plicata),
aspen (Populus tremuloides), choke cherry (Prunus virginiana var.
demissa) and willow (Salix sp.). One of the specimens in his collection
| UBC] is labeled as from hazel (Corylus sp).

More unusual are two males and a female |WSU] labeled ‘Moscow
Mt., Ida. May 1935. Ground squirrel burrow.” Unless the female fell in
while looking for an oviposition site, and the males followed her, this record
is hard to explain. Possibly they all pupated in the soil adjacent to the
rodent’s tunnel, and emerged into it.

ATTRACTION TO Licut. I suspect that all the western species of Centro-
dera are crepuscular or nocturnal, and know that at least five of them are
attracted to light. The coarsely faceted eyes of the eastern species suggest
that they too are nocturnal, and Professor J. N. Knull (in litt., 1962) states
that C. sublineata LeConte comes to light, while Engelhardt (1942, p. 38)
records a specimen so taken at Middletown, Virginia, April 13. Hatch

202 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER.

(1925, p. 579) lists C. decolorata as taken at light, and Leng and Davis
(1924, p. 59) cited C. picta.

Centrodera spurca has several times been recorded as attracted to light.
For instance Garnett (1918, p. 212) wrote “It flys [sic!] at night and is
attracted to light.” Hardy (1926A, p. C28; 1926B, p. 5) said “Taken only
at ‘light.’ Occasional. Latter part of May to July.” Doane ef al. (1936,
p. 176), “. . . frequently flies to light.” Moore (1937, p. 88), “Taken by
the author at lights in San Diego in April and July.” It is interesting to
compare the ratios of males to females in collections, by species, as shown
in the foregoing tables. This suggests either that females of C. autumnata
are much less attracted to light than are the males (and are females of
other species), or that they stay near the egg-laying sites while the males
range widely.

My own experience with C. spurca and C. autumnata is that they are
much attracted to light, especially of a rather low intensity. In season they
appeared on the windows of, or entered, rooms in which there were at
various times 1) one or more exposed regular 100-watt electrie light globes,
or 2) only shaded reading lamps with similar globes, or 3) lights on the
same walls as the windows and thus not directly visible from outside, or 4)
two 20-watt fluorescent light tubes. They were also attracted to an outdoor
light trap using a standard 300-watt globe. These species and C. dayi have
all been taken at single-mantle “Coleman” gasoline lanterns. I have had
examples of both C. autumnata and C. spurca fly to the same “Coleman”
lantern of an evening in places as widely separated as Mill Valley, Marin
County (altitude about 625 feet; mid-July), and just below the top of
Junipero Serra peak in the Santa Lucia mountains of Monterey County
(altitude approximately 5800 feet; mid-August). Examples of both C.
spurca and C. dayi from Oregon and California have been seen labeled as
taken at black (ultra violet) light, and I have so taken C. spurca at Mill
Valley, California, in June and July.

On the other hand, using a 500-watt “Photoflood” globe in a metal re-
flector from 9 to 10 P.M. on August 25, 1950, I watched examples of C.
autumnata on the trees by our house in Mill Valley. They were in all cases
flying to and settling on the branches of living madrono (Arbutus Menzresi)
trees, ten to fifteen feet above the ground; one landed on a spray of fruits
and investigated it for some time, but did not appear to eat any of the
berries. The beetles did not seem to be in any way disturbed by the bright
hght, and not one of them flew to it.

Mazkhin-Porshnyakov (1960) has proposed a most interesting theory
of why insects fly to hght at night. As he points out, “insects fly not only
to a radial source of light rays, for instance, a lamp, but also to diffuse light
reflected from a screen.”

Vou. XXXIT] LEECH: CENTRODERA SPURCA AND ALLIES 203

The following quotation is from his summary. “It is very probable that
light attracts insects only because it is a sign of open space, or an absence
of restraints. In nature open spaces are much better illuminated than
closed ones and they attract insects because there are no obstacles to avoid.
Thus they can orient themselves more easily in the carrying out of one or
another of their vital functions. During their life insects constantly make
use of brighter illumination as an indicator of open space (exit from eavi-
ties, from dense vegetation, ete.).

“Open space is characterized, above all, by an abundance of short-wave,
generally ultraviolet rays. The source of these rays during the night ap-
pears to be the sky—an orienting source which leads to open space. The
greater the quantity of short-wave rays (including ultraviolet rays) an
artificial hght contains, the more readily it attracts nocturnal insects. Such
illumination is similar to, but brighter than, natural light at night.”

ABILITY TO WALK ON VERTICAL PANES oF QGuass. There have been a
number of papers written on the abilities of certain insects to hold onto or
walk up very smooth surfaces. In the adephagous, silphoid, and some other
groups of beetles it is not uncommon to find isolated or grouped specialized
sucker-like hairs on the tarsi, especially on the fore tarsi, and often only
in the males. Miall (1903, pp. 53-59) gives an interesting discussion of
some of the problems involved in understanding how these suckers act.

One is surprised to find that the matter is still puzzling entomologists
and seems not to be resolved. Curran (1958, p. 85), in considering the
pulvilli of flies and how the insects land on ceilings, concludes ‘‘The con-
tention that the puvilli are sticky is almost certainly mistaken. What is
certain is that the pads act as suction cups, serving to anchor the fly firmly.”
The subject was fully discussed, with the same conclusions, by Kirby and
Spence nearly 150 years ago.

Not having investigated the matter in detail, I wish merely to draw
attention to the fact that both sexes of C. spurca and of C. autumnata are
able to land on and hold to a vertical window glass, from full flight. I do
not know whether the hairs of the tarsal pads have suction cup tips, or
exude an adhesive, but suspect the former type. I have seen a male C.
spurca so “stuck” to a window pane by one foot, by his own doing, that he
could not release himself, but could only shde slowly down the glass. One
is tempted to explain this by the analogy of sliding a rubber suction cup
on glass.

ADULTS ON FLowers. No records of C. spurca or allies having been taken
on flowers are known to me. In fact I do not recall flower records for any
west-coast nocturnal Cerambycidae, though many of the diurnal species,
especially in the Lepturini, are pollen feeders. It is thus interesting that

204 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

during the course of this study pollen has been found on examples of all
three species of the C. spurca group.

Some males of C. autumnata have earried enough of it to obscure the
surface of the prosternum, and the depression between the clypeus and the
antennal bases. Pollen has been noted on the head (base of elypeus and
near, genae, occiput, first three antennal segments, submentum—but not on
the mouthparts, which are perhaps most easily cleaned), sides of prothorax,
pro-, meso- and metasternum, fore and middle coxae, and the femora of all
legs. The distribution suggests that the beetles may have been probing or
feeding on floral parts, but not necessarily on pollen.

Mr. Robbin Thorp at the University of California, Berkeley, has been
so kind as to examine samples of the pollen which he removed from the
beetles. He reported that specimens of C. autumnata from Mill Valley,
Marin County, California (mid-August) carry a common type found in
several plant families. After eliminating groups not occurring in Marin
County, and those not in bloom while the beetles were active, he suggests
the following as probable sources of the pollen; my comments are in paren-
theses.

Fagaceae. Lithocarpus sp. (Tanbark oak, closely allied to Quercus.
There is only one species, L. densiflorus Rehd.; it is common at 427 Rose
Avenue, where the beetles were collected. )

Fagaceae. Castanopsis sp. (Chinquapin. A single species in the area,
C. chrysophylla, but much less common than tanbark oak.)

Caprifoliaceae. Sambucus sp. (Elderberry. Two species in Mill Valley,
S. callicarpa, red fruited, and S. coerulea, blue fruited. Neither species is
known to occur close to the collecting site.)

A female of C. dayi from Medford, Jackson County, Oregon, August 29,
earries pollen similar to that on the C. autwmnata males, as do a male and
a female of C. spurca from Mount Hermon, Santa Cruz County, California,
July 16. Both these latter have a good covering of pollen on the pro- and
mesosternum and adjacent parts, and dorsally on the head.

A male of C. spurca from Johnsville, Plumas County, California, July
21, has a seattering of pollen which Mr. Thorp identifies as Pinus sp.,
probably lodgepole pine, P. contorta var. murrayana, on the pro-, meso-
and metasternum and adjacent areas. Two other males and three females
from the same place, taken on three different dates in July, also have Pinus
sp. pollen. One female has a good deal, including on the femora, tibiae and
pronotum. Four males and four females from Twain Harte, Tuolumne
County, California, late June to late July, each show a little Pinus sp.
pollen ventrally.

Since all western species of Centrodera are to the best my knowledge
erepuscular or nocturnal, it was a surprise to read in Knull (1946, p. 175)

VoL. XXXIT] LEECH: CENTRODERA SPURCA AND ALLIES 205

that in the eastern C. picta “Adults frequent flowers of mountain maple
(Acer spicatum Lam.).” In a letter dated February 22, 1962, Professor
Knull verified this statement, remarking that he had so taken them in
southern Pennsylvania, and that he had collected isolated specimens by
beating foliage. On the basis of such diurnal habits, it is a surprise to find
that Leng and Davis (1924, p. 59) record examples of C. picta as having
flown to light at Clove Valley, Staten Island, New York, June 7. Is it pos-
sible there is a sibling species involved? Certainly the specimens of C. picta
I have examined differ from the rest of the species in the structure of the
prosternum, and perhaps do not belong in Centrodera.

ATTRACTION OF SOME WESTERN SPECIES TO Barts. Currie (1904, p. 28)
recorded having taken Pachyta spurca LeConte while “sugaringe” for moths
at Kaslo, Kootenay Lake, British Columbia, during the summer of 1903.
He used a mixture of 3 pounds of sugar and 1 pound of molasses, boiled
till the sugar was dissolved, then thinned with beer and a small glass of rum.
Among specimens from Quamichan Lake, Vancouver Island, British
Columbia (Hanham Collection, PMV) there is a male of C. spurca labeled
“at sugar.”

Mr. L. G. Gentner took the following specimens at Talent, Oregon: a
male of C. spurca “In ecodling moth bait pan” on 11.V.1936 [CIS]
and another on 19.VII.19388 [LGG], a female “In bait pan” on 3.VIT.1941
[LGG] and another on 31.VII.1941 [UCD]. Also a male [UCD] and a
female [LGG] of C. dayi on 24.VII.1938, both “In ¢. moth bait pan.” The
bait was made of 1 quart of Fleischmann’s Diamalt and 19 quarts of water,
giving 5 gallons of solution, to which 2 cakes of Fleischmann’s yeast were
added, and the mixture allowed to ferment. It was placed in open sauce
pans which were suspended in the upper third of the fruit trees (Gentner,
in litt., 1962).

One female of C. dayi from Monroe, Benton County, Oregon, 23.VII.
1931 (N. P. Larson, OSU) is labeled “In moth trap.” Judging by the con-
dition of this specimen, and of a male taken 17 days earlier, both were
collected from liquid bait traps.

STRIDULATION. Typical of the Cerambycinae, there is in C. spurca a
longitudinal tumid dark area on each side of the median line of the pre-
scutum, the two forming a pars stridens, the surface of which is uniformly
covered with fine transverse ridges.

Each time the prothorax is moved backward and forward a ridge
(plectrum) on the underside of its hind margin is drawn across these
striated plates, producing a squeaking noise. It can be heard easily when
live specimens of C. spurca and C. autumnata are held or restricted in their
movements, and equally by moving the parts of a dead, relaxed specimen.

Examination of the pars stridens in the three species of the C. spurca

206 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

group has shown too much variation in its form to offer specific separations.

CopuLation. On August 16, 1954 a male and a female of C. autumnata
were taken as they came to light separately at Novato, Marin County, Cali-
fornia. The next day they were released together; the male attempted
copulation immediately, and the act was observed with the aid of a stereo-
scopie microscope.

The male mounted the back of the female from behind and took a posi-
tion such that his head was almost at the basal one quarter of her elytra.
Bending the tip of his abdomen down to contact hers, he extruded his
parameres enough to catch against the tip of her fifth abdominal sternite,
then inserted the tip of the aedeagus between this sternite and her eighth
visible tergite (pygidium) and pulled upward.

At all times during the copulation the hairs of the parameres caught
against and pushed down the rim of the fifth sternite, while the aedeagus
pulled up on the pygidium. With these segments held apart, the intro-
mittent organ was inserted into the tip of the ovipositor, which the female
kept retracted except when she tried to prevent copulation, or terminated
it by extending the ovipositor beyond the point to which the intromittent
organ could reach.

These copulatory actions differ from any cited in Butovitsch’s summary
for the Cerambycidae.

Alexander (1962, p. 66) noted that the parameres of beetles have often
been interpreted as pries; but on the basis of his observation of a mating
in the carabid Pasimachus punctulatus Haldeman he is inelined (1959,
p. 485) toward Jeannel’s view that they are chiefly of use in producing
tactile sensory effects upon the female. There is surely no doubt that this
latter is true; but it is equally certain that at least in C. autumnata (and
on the basis of similarities in structures, probably in many Cerambycidae),
they are used not as true pries but as holders or pushers.

A pairing of C. spurca has also been observed. In this case the female
was more receptive; after a quick push down with the parameres and pull
upward with the tip of the aedeagus, the male extruded and inserted his
intromittent organ, then retracted the parameres and aedeagus. The action
up to this point was so fast that there was no chance for the parameres to
be effective as tactile exciters, and they were not so used thereafter. The
female remained quiescent; the intromittent organ was inserted almost up
to the paired armatures (for a comparable example see the excellent fig. 8
on p. 27 of Jeannel’s 1955 paper), at which point it became enlarged and
somewhat bulbous. The apex of the aedeagus and the parameres barely ex-
truded from the abdomen, and thus the tips of the abdomens of the ecopula-
ting beetles were separated by between one and two millimeters, the length

VoL. XXXIT] LEECH: CENTRODERA SPURCA AND ALLIES 207

of the everted intromittent organ between the paired armatures and the
ostium at the apex of the aedeagus.

The male mounted the female so that his mandibles touched the tip of
her seutellum. His front tarsi wrapped around her prothorax a little below
the lateral tubercles, the claws holding at the median line just in front of
her front coxae; his middle legs encircled her body, the tarsi closely in
front of her hind femora, and his hind legs clasped her abdomen. The pair
remained in copula for twelve minutes.

The positioning of the male with his mandibles at the female’s scutellum
may be significant. Michelsen (1958, p. 350) reported that in Rhagiwm
bifasciatum Fabricius and R. mordar DeGeer the males “lick” the scutellar
and adjacent areas of the females with their mouthparts, to calm them dur-
ing courtship. In R. bifasciatum, only “licking” near the scutellum was
effective, while in R. mordax any upper part of the elytra gave results.
Males of R. bifascratum never protrude the genital organs more than 1 milli-
meter, so that the abdominal tips of the sexes are close together during
copulation; thus a male larger than the female would be forced to “lick”
in front of the effective area, and fail to mate successfully. In R. mordax
the male genital organs are protruded several millimeters, giving more
latitude of movement, so that the size of the male relative to the female is
less important..

No such attempt to quiet the female occurred in the pairing of Centro-
dera spurca reported above, though the male’s mouthparts were adjacent
to the scutellum of the female. No “licking” was observed in C. autumnata,
but neither was it watched for. In C. spurca especially, there is such a
range of sizes in both sexes that I doubt a “licking” or similar process is es-
sential to successful matings.

Dusious REcORDS

Among the 1364 examples of C. spurca, C. autumnata and C. dayr
personally seen’, two short series carried what surely must be erroneous
locality labels. These were two specimens of C. spurca said to be from the
Mackenzie River, Northwest Territories, Canada, and four of C. autumnata
labeled Patagonia, Arizona. In addition two others are suspect. A single
male of C. autumnata is labeled ‘Orinda, Cal.” Contra Costa County, with
an illegible year date only, and no collector’s name. Of 209 specimens of
C. autumnata seen, this is the only one from the east side of San Francisco
Bay proper, an area which has had resident collectors for more than 75
years. Although the record seems distributionally probable, I prefer to

8. This figure does not tally with the totals from tables I, II, and III. This is because of specimens

which were adequate for study but lacked data as to month of collection, or were from places which I could
not find in any gazetteers.

208 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

await fully documented specimens before accepting it. A female of C.
spurca is labeled Salt Lake City, Utah; it has been included on the map
(fig. 14), but is a long way from other known localities for the species.

A few dates of collection are puzzling, and may be the results of labeling
from memory some time after the actual collecting. Such records are not
used in the tabulations of captures.

LITERATURE CITED

ALEXANDER, R. D.

1959. The courtship and copulation of Pasimachus punctulatus Haldemann
[sic!] (Coleoptera: Carabidae). Annals of the Entomological Scciety
of America, vol. 52, no. 4, p. 485, 2 figs.

1962. The role of behavioral study in cricket classification. Systematic Zoology,
vol. 11, no. 2, pp. 53-72, 19 figs.

AURIVILLIUS, PER O. C.

1912. Cerambycidae: Cerambycinae. In W. Junk, edited by S. Schenkling,
Coleopterorum Catalogus, Pars 39, pp. 1-574. Berlin, W. Junk.

BALFOUR-BROWNE, F.

1932. A Text-book of practical entomology. London, Edward Arnold & Co. Pp.
viii + 191, 116 text figs.

1940A. British water beetles. Volume I. London, The Ray Society. Pp. xx + 375,
89 text figs., pls. I-V.

1940B. Concerning the three species Haliplus lineolatus Mannerh., H. wehn-
ckei Gerh. and H. heydeni Wehncke (Col., Hydradephaga), with
some remarks upon the orientation of the male sexual armature in the
Hydradephaga. A reply to Dr. G. Falkenstrém. Entomologist’s
Monthly Magazine, vol. 76, no. 913, pp. 121-128, 4 text figs.

BAUMBERGER, P.

1921? [The result of a month’s collecting—from June 15th to July 15th—at
St. Helena, Napa County, California.] In: Proceedings of the Pacific
Coast Entomological Society, vol. 1, no. 6, third page.
[The meeting was held on August 20, 1910, but the date of publi-
cation of the Proceedings is uncertain, and may have been earlier
than 1921. See MacSwain, Pan-Pacific Entomologist, vol. 27, no. 3,
p. 109; 1951. The signature in which Baumberger’s list appeared
consisted of 8 unnumbered pages. ]

BEAULNE, J. I.

1932. Longicornes misibles aux vegetaux ligneux du Canada. Le Naturaliste
Canadien, vol. 59, no. 10, pp. 196—203.

BLATCHLEY, W. S.

1910. An illustrated descriptive catalogue of the Coleoptera or beetles (ex-
clusive of the Rhyncophora) known to occur in Indiana, with bibli-
ography and descriptions of new species. Indiana Department of
Geology and Natural Resources, Bulletin 1. Indianapolis, Nature Pub-
lishing Co. 1386 pp., 590 text figs., 1 map.

VoL. XXXIT] LEECH: CENTRODERA SPURCA AND ALLIES 209

BRADLEY, J. C.

1930. A manual of the genera of beetles of America north of Mexico. Keys
for the Determination of the Families, Subfamilies, Tribes, and Genera
of Coleoptera with a Systematic List of the Genera and Higher Groups.
Ithaca, New York. Daw, Illston and Co. xi + 360 pp.

BuGNON, E.

1931. L’appareil copulateur des longicornes du sexe male (avec 11 figures).
Bulletin Biologique de la France et de la Belgique, tome 65, fase. 2,
pp. 234-265, 10 text figs.

BUTOVITSCH, V. VON

1939. Zur Kenntnis der Paaring, Eiablage und Ernahrung der Cerambyciden.
Entomologisk Tidskrift, vol. 60, pp. 206-258.

Canova, M. F.

1936. An annotated list of the Lepturini of Oregon. Pan-Pacific Entomologist,
vol. 12, no. 3, pp. 126-132.

Casey, T. L.

1913. Further studies among the American Longicornia. Memoirs on the
Coleoptera, vol. 4, pp. 193—400.

CHAGNON, G.

1936. Contribution a l'étude des coléoptéres de la Province de Québec.
Famille XLI. Cérambycides. Le Naturaliste Canadien, vol. 63, nos.
8 and 9, pp. 201-210, pl. XV.

CHEVROLAT, L. A. A.

1860A. [As reported in the section entitled “Mélanges et nouvelles.”] Revue
et Magasin de Zoologie, ser. 2, vol. 12, p. 96.
[Following an editorial preamble on pp. 95-96, Chevrolat’s pre-
liminary diagnosis of Apatophysis toxvotoides, new genus and
species, is given. February issue of the journal.]

1860B. Description de Coléoptéres nouveaux d’Algérie. Revue et Magasin de
Zoologie, ser. 2, vol. 12, pp. 303-305.
[Formal description of Apatophysis torotoides. July issue of the
journal. }

CRAIGHEAD, F. C.

1923. North American cerambycid larve. A classification and the biology of
North American cerambycid larve. Canada Department of Agricul-
ture, Bulletin no. 27—-New Series, Technical (Entomological Bulletin
no. 23). 239 pp., 8 text figs., pls. I-XLIV.

[Plates I-V inserted between pp. 152-161, and pls. XXV-XXXII
between pp. 200-215 are differentiated and not included in page
numbering; pls. VI-XXIV and XXXIII-XLIV are on page stock
and included in the serial page numbering. |

1949. Family Cerambycidae. In: Craighead et al., Insect enemies of eastern
forests. U. S. Department of Agriculture, Miscellaneous Publication
no. 657. [2+] 679 pp., 197 figs. (Cerambycidae, pp. 228-271, figs.
50-57.)

210 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

CURRAN, C. H.
1958. Insect acrobat: feats of the housefly. Natural History, vol. 67, no. 2,
pp. 82-85, 9 figs.
Currie RE:
1904. An insect-collecting trip to British Columbia. Proceedings of the En-
tomological Society of Washington, vol. 6, no. 1, pp. 24-37.
DoANE, R. W., E. C. VAN Dyker, W. J. CHAMBERLIN, and H. BE. BuRKE

1936. Forest insects. A textbook for the use of students in forest schools, col-
leges, and universities, and for forest workers. New York and London,
McGraw-Hill Book Co., Ine. Pp. xii + 436, frontispiece, 234 text figs.

Dunn Hy AG Je

1953. A monograph of the immature stages of British and imported timber
beetles (Cerambycidae). London, British Museum (Natural History).
Pp. viii + 350, frontispiece, 292 figs., pls. I-VIII.

EHMARA, S.

1954. Comparative anatomy of male genitalia in some cerambycid beetles.
Journal of the Faculty of Science Hokkaido University, ser. 6, Zoology,
vol. 12, nos. 1-2, pp. 61-115, 199 figs.

E\NGELHARDT, G. P.

1942. In: C. G. Siepmann, Proceedings of the Brooklyn Entomological Society.
Bulletin of the Brooklyn Entomological Society, vol. 37, no. 1, p. 38.

Essie, EK. O.

1926. Insects of western North America. A manual and textbook for students
in colleges and universities and a handbook for county, state and fed-
eral entomologists and agriculturists as well as for foresters, farmers,
gardeners, travellers, and lovers of nature. New York, The Macmillan
Co. xi+ 3 + 1035 pp., 766 text figs.

IMAG, 1st, (Op

1901. List of the Coleoptera of Southern California, with notes on habits and
distribution and descriptions of new species. Occasional Papers of the
California Academy of Sciences, no. 8, pp. [4 +] 282.

Ios IDE ee

1906. Insects affecting park and woodland trees. New York State Museum,
Memoir 8, vol. 2, pp. [2 +] 333-877, figs. 64-223, + pls. 49-70.

ForBes, W. T. M.
1923. The wing-venation of the Coleoptera. Annals of the Entomological So-
ciety of America, vol. 15, no. 4, pp. 328-352, incl. pls. XXIX—XXXV.

[The December, 1922, number of the journal, was actually pub-
lished February 20, 1923.]

GAHAN, C. J.

1906. Cerambycidae. In: C. T. Bingham, ed., The Fauna of British India, in-
cluding Ceylon and Burma. London, Taylor & Francis. Coleoptera,
vol. 1, pp. xviii + 329, 107 text figs.

VoL. XXXII] LEECH: CENTRODERA SPURCA AND ALLIES 211

GARNETT, R. T.
1918. An annotated list of the Cerambycidae of California (Col). Canadian
Entomologist, vol. 50, no. 5, pp. 172-177; no. 6, pp. 205-213; no. 7,
pp. 248-252; no. 8, pp. 281-284.

GEMMINGER, M. and E. Von HAROLD
1872. Catalogus Coleopterorum hucusque descriptorum synonymicus et sys-

tematicus. Tom. IX. Scolytidae, Brenthidae, Anthotribidae, Ceram-
bycidae. Monachii, E. H. Gummi, pp. 2669-2988 [4+ 12].

GENTNER, L. G.
1962. (Letter of February 6, to H. B. Leech.)

Gipson, A.
1917. The entomological record, 1916. Notes of captures. 47th Annual Report
of the Entomological Society of Ontario (1916), pp. 137-171 (Coleop-
tera, pp. 145-152).

GILBERT, E. E.

1953. The homologies of the male genitalia of Rhynchophora and allied
Coleoptera. Annals of the Entomological Society of America, vol. 45,
no. 4, pp. 633-637, 8 text figs. (Issue for December, 1952, published
January 30, 1953.)

Gressirt, J. L.

1947. Notes on the Lepturinae (Coleoptera, Cerambycidae). Proceedings of the
Entomological Society of Washington, vol. 49, no. 7, pp. 190-192.

1951. Longicorn beetles of China. Longicornia, vol. 2, pp. [2 +] 667, frontis-
piece (map), pls. I-X XII with facing pp. of explanations.

GUIGNOT, F.

1931-1933. Les Hydrocanthares de France. Hygrobiidae, Haliplidae, Dytis-
cidae et Gyrinidae de la France continentale avec notes sur les espéces
de la Corse et de l’Afrique du Nord francaise. (Edition des Miscel-
lanea Entomologica.) Toulouse, Les Fréres Douladoure. Pp. xv
[— preface by M. P. de Peyerimhoff] + 1057, 558 text figs., 7 pls.

HALDEMAN, S. S., and J. L. LECONTE
1853. Catalogue of the described Coleoptera of the United States. By Friedrich
Ernst Melsheimer, M.D. Revised by S. S. Haldeman and J. L. LeConte.
Washington, Smithsonian Institution. Pp. xvi + 174.

Harpy, G. A.

1926A. Cerambycide of Vancouver Island. (Preliminary annotated list.)
Province of British Columbia. Report of the Provincial Museum of
Natural History for the year 1925. Victoria, Charles H. Banfield. Pp.
C24-C33, pl. IV and facing explanations between pp. C26 and C27, pl. V
and facing explanations between pp. C28 and C29, + glued in pink slip
of Corrigenda and Addenda after p. C38.
[This slip of corrigenda and addenda gives page references to the
Provincial Museum Report, as above, and to the separately paged
reprint, for which see next item. ]

212 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER.

Harpy, G. A.—Cont.
1926B. Cerambycide of Vancouver Island (Preliminary annotated list). Vic-
toria, Charles H. Banfield. Pp. 1-10 [+ 2] + pls. IV and V and facing
explanations (terminal) + bound in pink slip of Corrigenda and Ad-
denda to this and to the preceding item.

1942. Notes on some wood-boring beetles of Saanich, Vancouver Island, B. C.
(Coleoptera: Cerambycidae & Buprestidae). Proceedings of the En-
tomological Society of British Calumbia, vol. 39, pp. 9-13.

1948. (Letter of October 27, to H. B. Leech.)

HARVEY, R. V.

1907. The British Columbia list. Coleoptera. Family Cerambycidae. Bulletin
of the British Columbia Entomological Society, no. 6, pp. 3-4.

[Page 4 ends with “(To be continued.)” after Leptura obliterata
Hald. At some time, presumably between the issuing of no. 6 in
June 1907, and no. 7 in September 1907, a separately published
“Page 4 continued” was published. It completes the list of Ceram-
bycidae, and gives additions to earlier lists of Neuroptera and
Lepidoptera. It is all on one long page, 1°%4 inches longer than
p. 4. Reprinted in 1926; see next item.]

1926. The British Columbia list. Coleoptera. Family Cerambycidae. Quarterly
Bulletins of the Entomological Society of British Columbia (Incor-
porated), Numbers 1 to 10, March, 1906, to June, 1908. No. 6, pp. 3-6.

[A reprinting of the previous item, but because of differences in
types and page size, not all items appear on the equivalently
numbered pages in the reprint and in the original. ]

Hatcu, M. H.

1925. A list of Coleoptera from Charlevoix County, Michigan. Papers of the
Michigan Academy of Science, Arts and Letters, vol. 4, pp. 543-586.
[Usually listed as 1924, but the volume is marked as ‘‘Set up and
printed February, 1925.’’]

1939. Preliminary list of the Coleoptera of Washington. [2 +] 41 pp.

[This is mimeographed, on only one side of each sheet, thus the
page numbers given above are not comparable to those of a type-
set book. Issued by the Department of Zoology, University of
Washington, Seattle, October 1939. Sold for 50 cents. Dr. Hatch
does not consider it to constitute publication, but it appears to ful-
fill the requirements of the International Code of Zoological
Nomenclature as adopted by the XV International Congress of
Zoology and published in 1961; q.v. pp. 7, 9.]

Hatrcu, H., and T. Krincarw

1958. A list of Coleoptera from the vicinity of Willapa Bay, Washington.
Privately printed; The Calliostoma Co., 1904 East 52nd, Seattle 5. Pp.
4-+ 22+ map.

Hopkins, A. D.

*1894A. Insect enemies of the yellow poplar. The Timberman. (Issue for
July 21; 8 pp., 11 figs.)9

9: Reference not seen by H. B. L.

VoL. XXXIT] LEECH: CENTRODERA SPURCA AND ALLIES 213

Hopkins, A. D.—Cont.

1894B. Notes on some discoveries and observations of the year in West Vir-
ginia. Insect Life, vol. 7, no. 2, pp. 145-151.

1896. Some notes on insect enemies of trees. Canadian Entomologist, vol. 28,
no. 10, pp. 243-250.

1897. Report of the Entomological Department. In: Ninth Annual Report of
the West Virginia Agricultural Experiment Station for the fiscal year
ending June 30th, 1896. Fairmont, Index Steam Printing House. Pp.
72-164, 2 text figs., pls. I-III, VI-VII. 3 maps.

Hoppine, R.

1921? Untitled note in: F. E. Blaisdell, The sixty-second regular meeting of
the Society ... December 9, 1916. Proceedings of the Pacific Coast
Entomological Society, vol. 1, no. 12, 3 un-numbered pp.

[The dates of publication of the parts of vol. 1 of the Proceedings
are very poorly documented, as has been explained by MacSwain,
1951, pp. 105-109. However, it is clear from Blaisdell’s comments
in the Minutes of the 75th and 79th meetings that vol. 1, nos. 1, 2
and 6-15 had been distributed locally, prior to being gathered and
mailed to selected institutions and societies in early February,
1921 (see MacSwain’s Table I). But 1921 is the only known
quotable date. ]

1937. The Lepturini of America north of Mexico. Part II. National Museum
of Canada, bull. 85, Biological Series no. 22, pp. [4+] 42, incl. pls.
I-VI.

INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE
1961. International Code of Zoological Nomenclature adopted by the XV In-
ternational Congress of Zoology. London, International Trust for Zoo-
logical Nomenclature, pp. xvii + 176.

JAQUES, H. E.

1951. How to know the beetles. Pictured-Keys for identifying many of the
beetles which are most frequently seen, with aids for their study and
with other helpful features. Dubuque, Iowa; Wm. C. Erown Co., pp.
[6 +] 372, 865 text figs.

JEANNEL, R.
1955. L’édéage. Initiation aux recherches sur la systématique des Coléoptéres.
Publications du Muséum National d’Histoire Naturelle, no. 16, pp. 1-155,
94 text figs.

JEANNEL, R., et R. PAULIAN
1944. Morphologie abdominale des Coléoptéres et systématique de lOrdre.
Revue francaise d’Entomologie, tome 11, fase. 2, pp. 65-110, 131 text
figs.

Kirsy, W., and W. SPENCE
1823. An introduction to entomology: or elements of the natural history of
insects: with plates. London, Longman, Hurst, Rees, Orme, and
Brown. Vol. II, pp. frontispiece [-++ 4] + 529, 2 col. pls. with explana-
tions. (Pagination from a copy of the 3rd Edition.)

214 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER.

Know ton, G. F., and S. L. Woop

1950. An annotated list of Utah Cerambycidae. Bulletin of the Brooklyn En-
tomological Society, vol. 45, no. 1, pp. 10-13.

KNULL, J. N.

1932. Notes on Coleoptera. — No. 3. [continued.] Entomological News, vol.
43, no. 3, pp. 62-67.

1946. The long-horned beetles of Ohio (Coleoptera: Cerambycidae). Ohio
Biological Survey, vol. 7, no. 4 (=—bull. no. 39), pp. [2+] 133-354,
incl. 29 pls.

1962. (Letter of February 22 to H. B. Leech.)

LACORDAIRE, J. T.

1869. Histoire naturelle des insectes. Genera des Coléoptéres ou exposé
méthodique et critique de tous les genres proposés jusqu’ici dans cet
ordre d’insectes. Volume 8, Les familles des Tricténotomides et des
Longicornes. Vol. 8, pp. [4 +] 552.

LANE, F.
1961. (Letter of February to H. B. Leech.)
1962. (Letter of April 15, to H. B. Leech.)

LEConrTE, J. L.

1850. An attempt to classify the longicorn Coleoptera of the part of America
north of Mexico. Journal of the Academy of Natural Sciences of Phila-
delphia, ser. 2 (N. S.), vol. 1, pt. 4, pp. 311-340.

[According to the table of contents for vol. 1 N. S., part 4 was
published in January, 1850. LeConte’s paper was continued in vol.
2, pt. 1, pp. 5-38; in pt. 2, pp. 99-112 and 139-178. The table of
contents for vol. 2 N. S. shows that pt. 1 was issued in November,
1850, but a footnote on the table of contents states “Extra copies
printed for the Author, March, 1850.” Part 2 was published in
January, 1852, and the footnotes show that extra copies of pp.
99-112 were made for LeConte in October, 1851, and of pp. 139-178
in January, 1852. Presuming that LeConte distributed these pre-
prints as received, it is possible that the species in pp. 5-38 should
date from March, 1850, and those in pp. 99-112 from October, 1851,
not January 1852.]

1857. Report upon insects collected on the survey. Pp. 1-72, pls. I-II. Wash-
ington, D. C. (A preprint of his 1860 paper, q.v.)

1860. Report upon insects collected on the survey. Jn: Reports of explorations
and surveys, to ascertain the most practicable and economical route
for a railroad from the Mississippi River to the Pacific Ocean. Vol.
XII, bk. II, pt. III, Zoological Report, pp. 1-72, pls. I-II. Washing-
ton, D: C.

[The author’s name is incorrectly given as John T. LeConte.]

1862. Note on the classification of Cerambycide, with descriptions of new
species. Proceedings of the Academy of Natural Sciences of Phila-
delphia, 1862, pp. 38-43.

VoL. XXXIT] LEECH: CENTRODERA SPURCA AND ALLIES 215

LECoNTE, J. L.—Cont.
1869. List of Coleoptera collected in Vancouver’s Island by Henry and Joseph
Matthews, with descriptions of some new species. Annals and Maga-
zine of Natural History, ser. 4, vol. 4, no. 24, pp. 369-385.
1873. Classification of the Coleoptera of North America. Prepared for the

Smithsonian Institution. Part II. Smithsonian Miscellaneous Col-
lections, no. 265, pp. 279-348.

LeConte, J. L., and G. H. Horn

1883. Classification of the Coleoptera of North America. Prepared for the Smith-
sonian Institution. Smithsonian Miscellaneous Collections, no. 507,
pp. xxxviii + 567, + intercalary p. 7a; 1 numbered and 4 un-numbered
text figs.

[The lower half of p. 535, and pp. 536-552 are a list of “References
for specific identification,” and are by Samuel Henshaw.]

LENG, C. W.

1890. Synopses of Cerambycide. (Continued). lLepturini. Entomologica
Americana, vol. 6, no. 4, pp. 65-69, ++ Errata on a slip following p. 80;
no. 5, pp. 97-98.

1920. Catalogue of the Coleoptera of America, north of Mexico. Mount Vernon,
N.Y., John D. Sherman, Jr. Pp. viii + 470, folding chart, and p. of
Errata.

1928. Order Coleoptera. In: M. D. Leonard et al., A list of the insects of New
York with a list of the spiders and certain other allied groups. Cornell
University Agricultural Experiment Station Memoir 101. Pp. 1-1121,
map in pocket. (Coleoptera, pp. 203-520.)

LENG, C. W., and W. T. DAvis

1924. List of the Coleoptera of Staten Island, New York. Proceedings of the
Staten Island Institute of Arts and Sciences, vol. 2, part 1 (Special
Issue), pp. 1-82.
[Pages 83-89 constitute an index, prepared by P. Dowell; p. 90 is
titled “Corrections,” and is probably by Leng and Davis. These
pages 83-90 appear to have been issued separately from, and
presumably later than, the main list.]

LinprotH, C. H.

1957. The principal terms used for male and female genitalia in Coleoptera.
Opuscula Entomologica, Bd. 22, no. 2-8, pp. 241-256.

LinprotH, C. H., and E. PALMEN

1956. Coleoptera. In: Taxonomist’s glossary of genitalia in insects, Edited by
S. L. Tuxen. Copenhagen, Ejnar Munksgaard, pp. 284, 215 figs. (Lind-
roth and Palmén, pp. 69-76, figs. 73-82).

LINSLEY, E. G.

1942. Coleoptera: Cerambycidae. Jn: Contributions toward a knowledge of
the insect fauna of Lower California. No. 2. Proceedings of the Cali-
fornia Academy of Sciences, ser. 4, vol. 24, no. 2, pp. 21—96, pls. 4—5.

1961. The Cerambycidae of North America. Part I. Introduction. University
of California Publications in Entomology, vol. 18, pp. [4+] 97, 16
text figs., pls. 1-35.

216 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

LInsLEY, E. G.—Cont.
1962. The Cerambycidae of North America. Part II. Taxonomy and classifica-

tion of the Parandrinae, Prioninae, Spondylinae, and Aseminae. Loc.
cit., vol. 19, pp. vi + 1-102, 34 figs., 1 pl.

[Between p. 102 and the plate there is an un-numbered page titled
“Additions and Corrections to Part I.’’]

MacSwalIn, J. W.
1951. The Proceedings. In: Essig et al., The history of the Pacific Coast En-
tomological Society. Pan-Pacific Entomologist, vol. 27, no. 3, pp. 105—

109, 1 table.

MANK, EpirH W.
1934. The Coleoptera of Glacier Park, Montana. Canadian Entomologist, vol.

66, no. 4, pp. 73-81.

MAZKHIN-PORSHNYAKOY, G. A.

1960. Why insects fly to light by night. (Original in Entomologischeskoe
Obozrenie, vol. 39, no. 1, pp. 52-58; January—March 1960, issue. Trans-
lated from the Russian and published in Entomological Review [Wash-
ington, D. C.], with the above title, vol. 39, no. 1, pp. 32-36; July—
September 1960, issue.)

MraLt, L. C.
1903. The natural history of aquatic insects. London, Macmillan and Co., pp.

xii + 395, 116 figs.

MICHELSEN, A.
1958. Rhagiumarternes udbredelse i Danmark og _ parrings-biologien hos
Rhagium mordax De. G. (Ceramb.) (with an English Summary.)
Entomologiske Meddelelser, vol. 28, pp. 338-351, 9 figs

MICHENER, C. D.
1956. Hymenoptera. Jn: Taxonomist’s glossary of genitalia in insects, Edited
by S. L. Tuxen. Copenhagen, Ejnar Munksgaard, pp. 284, 215 figs.
(Michener, pp. 131-140, figs. 153-164.)

Moore, I.
1937. A list of the beetles of San Diego County, California. Occasional Papers,
San Diego Society of Natural History, no. 2, pp. 109 [+ 3].

Morris, F. J. A.
1916. Reports on insects of the year. Division No. 5, Port Hope District. 46th
Annual Report of the Entomological Society of Ontario, 1915, pp.
17-21.

PEYERIMHOFF, P. DE
1926. Notes sur la Biologie de quelques coléoptéres phytophages du Nord
Africain (quatriéme serie) avec les descriptions de quatre espéces
nouvelles et de quatre sous-espéces. Annales de la Société entomo-
logique de France, vol. 95, trim. 3 & 4. pp. 319-390.

ProcTer, W.
1946. Biological survey of the Mount Desert Region Incorporated. Part VII
being a revision of Parts I and VI with the addition of 1100 species.
The insect fauna, with references to method of capture, food plants,

VoL. XXXITJ LEECH: CENTRODERA SPURCA AND ALLIES 217

the flora and other biological features. Philadelphia, The Wistar In-
stitute of Anatomy and Biology, pp. 566, incl. folding map, 10 un-
numbered text figs.

PROVANCHER, L.

1877. Petite faune entomologique du Canada precedée d’un traite elementaire
d’entomologie. Volume I—Les Coleopteres. Québec, C. Darveau, pp.
786, 52 text figs.

SAALAS, U.

1936. Uber das Fliigelgeader und die phylogenetische Entwicklung der Ceram-
byciden. Annales Zoologici Societatis Zoologice—Botanice Fennice
Vanamo, vol. 4, no. 1, pp. 1-198, 28 text figs., pls. I-XVI, + large folded
pl. XVII.

SHARP, D., and F. Murr

1912. The comparative anatomy of the male genital tube in Coleoptera. Trans-
actions of the Entomological Society of London, 1912, pt. 3, pp. 477—
642, pls. XLII-LXXVIII, with facing pp. of explanations.

SMITH, J. B.

1900. Insects of New Jersey. A list of the species occurring in New Jersey,
with notes on those of economic importance. Supplement to the
Twenty-Seventh Annual Report of the State Board of Agriculture,
Trenton, 1899. (Order Coleoptera, pp. 167-367, figs. 78-156.)

1910. A report of the insects of New Jersey. Jn: Annual Report of the New
Jersey State Museum. Trenton, MacCrellish and Quigley. (Order
Coleoptera, pp. 195-406, figs. 85-170.)

SmirH, G. S.
1929. Coleoptera [a check list of the species collected at Copper Mountain,

British Columbia]. Museum and Art Notes [Vancouver, B. C.], vol. 4,
no. 2, pp. 69-74.

1930. Coleoptera. Part II. Museum and Art Notes, vol. 5, no. 1, pp. 22-25.
1948. (Letter of September 7, to H. B. Leech.)
1954. (Letter of June 15, to H. B. Leech.)

SNODGRASS, R. E.
1957. A revised interpretation of the external reproductive organs of male
insects. Smithsonian Miscellaneous Collections, vol. 135, no. 6, pp.
iv + 60, 15 figs.
SPANGLER, P. J.
1962. (Letter of February 23, to H. B. Leech.)

SwAINE, J. M., and R. Hoppine
1928. The Lepturini of America north of Mexico. Part I. National Museum of
Canada, bull. no. 52 (Biological Series no. 14), pp. [4+] 97, incl.
pls. I-XIII.

THOMSON, J.
1864. Systema cerambycidarum ou exposé de tous les genres compris dans la
famille des cérambycides et familles limitrophes. Mémoires de la
Société Royale des Sciences de Liége, vol. 19, pp. 1-540.

218 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4711 SER.

VAN DYKE, E. C.
1927. New species and subspecies of west American Cerambycide (Coleoptera).
Pan-Pacific Entomologist, vol. 3, no. 3, pp. 99-109.

VILLIERS, A.
1946. Coléoptéres cérambycides de l'Afrique du Nord. Faune de 1’Empire
Francais, vol. 5, pp. [4 +] 1538, 275 figs.

WALKER, F.
1866. Appendix. Jn: John K. Lord, The Naturalist in Vancouver Island and
British Columbia. London, Richard Bentley. Vol. 2, pp. 375, illus-
trated. (Walker’s Appendix = pp. 289-375.)

WICKHAM, H. F.
1897A. The Coleoptera of Canada. XXII. The Cerambycide of Ontario and
Quebec. Canadian Entomologist, vol. 29, no. 4, pp. 81-88, text figs.
12-14.
1897B. The Coleoptera of Canada. XXV. The Cerambycide of Ontario and
Quebec. Canadian Entomologist, vol. 29, no. 7, pp. 169-173, 2 text figs.

Woop, S. L.

1953. Observations on the homologies of the copulatory apparatus in male
Coleoptera. Annals of the Entomological Society of America, vol. 45,
no. 4, pp. 613-617, incl. 2 pls. (Issue for December 1952, actually pub-
lished January 30, 1953.)

WoopwortH, C. W.
1913. Guide to California insects. Berkeley, The Law Press, pp. vi + 360, 361
[+ 1] text figs.
WRIGHT, JULIA D. E., and K. R. CooLipGEe
1908. Notes on the Coleoptera of Placer County, Calif. Entomological News,
vol. 19, no. 2, pp. 66—69.

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

FOURTH SERIES

G Dallas Hanna Anniversary Volume

Vol. XXXII, No. 8, pp. 219-289, 4 figs. May 20, 1963

CONTRIBUTION TO THE BIOGEOGRAPHY
OF COCOS ISLAND, INCLUDING A
BIBLIOGRAPHY
by

Leo George Hertlein
Curator of Invertebrate Paleontology

California Academy of Sciences

CONTENTS
_/MUPE CITRIC? eee lteee ht anh Ae Meee a RC EI Cts Oe ee 220
Pee CRITINETRISE AY ee Art iy eS) Te oe ATS eee ead 220
WS ETL EIR EVSTTN OER or eg ae a RC gd en ee Rar en ae ee 224
UDA TEE 2 PED et, ee oe Sell See eee eR Ie Pl cole Ma a ae oe 229
Nn PE tee Cee ets ee a. 2 ae Oe ee 229
> TILT, EA OUCY AC ye Mn eel iy yO oe SSE rE ne dak AOR i 229
Punminm OClemnteratae s:: ssa fe she s oats akon ene heed oes 230
Pees a OZ Od: Fee fat ee te Oe tone hs Wom cke enn We, Wa Se A oe 202
Rebtel eM ies EMO OOS p< ¢c08 cc Ske wha Bed, 0 ane UR aae SR I ae AeA Dale 234
Poginmetenimodermata ) 2. 20s ll ote ssetees © wlye Gnade ond 234
L?ncwellniiy MG FTIS(C IR eae rte Ros, ee eke Ce a zr 237
LE nev mien WEST 6c aa eet oe ee ge oe 244
LELSVI ninety ZS 0) 61016 (2 eC, ok en en ee ee ee ee 245
Pll nit (CUNOTG ENG ae Ne oO me eg ee 252,
[2194

Tie
4 Marine Biological Laboratory
LIBRARY
WAY 29 1983
WOODS HOLE, MASS.

220 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 47TH SER.

CoNTENTS—Cont.

ROUAIY ss cca ce cs wa aves ob es See ee ne ee 254
Biblioeraphiy 0... 2. oe lee mela eee ee oe oe 254
ILLUSTRATIONS
Figures:
1. Cocos Island, at mouth of Arroyo del Genio................ 222
2. Cocos Island, just south of Chatham Bay. ...-... -2. sae 224

3. Map showing position of Cocos Island and the course of the
equatorial countercurrent during August-September of a nor-
Mal Wear «sa c% oi aye ewan Medel Gale Mae 227
4. Map showing position of Cocos Island and the course of the
current westward from along Central America during 1891, a
year of extreme displacement southward of the currents..... 228

INTRODUCTION

The author’s interest in Cocos Island was stimulated by an opportunity
to collect mollusks on the island in 1932, during the return of an expedition
to the Galapagos Islands on G. Allan Haneock’s motor cruiser Velero IIT.
The results of a study of these mollusks are contained in two papers, one on
marine species (Hertlein, 1932) and one dealing with non-marine and
brackish-water species (Hanna and Hertlein, 1938).

The present paper presents information concerning the species of mol-
lusks which I collected, including those reported on in various publications,
and summarizes what is known about the relationships and the zoogeographi-
eal significance of this assemblage.

In addition to this, during the course of this study, information was
accumulated concerning the occurrence and distribution of representatives
of other phyla of organisms reported from Cocos Island. This information
is included under each major category along with the names of authors and
dates of pertinent references, which are included in the bibliography. How-
ever, no attempt has been made to inelude all the phyla or all the species
reported from Cocos and seattered throughout the literature. I believe, how-
ever, that the biota here mentioned is representative of the island.

ACKNOWLEDGMENTS

The author gratefully acknowledges the aid of several persons without
whose cooperation this paper would not have appeared in its present form.
Dr. Alan E. Leviton, Department of Herpetology, California Academy of
Sciences, gave helpful criticism of the manuscript. In the same institution,
Mr. Hugh B. Leech and Dr. C. Don MaeNeill, Department of Entomology,

VoL. XXXII] HHRTLEIN: BIOGEOGRAPHY OF COCOS ISLAND 221

contributed information concerning the insects of Cocos Island. Dr. Franeis
X. Williams, Chula Vista, California, also furnished information coneern-
ing the insects which he observed on Cocos Island. Dr. Robert T. Orr, De-
partment of Ornithology and Mammalogy, California Academy of Sciences,
and Dr. Paul Sludd, Department of Natural Sciences, University of Florida,
aided with information on the birds of this island. Dr. Elizabeth MeClin-
tock, Department of Botany, California Academy of Sciences, gave advice
concerning the portion of this paper dealing with the botany; and Dr. John
C. Briggs, Department of Zoology, University of British Columbia, con-
tributed information concerning the fishes of Cocos Island. Dr. Robert
Robertson, Department of Mollusks, Academy of Natural Sciences of Phila-
delphia, cooperated by loaning specimens of some of the species which were
reported from Cocos Island by Pilsbry and Vanatta.

Several members of the Allan Hancock Foundation, University of South-
ern California, generously contributed their time to furnish information
and to read those portions of the manuscript dealing with their special field
of study. To these I express my sincere appreciation: Dr. John S. Garth
and Miss Janet Haig, Crustacea; Dr. John D. Soule, Bryozoa; Dr. Olea
Hartman, Annelida; Mr. Fred Ziesenhenne, Echinodermata.

Miss Veronica Sexton, Librarian, California Academy of Sciences, aided
in making available needed literature, some of which was lent by the Uni-
versity of California, or consulted in the Baneroft Library, University of
Washington, University of Missouri, University of Chicago, San Francisco
Public Library and Seattle Public Library.

Mr. William Old, Jr., American Museum of Natural History, New York
City, kindly checked certain references in the libraries available to him.
Mr. Robert I. Nesmith, Curator, Foul Anchor Archives, Rye, New York,
furnished information concerning manuscripts and published literature con-
cerning Cocos Island. Mr. William A. Coolidge, Cambridge, Massachusetts,
generously lent me a copy of a rare paper dealing with an expedition to
Cocos Island by his brother, Amory Coolidge. Dr. Bruce Halstead, World
Life Research Institute, Colton, California, and Mr. B. Joseph O’Neil, Bos-
ton Puble Library, also furnished useful information.

Photographs of the Island were made available by Dr. John S. Garth,
Allan Hancock Foundation, and Mr. Don Ollis, Santa Barbara, California.

GENERAL REMARKS

Cocos Island les at 5° 32’ 57” North Latitude and 86° 59’ 17” West
Longitude, about 500 kilometers (300 miles) west of Costa Rica and about
630 kilometers (350 miles) northeast of the Galapagos Islands. This island
was known to mariners and cartographers at least as early as the first half

CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

bo
bo
bo

of the sixteenth century for it was first shown as “Y® de Coques” on a map
by Nicholas Desliens, in 1541 (Anonymous, 1920, p. 15).

The name of this island appeared on many maps and in various publi-
cations during the following centuries. The abundance of fresh water, wood,
pigs, sea fowl, fish, and coconuts, and the ease with which they could be
obtained, made this small island a favorite stopping place of pirates, priva-
teers, and whaling vessels (Colnett, 1798, p. 73, mentioned placing 2000
coconuts on board the Rattler when he stopped there in 1793). The names
of various ships (see Hancock and Weston, 1960, pp. 300-3502) carved in
the rocks at Chatham Bay are reminders of these early visitors. Two bays
offer anchorage for ships. Wafer Bay is the more attractive for visitors but

Fic. 1. Wafer Bay, Cocos Island, at the mouth of Arroyo del Genio. View show-
ing sandy beach, dense vegetation, and a small house used by treasure hunters.
(Photograph by John Garth, 1931.)

Chatham Bay, on the northeast side of the island, is more sheltered from
the prevailing winds and offers more secure anchorage. In 1936, the gov-
ernment of Costa Rica issued stamps of several denominations on which
an outline map of the island appears.

Politically, Cocos Island belongs to Costa Rica which country exercised
sovereignty over it when, in 1888,1 August Gissler was nominated Governor
of the island (see Anonymous, 1920, p. 23) with a concession to search for
treasure. The circumference of Cocos is about 23.3 kilometers (13 nautical
miles or 14.6 statute miles), and the land area comprises approximately 46.6
square kilometers. The highest point, located in the western portion of the

1. Molina, 1851 (p. 27), mentioned that Cocos Island belonged to Costa Rica.

Vou. XXXII] AHERTLEIN: BIOGEOGRAPHY OF COCOS ISLAND 223

island, is reported to be 849.8 meters (2,788 feet) above sea level (see
Anonymous, 1935, U.S. Hydrographic Office chart 823, South Pacifie sheet
1; Anonymous, 1945, U.S. Hydrographie Office chart 1685, ed. 9).2, Much
of the coast is abrupt, and in places towering cliffs reach a height of 182.88
meters (600 feet). Chubb (1933, p. 27) remarked on these high cliffs and
stated: ‘It is clear that when the cliffs were being formed the island stood
some two or three hundred feet higher above sea level than it does now, but
that since then either it has subsided or the sea level has risen, with the
result that the lower part of the cliffs is submerged and the lower reaches
of the mature river valleys are flooded.” He also remarked that at some
places a shore-shelf a few feet above sea level is evidence of post-Pleistocene
fall in the sea level.

The island is composed of voleanic agglomerates, tuffs, and chiefly of
lava flows of labradorite-andesite and hornblende labradorite-andesite
(Chubb, 1933, p. 29). It is situated upon the southwestwardly trending
Cocos Ridge (Shumway, 1954) which in general les about 1829 meters
(1000 fathoms) below sea level. Chubb (1933, p. 30), coneluded from a
study of the rocks that “The Cocos Island analyses |of andesite] compare
fairly closely with those of the Galapagos basalts, but their alumina and
potash content is higher, and their lime content is lower. Except for their
abnormally high alumina content they compare better with labradorite-
andesites from Rapa and Tahiti.”” Macdonald (1949, p. 1588), however,
believed that knowledge of the composition of the rocks on Cocos Island
is not sufficient to justify placing it in the mid-Pacifie petrographie province.

Sedimentary rocks have been mentioned |Pittier, 1899, p. 144; Cha-
varria Mora (in Pittier), p. 158; Thomas, 1960, p. 34] as occurring on the
island but if present there, the details are vague.

Cocos Island, in general, lies within the path of the eastward flowing
equatorial counter current. Occasionally, however, southward shifting of
eurrents brings the island under the influence of currents sweeping west-
ward from the mainland (see figures 3 and 4). The temperature of the air
ranges between 20°C. (68°F.) and 33.3°C. (92°F.) (Stewart, 1912, p. 378),
and the months of greatest rainfall are reported to be May, June, and July,
although there is heavy rainfall throughout the year. The surface tempera-
ture of the sea water is reported to be about 26.7°C. (80°F.) but there is a
mean annual variation of about 5°C. (10°F .).

This beautiful and picturesque island is densely covered with vegetation
in which palms and ceecropia trees are conspicuous. Abundant rainfall gives
rise to numerous waterfalls which plunge over steep cliffs. Flocks of sea
birds (terns, noddies, Frigate-birds, and boobies), flying about the bays,
and others perched in the trees, add to the attractiveness of the scene.

2. According to Murphy (1936, p. 317), “The central hills rise to an altitude of 518 meters, which is
much less than is commonly credited to them.”

224 CALIFORNIA. ACADEMY OF SCIENCES [ Proc. 4TH SER.

Fig. 2. East side of Cocos Island, just south of Chatham Bay. View showing cliff,
waterfall, and dense vegetation. (Photograph by Don Ollis, December 26, 1952.)

Any attempt to penetrate inland, however, is accomplished with difficulty
because of the rough and irregular land surface which is covered by dense
rain forest. Attempts to follow the bed of the stream emptying into
Chatham Bay or the one in Arroyo del Genio (also cited as Canyon del
Infierno on some maps) at Wafer Bay, is fraught with difficulty. The rocks
in the stream and along the banks are very slick, and the stream bed is
frequently interrupted by cliffs forcing the traveler to make a torturous
detour. The presence of a particular fly, Lewcomelina pica, and a species
of ant, Wasmanma auropunctata, also add to the discomfort of such an
adventure. One shipwrecked adventurer (Palliser, Brawner and Stachwick,
1932, p. 134) described his attempt to eross the high ridge between Wafer
Bay and Chatham Bay, a distance over water, by boat, of about 1.6 kilo-
meters (1 mile). He lost his way in the dense jungle growth of trees, vines,
tall sharp-edged grass, was drenched by frequent torrential rains, and after
other harrowing experiences, found his way back to Wafer Bay only after

Vom. XXXII] HERTLEIN: BIOGEOGRAPHY OF COCOS ISLAND 225

several days had elapsed. Travel over the plateau-like top of the island
is reported to be less difficult.

Cocos Island is best known from traditional accounts of treasure reputed
to be buried there about 1819-1820, the value variously estimated at 60 to
100 million dollars. A large portion of this is supposed to have been placed
there by a pirate, Benito Bonito (alias Bennett Graham) of the Relampago,
who looted cities and ships along the coast of South America. Tradition: has
it that additional treasure was buried on the island by Captain William
Thompson.

According to this latter story, a revolution in Peru, in 1820, led authori-
ties in Lima to entrust for safe-keeping to Captain Thompson of the Mary
Dear the “treasure of Lima,” gold and jewels said to be valued at many
millions of dollars. The vessel made its way to Cocos Island where the
treasure was reputedly buried at Wafer Bay. The subsequent search with
expenditure of much time, energy, and money has led to over 400 expedi-
tions (Riesenberg, 1951, p. 5) of which at least 25 were well-equipped major
expeditions. Captain August Gissler spent nearly 18 years searching for
the treasure.

Although it has been rumored from time to time that these treasure-
troves have been found, so far as is definitely substantiated, no one has
found them, other than an occasional coin the source of which is unknown.
A vast amount of literature has grown up based upon these fascinating
legends of treasures and the search for them. Many references to this sub-
ject are contained in the bibliography.

For additional general information concerning Cocos Island see the
following: Anonymous (1920; 1939b; 1945; 1951) ; Beebe (1926) ; Belcher
(1848); Campbell (1932; 1934); Collenette (1926); Dampier (1729);
Fraser (1943); Gueydon (1948); Hancock and Weston (1960); Liévre
(1893) ; Nesmith (1958); Pittier (1899); Rogers (1931); Rose (1926) ;
Schmitt (1939a) ; Slevin (1931) ; Snodgrass and Heller (1902) ; Vancouver,
1798; Wafer [1699 (1903) ].

BIOGEOGRAPHY

An excellent summary of the zoogeography of the vertebrate fauna of
Cocos Island was published by Schmidt (1930). The general consensus of
most authors who have given careful consideration to this subject is that the
island received its fauna and flora by. transport over open water and that
the endemic species have arisen because of geographic isolation.

The vertebrate fauna of Cocos Island consists of two lizards and seven
land birds. Townsend reported a snake (not identified as to genus or
species) from this island, but this record has not been substantiated by any
subsequent collection from there. The lizards and three endemic land birds

226 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

differ from any other species, but their closest relatives live in Central
America. There are no native mammals. A rat, Rattus norvegicus, is re-
ported to have reached the island, undoubtedly from some of the many ships
which have anchored there or from an occasional shipwreck. Pigs have been
introduced, and evidence of their destructive influence is noticeable at
many places. There are also reports of other domestic animals including
cats, goats, deer, chickens, and even monkeys (Thomas, 1960, p. 34), intro-
duced on the island intentionally or otherwise, and many of these appear
to have become established as permanent members of the animal community.

The composition of the known invertebrate land and marine fauna is
predominantly eastern Pacific in affinities. The evidence as to the origin of
the sparse land snail fauna may be considered equivocal. Some authors
consider their affinity to be with Indo-Pacific forms, others consider the
relationship to be with American species. It is probable that some elements
of the land snail fauna were derived from Central America, and some from
Polynesia.

The most comprehensive study of the land plants of Cocos Island is by
Stewart (1912). He reported that eight species, 8.69 per cent of 77 vascular
plants, are endemic. Twenty-seven species also occur on the Galapagos
Islands where the endemicity was reported to be 40.9 per cent. Stewart
(1912, pp. 381-383) believed that the flora is that of an oceanic island, of
more recent origin than that of the Galapagos Islands, and that its flora was
derived chiefly from the mainland by chance agencies such as winds, ocean
currents, and migratory birds. He, furthermore, concluded that there is no
evidence to indicate that the island ever was connected by land with the
mainland or with the Galapagos Islands.

Svenson (1935, p. 259) following Johnston (1931, p. 35), mentioned
that the flora of Cocos Island consists of about 100 species of which about
10 are endemic, consisting overwhelmingly of ferns, melanostomes, and
orchids. He stated that with the exception of the ferns, virtually none of
these species occur in the Galapagos Islands.

More recently Vinton (1951), in discussing a possible explanation for
the derivation of the Galapagos fauna and flora, suggested a peninsula ex-
tending southwest from Costa Riea to within a hundred miles of the
Galapagos Islands. He postulated that such a land area existed during
Miocene time, and included the locality now occupied by Cocos Island (see
his fig. 1). It was also his belief that such a land area would have deflected
the currents at that time when an open seaway existed across Panama. He
remarked on the recent origin of Cocos Island and suggested a probable
Pleistocene age for it. He agreed in general with Stewart’s opinion concern-
ine derivation of the fauna and flora.

Schott (1931) published an excellent diseussion concerning the shifting
of oceanic currents, north and south, in the Panamie and northwestern

VoL. XXXII] HEHERTLEIN: BIOGEOGRAPHY OF COCOS ISLAND 227

Fic. 3. Copy of illustration by Schott (1931, plate 20, figure 1) showing position
of Cocos Island and the course of the equatorial countercurrent during August—
September of a normal year.

South American region. He reported that this shift takes place about every
seven years, and a decided displacement about every thirty-four years. The
effect which this phenomenon produced upon the climatic and biological
features in this region was described by Murphy (1936, pp. 102-108). Later
Garth (1946) discussed the effect of this shifting of oceanic currents upon
the composition of the brachyuran (crab) fauna of the Galapagos Islands.
This same shifting of currents whereby Cocos Island is at times in the path
of the Equatorial Countercurrent, and at other times under the influence of
westwardly directed currents from off the mainland, can be invoked to
explain the means of transportation by which the island received most of
its invertebrate marine fauna.

The percentage of Indo-Pacific species in the invertebrate fauna of
Cocos Island is small, except in the assemblage of corals and in those groups
in which the species have an exceptionally wide distribution, such as the
Holothurioidea. These species from the western Pacific may have reached
the island by the agency of transpacifie currents, directly, or conceivably
by progressing eastward from one atoll to another or to truncated submarine
islets (guyots) (see Ladd, 1960, pp. 143-145) which are known to exist in
some places in the eastern Pacifie.

228 CALIFORNIA ACADEMY OF SCIENCES [Proc. 471 Ser.

p Guayaquil
7

obitos

Fic. 4. Copy of illustration by Schott (1931, plate 21, figure 3) showing position
of Cocos Island and the course of the current westward from along the American
mainland of Central America during March, 1891, a year of extreme displacement
southward of the currents.

Very few species of invertebrates in the marine fauna of Cocos Island
are identical with Caribbean or Atlantic forms but many elosely related
species occur in that region.

The following tabular summary indicating the distribution of the species
of a few biologic categories (mostly those represented by a number of
species), although based on lists which undoubtedly do not inelude all the
species which may exist on the island, reveals the general interregional
relationship of these groups. Additional collecting, especially in this and
other tropical eastern Pacific islands where corals occur abundantly, may
result in changes in the pereentages of marine species shown in the various
regions. I believe, however, that the general affinities of the fauna are re-
vealed in this table. Additional remarks on the distribution and affinities
of the phyla accompany the lists of species included under their respective
categories in the following pages.

Krom the evidence of known distribution of the genera and species, I
am inclined to agree with Stewart, Schmidt, and Vinton that the biota of
Cocos Island was derived chiefly from the mainland, by the chance agencies
of wind, ocean currents, birds, or pelagic mammals.

Vou. XXXII] HERTLEIN: BIOGEOGRAPHY OF COCOS ISLAND 229

TABLE 1. Number and general distribution of species of particular groups of
animals reported from Cocos Island.

Number | Endemic | Mainland and Galapacos|) Indo—1" Garinvean:

os ao SEMEN Islands | Pacific Atlantic
Species | Island Region
Foraminifera iz 14 13 1 Mediter-
ranean
Coelenterata
Hydrozoa 5 5 3 1
Anthozoa
(corals) 18 1 12 10 10 il
Bryozoa 20 18 20 1 ze
Echinodermata
Asteroidea 6 4 5
Ophiurioidea 15 15 12 1
Echinoidea 1133 1 12 10 1
Holothurioidea 11 11 6 8 2
Mollusca
Marine and Many related
Brackish- Water 88 5 74 38 5 species
Land snails 9 a Lor 2
Annelida 9 4 4 i 1
Arthropoda
(Crustacea )
Brachyura 33 30 23 3 2
Anomura iL7 2 11 10 (or 11) 4
Myriopoda 7 3 4
Pisces 59 5 38 20 16
Reptilia
(lizards ) 7 2 Related
species
Aves
(land birds) q 3 4
ZOOLOGY

Phylum PROTOZOA
Class SARCODINA
Order Foraminifera

Ammodiscus pacificus Cushman and Valentine. Southern California to
Panama; Galapagos Islands.

230 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER.

Dentalina filiformis @Orbigny. Mainland; Galapagos Islands.

Dentalina ef. D. jugosa Williamson. Galapagos Islands. Monaco.

Eggerella advena Cushman. Cordova, Alaska, to Peru.

Haplophragmoides hancocki Cushman and McCulloch. Ketchikan, Alaska,
to Peru; Galapagos Islands.

Lagena striatopunctata var. excentricitas Cushman and McCulloch. Galapa-
gos Islands.

Nouria polymorphinoides Heron-Allen and Earland. West Mexico to South

America; Galapagos Islands.

Reophax agglutinatus Cushman. Mainland; Galapagos Islands.

Reophax excentricus Cushman. Mainland; Galapagos Islands.

Textularia articulata d’Orbigny. Southern California to Sechura Bay, Peru.
Textularia conica @Orbigny. Gulf of California to Colombia; Galapagos

Islands.

Textularia corrugata Heron-Allen and Earland. Gulf of California to EKeua-
dor; Galapagos Islands.

Textularia panamensis Cushman. Gulf of California to Peru.

Textularia schencki Cushman and Valentine. Southern California to Cen-
tral America; Galapagos Islands.

Trochammina charlottensis Cushman. San Franeiseo, California, to Co-
lombia; Galapagos Islands.

Trochammina ntida H. B. Brady. Cordova, Alaska, to Sechura, Peru.

Vaginulina exilis Cushman and McCulloch. Galapagos Islands.

These 17 species of foraminifera were reported from Cocos Island by
Cushman and McCulloch (1939; 1950) and Lalicker and MeCulloech (1940).
Fourteen of these also occur on the mainland and 13 are reported from the
Galapagos Islands.

Phylum COELENTERATA
Class Hyprozoa

The following species were reported from Cocos Island by Fraser (1938;
1948).
Clytia cylindrica A. Agassiz. Colombia to Peru; Galapagos Islands.
Gonothyraea gracilis Sars. Costa Riea to Peru; Galapagos Islands.
Halecium washingtoni Nutting. California to Peru.
Obelia commissuralis MeCrady. Mexico to Panama; Atlantic.
Thuriaria orisioides Lamouroux. Mexico to Colombia; Galapagos Islands.
These 5 species all live on the mainland coast, and 3 have been reported
from the Galapagos Islands. This class probably is represented on Cocos
Island by more species than shown in the present list assembled from only
a cursory search in papers published by Fraser.

Vou. XXXII] HERTLEIN: BIOGEOGRAPHY OF COCOS ISLAND 231

Class ANTHOZOA
Order Pennatulacea
Family Pennatulidae

Leioptilus undulatus Verrill [Boone (1933, pp. 16, 57, pl. 19) reported
Ptilosarcus gurneyi Gray (not P. gurneyt Gray, 1860), from Cocos Island.
Deichmann (194la, p. 13) later referred this record to Leioptilus
undulatus|. Magdalena Bay, Baja California, to the north end of the
Gulf of California and south to Panama.

Order Scleractinia
(Corals)

The following species have been reported from Cocos Island by Durham
and Barnard (1952) and Durham (1962). These, with their geographic
ranges follow:

Astrangia dentata Verrill (Durham, 1962, pp. 45, 46, 52). La Paz, Baja
California, to Panama.

Astrangia hondaensis Durham and Barnard (1952, p. 73; Durham, 1962,
pp. 44, 46). Questionably from Cocos Island and the Galapagos Islands.

Balanophyllia scheert Durham (1962, pp. 45, 46, 53). Known only from
Cocos Island.

Cladocera debilis Milne Edwards and Haime (Durham and Barnard, 1952,
p. 58; Durham, 1962, pp. 44, 46). Galapagos Islands; Atlantic.

Cycloseris mexicana Durham and Barnard (1952, p. 53; Durham, 1962, pp.
44,46). Gulf of California to La Libertad, Ecuador; Galapagos Islands.

Endopachys vaughani Durham and Barnard (1952, p. 103; Durham, 1962,
pp. 44, 46). Gulf of California; Galapagos Islands.

Leptoseris digitata Vaughan (Durham and Barnard, 1952, p. 36; Durham,
1962, pp. 44, 46). Questionably from Cocos Island; Gorgona Island,
Colombia, to La Plata Island, Eeuador; Hawaiian Islands; Andaman
Islands.

Pavona (Pavona) ef. P. (P.) explanulata Lamarek (Durham and Barnard,
1952, p. 42; Durham, 1962, pp. 44, 46). Clipperton Island; Indo-Pacific.

Pavona (Pavona) varians Verrill (Durham, 1962, pp. 45, 46, 50). Colombia ;
Hawaiian Islands; Great Barrier Reef; Red Sea.

Pavona (Polyastra) ponderosa Gardiner (Durham, 1962, pp. 45, 46, 50).
Bonin Islands; Maldive Islands.

Pocillopora damicornis Linnaeus (Durham, 1962, pp. 45, 46, 48). Panama;
ralapagos Islands; to Indian Ocean.

Pocillopora elegans Dana (Durham, 1962, p. 45, 46, 48). Gulf of California
to the Galapagos Islands; Indo-Pacific.

232 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER.

Pocillopora meandrina Dana (Durham, 1962, pp. 45, 46, 48). Gulf of Cali-
fornia; Indo-Pacifie.

Pocillopora verrucosa Ellis and Solander (Durham, 1962, pp. 45, 46, 48).
Gulf of California; Galapagos Islands; Clipperton Island; to Indian
Ocean.

Porites excavata Verrill (Durham, 1962, pp. 45, 46, 51). Panama.

Psammocora profundacella Gardiner (Durham, 1962, pp. 45, 46, 49).
Galapagos Islands; Fanning Island; Funafuti Island.

Psammocora (Stephanaria) stellata Verrill (Durham and Barnard, 1952,
p. 30; Durham, 1962, pp. 44, 45, 46, 49). Gulf of California to La
Libertad, Ecuador; Galapagos Islands; Hawaiian Islands; Fiji.

Tubastrea tenwlamellosa Milne Edwards and Haime (Durham, 1962, pp.
45, 46, 54). Gulf of California to Panama; Galapagos Islands.

Eighteen® species of corals (three identifications doubtful) have been
reported from Cocos Island and of these fourteen are reef-building (herma-
typic) forms. One species is known only from Cocos Island. Twelve species
(one identification doubtful) live along the eastern Pacific mainland or in
the Panamie province and 10 species (one identification doubtful) live in
the Galapagos Islands. Ten species (two identifications doubtful) also live
in Indo-Pacific waters. Only one of the 18 species occurs in Atlantic waters.

Phylum BRYOZOA

The following list of species of Bryozoa from Cocos Island, and their
occurrence elsewhere, was generously furnished by Dr. John D. Soule.
These species are all from Allan Hancock Foundation sample number
328, Chatham Bay, Cocos Island, in 25.6 meters (14 fathoms), February 13,
1938. Three of these species, as indicated in the list, also were represented
in Allan Haneock Foundation sample 330, Chatham Bay, Cocos Island, in
85.95 meters (47 fathoms), February 14, 1938.
A few species of Bryozoa from Cocos Island were mentioned by Osburn
(1950; 1952).
Aplousina filum (Jullien), 1903. Previously reported from the Gulf of
California; Cocos Island; Octavia Rocks, Colombia; Gal4pagos Islands.
Cellarva velerons Osburn, 1950. Originally reported from the Galapagos
Islands. This is the first record of its occurrence from Cocos Island.
Chaperiella condylata (Canu and Bassler), 1930. Common in the Galapagos
Islands. It ranges from southern California to the Galapagos Islands.
No prior record from Cocos Island.

Clerdochasma contracta (Waters), 1899. Previously reported from the

3. According to Dr. J. W. Durham (verbal communication, October 3, 1962), his record (1962, p. 51) of
the occurrence of Porites californica Verrill from Cocos Island, is incorrect.

Vou. XXXII] HHERTLEIN: BIOGEOGRAPHY OF COCOS ISLAND 233

Galapagos Islands and the Gulf of California. No prior record from
Cocos Island.

Colletosia radiata (Moll), 1803. Widely distributed from the cool temper-
ate waters off Oregon to the Galapagos Islands. No prior record from
Cocos Island.

Copidozoum tenwrostre (Hineks), 1880. Known from northern California
to the Galapagos Islands. No prior record from Cocos Island. (Also
present in sample 330.)

Crepidacantha poissoni (Audouin), 1826. Ranges from the Channel Islands,
California, to the Galapagos Islands. No prior record from Cocos Island.

Crepidacantha setigera (Smitt), 1873. Previously reported from Cocos
Island, Galapagos Islands and Florida.

Discoporella umbellata (Defranee), 1823. Common from Point Conception,
California, to Point Santa Elena, Ecuador, including the Gulf of Cali-
fornia, the coast of Costa Rica, and the Galapagos Islands.

Enantiosula manica Canu and Bassler, 1930. Reported from the Gulf of
California, Cocos Island, and the Galapagos Islands. (Also present in
sample 330. )

Holoporella brunnea (Hineks), 1884. Abundantly represented from off
British Columbia to the Galapagos Islands. No prior record from Cocos
Island. (Also present in sample 330.)

Micopora coriacea narmata Soule, 1959. Common in the Gulf of California.
Reported from the Galapagos Islands. No prior record from Cocos
Island.

Microporella ciliata (Pallas), 1766. Common from the coast of Oregon to
the Galapagos Islands. No prior record from Cocos Island.

Microporella marsupiata (Busk), 1860. Known from the Gulf of California
and the Galapagos Islands. No prior record from Cocos Island.

Parellisina curvirostris (Hineks), 1862. World wide in tropical and tem-
perate waters. No prior record from Cocos Island.

Reptadeonella violacea (Johnston), 1847. Previously reported from the
Gulf of California, coast of Mexico, Cocos Island, and Galapagos Islands.

Reteporellina denticulata gracilis Osburn, 1952. Recorded previously from
the Gulf of California, Cocos Island, and the Galapagos Islands.

Retevirgula areolata (Canu and Bassler), 1923. Ranges from southern
California to the Galapagos Islands. Common at the Galapagos Islands.
No prior record from Cocos Island.

Trypostega venusta (Norman), 1864. Previously reported from the Gulf of
California, the coast of Mexico, Cocos Island, and the Galapagos Islands.

Tubulipora fleruosa (Pourtales), 1867. Previously reported from the Gulf
of California and the Galapagos Islands. No prior record from Cocos
Island.

CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

bo
(J)
=

Eighteen of the 20 species in this list live along the mainland in tropical
eastern Pacific waters. All are known to live in the Galapagos Islands, 9
in California waters, 2 in the Caribbean area one of which occurs world-wide
in tropical and temperate waters.

Phylum BRACHIOPODA
Family Rhynchonellidae

Hispanirhynchia? craneana Dall [as Hemithyris craneana Dall, 1895, p.
717; 1920, p. 288; Hertlein and Grant, 1944, p. 60 as “Hispanirhynchia
[?]” craneana].

This species was originally dredged off Cocos Island in 2149 meters

(1,175 fathoms). It has not been reported elsewhere.

Phylum ECHINODERMATA
Class ASTEROIDEA

The following is a list of the species of Asteroidea from Cocos Island
and their occurrence elsewhere. The records of species not previously re
ported from this island were furnished by Mr. Fred Ziesenhenne.
Coronaster marchenus Ziesenhenne (1942, p. 212). Galapagos Islands.
LTinckia columbiae Gray (Ziesenhenne, 1962, written communication). Cali-

fornia to Peru; Galapagos Islands.

Narcissua gracilis A. H. Clark (Ziesenhenne, 1962, written communication).

Gulf of California; Galapagos Islands.

Oreaster occidentalis Verrill (H. L. Clark, 1940, p. 333). Mexico to Peru;

Socorro and Clarion Islands, Revillagigedo Islands; Galapagos Islands.
Pauliella aenigma Ludwig (Ziesenhenne, 1937, p. 215). Gulf of California

to Panama; Clarion Island, Revillagigedo Islands.

Tamaria obstipa Ziesenhenne (1942, p. 209). Galapagos Islands.

Four of the 6 species of this class reported from Cocos Island also live
along the mainland, 5 in the Galapagos Islands, and 2 in the Revillagigedo
Islands.

Class OPHIURIOIDEA

The following brittle stars, with their known ranges, have been reported
from Cocos Island. Records of those not previously reported from this
island were generously furnished by Fred Ziesenhenne.

Amphiodia violacea Liitken (A. H. Clark, 1939, p. 2). Los Coronados
Islands, off west coast of Baja California, Mexico, to Colombia (Ziesen-
henne, 1963, written communication).

Diopederma danianum Verrill (H. L. Clark, 1940, p. 343). Baja California
to Panama.

Vou. XXXII] HERTLEIN: BIOGEOGRAPHY OF COCOS ISLAND 235

Ophiacantha phragma Ziesenhenne (1940, p. 12). Southern California to
Ecuador; Galapagos Islands.

Ophiactis savignyt Miller and Troschel (Ziesenhenne, 1962, written com-
munication). Mainland; Galapagos Islands; Clipperton Island; world-
wide in tropical waters.

Ophiactis simplex LeConte (Ziesenhenne, 1962, written communication).
California to Peru; Galapagos Islands.
Ophiocoma aethiops Liitken (H. L. Clark, 1940, p. 341). Baja California

to Panama; Galapagos Islands.

Ophiocoma alexandri Lyman (H. lL. Clark, 1940, p. 341). Baja California
to Panama; Galapagos Islands.

Ophioderma panamense Liitken (H. L. Clark, 1940, p. 342; Ziesenhenne,
1955, p. 193). San Pedro, California, to Paita, Peru; Guadalupe Island;
Revillagigedo Islands; Galapagos Islands.

Ophioderma variegatum Litken (H. L. Clark, 1940, p. 348; Ziesenhenne,
1955, p. 198). San Diego, California, to Panama; Revillagigedo Islands;
Galapagos Islands.

Ophiomyxa panamensis Liitken and Mortensen (H. L. Clark, 1940, p. 336).
Gulf of California to Panama; Galapagos Islands; Socorro Island,
Revillagigedo Islands.

Ophionereis dictyota Ziesenhenne (1940, p. 30). Gulf of California, to
Panama.

Ophionereis nuda Liitken and Mortensen (H. L. Clark, 1940, p. 340). Isa-
bel Island, Mexico, to Panama; Galapagos Islands.

Ophiophragmus marginatus Liitken (Ziesenhenne, 1962, written communica-
tion). Mexico to Ecuador; Galapagos Islands.

Ophiophragmus paucispinus Nielsen (Ziesenhenne, 1962, written communi-
cation). Mexico to Costa Rica; Galapagos Islands.

Sigsbera lineata Liitken and Mortensen (H. L. Clark, 1940, p. 336).
Panama; Galapagos Islands.

Of the 15 brittle stars reported by H. L. Clark and by Ziesenhenne from

Cocos Island, all also live in mainland waters, 12 in the Galapagos Islands,

3 in the Revillagigedo Islands, Mexico, and 1 in Indo-Pacifie waters.

Class ECHINOIDEA

Hl. L. Clark (1940; 1948) recorded the occurrence of 13 species of
echinoids from Cocos Island. These, with their recorded geographic ranges
follow.

Centrechinus (= Diadema) mexicana A. Agassiz (H. L. Clark, 1948, p.

235). Consag Rock, Gulf of California, to La Plata Island, Ecuador;

Revillagigedo Islands; Galapagos Islands.

236 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Centrocidaris doederleini A. Agassiz (H. L. Clark, 1948, p. 226). Galapagos
Islands.

Clypeaster ochrus H. L. Clark (1947, p. 297). La Paz, Baja California, to
Ecuador; Galapagos Islands.

Clypeaster rotundus A. Agassiz (H. L. Clark, 1948, p. 296). Georges
Island, Gulf of California, to Santa Elena Bay, Ecuador; Revillagigedo
Islands; Galapagos Islands.

Clypeaster speciosus Verrill (H. L. Clark, 1940, p. 350). Magdalena Bay,
Baja California, and the Gulf of California; Revillagigedo Islands;
Galapagos Islands.

Echinometra van-brunti A. Agassiz (H. L. Clark, 1940, p. 349; 1948, p.
294). Angel de la Guardia Island, Gulf of California, to San Francisco
Bay, Ecuador; Revillagigedo Islands; Galapagos Islands.

Encope cocosi H. L. Clark [1948, p. 330; Mortensen (Monogr. Eehin., IV.2.
Clypeastroida, p. 448, 1948, as Encope micropora cocosi) |. Known only
from Cocos Island.

Eucidaris thouarsti Valenciennes (H. L. Clark, 1940, p. 347; 1948, p. 229).
Santa Catalina Island, California; Guadalupe Island, Baja California,
and the Gulf of California to La Plata Island, Ecuador; Revillagigedo
Islands; Galapagos Islands.

Hesperocidaris panamensis A. Agassiz (H. lL. Clark, 1948, pp. 226, 131).
Off Galera Point, Ecuador; Galapagos Islands.

Lovenia cordiformis A. Agassiz (H. L. Clark, 1940, p. 352; 1948, p. 348).
Santa Barbara, California, and the Gulf of California to Guayaquil,
Keuador; Revillagigedo Islands; Galapagos Islands. Hawaiian Islands
(Ziesenhenne, 1937, p. 236).

Lytechinus pictus Verrill (H. L. Clark, 1940, p. 349). Monterey, California,
to the Gulf of California.

Meoma grandis Gray (H. lL. Clark, 1948, p. 344). Angeles Channel, Gulf of
California, to Port Utria, Colombia; Revillagigedo Islands; Galapagos
Islands.

Plagiobrissus pacificus H. lL. Clark (1948, p. 342). Gulf of California to
Panama Bay and questionably from La Plata Island, Eeuador.

Of these 13 species, 1 is known only from Cocos Island. All the others
are known to occur also in the waters of the mainland of the Panamic
province. Six of these also occur in the Revillagigedo Islands and 10 occur
at the Galapagos Islands. One has been reported from the Hawaiian
Islands.

Class HoLoTHURIOIDEA

Brandtothuria arenicola Semper. Gulf of California to Eeuador; Galapagos
Islands; Revillagigedo Islands; West Indies. Almost cireumtropical.
Brandtothuria impatiens Forskal. Cedros Island, Baja California, and Gulf

Vout. XXXII] AHHRTLEIN: BIOGEOGRAPHY OF COCOS ISLAND 237

of California to Colombia; Galapagos Islands; Revillagigedo Islands;

West Indies. Almost cireumtropieal.

Jaegerothurva inhabilis Selenka. Gulf of California to Eeuador; Galapagos

Islands; Revillagigedo Islands; Hawaii to Australia.

Lessonothuria pardalis Selenka. Gulf of California to Colombia; Hawaii.

Almost cireumtropical but lacking in Atlantic and West Indies.
Ludwigothurva atra Jaeger. Galapagos Islands; Clipperton Island; Hawaii

to Mozambique.

Ludwigothuria keferstemt Selenka. Mexico to Peru; Galapagos Islands;

Revillagigedo Islands.

Microthele difficilis Semper. Gulf of California to Galapagos Islands; Clip-
perton Island; Australia; east coast of Africa to east coast of Pacifie.
Psolus diomedeae Ludwig. Gulf of California to Keuador; Galapagos

Islands.

Selenkothuria theelt Deichmann. Gulf of California to Zorritos, Peru;

Galapagos Islands; Revillagigedo Islands.

Semperothuria mmitans Ludwig. Gulf of California to Panama; Galapagos

Islands; Revillagigedo Islands; Samoa.

Theelothuria paraprinceps Deichmann. Gulf of California to Panama;

Clarion Island, Revillagigedo Islands.

These holothurians were reported from Cocos Island by Deichmann
(1941b; 1958). All occur in the Panamie region and all except three also
live in western Pacific waters. The percentage of these species reported
from both the eastern Pacific and western Pacific waters is much greater
than that of most classes of invertebrates recorded from Cocos Island.
However, many species in this group of animals, especially the Aspidoehi-
rota, are widely distributed (see Deichmann, 1958, pp. 253, 277).

Phylum MOLLUSCA
Marine and Brackish-Water Mollusks
Class PELECYPODA

Arca (Anadara) reinharti Lowe |Rost, 1955, p. 227, as Anadara (Sca-
pharca) reinharti. |

Cetoconcha scapha Dall (1902, p. 561). Off Cocos Island in 183 meters (100
fathoms).

Crenella divaricata dOrbigny (Soot-Ryen, 1955, p. 130).

Tsognomon (Melina) chemnitzianum d’Orbigny | Pilsbry and Vanatta, 1902,
p. 559 (as Perna chemnitzianum d’Orb. (?); Dall, 1908, p. 437 (as
Melina chemnitziana); Hertlein, 1932, p. 45 (as Pedalion chemnit-
zianum) |.

238 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Isognomon quadrangularis Reeve [von Martens, 1902b, p. 138 (as Perna
quadrangularis); Pilsbry and Vanatta, 1902, p. 137 (as Perna quad-
rangularis); Biolley, 1907, p. 25 (as Perna quadrangularis) |.

Lithophaga attenuata Deshayes (Bartsch and Rehder, 1939, p. 18).

Lithophaga (Myoforceps) aristata Dillwyn (Soot-Ryen, 1955, p. 141;
Turner and Boss, 1962, p. 108).

Ostrea palmula Carpenter [von Martens, 1902b, p. 1387 (as Ostrea ochracea
Sowerby) ; Biolley, 1907 (as Ostrea ochracea) ; Hertlein, 1932, p. 45 (also
cited as Ostrea callichroa Hanley) |.

Pecten (Pecten) sericeus Hinds (Grau, 1959, p. 142).

Pecten (Cyclopecten) cocosensis Dall (1908, p. 405); Hertlein (1935, p. 319.
Original locality only and pl. 18, figs. 7 and 8); Keen (1958, p. 72) ;
Grau (1959, p. 30) (Dall’s record).

Pecten (Cyclopecten) exquisitus Grau (1959, p. 36).

Pecten (Oppenhermopecten) hancochr Grau (1959, p. 155).

Class GASTROPODA
(Marine and Brackish-Water Forms)

Acanthina brevidentata Wood |von Martens, 1902b, p. 137 (as Monoceros
brevidentata); Biolley, 1907, p. 21 (as Monoceros brevidentatum);
Hertlein, 1932, p. 45].

Acmaea (Nomaeopelta) mesoleuca Menke [von Martens, 1902b, p. 137 (as
Acmaea striata Quoy and Gaimard) ; Pilsbry and Vanatta, 1902, p. 559
(as “Scarria” mesoleuca); Biolley, 1907, p. 26 (Record of Pilsbry and
Vanatta (1902), also p. 24 (as Acmaea striata); Tomlin, 1928, p. 188 (as
Collisella mesoleuca); Hertlein, 1932, p. 45 (as Acmaea (Collisella)
aerugimosa Middendorff); Hertlein, 1937, p. 306 (von Marten’s record
(1902b) of Acmaea striata cited) |.

Acmaea strigatella Carpenter (Pilsbry and Vanatta, 1902, p. 559).

Bursa caelata Broderip [von Martens, 1902b, p. 137 (as Ranella caelata);
Biolley, 1907, p. 21 (as Ranella caelata) |.

Caducifer cinis Reeve [von Martens, 1902b, p. 137 (as Pollia cinis); Pilsbry
and Vanatta, 1902, p. 559 (as Tritonidea cinis) ; Biolley, 1907, p. 21 (as
Pollia cinis); Keen (1958, p. 398, as Caducifer thalia Pilsbry and
Lowe). |

Cantharus (Gemmophos) gemmatus Reeve [Hertlein, 1932, p. 45 (as Can-
tharus gemmatus) |.

Cantharus sanguinolentus Duclos [von Martens, 1902b, p. 137 (as “Pollia
sanguinolenta Duclos 1832 == haemastoma Gray, 1839”): Pilsbry and
Vanatta, 1902, p. 559 (as Tritonidea sanguinolenta) ; Biolley, 1907, p.
21 (as Pollia sanguinolenta) |

VoL. XXXII] HHERTLEIN: BIOGEOGRAPHY OF COCOS ISLAND 239

Cerithium adustum Kiener (von Martens, 1902b, p. 1387; Biolley, 1907,
p. 22; Hertlein, 1932, p. 45.)

Cerithium maculosum Kiener (Pilsbry and Vanatta, 1902, p. 559; Dall,
1908, p. 487).

Conus brunneus Wood (von Martens, 1902b, p. 137; Biolley, 1907, p. 20;
Tomlin, 1927, p. 155; Hanna and Strong, 1949, p. 269).

Conus dalli Stearns (Hertlein, 1932, p. 45; Hertlein, 1937, p. 306; Hanna
and Strong, 1949, p. 305).

Conus gladiator Broderip (Calif. Acad. Sei. Coll., W. H. Ochsner collector,
1905).

Conus recurvus Broderip | Dall, 1910, p. 225 (as Conus scariphus Dall);
Hanna and Strong, 1949, p. 280 (Conus scartphus in synon.) |.

Conus tiaratus Broderip (Hertlein, 1932, p. 45; Hanna and Strong, 1949,
Date).

Cymatium vestitum Hinds (Hertlein, 1932, p. 45).

Cypraea isabella mexicana Stearns [von Martens, 1902b, p. 137 (as Cypraea
“wahrscheinlich” isabella Linnaeus) ; Biolley, 1907, p. 21 (‘°?”); Sehilder
and Schilder, 1938, pp. 176, 197 (as Cypraea controversa mexicana) ;
Demond, 1957, p. 304 (as Cypraea (Luria) isabella) }.

Cypraea moneta Linnaeus | Hertlein, 1932, p. 45; Hertlein, 1937, p. 307;
imeram: 19474, pp. 58 (16), 74 (32): Ingram, 1947b, p. 147 (11); In-
ram. 1048. p- 140: meram, 1951, p. 152 (28): Demond, 1957, p3304
(as Cypraea (monetaria) moneta) |.

Cypraea rashleighana Melvill |Ingram, 1945, p. 106; 1947a, p. 76 (34);
aio, p. 1438 °(12)- 1951, p. 155 (31) ].

Ellobium stagnalis d@’Orbigny [Biolley, 1907, p. 19 (as Auricula (?) stag-
nalis); Hanna and Hertlein, 1938, p. 32 (as Auricula stagnalis) |.

Fissurella virescens Sowerby (von Martens, 1902b, p. 1387; Pilsbry and
Vanatta, 1902, p. 559; Biolley, 1907, p. 23; Tomlin, 1928, p. 188; Hert-
lein, 1982, p. 45).

Harpa crenata Swainson (Hertlein, 1932, p. 45).

Hipponsx grayanus Menke | Pilsbry and Vanatta, 1902, p. 559 (as Amalthea
grayana) ; Biolley, 1907, p. 283; Hertlein, 1932, p. 45 “‘ef.’’].

Hippomax pilosus Deshayes [von Martens, 1902b, p. 187 (as Hipponix per-
haps barbatus Quoy and Gaimard) |.

Latirus tuberculatus Broderip (Dall, 1908, p. 436; Hertlein, 1932, p. 45).

Littorina aspersa Philippi [von Martens, 1902b, p. 137 (as Littorina aspera);
Biolley, 1907, p. 28 (as Inttorina aspersa) ; Hertlein, 1932, p. 45].

Inttorina conspersa Philippi [von Martens, 1902b, p. 137; Biolley, 1907,
p. 21; Dall, 1908, p. 437 (also, Malpelo Id.) ; Hertlein, 1932, p. 45].

Inttorina modesta Philippi (Tomlin, 1927, p. 168). [Keen (1958, p. 282)
pointed out that this species ‘has been cited from the Panamic province,

240 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

but it seems an indeterminate form with a type locality in Alaska.’’|

Marginella minor C. B. Adams (Bartsch and Rehder, 1939, p. 18).

Melampus tabogensis C. B. Adams | Dall, 1896, p. 452 (as Tralia panamensis
C. B. Adams); Dall, 1900, p. 97 (as Melampus panamensis); von
Martens, 1900, p. 561 (Dall’s record (1900) cited; von Martens, 1902b,
p. 137 (as Melampus tabogensis) ; Dall, 1908, p. 486 (as Melampus pana-
mensis); Morrison, 1946, p. 36) |.

Melampus trilineatus C. B. Adams (Hanna and Hertlein, 1938, p. 134).

Mitra fultoni E. A. Smith [ (Calif. Acad. Sci. Coll., W. H. Ochsner collec-
tor, 1905; Sphon, 1961, pp. 34, 35) J.

Mitra lens Wood | Dall, 1908, p. 436].

Mitra tristis Swainson (Stanford Uniy. Coll.).

Mitrella ocellata Gmelin [Pilsbry and Vanatta, 1902, p. 559 (as Columbella
cribraria Lamarck) ; Biolley, 1907, p. 26 (record of Pilsbry and Vanatta,
(1902))]..

Murex humilis Broderip (Dall, 1908, p. 436).

Nerita funiculata Menke [von Martens, 1902b, p. 137 (as Nerita bernhardi
Reeluz); Pilsbry and Vanatta, 1902, p. 559 (as Nerita fulgurans bern-
hardi Recluz) ; Biolley, 1907, p. 23 (as Nerita bernhardi) ; Hertlein, 1932,
p. 45 (as Nerita bernhardi) |.

Nerita ornata Sowerby [von Martens, 1902b, p. 137; Biolley, 1907, p. 23;
Dall, 1908, p. 437 (as Nerita scabricosta Lamarck); Hertlein, 1932, p. 45
(as Nerita scabricosta var. ornata) |.

Neritina pilsbryt Tryon | Biolley, 1907, p. 18 (as Neritina latissima Broderip
var. globosa Broderip ) ; Hertlein, 1902, p. 45].

Nitidella sertularium Orb.? (Tomlin, 1927, p. 162). [Reeord probably
incorrect. A Patagonian species].

Olivella (Olivella) cocosensis Olsson (1956, p. 180; Keen, 1958, p. 424).

Phos cocosensis Dall (Dall, 1917, p. 578; Strong and Lowe, 1936, p. 310;
Keen, 1958, p. 406.)

Planaaxis planaxis Wood (Dall, 1908, p. 436).

Planaxis planicostatum Sowerby [von Martens, 1902b, p. 137 (as Planawis
planicostatus); Pilsbry and Vanatta, 1902, p. 559 (as Planazis plani-
costatus); Biolley, 1907, p. 21 (as Planaxis planicostatus); Tomlin,
1927, p. 168; Hertlein, 1932, p. 45).

Polimices helicoides Gray [Tomlin, 1927, p. 170 (as Polinices glabella
Reeve) |.

Purpura patula pansa Gould [von Martens, 1902b, p. 137 (as Purpura
patula Linnaeus); p. 140 (as “Purpura patula L. (pansa Conr.)”;
Pilsbry and Vanatta, 1902, p. 559 (as Purpura patula); Biolley, 1907,
p. 21 (as Purpura patula) ; Dall, 1908, p. 436 (as Thais patula) ; Tomlin,

VoL. XXXII] HERTLEIN: BIOGEOGRAPHY OF COCOS ISLAND 241

1928, p. 163 (as Thais patula) ; Hertlein, 1932, p. 45 (as Thais patula) |.

Pyrene labiosa Sowerby [Pilsbry and Vanatta, 1902, p. 559 (as Columbella
labrosa) ; Biolley, 1907, p. 26 (as Columbella labrosa, record of Pilsbry
and Vanatta) |.

Siphonaria gigas Sowerby (Dall, 1896, p. 453; Dall, 1900, p. 97; von Mar-
tens, 1902b, p. 137; Pilsbry and Vanatta, 1902, p. 559; Biolley, 1907,
p. 25; Dall, 1908, p. 486; Dall, 1909, p. 205; Hertlein, 1932, p. 45).

Siphonaria gigas var. characteristica Reeve [Tomlin, 1927, p. 154 (as
Siphonaria characteristica); Hertlein, 1932, p. 45. |

Tegula cooksoni BK. A. Smith (Tomlin, 1928, p. 187; also Calif. Acad. Sei.
Coll., W. H. Ochsner collector, 1905).

Tegula maculostriata C. B. Adams [Pilsbry and Vanatta, 1902, p. 559 (as
Chlorostoma maculostriatum C. B. Adams) ; Biolley, 1907, p. 26 (record
of Pilsbry and Vanatta cited) |. [The type locality of 7. maculostriata
is Jamaica. The specimens so identified by Pilsbry and Vanatta from
Cocos Island resemble the illustration of the type specimen published
by Cleneh and Turner (Occasional Papers on Mollusks, Department of
Mollusks, Museum of Comparative Zodlogy at Harvard College, Vol. 1,
No.’ 15, p. 305, pl. 39, fig. 13). The spire is higher and the scupture
coarser than that of 7. cooksont E. A. Smith].

Tegula gallina multifilosa Stearns [Pilsbry and Vanatta, 1902, p. 559 (as
Chlorostoma gallinum multifilosum ) ; Biolley, 1907, p. 26 (as Chlorostoma
gallinum multifilosum, record of Pilsbry and Vanatta cited) |]. [Record
from Cocos Island doubtful. Not known with certainty south of Baja
California |.

Thais columellaris Lamarek |von Martens, 1902b, p. 137 (as Purpura
columellaris); Pilsbry and Vanatta, 1902, p. 559 (as Purpura columel-
laris); Biolley, 1907, p. 21 (as Purpura columellaris); Dall, 1908, p. 437;
Hertlein, 1932, p. 45) ].

Thais haemastoma biserialis Blainville | Hertlein, 1932, p. 45 (as Thais
biserialis) |.

Thais (Vasula) melones Duclos [von Martens, 1902b, p. 137 (as Purpura
melo); Pilsbry and Vanatta, 1902, p. 559 (as Purpura melones); Biolley,
1907, p. 21 (as Purpura melones); Dall, 1908, p. 436 (as Thais melones);
Hertlein, 1932, p. 45 (as Thais crassa Blainville) |.

Thais speciosa Valenciennes (Calif. Acad. Sei. Coll., W. H. Ochsner, col-
lector, 1905).

Thais triangularis Blainville (Calif. Acad. Sci. Coll., W. H. Ochsner, col-
lector, 1905).

Trivia pacifica Gray (Tomlin, 1927, p. 166.)

Vermetus sp. (von Martens, 1902b, p. 137; Biolley, 1907, p. 25).

242 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

(Land Forms)

Habroconus (Cocosconus) hopkinsi Dall (Baker, 1941, p. 225).
Habroconus (Pseudoguppya) pacificus Pfeiffer (Baker, 1941, p. 226).
Habroconus (Cocoslens) pallidus H. B. Baker (1941, p. 224).

Leptinaria biolleyi von Martens (Hanna and Hertlein, 1938, p. 128).
Nesopupa (Cocopupa) cocosensis Dall (Hanna and Hertlein, 1938, p. 129).
Ochrodermella biolleyi von Martens (Hanna and Hertlein, 1938, p. 127).
Ochrodermella cumingiana Pfeiffer (Hanna and Hertlein, 1938, p. 125).
Opeas gracile Hutton (Hanna and Hertlein, 1938, p. 130).

Succinea globispira von Martens (Hanna and Hertlein, 1938, p. 129).

The land snails from Cocos Island have been discussed in a paper by
Hanna and Hertlein (1938) who have summarized the literature pertaining
to them. Baker (1941, pp. 223-226) later made changes in the nomencla-
ture of some of the species.

Of the 9 species reported from Cocos Island, Opeas gracile Hutton is
widely distributed over the Indo-Pacific region and evidently is an intro-
duced species. It also is possible that the species of Leptinaria may have
arrived on the island adventitiously. The other 7 species, known only from
Cocos Island, belong to the genera Habroconus, Nesopupa, Ochrodermella,
and Succinea. Kobelt (1899) and Hanna and Hertlein (1938) suggested
Indo-Paecifie affinities of the endemic species of these genera on Cocos
Island. Germain (1934, p. 153), remarked on the small number of species,
the absence of Bulimulus, and the fact that Ochrodermella also oceurs in the
Caroline Islands. Baker (1941, p. 352) believes that the affinities of the
land snails from Cocos Island are with species on the American mainland.
It appears probable that some elements of the land snail fauna were derived
from the eastern Pacific and others from Polynesia.

For additional information concerning these land snails see Ancey
(1903); Baker (1945); Biolley (1907; 1908-1909) ; Dall (1896; 1900) ;
Gude (1903); von Martens (1890-1901; 1898; 1902a); Pilsbry and Cooke
(1920).

Order Pteropoda
Limacina wnflata VOrbieny (Howard, 1952, p. 13).
Creseus virgula Rang (Howard, 1952, p. 13).
Diacria quadridentata parva Howard (1952, p. 13).
Cavolina longirostris constricta Howard (1952, p. 13).

Class AMPHINEURA
Acanthochitona hirudiniformis Sowerby.
Chiton stokes’) ; Biolley, 1907, p. 24 (as Chiton (Radsia) stokesi) ; Dall,
Chiton (Chiton) stokesvi Broderip |Pilsbry and Vanatta, 1902, p. 559 (as

Vou. XXXII] HERTLEIN: BIOGEOGRAPHY OF COCOS ISLAND 243

1908, p. 436 (as Chiton stokesi); Tomlin, 1927, p. 154 (as Chiton
stokesii) ; Leloup, 1956, p. 246. |

Chiton (Radsia) goodallit Broderip [von Martens, 1902b, p. 137 (as Chiton
(Radsia) goodalli); Boone, 1933, p. 24 (as Chiton (Chiton) goodallii),
p. 201 (as Chiton goodallir) |.

Placiphorella blainvillei Broderip (Dall, 1908, p. 357; Dall, 1909, p. 246).

Class CEPHALOPODA

Abraliopsis hoylei Pfeffer (Robson, 1948, p. 118).

Argonauta expansa Dall (Robson, 1932, p. 197, in synonymy of Argonauta
cornuta Conrad, 1854, p. 332. Not Argonauta cornutus Bose, Hist. Nat.
des Coquilles, vol. 3, p. 262, AN X[1802]).

Helicocranchia beeber Robson (1948, p. 130).

Liocranchia reinhardti Steenstrup (Robson, 1948, p. 128).

Melanoteuthis beeber Robson (1932, p. 103).

Octopus (Octopus) pusillus Gould [ Dall, 1909, p. 194 (as Polypus pusillus);
Robson, 1929, p. 150].

Octopodoteuthis nielsent Robson (1948, p. 120).

Onykia (Teleoteuthis, Auctt.) sp. (Robson, 1948, p. 121).

Polypus januarti Hoyle (Dall, 1909, p. 194). [Locality records from the
Pacific doubtful according to Robson, 1932, pp. 235, 240. Type locality
northeast Brazil].

Pyroteuthis giardi Fischer (Robson, 1948, p. 118).

(?) Sthenoteuthis sp. (Rhynchoteuthis stage) (Robson, 1948, p. 125).

Symplectoteuthis oulaniensis Lesson (Dall, 1909, p. 195; Berry, 1912, p. 304;
Robson, 1948, p. 127).

Taonidium pacificum Robson (1948, p. 130).

One of the earliest references in which the occurrence of marine mol-
lusks at Cocos Island is mentioned is that by Colnett (1798, p. 71) who
stated “Shell-fish, were scarce, though we collected some very large limpets,
of a new kind, and a few dead conches. The latter were seen in great num-
bers on the beach, and mostly inhabited by the Diogenes erab.” It seems
possible that the large limpets mentioned by Colnett might be referable to
Siphonaria gigas Sowerby or S. gigas characteristica Reeve which occur
abundantly at some places on this Island.

The identification of some of the species in the present paper, based
upon records taken from the literature, may be doubtful, but the general
composition of the assemblage is in harmony with other island molluscan
faunas (except that of Clipperton Island) in the eastern Pacific. Further
collecting on Cocos Island would undoubtedly yield additional species. I
believe, however, that the general composition represented by the present
list is representative of the marine mollusean fauna of this Island.

244 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

The present list contains 88 species and subspecies (3 species identified
only as to genus, not included in this number) arranged as follows: Pele-
eypoda 12, Gastropoda 57, Pteropoda +, Amphineura 4, Cephalopoda 11.
Of the total only 4 species, 3 peleeypods and 2 cephalopods, described from
or near Cocos Island, have not been reported elsewhere. All but 14 species
have been reported occurring on or near the mainland. At least 38 species
listed here also have been reported from the Galapagos Islands and at
least 33 species occur in the Gulf of California. Two gastropods, belonging
to the genus Cypraea, and 3 cephalopods, also have been reported from
Indo-Pacific waters.

The 4 species of pteropods reported from Cocos Island by Howard
(1952, p. 18) also oceur at other localities in the eastern Pacific.

The marine molluscan fauna of Cocos Island contrasts decidedly with
that of Clipperton Island where nearly 50 per cent of the species are Indo-
Pacific forms or have affinities with species in the central or western Pacific
(see Hertlein and Emerson, 19538; 1957).

Phylum ANNELIDA

Chloeia entypa Chamberlin (Hartman, 1939a, p. 3). Southern California
to Panama.

Chloera viridis Schmarda (Hartman, 1940, pp. 190, 205. Also cited from
Cocos Island as C. euglochis Ehlers by Treadwell, 1928, p. 450). Gulf of
California to Panama and the Galapagos Islands; West Indies.

Eusigalion spinosum Hartman (1939b, pp. 10, 17, 59). Farallon Islands,
California, to Cocos Island, and the Galapagos Islands.

Lepidonotus furcillatus Ehlers (Hartman, 1939b, p. 16). Cited from Cocos
Island on page 16 but on page 2 cited from Wreck Bay, Chatham Island,
Galapagos Islands.

Nerine cirratulus hirsutus Treadwell (1928, p. 479, as Spio hirsuta); Hart-
man (1959, p. 390). Known only from Cocos Island.

Notopygos crinita Grube (Treadwell, 1928, p. 450). St. Helena Island.

Perinereis helleri Grube (Cited from Cocos Island as Neanthes obscura
Treadwell, 1928, p. 472). Philippine Islands.

Psammolyce spinosa Hartman (1939b, pp. 10, 17, 74). Clarion Island,
Revillagigedo Islands, Mexico.

Sthenelais fusca Johnson (Hartman, 1939b, pp. 10, 17, 61). Washington
to Panama and the Galapagos Islands.

A few species of this group have been reported from Cocos Island by

Treadwell (1928) and Hartman (1939a, 1939b, 1940). All except two of

these have also been reported from mainland waters. The nomenclature
followed here is that of Hartman (1959).

VoL. XXXII] HERTLEIN: BIOGEOGRAPHY OF COCOS ISLAND 245

Phylum ARTHROPODA
Class CRUSTACEA
Subelass CopEPopA

Gloiopotes ornatus Wilson.
Pennella instructa Wilson.

These two species, parasites on a sailfish, were reported from off Chat-
ham Bay, Cocos Island by Sehmitt (1939b, p. 27).

Subelass CrrriPpEDIA

Conchoderma, “very probably C. virgatum (Spengler)” (Schmitt, 1939b,
p. 27). Attached to a copepod, Pennella instructa Wilson (with sucker
fish, Bcheneis remora Linnaeus), taken from a sailfish from off Chatham
Bay, Cocos Island.

Tetrachta squamosa Bruguiere |von Martens, 1902b, p. 138, (as Tetraelita
(type. error) porosa Gmelin) |. World-wide in tropical and subtropical
seas (see Pilsbry, 1916, U.S. Nat. Mus., Bull. 98, p. 249).

Tetrachta squamosa milleporosa Pilsbry (Hertlein, 1932, p. 45). Originally
deseribed from Albemarle Island, Galapagos Islands (Pilsbry, 1916,
Wes Nat. Mus., Bull. 93, p. 257).

Subclass MALACOSTRACA
Order Amphipoda

Podoceropsis dubia Shoemaker (1942, p. 32, fig. 12). Originally deseribed
from Chatham Bay, Cocos Island.

Talorchestia fritza Stebbing (1903, p. 928). Originally deseribed from Cocos
Island.

Order Decapoda
Brachyura

Dr. John Garth furnished the following list of species of spider crabs
from Cocos Island:

Family Majidae

Euprognatha bifida Rathbun. Mainland.

Euprognatha granulata Faxon. Mainland; Galapagos Islands.
Herbstia tunida (Stimpson). Mainland.

Inachoides laevis Stimpson. Mainland.

Inssa aurwilliusi Rathbun. Mainland; Galapagos Islands.
Iassa tuberosa Rathbun. Mainland.

246 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Microphrys branchialis Rathbun. Mainland.

Microphrys triangulatus Lockington. Mainland; Galapagos Islands.
Mithrax (Mithrax) spinipes (Bell). Mainland; Galapagos Islands.
Podochela hemphilli (Lockington). Mainland.

Sphenocarcinus agassizi Rathbun. Mainland; Galapagos Islands.
Stenocionops ovata (Bell). Mainland; Galapagos Islands.
Stenorhynchus debilis Smith. Mainland; Galapagos Islands.
Teleophrys cristulipes (Stimpson). Mainland; Galapagos Islands.

Family Parthenopidae

Thyrolambrus glasselli Garth (formerly known as T. erosus Rathbun).

Mainland.

According to Dr. Garth, all these species also occur on the mainland
and none is found in the Indo-Pacific. He stated: “The only semi-endemies
among the brachyurans that I recall are Euprognatha granulata Faxon,
shared with the Galapagos Islands, and Portunus (Achelous) brevimanus
(Faxon) (family Portunidae), shared with the Revillagigedo Islands.”

The following records of Brachyuran crabs from Cocos Island, with
their general distribution, are taken from Boone, Rathbun, and others.

Family Majidae
Mithrax (Mithraculus) denticulatus Bell. Mainland; Galapagos Islands.
Paradasygius depressus Bell (Garth, 1958, p. 81). Mainland.
Family Parthenopidae

Parthenope (Platylambrus) exilipes Rathbun. Mainland; Galapagos Islands.

Family Portunidae

Portunus (Achelous) brevimanus Faxon. Cocos Island, type locality;
Revillagigedo Islands.

Family Xanthidae

Carpilodes cinctimanus White. Galapagos Islands. [According to Rathbun
(U.S. Nat. Mus., Bull. 152, p. 242, 1930), Liomera cocosana Boone, type
locality, Cocos Island, is a synonym of C. cinctimanus |.

Eriphia squamata Stimpson. Mainland; Galapagos Islands. [According
to Garth (Allan Hancock Pacifie Expeditions, Vol. 5, No. 10, p. 486,
1946) Boone’s record (1927) of Eriphia granulosa A. Milne Edwards
from Cocos Island is referable to E. squamata].

Vou. XXXII] HERTLEIN: BIOGEOGRAPHY OF COCOS ISLAND 247

Leptodius cooksom Miers. Galapagos Islands; Clarion Island, Revillagigedo
Islands. [According to Glassell (Trans. San Diego Soe. Nat. Hist., vol.

7, no. 38, p. 453, 1934), Boone’s record (1927) of Xanthodius lobatus
A. Milne Edwards, a female from Cocos Island, is based on a young
specimen of Leptodius cooksoni\.

Micropanope polita Rathbun [1902, p. 281 (as Xanthias politus)|. Gulf of
California; mainland; Galapagos Islands. :

Ozius tenuidactylus Loeckington [Boone (1927, p. 225) as Oztus agassizu
A. Milne Edwards, a species now believed to be a synonym of Ozius
tenwmidactylus Lockington (see Glassell, Trans. San Diego Soc. Nat. Hist.,
vol. 8, no. 14, p. 104, 1935, as Ozius tenurdactylos)|. Mainland; Gala-
pagos Islands.

Ozius verreauru de Saussure. Mainland; Galapagos Islands.

Trapezia cymodoce ferruginea Latreille. Mainland; Galapagos Islands;
Indo-Pacific.

Family Grapsidae

Grapsus grapsus Linnaeus. Mainland (east and west coasts); Galapagos
Islands. [According to Dr. Garth (written communication), Boone’s
record (1927) of Pachygrapsus crassipes Randall from Cocos Island,
was based upon a young specimen of Grapsus grapsus |.

Pachygrapsus transversus Gibbes. Mainland (east and west coasts); Gala-
pagos Islands.

Plagusia immaculata Lamarek. Mainland; Galapagos Islands; Indo-Pacific
| Also recorded from Cocos Island by Garth (Allan Hancock Pacifie
Expeditions, vol. 5, no. 10, p. 512, 1946) }.

Planes cyanea Dana. 60 miles south of Cocos Island; mainland; Galapagos
Islands; Indo-Pacific. [According to Chace (Proe. U. 8S. Nat. Mus., vol.
101, no. 3272, p. 70 et seq., 1950), records of Planes minutus Linnaeus
(such as that of Boone, 1927) from the eastern Pacific are referable to
P. cyanea}.

Family Gecarcinidae
Cardiosoma crassum Smith (Sehmitt, 1939b, p. 27). A land erab. San José,
Baja California, Mexico, to the mouth of the Tumbes River, Peru.

Family Ocypodidae

Uca panamensis Stimpson. Mainland; Galapagos Islands. [According to
Crane (Zoologica, New York Zool. Soe., vol. 26, pp. 205, and 178, 1941,
respectively), Boone’s record of Uca galapagensis Rathbun, based upon
a female from Cocos Island, is now referable to U. panamensis, and her

248 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47H Ser.

record of U. galapagensis, a male, from the Galapagos Islands, is re-
ferable to U. macrodactyla Milne Edwards and Lueas}.

Family Calappidae
Calappa convexra de Saussure. Mainland; Galapagos Islands.
Anomura

Miss Janet Haig furnished the following list (supplemented by three
species by Boone, 1932) of Anomuran crabs from Cocos Island with their
oeceurrences elsewhere.

Family Coenobitidae

Coenobita compressus H. Milne Edwards. Mainland; Galapagos; Indo.
Pacific.
Family Paguridae

Pagurus californiensis Benedict (Boone, 1932, p. 9). Mainland; Galapagos
Islands.

Calcinus explorator Boone (1932, p. 22; Chace, 1962, p. 624). Galapagos
Islands.

Pylopagurus hirtimanus Faxon. Galapagos Islands.

Pylopagurus longimanus Faxon. Known only from Cocos Island.

Family Porcellanidae

Petrolisthes cocoensis Haig (1960, p. 117). Cocos Island, probably endemie.
Petrolisthes edwardsu de Saussure. Mainland.

Petrolisthes marginatus Stimpson. Mainland.

Petrolisthes ortmanni Nobili. Mainland.

Petrolisthes tonsorius Haig. Mainland.

Family Hippidae

Hippa denticulatifrons Miers (Boone, 1932, p. 58). Galapagos Islands;
Indo-Pacific.

To these may be added the following Maecruran forms:

Family Palaemonidae

Brachycarpus biunguiculatus Lucas (Holthuis, 1952, pp. 6, 7). Gulf of Cali-
fornia to Colombia; East and West Americas; Galapagos Islands; Medi-
terranean; Indo-Pacific.

Macrobranchium americanum Bate (Holthuis, 1952, pp. 130, 131, 132. Also
cited from Cocos Island as M. jamaicense Herbst by Beebe, 1926, p. 435,

Vou. XXXII] HERTLEIN: BIOGEOGRAPHY OF COCOS ISLAND 249

and by Boone, 1930, pp. 20, 146). Baja California to Peru; Galapagos
Islands.

Macrobranchium hancocki Holthuis (1952, pp. 112, 118. Also cited from
Cocos Island as MV. olfersii Wiegmann by Beebe, 1926, p. 485, and Boone,
1930, pp. 20, 143). Costa Rica to Colombia; Galapagos Islands.

Palaemon (Palaemon) ritteri Holmes, Holthuis (1952, pp. 175, 176, 177).

San Diego, California, to Paita, Peru; Galapagos Islands.

Family Palinuridae

Panulirus gracilis Streets (Holthuis and Villalobos, 1961, p. 254). Gulf of
California to Paita, Peru; ?Galapagos Islands.

Panulirus penicillatus Olivier (Chace, 1962, p. 617). Galapagos Islands;
Socorro Island, Revillagigedo Islands; Hawaiian Islands; Tuamotu
Islands; Red Sea to South Africa.

From an inspection of the ranges of the species in the foregoing lists,
it is obvious that the affinities of the species are with those of the mainland
and the Galapagos Islands.

Fifty species of brachyuran, anomuran and macruran (shrimp-lke)
crustaceans are here listed from Cocos Island but additional species are
known to occur there. Of the 50 species, two are endemic, 41 occur on the
mainland, 33 (or 34) at the Galapagos Islands, seven in Indo-Pacific waters
and three (or four) in Atlantic waters.

Authors who have published papers which inelude crabs and shrimps
occurring at Cocos Island include: Boone (1927; 1930a; 1930b; 1932);
Chace (1962); Faxon (1893; 1895, list pp. 257-258); Garth (1946; 1958) ;
Haig (1960); Holthuis (1952); Holthuis and Villalobos (1961); Rathbun
(1902; 1930); Schmitt (1939b).

Class MyrIopopa

Newportia rogerst Pocock. Mainland.

Otocryptops melanostoma Newport. Mainland.

Otostigmus scabricauda Humbert and Saussure. Mainland.
Epinannolene pitticri Brolemann. Cocos Island only.
Leptodesmus folium Brélemann. Cocos Island only.
Orthomorpha coarctata Saussure. Cocos Island only.
Rhinocricus (Eurhinocricus) biolleyi Brélemann. Mainland.

Seven species of Myriopoda have been reported from Cocos Island by
Brélemann (1903; 1905). Of these, 3 chilopods and 1 diplopod also occur
on the mainland, and 3 diplopods are reported only from Cocos Island.

250 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SEr.

Class [INSECTA

A few members of this class from Cocos Island noticed in the literature
are mentioned here.

Arrhinotermes oceanicus Wasmann (1902, p. 189). A termite.

Eutermes sp. (prope ripperti Rambur) (Wasmann, 1902, p. 139). A termite.

Kalotermes (Neotermes) larsent Light (1935, p. 239). A termite. Known
only from Cocos Island. Said to be related to Neotermes castaneus Bur-
meister which occurs in the American tropics.

Leucotermes insularis Wasmann (1902, pp. 139, 140). A termite.

Historis odius Fabricius. Cited as Histrio probably odius by Williams,
(1911, p. 296). Hvistrio orion Fabricius, now believed to be a synonym,
also was reported from Cocos Island by Beebe (1926, p. 486). This
butterfly ranges throughout the American tropics.

Herse cingulata Fabricius | Williams, 1911, p. 317 (as Phlegathontius cin-
gulata)|. This hawk-moth is wide spread in the American tropies and
also oceurs in the Galapagos Islands.

Leucomelina pica Maequart (Coquillett, 1901, p. 375). This little fly is
abundant on Cocos Island at least at certain times in the vear. It was
originally described from Brazil and the American tropies.

Cicada sp. (Heidemann, 1901, p. 370). One species of cicada has been re-
ported from Cocos Island.

Odynerus (Pachodynerus) nasidens Latreille (Williams, 1926, p. 349).
Williams reported this wasp from Cocos Island.

Popilius lenzi Kuwert (1897, p. 301; van Doesburg, 1953, p. 203). This
beetle was deseribed from Cocos Island. Its affinities are with species
on the mainland.

Collenette (1926, p. 234) mentioned collecting a ‘‘longicorn beetle” and
“lady-birds” on Cocos Island. He also mentioned (p. 231) seeing an insect
there resembling Agraulis vanillae Linnaeus (a widely distributed species)
from the Galapagos Islands.

Order Hymenoptera

Atta cephalotes Linnaeus* (Forel, 1908, p. 40; Emery, 1919, p. 40).

Azteca emmae Forel* (1908, p. 62; Emery, 1919, p. 40).

Brachymyrmex longicornis Forel* (1909, p. 64; Emery, 1919, p. 40).

Camponotus (Myrmothriz) abdominalis stercorarius Forel* (1908, p. 71;
Emery, 1919, p. 40).

Camponotus (Myrmobrachys) biolleyi Forel [Forel, 1902, p. 177 (as Cam-
ponotus brolleyi) ; Wheeler, 1919, pp. 301, 305; Emery, 1919, p. 40) |.

Camponotus (Myrmobrachys) blandus F. Smith* [Forel, 1908, p. 72;
Kmery, 1919, p. 40, as Camponotus (Myrmocamelus) blandus}.

Vou. XXXII] HERTLEIN: BIOGEOGRAPHY OF COCOS ISLAND 251

Camponotus (Myrmaphaenus) cocosensis Wheeler (1919, pp. 301, 305; 1933,
p61).

Cryptocerus cristatus Emery (1919, p. 40).

Cyphomyrmex rimosus slavint Forel* (Forel, 1908, p. 43; Emery, 1919,

p. 40).

Eciton (Labidus) crassicornis F. Smith (Emery, 1919, p. 40).
Euponera (Trachymesopus) stigma Fabricius (Wheeler, 1919, pp. 301, 302;

Emery, 1919, p. 40).

Odontomachus haematodes Linnaeus* (Forel, 1908, p. 35; Emery, 1919,

p. 40).

Odontomachus haematodes insularis Guérin (Wheeler, 1919, pp. 301, 303).
Pheidole biconstricta bicolor Emery (1919, p. 40).

Pheidole punctatissima Mayr (Emery, 1919, p. 40).

Pheidole subarmata Mayr* (Forel, 1908, p. 52; Emery, 1919, p. 40).
Prenolepis (vividula) guatemalensis cocoénsis Forel |Forel, 1902, p. 178

(as Prenolepis guatemalensis var. cocoensis); Wheeler, 1919, pp. 301,

305; Emery, 1919, p. 40, as Prenolepis (Nylanderia) quatemalensis var.

cocoénsis |.

Pseudomyrma belti Emery, var. (1919, p. 40).

Solenopsis geminata Forel* (1908, p. 45; Emery, 1919, p. 40).

Solenopsis succinea Emery (1919, p. 40).

Tetramorium guineénse Fabricius (Wheeler, 1919, pp. 301, 303; Emery,

1919, p. 40).

Wasmannia auropunctata Roger (Wheeler, 1919, pp. 301, 504; Emery,

ee Ds 40))2
Wasmannia auropunctata var. rugosa Forel* (1908, p. 45; Emery, 1919,

p. 40).

The foregoing includes the combined lists of species and subspecies of
ants reported from Cocos Island by Forel (1902; 1908), by Wheeler (1919;
1933) and by Emery (1919).

Forel (1908) suggested that 10 species (those in the list indicated by
the symbol*) of the ant fauna of Cocos Island evidently arrived there with
cultivated plants. Emery (1919) discussed the ant fauna and remarked on
the preponderance of Central American forms. Apparently he favored the
theory that some of these insects reached the island by way of a land
bridge connected with the mainland during late Tertiary time.

Wheeler (1919, p. 301) remarked on seven species (which he listed) as
follows: ‘With the exception of the tropicopolitan ‘tramp’, T’etramorium
guineénse, of Old World origin, none of the forms is known to occur in the
talapagos Islands and all are neotropical or have strongly neotropical at-
finities.”” Further (p. 302), “It is evident that the Cocos ants are decidedly
tropical whereas those of the Galapagos are mainly such as belong to sub-

252 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

tropical or temperate regions or at any rate to the cooler or subalpine
regions in the New World tropies.”

Class ARACHNIDA

Argyroepeira nigriventris Keyserling. Reported from Cocos Island by Banks
(1902, p. 61) and by Heller (1902, p. 78). It has also been reported
from the Galapagos Islands, Central America, Colombia, Brazil, and
the West Indies.

Phylum CHORDATA
Class REPTILIA

Anolis townsendi Stejneger
Sphaerodactylus pacificus Stejneger

Only two species of lizards have been described from Cocos Island, both
by Stejneger (1900, p. 163; 1903, p. 3). Sphaerodactylus pacificus has heen
compared by herpetologists with S. lineolatus Lichtenstein which ranges
from Central America to Colombia (Stejneger, 1903, p. 4; Barbour, 1921,
p. 286, pl, fiea3, pl 13; figs, 1-4).

For additional information on this group see Heller (1903); Taylor
(1956).

Class PISCES

Fowler (1938, pp. 248-261) published lists of the fishes known to occur
at Cocos Island. He recorded 72 species from or in the general vicinity of
this island. Briggs (1961, pp. 552-554) reported 16 of 53 (30 per cent)
species of transpacific shore fishes at Cocos Island. This would be 22.2 per
cent of the 72 species reported from the island by Fowler. More recently
Briggs (written communication April 3, 1962) mentioned a total of 59
shore fishes ‘‘of which sixteen are trans-pacific, thirty-eight are American,
and five are endemic.”

A freshwater guppy, Cotylopus cocoensis Heller and Snodgrass (1903,
p. 211, pl. 11), described from streams at Chatham Bay, Cocos Island, was
said to be allied to Sicydiwm salvini Grant from Panama.

Other workers who have contributed information concerning fishes from
Cocos Island include: Beebe (1926, pp. 228, 435); Beebe and Tee-Van
[1941la; 1941b (sharks, rays, mantas, chimaeras)]; Fowler (1932); Hal-
stead and Bunker (1953) ; Halstead and Schall (1956) ; Heller and Snod-
grass (1903); Heere (1936); Klausewitz (1958); Myers (1941); Myers and
Wade (1941); Nichols and Breeder (1928) ; Schmitt and Schultz (1940) ;
Seale (1940) ; Snodgrass and Heller (1905).

Vou. XXXII] HHRTLEIN: BIOGEOGRAPHY OF COCOS ISLAND 253

Class AVES
(Land Birds)

Butorides virescens maculatus Boddaert. (Green heron). [Gifford, 1913,
p. 65 (as Butorides virescens); Sludd, 1962, written communication (as
Butorides virescens maculatus) ]. Mainland and Galapagos Islands.

Dendroica petechia aureola Gould. (Yellow warbler). Reported from Cocos
Island as Dendroica aureola Gould by Townsend (1895, p. 122) and as
Dendroeca petechia by Gifford (1919, p. 216) ; Beebe, 1926, p. 435 (as
Dendroica petechia aureola) ; Sludd, [1962, written communication (as
Dendroica petechia aureola) |. Mainland.

Mirundo rustica erythrogaster Boddaert. (Barn swallow). (Gifford, 1919,
p. 205). Mainland.

Nesococcyx ferrugineus Gould. (Cocos Island Cuckoo). [Gould, 18438, p.
105 (as Coccyzus ferrugineus) ; Gifford, 1919, p. 195 (as Coccyzus
ferrugineus) ; Beebe, 1926, p. 435 (as Coccyzus ferrugineus) ; Slaudd,
1962 [written communication (as Nesococcyx ferrugineus) |. Known
only from Cocos Island.

Nesotriccus ridgwayt Townsend. (Ridgway’s flyeatcher). (Townsend, 1895,
p. 124; Gifford, 1919, p. 200; Beebe, 1926, p. 435). Known only from
Cocos Island.

Pandion haliaétus Linnaeus. (Osprey). (Gifford, 1919, p. 193). Mainland.

Pinaroloxias mornata Gould. (Coeos Island finch). [Gould, 1843, p. 104 (as
Cactornis inornata) ; Townsend, 1895, p. 123 (as Cocornis agassizt) ;
Richmond, 1902, pp. 247-248 (discussion of the type locality and syn-
onymy of Pinaroloxias inornata) ; Gifford, 1919, p. 242 (as Pinaroloxrias
mornata); Beebe, 1926, p. 485 (as Cocornis agassizi); Swarth, 1931, pp.
268-271; Lack, 1945, pp. 19, 126, 129, and 1953, pp. 67, 72; Bowman
Gigaipp, 20, 92, 94 100) 104,109) 11S 5115, 129, 159; 161, 168; 177, 204,
ZO 216 28) 220) 225, 231,284,289) pls. 7, 12; 16,°20) |. Known only
from Cocos Island.

This list of 7 species and subspecies of land birds from Cocos Island with
their occurrence elsewhere, was compiled from the literature supplemented
by information received from Dr. Paul Sludd. Of the 7, 3 are known only
from this island. Wallace in 1876 (p. 60), remarked on the interesting
occurrence of “Coccyzus” [| Nesococcyx]| on this island.

Gifford (1919) discussed the land birds of Cocos Island and additional
remarks concerning members of this group may be found in papers by Beck
(1907) ; Murphy (1936); Rothehild (1902); Townsend (1895); see also Car-
riker (1910) and Eisenmann (1955).

Sea birds from Cocos Island have been discussed by Snodgrass and
Heller (1902; 1903); by Gifford (1913), whose account includes references
to other authors; and by Murphy (1936, p. 319).

254 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47H Ser.
BOTANY

The flora of Cocos Island has attracted the attention of several botanists.
A comprehensive report by Stewart (1912) recorded 77 species of vascular
plants, 20 of which are ferns. Only 8 of them (8.69 per cent), are endemic.
Twenty-seven of these species, 11 of them ferns, also occur in the Galapagos
Islands where the endemicity is reported to be 40.9 per cent

Stewart concluded from his studies that the flora of Cocos Island was
that of an oceanic island, younger than that of the Galapagos Islands, and
that there was no evidence to support the theory that the island was ever
connected by land with either the mainland or the Galapagos Islands. He
believed that the flora reached the island chiefly from the mainland for-
tuitously through the agencies of winds, ocean currents, and migratory
birds.

Svenson (1935, p. 259), referring to Johnston (1931, p. 35), stated that
the vegetation of Cocos Island consists of about 100 species, of which about
10 are endemie, “overwhelmingly of ferns, melanostomes, and orchids.” He
mentioned that there is no change in the flora to a height of at least 457
meters (1500 feet). He also stated that except for ferns, virtually none of
the species also occur on the Galapagos Islands.

Cook (1939) deseribed a mountain palm, Rooseveltia frankliniana, from
Cocos Island. It was said to be rather closely related to Plectis oweniana
Cook from Guatemala. Cook belheved that the coconut palms observed on
Cocos Island as early as 1699 by Wafer, were introduced there.

Various components of the Cocos Island flora have been discussed by
authors in several papers including: Bartram (1933. Mosses; see also list
by Stewart, 1912, p. 395) ; Clark (1953, Hepaticae) ; Cook (1910; 1939; 1940,
Palms) ; Cooke and Bonar (1961, Fungi) ; Howe (1934, Hepaticae) ; Pittier
(1899) ; Rose (1892) ; Stewart (1911, pp. 230, 233-235. He remarked on
the origin of the Galapagos flora and gave occasional reference to Cocos
Island) ; Svenson (1935, includes various authors in his bibliography).

BIBLIOGRAPHY
ANONYMOUS

1920. Malpelo, Cocos, and Easter islands. Handbooks prepared under
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1934a. Cocos Island: Lost treasure stirs an international feud. News-
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1934b. Treasure seekers arrested. Literary Digest, vol. 118, p. 8,
October 27.

VoL. XXXII] AHHLRTLEIN: BIOGEOGRAPHY OF COCOS ISLAND 255

1935. United States Hydrographic Office, Chart 823. South Pacifie
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1938a. Sailing directions for the west coasts of Mexico and Central
America. Hydrographic Office, no. 84, Washington, D.C.,
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1938b. Third presidential cruise of the U.S.S. Houston, Captain G. N.
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1939a. Cocos Island is still largely unknown world. Science News
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1945. Isla del Coco (Cocos Island). Hydrographie Office Chart no.
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AncEY, C. F.

1903. Faune malacologique terrestre de l’Ile des Cocos dans |’Océan

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ANDERSON, ISABEL
1936. Zigzagging the South Seas. Bruce Humphries, Ine., Publ., Bos-
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ANSON, G.
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ANTROBUS, E.
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Baker, H. B.
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1945. Some American Achatinidae. Nautilus, vol. 58, no. 3, pp. 84-92,
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Banks, N.

1902. Papers from the Hopkins Stanford Galapagos Expedition, 1898—
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Barpour, T.
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3ARTRAM, EK. B.
1933. The Templeton Crocker Expedition of the California Academy
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Vout. XXXII] HERTLEIN: BIOGEOGRAPHY OF COCOS ISLAND 257

Fourth Series, vol. 21, no. 8, pp. 75-86, figs. 1-3 in text,
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Bre, ih. H.
1907. Notes from Clipperton and Coccos islands. Condor, vol. 9, no. 4,
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1926. The Arcturus adventure. An account of the New York Zoologi-
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1926. Cocos—The isle of pirates. (In Beebe, W.) The Arcturus ad-

venture. G. P. Putnam’s Sons, New York and London, pp.
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1941a. Kastern Pacific expeditions of the New York Zoological Society.
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entific contributions of the New York Zoological Society, vol.
26, pt. 2, no. 15, pp. 93-122, pls. 1 and 2, figs. 1-34 in text,
September 8. [Concerning species from Cocos Island, see pp.
LO ET at. S20.)
1941b. Same reference, no. 26, part 3. Rays, mantas and chimaeras, pp.
245-280, pls. 1-4, figs. 1-40 in text, October 31. [Concerning
species from Cocos Island, see p. 274. |
BELCHER, E,
1843. Narrative of a voyage round the world, performed in Her
Majesty’s Ship Sulphur during the years 1836-1842. Vol. 1, pp.
I-X XXVIII, 1-387, frontispiece, 7 pls., 8 maps, 11 vignettes.
| Concerning Cocos Island, see pp. 186-191; 223, 253-254. |
Berry, S. S.
1912. <A review of the Cephalopoda of western North America. Bulletin
of the Bureau of Fisheries, vol. 30 (Document 761), Wash-

258 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER.

ineton, D.C., pp. 269-336, pls. 32-56, figs. 1-18 in text, 1910
(issued July 24, 1912).

BetTaGcH, W.

1728. Voyage round the world being an account of a remarkable enter-
prize, begun in the year 1719, chiefly to cruise on the Spaniards
in the great South Ocean. Relating the true historical facts of
that whole affair: Testifyd by many imployd therein; and
econfirmd by authorities from the owners. London, pages
[I-X not paginated| 1-342 [+ 3 not paginated], 1 folding
map. [Concerning Cocos Island, see pp. 146-147. |

BIoLLeEyY, P.

1907. Mollusques de L’Isla del Coco. Museum Nacional de Costa Rica,
San José, pp. 1-30, 2 pages of maps.

1908-1909. Los Moluseos de la Isla del Coco (Costa Rica). Anales
Museum Nacional, San Salvador, tome III, no. 22, pp. 327-
333, September 1, 1908; no. 23, pp. 384-387, January 1, 1909.
| Abbreviated data from paper of 1907, according to Calvert,
Amelia S., and Calvert, P. P., A year of Costa Rican natural
history. Macmillan Co., New York, 1917, p. 532. ]

Boone, LEE

1927. The littoral Crustacean fauna of the Galapagos Islands. Zoo-
logiea, Scientific Contributions of the New York Zoological
Society, vol. 8, no. 4, pp. 127-288, pl. C. (map), figs. 34-102D,
August 11.

1930a. Seientifie results of the cruises of the yachts Eagle and Ara,
1921-1928, William K. Vanderbilt, commanding. Crustacea:
Stomatopoda and Brachyura. Bulletin of the Vanderbilt Ma-
rine Museum, vol. 2, pp. 1-228, pls. 1-74, September 1. [Con-
cerning species from Cocos Island, see pp. 18, 19.]

1930b. Scientific results of the cruises of the vachts Hagle and Ara,
1921-1928, William K. Vanderbilt, commanding. Crustacea:
Anomura, Macrura, Schizopoda, Isopoda, Amphipoda, Mysi-
dacea, Cirripedia, and Copepoda. Bulletin of the Vanderbilt
Marine Museum, vol. 3, pp. 1-221, pls. 1-83, November 1.
| Concerning species from Coeos Island, see p. 20. |

1932. The littoral crustacean fauna of the Galapagos Islands. Zoo-
logiea, Scientific Contributions of the New York Zoological
Society, vol. 14, no. 1, pp. 1-62, figs. 1-19, January. [Con-
cerning species from Cocos Island, see pp. 9, 22, 26, 60. |

1933. Seientific results of cruises of the yachts Hagle and Ara, 1921—

Vout. XXXII] HERTLEIN: BIOGEOGRAPHY OF COCOS ISLAND 259

1928, William K. Vanderbilt, commanding. Coelenterata,
Echinodermata and Mollusea. Bulletin of the Vanderbilt
Marine Museum, vol. 4, pp. 1-217, pls. 1-133, April 30. [Con-
cerning species from Cocos Island, see pp. 16-24. |

Bowman, R. I.

1961. Morphological differentiation and adaptation in the Galapagos
finches. University of California Publications in Zoology, Vol.
58, pp. I-VI, 1-302, pls. 1-22, figs. 1-74 in text, August 22.
[| Concerning Cocos Island, see p. 6 and fig. 2 (p. 7). Coneern-
ing Pinaroloxis mornata, see pp. 20, 92, 94, 100, 104, 109, 113,
Size. tod lol 168) 177, 204 207-216, 218, 220-225, 231,
Zote Zoo pls: i. M2 16,-20.|

Brecs. J. C.
1961. The east Pacific barrier and the distribution of marine shore
fishes. Evolution, vol. 15, no. 4, pp. 545-554, figs. 1-3 in text,
December 30.

Briaas, MARIE
1950. Cocos Island venture. Borden Publishing Company, Los Ange-
les, California, pp. 1-214, 1 map (on lining paper).

BROLEMANN, H. W.

1903. Myriapodes recueillis a l’Isla de Cocos par M. le Professeur P.
Biolley. Annales de la Societe Entomologique de France, vol.
72, ler trimestre, pp. 128-143, 1 pl., figs. 1-10 in text, June.

1905. Myriapodes de Costa-Rica recueillis par Mr. le Professeur P.
Biolley. Annales de la Societe Entomologique de France, vol.
74, der trimestre, pp. 336-380, pls. 8, 9, 10, figs. 1-9 in text,
October. [Concerning species from Cocos Island, see pp. 337,
308, 339, 341, 372.]

BurNEY, J.

1803-1816. A chronological history of the discoveries in the South Sea
or Pacifie Ocean. London, pt. 1, pp. I-XII (4+ 8), 1-383, ap.,
pp. 385-391, 5 charts, 1803. [Concerning Cocos Island, see pp.
275-276.|; vol. 4, pp. I-X VI, 1-580, 5 charts, 1816. [Con-
cerning Cocos Island, see pp. 155, 189-190. ]

CALLANDER, J.

1768. Terra Australis Cognita: or, voyages to the Terra Australis, or
southern hemisphere, during the sixteenth, seventeenth, and
eighteenth centuries. Printed by A. Donaldson, Edinburgh,
vol. 2, pp. I-II (preface by the editor), 3-692. [In this sum-

260 CALIFORNIA. ACADEMY OF SCIENCES [Proc. 4TH SEr.

mary of voyages, Dampier’s remarks concerning Cocos Island
appear on pp. 578-579 and those of Wafer on pp. 674-676. |

CAMPBELL, MaLcoum, (Sir)

1931. My greatest adventure; searching for pirate treasure in Cocos
Island. T. Butterworth, Ltd., London, pp. 1-290, frontispiece,
plates.

1932. Searching for treasure in Cocos Island. Frederick A. Stokes
Company, New York, pp. I-VIII, 1-279, frontispiece colored,
pls. 1-4.

1934. Modern buried treasure. E. P. Dutton & Co., New York. | Con-
cerning Cocos Island, see pp. 100-138.] [Not seen. |

CARRIKER, M. A., JR.

1910. An annotated list of the birds of Costa Rica ineluding Cocos
Island. Annals of the Carnegie Museum, vol. 6, nos. 2-3, pp.
314-915, 1 map, August. [Concerning species from Cocos
Island, see pp. 563, 709, 813, 894. ]

CHACE, ine Aue Jin:

1962. The non-brachyuran decapod erustaceans of Clipperton Island.
Proceedings of the United States National Museum, vol. 118,
no. 3466, pp. 605-635, fies. 1-7 in text. [Concerning species
from Cocos Island, see pp. 617, 624. |

CHAMBERLIN, J.

1935. Treasure hunting, incorporated. Review of Reviews, vol. 91, no.
2, pp. 33-37, 68, 10 figs., February. [Concerning Cocos Island,
see pp. 37, 68.]

CHETWOOD, JJ.

1904. Our search for the missing millions (of Cocos Island) by one of
the searchers. Being an account of a curious cruise, and a
more than curious character. The South Sea Bubble Co. | ?San
Francisco], pp. 1-210, frontispiece and pls. 1-12. [Cocos
Island, pp. 17-30. |

CuHusp, L. J.

1933. Geology of Galapagos, Cocos, and Easter islands. Bernice P.
Bishop Museum, Bulletin 110, pp. 1-67, pls. 1-5, figs. 1-9 in
text. [Chubb, pp. 1-44, 65-67, pls. 1-5, figs. 1-8. “Petrology
of the Galapagos Islands” by Constance Richardson, pp. 45-64,
fig. 9.| [Concerning Cocos Island, see pp. 25-30, fig. 6, pl. 3,
fig. B.|

VoL. XXXII] HERTLEIN: BIOGEOGRAPHY OF COCOS ISLAND 261

CHURCH, G. E.
1897. [Cocos Island, in] Costa Rica. Geographical Journal, vol. 10,
no. 1, pp. 56-84, 1 map (opp. p. 128), July. [Cocos Island, p.
73. |

CLARK, A. H.

1939. Echinoderms (other than Holothurians) collected on the presi-
dential cruise of 1938. Smithsonian Miscellaneous Collections,
vol. 98, no. 11 (publ. 3536), pp. 1-18, pls. 1-5, June 2. [Con-
cerning species from Cocos Island, see p. 2.|

Cuark, H. L.

1940. Eastern Pacific expeditions of the New York Zoological Society.
XXI. Notes on the Echinoderms from the west coast of Cen-
tral America. Zoologica, Scientific Contributions of the New
York Zoological Society, vol. 25, pt. 3, no. 22, pp. 331-352,
pls. 1, 2, text figs. 1-4, November 14. [Concerning species from
Cocos Island, see pp. 333, 336, 341, 342, 3438, 347, 349, 350,
352. |

1948. A report on the Echini of the warmer eastern Pacific, based on
the collections of the Velero III. Allan Hancock Pacifie Ex-
peditions, vol. 8, no. 5, pp. I-XII (Introduction by Th. Mor-
tensen), pp. 225-352, pls. 35-71, text figs. 1-3, December 29.
[| Concerning species from Cocos Island, see pp. 226, 229, 235,
294, 296, 297, 330, 342, 344, 348.]

Cuiark, LoIs
1953. Some hepaticae from the Galapagos, Cocos, and other Pacific
Coast islands. Proceedings of the California Academy of
Sciences, fourth series, vol. 27, no. 18, pp. 593-624, pls. 37-41,
Mareh 30. [Concerning species from Cocos Island, see pp.
599, 601, 602, 603, 604, 606, 607, 608, 609, 610, 611. |

CLoveEr, S. T.
1932. A pioneer heritage. Saturday Night Publishing Company, Los
Angeles, pp. I-II, 1-291, pls. 1-97, 4 maps (front and back),
1 text fig. [Concerning Cocos Island, see pp. 143-144, 209-
Pl 2 /pls.l|

CoLLENETTE, C. L.

[1926]. Sea-girt jungles. The experiences of a naturalist with the Sf.
George Expedition. Hutchinson & Company, London, pp.
I-XIV, 15-275, frontispiece, 1 map, 34 illustrations. [Con-
cerning Cocos Island, see pp. 216-235. |

262 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

CoLNETT, J., (Captain)

1798. A voyage to the South Atlantic and round Cape Horn into the
Pacific Ocean, for the purpose of extending the spermaceti
whale fisheries, and other objects of commerce, by ascertaining
the ports, bays, harbours, and anchoring births, in certain
islands and coasts in those seas at which the ships of the
British merchants might be refitted. Printed for the author,
by W. Bennett, London, pp. I-X VIII, 1-179, 1 portrait, 6
charts and 3 pls. of illustr. [Concerning Cocos Island, see pp.
67-74, 1 ehart, 1 illustr. |

Cook, O. F.

1910. History of the Coconut Palm in America. Contributions from
the United States National Herbarium, vol. 14, pt. 2, pp.
I-VITI, 271-842, IX—XIII (index), pls. 53-66, December 31.
[See “The Palms of Cocos Island,” pp. 291-296. |
1939. A new Palm from Cocos Island collected on the presidential
eruise of 1938. Smithsonian Miscellaneous Collections, vol.
98, no. 7, (publ. 3532), pp. 1-26, pls. 1-26, May 29.
1940. An endemie Palm on Cocos Island near Panama mistaken for the
Coconut Palm. Science (new ser.), vol. 91, no. 2354, pp. 140-
142, February 9.
CooxkE, W. B., and L. Bonar
1961. Additional fungi from the Galapagos and other Pacifie Coastal
islands collected during the Templeton Crocker Expedition,
1932. Occasional papers of the California Academy of Sei-
ences, no. 29, pp. 1-5, January 30. [See Poris umbrinella
Bresadola, reported from Cocos Island, p. 5.|
Coouince, A.
1953. A visit to the Cocos and Galapagos islands on board the Blue
Dolphin. Boston, privately printed, 104 pp. [not paginated],
87 illustr., 1 chart, 2 vignettes. [16 pp. of text and 25 illustra-
tions taken at or near Cocos Island. |
COQUILLETT, D. W.
1901. Papers from the Hopkins Stanford Galapagos Expedition, 1898—
1899. II. Entomological results (2): Diptera. Proceedings
of the Washington Academy of Sciences, vol. 3, pp. 371-379,
November 7. [Leucomolina pica Macquart recorded from
Cocos Island, p. O79. |
CouLTER, J.

1847. Adventure on the western coast of South America and the in-

VoL. XXXII] HERTLEIN: BIOGEOGRAPHY OF COCOS ISLAND 263

terior of California. Longman, Brown, Green, and Longmans,
London, vol. 1, pp. I-XII, 1-288. [Concerning Cocos Island,
see pp. 95-126. |

CRAUFURD, Q. C. A.

1927. Concerning a hunt for treasure in the Pacific. A true yarn.
Blackwood’s Magazine, vol. 221, no. 1338, pp. 535-546, April.

CusHMAN, J. A., and IRENE McCuLuLocH

1939. A report on some arenaceous Foraminifera. Allan Hancock Pa-
cific Expeditions, vol. 6, no. 1, pp. 1-113, pls. 1-12, August
14. [Concerning species from Cocos Island, see pp. 60, 69,
79, 96, 104, 106, 112.]

1950. Some Lagenidae in the collections of the Allan Hancock Founda-
tion. Allan Haneock Pacific Expeditions, vol. 6, no. 6, pp.
295-364, pls. 37-48, February 23. |Concerning species from
Cocos Island, see pp. 315, 317, 327, 352. |

Dau, W. H.

1895. Scientific results of explorations by the U.S. Fish Commission
steamer Albatross. No. XXXIV. Report on Mollusca and
Brachiopoda dredged in deep water, chiefly near the Hawaiian
Islands, with illustrations of hitherto unfigured species from
northwest America. Proceedings of the United States National
Museum, vol. 17, no. 1032, pp. 675-733, pls. 23-32, July 2.
| Concerning species from Cocos Island, see p. 717.|

1896. Insular landshell faunas, especially as illustrated by the data
obtained by Dr. G. Baur in the Galapagos Islands. Proceed-
ings of the Academy of Natural Sciences of Philadelphia, vol.
48, pp. 395-459, pls. 15-17. [Pp. 395-456, issued September
15, pp. 457-459, September 22. |

1900. Additions to the insular land-shell faunas of the Pacifie Coast,
especially of the Galapagos and Cocos islands. Proceedings of
the Academy of Natural Sciences of Philadelphia, vol. 52, pp.
88-106, pl. 8. [Pp. 88-96, April 13; pp. 97-106, April 16.]
[‘“Coeos Island species,” pp. 96-99. |

1902. Illustrations and descriptions of new, unfigured, or imperfectly
known shells, chiefly American, in the U.S. National Museum.
Proceedings of the United States National Museum, vol. 24
(no. 1264), pp. 499-566, pls. 27-40, March 31. [Concerning
species from Cocos Island, see p. 561.|

1908. Reports on the dredging operations off the west coast of Central
America to the Galdpagos, to the west coast of Mexico, and in

264 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

the Gulf of California, in charge of Alexander Agassiz, ear-
ried on by the U.S. Fish Commission steamer Albatross,
during 1891, Lieut. Commander Z. L. Tanner, U.S.N., com-
manding. XXXVII. Reports on the scientific results of the
expedition to the eastern tropical Pacific, in charge of Alex-
ander Agassiz, by the U.S. Fish Commission steamer Albatross,
from October, 1904, to March, 1905, Lieut. Commander L. M.
Garrett, U.S.N., commanding. XIV. Reports on the Mollusea
and Brachiopoda. Bulletin of the Museum of Comparative
Zodlogy at Harvard College, vol. 43, no. 6, pp. 205-487, pls.
1-22, October. [Concerning species from Cocos Island, see pp.
405, 436-437. |

1909. Report on a collection of shells from Peru, with a summary of
the littoral marine Mollusea of the Peruvian Zoological Prov-
ince. Proceedings of the United States National Museum, vol.
37, no. 1704, pp. 147-294, pls. 20-28, November 24. [Con-
cerning species from Cocos Island, see pp. 194, 195, 205, 246. |

1910. Summary of the shells of the genus Conus from the Paeifie
Coast of America in the U.S. National Museum. Proceedings
of the United States National Museum, vol. 38, no. 1741, pp.
217-228, June 6. |Coneerning species from Cocos Island, see
[Oe 245)

1917. Summary of the mollusks of the family Alectrionidae of the
West Coast of America. Proceedings of the U.S. National
Museum, vol. 51, no. 2166, pp. 575-579, January 15. [Con-
cerning species from Cocos Island, see p. 578. |

1920. Annotated list of the recent Brachiopoda in the collection of the
United States National Museum, with descriptions of thirty-
three new forms. Proceedings of the U.S. National Museum,
vol. 57, no. 2314, pp. 261-877, June 24. [Concerning species
from Cocos Island, see p. 288.]

DAMPIER, W.

1699. A new voyage round the world. Printed for James Knapton,
London, vol. 1, edit. 4 corrected, pp. A2—A4 [6 pp.], 1-550,
4 maps. [Concerning Cocos Island, see pp. 111, 131.]

1729. A new voyage round the world (with an introduction by Sir
Albert Gray). Argonaut Press, London, pp. IL-XX XVII,
1-376, portrait, 6 pls. (4 maps). Reprint of Edit. 7, 1927.
| Concerning Cocos Island, see pp. 83, 111.]

Davis, W. M.

[1874] 1926. Nimrod of the sea or the American whaleman. Edition

Vout. XXXII] AHHRTLEHIN: BIOGEOGRAPHY OF COCOS ISLAND 265

1926 with an introduction and notes by John R. Spears (pp.
IX—XIX). Charles E. Lauriat Company, Boston, pp. 1-406,
frontispiece, 31 illustr. [Concerning Cocos Island, see pp.
115-124, 3 figs. |
DEICHMANN, ELIZABETH
1941a. Coelenterates collected on the presidential cruise of 1938. Smith-
sonian Miscellaneous Collections, vol. 99, no. 10, pp. 1-17,
1 pl, figs. 1-4 in text, January 27. [Concerning species from
Cocos Island, see p. 13.]
1941b. The Holothurioidea collected by the Velero III during the years
1932 to 1938. Part I. Dendrochirota. Allan Hancock Pacific
Expeditions, vol. 8, no. 3, pp. 61-195, pls. 10-30, June 20.
[Concerning species from Cocos Island, see pp. 69, 149, 150. |
1958. The Holothurioidea collected by the Velero III and IV during
the years 1932 to 1954. Part II. Aspidochirota. Allan Han-
cock Pacifie Expeditions, vol. 11, no. 2, pp. 253-349, pls. 1-9,
February 11. [Concerning species from Cocos Island, see pp.
261, 289, 292, 293, 296, 305, 312, 313, 318, 324, 326. ]
DEMOND, JOAN
1957. Micronesian reef-associated gastropods. Pacific Science, vol. 11,
no. 3, pp. 275-341, pls. 1-4, figs. 1-42 in text, July. [Concern-
ing species from Cocos Island, see pp. 301, 504. ]
DorsBurG, P. H. VAN
1953. On some neotropical Passalidae. Pan-Pacifie Entomologist, vol.
29, no. 4, pp. 203-205, October. [Concerning species from
Cocos Island, see p. 203. |
Dovueuas, A. J. A., (Major), and P. H. JoHNson
1926. The South Seas of today, being an account of the cruise of the
yaeht St. George to the South Pacific. Cassell and Company.
Ltd., London, New York, pp. i-xiv, 1-295, 39 illustrations and
3 charts.
. Driscouu, C. B.
1931. Pirates ahoy! The perennial treasure hunters of Cocos Island.
San Francisco Chronicle, vol. CXX XIX, no. 151, Sunday
magazine section [p. 8, not paginated], 1 illustr., December 13.

DurHam, J. W.
1962. Scientific results of the Galaépagos-Expedition 1953-54 of the In-

ternational Institute for Submarine Research, Vaduz (Liech-
tenstein), leader, Dr. Hans Hass. Corals from the Galapagos

266 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

and Cocos Islands. Proceedings of the California Academy of
Sciences, fourth ser., vol. 32, no. 2, pp. 41-56, figs. 1-9, June
29 [mailed July 13. |

DurHaAm, J. W., and J. L. BARNARD
1952. Stony corals of the eastern Pacific collected by the Velero III
and Velero IV. Allan Hancock Pacific Expeditions, vol. 16,
no. 1, pp. 1-110, pls. 1-16, August 18. [Concerning species
from Cocos Island, see pp. 30, 36, 42, 53, 58, 73, 103. ]

KISENMANN, E.
1955. The species of middle American birds. Transactions of the
Linnaean Society of New York, vol. 7, pp. I-VI, 1-128, April.
| Concerning species from Cocos Island, see pp. 68, 89, 102.]

Emery, C.
1919. Formiche dell’ Isola Cocos. Rendiconto delle sessioni della R.
Accademia della Scienze dell’ Instituto di Bologna, Classe di
Scienze Fisiche, Nuova serie, vol. 23 (1918-19), pp. 36-40.
| Concerning species from Cocos Island, see p. 40. |

Faxon, W.

1893. Reports on the dredging operations off the west coast of Central
America to the Galapagos, to the west coast of Mexico, and in
the Gulf of California. .. . by the U.S. Fish Commission
steamer Albatross, during 1891. ... VI. Preliminary descrip-
tions of new species of Crustacea. Bulletin of the Museum of
Comparative Zodlogy at Harvard College, vol. 24, no. 7, pp.
149-220, August.

1895. Reports on an exploration off the west coasts of Mexico, Central
and South America, and off the Galapagos islands, in charge
of Alexander Agassiz, by the U.S. Fish Commission steamer
Albatross, during 1891, Lieut. Commander Z. L. Tanner,
U.S.N., commanding. XV. The stalk-eyed Crustacea. Memoirs
of the Museum of Comparative Zodlogy at Harvard College,
vol. 18, pp. 1-292, pls.. A=K, 1-56, 57 (map), “Alp ior
a list of 32 species from off Cocos Island, see p. 257. |

FLEISCHMANN, J.

1935. Footsteps in the sea. G. P. Putnam’s Sons, New York, pp. 1-286,
frontispiece, 116 halftone illustr. (on 46 unnumbered plates),
maps on tront and back lining paper. [Concerning Cocos
Island, see pp. 21-38 and illustr. 5-11. ]

VoLt. XXXII] HEHERTLEIN: BIOGEOGRAPHY OF COCOS ISLAND 267

Fopor, N.
1962.

Foreu, A.
1902.

1908.

The Cocos Island mystery. Tomorrow, vol. 10, no. 1, pp. 71-80,

Winter [issued January |.

(Juatre notices myrmécologiques. Annales de la Société Ento-

mologique de Belgique, tome 46, pp. 170-182, May 1. [See
“TIT Fourmis de l’ile de Coco, ete.,” pp. 176-179. |

Fourmis de Costa-Riea récoltées par M. Paul Biolley. Bulletin

de la Société Vaudoise des Sciences Naturelle, ser. 5, vol. 44,
no. 162, pp. 35-72, March.

Fow ter, H. W.
1932. The fishes obtained by the Pinchot South Seas Expedition of

1929, with description of one new genus and three new species.
Proceedings of the U.S. National Museum, vol. 80, art. 6, publ.
no. 2906, pp. 1-16, figs. 1-4 in text, February 16. [Conecern-
ing species from Cocos Island, see pp. 1, 6, 9, 12, 15. |

1938. The fishes of the George Vanderbilt South Pacifie Expedition.

1937. Monographs of the Academy of Natural Sciences of
Philadelphia, no. 2, pp. I-V, 1-349, frontispiece (map), pls.
1-12, October 14. [List of species known from Cocos Island,
pp. 248-261.]

FRASER, C. McL.
1938. Hydroids of the 1932, 1933, 1935, and 1939 Allan Hancock Pa-

1943,

cifie Expeditions. Allan Haneock Pacifie Expeditions, vol. 4,
no. 3, pp. 129-153, pls. 19-21, October 20. | Concerning species
from Cocos Island, see pp. 132, 133. |

General account of the scientific work of the Velero IIT in the

eastern Pacific, 1931-41. Pt. II. Geographical and Biological
Associations. Allan Hancock Pacifie Expeditions, vol. 1, no. 2,
pp. 49-258, pls. 17-128, December. [Concerning Cocos Island,
see especially p. 161, pls. 78-80.| General account of the
selentifie work of the Velero III in the Eastern Paeifie,
1931-41. Pt. III. A ten-year list of the Velero IIT collecting
stations. Allan Haneock Pacifie Expeditions, vol. 1, no. 3,
pp. 259-431, charts 1-115, December, 1943. An appendix of
collecting stations of the Allan Hancock Foundation for the
year 1942. [Concerning Cocos Island, see pp. 279, 313, chart
78, p. 408. ]

1948. Hydroids of the Allan Hancock Pacifie Expeditions since March,

1938. Allan Hancock Pacific Expeditions, vol. 4, no. 5, pp.

268 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SEr.

179-335, April 21. [Concerning species from Cocos Island,
see pp. 226, 250. |

FREEMAN, O. W.
1951. Geography of the Pacific. John Wiley & Sons, Ine., New York,
pp. I-XII, 1-573, figs. 1-156, + map (frontispiece). [See
“Tslands of the Eastern and Northern Pacific,” pp. 522-544,
figs. 149-152. “Cocos Island,” pp. 532-533. |

GARTH, J. S.
1946. Distribution studies of Galapagos Brachyura. Allan Haneoek
Pacific Expeditions, vol. 5, no. 11, pp. 603-638, charts 1-10,
June 14. [Concerning species from Cocos Island, see pp. 621,
622, 623. |

1958. Brachyura of the Pacific Coast of America. Oxyrhynehia. Allan
Haneock Pacific Expeditions, vol. 21, pt. 1, I-XII, 1-499
[+ 1 blank] ; pt. 2, pp. 501-854, pls. A—Z 4, 1-55, July 25.

GERHARD, P.
1960. Pirates on the west coast of New Spain 1575-1742. Arthur H.
Clark Company, Glendale, California, pp. 1-274, 10 illustr.
| Concerning Cocos Island, see pp. 53, 104, 107, 113, 157, 163,
218, 222.)

GERMAIN, L.

1934. Etudes sur les faunes malacologiques insulaires de l’?Océan Pa-
cifique. [In] Contribution a ]’Etude du Peuplement Zoo-
logique et Botanique des Iles du Pacifique. Société de Biogéog-
raphie, mém. 4, pp. 89-153, ‘“‘eartes” 1-4. [Concerning the
mollusean fauna of Cocos Island, see p. 153, footnote 2. |

GirrorpD, E. W.
1913. Expedition of the California Academy of Sciences to the Galapa-
gos Islands, 1905-1906. No. VIII. The birds of the Galapagos
Islands, with observations on the birds of Cocos and Clipper-
ton islands (Columbiformes to Pelecaniformes). Proceedings
of the California Academy of Sciences, fourth ser., vol. 2, pt.
1, pp. 1-182, pls. 1-7, August 11.

1919. Expedition of the California Academy of Sciences to the Gala-
pagos Islands, 1905-1906. No. XIII. Field notes on the land
birds of the Galapagos Islands and of Cocos Island, Costa Riea.
Proceedings of the California Academy of Sciences, fourth
ser., vol. 2, pt. 2, no. 13, pp. 189-258, June 16. [Concerning
species from Cocos Island, see pp. 193, 195, 200, 205, 216, 242.]

Vout. XXXII] HERTLEIN: BIOGEOGRAPHY OF COCOS ISLAND 269

GISSLER, A.

My twenty years on Cocos Island. Manuscript in Foul Anchor
Archives, Rye, New York. [According to R. I. Nesmith,
1958. |
GODWIN, G.S.

1931. Vancouver. A life. 1757-1798. D. Appleton & Co., New York,
pp. I-XI, 1-308, frontispiece, 10 illustrations, 3 charts. [Con-
cerning Cocos Island, see pp. 138, 262. |

GOULD, J.

1843. Nine new birds, collected during the recent voyage of H.M.S.
Sulphur. Proceedings of the Zoological Society of London for
1843 (Part 11), pp. 103-108, December. [Concerning species
from Cocos Island, see pp. 104, 105. |

1959. Pectinidae of the eastern Pacific. Allan Hancock Pacific Expedi-
tions, vol. 23, pp. I-VIII, 1-308, pls. 1-57, September 25.
{Concerning species from Cocos Island, see pp. 30, 36, 142,
155. ]

GREY, Z.

1925. Tales of fishing in virgin seas. Harper & Brothers Publication,
New York and London, pp. I-VII, 1-216, pls. 1-112 (also
vignettes by Lilian W. Smith). [Cocos Island, pp. 12-42, pls.
1-13. ]

GupgE, G. K.

1903. Deseriptions of some new forms of helicoid land-shells. Proceed-
ings of the Malacological Society of London, vol. 5, no. 4, pp.
262-266, pl. 7, April. [Concerning species from Cocos Island,
see p. 265. ]

GurEypoN, H. L., (Comte de)

1948. Report on Cocos. Pacific Discovery. Published by the California
Academy of Sciences, vol. 1, no. 6, pp. 8-14, 4 halftone illustr.,
November—December. (Translation by T. R. Kelley).

Hac, JANET

1960. The Porecellanidae (Crustacea Anomura) of the eastern Pacific.
Allan Hancock Pacific Expeditions, vol. 24, pp. I-VII, 1-440,
pls. 1-41, March 1.
Hausteap, B. W., and N. C. BUNKER

1953. Report on an investigation of poisonous and venomous fishes at
Cocos, Galapagos and La Plata islands during December 4,

270 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SrEr.
1952 to 28th of January 1953. Office of Naval Research Con-
tract no. NONR-205 (00), pp. 1-14, February 16, 1953.
| Mimeographed.| [Concerning Cocos Island, see pp. 3-6. |

HausteaD, B. W., and D. W. SCHALL
1956. A report on the poisonous fishes captured during the Woodrow
G. Krieger Expedition to Cocos Island. Pacifie Science, vol.
10, no. 1, pp. 103-109, fig. 1 (map), January.

Hancock, R. H., and J. A. WESTON
1960. The lost treasure of Cocos Island. Thomas Nelson & Sons, Edin-
burgh; New York; Toronto, pp. 1-325, 16 halftone illustra-
tions. [Contains a summary of main expeditions to recover
treasure. |
Hanna, G D., and L. G. HERTLEIN
1938. Land and brackish water Mollusea of Cocos Island. Allan Han-
eock Pacifie Expeditions, vol. 2, no. 8, pp. 123-135, fig. 1,
August.

Hanna, G D., and A. M. Strone
1949. West American mollusks of the genus Conus. Proceedings of the
California Academy of Sciences, fourth ser., vol. 26, no. 9, pp.
247, 322, pls. 5-10, text figs. 1-4, Toney 28. [Concerning

species from Cocos Island, see pp. 269, 271, 272, 305. ]

HARTMAN, OLGA

1939a. The Polyehaetous Annelids collected on the presidential cruise
of 1938. Smithsonian Miscellaneous Collections, vol. 98, no.
13 (publ. 3538), pp. 1-22, figs. 1-3, June 9. [Concerning
species from Cocos Island, see especially p. 3. |

1939b. Polychaetous Annelids. Part I. Aphroditidae to Pisionidae.
Allan Hancock Pacifie Expeditions, vol. 7, no. 1, pp. 1-155,
pls. 1-28, August 28. [Concerning species from Cocos Island,
see pp. 10, 16, 17, 59, 61, 74.]

1940. Polychaetous Annelids. Part II. Chrysopetalidae to Goniadidae.
Allan Haneock Pacific Expeditions, vol. 7, no. 3, pp. 173-287,
pls. 31-44, February 20. [Concerning species from Cocos
Island, see pp. 190, 205.]

1959. Catalogue of the Polyechaetous annelids of the world. Part
I. Allan Haneock Foundation Publications, Occasional Paper
no. 23, pp. I-VI, 1-354; pt. II, pp. I-V, 855-628, December 30.

HEERE, A. W.
1936. Fishes of the Crane Pacifie Expedition. Reports on results of

VoL. XXXII] HHERTLEIN: BIOGEOGRAPHY OF COCOS ISLAND 271

the Crane Pacific Expedition. Field Museum of Natural His-
tory, Zoological Series, vol. 21, publ. 353, pp. 1-472, figs. 1-50
in text, April 15. [Concerning species from Cocos Island, see
pay]

HEMDEMANN, O.

1901. Papers from the Hopkins Stanford Galapagos Expedition, 1898—
1899. Entomological Results (1): 1. Hemiptera. Proceedings
of the Washineton Academy of Sciences, vol. 3, pp. 363-370
(Hemiptera, pp. 364-370), August 28. [Concerning species
from Cocos Island, see p. 370. |

HELLER, E.

1903. Papers from the Hopkins Stanford Galapagos Expedition, 1898—
1899. XIV. Reptiles. Proceedings of the Washington Acad-
emy of Sciences, vol. 5, pp. 39-98, February 26. [Concerning
species from Cocos Island, see p. 95. |

HELLER, E., and R. E. SNoperass

1903. Papers from the Hopkins Stanford Galapagos Expedition, 1898—
1899. XV. New Fishes. Proceedings of the Washington Acad-
emy of Sciences, vol. 5, pp. 189-229, pls. 2-20, 1 fig. in text,
September 12. [Concerning species from Cocos Island, see pp.
204, 210, 211.]

HERTLEI, L. G.

1932. Mollusks and barnacles from Malpelo and Cocos islands. Nauti-
lus, vol. 46, no. 2, pp. 48-45, October (issued October 22).

1935. The Templeton Crocker Expedition of the California Academy
of Sciences, 1932. No. 25. The Recent Pectinidae. Proceed-
ines of the California Academy of Sciences, fourth series, vol.
31, no. 25, pp. 301-528, pls. 18 and 19, September 26. [Con-
cerning species from Cocos Island, see p. 319. |

1937. A note on some species of marine mollusks occurring in both
Polynesia and the Western Americas. Proceedings of the
American Philosophical Society, vol. 78, no. 2, pp. 303-812, pl.
1, and map. [Concerning species from Cocos Island, see pp.
306, 307. |

HERTLEIN, L. G., and W. K. EMERSON

1953. Mollusks from Clpperton Island (eastern Pacific) with the
description of a new species of gastropod. Transactions of the
San Diego Society of Natural History, vol. 11, no. 13, pp. 345—
364, pls. 26, 27, July 22.

1957. Additional notes on the invertebrate fauna of Clipperton Island.

272 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

American Museum Novitates, no. 1859, pp. 1-9, December 6.

Hert Len, L. G., and U. 8. Grant, IV
1944. The Cenozoic Brachiopoda of western North America. Publi-
eations of the University of California at Los Angeles in
Mathematical and Physical Sciences, vol. 3, pp. I-VI, 1-236,
pls. 1-21, 34 figs. in text, November 4. [Concerning species
from Cocos Island, see p. 60. |

HoupHaus, K.

1934. Die Verbreitung der Insekten auf den Inseln des Pacifischen
Oceans. |In] Contribution a L’Etude du Peuplement Zoo-
logique et Botanique des Iles du Pacifique. Société de Biogéog-
raphie, mém. 4, pp. 201-218, figs. 1-3. [Concerning Cocos
Island, see pp. 204, 208. |

HoutuHuls, L. B.
1952. A general revision of the Palaemonidae (Crustacea Decapoda
Natantia) of the Amerieas. II. The subfamily Palaemoninae.
Allan Haneoeck Foundation Publeations, Occasional Paper
no. 12, pp. 1-396, pls. 1-55, June 19. [Concerning species
from Cocos Island, see pp. 6, 7, 112, 113, 130; 131i
TG, Atal

Hourtuuis, L. B., and F. ALEJANDRO VILLALOBOS
1961. Panulirus gracilis Streets y Panulirus inflatus (Bouvier), dos
especies de langosta (Crustacea, Decapoda) de la Costa del
Pacifico de América. Anales del Instituto de Biologia (Uni-
versity of Mexico), tomo 32, nos. 1 and 2, pp. 251-276, pls.

1-3, 1 fig. in text, 1 map.

Howarp, A. D.
1952. Pteropods collected by the Allan Hancock Foundation. Minutes
of the Conchological Club of Southern California, no. 121, pp.
12-14. July-August. [Concerning species from Cocos Island,

see p. 13.]

Howe, M. A.

1934. The Templeton Crocker Expedition of the California Academy
of Sciences, 1932. No. 17. The Hepaticae (chiefly Riecia and
Anthocerotaceae) of the Galapagos Islands and the coast and
islands of Central America and Mexico. Proceedings of the
California Academy of Sciences, fourth series, vol. 21, no. 17,
pp. 199-210, pl. 7, October 26. [Concerning species from

Cocos Island, see p. 206. ]

VoL. XXXII] HERTLEIN: BIOGEOGRAPHY OF COCOS ISLAND 273

INGRAM, W. M.
1945. An extension of the range of Cypraea rashleighana Melvill.
Nautilus, vol. 58, no. 3, pp. 106-107, January.

1947a. Fossil and recent Cypraeidae of the western regions of the
Americas. Bulletins of American Paleontology, vol. 31, no.
120, pp. 43(1)-124(82), pls. 5(1)-7(3), May 2. [Concern-
ing species from Cocos Island, see pp. 58(16), 74(32),
76(34).|

1947b. Check list of the Cypraeidae occurring in the western hemi-
sphere. Bulletins of American Paleontology, vol. 31, no. 122,
pp. 137(1)—161(25), May 23. | Concerning species from Cocos
Island, see pp. 147(11), 148(12).]

1948. The eypraeid fauna of the Galapagos Islands. Proceedings of
the California Academy of Sciences, fourth ser., vol. 26, no. 7,
pp. 185-145, June 28. [Concerning species from Cocos Island,
see p. 140. ]

1951. The living Cypraeidae of the Western Hemisphere. Bulletins of
American Paleontology, vol. 33, no. 136, pp. 125(1)-178(54)
pls. 21(1)-24(4), Mareh 24. [Concerning species from Cocos

Island, see pp. 180(6), 152(28), 155(31).]

JOHNSTON, I. M.
1931. The flora of the Revillagigedo Islands. Proceedings of the Cali-
fornia Academy of Sciences, fourth ser., vol. 20, no. 2, pp.
9-104, November 18. |Concerning species from Cocos Island,
see p. 35.|

Keen, A. Myra
1958. Sea shells of tropical west America. Marine mollusks from
Lower California to Colombia. Stanford University Press:
Stanford, California, pp. I-VIII, 1-624, illustr., 10 pls. col.
[| Concerning species from Cocos Island, see pp. 72, 406, 424. ]
KIRKENDALE, G.
1902-1903. In search of private treasure. Wide World Magazine, vol.

10, p. 452.
1904. In search of a treasure island. Wide World Magazine, vol. 12,
ps 612.

KLAUSEWITz, W.
1958. Ergebnisse der Galapagos—Expedition 1953-1954 des Instituts
fiir Submarine Forschung, Vaduz (Leitunge: Dr. H. Haas).
Fische aus dem Atlantik und Pazifik. Senckenbergiana Bio-

274 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser.

logiea, Bd. 39, no. 1-2, pp. 57-84, figs. 1-7, March 31. [Con-
cerning species from Cocos Island, see p. 73.]

KosBeut, W.
1899. Die Fauna der Cocosinsel. Nachrichtsblatt der Deutschen Mala-
kozoologischen Gesellschaft, Jahrg. 31, nos. 1 and 2, pp.
26-28, January and February.

Kuwert, A. [F.]
1897. Die Passaliden dichotomisch bearbeitet. Zweiter Theil. Die
Arten. Novitates Zoologicae, Tring, vol. 4, no. 2, pp. 274-306.
[| Coneerning species from Cocos Island, see p. 301. |

Lack, D.

1945. The Galapagos Finches (Geospizinae). A study in variation.
Occasional Papers of the California Academy of Sciences, no.
21, pp. I-VII, 1-158, pls. 1-4, map (frontispiece), figs. 1-4 in
text, May 30. [Concerning species from Cocos Island, see pp.
19126, £29) pl. 4 fies. 6.1

1953. Darwin’s Finches. Scientific American, vol. 188, no. 4, pp. 66-68,
70, 72, figs. 1-14 (numbered), 3 figs. (not numbered), April.
[Concerning species from Cocos Island, see pp. 67, 72, fig. 8
(p. 66). |

Lapp, H. 8.
1960. Origin of the Pacific island mollusean fauna. American Journal
of Science, Bradley Volume, vol. 258—A, pp. 1387-150, figs. 1-3.

LALICKER, C. G., and IRENE McCuLiocu
1940. Some Textulariidae of the Pacific Ocean. Allan Hancock Pacifie
Expeditions, vol. 6, no. 2, pp. 115-143, pls. 18-16, January 10.
[Concerning species from Cocos Island, see pp. 119, 126, 127,
137, 140. ]
Lamp, D., (in collaboration with June Cleveland)
1938. Enchanted vagabonds. Harper Bros.: New York; London,
I-VI{[+ 1] 1415, 1-23 halftone illustrations, maps 1-13.
[See Chapter 26; “The last adventure—Cocos Island,” pp.
370-409, 2 maps. ]

Lamp, D., and GINGER LAMB
1939. 16,000 miles on $4.20. Pt. 1. Popular Mechanies, vol. 72, no. 2,
pp. 226-229, 118A, 119A, 11 illustr., 1 map, August; pt. 2.
vol. 72, no. 3, pp. 381-383, 116A, 8 illustr., September.
LELouP, E.
1956. Reports of the Lund University Chile Expedition 1948-1949. 27.

Vou. XXXII] HERTLEIN: BIOGEOGRAPHY OF COCOS ISLAND 275

Polyplaecophora. Lunds Universitets Arsskrift, N.F., Avd. 2,
Band 52, no. 15 (Kungl. Fysiografiska Sallskapets I Lund
Handlingar, N.F., Band 67, no. 15), pp. 1-93(-+ 1), figs. 1-53
in text, September 27. [Concerning species from Cocos
Island, see p. 246. |

Ligvre, M. D.

1893. Une ile déserte du Pacifique. L’Ile des Cocos (Amérique). Re-
vue de Géographie (Paris), tome 32, Livraison de Mai, pp.
349-357, 2 Illustr.; Livraison de Juin, pp. 416-422; tome 33,
Livraison de Juillet, pp. 34-41.
Licut, S. F.

1935. The Templeton Crocker Expedition of the California Academy
of Sciences, 1932. No. 20. The termites. Proceedings of the
California Academy of Sciences, fourth ser., vol. 21, no. 20,
pp. 233-258, pls. 9-10, figs. 1-10 in text, April 3. [Concern-
ing species from Cocos Island, see especially pp. 254, 235, 239. |
LonestaFF, F. V., (Major)
1941. Esquimalt Naval Base. A history of its work and its defences.
Distributed by the Victoria Book & Stationery Company, Ltd.,
Victoria, British Columbia, pp. 1-189, frontispiece and _ pls.
I-XII. [Concerning Cocos Island, see remarks concerning
Henry St. Ledger Bury Palliser, p. 126. Palliser visited Cocos
Island in 1897 on his flagship Imperieuse ; in 1903 on a steam-
ship Lytton; and in 1905 on the yacht Veronique. |
MAcDONALD, G. A.
1949. Hawaiian petrographic province. Bulletin of the Geological So-
ciety of America, vol. 60, no. 10, pp. 1541-1596, figs. 1-11,
October. [Concerning Cocos Island, see pp. 1588, 1589, and
me IE
Manaspina, DoN ALEJANDRO

1885. Viaje politico-cientifico alrededor del mundo por las corbetas
descubierta y atrevida al mando de los eapitanes de navio D.
Alejandro Malaspina y Don José de Bustamente y Guerra
desde 1789 4 1794, publicado con una introduecién por Don
Pedro de Novo y Colson. .. . Madrid, impr. de la viuda é hijos
de Abienzo, 4 [title] pp., I-XXXI, [4 pp.], 1-681, 1 portrait,
6 pls., tables, fold. map. [Concerning Cocos Island, see pp.
120-121.] [Title on cover, “La Vuelta al mundo por las
Corbetas Descubierta y Atrevida al mando del Capitan de
Navio D. Alejandro Malaspina desde 1789 4 1794, publicado
con una introdueccion por Don Pedro de Novo y Colson.”’|

276 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4tH SER.

MARTENS, E. VON

1890-1901. Land and freshwater Mollusea. [Edited by Godman, F. D.,
and Salvin, O.] Biologia Centrali-Americana. London, pp.
I-XXVIII, 1-706, pls. 1-44.

1898. Landschencken von der Cocos-Insel. Sitzungs-Bericht der Gesell-
schaft Naturforschender Freunde zu Berlin, Sitzung vom 15
November, 1898, pp. 156-160, November.

1902a. Einige Sechnecken der Cocos Insel. Sitzungs-Bericht der Gesell-
schaft Naturforschender Freunde zu Berlin, Sitzung vom 18
Marz, 1902, pp. 59-62, June.

1902b. Die Meeres-Conchyhen der Cocos-Insel. Sitzungs-Bericht der
Gesellschaft Naturforschender Freunde zu Berlin, Sitzung
vom 17 Juni, 1902, pp. 187-141, June.
MERRILL, G. P.

1895. Reports on the dredging operations off the west coast of Central
America to the Galapagos, to the west coast of Mexico, and in
the Gulf of California, in charge of Alexander Agassiz ... V.
Report upon rocks collected from the Galapagos Islands.
Bulletin of the Museum of Comparative Zoology, vol. 16, no.
13 (Geol. Ser., vol. 2), pp. 235-237, July. [Concerning “Ande-
site (?)” from near Chatham Bay, Cocos Island, see p. 236. ]
MerepitH, DeWitrr
1939. Voyages of the Velero III. A pictorial version with historical
background of scientific expeditions through tropical seas to
equatorial lands aboard M/V Velero III. Published privately,
pp. 1-286, table of contents and index, illustr. [Concerning
Cocos Island, see pp. 86 (lower fig.), 171-173. |
Mo.uina, F'.

1851. Bosquejo de la Republica de Costa Rica, seguido de apuntamien-
tos para su historia. Con varios mapas, vistas y retratos.
Imprenta de S. W. Benedict, no. 16 Calle de Spruce, Nueva
York, pp. 1-128, 2 fold. pl., 3 port., 8 fold map, 1 fold. plan.
| Concerning Cocos Island, see p. 27.]
MonTMORENCY, H. DE

1904. On the track of a treasure. The story of an adventurous expe-
dition to the Pacific Island of Cocos in search of a treasure of
untold value hidden by pirates. Hurst & Blackett, London,
pp. I—X, 1-290, 22 illustr., 1 chart.

Morre., B., JR., (Captain)

28) aa yarne si. = . . 2 Z 1
1832. A narrative of four voyages, to the South Sea, north and south

VoL. XXXIT]

HERTLEIN: BIOGEOGRAPHY OF COCOS ISLAND 277

Pacific Ocean, Chinese Sea, Ethiopie and southern Atlantic
Ocean, Indian and Antarctic Ocean. From the year 1822 to
1831. Comprising critical surveys of coasts and islands, with
sailing directions. J. and J. Harper, New York, pp. I-X XVII,
[28 blank], 29-492, frontispiece (portrait). [Concerning
Cocos Island, see pp. 191-192. |

Morrison, J. P. E.
1946. The nonmarine mollusks of San José Island, with notes on those

of Pedro Gonzalez Island, Pearl Islands, Panama. Smith-
sonian Miscellaneous Collections, vol. 106, no. 6, (publ. 3850),
pp. 1-49, pls. 1-3, September 12. [See especially Melampus
tabogensis, p. 36. |

Mourpny, R. C.
1936. Oceanic birds of South America; A study of species of the re-

Myers, G. S.

lated coasts and seas, including the American Quadrant of
Antarctica, based upon the Brewster-Sanford Collection in the
American Museum of Natural History. New York, vol. 1, pp.
I-XXII, 1-640, pls. 1-38 and 6 pls. (col., unnumbered), figs.
1-61 in text. [Concerning Cocos Island, see pp. 317-319,
lower fig. on pl. 6 (opp. p. 171) ; see also maps, figs. 12, 13, 14,
and 48. |

1941. The fish fauna of the Pacific Ocean, with especial reference

MYERS, G.5S.,

to zodgeographical regions and distribution as they affect
the international aspects of the fisheries. Proceedings of the
Sixth Pacific Science Congress of the Pacific Science Associa-
tion, University of California press, vol. 3, pp. 201-210, 1940,
issued April 17, 1941. [Concerning Cocos Island, see espe-
cially pp. 204-205. |

and C. B. WADE

1941. Four new genera and ten new species of eels from the Pacific

Coast of tropical America. Allan Hancock Pacific Expedi-
tions, vol. 9, no. 4, pp. 65-111, pls. 7-16, June 25. [Concern-
ing species from Cocos Island, see pp. 75, 85.]

NesmiTH, R. I.

1958. Dig for pirate treasure. The Devin-Adair Company, New York,

pp. I-XIV, 1-302, pls. A-G, 9 line cuts including maps on
inside front and back covers. [Concerning Cocos Island, see
pp. 11, 18, 101, 125, 163, 206-256, 264, 287, map p. 251.]

278 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER.

NicHots, J. T., and C. M. BREEDER, JR.
1928. An annotated list of the Synentognathi with remarks on their
development and relationships collected by the Arcturus.
Zoologica, Scientific contributions of the New York Zoological
Society, vol. 8, no. 7, pp. 423-448, figs. 156-176 inclusive, June
11. [Concerning species from Cocos Island, see pp. 424, 426,
427, 443.]
Ousson, A. A.
1956. Studies on the genus Olivella. Proceedings of the Academy of
Natural Sciences of Philadelphia, vol. 108, pp. 155-225, pls.

8-16, October 3. [Concerning species from Cocos Island, see
p. 180.]

OsBURN, R. C.
1950. Bryozoa of the Pacific Coast of America. Part 1, Cheilostomata-
Anasea. Allan Hancock Pacific Expeditions, vol. 14, no. 1,

pp. 1-269, pls. 1-29, June 20. [Concerning species from Cocos
Island, see p. 47. |

po
No)
1
Lh)

Bryozoa of the Pacific Coast of America. Pt. 2, Cheilostomata-
Ascophora. Allan Haneock Pacifie Expeditions, vol. 14, no.
2, pp. 271-611, pls. 30-64, Mareh 20. [Concerning species
from Coeos Island, see pp. 281, 447, 468, 479. |

PALLISER, H., G. BRAWNER, and P. STacHWICK

1932. Six months on a deserted island. The American Magazine, vol.
113, no. 2, pp. 18-21, 182-135, 7 illustrations, February.

Passmore, R. Mc. C.

1895. Informe sobre la Isla del Coco. Memoria presentada al Congreso
Constitutional por el Seeretario de Estado en los despachos
de Guerra y Marina, Gral. don Juan B. Quirés, Tipografia
Nacional, San José, Costa Riea; Marina, pp. 30-84.

Piussry, H. A.

1900. Note on the anatomy of Guppya Hopkinsi Dall. See Dall, W. H.,
1900, p. 105, 4 figs. in text.

Pinsspry, H. A., and C. M. Cooke

1920. Pupillidae (Gastroecoptinae, Vertigininae). Manual of Con-
chology, Second Series, Pulmonata, vol. 25, pt. 100, pp. 225—
404, pls. 19-34, April. [See “Cocos Island Species,” pp. 322—
324. ]

Piuspry, H. A., and H. N. Lowe

1932. West Mexican and Central American Mollusks collected by H.

VoL. XXXII] HEHERTLEIN: BIOGEOGRAPHY OF COCOS ISLAND 279

N. Lowe, 1929-31. Proceedings of the Academy of Natural
Seiences of Philadelphia, vol. 84, pp. 33-144, pls. 1-17, figs.
1-6 in text, May 21. |Concerning species from Cocos Island,
see p. 67.]

Piussry, H. A., and E. G. VANATTA
1902. Papers from the Hopkins Stanford Galapagos Expedition, 1898—
1899. XIII. Marine Mollusea. Proceedings of the Washing-
ton Academy of Sciences, vol. 4, pp. 549-560, pl. 35, Septem-
ber 30. [Concerning species from Cocos Island, see p. 559. |
PINCHOT, G.
1930. To the South Seas. The cruise of the schooner Mary Pinchot to
the Galapagos, the Marquesas, and the Tuamotu Islands, and
Tahiti. The John C. Winston Company, Philadelphia, pp.
I-XIII, 1-500, frontispiece and 246 halftone illustr., 16
vignettes, 3 maps and maps inside front and back covers.
| Concerning Cocos Island, see pp. 56-97, 492. |
Pirrier, H.
1899. Apuntamientos preliminares sobre la Isla de Cocos, posesion
costarricense en el Oceano Pacifico. Ministerio de Fomento,
San José, Costa Rica, Memoria de Fomento presentada al
Congreso Constitucional de 1899, pp. 141-153, 1 map. [See
also N. Chavarria Mora, pp. 155-158, and A. Alfaro, pp. 159-
162. ]

PLUMPTON, J., (Commander)
[1936.] Treasure cruise ; the voyage of the Vigilant to Cocos Island... .
with an introduction by Adm. A. V. Campbell. H. F. and G.
Witherby, London, pp. 1-191, frontispiece, plates and 2 maps.

RATHBUN, Mary J.

1902. Papers from the Hopkins Stanford Galapagos Expedition, 1898—
1899. VIII. Braechyura and Macrura. Proceedings of Wash-
ington Academy of Sciences, vol. 4, pp. 275-292, pl. 12, June
20. [Concerning species from Cocos Island, see p. 281. |

1930. The eaneroid crabs of America of the families Euryalidae,
Portunidae, Atelecyclidae, Caneridae and Xanthidae. United
States National Museum, Bulletin 152, pp. I-X VI, 1-609, pls.
1-230, figs. 1-85 in text, May 29. [Concerning species from
Cocos Island, see pp. 69, 138, 242, 552. ]

RiIcHMOND, C. W.

1902. Note on Pinaroloxias inornata (Gould). Proceedings of the

280 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Biological Society of Washington, vol. 15, pp. 247-248, De-
cember 16.

RIESENBERG, H. E.
1951. The story behind Cocos Island and the “Loot of Lima.” Pacifie
Diseovery, published by California Academy of Scienees, vol.
4, no. 6, pp. 4-15, 15 illustr., November—December.

Rosinson, B. L.
1902. Contributions from the Gray Herbarium of Harvard University.
New Series.—No. XXIV. (Papers from the Hopkins-Stanford
Expedition to the Galapagos Islands.) Flora of the Galapagos
Islands. Proceedings of the American Academy of Arts and
Sciences, vol. 38, no. 4, pp. 77-269, pls. 1-8, October. [Con-
cerning plants from Cocos Island, see pp. 241, 261-263. |

Rosson, G. C.

1929. A monograph of the Recent Cephalopoda based on the collections
in the British Museum (Natural History). Part 1, Oetopo-
dinae. Printed by order of the trustees of the British Museum,
pp. I-XI, 1-236, pls. 1-7, figs. 1-89 in text, July 27. [Con-
cerning species from Cocos Island, see p. 150. |

1932. The same, Part 2, Octopoda, pp. I-XI, 1-359, pls. 1-6, figs. 1-79
in text, January 23. [Concerning species from Cocos Island,
see pp. 103, 197, 240.|

1948. The Cephalopoda Deeapoda of the Arcturus Oceanographic Ex-
pedition, 1925. Zoologiea, Scientific Contributions of the New
York Zoological Society, vol. 33, pt. 3, no. 7, pp. 115-182, figs.
1-15 in text, November 29. [Concerning species from Cocos
Island, see pp. 117, 118, 120, 121, 125, 127, 128, 1130:]

Rogers, 8. R. H.

1931. The Pacific. Thomas Y. Crowell Company, New York, pp. 1—254,

illustr. [Concerning Cocos Island, see pp. 100-105, 2 figs. |
Rose, J. N.

1892. List of plants from Cocos Island. Contributions from the United
States National Herbarium, vol. 1, no. 5(1), p. 135, September
20,

Rose, Ruta

1926. Cocos—A tale of treasure. [In Beebe, W.] The Arcturus ad-
venture, G. P. Putnam’s Sons, New York and London, pp.
250-281, figs. 40-41.

Vou. XXXII] HERTLEIN: BIOGEOGRAPHY OF COCOS ISLAND 281

Rost, HELEN
1955. A report on the Family Arcidae (Peleeypoda). Allan Hancock
Pacific Expeditions, vol. 20, no. 2, pp. 177-248, pls. 11-16,
figs. 79-95 in text, November 10. [Concerning species from
Cocos Island, see p. 227. |
RoTHscHiLp, W.
1902. [Exhibition of and comments upon| specimens of the land-birds
of Cocos Island, midway between the Galapagos Islands and
Costa Rica. Bulletin of the British Ornithologists’ Club, vol.
13, no. 91, p. 6, October 31.

SACHET, Marre-H&LENE, and F. R. FosBera
1955. Island bibliographies. (Micronesian botany, land environment
and ecology of coral atolls, vegetation of tropical Pacific
islands.) Compiled under the auspices of the Pacific Science
Board. Publication 335, National Academy of Sciences—Na-
tional Research Council, pp. I-V, 1-577.
SAVELL, M.
1934. Cocos, perennial goal of treasure seekers. Literary Digest, vol.
117, no. 15, whole no. 2295, pp. 42-43, 2 illustrs., April 14.

ScHILpER, F’. A., and Marta SCHILDER
1938. Prodrome of a monograph on living Cypraeidae. Proceedings of
the Malacological Society of London, vol. 23, pt. 3, pp. 119-
180, unnumbered figs. (p. 125), November 15; pt. 4, pp. 181-
231, 9 maps, Mareh 15. [Concerning species from Cocos
Island, see pp. 176, 197. |
Scumupt, K. P.
1930. Essay on the zoogeography of the Pacific islands. (In Shureliff,
S. N., Jungle Islands). Putnam Press, New York, pp. 270-
292. [Concerning Cocos Island, see pp. 278-280. ]
Scumirt, W. L.
1939a. The presidential cruise of 1938. Explorations and field-work of
the Smithsonian Institution in 1938. Smithsonian Institution,
publ. 3525, pp. 1-14, figs. 1-11 in text. [Concerning Cocos
Island, see p. 12, figs. 1, 6(2), 9, 14.] [Concerning a popular
account of the itinerary of this cruise see Anonymous, 1938b. |
1939b. Deeapod and other erustacea collected on the presidential cruise
of 1938 (with introduction and station data). Smithsonian
Miscellaneous Collections, vol. 98, no. 6, publ. 3531, pp. 1-29,
pls. 1-3, June 15. [Concerning Cocos Island, see especially pp.
Deedee pl. 2ihe; 2:

282 CALIFORNIA ACADEMY OF SCIENCES [Proc. 41TH SrEr.

Scumitt, W., and L. P. ScHULTZ

1940. List of the fishes taken on the presidential cruise of 1938. Smith-
sonian Miscellaneous Collections, vol. 98, no. 25, publ. 3551,
pp. 1-10, January 4. [Concerning species from Cocos Island,
see pp. 4, 6, 8, 9.]

ScHort, C.

1931. Der Peru-Strom und seine nordlichen Nachbargebiete in nor-
maler und anormaler Ausbildung. Annalen der Hydrographie
und Maritimen Meteorologie, Zeitschrift fiir Seefahrt- und
Meereskunde, Herausgegeben von der Deutschen Seewarte
Hamburg, Jahrg. 59, Heft 5, pp. 161-169, figs. 1-2 in text,
Taf. 17, May 15; Heft 6, pp. 200-2138, figs. 3, 4 in text, Taf.
20, 21, 22, June 15; Heft 7, pp. 240-253, July 15.

SEALE, A.

1940. Report on fishes from Allan Haneock Expeditions in the Cali-
fornia Academy of Seiences. Allan Haneock Pacifie Expedi-
tions, vol. 9, no. 1, pp. 1-46, pls. 1-5, April 29. [Concerning
species from Cocos Island, see pp. 16, 17, 22, 23, 24, 27, 29, 35,
45. |

SEYMOUR, GEORGE, (Sir)
1849. Notes on the Island of Cocos, and two of the Galapagos. Journal

of the Royal Geographical Society of London, vol. 19, pp.
20222:
SHOEMAKER, C. R.

1942. Amphipod crustaceans collected on the Presidential Cruise of
1938. Smithsonian Miscellaneous Collections, vol. 101, no. 11,
publ. 3677, pp. 1-52, figs. 1-17, March 5. [Concerning species
from Coeos Island, see pp. 4, 32.|

SHUMWAY, G.

1954. Carnegie Ridge and Cocos Ridge in the east equatorial Paeifie.
Journal of Geology, vol. 62, no. 6, pp. 573-586, figs. 1-7 in
text, November. [Concerning Cocos Ridge, see especially pp.
580-584, figs. 2, 6 and 7. |

SHURCLIFF, S. N.

1930. Jungle islands. The J//yria in the South Seas. The record of the
Crane Pacifie Expedition, Field Museum of Natural History,
Chicago, Illinois. With a scientific appendix by Karl Patter-
son Schmidt. G. P. Putnam’s Sons, New York, London, pp.
I-XV, 1-298, frontispiece (col.), 92 illustr., (8 col.; 2 charts).

Vou. XXXII] HERTLEIN: BIOGEOGRAPHY OF COCOS ISLAND 283

[Zoogeography by Schmidt, pp. 275-292] [Concerning Cocos
Island, see chapter IV, pp. 57-70, 4 pls. ]
SLEVIN, J. R.

1931. Log of the schooner Academy on a voyage of scientific research
to the Galapagos Islands 1905-1906. California Academy of
Sciences, Occasional Papers No. 17, pp. 1-162, pls. 1-16, 1
map, February 14. [Concerning Cocos Island, see pp. 27-32,
and pls. 2 and 3.|

1935. An equatorial wonderland. The Galapagos—a country of great
voleanoes, giant tortoises, and flightless birds. Natural History.
The Journal of the American Museum of Natural History, vol.
36, pp. 375-388, 14 illustrations, December. [Concerning
illustrations taken at Cocos Island, see pp. 383, 388. ]

Snoperass, R. EK.
1902. See Banks, N., 1902, pp. 71-80.
Snoperass, R. E., and E. HELLER

1902. Papers from the Hopkins Stanford Galapagos Expedition, 1898—
1899. XI. The birds of Clipperton and Cocos islands. Pro-
ceedings of the Washington Academy of Sciences, vol. 4, pp.
501-520, September 30. [Concerning species from Cocos
Island, see pp. 504-505. |

1903. Papers from the Hopkins Stanford Galapagos Expedition, 1898—
1899. XVI. Birds. Proceedings of the Washington Academy
of Sciences, vol. 5, pp. 231-372, January 28. [Concerning
species from Cocos Island, see pp. 239, 242, 246, 356. |

1905. Papers from the Hopkins Stanford Galapagos Expedition, 1898—
1899. XVII. Shore fishes of the Revillagigedo, Clipperton,
Cocos and Galapagos islands. Proceedings of the Washington
Academy of Sciences, vol. 6, pp. 333-427, January 31. [38
species reported from Cocos Island, p. 425.]

Soot—RYEn, T.

1955. A report on the family Mytilidae (Peleeypoda). Allan Hancock
Pacific Expeditions, vol. 20, no. 1, pp. 1-174, pls. 1-10, text
figs. 1-78, November 10. [Concerning species from Cocos
Island, see pp. 130, 141. |

SPHON, G. G., JR.

1961. Notes on the Mitridae of the eastern Pacific. 1—Mitra fultoni
K. A. Smith. Veliger, vol. 4, no. 1, pp. 32-86, pl. 7, 1 fig. in text
(map), July 1. [Species recorded from Cocos Island, pp. 34,
35. |

284 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

STEBBING, T. R. R.
1903. Amphipoda from Costa Rica. Proceedings of the United States
National Museum, vol. 26, no. 1341, pp. 925-931, pls. 60, 61,
July 9. [Concerning species from Cocos Island, see p. 928. ]

STEJNEGER, L.

1900. Reports on the dredging operations off the west coast of Central
America to the Galapagos, to the west coast of Mexico, and in
the Gulf of California, in charge of Alexander Agassiz, carried
on by the U.S. Fish Commission steamer Albatross, during
1891, Lieut. Commander Z. L. Tanner, U.S.N., commanding.
XXVIII. Deseription of two new lizards of the genus Anolis
from Cocos and Malpelo islands. Bulletin of the Museum of
Comparative Zoology at Harvard College, vol. 36, no. 6, pp.
161-164, 1 pl., November. [Anolis townsendi described from
Cocos Island, p. 163.]

1903. Description of a new species of gecko from Cocos Island. Pro-
ceedings of the Biological Society of Washington, vol. 16, pp.
3-4, February 21. [Sphaerodactylus pacificus described from
Cocos Island. |

STEWART, A.

1911. Expedition of the California Academy of Sciences to the Galapa-
gos Islands, 1905-1906. II. A botanical survey of the Galapa-
eos Islands. Proceedings of the California Academy of Scei-
ences, fourth ser., vol. 1, pp. 7-288, pls. 1 (map)-19, January
20. [Concerning Cocos Island, see p. 230. ]

1912. Expedition of the California Academy of Sciences to the Galapa-
gos Islands, 1905-1906. V. Notes on the botany of Cocos
Island. Proceedings of the California Academy of Sciences,
fourth ser., vol. 1, pp. 375-404, pls. 31-34, January 19. (Janu-
ary 16 on p. 375).
Strone, A. M., and H. N. Lowe
1936. West American species of the genus Phos. Transactions of the
San Diego Society of Natural History, vol. 8, no. 22, pp. 305-
320, pl. 22, December 7. [Concerning species from Cocos
Island, see p. 310. |
SVENSON, H. K.
1935. Plants of the Astor Expedition, 1930 (Galapagos and Cocos
islands). American Journal of Botany, vol. 22, no. 2, pp. 208-
277, pls. 1-9, February 22. [Includes “Catalogue of Plants,”
various families identified by E. B. Bartram, p. 260; A. Chase,

VoL. XXXII] HEHRTLEIN: BIOGEOGRAPHY OF COCOS ISLAND 285

pp. 260-261; L. B. Smith, p. 261; C. Schweinfurth, pp. 261—
263; H. A. Gleason, pp. 263-264. Concerning Cocos Island,
see pp. 259-265. |

SwartH, H. 8.
1931. The Avifauna of the Galapagos Islands. Occasional Papers of
the California Academy of Sciences, no. 18, pp. 1-299, pl. 1
(map), figs. 1-57 in text, June 29. [Concerning species from
Cocos Island, see pp. 268-271. |

TAMBS, E.
1934. The cruise of the Teddy. Harcourt Brace & Co., New York, pp.
I-XX, 1-237, 30 illustr. [Concerning Cocos Island, see pp.
89-101. ]

TANNER, Z. L.

1893. Report upon the investigations of the U.S. Fish Commission
steamer Albatross from July 1, 1889, to June 30, 1891. [In]
United States Commission of Fish and Fisheries, pt. XVII.
Report of the commissioner for 1889 to 1891, pp. 207-342, pl.
1. [Concerning Cocos Island, see pp. 260-262. |

TayLor, EK. H.

1956. A review of the lizards of Costa Riea. University of Kansas
Science Bulletin, vol. 38, pt. 1, no. 1, pp. 3-822, figs. 1-76,
December 20. [Concerning species from Cocos Island, see pp.
34-38, figs. 5, 6, and pp. 123-126, fig. 33.]

Tuomas, F. J.

1960. Treasure of Cocos Island. Américas, vol. 12, no. 5, pp. 31-34,
4 illustr., May.

1960a. The treasure of Cocos Island. Science Digest, vol. 48, no. 4, pp.
79-82, 2 illustr., October.

THomas, P.

1745. A true and impartial journal of a voyage to the South Seas, and
round the globe, in His Majesty’s Ship the Centurion, under
the command of Commodore George Anson, ... J. Newberry,
London, pp. A2 [1-14], 1-347, ap. 1-39. [Concerning Cocos
Island, see pp. 103, 110-111, ap. p. 31.]

ToMLIN, J. R. LEB.

1927-1929. The Mollusca of the St. George Expedition. (1). The Pa-
cifie Coast of S. America. Journal of Conchology, vol. 18, no.
6, pp. 153-170, December, 1927; vol. 18, no. 7, pp. 187-198,

286 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

May, 1928. (II). The West Indies, vol. 18, no. 11, pp. 807—
310, July, 1929.

TOWNSEND, C. H.

1895. Reports on dredging operations off the west coast of Central
America to the Galapagos, to the west coast of Mexico, and in
the Gulf of California, in charge of Alexander Agassiz, carried
on by the U.S. Fish Commission steamer Albatross, during
1891, Lieut. Commander Z. L. Tanner, U.S.N., commanding.
XVII. Birds from Cocos and Malpelo islands, with notes on
petrels obtained at sea. Bulletin of the Museum of Compara-
tive Zoology at Harvard College, vol. 27, no. 3, pp. 121-126,
2 pls. (colored), July.

TREADWELL, A. L.
1928. Polychaetous annelids from the Arcturus Oceanographie Expe-
dition. Zoologica, Scientifie Contributions of the New York
Zoological Society, vol. 8, no. 8, pp. 449-485, figs. 177-179
inclusive, December 5. [Concerning species from Cocos Island,
see pp. 450, 472, 478. |
TuRNER, RutH D., and K. J. Boss
1962. The genus lithophaga in the western Atlantic. Johnsonia, vol.
4, no. 41, pp. 81-116, pls. 57-75, September 7. [Concerning
species from Cocos Island, see p. 108. |

VANCOUVER, G., (Captain)

1798. A voyage of discovery to the north Pacific Ocean, and round the
world; in which the coast of north-west America has been
carefully examined and accurately surveyed. Undertaken by
his majesty’s command, principally with a view to ascertain
the existence of any navigable communication between the
north Pacifie and north Atlantic Oceans; and performed in
the years 1790, 1791, 1792, 1793, 1794, and 1795, im the Dis-
covery sloop of war, and armed tender Chatham, under the
command of Captain George Vancouver. London, vol. 3, [6
pp. not paginated] pp. 1-505, pls. XII-X VII. [Concerning
Cocos Island, see pp. 362-376. |

VANDERBILT, W. K.

1927. To Galapagos on the Ara 1926. The events of a pleasure-cruise
to the Galapagos Islands and a classification of a few rare
aquatic findings, including two specimens of a new species of
shark never caught before and here deseribed for the first
time. Privately printed, pp. I-XV, 1-161, 30 illustr. in color,

Vout. XXXII] HERTLEIN: BIOGEOGRAPHY OF COCOS ISLAND 287

65 illustr. in black and white, 2 maps. [Concerning Cocos
Island, see pp. 41-46, 159, and illustr. opp. p. 37 and on p. 45. |
VInTON, K. W.
1951. Origin of life on the Galapagos Islands. American Journal of
Science, vol. 249, no. 5, pp. 356-376, pls. 1-2, figs. 1 and 2 in
text, May. [Concerning Cocos Island, see especially pp. 363-
364, 366-374, and fig. 1.|
Waren, L.
[1699] 1903. A new voyage and description of the Isthmus of America
... by Lionel Wafer; reprinted from the original edition of
1699; ed. by George Parker Winship. The Burrows Brothers
Company, Cleveland, pp. 1-212, frontispiece, 3 pls., 1 map.
[Cocos Island, pp. 32-33, 194-195, reprint pp. 176-177. See
also pp. 379-380, in Volume of a collection of voyages pub-
lished by James and John Knapton, London, 1729. |

WALLACE, A. R.

1876. The geographical distribution of animals with a study of the re-
lations of living and extinct faunas as elucidating the past
changes of the earth’s surface. Maemillan and Company, Lon-
don, vol. 2, pp. I-VIII, 1-607, pls. XIV-XX, 2 maps. [Con-
cerning Cocos Island, see p. 60. |

WASMANN, E.

1902. Termiten, Termitophilen und Myrmekophilen, gesammelt auf
Ceylon von Dr. W. Horn 1899, mit anderem ost—indischen
Material Bearbeitet von E. Wasmann. Zoologische Jahrbicher,
Abtheilung fiir Systematik, Geographie und Biologie der
Thiere, Band 17, Heft 1, pp. 99-164, Tafel 4, 5, October 1.
| Concerning species from Cocos Island, see pp. 139, 140. |

WETMORE, A.
1939. Birds from Clipperton Island collected on the presidential
eruise of 1938. Smithsonian Miscellaneous Collections, vol.
98, no. 22, publ. 3548, pp. 1-6, August 11. [Concerning
species from Cocos Island, see pp. 4, 6.]
WHEELER, W. M.
1919. Expedition of the California Academy of Sciences to the Gala-
pagos Islands, 1905-1906. XV. The ants of Cocos Island.
Proceedings of the California Academy of Sciences, fourth
ser., vol. 2, pt. 2, no. 15, pp. 299-308, June 16.
1933. The Templeton Crocker Expedition of the California Academy
of Sciences, 1932. No. 6. Formicidae of the Templeton

288 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 47TH SrEr.

Crocker Expedition. Proceedings of the California Academy
of Sciences, fourth ser., vol. 21, no. 6, pp. 57-64, Mareh 22.
| Concerning species from Cocos Island, see p. 61. |
WILKINS, H. T.
1934. Modern buried treasure hunters. With a foreword by Sir Mal-
colm Campbell (p. IX), pp. I-X, 1-293, frontispiece and 8

illustr. {Concerning Cocos Island, see “The truth about Cocos
Island” pp. 100-188. |

1940. Panorama of treasure hunting. Romantic adventures in strange
lands and queer places. With a foreword by Admiral Bertram
Chambers, C. B. (pp. [X—XIII). E. P. Dutton & Co., Ine,
New York, pp. I-XX, 1-582, frontispiece, 37 illustr., also
charts and sketches. [Concerning Cocos Island, see pp. 105—
169, 1 chart, 4 halftone illustr. |}

WILLIAMS, F’. X.

1911. Expedition of the California Academy of Sciences to the Galapa-
gos Islands, 1905-1906. III. The butterflies and hawk-moths
of the Galapagos Islands. Proceedings of the California
Academy of Sciences, fourth ser., vol. 1, pp. 289-822, pls. 20,
21, October 7. [Concerning species from Cocos Island, see pp.
295-296) oii

1926. Expedition of the California Academy of Sciences to the Gala-
pagos Islands, 1905-1906. The bees and aculeate wasps of the
Galapagos Islands. Proceedings of the California Academy
of Sciences, fourth ser., vol. 2, pt. 2, no. 18, pp. 847-857, May
21. [Concerning species from Cocos Island, see p. 349. |

Woopty, W.J., and P. HINKLE

1880. History of the buried treasure on Cocos Island, and confession
of the pirates the night before execution, in Kingston, Ja-
maica. Taylor and Nevin, printers and engravers, 534 Com-
mercial Street, San Francisco, pp. 1-32, May 30.

ZIESENHENNE, F. C.

1937. The Templeton Crocker Expedition, X. Echinoderms from the
West Coast of Lower California, the Gulf of California and
Clarion Island. Zoologica, Scientific Contributions of the New
York Zoological Society, vol. 22, pt. 3, pp. 209-239, text figs.
1-2, October 7. [Concerning species from Coeos Island, see
p. 215.]|

1940. New Ophiurans of the Allan Hancock Pacifie Expeditions. Allan
Hancock Pacific Expeditions, vol. 8, no. 2, pp. 9-59, pls. 2-9,

VoL. XXXII] HERTLEIN: BIOGEOGRAPHY OF COCOS ISLAND 289

April 12. [Concerning species from Cocos Island, see pp. 12,
30. |

1942. New Eastern Pacific Sea Stars. Allan Hancock Pacific Expedi-
tions, vol. 8, no. 4, pp. 197-223, October 14. [Concerning
species from Cocos Island, see pp. 209, 212. ]

1955. A review of the genus Ophioderma M. & T. Essays in the
Natural Sciences in honor of Captain Allan Hancock on the
oceasion of his birthday, July 26, 1955. University of South-
ern California Press, Los Angeles, pp. 185-201, November 8.
| Concerning species from Cocos Island, see pp. 193, 198.]

~ a ny) pig . ;
=e awlehkonal

Hoge ey ered

oye

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

FOURTH SERIES

G Dallas Hanna Anniversary Volume

Vol. XXXII, No. 9, pp. 291-313, 18 figs., 1 table May 20, 1963

THE GOOSE LAKE FRAGMENTS
by

C. P. Butler
Physicist, U.S. Naval Radiological Defense Laboratory
Research Associate, Department of Astronomy,

California Academy of Sciences

INTRODUCTION

The Goose Lake meteorite was discovered in Modoe County, California,
by three hunters in the fall of 1938, removed from the site in May, 1939,
and brought to San Francisco, where it was exhibited at the Golden Gate
International Exposition. After the close of the fair it was shipped to the
Smithsonian Institution where it is now on display in the United States Na-
tional Museum.

The original site of the fall was re-examined in 1960, when a large
meteoritic fragment field was discovered in and around the impact point.
These fragments have raised several new questions to add to those already
associated with this remarkable meteorite. This paper is the first report
on the distribution, morphology, and metallurgy of these particles, and
some speculations on their relationships to the cavities and low terminal
velocity of the main mass.

[ 291 ]

Marine Biological Laboratory
a Bd BA = 92 5 a Se

MAY 29 1963
WOODS HOLE, MASS.

292 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47H Ser.
BACKGROUND STUDIES

The Goose Lake meteorite is unique among most of the existing meteor-
ites, principally because of its peculiar cavities, the origin and significance
of which have been controversial for a number of years. Several of these
are clearly shown in figure 1.

A typical cavity in this iron is 11 centimeters deep, 5 centimeters in
diameter at the aperture, with a slightly larger diameter at the bottom
of the hole. Width to depth ratios range from 0.25 to 1.10. In one ease, a
hole forms a tunnel completely through the mass. Around the edges of the
cavities, there is an overhanging lip of deformed metal with serrated radial
erooves extending back a short distance. These overturned edges are not
apparently related to the origin of the cavities but may be due to some
thermal action during flight through the atmosphere.

Three possibilities have been considered by Henderson and Perry

(1958a) to account for such cavity formation during flight:

Ficure 1. The Goose Lake meteorite, showing its cavities and two typical over-
turned-rim formations. This photograph was made soon after arrival at Mills
College, Oakland, California, and before exhibition at Treasure Island.

VoL. XXXIT] BUTLER: GOOSE LAKE FRAGMENTS 293

1. Burned out minerals. Troilite and carbon inclusions occur in rounded
masses in iron meteorites, often an inch or more in diameter, but none have
been found approaching the dimensions of the cavities. The melting tem-
peratures of troilite and schreibersite are close to 1000°C. but the micro-
structure of slices cut through a cavity does not indicate that temperatures
this high were ever reached.

2. The energy trap. If a small crevice or hole pre-existed, particles
striking this depression would erode the sides, thus deepening the hole since
they would not be deflected from the surface. If such a condition existed
on the leading edge or forward face of the meteorite when the stagnation
temperatures and adiabatic pressures were the highest, the rim of the holes
would certainly heat just as rapidly and erode away leaving a shallow de-
pression, instead of a deep hole.

3. Wave action within a cavity. It might be assumed that during the
ablation period, spallation of the surfaces would free small particles which
would be caught in a pre-existing hole and enlarge them by repeated im-
pacts around the interior. This is analogous to the holes formed in solid
rock on river bottoms, where stones are rolled around by the water currents
forming deep circular holes. It is hard to visualize how such a mechanism
could remove the volume of metal required in the short time available dur-
ing its fall.

There is ample evidence that the cavities are not due to weathering,
principally because the cavities are uniformly distributed over the surface
which would not be the case had the iron rested on the ground long enough
for oxidation on the under side to erode away large masses. Furthermore,
the Modoe Plateau is semi-arid, with an annual rainfall of about 12 inches,
approximately half of which falls as snow.

Henderson and Perry (1958a) concluded that the cavities existed be-
fore the iron entered the atmosphere and further that “this meteorite is
not much smaller now than when it formed in some primordial body ; that
no large piece broke off during flight and that this is probably not a portion
of the metallic core of the planetlike body where it was formed.”’

All the evidence available indicates that the iron landed gently. When
found, about half the mass protruded above the surface of the ground, just

-as though it had been dumped from a truck. Measurements of the crater
depth made in 1961 showed that the distance from the surface to bedrock
was about 9 inches. There are no discernible impact scars on the iron, even
though it fell on hard basaltic rock. The sand and detritus are so thin in this
area that one can easily scrape away the surface material by hand, exposing
the bedrock underneath.

Linsley (1939), who was present at the time it was recovered, reported:
“There was no evidence of shattered rocks indicating a recent fall and

294 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

there were no skid marks to show that the meteorite had swept along the
surface and come to rest where it was found. There was only a slight de-
pression in which it rested, which appeared to be due in part to wind erosion
as the air currents had eddied about it. A marmot seeking a bombproof
shelter had made a home under it.”

Something acted like a cushion at the end of its flight, and here it is
tempting to suggest that it fell in winter into a deep snow bank. The total
depth of accumulated snow for Modoe County is estimated to be 50 inches,
but it should be noted that the deep snows do not occur in flat denuded
windblown areas. Snow banks deep enough to cushion such a mass ean ac-
cumulate only in the lee of canyons or in heavily wooded regions.

The cavities may be related to its slow descent as suggested by Cornish
in Henderson and Perry (1958a), “The large cavity which made an opening
through this iron probably would give the body considerable spin during
its fall. This spin would generate enough lift to reduce the velocity of the
fall.” If an effect of this kind is possible, there is still the question of ex-
plaining the absence of intense ablation heating, for there are no evidences
of either a fusion crust on the surface, or granulation of the Neumann lines
which extend almost unbroken to the very edge.

MeETeEorITIC FRAGMENTS

At the time the Goose Lake meteorite was discovered, the concept that
molten droplets of iron from a glowing meteor would settle to the earth and
could be recovered from the soil was not widely recognized, although
Spencer (1933) had reported evidence of metallic rain of meteoric origin
in the Henbury Craters in Australia. At that time ablation heating and the
re-entry problems associated with missiles and satellites was not yet a
serious scientific problem.

Professor Leonard (1940), who was present at the time the Goose Lake
meteorite was removed, reported, “Although further and more conclusive
evidence of the impact as well as other meteorites, were diligently searched
for in the neighborhood, none was found.” Ninninger (1956), another mem-
ber of the recovery party apparently did not use the magnetie cane which
he says had “been an essential part of our field equipment since 1933.”

Since 1939, a great deal of work has been done on recovering meteoritic
particles from the soil, both in Arizona around the Canyon Diablo Crater
and at the site of the great Sikhote-Alin fall in 1947 in eastern Siberia.
Krinoy (1960) and his colleagues recovered large numbers of meteoritie
particles of this fall which were seattered over an area of several square
miles. Small globular droplets of nickel-iron ranging in size from 8 microns
to 0.10 millimeters in diameter were found seattered over a very wide area.

VoL. XXXIT] BUTLER: GOOSE LAKE FRAGMENTS 295

These are true ablation products, blown off the melting surface of large
masses during flight, and which subsequently condensed in little spheres.
Their original microstructure and composition are not identical with the
main masses because their temperatures were at least 1500°C. in an oxi-
dizing atmosphere. Krinov calls these ablation products ‘“‘meteorie dust.”
Besides the little droplets, they also found a great many fragmentation
particles, ranging in size from tiny flakes to pieces weighing several pounds.
Polished sections of these pieces showed that they retain both the micro-
structure and composition of the main body. These fragments are not abla-
tion products, but are produced by mechanical forces set up by shock
waves in the main mass during its flight. He calls these fragments “meteori-
tie dust,” and from his descriptions they are apparently identical to the
particles which Ninninger found around the Canyon Diablo Crater in
Arizona and which he ealls “sluglets.”” The Canyon Diablo is estimated to
have fallen some 50,000 years ago, a sufficiently long time so that one
would intuitively assume that tiny fragments of the nickel-iron would
either be completely rusted away, or would be so dispersed by weathering
that recovery would be very unlikely. Yet, in Ninninger’s words, “. . . the
idea that all small particles resulting from disintegration of large meteor-
ites would undergo immediate oxidation was seriously in error.”

An important difference between the Goose Lake and other fragment
producing falls is that there is no evidence that its impact velocity was
high. The Sikhote-Alin was seen to break up during flight and its frag-
ments were found seattered over several square miles. The impact velocity
of the Canyon Diablo was sufficient to produce complete fragmentation ac-
companied by an explosion which excavated a hole in the ground 570 feet
deep and 4000 feet across. While the main mass of the Goose Lake weighs
a little more than a ton, there is no evidence of any fragmentation of the
main body or crater formation at the place where it was found. It has
been suggested that it landed some distance away and bounced, coming to
rest where it was found. In this paper, the place where it was found will be
ealled the ‘‘impact site” as the simplest description of its terminal location.

I'irst Goosk LAKE EXPEDITION, 1960

In the light of the work done on recovering meteoritic particles during
the last two decades, it seemed odd that no particles had been reported from
the site of the Goose Lake fall. Inquiry showed, however, that no magnetic
survey had been made, and as far as known, no one visited the site between
1939 and 1960, except stockmen and hunters who would not have been
interested by a pole marker in the midst of a barren rock strewn area.

In 1960, the California Academy of Sciences approved a preliminary

296 CALIFORNIA ACADEMY OF SCIENCES [Proc, 471 SER

survey of the site, and in June of that year, the author and his son carried
out the first magnetic survey of the | toose Lake impact site.

Aside from the coordinates of the site given by Leonard (1956), the
only clue to its loc ation on current Forest Service maps is an excavated
reservoir designated by a sign reading “Meteorite Stock Tank,” which is

Ficure 2. Photograph of the Goose Lake meteorite in situ at the time of dis-
covery, October 13, 1938.

VoL. XXXII] BUTLER: GOOSH LAKE FRAGMENTS 297

almost half a mile south of the actual site. When the meteorite was removed
from the small crater in which it rested, the recovery party marked the
spot with a pole picturesquely described by Leonard, ‘. .. as a rude monu-
ment or marker, hewn from the trunk of a nearby sapling.”

With the able assistance of the U. S. Forest Service in Alturas, we
found this same pole intact and still erect, together with other small poles
lying about on the ground which were apparently used in hoistine the
heavy mass onto the wagon. From pictures made at the time of discovery
and before the iron had been moved, we were able to identify the same
trees on the horizon, even though they were somewhat taller than 21 years
ago. One very tall tree showing on the original print is now missine, but
we found it lying on the ground, its stump corresponding to its original
location.

Figure 2 shows a print of the Goose Lake meteorite before it was moved
at the time of discovery, October 13, 1938, and figure 3 is a picture taken
from about the same position in June, 1960, showing the pole marking the
impact site, and the same trees in the background.

Within a matter of minutes from the time we started searchine the
area with a hand-held magnet, it was clear that meteoritie particles lay

Ficure 3. Photograph of the impact site of the Goose Lake meteorite, made in
June, 1960.

298 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

about in great profusion. These were bottled, labeled, and returned to the
Academy for further study.

SEcoND GoosE LAKE EXPEDITION, 1961

The success of the first survey in recovering meteoritic fragments
prompted plans for a second survey of the field with more people. The
second Academy-sponsored expedition in September, 1961, included repre-
sentatives of the U. S. Geological Survey, the California State Division of
Mines, and the California Academy of Sciences. A much more thorough
search of the area was conducted, including partial excavation at the im-
pact point to determine the extent of bed-rock deformation. Many more
specimens were recovered, including large oxide fragments lying quite ex-
posed at some distance from the site.

MaGNeEtic RECOVERY METHODS

During the course of the first survey, no excavations were made, partly
beeause we wanted the evidence perfectly clear that particles of nickel-iron
and oxide were lying within a few millimeters of the surface of the ground.
A handful of soil was scraped from the ground and poured over the end of
a large conical magnet and during this operation magnetic particles in the
soil were drawn to and held by the magnet. Some of the material is mag-
netite, but this was readily separated from the nickel-iron by shaking the
magnet, the weaker magnetite falling off while the nickel-iron fragments
adhered as tightly as an iron nail. (The small magnetite particles must be
quite impure, because of their weak magnetism.) This method has the dis-
advantage that meteoritic oxide particles may not be recovered, because
they are less magnetic and may be lost.

Similar methods were used during the second expedition, but this time
the small magnet was replaced by a much larger double-pole magnet which
was provided with a long handle so it could be carried and lowered over a
given place. On this occasion, also, magnetic samplings of the soil were
bagged, marked, and returned to the Academy for analysis.

When the distribution of the large black massive-oxide fragments was
found to extend well beyond the limits of a few feet from the impact
point, areas of about 10 feet square were paced off at likely distances and
azimuths. Then a visual search was made by going over this area on hands
and knees, looking for the distinctive sheen of the oxide fragments. After
this visual search, a second scanning was made with a hand-held magnet.
The material from each plot was then bagged and labeled.

A more thorough search was conducted later on these samplings by
mounting the same magnet with the pole pieces facing down on a drill press

VoL. XXXIT] BUTLER: GOOSE LAKE FRAGMENTS 299

stand, spreading a small amount of the field material on a shallow aluminum
pan, and moving the pan around underneath the magnet. It was possible
to readily adjust the height of the magnet pole so that the magnetite
particles would not be drawn up as were the meteoritic particles. A thin
aluminum plate held to the magnet by a spring, was placed so as to extend
over both pole pieces, and each time a nickel-iron particle was separated, a
distinet ping could be heard as it struck the plate. When all the particles
had been removed, the magnet was. inverted bringing the aluminum plate
on the top with all the particles clustered around the two poles. Then, de-
taching the spring, the plate could then be lifted vertically and the particles
scooped into bottles. This method keeps the pole pieces clean, and allows
all the particles from one sample to be collected with no contamination
from previous sampling.

A photograph of the separator is shown in figure 4. This is a somewhat
simplified version of the automatic magnetic sampler used by Rhinehart
(1958) in his magnetic survey for meteoritic particles from the Canyon
Diablo area.

Ficure 4. Photograph of the magnetic separator used to sort out the nickel-iron
fragments from the soil.

Figure 5. Typical massive oxide fragments found lying on the surface of the
ground, and concentrated in a small area approximately 250 feet south of the
impact site. The largest piece measures about 3 centimeters in length.

300 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

After separating the particles, they were cleaned with a fine brass wire
brush, the heavy erusts and adhering soils removed with a dental scaler.
This latter operation must be done under a low power microscope.

After cleaning in this manner, some of the particles were mounted in
1-inch diameter lucite blocks, ground down with course emery paper until
an appropriate area for study had appeared and finished with 600-grit
paper. They were then carefully washed with water to remove any remain-
ing grits, and polished on a wheel with a felt lap saturated with AB alumina
polishing compound. The polishing was carried on until a good specular
finish was obtained with few seratch marks or comet tails. Since the lucite
is so much softer than the iron, it polishes away faster making the surfaces
slightly convex, especially at the edges where the nap of the polishing cloth
euts away the interface between the iron and lucite. For this reason, the
Neumann lines which extend to the edges are not quite in the same vertical
plane as those in the center. All etching was done with Nital. Alternate
polishing and etching was followed until the lines were clear.

MorPHOLOGY OF THE GOOSE LAKE FRAGMENTS

1. THE NICKEL-IRON PARTICLES. The shape and appearance of eight frag-
ments of the nickel-iron found during the first expedition in 1960 are shown
in figures 6 and 7. Their dimensions are indicated by seales and their in-

Figure 6. Typical nickel-iron fragments before cleaning. These were found
within a radius of 10 feet from the impact site. The scale shown along the lower
margin indicates millimeters. The weights of the fragments given in grams are
0.148, 0.162, 0.183, and 0.122 respectively.

Figure 7. Typical nickel-iron fragments which have been cleaned with a soft
wire brush. The scale shown along the lower margin indicates millimeters. The
weights of the fragments given in grams are 0.383, 0.250, 0.525, and 0.217 respec-
tively.

VoL. XXXII] BUTLER: GOOSE LAKE FRAGMENTS 301

dividual weights are given in each caption. Those in figure 6 are just as
they were removed from the magnetic separator, those in figure 7 have been
cleaned with a soft wire brush.

The color of the nickel-iron particles before cleaning is a light brown
or mahogany shade, identical with the soil. It is very difficult to distinguish
these by visual inspection from similarly shaped particles of rock. They
do not exhibit the characteristic brick red of freshly oxidized iron. When
cleaned, however, they look very much like many of the large iron meteor-
ites which exhibit the typical dull dark-grey sheen.

One feature common to almost all the nickel-iron particles is that they
are flattened. A rough estimate gives the average thickness to length ratio
of about 1/10. These dimensions were measured by placing each particle
between the jaws of a micrometer, so that the thickness figure includes any
protuberance or nodule. Some of the fragments excavated at the crater
during the second expedition measured 0.1 centimeter thick and 1.0 centi-
meter long. The nickel-iron fragments from below the surface show a
greater size range than those found on the surface. There is a marked
similarity between the shape of these flattened particles and the Algoma,
which Farrington (1915) ealls a peltoid or shield shape, even though the
dimensions differ by a factor of at least 50.

These pictures of the Goose Lake fragments should be compared to
Ninninger’s sluglets from Canyon Diablo. The hook shape is common to
both and the tiny hole in the lower right hand specimen in figure 7 is very
much lke the holes in some of the sluglets. The protuberances have the
same general appearance. The dimensions of the sluglets are about the
same as those just described from Goose Lake.

One of the smallest individual nickel-iron specimens of the Sikhote-Alin
shown by Krinov has the same weight as that of the lower right specimen
in figure 6, and shows much the same surtace topography. The Goose Lake
fragments, even after cleaning, do not appear to be as shiny and smooth
as those of the Sikhote-Alin.

2. THE MASSIVE OXIDE FRAGMENTS. The shape and appearance of the
Goose Lake meteoritic oxide fragments is entirely different from the nickel-
iron fragments. A few of these oxide fragments are shown in figure 5,
photographed soon after they were found. These are most easily recovered
by simply scanning the ground visually for their distinetly black sheen.
There are no other rocks in the vicinity which have quite the same color.
They are all magnetic, intermediate between magnetite and the nickel-iron,
and hence can be immediately checked in the field for meteoritie origin.
The fine powdery soil which was so adherent to the nickel-iron had ap-
parently been blown or washed from these fragments, leaving the surfaces

302 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser.

quite clean. No large pieces of the oxide were recovered from the pit exea-
vated at the impact site. However, there were many tiny oxide fragments
mixed with the nickel-iron fragments, not only on the surface but below as
well. Those found mixed with the soil did not have the characteristic black
sheen of the surface specimens, since they were covered with the light-
brown soil. However, they can often be recognized from their shape.

All the oxide fragments recovered show sharp fracture planes, including
the very smallest. The larger specimens taper toward the edges, giving an
elliptical cross-section. The sloping edges are not smooth, however, but
reveal steps formed by the lamellae planes.

The shapes of the ends of these oxide fragments suggest a block puzzle.
Of all these pieces of the oxide, only two were found which fitted togethe
just as though they had recently been broken. As shown in figure 8, it is
obvious that they were originally one piece. These large pieces are quite
homogeneous, are very hard and cannot be broken by hand. By contrast,
the thin oxide flakes ean be pinched in two with the fingers.

Figure 8. Two pieces of massive oxide which fit together perfectly. It is not
known how close together these lay. The larger weighs 8.19 grams, the smaller
3.70 grams. The scale indicates a length of 1 centimeter.

FicureE 9. Circular raised rim of hydrous iron oxide on one of the massive oxide
fragments. Weight 2.08 grams. The number (9) shown directly above the scale is
the figure number and is not related to the scale. The scale indicates a length
of 1 centimeter.

In the interstices of the lamellae, tiny deposits of salts and soil can be
seen under the microscope. The salts appear white when dry, green when
wet, can be removed easily with a pointed sealer.

One specimen of the oxide shows a small circular raised rim of brown
hydrous iron oxide, shown in figure 9. It has been suggested that this

VoL. XXXII] BUTLER: GOOSE LAKE FRAGMENTS 303

might also be the remains of an impact crater when the original metal was
in a near molten state. This example should be compared with a similar
structure found on one of the Canyon Diablo oxide fragments as given by
Buddhue (1957).

The oxide fragments collected from the surface of the ground only,
appear to increase in size from the crater or impact point out to about 250

TABLE 1

Weights of the massive oxide fragments found on the surface of the ground by
the second expedition to the Goose Lake site.

Azimuth from Crater Distance from Crater Weight

Specimen No. (degrees ) (feet ) (grams)

=
owmt KN OOF WD

weomnADO fe WwW hd

weomnondanrhrnwnnere &

=
o

170

125

5.44
2.75
3.61
1.00
1.00

2.61
LMG
hl
0.75
0.62
0.48
0.36
0.25
0.24
0.19

10.85
2.18
2.08
1.28
0.91
0.26
0.21
0.18
0.15
0.11

13.31
12.34
11.45
8.20
6.91
4.32
4.16
3.70
2.69
3.24

304 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER.

feet to the south and then terminate. There are also many small ones
mixed with these large pieces. The east and west distribution is roughly
20 feet on either side. It should be emphasized that the terrain slopes
gently to the south, so that the natural drainage is in the same direction
as the principal distribution of the surface oxide fragments. The prevailing
wind however, is at right angles to this, 7.e., from the west.

The individual weights of the large oxide fragments collected from the
four principal concentration areas to the south of the crater are given in
table 1. These were found only on the surface, during the second expedi-
tion and may or may not represent the true aerial distribution. This lst
includes only the ten largest specimens from each area, but does not in-
clude any of the nickel-iron particles.

An effort was made to estimate the total number of nickel-iron indi-
viduals and their weight distribution, but this was not satisfactory. It is a
simple matter to weigh and count the larger specimens, but as they get
smaller and smaller, their individual weights are more and more affected
by the thin oxide layer covering all the nickel-iron fragments. Besides this,
the amount of adhering soil and salts on each particle contributes more
and more to their weights.

A few of the larger nickel-iron fragments weighed about 0.200 grams,
but most weighed much less than this. When the weights approached 20
milligrams, the uncertainties mentioned above introduced very large errors.
We have not estimated how many of the smallest sizes were recovered, but
the number is certainly in the thousands.

During the course of both expeditions, it was anticipated that ablation
products in the form of spherical particles would be found. There are some
tiny, nearly spherieal particles, but under the microscope they are seen to
have a crystalline form suggesting magnetite. The fact that we found no
true ablation products adds further evidence to confirm Henderson’s theory
that this iron fell through the atmosphere at a low velocity.

METALLURGY OF THE FRAGMENTS

A number of the nickel-iron particles were examined under the micro-
scope to determine their internal structure and composition. These are
shown in figures 10 through 15, with some description of each. In all eases
examined, Neumann lines appeared. Since these lines appear only in
kamacite, it can be stated that all the nickel-iron particles reeovered and
examined are kamacite. A few thin threads of taenite were found traversing
the fragment. The Goose Lake iron is a coarse octahedrite, and if all the
nickel-iron fragments are kamacite, then there is no question that they are

VoL. XXXIT] BUTLER: GOOSE LAKE FRAGMENTS 305

: ‘+ ee a

Figure 10. Undistorted Neumann lines near the center of a nickel-iron frag-
ment. The number (10) shown directly above the scale is the figure number and
is not related to the scale. The scale indicates a length of 10 microns.

not fragments of the parent body, 7.e., the particles were not removed
either during flight, or mechanically by shock when it landed. However,
no nickel-iron particles have been found and examined which are greater
than the widest kamacite bands in the main mass, so if some mechanism
could selectively remove fragments like those we discovered, this would
support the possibility that the main mass is the parent body.

It is generally agreed that Neumann lines disappear at temperatures
above approximately 400°C, and since every nickel-iron fragment shows
these lines clearly, it seems evident that they were never heated above this
temperature and hence cannot very well be ablation products. If they were
not formed this way, then they must have been broken from a larger body
mechanically. Since there is no evidence that the main mass suffered a
severe shock on impact, we must conclude that the particles existed before
entry and were carried down through the atmosphere in the cavities or in
the wake of main mass.

In all specimens the nickel-iron particles show evidence of mechanical
strain or mechanical shock in the displacement and bending of the Neumann
lines. Many show the lines extending undistorted to the edge of one side,

306 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER.

while at the other side of the specimen the lines are bent and twisted and
in some cases completely granulated. The appearance of cross sections of
these particles suggests that they were projectiles at one time and that they
struck something hard. Their impact velocity was not sufficient to raise
their temperatures above 400°C, but was high enough to cause deformation
at the leading edge.

One ease in particuiar shown in figure 16 is that of an elongated frag-
ment which apparently suffered a head-on collision, bending completely
back on itself, in exactly the same way as a nail driven through a thin
board backed with an iron plate. One Neumann line could be followed all
the way around the end until it was nearly parallel to itself going in the
opposite direction. As in some of the other examples, the small end of this
specimen was apparently the leading edge, since all lines were obliterated at
the tip. At the other end, the lines appear in their normal undisturbed
form. This little spike was apparently traveling like an arrow when it
collided with a solid surface just hard enough to double it back.

The hypothesis that an extraterrestial shock effect can be observed in
meteoritic iron is not new. From analyses of the deformation structure in

Ficure 11. The same specimen as shown in figure 10, taken near one edge,
showing deformation of the lines. The scale shown in the upper part of the figure
indicates a length of 1 millimeter.

VoL. XXXIT] BUTLER: GOOSE LAKE FRAGMENTS 307

kamacite, Maringer and Manning (1962) tentatively conclude that a rela-
tively slow impact velocity between a large and a small body in space could
produce this kind of deformation. The gross forms as well as the internal
evidence of the Goose Lake fragments apparently confirm this suggestion.

One slice of the Goose Lake meteorite was available for making com-
parisons between its internal structure and that of the fragments. There
are some likenesses, but the long clean curving Neumann lines at the edge
of this slice have not been found among the nickel-iron fragments. In this
slice, there are many well formed rhabdite inclusions mixed with the
kamacite, but none have been found in the Goose Lake particles. Likewise,
no troilite inclusions have been found, although they are found in the main
mass. This may not be unusual, however, because the troilite inclusions are
roughly the same size as the fragments, and since this mineral is non-mag-
netic, it may not have been recovered from the soil by our techniques.

A few of the larger massive oxide fragments were ground and polished,
one of which is shown in figure 17. Etching was unnecessary to reveal
either the laminar structure, or the rhabdite inclusions. This characteristic
laminar form is almost identical to that of the massive oxide from Canyon
Diablo as can be seen in a similar specimen shown by Buddhue. Likewise,
the rhabdite erystals in the oxides appear identical to those found in the
Canyon Diablo as shown in figure 18.

THE ORIGIN OF THE GOOSE LAKE FRAGMENTS

The principal conclusion of these preliminary studies of the Goose Lake
fragments centers around the question of whether the particles we dis-
covered were once a part of the main body.

Since well developed Neumann lines have been found in all nickel-iron
particles so far examined, it is clear that they are not ablation products.
If they had been torn away mechanically from the surface of the main
body, either by violent vibrations during flight or by the shock of impact
with the ground, evidence of surface spallation should show on the surface
of the iron. None has been found.

Since taenite melts at a lower temperature than kamacite, Henderson
(1956a) has suggested that at the leading edge of the main mass where the
layers show some thermal deformation, ablation heating could loosen or at
least weaken the bonds between the taenite and kamacite plates. Then the
shearing action of the atmosphere during deceleration might tear off some
of the kamacite plates. If this occurred, it seems reasonable that some of
these particles of kamacite would be caught in the turbulent wake, or in the
cavities and be carried to the ground. There remains the question of how
this process could produce so many particles, and whether the heating
time required to weaken the bonds would also obliterate the Neumann lines

308 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 47TH Ser.

Ficure 12. Leading edge of a nickel-iron fragment showing three undistorted
Neumann lines completely granulated toward the edge. The number (12) shown
directly above the seale is the figure number and is not related to the scale. The
scale indicates a length of 100 microns.

Figure 13. The same specimen as shown in figure 12, taken from the opposite
side of the fragment. Note that the lines extend completely undisturbed to the
edge. These same lines connect with the three of figure 12. The number (13)
shown directly above the scale is the figure number and is not related to the scale.
The scale indicates a length of 100 microns.

in the kamacite plates. The outer surface of the main mass does not show
that such a phenomenon took place, but it is conceivable that subsequent
heating and pressure smoothed the leading edge where the fractures took
place. While this concept has some attractive features, it does not appear
to explain the occurrence of the fragments as simply as the following ac-
count.

Another alternative which seems more attractive, is that the fragments
pre-existed and accompanied the main mass when it was captured by the
eravitational field of the earth. If this third concept is tenable, then some
hypothesis is required to account for the high concentration of particles so
close to the point where the meteorite was found.

While we have no direct information on this point, it 1s tempting to
suggest that they were transported through the atmosphere in the cavities.
After reaching the ground, weathering on the top and gravity on the bot-
tom removed the nickel-iron particles from the holes, scattering them in the
immediate vicinity of the impact point. The highest concentration was ac-
tually in the bottom of the pit, directly under the place where it fell. So
far as we know, no one at the time of discovery thought to make a thorough
examination of those cavities facing upward to see if any of the fragments
were lying in the bottom.

VoL. XXXIT] BUTLER: GOOSE LAKE FRAGMENTS 309

It is quite certain that we have not recovered all the material at the
site, but the total volume of all we have found, including the large oxide
fragments could easily fit into the cavities, with room to spare. Cavity
transportation of the fragments would account for their coneentration
around the impact point.

If we assume that the main mass of the Goose Lake meteorite had passed
through swarms of tiny particles such as we have discovered, it seems
reasonable to suppose that during the thousands of years it was orbiting
through the solar system, there would have been hundreds of collisions, say

Figure 14. Lateral displacement in Neumann lines. The number (14) shown
directly above the scale is the figure number and is not related to the scale. The
scale indicates a length of 1 millimeter.

one a year. Further, if the orbits of the particles and of the main mass are
nearly identical so that the collisions made soft impacts, then there would
be no spallation of the surface of the main mass nor fusion of the particles
owing to heating by their sudden loss of kinetic energy. We will assume
that the impact velocity is just sufficient to partly deform the particles
which we have described. This would explain the deformation of the Neu-
mann lines on the leading edge of the particle, leaving those on the trailing
edge intact.

310 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SEr.

Figure 15. Large bend in a nickel-iron fragment, showing an inclusion of
taenite bent in a jagged Z-shape. The number (15) shown directly above the scale
is the figure number and is not related to the scale. The scale indicates a length of

1 millimeter.

Now, if the magnetic and gravitational forces between the main mass
and these particles is sufficient, those which have collided in this manner
will adhere to the main mass and be earried along with it, until it begins
to encounter atmospheric drag as it approaches the earth. At this time,
those particles which are on the outside will be swept away by the violent
turbulence of the air stream and will fall to the earth over very widely
scattered areas. Those others, however, which enter the apertures and
collide with the bottoms of the cavities will be protected from the streaming
atmosphere and will remain in their protected holes until arriving at the
surface of the earth.

The oxide fragments present another puzzling problem. Had these not
been found, we would have concluded that the large numbers of small
nickel-iron particles on the surface show further evidence that the Goose
Lake fall is of recent origin. Furthermore, if our preliminary conclusions
are correct, that these nickel-iron fragments are not a part of the main mass,

VoL. XXXIT] BUTLER: GOOSE LAKE FRAGMENTS 311

we have suggested a method by which they could have been transported
from the orbit of the main mass to the ground.

The thick oxide fragments, however, indicate that they have been sub-
jected to oxidation processes for a very long time. Their thicknesses are not
ereatly different from the massive oxidation fragments from the Canyon
Diablo. It is not clear how pieces of this meteoritic oxide ranging in size
from a few milligrams to 13 grams could still be clustered on the surface
of the ground so close to the impact point for even a fraction of the time
required to form this thick stable oxide. This area is wind whipped by
violent storms characteristic of the high Sierras, especially in winter. These
winds would certainly seatter particles like these over wide areas in a frac-
tion of the time presumably required to form these thick laminar oxide
layers.

It is possible, of course, that we are dealing with more than one fall.
In this case, there should be evidence of a fall nearby and many more of
these fragments throughout Modoe County.

However, if no more oxide fragments are discovered, and if the Goose
Lake meteorite did fall within the last fifty years or so, then we must con-
elude that the oxide fragments were transported in a manner similar to
that of the nickel-iron fragments, 7.e., in the wake of the main body or in its
cavities. The origin of pre-entry massive oxide meteoritic fragments re-
quires further study.

Figure 16. Hook-shaped nickel-iron fragment. Individual lines can be followed
all the way around the bend. The number (16) shown directly above the scale is
the figure number and is not related to the scale. The scale indicates a length of
1 millimeter.

FicurE 17. Flat side and edge of massive oxide fragment revealing laminar
structure. The number (17) shown directly above the scale is the figure number
and is not related to the scale. The scale indicates a length of 1 centimeter.

FicureE 18. Rhabdite crystal in one of the massive oxide fragments. No etching.
The number (18) shown directly above the scale is the figure number and is not
related to the scale. The scale indicates a length of 100 microns.

312 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

SUMMARY OF THE EVIDENCE

Two types of meteoritic particles have been discovered at the site where
the Goose Lake meteorite was found; nickel-iron fragments of kamacite
whose weights are in tenths of grams and massive laminar oxide fragments
weighing up to 13 grams. The former are concentrated close to the as-
sumed impact point, the latter distributed out to distances of approximately
300 feet.

The appearance of Neumann lines in the nickel-iron fragments indi-
cates that they were never heated above about 400°C. The bending and
twisting of the lines shows that they were deformed while cold.

The massive laminar oxide fragments, showing rhabdite inclusions, are
identical to those found at Canyon Diablo.

CONCLUSION

Evidence that the Neumann lines in the nickel-iron fragments are dis-
similar to those in the Goose Lake meteorite indicates that their origin is
independent of this mass of iron, that they were swept up by the meteorite
during its long life in orbit around the solar system. Only those fragments
which found their way into the cavities were recovered, any others were
blown off by the air stream during its final flight. Cavity transportation
explains their abundance at the impact site, as well as protection against
aerodynamic heating. Deformation of the nickel-iron fragments occurred
when they collided with the main mass in space.

The massive laminar oxide fragments suggest a second meteorite fall
in the same vicinity as old as Canyon Diablo.

ACKNOWLEDGMENTS

The author wishes to express his appreciation to the California Academy
of Sciences for its support in this work, especially in view of the fact that
all published data on the Goose Lake meteorite up to the time of the first
expedition were silent on the subject of fragments. A number of people
were most helpful in the early stages of this work before the problem had
been fairly defined. Dr. Max Hey of the British Museum and Dr. E. M.
Shoemaker of the Geological Survey made valuable suggestions as well as
providing me with specimens of other meteorites for comparison studies.
Special mention should be made of the voluminous correspondence and
personal discussions with E. P. Henderson of the Smithsonian Institution.
He very kindly made available a slice of the main mass, together with per-
mission to polish and etch it in the same way as the fragments.

For some 12 years, the author has been fortunate in having the advice
and counsel of Dr. G Dallas Hanna in problems relating to optical materials,

VoL. XXXII) BUTLER: GOOSE LAKE FRAGMENTS 315

eclipse instrumentation, and in meteorities. His enthusiastic and youthful
outlook has been an inspiration, and it was largely through his encourage-
ment that the first formal proposal was submitted to go in search of the
Goose Lake fragments.

LITERATURE CITED

BuppuHukE, J. D.
1957. The oxidation and weathering of meteorites, no. 3. 161 pp. University of
New Mexico Press, Albuquerque, New Mexico.
FARRINGTON, O. C.
1915. Meteorites. x + 233 pp. The Lakeside Press, Chicago, Illinois.
HENDERSON, E. P., and S. H. Perry
1958. (a) Studies of seven siderites. Proceedings of the United States National
Museum, vol. 107, no. 3388, pp. 339-403, pls. 1-22.
1962. (b) Personal communication.
Krinov, E. L.
1960. Principles of meteorites. xi + 535 pp. Pergamon Press Inc., London.
LINSLEY, E. G.
1939. The giant Goose Lake meteorite from Modoc County, California. Cali-
fornia Journal of Mines and Geology, vol. 35, no. 3, pp. 308-312.
LEONARD, F. C.
1940. (a) The Goose Lake siderite; the largest known meteorite of California.
Griffith Observer, no. 1, pp. 2-8.
1956. (b) A classificational catalog of the meteoritic falls of the world. Uni-
versity of California Publications in Astronomy. Vol. 2, no. 1, 80 pp.
University of California Press, Berkeley and Los Angeles, California.
MARINGER, R. E., and G. K. MANNING
1962. Researches on meteorites. Edited by Carleton B. Moore, 227 pp. John
Wiley and Sons, Inc., New York.
NINNINGER, H. H.
1956. Arizona’s meteorite crater. xv + 232 pp. World Press, Denver, Colorado.

RINEHART, J. S.

1958. Distribution of meteoritic debris about the Arizona meteorite crater.
Smithsonian Contributions of Astrophysics, vol. 2, no. 7, pp. 145-160.

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PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

FOURTH SERIES

G Dallas Hanna Anniversary Volume

Vol. XXXII, No. 10, pp. 315-338, 4 figs., 2 tables May 20, 1963

RELATIONSHIPS OF THE PERCOID FISH
PENTACEROS RICHARDSONI SMITH, WITH
DESCRIPTION OF A SPECIMEN FROM THE

COAST OF CALIFORNIA
by
W. I. Follett and Lillian J. Dempster

California Academy of Sciences

A specimen of the perecoid fish Pentaceros richardsoni Smith’ was col-
lected Mareh 10, 1960, off Pigeon Point, San Mateo County, California.
This capture extends the recorded range of this species more than 400 miles
southward on the Pacific coast of North America and adds to the known
fauna of California the western Pacific family Pentacerotidae.

The specimen (fig. 1), California Academy of Sciences no. 26759,
measures 252 mm. in standard length and 305 mm. in total length. It was
collected by the drag boat Henrietta Paladini in California Department of
Fish and Game Block 481 (387° 15’N., 122° 55’ W.) in one of three tows
made at depths of 265, 255, and 220 fathoms.

Other fishes taken in these tows were the pleuronectids Hopsetta jordani
(Lockineton), Parophrys vetulus Girard, Microstomus pacificus (Locking-
ton), and Glyptocephalus zachirus Lockington; the scorpaenids Sebastodes

1. See Generic Position, p. 324, and Nomenclature, p. 328.

[ 315 J

Marine Biological Laboratory
j EAA a Sia 54 Se f

MAY 2 9 1963

wnnne LnNiIc MACS

316 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Figure 1. Pelagic armorhead, Pentaceros richardsoni Smith, CAS 26759, stand-
ard length 252 mm., collected March 10, 1960, off Pigeon Point, San Mateo County,
California. Photograph by W. I. Follett.

paucispinis (Ayres), S. goodei Eigenmann and Kigenmann, and Sebastolo-
I ve ( As re ) S g l I oO oD )
bus alascanus Bean; and the anoplopomatid Anoplopoma fimbria (Pallas).

DISTRIBUTION

Pentaceros richardson has been recorded but onee from South Afriea,
its type-locality (Smith, 1849), and twice from New Zealand (Hutton,
1890; MeCulloch and Phillipps, 1923). Certain evidence suggests that its
center of abundance is in the vicinity of southern Japan: a number of
specimens have been taken south of Tokyo (Abe, 1957; Tomiyama and Abe,
1958), and a fishery for this species is conducted about 50 nautical miles
northeast of Hachijo Island, off central Honshu (Tokiharu Abe, personal
communication, November 21, 1961). Probably, the specimens recorded
from the North Pacific Ocean (Welander, Johnson, and Hajny, 1957) and off
British Columbia (Neave, 1959; Clemens and Wilby, 1961) and Oregon
(Wagener and Bond, 1962), and our specimen from central California, were
transported far to the east of the center of abundance of this species by the
North Pacific current system (Kuroshio, Kuroshio Extension, North Pacifie
Current, and California Current; see Sverdrup, Johnson, and Fleming,
1946, pp. 719-724; chart 7).

VoL. XXXIT] FOLLETT AND DEMPSTER: PENTACEROS 317

SYNONYMY

Pentaceros richardsonu Situ, 1849, p. [38] (listed in index to plates).
GUNTHER, 1859, p. 212 (reference; diagnosis; distribution; holotype in
British Museum). BLEEKER, 1876, p. 270 (type-species, by original
designation, of Pseudopentaceros, new genus). Hurron, 1890, p. 277
(specimens, from New Zealand, in Canterbury Museum). Gini, 1893a,
p. 116 (synonym of Pseudopentaceros richardsonu; reference). GIL-
CHRIST, 1902, p. 108 (references). WaiTr, 1907, p. 19 (reference).
THOMPSON, 1916, p. 1384 (synonymy).

Pentaceros richardsoni Smiru, 1849, pp. [51]—[52] (original description;
type-locality, “near to Cape Point,” South Afriea, “in very deep water”) ;
pl. 21. BuLreKer, 1860, p. 52 (listed, Cape of Good Hope). Hutton,
1904, p. 41 (reference). JoRDAN, 1907, p. 236 (specific name only; type
of Gilchristia, new genus; record). McCunuocn, 1915, p. 144 (type of
Pseudopentaceros Gill [sic], which = Gilchristia Jordan and, possibly,
Quinquarius Jordan; references). JORDAN, 1919, p. 382 (orthotype of
Pseudopentaceros Bleeker, which “replaces” Gilchristia Jordan).

Pseudopentaceros richardsonu. Gi, 1893a, p. 116 (Pentaceros richardson
a synonym; listed, New Zealand). Warrr, 1907, p. 19 (listed, New
Zealand; reference). PHILLIPPS, 1927, p. 13 (listed, New Zealand).

Histiopterus richardson. JORDAN, 1920, p. 524 (orthotype [sic] of Guil-
christia Jordan*, a synonym of Pseudopentaceros Bleeker).

Pseudopentaceros richardsom. Hutton, 1904, p. 41 (listed, New Zealand).
McCuu.tocu and Pumurprs, 1923, p. 18 (references; description of 70-
mm. specimen from Nelson, New Zealand); pl. 4, fig. 1. BarNnarp, 1927,
pp. 621-622 (references; description; distribution); 1937, pp. 56, 57
(comparisons; counts; reference). SmitH, 1951, pp. 873-875 (distribu-
tion; references; comparisons); 1961, p. 242 (references; diagnosis;
distribution) ; pl. 35, fig. 622. WELANDER, JOHNSON, and Hagny, 1957,
p. 244 (“boar fish”’) ; p. 245 (reference; counts; measurements ; descrip-
tion; comparisons; taxonomic notes; specimens from North Pacific,
45° 49’ N., 160° 03’ W., to 51° 00’ N., 150° 00’ W.) Aser, 1957, pp. 35-39,
71-73 (“kusakari-tsubodai”; references; description; counts; measure-
ments; comparisons; specimens from southern part of Bodsd Peninsula
(probably), from Hachij6 Island, and off Amatsu, Japan); fig. 1. Tomr-
YAMA and ABE, 1958, p. 165 (“kusakari-tsubodai”’; diagnosis; ‘fairly
palatable”; distribution; specimens from Hachij6 Island and off Amatsu;
col. fig.). NEAVE, 1959, p. 384 (“boar fish”; reference; specimen from
50° N., 145° W., off British Columbia). CLEMENS and WILBy, 1961, p. 42

2. Jordan (1920, p. 524) wrote ‘‘orthotype Histiopterus richardsoni Gilchrist’’—-surely a lapsus calami
for Pentaceros richardsoni Smith (see Jordan, 1919, p. 382).

318 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER.

(characters in key); p. 219 (‘“‘boarfish’; description; records; distribu-
tion) ; fig. 126. WaAGNER and Bonp, 1962, pp. 71-72 (references; counts;
measurements; distinguishing characters; age; specimens from Oregon
coast, between Umpqua and Alsea rivers, in 80-90 to 110 fathoms) ; fig.

1; fig. 2 (head).

Griffinetta nelsonensis WHITLEY and PHILLIPPS, 1939, p. 233 (new genus
and species, based on young specimen (total length 70 mm.) from Nelson,
New Zealand, described as Pseudopentaceros richardson by McCulloch
and Phillipps, 1928, p. 18).

In view of the extensive changes that occur with age in this family
(McCulloch, 1915, pp. 145-146, pl. 26; Waite, 1923, p. 143; Smith, 1951,
pp. 874-875), we follow MeCulloch and Phillipps (1923, p. 18) in referring
their young specimen from Nelson, New Zealand, to Pentaceros richardson.
Whitley and Phillipps (1939, p. 233) referred this specimen to a distinet
genus and species, Griffinetta nelsonensis, but expressed no basis for such
distinction.

DESCRIPTION

We confine our discussion to those respects in which our observations
differ from, or supplement, the excellent description of this species by
Abe (1957).

Counts and measurements are expressed as by Hubbs and Lagler (1958).
Principal caudal rays consist of all branched rays plus the upper and the
lower adjacent unbranched ray; all other unbranched caudal rays are re-
garded as procurrent rays. Predorsal bones (called auxiliary interneurals
by Starks, 1904, p. 613) are the median bones (rayless pterygiophores) that
are anterior to the dorsal pterygiophores (see Smith and Bailey, 1961, p.
345). Counts of the caudal rays, predorsal bones, and vertebrae were de-
termined from a radiograph.

Counts. Dorsal rays XIV,9. Anal rays IV,8. Caudal rays: principal
17 (9 upper, 8 lower) ; procurrent 11 (6 upper, 5 lower). Pectoral rays 19
on each side (first two and last two unbranched). Pelvie rays 1,5 on each
side. Seales (left side): 68 lateral-line pores, 14 rows above lateral line,
42 rows below lateral line. Seales (right side): 69 lateral-line pores, 14
rows above lateral line, 41 rows below lateral line. Cheek seales: horizontal
rows 7 on each side; vertical rows 13 on each side. Predorsal bones 2.
Branchiostegal rays 4 + 3 on each side. Vertebrae 25 (precaudal 12; caudal
13, ineluding urostylar vertebra).

nll rakers (counts inelude all rudiments; count of lower limb includes
raker in angle of arch) :

VoL. XXXIT] FOLLETT AND DEMPSTER: PENTACEROS 319

Anterior Posterior
Left Right Left Right
Lot 620 7+17 7+17 3+ 15 4+ 15
2G) 6g 3+ 15 3+ 15 2+ 12 2+ 12
So 30 2+ 12 2+ 13 1+ 11 1+11
J) 20 ne 1+ 11 1+ 11 0+ 7 0+ 7

MEASUREMENTS. These are given in hundredths of standard length
(252 mm.). Body depth: fifth dorsal spine to pelvic spine 40; fourteenth
dorsal spine to first anal spine 30; posterior end of dorsal base to posterior
end of anal base 15. Least depth of caudal peduncle 9. Body thickness:
greatest 16; between outer margins of pelvie origins 10.

Snout to dorsal origin 39. Dorsal base 53; spinous-dorsal base 40; soft-
dorsal base 13. Posterior end of soft-dorsal base to base of middle caudal
ray 15. Snout to pectoral origin 33. Snout to pelvic origin 45. Pelvie
origin to anal origin 29. Snout to anal origin 72. Anal base 17; spinous-
anal base 7; soft-anal base 10. Length of caudal peduncle (posterior end
of anal base to base of middle caudal ray) 16.

Head: length 33; depth 30; width 16; postorbital leneth 13. Snout
length 12. Suborbital width 1. Cheek: height 9; length 15. Orbit to angle
of preopercle 12. Interorbital width (least bony) 11. Orbit: horizontal
diameter 9; vertical diameter 8. Upper-jaw length 10. Mandible length 13.

Gape width 7.

DoRSAL-FIN SUPPORTS. The two predorsal bones and the first proximal
pterygiophore are shaped somewhat like the number “7.” The first and
second proximal pterygiophores are ankylosed, but they are distinguishable
from each other by their lateral keels (as we have confirmed by dissection).
Thus distinguished, the first and second pterygiophores support each one
dorsal spine. The first predorsal bone extends downward in front of the first
neural spine; the second predorsal bone, between the first and second neural
spines. The first and second pterygiophores extend downward between the
second and third neural spines; the third and fourth pterygiophores, be-
tween the third and fourth neural spines. The fifth pterygiophore extends
downward between the fourth and fifth neural spines. (Radiographs reveal
the same arrangement in the three other specimens of Pentaceros richard-
soni that we have examined (see table 1) and in a specimen of Quinquarius
(= Pentaceros) japonicus (Stanford University no. 18191) 184 mm. in
standard length.)

DorsSAL SPINES. The dorsal spines are heteracanth. Because of their
bilateral asymmetry, they fit closely together when depressed into the

320 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4vH SER.

ao
Sie

oni, CAS

Figure 2. Locking mechanism of dorsal spines of Pentaceros richards
26759. Spines erect. Radiograph by W. I. Follett.

80:

Figure 3. Locking mechanism of dorsal spines of Pentaceros richardsoni, CAS
26759. Spines partly depressed, indicating the manner in which the serrate lobe

engages the face of the preceding spine. Radiograph by W. I. Follett.

VoL. XXXIT] FOLLETT AND DEMPSTER: PENTACEROS 321

dorsal groove. (Jordan, 1907, p. 235, perhaps inadvertently, noted as a
familial character “dorsal... spines... not depressible in a groove.) As
in one of the specimens recorded by Abe (1957, p. 72), the first, third,
fifth, and successive odd-numbered spines incline to the right and the
second, fourth, sixth, and successive even-numbered spines incline to the
left. (In two other specimens recorded by Abe (1957, pp. 36, 71) the odd-
numbered spines inclined to the left and the even-numbered spines inclined
to the right.) The spines are longitudinally ridged. The ridges are more
conspicuous on the left side of the odd-numbered spines and on the right
side of the even-numbered spines.

The third to sixth spines, immediately distal to the articulation, are
each expanded into a vertical, laterally compressed anterior lobe (see figs.
24). (A lobe is faintly indicated on the second and the seventh spines,
but is not discernible on any of the others.) The front edge of the lobe is
bluntly serrate. The serrae are marginal protrusions of smooth ridges that

extend obliquely around the front edge of the lobe (see fig. 4). The lobe of
the third and fifth spines is at the left of the interradial membrane; that
of the fourth and sixth spines is at the right.

Figure 4. Fourth dorsal spine of Pentaceros richardsoni, CAS 26759. a. Lateral
view, left side. b. Anterior view. c. Lateral view, right side. Photographs by
W. I. Follett.

These lobate spines, which resemble those in Pentaceros capensis Cuvier
m Cuvier and Valenciennes, 1829, and those in Pentaceros japonicus
Doderlein am Steindachner and Déderlein, 1884, function as a locking

322 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

mechanism. This mechanism is similar to that in P. capensis as described
by Smith (1951, p. 878).

ANAL SPINES. The anal spines are heteracanth. The first and third
spines incline to the left; the second and fourth, to the right. (In two speci-
mens recorded by Abe, 1957, p. 72, the second spine inclined to the right.)
The spines are longitudinally ridged. The ridges are more conspicuous on
the right side of the first and third spines and on the left side of the second
and fourth spines.

The third spine, immediately distal to its articulation, is expanded into
a vertical, laterally compressed anterior lobe, the edge of which bears five
blunt serrae. There is a shght anterior lobe on the second spine, but none
is discernible on the first or fourth. These lobes are elements of a locking
mechanism.

PELVIC SPINES. Longitudinal ridges similar to those on the dorsal and
anal spines extend along both sides of the pelvie spines (and along the
basal portion of the lower side of the pelvic soft-rays).

BRANCHIOSTEGAL MEMBRANES. The inner fold of the right branchiostegal
membrane overlaps that of the left. This asymmetry is contrary to the
rule that, in fishes with overlapping branchiostegal membranes, the left
membrane generally overlaps the right (see Hubbs and Hubbs, 1945, p. 279;
Crossman, 1960, p. 368).

Variability in the asymmetry of the branchiostegal membranes may be
common in this species. Three specimens with the right branchiostegal
membrane overlapping the left, and four with the left overlapping the
right, were recorded by Abe (1957, pp. 38, 71, 72, 73). In the 254-mm.
specimen preserved by Welander, Johnson, and Hajny (1957, p. 245), the
right branchiostegal membrane overlaps the left; in their 240-mm. speci-
men, the left overlaps the right. In a 257-mm. specimen collected by
Richard C. Johnson, August 15, 1958, in the North Pacific Ocean, 49° 43’ N.,
146° 10’ W., the left branchiostegal membrane overlaps the right.

ASSOCIATION OF BILATERAL ASYMMETRIES IN SEVEN SPECIMENS

The bilateral asymmetries of the dorsal spines, anal spines, and branchi-
ostegal membranes in Pentaceros richardsoni are variously associated with
one another. The combinations of these characters in the four specimens
examined by us and in three specimens recorded by Abe (1957) are shown
in table 1.

VoL. XXXIT] FOLLETT AND DEMPSTER: PENTACEROS 323

Table 1. Bilateral asymmetries in seven specimens of Pentaceros richardsoni.

Side of spine expanded into an
anterior lobe

eae a ee Overlapping

Specimen Odd-numbered EKven—numbered branchiostegal
spines spines membrane

Dorsal Anal Dorsal Anal
OAS 26759, .... Left Right Right Left Right
Abe ’57-125 .... Right Right Left Left Lett
Abe (Masuda) .. Right Right Left Left Left
bemtOaD2.). . 2s: Left Right Right Left Right
MWe AL468) 0... Right Right Left Right Left
WetT469? .. Left Left Right Left Right
MVotob15® ....4 Left Right Right Left Left

1. Welander, Johnson, and Hajny (1957, p. 245), 240-mm. specimen.

2. Welander, Johnson, and Hajny (1957, p. 245), 254-mm. specimen.

3. Specimen collected in a surface gill net by Richard C. Johnson, August 25, 1958, in the North
Pacific Ocean, 49° 43’ N., 146° 10’ W., water temperature 11.8° C.

DENTITION. The teeth on the premaxillaries and dentaries are in a band.
They are small, conical, and slightly curved—those of the outer row ir-
regularly enlarged. There are 24 small, scarcely curved teeth on the head
of the vomer. No teeth are present on the shaft of the vomer or on the
palatines or tongue. The teeth of the upper and lower pharyngeals are
similar in shape to the enlarged teeth of the premaxillaries and dentaries,
but are somewhat larger. (The lower pharyngeals are not coalesced. )

CAUDAL SKELETON. Characters revealed by a radiograph of our speci-
men indicate that the caudal skeleton of Pentaceros richardsoni resembles
the most primitive perciform type as defined by Gosline (1961). In the
following discussion, the terminology follows that of Gosline (1961), except
as noted.

Epurals. There are three epurals, similar in shape and size. Their
rounded anterior (lower) ends lie above uroneural 1, the anterior end of
epural 1 extending slightly ahead of uroneural 1, to a point above the
neural process of the penultimate (12th) vertebra.

Uroneurals. There appear to be two uroneurals. Uroneural 1 is similar
in shape to that figured by Gosline (1961, fig. 1). The presence of uroneural
2 is suggested by a projection above the dorsal margin of hypural 6 (the
uppermost hypural) and by a faint line extending obliquely downward
and forward from that projection.

Urostyle. There is a single urostylar ossification.

324 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Hypurals. The sutures separating all six hypurals are distinet. The
hypural spine (defined by Merriman, 1940, p. 63) is well developed.

Hemal arches. The hemal arch of the antepenultimate vertebra, as well
as that of the penultimate vertebra, is autogenous (separate from the
centrum ).

Caudal rays. There are 17 principal caudal rays (9 upper, 8 lower)
and 11 procurrent rays (6 upper, 5 lower), their articulation with the
epurals, hypurals, and hemal spines, respectively, appearing to be as follows:

Epurals 1 and 2 support each one procurrent ray; epural 3 supports
four procurrent rays.

Hypural 6 supports one unbranched ray (the uppermost principal ray) ;
hypural 5, six branched rays; hypurals 5 and 4 together, one branched ray;
hypural 4, one branched ray (the lowermost ray of the upper lobe) ; hypural
3, one branched ray (the uppermost ray of the lower lobe); hypural 2, four
branched rays; and hypural 1, two branched rays (ineluding the lowermost
branched ray of the lower lobe).

The hemal spine of the penultimate vertebra supports the lowermost
(unbranched) principal ray and three procurrent rays. The hemal spine
of the antepenultimate vertebra supports two procurrent rays.

GENERIC POSITION

We find no characters that justify the distinction of Pseuwdopentaceros
Bleeker, 1876, from Pentaceros Cuvier in Cuvier and Valenciennes, 1829.

When Bleeker (1876, p. 270) erected the genus Pseudopentaceros (type-
species, Pentaceros richardsoni), he diagnosed it as having small teeth on
maxillaries and vomer, 14 dorsal spines, 4 anal spines, and no erests or
prominent osseous tubercles on the upper surface of the head. He diagnosed
the genus Pentaceros Cuvier and Valenciennes (type-species, Pentaceros
capensis) as having small teeth on maxillaries and vomer, 12 dorsal spines,
5 anal spines, and crests or prominent osseous tubercles on the upper sur-
face of the head (Bleeker, 1876, p. 269).

The supposed distinction based on 14 dorsal spines and 4 anal spines in
Pseudopentaceros in contrast with 12 dorsal spines and 5 anal spines in
Pentaceros is dispelled by the following material :

1) A specimen of Pentaceros capensis with 4 anal spines (Fowler, 1935,
pp. 595-394 and fig. 26, as Quinquarius capensis). (Fowler’s seale counts
negate the possibility considered by Barnard, 1937, p. 57, that this speci-
men might represent Pentaceros richardsoni. )

2) A specimen of Pentaceros capensis with 13 dorsal spines (Barnard,
1937, p. 56, as Quinquarius capensis).

3) A specimen of Pentaceros richardsoni with 13 dorsal spines (Clemens
and Wilby, 1961, p. 219, and fig. 126, as Pseudopentaceros richardsoni).

VoL. XXXIT] FOLLETT AND DEMPSTER: PENTACEROS 325

The presence or absence of cephalic crests or osseous tubercles appears
of doubtful significance. This distinction was proposed by Bleeker on the
basis of a juvenile of one form and an adult of the other, and apparently
without knowledge that these structures are known only in juveniles. (See
McCulloch, 1915, pp. 145-146, pl. 26; Waite, 1923, p. 143; Smith, 1951, pp.
874-875. )

In the following summary of noteworthy similarities of the two forms,
the characters of Pentaceros richardsoni are based on our specimen, 305 mm.
in total leneth, and those of P. capensis are based on the description by
Smith (1951, as Quinquarius capensis) of a specimen 320 mm. in total
length.

Bones of the head exposed, bearing rugulose ridges. Lips and chin
villous. Dorsal spines heteracanth, longitudinally ridged on alternate sides;
on the ridged side, a serrate anterior lobe near the base of the third, fourth,
and fifth spines, functioning as an element of a locking mechanism. Anal
spines heteracanth, longitudinally ridged on alternate sides; on the ridged
side, a serrate anterior lobe near the base of the third spine, functioning
as an element of a locking mechanism. Teeth on head of vomer*, none on
palatines or tongue. Lower pharyngeals not coalesced. Preorbital, cireum-
orbitals, preoperecle, and ventral surface of mandible, with large sensory
pores roofed by membrane. Six large pores across the chin. Lateral line
arching toward base of fourth dorsal spine, thence roughly parallel to
dorsal base, curving down to caudal pedunele, and thence extending hori-
zontally to caudal base. Seales of throat, breast, cheeks, belly, and nape
scutelike, slightly or not at all imbricate. Vertebrae 12 + 13—25. (The
vertebral count of 12 + 12 noted by Welander, Johnson, and Hajny (1957,
p. 245) in two specimens of Pentaceros richardsoni must have excluded
the urostylar vertebra. We have X-rayed these two specimens and we find
that each has a vertebral count of 12 -+ 13, including the urostylar vertebra.
Smith (1951, p. 877) noted the number of vertebrae in his specimen of
Pentaceros capensis as 13 + 12. His demarcation between precaudal and
caudal vertebrae may be different from ours, since his radiograph (cbid.,
pl. 18) shows 138 vertebrae that we regard as caudal, and seems to show 12
vertebrae that we regard as precaudal. We therefore believe that both
forms have the vertebrae, according to our method of counting, 12 + 13 =
25.)

Smith (1951, p. 874) stated that if only the adult stadia of Psewdopen-
taceros and Quinquarius (== Pentaceros) were compared, ‘one might al-

3. In the original diagnosis of the genus Quinquarius, Jordan (1907, p. 238) stated that there are ‘‘no
teeth on vomer ... ,’’ but in his key to the genera of the Quinquariinae (ibid., p. 236), he stated that
there are ‘‘teeth on vomer’’ in Quinquarius. In a 154-mm. specimen of Quinquarius (= Pentaceros) japoni-
cus (Stanford University no. 18191), which we have examined, teeth are present on the head of the vomer,
but not on its shaft.

326 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

most be justified in uniting them.” But he distinguished them on the fol-
lowing basis (ibid., p. 876) :

Pseudopentaceros Quinquarius (== Pentaceros)
Seales in more than 70 series Seales in fewer than 60 series
Exposed bones of head without Exposed bones of head with large
marked sensory pores sensory pores
In juveniles: In juveniles:
No horns above eye Horns above eye
Pelvies not very widely separated Pelvies widely separated
Depth of body less than half Depth of body more than half
leneth length

The difference in scale counts does not impress us as of generic signifi-
cance.

In our specimen of Pentaceros richardsoni, the exposed bones of the
head have large sensory pores. The presence of such pores in this species
has been noted also by Welander, Johnson, and Hajny (1957, p. 245) and
Whe (957, p37).

The presence or absence of horns above the eye in juveniles may not be
a reliable character in this family. In another species of this family, these
structures, which had been considered characteristic of the young of that
species, were absent in one young specimen (Barnard, 1927, p. 621).

The distance between pelvics has not been determined for juveniles of
Pentaceros richardsoni. (In the four adults that we have examined (see
table 1), this distance ranges from 2.5 to 3.3 in length of head.)

The difference between a body depth of 2.3 in a juvenile of Pentaceros
richardsont (MeCulloch and Phillipps, 1923, p. 18) and a body depth of 1.8
in juveniles of P. capensis of comparable size (Smith, 1951, p. 881) does
not impress us as of generic significance.

We therefore concur with Welander, Johnson, and Hajny (1957, p. 245)
in regarding Pseudopentaceros Bleeker, 1876, as a subjective synonym of
Pentaceros Cuvier in Cuvier and Valenciennes, 1829.

SPECIFIC DISTINCTION

A lower number of dorsal soft-rays and a higher number of seales dis-
tinguish Pentaceros richardsoni from P. capensis, as well as from P.
japonicus Doderlein in Steindacher and Déderlein, 1884, and from P.
hendecacanthus McCulloch, 1915; a lower number of dorsal soft-rays dis-
tinguishes P. richardsoni from P. decacanthus Giinther, 1859 (based on a
dried specimen less than two inches long). See table 2.

(The data in table 2 suggest that P. japonicus and P. hendecacanthus,
and possibly even P. decacanthus, may not be specifically distinet from
P. capensis.)

327

PENTACEROS

FOLLETT AND DEMPSTER

VoL. XXXIT]

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328 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER.

NOMENCLATURE

GENERIC NAME. The generic name Pentaceros Cuvier in Cuvier and
Valenciennes (1829, p. 30) is valid. For some time it has been thought to
be preoccupied either by Pentaceros Schulze (1760, p. 50) or by Pentaceros
Schroter (1782, p. 58), but it is evident that under the present rules neither
of those earlier names is nomenclaturally available.

For a name to be available, the International Code of Zoological Nomen-
clature requires that its author must have consistently applied the principles
of binominal nomenclature in the work in which the name is published
(International Commission on Zoological Nomenclature, 1961, Art. 11e).
Binominal nomenclature is the system under which each species receives a
name consisting of two words, of which the first is the generic name and
the second is the specific name (ibid., Glossary, p. 148).

Schulze (1760) did not consistently, if at all, apply the principles of
binominal nomenclature (see Clark, 1908, p. 517; Springer, 1909, p. 183;
Bather, 1909, p. 40; and Jordan and Richardson, 1909, p. 192). It cannot
be demonstrated that the names used by Schulze are even names of genera
or species. Pentaceros Schulze (1760, p. 50) is a deseriptive term applied
to a group of starfishes of indeterminate rank. It includes two subgroups,
also of indeterminate rank, the planae (plural) and the gibbae (plural).
Jordan and Evermann (1917, p. 126) remarked that “obviously this is not
scientific nomenclature,” and Fisher (1908, p. 91) stated that “there is no
evidence that Schulze knew anything about binomial nomenclature, for
he does not conform to the Linnaean system . . .”

Any doubt regarding the nomenclatural status of Pentaceros Schulze,
1760, has now been eliminated by Opinion 636, which placed Schulze, 1760
(misspelled “Schultze”), on the Official Index of Rejected and Invalid
Works in Zoological Nomenclature, on the ground that the author did not
apply the principles of binominal nomenclature (International Commission
on Zoological Nomenclature, 1962, p. 263).

Schroter (1782), the next author to use the name Pentaceros, did not
consistently apply the principles of binominal nomenclature. In his only
mention of Pentaceros (ibid., p. 58), he merely cited the polynominal term
that had been applied to the “five-horned star” by the pre-Linnaean author
Linck :

“Fig. ITT. ist auf der Seite des Riickens vorgestellt, und kommt vor im
Link [sic] de stellis mar. tab. III. fig. 3. auf der Riickenseite, und Tab. II.
fig. 3. auf der Seite der Miindung. S. 21. 22. stehet er unter dem Geschlecht
Pentaceros, der finfhornige Stern, und heisst Pentaceros gibbus turritus
pluribus velut turriculis munitus. Die Seepastete Rumph hollind. p. 39.
Beym Linné ist er ed. XII. p. 1100. Gen. 298, sp. 7. Asterias nodosa oder

Asterias stellata, radius converis longitudinaliter elevatis muricatis. .. .

VoL. XXXIT] FOLLETT AND DEMPSTER: PENTACEROS 329

We concur with Fisher (1908, p. 98), who stated, “Schroéter, in 1782
(Musei Gottwaldiani Testaceorum, Stellarum marinum, ete., Niirnberg, 58),
used Pentaceros, but he is not a consistent binomialist, and his ‘generic’
names are not tenable.”

Since Pentaceros Cuvier in Cuvier and Valenciennes is not preoccupied,
the name Quwinquarius Jordan, 1907, which was proposed “to replace
Pentaceros Cuv. & Val., regarded as preoccupied” (Jordan, 1920, p. 524),
is relegated to synonymy.

The name Pentaceros Cuvier in Cuvier and Valenciennes has been used
in the primary zoological literature within the past fifty years (Thompson,
1916, p. 184), and is therefore not to be considered a forgotten name
(nomen oblitum); see International Commission on Zoological Nomencla-
ture (1961, Art. 28b).

FAMILY NAME. The oldest available family-group name based on the
nominal genus Pentaceros is Pentacerotinae (correction by Guill, 1893b,
p. 134, of Pentaceratina Giinther, 1859, p. 212). The name Pentacerotinae
fulfils the requirements of availability of family-group names (International
Commission on Zoological Nomenclature, 1961, Art. lle): it was based on
the name then valid for a contained genus (Pentaceros) and it was a noun
in the nominative plural; it was clearly used to denote a suprageneric taxon
(it was designated “group” but was used in the sense of subfamily), and
was not merely employed as a plural noun or adjective referring to the
members of a genus. The subfamily name Pentacerotinae, so used, makes
available the family name Pentacerotidae Giinther, 1859 (see International
Commission on Zoological Nomenclature, 1961, Arts. 29, 36). The name
Pentacerotidae, which has priority over the name Histiopteridae Jordan
(1905, p. 398, and footnote to p. 585), is the valid name of this family.
(Under the present rules, the valid name of a family is the oldest available
name applied to it. See International Commission on Zoological Nomenela-
ture, 1961, Art. 23.)

The correct form of the family name based on Pentaceros is Pentace-
rotidae, not Pentaceridae or Pentaceratidae. The genitive singular of Penta-
ceros, an adjectival form (used as a noun) based on -ceras, is Pentacerotos,
from which is derived the family name Pentacerotidae (LL. W. Grensted, clas-
sical adviser to the International Commission on Zoological Nomenclature,
m litt., February 9, 1962).

Family-group names based on Pentaceros have been used by a number
of authors: the family name Pentacerotidae, by Bleeker (1860, p. 52, cor-
rection by Gill, 1885, p. 210, of Pentacerotoidei), by Gill (1893a, pp. 105,
116; 1893b, p. 134), by Jordan and Evermann (1902, p. 351), by Hutton
(1904, p. 41), by Jordan (1905, p. 333), by Smith and Pope (1906, p. 479),
and by Jordan and Richardson (1909, p. 192); the subfamily name Penta-

330 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

cerotinae, by Giinther (1859, p. 212, correction by Gill, 1893b, p. 134, of
Pentaceratina), by Bleeker (1876, pp. 269, 287, correction by Gill, 1893b,
of Pentacerotiformes), by Boulenger (1910, p. 660), and by Thompson
(1916, p. 134, correction of Pentaceratinae).

SPECIFIC NAME. The specific name of this fish was spelled in two ways
in the original publication (Smith, 1849): richardsonw in the index to
the plates, and richardsoni in the text and on plate 21.

If a name is spelled in more than one way in the original publication,
the spelling adopted by the first reviser is to be accepted as the correct
original spelling (International Commission on Zoological Nomenclature,
1961, Art. 32b). An author, to be a first reviser, must cite the names in
question, make it clear that he believes them to represent the same taxon,
and choose one as the name of that taxon (ibid., Art. 24a(i) ).

Since we have found no publication, other than the original deseription,
in which both original spellings of this specifie name were cited, we con-
elude that neither spelling has been adopted by a “first reviser.”

The Code recommends that a specific name based on a modern personal
name that is neither Latin nor latinized nor of Greek origin should end in
-t rather than -a (abid., Appendix D, Recommendation 16). As first revisers,
we therefore choose the spelling richardsoni in preference to the spelling
richardsonu for the specific name of this fish.

CoMMON NAME

We propose the common name “pelagic armorhead” for Pentaceros
richardson.

Although the name “boarfish” has been applied to this and other species
of the Pentacerotidae by a number of ichthyologists (see, for example,
Giinther (1880, p. 388), Tenison-Woods (1883, p. 183), Waite (1898, p. 33;
1911, p.216; 1921, p. 117), Roughley (1916, pp. 9° 127); PhillippessGeen
p. 18), MeCann (1953, p. 12), Welander, Johnson, and Hajny (1957,
p. 244), Smith (1961, p. 242), and Clemens and Wilby (1961, p. 219) ), we
believe that “boarfish” should be restricted to the species of the Caproidae.
Such a restriction appears justified on the following grounds:

1) Capros (the name of the type-genus of the family Caproidae) itself
means “boar.” The vernacular “aper,” meaning “wild boar,’ was applied
to the fish currently known as Capros aper (Linnaeus, 1758) more than 400
years ago (Boussuet, 1558, p. 28)—and more than 300 years before “boar-
fish” was applied to any species of the Pentacerotidae.

2) Use of the name “boarfish” for Capros aper and other species of
the Caproidae is deeply ingrained in the literature of the past century
(see Yarrell, 1859, p. 258; Couch, 1869, p. 142; Buckland, 1880, p. 77;
Giinther, 1880, p. 449; Tenison-Woods, 1883, p. 188; Gill, 1885, p. 209;

VoL. XXXIT] FOLLETT AND DEMPSTER: PENTACEROS 331

Jordan and Evermann, 1898, p. 1663; Jordan and Fowler, 1902, p. 521;
Aflalo, 1904, pp. 129, 137; Jordan, 1905, p. 398; Boulenger, 1910, p. 666;
Meek, 1916, p. 305; Barnard, 1925, p. 380; Norman, 1931, p. 378; Gregory,
1933, p. 272; Munro, 1938, p. 77, and 1955, p. 89; Jenkins, 1950, p. 81;
Berry, 1959; Food and Agriculture Organization, 1960, p. 99; Myers, 1960,
pp. 89, 96; Herald, 1961, p. 160).

3) The name “boarfish” has been adopted for species of the Caproidae
by the American Fisheries Society Committee on Names of Fishes (1960,
p. 24).

No English name other than ‘“‘boarfish” has been widely used for species
of the Pentacerotidaet. It seems desirable therefore to apply a new common
name to those species. Our choice of such a name, “armorhead,” is based
on the rough, bony plates of the head—a conspicuous and unique character.

The attributive “pelagic” in the proposed name, “pelagic armorhead,”
refers to the occurrence of Pentaceros richardson far offshore.

SUMMARY

1) A specimen of Pentaceros richardson Smith from the coast of Cali-
fornia is described, with meristic and morphometric data and notes on the
dorsal-fin supports, the locking mechanism of the dorsal and anal spines, and
the osteology of the caudal region. The bilateral asymmetries of the dorsal
spines, anal spines, and branchiostegal membranes of seven specimens of this
species are shown to be variously associated.

2) Distributional notes and an annotated synonymy of Pentaceros
richardsont are provided.

3) Pseudopentaceros Bleeker, 1876 (type species, Pentaceros richard-
soni) is shown to be a subjective synonym of Pentaceros Cuvier in Cuvier
and Valenciennes, 1829 (type species, Pentaceros capensis).

4) The specific distinction of Pentaceros richardsoni from P. capensis
Suvier mm Cuvier and Valenciennes, P. decacanthus Giinther, P. japonicus
Doderlein 7 Steindachner and Doderlein, and P. hendecacanthus MeCulloch
is confirmed. Pentaceros japonicus and P. hendecacanthus, and possibly
even P. decacanthus, are regarded as questionably distinet from P. capensis.

5) The generic name Pentaceros Cuvier in Cuvier and Valenciennes,
1829, thought to be preoccupied either by Pentaceros Schulze, 1760, or by
Pentaceros Schroter, 1782, is shown to be valid. Quinquarius Jordan, 1907,
proposed to replace Pentaceros Cuvier in Cuvier and Valenciennes, is rele-
gated to synonymy. The family name Pentacerotidae Giinther, 1859, which
has priority over Histiopteridae Jordan, 1905, is shown to be valid.

6) Of the two original spellings, richardsoni is adopted as the correct

4. The vernacular “‘porgy,” used for species of this family by Jordan (1907), has been adopted for cer-
tain species of the Sparidae by the American Fisheries Society Committee on Names of Fishes (1960, p. 32).

332 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

original spelling in preference to richardsonu for the name of this species.
7) The common name “pelagic armorhead” is proposed for Pentaceros
richardson.

ACKNOWLEDGMENTS

We wish to express our gratitude to Mr. David Moss of A. Paladini, Inc.,
for enabling us to obtain the first specimen of Pentaceros richardson
known from California; to Dr. George S. Myers of Stanford University and
Dr. Arthur D. Welander of the University of Washington, for the loan of
specimens; to Dr. Tokiharu Abe of the Tokai Regional Fisheries Research
Laboratory, Dr. James E. Bohlke of the Academy of Natural Sciences of
Philadelphia, Mr. Nicholas Camarda of A. Paladini, Inc., Prof. L. W. Gren-
sted, classical adviser of the International Commission on Zoological Nomen-
clature, Mr. John D. Hopkirk of the University of California, Mr. Walter W.
Schneebeli of the Steinhart Aquarium, and Dr. Norman J. Wilimoysky of
the University of British Columbia, for information; to Mr. C. Edward
Crompton and Mr. Maurice C. Giles of the California Academy of Sciences,
for prints of the photographs; and to Dr. Carl L. Hubbs of the University
of California, for a critical review of the manuscript.

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336 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

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338 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

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PROCEEDINGS
OF THE

CALIFORNIA ACADEMY OF SCIENCES
FOURTH SERIES

G Dallas Hanna Anniversary Volume

Vol. XXXII, No. 11, pp. 339-362; 7 figs. September 10, 1963

CONTRIBUTIONS TO THE PETROGRAPHY
of the
GALAPAGOS, COCOS, MALPELO, CEDROS,
SAN BENITO, TRES MARIAS, AND
WHITE FRIARS ISLANDS.

| Marine Biological Laboratory
By i icr AR Y
CED 1062
Charles W. Chesterman | SEF 53
Honorary Curator of Mineralogy | VOODS HOLE. MA as
California Academy of Sciences WUUDS HULE, MASS bax

CONTENTS

Page

LL EINES TOV CT DOV os Ss OA aa Pe Ue 340
ReeeeAOsml Gl amGs. Se. tees ai he Lea, RO eel 340
ocipiom and General Geoloey® 6.20. oi0css cose Sen ne ee. Oe 540

Re CULMeLeN MANO Lert eres iG, 2, Seon 2 Not SAS nS ee eA aes La +1
EMMA OMOTIVAL Set Clee oe 2 Ie PNR SO een ee Fae Wl 541

FepsTa ee es LAIN ese Sh Sg. Sh Se eae cl glow eee ee eh Si 342
Ronweralighand estar. 22%. 40/2 $5 Pk 84 Rs ee ee 4 ad43

a! CASSIE 0 MAA ae em ae RO OR amy 1 Mis, Pei Ea Sa Pt | eae 544
DeKmMOtmemliglime nk tess cet te te EES Oe ek O44
indetaiioable island! 0: 10% Mes oe ed Di eu 345

Pu beIMevelems lamers. oxo y woes Ana N ek aco Poe 5 Ack a 345

is tate OAD S| ANNO te Ae sas eeu A ay ls eos, Gh weve ee 2 2 346
Whitihamnislsl ancl Ae as Ae Wee le pene ne 5 atten ce ncenes) Sle depen ou 347

340 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SrEr.

CONTENTS— Cont:

Galapagos Islands—Cont. Page
Charles Island .2.. S. 22e 0502" 0 See oo ee eee 349

Hood sland. \) 0.8 eee Oe eee eee 350

Cocos: Tseland von. hssage eed, acs 5 ee OE bes eee 350
Malpelo: Islam) ..2(27. os x da75. as Oe © obuaere Ok oe ee 354
Cedros Island siete d Bios. otc se Oe oe eee 395
nan: Benito. Islands's.2¢. 5 advance tase ge ne ees ee oe ee 355
Tres Marias Islands» wee 3.3.5 {324 oo fos oe Bs on eee 396
White Friars. Islands. 2.6.0) .ck. 52% Zaelee eks ooh eee 359
Phosphate Rock |. .855 . 2262.45.02... UL eee 360
Bibhography “02 22.653. ve ae eee ee ee eee 361

INTRODUCTION

For a period of several vears, between 1905 and 1932, the California
Academy of Sciences conducted extensive scientific expeditions to the Gala-
pagos, Cocos, Malpelo, Cedros, San Benito, Tres Marias, and White Friars
islands.

During the course of studies of these eastern Pacific Ocean islands,
numerous volcanic rocks were collected by Washington Henry Ochsner from
the Galapagos Islands (ineluding Abingdon, Albemarle, Barrington, Bind-
loe, Charles, Chatham, Gardner, Hood, Indefatigable, Jervis, South
Seymour and Tower islands) ; from Cocos, Malpelo, Tres Marias and White
Friars islands by C. B. Perkins, Leo G. Hertlein, and G Dallas Hanna;
and metamorphic and plutonic rocks from Cedros, San Benito and Tres
Marias islands, by G Dallas Hanna. The voleanie rocks that were col-
lected on these islands range in composition from basalt to rhyolite. Those
lavas on the Galapagos Archipelego and Cocos Island are principally
basaltic in composition, although a few are of andesitic composition (Rich-
ardson, 1933, p. 46). One lava from Cocos Island is a latite, and several
collected from Maria Madre of the Tres Marias islands are rhyolitie in
composition. Glaucophane schists were collected on Cedros and San Benito
islands and several plutonic rock types were collected on Maria Madre
Island.

Special thanks are due to Dr. Leo G. Hertlein for his encouragement
and suggestions during the course of this investigation, and especially to
Dr. G Dallas Hanna who made available the rock specimens and suggested
that a petrographic study be made of them.

GALAPAGOS ISLANDS

LOCATION AND GENERAL GrEoLOGY. The Galapagos Islands are on the
equator about 650 miles west of Eeuador. They form an archipelago con-

Vout. XXXII] CHESTERMAN: EASTERN PACIFIC PETROGRAPHY 341

GALAPAGOS ISLANDS

_ BDCOLLECTING LOCALITY (shown when t Abingdon

stated in Ochsner's notes) 65
Bindloe

6 Jervis ee Seymour

1
South Seymour

Narborough

Duncan

(2-25{71)}3)

Barrington

Albemarle

9)
)
e

Chorles

Bose from U S Hydrographic Chart No 1798

Figure 1. Map of the Galapagos Archipelago showing the major islands and the
localities where rock specimens were collected.

sisting of 13 islands and many islets and projecting rock masses. The
islands, except for minor amounts of sedimentary beds, are wholly of
voleanie origin and contain at least 2000 cones and eraters (Darwin, p.
110-131, 1891). This is a very conservative estimate for it has been shown
that there are at least 2500 cones and craters alone on Albemarle Island
(Banfield et al., 1956, p. 222) which is the largest island in the archipelago.
Some of the voleanoes have erupted during the last hundred years (Sapper,
1917, p. 95). In addition to the many eraters, there are lava flows and
deposits of tuffs and tuff-breccias.

PeTrRoGRAPHY. Most of the crystalline rocks that make up the Galapagos
Islands are basaltic in composition, and porphyritie with phenocrysts of
basic plagioclase set in a microcrystalline groundmass. Olivine is not com-
mon, but when it is present it is more readily visible in lavas collected on
Chatham, Charles, Gardner, Hood, Jervis, and Indefatigable islands.

ABINGDON IsLAND (Pinta; Geraldino). Two specimens of basalt were
collected on Abingdon Island, the northernmost island in the archipelago.

342 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER.

Specimen no. 88 is typical of a great mass which forms an extensive flow
that covers much of the island. Specimen no. 90 is from a more recent flow,
and is less extensive in its occurrence than specimen no. 88 over which it
hes.

Specimen no. 88 is a dark, porphyritic, vesicular rock containing pheno-
erysts of plagioclase up to one-half inch across enclosed in a fine-grained
groundmass of feldspar laths, grains of colorless pyroxene and black opaque
olass. The feldspar phenoerysts are subhedral in shape, and show well-
developed zoning and twinning. They are unaltered and have a composition
in the range of intermediate labradorite (An;.). The groundmass feldspar
is in irregular laths and is seldom more than .2 mm. in length. They, too,
are fresh, twinned and have a composition slightly more sodie than the
feldspar phenocrysts. The augite is fresh and occurs in subhedral color-
less grains. The principal constituent of the groundmass is dark, opaque
basaltic glass which derives its dark color in part from small dust-like
particles of magnetite. Specimen no. 90 is not porphyritic, but shows a well-
developed hyaloophitie texture in thin-section. It is a fine-grained rock
and under the microscope one ean see laths of feldspar and grains of augite
enclosed in a dark glassy groundmass. The feldspar laths are small and
not more than .7 mm. in length, but reasonably accurate determinations
indicate a composition in the range of intermediate labradorite (An,;-50).
This mineral constitutes about 35 per cent of the rock. The augite is colorless
and is in twinned, subhedral grains that constitute about 10 per cent of
the rock. The remainder of the rock is dark basaltic glass.

BINDLOE ISLAND (Marchena; Torres). Of the three rock specimens col-
lected on Bindloe Island, only one, no. 86, was available for study. This
specimen was collected from a lava flow well exposed along the west coast
of the island which appears to consist largely of pyroclastic deposits with
a few narrow, smooth-surfaced lava flows.

Specimen no. 86 is basalt. It is porphyritic, slightly vesicular, and con-
tains phenocrysts of feldspar up to one-half inch across enclosed in a fine-
grained, medium-gray colored groundmass.

The feldspar phenoerysts are euhedral to subhedral in shape, unaltered
and constitute from 15-20 per cent of the rock. This mineral shows zoning,
a combination of twinning, and a composition in the range of intermediate
labradorite (An;.). The groundmass feldspar is in irregular-shaped laths
that range up to .6 mm. in length. The laths are fresh and constitute as
much as 45-50 per cent of the rock. They are twinned, but not zoned, and
have a composition slightly more sodie than the feldspar phenocrysts, and
are in the range of An,; to Anso. The augite is in angular grains which are
seldom more than .2 mm. across. The mineral is slightly pleochroie in pale
pinkish to pale greenish colors, unaltered, and is untwinned and unzoned.

VoL. XXXII] CHESTERMAN: EASTERN PACIFIC PETROGRAPHY 343

COCOS, MALPELO, CEDROS, SAN BENITO,
TRES MARIAS AND WHITE FRIARS
ISLANDS

0) Tres 4%

Morio Madre . Nate e ae
= OE

Moria <P)
Magdalena

White Friars ;
Maria Cleofos(y Islands
TRES MARIAS ISLANDS

Cocos |

o
5

Cocos ISLAND MALPELO ISLAND

Figure 2. Map of the Pacific Ocean off the west coast of Central America show-
ing the location of Cocos, Malpelo, Cedros, San Benito, Tres Marias and White
Friars islands.

It constitutes about 35 per cent of the rock. Magnetite which occurs in
irregular grains, scattered among the other minerals, constitutes about 5
per cent of the rock.

TowER ISLAND (Genovesa, Ewres). Only one specimen was collected on
Tower Island, and this is a basalt from one of several prominent lava flows
well exposed in Darwin Bay. The basalt from Darwin Bay, is moderately
vesicular and porphyritic, with phenocrysts of feldspar up to one-quarter
inch across set in a fine-grained, dense, gray groundmass.

The feldspar phenocrysts are euhedral and subhedral in shape, and
show well developed zoning and twinning. They are in the composition
range of intermediate labadonite An;,-;;. The erystals are slightly altered
and contain small, dark, angular and irregular shaped inclusions. The
groundmass feldspar occurs in poorly developed lath-shaped erystals. They
are fresh and range in composition from Any; to Ansp.

Augite occurs as euhedral phenocrysts and as anhedral grains in the
groundmass. This mineral is common, fresh and lacks zoning and twinning.

344 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER.

Olivine occurs sparingly as euhedral grains in the groundmass. It is color-
less where unaltered, but commonly shows alteration to golden brown id-
dingsite.

JERVIS ISLAND (Rabida). Four rock samples were collected on Jervis
Island, and only one of these, a lava, was collected from loose boulders
which occur on the north side of the island.

The rock specimen no. 38, collected from Jervis Island is an olivine
basalt. It is porphyritie and contains phenocrysts of olivine and feldspar,
both as much as one-quarter inch across, set In a medium-grained gray
eroundmass. In thin section the rock has a xenomorphic-granular texture,
vet with phenocrysts of olivine and feldspar set in a granular groundmass
composed of the same minerals plus augite and a small amount of glass.

The plagioclase in the phenocrysts and in the groundmass appear to be
identical in all respects except in size, of which the phenocrysts are as much
as one-quarter inch across and the groundmass grains rarely more than .3
mm. across. The feldspar is fresh and constitutes between 25 and 30 per
cent of the rock. The erystals are twinned and a few of the phenocrysts
show zoning. Inclusions of dark glass are common and appear to be con-
eentrated along the margins of the erystals where they are elongated and
arranged in rows parallel to cleavage and twinning planes. The feldspar is
intermediate labradorite, Anso_;5.

Augite is pale greenish in color, and occurs as anhedral erystals as
mueh as .5 mm. across. This mineral, too, has dark glass inclusions, but in
lesser amounts than the feldspar. The augite is fresh and constitutes about
50 per cent of the rock.

Olivine is in colorless anhedral crystals that show zones of yellow which
are due to limonite and not the result of alteration to iddingsite.

SoutH Seymour ISLAND (Baltra). Three specimens were collected on
South Seymour Island, and only two of these, nos. 27256 and 32, were
available for study.

Specimen no. 27256 is fossiliferous limestone from one of beach terraces
near the southern part of the island (no. 2 of Ochsner), and specimen no.
32 is from a lava flow that rests upon the fossiliferous terrace deposits, also
in the southern part of the island.

Specimen no. 27256 should properly be called a fossiliferous tuffaceous
sandstone. It contains white fossil shell fragments and dark brown, dark
red and buff colored voleanic rock fragments enclosed in a medium-gray,
fine-grained, dense groundmass.

Under the microscope one can see rounded grains of orange-red pumice
containing phenocrysts of plagioclase, An;o, rounded grains of plagioclase,
Any;-50, Olivine altered almost wholly to orange-red iddingsite, and white
shell fragments all enclosed in a murky matrix of secondary ealcite.

VoL. XXXII] CHESTERMAN: EASTERN PACIFIC PETROGRAPHY 345

Specimen no. 32 is andesite. It is a fine-grained, weakly prophyritie,
medium- to dark-gray colored rock. Phenoerysts of plagioclase are un-
common, and range in size from .6 to 1.0 mm. The erystals are lath-shaped,
fresh, and show simple albite and carlsbad twinning. They are in the com-
position range of An,, to Any. The groundmass plagioclase occurs as
microlites which are less than .5 mm. in length. They are abundant and
constitute about 30 per cent of this rock. The microlites of plagioclase
have the same range in composition as the plagioclase phenocrysts. Augite
is also very common, and it occurs in two distinct sizes: as small anhedral
grains less than .1 mm. in diameter scattered throughout the rock and as
larger subhedral grains up to .5 mm. across. The smaller grains are nearly
colorless and the larger grains show pale pinkish and greenish pleochroie
colors. All of the augite is fresh and untwinned. Magnetite is common and
constitutes about 10 per cent of the rock.

INDEFATIGABLE ISLAND (Santa Cruz; Bolivia; Norfolk; Porter; Valdéz:
Chavez; San Clemente). Five specimens were collected on Indefatigable
Island. Of these specimens, no. 28, was the only specimen available for
study. It is a dense, fine-grained, dark gray, slightly vesicular olivine
basalt. Under the microscope, the olivine basalt has a diabasic-intereranular
texture. Plagioclase, which constitutes approximately 40 per cent of the
rock, occurs in lath-shaped erystals that range up to 1.0 mm. in leneth. It
is twinned, fresh and has a composition of An; o. Augite oceurs in large
irregular-shaped crystals and constitutes approximately 45 per cent of the
rock. It is weakly pleochroic in shades of pale purple and light ereen.
Olivine is colorless when fresh, but golden-yellow when altered to iddingsite.
It oceurs in subhedral-shaped grains and constitutes about 15 per cent of
the rock. Both magnetite and ilmenite are present. Magnetite is in aneular
and rounded grains whereas the ilmenite is in small skeletal crystals,
usually enclosed in the feldspar.

ALBEMARLE ISLAND (Isabela; Santa Gertrudis). Eleven specimens were
collected on Albemarle Island. Specimens numbered 44, 47, 48, 51, 52, and
54+ are voleanie rocks, and of these specimens only nos. 48, 49, 50, and 52
were available for study. Specimen no. 48 is basalt. It is non-vesieular,
porphyritic, and contains phenocrysts of feldspar and augite up to one-
eighth inch in length enclosed in a dense, fine-grained, medium-gray colored
eroundmass. The phenocrystic feldspar, in the composition range of An,;
to Ang, occurs as anhedral and subhedral shaped grains. They are com-
pletely twinned, but not zoned or altered. Augite occurs as small granular
inclusions in the larger feldspar phenocrysts. The groundmass feldspar,
whose composition is approximately An; , occurs as irregular-shaped laths
that are slightly more sodic than the feldspar phenocrysts. The groundmass
feldspar constitutes about 25 per cent of the rock. Augite, although ocur-

346 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

ring as phenocrysts and in the groundmass too, is more abundant in the
eroundmass. It is pale-greenish in color and shows twinning but no zoning.
The phenocrysts are euhedral in shape, whereas the groundmass grains are
subhedral and anhedral in shape. Few of the phenocrysts of augite are
rimmed with dust-like particles of magnetite, and a few of the grains in
the groundmass are almost completely clouded by magnetite.

Specimen no. 49 is a erystal-lithic-vitrie tuff. It contains pyroclasts of
plagioclase, porphyritie basalt and andesite, and pumice set in a ground-
mass of fine-grained, partly altered glass. The feldspar pyroclasts are
subhedral and anhedral in shape, and range in composition from Any to
Ans). The larger grains contain bleb-shaped inclusions of pale-yellowish
glass, arranged parallel to twinning planes. The larger grains are un-
altered, but a few of the smaller feldspar grains have a narrow rim of
pale-yellowish opal. Augite pyroclasts are colorless, and are rimmed by
yellowish nontronite. The lithic pyroclasts consisting essentially of andesite
and basalt, are rounded and also rimmed by opaline silica. The pumice
pyroclasts are rounded and contain a few erystals of potash feldspar
(sanidine) as well as some yellowish colored nontronite. A small amount
of secondary calcite is present.

Specimen no. 50 is tuff-breecia. It consists of angular and rounded,
dark-gray pumice fragments up to one-half inch across enclosed in a
medium- to fine-grained, grayish- and greenish-brown matrix of partly al-
tered volcanic ash.

Specimen no. 52 is a porphyritie basalt from Tagus cove. It contains
phenoerysts of plagioclase and augite up to one-quarter inch across en-
elosed in a fine-grained, dense, dark-gray, vesicular groundmass. The
plagioclase phenocrysts are in the composition range of An;; to Ango, and
occur in subhedral and anhedral grains that are well twinned and show
zoning. Inclusions of dark glass are present, and especially noticeable at
the interior of the feldspar grain where they are arranged in elongated
clots parallel to the twinning planes. The groundmass feldspar oceurs in
microlites that constitute about 30 per cent of the rock. They are twinned
and range in composition from An;, to An;;. Augite occurs as euhedral
phenoerysts and as anhedral grains scattered among the feldspar microlites.
The augite is fresh, though in places it is stained yellowish-orange by
limonite. It constitutes about 30 per cent of the rock. The remainder of
the rock is dark basaltic glass.

BARRINGTON ISLAND (Santa Fé). Three specimens were collected on
Barrington Island; and only one of these (no. 73), a specimen of olivine
basalt, was available for study. It is said to be characteristie of the lava
flows in the northern part of the island. This olivine basalt is a fine-grained,
slightly vesicular rock containing phenocrysts of olivine and plagioclase set

VoL. XXXII] CHESTERMAN: EASTERN PACIFIC PETROGRAPHY 347

Figure 3. Olivine basalt from Barrington Island, Galapagos Achipelago. Pheno-
erysts of olivine are rimmed by yellow iddingsite and are enclosed in a groundmass
composed of plagioclase laths (Angy) and granular augite. Plain light. x 30.

in a groundmass of feldspar laths and augite grains. The plagioclase
phenocrysts are labradorite (Ang) and occur as slightly rounded, twinned,
unzoned erystals as much as .2 mm. across. The feldspar phenocrysts are
fresh and constitute about 5 per cent of the rock. The groundmass feldspar
oceurs in lath-shaped crystals not more than .5 mm. in length. They are
fresh, twinned, not zoned and are in the composition range of An,; to Anso.

Olivine occurs as subhedral and anhedral grains up to .2 mm. in
diameter. They are colorless and slightly altered to golden-yellow idding-
site. Olivine also makes up part of the groundmass where it occurs in
rounded grains scattered among the feldspar laths. It constitutes approxi-
mately 20 per cent of the rock. Augite occurs in irregular-shaped erystals
that include small laths of feldspar. It is weakly pleochroic from pale
pinkish-purple to pale greenish, and constitutes about 40 per cent of the
rock.

CHatTHAM ISLAND (San Cristébal; Dassigney; Grande). Five rock
specimens were collected on Chatham Island. Only one of these specimens,

348 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

y . av en on me + «. - ax . Re’
Figure 4. Olivine basalt from Chatham Island, Galapagos Archipelago. Pheno-
crysts of olivine enclosed in a groundmass of plagioclase (An-=,)) and augite. Plain

light. x 30.

no. +3, an olivine basalt collected at the terminus of a lava flow south of
Sappho Cove, was available for study. The olivine basalt from Sappho Cove
is a fine-grained, finely-vesicular, medium-gray porphyritie rock which con-
tains phenocrysts of augite and plagioclase enclosed in an intergranular
eroundmass of feldspar laths and grains of augite and olivine. The feld-
spar phenocrysts, intermediate labradorite An;;—Ang., occur in subhedral
and anhedral crystals up to .5 mm. across. They are fresh, well-twinned
and constitute about 10 per cent of the rock. The groundmass plagioclase
(An;,) 1s shehtly more sodie than the feldspar phenocrysts, and oceurs as
lath-shaped crystals partly enclosed in the augite and partly forming a
mesh structure which is infilled by smaller augite crystals. The ground-
mass feldspar constitutes about 50 per cent of the rock. The augite occurs
in colorless and pale-purplish, irregular-shaped erystals that measure as
much as .6 mm. across. This mineral is unaltered and constitutes between
35 and 40 per cent of the rock. Olivine is in euhedral to subhedral grains.
It is colorless where fresh, but is altered in part to golden-yellow iddingsite,

Vou. XXXII] CHESTERMAN: EASTERN PACIFIC PETROGRAPHY 349

especially around the margins of the crystals. Olivine constitutes between
10 and 15 per cent of the rock.

CHARLES ISLAND (Santa Maria; Floreana). Eight rock specimens were
collected on Charles Island. Two of these specimens numbered 59 and 63,
were available for study. Specimen no. 59 was collected from an exposure
on Cormorant Bay and is a erystal-vitric-lthie tuff. In hand specimen
this rock is dark, greenish-black to black in color and contains angular
pyroclasts set in a fine-grained, dense matrix of dark-brown colored glass.

Under the microscope one can see angular pyroclasts of colorless augite
and irregular grains of basic andesine An,;, and rounded pyroelasts of
slightly vesicular, amygdaloidal, and vitrophyrie glass. These pyroclasts of
olass constitute a bulk of the rock. They are yellow in color, with the interior
part being canary-yellow and the rims a golden-yellow. The amyedules
are ovoid in shape and more or less completely filled with secondary calcite.
Augite and andesine are the crystalline constituents in the glassy pyroelasts.
The augite is colorless and is in angular erystals. The andesine (An, to
An,;) occurs in lath-shaped as well as in irregular-shaped erystals. The
matrix is a glass. It is grayish in color and contains finely divided magnetic
as well as microlitic laths of andesine and angular erains of colorless augite.

This tuff is similar in many respects to the tuffs found on Albemarle
Island, excepting the tuff from Cormorant Bay on Charles Island contains
very small amounts of lithic material such as pyroclasts of andesite and
basalt. Specimen no. 63 is said to be representative of a lava flow which
reached the ocean at Black Beach Road (Ochsner, unpublished manu-
seript). The rock is basaltic in composition. It is porphyritie and contains
phenocrysts of plagioclase and pyroxene set in a sub-ophitie groundmass
of plagioclase microlites and intergranular augite.

The plagioclase phenocrysts (An;; to Any) occur in subhedral and
anhedral grains. They are completely twinned, zoned and show a slight
amount of alteration. The larger phenocrysts, up to .5 mm. across, contain
numerous inclusions of glass, augite, and alteration products of the pyroxene,
all localized more toward the interior of the crystal and oriented parallel to
the twinning planes. The plagioclase phenocrysts constitute about 25 per
cent of the rock.

The plagioclase in the groundmass has the composition of An;; and
occurs in subhedral and irregular-shaped crystals up to .4 mm. in length.
They are twinned, but not altered, and constitute about 30 per cent of the
rock. The pyroxene is hypersthene which occurs in slightly colored euhedral
grains that exhibit weak pleochroice colors. Many of the larger erystals of
hypersthene have a rim of iddingsite; the smaller grains are almost com-
pletely altered to iddingsite.

Small amounts of magnetite and colorless glass are present in the rock.

350 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Hoop Isuanp (Espanola). Three rock specimens were collected on Hood
Island. Specimen no. 65 is from a lava flow exposed in the eliff south-
west of Gardner Bay, and specimen no. 69 from a lava flow exposed due
west of Gardner Bay. Specimen no. 65 is olivine basalt. It is porphyritie
and contains phenocrysts of olivine and plagioclase set in a dense, fine-
erained, dark-gray groundmass composed of plagioclase microlites and
erains of augite. The plagioclase phenocrysts are in rounded, zoned erystals
that range in composition from An,; to An;;. They contain numerous in-
clusions of very small crystals of apatite and augite which tend to be con-
centrated in the outer zone of the phenocrysts. The plagioclase microlites
are slightly more sodie than the phenocrysts and have a composition of ap-
proximately An,;.

Augite is in small, colorless, subhedral and anhedral grains. It is very
common and constitutes about 25 per cent of the rock. Olivine oeceurs as
anhedral crystals, of which many of them have been altered in part to id-
dingsite. Olivine constitutes about 20 per cent of the rocks. Specimen no.
69, too, is olivine basalt, and is similar in many respects to specimen no.
65, excepting the augite which in specimen no. 69 is pale purplish in color,
indicates the presence of titanium in more than normal amounts.

Cocos ISLAND

Cocos Island, governed by Costa Riea, is located on latitude 5° 32’N.,,
about 650 miles west of Cape Corrientes, Colombia. It is a voleanie island
approximately 4 miles long and slightly more than 2 miles wide, with a
maximum elevation of 1932 feet* above sea level. Six rock specimens were
collected on Cocos Island by W. H. Ochsner in 1905, and by Dr. Leo G.
Hertlein, California Academy of Sciences, February, 1932. Specimen nos.
14-1 and 17 are basalt, no. 12-1 is latite, no. 13-1 is tuff-breccia, and no.
18-1 is andesite (oligophyre).

Specimen no. 14-1 is basalt. It is a dense, fine-grained, medium-gray,
slightly porphyritie rock containing phenocrysts of plagioclase and hyper-
sthene enclosed in a sub-ophitie groundmass of plagioclase laths and
microlites and intergranular hypersthene. The plagioclase phenocrysts are
anhedral in shape and are in the composition range of An,;; to Ango. Both
zoning and twinning are common, and a few of the erystals are partly
altered. The larger phenocrysts contain many inclusions of glass, hyper-
sthene, and its alteration products. The inclusions tend to be localized
more at the central portion of the ervstal and in a few erystals they are
oriented somewhat parailel to the twinning planes. The plagioclase pheno-
crysts constitute about 25 per cent of the rock. The groundmass plagioclase

*This elevation is given in Sailing Directions for the west coasts of Mexico and Central America. United
States Naval Oceanographic Office, H.O. No. 26, p. 26, January, 1962.

Figure 5. Oligophyre from Cocos Island. The euhedral phenocryst of olivine
which has been altered entirely to iddingsite, is enclosed in a trachytic groundmass
of oligoclase laths and grains of olivine. Plain light. x 30.

occurs in microlites and subhedral, irregular-shaped laths, of which the
latter are no more than .4 mm. in leneth. The laths are twinned, but not
altered and are slightly more sodic than the phenocrysts. The groundmass
feldspar constitutes about 30 per cent of the rock. Hypersthene occurs in
euhedral crystals, of which a majority are phenocrysts. The large hyper-
sthene erystals have a narrow rim of yellow iddingsite, whereas, the small-
grains ot hypersthene are altered largely to yellowish-orange iddingsite.
Minor accessory materials include colorless glass and grains of black magni-
tite. Specimen no. 16, olivine basalt, is porphyritic and contains pheno-
erysts of olivine and plagioclase set in a groundmass of plagioclase laths
and grains of augite.

The plagioclase phenocrysts are subhedral to euhedral and exhibit
well developed twinning and zoning. Their composition is in the range of
An,;) to An;; and most of the alteration is at the interior of the crystals
where they are more ealeic. The groundmass plagioclase occurs in irregular,
lath-shaped erystals. Because of their small size their composition is dif-
ficult to determine, but it is in the range of An,;—-An; 9. The olivine is in

352 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER.

euhedral-shaped crystals which are altered largely to iddingsite, especially
around the margins of the crystals. Augite is present in small colorless grains.
Secondary materials include iddingsite, limonite, and sericite. A small
amount of dark glass occurs interstitially between the feldspar laths.

Specimen no. 17 is also an olivine basalt. It has a porphyritie texture
and contains phenocrysts of plagioclase and hypersthene enclosed in a fine-
erained groundmass of feldspar laths and grains of olivine and hyper-
sthene.

The plagioclase phenocrysts are in anhedral and subhedral-shaped
erystals and are in the composition range of An,, to Ans). They are com-
pletely twinned, zoned, and remarkably fresh. The groundmass plagioclase
is in small, lath-shaped erystals and is in the composition range of Any, to
An,;. They are twinned, unaltered and show no zoning.

Hypersthene occurs in euhedral-shaped erystals that show faint pleoch-
roism, and are rarely twinned. Olivine is in euhedral-shaped erystals that
show almost complete alteration to yellowish-orange iddingsite.

Specimen no. 12-1 from Cocos Island is latite. It is shehtly vesicular,
purplish-gray in color, and porphyritie with phenocrysts of plagioclase set
in a fine-grained groundmass. The plagioclase phenocrysts occur as euhedral
and subhedral grains of which some are skeletal and contain inclusions of
magnetite and glass. Their composition is oligoclase, in the range of Ans,
to An.;. The erystals are zoned and complexly twinned and constitute
about 10 per cent of the rock. The plagioclase in the groundmass is slightly
more sodie than the phenocrysts and have an An content of about 20 per
cent. The grains are anhedral and show shght alteration, twinning, and
zoning. They are very abundant and constitute about 80 per cent of the
rock. The pyroxene is diopsidic-augite and it occurs in pale, yellowish-
ereen anhedral grains. It makes up about 5 per cent of the rock. Magnetite
oceurs in irregular-shaped grains which are commonly surrounded by
yellow limonite.

Specimen no. 13-1 is a lapilli-tuff-breecia. This rock is well consolidated
and consists of angular and rounded fragments of dark-gray and brownish
vesiculated glass, lithic material, and mineral grains set in a fine-grained
ashy matrix. The vesiculated glass fragments range up to three-eighths of
an inch across. They are relatively unaltered, and contain very few pheno-
erysts, but refractive index determinations made on the glass indicate an
andesitie composition. In addition to the vesiculated glass fragment, the
rock contains also dark pyroclasts of vitrophyre which have phenocrysts
of intermediate andesine, (An,;) enclosed in a dark glass. The lithie frag-
ments are of three types: hornblende andesite, andesite, and basalt. The
hornblende andesite contains strongly pleochroic, needle-shaped crystals
of hornblende and fresh andesine (An, 4;). The andesite is composed of
andesine phenoerysts (An,;-50), set in a groundmass of small andesine

VoL. XXXII] CHESTERMAN: EASTERN PACIFIC PETROGRAPHY 353

vihew of ¢« ry

. iti’. F

Figure 6. Olivine basalt from Cocos Island. Phenocrysts of plagioclase (An,;
to An;,) and olivine enclosed in a fine-granied groundmass of feldspar laths and
grains of olivine and hypersthene. Plain light. x 30.

erains and colorless diopsidicaugite. The basalt pyroclasts, on the other
hand, are much darker in color than either of the andesite pyroclasts and
eontains phenocrysts of labradorite (Ango-«;), enclosed in a fine-grained
eroundmass of plagioclase microlites and small, anhedral grains of augite.
Olivine, too, is present in the basalt and this mineral is in small, anhedral
grains which are altered largely to iddingsite. Mineral pyroclasts include
euhedral grains of plagioclase in the composition range of An, to An,s,
subhedral, fresh and colorless grains of hypersthene, rounded and corroded
erains of brown hornblende, and a few rounded grains of quartz.
Specimen no. 18-1 is by far the most interesting rock collected on Cocos
Island. The rock is called an oligophyre for want of a better name. It is
fine-grained, buff-colored, and has a poorly defined flow banding. Under
the microscope one can see that the rock is weakly porphyritic. It is com-
posed largely of oligoclase. The phenocrysts of oligoclase are anhedral and
are in the composition range of Anz, to An,;. The egroundmass oligoclase

(Anso25) 18 In lath-shaped erains that are arranged in zones around the

354 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SEr.

oligoclase phenocrysts. Diopsidic-augite is present in colorless, subhedral
grains. Biotite is in dark brown, strongly pleochroic, anhedral grains.
Both of these ferromagnesian minerals are present in about equal amounts.
Maegnetite occurs in cubie-shaped grains. Secondary minerals include
limonite, which is probably derived either from the biotite or from the
small amount of pyrite that is present in the rock.

MALPELO ISLAND

Malpelo Island is on latitude 3° 59’ N., and lies about 250 nautical
miles westerly from Buenaventura, Colombia. It is a territory of the Re-
publie of Colombia and was known as early as 1530. The island is of
voleanie origin and is approximately one and one-half miles long, one-half
mile wide, and has a maximum elevation of 854 feet above sea level. Very
little has been written regarding the geology of Malpelo Island, and only
a limited amount is known about the rocks that are exposed there. Me-
Connell (1948) gives an excellent description of the fresh lava and phos-
phatized rocks that were collected by Comandante Eduardo Fallon of the
Colombian Navy in December, 1940 (Murphy, 1941, p. 57).

Two rock specimens were collected by C. B. Perkins, herpetologist, San
Diego Zoo, in December, 1931, and the following comments are based upon
a study of these specimens. One is pyroxene andesite and the other is
phosphate rock.

The pyroxene andesite is greenish-gray in color, dense, and fine-grained.
Under the microscope the rock is holoerystalline, microporphyritie, and
has a pilotaxitie texture. The plagioclase phenocrysts are fractured,
twinned, slightly altered, and range in composition from An,; to Any. The
plagioclase that comprises a bulk of the groundmass is in microlites whose
composition based upon limited determination is in the range of Ans, to
An,;. The plagioclase phenocrysts and groundmass microlites combined
constitute about 50 per cent of the rock. The pyroxene, diopsidic-augite,
occurs in colorless, anhedral grains usually no more than .05 mm. aeross,
and oceasionally in larger grains up to .1 mm. across. It constitutes about
20 per cent of the rock. Chloritie material is present, especially in close
association with the plagioclase phenocrysts and the larger pyroxene grains.
It is grass-green in color, moderately pleochroic, and locally stained brown
by limonite. Both quartz and chaleedony are present, but as secondary
minerals, and in small amounts. Seattered uniformly throughout the roek
are black, slightly angular, needlelike crystals. They are opaque and could
possibly be ilmenite. The phosphate rock is dark brownish-black, mottled,
and fine- to medium-grained. Under the microscope it is dark brown in
color with scattered irregular, colorless areas, composed of quartz and
chaleedony. Calcite is present in narrow, curved veinlets.

VoL. XXXII] CHESTERMAN: EASTERN PACIFIC PETROGRAPHY 355

Because of the lack of relict mineral grains in the phosphate rock, it is
exceedingly difficult to determine whether it was originally a pyroclastic
or flow rock. One could speculate, on the basis of lack of crystalline con-
stituents, that the original rock was a vitric-crystal tuff in which the
erystalline constituents were titanium and iron-bearing minerals, or titani-
ferous magnitite.

(CEpROS ISLAND

Cedros (Cerros) Island is situated just outside of Sebastian Vizeaino
Bay, on the west side of Baja California, midway between San Diego and
Cape San Lucas. The island is elongated in a north-south direction, and
measures about 22 miles long and 7 miles wide. The highest peak is Cedros
Mountain whose elevation is 3950 feet. Very little is known about the
geology of the island, but from observers (Hanna, 1925, p. 268) who have
visited the island we learn that it is composed largely of cherts, sandstones,
schists, and serpentine, all similar in many respects to the Franciscan rocks
(upper Jurassic) that are exposed so well in the San Francisco Bay area.
In addition there are Miocene and Pliocene sedimentary rocks on the eastern
side of the island and some voleanic rocks of presumably Tertiary age at the
southwest corner of the island.

Two rock specimens were collected on Cedros Island: Specimen no. 7
is tuff-breecia and specimen no. 9 is glaucophanized voleanie rock.

Specimen no. 7 tuff-breccia, is orange red in color, somewhat friable,
and contains partly altered pumice fragments, pyroclasts of basalt. and
grains of augite, altered olivine, biotite, hornblende, and plagioclase (Ans»),
all enclosed in a matrix of partly devitrified, brownish colored glass. The
basalt pyroclasts are of two types, one which is composed wholly of a felted
mat of plagioclase (An;.) laths, and the other which contains plagioclase
microlites (An; -;;) set in a dark brownish elass which constitutes at least
50 per cent of the rock.

Specimen no. 9, glaucophanized volcanic rock, the first of two meta-
morphie rocks to be mentioned in this paper, is a dense, fine-grained bluish
gray rock. Distinct foliation is lacking and the rock has a eataclastie
texture. Glaucophane is common and contributes largely to the bluish colora-
tion imported to the rock. It occurs in ragged and irregular-shaped erains
that are strongly pleochroic with & == pale greenish-colorless, y = lavender
blue, z—=blue. The glaucophane appears to have formed in part from a
colorless pyroxene, and it is altered locally to a fibrous mineral of low
birefringence. Tremolite is very abundant and constitutes at least 45 per

cent of the rock. It occurs in small, lath-shaped erystals and fibrous needles,
usually arranged in angular and subangular clots.
Both calcite and aragonite are present. The calacite is colorless and is

356 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SEr.

in veins, whereas the aragonite, which, too, is colorless, is present as discrete
erains forming a mineral phase in the rock. A pyroxene of diopsidic-augite
composition occurs as rounded grains that have altered in part to glauco-
phane and to pale-green chlorite. Quartz is not abundant, but this mineral
is present as small rounded grains scattered irregularly throughout the
rock.

Because of the lack of distinct foliation, the irregular distribution of the
elaucophane and tremolite, and the presence of relict pyroxene, it is be-
lieved that the glaucophane rock was derived from a basic voleanic rock.

San Benito ISLANDS

The San Benito Islands are about 18 miles west of Cedros Island. There
ave three islands in the group. West San Benito (the largest), Middle
San Benito (smallest), and Kast San Benito (next largest); all he close
together in an east west line. West and middle San Benito are composed
largely of Francisean chert, and Hast San Benito, the most rugged of the
three islands, contains schist and marble in addition to chert (Hanna, 1924,
pp. 373-374) (Van West, 1959, pp. 8-13). Only one specimen was available
from San Benito Islands for study, and it is a glauecophane schist, collected
on East San Benito Island.

The glaucophane schist is a medium-to-fine-grained, grayish-blue rock
that shows well developed foliation. Under the microscope one can see
wispy muscovite plates curved around earlier formed grains of glaucophane
and plagioclase. Glaucophane occurs in irregular-shaped to xenoblastie
erystals that show distinct pleochroism: = colorless, y= purplish blue,
z—=ultramarine blue. The crystals appear broken as though they had de-
veloped early and were fractured and disrupted during shearing stages of
metamorphism. Plagioclase (An;_;)) occurs as rounded and angular grains,
usually closely associated with quartz which, too, is in rounded grains. The
quartz shows undulatory extinction. Muscovite is present as wispy-shred-like
ageregates of small plates. It apparently developed late. Van West (1958,

pp. 535-87) deseribes glaucophane schists from San Benito Islands similar
to that mentioned immediately above and states that this is “. . . a common
variety among the elaucophane schists formed on East San Benito Island.”

Tres Marias ISLANDS

The Tres Marias Islands, consisting of Maria Madre, Maria Magdalena,
and Maria Cleofas, lie about 55 miles off the west coast of Mexico, west of
San Blas, Nayarit, between latitudes 21° and 22° N., and longitudes 106°
and 107° W. (see fig. 2).

Much has been written of the paleontology of these islands as a result

VoL. XXXII] CHESTERMAN: EASTERN PACIFIC PETROGRAPHY 357

of expeditions by the California Academy of Sciences. However a few re-
marks about some of the rocks collected in 1925 by G@ D. Hanna and E. K.
Jordon, and other rocks collected and reported upon by Hertlein and
Emerson (1959, p. +) will help to explain further the geology of these
islands.

All of the rock specimens were collected on Maria Madre Island which,
according to Hanna (1926, p. 69) is stated as consisting “. . . of granite,
chiefly, with a rim of diorite around the edges.” It is apparently from this
granitic core and dioritic rim that the specimens of biotite granite and
hornblende-quartz diorite porphyry were collected. The biotite granite
(CAS-35) is a medium-grained, light-colored rock which is speckled by
biotite and hornblende. Under the microscope the rock has a hypidiomorphie-
granular texture, the texture common to most plutonic rocks of this com-
position. Both orthoclase and plagioclase feldspars are present and where
they occur adjacent to another a myrmeketic border has developed on the
orthoclase. The orthoclase is in anhedral grains which constitute about 40
per cent of the rock. It is fresh for the most part and contains a small
amount of sericite along fractures. Plagioclase, in the composition range of
An, to An,;, occurs in subhedral and anhedral grains which, too, contain
some sericite. Zoning is absent, but the twinning in the plagioclase is com-
plex. This mineral constitutes about 20 per cent of the rock. Quartz is in
rounded, slightly fractured grains. It constitutes about 10 per cent of the
rock. Biotite is present in strongly, pleochroic, dark greenish-brown sub-
hedral grains. This mineral has been altered locally to deep-green chlorite
and contains small inclusions of zircon and apatite. Hornblende is in short
prismatic crystals. It is fresh, grass-green in color, and too contains in-
clusions of apatite and zircon.

CAS-—39, the hornblende-quartz diorite porphyry, is considerably darker
colored than the biotite granite. It has a distinct porphyritic texture and
contains phenocrysts of plagioclase, quartz, and hornblende enclosed in fine,
eranular groundmass of the same materials.

The plagioclase phenocrysts are subhedral in shape and range up to .4
mm. in size. They are zoned and complexly twinned and range in com-
position from core to rim, from An;, to An.;. Alteration is present, prin-
_eipally to sericite and calcite, with the cores being more highly altered than
the rims. This mineral constitutes about 40 per cent of the rock. The ground-
mass feldspar is in small lath-shaped crystals that average less than .5 mm.
in length. They range in composition from An,,—An.;, show slight altera-
tion to sericite, and locally form diabasic intergrowths with the quartz.
The groundmass plagioclase constitutes about 20 per cent of the rock. Quartz
occurs as angular phenocrysts up to .4 mm. across and as rounded grains
in the groundmass. It constitutes about 10 per cent of the rock. The horn-

358 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

blende is in pale-green crystals that have a moth-eaten appearance. It is
weakly pleochroic, contains inclusions of apatite and magnetite, and con-
stitutes about 5 per cent of the rock. A small amount of deep-green biotite
is present. It is strongly pleochroie in shades of green and brown, and
contains inclusions of apatite and magnetite. Several specimens of rhyolite
(CAS-37 and 38) were collected as float in the first main canyon southwest
of the village on Maria Madre Island. The rhyolite is a fine-grained, dense
rock and ranges in color from purplish-gray to gray. It shows flow banding
and contains white, rounded phenocrysts and small cavities of which the
latter now contain small, well-formed quartz crystals. The rock is porphyritie
and contains altered phenocrysts of plagioclase and green biotite enclosed
in a fine-grained groundmass which shows a micrographie structure. The
plagioclase phenoerysts are subhedral in shape, show simple twinning, and
are in the composition range of An,; to Ano. Alteration of the plagioclase
is principally to sericite and calcite. The phenocrysts constitute about 10
per cent of the rock. The groundmass feldspar is orthoclase and is in an-
hedral, untwinned, slightly altered grains. It forms microscopic intergrowths
with quartz and constitutes about 50 per cent of the rock. Excepting for a
few scattered ragged-appearing, green-colored phenocrysts of biotite, quartz
comprises the remainder of the rock. It occurs both as distinct anhedral
grains and in micrographic intergrowths with the groundmass feldspar.
In the same canyon where the rhyolite was collected and upstream from
the contact between the Pliocene sedimentary rocks and the underlying
eranite, a specimen (CAS no. 36) of rhyolite granophyre was collected
from an outerop of this rock. The granophyre resembles the rhvolite in
hand specimen. It lacks the flow banding which is so characteristic of the
rhyolite, but does show phenocrysts of feldspar and has a dull gray color.
The texture of the rock is granophyric and porphyritic, and the rock
contains rounded phenocrysts of plagioclase and quartz set in a grano-
phyrie intergrowth of quartz and orthoclase. The plagioclase, though
slightly altered is in the composition range of An,, to An,;. It is subhedral
in shape and measures up to .8 mm. in size. The groundmass feldspar,
orthoclase is less altered than the phenocrystic, and constitutes about 45
per cent of the rock. Quartz occurs sparingly as rounded phenocrysts.
However, it is more abundant in the groundmass where it is much coarser
erained when intergrown with the orthoclase. The quartz constitutes about
40 per cent of the rock. The biotite is green, strongly pleochroic, and occurs
in sponge-like crystals with irregular outlines.

Specimen CAS—40 is fossiliferous siltstone and was obtained from a
large boulder in the first main canyon southwest of the village on Maria
Madre Island. The siltstone is fine-grained and contains rounded and
angular grains of quartz and shell fragments, and fossil foraminifera in a
matrix consisting of calcite and chaleedonic siliea. C. C. Chureh (written

Vou. XXXII] CHESTERMAN: EASTERN PACIFIC PETROGRAPHY 359

communication, January, 1963) kindly examined the siltstone and identified
the following genera:

Peneroplis sp.

Bolwina sp.

Bulimina sp. similar to B. ovata

Frondicularva sp.

Globigerina sp.

These fossils indicate late Tertiary (possibly Plocene) age for the

siltstone.

WuitE Friars ISLANDS (Morro DE PAPANOA)

During the course of later excursions off the west coast of Mexico by the
California Academy of Sciences, several rock specimens were collected by
Dr. Leo G. Hertlein from a group of small islands which lie some 5 to 6
miles south of Zihuatenejo, State of Guerrero. These islands are referred
to as the White Friars and are said to resemble members of that monastic
order in a kneeling position.

‘ rete a r
ag OTE

~~ ee

+ £

s Islands. Veins of white meta-

variscite in collophane. Plain light. x 30.

360 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4111 SER

Most of the rock specimens collected by Dr. Hertlein are highly phos-
phatized voleanie rock, but one (CAS-42) is reasonably unaltered and is
identified as hornblende andesite. The andesite is porphyritic and contains
plagioclase phenocrysts set in a granular matrix of feldspar, hornblende,
voleanic glass, and minor quartz. The plagioclase phenocrysts are sub-
hedral, twinned, and zoned; a few are bent. Those phenocrysts that show
the best zoning are euhedral in shape. Many of the phenoerystice plagioclases
are of a pale, pinkish-brown color. These colored feldspar erystals have a
composition in the range of Ans) to An,;. The remaining, non-colored
plagioclase phenocrysts are in a composition range of An,; to An;,. The
phenocrystic plagioclase constitutes about 45 per cent of the rock. The
eroundmass plagioclase is in subhedral and anhedral grains which show
twinning, but lack zoning. They are relatively unaltered, range in com-
position from An,, to Any, and constitute about 40 per cent of the rock.

Hornblende occurs as pale-green, weakly pleochroie, prismatic and
skeletal erystals. Biotite is in irregular pleochroie erystals. The hornb!ende
and biotite constitute less than 10 per cent of the rock. The quartz present
in this rock is searee, and oceurs in rounded grains that show wavy ex-
tinetion and contain tiny prismatic crystals of zirion. Both apatite and
magnetite are present. The apatite occurs as needle-like inclusions in the
plagioclase and the magnetite usually in close proximity to the biotite and
hornblende. A dark, brown-colored voleanic glass occurs in irregular areas
seattered irregularly throughout the rock.

PHOSPHATE Rock. Several samples of phosphate rock were available
for study and they appear to be the same in all respects. Specimen no.
CAS—44 is the largest and offers the greatest opportunities for detailed
petrologic and mineralogic study. It is relatively smooth, dense, and light
brownish-gray to beige in color. Cut surfaces show irregularly distributed
open voids as well as a well developed colloform banding which in some cases
conforms to the outline of the void. Under the microscope the colloform
banding shows up fairly well and one ean see that the rock consists prin-
cipally of two distinct materials: (1) a pale-buff to brown-colored, weakly
birefringent isotropic mineral whose refractive index and structure indi-
cate collophane and (2) a erystalline, colorless mineral which has moderate
to strong birefringence, moderate axial angle, optically positive and an ex-
tinetion of Z’/)\C = 27°. This latter mineral is metavariseite in which a
small amount of iron isomorphously replaces some of the aluminum. The
metavariscite occurs as small prismatic erystals and in erusts that show
a radial-fibrous structure.

The phosphate rock from the White Friars resembles in many respects
the phosphate rock found on Malpelo Island, yet they differ somewhat in

their mineralogical content. The phosphate rock on Malpelo consists largely

Vom. XXXII] CHESTERMAN: EASTERN PACIFIC PETROGRAPHY 361

of phosphosiderite and strengite (MeConnell, 1943, p. 713) whereas that on
the White Friars Islands is collophane and metavariscite.

BIBLIOGRAPHY

BANFIELD, ARMINE FREDERICK; CHARLES HENRY BEHRE, JR.; and Davin Sr. CLAIR

1956. Geology of Isabela (Albemarle) Island, Archipielago de Colon (Gala-
pagos). Bulletin of the Geological Society of America, vol. 67, no. 2,
pp. 215-234, pls. 1-4, figs. 1-4 in text, February.

CnuuBsB, JOUN LAWRENCE

1933. Geology of the Galapagos, Cocos and Easter Islands. Bernice P. Bishop
Museum, Bulletin 110, pp. 1-44, pls. 1-5, figs. 1-8 in text.

DARWIN, CHARLES

1869. Geological Observations on the Volcanic Islands and parts of South
America visited during the Voyage of H.M.S. Beagle. Third edition.
D. Appleton and Company, New York, pp. I-XIII, 1-648, pls. 1-5 (fold),
2 maps.

EHRENBERG, CHRISTIAN GOTTFRIED

1853. Das Mikroskopische Leben der Galapagos-Inseln und iiber die organische
Mischung der dortigen vulkanischen Gebirgsarten, besonders der Pala-
gonits. Monatsberichte der Berliner Akademie der Wissenschaften,
1853, pp. 178-182, 1 Tafel (table).

1854. Die Galapagos-Inseln. Mikrogeologie (Verlag von Leopold Voss: Leip-
zig), pages XXVII, 346-349, Atlas, Tafel XX XVIII, fig. XVIII.

Foose, Ricuarp M.

1962. Reconnaissance Geology of Maria Cleopha Island, Tres Marias Islands,
Mexico. Bulletin of the American Association of Petroleum Geologists,
volume 46, number 9, pp. 1740-1745, figs. 1-6, September.

HANNA, G DALLAS

1925. Expedition to Guadalupe Island, Mexico, in 1922. Proceedings of the
California Academy of Sciences, fourth ser., vol. 14, no. 12, pp. 217-
275, pls. 15-19, figs. 1-2 in text, September 5.

1926. Expedition to the Revillagigedo Islands, Mexico, 1925. General Report.
Proceedings of the California Academy of Sciences, fourth ser., vol.
15, no. 1, pp. 1-113, pls. 1-10, figs. 1-7 in text, March 30.

HER?tTLEIN, LEO GreorGE, and WILLIAM K. EMERSON

1959. Results of the Puritan-American Museum of Natural History Expedition
to Western Mexico. 5. Pliocene and Pleistocene megafossils from the
Tres Marias Islands. American Museum of Natural History, Novitates
no. 1940, pp. 1-15, figs. 1-5 in text, June 5.

McCoNNELL, DUNCAN

1943. Phosphatization at Malpelo Island, Colombia. Bulletin of the Geological
Society of America, vol. 54, no. 5, pp. 707-716, pls, 1-2, May 1.

362 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4v1r Ser.

MurpuHy, Rospert CUSHMAN
1941. The Askoy Expedition of the American Museum of Natural History in
the Eastern Tropical Pacific. Science, new ser., vol. 94, pp. 57—58,
July 18.

OCHSNER, WASHINGTON HENRY
1906. Geology of the Galapago Islands. Unpublished notes in the California
Academy of Sciences.
RICHARDSON, CONSTANCE
1933. Petrology of the Galapagos Islands. Bernice P. Bishop Museum, Bulletin
110, pp. 45-64.
SAPPER, KARL
1917. Katalog der geschichtlichen Vulkanausbruche. Schrift 27 Wissengesell-
schaft in Strassburg (K. Tuber), pp. 1-358.
SLEVIN, JOSEPH RICHARD

1959. The Galapagos Islands. A History of their Exploration. Occasional
Papers of the California Academy of Sciences, no. XXV, pp. 1-150,
figs. 1-31 in text, December 22.

VAN WEsT, OLAF

1959. Geology of the San Benito Islands and the southwest part of Cedros
Island, Baja California, Mexico. Unpublished thesis, Pomona College.
Claremont, California, pp. 1-28.

PROCEEDINGS

OF THE
CALIFORNIA ACADEMY OF SCIENCES
FOURTH SERIES

G Dallas Hanna Anniversary Volume

Volume XXXII, No. 12, pp. 363-375; 2 tables July 28, 1965

STUDIES ON THE ATLANTIC AMERICAN
PIPEFISHES WITH DESCRIPTIONS OF
NEW SPECIES
By

Earl S. Herald

Steinhart Aquarium, California Academy of Sciences

The Atlantic American Syngnathid fauna seems to be in the process of
evolving many new forms. Some of the divergencies from the parent stock are
as yet so slight that a sizable collection of comparative material is often re-
quired to distinguish these trends. In part this has been responsible for some of
the confusion in the classification within this group. The material herein pre-
sented is an attempt to clarify some of the problems.

The descriptions of four new forms in this paper as well as the resurrection
of two old names brings the known Atlantic American pipefishes to a total of 29
forms representing 25 species. In the following list, the common names with
asterisks appear for the first time. Those without asterisks are from the Ameri-
~ can Fisheries Society Checklist of Common and Scientific Names (second edi-
tion, 1960):

Opossum Pipefish Oostethus lineatus (Valenciennes, 1856)
Whitenose Pipefish Corythoichthys albirostris Heckel, 1853
Crested Pipefish Corythoichthys brachycephalus (Poey, 1867)
*Deepwater Pipefish Corythoichthys profundus, new species
Northern Pipefish Syngnathus fuscus fuscus Storer, 1839

*Relict Northern Pipefish Syngnathus fuscus affinis Gunther, 1870

[ 363 ]

364 CALIFORNIA ACADEMY OF SCIENCES

Gulf Pipefish

Sargassum Pipefish

*“Chesapeake Dusky Pipefish

Dusky Pipefish

*Key Dusky Pipefish

*Bermuda Dusky Pipefish

*Caribbean Pipefish
*Pivothead Pipefish
Chain Pipefish
Bull Pipefish
*Southern Pipefish
Shortfin Pipefish
*Ocellated Pipefish
Pugnose Pipefish
*Dwarf Pipefish
*Freshwater Pipefish

*Finless Pipefish
*Lost Pipefish
*Deep-bellied Pipefish

Banded Pipefish
Fringed Pipefish

Insular Pipefish

*Seahorse Pipefish

[Proc. 4TH SER.

Syngnathus scovelli (Evermann and Kendall,
1895)

Syngnathus pelagicus Linnaeus, 1758

Syngnathus floridae hubbsi, new subspecies

Syngnathus floridae floridae (Jordan and Gil-
bert, 1884)

Syngnathus floridae mckayi (Swain and Meek,
1884)

Syngnathus floridae nesiotes Herald, 1942

Syngnathus rousseau Kaup, 1856

Syngnathus fistulatus Peters, 1868

Syngnathus louisianae Gunther, 1870

- Syngnathus springeri Herald, 1942

Syngnathus folletti Herald, 1942

Syngnathus elucens Poey, 1867

Syngnathus randalli, new species

Syngnathus dunckeri Metzelaar, 1919

Syngnathus hildebrandi, new species

Pseudophallus mindii (Meek and Hildebrand,
1923)

Penetopteryx nanus (Rosen, 1911)

Ichthyocampus pawneei Herald, 1950

Leptonotus blainvilleanus (Eydoux and Gervais,
1837)

Micrognathus vittatus (Kaup, 1856)

Micrognathus (Anarchopterus) crinigerus (Bean
and Dressell, 1884)

Micrognathus (Anarchopterus) crinitus (Jenyns,
1842)

Amphelikturus dendriticus (Barbour, 1906)

Since the primary purpose of this paper is to describe new forms that will be

treated in more detail in a forthcoming volume of the Sears Foundation, Fishes
of the Western North Atlantic, a key will not be published herein. However,
during this interim period, a mimeographed key, modified from Herald, 1942,
will be available upon request from the author.

Deepwater Pipefish, Corythoichthys profundus Herald, new species.

Ho.otyprE. USNM 198096, male 198.5 mm. standard length (204 mm. total
length); Silver Bay Station 3466: About 90 miles slightly south of due east
from Melbourne, Florida (Lat. 27°56’ N.; Long. 79°05’ W.); 100 fathoms; 6-
foot tumbler dredge; October 25, 1961.

Dracnosis. Dorsal fin rays 27; pectoral 14-14; anal 3; caudal 10; dorsal

Vou. XXXII] HERALD: ATLANTIC AMERICAN PIPEFISHES 365

fin covering 2 trunk ring and 5" tail rings; trunk rings 18; tail rings 38; head
21.8 mm.; snout 12.6 mm.; dorsal fin base 16.0 mm.; pectoral base 2.2 mm.;
pectoral length 3.6 mm.; head-in-standard length 9.12; snout-in-head 1.73;
dorsal fin-base-in head 1.36; pectoral base-in-pectoral length 1.63; brood pouch
covering 20 tail rings; eggs in anterior section only, starting at second tail ring
and extending for an additional 9 rings; eggs arranged 1 layer in thickness and
maximum of 6 rows in width with an approximate total of 131; brood pouch
closure of open type with flaps not meeting in center (see Herald, 1959); brood
pouch protecting plates slightly developed. Body ridges of Corythoichthys type.

DescripTIon. All ridges of head and body strongly accentuated showing
minute crenulations when viewed under low-power microscope. Median snout
ridge extends over posterior half of snout ending between eyes. Anterior orbital
projection accentuated; superior orbital ridge begins above center of eye and
extends posteriorly for about one eye diameter. Opercular ridge extends over
anterior fourth of opercle; single supra-opercular ridge is half length of opercle.
Median head crest trilobed with small additional projection just behind eyes.
Pectoral cover plate with both superior and inferior ridges. Color overall light
tan with small flecks of black pigment when viewed under scope. Small blackish
pigment spots in web between individual rays of caudal fin. Dorsal fin with
pigment spots along base of rays and extending out for short distance on indi-
vidual rays. Pectoral and anal fins colorless.

Discussion. The combination of characters: (a) ring counts, (b) dorsal
count, (c) position of dorsal fin, (d) head-in-standard length, and (e) snout-in-
head values serve to segregate Corythoichthys profundus from all other Atlantic
American pipefishes. Corythoichthys profundus comes from a greater depth
than that at which other pipefishes are found (100 fathoms: 600 feet).

The head of Corythoichthys profundus has the typical appearance of the
Indo-Pacific group of Corythoichthys, all of which have long snouts but lack
protective plates along the sides of the brood pouch. Corythoichthys profundus
does have pouch protecting plates, although only slightly developed. This has
been one of the main features used to separate the Atlantic species (subgenus
Corythoichthys) and the Pacific species (subgenus Bhanotichthys), so that now
the validity of the subgenera becomes questionable.

Hasitat DATA. The detailed data carefully recorded in the log of Silver Bay
(Station 3466) show that the type of Corythoichthys profundus was taken in a
55-minute drag over coral and sand bottom. During this time the 6-foot tumbler
dredge traveled a distance of about 4 knots all on a flat bottom of 100 fathom
depth. Surface water temperature was 78° F.; bottom temperature was not re-
corded; and air temperature was 75° F. Fifteen pounds of material was found
in the dredge, of which two pounds was inert calcareous bottom material and 13
pounds consisted of animals of 17 kinds: 5 crinoids, 15 sand dollars, 100 miscel-

366 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser.

laneous crabs, 50 hermit crabs, 5 Sconsia sp., 2 Fusinus sp., 25 Murex callati,
1 cat-shark egg case, 7 Kathetostoma cubana, 1 wrasse, 2 Antigonia capros, 3
Trichopsetta ventralis, 3 Achirus inscriptus, 10 Syacium sp., 15 other flatfish,
1 Prionodes phoebe, and 1 pipefish (the type described above).

Dwarf Pipefish, Syngnathus hildebrandi Herald, new species.

Syngnathus elucens Lonctry and Hitpesranp, 1941, Carnegie Inst. Wash., Tortugas Lab.,
vol. 34, pp. 61 (holotype doubtfully identified as S. elucens).

Hototype. USNM 117251, female 86.5 mm. standard length; Tortugas,
Florida; W. H. Longley.

Paratypes. USNM 73235, female 72.5 mm.; Fish Hawk Station 7165;
Pepperfish Key, SW. Florida (Lat. 29°13.25’ N.; Long. 83°32.5’ W.); 7%
fathoms, rocky bottom; oyster dredge; November 21, 1901.

USNM 73239, two females 61.5 and 52.5 mm.; Fish Hawk Station 7216:
St. Martin’s Reef, West Florida (Lat. 28°26.5’ N.; Long. 83°08’ W.); 10
fathoms, sandy—grassy bottom; oyster dredge; January 15, 1902.

USNM 109826, female approximately 67 mm. (tail broken); Fish Hawk
Station 7217: St. Martin’s Reef, West Florida (Lat. 28°27’ N.; Long. 83°13’
W.); 11 fathoms, rocky—sandy bottom; oyster dredge; January 15, 1902.

USNM 134312, female 59 mm.; Fish Hawk Station 7220: St. Martin’s
Reef, West Florida (Lat. 28°34’30” N.; Long. 83°15’45” W.); 7’4 fathoms;
oyster dredge; January 15, 1902.

Diacnosis. Dorsal fin rays 19-21; pectoral 10-12; anal 2; caudal 10;
dorsal fin covering 0 to '2 trunk ring and 41% to 5 tail rings; trunk rings 17; tail
rings 33-34; head-in-standard length 9.5—10.7; snout-in-head 2.53—2.87; dorsal
base-in-head 0.97—1.22; length of brood pouch unknown; color light tan; cirri
sometimes present on head and body; very similar in appearance to Syngnathus
dunckeri. Body ridges typical for genus Syngnathus.

Discussion. Syngnathus hildebrandi has a small anal fin which instantly
serves to segregate it from Syngnathus dunckeri, with which it would otherwise
be easily confused. In ring counts it is similar to Syngnathus elucens, but its
much shorter head will separate it from that species (head-in-standard length
9.5—10.7 for S. hildebrandi versus 7.08-8.13 for S. elucens). Syngnathus hilde-
brandi also has fewer dorsal fin rays than other Atlantic American pipefishes
with the same lateral ridge pattern. It may be noted that there are four addi-
tional species with a different lateral ridge pattern, i.e., three of Micrognathus
and one of Amphelikturus, that also have low dorsal fin counts in the 16-22
range.

NAMED hildebrandi in honor of the late Dr. Samuel F. Hildebrand who rec-
ognized that the holotype was somewhat different from other previously known
American pipefishes.

VoL. XXXII] HERALD; ATLANTIC AMERICAN PIPEFISHES 367

THE FLORIDAE COMPLEX

Along the Atlantic coast of North America from Chesapeake Bay to Panama
there lives in many shallow areas a grass-inhabiting pipefish which has been
variously called Syngnathus floridae and Syngnathus mckayi. This population
complex presents an interesting study in that it must have had a continuous
shoreline distribution during earlier geological periods, although the present dis-
tribution is discontinuous. In the Caribbean the insular member of this complex
is Syngnathus rousseau.

One population is now restricted to the region between Chesapeake Bay and
Seabrooks Beach, South Carolina. This isolated form has previously been
known as Syngnathus floridae, but herein is given a new name (Syngnathus
floridae hubbsi). From South Carolina to Biscayne Bay on the southeast Flor-
ida coast there is a break in distribution in which no member of the floridae
complex is known to occur. At Biscayne Bay the floridae-type pipefishes are
very different from those of the Chesapeake-South Carolina area. This Bis-
cayne Bay member of the complex (Syngnathus floridae mckayi) is found from
the Miami area south along the keys to Tortugas. Another population (Syng-
nathus f. floridae) is found along the west coast of Florida from Cape Sable
(SW. Florida) to Corpus Christi, Texas. From the coastline distances involved,
it would be suspected that the Chesapeake S. f. Aubbsi and the southern Florida
S. f. mckayi would be most closely related. Surprisingly, this is not the case, for
it is the west Florida to Texas component that shows the closest relationship
with S. f. hubbsi. This similarity is most striking when pipefishes from the
Pensacola to Corpus Christi area are compared with the Chesapeake group. It
is suspected that these last two populations were continuous at an earlier time
by means of a wide seaway or canal which geologists tell us formerly existed
across the north-central section of Florida.

It may be noted that Ginsburg (1937) has shown a similar relationship for
the common Atlantic American seahorse, Hippocampus erectus (sometimes
called H. hudsonius), in which the populations north and west of Florida are
considered to be the same whereas those found in southern Florida and Cuba are
assigned to a separate subspecies.

Between southern Texas and Panama only a few specimens of S. floridae are
known, undoubtedly because of limited collecting. At Panama records indicate
a sizable population which meristically is more closely related to the Florida
keys’ S. f. mckayi than it is to the adjacent S. f. floridae. This may be a natural
result either of temperature or of ocean current patterns moving through the
Cayman Sea northeastward through the Straits of Florida. When more material
is available it is probable that the Panamanian population can stand as a sepa-
rate subspecies, and this is probably also true of the Syngnathus floridae group
on the west coast of Florida. In fact, the writer at one time had these written up
in this manner, but has withheld such until the case is stronger.

368 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47H Ser.

In the Caribbean, the inshore weed habitat of Syngnathus floridae is taken
over by the closely related but less abundant Syngnathus rousseau. Specimens
of the former have not been collected in grassy habitats within the insular area
although the two species have been taken together in Panama. Even in this
locality it is not easy to separate them since most of the meristic characters are
similar with the exception of the tail ring count (32—34 for S. rousseau and 35—
37 for S. floridae). Were it not for the fact that they have been collected at the
same place, they would otherwise have to be considered as mutual subspecies
rather than full species. Unfortunately this tail ring separation breaks down in
the Caribbean so that the many small floating pipefishes found at sea are often
impossible to assign exactly to one species or the other.

Although there is a large S. floridae population of dwarf forms in Bermuda
(S. f. nesiotes), only four S. floridae specimens have been collected in the inter-
vening Bahamas. Indications are, however, that some grassy shallow areas in
the Bahamas do support a modest S. floridae population, for example, Little
Bahama Bank. Detailed analyses of the various S. floridae groups will be pre-
sented later; for the present the Chesapeake Bay population will be recognized
under a new name, described as follows:

Chesapeake Dusky Pipefish, Syngnathus floridae hubbsi Herald, new sub-
species.

HototyPr. USNM 91321, gravid male 177 mm. standard length; Lower
York River, Virginia; Wm. C. Schroeder, July 8-12, 1921.

ParATyPES. USNM 133053, 33 specimens including 6 males and 27 females
and subadults (44-172 mm.) ; same data as holotype.

OTHER SPECIMENS EXAMINED. (Total 133, excluding types; mostly in
USNM); Maryland: Crisfield (4); Plum Point (1); Virginia: Cape Charles
City (33); Lewisetta (2); Lower Rappahannock River (49); Buckroe Beach
(5); Mouth of Hampton Creek (4); Norfolk (3); Cape Henry (1); North
Carolina: Beaufort (28); Cape Lookout (2); Wilmington (1).

Diacnosis. Dorsal fin rays 27-31; pectoral 14-15; anal 3; caudal 10;
dorsal fin covering ’2—-2 trunk rings and 5—6% tail rings, usually 1 + 6; trunk
rings 16-18, usually 17-18; tail rings 31-34; head-in-standard length 5.2—-6.8,
usually 5.4—5.9; dorsal-in-head 1.35—1.85, usually 1.45—1.7; snout-in-head 1.6—
1.88; brood pouch covering 18-21 tail rings; brood pouch closure of inverted
type, 7.e., contacting brood pouch folds turning inwardly dorsad; eggs averaging
0.9 mm. in diameter and arranged 1 or 2 layers in depth and 3 or 4 rows in
width on each side of pouch; protecting plates of pouch moderately developed;
largest female 206 mm.; largest male 180 mm.; smallest sexually mature male
103 mm.; adult females slightly V-bellied.

VoL. XXXII] HERALD: ATLANTIC AMERICAN PIPEFISHES 369

DISCUSSION AND COMPARISON. Syngnathus floridae hubbsi has been recog-
nized since 1882 when Jordan and Gilbert stated in their original description of
the Pensacola types of Syngnathus floridae: ‘In our paper on the Fishes of
Beaufort Harbor (Proc. U. S. Nat. Mus., 1878, 368), we have recorded a
‘Siphostoma fuscum’ from that locality. The specimens referred to under that
name belong to Siphostoma louisianae chiefly; among them are examples of the
present species.”

Although at the time of his 1942 paper the writer considered the Chesapeake
Bay population as distinct from that existing in Florida, Texas, Panama, and
Bermuda, it was nevertheless felt that there was reason to suspect that some of
the type material of Syngnathus floridae from Pensacola had been mixed with
males of the Chesapeake form. Consequently the use of names in the 1942
publication is different from that in this paper. The change is due to the addi-
tional material which has become available since that time and to the conclusion
that the Chesapeake Bay form, although recognized, had never actually been
described.

In the region in which Syngnathus floridae hubbsi occurs, it can be segre-
gated from the other two species which are present (S. fuscus and S. louisianae )
by the head-in-standard length value (S. f. Aubbsi 5.4—5.9 vs 6.3-9.5 for oth-
ers); also by the number of rays in the dorsal fin (S. f. Aubbsi 28-31 vs 32-41
for others) ; and by the position of the dorsal fin on trunk and tail (S. f. hubbsi
usually 1 + 6 vs 3-5 + 4-6 for others). From the members of the floridae
complex, the mature males of S. f. kubbsi can be separated without difficulty on
the number of rings covered by the brood pouch (S. f. Awbbsi 18-21 vs 12-17
for others). Although geographically the females of S. f. hubbsi can be segre-
gated from the other subspecies of S. floridae on the basis of locality data, there
are nevertheless many females of S. f. Awbbsi which cannot be segregated from
the other subspecies of S. floridae on meristic characters. This is especially true
of the type subspecies, S. f. floridae. One who is thoroughly familiar with the
two subspecies can sometimes segregate typical females by means of their gen-
eral appearance, but as yet there are no objective criteria which can be applied
despite a considerable amount of time that has been expended on the problem.

Namep hubbsi in honor of Dr. Carl L. Hubbs whose helpful interest in the
S. floridae complex has been of considerable aid.

LECTOTYPES
Since the original descriptions of two of the S. floridae complex were based
on series material without designation of holotype, it is advisable to select lecto-
types as follows:
Syngnathus floridae floridae (Jordan and Gilbert, 1884). Siphostoma flori-
dae Jordan and Gilbert, 1884, Proc. U. S. Nat. Mus., vol. 5, p. 263.
LecToTyPE (herein designated). MCZ 35958, originally from USNM

370 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

30826: 148 mm. gravid male with brood pouch covering 17% tail rings; Pensa-
cola, Florida; Silas Stearns and David Starr Jordan.

SyntyPEsS. USNM 30826, six females 121-155 mm.; same data as lectotype
above.

Syngnathus floridae mckayi (Swain and Meek, 1884). Siphostoma mckayi
Swain and Meek, 1884, Proc. U. S. Nat. Mus., vol. 7, p. 239.

LEecTOTYPE (herein designated). SNHM 1894; 138.5 mm. gravid male with
brood pouch covering 13 tail rings; Key West, Florida; David Starr Jordan;
December, 1884.

SyntypEs. SNHM 1894, three females, 170, 178, and 211 mm.; same data
as lectotype; USNM 34989, two females, 158 and 163 mm.; same data as
lectotype.

Relict Northern Pipefish, Syngnathus fuscus affinis Giinther, 1870.

Syngnathus affinis GUNTHER, 1870, Cat. Fishes, Brit. Mus., vol. 8, p. 163; holotype 145.5
mm. male from “Louisiana”; Brit. Mus. 1854.7.32.

OTHER MATERIAL EXAMINED. USNM 132675; 174 mm. male and 117 mm.
female from Corpus Christi Pass, Oct. 14, 1926, J. C. Pierson; CNHM 40309;
two females, 206 mm. and 218 mm., from Corpus Christi, C. T. Reed.

Dracnosis. Dorsal fin rays 36-39; dorsal covering 4—5 trunk rings and
4—5 tail rings; pectoral 14-16; anal 3; caudal 10; trunk rings 18-19; tail rings
33-36; head-in-standard length 8.55—9.8:; snout-in-head 2.06—2.36; females flat-
bellied as is characteristic of Syngnathus fuscus in the southern portion of the
range (South Carolina to Florida). Brood pouch covering 12% to 15 tail rings.

Discussion. Since the time of the original description of Syngnathus af-
finis, no specimens have been known which could definitely be assigned to this
species. At the time of his 1942 paper, the writer thought that the holotype of
S. affinis was a specimen of S. fuscus from an erroneous locality. However the
four specimens indicated above demonstrate that there is a relict S. fuscus
population in the Corpus Christi area. Although the characters of these speci-
mens are slightly different from those of the type of S. affinis, their variation is
not sufficiently great to prohibit their being assigned to subspecific status under
that name. The holotype of Syngnathus affinis was re-examined for the writer
by the late Mr. V. Tchernavin.

Further study and more material will be needed to determine the relationship
of this relict flat-bellied population to the similar flat-bellied group of S. fuscus

from South Carolina to Florida.
Bull Pipefish, Syngnathus springeri Herald, 1942

At the time the writer described this short-nosed, long-trunked cousin of
Syngnathus louisianae, only four specimens were known. Since then, largely

VoL. XXXII] HERALD: ATLANTIC AMERICAN PIPEFISHES 371

because of the extensive exploratory work of the U. S. Fish and Wildlife Service
ships, Silver Bay, Oregon, Pelican, Combat, and Gill, a representative group of
specimens has become available. It now appears that Syngnathus springeri is
truly different in habitat from the other American pipefishes. Although moder-
ately abundant, it generally does not occur in the intertidal and other very shal-
low water, but prefers slightly deeper water from 10 to 70 fathoms, and it may
occur as much as 100 miles offshore. In the young stages it may be a floater and
be picked up by dipnet under the night light (eight localities) or eaten by tuna
(one locality). The species apparently has a higher reproductive potential than
most other pipefishes: one male, 274 mm., had 1,390 eggs in the 16'2-ring brood
pouch. The range, based on the 65 specimens and 48 localities examined either
directly by the writer or from data sent to him, now extends from Cape Lookout
and Morehead City, North Carolina, to Pensacola, Florida. It occurs in the
Bahamas (four localities) where its relative, Syngnathus louisianae, does not
occur; but it is missing at Bermuda where the latter is found. It has been taken
in the same offshore haul with Syngnathus louisianae (30 miles south of Pensa-
cola, 70 fathoms), but this latter species, by contrast, is found as well in very
shallow water. These two species are the largest in the Atlantic American Syng-
nathid fauna, with the short-snouted S. springeri reaching 355 mm., and the
long-snouted S. louisianae, 326 mm. Syngnathus springeri also has the distinc-
tion of having the largest number of trunk rings—23-—24, usually 23, as con-
trasted with the next highest number for S. lowisianae—19-21, usually 20.

Insular Pipefish, Micrognathus (Anarchopterus) crinitus (Jenyns, 1842).

Pipefishes previously assigned to Micrognathus (Anarchopterus) crinigerus
have upon re-analysis been found to represent two species whose appearances
are identical, but whose meristic characters allow quick separation on the follow-
ing data:

No. Dorsal Pectoral Trunk Tail
Specimens Rays Rays Rings Rings

M. crinitus 28 18-20 10-11 17-18 32-35
M. crinigerus 86 16-18 8-9 14-16 37-39

Micrognathus crinitus has been taken at Tortugas, Florida, and between
- Vieques and Culebra islands, Puerto Rico; other records are from the Bahamas:
Royal Island, New Providence Island, and Little Bahama Bank. Records for
five additional Bahaman localities are contained in the collections of the Acad-
emy of Natural Sciences of Philadelphia, and will be reported by Dr. James
Bohlke. Surprisingly, only one specimen of Micrognathus crinigerus has been
taken in the Bahamas, and yet it is not an uncommon species along many sec-
tions of the Florida coastline from Biscayne Bay southward as well as on the
entire west coast of Florida. On the other hand, Micrognathus crinitus is strictly

SUP CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser.

insular in habitat, at least insofar as the Caribbean is concerned. A total of 28
specimens including the type are known (January, 1965).

The late Mr. V. Tchernavin of the British Museum re-examined the type of
Syngnathus crinitus for the author, and in spite of the fact that it was collected
at a considerable distance from the Caribbean, 7.e., Bahia Blanca, northern
Patagonia, its characters are sufficiently close to those of the North American
specimens that the latter can be assigned to M. crinitus.

The subgenus Anarchopterus of Micrognathus has only the two species men-
tioned above. This subgenus is characterized by smooth body ridges and ab-
sence of the anal fin. By contrast, the subgenus Micrognathus is characterized
by sharp body ridges, and the presence of the anal fin. It has one Atlantic
species, M. vittatus, and eight species in the Pacific.

Banded Pipefish, Micrognathus vittatus (Kaup, 1856) versus

Micrognathus ensenadae (Silvester, 1916).

There are two color variations of the banded pipefish. If these were to be
recognized as separate species, the above two names would be applicable. The
two forms are not separable on the basis of meristic data; however, some of the
writer’s colleagues feel that they are separable on ecological data and should
thus be accorded specific rank. Hence a re-examination of the approximately 42
known specimens has been made. These specimens came from 25 localities
ranging from Bermuda and the Bahamas southward through the Florida keys to
Cuba, Jamaica, Haiti, Puerto Rico, Arcas Cay (75 miles off Campeche), Vene-
zuela, and Brazil. Obviously, with this distribution the banded pipefish is pri-
marily insular in habitat. On the basis of color, the specimens available separate
into a group of 31 with the vittatus pattern and 11 with the ensenadae pattern.
It is interesting to note that the esenadae pattern is duplicated in the smaller
but not closely related Micrognathus nitidus of the Pacific.

The typical vittatus coloration is usually rather dark, variable, and nonde-
script; there may be 3-5 dark bands around the trunk and 8—9 around the tail.
These bands are more distinct on the upper surface, often fading ventrally. By
contrast, the ensenadae pattern is quite spectacular with a series of rich brown
bands around the head and body. These are about a ring or more in width. The
head has a variable mottled pattern with the first distinct band just ahead of
the pectoral fins, followed by 6 bands on the trunk and 13-16 on the tail. Be-
tween the brown rings is a much lighter color ranging from yellow to off-white.
Although the type of Corythoichthys ensenadae is lost, Silvester did publish an
excellent color plate two years after his original description (Carnegie Inst.
Wash., Publ. 252, Dept. Mar. Biol., vol. 12, p. 21, fig. 3, 1918). The type of
Corythoichthys vittatus Kaup is in good condition at the Paris Museum (no.
602), where it was examined by the writer.

VoL. XXXII] HERALD: ATLANTIC AMERICAN PIPEFISHES 373

The range in size of the 11 M. ensenadae specimens is from 54 mm. to 116
mm., whereas that of the 31 M. vittatus specimens is from 52 mm. to 140 mm.
Unfortunately, there are no really small specimens, hence the size at which the
patterns first appear cannot be determined. Not one of the patterns is truly
intermediate between M. vittatus and M. ensenadae although there is an occa-
sional indication that intermediates might occur in some individuals of MW. vit-
tatus. Micrognathus vittatus—ensenadae appears to be a solitary species, as
demonstrated by the collection records, most of which consist of single individu-
als. In four cases, two specimens were taken at the same time, and from Ber-
muda there is one collection of five individuals which might, however, be an
accumulation of specimens. From Albrolhos Islands, Brazil, there are two
specimens, both 92.5 mm., one of which exhibits the MW. ensenadae pattern and
the other the MW. vittatus pattern. This is the only time that both color patterns
were taken at the same site, apparently at the same time. In the Bahamas the
two color patterns have been taken at closely adjacent localities.

My colleagues, Drs. James Bohlke and Richard Robins, have postulated
that the 7. ensenadae type is always found in the area of sea fans and gorgonian
corals, whereas the M. vittatus type is characteristically found in grassy areas.

In summation, Micrognathus vittatus and Micrognathus ensenadae must be
considered as one from the standpoint of anatomy, but from the standpoint of
coloration they are definitely two distinct varieties which as yet have no inter-
grades. More material will be required to make the final determination whether
we are dealing with two full species, subspecies or color variants.

While this manuscript was in page proof, a strange new Venezuelan pipefish
was received from Dr. John Randall. Fortunately it was possible to add its
description to this paper.

Ocellated Pipefish, Syngnathus randalli, new species.

HoLotyePe. USNM 198903; female 93.5 mm. standard length (95.3 mm.
total length) ; South side of Isla Venados, about 28 kilometers WSW of Cumana,
Venezuela; mangroves sparse on rocky shore; bottom, Porites furcata; depth 1
to 2 feet; visibility 20 ft.; temperature 22.5° C. (72.5° F.); ichthyocide sta-
tion; January 27, 1965; John E. Randall.

Dracnosis. Dorsal fin rays 24; pectoral 13-14; anal ABSENT; caudal
10; dorsal fin covering ’% trunk ring and 4’ tail rings; trunk rings 17; tail
rings 32; snout 3.9 mm.; head 9.4 mm.; trunk 31 mm.; tail 53.1 mm.; dorsal
fin base 8.3 mm.; pectoral base 1.3 mm.; pectoral length 1.5 mm.; head-in-
standard length 9.84; snout-in-head 2.41; head-in-dorsal fin base 0.88; brood
pouch details unknown; lateral trunk ridge of typical Syngnathus type, 1.e.,
interrupted at anal ring and not continued with lateral tail ridge; belly slightly

374 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

V-shaped. Color brown with regular series of round to oval spots on trunk
giving way to band markings on tail.

DescripTION: Median snout ridge smooth, slightly raised, beginning at
interorbital area and extending forward, three-quarters length of snout. Su-
perior ocular ridge extends posteriorly from center top of orbit for about one
eye diameter. Supraopercular ridge present, in posterior position; midopercular
ridge short, equal to one-quarter length of opercle. Pectoral cover plate with
single ridge in inferior position. Nuchal ridge slightly raised, trilobed. Body
ridges distinct, rounded, not sharp. Intermediate plates between rings fairly
large, about equal in plate width to distance between plates.

Color pattern quite startling and from Dr. Randall’s Field Notes: “Color
when fresh, coarse network of greenish brown, enclosing dark-edged spots of
light yellowish gray; posteriorly the ground color is light yellowish gray and
the reticulatum gives way to interconnected spots along side and ultimately to
discrete bars; upper part of snout light greenish yellow, the lower part reddish
brown; cheek area olive brown with [four] dark edged light yellow lines (most
of which are diagonal); dorsal fin pale; caudal fin dusky yellow; iris red with
spoke-like lines of pale yellow.”” Some appear to continue outward onto throat
(five), behind eye (one), and above eye (one).

With preservation, greens and yellows gradually disappear leaving dark
brown ground color punctuated with regularly arranged series of light spots,
each bordered by a thin dark line. All seven intermedial plates between each
trunk ring covered by individual dark-ringed light spots; one on upper surface,
two on each side surface, and two on belly. Center of each trunk ring with
additional six spots: one at lateral trunk ridge; one at upper trunk ridge ex-
tending onto dorsal surface of trunk, and one at lower trunk ridge extending
onto abdomen. This middle series of spots begins coalescing at anal fin area,
giving rise on tail to complete dark bands between each ring.

OTHER SPECIMENS. USNM 164831, 2 juveniles, 31.3 and 32 mm. standard
length; Haiti; William Beebe. These are referred questionably to Syngnathus
randalli, but are not paratyped. Although both lack the anal fin and have
nearly identical meristic counts when compared with the holotype, the head-in-
standard length values are quite different: 9.84 for Syngnathus randalli and
7.35—7.45 for the Haitian specimens. In addition, the head profiles of the Hai-
tian and Venezuelan specimens are more dissimilar than would be expected if
they were size variables of the same species.

Discussion. Syngnathus randalli is best described as a short-nosed cousin
of Syngnathus elucens, but it also differs from S. elucens in that it lacks the
anal fin. Absence of the anal is important because the only other Atlantic
American pipefish with the same lateral ridge pattern that also lacks the anal

Vor. XXXII] HERALD: ATLANTIC AMERICAN PIPEFISHES 375

fin is Syngnathus dunckeri. However, this latter species has a much shorter
snout (snout-in-head 2.5—3.4 for S. dunckeri versus 2.41 for S. randalli), and
also the dorsal fin of S. dunckeri covers 6 or 7 rings rather than 5 rings and is
located entirely on the tail.

NAMED in honor of Dr. John E. Randall whose extensive underwater eco-
logical studies have made him a leader in the new approach to ichthyology.

LITERATURE CITED
GINSBURG, ISAAC
1937. Review of the seahorses (Hippocampus) found on the coasts of the American
continents and of Europe. Proceedings of the United States National Museum,
vol. 83, no. 2997, pp. 497-594, figs. 54-71.
HERALD, Eart S.
1942. Three new pipefishes from the Atlantic coast of North and South America with a
key to the Atlantic American species. Stanford Ichthyological Bulletin, vol. 2,
no. 4, pp. 125-134.
1959. From pipefish to seahorse—a study of phylogenetic relationships. Proceedings of
the California Academy of Sciences, 4th ser., vol. 29, no. 13, pp. 465-473, 3 figs.

= Ayu. Wem «K* oe ern Deere,

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OF THE

CALIFORNIA ACADEMY OF SCIENCES
FOURTH SERIES

G Dallas Hanna Anniversary Volume

Vol. XXXII, No. 13, pp. 377-404; 12 figs.; 3 tables October 8, 1965

THE PINNIPED POPULATION OF
ANO NUEVO ISLAND, CALIFORNIA
By

Robert T. Orr

California Academy of Sciences, San Francisco, California 94118

and
Thomas C. Poulter

Stanford Research Institute, Menlo Park, California

Agstract: <A 3-year study on Ano Nuevo Island, California, showed that this area was
regularly used by four species of pinnipeds, the Steller sea lion (Eumetopias
jubata), the California sea lion (Zalophus californianus), the northern elephant
seal (Mirounga angustirostris), and the harbor seal (Phoca vitulina). Steller sea
lions were found there throughout the year although a minimum population, con-
sisting of several hundred cows and young, occurred in late winter and early spring.
In Jate April the first large males arrived. Maximum numbers of about 100 bulls
were present by the end of May. Bachelors also began arriving in late April and
reached maximum numbers by late June. There was a great increase in the number
of females in late May and early June as the harems formed. By the end of June
the female population was about 1,100. An estimated 1,000 pups were born in June
and early July. The bulls and bachelors began to leave in the latter part of July at
which time the harems broke up. They were essentially all gone by late August or
early September. In midwinter another exodus of many of the females and young
occurred.

Male California sea lions were present in numbers except from early June to
late July when the species was essentially absent from the island. No females were
noted here. Approximately 80% of the males present were adults. There were two
population peaks, one in mid-May and the other around September 1. The latter
peak was the higher and was found to double each year.

Elephant seals were present throughout the year; breeding occurred from late
December to early March. Greatest numbers were noted in early May and during

[ 377 ]

CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

wW
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ao

October and early November. The spring peak was the higher of the two. Total
numbers recorded on May 7, 1961, May 15, 1962, May 8, 1963, and May 7, 1964
were 86, 157, 450, and 483, respectively. These figures indicated a marked annual
increase in numbers. Bulls were present only in midwinter and midsummer. In
spring and autumn the population consisted almost entirely of immature animals.
There was a resident population of about 100 harbor seals in the vicinity of Ano
Nuevo Island.

INTRODUCTION

In May, 1961, we began a study of the pinniped population on Ano Nuevo
Island which is one-half mile off the central California coast approximately 40
miles south of San Francisco. It was originally anticipated that observations
would be concerned almost exclusively with the Steller sea lion (Eumetopias
jubata) since the island has long been known to have one of the largest rookeries of
these animals south of Alaska. However, it was soon discovered that it was also
used regularly by three other species of pinnipeds, the California sea lion
(Zalophus californianus), the elephant seal (Mirounga angustirostris), and the
harbor seal (Phoca vitulina). A fifth species, the northern fur seal (Callorhinus
ursinus ), was also recorded here.

Between May 25, 1961, and May 7, 1964, 60 visits were made to the island
primarily for the purpose of observing seasonal changes in its pinniped popula-
tion and composition as well as to conduct behavioral studies on the several
species concerned. There were a few occasions, however, when unfavorable
conditions or disturbances made it impossible to secure accurate counts. Ad-
ditional trips were made for other purposes at which times incidental popu-
lation data were occasionally secured. All crossings of the half-mile channel
separating Ano Nuevo Island from the adjacent mainland were made in a 14-
foot fiberglass boat with an outboard motor. As the result of unfavorable
weather for considerable periods of time between October until April more
days were spent on the island in summer than in winter (tables 1 to 3).

Counts were made with the aid of binoculars and were usually checked
later against extensive photographs which were made on each visit. On-the-spot
counts were frequently made from the top of the lighthouse which affords a
view of most of the areas occupied by pinnipeds as well as by careful approach
to each of the rookery or hauling-out areas. Effort was made to determine the
sex and age composition for each species whenever possible.

HISTORY OF THE SEA LIONS IN CENTRAL CALIFORNIA

For the past century the specific composition of the sea lion population
along the central California coast has presented many confusing aspects. This
is an area where the ranges of two genera, Zalophus and Eumetopias, overlap.
The breeding range of the northern Steller sea lion (Eumetopias jubata) extends
from Japan northward to the Bering Sea, then down the coast of North America

VoL. XXXII] ORR and POULTER: ANO NUEVO PINNIPEDS 379

TaBLe 1. Total Steller sea lion population on Ano Nuevo Island from May,
1961, to May, 1964.

a ————————

Date Number Date Number
1961 1963
May 25 1,500 April 13 268
June 27 1,934 April 19 286
July 13 1,586 April 21 305
July 28 1,596 April 22 300
August 11 1,632 April 25 315
September 7 1,395 April 29 524
September 23 Ay May 8 350
1962 May 23 672
February 24 225 May 27 418
March 24 451 May 29 428
May 15 193 June 4 1,045
June 5 839 June 6 954
July 3 1,822 June 11 1,166
July 16 2,226 July 12 2,395
August 4 255 July 23 1,981
August 18 2,629 July 30 2,625
September 1 2,207 August 19 1,605
September 21 2,009 August 23 1,509
October 24 1,188 August 30 1,510
November 26 940 September 6 1,070
December 20 1,543 September 13 1,186
December 29 15220 September 27 1,602
1963 October 12 1,449
January 9 927 October 25 1,140
January 19 690 November 13 1,093
February 15 624 December 28 815
February 16 603 1964
February 17 583 January 8 445
_ March 2 ZN, March 30 244
March 3 240 May 7 477

SS SS ee eee

from Alaska to the Channel Islands off southern California (Scheffer, 1958).
The California sea lion, Zalophus californianus, is represented by three separate
breeding populations: Z. c. wollebaeki in the Galapagos Islands, Z. c. japonicus
in the Sea of Japan, and Z. c. californianus extending from the west coast of
northern Mexico northward to the Channel Islands off southern California. The

380 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser.

distribution of Eumetopias and Zalophus outside of the breeding season, plus
periodic decimation of large numbers of these animals by man in the past, has
been responsible for much of the confusion apparent in the literature.

Rowley (1929) refers to some of the presumed errors and inconsistencies of
early observers like Scammon (1874) and H. W. Elliott (1875). As Rowley
points out, Captain Scammon’s measurements and descriptions obviously indi-
cate that he confused Eumetopias and Zalophus in the area where the two species
overlap. Elliott (sup. cit.) reported both Eumetopias and Zalophus around San
Francisco and the Farallon Islands, 25 miles to the west, yet indicated that in
numbers Zalophus greatly predominated. Rowley implied that these early
observers were in error and that Zalophus regularly came only as far north as
Monterey. Bonnot (1928), however, who was also an authority on this group,
stated that “In the past, the California sea lion was fairly abundant and prob-
ably maintained rookeries as far north as the Farallons. They have been recorded
as far north as Puget Sound.”

Sea lions of both species periodically were hunted and killed for their oil and
hides from around 1860 until the early part of the present century (Rowley,
op. cit.; Bonnot, 1951). This resulted in the elimination of certain breeding
rookeries, including that at Seal Rocks, San Francisco, where Steller sea lions
formerly bred, and a general reduction in the total number of sea lions of both
kinds along the coast of California and Baja California. It also makes it diffi-
cult to ascertain the distribution and relative numbers of the two species of sea
lions in this region prior to their disturbance by man.

In 1927 the California Department of Fish and Game, at the instigation of
fishermen who believed that pinnipeds were increasing and endangering com-
mercial interest, made a census of the sea lions along the coast between the
Mexican border and the Oregon state line. This and subsequent censuses in
1928, 1930, 1936, 1938, 1946, and 1947 were conducted primarily by the late
Paul Bonnot (Bonnot, 1928a, 1928b, 1931, 1937; Bonnot, Clark, and Hatton,
1938; Bonnot and Ripley, 1948). The results of these censuses are summarized
in the last-cited publication. These counts were made in June or July, the
breeding season for both species concerned. Pups, in most instances were not
included. The northernmost point any California sea lions were noted was Pt.
Reyes where nine were seen in 1936. Proceeding geographically southward, 6
were seen on the Farallon Islands in 1927, 28 in 1930, 25 in 1936, and 90 in
1938. None was noted during these years at Purissima Rock which is several
miles south of Half Moon Bay. In 1936, 200 were counted on Afio Nuevo Is-
land. This was the first time this species was recorded here. No observations
were made between Ano Nuevo Island and the Monterey County coast because
of the lack of either suitable hauling-out areas or rookeries.

The Steller sea lion counts over these areas at the same time were much
greater, especially since Ano Nuevo constitutes what is believed to be one of the

VoL. XXXII]

ORR and POULTER: ANO NUEVO PINNIPEDS

381

TABLE 2. California sea lion population on Ano Nuevo Island from May,

1961, to May, 1964.

Date

1961
May 25
June 27
July 13
July 28
August 11
September 7
September 23
1962
February 24
March 24
May 15
June 5
July 3
July 16
Waly 25
July 30
August 4
August 18
September 1
September 21
October 24
November 26
December 20
December 29
1963
January 9
January 19
February 15
February 16
February 17
March 2

Number

1,500

232
417

Date

1963
March 3
April 13
April 19
April 21
April 22
April 25
April 29
May 8
May 23
May 27
May 29
June 4
June 6
June 11
July 12
July 23
July 30
August 3
August 19
August 23
August 30
September 13
September 27
October 12
October 25
November 13
December 28
1964
January 8
March 30
May 7

Number

186
1,289
1,254
1,146
1,966
1,488
1,345
2,000
1,183
1,258
1,090

482
309
185
15
198
3,203
4,650

10,275
5,874

13,367
Sis
8,119
4,000
Sooo
6,285
2,899

250
915
3,428

largest breeding rookeries for this species south of Alaska. At Pt. Reyes 45 Steller
sea lions were counted in 1936, 6 in 1938, and 2 in 1947. On Farallon Islands the
number was 700 in 1927, 540 in 1928, 900 in 1930, 500 in 1936, 357 in 1938,
and 750 in 1947. At Purissima the count was 150 in 1927, 42 in 1928, 4 in 1936,

382 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

2 in 1938, and 50 in 1947. The population on Ano Nuevo was 1,500 in 1927,
1,500 in 1928, 2,500 in 1930, 1,000 in 1936, 2,000 in 1938, and 2,050 in 1947.

A census was taken along the California coast in 1958 and again in 1960 and
1961 by the Department of Fish and Game (Ripley, Cox, and Baxter, 1962).
Pups were included in these censuses but no segregation was made as to species.
During these 3 years aerial counts showed the Ano Nuevo population to be 1,170,
1,350, and 2,342, respectively. Smaller aggregations, possibly breeding, were
noted on the Farallon Islands and occasionally small groups were found hauled
out at Pt. Reyes and Purissima. The latter animals were no doubt nonbreeding
individuals. Earlier observations in 1920 by Evermann (1921), and in 1924 by
Evermann and Hanna (1925), showed that in those years the Steller sea lion
population was around 2,000 on Ano Nuevo Island and no California sea lions
were observed there.

The numbers of California sea lions recorded were small and sporadic for
the years in which the censuses were made and seemed to lend support to Row-
ley’s (1929) conclusions that Scammon, Elliott, and other early observers con-
fused the two species.

Probably the first real clue to the reason for this seeming confusion was pre-
sented by Fry (1939) who made a population study of sea lions for the Califor-
nia Division of Fish and Game in March of that year on some of the Channel
Islands. The purpose was to determine the numbers of these animals along the
the California coast outside of the breeding season when they are rather widely
dispersed and many individuals may be overlooked. Despite this the March count
showed the number of California sea lions to be twice that obtained during the
census the previous summer. Fry (op. cit.) also noted that between the Channel
Islands and San Francisco fewer Steller sea lions were seen than during the
previous summer. He concluded that the increase in the number of California
sea lions was probably the result of an influx of animals from Baja California.
This, he stated, ‘could have been caused by the activities of a dog-food manu-
facturer who has been using sea lions from Lower California as a meat supply.
The creatures may possibly have decided that their old haunts were no longer
healthy, but is seems more likely that the migration is a regular annual event.”

With regard to the decrease in the number of Steller sea lions in March over
the previous summer’s population (Fry op. cit.) suggests: “If California sea
lions show a northward migration, it seems within reason that the Stellers would
show a corresponding movement.”

Bartholomew and Hubbs (1952) published an account of winter observations
on pinnipeds made late in January and early in February, 1950, on Guadalupe,
the San Benito, and Cedros islands off the northwest coast of Baja California.
The California sea lion populations on both Guadalupe and Cedros islands
were small (less than 200 and 340, respectively). On the San Benitos they were
extremely high, 9,714 individuals being recorded. Most interesting, however, is

VoL. XXXII]

ORR and POULTER: ANO NUEVO PINNIPEDS

TaBLeE 3. Elephant seal population on Ano Nuevo Island from March, 1961,

to May, 1964.

Date

1961
March 11
May 7
May 25
June 27
July 13
July 28
August 11
September 7
September 23
November 19

1962
February 24
March 24
May 15
June 5
July 3
July 16
August 4
August 18
September 1
September 21
October 24
November 26
December 20
December 29

1963
January 9
January 19
February 15
February 16
February 17
March 2

Number

24
86

70
71
a2

Date

1963
March 3
April 13
April 19
April 21
April 22
April 25
April 29
May 8
May 23
May 27
May 29
June 4
June 6
June 11
July 12
July 23
July 30
August 19
August 23
August 30
September 6
September 13
September 27
October 12
October 25
November 13
December 28
1964
January 4
March 30
May 7

Number

_ 243
276
276
325
3)5))
429
450
400
200
125
107

the fact that most of these animals were less than 3 years old and the majority
were yearlings. These writers state (op. cit.): “Of the 9,714 sea lions which
we counted on the San Benitos, only 11 were identified as adult males.” They
concluded, as did Fry (op. cit.) that there was a northward postbreeding move-

384 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

ment of California sea lions from Baja California. They indicated that this
northward migration was essentially confined to adult males. They further state
(op. cit.) regarding adult male California sea lions as follows: “That they may
migrate northward as far as California seems possible. A very high proportion
of adult males has been observed in nonbreeding aggregations in that state.
Thus on April 22, 1950, Bartholomew estimated that fully 80% of the 956
sea lions on a sandy beach on San Nicolas Island were adult males. The over-
wintering population of sea lions on a beach in Monterey County south of Point
Sur, with numbers estimated by Hubbs as high as 1,800 and by others still higher,
consists largely of adult males of this species, with a few Steller sea lions.”

Bartholomew and Boolootian (1960) made a study of the summering and
wintering populations of pinnipeds on the Channel Islands off southern Cali-
fornia so that comparisons could be made with data obtained since 1927 when
the first comprehensive census was made by the California Department of Fish
and Game. The Steller sea lion population, which reached a peak of about
2,000 animals in that area in 1938, was found to be almost completely replaced
by California sea lions by 1958. Only 50 individuals of the former species were
recorded the latter year and no bulls were noted in winter. The increase in
California sea lions was logarithmic. The nonbreeding population for the Chan-
nel Islands was between 9,000 and 10,000 in 1959 and the breeding population
that year was more than 13,000. In 1927 the total count for this species for
the entire state of California was only 915 in summer. Furthermore, the winter
population of California sea lion bulls in 1958 was greater than the summer
population. This tended to confirm the theory that there is a northward
movement of California sea lion bulls in winter and possibly a southward
movement of cows and immatures. The absence of Steller sea lion bulls in winter
also indicates the possibility of a northward movement of adult males of this
species after the breeding season.

The data from our studies confirm the suggestions made by Fry (1939),
Bartholomew and Hubbs (1952), and Bartholomew and Boolootian (1960)
concerning a postbreeding northward migration of male California sea lions,
and also support the idea that there is a similar northward postbreeding migra-
tion on the part of male Steller sea lions in central California. Seasonal move-
ments of Steller sea lions have been noted in British Columbia by Pike and
Maxwell (1958) and in Alaska by Kenyon and Rice (1961).

ANo Nuevo IsLAND

The island (fig. 1) is irregular in shape with its main axis in a northwest—
southeast direction. It has a land mass of slightly less than 12 acres but an ad-

Figure 1. Aerial view of Ano Nuevo Island with Afio Nuevo Point in the distance.
Photograph by John Gorman, April 7, 1961.

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Q
es
aX
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=
=
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ie
S
SI
Le)
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z
x
Re
cs
WN
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=)
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a
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8
Re
Re
i)

VoL. XXXII]

386 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 2. Map of Aho Nuevo Island. The areas most frequently used by pinnipeds
are referred to by number.

ditional 4 acres is provided by adjacent rocks and reefs. These areas were com-
puted at low tide level. The greatest length along the main axis from water line
to water line is about 1,300 feet and the greatest width 850 feet. At its narrowest
point it is 36 feet. On its seaward side, from the northwestern end to the south
end, there are numerous slabs of rock rising out of the water and separated from
the main mass of the island. Some of these lie 450 feet from the is-
land. The larger of these rocks, whose exposed surfaces are 300 to 400 feet
long, are used extensively by Steller sea lions. For convenience and quick ref-
erence, numbers were given to parts of the island and to those rocks that were
utilized to a significant extent by pinnipeds (fig. 2).

The island is composed of layered beds of Miocene cherty shale which gen-
erally slope down toward the southwest. The most extensive reefs and exposed
slabs of shale are on the seaward side. There are two fairly extensive sandy beaches,
one on either side of the southeastern half of the island, each over 300 feet in
length. The main mass of the island above the beaches and reefs is covered with
sand on which a number of kinds of plants grow. The movement of
large numbers of sea lions has essentially eliminated vegetation over the
southeastern half of the island, but these animals, even when abundant along the
shoreline, rarely come onto the top of the northwestern part of the island. Here
Distichlis spicata, Lupinus arboreus, Eriophyllum staechadifolium, Cakile mari-
tima, Oenothera cheiranthifolia, Amsinckia spectabilis, Franseria bipinnatifida,
Chenopodium californica, Spergularia macrotheca, and Echeveria farinosa are
among the most common plants found.

For a history of the island and its former utilization as a lighthouse station

VoL. XXXII] ORR and POULTER: ANO NUEVO PINNIPEDS 387

Ficure 3. A Steller sea lion harem in area 7. The bull (slightly right of center) is
surrounded by cows with pups in the foreground. Photograph by Robert T. Orr, July 9,
1964.

by the United States Coast Guard, see Orr and Poulter (1962). Although Ano
Nuevo Island is the most important area for pinnipeds in central California,
certain supplemental data obtained from other localities in this region, including
the Farallon Islands, Pt. Reyes in Marin County, Seal Rocks off San Francisco,
and coastal San Mateo County, are included here.

STELLER SEA LION

AREAS OCCUPIED. In general the Steller sea lions showed preference for the
large, outlying rocks off the northwest end of Ano Nuevo Island. Those rocks,
designated as 11 and 12, were found to have some of these sea lions on them
throughout the year. On only one occasion, December 20, 1962, was rock 11
unoccupied. On this day, however, it was completely awash because of very
large swells. Two of the three largest aggregations of breeding sea lions occupied
these rocks during the summer months. The other most utilized rock was 10,
which, because of its partial division by a surge channel, was designated as 10
E (east) and 10 W (west). Rock 14 was utilized as a hauling-out area through-

388 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

200 [a
it N
2400 — IN /\ i \
1964 \
Pos
ge Pease Gee [Aj \ SS
veal 1962 / \ N ‘
< EEG rok z \ I
a | ——-—— 196! Ha Ae 7 :
> 1600}— WA / rie \
2 4 / \ //
WL v
2) | 200) = /
ui pe it V
g /
= |
> g00-\ /

Ficure 4. The total Steller sea lion population on Ano Nuevo Island from May 25,
1961, to May 7, 1964.

out most of the year by a small number of Steller sea lions but it was of con-
siderably greater importance to California sea lions and harbor seals. A few
harems of Steller sea lions were noted here in the breeding season. Areas 1 and
2 were used to a large extent by bachelors or nonbreeding animals during the
breeding season, and to a limited extent by a few females and young during other
times of the year when the California sea lion population was low. Area 7 con-
tained an active breeding rookery with 11 harems in the summer of 1961. This
is the only breeding group recorded on the main island, either in 1961 or 1962. In
1962 area 7 was occupied by a number of solitary bulls, but no females were
present. In 1963 and 1964, however, both area 7 (fig. 3) and area 9 contained
breeding groups. Toward the end of the breeding season the smaller rocks
around the periphery of reef 7, including 6a, were frequently used as hauling-
out areas for cows, pups, and even subadults and occasionally bulls. Solitary
bulls were frequently seen on area 8 from late May to early July; and during
most of the breeding season some females and pups were found on 9a.

The large sandy beach on the eastern side of the island was not used by
Steller sea lions although the sandy beach on the west side (no. 3) was oc-
casionally occupied by bachelors at the height of the breeding season. It ap-
peared that they were forced to occupy it then because the more favorable situa-
tions were preempted by the breeding bulls who were rather intolerant of the
bachelors from early May until the middle of July.

BREEDING POPULATION. Although there was marked seasonal fluctuation in

VoL. XXXII] ORR and POULTER: ANO NUEVO PINNIPEDS 389

the total number of Steller sea lions occupying Ano Nuevo Island (fig. 4, table
1), as well as changes in the sex and age composition throughout the year,
there appears to have been relatively little change in the number of animals
breeding here for at least the past 42 years. Evermann (1921), in his report on
the Ano Nuevo Steller sea lion rookery as of June 27 and 28, 1920, estimated
the total population as “between 1,500 and 2,000, with the probability that
2,000 is more nearly correct.” His total pup count for the reefs and island
was “‘not more than 100” with another 106 dead or dying pups recorded on the
beaches and floating in the water. Counts made on June 22 and 29, 1924, by
Evermann and Hanna (1925) again resulted in an estimate of approximately
2,000 animals of all ages. Of this number 954 were adults. Another 150 were
estimated to have been overlooked. This would indicate a total adult popula-
tion of about 1,100 animals. Mention is made of the absence of subadult males
and females at this time. Such nonbreeding animals may have been disturbed
by human beings, as the island was then being used as a lighthouse station, and
nonbreeding animals are easily flushed into the water in contrast to those on
breeding rookeries.

These figures are not too different from those obtained by us during the
breeding seasons of 1961, 1962, and 1963. On May 25, 1961, a total of 1,500
adults or subadults, including yearlings, was estimated to be present. No separa-
tion by age and sex was attempted, but this was just prior to the birth of the
first pups. On June 27, 1961, the total population was estimated to be 1,934.
On July 3, 1962, the adult and subadult population was estimated to be about
1,500, consisting of approximately 100 bulls, 275 bachelors and subadults, and
1,125 adult females. In addition 750 pups were counted. On July 12, 1963,
the Steller sea lion population consisted of about 100 bulls, 266 bachelors and
subadults, 1,444 cows, and 585 pups. The pup count was too low because of
poor visibility.

The various censuses taken by the California Department of Fish and Game
from 1927 to 1961 tend to substantiate the relative uniformity of the breeding
population. The summary given by Ripley, Cox, and Baxter (1962) shows a
range in numbers counted from 1,200 to 2,500. However, the time, as previously
noted, varied from June to July and no effort was made to distinguish species.
Furthermore, some counts were made on the ground and others by means of
aerial photography.

Butts. Adults bulls spend a relatively short time on the island. They first
arrive in the latter part of April, reach maximum numbers by the first of June,
begin to leave late in July, and are mostly gone by the middle of August. On
April 25, 1963, 10 bulls or large subadult males were observed on the island.
Approximately 2 weeks later (May 8) 52 large males were present. Forty-
seven were counted on May 7, 1964. Twenty-two of these were bulls and all
except one, which was sleeping on beach 3 with a group of California sea lions,

390 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

were established on rookery areas. Twelve were associated with cows and
yearlings. The remaining 27 males were in bachelor groups. By the last week
in May in 1961, 1962, and 1963, approximately 100 harem bulls were present.

By July 28, 1961, the harems had largely broken up and a marked decline
in the number of bulls was apparent. In area 7 only two lone bulls were seen
on rocks that had contained 10 bulls, 120 cows, and 100 pups 15 days earlier.
Aggregations composed of females, pups, subadults, and several bulls were
seen on some of the larger islets. On August 4, 1962, and August 11, 1961, only a
few bulls were seen in the island area and on August 18, 1962, only one large
bull was seen among the 2,427 Steller sea lions counted. The August population
consisted almost entirely of adult females, pups, yearlings, and subadults.

FEMALES. Evermann and Hanna (1925) reported that the lighthouse
keeper on Ano Nuevo Island informed them that some sea lions (presumably
Eumetopias jubata) are present on the island throughout the year. They suggest
that “it is possible that the young males and females haul out there after the
breeding season is over.”’

At no time during this study was the island devoid of adult female Steller
sea lions, although there were marked seasonal changes in numbers. Maximum
numbers were recorded from shortly after the beginning of the breeding period
in early June, through September, and minimum numbers in late winter and
spring.

On May 15, 1962, the total population for this species consisted of about
200 individuals of which less than 100 were females. On this date some bulls
were noted. The arrival of large numbers of females followed the arrivals of
the bulls. On June 5, 1962, 558 adult females were counted and the number
reached 1,125 on July 3, 1962. The reason for the small number on June 5 of
that year is not understood. Some of the females present had young by this date.
Furthermore, the presence of harbor seals, elephant seals, and California sea
lions indicated that the island had not recently been disturbed by human
activities. The previous year, on June 27, the total adult female population was
1023.

From September until early May the composition of the Steller sea lion
herd consists entirely of adult females, young of the year, and a few animals
judged to be between 1 and 2 years old. There was a marked decline each year
in midwinter in the number of animals present (table 1).

Little is known about the movements of cows or young in winter and spring
when the Ano Nuevo population shows a marked decline. Some evidence indi-
cates, however, that it may be largely local, perhaps confined to the central
California area. During these seasons some females and immatures are usually
to be found on Seal Rocks and Pt. Reyes. The following numbers of Steller
sea lions, thought to be females and immatures, were observed on Seal Rocks
on each of the following dates: January 27, 1962, 10; March 3, 1962, 19;

—————

‘VoL. XXXII] ORR and POULTER: ANO NUEVO PINNIPEDS 391

Ficure 5. Adult and immature male California sea lions on beach 3. Some may also
be seen on the edge of area 19 (upper left) next to the abandoned Coast Guard houses and
on area 2 (extreme upper right). A few elephant seals are on the upper beach just below
the houses. Photograph by Robert T. Orr, November 13, 1963.

May 26, 1962, 12+; February 14, 1963, 45; February 16, 1963, 5. A visit to Pt.
Reyes on April 15, 1962, showed 10 females and immatures hauled out with
a group of 90 California sea lion bulls. We were not able to visit the Farallon
Islands where sea lions are present throughout most of the year, but it is
probable that a considerable number of Steller females and immatures haul out
there.

BacHELors. Nonbreeding males begin to appear on the island at about the
same time as the bulls. The first individuals were noted by the latter part of
April. On May 8, 1964, 25 bachelors (males not sufficiently old to secure a
harem) were present. On June 5, 1962, when 101 bulls were counted, 154
bachelors were recorded. On June 27, 1961, about 200 bachelors and bulls
without harems were observed on one beach alone. Others were scattered
along the shore, on reefs, and in the water. Two hundred and sixty-four bachelors
were counted on July 3, 1962. In 1961, 1962, and 1963, the maximum
bachelor and yearling population during the breeding season was between
275 and 300 individuals. By the middle of July there appeared to have been a
decline in the number of these nonbreeding sea lions.

Evidence that a northward migration starts about mid-July was obtained

392 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 6. California sea lions hauled out on areas 19, 3, and 2 of Ano Nuevo Island.
Photograph by Robert T. Orr, August 30, 1963.

on July 20, 1961, when one of us (Orr) visited Pt. Reyes, approximately 70
miles northwest of Ano Nuevo Island. On this occasion 32 large Steller sea
lions, believed to be subadult males or young bulls, were seen hauled out on
exposed rocks in the water or on ledges at the base of a cliff. Since Ano Nuevo
Island is the only known breeding rookery along the central California coast
it was suspected that these animals were recent arrivals from there. Nineteen bull
California sea lions, also believed to be recent migrants from southern California
or northern Baja California, were intermingled with the larger Stellers. A visit
to Ano Nuevo Island 8 days later revealed a marked decline in both the
number of bachelor and bull Steller sea lions and a corresponding increase
in male California sea lions. The departure of all of the bachelors was not
complete until early September. Not one of this age group was seen on September
7, 1961, while on September 1, 1962, a few remained.

Younc. On May 25, 1961, two dead pups were found on one of the
sandy beaches but no living young were seen. The pups, which had died very

VoL. XXXII] ORR and POULTER: ANO NUEVO PINNIPEDS 393

recently, were thought to have been born prematurely. On June 14, 1961,
numerous young were observed. In 1962 no young were present on May 22,
but on June 5 there were 18 newborn young. By July 3 the number had
increased greatly and 750 were counted. This was considered a very conservative
figure since the pups are difficult to count and many are overlooked, especially
on the outermost reefs, when they are lying on the far side of adult sea lions.
It is probable that the actual figure was closer to 1,000. At the peak of the
breeding season in 1961, 1962, and 1963, the total number of adult females
was estimated to be approximately 1,100, and probably most of these bore young.

There was a high pup mortality during the first month following birth.
Likewise, in midwinter the number of young showed a marked decline. The
latter, however, appeared to be correlated with the disappearance of many
of the females. It was thought that either many of the young and their mothers
left the island or else both spent more time at sea, thus leaving the island
relatively depopulated. However, some young were always to be found there
throughout the winter and spring months.

A small percentage of yearlings was present during the breeding season
and some were observed until the following January at which time they were
19 or 20 months of age. Mathisen, Baade, and Lopp (1962) estimated that
yearlings constituted about 15 per cent of the Steller sea lion population on
Chernabura Island, Alaska, in the summer. This is much higher than we
observed on Ano Nuevo Island.

CALIFORNIA SEA LION

AREAS OCCUPIED. Members of this species, unlike the larger Steller sea
lions, show preference for sandy beaches or inner reefs and rocks. The greatest
numbers of California sea lions were found on the southern part of Ano Nuevo
Island because of the presence there of these habitats. The area most frequently
used for hauling out throughout much of the year was the sandy, seaward beach
designated as 3 (fig. 5). This beach is approximately 350 feet long and slopes
gently back 50 to 75 feet to the base of the bluff marking the edge of the main
body of the island. The few individuals present in midsummer could be found
here. This was also true during the population low of late winter and early
spring. On September 27, 1963, when the population was high, 2,125 California
sea lions and 75 elephant seals were counted on this beach. The greatest number
seen here was on May 7, 1964, when 2,700 California sea lions and 483
elephant seals were recorded. Since the area of this beach is only 0.56 acre,
an average of about 7 square feet was available for each of the 3,183 animals.

During the population peak in late summer and early autumn area 19, com-
prising the top of the south end of the island, was frequently utilized as a
resting area by the majority of the California sea lions present (fig. 6). For

394 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

14,000

12,000

1964

— 1963

10,000 —------- 1962

HONS

8000

6000

NUMBER OF INDIVIDUALS

4000

2000

J B M A M J J A S (0) N D

Ficure 7. Seasonal fluctuation and annual increase in the California sea lion population
on Ano Nuevo Island from May 25, 1961 to May 7, 1964.

example, on August 30, 1963, about 9,000 individuals were observed here. The
total count for the island on this day was slightly more than 13,000. On
September 13, 1963, when the total count for this species for the island was
approximately 8,000, about 5,000 were on area 19. At other times of the year
area 19 was rarely used. Beach 17 on the leeward side of the island was another
hauling-out area found to be important when the population was high or when the
animals had been disturbed elsewhere. It is 0.95 acre in extent and, unlike the
seaward beaches, consists of coarse gravel, shells, and rocks in addition to sand.

VoL. XXXII] ORR and POULTER: ANO NUEVO PINNIPEDS 395

In late August and September 1963, over 1,000 individuals were recorded here on
several occasions.

Areas 1, 2, 3a, 6a, 9, 9a, 14, 15, 16, and 18 were also frequently used by
California sea lions. Rocks designated as 4, 5, and 6 were of little significance
because of their small size. Less frequently used as hauling-out areas by mem-
bers of this species were areas 7, 1OE, 1OW, 11, and 12. These provided the major
breeding areas for Steller sea lions. Area 11 was also occupied throughout
the year by Steller sea lions. Only on rare occasions did California sea lions
haul out here. Their essential absence from this rock, however, was attributed
to its exposed seaward position rather than to the presence of the larger Steller
sea lions (Orr, 1965).

SEASONAL AND ANNUAL CHANGES IN NUMBERS. As noted previously, there
is little information available on the presence of this species in central California.
Several aerial photographs of Ano Nuevo Island taken in 1953 and made
available to us by the United States Coast Guard reveal the presence of about
1,000 California sea lions on beach 3. From our present knowledge of the
habits of these animals we judge that the pictures were taken sometime during
the winter season.

Two of the most interesting discoveries made during this study were the
marked seasonal changes that occurred in the numbers of California sea lions
on Ano Nuevo Island and the great increase in the population each successive
year. There were two population peaks during each year, one just prior to the
middle of May and the other about the first of September (fig. 7, table 2). The
September peak, however, was by far the higher. Both population highs were
of rather brief duration. Following the spring peak there was a steady decline
in numbers until the latter part of June when practically no California sea lions
were present. They reappeared in the latter part of July and within 5 or 6
weeks reached maximum numbers. By October 1 the population declined to
about one-half that present in early September and then gradually leveled off
until the first of the year at which time a second low of 1,000 or less was found.
Around April 1 there was an increase in numbers again which culminated in
the spring peak. The population during the September peak in 1962 was essential-
ly twice that of 1961, and that of September 1963 was twice that of 1962 (fig.
7, table 2). This is even greater than the rate of increase recorded from the
Channel Islands by Bartholomew and Boolootian (1960).

The absence of California sea lions throughout most of June and July
in Central California was correlated with the breeding season for this species
and we presume these animals migrated south to rookeries on the islands off
southern California and the west coast of Baja California. A possible explana-
tion to account for the fall and spring peaks at Afio Nuevo Island is that many
sea lions migrating to and from nonbreeding areas north of central California
rest at the island for a while en route. This might be an especially important

396 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 8. Immature elephant seals on beach 3. Male California sea lions may be seen
in the distance near the water’s edge. Photograph by Richard Jennings, early May, 1963.

resting place in late August and September after the rigors of the reproductive
season farther south.

SEX AND AGE COMPOSITION. In no instance during the 3 years of this study
was the total California sea lion population found to consist of less than 80 per
cent adult males. The others were immatures ranging from subadults down to
individuals that were thought to be yearlings. Every individual in this immature
class whose sex could be determined proved to be a male. This was also true
of all young California sea lions found dead on the island.

These observations substantiate the suggestion made by Bartholomew and
Hubbs (1952) that adult male California sea lions move north to California
after the breeding season. These authors found males of this species essentially
absent in winter from the islands where they breed along the west coast of Baja
California. They also comment on the fact that on April 22, 1950, fully 80 per
cent of the 956 California sea lions on San Nicolas Island off the coast of
southern California were adult males.

NORTHERN ELEPHANT SEAL

AREAS occuPIED. The history of the development of the elephant seal
colony on Ano Nuevo Island has already been described (Radford, Orr, and
Hubbs, 1964). Elephant seals were most often seen on beach 3 where they

Vor. XXXII] ORR and POULTER: ANO NUEVO PINNIPEDS 397

Sa Beet M

eR ee

Ficure 9. A young bull elephant seal in process of molt in summer on Ano Nuevo
Island. Photograph by Robert T. Orr, July 9, 1964.

tended to stay on the upper dry sandy parts. This is in contrast to California
sea lions which were usually found closer to the water’s edge (fig. 8). This
habitat preference by members of this species is thought to be associated with
their habit of throwing dry sand over the body with the front flippers.

Beach 3a was also used occasionally by a few immature animals. Rarely
single individuals were seen on top of the island in area 19. Small numbers of
immatures were sometimes noted on beach 17 in spring and fall. This area,
however, was regularly used by solitary bulls during the breeding season. These
large males might also be found anywhere about the periphery of the island
from late December until early March.

All breeding activity was confined to beaches 3 and 17. Cows and pups
were not noted elsewhere. Although most of the young were observed on beach
3, three were born on beach 17 in January, 1963, and 16 during the 1964
breeding season.

Mates. Large males, ranging in age from old bulls to subadults or
bachelors, were regularly observed from December until March and again
during July and August. A few large subadults were seen at other times. Each

398 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 10. An elephant seal bull with a harem of cows and pups on beach 3. Photo-
graph by Richard Jennings in late January, 1963.

year the population appeared to increase over that of the preceding year. Four
large males were recorded on July 28, 1961, 6 on February 24, 1962, 8 on July
16, 1962, 23 on December 29, 1962, 22 on July 23, 1963, and 20 on December
28, 1963. Even though this paper relates primarily to observations made up to
May 7, 1964, 30 big males were counted on July 9, 1964. The large males all
appeared to undergo a molt in summer (fig. 9).

FEMALES. Adult females are present during the reproductive season which
is from the latter part of December until March (fig. 10). Not one was observed
during July and August. Their presence in spring and fall was questionable and
difficult to ascertain because of the large number of immatures of both sexes
that were present then.

Pups. The young are born in January and February. Each winter during
this study the number of births increased. Twenty-three were recorded in 1962,
32 in 1963, and 60 in 1964. These young began leaving by the end of May and
by midsummer were mostly gone, although a few were noted later. Since metal
flipper bands were attached to 12 of the 23 pups born in the winter of 1962
and to all but one or two in 1963 and 1964, it was easy to determine the
presence or absence of the young subsequently. Some information was obtained
regarding the movements of these animals. One male, tagged as a pup on Afio
Nuevo Island on March 28, 1962, was observed by Dr. David Regnery and

VoL. XXXII] ORR and POULTER: ANO NUEVO PINNIPEDS 399

NUMBER OF INDIVIDUALS

Ficure 11. Seasonal fluctuation and the annual increase in the elephant seal population
on Ano Nuevo Island from May 7, 1961 to May 7, 1964.

Theodore C. Pinney on South Farallon Island on September 20, 21, and 22,
1962. In late May, 1964, three of five elephant seals seen by the same observers
on South Farallon bore Afo Nuevo Island tags. The Farallon Islands are ap-
proximately 55 miles northwest of Ano Nuevo Island. A male elephant seal
that had been tagged as a pup on Ano Nuevo Island on February 17, 1962, was
found dying on a beach 5% miles south of Cape Sebastian, Oregon, by Dan
Snook on May 30, 1964, and reported to the Oregon Fish and Game Com-
mission. The distance between these two localities is nearly 400 miles.

Immatures. The majority of elephant seals on Ano Nuevo Island come
under this category, here used to include animals ranging from about 6 months
of age up to, but not including, subadults. Males classified as subadult were
those with a fairly well developed, yet not markedly, pendulous proboscis and
which lacked the gray and pink coloration on the thickened epidermal shield
of the chest and neck. With the females age grouping was more difficult so
relative size alone was relied upon.

400 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser.

Because the young are born in January and February, the influx of im-
mature elephant seals which began in early September and reached a peak in
October (table 3) was composed of animals over 6 months of age. No adults or
subadults were noted until late in November when the immature population
began to decline. A few immatures were still present in January, but none was re-
corded in February. In March immatures again appeared on the island. This
influx continued through April, with the greatest number of individuals for the
year being recorded about the beginning of the second week of May (fig. 11).
Maximum numbers recorded for 4 years are as follows: 86 on May 7, 1961
(Radford, Orr, and Hubbs, 1964, table 1), 157 in 1962, 450 on May 8, 1963,
and 483 on May 7, 1964. On only one of these dates were any individuals other
than immatures and young of the year noted. This was on May 15, 1962, when
two subadult males were seen. Bartholomew and Hubbs (1960) noted a complete
absence of adults in April on Guadalupe Island.

After the middle of May the number of immatures declined, and by July
they were mostly replaced by large males. The number of males exceeded the
number of females during the spring peak but in late September and October
the sexes were present in about equal numbers.

HARBOR SEAL

AREAS OCCUPIED. The harbor seal, unlike the Steller sea lion, the California
sea lion, and the elephant seal, is a shoreline, bay, and estuarine species. In the
Ano Nuevo area it is not limited to the island but is also found along the
adjacent rocky mainland. There appeared to be a movement of individuals
between these two areas which are separated by only one-half mile of shallow
water.

On Ano Nuevo Island harbor seals were observed in the vicinity of the northern
and northeastern parts. During low tide area 13 was a favorite resting place.
Here they were scattered on rocks that were underwater at high tide and with
the incoming tide they floated or swam off. Some would remain in the protected
cove formed by rocks 11, 12, 13, 14, and 15, while others frequently moved onto
a ledge on the south end of 14 (fig. 12). A number of individuals could usually
be observed in the water east of both 14 and the northern part of 17 during high
tide.

On the mainland the principal hauling-out areas at low tide were offshore
rocks about one-quarter of a mile north of Afio Nuevo Point and a large off-
shore rock about 300 yards east of the point in Afio Nuevo Bay. At high tide
most of the seals appeared to stay in the water between the point and the island
or in the Ano Nuevo Bay just east of the point.

PoPULATION. The maximum number of harbor seals in the area extending
from the northwestern part of Afio Nuevo Bay to the end of the reefs about 1

VoL. XXXII] ORR and POULTER: ANO NUEVO PINNIPEDS 401

Ficure 12. Three harbor seals on a ledge near the water’s edge on reef 14. Photograph
by Robert T. Orr, July 23, 1963.

mile north of Ano Nuevo Point and out to the island itself was estimated
to be about 100. The greatest number counted at low tide was 86 on July 23,
1963. Not all potential hauling-out areas were examined at this time and some
individuals may have been overlooked in the water.

There was no indication, locally, of any migratory movement on the part of
harbor seals; consequently the population showed only minor fluctuations
throughout the year. Since the young are born in this region from late March
through May, somewhat greater numbers were noted in June and July.

NORTHERN FuR SEAL

On July 27, 1962, James H. Miller observed a fur seal periodically occupying
a rock in area 18. The following day one of us (Poulter) photographed it with
a 24-inch telephoto lens. Pictures were sent to Dr. Victor B. Scheffer and Dr.
Carl L. Hubbs, both of whom are familiar with fur seals. Each wrote that
without doubt is was a northern fur seal (Callorhinus ursinus); Dr. Scheffer
further concluded from the appearance of the animal that it was a male “at
least five years old and probably much older.”

402 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Through the courtesy of Mr. Raymond Bandar we also examined a skull
from a northern fur seal that washed ashore on Waddell Beach in the spring of
1959. This locality is about 3 miles southeast of Ano Nuevo Island.

On March 30, 1964, we found the remains of a newborn northern fur seal
on beach 17. The skin, vertebral column, and shoulder girdles were all that
remained, but this was sufficient for identification. It was suspected that gulls
had attacked the carcass. We do not know whether this seasonally early birth
occurred on this beach or at sea.

Northern fur seals regularly winter in small numbers off the California
coast but these are mostly females (Taylor, Fujinaga, and Wilkie, 1955). Since
many of the females are pregnant, an occasional birth prior to the northward
migration to the Pribilof Island rookeries does not seem too unusual. Much
more surprising is the presence of the adult male in the midsummer of 1962.
Males recorded along the California coast have been young and were not found
in midsummer.

ACKNOWLEDGMENTS

Many persons visited Ano Nuevo Island with one or both of us during the
course of this project. To these persons we wish to express our sincere thanks
for numerous valuable suggestions. We are indebted to the California Division
of Beaches and Parks for permission to make these studies on property under its
administration. Personnel of that Division in Sacramento, as well as in the
Half Moon Bay office, were extremely generous in providing maps, photo-
graphs, and other valuable data bearing on the region; frequently one or more
staff members accompanied us on our visits.

LITERATURE CITED

BARTHOLOMEW, GEORGE A., and RicHArp A. BOOLOOTIAN
1960. Numbers and population structure of the pinnipeds on the California Channel
Islands. Journal of Mammalogy, vol. 41, pp. 366-375.
BARTHOLOMEW, GEorGE A., and Cart L. Hupss
1952. Winter population of pinnipeds about Guadalupe, San Benito, and Cedros
islands, Baja California. Journal of Mammalogy, vol. 33, pp. 160-171.
1960. Population growth and seasonal movements of the northern elephant seal,
Mirounga angustirostris. Mammalia, vol. 24, pp. 313-324.
Bonnot, Pau
1928a. The sea lions of California. California Fish and Game, vol. 14, pp. 1-16.
1928b. Report on the seals and sea lions of California. California Division of Fish
and Game, Fish Bulletin no. 14, pp. 1-62.
1931. The California sea lion census for 1930. California Fish and Game, vol. 17, pp.
150-155.
1937. California sea lion census for 1936. California Fish and Game, vol. 23, pp.
108-112.
1951. The sea lions, seals and sea otter of the California coast. California Fish and
Game, vol. 37, pp. 371-389.

VoL. XXXII] ORR and POULTER: ANO NUEVO PINNIPEDS 403

Bonnort, PaAut, G. H. Crark, and S. Ross Hatton
1938. The California sea lion census for 1938. California Fish and Game, vol. 24,
pp. 415-419.
Bonnot, Pau, and WILLIAM ELLIs RIPLEY
1948. The California sea lion census for 1947. California Fish and Game, vol. 34, pp.
89-92.
ELLiotT, HENRY Woop
1875. A report on the condition of affairs in the Territory of Alaska. Washington,
United States Government Printing Office. 277 pp.
EVERMANN, BARTON WARREN
1921. The Afto Nuevo Steller sea lion rookery. Journal of Mammalogy, vol. 2, pp.
16-19.
EVERMANN, BARTON WARREN, and G DaLrias Hanna
1925. The Steller sea lion rookery on Ano Nuevo Island, California, in 1924. Journal
of Mammalogy, vol. 6, pp. 96-99.
Fry, Donan H., Jr.
1939. A winter influx of sea lions from Lower California. California Fish and Game,
vol. 25, pp. 245-250.
GRINNELL, JOSEPH
1933. Review of the recent mammal fauna of California. University of California
Publications in Zoology, vol. 40, pp. 71-234.
Kenyon, Kart W., and Date W. RIcE
1961. Abundance and distribution of Steller sea lions. Journal of Mammalogy, vol. 43,
pp. 223-234.
MArTHIsEN, OLE A., RoBert T. BAADE, and Ronatp J. Lope
1962. Breeding habits, growth and stomach contents of the Steller sea lion in Alaska.
Journal of Mammalogy, vol. 43, pp. 469-477.
Orr, RosBert T.
1965. Interspecific behavior among pinnipeds. Zeitschrift fiir Saugetierkunde, vol. 30,
pp. 163-171.
Orr, Rorert T., and THomas C. POULTER
1962. Ano Nuevo Marine Biological Park. Pacific Discovery, vol. 15, no. 1, pp. 13-19.
PIKE, Gorpon C., and Brian E. MAXWELL
1958. The abundance and distribution of the northern sea lion (Eumetopias jubata) on
the coast of British Columbia. Journal of the Fisheries Research Board of
Canada, vol. 15, pp. 5-17.
RaprorpD, KeirH W., Ropert T. Orr, and Cart L. Huss
1965. Reestablishment of the northern elephant seal (Mirounga angustirostris) off
central California. Proceedings of the California Academy of Sciences, 4th
ser., vol. 31, pp. 601-612.
Rrerey, WILLIAM E tis, Keiru W. Cox, and Joun L. BAXTER
1962. California sea lion census for 1958, 1960 and 1961. California Fish and Game,
vol. 48, pp. 228-231.
Row Ley, JOHN
1929. Life history of the sea lions on the California coast. Journal of Mammalogy,
vol. 10, pp. 1-36.
SCAMMON, CHARLES MELVILLE
1874. The marine mammals of the north-western coast of North America. John H.
Carmandy and Company, San Francisco. 319 pp.

404 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

SCHEFFER, VICTOR B.
1958. Seals, sea lions and walruses; a review of the Pinnipedia. Stanford University

Press, Stanford, California. x + 179 pp.
Taytor, F. H. C., M. Fuyinaca, and Forp WILKIE
1955. Distribution and food habits of the fur seals of the North Pacific Ocean. Re-
port of Cooperative Investigations by the Governments of Canada, Japan,
and the United States of America, February—July 1952. United States Fish
and Wildlife Service. x + 86 pp.

fPas6eo SSS - — ——-—
— oe ere

Marine Blolc gical Laboratory
Li @RAR *
OCT 191965 |

OODS HOLE, MASS

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES
FOURTH SERIES

G Dallas Hanna Anniversary Volume

Vol. XXXII, No. 14, pp. 405-432; 2 figs. October 8, 1965

BREEDING CHARACTERISTICS OF YELLOW.

BILLED CUCKOOS IN ARIZONA
=
William J. Hamilton II and Marian E. Hamilton

California Academy of Sciences*

The yellow-billed cuckoo is widely distributed throughout the United States,
southern Canada, and northern Mexico. The nominate race, Coccyzus a. ameri-
canus, is found in the most suitable habitat and is one of the commonest eastern
birds. There are no striking distributional gaps. Nevertheless, in eastern North
America this species has received only casual attention and the only account of
its life history consists of various remnant materials gathered by Bent (1940).
The western subspecies, Coccyzus a. occidentalis, at best a weakly defined race, is
more scattered, occurring only in certain relatively humid regions in the West,
especially along river bottoms in the southerly parts of its range. In California
some of the habits of this race have been reported by Shelton (1911) who de-
scribed a population nesting along a slough in Sonoma County in northern Cali-
fornia, and Jay (1911) and Hanna (1937) who reported characteristics of popu-
lations in Los Angeles and Riverside counties in southern California.

In Arizona the only accounts are Bendire’s (1895) report of a number of
~ nests along Rillito Creek near Tucson, and Brandt’s (1951) comments on the
status of the species in the San Pedro Valley near Hereford.

ACKNOWLEDGMENTS
Special thanks are due Clark Ross who participated in the field investi-
gations during August and made many significant observations. Adeline Gun-
* Present address, University of California, Davis, California.

[ 405 ]

406 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

tern typed all field notes taken on recorder tape; her rapid and accurate execu-
tion of this task greatly facilitated this research. In addition, we are grateful for
her help in preparing the manuscript. Mr. J. T. Howell, California Academy of
Sciences, supplied the plant indentifications.

LocaL ENVIRONMENT

The observations upon which this account are based were made along the
San Pedro River, Cochise County, Arizona, 4 miles downstream from Fairbank,
Arizona (31°43’ N., 110°11’' W.) at an elevation of 4,100 feet, and along
Sonoita Creek, Santa Cruz County, Arizona, southwest of Patagonia, Arizona
(31°33' N., 110°45’ W.) at 3,800 feet. A brief visit was made to the Patagonia
area in mid-June, 1963, and an intensive investigation was made during the
first 3 weeks of August, 1963, at both localities.

Brief additional observations were made at the Arizona—California border in
mid-June, 1964, near Laguna Dam on the Colorado River.

PHYSICAL ENVIRONMENT. ‘These two Arizona locations were of consider-
ably different physiognomy. At Patagonia a small stream, Sonoita Creek, cuts
through a narrow mountain valley. On either side of the river bottom, never
more than a half-mile wide and eventually narrowing to a gorge, rocky out-
crops and steep slopes give way to the arid jagged terrain of the Santa Rita and
Patagonia mountains. Sonoita Creek floods with seasonal rain, but the under-
lying rock basin forces water to the surface to maintain a permanent water
flow beginning about 2 miles below Patagonia. Upstream the flow is sub-
terranean except during the rainy season, and it is emergent only for a few
hours after storms.

The San Pedro River has, by contrast, a broad river bottom, in some places
over a mile wide. It centers in a sloping valley many miles wide. The drainage,
from a considerable area, maintains a permanent flow. During the rainy season
the shallow sandy river is over 100 feet wide in places.

These two areas, approximately 50 miles apart, share a similar climate. By
late June or early July thunderstorms originating in the nearby mountains
sweep across the intermountain plains, creating intermittent flood conditions
in streambeds. The rivers often rise under sunny skies, the result of runoff
from heavy rainfall elsewhere in the watershed. In consequence the sum-
mer climate during the yellow-billed cuckoo’s stay is regularly quite humid.
Only a few yards away from the river, however, the terrain becomes very
dry, and the humidity drops sharply shortly after a storm passes.

BIOLOGICAL ENVIRONMENT. No attempt is made here to describe the faunal
of these two localities. Swarth (1929) has surveyed the vertebrate fauna of
the Patagonia area.

Broad-topped tall cottonwoods (Populus fremontii) line the stream edge
and scatter across the stream bottom of Sonoita Creek. Tree willows (Salix

Vor. XXXII] HAMILTON and HAMILTON: ARIZONA CUCKOOS 407

gooddingii) edge the creek and their roots, together with occasional large rock
outcroppings, fix the stream meander. Additional willows occur irregularly in
the stream bottom and along lateral tributaries of sufficient drainage. These
willows often rise from several main trunks and may be as tall as 40 feet. Under
the cottonwoods is a ragged cattle-grazed understory of elderberries (Sambucus
glauca), walnuts (Juglans major), and mulberries (Morus microphylla). In
gravelly bars in the actual river bottom, arrowwood (Celtis reticulata) occurs
in dense stands, a favored nesting place of the yellow-breasted chat (J/cteria
virens). Four miles below Patagonia the river bottom widens, and in this flat
the ash (Fraxinus pennsylvanica) is exceeded in abundance only by the tree
willows. At the edge of this creek mesquite (Prosopis juliflora) flourishes,
shaded in places by tall cottonwoods. As mesquite extends from the river
bottom, individual bushes are smaller and more widely scattered. They extend,
however, well up on the rocky slopes before giving way to mesic vegetation such
as ocotillo, century plant (A gave sp.), and Opuntia.

The woody vegetation along the San Pedro River resembles that at Pata-
gonia. Cottonwoods, tree willow, and mesquite occupy comparable habitats.
However, in the broad river bottom the Eurasian tamarisk (Tamarix pentan-
dra), absent at Sonoita Creek, is a prominent feature of the broad river plain.
The Juglans, Sambucus, Morus, Celtis, and Fraxinus species, so characteristic
of the Sonoita Creek locality, were not found on the San Pedro below Fairbank.

One of the most important floral features of both areas was Condalia
lycioides, a spiny shrub scattered amongst the understory of the often continuous
stands of mesquite and spottily but regularly distributed amongst the river
bottom vegetation. A caterpillar hosted by this plant was an important food of
the yellow-billed cuckoo at both study areas in the summer of 1963.

DEMEANOR AND HUNTING BEHAVIOR

Cuckoos observed during the breeding season in August were not as vocal
as most breeding songbirds. Possibly these birds become more vocal with the
onset of July rains. In June they were practically silent. At that time we
each spent over 30 daylight hours encamped and actively searching an area
occupied by cuckoos. Yet we each heard but a single “kowlp” call. A 4-mile
- stretch of creek bottom was covered, listening carefully for 20- to 30-minute
periods, without hearing a cuckoo. But when we played a recorded yellow-
billed cuckoo call, we were at once able to locate an adult bird which had
apparently been lurking nearby. Subsequent use of this technique produced
two additional birds in less than 2 hours in the same area that had been so
carefully worked earlier. These observations point out the inadequacy of
attempting to determine presence or absence, much less abundance at this
season, by observation or listening-post techniques. A more casual observer,

408 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

spending 3 hours in the field in a morning, could expect to hear less than one
call a week at this season and in this area.

SKULKINEsSS. During the course of the fieldwork in August, it gradually
occurred to us that cuckoos were deliberately avoiding movement in our
presence, especially nesting individuals. Continued observation confirmed
this matter. When foraging undisturbed by the human observer, cuckoos would
move about in the willow and ash vegetation with no apparent predilection to
any particular level. High posts in cottonwoods were largely avoided. But
when a bird was surprised or approached incautiously, it often retreated to a
high post in the leafy lobes of cottonwood vegetation, holding this post
for a remarkable period of time without additional movement. On some oc-
casions the initial flight would take the cuckoo into the deep willow or mes-
quite understory, especially in areas lacking cottonwoods.

The positioning of these skulky birds was usually the same, backside to
the observer. The back was arched, the body held low, the head turned slightly,
watching the intruder. This performance, with the cryptically colored back
oriented to the witness, the white front shaded, was a regular response to the
observer. Once a bird in a willow was circled and during the periods when this
bird was in view, its position relative to the observer was the same.

During these occasions when the human intruder was holding the attention
of the cuckoos, they would not feed but simply devoted their time to watching
and to evasive retreat. Only after repeated association with a bird, or in areas
such as much used picnic grounds, would the cuckoo ignore the observer to
carry out more routine activities.

HUNTING BEHAVIOR. Occasionally we were able to follow a bird without
apparently disrupting its normal feeding routine for 2 hours or more. At
first impression these birds seemed to have time to spare, being mainly con-
cerned with scratching the head or other minor comfort movements, with much
leisure time left over. For nonincubating birds this may indeed have been the
case. However, it soon became apparent that what we were witnessing was
the hunting mode of this bird. They are, in fact, no less alert to potential food
reserves than a sparrow hawk sitting quietly on a wire over a field. Many of
the actions of the hunting cuckoo are indeed quite hawklike. Much time is spent
quietly waiting for the prey to reveal itself by movement.

Such a preying stratagem is suited only to predation upon large items,
i.e., items with a unit intake value approximately equivalent to the amount
of food which could be obtained by active search in an equivalent amount of
time minus the extra energy required for active versus passive prey search. The
items taken were indeed quite large, consisting, so far as we could indentify
them, of large moth larvae and katydids. Apparently numerous smaller insects,
many perhaps belonging to the same groups as those of the larger prey items
selected, were being ignored.

Vor. XXXII] HAMILTON and HAMILTON: ARIZONA CUCKOOS 409

An account of the relatively slow feeding tempo of a hunting cuckoo on
August 6 for 35 foraging minutes, spoken to a tape recorder, was as follows:

(1) At 0943 it hopped through a tree willow at the 15-foot level, then
(2) flew 60 feet to another tree willow 20 feet up, changed its perch three times,

(3) flew 40 feet to another tree willow, 10 inches from the outer limbs, and 20 feet up,
took a new perch 10 feet higher in the same tree, again changed perches at this level,
capered along a bare limb with tail cocked, stopped at this spot, then moved 5 feet
across the open heart of the tree, to take a new post,

(4) flew across an opening to an ash, 18 feet up, and almost at once

(5) flew on 100 feet to the outer limbs of a cottonwood 45 feet up, moved 30 feet across
the crown to a new post in the same tree, gave a “kowlp” call, and disappeared at 1018.

It thus changed posts 13 times in 35 minutes, with the feeding circuit including
five trees, one of them only momentarily. No prey was taken during this period.

VOCALIZATIONS

“KOWLP” CALLS AND SPATIAL RELATIONSHIPS. This characteristic call is the
species identification tag, often the field observer’s only indication of the
presence of this species. It has been variously described as ‘‘an uncouth guttural
sound or note, resembling the syllables kowe, Rowe, kowe, kowe, kowe!
beginning slowly, but ending so rapidly, that the notes seem to run _ in-
to each other, and vice versa. . . ” (Wilson and Bonaparte, 1878), and as
resembling a rapid pulling of corks from a bottle (Hanna, 1937). We have
already mentioned the almost complete absence of this or any other call by
cuckoos at Patagonia in June prior to the actual breeding season. In August
these calls were considerably more prevalent.

On several occasions “kowlp” calls were answered by ‘“kowlp” calls from
other individuals. In these instances when the calling bird was observed, it
faced toward the other calling individual and sometimes flew off in the direction
of it.

Each pair clearly ranged over several acres and it was impossible to follow
any individual long enough to effectively determine its spatial relationship
to its neighbors. The response to recorded calls varied from individual to
individual. The observer with the recorder was approached only occasionally.
Then the bird moved in with a swooping flight, tail and wings spread, and
- flight slowed as the bird moved past the observer and instrument to a distance
beyond. Frequently the responding bird either called back at the recording or
simply moved into the vicinity of the playback device, remaining silent and
perched at a high lookout post.

On August 11 at 1610 a cuckoo was heard giving repeated ‘“‘knocker”’ calls
(see below) at the mouth of a dry gulch entering the San Pedro River. This
bird was perched in the top of a 40-foot willow on the bank of the San Pedro.
Moments later another cuckoo flew into the adjoining willow only 15 yards

410 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

from the bird giving the “knocking” call. Shortly the original bird again gave the
“knocker” call. Almost at once the second bird gave a full “kowlp” call, then
flew over and supplanted the original bird, almost landing on top of him.
During the supplanting performance there was no obvious plumage or postural
display. The supplanted bird flew off downstream. The supplanting bird found
a small prey item near the top of this tree where the original bird had been,
secured it, and ate it almost at once. This individual was followed as it hunted in
the treetops here for the next 28 minutes. During this time it gave one additional
“kowlp” call at 1635. At 1640 it flew 75 yards upstream, to within 40 yards
of its nest, giving a full “kowlp” call as it landed. This and other similar field
observations suggest that the ‘““kowlp”’ call is associated with the species spacing
mechanism.

Since birds had probably been on these areas for 2 months, it is perhaps
not surprising that no clear territorial situation was determinable. If territoriality
is characteristic of this species, the wide range of each pair, covering many
acres, and their secretive and elusive habits will make the description of the
spatial characteristics of pairs a formidable task.

On numerous occasions birds gave “kowlp” calls after a flight, immediately
upon landing. On August 7 below Patagonia, at 0901, we heard a complete
“kowlp” call. It was possible to visually surround the tree from which this
bird called as we approached. Presently, the bird flew out of this willow, directly
overhead, with labored wingbeat and in full song. This vocalization approximated
the usual “kowlp” call, but the delivery was more deliberate. While it was not
the typical two-parted “kowlp” call, it was obviously composed of the same
notes. On no other occasion did I hear this peculiar assemblage of notes. This
bird landed within sight high in a cottonwood, went through a considerable
series of preening movements, and hopped to a high open perch and at once
gave a full and more characteristic ‘“‘kowlp”’ call.

On all occasions when a bird was observed giving a “kowlp” call, there was
never any indication of an extension of the chest or throat.

During August we heard over 200 “kowlp” calls. Since none was ever heard
beyond 150-200 yards, this was the carrying limit of this call to our ears.
“Kowlp” calls were given most frequently in the early morning hours before
0700. However, additional calls were given from time to time throughout the
day.

It is possible that the “kowlp” call may have a mating function as well.
On June 17, 1964, along the Colorado River, what was probably a male cuckoo
flew into a willow tree directly over our heads, 40 minutes before sunset. Mo-
ments later he flew to an upper position in the tree willows 50 meters from the
river and gave two complete “kowlp” songs. A female joined him in the dense
upper branches of this willow tree. This female took what seemed to be the pre-
flight intention movement (fig. 1c), but exaggerated, so that her tail pointed

Vor. XXXII] HAMILTON and HAMILTON: ARIZONA CUCKOOS 411

straight up. The male flew down, hovered over her with legs dangling, landed on
her back, and either attempted to or did mate with her. This all took place
quite quickly and the male at once flew off. The female remained in the vicinity
for awhile, then flew off in a different direction. Some minutes later the male
gave the “kowlp” call again, and was apparently rejoined by the female.

At this season (June 17-18) along the Colorado it was apparent that nesting
was not yet under way, for we saw several pairs moving about as a pair, often
separated by 50 meters or more, but occasionally coming together. Territorial
boundaries seemed to be in a state of flux, but with the general spatial features
already established. The frequent songs (“‘kowlp” calls) suggested that this
population was at a stage approaching that of the southern Arizona populations
we had observed on August 1 the year before. One pair was frequently observed
in a tree willow overhanging the river, a site to which they repeatedly returned.
Perhaps this was the potential nest site, but on June 18 there was no trace of
a nest in this dense vegetation.

On these occasions when two birds were under observation for a considerable
period of time only one bird of the pair was ever noted to give the “kowlp” call.
It seems probable, therefore, that the ‘“kowlp” call is limited to the male.

The loudness and ringing character of this call is probably adapted to the
large territory size of this species. Little attention seems to have been paid to
the volume characteristics of territorial proclamation. Increased volume should
be characteristic of sizable territories, the natural sound environment playing a
modifying role. This may be one reason why early morning song is so characteris-
tic, for it is a time of environmental stillness, generally windless. Selection for
lowered volume in territorial proclamation calls would come from factors of
energy conservation and reduction in the number of territorial boundary en-
counters.

At the Colorado River area this call seemed much more variable than in the
Patagonia or San Pedro areas. Here this call was often not as prolonged and
sometimes terminated with but a single or two or three ringing notes. Since
this population was decidedly more dense, it is possible that the more varied
vocal repertoire was related to this aspect of the population.

“KNOCKER” CALL. This call, a harsh rattled call, was quite different in
character from the clear “kowlp” call. Often repeated once initiated, this call
consists of a series of notes blended together; each quickly follows the other to
form an integrated call which sounds somewhat like a mechanical door knocker
allowed to drop freely against a striker plate. Unlike the “kowlp” call, this call
appeared to be limited to social situations; when we heard it we could be sure
that the mate was nearby. The volume was considerably less than for the
“kowlp” call and we rarely were able to hear it beyond 40 meters. A very
variable call, it seemed less ritualized than the ‘“kowlp” call, yet it was usually
quite identifiable as this particular vocalization. This call is given by both mem-

412 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

bers of the pair, often several calls to the sequence, again in contrast to the
‘“kowlp” call which was seldom repeated as frequently as once every 10 minutes.
The communicatory significance of this call was never well understood by us, but
the most frequent response of a bird hearing it was to return the call and fly off
in the direction of it if it were not already in the presence of the mate. In close
proximity return calls were regularly given. Yet we should not create the im-
pression that these birds chatter back and forth with this call in the normal
course of activity. Normally they are silent. Occasionally they give the ‘““kowlp”
call, and when disturbed or in other unidentified special circumstances the
members of a pair may vocalize with a ‘‘knocker”’ call.

“Coo” CALL. On several occasions during August a soft many-noted cooing
call was heard. These resonant sequences of cooing were repeated several
times per minute, often for an hour or more. Cooing birds held high to the
treetops, usually at exposed posts in dead snags 40 feet or more above the
ground. Sometimes these cooing birds would range widely to the surrounding
slopes several hundred yards from the river bottom, taking posts on bare oak
limbs. All the while these individuals were incredibly exposed by comparison with
the usually secretive demeanor of the birds in the river bottom. Moreover they
did not attempt to shelter the white breast, which was sometimes visible 100
yards or more in the slanting morning light. Exposure was indeed typical of
this behavior, and cooing birds moved openly along the river bottom, venturing
freely into areas occupied by other birds. Occasionally other cuckoos would
approach, to give the “kowlp” call once or twice, then back off. These cooing
performances were persistent, with activity located in certain broad areas for
days. We saw this performance from only three birds, out of a total population
of more than 30 taken under observation. One possible explanation, based upon
the persistence of the behavior, the response of other birds, the openness sug-
gesting advertisement, and the visual component described below, is that this
display is a function of unmated males.

Accompanying this call, with every note of each series of coos, the gular
area is remarkably inflated, filling to about the size and shape of a golf ball
(fig. 1). The sac is inflated and collapsed with each note. The bill is not opened,
but the head falls lower and lower with each successive coo.

The number of cooing notes per sequence varied from individual to
individual. A typical observation was as follows: “During the past 28 minutes,
starting at 0903 (August 7, 1963) the bird has given 4 to 6 series of these coo
notes per minute. The number of notes in each series does not vary to any great
extent. The following count is the total notes per series for a sample run: 7, 9,
8, 6, 8, 10,9, 11,9, 7, 5, 9, 9, 7, 6, 6.” Each sequence takes from 3 to 7 seconds
to deliver and the pause between sequences was from 7 to 10 seconds.

In June of 1964 along the Colorado River we heard this call repeatedly,
coming from at least three different birds. At that time the birds in that area

VoL. XXXII] HAMILTON and HAMILTON: ARIZONA CUCKOOS 413

Ficure 1. Postures of the yellow-billed cuckoo. The upper left shows a “coo’’-calling
cuckoo with throat sac inflated, the upper right the same bird at the end of a series of
“coo” notes. The lower left is the flight intention movement (but see text) and the lower
right is a hunting and alert (to man) posture. Drawings by Bob L. Olson.

moved about in pairs, apparently defending large territories together. The ob-
servation of coition and the frequency of ‘“‘kowlp” calls suggested that nesting
was imminent but not yet under way. The context of these ‘‘coo”’ calls was less
apparent in this region since we were unable to extend observations over a
sufficient period to evaluate the status of individual birds. However, these
observations seem to offer additional evidence that these calls are characteristic
of prebreeding behavior.

SCREAM. On August 18 we attempted to net the birds at a nest with nestlings.
In spite of the presence of a mist net near the nest, the female at once returned
to brood the young and was secured in a fold in the net when it left the nest. As
one of us was climbing to within about 12 feet of the netted bird, it ceased its
struggles and began to scream continuous loud, harsh, rasping cries. Almost at
once the male dashed through a gap in the crown, struck the net full force, and
was captured. This call as never heard on other occasions in the natural situation.
These individuals gave this call in captivity on other occasions when attempts
were made to grab them inside a cage.

414 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER.

“Mew” cart. A unique mewing sound accompanied the distraction per-
formance of these birds under certain circumstances (see Incubation, below).
Like the scream, this call is reserved strictly for predation situations.

CALLS OF THE YOUNG. Some of the vocalizations of the young are described
below under Development of the Young. It is interesting to note that in the
young there is an additional unique vocalization which has apparently evolved
in the context of potential predation.

INCUBATION

Data on the incubation period were obtained at nest 2. This nest was located
on August 6, 1963, at 0904. It contained one cold egg. No bird was on or close
to the nest. The egg was marked. On August 8 at 1025 a bird was on the nest
incubating two eggs. The new egg was noticeably smaller and more evenly
colored than the first. It was also marked. Daily visits on the following days
confirmed that the clutch was complete at two eggs. On August 17 at 1045 the
nest contained one egg and one freshly hatched young. The following morning
at 1030 the second egg had hatched.

An approximate determination of the incubation period is possible based
upon these data. Since the eggs hatched on separate days, incubation must have
started on separate days, assuming the incubation period of both eggs was the
same. The earliest incubation could have started would be the morning of August
6 for the first egg and August 7 for the second. It could have been a day later for
both. This would make the incubation between 10 and 11 days. Perhaps the
first egg was laid on August 5 and incubated during the nights of August 5—6
and August 6—7, with the second egg being laid the morning of August 7 and
continuous incubation starting at that time. This would still leave the incuba-
tion period at 10 or 11 days.

With this incubation period as a basis, it is possible to fairly accurately
estimate the timing of the start of the other nests. Nest 1 contained two young
and one egg on August 13 at 1030. One of these young gaped freely when the
nest was jiggled, the other kept its head down. Probably they had hatched on
separate days, one on August 12 and one the morning of August 13. The third
young had hatched when the nest was checked at 1255 on August 14. Presumably,
then, the final egg was laid August 3 and the others on August 1 and July 29, if
laying on separate days is assumed.

Nest 3 had one egg and one young when it was discovered on August 12.
This egg never hatched. The eyes of the young bird opened the following day,
which would make it about 3 days old when first discovered, putting hatching
at August 9 and the incubation start at July 30 for this egg.

It is revealing to review historical statements concerning the incubation
period of the cuckoo. The duration is currently quoted in most references as
14 days for both the yellow-billed and black-billed cuckoos. This erroneous

VoL. XXXII] HAMILTON and HAMILTON: ARIZONA CUCKOOS 415

figure has been repeated in scores of state bird compendia and other references.
The first reference to the incubation period of the yellow-billed cuckoo was
by Bendire (1895): “Incubation, I think, lasts about 14 days. .. .” This
qualified statement was enlarged upon by Burns (1915) who did not cite his
source but indicated that all the sources of his information were “reliable.”
He gave the incubation period for the yellow-billed and black-billed cuckoos
as 14 days, without qualification. It is apparent that later workers have derived
their figures from this source or from one another.

The incubation period of the black-billed cuckoo was determined in 1943 by
Spencer as 11 days. It thus seems likely that the incubation period of both
North American cuckoos is 10 or 11 days.

BEHAVIOR AT THE NEST. The only persistent pattern to the incubation
position was that the bird never faced the heart of the tree, but always oriented
at least generally away from the trunk. The long tail poses a considerable
concealment problem during incubation. Held horizontally it would hang over
the side of the nest, and often it is held in such a position. On other occasions
the bird aligned the tail along the nest limb, but this apparently does not exceed
chance. This was true at both nest 1 and nest 2. The posture of the bird at nest
2 differed from nest 1. The incubating bird at nest 2 would often sit with the
tail cocked at a 45° angle with the neck and bill pointing up at the same angle.
Perhaps this was simply an alert position induced by close approach to this
low nest.

Incubating birds did not sit especially close. On August 10 we raised a pole
with a mirror to the 32-foot-high nest 1. The bird, sitting on well-incubated
eggs, flushed when the pole was 2 feet away. At this same nest on August
14, with three newly hatched young, the adult bird did not flush when one of
us climbed to within 5 feet of the nest. Instead, it stood at the edge of the nest,
back to the hot midday sun, wings spread, sheltering the nest. It made no move
to depart until a gesture was made to approach closer. At the more strongly
supported nest 2 no such approach was tolerated at any stage of the nesting
cycle.

Shading the young was noted again on August 13 at 1400. The day was hot
and this crown nest was shaded from the sun only by a thin sprig of willow leaves.
The heat was intense. An adult stood at the edge of this nest, back to the sun,
wings cupped and partially spread, but flew off when the nest was approached
to within 15 feet. By the time the nest was reached, the unprotected young,
pinfeathered bird was panting, mouth continuously open.

DISTRACTION ELEMENTS. A distraction display was noted at nest 1 on
August 10 when we put off an incubating bird. When the bird flushed it dropped
through the open undercanopy of the tree willow in which the nest was located.
The wings were held high over the back, sharply cocked at the carpal joint, and
opened, so that the copper red of the primaries and secondaries flashed. It

416 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

alighted first on a heavy trunk in the upper canopy 18 feet from the ground,
then immediately continued its downward flutter, landing again on the trunk
of the adjacent tree willow, now with the wings held more fully to the side. The
tail was half fanned, the dorsal aspect oriented towards us. This away-facing
posture was held, so that we could view the upper side of the wings and the dorsal
side of the tail. The entire effect was one of a greater show of the coppery-red
plumage in the wings than could have been presented in any other manner.
The limb on which it now perched was nearly on the ground. It cocked its head
towards us, then moved off into the deeper streamside vegetation. A few moments
later we heard a modified ‘‘kowlp” call less than 50 yards away. On other
occasions these same birds remained in the adjacent trees giving repeated
“knocker” calls.

On August 16 the distraction attempts at nest 1 were more vigorous. The
bird dropped almost vertically through the upper stay of the host willow’s
vegetation, all the while holding the wings forward and fanned. The tail was
spread wide, and for the first time the bird gave an audible sound to accompany
the performance, a whining ‘‘mew”’ interspersed with “knocker” calls. Now the
mate appeared, flashing its wings. This second bird gave a modified ‘“‘kowlp”
call. On subsequent visits this mewing call always accompanied the distraction
display.

The birds at nest 2 never performed this sort of distraction. On August 8
the bird flew directly away in a wavering flight when flushed, only 2 or 3 feet
from the ground. The following day it left when we were 12 yards away. As it
departed it dropped low, remaining no more than 3% feet above the level
clear ground under the mesquite canopy. The flight was slightly erratic, the
tail flashed considerable white, and the red in the wings was more noticeable
than in ordinary flight. On this and other occasions the departure flight was
slowed. Once, a bird leaving this nest actually soared for 8 or 10 feet in much
the manner and posture of a sailing nighthawk. On other occasions the wingbeat
was deliberate, the deep strokes could easily be counted, and at least once were
alternate.

At nest 3 the performance closely resembled that at nest 1. Like nest 1 this nest
was in the crown of a sizable tree willow. These birds displayed with the wings
and tail, and were very vocal with “knocker” and occasionally “kowlp” calls.
On August 12, while we were examining their young, one of the birds flew from
perch to perch in the nearby trees giving repeated “knocker” calls at approxi-
mately 10-second intervals.

Distraction seems, therefore, to be adjusted to the environmental situ-
ation. Under the closed canopy of the nest tree at nest 1, a slowed flight directly
away from the nest would have been meaningless to a ground-traveling predator.
At nest 2 this might have induced pursuit.

The distraction elements noted were, then: (1) display of bright plumage

—

VoL. XXXII] HAMILTON and HAMILTON: ARIZONA CUCKOOS 417

features, including the coppery red of the wings and the white in the tail,
(2) sound production, including a special mewing call heard only in this context,
frequent “knocker” calls, and occasional “kowlp” calls, and (3) slowed loco-
motion away from the nest.

The distraction elements noted may be broadly categorized as auditory
and visual. Auditory distraction was noted where a close canopy limited the
opportunity for effective visual displays.

The visual distraction display includes the following elements: (1) the dis-
tracting bird drops almost vertically from the nest position, (2) it moves
away from the vicinity of the nest at the same time, (3) spreading the wings and
tail, and (4) accompanying this visual performance with a mewing call.

During auditory distraction this mewing call is not utilized. Instead, the
bird may (1) emit “knocker” calls at intervals much more frequently than
would ordinarily be produced, along with (2) a similarly accelerated pace of
“kowlp” calls, while (3) circling the potential predator out of sight, but not
moving away.

While yellow-billed cuckoos in most areas nest in fairly deep vegetation, they
are often at the edge of openings. This is particularly true of western yellow-
billed cuckoos which may nest in deep riparian vegetation, but are often close
enough to openings over watercourses to permit an injury-feigning arena. In
the case of the nests reported here, this adaptability to the particular local situa-
tion was prominent. In the case of nests 1 and 3, which were in deep-willow vege-
tation, the response tended toward auditory distraction. Nest 2, in a compara-
tively open area, lacked auditory distraction altogether. At nest 1 the open
understory of the tree was the visual distraction arena. This again emphasizes the
latitude of the response in closely adapting to the immediate situation. The
distraction at this site could not be elicited simply by open space and a clear
view ahead: the tree canopy precluded this visual stimulus.

DEVELOPMENT OF THE YOUNG

Abbreviated incubation is followed by a similarly rapid nestling develop-
ment. The eyes open 3 days after hatching. On the 6th or 7th day the feather
sheaths are methodically pulled off by the nestlings and the whole appearance
of the bird changes from a quilled, dark-skinned lump, to a feathered bird
with brown back and white breast. The tail is still stumpy and takes several
days to grow out, but after the 8th day these birds can easily move away from
the nest and, if forced, will fly to another perch. They are not easily caught.

SOUND PRODUCTION. By the time the young are a day old they begin to
make a persistent buzzing call which remains in the vocal repertoire until
fledging. This call becomes louder during the 2nd and 3rd days, but thereafter
its intensity no longer increases noticeably. Basically a series of very closely

418 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Serr.

spaced clicks, these calls follow one upon the other. When there is more than
one bird in the nest the effect is a continuous insect-like buzz.

In captive young this buzzing is continuous during agitation. With artifical
brooding, i.e., a warm covering, the buzzing becomes less frequent and each
burst is less prolonged, until the sound fades away. This sound accompanies
begging. Nevertheless, it is not necessarily an amplification of the begging re-
sponse, since it is also continued when the young are satiated and not begging.
Nestling black-billed cuckoos from Ithaca, New York, made a very similar
sound, but louder and crisper. There we thought this sound resembled and
perhaps mimicked the solitary paper wasps which make a similar buzz before
exiting their homes. The sound is well adapted to predation situations, since it
is continuous, permitting few location cues, and is faint, audible at only 4 or
Seeet-

PREDATOR RESPONSES. In their first days the young show no awareness of
potential predation by a human intruder. When the eyes open this at once
changes. On August 16 the two older young at nest 1, now aged 4 and 5 days,
hunched down when we appeared a yard away. Their eyes were wide open, but
they remained motionless. At the same time the 2-day-old birds at this nest
betrayed their silence and stillness, rising high and begging vigorously and noisily
when the nest branch was disturbed. The same thing happened the following day.
Lack of uniformity of action is obviously one of the prices paid for the advantage
derived from asynchronous hatching.

By the time the young are a week old the alert posture of the adults, with
tail up and wings down, is attempted by nestlings, and they may try to scramble
away. From their 3rd day the young clutch at the lining or framing material of
the nest when they are picked up and it is difficult to remove them without
damaging the nest. The young at nest 3, about 7 days old on August 17, per-
formed the wings down, tail up, posture and gave a hard rasping squawk. It
was 2 feet distal to the nest on a large, deeply furrowed willow limb. When we
started to pick it up, it dug its claws deeply into the fissures of the willow bark
and hung on tightly with its toenails.

There was no opportunity to follow nestling development more than a day
beyond the sheath-breaking stage in the field. From the fully feathered phase
there is surely a considerable period of additional wing and tail feather
development. At this stage the young weigh only about 20 grams, and they
will reach 50 at maturity. But the spurt in 17 days from the start of incubation
to freedom from the nest is one of the shortest for any bird, precocial or altricial.
Predation might be a selective pressure favoring a rapid development, but there
was no evidence of unusual predation factors in either area, and a more likely
explanation is discussed below under Food.

Vor. XXXII] HAMILTON and HAMILTON: ARIZONA CUCKOOS 419

Foop

From June 12-16, 1963, the river bottom at Sonoita Creek was populated
with cuckoos. At that season they were secretive (see Vocalizations: ‘“kowlp”
call), but the breeding population had apparently arrived. No observations on
feeding behavior were made then and it was possible to evaluate only in general
the availability of cuckoo food items.

In August the longer observation period and the breeding activity of cuckoos
permitted direct observation of feeding behavior and an attempt was made to
determine food items taken.

Foop HABITS OF CUCKOos. The American cuckoo species seem to specialize
on insects of fairly sizable proportion available locally in abundance. In North
America at times other than the breeding season, a comparatively wide variety
of insect and fruit species may be taken. Among the insect prey of the cuckoo
are several species not usually selected by other insectivorous birds. For example,
Audubon (1849) illustrates the yellow-billed cuckoo holding a swallowtail
butterfly (Papilio turnus), and Dawn (1955) noted a black-billed cuckoo
taking an adult monarch butterfly (Anosia plexipus), during the period of fall
migration, then returning to attempt to capture another individual of the same
species. An examination of a large series of stomachs revealed an entire tree
frog in one (Beal, 1897), and Clay (1929) saw one hunt down a tree frog on
the ground and swallow it. Lizards occasionally enter the dietary. Swarth (1929)
found a whole lizard in a yellow-billed cuckoo stomach and we have seen one
take a sizable Sceloporus lizard in central California. Reptiles are staple fare
for many tropical cuckoos, so these observations are, perhaps, not surprising.

But by far the most regular and characteristic category of food items of
cuckoos consists of caterpillars. These the cuckoo takes regardless of whether
they are smooth, hairy, or spiny. In the eastern United States where massive
outbreaks of certain tent-building caterpillars are periodic, it has frequently been
suggested that cuckoos are especially abundant during these irruptions (Clay,
1929; Forbush, 1927; many others). Presumably cuckoos are recruited to areas
of high caterpillar density during outbreaks such as these, suggesting a relatively
nomadic phase during the period of spring arrival. In western river bottoms
there may be a similar dependence upon a small number of caterpillar species
during the breeding season.

Foop IN SOUTHERN CALIFORNIA. Cottonwood and willow bottoms along
permanent freshwater courses are the habitat of the yellow-billed cuckoo in
southern California (Jay, 1911; Hanna, 1937). Along the Santa Ana River and
its tributaries near Riverside, Hanna (1937) found 30 nests over a period of
years, with egg dates extending from May 29 to July 10.

During the first 11 days of June, 1963, we intensively worked the river-
bottom areas along this river in the area where Hanna had been so successful
in locating nests. Eugene Cardiff (personal communication) had heard cuckoos

420 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

in this area in recent years. But in 1963 we were unable to locate a cuckoo.
Available habitat for cuckoos along the Santa Ana River has been greatly
reduced owing to increasing urbanization and water-use changes. However,
considerable stretches of seemingly favorable habitat remain, and most of the as-
sociated species mentioned by Hanna (op. cit.) are still to be found. Since the
cuckoo is notably erratic in local abundance, no firm conclusions concerning its
status here can be made. During this period of field investigation we were
struck by the abundance of a spiny caterpillar (Hemileuca sp.). This caterpillar
reaches a length of approximately 40 mm. before descending to the ground to
pupate. In its early stages of development it is colonial and obvious. It feeds
upon the cottonwood (Populus sp.), and several species of willow (Salix sp.)
which flourish along the permanent sections of this stream. During June these
caterpillars mature. The last instar is solitary and the coloration becomes
lighter, making them comparatively cryptically colored. It seems possible that
the June and July breeding season of the cuckoo, so clearly established by
Hanna’s large series of nests, may be timed to this caterpillar species here.
Most other species in this area, insectivorous and seed eating, complete their
breeding cycle considerably earlier.

NESTS

The three Arizona nests upon which this account is based were examined
entire, then turned upside down and taken apart. Since there was practically no
intertwining of components, nests came apart piece by piece in about the order
of original construction. While these nests were basically similar to those de-
scribed above and to one another, several differences indicate a certain versatility
adapted to the nest site and to available materials.

ANCHORING. Versatile adaptation of the nest to its site was well illustrated
by nest 3. This nest was centered in the uppermost and outermost branches of
a 35-foot tree willow at the fork of a 1-inch branch. The first material laid down
was a series of 51 dead willow twiglets close to the fork. The nest was not cen-
tered over this start, however, but was 65 mm. more distal. It seems that in
order to properly secure the nest this preliminary anchor was constructed. By
comparison, the anchoring and framing materials of nest 2 were indistinguishable.
This nest was also placed in a fork, but the broader branches were larger and
the nest was nearly centered over the fork. The anchoring material of nest 1
could not be examined. In removing it from its treetop position 32 feet over a
stream this material pulled away.

The anchoring branchlets and the framing materials of all nests were all
either mesquite or willow twiglets except for one tamarisk branchlet in nest
3. In nest 3 the first 51 twiglets were measured (table 1). The uniform diameter
and fairly uniform length of these twiglets is probably not simply a reflection
of the size twig which is easily broken off since there were a few which were

——

VoL. XXXII] HAMILTON and HAMILTON: ARIZONA CUCKOOS 421

TABLE 1. Consecutive measurements in millimeters of the first 50 willow twigs laid down
in cuckoo nest number 3.

No. Diam. Len. No. Diam. Len. No. Diam. Len. No. Diam, Len.
ile 1.8 110 14. 1.9 190 DT. 1.8 100 39. 3.4 110
wR 2a 130 15. os) 140 28. Ded 240 40. 2.8 100
or IES 90 16. 2.3 170 29. 2.0 90 41. Phe | 170
4. 1.9 170 7 zal 130 30. Des 200 42. ee) 90
oe I of? 110 18. 2.4 160 Sil, 1.9 60 43. The) 130
6. 2.0 190 19. Dra. 150 322 8 140 44. Dall 120
he AA 110 20. 1.3 80 (Tamarisk Branch) 45. D5) 140
8. 1S 90 Zils 1.9 120 335 Dee 190 46. 2.6 140
9. e/ 120 Don 2.9 180 34. Ns} 160 47. 1.8 70

10. 1.7 150 Zon 1.0 50 5). 1.8 110 48. 2.4 280

Le 1.8 130 24. 1.9 100 36. 1.9 150 49. 15 90

WA 1.8 110 25. 2.0 100 Sila Bl 270 50. 1.9 70

ile 2p 100 26. ef) 160 38. 2.5 270 Sie 1.8 180

considerably larger. In Ohio, Clay (1929) observed a yellow-billed cuckoo
breaking off twigs for nesting material with the bill.

These anchoring branchlets are laid across one another in a random pattern,
some extending well to the side of the main nest as supporting outrigging ex-
tending over the hosting limbs. All of the branchlets used in nest 3 were terminal
tips. The majority were placed with the butt end outward. The outermost anchor-
ing branchlets and the outermost framing pieces above them which did not cross
under the center of the nest were the longest. Several of these long branchlets
were curved, the contour of the curve forming the outline of the nest.

When I attempted to make a similar structure I had considerable difficulty
at the start in keeping it from dropping through the crotch of the nest site. This
was a lesser problem if branched twiglets were used, and the most complete nest,
number 3, was examined with this in mind. Of the 51 twiglets listed in table 1,
9 of the first 10, 6 of the second 10, and 5 of the third and fourth 10’s, and 4 of
the final 10, were branched. Branch 32, of a different material, was not included
in this tally. Evidently cuckoos resolved the problem of having the initial
structure collapse some time ago, and tend to use branched twiglets when start-
ing their nest.

The first item laid down in nest 1 was a bunch of three 90-mm. willow
leaves. This nest, unlike nests 2 and 3, was placed on the top of a limb. A leafy
start would, of course, only be possible at such a site. Above these three willow
leaves were four cottonwood leaves, then a nest frame of willow branchlets
much like nest 3. These leaves appear to have come from the tents of a hairy
caterpillar whiched flourished in the river-bottom cottonwoods during June, 1963.
These communal caterpillars, under the protection of web tents, eat both sides

422 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Serr.

of the surface flesh of cottonwood leaves, leaving the vein structure of the leaf
intact. It was four of these leaves that followed the willow leaves into nest 1.
Their presence in this nest confirms the presence of cuckoos about these tents.
It may also suggest that actual nest construction was initiated, at least in a
preliminary way, by late June or early July. They may have been placed in the
nest along with the willow leaves when nest-site selection was under way and
the actual process of nest construction had not yet started. However, some of
these leaves persisted into August, and could have been obtained then.

FRAMING. Above the anchoring material in nest 3 there were 92 additional
willow twigs. As with the anchoring pieces, the butt ends are most frequently
pointed outward. These pieces are on an average shorter than the anchoring
members and only a few spikes stick out from the nest to give it its pincushion
appearance.

In nest 2 the anchoring material and the framing stuff were indistinguish-
able. In this nest these parts were all mesquite, the species of the host tree, in
spite of an abundance of nearby willows. Perhaps this is more a concession to
concealment than necessity. These larger branches conformed neatly to the host
tree, hiding the nest much more effectively than if it had been made of willows.
These twigs were of much greater diameter than the willow twigs used in nests
1 and 3. There were 137 of these mesquite twigs below the lining cup.

Throughout the anchoring and framing portion of the nest, there seems to
be no indication of weaving. However, the twigs do seem to be poked into place
at the higher levels, and the precise order of insertion was difficult to deter-
mine.

LINING THE NEST. Above the frame the construction materials change
abruptly to lining items. In nest 3 the material from the lining was about 6 mm.
thick and consisted of strips of bark, leaves, and a great number of small twigs
from the river cedar. A count of this latter material did not seem very useful
since it was probably gathered in mouthfuls rather than as individual items.
There were several hundred individual twiglets of this material and several
larger much branched pieces. The whole of this material could have been
gathered very quickly near the nest tree.

In addition there were 7 pieces of stripped bark in this nest, to 260 mm. in
length and 15 mm. wide. The rest of these pieces were less than half as large.
Two of these straps were wrapped over a framing willow twig and pushed back
into the cup material. This was the only evidence of any weaving in any of the
three nests examined.

Additional material in this nest included eight small separate willow leaves,
a single small cottonwood leaf, and four mesquite leaves which had broken apart
to a considerable extent. It seems likely that the mesquite leaves which Brandt
(1951) found in his nest fell apart on drying rather than being stripped as he
suggests.

Vor. XXXII] HAMILTON and HAMILTON: ARIZONA CUCKOOS 423

The lining of nests 1 and 2 was similar but the area of these nests near
Patagonia lacked the tamarisk material. We did not find this plant at Pata-
gonia. The 6 mm. lining of nest 1 started with a flat mat of 6 willow leaves.
These may have been placed fresh, since they were matted quite flat. These
leaves were oriented so that the long axis of the leaves followed the long axis
of the nest. Above these leaves there were additional mesquite leaves (entire
compound leaf) and a number of cottonwood bark strips. These could have
been obtained easily from almost any nearby cottonwood tree since almost all
of these trees had a number of large dead branches with bark stripping away.

The lining of nest 2 was more than twice as deep as in the other two nests.
This lining was clearly stratified. As in nest 1, broad leaves start the lining, in
this case 17 caterpillar-eaten cottonwood leaves. Above these followed a mixture
of broken mesquite leaves, 9 willow leaves, 11 additional pieces of mesquite
twigs all about 50 mm. in length, a branchlet of ash seeds, and a small packet
of unidentified capsular seeds. Finally, there was a lining of several strips of
bark, more than a dozen rootlets, and two oak leaves.

All of the lining above the cottonwood leaves seems to have been added after
the first egg was laid since my notes indicate that on the day the nest was dis-
covered, August 6, it was lined with three cottonwood leaves, and it was only
by the grace of these three leaves that the egg was not visible from below.

OVERALL FEATURES OF NESTS. The greatest overall dimensions of these nests
were as follows: The frame of nest 1 was not measured. The cup was 102 by 275
mm. Nest 2 had a frame 275 X 360 mm. with a cup 84 X 112 mm. Nest 3 was
290 X 365 mm. and had a cup 115 X 140 mm. These dimensions emphasize the
oblong and somewhat unsymmetrical appearance of the nest. From the nest
cup the brooding bird overhangs the cup but only the tail exceeds the frame.
The frame material effectively breaks up the outline of the bird which would
otherwise, in a nest of more modest dimension, be silhouetted against the back-
ground.

BEHAVIOR OF ADULTS RELATIVE TO THE NEST. That this may indeed be a
very adaptive structure is illustrated by an experience with the adults of nest
1. After the eggs hatched in this nest, the nest was shifted to a lower position
in the tree in order to facilitate netting the adults at a later date. The entire
nest was placed on a rimmed wooden platform slightly larger than the anchoring
branchlet lengths. The sitting bird could thus not see through the nest to the
ground. On August 16 at 1545, when I approached the tree to check the progress
of the nest, a stiff wind was blowing, the promise of an oncoming thunderstorm.
The major trunks of the tree, including the displaced nest, were swaying con-
siderably. I was anxious to flush the bird so it would not be startled and perhaps
injure the young. So when I reached the branching trunk 8 feet below the bird, I
banged the trunk and shook it. The bird was unperturbed, and remained on the
nest, with the tail hanging over the edge of the platform. It finally flushed when

424 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

I was only 4 feet below and in sight. On other occasions these birds would leave
when we were over 20 feet away. The advantage of the open nature of the
framing material is demonstrated by this experience. It allows a careful watch
of movement below the nest and at the same time conceals the brooding bird.

NEST BUILDING. Our arrival on August 3 was probably too late to witness
nest construction activity. However, on August 4 at 1545, following our 5
hours of intermittent rain, Clark Ross observed a cuckoo, probably one of the
nest 1 birds, in the top of a mesquite tree along a nearby tributary creek. First
it flew into a mesquite tree and gave the ‘“‘kowlp” call. It then picked off a
short dead leaf from a mesquite tree, dropped this item, did the same with
another leaf, and finally selected a larger dead leaf and flew off with it to the
base of a nearby hill. This would be in the direction of nest 1, which contained
three eggs at that time, with incubation just under way. There are several
possible interpretations of this observation. It is possible (1) that nest improve-
ments continue after incubation has started, (2) that the continuing rain stimu-
lated an attempt to dry or improve the nest lining, (3) that the nest building
tendency had not completely subsided in spite of the recent completion of the
nest, or (4) that the nest material gathering movements were a displacement
activity resulting from some social encounter not sensed by the observer.

INCOMPLETE NESTS AND THEIR INTERPRETATION. The entire lining of the
nest seems to be omitted on certain occasions, while at other times it is less
complete (Hanna, 1937). Since the lining phase of nest construction is a distinct
step, its omission may reflect an inherited trait, perhaps favored in areas of
high predation or moderate climate during incubation.

MIGRATION

In Arizona the yellow-billed cuckoo is strictly a summer resident. The win-
ter quarters for this population are unknown, but probably include the jungles
of South America where the species winters, from Venezuela to Argentina
(ALOU Checklist, 1957).

SPRING ARRIVAL. For a bird as unobtrusive as the cuckoo, migratory schedules
are best established by workers who are in the field continuously through
the spring and after the breeding season. In 1927 Swarth (1929) was in the
vicinity of the Patagonia study area from May 10. He first observed yellow-
billed cuckoos at Patagonia on May 25 and “others were seen and heard several
times during the next few days, and it seemed evident that they were just
arriving from the south.”

Other evidence suggests that cuckoos are on the move later than June 1,
however. At Sycamore Canyon, Arizona, Miller (1950) observed what he inter-
preted as a migratory wave on June 30, 1945. He had been encamped at that
locality 2 days before this species was observed. They were quite vocal for a
day or two, then none were seen.

VoLt. XXXII] HAMILTON and HAMILTON: ARIZONA CUCKOOS 425

At some western localities these cuckoos apparently occupy upland country
prior to actual breeding, invading riparian woodland only when it is time to
breed. In Sonoma County, California, Shelton (1911) states that “this bird
keeps to the higher land, among the oaks and other timber, for a period of two
or three weeks before retiring to the willow bottoms to breed.” And in southern
California Baumgardt (1951) watched a yellow-billed cuckoo eating caterpillars
on manzanita in the San Bernardino Mountains at 5,000 feet on June 2, 1950.
This is a habitat completely different from the lowland riparian habitat occupied
by the species during the breeding season. In the Cape Region of Baja California
the species has been reported in the mountains in midsummer. This may be the
retreat of this species prior to its arrival in lower riparian situations to breed
later in the summer (Brewster, 1902). But other workers have subsequently
searched these mountain areas without obtaining evidence of cuckoo breeding
populations (Van Rossen, 1945).

FALL DEPARTURE. Swarth (1929) continued fieldwork through the period of
fall migration. He notes: “During the last week in August cuckoos were seen
in fair abundance about Patagonia, and in lesser numbers somewhat later, the
last on September 11.” I have examined five specimens taken by his group at
that time, now in the collection of the California Academy of Sciences. A
specimen taken in the Huachuca Mountains, Arizona, August 31, was an adult
as were two birds taken at Patagonia on September 5 and 9. Two additional
specimens taken at Patagonia on August 31 and September 11 were juveniles,
easily identified by the less strikingly patterned tail feathers. It would appear
from this scant evidence that the adults do not precede the departure of the
young by any considerable degree.

DISCUSSION

LIMITING FACTORS, BIOLOGICAL. The overall picture which this study produced
was one of cuckoos breeding at a density considerably below that which the
apparent food supply would permit. It is possible that food, in fact, is not a
factor limiting the abundance of this species, but in Arizona we found no evidence
of other environmental population-limiting mechanisms.

In considering the food supply with respect to the cuckoo population, three
possibilities become evident. First is that breeding densities and spatial

distribution of breeding pairs are adapted to the average or lowest year of food

abundance. In 1963 the caterpillar which formed the major part of the diet of
the young might have been unusually abundant. At the San Pedro River
location there were, in fact, fewer caterpillars than along Sonoita Creek.
Nevertheless, even there they seemed to be superabundant.

Second, what seems to be a superabundant food supply may be basically
marginal to begin with. The caterpillar of the Condalia bush which is the main
item taken to the young, is a relatively small prey item for the cuckoo, and

426 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser.

they are taken one at a time. Since each caterpillar it treated before feeding
to the young or before being eaten by the adults, the labor involved in flying
out to a Condalia bush, obtaining a caterpillar, and returning to the nest, would
be considerable. Thus, it may be that several of the breeding adaptations of the
yellow-billed cuckoo in Arizona are adaptations to time limitations with respect
to this caterpillar.

A third possibility is that the spatial distribution characteristic of the
yellow-billed cuckoo is adapted to other regions of quite different nutritive
characteristic. There is some evidence to support the contention that gene flow
from other areas is significant. The western race of the yellow-billed cuckoo
is poorly defined and additional races from either Mexico or other regions have
not been suggested. In other parts of North America the spatial requirement
for successful breeding is probably greater. If the relatively small populations
in Arizona are swamped by gene flow from these cuckoo populations, the
yellow-billed cuckoos of Arizona might have no opportunity to evolve spacing
mechanisms locally adapted to a particularly favorable food resource. It seems
reasonable, therefore, to suggest that the density regulation mechanism of the
Arizona populations may, in fact, reflect the requirements of the species
population center in the eastern United States.

LIMITING FACTORS, PHYSICAL. Possibly the location of nest sites is restricted
to river bottoms because of humidity requirements for successful hatching
and rearing of the young, regardless of the proximity of food. In the San Pedro
River locality there are numerous stock ponds which have large cottonwoods
and willows about them. Some of these ponds have permanent water.
Near Saint David, Arizona, yellow-billed cuckoos were seen about these ponds
in mid-August. While no nests were found, it seems likely that the species
breeds there. Except for these ponds, no cuckoos were seen other than in the
immediate vicinity of river bottoms. Along Sonoita Creek no cuckoos were
noted above the region of permanent water. These observations suggest that
permanent water or some environmental factor closely correlated with it are a
basic requirement for the yellow-billed cuckoo. At the San Pedro River locality
the Condalia shrub and its caterpillar extended several hundred yards beyond
the river bottom. Cuckoos flew out to these areas to feed, but nests were
apparently all located along the river bottom. The denser cover in the river
bottom would not seem to be the sole factor dictating this choice since nest
2 at Sonoita Creek was in a mesquite tree, a few yards from the river bottom.
Mesquite is the predominant shrub in the area for a considerable distance
beyond the river along the San Pedro. The nest along the San Pedro River
is on interesting example with respect to this hypothesis. Only 30 yards from
the tree willow which supported this nest the humidity became strikingly lower.
The extremely arid region adjacent to the river-bottom area was consistently
less humid.

VoL. XXXII] HAMILTON and HAMILTON: ARIZONA CUCKOOS 427

This suggestion again considers the genetic adaptation of the species as a
whole throughout North America. The species population center in the eastern
United States is concentrated in the deciduous forests and meadows which are
consistently humid during the early summer breeding season of the species
there. River bottoms in the west could be easily adapted as geographic barriers
were penetrated. However, the utilization of a breeding terrain strikingly
drier than that used by the species as a whole might be precluded by the lack
of an adaptation permitting successful egg hatching under these conditions.

ADAPTATIONS TO SEASONAL BREEDING. Compared with most temperate region
songbirds, the yellow-billed cuckoo has an exceptionally long breeding season
in the eastern part of its range. In the southeast and West Indies breeding begins
in April, but may be delayed until June in the northen part of the range. In the
southeastern United States the species may be double brooded, but this has only
been inferred from the discovery of a progression of nests not necessarily by the
same individual, as the season progresses. In every part of the species range
exceptionally late nests have been located. Extremes are a nest with two young
on September 9 in Missouri (Adams, 1933), incubated eggs in Alabama on
August 11 (Golsan and Holt, 1914), eggs as late as August 15 in New Jersey
(Harlow, 1918), four eggs in a nest in Illinois on September 5 (Hess, 1910),
eggs in Alabama on August 10, 11, and 14 (Holt, 1925), fresh eggs in Florida
on August 11 (Williams, 1904), and eggs in Michigan on August 27 (Swales,
1903). The latest date appears to be for the black-billed cuckoo, which was
found sitting on four fresh eggs on September 14 in Michigan (Barrows, 1912)
and with young in New York on September 10 (Bendire, 1895). These extremes
point to the fact that in the breeding range as a whole reproduction may occur
almost any time the species is present. This wide range of the reproductive
season suggests at least a partial role of environmental control of reproduction.
In addition it suggests a preadaptation to taking advantage of an abundant
food source when it becomes available at a season not in phase with the usual
breeding season. In the case of the yellow-billed cuckoo, however, the range of
variability of the eastern populations includes the adaptive season in Arizona,
and adaptation to this area would have been possible by selection from the
genetic pool of the species: no new mutations would be necessary.

The wide range of breeding seasons presents an interesting problem with
respect to the basis of their timing. If we accept Lack’s (1954) hypothesis that
the breeding season of altricial birds is timed to the maximum abundance of
food, a well-supported hypothesis in general and one which seems to be supported
in the case of these several populations of cuckoos, then the regulation of the
timing of the breeding season presents a particularly interesting problem for
this species. Several aspects of the breeding cycle in southern Arizona and
southern California are compared in figure 2. The difference in breeding season

428 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

of these two populations could be based either upon local adaptations of
mechanisms which permit the initiation of egg laying in anticipation of the
maximum food supply or actual timing by the appearance of a satisfactory
food supply.

The tent-building caterpillars in Arizona which were present in June
provide interesting evidence with respect to these alternatives. If the mere
presence of a rather abundant favorable food species triggered reproduction,
this comparatively small outbreak could conceivably have triggered a flurry of
breeding at a very inappropriate time. Yet breeding bypasses this false
opportunity. During the latter rainy season in July and August when the flush
of breeding occurs, not only are caterpillars available but alternate food
resources such as other caterpillar species, grasshoppers and beetles would provide
at least a partial substitute should the caterpillar crop fail.

It would be reasonable to suggest that breeding was timed by the seasonal
rains here only if one assumed that this timing is a local population
characteristic, since the nearby southern California populations breed at a
season when there is no likelihood of any rain during their breeding season. And,
lacking any evidence of large-scale genetic adaptations to local environments,
it seems too much to expect that behavioral characteristics would be so different
in these nearby populations.

Several features of the breeding cycle point to the possibility of adaptation
to exogenous regulation. In particular, the tremendously abbreviated incubation
period and rapid nestling development point in this direction. However, this
could be an equally effective adaptation to a genetically timed cycle which
takes advantage of the food supply available for only a brief period of time. If
the clutch size of the Arizona population is actually reduced, as it seems to be,
this could again be construed as evidence for genetic adaptation to a sharply
resticted period of adequate food supply for the young.

ADAPTATIONS OF THE SPECIES. The field observations reported here and
the general ecological situation suggest several aspects of the overall adaptation
of American species of cuckoos to their niche not previously discussed elsewhere.
A part of these imply specialization, but in general these adaptations seem rela-
tively broad. In comparison with other species of similar size, the yellow-billed
cuckoo is usually less abundant. This probably reflects the role of the cuckoo as
a predator on sizable prey species.

A major adaptation which the yellow-billed cuckoo has made is with respect
to a seasonally available, and comparatively abundant, source of food which
has not been fully utilized by other species. In particular, the hairy tent-
building caterpillars in the eastern portion of the range and certain species of
spiny caterpillars which are rejected by other species are favored food items of
the cuckoo. Combined with this predilection to secure food items which are
not utilized by other species is an ability to time the breeding season to local

Vor. XXXII] HAMILTON and HAMILTON: ARIZONA CUCKOOS 429

SOUTHERN CALIFORNIA #
ARRIVAL DEPARTURE =
<2 Zz
= —
WwW zy
= 2
ue WL
o Fz
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a ro
5 af
Fa ae
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5 158) 25 45 15g eeZo 5 sn 75S sy 25) 5 la 7 5
May June July August September October
DATE FIRST EGG HATCHES
SOUTHERN ARIZONA ”
ARRIVAL DEPARTURE eS
eB z
rf rr
Zz PATAGONIA meas =
(46 YR. : <
u (40 YR. AVG.) 9 (1S VR. AVG.) fh
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5 15a 25 5 15 25 5 a) 25 5 1525 5 15? S252. 55
May June July August Se ptember October
DATE FIRST EGG HATCHES

Ficure 2. Breeding characteristics of yellow-billed cuckoos in southern California and
southern Arizona. These dates are compiled from the literature, collections, and original
observations. Five days were added to egg dates unless stage of incubation could be deter-
mined. The number of nests for southern California reflects the greater amount of study
directed to cuckoos in that area. The departure dates are approximate and it is obvious
that birds which hatch during the week of September 5 in Arizona can not leave that early.
This discrepancy reflects inadequate data and a span of departure times; the entire population
does not leave at once. Weather data are from U. S. Weather Bureau records.

conditions of food abundance. The observation of Bendire (1895) of cuckoos
carrying a large number of sizable crickets (Anabus) to nestling young
emphasizes that the yellow-billed cuckoo is adaptable, and not, at least as a
' species, restricted to a limited diet.

TIMING OF MIGRATION. In spite of the sharply different breeding seasons
of the southern California and southern Arizona breeding populations, migration
is accomplished by both populations at approximately the same time (fig. 2).
In the case of the Arizona population this takes the species to the breeding
grounds well in advance of the breeding season. There seem to be two plausible
explanations for this characteristic.

First, it is possible that the time of migration of the species is adapted to

430 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER

anticipate the season of maximum opportunity for the species as a whole. Since
a relatively small part of the whole breeding population is breeding in a place
with the late summer thunderstorm ecology of Arizona and Mexico, the fixed
timetable of the vast majority of the population may swamp any tendency to
adapt the migratory schedule more closely to the period of maximum food
abundance.

Another possibility is that the premature arrival at the Arizona breeding
locality simply reflects the necessity to vacate feeding grounds on the winter
quarters which become inadequate, or at least less favorable, compared with
what is available at the prebreeding Arizona environment.

INCUBATION PATTERN. Audubon (1849) was shown a yellow-billed cuckoo
nest at Charlestown, South Carolina, which had two feathered young able to
fly and three additional young of different sizes. In addition, the nest contained
two eggs, one containing an embryo, the other fresh. None of the young were of
the same size. Based upon the comments of the discoverer of the nest, Audubon
felt that laying in the same nest continued over a prolonged period of time.
It is now apparent, however, that the eggs of the American cuckoo species are
laid in clutches. Incubation apparently begins soon after the first egg is laid.
Thus, the young hatch either daily or every other day. Perhaps occasionally a
longer span may separate hatching dates. This staggered incubation pattern
is characteristic of owls, some hawks, coots, and a number of other species. It
has often been suggested that the adaptive significance of this pattern is to
insure that at least a part of the clutch receives adequate food. Thus, the first-
born will, with even a minimum amount of food, be vigorous and obtain food
at every nest visit by the parent. During times of food shortage the last young
to hatch will be neglected and will not limit the survival potential of the first-
born.

An alternative hypothesis is suggested by the limitations which seem to
be imposed on cuckoos by the nature of their caterpillar food supply which
they depend upon to such a great extent. If the amount of food which can be
delivered to the young is restricted by time, i.e., the time required to fly out,
obtain a caterpillar, return to the nest, treat the caterpillar, and deliver it to
the young, rather than being limited by the overall abundance of food, then a
staggered hatching sequence would be of considerable advantage. This would
permit a greater number of trips for those young at a stage with maximum food
requirements and would extend the period of productive food gathering.

The early fledging of the young and their especially early departure from
the nest are also adaptive in connection with the considerable pretreatment
which each caterpillar receives before it is ingested. If the young mature to
a state of being able to move to the caterpillar source and do their own food
manipulation at an early stage, a considerable time economy would be achieved.

Vout. XXXII] HAMILTON and HAMILTON: ARIZONA CUCKOOS 431

BIBLIOGRAPHY

ApAMs, I. C., Jr.
1933. Late nesting of the yellow-billed cuckoo in Missouri. Auk, vol. 50, p. 107.

AMERICAN ORNITHOLOGISTS’ UNION
1957. Check-list of North American birds. Published by the Union.

AUDUBON, J. J.
1849. Ornithological biography. Vol. 5. Edinburgh, Adam and Charles Black. 664 pp.
Barrows, W. B. .
1912. Michigan bird life. Michigan Agricultural College. Department of Zoology and
Physiology. Special Bulletin. xiv + 822 pp.

BauMGarnt, J. H.
1951. Yellow-billed cuckoo in the San Bernardino Mountains, California. Condor, vol.
53, p. 207.

BEAL, F. E. L.
1897. Some common birds in their relation to agriculture. U. S. Department of
Agriculture Farmers’ Bulletin no. 54, p. 40.

BENDIRE, C.
1895. Life histories of North American birds. Smithsonian Institution, U. S. Museum
Special Bulletin 3.
Bent, A. C.
1940. Life histories of North American cuckoos goatsuckers, hummingbirds and their
allies. U. S. National Museum Bulletin 176. viii + 506 pp.
Branpt, H.
1951. Arizona and its bird life. The Bird Research Foundation, Cleveland, Ohio. xvi
+ 725 pp.
BREWSTER, W.
1902. Birds of the Cape Region of Lower California. Harvard University. Bulletin
of the Museum of Comparative Zoology, vol. 41, pp. 1-241.
Burns, F. L.
1915. Comparative periods of deposition and incubation of some North American
birds. Wilson Bulletin, vol. 22, pp. 275-286.

Cray, M. B.
1929. The yellow-billed cuckoo. Bird Lore, vol. 31, pp. 189-190.
Dawy, W.
1955. Black-billed cuckoo feeds on monarch butterfly. Wilson Bulletin, vol. 67, pp.

133-134.
Forsusu, E. H.
1927. Birds of Massachusetts and other New England states. Part 2. Norwood
Press, 1 + 461 pp.
Gotsan, L. S., and E. G. Hort
1914. Birds of Autauga and Montgomery counties, Alabama. Auk, vol. 31, pp. 212-235.
Hanna, WItson C.
1937. California cuckoo in the San Bernardino Valley, California. Condor, vol. 39,
pp. 57-59.
Hartow, R. C.
1918. Notes on the breeding birds of Pennsylvania and New Jersey. Auk, vol. 35,
pp. 18-29.

432 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Hess, I. E.
1910. One hundred breeding birds of an Illinois ten-mile radius. Auk, vol. 27, pp.
19-32.
Horr, Ey. |G:
1925. Early shooting and some late breeding records for Alabama. Auk, vol. 42,
pp. 147-148.
Jay, A.

1911. Nesting of the California cuckoo in Los Angeles County, California. Condor,
vol. 13, pp. 69-73.

ILI, 1D).
1954. The natural regulation of animal numbers. Oxford, 343 pp.
Murer, L. H.
1950. Lifelong boyhood. University of California Press, Berkeley and Los Angeles.

ix + 226 pp.
SHELTON, A. C.
1911. Nesting of the California cuckoo. Condor, vol. 13, pp. 19-22.
SPENCER, O. R.
1943. Nesting habits of the black-billed cuckoo. Wilson Bulletin, vol. 55, pp. 11-22.
Swates, B. H.
1903. A list of the land birds of south-eastern Michigan. Part 1. Bulletin of the Michigan
Ornithological Club, vol. 4, pp. 14-17.
SwartTH, H.S.
1929. The faunal areas of southern Arizona: a study in animal distribution. Proceedings
of the California Academy of Sciences, 4th ser., vol. 18, pp. 267-370.
Van Rosse, A. J.
1945. A distributional survey of the birds of Sonora, Mexico. Occasional Papers
Museum of Zoology, Louisiana State University, no. 21.
WILiiaMs, R. W., Jr.
1904. A preliminary list of the birds of Leon County, Florida. Auk, vol. 21, pp. 449-
462.
Wutson, A., and C. L. BONAPARTE
1878. American ornithology, Volume 1. Porter and Coates, Philadelphia. cxxxii + 214
pp.

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

G Dallas Hanna Anniversary Volume

Volume XXXII, No. 15, pp. 433-446; 11 figs. October 24, 1966

THE LARVAL DEVELOPMENT OF CHITONS
(AMPHINEURA)
By
Allyn G. Smith

Associate Curator, Department of Invertebrate Zoology
California Academy of Sciences, San Francisco 94118

Most species of chitons, as indicated by previous studies, develop in much
the same manner as gastropod mollusks. Normally, the sexes are separate and
during spawning periods the sex products are extruded into the surrounding
sea water, fertilization being largely a matter of chance. The fertilized ova
divide rapidly and in a period of a few hours hatch into free-swimming larval
stages consisting of ciliated trochophores. In a few days to a week or more
the trochophore starts to take on the form of an adult, developing a creeping
foot and the beginnings of the shell structure. At this stage a larval chiton
is said to “metamorphose” and moves to the bottom to carry on further
development as a young individual with all eight plates or valves completely
formed. While this is the usual procedure, there are exceptions upon which
the purpose of this paper is intended to focus particular attention.

Because the literature on chiton development is scattered, a brief enumera-
tion of studies made by both earlier and later workers in this field will be
helpful. Although the spawning habits of chitons and their embryological and

_ subsequent larval development have been reported by relatively few investi-
gators, some of the earlier work has been thorough for the particular species
studied.

Earlier workers dealt with /schnochiton cinereus (Linné), a fairly common
small chiton with a distribution in British and Scandinavian seas and extend-
ing into the Mediterranean. Some applied other species names because taxonomic
relationships were not well understood at the time. Clark (1855), Lovén (1856),
Garnault (1888), Schweikart (1905), Knorre (1925), Hoffman (1931), and

[ 433 ]

Marine Biotogtcal Laboratory
LIBRARY

NOV2 1966
WOODS HOLE, MASS.

434 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Thorson (1946) all worked with Ischnochiton cinereus. Other European students
of chiton development include Kowalevsky (1833), who dealt with Chiton poli
Philippi |= Nuttallina (Middendorffia) cinerea (Poli, 1791)|; Plate (1899—
1901), who reported on the embryology of Nuttallochiton hyadesi (Roche-
brune); Lyngnes (1924), who studied both /schnochiton cinereus and Lepido-
pleurus asellus (Spengler); Thiele (1906), who investigated Notochiton
mirandus (Thiele); and Hammarsten and Rundstrom (1925), who did an
excellent piece of research on Acanthochitona discrepans (Brown). Christiansen
(1954) reported on the life history of Lepidopleurus asellus. Her paper on the
methods used and the results achieved represents not only an important
contribution to the subject of chiton development, but also contains by far
the best review of the subject to date.

Heath (1899) started the study of New World chitons with a detailed
investigation of the development of /schnochiton magdalenensis (Hinds) [=
Stenoplax heathiana Berry]. He also studied the breeding habits and develop-
ment of Lepidozona mertensi (Middendorff), LZ. cooperi (Dall), Mopalia mus-
cosa (Gould), Katharina tunicata (Wood), Trachydermon raymondi (Pilsbry )
| = Cyanoplax dentiens (Gould)|, Nuttallina thomasi Pilsbry, and the giant
chiton, Cryptochiton stelleri (Middendorff), all relatively common species in the
vicinity of Stanford University’s Hopkins Marine Station, where Heath lived
and worked.

Metcalf’s work (1892-93) on the Caribbean species Chiton squamosus
Linné and C. marmoratus Gmelin was followed by that of Crozier (1918) on
Chiton tuberculatus Linné, type species of the genus Chiton. Grave (1932)
reported on the development of the western Atlantic species, Chaetopleura
apiculata (Say); Okuda (1947) also worked with Cryptochiton stelleri. The
most recent published report is by Thorpe (1962), who made a study of the
spawning habits of a number of California species, including specific infor-
mation on the larval development of Mopalia ciliata (Sowerby).

The investigations of all workers mentioned cover a total of 19 species,
in 14 presently recognized genera, grouped into 7 families, which is a fairly
representative coverage of the order Polyplacophora. Nearly all of the chitons
studied have a developmental pattern consistent with that already briefly
sketched. In a few instances, however, exceptions to the normal pattern have
been found to occur.

Kowalevsky in 1883 and Plate in 1898 noticed deviations in the develop-
ment of the larval stages based, respectively, on studies of Jschnochiton
cinereus and I. imitator (E. A. Smith). Heath, reporting in 1905, called

attention to similar deviations as a result of his work with Cyanoplax dentiens
and Nuttallina thomasi, saying these species:

. carry their young until they are in an advanced trochophore stage. These, to
the number of about 200 in the case of Trachydermon [= Cyanoplax], are held

VoL. XXXIT] SMITH: LARVAL DEVELOPMENT OF CHITONS 435

Ficure 1. Hemiarthrum setulosum (Dall). Female from below showing metamorphosed
young in the branchial cavities. Length, about 10 mm. (After von Martens and Pfeffer.)

Ficure 2. Same. Enlarged sketch of a single metamorphosed young animal. (After von
Martens and Pfeffer.)

Ficure 3. Callistochiton viviparus Plate. Enlarged sketch of a metamorphosed young
specimen showing seven valves. (After Plate.)

Ficure 4. Schizoplax brandti (Middendorft). Enlarged sketch of a female from below
showing metamorphosed young in the branchial cavity. (After Kussakin.)

FicureE 5. Same. (5a) Embryo at the stage of formation of the shell plates. (5b) Shell
plates already formed but not fully developed. (5c) Young individual showing the divided
intermediate shell plates and at a stage when ready to leave the mantle cavity of the mother.
Length, about 0.75 mm. (After Kussakin.)

436 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

in the mantle cavity on each side of the foot. Plate speaks of their extending over
the neighboring regions of the mantle and to some extent on the foot, but in the
present case this removal of the eggs from the mantle cavity never occurs until the
animal has been disturbed... .

Nuttallina thomasi, which breeds during the summer, carries its eggs in two rod-
shaped masses, each approximately 80 eggs, held in the mantle cavity on each side
of the foot.

Commenting on the dioecious condition in chitons, Heath (1907) found in
his work with the development of Cyanoplax dentiens that this species is
normally hermaphroditic and that the young “are brooded by the parent as in
the case of Chiton polii [= Nuttallina cinerea], Ischnochiton imitator, and
two or three other species.’

Iredale and Hull (1923, p. 191) state that: “One of us took specimens of
Heterozona subviridis at King Island, Bass Strait, with clusters of eggs
disposed along the outer margin of the foot.” Dell (1962), while examining
specimens of /schnochiton constanti (Vélain) collected by the William Scoresby
off Gough Island, Tristan da Cunha Group, in 40 to 60 fathoms, noted that
“one specimen had a clump of eggs fixed to the branchial groove on each
Sicleperaw

Martens and Pfeffer (1886) apparently were the first to observe a further
deviation from the normal whereby the eggs and the hatched larvae were
retained in the mantle cavity through metamorphosis, including the develop-
ment of fully formed shell plates, a phenomenon they termed brutpflege. The
species in this instance was Hemiarthrum setulosum Dall from Antarctic seas.
In his discussion of this species in the Manual of Conchology (1892, pp.
19-21) Pilsbry does not mention its larval development. His figure 1 on plate
5 of the Manual is a 1:1 reproduction of Martens and Pfeffer’s original
figure, which shows the ventral side of a female specimen with metamorphosed
young arranged along the sides of the foot under the girdle. This would have
been made clear had Pilsbry used the original caption for this illustration.
Also, Pilsbry did not include the enlarged drawing of a metamorphosed young
specimen of H. setulosum supplied by Martens and Pfeffer. In consequence,
both original illustrations are included herein as figures 1 and 2.

Plate, in 1898, was the next to report on a chiton actually brooding its
young through metamorphosis. He also commented on the possibility that
some might be viviparous. He named no species at that time but in his
comprehensive work on the South American chitons contained in the Fauna
Chilensis, published in 1902, he described Callistochiton viviparus as a new

‘Heath mentions his success in raising the young of Cyanoplax dentiens and Nuttallina thomasi to sexual
maturity by removing adults to isolated tide pools where the development of the young could be observed
with facility over a considerable period of time. Christiansen used the method followed by Grave (1937) to
induce larval development, moving the larvae to clean dishes with filtered sea water. Seven days after fer-
tilization the larvae were supplied with the diatom WNitschia closterior as food. but her efforts to continue
development of the young chitons after metamorphosis were unsuccessful.

VoL. XXXIT] SMITH: LARVAL DEVELOPMENT OF CHITONS 437

species from Isla de Pacheros, 12 nautical miles from Coquimbo, Chile. His
figure of the young stage after metamorphosis is reproduced in figure 3.

In 1910, Thiele added another species to the deviate list, Svpharochiton
nigrovirens (Blainville). Barnard (in Ashby, 1931) notes that: ‘Thiele refers
to the presence of young under the mantle (girdle) edge. The same fact was
observed in the case of specimens collected by me at Smitswinkel Bay, False
Bay, in July 1912. The young are about 0.75 mm. in length.” Dell (1962)
states that a specimen of S. nigrovirens from Saldhana Bay, South Africa,
has fully developed young clustered under the girdle edge, about 33 on the
left and 22 on the right side. The Spencer Thorpe Collection has a series of
62 specimens, preserved dry, from Kleinmond, South Africa, 20 of which have
similar metamorphosed young. One specimen measuring 16.9 mm. in length
has a total of 42, 23 on the right and 19 on the left side. The females bearing
young range in length from about 9.5 to 18.5 mm., the young themselves being
about 0.7 mm. long.

Another report of a chiton brooding its young was made in 1960 by Dr.
O. G. Kussakin of the Zoological Institute of the USSR, Department of
Hydrobiology, Leningrad. In this instance the species is the peculiar Schizoplax
brandti (Middendorff), in which the intermediate shell plates are separated
along the median line by a narrowly triangular wedge of cartilaginous material
similar to that forming the hinge of a bivalve. This species occurs from the
Okhotsk Sea, along the Aleutian Islands, and south on the Alaskan and
Canadian side of western North America to at least as far as Graham Island
off the coast of British Columbia. Kussakin’s work was based on a year-
round study of the development of S. brandti in the southern Kurile Islands
in 1954-55. Because his paper may not be readily available to western
specialists the somewhat revised English summary of it follows:

A peculiar feature of the biology of S. brandti is the complete lack of a plankton
larval stage; the whole development proceeds in the mantle (gill) cavity of the
female from which the juveniles emerge with a completely formed shell resembling
the adults. In this connection the development of S. brandti is considerably altered
and simplified. Comparison of the development of this species with data available
in the literature on other species indicates a trend in some chiton families toward a
transition from a more complex to a more direct developmental process and toward
viviparity.

Dr. Kussakin’s drawings are indicative of the development of larval Schizo-
plax and they are reproduced with his permission as figures 4 and 5 of this
report. Also, Dr. Kussakin has kindly furnished specimens in alcohol of
S. brandti from the Kurile Islands, which contain both trochophore larvae and
metamorphosed young. These have been deposited in the collection of preserved
invertebrates in the Academy’s Department of Invertebrate Zoology.

438 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Mr. Spencer Thorpe informs me in connection with his study of the breed-
ing habits of California chitons that he has found Nuttallina thomasi to be
another species that mothers its young through metamorphosis, thus adding
a fifth species to this category. Although Heath reported this species as carry-
ing its eggs in the mantle cavity he made no mention of the stage at which
the larvae emerged or were released from beneath the mantle.

To the five chiton species that follow a special development pattern, a
sixth, may now be added, the particulars regarding which follow:

During a collection trip to Guadalupe Island, Mexico, in 1946, Mr. Wood-
bridge Williams made an interesting collection, principally in tide pools, of
chitons and other species of mollusks (Smith, 1963). Among these was a
large series of a small chiton that provides ample evidence of a propensity
for mothering its young. Although original preservation of the specimens was
in alcohol, unfortunately this particular lot was allowed to dry out. Careful
microscopic examination of them disclosed that of 71 specimens, minute, white,
perfectly-formed metamorphosed young were lying between the ctenidia and
the side of the foot in 21 of them. Some of the specimens had from three to
six young adhering to the dried-out animals, all with at least seven fairly
well formed valves and an extremely narrow girdle with an outer fringe of well
developed spicules. Although somewhat curled because of their dried-out condi-
tion, approximate measurements show a length of 0.25 to 0.35 mm., which is
considerably smaller than for comparable young specimens of Sypharochiton
nigrovirens owing, no doubt, to the much smaller size of adults of the Guadalupe
Island species. In size and other characters they resemble a specimen figured by
Christiansen for an early stage of Lepidopleurus asellus (fig. 6).

Examination under higher magnification of some of the Guadalupe Island
specimens with metamorphosed young showed as many as six or eight additional
roundish bodies between the sides of the foot and the gill rows. Undoubtedly
these are trochophore larvae in various stages of development. One of them,
removed after careful softening, showed some of the characters of a typical
trochophore larva of L. asellus at the end of eight or nine days’ development,
with three ciliary processes or flagella still intact projecting from the anterior
ence (iigan7s)*

At first, the above evidence seemed impelling reason to describe this unusual
little chiton species as new. However, in discussing the situation with Dr. S.
Stillman Berry of Redlands, California, he mentioned finding very young
examples under the girdles of his Lepidozona asthenes, making a check with
his species imperative. It happens that I collected the original lot of ZL.
asthenes in 1916 at White’s Point, Los Angeles County, California, consisting
of about 70 specimens. The Academy’s collection has 28 specimens from this
type lot, enough to make an adequate comparison with the series from
Guadalupe Island. Observation under a magnification of X 90 showed six

VoL. XXXII] SMITH: LARVAL DEVELOPMENT OF CHITONS 439

with metamorphosed young still adhering to the under sides of the girdles in
spite of the fact that in many specimens a crude attempt at removing the foot
and the underlying viscera had been made.

Further comparison between the two series of specimens (Smith, 1963)
led to the conclusion that the Guadalupe Island series unquestionably belongs
to L. asthenes based on the criteria of valve sculpture and configuration, and
on the relative size, placement, and striated nature of the girdle scales. _

Still more evidence that this species mothers its young through meta-
morphosis was discovered recently while studying a series of small chitons
taken off the coast of southern California by Dr. William E. Ritter and
Professor William J. Raymond in the summer of 1901 under the auspices of
the San Diego Marine Biological Association (a predecessor of the Scripps
Institution of Oceanography). Among the various lots was one containing 18
specimens of L. asthenes collected intertidally at White’s Point, the type
locality. Three of these also had metamorphosed young beneath the mantles;
one still has eight of them plainly visible on both sides of the foot of the dried
animal, along with nine or ten unmetamorphosed larvae.

DEVELOPMENT OF CHITON SHELL PLATES

Although perhaps ancillary to the main subject of general chiton develop-
ment in the early stages, the manner in which the shell plates form is of con-
siderable interest. A number of accounts have been published about this but
the reports of two or three specialists will serve to review our knowledge of
the process.

Dall, writing in 1879 (p. 291), says: “The valves are first irregular, but
increase from below, and deep notches, persistent in the adult, are formed
on the front edges, one on each side. It will be seen that the valves are formed
each in one piece, and by the coalescence of parts corresponding to the various
areas of the adult valve. There are eight valves in all Chitons .. .” (fig. 8).

Heath’s more detailed account (1899, Thesis, pp. 65-66) is as follows:
The first clearly marked indications of the shell [in Stenoplax heathiana| occur
usually about the fourth or fifth day when a band of somewhat clear cells appears
parallel with and a little posterior to the prototroch. Very soon after another ap-
pears a short distance behind the first and the process is continued until seven such
bands alternate with six narrow rows of darkly staining cells. . . . When treated
with methyl green they stain intensely and this reaction continues as far as I have
traced the development of the shell, when these cells come to occupy positions be-
tween the valves of the shell. The narrow bands of cells apparently but little dif-
ferentiated which alternate with these mucous (?) cells gradually increase in breadth
. and ultimately the calcareous portions of the shell appear above them.

The calcareous salts are deposited in the cuticle covering the region of the shell
and the presence of the tegmental sense organs (aesthetes) makes it evident that the
first part of the shell to form is the tegmentum. The articulamentum appears later
but the manner in which it forms is as yet unknown.

440

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VoL. XXXII] SMITH: LARVAL DEVELOPMENT OF CHITONS 441

Kowalevsky (’83) has accurately described the formation of the calcareous portions

of the shell and Jschnochiton [i.e. Stenoplax] affords no further important points on

this subject, (figs. 10, 11).

In Lepidopleurus asellus during metamorphosis, Christiansen says the
rudiments of the shells extend into the dorsal region of the cephalic portion
of the developing larva and the calcareous plates themselves appear very
irregularly, each plate being composed of three parts. The eighth shell plate
is the last to appear. In a nearly two-month-old larva the eight calcareous
plates cover the back and appear to be well developed.

Thus, from these accounts, the formation of the first six shell plates seems
reasonably clear, allowing for possible minor differences between species.
There appears to be no specific information, however, on just how and when
the last two (valves vii and viii) are formed. Plate’s illustration of the meta-
morphosed larva of Callistochiton viviparus (fig. 3, herein) shows but seven
complete shell plates. Likewise, Kowalevsky’s figure of a well developed larva
of “Chiton polii’”’ shows seven (fig. 9), while his figure of a “young form”
shows eight, with the eighth just beginning to develop. Christiansen says the
tail plate (valve viii) of Lepidopleurus asellus is the last to appear. In my
detailed examination of the metamorphosed young of both Sypharochiton
nigrovirens and Lepidozona asthenes, however, the line of demarcation between
the last two is extremely difficult to make out. For L. asthenes, in fact, some
of the young appear to have only seven valves at the stage of development
when they were collected, valve vii not being well developed.

SUMMARY AND OBSERVATIONS

To sum up from a survey of published accounts, the chiton species of the
following list have been observed to deviate from the normal pattern of larval

Ficure 6. Lepidopleurus asellus (Spengler). Enlarged sketches of dorsal and ventral sides
of a nearly two-month-old specimen after metamorphosis. (After Christiansen.)

Ficure 7. Same. Enlarged sketches of dorsal and side views of a trochophore larva at
8°%4 days. (After Christiansen.)

Ficure 8. Ischnochiton cinereus (Linné). (8a) Dorsal view of larva, the valves begin-
ning to be formed. (8b) Same. Ventral view, showing foot, and eyes at the sides of the
head. (8c) Older individual, showing diminished size of the anterior tuberculate lobe, or
head. (Dall, after Lovén.)

Ficure 9. Nuttallina (Middendorffia) cinerea (Poli, 1791). Enlarged sketch of young
form showing seven valves in process of formation. (After Kowalevsky.)

Ficure 10. Stenoplax heathiana Berry. Much enlarged sketch of young stage at end of
fourth day, showing fusion of the three portions of the valves. (After Heath.)

Ficure 11. Same. Young stage fifteen days after hatching, with metamorphosis complete.
(After Heath.)

442 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

development, with the females depositing their eggs in the branchial cavity and
retaining them there at least to the trochophore larval stage:

SPECIES FAMILY AUTHORITY

Ischnochiton cinereus (Linné) Ischnochitonidae Thorson (1946) and others

i" imitator (E. A. Smith) " Plate (1897-1901)

" subviridis (Iredale and May) " Iredale and Hull (1923)

" constanti (Velain) " Dell (1962, 1964)
Cyanoplax dentiens (Gould) " Heath (1905)
Nuttallina cinerea (Poli) Callistoplacidae Kowalevsky (1883)
Chiton barnesi Gray Chitonidae Plate (1897-1901)

Species that mother their young through the final stage of metamorphosis,
including the development of fully-formed shell plates are:

SPECIES FAMILY AUTHORITY
Hemiarthrum setulosum Dall Lepidopleuridae Martens and Pfeffer (1886)
Lepidozona asthenes (Berry) Ischnochitonidae This report
Schizoplax brandti (Middendorff) Schizoplacidae Kussakin (1960)
Callistochiton viviparus Plate Callistoplacidae Plate (1902)
Nuttallina thomasi Pilsbry Callistoplacidae Thorpe (unpubl.)
Sypharochiton nigrovirens (Blainville) Chitonidae Thiele (1910); Barnard (in

Ashby, 1931) ; Dell (1962) ;
this report.

The geographical distribution of these species, taken as a group, is world-
wide. It does not conform to any particular pattern. Neither do their presently
assigned systematic positions follow a pattern as the family-groups represented
range from the more primitive (Lepidopleuridae) to species considered to have
a high evolutionary development (Chitonidae).

The fact that a small number of totally unrelated species of chitons are
now known to follow a type of larval development that deviates from the
normal process indicates that studies of other species, whose life histories
are unknown, should be investigated. Additionally, for some of the chiton
species listed above, we need to be sure that the reported condition character-
izes a species as a whole and is not merely an aberrant situation exhibited by
a single or a few individuals induced, perhaps, by unusual or abnormal
ecological conditions or other circumstances. Obviously, the possibilities for
such studies are enormous when one considers that the total Recent chiton
fauna of the world comprises something like 500 described forms. An interesting
point, at present apparently undescribed, would be to determine specifically
when and how the trochophore larvae of some species escape from the branchial
cavity of the female as free-swimming veligers before metamorphosis occurs.
Of equal interest would be the circumstances surrounding the eventual escape

of the metamorphosed larvae of those species that mother their young through
this stage.

VoL. XXXIT] SMITH: LARVAL DEVELOPMENT OF CHITONS 443

This brief account of chiton development is dedicated with the respect
and sincere good wishes of the author to his long-time friend and close asso-
ciate, Dr. G Dallas Hanna, whose helpful encouragement and sound advice
have been a constant source of inspiration over many years of personal contact.
Such dedication is all the more fitting in view of Dr. Hanna’s many contribu-
tions to the Academy’s Recent mollusk collection over a long period, including
the large suites of chitons he has collected from Pt. Barrow, Alaska, to Cape
San Lucas at the tip of the Baja California peninsula. .

Appreciation is due to Mr. Spencer R. Thorpe, Jr., of El Cerrito, California,
for loaning his set of Sypharochiton nigrovirens for study, and to Mr. Maurice
Giles, California Academy photographer, for his preparation of the illustrations.

AsHBY, EDWIN
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1879. Report on the limpets and chitons of the Alaskan and Arctic regions, with de-
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GRAVE, BENJAMIN H.
1932. Embryology and life history of Chaetopleura apiculata. Journal of Morphology,
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1937. Culture methods for invertebrate animals. Chaetopleura apiculata, pp. 519-520.

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HicLey, Rose M., and Harotp HeatH

1912. The development of the gonad and gonoducts in two species of chitons. Biologi-

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VoL. XXXII] SMITH; LARVAL DEVELOPMENT OF CHITONS 445

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1946. Reproduction and larval development of Danish marine bottom invertebrates,
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Plankton, vol. 4, pp. 1-523.

PROCEEDINGS
CALIFORNIA ACADEMY OF SCIENCES

G Dallas Hanna Anniversary Volume

Volume XXXII, No. 16, pp. 447-513; 65 figs. October 24, 1966

REVISION OF THE NEARCTIC SPECIES OF SILIS
(CANTHARIDAE: COLEOPTERA)
By

John Wagener Green
California Academy of Sciences, San Francisco, California, 94118

The latest revision of the Nearctic species of Silis was published in 1918
by Van Dyke, who recognized 19 named taxa in the subgenus Sis and 8 in
the subgenus Ditemnus. Since then only 2 new ones have been added, by
Fender and Brown. Van Dyke separated the species solely on the basis of the
pronotal armature of the males. It has been found that a number of his so-
called species of Silis in the strict sense are actually evolutionary complexes
depending almost entirely on the male genitalia for the identification of their
components. A preliminary analysis of this totally unexplored field is the
primary objective of this study. It has proved to be a taxonomic problem of
very great difficulty.

The genus Silis, as at present constituted, is defined, in addition to the
longitudinally divided eighth ventral segment common to all Silini, by the
excised and modified lateral margins of the male pronota. The ramifications
of this structure are so elaborate and varied that no rigid statement can be
made that would include all of them. In general, the region of the posterior
angles is excised, retracting these angles, and forming an angular prominence
in front; while the excision is occupied by a posterior process arising from the
hypomera. The angular prominence will be designated hereinafter as the
anterior process. It is strictly marginal, and the term does not apply to any
prominence or elevated area not extending to the lateral margin. Very little
external structural diversity occurs elsewhere in the body.

Two subgenera are found in the Nearctic fauna: Si/is (strict sense) and
Ditemnus. Plectonotum, of the Leng catalogue, is not a subgenus of Silis.
The Arizona species assigned thereto by Schaeffer is a member of the Discodon-
Polemius association, with numerous related forms in the Mexican fauna.

Marine Biological Laboratory.
LIBRARY
‘NOV 2 1966

WOODS HOLE, MASS.

448 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Silis (strict sense) occurs in Europe and Canada, and throughout most of
the United States, but apparently not south of the Mexican border. Ditemnus,
except for two species in eastern United States, is of southern distribution:
Texas to California; and south through Mexico, and Central and South America.
The propriety of assigning subgeneric status to Ditemnus is seriously questioned,
but no change in the generic structure of the Silini is proposed at this time.
A new, and perhaps significant, differential is given in the subgeneric key
that follows. The species are quite constant in color pattern and structure,
and are easily identified.

The male genitalia of Ditemnus are small, usually fragile, and structurally
difficult to visualize. They have not been investigated in this study. In the
genitalia of Silis (strict sense), as in most of the Cantharidae, the aedeagus
arises within a largely sclerotized tegmen that has been considered a modifi-
cation of the basal piece. It is suggested that the tegmen is an invaginated
ninth abdominal segment that is visible externally only in the exposed tip of
ventral segment nine, sometimes called the genital segment. In support of this
contention, it should be noted that ventral segment 9 is attached to the under-
side of the tegmen near the base, and no dorsal or pleural parts of segment
9 are present, other than the tegmen itself. Comparing this to the structure
of segment 9 in the related families Lampyridae and Lycidae, it is seen that
in these two families ventral segment 9 unites with a dorsopleural portion to
form a homogeneous structure completely inclosing the aedeagus.

The upper surface of the tegmen, designated hereinafter as the dorsal plate,
is membranous except for a sclerotized distal portion. Its posterior margin is
usually emarginate, with the emargination more or less unstable intraspecifically,
and rarely entirely lacking. Sometimes the emarginate area is abruptly inflexed
so the notch is posterior and does not show from a direct dorsal viewpoint.
Lateral incisures separate a ventral lobe from the main body of the tegmen.
The ventral lobe is not prominently emarginate, usually rounded or subangulate
apically, or with at most a small median notch. In several specimens a deep
and narrow emargination has been noted in the ventral lobe, supposedly
fractures resulting from pressure. The tegmen is subject to considerable intra-
specific variability in many of the details of its conformation, and differential
characters derived therefrom cannot be rigidly interpreted.

In all the species there arise from the inner base of the tegmen, above
the aedeagus, strongly sclerotized dual processes that may be separated through-
out; or united basally, separating at or beyond the middle of their length;
or completely united to form a single median process, or sometimes a broad
plate. These processes, or basal apophyses, will be designated hereinafter as
basophyses. They are directed backward and upward, their tips usually attain-
ing the underside of the dorsal plate, or sometimes passing through its
emargination. It is impossible to get a direct or completely unobstructed view

VoL. XXXII] GREEN: REVISION OF THE NEARCTIC SILIS 449

of these structures, so their delineation in the drawings is more or less
diagrammatic.

The parameres of the aedeagus are of two radically different types, which
would seem to constitute subgeneric division except that no external differentials
are available. In one type, exemplified by S. difficilis, the parameres extend in
a partially sclerotized process along either side of the median lobe nearly to
its apex, and appearing to constitute an integral part thereof. In the second
type, exemplified by S. cava, the parameres are shorter and visible, if at all,
only toward the base of the median lobe, but are provided each side with a
long and usually slender sclerotized process extending to or beyond the tip
of the median lobe. These processes, or lateral apophyses, will be designated
hereinafter as the laterophyses. Apparently they are capable of a limited
rotary motion around a longitudinal axis, thereby altering their appearance
from a fixed viewpoint.

Complete eversion of the internal sac of the aedeagus is highly desirable in
cantharid studies, but this situation is rarely found in cabinet specimens.
Drying distortion and varying degrees of expansion produce varying aspects
of the median lobe, and these are responsible for most of the major difficulties
encountered in a taxonomic analysis. The author has endeavored to limit the
announcement of new species to those taxa in which the genitalia are
positively distinctive; and to those more closely related forms in which
apparent differences in the median lobe seem to be conclusive, provided that
other structural characters and geographic isolation also are involved. A
difficult situation, constantly recurring in all the evolutionary complexes,
is the necessity of deciding whether an observable difference is due to normal
intraspecific variability, or to valid interspecific differentiation. Such decisions
are herein based on a study of sizable series, and on the general experience
gained from endless observations throughout the genus. It seems likely, how-
ever, that final solutions will be found only when a practical method is dis-
covered for everting the internal sac of the aedeagus in dried specimens.

In extracting the genitalia, it has been found impractical to try to press
them out through the anus, except for recently collected individuals in a
perfectly relaxed condition. The specimens must be thoroughly relaxed, three
days in a very wet relaxing dish is not too much time for older ones. Abdominal
segments 6 to 9 are removed, and dorsals 6 and 7 are opened with a chisel-
pointed needle. Water should be applied as needed, to prevent drying and to
act as a lubricant. By prying with the needle and pressing on both sides with
the tweezers, the genitalia will usually come out readily, backwards. It is
essential not to apply pressure along the median line because of the danger
of breaking the fragile processes of the aedeagus. It is the author’s practice
to mount the genitalia upright on a paper point, together with the detached

450 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

abdominal part, and to pin this below the specimen. Upright mounting insures
an unobstructed dorsal, lateral, and ventral view.

In addition to the extensive collections of the California Academy of
Sciences (indicated as CAS), valuable material and other assistance was re-
ceived from the following institutions and individuals, to all of whom the
author extends his sincere thanks and appreciation. The abbreviation symbols
in parentheses are used in the text to indicate the present location of certain
specimens.

(ANSP)—Academy of Natural Sciences of Philadelphia, H. J. Grant, Jr.
(AMNH)—American Museum of Natural History, J. G. Rozen, Jr.
(CNC)—Canadian National Collection, H. F. Howden.
(CU)—Cornell University, Henry Dietrich.

(LAM )—Los Angeles County Museum, R. R. Snelling.
(MCZ)—Museum of Comparative Zoology, P. J. Darlington, Jr.
(OSU)—Ohio State University, J. N. Knull, P. H. Freytag.
(OrSU)—Oregon State University, J. D. Lattin.

(SDNH)—San Diego Natural History Museum, C. F. Harbison.
(USNM)—U. S. National Museum, T. J. Spilman.
(UBC)—University of British Columbia, G. J. Spencer.
(UCB)—University of California at Berkeley, Jerry Powell.
(UCD)—University of California at Davis, A. T. McClay.
(UCR)—University of California at Riverside, Saul Frommer.
(UId)—University of Idaho, W. F. Barr.

(UK)—University of Kansas, G. W. Byers.

(UW )—University of Washington, M. H. Hatch.

Jane C. Dirks-Edmunds.

J. Gordon Edwards.

Kenneth M. Fender.

Gayle L. Nelson.

Joe Schuh.

P. H. Timberlake.

Vladimir Vosyka, Czechoslovakia.

All the keys that follow apply to males only. No attempt has been made
at this time to identify females of the subgenus Silis. Usually no trouble will
be encountered in placing Ditemnus females. The length-width ratio given in
some species descriptions is intended as a rough indication of the body form.

The head, usually deflexed, is not included in length measurements, which
are more or less approximate.

Key TO NEARCTIC SUBGENERA OF SILIS

Hypomera completely divided by a more or less undefined concavity or depression that
follows a production of anterior margin of posterior process, the front part flat or con-

Vou. XXXII] GREEN: REVISION OF THE NEARCTIC SILIS 451

vex, punctate and pubescent. Notch formed by longitudinal division of ventral segment

8 closed, its sides parallel and largely contiguous along median line _.. __. Ditemnus LeConte
Hypomera not so divided, smooth, shining, and virtually glabrous, longitudinally concave.

Notch formed by longitudinal division of ventral segment 8 open, V- or U-shaped

psoeressceeruse eS ee ee eee eee Silis Charpentier

SUBGENUS DITEMNUS

The Nearctic species of this subgenus divide into three well defined
categories, which should all be accorded generic rank if that status is given to
Ditemnus. Number I, containing two species, is Ditemnus LeConte (1861),
with D. bidentata Say as the genotype. Category number II, numerously
represented in the Neotropical fauna, and number III, largely Nearctic, are
at present nameless. Two new species of the third category have the modified
tarsal claws of the male finely cleft, while in all the other species of Nearctic
Ditemnus these claws are appendiculate. Cleft claws have also been noted in
the Cuban D. marginella DuVal. This alone is not regarded as a character
of generic weight. The punctate and pubescent hypomera characteristic of
this subgenus occur also, predominantly, in the Discodon-Polemius association,
and have been noted in the Palearctic Silas ruficollis Fabricius. One of the
new species, S. howdent, will no doubt be otherwise disposed when the generic
structure of the Silini is revised.

Key To NEARCTIC SPECIES OF THE SUBGENUS DITEMNUS, MALES

1. Form shorter and broader. Antennae stout. Disk of pronotum with median impression

2
Form elongate. Antennae slender, subfiliform. Disk of pronotum without median im-
JOHESSI Ol mpe ean enone patos oes TI he AE et ee te ee 7
2. Disk of pronotum with shallow subreniform impression, deeper each side. Antennae
subserrate. Fourth segment of front tarsi normal. Eastern and midwestern species
ee aa er Pe Se a a ge a 3)
Disk of pronotum with deep median fossa. Antennae subfiliform. Fourth segment of
front tarsi larger than usual, subquadrate. Southwestern species —. 4
Category I
3. Posterior process of pronotal armature arising in advance of hind angles, which are
subrectangular and rather well defined = (1) S. (D.) bidentata Say
Posterior process of pronotal armature broader, overlapping hind angles which are
roundedrand) indistinct) = eee NS EE Ee ee (2) S. (D.) latiloba Blatchley
Category II
Pemelecdteraleborder Olelytra testaceOus) 2. 5
Hl voram IDI COlOLOUS MD la Cke semeeeee ae eee eee ey RUE ewe | Pe) OR 6
PeESTcal border ol elytra) testaceous' =.= (3) S. (D.) freemani Brown
Sutural border of elytra not testaceous (4) S. (D.) fossiger LeConte
6. Pronotum pale, or with median dark area (5) S. (D.) tricornis Van Dyke

Pronotum pale with all borders black (6) S. (D.) obtusa LeConte

452 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Category III

7. Posterior process of pronotal armature continuous with base of pronotum, obscuring
hind an 2] @sy eee 1248
Posterior process arising in front of hind angles, which are well detmed and sub-
T@CtAN GUAT <9... e S — 9

8. Body entirely black. Pronotal incisure feeble. Modified tarsal claws Pe romneniate
RET ee. SR ee ee (7) S. (D.) howdeni Green, new species
Pronotum pale, incisure normal. Modified tarsal claws cleft
are eo En (8) S. (D.) vandykei Green, new species

Oe Body: entirely black- 4) 23 "ee (9) S. (D.) nigerrima Schaeffer
Pronotum pale, abdomen rufous or partly so)= ae 10
10. Antennal vestiture erect and bristling —-____________ (10) S. (D.) perforata LeConte
Antennal vestiture decumbent -_..___—._- - see 11

11] Mees black. Modified tarsal’claws cleft) Bastem Texas eee
get atte AP aE len Ee, Set Bem | RE RP ws RN EEN et (11) S. (D.) knulli Green, new species
Legs partly pale. Modified tarsal claws appendiculate. Arizona __--_.___

(1) Silis (Ditemnus) bidentata Say.

Cantharis bidentata Say, 1825, Jour. Acad. Nat. Sci. Philadelphia, vol. 5, p. 169.
Silis lepida LeConte, Dejean Cat., 3rd ed., p. 121 (nomen nudum).

Black, prothorax rufous, apical borders of ventral segments pale. Varies
with head and abdomen more or less pale.

Length-width ratio about 2.2 : 1. Antennae stout, subserrate, about three-
fourths as long as body, intermediate segments about one and one-third times
as long as wide, vestiture short and decumbent. Pronotum as in figure 1P,
posterior process narrower than in S. latiloba, arising in advance of hind
angles which are subrectangular and rather well defined; disk with subreniform
median impression deepest each side. Anterior claw of front tarsi with blunt
exterior basal appendix, posterior claw simple; middle tarsi similar, appendix
of anterior claw small and inconspicuous; hind tarsi with both claws simple.
Length 3-3.5 mm.

DISTRIBUTION. MAINE, NEW HAMPSHIRE, MASSACHUSETTS,
RHODE ISLAND, CONNECTICUT, NEW YORK, PENNSYLVANIA,
OHIO, NEW JERSEY, MARYLAND, DISTRICT OF COLUMBIA, VIR-
GINIA, WEST VIRGINIA, TENNESSEE, NORTH CAROLINA, SOUTH
CAROLINA, GEORGIA, ALABAMA, MISSISSIPPI, FLORIDA, and
OKLAHOMA (the last possibly an error).

(2) Silis (Ditemnus) latiloba Blatchley.
Silis latiloba BLATCHLEY, 1910, Coleop. of Indiana, p. 837.

Black, prothorax rufous, apical borders of ventral segments pale. Varies
with scutellum rufous; abdomen paler.

VoL. XXXII] GREEN: REVISION OF THE NEARCTIC SILIS 453

Length-width ratio about 2.2 : 1. Antennae stout, subserrate, about three-
fifths as long as body, intermediate segments about one and one-half times
as long as wide, vestiture short and decumbent. Pronotum as in figure 2P,
posterior process broad, overlapping hind angles which are rounded and
indistinct; disk with shallow subreniform median impression deepest each side.
Anterior claw of front tarsi with blunt exterior basal appendix, posterior
claw simple; middle tarsi similar, appendix of anterior claw small and _ in-
conspicuous; hind tarsi with both claws simple. Length 3.7-5 mm.

DISTRIBUTION. KENTUCKY, OHIO, MICHIGAN, INDIANA, ILLINOIS,
IOWA, WISCONSIN, MINNESOTA, MISSOURI, ARKANSAS, KANSAS,
NEBRASKA, NORTH DAKOTA, TEXAS, MANITOBA.

(3) Silis (Ditemnus) freemani Brown.

Silis freemani Brown, 1940, Canadian Ent., vol. 72, p. 163.

Antennae and tips of palpi black. Head black posteriorly, pale in front.
Pronotum and scutellum pale rufous. Elytra black; sutural borders narrowly,
and lateral borders more widely, pale; pale color not extending around apex.
Underside rufous anteriorly, metathorax black, abdomen largely black. Legs
black, anterior femora pale basally; trochanters and coxae pale, the posterior
more or less blackish.

Length-width ratio about 2.5: 1. Antennae rather stout, about two-thirds
as long as body, intermediate segments about twice as long as wide, vestiture
short and decumbent. Pronotum as in figure 3P, with broad median fossa,
median line impressed or excavated basally. Elytral pubescence somewhat
longer and more erect distally. Anterior claw of front tarsi with blunt exterior
basal appendix, all other claws simple. Tarsi stout, fourth segment of front
tarsi larger than usual, subquadrate. Length 4-5 mm.

DisTRIBUTION. TEXAS: Cameron County: Brownsville; Hidalgo; Laredo;
Mission.

Specimens from Linares, Nuevo Leon, Mexico, identified as Silis biau-
riculata Champion, seen to be conspecific, differing externally in the backward
extension of the anterior pronotal process being less developed. The genitalia
of Mexican and Texas specimens seem to be similar, but they are so constructed
that a comparison is difficult and inconclusive. H. S. Barber recognized this
‘species as new to the United States fauna. He attached “glicki” type labels to
USNM material, but did not publish on this. His specimens were intercepted
at Brownsville and Laredo on agricultural imports from Mexico.

(4) Silis (Ditemnus) fossiger LeConte.
Silis fossiger LECONTE, 1881, Trans. Amer. Ent. Soc., vol. 9, p. 58.

Antennae and tips of palpi black. Head black posteriorly, pale in front.
Pronotum pale rufous, scutellum black. Elytra black, lateral borders narrowly

454

CALIFORNIA ACADEMY OF SCIENCES

[Proc. 4TH SER.

VoL. XXXII] GREEN; REVISION OF THE NEARCTIC SILIS 455

pale. Underside rufous, metathorax black, abdomen more or less dusky medially.
Legs black, front coxae pale.

Length-width ratio about 2.5: 1. Antennae rather stout, about three-
fourths as long as body, intermediate segments about twice as long as wide,
vestiture short and decumbent. Pronotum as in figure 4P, disk with broad
median fossa. Anterior claw of front tarsi with blunt exterior basal appendix,
all other claws simple. Tarsi stout, fourth segment of front tarsi larger than
usual, subquadrate. Length 4-5 mm.

DIsTRIBUTION. TEXAS: Devils River, Del Rio, Pleasanton.

This species is very much like the Mexican S. dilacerata Gorham, but the
genitalia do not agree. It is probable that they are members of a more extensive
complex of Neotropical species. A third one, from Colombia, is in the collection
of the California Academy of Sciences.

i
Ficures 1P-30P. Pronota of Silis species.

Ficure 1P. Silis (Ditemnus) bidentata Say.
FicurRE 2P. Silis (Ditemnus) latiloba Blatchley.
Ficure 3P. Silis (Ditemnus) freemani Brown.
FicuRE 4P. Silis (Ditemnus) fossiger LeConte.
Ficure 5P. Silis (Ditemnus) tricornis Van Dyke.
FicureE 6P. Silis (Ditemnus) obtusa LeConte.
Ficure 7P. Silis (Ditemnus) howdeni Green.
Figure 8P. Silis (Ditemnus) vandykei Green.
Ficure 9P. Silis (Ditemnus) nigerrima Schaeffer.
Ficure 10P. Silis (Ditemnus) perforata LeConte.
FicureE 11P. Silis (Ditemnus) knulli Green.
Ficure 12P. Silis (Ditemnus) abdonvinalis Schaeffer.
Ficure 13P. Silis (Silis) spinigera LeConte.
Ficure 14P. Silis (Silis) rugosa Van Dyke.
Ficure 15P. Silis (Silis) percomis Say.

Ficure 16P. Silis (Silis) spathulata LeConte.
Ficure 17P. _—Silis (Silis) lutea group.

Figure 18P. = Silis (Silis) filicornis Van Dyke.
Ficure 19P. = Silis (Silis) filicornis variation.
Ficure 20P. Silis (Silis) tardella Green.

Ficure 21P. Silis (Silis) difficilis group.

Ficure 22P. Silis (Silis) atra LeConte.

Ficure 23P. Silis (Silis) deserticola group.
Ficure 24P. Silis (Silis) emarginata Green.
Ficure 25P. Silis (Silis) cava complex.

Ficure 26P. Silis (Silis) arizonica Van Dyke.
Figure 27P. Silis (Silis) fenestrata Van Dyke.
Ficure 28P. Silis (Silis) recta Green.

Ficure 29P. Silis (Silis) vulnerata group.

Ficure 30P. — Silis (Silis) pallida group.

456 CALIFORNIA ACADEMY OF SCIENCES | Proc. 4TH SER.

(5) Silis (Ditemnus) tricornis Van Dyke.
Silis tricornis VAN DyKe, 1918, Jour. New York Ent. Soc., vol. 26, p. 177.

Antennae and palpi black. Head black posteriorly, pale in front. Pronotum
entirely pale, or with median black area. Scutellum and elytra black. Under-
side black, prothorax pale. Legs black, trochanters pale.

Length-width ratio about 2.3: 1. Antennae rather stout, filiform, about
three-fourths as long as body, intermediate segments about two and one-third
times as long as wide, vestiture short and decumbent. Pronotum as in figure
5P, disk with broad median fossa. Anterior claw of front tarsi with blunt
exterior basal appendix, all other claws simple. Tarsi stout, fourth segment
of front tarsi larger than usual, subquadrate. Length 4.5 mm.

DISTRIBUTION. CALIFORNIA: Calipatria; Thousand Palms. ARIZONA:
Bill Williams Fork, Globe.

(6) Silis (Ditemnus) obtusa LeConte.

Silis obtusa LeConteE, 1874, Trans. Amer. Ent. Soc., vol. 5, p. 62.

Body black, pronotum pale rufous with all borders narrowly black, apical
margins of ventral segments pale.

Length-width ratio about 2.4: 1. Antennae stout, subserrate, about seven-
tenths as long as body, intermediate segments about twice as long as wide,
vestiture short and decumbent. Pronotum as in figure 6P, with broad median
fossa. Anterior claw of front tarsi with small blunt exterior basal appendix,
all other claws simple. Tarsi stout, fourth segment of front tarsi larger than
usual, subquadrate. Length 4.5—5 mm.

DIstRIBUTION. CALIFORNIA: Bakersfield; Camp Baldy (San Bernardino
Mountains); /ndian Canyon, San Diego County; Inyo County; Keeler; Lake
Henshaw, San Diego County; Mojave Desert; Oil City, Kern County; Olanche,
Los Angeles County; Oro Grande; Pasadena; Riverside; Saboda Springs,
Riverside County; Victorville; Warners; Waterman Canyon.

(7) Silis (Ditemnus) howdeni Green, new species.

HoLotyPeE, male; Southwestern Research Station, Portal, Arizona, VI-17-56,
H. and A. Howden. In Canadian National Collection.

Body and appendages entirely black.

Length-width ratio about 3.0: 1. Antennae moderately slender, filiform,
about seven-tenths as long as body, intermediate segments about two and
one-half times as long as wide, vestiture erect and bristling. Pronotum as in
figure 7P, base arcuate each side, region of hind angles not excised, disk with
shallow undefined median impression extending from base nearly half way
to apex. Anterior claw of front tarsi with blunt exterior basal appendix,

Vor. XXXII] GREEN: REVISION OF THE NEARCTIC SILIS 457

posterior claw simple; middle and hind tarsi similar to front, with basal appen-
dix progressively smaller. Length 4.5 mm.

VariATIons. Nothing of importance noted. Length 3.5—-4.5 mm.

DISTRIBUTION. ARIZONA: Chiricahua Mountains, Southwestern Research
Station, Portal, H. and A. Howden, VI-17-56, holotype (CNC); VII-7-56,
1 paratype (CAS). Cochise County, Southwestern Research Station, 5 mi. W.
of Portal, 5,400 ft., V-19-56, M. Statham, 1 paratype; V-31-56, E. Ordway,
1 paratype (AMNH).

This species, lacking the excision of the hind angles of the pronotum, and
apparently without a definite posterior process, probably requires a new
genus for its reception. It is assigned here tentatively, pending clarification of
the confused generic conceptions involving Silis, Ditemnus, Polemius, and
Discodon.

(8) Silis (Ditemnus) vandykei Green, new species.

Hototyre, male; Jeff Davis County, Texas, VII-4-53, D. J. and J. N.
Knull. In collection of Ohio State University.

Antennae and palpi black. Head black, mandibles and adjacent area dusky
rufous; beneath pale rufous, black each side of gula. Pronotum and scutellum
pale rufous. Elytra black. Ventral surface of meso- and meta-thorax black;
abdomen dusky, lateral and apical borders of segments irregularly pale. Legs
black.

Length-width ratio about 2.6: 1. Antennae slender, filiform, about three-
fourths as long as body, intermediate segments about three times as long as
wide, vestiture erect and bristling. Pronotum as in figure 8P, base arcuate
each side obliterating hind angles, disk without distinct median impression.
Anterior claw of front and middle tarsi finely cleft, parts of divided tip parallel
and subequal in length; anterior claw of hind tarsi more widely cleft, tooth
much shorter than apical part; all teeth exterior. Posterior claw of all tarsi
simple. Length 4.5 mm.

VARIATIONS. In one example the head in iront of the antennal sockets is
entirely pale rufous. Length 3.5—5 mm.

DISTRIBUTION. TEXAS: Jeff Davis County, VI-20-52, D. J. and J. N.
Knull, 3 paratypes (OSU 2, CAS 1); VII-4-53, D. J. and J. N. Knull, holo-
.type, female (OSU). Davis Mountains, V1-24-56, VI-14-56, D. J. and J. N.
Knull, 2 paratypes (OSU). Ft. Davis, VI-1-59, Howden and Becker, 2 para-
types (CNC). Davis Mountains, 1X-3-49, Werner-Nutting, 1 paratype (Fend-
er). Chisos Basin, Big Bend, VII-16-56, H. and A. Howden, 1 female
(Howden). Big Bend National Park, Chisos Basin: Boat Springs, 7,000 ft.;
Pine Canyon, 5,000 ft.; and Green Gulch, 5,300 ft., V-4-59 to V-29-59,
Howden and Becker, 20 paratypes, 4 females (CNC), 2 paratypes (CAS).
Chisos Mountains, VII-18, J. W. Green, 1 paratype, 4 females (CAS);

458 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

VI-20, 21-61, VII-28-62, D. J. and J. N. Knull, 5 paratypes, 5 females (OSU),
1 paratype (CAS).

(9) Silis (Ditemnus) nigerrima Schaeffer.
Silis nigerrima SCHAEFFER, 1908, Jour. New York Ent. Soc., vol. 16, p. 66.

Body entirely black except mandibles pale.

Length-width ratio about 2.8 : 1. Antennae slender, as long as body, inter-
mediate segments about four times as long as wide, vestiture erect and bristling.
Pronotum as in figure 9P. Anterior claw of front and middle tarsi with blunt
exterior basal appendix, anterior claw of hind tarsi with appendix much
reduced; posterior claws of all tarsi simple. Length 6-6.5 mm. Closely re-
sembling the Mexican S. armitage Pic.

DistrRIBUTION. ARIZONA: Chiricahua Mountains: Pinery Canyon; Onion
Saddle; Rustlers Park. Huachuca Mountains: Carr Canyon; Ramsey Canyon;
Miller Canyon. Santa Rita Mountains: Madera Canyon. Santa Catalina
Mountains: Bear Wallow; Summerhaven, Pine County. White Mountains.

(10) Silis (Ditemnus) perforata LeConte.
Silis perforata LEConTE, 1881, Trans. Amer. Ent. Soc., vol. 9, p. 57.

Antennae and palpi black. Head black posteriorly, pale in front. Pronotum
and scutellum pale rufous. Elytra black. Underside black except prothorax and
abdomen rufous, varying with abdomen dark medially. Legs black, front
coxae pale.

Length-width ratio about 3.3: 1. Antennae slender, filiform, about three-
fourths as long as body, intermediate segments about four times as long as
wide, vestiture erect and bristling. Pronotum as in figure 10P, backward
extension of anterior process overlapping posterior process, hind angles dis-
tinct. Anterior claw of front tarsi with blunt exterior basal appendix, posterior
claw simple; middle and hind tarsi similar to front, with basal appendix pro-
gressively smaller. Length 5—6.5 mm.

DISTRIBUTION. TEXAS: Davis Mountains; Cypress Mills; Big Bend
National Park, Chisos Basin; Kerrville; Austin; Gillespie County; Comal
County; Randall County, Palo Duro Canyon; Bexar County, Fort Sam
Houston, and Salada Creek; Real County, 26 mi. N. of Leakey; Rock Springs.

(11) Silis (Ditemnus) knulli Green, new species.

Ho.otypPe, male; Lake Corpus Christi, Texas, III-26-53, D. J. and J. N.
Knull. In collection of Ohio State University.

Antennae and palpi black. Head black posteriorly, pale in front. Pronotum
and scutellum rufous. Elytra black. Underside of metathorax and tip of abdo-
men black, balance of ventral surface pale rufous. Legs black, front and middle
coxae pale.

VoL. XXXII] GREEN: REVISION OF THE NEARCTIC SILIS 459

Length-width ratio about 2.3: 1. Antennae slender, filiform, about three-
fourths as long as body, intermediate segments about three times as long as
wide, vestiture short and decumbent. Pronotum as in figure 11P, similar to
S. perforata, backward extension of anterior process shorter, scarcely over-
lapping posterior process; hind angles distinct. Anterior claw of front and
middle tarsi finely cleft, parts of divided tip parallel, subequal in length;
anterior claw of hind tarsi more widely cleft, tooth much shorter than apical
part; all teeth exterior. Posterior claw of all tarsi simple. Length 4 mm.
Closely resembles S. perforata.

VariATIons. Nothing of importance noted. Length 3.5-5 mm.

DISTRIBUTION. TEXAS: Corpus Christi and Lake Corpus Christi, I-17
to III-30, 1952 and 1954, D. J. and J. N. Knull, holotype, 39 paratypes (OSU
30, MCZ 1, CAS 8), 24 females (OSU 20, CAS 4). Cameron County, II1-24-60,
D. J. and J. N. Knull, 6 paratypes, 3 females (OSU).

(12) Silis (Ditemnus) abdominalis Schaeffer.
Silis abdominalis SCHAEFFER, 1908, Jour. New York Ent. Soc., vol. 16, p. 66.

Antennae black, first segment largely pale; tips of palpi black. Pronotum
and scutellum rufous. Elytra black. Underside rufous, metathorax black,
abdomen variably dusky baso-medially. Legs rufous, tibiae in part black.

Length-width ratio about 2.7: 1. Antennae moderately slender, filiform,
about seven-tenths as long as body, intermediate segments nearly three times
as long as wide, vestiture short and decumbent. Pronotum as in figure 12P.
Anterior claw of front tarsi with blunt exterior basal appendix, posterior claw
simple; other tarsi similar to front, with basal appendix progressively smaller.
Length 4-5 mm.

DISTRIBUTION. ARIZONA: Huachuca Mountains; Chiricahua Mountains,
Southwestern Research Station, 5 mi. W. of Portal; Santa Rita Mountains,
Madera Canyon.

SUBGENUS Silis

The genus Silis was proposed by Charpentier (1825) with S. spinicollis
Charpentier, a synonym of S. nitidula Fabricius, as the genotype. Specimens of
S. nitidula, generously donated by Vladimir Vosyka of Czechoslovakia, have
genitalia similar to those found in the ‘“‘vulnerata’” group, confirming the
generic assignment of the Nearctic species. They also have the same smooth,
shining, and glabrous hypomera.

Modification of the tarsal claws in the males is of the same type through-
out, and mention of it will be omitted from the species descriptions that follow.
The anterior claw of all tarsi is provided with a blunt exterior basal appendix,
the posterior claw of all tarsi being simple. The antennal structure is also
fairly constant: slender, subfiliform, about four-fifths as long as the body,

460

CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER.

intermediate segments three to four times as long as wide. Length-width ratios
are likewise omitted, unless they vary appreciably from the normal 2.75: 1
The color of the mandibles is entirely or largely pale in all species, and the
apical margins of the ventral segments are more or less distinctly pale. The
most essential descriptive items are the genitalic and pronotal drawings, and
these alone should be sufficient for species identification. It is not to be ex-
pected that every specimen is identifiable. Variability is often baffling, and
unrecognized species are probably still numerous.

In the females the hypomera, unlike the males, are flat or convex, and

usually punctate and pubescent. The females of those species with bicolored
pronota have the black areas much reduced, and occasionally absent altogether.

Key TO NEARCTIC SPECIES OF THE SUBGENUS SILIS, MALES

Prothorax small, narrower than elytra at base. Posterior process of pronotal armature

broad, bidentate, anterior tooth long and acute _.. (1) S. (S.) spinigera LeConte
Prothorax normal, as wide as elytra at base. Posterior process not bidentate 2
Antennal vestiture erect aime brs blir ps eee ee ee nn 3
Antennal -vestiture decumbent —_ eee 5
California species. Aedeagus with laterophyses (2) S. (S.) rugosa Van Dyke

Eastern or midwestern species. Aedeagus without laterophyses
Acute lateral margin of pronotum attaining tip of anterior process
Eek Chg ope: Vans Sed eS POE Petey Pt SNe wee che oe Pee. b+. (3) S. (S.) percomis Say
Acute lateral margin of pronotum not attaining tip of anterior process —..
BBs Pe RSS ER See in Re: Sand oe a, ee ee a _ (4) S. (S.) spathulata LeConte
Posterior process of pronotum, viewed ventrally, rapidly narrowing and terminating

invay setitorm’ spine directed amtersorly, se ee ee enn Sn 6
Posterior process not as above, of substantial width throughout — 7
Posterior process abruptly curving forward, attaining tip of anterior process. Excision

at hind angles large and open (S. (S.) lutea. Group) =.= = ee 12

Posterior process terminating in a small seta-like spine not attaining tip of anterior
process. Excision at hind angles more or less completely closed by dorsal surface of
extended hypomeron. Southern California (8) S. (S.) filicornis Van Dyke
Posterior process combining with laterally produced hind angle of pronotum to form
a bidentate structure in dorsal view. Basophyses united, forming a broad deeply

(Hoifavornie jokes ee (9) S. (S.) tardella Green, new species
INOtyaSsabovie 2. ee epee eee Be 8
Posterior process nearly Pecos subspiniform, directed outward and slightly backward 9
Not.as above 0 ee 10
Pronotum pale with dark borders which are rarely lacking. Redeagee without latero-

physes (S. (S.) difficilis Group) ___.._... . | ee 14
Pronotum unicolorous, fulvous or black. Aedeagus with laterophyses (S. (S.) deserti-

Posterior process simple, narrow, directed outward. Tips of anterior and posterior

processes widely separated _. rer Il!
Posterior process complex, broad (S. (S.) cava : Group) eos 29
Posterior process curving slightly forward (S. (S.) vulnerata Group) 47

Posterior process bending abruptly forward acta (S. (S.) pallida Group)

Vor. XXXII] GREEN: REVISION OF THE NEARCTIC SILIS 461

In the following part of the key, all characters, except where otherwise stated, refer
to the male genitalia.

S. (S.) LUTEA GROUP

12. Dorsal plate with small median emargination not entering membraneous area. Figure 5
oe raun sents nei Ge sale a tel SRR i ll ane ed (5) S. CS.) carmelita Green, new species
Dorsal plate with wide and deep emargination entering membraneous area —- 13

13. Basophyses moderately bent inward submedially, exposed tips straight in lateral view.
Emargination of dorsal plate narrower. deeper than wide. Pronotum with median
dark area when elytra are black. Figure 6 _._________ (6) S. (S.) lutea LeConte
Basophyses strongly bent inward submedially, exposed tips slightly recurving or hooked
in lateral view. Emargination of dorsal plate with depth and width subequal. Pro-
notum entirely pale, elytra black. Figure 7 —.____ (7) S. (S.) filigera LeConte

S. (S.) DIFFICILIS Group

In some of the species of this group, the upper margins of the lateral incisures of the
tegmen, viewed ventrally, are more or less prominently angulate at about the distal third, the
angulation sometimes convex and subcarinate, varying to a simple flat extension of the sur-
face. When the angulation is fully developed, the margin of the incisure deflects around the
tip of the angle, which is actually a small oblique lamina appearing acute from a direct ven-
tral viewpoint. Nearly all of the S. difficilis group species with this modification are Cali-
fornian, the only known exception being S. nevadica. The emargination of the dorsal plate
follows a general plan for each species, as shown in the drawings, but it is usually quite
variable. There are no laterophyses. The antennal vestiture is short and decumbent.

14. Emargination of dorsal plate narrowing to apex, basophyses emerging close together
and subparallel. In lateral view, tips of basophyses recurving or enlarged on proxi-
mall Hele, TSK REN Gye ee ee ee 15

15. In ventral view, upper margins of lateral incisure of tegmen subangulate at about
distal third. Emargination of dorsal plate usually with sides sinuate distally. Figure 10
copier A ee a ee ee eee (10) S. (S.) flavida LeConte
In ventral view, upper margins of lateral incisure of tegmen not subangulate at distal
third. Emargination of dorsal plate usually with sides straight, not, or very feebly,

Suatexdistallys, Bircune Wil <2 (11) S. (S.) lasseni Green, new species
16. In ventral view, upper margins of lateral incisure of tegmen subangulate at about
eles reals Gl air mes 8 ae eM TREN ey SEC eg Se a ee eee ee ey 17
In ventral view, upper margins of lateral incisure of tegmen not subangulate at distal
(HOmRG! ont te ee ee ek Aor. Dawes Ser een EA) ba oe Eee ee 19
17. In lateral view, tips of basophyses recurved or prominently enlarged on proximal side.
LENVSGUST@ TWA ca a a a oh Rp a Le een (12) S. (S.) nevadica Green, new species
In lateral view, tips of basophyses not recurved or prominently enlarged on proximal
1 ee vs ee So ee eh Se ee eee 18

18. Exposed tips of basophyses very slender. In dorso-ventral view, median lobe of
aedeagus broad, not narrowing distally, sides not striate. Ventral lobe of tegmen
wider, with distinct apical notch. Figure 13 (13) S. (S.) tenuata Green, new species
Exposed tips of basophyses normally wide. In dorso-ventral view, median lobe of
aedeagus narrowing to apex, sides striate. Ventral lobe of tegmen narrower, without,
or with minute, apical notch. Figure 14 (14) S. (S.) carbo Van Dyke

19. In dorso-ventral view, median lobe of aedeagus broad, with small apical notch, sides
MoOtestilates Eup UTE MS; me 5 ee SAE eS (15) S. (S.) oregonensis Green, new species

462 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

In dorso-ventral view, median lobe of aedeagus elongate, narrowing to apex, sides in
part sclerotized and with irregular longitudinal striation —_—_____________ 20
20. Emargination of dorsal plate less broadly U-shaped, deeper than wide, sides subparallel
or feebly diverging, usually not curving inward distally. Emargination not attaining
membraneous area by a conspicuous distance. Figure 16 —.
pa ae ee eS _ (16) S. (S.) striatella Green, new species
Emargination of dorsal plate broadly U- or V-shaped, about as deep as wide, sides
curving inward distally —.....2. 2. eee 21
21. Elytra in part usually more or less flavate, this not apparent in darker individuals un-
less examined under brilliant illumination. Basophyses typically extending very little
beyond emargination of dorsal plate, narrowing apically as seen in lateral view.
Emargination of dorsal plate usually not attaining membraneous area by a conspicuous
GistanceseE1oure ir eS ee ee ee (17) S. (S.) difficilis LeConte
Elytra usually uniformly deep black. Basophyses typically extending conspicuously
beyond emargination of dorsal plate, somewhat expanded apically as seen in lateral
view. Emargination of dorsal plate usually attaining membraneous area. Figure 17a
Maco bn wit Ai f oe ... (17a) S. (S.) difficilis occidens Green, new subspecies

S. (S.) DESERTICOLA GROUP

In this group the laterophyses are provided with a coriaceous and sparsely setiferous distal
appendix. The appendix may be small and inconspicuous, or it may involve the entire tip
of the laterophysis. In the latter case, the actual tip of the laterphysial rod is recognized by
being strongly sclerotized and glabrous. This structure is also found in the S. vulnerata
group and in the Palearctic genotype, S. nitidula Fabricius. The basophyses are united basally
and furcate at about the middle of their length. A unique feature of this group, S. atra ex-
cepted, is the trilobed dorsal plate, with the lobes on each side at a level below that of the
median part. This complicates the genitalic drawings, but it may be clarified to some extent
by the posterior view diagrams of the tegmen. All of the Silis species with appendiculate
laterophyses are apparently well established and not involved in current evolutionary de-
velopment. The species allied to S. deserticola, having fulvous pronota, are rare in collections.
Only eight males, representing seven species, are known at present. They are identical exter-
nally, differing radically in the male genitalia. All occur in southern California. The antennal
vestiture is short and decumbent.

22. Dorsal plate simple, with deep and very wide emargination. Body entirely black
ADOVE Sigur gS) <= Pees so ee Pa ee ee (18) S. (S.) atra LeConte
Dorsal plate trilobed. Body, black, pronotumm fullvovs) 23

23. Ventral lobe of tegmen extending well beyond dorsal plate. Posterior orifice of tegmen
not constricted below dorsal plate. Figure 19 (19) S. (S.) egregia Green, new species
Ventral lobe of tegmen not extending beyond dorsal plate. Posterior orifice of tegmen

constricted (below dorsal plate 24
24. Laterophyses with tips squarely truncate 25

Laterophyses with tips not squarely truncate, more or less acute 26
25. Median part of dorsal plate feebly lobed, lobes on each side extending far beyond it.

Figure 20 a -. _ (20) S. (S.) singularis Green, new species

Median part of aareal iaiese not pdlahede Ww math shallow and broadly angulate emargina-

CLONE UT Opole een NS as PLS Se ses (21) S. (S.) eximia Green, new species

26. Median part of dorsal plate deeply ‘aaa emarginate. Figure 22

ee Seat (22) S. (S.) abstrusa Green, new species
Median part of dorsal plate not emarginate __ wove eo) eee 27

Vor. XXXII] GREEN: REVISION OF THE NEARCTIC SILIS 463

Bile

29.

30.

Oye

32°

SS.

34.

Bo.

36.

38.

Median part of dorsal plate with prominent lobe about half as long as wide. Figure 23
cn neese clits See tee eae eS SOS ee a (23) S. (S.) arida Green, new species
Me dianepartsonidorsdleplatessub truncal cys eee ee et ee eee 28

. Lobes each side extending well beyond median part of dorsal plate. Median lobe of

aedeagus with apical process. Figure 24 ______ (24) S. (S.) ursina Green, new species
Lobes each side extending only slightly beyond median part of dorsal plate.
Median lobe of aedeagus without apical process (25) S. (S.) deserticola Van Dyke

S. (S.) CAVA Group

Genitalia with single basophysis, which may be furcate or not — 30

Genitalia with two basophyses S. (S.) emarginata Section 31

Basophysis furcate at about middle of length S. (S.)cava Section 39

Basophysis not furcate, tip notched S. (S.) californica Section —__ oe NG Bone Meal nde, 40
S. (S.) EMARGINATA SECTION

Dorsal plate not emarginate, posterior margin convex. Figure 26 ee NA

co eee SE a a ED (26) S. (S.) imcongrua Green, new species
Dorsal plate emarginate; or subtruncate in dorsal view, emarginate in posterior view —_ 32
Laterophyses with prominent exterior distal tooth. Figure 27 —-._______________-

caeele renee st EE es En (27) S. (S.) angulata Green, new species

NPA eEOp My CeSpawatl OU tre O OC) soe ee Sa a ee eet 33
Laterophyses with tips somewhat widened and everted in dorso-ventral view 34
Laterophyses slender distally, tips straight or feebly diverging — 36

Basophyses extending posteriorly as far as, or beyond, apex of deep emargination of
dorsal plate. Elytra flavate, tips sometimes dark. Legs largely pale. Figure 28
Seeoine Ser esi res) et ahs Arius St lees (28) S. (S.) emarginata Green, new species
Basophyses not extending posteriorly as far as apex of shallower emargination of
dorsal plate. Elytra flavate with sutural bead and apex dark. Legs largely dark 35
Dorsal plate with rather deep subtriangular emargination usually attaining mem-
braneous area. Tips of laterophyses everted for a considerable distance. Figure 29
eNUENNERIE Naaa A ee ig Tk ot EA (29) S. (S.) fenderi Green, new species
Dorsal plate with shallow arcuate emargination not attaining membraneous area by a
conspicuous distance. Everted tips of laterophyses very short. Figure 30
careaanc iit (30) S. (S.) subtruncata Green, new species

In lateral view, median lobe of aedeagus not subtriangular, upper margin broadly
LRG ACC OTEM CATV ELS BAN CIN (ey ete es nee ane eb fo ee ae a ee 37
In lateral view, median lobe of aedeagus elongate subtriangular, widest at about distal
tavinGl, ANS (yay Kein Tae xeat ny 1) Gyo ob oyeq hl Eye) ee ee 38

. Lobes of dorsal plate emargination not, or only feebly, deflected: in direct posterior

view, emargination completely flattened or very broadly V-shaped. Laterophyses
moderately slender = elt, lees, setae (31) S. (S.) macclayi Green, new species
Lobes of dorsal plate emargination strongly deflected, forming a deep U-shaped emar-
gination in direct posterior view. Laterophyses distinctly broader as compared to
preceding species, Bigure 32) 2 7 (32) S. (S.) crucialis Green, new species
Laterophyses broad except apically. Figure 33 (33) S. (S.) thermalis Green, new species
Materophyses rather slender throughout. Bicure 34 22

Genitaliagasninptl ome ys Ss ewes oes ee ee ee (35) S. (S.) cava LeConte
Genitalia similar but differing in some respects — S. (S.) cava complex

464

40.

41.

42.

43.

44.

45.

46.

47.

48.

49.

Sule

CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER.

S. (S.) CALIFORNICA SECTION

Dorsal plate not emarginate. Pronotum entirely pale fulvous. Figure 36
Se Ap er ete, A ane CE Soe eae aS ln AU a Dah (36) S. (S.) arizonica Van Dyke
Dorsal plate emarginate, emargination sometimes lacking in S. (S.) fenestrata, in
which’ the pronotall borders are) black Se 41
Dorsal plate with small variable emargination, rarely lacking. Pronotum pale fulvous
with black borders; anterior process with narrow extension deflected below acute
angle of posterior process, completely closing marginal incisure in direct dorsal view.

IEG Pe Ue ghia es eee Pee oe ae ee (37) S. (S.) fenestrata Van Dyke
Dorsal plate with normal, usually deep, emargination. Pronotum entirely pale fulvous
on tlavates swith) distinct manera eames une sea i3t 42
Elytra dark, partly fusco-flavate under brilliant illumination __._.___ 43
Elytrapale flavate. 2.20 he ee 44
Laterophyses shorter, stout except apically, curving downward distally. Basophysis
expandedeapicallyzmEr one ns eee (38) S. (S.) californica Fender
Laterophyses longer, slender throughout, curving upward distally. Basophysis not ex-
pandedtapicallly sbi eure 23,9) eee ee (39) S. (S.) reversa Green, new species

Laterophyses broad, tips bent strongly inward and downward. Emargination of dorsal
plate wide, moderately deep, apex subtruncate. Figure 40 _..
pete Sete «piano aa. Se A a (40) S. (S.) solitaria Green, new species
In dorso-ventral view, tips of laterophyses subparallel or diverging _. 45
Laterophyses slender throughout, tips not diverging. Figure 41
BER ea Ne Ste Sie 2: ee ee Oe eee (41) S. (S.) recta Green, new species

iaterophyses, broad except apicalliya) tips) diver im pe ee eee 46
In dorso-ventral view, laterophyses curving inward at about distal fourth, lyre-shaped.
Bigunes42% ee aes es Se ee ee (42) S. (S.) angelica Green, new species

In dorso-ventral view, laterophyses cubparallel Figure 432... 44 ee eee
_ (43) S. (S.) latestyla Green, new species

S. (S.) VULNERATA GROUP

With single basophysis, bilaterally enlarged at apex. Laterophyses slender, lyre-shaped,
without distal appendix. Pronotum without distinct median dark area. Figure 44
EEE co Rat Las Bea anne es each wa eS (44) S. (S.) fabulosa Green, new species
W tin two a ecooine S@S 22230 ee 48
Posterior margin of dorsal plate not nile or thickened, with at most only a minute
denticulation each side
Posterior margin of dorsal plate inflexed or thickened = 50
Basophyses widely divaricate, tips broadly visible each side of ventral bhe in direct
Wena Yate, lieguns Gy) _ (45) S. (S.) divaricata Green, new species
Basophyses subparallel, nee not nts ace Sade of ventral lobe in direct ventral view
= (46) S. (S:) montanica Green, new species
Bperenier margin of aera pate not bieutate: a short inflexed median lobe separating
tips of basophyses. Laterophyses apparently without distal appendix. Figure 47 __
Wr ee (47) S. (S.) lecontei Green, new species

Borierion margin of nin pinion more or less strongly bidentate, teeth inflexed, acute
or obtuse _ : to te : Pra tesroyiL

Marginal teeth of dorsal slate very are wade orominent plains visible feat ‘direct

lateral viewpoint. Basophyses not extending to dorsal plate, their tips separated
therefrom by a considerable distance ee Fe 3 52

VoL. XXXII] GREEN: REVISION OF THE NEARCTIC SILIS 465

sya

54.

35y

56.

58.

60.

61.

62.

63.

64.

Marginal teeth of dorsal plate of normal size, not plainly visible from direct lateral
SW TERME a a a Sc kc a Es ee Ne Ne Oa
Basophyses long, slender in more than distal half, curving outward and upward. Mar-
ginal teeth of dorsal plate extremely large, much retracted. Pronotum with black
borders, disk entirely fulvous. Figure 48 (48) S. (S.) dentigera Green, new species
Basophyses short and stout. Marginal teeth of dorsal plate smaller. Pronotum largely
DlackabicurerAQhs ee Se (49) S. (S.) disjuncta Green, new species
Basophyses long, extending for a considerable distance beyond posterior margin of

GL NES etl YO cht een Se a ee Re ee ee 8 GA
Basophyses of normal length, at most with tips curving over posterior margin of
dorsal plate, but not extending beyond for any appreciable distance — 58
Appendix of laterophyses small and inconspicuous, sometimes not apparent ~ 55
Appendix of laterophyses larger, conspicuous, not lineate 56
In ventral view, tips of basophyses curving slightly inward. Laterophyses feebly
sinuate in lateral view, curving downward distally; parallel and straight in dorsal view.
PHpUn®. RO) sn Re Se Si ee a ee ene (50) S. (S.) triplicata Green, new species
In ventral view, tips of basophyses curving strongly inward. Laterophyses nearly
straight in lateral view, curving inward distally in dorsal view. Figure 51
Se eB OI ss (51) S. (S.) protracta Green, new species
Appendix larger than contiguous portion of laterophysis, which it may envelop and
SS CU Ts 2 nce ec ee ea PB eS 59
PD DENC ina OLAS (AD OVC = ss 5 ee ee ee el FE ee ee
In lateral view, laterophyses slender throughout. Figure 5la
Unidentified, near S. protracta
In lateral view, laterophyses much expanded on dorsal side. Figure 52 —
(52) S. (S.) simulata Green, new species

In lateral view, appendix large and prominent, nearly as large as, to larger than, con-

tiguous portion of laterophysis _— er re ctw EN SO ee ie 59
In lateral view, appendix small and inconspicuous, sometimes not apparent. Pronotum
OGTR TN © CLA CH TD Wilt ct ee we eek ake Ske oe ee A 65
In lateral view, appendix foliate, considerably broader and extending far beyond
Send Gat Tp mOlmslatehO ply SiG gees pene e fe ek tA ee ee eee 60
In lateral view, appendix extending little or not at all beyond tip of laterophysis 62

In ventral view, basophyses straight, divergent, tips visible each side of ventral lobe __ 61
In ventral view, basophyses diverging, abruptly converging from about distal fourth;
tips not visible each side of ventral lobe. Figure 53 —
cence ee 2 Ee ee (53) S. (S.) introversa Green, new species
In dorsal view, diverging tips of laterophyses short. abruptly everted. Figure 54
er cesgee ee Ea ee ee eee ee _ (54) S. (S.) perfoliata Green, new species
In dorsal view, diverging tips of laterophyses longer, regularly arcuate. Figure 55
(55) S. (S.) proxima Green, new species

In ventral view, basophyses divergent, straight, tips visible each side of ventral lobe __- 63
In ventral view, basophyses feebly arcuate, subparallel distally, tips not visible each
SHG Oi. Wyuna) | 10 oe ee a ee ee ee, ee eee ee 64

Diskwotepronotumy without median) vittas, Kigure 56) 2 ee
nec ec Og PE 8 oe _. (56) S. (S.) abrupta Green, new species
Disk of pronotum with median vitta. Figure 57 __ (57) S. (S.) lobata Green, new species
In lateral view, laterophysis asymmetric, somewhat L-shaped, its lower margin bend-
ing rather abruptly upward at about distal third, forming an obtuse angle; its upper
margin concavely arcuate. In lateral view, basophyses with lower margin uniformly

Vor. XXXII] GREEN: REVISION OF THE NEARCTIC SILIS 467

ALCUALC MOP IPT EERO Se eees a! AA le ok et eee ee daw 2 hho) (58) S. (S.) vulnerata LeConte

In lateral view, laterophyses not as above, nearly symmetrical. In lateral view, baso-

physes with lower margin usually subangulate. Figure 59
censrnanaistn le Se ee (59) S. (S.) parallela Green, new species

65. In lateral view, basophyses very stout, constricted near tip. Figure 60
penne re oe a FDIS EE 2s VARI (60) S. (S.) constricta Green, new species
BAS OMY Sesuh OLtaStADON GC) o-oo a eee ee oe ee eee ee Pe 66

66. In lateral view, laterophyses abruptly bending downward at about middle of length;
appendix setiform:. Bicure 61 22 = 2 22 (61) S. (S.) barri Green, new species

In lateral view, laterophyses curving feebly downward near tip; appendix not appar-
SiR CULCtO2: see eee ee Ee ee (62) S. (S.) insolita Green, new species

S. (S.) PALLIDA GROUP

67. Median lobe of aedeagus narrowly subtriangular in dorso-ventral view, slender
apically. Ventral lobe of tegmen subtriangular, apex more or less acute —-------_---__. 68
Median lobe of aedeagus broad in dorso-ventral view, sometimes narrowing toward
PYDET, caper EN ON 5 Oi lle Ee ee Re bs Ee EE ee ee ee ee 69

68. Basophyses shorter, not attaining posterior margin of dorsal plate. Figure 63 —
eee cet ast EE ai Be | (63) S. (S.) pallida Mannerheim
Basophyses attaining posterior margin of dorsal plate. Figure 64 —
cone cents Es ne (64) S. (S.) imsperata Green, new species

69. Elytra black. Laterophyses curving downward at about distal third or fourth. Figure
ONS) ps a ee 2 ee fig el as (65) S. (S.) maritima Van Dyke
Elytra flavate, sides and apices dark. Laterophyses various Unidentified complex

(1) Silis (Silis) spinigera LeConte.

Silis spinigera LECONTE, 1874, Trans. Amer. Ent. Soc., vol. 5, p. 61.
Silis munita LECONTE, 1881, Trans. Amer. Ent. Soc., vol. 9, p. 56.

Black; prothorax flavate, elytra flavate with apices more or less broadly
dusky, legs partly pale. Varies with elytra and legs entirely black.

Ficures 1-12. Male genitalia; ventral, lateral, and dorsal views, arranged in that order
from left to right.

Ficure 1. Silis (Silis) spinigera LeConte, dorsal view omitted.
FicureE 2. Silis (Silis) rugosa Van Dyke.

Ficure 3. Silis (Silis) percomis Say.

-Ficure 4. Silis (Silis) spathulata LeConte.

Ficure 5. Silis (Silis) carmelita Green, holotype.

Ficure 6. Silis (Silis) lutea LeConte.

Ficure 7. Silis (Silis) filigera LeConte.

Ficure 8. Silis (Silis) filicornis Van Dyke.

Ficure 9. Silis (Silis) tardella Green, holotype, dorsal view omitted, basophysial plate
inserted.

Figure 10. Silis (Silis) flavida LeConte.

Ficure 11. Silis (Silis) lasseni Green, holotype.

Ficure 12. Silis (Silis) nevadica Green, holotype.

468 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Form elongate, length-width ratio about 3.2 : 1. Antennal vestiture short
and decumbent. Pronotum as in figure 13P, small, narrower than elytra at
base. Genitalia as in figure 1, no laterophyses; median lobe large and stout,
as wide as ventral lobe of tegmen. Length 7—8.5 mm.

DisTRIBUTION. CALIFORNIA, OREGON, BRITISH COLUMBIA,
IDAHO, UTAH, WYOMING, COLORADO.

This species, occurring over a wide range, is remarkably constant in
structure, regardless of color. It probably should be removed from Szlis to a new
monobasic genus because of its radically different genitalia and facies.

(2) Silis (Silis) rugosa Van Dyke.
Silis rugosa VAN Dyke, 1918, Jour. New York Ent. Soc., vol. 26, p. 169.

Body and appendages deep black; pronotum fulvous with all borders black,
the lateral borders narrowly. Underside black except prothorax fulvous with
lateral borders black.

Antennal vestiture erect and bristling. Pronotum as in figure 14P. Geni-
talia as in figure 2, basophyses widely divergent, surpassing sides of tegmen.
Laterophyses each with acute external tooth distally. Dorsal plate shallowly
emarginate throughout its width. Ventral lobe with apex acute. Length
4—5 mm.

DiIsTRIBUTION. CALIFORNIA: Marin County: Mt. Tamalpais, Mill
Valley; Lake County: Anderson Springs; Napa County: Oakville; Santa Cruz;
Alameda County; Mendocino County: Ryan Creek, Fort Bragg.

All dates of capture are for February and March, except for a Fort Bragg
specimen taken in December.

(3) Silis (Silis) percomis Say.

Cantharis percomis SAY, 1835, Boston Jour. Nat. Hist., vol. 1, p. 159.
Podabrus curtus LEContTEe, 1850, In: Lake Superior, by Louis Agassiz, p. 229 (female).
Silis longicornis LEContTE, In: Lake Superior, by Louis Agassiz, p. 230 (male).

Black; pronotum pale rufous with anterior explanate border, and basal
border medially, more or less blackish.

Antennal vestiture erect and bristling. Pronotum as in figure 15P, acute
lateral margin continuing from apex to tip of anterior process. Genitalia as in
figure 3, no laterophyses, ventral lobe of tegmen extending posteriorly beyond
dorsal plate. Length 5-6 mm.

DistripuTION. NEW BRUNSWICK, NOVA SCOTIA, ONTARIO, QUE-
BEC, MAINE, NEW HAMPSHIRE, MASSACHUSETTS, CONNECTICUT,
NEW YORK, PENNSYLVANIA, NEW JERSEY, MARYLAND, VIRGINIA,
KENTUCKY, TENNESSEE, NORTH CAROLINA, SOUTH CAROLINA,
ALABAMA, OHIO, INDIANA, ILLINOIS, MICHIGAN, WISCONSIN,
IOWA, MINNESOTA.

Vor. XXXII] GREEN: REVISION OF THE NEARCTIC SILIS 469

(4) Silis (Silis) spathulata LeConte.

Silis spathulata LeContTE, 1881, Trans. Amer. Ent. Soc., vol. 9, p. 57.

Black, pronotum pale rufous with anterior border more or less blackish.

Antennal vestiture erect and bristling. Pronotum as in figure 16P, acute
lateral margin not extending to tip of anterior process. Genitalia as in figure 4,
ventral lobe of tegmen not extending posteriorly beyond dorsal ie No
laterophyses. Length 5 mm.

DIsTRIBUTION. NEW YORK: J/thaca; TENNESSEE: Knoxville; NORTH
CAROLINA: Chapel Hills; SOUTH CAROLINA: Clemson.

This species is rare in collections, most of the specimens so identified
belonging to S. percomis. The genitalia of the two species are quite distinct,
but externally the two are not so readily separated. The best character for
that purpose is the acute lateral margin of the pronotum, which in S. percomis
extends to the tip of the anterior process, while in S. spathulata the acute
margin obsolesces at an appreciable distance before the tip.

(5) Silis (Silis) carmelita Green, new species.

HOLOTYPE, male; Carmel, Calif., Monterey County, IV-9-11, L. S.
Slevin. In collection of California Academy of Sciences. CAS type number 9081.

Head and appendages black. Pronotum pale fulvous with entire median
black vitta about one-third pronotal width, widest at base, narrowing an-
teriorly, expanding abruptly over anterior explanate border, margins of vitta
irregular. Scutellum and elytra black. Underside, except prothorax, dark.
Legs black.

Pronotum as in figure 17P. Genitalia as in figure 5, no laterophyses;
emargination of dorsal plate small, not entering membraneous area. Basophyses
feebly bending inward near middle of length. Length 5.5 mm.

VARIATIONS. The median pronotal vitta may be much constricted or en-
tirely eliminated in anterior third or half. This occurs mostly in females, only
one male showing a pronounced constriction. In two males the elytra are
flavate with the sutural bead dark, and the lateral borders except basally, and
the apex more broadly, infuscate. In one of these males the pronotum is
entirely flavate, in the other it is flavate with a large medio-basal dark area.
_Two females also have similarly flavate elytra, but without pronotal maculation.

DISTRIBUTION. CALIFORNIA: Carmel: holotype, 5 paratypes, 6 females.
IV-9 to V-20, E. C. Van Dyke, or F. E. Blaisdell, or L. S. Slevin (CAS); 1
paratype (elytra flavate) II-22-16, Slevin (CAS); 1 female (elytra flavate)
III-15-31, Slevin (CAS). Del Monte, 1 paratype, V-5-23, Slevin (CAS). San
Simeon, 1 paratype, V-22-54, O. Bryant (CAS). Soquel Creck, Santa Cruz
County, 1 paratype, 1 female (both with flavate elytra) V-30-19, Van
Dyke (CAS). Seaside, Monterey County, 1 female, IV-11-13, Slevin (CAS).

470 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

(6) Silis (Silis) lutea LeConte.

Silis lutea LeContE, 1853, in Catalog-desc. Coleop. of U.S., by Melsheimer, Smithsonian

Inst., p. 78.
Silis pallens LeConte, 1851, Proc. Acad. Nat. Sci. Philadelphia, vol. 5, p. 339 (preoccupied).

Head and appendages black. Pronotum pale fulvous with entire median
black vitta about one-third pronotal width, widest at base, narrowing anteriorly,
expanding abruptly over anterior explanate border, margins of vitta irregular ;
varying with partial elimination of dark area anteriorly, progressing posteriorly
to pronotum entirely pale. Scutellum black. Elytra flavate with sutural bead
dark, apex and lateral borders, except basally, usually somewhat infuscate.
Varying with elytra entirely black, which is usually seen to be in part fusco-
flavate or brunneous when examined under brilliant illumination. Pronotal
vitta always present when elytra are dark.

Antennal vestiture short and decumbent. Pronotum as in figure 17P.
Genitalia as in figure 6, no laterophyses; emargination of dorsal plate large,
deeper than wide, entering membranous area. Basophyses feebly bending inward
near middle of length. Length 4.5—-6.5 mm.

DISTRIBUTION. BRITISH COLUMBIA, WASHINGTON, OREGON,
northern half of CALIFORNIA.

The color of the elytra is of no taxonomic importance. The dark form
extends from Humboldt County, in northern California, to British Columbia
where it predominates over the pale phase. Pale and dark ones occur together
in the same locality. Examples of S. dutea with dark elytra were considered by
Van Dyke to be a color phase that he identified as S. filigera LeConte. The
latter is a valid species quite different in genitalic structure, and occupying
a different geographic range.

A specimen from Mt. Madonna, Santa Cruz County, California (UCD),
having the elytra black and the pronotum entirely pale, is referred here
tentatively. The basophyses differ slightly in being more slender, and ap-
parently do not bend inwardly near middle of length. This feature is some-
what variable and cannot be considered conclusive without additional material.

(7) Silis (Silis) filigera LeConte.
Silis filigera LeConte, 1874, Amer. Ent. Soc., Trans., vol. 5, p. 62.

Body and appendages black, pronotum entirely pale rufous.

Pronotum as in figure 17P. Genitalia as in figure 7, no laterophyses,
emargination of dorsal plate very large, as deep as wide, entering mem-
braneous area, basophyses bending strongly inward near middle of length.
Length 4.5—5 mm.

DistRIBUTION. CALIFORNIA: Pasadena, IV, Fenyes, 1 male, 1 female
(CAS). Sierra Madre, VI, Fenyes, 1 male (CAS).

Vor. XXXII] GREEN: REVISION OF THE NEARCTIC SILIS 471

This is definitely a valid species and not a color phase of S. dutea. Only
two males and one female have been seen. They are constant in color pattern,
agreeing with LeConte’s description, but probably additional material will
show the color variability characteristic of the S. lutea group.

(8) Silis (Silis) filicornis Van Dyke.
Silis filicornis VAN Dyxer, 1918, Jour. New York Ent. Soc., vol. 26, p. 172.

Body and appendages black, pronotum pale fulvous.

Length-width ratio about 3:1. Form narrow, parallel-sided. Antennae
slender, filiform, about as long as body, intermediate segments about four
times as long as wide, vestiture short and decumbent. Pronotum as in figure
18P, excision at hind angles more or less completely filled by dorsal surface
of hypomeron which joins lateral margin of pronotum at tip of anterior
process, and at its angular rear extremity is produced in a small extension
(the posterior process) terminating in a forward-curving spiniform seta.
Genitalia as in figure 8. Basophyses united to form a broad _ parallel-sided
plate about half as wide as tegmen, shallowly emarginate distally. Latero-
physes straight, spiniform. Median lobe of aedeagus with dual short spiniform
processes at apex. Dorsal plate with raised median lobe extending backward,
its tip meeting posterior margin of basophysial plate. Length 4-5 mm.

A specimen from Pine Valley, San Diego County (CAS), varies in having
the pronotum as in figure 19P. It is referred to this species as the genitalia
seem not to differ in any way.

DISTRIBUTION. CALIFORNIA: San Jacinto Mountains. Palm Springs;
same, Andreas Canyon; same, Palm Canyon. White Water; same, Snow
Creek. Thousand Palms. East Highlands. San Bernardino Mountains, Devil
Canyon. 2 mi. W. of Jacumba. San Diego County, Pine Valley.

(9) Silis (Silis) tardella Green, new species.

Ho.otyPe, male; California, no definite locality, Horn collection, H 4923,
with identification label “‘S. filigera Lec.” In collection of Academy of Natural
Sciences of Philadelphia.

Head black, pale in front; antennae and palpi dark. Pronotum entirely
_fulvous. Scutellum and elytra black. Underside: head fulvous, dark each side
of gula; prothorax fulvous; balance of ventral surface dark except apical
borders of ventral segments pale. Legs and coxae dark.

Length-width ratio about 2.75: 1. Antennae filiform, about four-fifths as
long as body, intermediate segments about three and one-half times as long
as wide, vestiture short and decumbent. Pronotum as in figure 20P, lateral
explanate borders thin, subhyaline. Anterior claw of all tarsi with blunt ex-
terior basal appendix, other claws simple. Genitalia as in figure 9, dorsal plate

472 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

not emarginate, no laterophyses, basophyses united in a broad deeply emar-
ginate plate attaining posterior margin of dorsal plate. Length 5 mm.
DIsTRIBUTION. CALIFORNIA, no definite locality (ANSP).
This species is represented only by the holotype from the Horn collection.
The possibility that it might be a mislabeled European specimen should be
investigated.

(10) Silis (Silis) flavida LeConte.
Silis flavida LEContr, 1874, Trans. Amer. Ent. Soc., vol. 5, p. 61.

Antennae black, pale beneath basally; palpi black. Head black, pale in
front. Pronotum flavate; varying with basal, apical, and sometimes lateral,
borders black. Scutellum black. Elytra flavate with apices broadly, and sides
except basally, black or dusky; sutural bead usually brunneous, varying to
partly or entirely flavate. Underside, except prothorax, dark. Legs pale, coxae
and base of femora black, metafemora largely black.

Pronotum as in figure 21P. Genitalia as in figure 10, emargination of
dorsal plate narrowing to apex, its sides usually sinuate distally; basophyses
emerging close together, subparallel, tips recurving or enlarged on proximal
side. In ventral view, upper margins of lateral incisure of tegmen subangulate
at about distal third. Length 5—6.5 mm.

DisTRIBUTION. CALIFORNIA: Nevada County: Sagehen Creek near
Hobart Mills; Truckee. El Dorado County: Tahoe; Grass Lake; Echo Lake;
Lake Tahoe; Fallen Leaf Lake; Mt. Tallac; Tallac; Wright’s Lake; Angora
Lake. Alpine County: Hope Valley; Ebbets Pass. Calaveras Big Trees. Sonora
Pass; Tuolumne Meadows; Yosemite National Park; Yosemite Valley; Saylor
Lake; May Lake; Strawberry; Summerdale (Fish Camp). Mono County:
Tioga Crest; Sardine Creek; Sonora Pass. NEVADA: Lake Tahoe.

This species is quite variable and may be complex. Material from the
Yosemite region usually has the emargination of the dorsal plate V-shaped
with straight sides and often with the tips turning inward. A specimen labeled
“Nev.” (ANSP) has the tegminal structure of S. flavida and the aedeagus of
S. lasseni, and is possibly a hybrid.

(11) Silis (Silis) lasseni Green, new species.

HoLotyPe, male; Duck Lake, Lassen County, California, V-8-21, J. O.
Martin. In collection of California Academy of Sciences. CAS type number
9082.

Antennae black, three basal segments pale beneath; palpi dark. Head black,
pale in front. Pronotum flavate with all borders narrowly black. Scutellum
black. Elytra flavate, becoming darker rufo-brunneous in more than distal

VoL. XXXII] GREEN: REVISION OF THE NEARCTIC SILIS 473

half. Underside, except prothorax, dark. Legs dark, tibiae and tips of femora
pale.

Pronotum as in figure 21P. Genitalia as in figure 11, emargination of
dorsal plate narrowing to apex, sides not sinuate distally; basophyses emerging
close together, subparallel, tips recurving or enlarged on proximal side. In
ventral view, upper margins of lateral incisure of tegmen not subangulate
distally. Length 5 mm.

VaRIATIONS. The dark pronotal borders vary by reduction, leaving only
the apical and basal narrowly dusky. The holotype, described above, was
selected because of the clear visibility of the genitalic structure. Normally the
elytra are more or less broadly black or dusky at apex and sides, the lateral
dark area narrowing anteriorly and not reaching the base. The sutural bead is
usually dark except near the scutellum.

The genitalia vary considerably. The emargination of the dorsal plate,
normally V-shaped, sometimes has the sides sinuate as in S. flavida. Occasional
examples occur in which a slight angulation is apparent at about distal third
of the upper margins of the lateral incisure of the tegmen, viewed ventrally.
The formation of the median lobe of the aedeagus offers the most decisive
character for separating this species from S. flavida, but unfortunately this
structure is often distorted and inconclusive. Length 5.5—6.5 mm.

Typical specimens of S. ldasseni, from the Cascade Range, and _ typical
examples of S. flavida, from the Sierra Nevada Mountains, are readily identi-
fiable. Possibly cross breeding has occurred where their habitats overlap,
resulting in hybrids that cannot be confidently assigned to either species.

DisTRIBUTION. CALIFORNIA: Modoc County: Blackmore, VII-3-50, M.
Wasbauer, 1 paratype (UCB). Shasta County: Shingletown, V-24-11, C. D.
Michener, 1 paratype (UCB). Lassen National Park, V1-28, 29-60, D. J. and
J. N. Knull, 5 paratypes, 6 females (OSU); same, Kelly’s Resort, VI-14-31,
Van Dyke, 1 paratype (CAS); same, Manzanita Lake, VI-8-41, 1 paratype
(Fender); same, Manzanita Camp, VI-18-60, 1 paratype (Edwards). Mt.
Lassen, VII-5-63, D. J. and J. N. Knull, 2 females (OSU); same, 3 mi. SE.,
VII-8-55, D. L. Dahlston, 1 paratype (UCD). Manzanita Lake, VI-12-41, C.
Michener, 1 paratype (CNC). Facht, VI-24, 27-22, V-3-23, J. O. Martin,
4 paratypes (CAS). Duck Lake, V-8-21, J. O. Martin, holotype (CAS).
Plumas County: Bucks Lake, VII-1-49, W. R. Schreader, 1 paratype (UCD);
same, Clover Valley, VI-17-23, J. O. Martin, 1 paratype (CAS); same,
Chester, VI-7, 13-60, D. J. and J. N. Knull, 4 paratypes, 11 females (OSU).
Mineral, 5 mi. E., VI-9, 26-60, D. J. and J. N. Knull, 24 paratypes (OSU 22,
CAS 2), 46 females (OSU). Sierra County: Webber Lake, VII-2-59, Linsley,
1 paratype (UCB). Placer County: Baxters, V1-3-45, A. T. McClay, 1 para-
type (UCD).

474 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

(12) Silis (Silis) nevadica Green, new species.

Hototypr, male; Carson City, Nevada, V-12-41, A. T. McClay. In col-
lection of University of California at Davis.

Body and appendages black, pronotum pale fulvous with all borders black.

Pronotum as in figure 21P. Genitalia as in figure 12, dorsal plate with
broad U-shaped emargination, basophyses emerging widely separated, con-
verging apically, tips prominently enlarged on proximal side in lateral view.
In ventral view, upper margins of lateral incisure of tegmen strongly angulate
at about distal third. Length 6.5 mm.

DistRIBUTION. NEVADA: Carson City, V-12-41, A. T. McClay, holotype.
1 paratype (UCD).

(13) Silis (Silis) tenuata Green, new species.

Hototype, male; Giant Forest, Tulare County, California, 6,500 ft.,
VII-1915, G. Hopping. In collection of California Academy of Sciences. CAS
type number 9083.

Antennae dark, four basal segments pale beneath; palpi dark. Head black,
pale in front. Pronotum flavate, anterior border blackish medially, posterior
border narrowly dusky. Scutellum black. Elytra flavate, broadly dusky at
apex and sides, lateral dark area narrowing anteriorly, not reaching base.
Underside black, prothorax pale, ventral segments with paler lateral and apical
borders. Legs pale, coxae and femora, except at apex, black; profemora largely
pale, metafemora largely black.

Pronotum as in figure 21P. Genitalia as in figure 13, basophyses with
exposed tips narrowing to very slender apically. In ventral view, upper margins
of lateral incisure of tegmen strongly angulate at about distal third. Median
lobe of aedeagus broad in dorso-ventral view, sides not striate. Ventral lobe
of tegmen with distinct apical notch, sides subparallel. Length 6.5 mm.

VARIATIONS. The pronotum may have the lateral borders also dark. Speci-
mens occur at high altitudes that are black with only the mandibles pale, or
with the elytra partly fusco-flavate. Length 5.5—6.5 mm.

DisTRIBUTION. CALIFORNIA: Fresno County: Huckleberry Meadow,
6,500 ft., V-25-10, R. Hopping, 1 paratype; V-VII, 1 paratype (CAS).
Round Meadow, Giant Forest, 6,400 ft., VII-1915. R. Hopping, 1 paratype
(CAS). Fresno County: Piute Creek, VII-6-52, 6,200 ft., Peter Raven, 1
paratype (CAS). Fresno County: Cedar Grove, V-25-11, R. Hopping, 1
paratype (CAS). Tulare County: Giant Forest, 6,500 ft., VII-1915, G. Hopping,
holotype (CAS). Fresno County: McGee Creek, VII-10-52, 11,000 ft., Peter
Raven, 1 male, 1 female (CAS). Huntington Lake, V1-26-46, VI-28-40, A. T.
McClay, 2 paratypes (UCD); same, Badger Flat, VII-20, 22-37, VII-27-48,
A. T. McClay, 3 paratypes (UCD). Tulare County: Dinuba, VI-19-57, A. T.
McClay, 1 paratype (UCD). Above Lundy, VII-9, 9-11,000 ft., Wickham, 1

VoL. XXXII] GREEN: REVISION OF THE NEARCTIC SILIS 475

paratype (USNM). Shaver Lake, VI-3-37, 1 paratype (Fender). /nyo County:
Mono Pass, VIII-11-63, R. M. Brown, 1 paratype (CAS).

(14) Silis (Silis) carbo Van Dyke.
Silis difficilis carbo VAN Dyke, 1918, New York Ent. Soc., Jour., vol. 26, p. 168.

Body black, underside of prothorax more or less pale anteriorly each side.
Varying with pronotum flavate with black borders, disk sometimes partly
blackish, elytra dark fuscous; or with both pronotum and elytra in large
part flavate, as in S. flavida.

Pronotum as in figure 21P. Genitalia as in figure 14, exposed tips of baso-
physes normally broad. In ventral view, upper margins of lateral incisure of
tegmen angulate at about distal third. In dorso-ventral view, median lobe of
aedeagus narrowing to apex; sides longitudinally striate. Ventral lobe of
tegmen without, or with very minute, apical notch; sides converging apically.
Length 5—6.5 mm.

DIsTRIBUTION. CALIFORNIA: Fresno County: Rea Lake, 10,500 ft.;
Bubbs Creek Canyon, Kings River, 9,700 ft.; 60 Lake Basin, 10,500 ft.;
Bullfrog Lake, 10,600 ft.; East Lake, 10,000 ft.; Huntington Lake, 7,000 ft.
Tulare County: Rattlesnake; Atwells Mill; Colony Road; Siberian Outpost,
10,500 ft. Kaweah, 7,500 ft. Monarch Lake. Mt. Mitchell. Sequoia National
Park, Alta Peak, 9,500 to 11,000 ft. Kings Canyon National Park, Deadman
Canyon. Mt. Whitney, 11,000 ft. Inyo County: Lone Pine Lake; Monache;
Whitney Portal, 11,500 ft.

This species was described by Van Dyke as a color variety of S. difficilis.
It is raised to specific rank because of its isolated habitat, accompanied by
apparently constant structural differences in the tegmen, namely, the narrower
ventral lobe lacking the apical notch, and the subangulate upper margins of the
lateral incisure. The available data indicates that the all-black phase occurs
only at altitudes of ten thousand feet or higher.

(15) Silis (Silis) oregonensis Green, new species.

HoLotyrr, male; Warner Mountains, Lake County, Oregon, VI-18-22,
Van Dyke. In collection of California Academy of Sciences. CAS type number
- 9084.

Black, pronotum fulvous with all borders narrowly black.

Pronotum as in figure 21P. Genitalia as in figure 15. In lateral view, tips
of basophyses straight, rather broad, not narrowing apically. In ventral view,
upper margins of lateral incisure of tegmen not angulate at distal third. Emargi-
nation of dorsal plate broadly V-shaped, attaining membraneous area. In
dorsoventral view, median lobe of aedeagus broad, with small apical notch,
sides not striate. Length 6.5 mm.

476 CALIFORNIA ACADEMY OF SCIENCES | Proc. 47H Ser.

VoL. XXXII] GREEN: REVISION OF THE NEARCTIC SILIS 477

VARIATIONS. The elytra may be partially flavate, as in many S. difficilis ;
or they may be largely flavate, as in S. flavida. The apical notch of the median
lobe of the aedeagus is often not apparent, due to drying distortion.

DISTRIBUTION. OREGON: Lake County: Warner Mountains, VI-16-22,
Van Dyke, holotype, 3 paratypes (CAS); Klamath County: Williamson River
near head, V-30-59, Joe Schuh, 1 paratype (Fender); 11 mi. NE. of Bly,
VI-7-59, Joe Schuh, 5 paratypes (Fender); 20 mi. N. of Beatty, Sycan River,
V-30-59, Joe Schuh, 6 paratypes (Fender); 7 mi. W. of Kano, 3,000 ft.,
V-24-58, K. M. Fender, 1 paratype (Fender); 11 mi. NE. of Bly, V-5-62,
Scott and Schuh, 3 paratypes (Schuh); Pelican Butte, VII-8-60, 2 paratypes
(Schuh); Sun Creek, VI-14-62, J. D. Vertrees, 1 paratype (Schuh); Fremont
National Forest, 5,000 ft., VI-17-22, Van Dyke, 1 paratype (CAS); Sun
Mountain, VI-11-56, J. D. Vertrees, 1 paratype (Schuh); 1 mi. N. Ft. Klamath
Junction, V-25-58, R. K. Eppley, 1 paratype (Edwards). Upper Klamath Lake,
Three Mile Creek, V-30-60, Joe Schuh, 3 paratypes (Schuh). Ochoco National
Forest, V1-14-41, Fender, 1 paratype, 1 male (Fender); same, Wildwood,
V-26-50, 1 paratype (Fender). Crater Lake, 7,000 ft., VII-16-22, Van Dyke,
1 paratype (CAS); VII-14-38, A. T. McClay, 1 paratype (UCD); same,
Park HQ. D. H. Huntzinger, 3 paratypes (Edwards); same, Garfield Peak,
VI-27-57, D. H. Huntzinger, 2 paratypes (Edwards). Jackson County: Butte
Falls, V-22-41, 1 paratype (UCD). Douglas County: Kelsey Valley, V1-20-62,
J. D. Vertrees, 2 paratypes (Schuh). Anthony Lake, VI-12-39, 1 paratype
(Fender). Bear Springs, V1-6-39, V-4, 26-40, Fender, 4 paratypes (Fender).
Odell Lake, 3 mi. SE., VI-8-41, Schuh and Gray, 1 paratype (Schuh). CALI-
FORNIA: Trinity County: Carville, VI-16-13, Van Dyke, 1 paratype

Ficures 13-21. Male genitalia; ventral, lateral, and dorsal views, arranged in that
order from left to right.

FicureE 13. Silis (Silis) tenuata Green, holotype.

Ficure 14. Silis (Silis) carbo Van Dyke.

Ficure 15. Silis (Silis) oregonensis Green, holotype.

Ficure 16. Silis (Silis) striatella Green, holotype.

_ Ficure 17. Silis (Silis) difficilis LeConte.

Ficure 17a. Silis (Silis) difficilis occidens Green, holotype.

Figure 17b. Silis (Silis) difficilis subspecies ?

Ficure 18. Silis (Silis) atra LeConte.

Ficure 19. Silis (Silis) egregia Green, holotype. Diagram of posterior orifice of tegmen
inserted.

Ficure 20. Silis (Silis) singularis Green, holotype. Diagram of posterior orifice of teg-
men inserted.

Ficure 21. Silis (Silis) eximia Green, holotype. Diagram of posterior orifice of tegmen
inserted.

478 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

(CAS); Siskiyou County: Paynes Meadow, VII-5-60, D. Q. Cavagnaro, 1
paratype (UCD).

(16) Silis (Silis) striatella Green, new species.

Hototyee, male; Paradise Valley, Mt. Rainier, Washington, VII-24-20,
Van Dyke. In collection of California Academy of Sciences. CAS type number
9085.

Head and antennae black. Pronotum pale fulvous with all borders black.
Scutellum black. Elytra bicolored, broadly fulvous at base, pale area narrow-
ing distally, extending to about apical fourth, its margins obscure; sutural
bead, sides, and apex dark fuscous. Underside and legs mostly dark.

Pronotum as in figure 21P. Genitalia as in figure 16. Emargination of
dorsal plate rather narrowly U-shaped, deeper than wide, sides subparallel or
somewhat diverging, not curving inward distally, emargination not attaining
membraneous area by a conspicuous distance. Upper margins of lateral incisure
of tegmen not angulate at distal third. Median lobe of aedeagus with sides
longitudinally striate. Length 6 mm.

VariaTIONS. The color of the elytra varies from entirely black to almost
entirely fulvous. Rarely the sides of the dorsal plate emargination curve some-
what inward distally. Length 6—7.5 mm.

DISTRIBUTION. BRITISH COLUMBIA: Midday Valley, Merritt, V-12,
27-26, E. Rendell, 2 paratypes; V-17-25, J. Stanley, 1 paratype; VI-4-23,
R. Hopping, 1 paratype; V-24-23, R. Hopping, 1 paratype (CAS). Same,
VI-4, 7-23, R. Hopping, 2 paratypes (CNC). WASHINGTON: M¢. Rainier:
Paradise Valley, VII-24-20, Van Dyke, holotype, 12 paratypes (CAS);
Paradise Park, 6,000 ft., VII-5, 31-05, Van Dyke, 2 paratypes (CAS);
Paradise, VII-1, 4-34, O. Bryant, 8 paratypes (CAS); Longmire Springs,
2,500 ft., VII-25-19, Blaisdell, 1 paratype (CAS). Mt. Rainier, VII-22-40,
1 paratype (Fender). Rainier National Park, Sunrise Peak, VI1-24-36, Van
Dyke, 1 paratype (CAS). Pierce County, Fort Lewis, V-3-46, P. H. Arnaud,
2 paratypes (CAS). Rimrock, VI-8-54, H. P. Lanchester, 2 paratypes (Fender).
Spirit Lake, VII-22-56, 3,800 ft., J. D. Lattin, 1 paratype (OSU).

OREGON: Bear Springs, VI-6, 7-39, V-26-40, K. M. Fender, 6 paratypes
(Fender). Mt. Hood, Homestead Inn, VI-30-27, Van Dyke, 1 paratype
(CAS). Mt. Hood, 3,000-6,000 ft., VI-23-25, Van Dyke, 3 paratypes (CAS).
Kirby, 1V-27-37, Fred Lawrence, 1 paratype (CAS). Ochoca National Forest,
VI-13-41, 1 paratype (Fender). Swim, VII-2-42, Schuh and Gray, 1 paratype
(Schuh). Parkdale, VII-1-38, Schuh and Gray, 1 paratype (Schuh). Klamath
County, Little Deschutes River, VI-5-58, 5 paratypes (Schuh). Lamola-
Tokatec Falls, North Umpqua River, V1-26-62, 1 paratype (Schuh).

(17) Silis (Silis) difficilis LeConte.

Silis difficilis LeContr, 1850, In: Lake Superior, by Louis Agassiz, p. 230.

VoL. XXXII] GREEN: REVISION OF THE NEARCTIC SILIS 479

Head black. Pronotum flavate or fulvous with all borders black, varying
by reduction but never entirely pale in the male. In one example median
extensions of front and rear black borders extend on disk. Scutellum black.
Elytra flavate with apex and sides more or less black or fuscous; varying to
entirely dark with more or less extensive pale color nearly always evident
under brilliant illumination. Underside and legs mostly black.

Pronotum as in figure 21P. Genitalia as in figure 17. Emargination of
dorsal plate broadly V- or U-shaped, about as deep as wide, not attaining
membraneous area, sides curving inward distally. Basophyses usually ex-
tending only slightly through emargination of dorsal plate, narrowing apically
in lateral view. Upper margins of lateral incisure of tegmen not angulate at
distal third. Median lobe of aedeagus with sides longitudinally striate. Length
5—6.5 mm.

DISTRIBUTION. QUEBEC: Gaspe County; Duparquet. SASKATCHEWAN:
Rockglen; Attons Lake; Saskatoon; Cypress Hills; Prince Albert; Bounty;
Beaver Creek; Katepwa. ALBERTA: McMurry; Lethbridge; Waterton, Elk-
water; Banff; Edmonton; Olds; Pinehen Creek. BRITISH COLUMBIA:
Atlin; Trinity Valley; Oliver; Creston; Cranbrook; Vernon; Adams Lake.
MICHIGAN: Marquette. WISCONSIN: Sauk County. SOUTH DAKOTA:
Custer; Spearfish Canyon; Blue Bell, NEW MEXICO: Jemez Mountains;
Lincoln County. WYOMING: Yellowstone National Park; Curtis Canyon near
Jackson; Teton National Park; Sublette County; Wind River Range. MON-
TANA: Glacier National Park; Mineral County; Fergus County; Big Snowy
Peak. IDAHO: Wallace; Twin Creek Camp; Moscow; Stanley; Targhee Pass;
Baker Creek NW. of Ketchum. WASHINGTON: Easton. OREGON: Lake
Wallowa; Meacham; Pine Creek, Baker County, Tollgate, Blue Mountains;
Durkee; Jackson County; Pinehurst; Butte Falls; Ashland; Buckhorn Mineral
Springs; Deschutes County between Suttle Lake and Sisters.

This very widely distributed species is undoubtedly a complex as it is here
recognized. It has not been found possible to segregate definitely recognizable
taxa, with the exception of the somewhat unstable subspecies following. Ex-
ceptions to the bicolored elytra of S. difficilis are rare, and occur mostly in
Idaho, Montana, and Wyoming. Some Oregon examples with an unusually wide
smooth band separating the apex of dorsal plate emargination from the mem-
braneous area, have the elytra usually entirely black.

(17a) Silis (Silis) difficilis occidens Green, new subspecies.
HOLOTYPE, male: 7 mi. W. of Westgard Pass, Inyo County, California,
VI-26-53, W. D. McClellan. In collection of University of California at Davis.

Deep black, pronotum fulvous with all borders black, elytra without trace
of paler coloration.

480 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Pronotum as in figure 21P. Genitalia as in figure 17a, similar to S. diffi-
cilis, differing in basophyses extending well beyond emargination of dorsal
plate, somewhat expanding apically as seen in lateral view, emargination
attaining membraneous area. Length 5.5 mm.

Variations. No significant variation is noted in this subspecies as it
occurs in California east of the Sierra Nevada Range. Progressing eastward
through northern Arizona, Utah, and Colorado, the characteristic genitalic
distinctions become less obvious, and no paratypes have been designated except
from the California area. There are few exceptions to the uniform deep black
coloration of the elytra. Length 4.25-6 mm.

DIsTRIBUTION. CALIFORNIA: /nyo County: 7 mi. W. of Westgard Pass,
VI-28-53, W. D. McClellan, holotype, 4 paratypes (UCD); Big Pine Creek,
8,000 ft., V-19-47, R. M. Bohart, 2 paratypes (UCD); Lone Pine, VI-18-37,
N. W. Frazier, 2 paratypes (UCB); Upper Big Pine Creek, 8,500—9,500 ft.,
VII-1-60, W. F. Barr, 2 paratypes (UId). Mono County: Blanco’s Corral,
White Mountains, 10,000 ft., VI-20, 23-53, VII-7-53, J. W. McSwain, 8 para-
types (UCB); same, VI-23, 29-53, 5 paratypes (UCD); same, VI-23-53, H.
Nakahihara, 2 paratypes (UCR); White Mountains, 10,000 ft., VI-19, 26-61,
Buckett, Miller, and Lange, 7 paratypes (UCD 6, CAS 1). Siskiyou County:
Macdoel, 1V-28-49, Joe Schuh, 1 paratype (Schuh). Gilbert Lake to Indepen-
dence, VII-17, Wickham, 1 paratype (USNM). Bubbs Creek, VII-29, 8,000
ft., Wickham, | paratype (USNM). ARIZONA: White Mountains; Diamond
Creek; Grand Canyon; San Francisco Mountains; 20 mi. S. of Jacobs Lake;
Williams; Flagstaff. NEVADA: Elko County: Green Mountain Creek; Angel
Lake, 12 mi. SW. Wells. UTAH: Springville; Ft. Duchesne; Kanesville; Mill-
ville; Indian Canyon; Utah Experiment Station; Mapleton; Providence;
Daniel Pass, 2 mi. E. of Strawberry; Salt Lake County: Dry Canyon; Morgan
County: E. Canyon Dam; Wasatch County: Heber; Salt Lake City; Logan
Canyon; Cowley Canyon; Card Canyon; Y Mountain, Utah County; Vine-
yard; Navajo Mountain. COLORADO: Coal Creek; Longs Peak Inn; Colorado
Agricultural College; Denver; Colorado Springs; Boulder; Camp Creek Rail-
road Station; Chimney Gulch; Glenwood.

In a series from French Glen, Oregon (Fender), the tips of the basophyses
are variably recurved or enlarged on proximal side, as viewed laterally. They
may represent another subspecies, and have been labeled “Silis near occidens.”

(18) Silis (Silis) atra LeConte.

Silis atra LECONTE, 1884, Trans. Amer. Ent. Soc., vol. 12, p. 22.

Black, elytra varying to dark rufo-brunneous.

Pronotum as in figure 22P. Genitalia as in figure 18. Dorsal plate simple,
with deep and very wide emargination. Laterophyses with small inconspicuous
appendix. With single basophysis furcate submedially. Length 4—5 mm.

VoL. XXXII] GREEN: REVISION OF THE NEARCTIC SILIS 481

DISTRIBUTION. OREGON: Odell Lake; Mt. Hood; Tilly Jane Creek;
near Grant Camp; Summit Meadow, Homestead Inn; Hood River Meadow,
Three Sisters, Frog Camp; Diamond Lake; Klamath County, Summit Lake ;
Wascoe County, Bear Springs; Marion County, Park Butte; Lake County,
Beaver Marsh; Mt. Jefferson. WASHINGTON: Mt. Rainier, Sunrise; Mt.
Adams, Bird Creek; Rimrock; Lake Cle Elum. NORTHWEST TERRITORY:
Horse Lake Area.

(19) Silis (Silis) egregia Green, new species.

Hototyper, male; Warners Hot Springs, California, IV-6-40, G. P. Mac-
kenzie. In collection of California Academy of Sciences, on indefinite loan
from K. M. Fender. CAS type number 9086.

Black, prothorax pale fulvous above and _ beneath.

Pronotum as in figure 23P. Genitalia as in figure 19. Dorsal plate feebly
trilobed, posterior orifice of tegmen not constricted below it. Ventral lobe
extending well beyond dorsal plate. Laterophyses with small inconspicuous
distal appendix. With single basophysis furcate submedially. Length 5.5 mm.

DiIsTRIBUTION. CALIFORNIA: Warners Hot Springs, 1V-6-40, G. P. Mac-
kenzie, holotype (CAS). Mission Vallev, I1I-8-28, 1 paratype (SDNH).

(20) Silis (Silis) singularis Green, new species.

HoLotype, male; Lebec, California, altitude 4,000 ft., V-15-25, J. O.
Martin. In collection of California Academy of Sciences. CAS type number
9087.

Black, prothorax pale fulvous above and beneath.

Pronotum as in figure 23P. Genitalia as in figure 20. Dorsal plate trilobed,
side lobes prominent, extending far beyond feebly lobed median part. Posterior
orifice of tegmen strongly constricted below dorsal plate. Ventral lobe of
tegmen and dorsal plate subequal in length. Laterophysis (one missing in
holotype) with tip everted and squarely truncate, surpassed by foliately ex-
panded (in lateral view) distal appendix. With single basophysis furcate sub-
medially. Length 8 mm.

DISTRIBUTION. CALIFORNIA: Lebec, 4,000 ft., V-15-25, J. O. Martin,
_ holotype (CAS).

(21) Silis (Silis) eximia Green, new species.

Hoiotyper, male; Lockwood Creek near Stauffer Post Office, Ventura
County, California, V-7-59, G. I. Stage. In collection of California Academy
of Sciences, on indefinite loan from the University of California at Berkeley.
CAS type number 9107.

Black, prothorax pale rufous above and beneath.

482 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Pronotum as in figure 23P. Genitalia as in figure 21. Dorsal plate trilobed,
side lobes extending moderately beyond median part, which has a shallow
broadly angulate emargination. Posterior orifice of tegmen constricted below
dorsal plate. Ventral lobe of tegmen not extending as far posteriorly as dorsal
plate. Laterophyses with tips everted and squarely truncate, surpassed by tip
of foliately expanded (in lateral view) distal appendix. With single basophysis
furcate submedially. Length 6.5 mm.

DISTRIBUTION. CALIFORNIA: Lockwood Creek, near Stauffer Post
Office, Ventura County, V-7-59, G. I. Stage, holotype (CAS). Pine Creek,
Alamo Mountain, Ventura County, V-6-59, C. W. O’Brien, 1 female, pre-
sumably of this species (UCB).

(22) Silis (Silis) abstrusa Green, new species.

Hototype, male; San Bernardino County, California, 6 mi. NNE. of
Hesperia, IV-16-62, MacNeill, Rentz, Brown, and Lundgren. In collection of
California Academy of Sciences. CAS type number 9088.

Black, prothorax pale fulvous above and beneath.

Pronotum as in figure 23P. Genitalia as in figure 22. Dorsal plate trilobed,
side lobes extending far beyond median part which is deeply triangular emar-
ginate. Posterior orifice of tegmen constricted below dorsal plate. Ventral lobe
of tegmen slightly shorter than dorsal plate. Laterophyses with tips acute,
slightly diverging, surpassed by small lineate distal appendix. With single
basophysis furcate submedially. Length 6.5 mm.

DISTRIBUTION. CALIFORNIA: San Bernardino County, 6 mi. NNE. of
Hesperia, 1V-16-62, MacNeill et al., holotype (CAS).

(23) Silis (Silis) arida Green, new species.

HoLotype, male; Isabella, California, IV-5-14. In collection of American
Museum of Natural History.

Black, prothorax pale fulvous above and beneath.

Pronotum as in figure 23P. Genitalia as in figure 23. Dorsal plate trilobed,
side lobes extending well beyond median part which is prominently lobed.
Posterior orifice of tegmen constricted below dorsal plate. Ventral lobe of
tegmen slightly shorter than dorsal plate. Laterophyses with tips acute, not
diverging, extending about as far posteriorly as tips of large foliately expanded
(in lateral view) distal appendix. With single basophysis furcate submedially.
Length 7 mm.

DiIsTRIBUTION. CALIFORNIA: J/sabella, IV-5-14, holotype (AMNH).
(24) Silis (Silis) ursina Green, new species.

HoLotypPe, male; Bear Valley, California, VI-6-14, R. S. Woglum. In
collection of California Academy of Sciences on indefinite loan from the Uni-
versity of California at Riverside. CAS type number 9165.

VoL. XXXII] GREEN: REVISION OF THE NEARCTIC SILIS 483

Black, prothorax pale fulvous above and beneath.

Pronotum as in figure 23P. Genitalia as in figure 24. Dorsal plate trilobed,
deeply emarginate, side lobes extending well beyond subtruncate median part.
Ventral lobe of tegmen slightly shorter than dorsal plate. In dorso-ventral
view, laterophyses arcuately converging distally, tips reversing, acute; appendix
inconspicuous, setiform. With single basophysis furcate submedially. Median
lobe of aedeagus with horizontally flattened apical process. Length 7 mm.

DISTRIBUTION. CALIFORNIA: Bear Valley, VI-6-14, R. S. Woglum,
holotype (UCR). The exact location of Bear Valley, in southern California,
could not be determined.

(25) Silis (Silis) deserticola Van Dyke.
Silis deserticola VAN Dyxe, 1918, Jour. New York Ent. Soc., vol. 26, p. 173.

Black, prothorax pale fulvous above and beneath.

Pronotum as in figure 23P. Genitalia as in figure 25. Dorsal plate trilobed,
feebly emarginate, side lobes extending but little beyond truncate median part.
Posterior orifice of tegmen prominently constricted below dorsal plate. Ventral
lobe of tegmen slightly shorter than dorsal plate. Laterophyses with tips acute,
not diverging, surpassed by moderately expanded (in lateral view) distal
appendix. With single basophysis furcate submedially. Length 7 mm.

DISTRIBUTION. CALIFORNIA: Argus Mountains, May, holotype, and 1
female (USNM). The holotype is USNM number 21695.

(26) Silis (Silis) incongrua Green, new species.

HoLotype, male; Santa Cruz Mountains, California, Koeble. In collection
of California Academy of Sciences. CAS type number 9089.

Head blackish, paler in front, antennae and palpi dusky. Pronotum flavate.
Scutellum black. Elytra flavate, sutural bead slightly darker, apices black.
Beneath, except prothorax, dark. Legs flavate, middle and hind femora
basally, and hind tibiae apically, dark.

Pronotum similar to figure 25P except front margin of posterior process
not angulate. Genitalia as in figure 26. Dorsal plate not emarginate, posterior
margin convex, preceded by broad sclerotized area. Laterophyses long, slender
_ throughout, curving strongly downward and feebly inward. With two baso-
physes. Length 5.25 mm.

DISTRIBUTION. CALIFORNIA: Santa Cruz Mountains, Koeble, holotype
(CAS).

(27) Silis (Silis) angulata Green, new species.

Hototyere, male; Adams Springs, Lake County, California, F. E. Blaisdell.
In collection of California Academy of Sciences. CAS type number 9090.

VoL. XXXII] GREEN: REVISION OF THE NEARCTIC SILIS 485

Head black, clypeus pale, antennae dusky; palpi dark (ex paratype).
Pronotum flavate. Scutellum black. Elytra flavate, tips black, sutural bead
brunneous. Beneath, except prothorax, dark. Legs flavate, femora largely
dark, pale apically.

Pronotum similar to figure 24P. Genitalia as in figure 27. Dorsal plate
with broad shallow emargination attaining membraneous area. Laterophyses
long, moderately stout, nearly straight, with prominent exterior tooth at about
distal third, tips diverging. With two basophyses failing by far to attain apex
of dorsal plate emargination. Length 6 mm.

DISTRIBUTION. CALIFORNIA: Lake County: Adams Springs, F. E. Blais-
dell, holotype (CAS). Mark West Springs, V-11-30, E. P. Van Duzee, 1
paratype (CAS). Mt. St. Helena, V-12-26, E. P. Van Duzee, 1 paratype
(CAS); same, V-10-20, S. F. Bailey and E. J. Taylor, 1 paratype (UCD).

(28) Silis (Silis) emarginata Green, new species.

Hototypr, male; Yosemite Valley, California, V-26-25, Blaisdell. In
collection of California Academy of Sciences. CAS type number 9091.

Head black, pale in front; antennae dusky, paler beneath basally; terminal
palpal segments dark. Pronotum flavate. Scutellum black. Elytra entirely
flavate. Beneath dark, front of head and prothorax pale. Legs flavate, tarsi
darker, front coxae pale, others largely dark.

Pronotum as in figure 24P. Genitalia as in figure 28. Dorsal plate with
large deep U-shaped emargination entering membraneous area. Laterophyses
moderately long, curving upward and inward, tips widened and everted. With
two basophyses extending beyond apex of dorsal plate emargination. Length
6.5 mm.

Ficures 22-31. Male genitalia; ventral, lateral, and dorsal views, arranged in that
order from left to right.

Ficure 22. Silis (Silis) abstrusa Green, holotype. Diagram of posterior orifice of tegmen

inserted.

Ficure 23. Silis (Silis) arida Green, holotype. Diagram of posterior orifice of tegmen
inserted.

Ficure 24. Silis (Silis) ursina Green, holotype. Diagram of posterior orifice of tegmen
inserted.

Ficure 25. Silis (Silis) deserticola Van Dyke, holotype. Diagram of posterior orifice of
tegmen inserted.

Ficure 26. Silis (Silis) incongrua Green, holotype.

Figure 27. Silis (Silis) angulata Green, holotype.

Ficure 28. Silis (Silis) emarginata Green, holotype.

Ficure 29. Silis (Silis) fenderi Green, holotype.

Ficure 30. Silis (Silis) subtruncata Green, holotype.

Ficure 31. Szlis (Silis) macclayi Green, holotype.

486 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Variations. Rarely the elytra are tipped with black, the sutural bead re-
maining pale almost without exception. Only one individual has been seen
with the sutural bead slightly darkened. The middle and hind femora and
hind tibiae are sometimes partly dark. The dorsal plate emargination varies
greatly to widely V-shaped with the lobes each side rather acutely angulate.
Length 4.5—-6.5 mm.

DisTRIBUTION. CALIFORNIA: Plumas County: Quincy, VI-5-63, G.
Leskey, 4 paratypes (UCD). Placer County: Emigrant Gap, V1-13-39, Cazier,
1 paratype (AMNH). &/ Dorado, 1 paratype (ANSP). Tallac, VI-1899, Van
Dyke, 1 paratype (CAS). Calaveras County: Murphy’s Canyon, 2,500 {ft.,
V-15-36, Blaisdell, 1 paratype (CAS). Tuolumne County: Basin Creek, 5 mi.
S. of Tuolumne City, V-31-64, Lundgren, 1 paratype (CAS). Yosemite, VI,
1 paratype (Fender); same, 4,000 ft., VI-17-28, VI-12-31, V-20-31, E. O.
Essig, 3 paratypes (UCB). Yosemite National Park, V1-27-57, P. J. Santana,
1 paratype (Edwards). Yosemite Valley, V-22-21, 2 paratypes (UCB); same,
V-26-25, Blaisdell, holotype (CAS). Akwahnee, V, Fenyes, 1 paratype (CAS).
Mariposa County: Miami Ranger Station, V-17, 23-42, C. Kennett, 2 para-
types (UCB); same, V-27-42, W. W. Allen, 2 paratypes (UCB). Mariposa,
VI, Wickham, 1 paratype (USNM). Madera County, Coarsegold, V-12-42,
C. Kennett, 1 paratype (UCB). Fresno County: Cedar Grove, V-25-11, R.
Hopping, 2 paratypes (CAS); Huckleberry Meadows, 6,500 ft., VII-19-17, R.
Hopping, 1 paratype (CAS); Sierra National Forest, Stevenson Creek, V-29-
15, R. Hopping, 1 paratype (CAS); Dalton Creek, 4,800 ft., V-6-20, H.
Dietrich, 1 paratype, 1 female (CU); Bubbs Creek Canyon, Kings River,
9,700 ft., VI-8-10, Van Dyke, 1 paratype (CAS). Tulare County, VI-12-39,
Nunenmacher, 1 paratype (CAS); same, Kaweah, IV-12-31, 1 paratype
(Fender). Kaweah, R. Hopping, 1 paratype (CAS); same, VII-1-35, Fred
Lawrence, 3 paratypes (CAS); same, IV-12-31, 3 paratypes (SDNH); same,
1 paratype, 1 male, 3 females (UK). Sequoia National Park, V-30-29, VI-13-29,
A. T. McClay, 2 paratypes (UCD); same, VI-27, Edith Mank, 1 paratype,
3 females (CU). North Fork, V-23-20, H. Dietrich, 1 paratype (CU). Kern
County, Hubbard and Schwarz, 1 paratype (USNM). Monarch Lake, VI-13-
13, Van Dyke, 1 paratype (CAS). Sugar Pine, Fenyes, 1 paratype (CAS).
Atwell Mill, V-30-29, A. T. McClay, 1 paratype (UCD). Mono County:
W. Walker River, 6,000 ft., N. W. Frazier, 1 paratype (UCB). Southern Cali-
fornia, Horn collection, 1 paratype (ANSP). Monterey County: Stone Canyon,
IV-21, 27-19, Van Duzee, 3 paratypes (CAS). San Benito County: Waltham
Creek, V-11-09, R. Hopping, 2 paratypes (CAS). Diablo Range, 2,000—4,000 ft.,
V-12-52, O. Bryant, 1 paratype (CAS).

(29) Silis (Silis) fenderi Green, new species.

Ho.otypPer, male; Peavine Ridge, near McMinnville, Oregon, VI-9-48, K.

VoL. XXXII] GREEN: REVISION OF THE NEARCTIC SILIS 487

M. Fender. In collection of California Academy of Sciences on indefinite loan
from K. M. Fender. CAS type number 9092.

Head black, pale in front; antennae and terminal palpal segments black.
Pronotum flavate. Scutellum black. Elytra flavate, tips black, sutural bead
brunneous. Beneath dark, except prothorax and head in front. Legs and coxae
largely black, tips of femora, protibiae, and base of other tibiae, pale.

Pronotum similar to figure 24P. Genitalia as in figure 29. Dorsal plate
with rather deep triangular emargination attaining membraneous area. Latero-
physes extending somewhat beyond dorsal plate, tips subrectangularly everted
for a considerable distance. With two basophyses not extending as far poster-
iorly as apex of dorsal plate emargination. Length 5.5 mm.

VARIATIONS. The dorsal plate emargination varies in depth, and sometimes
does not attain the membraneous area. The everted tips of the laterophyses
may be either acute or blunt, and the angle of eversion may be somewhat
obtuse. In one example the sutural bead of the elytra is not appreciably darker.
Length 5—6 mm.

DISTRIBUTION. OREGON: Peavine Ridge, near McMinnville, V1-9-48, K.
M. Fender, holotype (CAS); same, VI-11-46, VI-5-45, 2 paratypes (Fender).
Matulius River, VI-13-47, 1 paratype (Fender). Corvallis, VI-12-25, Van
Duzee, 1 paratype (CAS); same, VII-3-46, K. R. Hobbs, 1 paratype (OrSU).
Josephine County: Selma, 3.5 mi. W., V-27-62, Joe Schuh, 1 paratype (Schuh).
CALIFORNIA: Blocksburg, V-18-35, E. W. Baker, 1 paratype (AMNH).
Humboldt County, V1-47, Bryant, 1 paratype (CAS).

(30) Silis (Silis) subtruncata Green, new species.

Ho.otype, male; Fairfax, Marin County, California, V-7-11, Van Dyke.
In collection of California Academy of Sciences. CAS type number 9093.

Head black, clypeus pale, antennae and palpi dusky. Pronotum flavate.
Scutellum black. Elytra flavate, tips black, sutural bead brunneous. Beneath,
except prothorax, dark. Legs largely dark, front tibiae and base of other
tibiae pale.

Pronotum similar to figure 24P. Genitalia as in figure 30. Dorsal plate with
very shallow arcuate emargination, as seen in direct dorsal view, failing to
- attain membraneous area by a conspicuous distance. Laterophyses not sur-
passing dorsal plate, everted tips very short. With two basophyses not extend-
ing as far posteriorly as apex of dorsal plate emargination. Length 5.5 mm.

VARIATIONS. The laterophyses vary in length, often surpassing the dorsal
plate by a short distance; and the everted tips are not at all constant in size
or shape. In one example from Fort Seward, tentatively assigned, the latero-
physes are more nearly as in S. fenderi, while the other diagnostic characters
indicate its position here. Specimens from Berkeley and Oakland, and one from

488

CALIFORNIA ACADEMY OF SCIENCES

[Proc. 4TH SER.

VoL, XXXII] GREEN: REVISION OF THE NEARCTIC SILIS 489

Mill Valley, all unidentified at this time, suggest that several additional species
may be involved. Length 5.5—6.5 mm.

DISTRIBUTION. CALIFORNIA: Marin County: Fairfax, V-7-11, Van
Dyke, holotype (CAS); V-20-11, Van Dyke, 1 paratype (CAS); V-7, 29-
11, Blaisdell, 2 paratypes (CAS); V-11, 25-19, Van Duzee, 2 paratypes
(CAS); Fairfax Canyon, V-18-46, J. J. Dubois, 1 paratype (UCB); Phoenix
Lake, 1V-28-40, C. D. Duncan, 1 paratype (Edwards); Mill Valley, V-10-58,
H. B. Leech, 1 paratype (CAS); Mt. Tamalpais, V-23-09, Van Dyke, 1
paratype (CAS); same, V-7-11, Blaisdell, 1 paratype (CAS). Sylvania
(Camp Meaker, Sonoma County), May, R. Ecker, 3 paratypes (CAS); same,
V-24-1895, 1 paratype (UK). (?)Humboldt County: Fort Seward, V-18-35,
E. O. Essig, 1 male, tentative identification (UCB).

(31) Silis (Silis) macclayi Green, new species.

Ho.otypr, male; Arbuckle, Colusa County, California, IV-20-62, P. M.
Marsh. In collection of the University of California at Davis.

Head black, pale in front; antennae and tips of palpi dark, antennae pale
beneath basally. Pronotum flavate. Scutellum black. Elytra flavate, tips black,
sutural bead brunneous. Beneath, except head in front and prothorax, black.
Front and middle legs, including coxae, flavate; tarsi and base of mesofemora
dark. Hind legs and coxae black, knees pale.

Pronotum similar to figure 24P. Genitalia as in figure 31. Median lobe of
aedeagus not subtriangular in lateral view, its upper margin nearly straight.
Dorsal plate with small median V-shaped emargination, lobes each side scarcely
deflected, emargination appearing nearly flat transversely in direct posterior

Ficures 32-41. Male genitalia; ventral, lateral, and dorsal views, arranged in that order

from left to right.

Ficure 32. Silis (Silis) crucialis Green, holotype.

Ficure 33. Silis (Silis) thermalis Green, holotype.

Ficure 34. Silis (Silis) acuta Green, holotype.

Ficure 35. Silis (Silis) cava LeConte, Cave Junction, Josephine County, Oregon. Dorsal
view omitted.

_ Ficure 35a. Silis (Silis) cava complex, Miami Ranger Station, Mariposa County, Califor-
nia. Dorsal view omitted.

Ficure 35b. Szlis (Silis) cava complex, Jackson County, Oregon. Dorsal view omitted.

Ficure 36. Silis (Silis) arizonica Van Dyke.

Ficure 37. Silis (Silis) fenestrata Van Dyke.

Ficure 38. Silis (Silis) californica Fender. Dorsal view omitted.

Ficure 39. Silis (Silis) reversa Green, paratype, Keen Camp, California. Dorsal view
omitted.

Ficure 40. Silis (Silis) solitaria Green, holotype. Dorsal view omitted.

Ficure 41. Silis (Silis) recta Green, holotype. Dorsal view omitted.

490 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47H SER.

view. Laterophyses moderately slender, curving downward distally; tips slender,
not everted. With two basophyses not extending posteriorly as far as apex
of dorsal plate emargination. Length 6 mm.

VARIATIONS. The lobes each side of the dorsal plate emargination may be
slightly deflected so that the emargination is very broadly and shallowly V-
shaped in direct posterior view. The tips of the laterophyses may be straight
or diverging in dorsoventral view. Specimens not from the type locality are
believed to be conspecific, but they are atypical and have not been designated
paratypes. In these the emargination of the dorsal plate is larger and deeper,
broadly V-shaped, with the lobes each side somewhat deflected; and the legs
are more extensively black. Length 5.5—6.5 mm.

DistTRIBUTION. CALIFORNIA: Colusa County: Arbuckle, 1V-20-62, P. M.
Marsh, holotype (UCD); IV-13 to V-4-62, P. M. Marsh, 6 paratypes (UCD).
Yreka, V-10-32, E. O. Essig, 3 males (UCB). OREGON: Klamath County:
20 mi. N. Beatty, Sycan River, V-30-59, Joe Schuh, 4 males (Fender); Keno,
V-25-58, J. D. Vertrees, 1 male (Schuh). Jackson County, Pinehurst, V-21-61,
Joe Schuh, 1 male (Schuh).

(32) Silis (Silis) crucialis Green, new species.

Hototyper, male; Ben Lomond, Santa Cruz County, California, V-16-31,
Van Dyke. In collection of California Academy of Sciences. CAS type number
9094.

Head black, pale in front; antennae and tips of palpi dark, antennae pale
beneath basally. Pronotum flavate. Scutellum black. Elytra flavate, tips black,
sutural bead dark brunneous. Beneath, except head in front and prothorax,
black. Legs including coxae largely dark, protibiae pale, other tibiae pale
basally, tips of femora pale.

Pronotum similar to figure 24P. Genitalia as in figure 32. Median lobe of
aedeagus not subtriangular in lateral view, its upper margin broadly arcuate.
Dorsal plate with large broadly V-shaped emargination, lobes each side
strongly deflected, forming a deep U-shaped emargination in direct posterior
view. Laterophyses stout, curving downward distally, tips slender, not everted.
With two basophyses not extending as far posteriorly as apex of dorsal plate
emargination. Length 6 mm.

VARIATIONS. The posterior aspect of the dorsal plate emargination is
usually more or less V-shaped. The legs vary in color to almost all black, only
the knees pale.

DISTRIBUTION. CALIFORNIA: Ben Lomond, V1-7-24, 8 paratypes, 16
females (LAM); IV-31, Saylor, 2 paratypes (UA). Santa Cruz County: Ben
Lomond, V-16-31, Van Dyke, holotype (CAS). Santa Cruz, VI-1-19, Van

Vor. XXXII] GREEN: REVISION OF THE NEARCTIC SILIS 491

Duzee, 1 paratype (CAS). Felton, Santa Cruz Mountains, V-15 to 19-07,
Bradley, 1 paratype (CU). La Honda, V-20-52, O. Bryant, 1 paratype (CAS).
Santa Clara County, Alma College, V-10-51, H. B. Leech, 1 paratype (CAS).

(33) Silis (Silis) thermalis Green, new species.

Hototype, male; Paraiso Springs, California, IV-22-14, L. S. Slevin.
In collection of California Academy of Sciences. CAS type number 9095.

Head black, pale in front; antennae and palpi dusky. Pronotum flavate.
Scutellum black. Elytra flavate, tips black, sutural bead brunneous. Beneath,
except prothorax and head in front, dark. Legs largely dark, tibiae paler.

Pronotum similar to figure 24P. Genitalia as in figure 33. Median lobe of
aedeagus elongate subtriangular in lateral view, widest at about distal third
where upper margin is subangulate. Dorsal plate with large V-shaped emar-
gination. Laterophyses stout, nearly straight in lateral view, tips slender, not
everted. With two basophyses not extending posteriorly as far as apex of dorsal
plate emargination. Length 6 mm.

VarIATIONS. Nothing of importance noted. Length 5.5-6 mm.

DISTRIBUTION. CALIFORNIA: Paraiso Springs, IV-22-14, L. S. Slevin,
holotype (CAS); IV-8-34, IV-18-32, V-6-28, VI-31-23, L. S. Slevin, 4 para-
types (CAS). Paraiso Hot Springs, V-4-22, L. S. Slevin, 2 paratypes (CAS).

(34) Silis (Silis) acuta Green, new species.

Howotype, male; Carrville, Trinity County, California, VI-26-14, Van
Dyke. In collection of California Academy of Sciences. CAS type number 9096.

Head black, clypeus pale; antennae and palpi dusky. Pronotum flavate.
Scutellum black. Elytra flavate, tips black, sutural bead brunneous. Beneath,
except prothorax and head in front, dark. Front and middle legs flavate,
femora basally and mesotibiae distally, dark; hind legs largely dark.

Pronotum similar to figure 24P. Genitalia as in figure 34. Median lobe
of aedeagus elongate subtriangular in lateral view, widest at about distal third
where upper margin is subangulate. Dorsal plate with large V-shaped emar-
gination. Laterophyses long, rather slender throughout, nearly straight in
lateral view, tips feebly diverging. With two basophyses extending posteriorly
_ about as far as apex of dorsal plate emargination. Length 6 mm.

VARIATIONS. The upper margin of the median lobe of the aedeagus, beyond
the subangulation, may be abruptly re-entrant, forming a protuberance in
lateral view. Length 4.5—-6 mm.

DiIsTRIBUTION. CALIFORNIA: Trinity County: Carrville, V1-28-14,
Van Dyke, holotype (CAS); VI-4, 16-13, Van Dyke, 2 paratypes (CAS).
Humboldt County: Green Point, VI-5-16, Blaisdell, 1 paratype (CAS).
Humboldt County: Redwood Canyon, V1I-4-16, Blaisdell, 1 paratype (CAS).

492 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

(35) Silis (Silis) cava LeConte.
Silis cava LEContE, 1874, Trans. Amer. Ent. Soc., vol. 5, p. 61.

Head flavate in front, black posteriorly; antennae dusky, paler beneath
basally; terminal palpal segments black. Pronotum flavate. Scutellum black.
Elytra flavate tipped with black, sides dark apically, sutural bead brunneous.
Beneath, except head and prothorax, dark. Front and middle legs, including
coxae, largely pale; hind legs largely dark.

Pronotum similar to figure 25P. Genitalia as in figure 35. With single
basophysis furcate submedially. Laterophyses slender, not extending posteriorly
beyond median lobe of aedeagus, turning more or less abruptly downward for
a short distance apically. Dorsal plate subtruncate in direct dorsal view.
Length 4.5—5.5 mm.

DISTRIBUTION. OREGON: No definite locality, LeConte type (MCZ).
Josephine County: Cave Junction, 1V-25-38, A. T. McClay, 8 males (UCD 7,
CAS):

Through the courtesy of Dr. P. J. Darlington, Jr., the author was able to
study LeConte’s type of this species. In the abundant material available, only
the Josephine County series, cited above, agrees accurately in genitalia with
the LeConte type. They have been labeled ‘‘Silis cava Lec., typical.”

All others keying here have been labeled ‘‘Szlis cava complex.” Some of
these, representing variability, may be conspecific with S. cava, while others
undoubtedly comprise several valid species. All attempts at segregating these
have failed. The specimens all have in common a single basophysis furcate
submedially, and the dorsal plate subtruncate in direct dorsal view. They
differ confusingly in the length and curvature of the laterophyses, the length
of the ventral lobe compared with the dorsal plate, and the posterior formation
of the latter (figures 35a, 35b). Their distribution follows.

DISTRIBUTION. OREGON: Yamhill County, Meadow Lake. Peavine Ridge,
near McMinnville. Josephine County, Wilderville. Jackson County: Butte
Falls; Griffin Creek; summit Green Springs Highway, Jacksonville. Ashland.
Klamath County, Rocky Point. Klamath Falls: above Geary Ranch; Algoma.
Upper Klamath Lake. CALIFORNIA: Trinity County: Eagle Creek. Men-
docino County: Vorkville; Big Dam Creek. Lake County: Lakeport. Yolo
County, Cache Creek Canyon. Plumas County: Belden; Quincy; Johnsville.
Oroville. Placer County, Baxters. Applegate. El Dorado County: Fallen Leaf;
Pollock Pines. El Dorado. Tallac. Mokelumne Hill. Calaveras County, near
Dorrington. Vosemite. Yosemite National Park. Mariposa County: Miami
Ranger Station. San Diego County.

A single specimen from a series of “S. cava complex” collected at Quincy,
Placer County, California, has the ventral lobe of the tegmen longitudinally
divided by a deep and narrow emargination. The emargination is perfectly

VoL. XXXII] GREEN: REVISION OF THE NEARCTIC SILIS 493

symmetrical and shows no indication of having been caused by a fracture.
A somewhat similar deformity has been noted in several specimens of the Sz/is
pallida group.

(36) Silis (Silis) arizonica Van Dyke.
Silis arizonica VAN Dyke, 1918, New York Ent. Soc., Jour., vol. 26, p. 174.

Black, prothorax pale fulvous above and beneath.

Pronotum as in figure 26P. Genitalia as in figure 36. Dorsal plate not
emarginate. Laterophyses moderately slender, curving downward distally,
shorter than median lobe of aedeagus. With single basophysis, tip expanded
and notched. Length 6—7 mm.

DistTRIBUTION. ARIZONA: Williams, V-27 to VI-7, Barber and Schwarz,
8 males, 1 female (USNM). Ashfork, V-31-62, G. H. Nelson, on Pinus
ponderosa, 2 males (Nelson). Prescott, holotype (CAS).

(37) Silis (Silis) fenestrata Van Dyke.
Silis fenestrata VAN Dyke, 1918, Jour. New York Ent. Soc., vol. 26, p. 175.

Black, pronotum pale fulvous with all borders black. Beneath, except
prothorax, black.

Pronotum as in figure 27P, lateral incisure closed by backward extension
of anterior process. Genitalia as in figure 37. Dorsal plate with small variable
emargination, sometimes lacking. Laterophyses long, very slender throughout,
turning downward distally, sometimes rotated to form a lyriform pattern as
shown in figure. With single basophysis, tip expanded and notched. Length
5—5.5 mm.

DISTRIBUTION. CALIFORNIA: Bear Lake, VI-i15-17, Van Dyke, 1
male (CAS). San Bernardino County: Barton Flats, V-30-58, M. E. Erwin,
2 males (UCD). San Jacinto Mountains, Marion Mountain Camp, VII-1-52,
A. T. McClay, 1 male (UCD). Tahquitz Valley, VI-3-40, H. T. Reynolds, 2
males (UCB, Fender). San Bernardino Mountains, Mill Creek, V-6, 16, 23-48,
Timberlake, 4 males, 1 female (Timberlake); same, 6,200 ft., V-22-38, 1 male
(Timberlake) Mt. San Jacinto, holotype (CAS).

The emargination of the dorsal plate is more or less variable in most
_ Silis species, but nowhere else has there been encountered such extreme vari-
ation as that occurring in S. fenestrata. In the female the black pronotal borders
are reduced to the anterior border and the median half of the basal border.

(38) Silis (Silis) californica Fender.
Silis californica FENDER, 1948, Wasmann Collector, vol. 7, p. 119.

Head black, pale in front, antennae and palpi dark. Pronotum fulvous.
Scutellum black. Elytra appearing totally black, but disk usually showing

494 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

partly fusco-flavate under brilliant illumination. Beneath, except prothorax,
dark. Legs and coxae black, anterior pair partly pale.

Pronotum similar to figure 28P. Genitalia as in figure 38. Dorsal plate
with large deep emargination. Laterophyses broad except apically, curving
downward distally for a variable part of their length. With single basophysis,
tip expanded and notched. Length 5—7 mm.

DistTRIBUTION. CALIFORNIA. J/nyo County: Panamint Mountains ; Whit-
ney Portal. San Jacinto Mountains; Idyllwild; Tahquitz Valley. Keen Camp.
Laguna. San Diego County: Otay Mountains. NEVADA: Charleston Moun-
tains, Kyle Canyon.

A specimen from Wrightwood, San Bernardino County, California, agrees
with the description of S. californica, except the tip of the basophysis is
not expanded, like in S. reversa. It has been labeled “near californica” (UCD).

(39) Silis (Silis) reversa Green, new species.

Ho.totype, male; Santa Rosa Mountain, California, VI-15-46, D. J. and
J. N. Knull. In collection of Ohio State University.

Head black, clypeus pale; antennae black, paler beneath basally; terminal
palpal segments black. Pronotum pale fulvous. Scutellum black. Elytra appear-
ing totally black, but disk showing partly fusco-flavate under brilliant illumi-
nation. Beneath, except prothorax, dark. Legs black, anterior coxae partly pale.

Pronotum similar to figure 28P. Genitalia as in figure 39. Dorsal plate
with large deep emargination. Laterophyses long, slender, curving upward
distally. With single basophysis, tip notched but not expanded. Length 6.5 mm.

VarRIATIONS. The color of the elytra, under brilliant illumination, varies
from almost entirely dark to fusco-flavate with base, sides, and apex blackish.
The front legs may be largely pale. Length 5.5—6.5 mm.

DISTRIBUTION. CALIFORNIA: Santa Rosa Mountain, VI-15-46, D. J.
and J. N. Knull, holotype, 1 paratype (OSU), 1 paratype (Fender). Keen
Camp, V-14-46, D. J. and J. N. Knull, 4 paratypes (OSU 2, Fender 2);
same, V-24-46, | paratype (OSU). San Jacinto Mountains, Pine Cove, V1-4-39,
E. S. Ross, 1 paratype (UCB).

(40) Silis (Silis) solitaria Green, new species.

HoLotyre, male; Sequoia National Park, California, 2,000-3,000 ft., V-17-
29, A. T. McClay. In collection of California Academy of Sciences. CAS type
number 9097.

Head black, pale in front, terminal segment of maxillary palpi black; basal
segment of antennae pale, others missing. Pronotum flavate. Scutellum black.
Elytra flavate, tips black, sutural bead not darker. Beneath, except prothorax,
dark. Legs, including coxae, largely pale; tarsi, basal half of middle and hind
femora, and metacoxae, dark.

VoL. XXXII] GREEN: REVISION OF THE NEARCTIC SILIS 495

Pronotum similar to figure 28P. Genitalia as in figure 40. Dorsal plate
with wide subtruncate emargination. Laterophyses broad, tips bent strongly
inward and downward. With single basophysis, tip expanded and notched.
Length 6.5 mm.

VARIATIONS. The two paratypes have the inverted tips of the laterophyses
longer and more acute than in the holotype. Length 6.5-7 mm.

DISTRIBUTION. CALIFORNIA: Sequoia National Park, 2,000-3,000 ft.,
V-17-29, A. T. McClay, holotype (CAS). Kings River Canyon, V-25, 26-48,
Ae. McClay, 2 paratypes (UCD).

(41) Silis (Silis) recta Green, new species.

Hototypr, male; Ahwahnee, California, May, A. Fenyes. In collection of
California Academy of Sciences. CAS type number 9098.

Head black, pale in front; antennae and tips of palpi black. Pronotum
flavate. Scutellum black. Elytra flavate, tips black, sutural bead not darker.
Beneath, except prothorax, dark. Legs and coxae largely pale; tarsi, base of
middle and hind femora, tips of hind tibiae, and metacoxae, dark.

Pronotum as in figure 28P. Genitalia as in figure 41. Dorsal plate with
deep rounded emargination. Laterophyses nearly straight, slender throughout,
tips not diverging. With single basophysis, tip expanded and notched. Length
7 mm.

VARIATIONS. The laterophyses may curve more obviously inward and down-
ward than in the holotype, but always they are slender with the tips not
diverging. The emargination of the dorsal plate may have the apex subtruncate.
Length 5.25—7 mm.

DISTRIBUTION. CALIFORNIA: Ahwahnee, May, A. Fenyes, holotype, 1
paratype, 1 female (CAS). Yosemite Valley, VII-8-30, F. E. Blaisdell, 1 para-
type (CAS); VI-27-21, 1 paratype (CAS). Lebec, V-30-37, Andrews and
Martin, 1 paratype (LAM). No definite locality, 2 paratypes (ANSP).

(42) Silis (Silis) angelica Green, new species.

HoLotypr, male; Waterman Canyon, California, V-28-16, J. O. Martin.
In collection of California Academy of Sciences. CAS type number 9099.

Head black, pale in front; antennae and palpi dark. Pronotum flavate.
Scutellum black. Elytra flavate, tips black, sutural bead slightly darker. Be-
neath, except prothorax, dark. Front legs and coxae largely pale, others largely
dark.

Pronotum similar to figure 28P. Genitalia as in figure 42. Dorsal plate
with rather small rounded emargination. Laterophyses moderately broad,
curving inward at about distal fourth, tips diverging forming a lyre-shaped
pattern. With single basophysis, tip expanded and notched. Length 6 mm.

VaRIATIONS. In one example the tips of the laterophyses are almost

VoL. XXXII] GREEN: REVISION OF THE NEARCTIC SILIS 497

rectangularly everted, and the dorsal plate emargination is wide and subtrun-
cate (LAM). Length 5.5-6 mm.

DISTRIBUTION. CALIFORNIA: Waterman Canyon (south slope, San
Bernardino Mountains, San Bernardino County), V-28-16, J. O. Martin, holo-
type; same, V-27-16, 1 paratype (CAS). San Bernardino Mountains, 6,000 ft.,
VI-7-1898, 1 paratype (LAM); same, 4 mi. S. of Camp Angelus, V-24-58, G. H.
Nelson, 1 paratype, 3 females (Nelson). Los Angeles County: San Gabriel
Canyon, V-18-32, 1 paratype (AMNH). Mt. San Antonio (Los Angeles
County), 10,700 ft. (corrected, 10,059 ft.), VII-3-11, P. H. Timberlake, 1
paratype (USNM).

(43) Silis (Silis) latestyla Green, new species.

Ho.totyrE, male; Fort Tejon, Kern County, California, V-14-28, Van
Dyke. In collection of California Academy of Sciences. CAS type number 9100.

Head black, pale in front; antennae and terminal palpal segments dark.
Pronotum flavate. Scutellum black. Elytra flavate, tips and sides distally black,
sutural bead not darker. Beneath, except prothorax and metasternum, dark.
Legs and coxae largely pale, tarsi and basal half of metafemora black.

Pronotum similar to figure 28P. Genitalia as in figure 43. Dorsal plate
with wide subtruncate emargination. Laterophyses broad, curving feebly
downward, tips somewhat diverging. With single basophysis, tip notched, not
appreciably expanded. Length 6.5 mm.

VARIATIONS. The dark color of the sides of the elytra distally is variable
in extent, sometimes lacking; the sutural bead varies to more or less dark.
The color of the legs varies to largely dark. Length 6—7 mm.

DISTRIBUTION. CALIFORNIA: Fort Tejon, Kern County, V-14-28, Van
Dyke, holotype, 2 paratypes (CAS); same, V-29-27, 1 paratype (USNM).
Ventura County, Mt. Pinos, V1-12-04, F. Grinnell, 2 paratypes (CAS). Lebec,

Figures 42-52. Male genitalia; ventral, lateral, and dorsal views, arranged in that order
from left to right.
Ficure 42. Silis (Silis) angelica Green, holotype. Dorsal view omitted.
Ficure 43. Silis (Silis) latestyla Green, holotype. Dorsal view omitted.
-Ficure 44. Silis (Silis) fabulosa Green, holotype.
Ficure 45. Silis (Silis) divaricata Green, holotype.
Ficure 46. Silis (Silis) montanica Green, holotype.
Ficure 47. Silis (Silis) lecontez Green, holotype.
Ficure 48. Silis (Silis) dentigera Green, holotype.
Ficure 49. Silis (Silis) disjuncta Green, holotype.
Ficure 50. Silis (Silis) triplicata Green, holotype.
Ficure 51. Silis (Silis) protracta Green, holotype.
Ficure 51a. Silis (Silis) sp. near protracta?
Ficure 52. Silis (Silis) simulata Green, holotype.

498 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

4,000 ft., V-13-28, J. O. Martin, 1 paratype (CAS). Los Angeles County: Camp
Baldy, V1-26-50, J. D. Paschke, 1 paratype (UCB). San Bernardino County:
Camp Baldy, V1-20-37, A. T. McClay, 1 paratype (UCD); same, VI-29-56,
H. R. Moffitt, 1 paratype (UCD). San Bernardino Mountains, 6,000 ft.,
VI-7-1898, 1 paratype (LAM).

(44) Silis (Silis) fabulosa Green, new species.

Hototype, male; Grand Coulee, Washington, IV-4-34, L. T. Turner. In
collection of California Academy of Sciences. CAS type number 9101.

Body and appendages deep black; pronotum fulvous with all borders black,
basal black border extending indefinitely forward, no median maculation of
disk. Underside of head in front, and of prothorax except lateral borders,
fulvous.

Pronotum similar to figure 29P. Genitalia as in figure 44. Dorsal plate with
small semicircular emargination in the inflexed sclerotized posterior border.
Laterophyses long, slender, lyrate, without distal appendix. With single baso-
physis bilaterally enlarged at tip. Length 5.5 mm.

VARIATIONS. The lateral black border of the prothorax may be much re-
duced above and beneath; the basal black border may not extend forward.
Length 5—6 mm.

DISTRIBUTION. WASHINGTON: Grand Coulee, IV-4-34, L. T. Turner,
holotype (CAS). Grand Coulee, Dry Falls, 1V-20-35, M. H. Hatch, 6 para-
types (UW). Grant County: Electric City, IV-12-42, Rogers, 2 paratypes
(UW).

(45) Silis (Silis) divaricata Green, new species.

Ho.otypr, male; Lind, Washington, IV-23-19, F. R. Cole. In collection
of California Academy of Sciences. CAS type number 9102.

Pronotum similar to figure 29P. Genitalia as in figure 45. Basophyses widely
divaricate, attaining posterior margin of dorsal plate exterior to a minute
marginal denticulation each side, margin between denticulations not inflexed
or thickened. Laterophyses without distal appendix. Length 5.25 mm.

DistRIBUTION. WASHINGTON: Lind, IV-23-19, F. R. Cole, holotype
(CAS):

(46) Silis (Silis) montanica Green, new species.

HoLotyPE, male; Florence, Montana, V-16-13, H. P. Wood. In collection
of U. S. National Museum.

VoL. XXXII] GREEN: REVISION OF THE NEARCTIC SILIS 499

Body and appendages deep black; pronotum fulvous with all borders
black, a black median vitta, narrow in front and broad at base, connecting
anterior and basal black borders. Underside of prothorax, except lateral borders,
fulvous.

Pronotum similar to figure 29P. Genitalia as in figure 46. Basophyses
arcuately subparallel, not surpassing posterior margin of dorsal plate, this
not inflexed and without marginal teeth. Laterophyses moderately slender,
feebly curving upward, appendix large and prominent, surpassing tip of latero-
physis. Length 5.5 mm.

VARIATIONS. The median black vitta of the pronotum may be narrowly
interrupted in front, not quite attaining the anterior black border. Length
5.5—6 mm.

DISTRIBUTION. MONTANA: Florence, V-16-13, H. P. Wood, holotype,
2 paratypes (USNM).

(47) Silis (Silis) lecontei Green, new species.

This name is proposed for a specimen in the LeConte collection, at the
Museum of Comparative Zoology, bearing labels ‘““Van., male” and “‘vulnerata
TYPE 3379,” the latter an MCZ label. LeConte’s locality designation pre-
sumably refers to Vancouver, British Columbia, which disqualifies the speci-
men as the original type of S. vulnerata, described from Oregon.

Body and appendages black, disk of pronotum with large fulvous spot
each side.

Pronotum similar to figure 29P. Genitalia as in figure 47. Basophyses
attaining posterior margin of dorsal plate, tips not visible beyond sides of
ventral lobe in direct ventral view. Posterior margin of dorsal plate without
distinct marginal teeth, a single short inflexed median lobe separating tips of
basophyses. Laterophyses apparently without distal appendix. Length below
average (not measured).

DISTRIBUTION. “Van.,” presumably Vancouver, British Columbia, holotype
(MICZ).

(48) Silis (Silis) dentigera Green, new species.

HoLotyPer, male; Hood River, Oregon, Childs Collection, 4-15 (mounted
- with female). In collection of California Academy of Sciences on indefinite
loan from Oregon State University. CAS type number 9051.

Body and appendages deep black, pronotum fulvous with all borders black,
disk without median vitta. Underside of head in front, and of prothorax,
except lateral borders, fulvous.

Pronotum similar to figure 29P. Genitalia as in figure 48. Basophyses long,
slender in more than distal half, curving outward and upward but not attain-
ing dorsal plate (taken from copulating pair, may not be normal position).

500 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Marginal teeth of dorsal plate very large and much retracted, plainly visible
from direct lateral viewpoint. Laterophyses slender, ascending, appendix
inconspicuous. Length 5 mm.

DISTRIBUTION. OREGON: Hood River, Childs, 4-15, holotype, 1 female
(GAS)e

(49) Silis (Silis) disjuncta Green, new species.

Hototyre, male; Gifford, Idaho, 2,900 ft., V-6-49, W. F. Barr. In col-
lection of California Academy of Sciences on indefinite loan from the University
of Idaho. CAS type number 9052.

Body and appendages deep black, pronotum largely black, disk with
dusky fulvous area each side. Underside of prothorax, except lateral borders,
fulvous.

Pronotum similar to figure 29P, paler areas irregularly elevated, forming
a rough semicircle (perhaps not normal). Genitalia as in figure 49. Baso-
physes short and stout, curving upward but not attaining dorsal plate. Marginal
teeth of dorsal plate larger than usual, retracted, plainly visible from direct
lateral viewpoint. Laterophyses straight, appendix inconspicuous. Length 5 mm.

DistriBpuTION. IDAHO: Gifford, V-6-49, 2,900 ft., W. F. Barr, holotype
(CAS).

(50) Silis (Silis) triplicata Green, new species.

HoLotypPE, male; Moscow, Idaho, May 13, 1928, altitude 2,800 ft. In
collection of California Academy of Sciences on indefinite loan from the Uni-
versity of Idaho. CAS type number 9059.

Pronotum similar to figure 29P. Genitalia as in figure 50. Basophyses
long, divergent, extending well beyond posterior margin of dorsal plate, tips
curving slightly inward in ventral view. Marginal teeth of dorsal plate large
and acute. In lateral view, laterophyses feebly sinuate, curving downward
distally, parallel and straight in dorsal view; appendix not apparent. Length
5225) mim.

DistripuTIon. IDAHO: Moscow, V-13-28, 2,800 ft., holotype (CAS); 5
paratypes (UId). Moscow, Paradise Ridge, V-7-32, 3,000 ft., J. Gillette, 2
paratypes (UId). Lenore, V-7-38, 1,000 ft., E. Ritzheimer, 1 paratype (UId).
WASHINGTON: Uniontown, V-3-60, V-20-39, 2 paratypes (Fender). Pull-
man, August, 1 paratype (USNM).

Vou. XXXII] GREEN: REVISION OF THE NEARCTIC SILIS 501

(51) Silis (Silis) protracta Green, new species.

HoLotypet, male; Spalding, Idaho, Nez Perce County, IV-18-48, W. F.
Barr. In collection of California Academy of Sciences on indefinite loan from
the University of Idaho. CAS type number 9057.

Pronotum similar to figure 29P. Genitalia as in figure 51. Basophyses long,
divergent, extending well beyond posterior margin of dorsal plate, tips curving
strongly inward in ventral view. Marginal teeth of dorsal plate large and
acute. Laterophyses nearly straight in lateral view, curving inward distally
in dorsal view, appendix small and inconspicuous. Length 5.25 mm.

VARIATIONS. The basophyses vary in length, and the inward curvature of
the tips varies in degree but is always evident. A single specimen of doubtful
position, from Lenore, Idaho, figure 5la (Nelson), differs from S. protracta
mainly in the expanded appendix of the laterophyses, plainly visible in dorso-
ventral view. Both the laterophyses and basophyses are more widely divergent.
It is possible that these differences may be explained as being preliminary to,
or immediately following, copulation. Length 5—5.75 mm.

DISTRIBUTION. IDAHO: Spalding, Nez Perce County, IV-18-48, W. F.
Barr, holotype (CAS); 6 paratypes (UId); same, III-28-53, C. J. Tarhaar,
1 paratype (UId). Lewiston, Lewiston Grade, IV-23-38, J. W. Zuckel, 1 para-
type (UId). Lewiston, IV-23-38, M. D. Bentley, 1 paratype (UId). WASH-
INGTON: Colton, V-3-60, W. W. Cone, 2 paratypes, 1 female (UW).

(52) Silis (Silis) simulata Green, new species.

HoLotypr, male; Lenore, Idaho, V-7-38, altitude 1,000 ft., H. C. Eig. In
collection of California Academy of Sciences on indefinite loan from the
University of Idaho. CAS type number 9058.

Body and appendages deep black; pronotum fulvous with all borders black,
a black median vitta of nearly uniform width, except for a submedian con-
striction, connecting anterior and basal black borders. Underside of prothorax,
except lateral borders, fulvous.

Pronotum similar to figure 29P. Genitalia as in figure 52. Basophyses long,
divergent, extending well beyond posterior margin of dorsal plate, tips not
appreciably curving inward in ventral view. Marginal teeth of dorsal plate
large and acute. Laterophyses straight, much expanded on dorsal side in lateral
view, appendix conspicuous. Length 5 mm.

DISTRIBUTION. IDAHO: Lenore, V-7-38, altitude 1,000 ft., H. C. Eig,
holotype (CAS).

502

CALIFORNIA ACADEMY OF SCIENCES

[Proc. 4TH SER.

Vor. XXXII] GREEN: REVISION OF THE NEARCTIC SILIS 503

(53) Silis (Silis) introversa Green, new species.

HoLotyPE, male; 4 mi. S. of Whitebird, Idaho County, Idaho, IV-4-60,
A. R. Gittins. In collection of California Academy of Sciences on indefinite
loan from the University of Idaho. CAS type number 9053.

Body and appendages deep black; pronotum fulvous with all borders black,
a black median vitta, narrow in front and broad at base, connecting anterior
and basal black borders. Underside of prothorax, except lateral borders, fulvous.

Pronotum similar to figure 29P. Genitalia as in figure 53. Basophyses di-
verging, abruptly converging from about distal fourth, not visible each side
of ventral lobe of tegmen in direct ventral view, tips resting on apices of
small closely placed marginal teeth of dorsal plate. In lateral view, appendix
prominent, foliate, much larger than contiguous portion of laterophysis and
surpassing its tip. Length 6 mm.

DIstTRIBUTION. IDAHO: 4 mi. S. of Whitebird, Idaho County, IV-4-60,
A. R. Gittins, holotype (CAS), 1 paratype (UId).

(54) Silis (Silis) perfoliata Green, new species.

HoLotypPe, male; Lewiston, Idaho, IV-30-32, J. Gillette. In collection of
California Academy of Sciences on indefinite loan from the University of
Idaho. CAS type number 9055.

Body and appendages deep black; pronotum fulvous with all borders black,
a black median vitta, narrow in front and broad at base, connecting anterior
and basal black borders. Underside of head in front, and of prothorax except
lateral borders, fulvous.

Pronotum similar to figure 29P. Genitalia as in figure 54. Basophyses di-
vergent, straight, tips visible each side of ventral lobe of tegmen in direct
ventral view, extending around posterior margin of dorsal plate exterior to

Ficures 53-64. Male genitalia; ventral, lateral, and dorsal views arranged, in that order
from left to right.

Ficure 53. Silis (Silis) introversa Green, holotype.
Ficure 54. Silis (Silis) perfoliata Green, holotype.
Ficure 55. Silis (Silis) proxima Green, holotype.
Ficure 56. Silis (Silis) abrupta Green, holotype.
Ficure 57. Silis (Silis) lobata Green, holotype.
Ficure 58. Silis (Silis) vulnerata LeConte.

Ficure 59. Silis (Silis) parallela Green, holotype.
Ficure 60. Silis (Silis) constricta Green, holotype.
Ficure 61. Silis (Silis) barri Green, holotype.
Ficure 62. Silis (Silis) inmsolita Green, holotype.
Ficure 63. Silis (Silis) pallida Mannerheim, Sitka, Alaska.
Ficure 64. Silis (Silis) insperata Green, holotype.

504 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

marginal tooth each side. In lateral view, appendix prominent, foliate, much
larger than contiguous portion of laterophysis and surpassing its tip. In dorsal
view, diverging tips of laterophyses short, abruptly everted. Length 4.5 mm.

VARIATIONS. In one paratype the basophyses extend beyond the dorsal
plate, as in S. protracta and allied species. Any individuals keying to that
section are removed by couplet 54. Length 4.5—5.5 mm.

DIstRIBUTION. IDAHO: Lewiston, IV-30-32, 550 ft., J. Gillette, holotype
(CAS). Nez Perce County, Central Grade, IV-13-60, A. R. Gittins, 2 paratypes
(UId).

(55) Silis (Silis) proxima Green, new species.

Ho.otype, male; Webb, Nez Perce County, Idaho, IV-8-49, W. F. Barr.
In collection of California Academy of Sciences on indefinite loan from the
University of Idaho. CAS type number 9056.

Body and appendages deep black; pronotum fulvous with all borders black,
a black median vitta, narrow in front and broad at base, connecting anterior
and basal black borders. Underside of head in front, and of prothorax except
lateral borders, fulvous.

Pronotum similar to figure 29P. Genitalia as in figure 55. Basophyses
divergent, straight, tips visible each side of ventral lobe of tegmen in direct
ventral view, extending around posterior margin of dorsal plate exterior to
marginal tooth each side. In lateral view, appendix prominent, foliate, much
larger than contiguous portion of laterophysis and surpassing its tip. In dorsal
view, diverging tips of laterophyses longer than in S. perfoliata, regularly
arcuate. Length 5.5 mm.

DISTRIBUTION. IDAHO: Webb, Nez Perce County, IV-8-49, W. F. Barr,
holotype (CAS). Gifford, V-27-49, 2,900 ft., W. F. Barr, 2 paratypes (UId).
(56) Silis (Silis) abrupta Green, new species.

HoLotyPe, male; 33 mi. W. of Yakima, Washington, V-8-49, G. H. Nelson.
In collection of California Academy of Sciences. CAS type number 9103.

Body and appendages deep black; pronotum fulvous with all borders
narrowly black, disk without median vitta. Underside of prothorax, except
lateral borders, fulvous.

Pronotum similar to figure 29P. Genitalia as in figure 56. In ventral view,
basophyses divergent, straight, tips visible each side of ventral lobe of tegmen,
attaining posterior margin of dorsal plate exterior to marginal tooth each side.
In lateral view, basophyses turning abruptly upward distally. Laterophyses
straight and rather slender in lateral view, appendix broader than, but not
surpassing, tip of laterophysis. Length 6 mm.

VARIATIONS. The laterophyses vary somewhat in width, and in one example
they curve slightly downward distally. Length 5—6 mm.

VoL. XXXII] GREEN: REVISION OF THE NEARCTIC SILIS 505

DISTRIBUTION. WASHINGTON: Yakima, VI-8-54, H. P. Lanchester, 1
paratype (Fender). 33 mi. W. of Yakima, V-8-49, G. H. Nelson, holotype, 2
paratypes, 1 female (CAS); same, C. Chastain, 1 paratype (Uild)jiesatemn2:, W.
of Yakima, V-8-49, Francis Ritz, 1 paratype (UId). Mt. Adams, V1-11-41,
4 paratypes (Fender). Ellensburg, V-5-41, Lewis, 1 paratype (UW). Virden,
IV-23-36, J. Wilcox, 1 paratype (OrSU).

(57) Silis (Silis) lobata Green, new species.

HototyreE, male; Gifford, Idaho, 2,900 ft., V-27-47, W. F. Barr. In col-
lection of California Academy of Sciences on indefinite loan from the Uni-
versity of Idaho. CAS type number 9054.

Body and appendages deep black; pronotum fulvous with all borders biack,
a black median vitta, narrow in front and broad at base, connecting anterior
and basal black borders. Underside of prothorax fulvous each side, lateral
borders black.

Pronotum similar to figure 29P. Genitalia as in figure 57. In ventral view,
basophyses divergent, straight, tips visible each side of ventral lobe of tegmen,
attaining posterior margin of dorsal plate exterior to marginal tooth each
side. In lateral view, basophyses curving upward distally. Laterophyses straight
and rather stout in lateral view; appendix conspicuous, not surpassing tip of
laterophysis. Median lobe of aedeagus broadly subtriangular in lateral view.
Length 5.5 mm.

DIsTRIBUTION. IDAHO: Gifford, 2,900 ft., V-27-47, W. F. Barr, holotype
(CAS).

(58) Silis (Silis) vulnerata LeConte.

Silis vulnerata LEContTE, 1874, Trans. Amer. Ent. Soc., vol. 5, p. 61.

The type of this species, described from a single Oregon specimen without
a more definite locality, is presumably lost, for it is not in the LeConte or in
the Horn collection. It is accordingly necessary to select a neotype so the
name may be retained in the list of recognizable species. This does not need to
be a random choice, because apparently only one of the numerous species
at hand, having the pronotum bimaculate as described by LeConte, occurs in
Oregon. Nothing else in LeConte’s description is of any identification value,
the only known differentials being derived from the male genitalia. One other
species of the “valnerata’ group has been taken in Oregon, S. dentigera. It
has the disk of the pronotum entirely fulvous. This character is not invariable,
but it is the only available evidence of identity.

NEOTYPE, male; Tumalo, Oregon, V-1-49, from the K. M. Fender collection,
Deposited in the LeConte collection at the Museum of Comparative Zoology.

Body and appendages deep black; pronotum fulvous with all borders black,
a black median vitta, broad at base and narrowing anteriorly, not quite attain-

506 CALIFORNIA ACADEMY OF SCIENCES [| Proc. 4TH SER.

ing anterior black border. Underside of head in front, and of prothorax except
lateral borders, fulvous.

Pronotum similar to figure 29P. Genitalia as in figure 58. In ventral view,
basophyses diverging, curving slightly inward distally, tips not visible each
side of ventral lobe of tegmen, attaining posterior margin of dorsal plate
exterior to marginal tooth each side. In lateral view, lower margin of baso-
physes uniformly arcuate. In lateral view, laterophyses stout, somewhat L-
shaped, lower margin bending rather abruptly upward at about distal third
forming an obtuse angle, upper margin concavely arcuate, tip acute; appendix
large, surpassing tip of laterophysis. Length 5.75 mm.

VARIATIONS. The median pronotal vitta varies by reduction, and may
occasionally extend less than half way to the apex and be confined to the
median basal impression. Length 5—6 mm.

DISTRIBUTION. OREGON: Tumalo, V-1-49, neotype (MCZ); 33 males,
2 females (Fender, CAS 1 male). Lake County: Fossil Lake, V-16-57, 2 males
(Fender); 2 males (OrSU). Redmond, IV-20-39, Schuh and Gray, 1 male
(Schuh).

(59) Silis (Silis) parallela Green, new species.

Hoxrotyper, male; Richter Pass, British Columbia, III-29-41, H. Leech.
In collection of California Academy of Sciences. CAS type number 9104.

Body and appendages deep black; pronotum fulvous with all borders black,
a black median vitta, narrow and nearly interrupted in front, broad at base,
connecting anterior and basal black borders. Underside of prothorax, except
lateral borders, fulvous.

Pronotum similar to figure 29P. Genitalia as in figure 59. In ventral view,
basophyses subparallel, feebly arcuate, outer margins convex, tips not visible
each side of ventral lobe of tegmen, attaining posterior margin of dorsal plate
exterior to marginal tooth each side. In lateral view, lower margin of baso-
physes subangulate. In lateral view, laterophyses stout, nearly straight,
symmetrical, narrowing to apex; appendix large, slightly surpassing tip of
laterophysis. Length 6 mm.

VARIATIONS. In one example, from Oliver, the subangulation of the lower
margin of the basophyses, in lateral view, is much less apparent. Length
4.5—6 mm.

DisTRIBUTION. BRITISH COLUMBIA: Richter Pass, I11-29-41, H. Leech,
holotype, 1 paratype, 1 female (CAS). Osoyoos, V-23-25, E. R. Buckell, 1
paratype (CAS); same, V-3-54, Roy Scott, 2 paratypes, 1 female (UBC).
Oliver, V-19-59, E. E. MacDougall, 1 paratype (CNC).

(60) Silis (Silis) constricta Green, new species.
HoLotype, male; Republic, Washington, V-7-36, G. R. Hopping. In col-
lection of California Academy of Sciences. CAS type number 9105.

VoL. XXXII] GREEN: REVISION OF THE NEARCTIC SILIS 50

Body and appendages deep black; pronotum fulvous with all borders black,
a black median vitta, narrow in front and broad at base, connecting anterior
and basal black borders. Underside of prothorax, except lateral borders, fulvous.

Pronotum similar to figure 29P. Genitalia as in figure 60. In ventral view,
basophyses feebly diverging, nearly straight, tips not visible each side of
ventral lobe of tegmen, attaining posterior margin of dorsal plate exterior to
marginal tooth each side. In lateral view, basophyses very stout, constricted
at tip. In lateral view, laterophyses slender, curving downward at about distal
third; in dorsal view, slender, not expanding distally; appendix minute, seti-
form. Length 5 mm.

VARIATIONS. In the paratype the pronotum is nearly all black, the fulvous
areas each side much reduced. Length 4.5—5 mm.

DISTRIBUTION. WASHINGTON: Repudlic, V-7-36, G. R. Hopping, holo-
type, 1 paratype (CAS).

(61) Silis (Silis) barri Green, new species.

HoLotyPe, male; Wawawai, Whitman County, Washington, IV-4-49, Ralph
Schopp. In collection of California Academy of Sciences on indefinite loan from
the University of Idaho. CAS type number 9050.

Body and appendages deep black; pronotum fulvous with all borders black,
a black median vitta, narrow in front and broad at base, connecting anterior
and basal black borders. Underside of prothorax, except lateral borders,
fulvous.

Pronotum similar to figure 29P. Genitalia as in figure 61. In ventral view,
basophyses divergent, straight, attaining posterior margin of dorsal plate ex-
terior to marginal tooth each side. In lateral view, lower margin of basophyses
feebly arcuate. In lateral view, laterophyses slender, widest and bending sub-
rectangularly downward near middle of length; in dorsal view, slender, not
expanded distally; appendix minute, setiform. In lateral view, median lobe of
aedeagus broadly subtriangular. Length 5 mm.

VARIATIONS. The subtriangular outline of the median lobe of the aedeagus
varies by the withdrawal of the shorter upper part within a cavity formed by
the overlapping sides of the longer lower part.

DISTRIBUTION. WASHINGTON: Wawawai, Whitman County, IV-4-49,
Ralph Schopp, holotype (CAS); 2 paratypes (UId). Wawawai, V-12-48, 1
paratype (USNM). Pullman, IV-32, Bales, 1 paratype (UW).

(62) Silis (Silis) insolita Green, new species.

Ho.otyrPe, male; “W. T.” (Washington Territory). In collection of Acad-
emy of Natural Sciences of Philadelphia.

508 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Body and appendages deep black; pronotum fulvous with all borders black,
a black median vitta, narrow in front and broad at base, connecting anterior
and basal black borders. Underside of prothorax, except lateral borders, fulvous.

Pronotum similar to figure 29P. Genitalia as in figure 62. In ventral
view, basophyses subparallel, feebly arcuate, outer margins convex, tips not
visible each side of ventral lobe of tegmen. In lateral view, basophyses strongly
arcuate, failing to reach posterior margin of dorsal plate (possibly not a
normal position). Dorsal plate with usual marginal tooth each side. In lateral
view, laterophyses very slender, curving slightly downward distally; in dorsal
view, nearly straight, somewhat expanded distally; appendix not apparent.
Length 4.5 mm.

DistrRIBuTION. “W. T.”’ (Washington Territory, includes Oregon), holotype
(ANSP).

(63) Silis (Silis) pallida Mannerheim.

Silis pallida MANNERHEIM, 1843, Bull. Moscou, vol. 2, p. 246.

Head and appendages black. Pronotum flavate, anterior and basal borders
black, lateral borders more or less dusky. Scutellum black. Elytra flavate;
sutural bead, lateral borders narrowly, and apices, dark. Underside black,
prothorax pale with lateral borders dark. Legs largely black.

Pronotum similar to figure 30P. Genitalia as in figure 63. In ventral view,
basophyses subparallel or somewhat diverging, not attaining posterior margin
of dorsal plate. In posterior view, emargination of dorsal plate with prominent
acute basal angles, these partly visible in direct lateral view as an obtuse
anteapical projection extending below upper margin of lateral incisure of
tegmen. Laterophyses rather stout, curving slightly upward in lateral view,
nearly straight in ventral view. Ventral lobe of tegmen subtriangular, apex
more or less acute, usually extending as far posteriorly as dorsal plate. In
dorso-ventral view, median lobe of aedeagus narrowly subtriangular, slender
distally. Length 5.25—6.5 mm.

Rare in collections, only fifteen males of this species have been seen. It
was described by Mannerheim from Sitka, Alaska. A National Museum
specimen from that locality constitutes the determining factor in its identifi-
cation. It differs from all other members of the “pallida” group in the shorter
basophyses not attaining the posterior margin of the dorsal plate. No variation
of importance has been noted.

DistriBuTION. ALASKA: Sitka, VI-16, Harriman Expedition, T. Kin-
caid, 1 male (USNM). BRITISH COLUMBIA: Metlakatla, Wickham, 2
males (USNM). Massett, Queen Charlotte Island, Rev. Keene, 1 male
(USNM); same, VI-3-57, E. E. MacDougall, 1 male (CNC). Zymagotitz River,
6 mi. W. of Terrace, VI-20-60, W. W. Moss, 1 male (CNC). Tyee, 27 mi. E. of
Prince Rupert, V1-24-60, B. S. Heming, 1 male (CNC). Prince Rupert, VI-15-

VoL. XXXII] GREEN: REVISION OF THE NEARCTIC SILIS 509

59, G. G. Scudder, 3 males (UBC); same, VI-1926, R. W. Pillsbury, 1 male
(UBC). Kwina Point, Sandspit, Queen Charlotte Island, V1-26-46, M. G.
Thomson, 1 male (UBC). OREGON: Olney, VI-15-25, Van Dyke, 3 males
(CAS).

(64) Silis (Silis) insperata Green, new species.

HoLotyPE, male; Homestead Inn, Mt. Hood, Oregon, VII-2-27, Van
Dyke. In collection of California Academy of Sciences. CAS type number 9106.

Head and appendages black. Pronotum flavate with all borders black,
basal black border extending forward in median impression, not attaining
anterior black border. Scutellum black. Elytra flavate; sutural bead, lateral
borders, and apices, black; lateral black border expanding toward apex. Under-
side black, prothorax pale with lateral borders black. Legs and coxae black.

Pronotum similar to figure 30P. Genitalia as in figure 64. In ventral view,
basophyses subparallel, attaining posterior margin of dorsal plate. Laterophyses
slender, nearly straight in lateral view, curving somewhat inward and reversing
distally in dorso-ventral view. Ventral lobe of tegmen extending about as far
posteriorly as dorsal plate. In dorso-ventral view, median lobe of aedeagus
narrowly subtriangular, slender distally. Length 5 mm.

VARIATIONS. The elytra are entirely black in a few specimens, otherwise
normal, from Siltcoos Lake and Gardiner, Oregon. The forward extension of
the basal black border of the pronotum may be much reduced, or entirely
lacking. Rarely the lateral black borders of the elytra are narrow throughout.
Three males have been noted in which the ventral lobe of the tegmen is deeply
and narrowly emarginate. This is an accidental deformity that has been found
also in the “cava” group. The ventral lobe, typically subtriangular with the
apex acute, varies in length and shape with the sides sometimes convex and
the apex more or less obtuse. The median lobe of the aedeagus may be partly
expanded, obscuring its normal shape. In this case there is usually visible
in ventral view a narrow median longitudinal convexity each side of which
rest the laterophyses. The laterophyses may curve definitely downward in
lateral view, reversing slightly upward distally. Length 4.5-5.75 mm.

DISTRIBUTION. OREGON: Homestead Inn, Mt. Hood, VII-2-27, Van
Dyke, holotype, 9 paratypes (CAS). Elk Lake, VII-3-38, K. Fender, 3 para-
types (CAS); VII-9-39, K. M. Fender, 2 paratypes (Fender). Cannon Beach,
VI-16-27, Van Dyke, 1 paratype (CAS). Corvallis, VI-11-25, Van Dyke,
1 paratype (CAS). Siltcoos Lake, V1-20-34, Bryant, 6 paratypes (CAS).
McMinnville, V1-25-42, 1 paratype (Fender). Sheridan Peak, Yamhill County,
VII-12-42, 8 paratypes (Fender). Still Creek, Mt. Hood, VII-17-47, VII-15-54,
4 paratypes (Fender). Government Camp, VII-5-42, 3 paratypes (Fender).
Mt. Hood, 3,000-6,000 ft., VI-20-25, 1 paratype (Fender). Glenada, Lane
County, VI-16-52, B. Malkin, 1 paratype (Fender). Florence, Lane County,

510 CALIFORNIA ACADEMY OF SCIENCES [| Proc. 4TH SER.

aural

Ficures 65-65b. Male genitalia; ventral, lateral, and dorsal views arranged, in that

order from left to right.

Ficure 65. Silis (Silis) maritima Van Dyke.

Ficure 65a. Silis (Silis) maritima complex, Woods, Tillamook County, Oregon.
Ficure 65b. Silis (Silis) maritima complex, Newport, Oregon.

VI-14-50, Malkin and Leeper, 1 paratype (Fender). Gardiner, Douglas County,
V-14-49, F. M. Beer, 1 paratype (Fender). /ndependence, VI-6, 8-34, N. P.
Larson, 2 paratypes (Schuh). Eagle Creek, VII-4-40, Joe Schuh, 1 paratype
(Schuh). Linn County, Monument Park, VI-16-60, J. D. Lattin, 1 paratype
(OrSU). Wasco County, Mayer State Park, V-23-59, 1 paratype (OrSU).
Portland, V-28, 1 paratype (USNM). WASHINGTON: Near Snowqualmie
Pass, VII-9-36, Van Dyke, 1 paratype (CAS). Paradise Valley, Mt. Rainier,
6,000-8,000 ft., VITI-2-19, Blaisdell, 1 paratype (CAS). Paradise, Mt. Rainier,
VII-4-34, Bryant, 2 paratypes (CAS). Longmires, Mt. Rainier National Park,
VII-12-36, Van Dyke, 2 paratypes (CAS). Mt. Rainier National Park, Long-
mire Springs, VI1-26-53, K. M. Fender, 1 paratype (Fender). Mt. Rainier
National Park, V1-26-62, 2 paratypes (Edwards). Puyallup, 40 ft., V-14-31,
A. J. Hanson, 2 paratypes (CAS). Seattle, V-23-14, 7 paratypes; V-14-07, 2
paratypes (CAS); same, V-1951, B. Malkin, 1 paratype (Fender). Sol Duc
Hot Springs, V1-24-36, Van Dyke, 1 paratype (CAS). Skye, VI-12, 19-36, 2
paratypes (Fender). Berkeley Park, Mt. Rainier National Park, VI1-20-41,
I paratype (Fender). Factoria, King County, V1-4-49, E. C. Johnston, 1 para-
type (CNC). Bremerton, V-27-48, Don Frechin, 1 paratype (CNC). Fort
Lewis, Pierce County, V-29-51, R. Schuster, 3 paratypes (UCB). Bothell,
V-25 to VI-3-49, Geo. Schenk, 7 paratypes (UId). Tenino, Hubbard and
Schwarz, 4 paratypes (USNM). Easton, 1 paratype (USNM). “W. T.” (Wash-

VoL. XXXII] GREEN: REVISION OF THE NEARCTIC SILIS 511

ington Territory) 2 paratypes (AMNH). BRITISH COLUMBIA: Steelhead,
V-31 to VI-20-33, H. B. Leech, 8 paratypes (CAS). Pender Harbor, G. R.
Hopping; same, R. T. Turner, 7 paratypes (CAS, UBC 1). Vancouver, V1-3-
30, V-22-30, H. B. Leech, 2 paratypes (CAS); same, V-10-31, H. B. Leech,
2 paratypes (Fender); same, V-31-31, H. B. Leech, 1 paratype (CNC); same,
VI-12-30, V-31-31, H. B. Leech, 3 paratypes (UBC); same, VI-29-32, R.
Hopping, 1 paratype (UBC); same, VI-16-55, G. Stace Smith, 1 paratype
(UBC); same, VI-11-27, G. J. Spencer,.1 paratype (UBC). Mission, V-25-30,
E. E. Peden, 1 paratype (Fender). North Bend, Chas. Palm collection, 1
paratype (AMNH); same, VI-6, Hubbard and Schwarz, 4 paratypes (USNM).
Forestry Station, New Westminster, V1-6-39, R. H. Longmire, 1 paratype
(CNC). Hope Trail, VI1-4-30, G. Stace Smith, 2 paratypes (UBC). Langley
Pr., V-20-30, K. Graham, 1 paratype (UBC). B. A., Chas. Palm collection,
1 paratype (AMNH). No definite locality, 1 paratype (Fender).

This species may be composite, or, more likely, in a state of continuing
evolutionary development. Typical specimens are easily recognized by the male
genitalia; but in many others the variable shape of the ventral lobe of the
tegmen, and supposed partial expansion of the median lobe of the aedeagus,
produce a more or less confusing atypical appearance. The fact that these
possibly atypical forms occur with typical ones in series from the same locality,
and that no definable line of demarcation separates them, seems to justify
the inclusion of all under this heading. Some material at hand has not been
so included, but is labeled simply “pallida group.” Among these are two
specimens from near Orick, Humboldt County, California (CAS), the south-
ernmost record for the group.

(65) Silis (Silis) maritima Van Dyke.
Silis pallida maritima Van Dyke, 1918, Jour. New York Ent. Soc., vol. 26, p. 170.

Body and appendages black, pronotum pale fulvous with all borders black,
basal black border extending forward a variable distance in median depression,
not attaining apical black border. Underside of prothorax, except lateral
borders, fulvous.

Pronotum similar to figure 30P. Genitalia as in figure 65. In ventral view,
basophyses subparallel, attaining posterior margin of dorsal plate. In ventral
view, laterophyses subparallel, tips acute; in lateral view bending downward
at about distal third or fourth, tips usually not at all reversing upward. In
ventral view, median lobe of aedeagus broad, tip sometimes contracted to a
triangular shape, then more or less abruptly widening. Ventral lobe of tegmen
usually not extending as far posteriorly as dorsal plate. Length 4.25—6 mm.

DISTRIBUTION. OREGON: Marshfield, IV-14-14, VI-11-14, Van Dyke
(CAS). Charleston, Coos County, VI-17, 20-57, Fender, VI-17-52, Malkin
(Fender).

512 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

One specimen, not differing in any particular from the above description,
is labeled “Carmel, Cal., V-19-13, Van Dyke” (CAS). This is obviously a
labeling error. A specimen from Humboldt County, California (Fender), is
black with the pronotum entirely pale. It has been labeled ‘‘near maritima.”

All specimens with flavate elytra keying here have been labeled “‘Silis
maritima complex.’ Extremes in genitalic structure, figures 65a and 65b,
indicate that at least two species are involved. Neither can be positively
defined because of the occurrence of numerous intermediates of doubtful
position. Members of this complex have been taken in the following localities.

DISTRIBUTION. OREGON: Waldport; Marys Peak, Benton County; New-
port; Corvallis; Woods, Tillamook County; Sandlake; Baker Creek; Mc-
Minnville; Depot Bay; Xena; Little Nestuca River; Boyer; Gold Beach.
WASHINGTON: Long Beach, Pacific County.

LITERATURE CITED

BLATCHLEY, WILLIS S.
1910. An illustrated descriptive catalogue of the Coleoptera or beetles (exclusive of the
Rhynchophora) known to occur in Indiana. Indiana Department of Geology
and N. R. Bulletin 1, pp. 1-1386.
Brown, W. J.
1940. Some new species of Cantharidae and Chrysomelidae. Canadian Entomologist,
vol. 72, pp. 161-166.
CHARPENTIER, TOUSSAINT DE
1825. Horae Entomologicae. Wratislawiae, pp. xvi + 260.
FENDER, KENNETH M.
1948. Some new and little known Cantharidae. The Wasmann Collector, vol. 7, pp.
117-123.
LreConte, JOHN L.
1850. General remarks upon the Coleoptera of Lake Superior. Jn: Lake Superior, by
Louis Agassiz, Boston, Massachusetts, pp. 201-242.
1851. Synopsis of the Lampyrides of temperate North America. Proceedings of the
Academy of Natural Sciences of Philadelphia, vol. 5, pp. 331-347.
1853. Catalogue of the described Coleoptera of the United States, by Melsheimer, re-
vised by Haldeman and LeConte. Smithsonian Institution, Washington, D. C.,
pp. xvi + 174.
1861. Classification of the Coleoptera of North America, Part I. Smithsonian Miscel-
laneous Collections, pp. 1-214.
1874. Descriptions of new Coleoptera chiefly from the Pacific slope of North America.
Transactions of the American Entomological Society, vol. 5, pp. 43-72.
1881. Synopsis of the Lampridae of the United States. Transactions of the American
Entomological Society, vol. 9, pp. 15-72.
1884. Short studies of North American Coleoptera, number 2. Transactions of the
American Entomological Society, vol. 12, pp. 1-32.
MANNERHEIM, Cart G. von
1843. Beitrag zur kaefer-fauna der Aleutischen Inseln, der Insel Sitka, und Neu Cali-
forniens. Société Imperiale Naturalistes Moscou, Bulletin vol. 16, pp. 175-314.

Vor. XXXII] GREEN: REVISION OF THE NEARCTIC SILIS Sule

Say, THOMAS
1825. Descriptions of new species of coleopterous insects. Journal of the Academy of
Natural Sciences of Philadelphia, vol. 5, pp. 160-204.
1835. Descriptions of new North American coleopterous insects, and observations on
some already described. Boston Journal of Natural History, vol. 1, pp. 151-203.
SCHAEFFER, CHARLES F. A.
1908. List of the Lampyridae from the Huachuca Mountains, Arizona, and descriptions
of new species. Journal of the New York Entomological Society, vol. 16, pp.
61-67.
Van Dyke, Epwin C.
1918. A review of the species of the coleopterous genus Silis Latreille which are found
in America north of Mexico. Journal of the New York Entomological Society,
vol. 26, pp. 161-179.

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PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES
FOURTH SERIES

G Dallas Hanna Anniversary Volume

Vol. XXXII, No. 17, pp. 515-521; 4 figs. November 22, 1967

OBSERVATIONS ON PLEUROBRANCHAEA
CALIFORNICA MACFARLAND, 1966
(OPISTHOBRANCHIA, NOTASPIDEA)

Dustin D. Chivers
Department of Invertebrate Zoology, California Academy of Sciences

The large notaspidean opisthobranch Pleurobranchaea californica was de-
scribed by MacFarland (1966). In October, 1964, two large specimens of this
species were brought to Steinhart Aquarium, San Francisco, by Richard Poole
of the California Department of Fish and Game. The larger individual mea-
sured 355 mm. in length by 125 mm. in width and was nearly twice the size
of the other specimen. These notaspideans, taken from commercial crab pots
set in waters of 30 to 40 fathoms near the southeast Farallon Island, were
placed in a 55-gallon aquarium containing circulating refrigerated salt water
(10-13°C.). Because the entire animal was not illustrated in connection with
the original description of this species, it was felt desirable to provide a photo-
graph of one of these fine specimens, as well as to record certain observations
relating to them and to several other specimens (fig. 1).

Unfortunately, within 2 weeks of the time of capture the smaller individual
had apparently disintegrated or had been eaten by the larger. Coan (1964)
observed that this species is carnivorous and that its members will attack any
animal offered, including those of its own species. The author has observed a
large specimen of P. californica attack and almost completely devour a medium-
sized anemone (Anthopleura elegantissima) in less than 10 minutes. The buccal
armature and the extremely large radula would seem to lend themselves well
to a predatory habit.

COLOR VARIATION

The over-all color of eight living specimens of Pleurobranchaea californica
observed by the author showed little variation. These specimens were all taken
in the vicinity of the Farallon Islands and Bodega Bay, California. The south-
east Farallon Island individual (now faded) was quite typical of the northern
color phase. The warty dorsal surface of the mantle was covered with a mottled

[515]
Marine Biological Laboratory
LIBRARY

DEC 4 1967
WOODS HOLE, MASS.

516 CALIFORNIA ACADEMY OF SCIENCES [Proc, 4TH. SER.

Ficure 1. Pleurobranchaea californica MacFarland, 1966. Southeast Farallon Island,
California, in 30-40 fathoms, 355 mm. in length.

pattern of brown pigment. Areas lacking pigment appeared as a series of rather
large, irregular, translucent white patches. The dorsum of the foot and oral
veil were lighter in color. An irregular network of fine wrinkles gave the entire
dorsum a crepe-like appearance. A single living southern California specimen
observed exhibited a much deeper shade of pigmentation, although the pattern
was essentially the same (fig. 2).

Ecc MASS AND VELIGERS

On 13 November, an egg mass was noted attached to a stone in the aquarium
(fig. 3). The entire mass had been laid the previous night. Subsequently, the
remaining large animal and its egg mass were placed in a 30-gallon tank and
photographed. The veligers began to hatch from the gelatinous matrix of the
egg ribbon 22 November, and by the next day the ribbon had disintegrated.
At the time of hatching, alternating rows of egg capsules were observed within
the egg ribbon, and each capsule contained many active veligers. The number
of veligers per capsule appeared to range from 10 to 15 (fig. 4). The free-
swimming larvae were so numerous on the morning of 23 November that the
water in the tank became quite murky. Samples of the veligers were taken over
a period of 2 days, until living individuals could no longer be found in the

VoL. XXXIT] CHIVERS: PLEUROBRANCHAEA CALIFORNICA

on
—_
ba f

Ficure 2. Pleurobranchaea californica MacFarland, 1966. Santa Cruz Island, California,
in 30-60 feet, 145 mm. in length, showing the gill on the right side.

aquarium. Upon hatching, the tiny shells measured 150 and consisted of one
and one-half whorls.

NARCOTIZATION

Considerable difficulty was encountered in the narcotization of the large
adult of P. californica. On 27 November the specimen was placed in a solution
of magnesium chloride and sea water. Five hundred ml. of 7 percent magnesium
chloride solution were added to 8 liters of sea water and placed under refrigera-
tion at approximately +5°C. After 7 hours the animal appeared to be unaffected,
except for the production of large quantities of viscous mucus. That this mucus
may have defensive qualities for the organism is suggested by the report of
Paine (1963) who found that the mucus secretions of an individual belonging
to Pleurobranchaea sp. (in all probability P. californica) collected in southern
California waters, had a pH of 1.0. The morning of 28 November the magnesium
chloride appeared to have had no narcotizing effect; therefore, 10 gms. of chloral
hydrate crystals were added and the temperature reduced to approximately —1°C.
Twenty-four hours later the animal still actively responded to touch by retracting
its rhinophores and oral veil. An additional 10 gms. of chloral hydrate was
added at this time.

518 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47H SER.

-
e

Ficure 3. Egg ribbon of Farallon Island specimen shown in figure 1. Length about 6 inches.

os ba *

Ficure 4. Egg capsules with unhatched veligers of Farallon Island specimen.

VoL. XXXIT] CHIVERS: PLEUROBRANCHAEA CALIFORNICA 519

On 30 November, there was no change in the specimen’s ability to respond
to mechanical stimulation. Consequently, 70 percent ethyl alcohol was slowly
added, 15 ml. at a time over a 4 day period. By 4 December, 270 ml. of 70
percent ethyl alcohol had been added and although the solution had become
quite murky, the opisthobranch retracted its rhinophores and oral veil when
prodded. The solution was maintained at this concentration until mechanical
stimulation failed to cause visible contraction of any part of its external anatomy.
On 7 December, 11 days after the beginning of the narcotizing process, the ani-
mal failed to respond.

GEOGRAPHIC DISTRIBUTION

Acquisition of this fine specimen of Pleurobranchaea californica brings the
total number of specimens of this species contained in the Academy’s preserved
invertebrate collection to 39. The over-all length of individuals ranges from
165 mm. down to 8 mm. The geographic range of the Academy’s specimens is
from lat. 41°27’N.; long. 124°29’W. to lat. 32°43’N.; long. 117°14’W., or ap-
proximately from off the mouth of the Klamath River to San Diego, California.
All specimens which carried data regarding bottom conditions were found on
a substrate consisting predominantly of fine “green mud.” The bathymetric
range of our specimens is from 30-60 feet to approximately 200 fathoms.

Following is a list of the 20 specimen lots of P. californica contained in the
collection of the California Academy of Sciences, Department of Invertebrate
Zoology. All are from the California coast:

Lot no. 1, 115-119 fms., WSW. of the mouth of the Klamath River, lat.
41°27’N., long. 124°29’W., Peter Isaacson, collector, 10 September 1964.
Single specimen.

Lot no. 2, 70-74 fms. off Bodega Bay, California Fish and Game Block no.
249. Phil Hanson and Dan Clark, collectors. May, 1949. Single specimen.

Lot no. 3, 68 fms., fine green sand, Farallon Light, N. 11’2° East, 3.8 miles.
United States Fish Commission Steamer Albatross, collector. Station no.
D-5788, 21 October 1912. Two specimens. Syntypes.

Lot no. 4, 46 fms., fine green sand, Farallon Light, N. 421%2°W., 3.9 miles.
United States Fish Commission Steamer Albatross, collector. Station no.
D-5789, 21 October 1912. Single specimen.

Lot no. 5, 60 fms., Farallon Light, 4 miles, 10°W. California Departmen:
of Fish and Game R/V. XN. B. Scofield, collector, 21 April 1948. Single
specimen.

Lot No. 6, 60-80 fms. Farallon Islands, area of the southeast Farallon. Richard
Poole, California Department of Fish and Game, collector. Taken in com-
mercial crab pot. October, 1964. Single specimen.

Lot no. 7, 15-20 fms. off Ocean Beach, San Francisco, near Fleishhacker Pool.
Dennis Sullivan, collector, 7 December 1962. Single specimen,

520 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Lot no. 8, 100 fms. off Davenport, Santa Cruz County. Captain C. Davies
of the drag boat Warrior, collector. June, 1962. Single specimen.

Lot no. 9, 76 fms. Dark green mud and sand, off of Santa Cruz light, Monterey
Bay. N. 31°W., 6.1 miles. United States Fish Commission Steamer, Alba-
tross, collector, 12 May 1904. Station no. D-4480. Single specimen.

Lot no. 10, 37 fms. On dark green mud, fine sand, black specks. Off Monterey,
Point Pinos Light House. Six miles S. 35°W. United States Fish Com-
mission Steamer Albatross, collector. Station no. D-4458, 12 May 1904.
Single specimen.

Lor no. 11, 92 fms. Off Avila, San Luis Obispo County. Ten miles, 348° True
SW. of Avila. G D. Hanna, collector, 25 February 1951. Two specimens.

Lot no. 12, 30-60 ft. Off Santa Cruz Island. Dr. James Case, collector.
Taken with SCUBA. Single specimen. Figure 2.

Lot no. 13, 80-92 fms. Off Santa Barbara. Ten miles, 65° True. G D.
Hanna, collector, 19 February 1951. Single specimen.

Lot no. 14, About 200 fms., green mud, 198° True from Point Fermin, 8.5
miles. Crocker-Stanford 1938 Deep Sea Expedition. G. S. Myers, R. L.
Bolin e¢ al., collectors. Station no. 28, 17 September 1938. Thirteen speci-
mens all less than 25 mm. in length.

Lot no. 15, 440-470 meters. Off San Pedro, lat. 33°38’ to 33°41’N.; long.
118°17.6’ to 118°19.5’°W. W. E. Ritter and W. J. Raymond, collectors.
San Diego Marine Biological Association. Station no. XIV-H2, 13 June 1901.
Single specimen less than 25 mm. long.

Lot No. 16, 60 fms. Santa Catalina Island. One and three-quarters miles,
105° True from Long Point. Lat. 33°22’N.; long. 118°20’°W. Crocker-
Stanford 1938 Deep Sea Expedition. G. S. Myers, R. L. Bolin eé al.,
collectors. Station no. 47, 20 September 1938. Three specimens, one less
than 25 mm. long.

Lot no. 17, 178 fms. On fine gray sand and rock off Santa Catalina Island.
N. 79°W., 2.8 miles off Long Point. United States Fish Commission Steamer
Albatross, collector. Station no. D-4410, 11 April 1904. Single specimen.

Lot No. 18, 80 fms. On dead and bleached corallines. Santa Catalina Island,
33°23’N., 118°20’W., off Avalon. Crocker-Stanford 1938 Deep Sea Expedi-
tion. G.S. Myers, R. L. Bolin e¢ al., collectors. Station no. 30, 17 September
1938. Two specimens.

Lor no. 19, 110 fms. Green mud and shale, off La Jolla, 3.2 miles S. 34°E.
from Soledad Hill Point. United States Fish Commission Steamer Albatross,
collector. Station no. D-4322, 7 March 1904. Two specimens.

Lot No. 20, 35-38 fms. Green mud and sand. Off Point Loma. M. W. Williams
and K. W. Kenyon, collectors, 9 October 1946. Two specimens under 25 mm.
long.

Vor. XXXII] CHIVERS: PLEUROBRANCHAEA CALIFORNICA 5

Les)
—

ACKNOWLEDGMENTS

I would like to thank Mr. Allyn G. Smith, Associate Curator of the Depart-
ment of Invertebrate Zoology, for the photographs used. I wish also to express
thanks to Mr. Richard Poole and Mr. Eugene Coan for supplying specimens,
and to Mr. Maurice Giles of the Academy for preparation of the illustrations
from the original Kodachrome slides.

LITERATURE CITED

Coan, EUGENE
1964. A note on the natural history of Pleurobranchaea species (Gastropoda:
Opisthobranchia). Veliger, vol. 6, no. 3, p. 173. January 1.
MaAcFarranp, FRANK MACE
1966. Studies of Opisthobranchiate Mollusca of the Pacific Coast of North America.
Memoir VI, California Academy of Sciences. i-xvi, 1-546 pp., plates 1-72.
April 8.
PAINE, ROBERT T.
1963. Food recognition and predation of Opisthobranchs by Navanax inermis. Veliger,
vol. 6, no. 1, pp. 1-9. July 1.

PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES
FOURTH SERIES

G Dallas Hanna Anniversary Volume

Vol. XXXII, No. 18, pp. 523-580; 8 plates February 23, 1968

LOWER CRETACEOUS FORAMINIFERA OF
THE ORCHARD PEAK-DEVILS DEN AREA,
CALIFORNIA

By
Clifford C. Church

Research Associate, California Academy of Sciences

The microfaunas of the lower Cretaceous of California have received very
little attention from micropaleontologists in the past and only in recent years
has anything been published about them. This has been partly because of a
lack of economic interest in this part of our extensive Cretaceous section, in
part to its limited areal extent when compared to the more widespread upper
Cretaceous, and possibly, in a large measure, to the fact that well-preserved
faunas are scarce in many of the better known outcrop sections. With regard
to the macro fossils the opposite of this situation has prevailed. Numerous
papers on the stratigraphy and paleontology of the lower Cretaceous have ap-
peared from the beginning of geologic work in California. The most recent
paper describing lower Cretaceous beds in the area under consideration, is by
Owen T. Marsh (1960). In it Marsh names and describes in detail seven
new formations. One of the lower formations, the Badger shale, is questionably
listed as of lower Cretaceous age. A limited outcrop of serpentine is within or in
fault contact with the Badger at its base. Below the Badger shale in structural
discordance, is the Hex formation which Marsh, with considerable doubt, assigns
to the late Jurassic. The doubtful assignment was based on the identification of
belemnites by Professor Leslie Baristow of the British Museum of Natural
History and Dr. J. A. Jeletzky of the Geological Survey of Canada. Professor
Baristow suspected the belemnites were of late Jurassic age but stated that his
conclusions were only provisional. Dr. Jeletzky’s opinion was quoted as follows,
“ _ . the following suggestions are to be considered quite tentative. It would

[523]

524 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

appear to me that the fragment belongs to a large and sturdy Belemnopsis
probably of the group of Belemnopsis sulcatus Phillips or Belemnopsis gerardi
_.. Its being of the group of Indo-Pacific Belemnopsis gerardi Oppel, 1863 . . .
appears to be more probable . . . Should the above assumption be correct . . . up-
per Jurassic age would be indicated for the beds containing it.” Marsh further
stated that a different opinion of the age was reached by John P. Wagner
whose MS. thesis (University of California, 1947) deals with the geology of
the Sawtooth Ridge Quadrangle. One of the several species of belemnites found
in the Hex formation was identified by Dr. J. Wyatt Durham of the University
of California at Berkeley, as Acroteuthis winslowensis Anderson, 1938. This
identification was subsequently confirmed by Dr. J. A. Jeletzky who stated
that the form should be, “‘tentatively assigned to early to mid-lower Cretaceous
(not younger than Barremian, though).”

In a further attempt to determine the age of the Hex formation, Marsh col-
lected samples for microfossil determinations and submitted them to Mr. J. D.
Bainton of the Standard Oil Company laboratory for study. Both radiolarians
and foraminifera were found and these, mostly long ranging species, were
tentatively considered to be of late Cretaceous age. However, the concluding
statement by Bainton was that, “‘the fauna is not definitely definitive of age and
therefore the age assignments . . . should not be considered conclusive.” Thus
faced with three possible age assignments for the Hex, Marsh concluded, “It is
possible that all three ages are represented by the Hex, but this possibility cannot
as yet be proved because none of the age assignments can be stated as being con-
clusive. For the sake of convenience, the Hex formation will be designated pro-
visionally in this report as upper Jurassic (?).”

In the Devils Den outcrop the Hex formation is in fault contact with Eocene
and Miocene at its eastern extremity and probably Eocene and Paleocene at the
west where its limits are not so clearly defined. In the Hex Hill occurrence both
folding and faulting appear to account for the outcrop which extends over a
much wider area than the more easterly outcrop. In describing the Hex Hill
exposure Marsh stated, ““The Hex formation crops out east of Orchard Peak
as a single band about a half mile wide and more than four miles long, parallel-
ing the base of the scrap formed by Avenal Ridge. The unit occupies the core
of the Avenal Ridge piercement anticline. Apparently the soft, plastic shales of
the Hex formation, originally at an unknown depth below their present position,
were squeezed up through the overlying formation during folding and thrust
faulting of the Avenal Ridge anticline.” In whatever way this extensive body
of plastic clay arrived at its present position, there are very few contrasting
beds of silt or sand to suggest a structural pattern. In only one small area, in a
deep gully in the central part of the Devils Den proper, have beds been found
which show definite stratification. Here the strike is approximately north-south
which is at right angles to the Eocene and Miocene on its north and south

VoL. XXXII] CHURCH: CRETACEOUS FORAMINIFERA

On
bo
UL

flanks. As most of the Hex formation is a massive claystone with little evidence
of bedding, the only relief from its sameness are the occasional outcrops of
lenses and pods of impure, whitish to buff limestone which, because of its su-
perior resistance to weathering, stands out above the clays. It is these limy
masses which have yielded the few ammonites found. They also contain numer-
ous radiolaria and occasional foraminifera. Belemnites have also been found in
them in rare instances but most of these weather out of the clay and occur as
fragments on the rain-washed surface.

Foraminifera were found in virtually all fresh samples of the clay but in many
of them only arenaceous species were present.

HISTORY OF PRESENT WORK

From August to October of 1932, Mr. F. A. Menken, then a field geologist
with the Associated Oil Company (later Tidewater Oil Company), made a geo-
logic study of the Devils Den—Orchard Peak area and mapped the various
geologic units. From time to time during the course of his work he submitted
macrofossil and microfossil samples to the laboratory for identification and age
determination. Among the samples were a few soft claystone samples from the
deeply dissected, low, rounded hills long known as Devils Den, and from the
smoothly contoured, flat-topped clay hill on the southeast slope of Orchard
Peak, more recently named Hex Hill by Marsh. These samples yielded forami-
nifera which were new to our experience in California foraminifera, but from
their similarities with faunas from the lower Cretaceous of Europe and Texas
and the association with the belemnites, they were tentatively considered to be
of lower Cretaceous age. Because of our uncertainty as to the age of the fauna
and its unusual character, Dr. G D. Hanna, then senior Paleontologist for
the company, and the author, accompanied by J. B. Stevens, went to the Devils
Den—Orchard Peak area with the hope of finding more conclusive evidence for
the age. Samples of claystone were collected at Devils Den and from the clay-
stone hill on the eastern flank of Orchard Peak. These samples later yielded
foraminifera similar to those collected by Mr. Menken. In the Hex formation
belemnites like those from Devils Den were found along with a single ammonite
of the genus Lytoceras which Dr. F. M. Anderson later named Lytoceras
saturnale. A few aucellas were also found which Dr. Anderson identified as
Aucella solida Lahusen and A. inflata Toula, all known to him from the lower
Cretaceous Paskenta beds of the Shasta Series. This was our first definite indi-
cation of the true age of these beds. In his paper (Anderson, 1938, p. 105) on
the lower Cretaceous deposits in California and Oregon, he made the following
statement regarding Aucella solida. ‘This species has been found with A. inflata,
A. uncitoides and Lytoceras saturnale nov., in the lower beds of the Paskenta
group in the type district, and in the same horizon in the Devils Den district,
Kern County, where it was collected by Hanna and Church.”

526 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER.

More recently (1962-1964), the author made several visits to the area which
permitted a more detailed examination of the outcrop and more extensive and
selective sampling of the clays and fossils. Two more specimens of the ammo-
nite species, Lytoceras saturnale, were found in the Devils Den outcrop as well as
an aucella and a better preserved collection of belemnites. A good series of the
clays were collected from both Devils Den and Hex Hill which furnished a wide
variety of well preserved foraminifera. These faunas were similar to those col-
lected at the earlier date but in the variety of species, far exceeded any previously
collected. The microfaunas from the two areas were studied and compared and
while many species are common to the two areas, there are some notable differ-
ences in the two faunas. A number of distinctive species from no. 7 of Hex
Creek have not been found at Devils Den and some equally distinctive species
found at Devils Den have not been found at Hex Creek, a circumstance possibly
due to insufficient sampling. It does seem quite certain, however, that with the
number of species, both calcareous and arenaceous, common to the two areas,
the formations are of essentially the same age. This belief is further strengthened
by the fact that such uncommon species as the new genus Menkenina and cer-
tain species of Citharina are found at both localities. Also, as has been mentioned
earlier in the text, the same species of macrofossils, have been found at both
outcrops and these well authenticated time markers are the key to the age of the
Hex. As indicated earlier, the isolated nature of the Hex formation offers little
opportunity for an age assignment from its relationships with contiguous beds
of established zones and ages. The beds in fault contact with it are from upper
to possibly lower Cretaceous but Marsh had no definite proof of the true age
of the Badger shale which is the oldest of the strata pierced by the Hex. Forami-
nifera from the lowest part of the Serpiente formation in fault contact with the
Hex are G-2 (Turonian) in age but according to Marsh’s mapping this would
be in the upper few hundred feet of the formation. The determination of the
exact age of the Hex formation therefore must be decided entirely on the basis
of the fossils found in it. Marsh placed the age of the Hex formation in the
Jurassic with considerable doubt, for reasons already stated but apparently was
not aware of Anderson’s identification of the ammonite and the aucellas, all of
which had been found previously in the lower Paskenta group of the type district
in northern California. In the light of Anderson’s age assignment, it now seems
more significant that J. Wyatt Durham identified the belemnite found by John
P. Wagner in the Hex formation, as Acroteuthis winslowensis Anderson, a lower
Cretaceous species. In the present study of the Foraminifera many species
were identified from the Valanginian of Europe but some do range much higher.
It was apparent early in the study that a Jurassic age could be ruled out and
further comparisons indicated rather strongly that a correlation with the
Valanginian, as suggested by the macrofossils, was the most likely possibility.

VoL. XXXII] CHURCH: CRETACEOUS FORAMINIFERA Seat!

This view is supported in a recent study of the aucellas (now called buchias),
by Dr. Ralph W. Imlay of the United States National Museum. He critically
examined numerous suites of the genus from Pacific Coast states, particularly
Alaska and California, the express purpose of his study being to determine the
validity of the numerous named species, many of which were described from
single localities. Of particular import to the present problem, to determine
the age of the Hex formation, is his conclusion concerning the species Aucella
crassicollis Keyserling. As stated earlier, Anderson identified the two species of
Aucella from the beds containing the ammonite, Lytoceras saturnale, as A. solida
Lahusen and 4d. inflata Toula. Both of these species, as well as A. uncitoides
Pavlow, are considered by Imlay to be named variations of a single species, A.
crassicollis Keyserling. In his range chart of the various species of Aucella,
Imlay places this species in the upper two-thirds of the Valanginian of the lower
Cretaceous, where it is often abundant, but he does not show it ranging above
this zone into the Hauterivian. This opinion of the age level of A. crassicollis
appears to be shared by Dr. J. A. Jeletzky who examined the Hex Hill-Devils
Den macrofossils at the California Academy of Sciences in San Francisco, Cali-
fornia, 23 November 1966. He identified the buchias |aucellas| from the Hex
formation as being of the B. crassicollis group and illustrated, by means of a
chart (information received by Dr. L. G. Hertlein, of the Academy’s Depart-
ment of Geology), that Buchia crassicollis, B. inflata, and B. solida were of the
uppermost Valanginian but not extending above that stage. Anderson (1932,
p. 103) also stated that, ‘“They [Awcella crassicollis| are not known to occur
in any part of the Knoxville series (upper Jurassic) or in any beds later than
Paskenta,” again placing an upper limit on the species. If these specialists in
the field of lower Cretaceous fossils are correct in their opinions, that Awcella
crassicollis Keyserling is confined to the Valanginian stage, at least on the Pa-
cific Coast of North America, and there seems little reason to doubt it, then
the Hex formation must be accepted as being of upper Valanginian equivalence.
From a historical viewpoint the finding of fossils of lower Cretaceous age here
should cause little surprise because, as far back as 1910, Paskenta fossils (then
called Knoxville), were reported found in McLure Valley, only a few miles north-
east of Devils Den by Arnold and Anderson (1910). The most significant
fossil reported found was Aucella crassicollis Keyserling. A few years later
Dr. G D. Hanna found the same species in this locality. The clay shale in which
the aucellas were found is very similar to that at Devils Den and Hex Hill but
it yielded no foraminifera.

RECENT ZONATION OF THE LOWER CRETACEOUS

At the time of the first reading of this paper in Bakersfield, California, 1 April
1965 (Fortieth Annual Meeting of the Pacific Section, American Association of
Petroleum Geologists, Society of Exploration Geophysicists and Society of Eco-

8 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

UL
bo

FORAMINIFERAL FM ANDERSON SOUTH AND CHUBER NORTH END
Bauigea CENTRAL SACRAMENTO REDDING
ZONATION WESTSIDE CLASSIFICATION yi)

SAGE SACRAMENTO
GOUDKOFF, A-H ZONES TERMINOLOGY FRUTO ONO AREA
CCAS SIBLCALLON on VALLEY MURPHY, RHODDA POPENOE
y -M
ee he ees a AEG EME QUAD. AND PETERSON

DDD DDD OOO
CAMPANIAN FORBES
FM f
F-
GUINDA FM
— SEES

2
CONIACIAN ZONE FUNKS FM ae Seu
ea ae) ¥
TURONIAN Hw"
BIOFACIE HED
BIOFACIES

AREA

SENONIAN
PANOCHE

CHICO SERIES

MEMBE
YOLO FM ?. ?
SITES FM
MEMBER IIT SAN
YOLO FM

——_—
VENADO FM ~
VENADO AND MEMBER II SHALE
FRUTO SS TONGUE
FM

UPPER CRETACEOUS
CHICO SERIES

PIONEER
FORMATION

CANYON

CHICKABOLLY
MEMBER

BUDDEN

LOWER CRETACEOUS

NEOCOMIAN

SHASTA SERIES
PASKENTA HORSE TOWN

SHASTA SERIES
PASKENTA HORSETOWN

CLARK VALLEY
I "ANTELOPE MUDSTONE
SHALE"
CENOMANIAN HOME (TALIAFERRO) | JULIAN ROCKS
PLECTINA SALT. FORMATION MEN
SUBZONE CREEK" CONGLOM
NEW GENUS ee
= SP 19
SUBZONE
ALBIAN EPISTOMINA ASPILCHE
SP 10 FORMATION
SUBZONE
? 2:
APTIAN
? WINTUN
2, '
BARREMIAN FORMATION ;
?
HAUTERIVIAN
nECTOR CONBLOW
:
VALANGINIAN
? SANHEDRIN
BERRIASIAN ? eae
; :
TITHONIAN aya
K
& NOXVILLE KNOXVILLE eT L
SERIES SERIES COMPLEX
PORTLANDIAN

Ficure 1. Chart showing foraminiferal zonation, late Mesozoic of the Sacramento
Valley, California. Chart reproduced through the courtesy of Keith D. Berry, Standard
Oil Company, Oildale, California.

nomic Paleontologists), a paper was presented by Mr. Keith D. Berry on, “New
Foraminiferal Zonation, upper Mesozoic, Sacramento Valley, California.” This
paper was based on a detailed study of a large number of surface and well
samples from the upper Jurassic Knoxville to the base of the upper Cretaceous
where the established zones of Goudkoff were encountered. In this study Berry
described and named six new zones, I, J1, J2, K, L, M, extending the zonation of
Goudkoff downward from the base of the upper Cretaceous to the upper Jurassic.
He also established more definite criteria for the ““H” Zone of Goudkoff and de-
fined its limits, placing it on a par with the other more adequately defined zones.
In naming the new zones he continued the alphabetical system begun by Goudkoff.
Zone I to L embrace the lower Cretaceous and M the upper Jurassic, Tithonian.

VoL. XXXII] CHURCH: CRETACEOUS FORAMINIFERA 529

In the abstract of his paper he stated that the K, L, and M Zones are ‘“‘more
difficult to differentiate, because of the absence of planktonics and the predomi-
nance of many similar Nodosariidae.” The latter part of this statement is
strikingly descriptive of the Devils Den faunas which are largely composed of
Nodosariidae. The statement about the “absence of planktonics’’ however, does
not apply as several species were found in the Hex Hill fauna. This may be
because of better preservation of the fossils in the Hex Hill and its quick and
complete disintegration in water in preparing samples for study. Berry’s new
zones are approximately correlated with the European stages on the basis of
both benthonic and planktonic foraminifera and where possible, tied in to the
known macrofossil localities which had been equated with the European standard
section. Where uncertainties existed between zones he inserted question marks
to allow for them. This new zonation, based on normal, relatively undisturbed
lower Cretaceous and upper Jurassic sections, presented the first available oppor-
tunity for a comparison of the Hex Hill-Devils Den faunas with an established
lower Cretaceous stratigraphic and faunal sequence. Through the considerate
cooperation of Mr. Berry, a comparison was made between faunas of the Hex
formation and those from the Sacramento Valley sections. In this examination
a fauna was found which contained some of the more distinctive species common
to the Devils Den faunas. It is representative of Berry’s K Zone which he
believed to be of Hauterivian, Barremian equivalence. The rich fauna of Hex
Creek no. 7 was not recognized among Berry’s slides but he was of the opinion
that it represented beds younger than those at Devils Den. This determination
of the age is at variance with Anderson’s and Jeletzky’s placement of the macro-
fossils of the Hex Hill in the Valanginian, but the foraminiferal comparisons
were admittedly brief and more detailed work and comparisons of these lower
Cretaceous faunas may reveal the reasons for the apparent difference in correla-
tion. From the variations now known to exist in the Hex Hill faunas it may be
found that some part of it is younger than the upper Valanginian.

DISTINGUISHING FEATURES OF THE HEX HILL FAUNA

There are many features about this fauna which serve to distinguish it from
the later Cretaceous of the Pacific Coast, some of the more important being, (1)
The great preponderance of the Nodosariidae (Lagenidae), number at least 50
of the listed species and many more if all the nodosarias not listed were to be
included; (2) The small number of planktonic species present and their small
size; (3) The presence of genera not found in the upper Cretaceous of California;
(4) The similarity of the fauna with that of the lower Cretaceous Valanginian
of Northwestern Germany and Trinidad, British West Indies; (5) Also the
presence of genera and species heretofore not described from California. Of the
88 species listed, 69 are calcareous and 19 arenaceous. The few, small pelagic
species were found in only a few places but are usually quite common when

530 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

present. There are at least seven rotaloid genera present and where they occur
they are usually quite common.

Of the other microfossils, radiolarians are quite abundant in the limestone
masses and more rarely they may be common in the clays. Ostracods are very
rare and small. Minute carbonaceous fragments are occasionally quite abundant
so it is probable that spores and pollen are also present. The claystones have
not, to my knowledge, been examined for the smaller microfossils such as dis-
coasters, coccoliths or hystrichospherida. The limestone was examined for dia-
toms by Dr. G D. Hanna but not one was found.

ACKNOWLEDGMENTS

In the preparation of this paper I am especially indebted to Dr. and Mrs.
G D. Hanna of the California Academy of Sciences. This applies not only for
the preparation of photographic plates but to never-failing encouragement and
assistance in many other ways. To Mrs. Hanna I owe my special thanks for
her expert illustration of the fauna. Her accurate and detailed drawings of
foraminifera have long been known to most paleontologists and recognized as a
standard of excellence among illustrators. Without her generous assistance in
making the drawings this paper would not have been attempted. Dr. L. G.
Hertlein, also of the Geological Department of the California Academy of
Sciences, was of great assistance in editorial and technical advice.

I further wish to acknowledge the invaluable assistance of Dr. J. J. Graham
of Stanford University, Mr. Keith D. Berry of the Standard Oil Company
Laboratory of Oildale, California, and Mr. Andrew W. Marianos of the Humble
Oil Company, Los Angeles, California for the loan of rare and hard-to-obtain
papers on the lower Cretaceous of Europe, and Mr. Berry and Mr. Marianos for
the opportunity of comparing the Hex formation fauna with their large series
of slides of the lower Cretaceous from the west side of the Sacramento Valley,
California. Mr. E. H. Stinemeyer of the Shell Oil Company, Oildale, California,
was of assistance in making a number of color photographs of foraminifera and
Mr. A. A. Almgren has been of assistance on field trips and in the loan of com-
parative material. For the loan of the excellent geologic map of the Devils Den
area which was reproduced in their article in Bulletin 118, California Division of
Mines and Geology, part 3, I am indebted to Mr. Martin Van Couvering and
H. B. Allen. For providing me with an excellent set of foraminiferal samples
from the Grayson formation, Grayson Bluff on Denton Creek, Denton County,
northern Texas, the locality from which Helen Tappan obtained most of the
foraminifera described in her paper on the Grayson formation, I owe my thanks
to my nephew and niece, Max and Joan Church of Fort Worth, Texas. Finally,
I owe much to the support of the Tidewater Oil Company through many years
and for the friendly cooperation of its geologic staff.

VoL. XXXII] CHURCH: CRETACEOUS FORAMINIFERA Soll

CALIFORNIA ACADEMY OF SCIENCES LOCALITIES
Devits DEN

Locality 27502 (CAS). Number 49. About 150 feet west and 400 feet
north of the south quarter corner of the northeast quarter of Section 20. Town-
ship 25 south, Range 18 east of the Mount Diablo base and meridian north-
western Kern County, California. F. A. Menken, collector.

Number 50. About 600 feet west and 100 feet north of center of Section 20,
Township 25 south, Range 18 east of the Mount Diablo base and meridian.
F. A. Menken, collector.

Locality 40081 (CAS). Head of east-west trending gully, about 800 feet
northwest of the center of Section 21, Township 25 south, Range 18 east of the
Mount Diablo base and meridian, Kern County, California, Sawtooth Ridge
quadrangle, 7.5 minutes, 1935 Edition. C. C. Church, collector.

Number 47. North side of east-west gully above and about 500 feet
southwest of first sample. Earlier localities 16196 and 27501 (CAS) same
locality, described as ‘“‘center Section 21, Township 25 south, Range 18 east of
the Mount Diablo base and meridian, Devils Den area.” F. A. Menken,
collector.

Number 48. About 300 feet northwest of no. 47 near head of small side
gully. Also listed under 16196 and 27501 (CAS). F. A. Menken, collector.

Locality 40091 (CAS). Station numbers 1 to 8. From deep north-south
trending V-shaped gully beginning 500 feet south and 700 feet west of the
east quarter corner of Section 20, Township 25 south, Range 18 east of the
Mount Diablo base and meridian, Kern County, California. C. C. Church,
collector.

Number 1. East bank of gully near base, approximately 75 feet north of
gully mouth.

Number 2. About 75 feet up stream north of no. 1, in a straight line.
Number 3. About 30 feet north of no. 2.
Number 4. About 25 feet north of no. 3.
Number 5. About 20 feet north of no. 4.
Number 6. About 50 feet north of no. 5.
6

Number 7. About 75 feet north of no.
Number 8. From steep east bank of gully about 20 feet south of junction
with first northeast trending branch; this is about 600 feet due west of the east
quarter corner of Section 20, Sawtooth Ridge quadrangle, 1953 Edition.
Lytoceras saturnale was found on the side of this gully near station no. 7.

Hex HILu

This flat-topped hill with deeply dissected sides occupies much of the north-
east quarter of Section 31 and the northwest quarter of Section 32, Township 25
south, Range 18 east of the Mount Diablo base and meridian, northwestern

532 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Kern County, California. It is terminated on the west side by a deep V-shaped
gully which is dry most of the year. This creek has no official name on the
topographic sheet but is referred to here as “Hex Creek.”

The mouth of this gully is at about the exact center of Section 31 and the
creek trends due north upstream a quarter mile, then turns northeast and divides
into two branches near the fault contact of the Hex formation and the overlying,
well bedded and steeply dipping shale and sandstone of the Serpiente formation
of late Cretaceous age. The Sawtooth Ridge, 7.5 minute quadrangle, is the
basis for all measurements.

Locality 40092 (CAS). Station no. 1, from west slope of the most easterly
deep gully of Hex Hill, about 200 feet north and 400 feet west of center of
Section 32, Township 25 south, Range 18 east of the Mount Diablo base and
meridian, northwestern Kern County, California. C. C. Church, collector.

Locality 40093 (CAS). Station no. 15, from the upper end of the second
west trending side gully from the mouth of the most easterly deep gully of Hex
Hill, about 1000 feet west and 200 feet north of center of Section 32. Good
foraminiferal fauna similar to no. 7 of Hex Creek, 40095 (CAS). C. C. Church,
collector.

Locality 40094 (CAS). Clay test pit, southwestern extremity of Hex Hill
about 75 feet east of the mouth of Hex Creek and the center of Section 31. C.C.
Church, collector.

Locality 40095 (CAS). Station no. 7, Hex Creek. From the lower slope of
a south-facing small ridge projecting into Hex Creek from the west causing it to
veer to the northeast, about 1150 feet south and 200 feet east of north quarter
corner, Section 31, Township 25 south, Range 18 east of the Mount Diablo base
and meridian calcareous and arenaceous foraminifera. Station no. 8, from
creek bed about 100 feet north of no. 7 is also included in Locality 40095 (CAS).
Fauna same as no. 7. C. C. Church, collector.

Locality 40096 (CAS). Field numbers 10 and 11, late Cretaceous from
clay shale bed 3 feet thick above thick bedded sandstone, dipping about 85
degrees north, south side of gully from northwest near bend in Hex Creek and
70 feet up gully from creek. About 1000 feet south and 600 feet east of the
north quarter corner, Section 31. Turonian, G-2 stage. C. C. Church, collector.

DESCRIPTION OF FORAMINIFERA

Order FORAMINIFERIDA

Suborder TEXTULARIINA

Superfamily AMMODISCACEA
Family AMMOoDISCIDAE Reuss, 1862
Subfamily AMMODISCINAE Reuss, 1862

Genus Ammodiscus Reuss, 1862

VoL. XXXII] CHURCH: CRETACEOUS FORAMINIFERA 533

Ammodiscus glabratus CUSHMAN and JARVIS, 1928.
(Plate 7, figure 9.)

Ammodiscus glabratus CUSHMAN and JARvIS, 1928, Contrib. Cushman Lab. Foram. Res.,
vol. 4, p. 86, pl. 12, figs. 6 a, b.

This species occurs commonly in strata of upper Cretaceous and of Paleocene
age and seems to be very much the same in the Devils Den-Hex Hill area. It
was found at sample localities nos. 1, 2, and 5 of Devils Den and no. 46 (F. A.
Menken). At Hex Hill it was found at nos. 1 and 15 and the clay pit near the
mouth of Hex Creek.

The same species was described by Lewis Martin in his paper on the upper
Cretaceous of the Panoche Hills, Fresno County, California (1964) but he ap-
plied the generic name /nvolutina. This use of the generic name, /nvolutina,
was, no doubt, due to the confusion which involved the two names for so long.
This appears to have been straightened out by the research of A. R. Loeblich
and Helen Tappan (1961, p. 187). They conclude that the name Ammodiscus
applies only to the arenaceous species while /nvolutina includes the calcareous
species. This point is stressed here because both genera are present in the Devils
Den material and, to this author’s knowledge, true /nvolutina has not been
previously reported from the early Cretaceous of California. Length .61 mm.

Superfamily LITUuOLACEA
Family TROCHAMMINIDAE Schwager, 1877
Subfamily TROCHAMMININAE
Genus Trochammina Parker and Jones, 1859

Trochammina species.
(Plate 1, figures 1 a, b, c.)

A fairly common species at nos. 7 and 8 of Hex Creek and no. 15 of Hex
Hill but not found at Devils Den.

Trochammina species.

This questionable arenaceous species .84 mm. long, is trochospiral and in
shape and arrangement of the chambers similar to the genus Gyroidina. It is
irregularly flattened on the dorsal side and tapers to a conical point at the
base. The chambers are broad and narrow and somewhat longer than wide
and the apertural face correspondingly narrow. There are five chambers in the
outer whorl and their overlapping at the base forms a conical depression.
There is also a rounded pit or depression in the center of the dorsal side. There
is a suggestion of a small arched opening or aperture about midway of the
base of the apertural face in one or two specimens but in most of them no
aperture can be seen.

534 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

This is one of the more distinctive species in this fauna but in the literature
consulted, nothing like it was found. It was found in only two of the localities
at Hex Hill, one at the mud pit near the mouth of Hex Creek in the southwest
corner of the northeast quarter of Section 31, Township 25 south, Range 18
east and the other on the west face of a deep gully in the southeast corner of
northwest quarter of Section 32, Township 25 south, Range 18 east of the
Mount Diablo base and meridian.

Trochammina orchardensis Church, new species.
(Plate 2, figures 11 a, b, c.)

Test free, trochospiral, agglutinated, dorsal side concave with the rim of the
final whorl curving upward, surface roughened by irregularly raised chambers
and slightly depressed sutures giving the edge a gently scalloped effect; sutures
composed of more transparent cementing material, slightly depressed near
central areas on the umbilical side, flush with surface or slightly raised on the
dorsal side, sutures straight on umbilical side, sharply curved on the dorsal
side. The thickness of the test and degree of concavity varies, the concavity on
the umbilical side being smoothly rounded and shallow, the dorsal side concave
to nearly flat with a rugose surface, there are ten chambers in the final whorl,
the aperture appears to be a narrow opening at the base of the final chamber.
Length .59 mm. Width .45 mm.

Ho.ortype no. 12968 (California Academy of Sciences, Department of Geol-
ogy Type Collection), from Locality 40092 (CAS), from west slope of the most
easterly deep gully on Hex Hill, about 200 feet north and 400 feet west of the
center of Section 32, Township 25 south, Range 18 east of the Mount Diablo
base and meridian, northwestern Kern County, California; C. C. Church, col-
lector; early Cretaceous.

This species has been found at Hex Hill no. 1 and nos. 1 and 5 of Devils
Den, also more easterly, no. 46 (F. A. Menken). At Hex Hill it is quite com-
mon.

Genus Glomospira Rzehak, 1885

Glomospira gordialis (Jones and Parker).
(Plate 1, figure 2.)
Trochammina squamata var. gordialis JoNEs and PARKER, 1860, Quart. Journ. Geol. Soc.,

vol. 16, p. 304.

Glomospira gordialis CusHMAN, 1918, U. S. Nat. Mus., Bull. 104, pt. 1, p. 99, pl. 36, figs.
7-9.

Of all the species found in this fauna, the present species is one of the most
common. It is also probably one of the longest ranging species as it has been
reported from the Silurian to the Recent. It was found in most samples from
Devils Den, particularly from stations nos. 1, 5, 8, and no. 46 (F. A. Menken).

VoL. XXXII] CHURCH: CRETACEOUS FORAMINIFERA 53

It was found at Hex Hill at nos. 1 and 15 and the Clay pit, also no. 3 of
Hex Creek. Diameter .25 mm.

Family Lituotmaer de Blainville, 1825
Subfamily LITUOLINAE de Blainville, 1825
Genus Bulbophragmium Maync, 1952

Bulbophragmium species.
(Plate 2, figure 2.)

This small, arenaceous species would seem to fall in the above genus since
it appears to have a cribrate aperture and a bulbous, streptospiral initial stage,
the transitional stage to the uniseral is very short and the biserial stage not
clearly demonstrated. There are eight to ten uniseral chambers, all very thin
and much broader than high, they are rounded to oval in cross section, the
increase in diameter with growth is very slight. The test in general, is very
similar in appearance to Ammobaculoides romaensis Crespin from the lower
Cretaceous of Australia except for the aperture.

This is not a common species and so small that it is easily overlooked. It
was found in only one sample of the Hex Formation of Hex Hill, Station no. 1
and at nos. 1 and 5 of Devils Den. Length .59 mm.

Genus Triplasia Reuss, 1854
Triplasia species.

This very small, arenaceous species is not common in the Hex Formation but
was found at stations no. 1 of Hex Hill and no. 46 (F. A. Menken) of Devils
Den. It also was found at no. 5 and above in the same canyon. It appears to be
uniserial and concave triangular from the beginning and the three edges are
beaded and irregular with most tests having a curved and somewhat twisted
shape. There is no apparent neck or projection at the apertural end.

Subfamily HAPLOPHRAGMOIDINAE Maync, 1952

Genus Cribrostomoides Cushman, 1910
Cribrostomoides species.

A rather robust species, fairly common where it does occur as at stations
nos. 3, 4, 6, 7, 8, and 15 of Hex Creek, not found at Devils Den.

This is a very thick, close coiled species, almost completely involute, rounded
in cross section, sutures at right angles to periphery, very slightly to moderately
depressed, seven to nine chambers to a whorl, aperture obscure but appears to
be cribrate, .6 mm. to .7 mm. in diameter composed to fairly coarse sand grains
but smoothly finished,

536 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Genus Haplophragmium Reuss, 1860

Haplophragmium aequale (Roemer) 1841.
(Plate 2, figure 10.)
Spirolina aequalis RoEMER 1841, Kreidegeb. 98, pl. 15, fig. 27.
Haplophragmium aequale Roemer sp., 1863, Reuss, Hils u. Gault, p. 29, pl. 1, figs. 1-7.
Haplophragmium aequale (Roemer) 1841, BARTENSTEIN, 1952, Senckenbergiana, vol. 33,
nos. 4-6, p. 325, pl. 1, figs. 2-11; pl. 2, figs. 17-26; pl. 3, figs. 1-6; pl. 6, figs. 6-8;
pl. 7, figs. 1-2.
Of the arenaceous species this one is very common at Devils Den, Stations
1 to 7 and at Hex Hill no. 1. In the paper by Bartenstein (1952), a wide varia-
tion in this species is shown. In the present instance the species shows very little
variation. Length .48 mm.

Genus Haplophragmoides Cushman, 1910

Haplophragmoides species.

(Plate 1, figures 7 a, b.)

This species is irregular, trochospiral, somewhat compressed and distorted,
aperture not apparent, not a common species where it occurs at Hex Hill no. 1.
Length .53 mm.

Family ATAXOPHRAGMIIDAE Schwager, 1877
Subfamily GLOBOTEXTULARIINAE Cushman, 1927

Genus Dorothia Plummer, 1931

Dorothia PLUMMER, 1931, Bureau Econ. Geol., Texas Univ., Bull. no. 3101, p. 130.
Marssonella CUSHMAN, 1933, Cushman Lab. Foram. Res., Contrib., vol. 9, pt. 2, p. 136.

Dorothia oxycona (Reuss).

(Plate 1, figures 4, 6 a, b.)

Gaudryina oxycona Reuss, 1860, K. Akad. Wiss. Wien, Math.-Naturw. Cl. Sitzungsber, vol.
AKO) jo), BS) joll, WA, ines, Si

Marssonella oxycona CUSHMAN, 1933, Cushman Lab. Foram. Res., Contrib., vol. 9, p. 36,
jouh, 24 aire, 113.

Dorothia oxycona (Reuss), TRUJILLO, 1960, Jour. Paleo., vol. 34, no. 2, p. 309, pl. 44, figs.
5 fly lek

This widespread and generally long range species is one of the most common
forms found in the Devils Den section, particularly in the part from nos. 2 to 6.
There is considerable variation in the species from the typical D. oxycona to
specimens with a wide, flaring, final two chambers and figured as M. trochus
(d’Orbigny) in a number of papers, e.g., lower Cretaceous of Trinidad by Bar-
tenstein, Bettenstaedt, and Bolli, plate 3, figure 44. Smaller and less typical

VoL. XXXII] CHURCH: CRETACEOUS FORAMINIFERA 537

specimens were found at station no. 1 of Hex Hill. Length .92 mm. (fig. 4).
Length .49 mm. (figs. 6 a, b).

Dorothia species.

(Plate 2, figure 5.)

Test small with a very short triserial initial stage followed by nine pairs of
short, slightly inflated chambers very faintly outlined by the very gently de-
pressed sutures, test slightly curved and with a partial twist, oval in cross section,
composed of very fine sand grains and having a quite smooth surface. One of
the commoner species at no. 5 at Devils Den and no. 40 (F. A. Menken).
Length .71 mm.

Genus Eggerella Cushman, 1933

Eggerella species B, Stelck, Wall, Bahan, and Martin.
(Plate 1, figure 5.)

Eggerella sp. B, STELCK, WALL, BAHAN, and Martin, 1956, Res. Council Alberta, Canada,
Repanow/5 py ois ple 4 tig: 7:

The species with which the Devils Den form is identified is from the middle
Albian of western Canada. The Devils Den species is one of the rarer forms in
that fauna having been found at only one station, no. 5 where only a few were
found. As in many of the other arenaceous species in this fauna, the aperture is
obscure. Length .30 mm.

Subfamily VERNEUILININAE Cushman, 1911

Genus Pseudoreophax Geroch, 1961

Pseudoreophax cisovnicensis Geroch.
(Plate 2, figure 1.)

Pseudoreophax cisovnicensis GEROCH, 1961, Polskiego Towarzystwa, Rocznik., vol. 31, pt. 1,
pp. 159-167, pl. 17, text figs. 1, 2.

Where it was found this was one of the common species and while it is small
it is very distinctive. The five uniserial chambers are, in general, shorter than
they are broad and variable in shape and size. The test is usually curved and
the chambers increase very little in diameter with growth.

The aperture is indicated by a short, pointed projection, usually well to
one side of the final chamber or just off center. The test varies in cross section
from round to oval and the initial end is rounded and bulbous.

It was found at stations no. 1 at the east end of Hex Hill and at no. 46
(F. A. Menken) of Devils Den, the farthest east of the samples in that locality.
Length 53 mm.

538 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Plate 1

All specimens illustrated on this plate are in the California Academy of Sciences, Depart-
ment of Geology Type Collection.

Ficure 1. Trochammina species. Length .84 mm.; (a) ventral view; (b) dorsal view;
(c) peripheral view. Hypotype no. 12951 (CAS). From Locality 40093 (CAS), southeast
side of Hex Hill, northwestern Kern County, California. Page 533.

Ficure 2. Glomospira gordialis (Jones and Parker). Length .25 mm.; side view. Hypo-
type no. 12952 (CAS). From Locality 40092 (CAS); east side of Hex Hill, NW. Kern
County, California. Page 535.

Ficure 3. Gaudryinella almgreni Church, new species. Length 1.24 mm., width .59 mm.;
side view. Holotype no. 12953 (CAS). From Locality 40095 (CAS), sample 7, Hex Creek,
NW. edge of Hex Hill, NW. Kern County, California. Page 540.

Ficure 4. Dorothia oxycona (Reuss). Length .92 mm.; side view. Hypotype no. 12954
(CAS). From Locality 40091 (CAS), sample 5, deep north and south gully, east side of
Devils Den, NW. Kern County, California. Page 536.

Ficure 5. Eggerella species B, Stelck, Wall, Bahan and Martin, Length 30 mm.; side
view. Hypotype no. 12955 (CAS). From Locality 40091 (CAS), sample 5, deep north-
south gully, east side of Devils Den, NW. Kern County, California. Page 537.

Ficure 6. Dorothia oxycona (Reuss). Length .49 mm.; (a) side view; (b) apertural
view. Hypotype no. 12957 (CAS). From Locality 40092 (CAS), most easterly deep gully
of Hex Hill, NW. Kern County, California. Page 536.

Ficure 7. Haplophragmoides species. Length 53 mm.; (a) side view; (b) peripheral
view. Holotype no. 12957 (CAS). From Locality 40092 (CAS), most easterly deep gully
of Hex Hill, NW. Kern County, California. Page 536.

Vol. XXXII CHURCH: CRETACEOUS FORAMINIFERA Plate 1

VoL. XXXIT] CHURCH: CRETACEOUS FORAMINIFERA

UL
WwW
Ne}

Family ASTRORHIZIDAE Brady, 1881
Subfamily HIPPOCREPININAE Rhumbler, 1895
Genus Hyperammina Brady, 1878

Hyperammina elongata Brady, 1878.

Hyperammina elongata Brapy, 1878, Ann. and Mag. Nat. Hist., ser. 5, vol. 1, p. 433, pl. 20,
figs. 2 a, b. CusHMAN, 1946, U. S. Geol. Surv., Prof. Pap. 206, p. 15, pl. 1, figs. 12, 13.

This long-range species occurs at stations no. 7 of Hex Creek and nos. 5 and 6
of Devils Den. From its long range it could be expected to occur in any of the
samples where other arenaceous species are preserved. In its present occurrence
it was not a common species.

Family TEXTULARIIDAE
Subfamily TEXTULARIINAE
Genus Bigenerina d’Orbigny, 1826

Bigenerina antiquissima Bartenstein and Brand.
(Plate 2, figure 4.)

Bigenerina antiquissima BARTENSTEIN and BRAND, 1951, Abh. senckenb. naturf. Ges., no. 485,
p. 275, pl. 3, figs. 73, 74.

The first few chambers are irregularly biserial and twisted, short and closely
set (microspheric form), with the next few chambers larger and longer and
quickly becoming cuneate before the final one or two uniserial chambers. The
megalospheric form is inflated and bulbous in the initial stages with the biserial
stage short and less distinct, the test curved and twisted, aperture centrally lo-
cated, rounded and projecting with a short neck having a rough, irregular edge.
In the later chambers the degree of inflation varies so much that the tests have
a knobby, twisted shape. The test is composed of a high percentage of cement
and fine sand resulting in a smooth and somewhat translucent surface.

This species was found most abundant in the most northerly of the samples,
near the center of the Devils Den, its is also found at stations nos. 5, 6, and
no. 46 (F. A. Menken) of Devils Den, also at no. 1 at the easterly end of Hex
Hill. Length .47 mm.

Bigenerina deciusi Church, new species.
(Plate 2, figure 9.)

Test finely arenaceous, biserial for the first four pairs of chambers, chambers
enlarge very rapidly with growth becoming inflated and cuneate and finally
uniserial in mature specimens, the uniserial part consisting of one or two
chambers, test slightly twisted in early portion and curving with the change to
the uniserial stage, biserial stage best developed in the microspheric form, in the
megalospheric form only one or two biserial pairs of chambers can be recognized,

540 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

initial chamber blunt and rounded, aperture rounded with a ring-like neck raised
very slightly above the spherical surface at the apex of the final chamber, the
position remaining the same for both biserial and uniserial stages, in its early
stages it is similar in appearance to Bimonilina variana Eicher but that species
has an elongate, slit-like aperture and does not develop a uniserial stage, it
also closely approximates the genus Haeus/arella in its cuneate chamber develop-
ment but the aperture is more centrally located and mature specimens are uni-
serial. Length .43 mm.

HototyrPE no. 12966 (California Academy of Sciences, Department of
Geology Type Collection), from Locality 40092 (CAS), from west slope of the
most easterly deep gully on Hex Hill, about 200 feet north and 400 feet west of
the center of Section 32, Township 25 south, Range 18 east, Mount Diablo base
and meridian, northwest Kern County, California; C. C. Church collector; early
Cretaceous.

Bigenerina deciusi was found at Locality 40092 (CAS) and Locality 40081
(CAS) in Devils Den area.

This species is named in honor of Mr. L. C. Decius, Geologist, San Francisco,
California.

Subfamily VERNEUILININAE

Genus Gaudryinella Plummer, 1931

Guadryinella almgreni Church, new species.
(Plate 1, figure 3.)

Test finely arenaceous, triserial and triangular in the initial stage with flat
to gently concave faces, biserial and compressed laterally for the next three
paired chambers and finally uniserial for one or two chambers, the biserial por-
tion may be less than three paired chambers or not apparent at all, in the uni-
serial stage the chambers may be irregularly rounded, triangular or quadrate
and compressed with distinct downward curving lobes which project as knobs,
giving the upper one-half or two-thirds of the test a knobby, irregular surface
which from test to test, follows no set pattern, the uniserial stage is attained in
less than half the specimens, the greater percentage reaching the somewhat
cuneate, biserial second stage only with the aperture near or at the inner edge
of the last chamber, whereas in the uniserial stage it is a central depression and
approximately round with a slightly raised corona in some specimens. Length
1.24 mm., width, .59 mm.

HoLotyPeE no. 12953 (California Academy of Sciences, Department of Geol-
ogy Type Collection), from Locality 40095 (CAS), sample no. 7 from the
lower slope of a south-facing small ridge projecting into Hex Creek from the
west causing it to veer to the northeast, about 1150 feet south and 200 feet
east of the north quarter corner of Section 31, Township 25 south, Range 18

VoL. XXXII] CHURCH: CRETACEOUS FORAMINIFERA 541

east, Mouth Diablo base and meridian, Hex Hill area, Kern County, California;
C. C. Church collector; early Cretaceous.

The species here described differs from the original description of Gaudryi-
nella in having a more compressed, distinct, biserial stage and appressed uniserial
chambers which are only slightly inflated but the overall description best places
it in this genus.

This species was found at station no. 15 on Hex Hill and nos. 7 and
8 on Hex Creek where it is one of the most common species.

This species is named in honor of Mr. Alvin A. Almgren of Bakersfield,
California.

Superfamily NoposARIACEA
Family NopDOSARIIDAE
Subfamily NODOSARIINAE
Genus Nodosaria Lamarck, 1812

Nodosaria elegantia Lalicker.
(Plate 5, figures 4, 5.)

Nodosaria elegantia LALICKER, 1950, Univ. Kansas, Paleo. Contrib., art. 2, p. 15, pl. 2, figs.
9 a-d.

The above species occurs typically at stations 3, 5, and 6 at Devils Den
with less typical but similar forms at stations 15 of Hex Hill and no. 7 of Hex
Creek. It is usually present in small numbers.

The species was originally described from the Ellis group of Kansas of
middle and late Jurassic age. It has six straight, round-bottomed, intercostate
channels and six sharply keeled, straight costae extending unbroken from the
rounded upper end to its bluntly pointed, apiculate initial end. The Devils Den
species has a well defined spine at the initial end but this is sometimes broken
off giving it the rounded appearance which Lalicker describes as characteristic
of the Jurassic species. Length .55 mm., (fig. 4), .80 mm., (fig. 5).

Nodosaria humilis Roemer.
(Plate 7, figure 7.)

Nodosaria humilis RoEMER, 1841, Kreidebirges, Hannover, p. 95, pl. 15, fig. 6.
Glandulina humilis RoEMER, 1934, Eichenberg, Hauterive, p. 174, pl. 16, fig. 9; pl. 11, fig. 15.

Pseudoglandulina humilis (Roemer), BARTENSTEIN and BRAND, 1951, Abh. senckenb. naturf.
Ges., no. 485, p. 315, pl. 10, figs. 266-271.

Cushman gave the new generic name Pseudoglandulina to this type of nodo-
sarian previously classed as Glandulina to separate them from the Glandulinas
which were derived from the Polymorphinidae. More recently, in the Treatise,
Loeblich, and Tappan included the genus under Nodosaria which places it

542 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Plate 2

All specimens illustrated on this plate are in the California Academy of Sciences, Depart-
ment of Geology Type Collection.

Ficure 1. Pseudoreophax cisovnicensis Geroch. Length .53 mm.; side view. Hypotype
no. 12958 (CAS). From Locality 40092 (CAS), most easterly deep canyon of Hex Hill,
NW. Kern County, California. Page 537.

Ficure 2. Bulbophragmium species. Length .59 mm.; side view. Hypotype no. 12959
(CAS). From Locality 40092 (CAS), most easterly deep gully of Hex Hill, NW. Kern
County, California. Page 535.

Ficure 3. Dorothia cf. D. oxycona (Reuss) var. Length 37 mm.; (a) side view; (b)
apertural view. Hypotype no. 12960 (CAS). From Locality 40091 (CAS), sample 8, deep
gully, east side of Devils Den, NW. Kern County, California.

Ficure 4. Bigenerina antiquissima Bartenstein and Brand. Length 47 mm.; side view.
Hypotype no. 12961 (CAS). From Locality 40092 (CAS), most easterly deep gully of Hex
Hill, NW. Kern County, California. Page 539.

Ficure 5. Dorothia species. Length .71 mm.; side view. Hypotype no. 12962 (CAS).
From Locality 40091 (CAS), sample 5, deep canyon, east side of Devils Den, NW. Kern
County, California. Page 537.

Ficures 6, 7, 8. Identity unknown; side views. Hypotype nos. 12963, 12964, 12965
(CAS). Figure 6. Length 57 mm. Figure 7. Length 55 mm. Figure 8. Length 49 mm.
From Locality 40091 (CAS), sample 5, deep gully east side of Devils Den, NW. Kern
County, California.

Ficure 9. Bigenerina deciusi Church, new species. Length .43 mm.; side view. Holo-
type no. 12966 (CAS). From Locality 40092 (CAS), most easterly gully of Hex Hill,
NW. Kern County, California. Page 539.

Ficure 10. Haplophragmium aequale (Roemer). Length .48 mm.; side view. Hypotype
no. 12967 (CAS). From Locality 40092 (CAS), most easterly deep gully of Hex Hill, NW.
Kern County, California. Page 536.

Figure 11. Trochammina orchardensis Church, new species. Length .59 mm., width
45 mm.; (a) ventral view; (b) dorsal view; (c) peripheral view. Holotype no. 12968 (CAS).
Locality 40092 (CAS), most easterly deep gully of Hex Hill, NW. Kern County, California.
Page 534.

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VoL. XXXII] CHURCH: CRETACEOUS FORAMINIFERA 543

back under its original name. It occurs rather sparingly at station no. 7 of Hex
Creek. The species was listed and figured in the paper on the lower Cretaceous
of Trinidad, British West Indies by Bartenstein, Bettenstaedt, and Bolli. Length
.69 mm.

Nodosaria mutabilis (Reuss).

Glandulina mutabilis Reuss, 1863, K. Akad. Wiss. Wien, math.-naturwiss. Cl., Sitzungsber,
vol. 46, p. 58, pl. 5, figs. 7, ? 8.

Nodosaria (G.) mutabilis Reuss, CHAPMAN, 1893, Jour. Roy. Microsc. Soc. London, no. 4,
p. 585, pl. 8, figs. 19, 20.

Pseudoglandulina mutabilis mutabilis (Reuss, 1863), BARTENSTEIN and Branp, 1951, Abh.
senckenb. naturf. Ges., no. 485, p. 315, pl. 15 C, fig. 10; pl. 14 C, fig. 36.

Only a few of this species were found at station no. 7 of Hex Creek. None
were found at Devils Den. It is apparently a long range species as it occurs in
the upper Cretaceous.

Nodosaria sceptrum Reuss.

Nodosaria sceptrum Reuss, 1863, Hils u. Gault, p. 37, pl. 2, fig. 3. BaRTENSTEIN, BETTEN-
sTAEDT and Bort, 1957, Eclog. Geol. Helv., vol. 50, no. 1, p. 35, pl. 7, figs. 150 a, b.

This is one of the rarer species and was found only at Devils Den, stations
nos. 1 to 5. It varies somewhat in size but retains the shape and costation typical
of the species. It has been figured from the lower Cretaceous of Trinidad, British
West Indies, and northwest Germany where it occurs in the Valanginian. It is
very similar to NV. amphioxys Reuss as figured by Helen Tappan from the
Grayson formation of north Texas (Albian).

Nodosaria cf. N. tenuicosta Reuss.
(Plate 6, figures 3, 5)

Nodosaria cf. N. tenuicosta Reuss, 1845, Verstein. bohmischen Kreide-formation. Stuttgart,
Germmrn Ee schweiz.) Abths isp. 25. pl. 13. tess 5.16:

The present species is almost straight-sided with very little restriction at the
sutures to set the chambers apart. There are six continuous costae running the
length of the test but they are low and narrow. The megalospheric form has a
round proloculus with a short spine. The initial chamber is larger than the
succeeding chambers. In the microspheric form the initial chambers are smaller
and the additional chambers enlarge very gradually, the costae are very fine
and low. The species occurs only at Devils Den at stations 5 and 6. Another
form, at first thought to be a separate species, is now thought to be the micro-
spheric form of this species. Length 1.15 mm. (fig. 3), length .92 mm. (fig. 5).

544 CALIFORNIA ACADEMY OF SCIENCES [PRoc. 4TH SER.

Nodosaria species.

A distinctive but rare species the first two or three chambers are small,
straight-sided, and smooth, the following three or four chambers growing much
larger but with no restriction at the sutures and no ornamentation. It was found
at stations nos. 4 and 5 at Devils Den.

Nodosaria hexensis Church, new species.
(Plate 6, figure 13.)

Test free, multilocular, rectilinear, calcareous, composed of five ovoid cham-
bers each slightly longer than wide, round in cross section, surface smooth,
chambers enlarge gradually and regularly from the initial spheroid chamber,
aperture small, round, at the end of a short, tube-like neck at the apex of the
last chamber, sutures marked by a moderate restriction with a narrow welt at
the juncture, chambers with moderate overlap of previous chambers, similar in
general appearance to Pseudoglandulina tenuis (Bornemann) but with different
aperture, found only at stations 5 and 6 at Devils Den. Length 1.01 mm., width
.24 mm.

HototypPe no. 13015 (California Academy of Sciences, Department of
Geology Type Collection), from Locality 40091 (CAS), from deep north-south
trending V-shaped gully beginning 500 feet south and 700 feet west of the east
quarter corner of Section 20, Township 25 south, Range 18 east, Mount Diablo
base and meridian, Kern County, California. Station no. 5, east bank of gully
approximately 150 feet north of the mouth of the gully; C. C. Church collector;
early Cretaceous.

Dentalina catenula Reuss.

Dentalina catenula Reuss, 1860, Akad. Wiss. Wien, Math.-naturwiss. KI., Sitzungsber., vol. 40,
p. 185, pl. 3, fig. 6. CusHMan, 1940, Contrib. Cushman Lab. Foram. Res., vol. 16, pt. 4,
p. 81, pl. 13, figs. 29-34.

This is a variable and relatively common form at Hex stations nos. 7 and 8.
A similar species is present at no. 4 of the Devils Den outcrop.

Dentalina communis d’Orbigny.

Dentalina communis v’ORBIGNY, 1840, Soc. Géol. France, Mém., vol. 4, p. 13, pl. 1, fig.
4. BARTENSTEIN, BETTENSTAEDT and Botti, 1957, Eclog. Geol. Helv., vol. 50, no. 1, p. 34,
pl. 7, figs. 144, 145.

One of the more common species of the Devils Den section where it is found
in samples from stations nos. | to 6.

VoL. XXXII] CHURCH: CRETACEOUS FORAMINIFERA

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Dentalina grahami Church, new species.
(Plate 3, figures 6 a, b.)

Test short, consisting of five chambers, each nearly circular in cross-section,
gently arcuate, slight flattening on the inner curved side, chambers somewhat
wider than high, sutures almost at right angles in the first three chambers,
slightly oblique in the last two, the aperture projects as a tapered, tubular neck
flush with the inner curve of the test and forms a continuous arch with the
inner edge of the test, sutures slightly depressed, sides of test ornamented with a
few, longitudinal, more or less discontinuous, low costae which do not extend
over the last chamber, initial chamber large, globular with second chamber
slightly smaller but succeeding chambers increasing in width very gently to the
last chamber. Length .66 mm., width .13 mm.

HoLotyre no. 12974 (California Academy of Sciences, Department of
Geology Type Collection), from Locality 40091 (CAS), from deep north-south
trending V-shaped gully beginning 500 feet south and 700 feet west of the east
quarter corner of Section 20, Township 25 south, Range 18 east, Mount Diablo
base and meridian, Devils Den area, Kern County, California. Sample no. 1,
east bank of gully near base, approximately 75 feet north of the mouth of the
gully; C. C. Church collector; early Cretaceous.

This species was taken at stations nos. 1 and 5 in the Devils Den area,
Locality 40091 (CAS).

This species is named in honor of Dr. Joseph J. Graham, Professor of
Micropaleontology at Stanford University, Stanford, California.

Dentalina species.
(Plate 6, figure 2.)

Only one complete specimen of this species was found. The early chambers
are compressed but the later ones are rounded in cross section and separated
by a small degree of inflation and a restriction at the sutures. The entire test
is marked by fine longitudinal, closely spaced costae.

Found only at the most northerly of the Devils Den canyon series which
is near the center of the group of hills known as ‘“‘Devils Den.” Length 1.90 mm.

Genus Vaginulina d’Orbigny, 1826

Vaginulina recta Reuss.
(Plate 5, figures 1, 3.)
Vaginulina recta Reuss, 1863, Hils u. Gault, Akad. Wiss. Wien, Sitz., vol. 46, p. 48, pl. 3,
figs. 14, 15.
A rather rare species at station no. 7 on Hex Creek but more common
at station no. 2 in Devils Den. This is a long and narrow species which remains
about the same width throughout its growth after the first globular chamber

546 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER.

and the one following, test straight to gently curved, chamber edges raised into a
sharp ridge forming an unbroken rim on each side of the test. This is one of
the more distinctive species of the fauna but it is believed to range upward into
the upper part of the lower Cretaceous. Length .72 mm. (fig. 1), length .57 mm.

(tige 3))):

Vaginulina octocostata Church, new species.

(Plate 5, figure 2; plate 6, figure 6.)

Test free, calcareous, rectilinear, gently curved, compressed ovate in section,
surface marked by eight vertical, more or less continuous costae two of which are
at the outer edges of the test and three equally spaced on each side, all irregularly
carinate and extending from the initial end to the edge of the final chamber
where they merge around the apertural face, often forming a slight depression
and extending to the aperture at the peripheral angle making it more apiculate,
chambers enlarge very gradually for the first two or three then continue with
little change for the final two or three chambers which are usually five or six
in number, initial chamber small and pointed with a short basal spine in the
microspheric form and compressed spherical in the megalospheric form, addi-
tional chambers compressed to gently inflated and defined by gentle depressions
at the sutures. In the more extreme developments, as in figure 6, plate 6, the
species has a more rounded final chamber and more restricted suture as in
Dentalina. Without the intermediate forms one might easily identify the two
extremes as different species or genera. Length .86 mm.

Hototype no. 13009 (California Academy of Sciences, Department of
Geology Type Collection), from Locality 40091 (CAS), from deep north-south
trending V-shaped gully beginning 500 feet south and 700 feet west of the
east quarter corner of Section 20, Township 25 south, Range 18 east, Mount
Diablo base and meridian, Kern County, California. Sample from station no. 8,
from steep east bank of gully about 20 feet south of junction with first northeast
trending branch; this is about 600 feet due west of the east quarter corner of
Section 20, Sawtooth Ridge quadrangle, 1953 edition; C. C. Church collector;
early Cretaceous.

This species was found at stations 5 and 8 at Locality 40091 (CAS), Devils
Den area.

Vaginulina debilis (Berthelin).

(Plate 6, figure 4.)

Marginulina debilis BERTHELIN, 1880, Soc. Géol. France, Mém., ser. 3, vol. 1, Mem. 5, p. 35,
pl. 3, (26) fig. 28.

Vaginulina debilis (Berthelin), TAPPAN, 1940, Jour. Paleo., vol. 14, no. 2, p. 108, pl. 16, figs.
26 a, b.

Vou. XXXII] CHURCH: CRETACEOUS FORAMINIFERA 547

The original description is of a smooth form but a similar species from the
Grayson formation has both smooth and sparsely costate types. The ones with
the costation are very similar to those from the more northerly of the Devils Den
samples but smooth forms are present here also. A similar form without the early
chambers was figured by Bartenstein, Bettenstaedt and Bolli from the lower
Cretaceous of Trinidad. In this instance it was classified as ‘‘Dentalina” debilis
(Berthelin). The Trinidad species appears to be more sharply angled and
flatter on the side. It is also without any vertical costae. Some of the more
attenuated and less angled fragments from Devils Den could very readily be
considered as Dentalina but where perfect specimens were found, the test was
wedge-shaped and compressed. Found only at Devils Den. Length 1.64 mm.

Vaginulina striolata Reuss.
(Plate 4, figure 9; plate 6, figure 14).
Vaginulina striolata Reuss, 1863, Akad. Wiss. Wien, Sitz., vol. 46, pt. 1, p. 46, pl. 3, fig. 7.

Vaginulina kochii var. striolata CUSHMAN and ALEXANDER, 1930, Cushman Lab. Foram. Res.,
Contr., vol. 6, pt. 1, p. 4, pl. 1, figs. 10-16.

Vaginulina striolata Reuss, 1863, BARTENSTEIN and Brann, 1951, Abh. senckenb. naturf., Ges.,
no. 485, p. 294, pl. 6, figs. 161-164.

This may be a variety of Vaginulina kochii as it was considered by Cushman
and Alexander and by Helen Tappan. It is a rather rare species here but was
found at Stations 5 and 6 of Devils Den and at no. 7 of Hex Creek. Length
1.06 mm. (pl. 4, fig. 9). Length 1.07 mm. (pl. 6, fig. 14).

Vaginulina kochii Roemer.
(Plate 4, figure 11.)
Vaginulina kochii RoEMER, 1840-1841, Verst. norddeutsch. Kreide, p. 96, pl. 15, fig. 10.

Tappan, HELEN, 1940, Jour. Paleo., vol. 14, no. 2, p. 109, pl. 17, figs. 2-4.

This is one of the more commonly occurring species at both Hex Creek and
Devils Den but it is not abundant in any of the samples. It is listed from the
lower Cretaceous of Europe, the lower Cretaceous of Trinidad, southeastern
United States and the upper Cretaceous of the Sacramento Valley, California.
Length .96 mm.

Vaginulina riedeli riedeli Bartenstein and Brand.
(Plate 4, figures 7, 8.)
Vaginulina riedeli riedeli BARTENSTEIN and BRAND, 1951, Abh. senckenb. naturf. Ges., no. 485,
p. 295, pl. 7, figs. 165 a, b, c.
One of the less common but very distinctive species of the Hex Hill fauna.
Found at station no. 15, Hex Hill. Length 1.11 mm. (fig. 7), length 1.00 mm.
Gigees:):

548 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Vaginulina truncata Reuss.

Vaginulina truncata Reuss, 1863, Hils u. Gault. p. 47, pl. 3, fig. 9. BARTENSTEIN and BRAND,
1951, Abh. senckenb. naturf. Ges., no. 485, p. 293, pl. 12B.

This rare variety may be only a more flaring variation of V. kochii as similar
species have been described as V. kochii from the German lower Cretaceous by
Bartenstein and Brand. Found at stations no. 5 and no. 8 of Devils Den.

Genus Vaginulinopsis Silvestri, 1904

Vaginulinopsis pachynota Ten Dam.
(Plate 4, figure 3.)

Vaginulinopsis pachynota TEN Dam, 1946, Jour. Paleo., vol. 20, no. 6, p. 575, pl. 88, figs. 5,
6 a, b.

A common and variable form at stations nos. 5 and 6 of Devils Den but not
found in the Hex Hill samples. This strongly sutured species is similar in many
respects to some of the later Eocene forms. Length 1.20 mm.

Genus Marginulina d’Orbigny, 1826

Marginulina bullata Reuss.

Marginulina bullata Reuss, 1845, Verstein. bohm. Kreideformation, pt. 1, p. 29, pl. 13, figs.
34-38. CUSHMAN, 1937, Cushman Lab. Foram. Res., Contrib., vol. 13, p. 96, pl. 14, figs.
9-15. CusHMAN, 1944, Cushman Lab. Foram. Res. Contrib., vol. 20, p. 6, pl. 1, fig. 21.

This species occurs very sparingly in samples nos. 7 and 8 of Hex Creek.
Very similar species are, M. curvitura Cushman and M. texasensis Cushman,
both from the upper Cretaceous of Texas.

Marginulina cf. M. parkeri (Reuss).

(Plate 3, figure 3.)

Lenticulina parkeri Reuss, 1863, Hils u. Gault, p. 59, pl. 5, fig. 14.
Lenticulina (Marginulinopsis) parkeri (Reuss), BARTENSTEIN and Branp, 1951, Abh. sencken.
naturf. Ges., no. 485, p. 288, pl. 6, figs. 136, 137.

This appears, from the figures, to be a much smoother form than the species
to which it is compared. The test is without ornamentation and the sutures
flush with the surface although very distinctly outlined. Even the initial cham-
bers are clearly displayed showing the half coiled nature of the early chambers
which gives to the early portion of the test its distinct but gentle curve. Follow-
ing the first two or three chambers the oval section continues its growth without
appreciable change with parallel, oblique sutures outlining the chambers which
are wider than they are high, the test itself gently curved or almost straight with
smooth, parallel sides. Length 1.74 mm.

Vor. XXXII] CHURCH: CRETACEOUS FORAMINIFERA 549

Marginulina robusta Reuss.
(Plate 4, figure 6.)

Marginulina robusta REUSS, 1863, Hils u. Gault, p. 63, pl. 6, figs. 5, 6.

Lenticulina (Marginulinopsis) robusta (Reuss), 1863, BARTENSTEIN and BRAND, 1951, Abh.
senckenb. naturf. Ges., no. 485, p. 289, pl. 6, figs. 142, 143.

Marginulina robusta (Reuss), TEN DAM 1948, Neocom., Jour. Paleo., vol. 22, no. 2, p. 185,
pl. 32, fig. 6.

This species was found sparingly at stations nos. 1, 4, and 5 of Devils Den
area but not at Hex Creek or Hex Hill. From a comparison with figures and
actual specimens from the European section, it appears to be very close to the
species described by Reuss. Length .67 mm.

Marginulina pyramidalis Koch.
(Plate 6, figure 12.)

Nodosaria pyramidalis Kocu, 1851, Palaeontographical, p. 169, pl. 24, fig. 8.
Marginulina pyramidalis (Koch), BARTENSTEIN and BRAND, 1951, Abh. senckenb. naturf. Ges.,
no. 485, p. 307, pl. 9, figs. 221-223.

A common species at station 5 of Devils Den and found from nos. 2 to 6.
It is quite variable in size but retains its characteristics of strong, continuous
carinate costae and initial spine and prominent, off-center, spout-like aperture,
final chamber noticeably larger and more inflated, initial chamber also somewhat
larger and more bulbous than those immediately following. It is one of the
distinctive species of the Devils Den section but not found in the Hex Hill
area. Length 1.09 mm.

Marginulina sigali Bartenstein, Bettenstaedt, and Bolli.
(Plate 6, figure 8.)

Lenticulina (Marginulina) sigali BARTENSTEIN, BETTENSTAEDT and BOLL, 1957, Eclog. Geol.
Helv., vol. 50, no. 1, p. 32, pl. 5, fig. 99; pl. 6, figs. 130, 131 a, b.

This is a quite common species at station 5 of Devils Den but was not found
in other parts of the section or at Hex Hill. It is a species with distinctive
features and appears from Bartenstein, Bettenstaedt, and Bolli, plate 5, figure 99
(see above) to be the same as their Trinidad species. Length .61 mm.

Marginulina species.
(Plate 4, figure 12.)

This is one of the rarer species of the Hex formation. It was found only at
stations no. 7 of Hex Creek and no. 5 of Devils Den but most likely will be found

CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

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Plate 3

All specimens illustrated on this plate are in the California Academy of Sciences, Depart-
ment of Geology Type Collection.

Ficure 1. Lingulina tenera Bornemann. Length 59 mm.; side view. Hypotype no.
12969 (CAS). From Locality 40091 (CAS), sample 5, deep north-south gully east side of
Devils Den, NW. Kern County, California. Page 560.

Ficure 2. Saracenaria spinosa (Eichenberg). Length .94 mm.; side view. Hypotype no.
12970 (CAS). From Locality 40095 (CAS), sample 7, Hex Creek, NW. edge of Hex Hill,
NW. Kern County, California. Page 552.

Ficure 3. Marginulina cf. M. parkeri (Reuss). Length 1.74 mm.; side view. Hypotype
no. 12971 (CAS). From Locality 40095 (CAS), sample 7, Hex Creek, NW. edge of Hex
Hill, NW. Kern County, California. Page 548.

Ficure 4. Citharina species. Length 1.12 mm.; side view. Hypotype no. 12972 (CAS).
From Locality 40091 (CAS), sample 5, deep north-south gully, east side of Devils Den, NW.
Kern County, California. Page 557.

Figure 5. Citharina acuminata (Reuss). Length 1.27 mm.; side view. Hypotype no.

12973 (CAS). From Locality 40091 (CAS), sample 5, from deep north-south gully, east
side of Devils Den, NW. Kern County, California. Page 556.

Ficure 6. Dentalina grahami Church, new species. Length .66 mm.; width .13 mm. (a)
side view; (b) side view. Holotype no. 12974 (CAS). From Locality 40091 (CAS), sample
5, from deep north-south gully, east side of Devils Den, NW. Kern County, California.
Page 545.

Ficure 7. Frondicularia frankei Cushman. Length 1.21 mm.; side view. Hypotype no.
12975 (CAS). An immature specimen from Locality 40091 (CAS), sample 5, deep north-
south gully, east side of Devils Den, NW. Kern County, California. Page 557.

Ficure 8. Astacolus perobliqua (Reuss). Length .82 mm.; side view. Hypotype no.
12976 (CAS). From Locality 40091 (CAS), sample 6, deep gully, east side of Devils Den,
NW. Kern County, California. Page 556.

Ficure 9. Menkenina berryi Church, new species. Length 1.17 mm.; width 37 mm.;
(a) dorsal view; (b) ventral view. Holotype no. 12977 (CAS). From Locality 40091 (CAS),
sample 6, deep north-south gully, east side of Devils Den, NW. Kern County, California.
Page 561.

Ficure 10. Citharina species “3,” (Bartenstein and Brand). Length .83 mm.; side view.
Hypotype no. 12978 (CAS). From Localities 27501 (CAS), 47 (FAM). Near central area
of Devils Den, NW. Kern County, California. Page 556.

Ficure 11. Astacolus perobliqua (Reuss). Length 1.65 mm.; side view. Hypotype no.
12979 (CAS). From Locality 40091 (CAS), sample 6, deep north-south gully, east side of
Devils Den, NW. Kern County, California. Page 556.

CHURCH: CRETACEOUS FORAMINIFERA Plate 3

Vol. XXXII

Se ee ee Kies
DARA 21a Vase

VoL. XXXII] CHURCH: CRETACEOUS FORAMINIFERA 551

elsewhere in the section. It is almost round in section and with parallel sides and
only a suggestion of a beginning coil, chambers few, usually not more than six
which indicates the nature of the short, thick species, surface smooth and only
slightly indented at the last suture, sutures flush with surface, almost at right
angles to dorsal side in later chambers, aperture at dorsal angle of last chamber.
Length 1.03 mm.

Genus Marginulinopsis Silvestri, 1904

Marginulinopsis gracillissima (Reuss).
(Plate 6, figure 9.)

Cristellaria gracillissima Reuss, 1862, Sitz. Ber. Akad. Wiss. Wien, vol. 46, p. 64, pl. 6, figs.
9, 10.

Marginulinopsis gracillissima (Reuss), TEN Dam, 1948, Jour. Paleo., vol. 22, no. 2, p. 184, pl.
Ss sie 1 tke

This species was found sparingly but consistently through most of the Devils
Den series from 1 to 8. Most specimens do not have the pronounced initial coil
as figured in the original type, but in all other respects is very similar. In
some of the smaller varieties the aperture is drawn out into a distinct neck.
Length .49 mm.

Marginulinopsis collinsi Mellon and Wall.

Marginulinopsis collinsi MELLON and WALL, 1956, Res. Council Alberta, Report no. 72, pt. 1,
PaZOsply 2. tigs. 17:

There is a tendency for this species, from the Hex Creek occurrence, to de-
velop the flattened apertural face of a Saracenaria reminiscent of Lenticulina
valanginiana Bartenstein and Brand of the lower Cretaceous of Germany. Our
Hex Creek, no. 7 occurrence is quite rare but it is possible that it may be less so
with further sampling. The species named by Mellon and Wall from Alberta,
Canada, is from the basal Clearwater formation, considered to be of middle
Albian in age on the basis of its ammonite fauna. A few specimens were also
found at station no. 48 (F. A. Menken) of Devils Den which are very similar to
the species described by Mellon and Wall.

Marginulinopsis species.
(Plate 4, figure 13.)

This species is similar in general shape and degree of uncoiling to Cristellaria
hamata Franke which was described from the Oligocene of Denmark but has
fewer and wider chambers and greater inflation in the final chamber. It is one
of the more common species found in samples from stations 7 and 8 of Hex
Creek, Locality 40095 (CAS) but was not found in the Devils Den samples.
Length 1.03 mm,

CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

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Genus Saracenaria Defrance in de Blainville, 1824

Saracenaria italica Defrance.
(Plate 4, figure 5.)

Saracenaria italica DEFRANCE, 1824, Tableau, p. 176 (vol. 32), pl. 13, fig. 6 (vol. 12).
CHAPMAN, 1894, Jour. Roy. Micr. Soc. London, no. 7, p. 653, pl. 10, fig. 10.

This is not one of the commoner species but it was found at station no. 7
of Hex Creek and later more commonly at nos. 2 and 5 of Devils Den. It is a
large robust form. Length .78 mm.

Saracenaria spinosa Eichenberg, 1935.

(Plate 3, figure 2.)

Saracenaria spinosa EICHENBERG, 1935, Niedersachs. Geol. Ver., Jahesberichts 27 (Mitt.
Roemer-Museum, Hildesheim, no. 37, Teil 1, Folge 4), p. 10, pl. 4, fig. 5 a-d. Aptian,

Germany. TAPPAN, HELEN, 1962, U. S. Geol. Surv., Prof. Paper 236C, p. 165, pl. 41,
figs. 18, 19.

Occurs rather sparingly at station no. 7 of Hex Creek and at station no. 4 of
Devils Den. It is much more slender and elongate than S. italica and a sharp
spine terminates the base of the chambers at the two side angles where the face
and lateral sutures meet. Specimens from the Aptian of northern Germany are
very similar to the Hex Hill specimens. The species figured from Alaska is
shorter and less spinose. Length .94 mm.

Genus Lenticulina Lamarck, 1804

Lenticulina saxonica saxonica Bartenstein and Brand.

(Plate 5, figures 11 a, b.)

Lenticulina saxonica saxonica BARTENSTEIN and Branp, 1951, Abh. senckenb. naturf. Ges.,
no. 485, p. 284, pl. 5, figs. 115 a, b.
This is one of the common forms at stations nos. 7 and 8 of Hex Creek.

It is usually very well preserved as it is large and has a sturdy test. Length
1.29 mm.

Lenticulina (Lenticulina) miinsteri (Roemer).

(Plate 5, figures 7 a, b.)

Robulina miinsteri ROEMER, 1839, Oolith.-Geb., p. 48, pl. 20, fig. 29.

Lenticulina (Lenticulina) miinsteri (Roemer), BARTENSTEIN and BRAND, 1951. Abh. senckenb.
naturf. Ges., no. 485, p. 283, pl. 5, fig. 109.

This is one of the largest and most common of the many species of Lenticu-
lina at stations nos. 7 and 8 of Hex Creek. It varies somewhat in the raised
or depressed condition of the suture but in general it is a quite smooth form with

VoL. XXXII] CHURCH: CRETACEOUS FORAMINIFERA 553

seven to nine chambers, sutures curved, slight depression at the center due
to the overlapping of the last few chambers and concealing a possible umbo,
outer periphery rather sharply keeled and uniform. Length 1.34 mm.

Lenticulina species.
(Plate 4, figure 2.)

A few specimens of this unidentified species were found at station no. 7 of
Hex Creek. It has seven to nine chambers with prominent, sharp raised sutures
which becomes less prominent toward the periphery, each one gently curved,
periphery sharply keeled with a thin carina of clear shell material, raised
sutures may end abruptly near center without coalescing or blend with the
others to form a slightly higher center, height of test from .8 to 1 mm.; width
.5 mm. Length .96 mm.

Lenticulina species.
(Plate 4, figure 1.)

This large, heavily ornamented species is one of the less common of the
large lenticulinas at stations nos. 5 and 6 of Devils Den. From among the
many at this point the present species may be distinguished by the presence of
faint to prominent spiral ridges which generally follow the outer curve of the
test, often being higher and noded over the sutures and strongest near the
umbonal area, sutures may be raised into welts near the center but usually are
flush or depressed near the periphery. In the specimen illustrated the curved
welts are strongest between sutures and more regular and prominent than usual.
Test large, calcareous, free, thick, lenticular, often reaching a diameter of 1.5
mm. with a height only slightly greater than the width, periphery strongly
keeled throughout, surface generally uneven, chambers six to eight in final whorl,
sutures gently curved, aperture typically radial. In rare instances the final
chamber may depart from the curve and extend out from the test in a straight
line resembling a blunt spine in the final chamber. This may be an aberrant
form.

The variability of the species suggests that it may be a variation of one of
the large species with which it is associated but the smaller number of chambers,
the strong peripheral flange and the rugose surface of the test sets it apart.
Length 1.46 mm.

Lenticulina kugleri Bartenstein, Bettenstaedt, and Bolli.

Lenticulina kugleri BARTENSTEIN, BETTENSTAEDT, and BOLti, 1957, Eclog. Geol. Helv., vol. 50,
NOs a2, plo, tie. 95> pl 6, figs. 11/6) a, b:

This species was found only at station no. 47 (F. A. Menken) where it

554 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 2. Lenticulina eichenbergi Bartenstein and Brand. Hypotype no. 12950 (Calif.
Acad. Sci., Dept. Geol. Type Coll.), from the same locality as the specimen shown in figure 3.

Ficure 3. Lenticulina ci. L. eichenbergi Bartenstein and Brand. Hypotype no. 12949
(Calif. Acad. Sci., Dept. Geol. Type Coll.), from Locality 27502 (CAS), no. 50, about 600
feet west and 100 feet north of the center of Section 20, Township 25 south, Range 18 east,
Mount Diablo base and meridian, Kern County, California.

was not a rare species. It is a compressed species with the sutures thickened and
raised and the test longer than broad.

Lenticulina (L.) eichenbergi Bartenstein and Brand.
(Figure 2.)

Lenticulina (Lenticulina) eichenbergi BARTENSTEIN and BRAND, 1951, Abh. senckenb. naturf.
Ges., no. 485, p. 285, pl. 5, figs. 118, 119.

In the earlier samples from Devils Den this species was found at stations
nos. 46 and 50 (F. A. Menken) and later from 5 to 8 of the Devils Den series.
Also at Locality 27605 (CAS), south slope of Hex Hill. It is a large, well-
preserved form with considerable variation in the number and arrangement of
nodes or bosses in the central area and along the curved sutures where they may
at times coalesce to form a welt, both nodes and broken sutures diminish toward
the periphery, usually leaving a smooth outer edge, periphery sharply keeled
to carinate, number of chambers in outer whorl, seven to ten, average height 1
mm., width .8 mm, length 1.02 mm.

Lenticulina cf. L. eichenbergi Bartenstein and Brand.
(Figure 3.)

Of the several species of large lenticulinas of the Devils Den 40091 (CAS),
station 5 locality, this one is probably a variety of L. eichenbergi Bartenstein
and Brand, but in the character of its ornamentation it also has much in

VoL. XXXII] CHURCH: CRETACEOUS FORAMINIFERA

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common with a species described by Bartenstein, Bettenstaedt, and Bolli from
the lower Cretaceous of Trinidad, British West Indies as L. ouachensis (Sigal,
1952). These authors also described three subspecies of L. owachensis, suggest-
ing a proliferation of varieties of the species at this time or possibly a wide
variation in a single species. Regardless of the true or accepted nature of the
species, a similar wide variation is evident among the Devils Den specimens.
The species and its variations are well represented at station 5 at Localities
40091 and 27502 (CAS), station no. 50 (F. A. Menken) both from Devils Den.
Length .88 mm.

Genus Darbyella Howe and Wallace, 1932

Darbyella species.
(Plate 7, figures 10 a, b, c.)

A species from the Jackson Eocene described by Howe and Wallace is very
similar to the lower Cretaceous species except that, in perfect specimens, the
Devils Den species has a typical conical, radiate aperture. The Eocene species
has a slit-like aperture with a calcareous lip on either side.

The authors of the Treatise consider Darbyella as an aberrant form of Len-
ticulina but it is so distinct and constant in its characters that its occurrence
here seems worthy of mention. It was found at stations nos. 4 and 5 of Devils
Den. Length .69 mm.

Genus Astacolus de Montfort, 1808

Astacolus grata (Reuss).

(Plate 4, figure 10.)

Cristellaria grata REusS, 1862, Sitz. Akad. Wiss. Wien, vol. 46, pt. 1, p. 70, pl. 7, figs. 14 a, b.
CusHMAN, 1926, Bull. Amer. Assoc. Petr. Geol., vol. 10, p. 598, pl. 19, figs. 1 a, b.

Marginulina grata (Reuss), CUSHMAN and Jarvis, 1932, Proc. U. S. Nat. Mus., no. 2914,
VOlEGO mat 14 pao pled tiess 7 abe ple Setics. 3) an ib:

This is a very smooth form but the sutures are very distinct as fine lines.
The present species differs slightly from the figured specimen of Cushman and
Jarvis in that the last few chambers do not tend to extend down to the initial
coil but strike across at a more gentle angle. It is quite common at stations nos.
7 and 8 of Hex Creek. Length 1.36 mm.

Astacolus incurvata (Reuss).

Cristellaria incurvata REUsS, 1863, K. Akad. Wiss. Wien, p. 66, pl. 6, fig. 18.
Lenticulina (Astacolus) incurvata (Reuss), BARTENSTEIN, BETTENSTAEDT, and BoLtrt, 1957,
Eclog. Geol. Helv., vol. 50, no. 1, p. 30, pl. 3, figs. 57 a, b; pl. 4, fig. 86.

This species occurs at stations no. 7 of Hex Creek and no. 15 of Hex Hill,
also at no. 3 of Devils Den. The species is very similar to other uncoiled,

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CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

elongate species in these faunas and it is often a question of whether there are
a number of species or if they are variants of a single species.

Astacolus perobliqua (Reuss).
(Plate 3, figures 8, 11.)

Cristellaria (Cristellaria) perobliqua Reuss, 1863, K. Akad. Wiss. Wien, Math., Naturw.
Cl., Sitzungsber., vol. 46, pt. 1, p. 67, pl. 7, fig. 3.

There is considerable variation in this species, especially in the initial cham-
bers which may form a sharply curved, pointed, primary part or start with a
gentle, rounded curve which emerges quickly into a larger slightly curved but
generally parallel sided test with highly oblique sutures and more inflated,
slightly wedge-shaped chambers in section, usually seven chambers.

Fairly common at station no. 6 of Devils Den but found also at nos. 4 and 5
of the same section. Length .82 mm. Another specimen (figure 11) was found
at station 6 which had a length of 1.65 mm.

Genus Citharina d’Orbigny in de la Sagra, 1839

Citharina acuminata (Reuss).
(Plate 3, figure 5.)

Vaginulina acuminata Reuss, 1863, K. Akad. Wiss. Wien, Math.-Naturw. Cl., Sitzungsber.,
vol. 46, pt. 1, p. 49, pl. 4, fig. 1.

Citharina acuminata (Reuss), BARTENSTEIN, BETTENSTAEDT and Botti, 1957, Eclog. Geol.
Helv., vol. 50, no. 1, p. 39, pl. 7, figs. 159 a, b.

One of the rarer species at Devils Den, it was found only at stations nos. 1
and 5. It was not found in the Hex Hill area. Length 1.27 mm.

Citharina species “3,” (Bartenstein and Brand).
(Plate 3, figure 10.)

Vaginulina sp. 3, BARTENSTEIN and BrAnp, 1951, Abh. senckenb. naturf. Ges., no. 485, p. 292,
pl. 6, fig. 153.

There is considerable variation in this species which corresponds very closely
to the species figured by Bartenstein and Brand. It occurs rather commonly at
the most northerly of the Devils Den series of samples and F. A. Menke no. 47
and is scarce but present at station no. 8 of Hex Creek. What appears to be
a Closely related species occurs at station no. 7 on Hex Creek. Length .83 mm.

Citharina kernensis Church, new species.
(Plate 4, figure 4.)

Test small, calcareous, compressed, wedge-shaped in cross section, more
compressed in early chambers, becoming slightly inflated in later ones, early

VoL. XXXII] CHURCH: CRETACEOUS FORAMINIFERA 557

chambers develop a sharp curve, later ones become more oblique and terminate
in a more or less straight line at the back, later chambers tend to overlap the
previous chambers, extending down almost to the curved initial part of the
chambers, aperture a small opening at the apex of the final chamber where the
somewhat angular suture and the sharply angled back side of the test converge
to form a slight neck for the aperture, sutures of clear shell material may be
depressed or raised in a definite welt, in some specimens forming a sharp edge
to the more inflated part of the chambers. This species is definitely narrower
than Citharina species 3 and more inflated but the wide variation in both
species and their general similarities suggests the possibility that they are a
single species. If further work and more detailed comparisons of the species
should support the above possibility, it is this authors suggestion that the name
C. kernensis be retained as the name for both variations. Length .52 mm.,
width .15 mm.

HoLotyPE no. 12983 (California Academy of Sciences, Department of
Geology Type Collection), from Locality 40095 (CAS), sample no. 7, from the
lower slope of a south-facing small ridge projecting into Hex Creek from the
west causing it to veer to the northeast, about 1150 feet south and 200 feet
east of the north quarter corner of Section 31, Township 25 south, Range 18
east, Mount Diablo base and meridian; C. C. Church collector; early Cretaceous.

Citharina species.
(Plate 3, figure 4.)

This very small species may be a variation of C. acuminata but it is much
narrower and the oblique costae are fewer and less continuous than those in
C. acuminata. It is very similar in shape and markings to the much larger
species from the Duck Creek, lower Cretaceous of Texas, which was referred
to as Vaginulina raristriata (Chapman) by Helen Tappan in her paper on the
foraminifera of that formation. It is rare in the Devils Den samples having
been found only at stations nos. 1 and 5. It probably occurs at other points
in the section but its relatively small size and delicate construction renders it
more difficult to detect and less likely to be found whole. Length 1.12 mm.

Genus Frondicularia Defrance in d’Orbigny, 1826

Frondicularia frankei Cushman.

(Plate 3, figure 7; plate 6, figures 1, 10.)

Frondicularia angusta Reuss, 1860, Akad. Wiss. Wien, Math.-naturwiss. K]., Sitzungsber.,
vol. 40, p. 196, pl. 4, fig. 5.

Frondicularia frankei CUSHMAN, 1936, Contrib. Cushman Lab. Foram. Res., vol. 12, pt. 1,
p. 18, pl. 4, figs. 6, 7.

Found only at Devils Den in samples from stations nos. 2 to 6. In all of

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CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Plate 4

All specimens illustrated on this plate are in the California Academy of Sciences, Depart-
ment of Geology Type Collection.

Ficure 1. Lenticulina species. Length 1.46 mm.; side view. Hypotype no. 12980
(CAS). From Locality 40091 (CAS), sample 5, deep north-south gully, east side of Devils
Den, NW. Kern County, California. Page 553.

Ficure 2. Lenticulina species. Length .96 mm.; side view. Hypotype no. 12981 (CAS).
From Locality 40091 (CAS), sample 5, deep north-south gully, east side of Devils Den,
NW. Kern County, California. Page 553.

Ficure 3. Vaginulinopsis pachynota Ten Dam. Length 1.20 mm.; side view. Hypotype
no. 12982 (CAS). From Locality 40091 (CAS), sample 5, deep north-south gully, east side of
Devils Den, NW. Kern County, California. Page 548.

Ficure 4. Citharina kernensis Church, new species. Length .52 mm.; width .15 mm.;
side view. Holotype no. 12983 (CAS). From Locality 40095 (CAS), sample 7, Hex Creek,
NW. edge of Hex Hill, NW. Kern County, California. Page 556.

Ficure 5. Saracenaria italica Defrance. Length .78 mm.; side view. Hypotype no.
12984 (CAS). From Locality 40091 (CAS), sample 5, deep north-south gully, east side of
Devils Den, NW. Kern County, California. Page 552.

Ficure 6. Marginulina robusta Reuss. Length .67 mm.; side view. Hypotype no.
12985 (CAS). From Locality 40095 (CAS), sample 7, Hex Creek, NW. edge of Hex Hill,
NW. Kern County, California. Page 549.

Ficure 7. Vaginulina riedeli riedeli Bartenstein and Brand. Length 1.11 mm.; side view.
Hypotype no. 12986 (CAS). From Locality 40095 (CAS), sample 7, Hex Creek, NW. edge
of Hex Hill, NW. Kern County, California. Page 547.

Ficure 8. Vaginulina riedeli riedeli Bartenstein and Brand. Length 1.00 mm.; side
view. Hypotype no. 12987 (CAS). From Locality 40095 (CAS), sample 7, Hex Creek, NW.
edge of Hex Hill, NW. Kern County, California. Page 547.

Ficure 9. Vaginulina striolata Reuss. Length 1.06 mm.; side view. Hypotype no.
12988 (CAS). From Locality 40095 (CAS), sample 7, Hex Creek, NW. edge of Hex Hill,
NW. Kern County, California. Page 547.

FicurE 10. Astacolus grata (Reuss). Length 1.36 mm.; side view. Hypotype no. 12989
(CAS). From Locality 40095 (CAS), sample 7, Hex Creek, NW. edge of Hex Hill, NW.
Kern County, California. Page 555.

Ficure 11. Vaginulina kochii Roemer. Length .96 mm.; side view. Hypotype no. 12990
(CAS). From Locality 40095 (CAS), sample 7, Hex Creek, NW. edge of Hex Hill, NW.
Kern County, California. Page 547.

Ficure 12. Marginulina species. Length 1.03 mm.; side view. Hypotype no. 12991
(CAS). From Locality 40095 (CAS), sample 7, Hex Creek, NW. edge of Hex Hill, NW.
Kern County, California. Page 549.

Ficure 13. Marginulinopsis species. Length 1.03 mm.; side view. Hypotype no. 12992
(CAS). From Locality 40095 (CAS), sample 7, Hex Creek, NW. edge of Hex Hill, NW.
Kern County, California. Page 551.

CHURCH: CRETACEOUS FORAMINIFERA Plate 4

Vol. XXXII

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VoL. XXXII] CHURCH: CRETACEOUS FORAMINIFERA 559

its occurrences it was found sparingly and usually broken. Length 1.21 mm.
(pl. 3, fig. 7), length 3.19 mm. (pl. 6, fig. 1). [Specimen illustrated in pl. 6,
fig. 10 missing. |

Frondicularia concinna Koch.

Frondicularia concinna Kocw, 1851, Palaeontographica, 1, p. 169, Cassel. REuss, 1863, Sitz.
Akad. Wiss. Wien, p. 54, pl. 4, fig. 13. Hecut, 1938, Abh. senckenb. naturf. Ges. 443,
p. 28 (as Frondicularia D.6), pl. 18b, figs. 116-119.

This is a very distinctive species but also very rare as only two imperfect
specimens were found. One of these has only the first few initial chambers
missing. It was found at station no. 5 of Devils Den.

Frondicularia species.
(Plate 5, figure 10.)
Frondicularia sp. 2, 1957, BARTENSTEIN, BETTENSTAEDT and Botti, Eclog. Geol. Helv., vol.

50, no. 1, p. 40, pl. 5, fig. 109; pl. 6, figs. 138 a, b.

This species, from stations 7 and 8 of Hex Creek, is so similar to the species
from the lower Cretaceous of Trinidad referred to above, that I have used their
names and number. Faint vertical costae are visible on some specimens. It is a
large, robust species, some specimens measuring over 1 mm. in length. In the
earlier half of the test the edge is flattened and forms a distinct right angle
with the two faces. Length 1.22 mm.

Frondicularia species.

This species is similar in some respects to Frondicularia species 2 and may
be a variation of that species. It differs in being flatter and without inflation
of the chambers. Also in many of the specimens the first few chambers are
uniserial. The test as a whole is quite compressed and thin and the edges at
right angles to the sides, the test itself of medium size and the chevron-shaped
chambers have a moderate flair.

Frondicularia species.
(Plate 5, figure 6.)

A very rare species, only two specimens were found and the second one is
probably immature. This species is almost parallel sided as it tapers very
gently with the addition of each chamber, the cross section is a flattened oval
and the edges taper to a thin, narrow carina, the final chamber is attenuated
and the aperture at the end of a rather long, narrow neck, the initial chamber

560 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

is blunt and rounded and compressed into a flat oval which changes little with
growth, surface marked by faint, broken costae. Length .73 mm.

Subfamily LINGULININAE Loeblich and Tappan, 1961

Genus Lingulina d’Orbigny, 1826

Lingulina californiensis Trujillo.

(Plate 6, figure 7.)
Lingulina californiensis TRUJILLO, 1960, Jour. Paleo., vol. 34, no. 2, p. 314, pl. 45, figs. 8 a, b.

This is a rare species in the Hex formation and was found only at no. 7
of Hex Creek. It appears to be identical to Trujillo’s species from the upper
Cretaceous Coniacian of the Sacramento Valley, California. Length .45 mm.

Lingulina tenera Bornemann.

(Plate 3, figure 1.)

Lingulina tenera BORNEMANN, 1854, Gottingen, p. 38, pl. 3, fig. 24. BARTENSTEIN and BRAND,
1951, Abh. senckenb. naturf. Ges., no. 485, p. 301, pl. 8, fig. 194 a, b.

Of infrequent occurrence in the Devils Den series from stations nos. 2 to 5.
Not found at Hex Hill. Reported from as low as the Lias in Europe. Length
.59 mm.

Subfamily MENKENINAE Church, new subfamily

Genus Menkenina Church, new genus
Type Species: Menkenina berryi Church, new species.

The new genus has the usual characters of the uncoiled members of the
family. Test calcareous, perforate with a linear arrangement of the chambers,
differs from Frondicularia in having rounded sutures with a gentle upward
curve on the ventral side and curving upward in a broad, inverted “V” on the
dorsal side, test differs from Vaginulina in being compressed dorso-ventrally
rather than laterally and in having a rounded aperture at the end of a tapered,
tubular neck located at the extreme edge of the dorsal side the trace of which
persists as a central welt on the dorsal side, the genus may have evolved from a
Vaginulina-like ancestor as suggested by some specimens which are more
rounded and inflated on the ventral side, test normally ornamented with a
variable number of broken and continuous, longitudinal, carinate carinae extend-
ing from the initial end to the penultimate chamber, becoming less prominent

Vor. XXXII] CHURCH: CRETACEOUS FORAMINIFERA 561

with growth, one of these forming a thin, translucent flange on the edges of the
test. The new name is in honor of the late Mr. F. A. Menken! who first brought
this fauna to my attention.

Menkenina berryi Church, new species.
(Plate 3, figures 9 a, b.)

Test free, calcareous, perforate, lanceolate, uniserial, compressed from front
to back in contrast to Vaginulina, generally flattened on the dorsal side gently
rounded on the ventral side, sutures curved upward on the ventral side, gently
depressed in last and next to last chambers, flush in earlier ones, chambers lunate
on ventral side, chevron-shaped on dorsal side with sutures merging with the
trace of the tubular neck of the previous chamber, apertural end drawn out
into a short, tapering neck at the extreme edge of the dorsal side, aperture
rounded and simple at the end of the tubular neck which is, at least in part,
retained in each added chamber and forms what appears to be a continuous tube
near the surface, a thin transparent carina extends along each side of the test
from the base of the final chamber to the initial end, usually broken and
irregular, in the megalospheric form the initial chamber is large and globular
followed by six lunate, alightly inflated chambers which increase regularly with
growth, the initial chamber of the microspheric form is also spherical but much
smaller resulting in a more sharply pointed initial end, several strong, broken
to continuous costae extend longitudinally over both sides of the test from the
penultimate chamber to the initial end, becoming more prominent and carinate
toward the initial end but absent from the final, more inflated chamber, test
straight but with a suggestion of a gentle upward curve at the initial end on
the ventral side. Length 1.17 mm., width .37 mm.

HoLotypeE no. 12977 (California Academy of Sciences, Department of
Geology Type Collection) from Locality 40091 (CAS), from a deep north-south
trending V-shaped gully beginning 500 feet south and 700 feet west of the east
quarter corner of Section 20, Township 25 south, Range 18 east. Mount Diablo
base and meridian, Devils Den area, Kern County, California. Sample no. 6,
east bank of gully near base, approximately 200 feet north of the mouth of the
gully; C. C. Church collector; early Cretaceous.

Remarks. This species was found at stations 1, 2, 5, 6, of Localities 40091
(CAS) and no. 49 (F. A. Menken), Devils Den, also at Locality 27605 (CAS),
south slope of Hex Hill, Kern County, California. It does not occur in abun-
dance in any of these localities.

The specific name is in honor of Mr. Keith D. Berry of the Standard Oil
Company, Oildale, California, author of new stage names for the early Creta-
ceous of California.

1 Geologist and later Vice President in charge of Exploration for Tidewater Oil Company. Died in San
Francisco, California, May 4, 1965.

CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

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Plate 5

All specimens illustrated on this plate are in the California Academy of Sciences, Depart-
ment of Geology Type Collection.

Ficure 1. Vaginulina recta Reuss. Length .72 mm.; side view. Hypotype no. 12993
(CAS). From Locality 40091 (CAS), sample 6, deep north-south gully, east side of Devils
Den, NW. Kern County, California. Page 545.

Ficure 2. Vaginulina octocostata Church, new species. Length .78 mm., width .17 mm.;
side view. Hypotype no. 12994 (CAS). From Locality 40091 (CAS), sample 8, deep north-
south gully, east side of Devils Den, sample farthest north from mouth, NW. Kern County,
California. Page 546.

Ficure 3. Vaginulina recta Reuss. Length .57 mm.; side view. Hypotype no. 12995
(CAS). From Locality 40095 (CAS), sample 7, Hex Creek, NW. edge of Hex Hill, NW.
Kern County, California. Page 545.

Ficure 4. Nodosaria elegantia Lalicker. Length .55 mm.; side view. Hypotype no.
12996 (CAS). From Locality 40091 (CAS), sample 6, deep north-south gully, east side of
Devils Den, NW. Kern County, California. Page 541.

Ficure 5. Nodosaria elegantia Lalicker. Length .80 mm.; side view. Hypotype no.
12997 (CAS). From Locality 40095 (CAS), sample 7, Hex Creek, NW. edge of Hex Hill,
NW. Kern County, California. Page 541.

Ficure 6. Frondicularia species. Length .73 mm.; side view. Hypotype no. 12998 (CAS).
From Locality 40091 (CAS), sample 6, deep north-south gully east side of Devils Den, NW.
Kern County, California. Page 559.

Ficure 7. Lenticulina (Lenticulina) miinsteri (Roemer). Length 1.34 mm.; (a) side
view; (b) apertural view. Hypotype no. 12999 (CAS). From Locality 40095 (CAS), sample
7, Hex Creek, NW. edge of Hex Hill, NW. Kern County, California. Page 552.

Ficure 8. Quadratina strombecki (Reuss). Length .51 mm.; side view. Hypotype no.
1300 (CAS). From Locality 40095 (CAS), sample 7, Hex Creek, NW. edge of Hex Hill.
NW. Kern County, California. Page 563.

Ficure 9. Tristix acutangulum (Reuss). Length .78 mm.; side view. Hypotype no.
13001 (CAS). From Locality 40095 (CAS), sample 7, Hex Creek, NW. edge of Hex Hill,
NW. Kern County, California. Page 563.

Ficure 10. Frondicularia species 2, Bartenstein, Bettenstaedt, and Bolli. Length 1.22
mm.; side view. Hypotype no. 13002 (CAS). From Locality 40095 (CAS), sample 7, Hex
Creek, NW. edge of Hex Hill, NW. Kern County, California. Page 559.

Ficure 11. Lenticulina saxonica saxonica Bartenstein and Brand. Length 1.29 mm.;
(a) side view; (b) apertural view. Hypotype no. 13003 (CAS). From Locality 40095
(CAS), sample 7, Hex Creek, NW. edge of Hex Hill, NW. Kern County, California. Page 552.

Vol. XXXII CHURCH: CRETACEOUS FORAMINIFERA Plate 5

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VoL. XXXII] CHURCH: CRETACEOUS FORAMINIFERA 563

Family GLANDULINIDAE Reuss, 1860
Subfamily GLANDULININAE
Genus Quadratina Ten Dam, 1946

Quadratina strombecki (Reuss).

(Plate 5, figure 8.)

Quadratina strombecki (REusSS), 1863, Hils u. Gault, p. 56, pl. 5, fig. 3. BARTENSTEIN and
Branp, 1951, Abh. senckenb. naturf. Ges., no. 485, p. 314, pl. 10, figs. 264 a, b, c.
Only a few of this species were found and at one station, no. 7 of Hex Creek.

Bartenstein and Brand reported it from the upper Valanginian of northwest

Germany. Loeblich and Tappan believe this form to be a quadrate varient of

the genus T77istix which it may be as both were found at station no. 7 of Hex

Creek and except for the difference in the number of sides, are very similar in

size and in being composed of clear shell material. Length .51 mm.

Genus Tristix Macfadyen, 1941

Tristix acutangulum (Reuss).
(Plate 5, figure 9.)
Tristix acutangulum (Reuss), TEN DAM, 1948, Jour. Paleo., vol. 22, no. 2, p. 181, pl. 32, figs.

9, 10.

This is one of the rarer species in this fauna but the few found are very
well preserved. The angles are sharply carinate with thin transparent flanges
extending the length of the test. The sutures are well incised and the chambers
moderately inflated. The more mature specimens have six chambers with the
greatest width at the fourth chamber. The largest specimens are about 1 mm.
in length and .3 mm. in width. The species was found only at station no. 7
of Hex Creek. Length .78 mm.

Family POLYMORPHINIDAE d’Orbigny, 1839
Subfamily POLYMORPHININAE d’Orbigny, 1839
Genus Guttulina d’Orbigny in de la Sagra, 1839

Guttulina species.
(Plate 7, figures 3 a, b.)

Individuals representative of this family are few in number in the Hex forma-
tion and as a rule quite small. Some of the smaller, more rounded species are
so similar to the small, Nodosaria-Glandulina-like species that they are difficult
to segregate with any assurance. A few specimens were found at station no. 5
of Devils Den and a few at no. 7 of Hex Creek.

564 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER.

Plate 6

All specimens illustrated on this plate are in the California Academy of Sciences, Depart-
ment of Geology Type Collection.

Ficure 1. Frondicularia frankei Cushman. Length 3.19 mm.; side view. Hypotype no.
13004 (CAS). From Locality 40091 (CAS), sample 5, deep north-south gully, east side of
Devils Den, NW. Kern County, California. Page 557.

Ficure 2. Dentalina species. Length 1.90 mm.; side view. Hypotype no. 13005 (CAS).
From Locality 40091 (CAS), sample 6, deep north-south gully, east side of Devils Den, NW.
Kern County, California. Page 545.

Ficure 3. Nodosaria cf. N. tenuicosta Reuss. Length 1.15 mm.; side view. Hypotype no.
13006 (CAS). From Locality 40091 (CAS), sample 5, deep north-south gully, east side of
Devils Den, NW. Kern County, California. Page 543.

Ficure 4. Vaginulina debilis (Berthelin). Length 1.64 mm.; side view. Hypotype no.
13007 (CAS). From Locality 40091 (CAS), sample 8, deep north-south gully, east side of
Devils Den, sample farthest north from mouth, NW. Kern County, California. Page 546.

Ficure 5. Nodosaria cf. N. tenuicosta Reuss. Length .92 mm.; side view. Hypotype no.
13008 (CAS). From Locality 40091 (CAS), sample 8, deep north-south gully, east side of
Devils Den, sample farthest north from mouth, NW. Kern County, California. Page 543.

FicureE 6. Vaginulina octocostata Church, new species. Length .86 mm.; side view.
Hypotype no. 13009 (CAS). From Locality 40091 (CAS), sample 8, deep north-south gully,
east side of Devils Den, sample farthest north from mouth, NW. Kern County, California.
Page 546.

Ficure 7. Lingulina californiensis Trujillo. Length .45 mm.; side view. Hypotype no.
13010 (CAS). From Locality 40081 (CAS), and 27501, 47 (FAM). East-west gully 1% mile
east of Devils Den, NW. Kern County, California. Page 560.

Ficure 8. Marginulina sigali Bartenstein, Bettenstaedt and Bolli. Length .61 mm.;
side view. Hypotype no. 13011 (CAS). From Locality 40091 (CAS), sample 5, deep north-
south gully east side of Devils Den, NW. Kern County, California. Page 549.

Ficure 9. Marginulinopsis gracillissima Reuss. Length 49 mm.; side view. Hypotype
no. 13021 (CAS). From Locality 40091 (CAS), sample 6, deep north-south gully, east side
of Devils Den, NW. Kern County, California. Page 551.

Ficure 10. Frondicularia franket Cushman. Side view. Hypotype (CAS). From
Locality 40091 (CAS), sample 5, deep north-south gully, east side of Devils Den, NW.
Kern County, California. Page 557.

Ficure 11. Lingulina species. Length .52 mm.; side view. Hypotype no. 13013 (CAS).
From Locality 40091 (CAS), sample 6, deep north-south gully, east side of Devils Den,
NW. Kern County, California.

Ficure 12. Marginulina pyramidalis Koch. Length 1.09 mm.; side view. Hypotype no.
13014 (CAS). From Locality 40091 (CAS), sample 5, deep north-south gully, east side of
Devils Den, NW. Kern County, California. Page 549.

Ficure 13. Nodosaria hexensis Church, new species. Length 1.01 mm.; width .24 mm.;
side view. Holotype no. 13015 (CAS). From Locality 40091 (CAS), sample 5, deep north-
south gully, east side of Devils Den, NW. Kern County, California. Page 544.

Ficure 14. Vaginulina striolata (Reuss). Length 1.07 mm.; side view. Hypotype no.
13016 (CAS). From Locality 40091 (CAS), sample 5, deep north-south gully, east side of
Devils Den, NW. Kern County, California. Page 547.

Vol. XXXII CHURCH: CRETACEOUS FORAMINIFERA Plate 6

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VoL. XXXII] CHURCH: CRETACEOUS FORAMINIFERA 56

Genus Globulina d’Orbigny in de la Sagra, 1839

Globulina cf. G. ampulla (Jones).

(Plate 8, figure 4.)

Polymorphina ampulla Jones, 1852, Quart. Jour. Geol. Soc., vol. 8, p. 267, pl. 16, fig. 14.

Polymorphina lactea PLUMMER, 1927, Bull. 2644, Univ. Texas, p. 121, pl. 6, figs. 7 a—c.

Globulina ampulla (Jones), CUSHMAN and OzAwa, 1930, Proc. U. S. Nat. Mus., vol. 77, art.
6, no. 2829, p. 79, pl. 19, figs. 9 a-c.

There is considerable variation in these lower Cretaceous species. Most
specimens are well rounded and globular but some compressed. The lower
initial end is distinctly pointed in some but only suggested in most of them.
It is a fairly common species at station no. 5 of Devils Den. Length .48 mm.

Globulina prisca Reuss.

Globulina prisca REUSS, 1862 (1863), Stiz. Akad. Wiss. Wien, vol. 46, pt. 1, p. 79, pl. 9,
fig. 8. CUSHMAN and Ozawa, 1930, Proc. U. S. Nat. Mus., vol. 77, art. 6, no. 2829, p. 73,
pl. 12, figs. 6 a—c.

This is not a common species here and was found only at station no. 5 of
Devils Den.

Genus Pseudopolymorphina Cushman and Ozawa, 1928

Pseudopolymorphina cf. P. leopolitana (Reuss).

Polymorphina leopolitana Reuss, 1851, Haidinger’s Nat. Abhandl., vol. 4, p. 28, pl. 4, fig. 11.
Pseudopolymorphina leopolitana (Reuss), CUSHMAN and Ozawa, 1930, Proc. U. S. Nat. Mus.,
vol. 77, no. 2829, art. 6, p. 108, pl. 28, figs. 4 a—c.

This is an extremely rare species as only one was found at station no. 7 of
Hex Creek.

Genus Pyrulina d’Orbigny, 1826
Pyrulina species.
(Plate 7, figure 5.)

A single, well preserved specimen of the genus was found at station no. 5 of
Devils Den. Length 1.16 mm.

Subfamily RAMULININAE, Brady, 1884
Genus Ramulina Jones in Wright, 1875

Ramulina spandeli Paalzow.

Ramulina spandeli Paatzow, 1917, Schwammergel, p. 46, pl. 47, fig. 15. BARTENSTEIN, BET-
TENSTAEDT, and Bortt, 1957, Eclog. Geol. Helv. vol. 50, no. 1, p. 42, pl. 5, fig. 106.

This is not an uncommon species at Devils Den stations nos. 1, 2, and 5 but in
most of its occurrences it is found in fragments.

566 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Family NONIONIDAE Schultze, 1854
Subfamily CHILOSTOMELLINAE

Genus Quadrimorphina Finlay, 1939

Quadrimorphina cf. Q. ruckeri (Tappan).

(Plate 8, figures 1 a, b, c; 2 4a, b.)

Pallaimorphina ruckeri TAPPAN, 1957, U. S. Nat. Mus., Bull., 215, p. 221, pl. 71, figs. 1-9.
This very small species is very similar to Q. albertensis Mellon and Wall

but the chambers are much less inflated. It was found rather commonly at
stations 4 and 5 of Devils Den but not at Hex Hill. Length .18 mm.

Family ALABAMANIDAE Hofker, 1951
Genus Gyroidina d’Orbigny, 1826

Gyroidina globosa (Hagenow) Cushman.
(Plate 8, figures 3 a, b, c.)

Nonionina globosa HAGENOw, 1842, Neues Jarb., p. 574.

Rotalia globosa Reuss, 1861, Akad. Wiss. Wien, Math. naturwiss. Kl. Sitzungsber., vol. 44,
joie Hs jos SOL fol 7, ES, Bel, 10).

Gyroidina globosa CUSHMAN, 1931, Jour. Paleo., vol. 5, no. 4, p. 310, pl. 35, figs. 19 a—c.

This long-ranging species is one of the commoner forms at stations nos. 7 and
8 of Hex Creek and no. 15 of Hex Hill. Length .40 mm.

Superfamily CAssmpULINACEA d’Orbigny, 1839
Family ANOMALINIDAE
Subfamily ANOMALININAE

Genus Gavelinella Brotzen, 1942

Gavelinella barremiana Bettenstaedt.
(Plate 8, figures 6 a, b, c.)

Gavelinella barremiana BETTENSTAEDT, 1952, Senckenbergiana, vol. 33, no. 4/6, p. 275, pl. 2,
figs. 26-29. BARTENSTEIN, BETTENSTAEDT, and BOLI, 1957, Eclog. Geol. Helv., vol. 50,
no. 1, p. 47, pl. 7, figs. 168 a-c; 169 a-—c.

This is one of the common species in samples 7 and 8 of Hex Creek. It was
also found at no. 15 of Hex Hill. The species was described from the middle
Barremian of Germany by Bettenstaedt but ranges somewhat higher. It has
also been described from the lower Cretaceous of Trinidad, British West Indies.
Length .47 mm.

VoL. XXXII] CHURCH: CRETACEOUS FORAMINIFERA 567

Family CrBici1pAE Cushman, 1927
Subfamily PLANULININAE Bermudez, 1952
Genus Planulina d’Orbigny, 1826

Planulina andersoni Church, new species.
(Plate 8, figures 7 a, b, c.)

Test trochoid, biconvex, dorsal side gently convex, ventral side compressed
but with a slight convexity and central depression, periphery subacute, partially
evolute on dorsal side, involute or nearly so on the ventral side, periphery only
faintly lobulate, chambers distinct throughout, 12 in final whorl, uninflated
and increasing gradually and uniformly as added, sutures broad, strongly re-
curved and prominent because of lighter color and elevation above the chamber
surface near the outer curve of test, sutures tend to merge at outer edge to form
a smooth edged keel, wall perforate, smooth, aperture a low arch with narrow
bordering lip at the peripheral margin, extending a short distance into the
umbilical area on the ventral side. Length .42 mm., width .33 mm.

HoLotyre no. 13033 (California Academy of Sciences, Department of
Geology Type Collection), from Locality 40091 (CAS), from deep north-south
trending V-shaped gully beginning 500 feet south and 700 feet west of the east
quarter corner of Section 20, Township 25 south, Range 18 east, Mount Diablo
base and meridian, Devils Den area, Kern County, California. Sample no. 5,
east bank of gully near base, approximately 150 feet north of the mouth of the
gully; C. C. Church collector; early Cretaceous.

REMARKS. This species is fairly common where it occurs at the Devils Den
locality at stations nos. 1 to 5 but was not found at Hex Hill. It is somewhat
similar to Anomalina popenoei Trujillo from the upper Cretaceous of the Sacra-
mento Valley, California and also bears a general resemblance to Planulina
spissocostata Cushman from the Taylor marl of Texas, U. S. A. From the
former it differs in having more chambers, a smoother periphery and being
less sharply keeled and from the latter in having fewer chambers per whorl,
a more rounded keel and no buildup of calcite near the umbilicus. Named in
honor of the late Dr. F. M. Anderson.

Superfamily ORBITOIDACEA Schwager, 1876
Family EponipipaE Hofker, 1951
Genus Eponides de Montfort, 1808

Eponides species.

This is one of the rare species in the Devils Den assemblage and the only
representative of the genus found in this lower Cretaceous fauna. It is of mod-
erate size, trochospiral, about equally biconvex, four slightly lobed chambers
visible on the ventral side, edge smoothly rounded. Found only at stations nos.
4 and 5 of the Devils Den series.

568 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SEr.

Plate 7

All specimens illustrated on this plate are in the California Academy of Sciences, Depart-
ment of Geology Type Collection.

Ficure 1. Praebulimina species. Length 32 mm.; (a) side view; (b) end view.
Hypotype no. 13017 (CAS). From Locality 40091 (CAS), sample 4, deep north-south
gully, east side of Devils Den, NW. Kern County, California. Page 570.

Ficure 2. Globanomalina hexensis Church, new species. Length .28 mm.; width .22 mm.;
(a) side view; (b) apertural view. Holotype no. 13018 (CAS). From Locality 40095 (CAS),
sample 7, Hex Creek, NW. edge of Hex Hill, NW. Kern County, California. Page 571.

Ficure 3. Guttulina species. Length .71 mm.; (a) side view; (b) apertural view.
Hypotype no. 13019 (CAS). From Locality 40095 (CAS), sample 7, Hex Creek, NW. edge
of Hex Hill, NW. Kern County, California. Page 563.

Ficure 4. Hedbergella planispira (Tappan). Length .21 mm.; (a) ventral view: (b)
dorsal view; (c) apertural view. Hypotype no. 13020 (CAS). From Locality 40095 (CAS),
sample 7, Hex Creek, NW. edge of Hex Hill, NW. Kern County, California. Page 571.

Ficure 5. Pyrulina species. Length 1.16 mm.; side view. Hypotype no. 13021 (CAS).
From Locality 40091 (CAS), sample 5, deep north-south gully, east side of Devils Den,
NW. Kern County, California. Page 565.

Ficure 6. Involutina stinemeyeri Church, new species. Length 44 mm.; width 41
mm.; side view. Holotype no. 13022 (CAS). From Locality 40091 (CAS), sample 6, deep
north-south gully, east side of Devils Den, NW. Kern County, California. Page 565.

Ficure 7. Nodosaria humilis Roemer. Length .69 mm.; side view. Hypotype no. 13023
(CAS). From Locality 40091 (CAS), sample 5, deep north-south gully, east side of Devils
Den, NW. Kern County, California. Page 541.

Ficure 8. Gubkinella californica Church, new species. Length .19 mm., width .15 mm.;
side view. Holotype no. 13024 (CAS). From Locality 40091 (CAS), sample 6, deep north-
south gully, east side of Devils Den, NW. Kern County, California. Page 573.

FicureE 9. Ammodiscus glabratus Cushman and Jarvis. Length .61 mm.; side view.
Hypotype no. 13025 (CAS). From Locality 40092 (CAS), west side of most easterly deep
gully of Hex Hill, NW. Kern County, California. Page 533.

Ficure 10. Darbyella species. Length .69 mm.; (a) dorsal view; (b) ventral view; (c)
apertural view. Hypotype no. 13026 (CAS). From Locality 40091 (CAS), sample 5, deep
north-south gully, east side of Devils Den, NW. Kern County, California. Page 555.

Vol. XXXII CHURCH: CRETACEOUS FORAMINIFERA Plate 7

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VoL. XXXII] CHURCH: CRETACEOUS FORAMINIFERA 569

Superfamily ROBERTINACEA Reuss, 1850
Family CERATOBULIMINIDAE
Subfamily EPISTOMININAE Wedekind, 1937
Genus Epistomina Terquem, 1883

Epistomina cf. E. caracolla caracolla (Roemer).

(Plate 8, figure 5.)

Gyroidina caracolla RoEMER, 1841, Kreidegeb., p. 97, pl. 15, fig. 22.
Epistomina djaffaensis StGAL, 1952, Crétacé, p. 14, fig. 7.

Epistomina (Hoglundina) caracolla caracolla (Roemer, 1841), S1GAL, 1957, Eclog. Geol. Helv.,
vol. 50, no. 1, p. 46, pl. 5, figs. 113, 114; pl. 6, fig. 142.

In the state of preservation of this species it is difficult to determine internal
structure and apertural characters but because of the outer similarity to the
species E. caracolla caracolla, it is compared to that species. It is one of the
common species of the Devils Den area where it was found at stations nos. 3 to
6. There is quite a range in the size but the largest ones measure up to one
millimeter in maximum width. Length .62 mm.

Superfamily CAssmwuLinacea d’Orbigny, 1839
Family INVoLUTINIDAE Biitschli, 1880
Genus Involutina Terquem, 1862

Involutina stinemeyeri Church, new species.
(Plate 7, figure 6.)

Test free, calcareous, perforate, lenticular, consisting of a proloculus followed
by a nonseptate, planispirally, coiled tubular second chamber, sides of equal
thickness and curvatures as a result of the growth of secondary calcite over
the primary coil which is slightly thicker over central area and becoming thinner
near the edge, surface deeply pitted from the termination of small, calcite pillars
which are part of the secondary deposit, the overgrowth covering all coils except
the last which may be partially covered, diameter of the tube increases very
gradually, most specimens having six coils, the tube being lunate in cross
section. Length .44 mm. Width .41 mm.

HoLotyPe no. 13022 (California Academy of Sciences, Department of
Geology Type Collection), from Locality 40091 (CAS), from deep north-south
trending V-shaped gully beginning 500 feet south and 700 feet west of the east
quarter corner of Section 20, Township 25 south, Range 18 east, Mount Diablo
base and meridian, Devils Den area, Kern County, California. Sample no. 6,
east bank of gully near base, approximately 150 feet north of the mouth of the
gully; C. C. Church collector; early Cretaceous.

REMARKS. The earlier described species of /nvolutina were largely from the
upper Triassic and lower Jurassic (Lias). I have found only one listed from
California and it was from the upper Cretaceous Panoche formation of Fresno

570 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

County. It was described by Lewis Martin. This species is finely arenaceous
and from the more recent analysis of the history of the genus by Loeblich and
Tappan (1961), it would have to be classified as an Ammodiscus. Our species
was found rather common at Devils Den Localities nos. 2 and 6. It is a very
distinctive species and should prove to be a good marker fossil.

Superfamily BULIMINACEA Jones, 1875
Family TuRRILINIDAE Cushman, 1927
Subfamily TURRILININAE Cushman, 1927
Genus Praebulimina Hofker, 1953

Praebulimina species.
(Plate 7, figures 1 a, b.)

The generic assignment of this species is very much in question. As in
Bulimina and Praebulimina, it is high spired and triserially arranged, but as the
test has been filled with calcite the aperture is not clearly defined. The apertural
face is flattened and near the top edge of the final chamber there is a narrow,
slit-like channel which follows the curve of the apertural face over a third to
one half the distance from the top to its base. This may be the aperture but it
could be a growth feature. The only species which I have seen figured which
has an aperture somewhat similar to it is Praebulimina seabeensis Tappan from
the Seabee formation of the upper Cretaceous of Alaska. The aperture of this
species is slit-like but lower down on the face. On the present species there is a
darker line which seems to be wider than a suture at the base of the apertural
face. It is possible that this feature may have been the aperture rather than the
upper facial channel. This species was found only at station no. 4 at Devils
Den. Length .32 mm.

Superfamily ROBERTINACEA Reuss, 1850
Family CERATOBULIMINIDAE Cushman, 1921
Subfamily CERATOBULIMININAE Cushman, 1927
Genus Conorboides Hofker in Thalmann, 1952

Conorboides species.

A number of what appears to be this genus were found at sample Locality
no. 4 of Devils Den. In its chamber arrangement and shape it is very similar
to Conorboides umiatensis (Tappan) from the Grandstand formation of northern
Alaska of Albian age. The Devils Den species differs in having a much higher
spire, a somewhat flatter and more deeply cleft ventral side. The apertural
characters are concealed by a secondary extraneous deposit and the complete
crystallization of the test. Length .18 mm.

VoL. XXXII] CHURCH: CRETACEOUS FORAMINIFERA 571

Superfamily GLOBIGERINACEA Carpenter, Parker and Jones, 1862
Family HANTKENINIDAE Cushman

Genus Globanomalina Haque, 1956

Globanomalina hexensis Church, new species.
(Plate 7, figures 2 a, b.)

Test minute, free, calcareous, planispiral, partially involute, biumbilicate,
chambers globose to broadly oval, smooth, finely perforate, sutures depressed,
radial, chambers enlarge gradually and evenly with growth, seven in final whorl,
aperture a low equatorial arch at base of final chamber with narrow lip which
ends abruptly at side of previous whorl with no lateral openings. Diameter .35
mm; thickness of final chamber .15 mm. Found only in the Hex Hill area at
station no. 7 where it was fairly common and at station no. 15 where it was
quite rare. Length .28 mm., width .22 mm.

HoLotype no. 13018 (California Academy of Sciences, Department of Geol-
ogy Type Collection), from Locality 40095 (CAS), sample no. 7, from the
lower slope of a south-facing small ridge projecting into Hex Creek from the
west causing it to veer to the northeast, about 1150 feet south and 200 feet
east of the north quarter corner of Section 31, Township 25 south, Range 18
east, Mount Diablo base and meridian, Hex Hill area, Kern County, California;
C. C. Church collector; early Cretaceous.

Family RoTALIPoRIDAE Sigal, 1958
Subfamily HEDBERGELLINAE Loeblich and Tappan, 1961

Genus Hedbergella Bronniman and Brown, 1958

Hedbergella planispira (Tappan).
(Plate 7, figures 4 a, b, c.)

(SS)

Globigerina planispira TAPPAN, 1943, Jour. Paleo., vol. 17, no. 5, p. 513, pl. 83, fig. 3.
Praeglobotruncana planispira (Tappan), Botti, LorBLicu, and TAppaAn, 1957, U. S. Nat.
Mus., Bull. 215, p. 40, pl. 9, fig. 3.

Hedbergella planispira (Tappan), LorsricH and Tappan, 1961, Micropaleontology, vol. 7,

no. 3, p. 276, pl. 5, figs. 4-11. NeraGu, 1965, Micropaleontology, vol. 11, no. 1, p. 36,

pl. 10, figs. 1-4.

This very small species was found at station no. 7 of Hex Creek in some
abundance and at station no. 8 in fewer numbers. At Station no. 15 of Hex
Hill they were present but rare. The species was also found at no. 4 of Devils
Den but was not of common occurrence. This species is also very similar to
Globigerina infracretacea Glaessner. Length .21 mm.

CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER.

Ul
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bo

Plate 8

All specimens illustrated on this plate are in the California Academy of Sciences, Depart-
ment of Geology Type Collection.

Figure 1. Quadrimorphina cf. Q. ruckeri Tappan. Length .18 mm.; (a) dorsal view;
(b) ventral view; (c) peripheral view. Hypotype no. 13027 (CAS). From Locality 40091
(CAS), sample 5, deep north-south gully, east side of Devils Den, NW. Kern County,
California. Page 566.

Ficure 2. Quadrimorphina cf. Q. ruckeri Tappan. Length .18 mm; (a) dorsal view;
(b) ventral view. Hypotype no. 13028 (CAS). From Locality 40091 (CAS), sample 4,
deep north-south gully east side of Devils Den, NW. Kern County, California. Page 566.

Ficure 3. Gyroidina globosa (Hagenow). Length .40 mm.; (a) dorsal view; (b) ventral
view; (c) peripheral view. Hypotype no. 13029 (CAS). From Locality 40095 (CAS), sam-
ple 7, Hex Creek, NW. edge of Hex Hill, NW. Kern County, California. Page 566.

Ficure 4. Globulina cf. G. ampulla (Jones). Length 48 mm.; side view. Hypotype no.
13030 (CAS). From Locality 40091 (CAS), sample 8, deep north-south gully, east side of
Devils Den, mostly northerly sample from mouth, NW. Kern County, California. Page 565.

Ficure 5. Epistomina cf. E. caracolla caracolla (Roemer). Length .62 mm.; dorsal view.
Hypotype no. 13031 (CAS). From Locality 40091 (CAS), sample 5, deep north-south gully,
east side of Devils Den, NW. Kern County, California. Page 569.

Ficure 6. Gavelinella barremiana Bettenstaedt. Length 47 mm.; (a) dorsal view; (b)
ventral view; (c) peripheral view. Hypotype no. 13032 (CAS). From Locality 40095
(CAS), sample 7, Hex Creek, NW. edge of Hex Hill, NW. Kern County, California. Page 566.

Ficure 7. Planulina andersoni Church, new species. Length 42 mm., width 33 mm.;
(a) dorsal view; (b) ventral view; (c) peripheral view. Holotype no. 13033 (CAS). From
Locality 40091 (CAS), sample 5, deep north-south gully, east side of Devils Den, NW.
Kern County, California. Page 567.

Vol. XXXII CHURCH: CRETACEOUS FORAMINIFERA Plate 8

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aw

Family HETEROHELICIDAE Cushman, 1927
Subfamily GUEMBELITRIINAE Montanaro Gallitelli, 1957
Genus Gubkinella Suleymanov, 1955

Gubkinella californica Church, new species.
(Plate 7, figure 8.)

This species is somewhat similar to G. asiatica Suleyma (1955) except’ that
in the final four chambers of the present species, the chamber size increases
very slightly from the first to the fourth of the last four chambers while the
change from the first to the second whorl is so great that in some specimens the
first whorl appears to be almost a separate individual attached to the four
globular chambers. Gubkinella californica also resembles Giimbelitria harrisi
Tappan from the Grayson formation of northern Texas, but is shorter and the
globular chambers more compact; it also has four chambers to a whorl.

Gubkinella californica Church is very small, having a height of .15 mm. to
.2 mm. and a width of .2 mm. In a few specimens the width may be greater
than the height, four chambers to a whorl with three whorls, the aperture is a
low arch at the base of the last-formed chamber, test calcareous, finely perforate,
trochospiral; the inflated chambers suggest that the species was pelagic. It was
found only at Devils Den where it was of quite common occurrence at stations
nos. 4 and 6 but since the two samples were collected at different times and
the first not located by a marker, they may be from the same spot. Length .19
mm. Width .15 mm.

HoLotypPe no. 13024 (California Academy of Sciences, Department of
Geology Type Collection), from Locality 40091 (CAS), from deep north-south
trending V-shaped gully beginning 500 feet south and 700 feet west of the east
quarter corner of Section 20, Township 25 south, Range 18 east, Mount Diablo
base and meridian, Devils Den area, Kern County, California, sample no. 6,
east bank of gully near base, approximately 200 feet north of the mouth of the
gully; C. C. Church collector; early Cretaceous.

SELECTED BIBLIOGRAPHY

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574 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

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VoL. XXXII] CHURCH: CRETACEOUS FORAMINIFERA 57

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1.
~I
oO

CALIFORNIA ACADEMY OF SCIENCES [ Proc. 47TH SER.

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VoL. XXXII] CHURCH: CRETACEOUS FORAMINIFERA 577

GRAHAM, J. J.

1961. An annotated bibliography of California Cretaceous microfossils. Special Report

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1963, Campanian Foraminifera from the Stanford University campus, California.
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HEcnutT, FrAnz E.

1938. Standard-Gliederung der Nordwest-deutschen Unterkreide nach Foraminiferen.
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IMLAY, RALPH W.

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1964. Illustrations of Canadian fossils. Early lower Cretaceous (Berriasian and Valan-
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Kuan, Mousin H.

1962. Lower Cretaceous index Foraminifera from northwestern Germany and England.

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1955. Upper Cretaceous Foraminifera from the “Franciscan Series,’ New Almaden
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LALICKER, C. J.

1950. Foraminifera of the Ellis group, Jurassic at the type locality. University of

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LoesiicyH, A. R., and HELEN TAppAN

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1950a. Foraminifera of the type Kiowa shale, lower Cretaceous of Kansas. University of
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578 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

LOETTERLE, GERALD J.

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VoL. XXXII] CHURCH: CRETACEOUS FORAMINIFERA 579

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1965. Upper Cretaceous planktonic Foraminifera from the Putah Creek subsurface
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PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES
FOURTH SERIES

G Dallas Hanna Anniversary Volume

Vol. XXXII, No. 19, pp. 581-586 June 27, 1968

THE PLANT GENUS POLYGALA IN THE
GALAPAGOS ISLANDS

John Thomas Howell

Curator of Botany, California Academy of Sciences

and

Duncan M. Porter

Assistant Professor of Biology, University of San Francisco

The five taxa of Polygala Linnaeus on the Galapagos Islands are endemics
and all are similar morphologically. They perhaps represent a radiation from
a single past introduction from mainland South America, similar to those radia-
tions postulated for the justly famous Darwin’s finches and the plant genus
Scalesia. They are members of Polygala section Polygala series Tenues Chodat,
and appear to be most closely related to P. paludosa St. Hilaire or P. paniculata
Linnaeus, two widespread species of the New World tropics.

Both authors have had field experience in the Galapagos Islands. Howell
was botanist on the Templeton Crocker Galapagos Expedition of the California
Academy of Sciences from April to June, 1932, and later was able to examine
type material of the taxa in question in several European herbaria. Porter visited
the archipelago in January and February, 1967, while assisting Dr. Ira L.
Wiggins in a study of its flora under a National Science Foundation grant to
the California Academy of Sciences. Both authors wish to express their gratitude
to Dr. Wiggins and to the California Academy of Sciences for making this
study possible.

[581 ]

582 CALIFORNIA ACADEMY OF SCIENCES [Pcoc. 4TH SER.

KrEY TO THE SPECIES AND VARIETIES OF POLYGALA IN THE GALAPAGOS ISLANDS

1. Stems puberulent; wings 5—7-nerved; aril 4 as long as seed —_____ 1. P. anderssonii
1. Stems glabrous; wings 3-
2. Apical crest of keel inconspicuous, not showy or petaloid, divisions simple or

2-lobed, final lobes 6--8; wings 3.5-5 mm. long, 3-nerved.
3. Leaves linear to oblanceolate-elliptic, acute to cuspidate; racemes becoming
elongate, acute, 1.5-12 cm. long; wings 3.5-4 (or 5) mm. long —_____
ON es ee EN Nn 2a. P. galapageia var. galapageia

5-
5-nerved; aril about 15 as long as seed.

3. Leaves Sia mecains to linear-lanceolate, obtuse to acute; racemes more
congested, oblong or rounded, obtuse to subacute, 1-4 cm. long; wings
4-5 mm. long ____________.__..._........_. 2b. P. galapageia var. msularis
2. Apical crest of eile conspicuous, Nome and petaloid, divisions frequently 3-
lobed, final lobes 12-16; wings 4-6.5 mm. long, 5-nerved.
4. Leaves broadly spathulate to rotundate, 6-13 mm. long, 3-8 mm. wide
_ 3a. P. sancti-georgi var. sancti-georgii

4. Tees eS to ieee Vtshecblate, 8-25 mm. long, 1-4 mm. wide
3b. P. sancti-georgii var. oblanceolata

1. Polygala anderssonii Robinson.

Polygala puberula ANDERSSON, Kongliga Vetenskaps-Akademiens Handlingar, 1853, p. 232,
1855. Anpersson, Kongliga Svenska Fregatten Eugenies Resa omkring jorden . . . aren
1851-1853, Botanik, vol. II, p. 100, 1861. Roprnson AND GREENMAN, American Journal
Science, series 3, vol. 50, p. 145, 1895. Not P. puberula Gray, Plantae Wrightianae Texana
—Neo-Mexicanae part I, p. 40, 1852 (Smithsonian Contributions to Knowledge, vol. 3,
aint, 5))).

Polygala galapageia A. W. BENNETT in part as a synonym, Journal Botany, vol. 17, p. 204,
1879.

Polygala anderssonii Ropinson, Proceedings American Academy, vol. 38, p. 160, 1902.
Stewart, Proceedings California Academy Sciences, series 4, vol. 1, p. 85, 1911.
CHRISTOPHERSEN, Nyt Magazin for Naturvidenskaberne, vol. 70, p. 80, 1932.

Suffruticose (perennial?), to 1 m. high, rarely branched basally; stems
spreading, puberulent, becoming glabrate below, yellowish or rarely reddish;
leaves alternate, linear-lanceolate to oblanceolate, acuminate, puberulent to
subglabrate, coriaceous, 4-17 mm. long, 1-1.5 mm. wide; petioles less than
1 mm. long; racemes becoming elongate, 3-9 cm. long, about 1 cm. wide, axis
puberulent; flowers white to pale purple; pedicels 1 mm. long, glabrous;
bracteoles caducous; outer sepals ovate, acute, glabrous, green, margins white,
more or less equal, 1.5-2 mm. long, 1 mm. wide; wings petaloid, obovate-
elliptic, obtuse, glabrous, 5—7-nerved, 3.5-4.5 mm. long, about 2 mm. wide, longer
than capsule; corolla about 3 mm. long; apical crest of keel 12—14-lobed, less
than 1 mm. long; capsule oblong, glabrous, equally 2-loculed, loculicidally
dehiscent, 3.5-4 mm. long, about 2 mm. wide; seeds 1 per locule, obconical,
apex obtuse and more or less apiculate, base attenuate, dark, shortly lanate,
trichomes more or less appressed apically, 2.5—3 mm. long; aril elliptical, 2-lobed,
obtuse, white, ’ as long as seed.

Vor. XXXII] HOWELL & PORTER: POLYGALA IN GALAPAGOS ISLANDS 583

SPECIMENS EXAMINED. Galapagos, Cuming 103 (CGE,K). IsABeLa: Cowley
Bay, not abundant, Stewart 1775 (CAS,K). SAN SALVADOR: west of lava flow
at James Bay, Howell 9720 (flowers white, more or less tinged with lavender;
CAS). Santa Cruz: Andersson (S, type of P. puberula Andersson); north
shore, Snodgrass & Heller 668 (DS); north side, abundant in light ashy soil near
shore, Stewart 1778 (CAS); northwest side, abundant in tufaceous soil near
shore, Stewart 1779 (CAS); in more open spaces in restricted area of Croton
forest northwest of Academy Bay, 50 feet, G. Taylor 7 (CAS); south of Conway
Bay, Baur 13 (DS,K).

2. Polygala galapageia Hooker fil.

2a. Polygala galapageia var. galapageia.

Polygala galapageia Hooker fil., Transactions Linnaean Society, Botany, vol. 20, p. 233, 1847.
A. W. BENNETT, Journal Botany, vol. 17, p. 204, 1879. RoBrnson AND GREENMAN, Ameri-
can Journal Science, series 3, vol. 50, p. 145, 1895. Ropinson, Proceedings American

Academy, vol. 38, p. 160, 1902. Stewart, Proceedings California Academy Sciences,
series 4, vol. 1, p. 85, 1911.

Polygala galapageja ANDERSSON, Kongliga Vetenskaps-Akademiens Handlingar, 1853, p. 232,
1855. Anpersson, Kongliga Svenska Fregatten Eugenies Resa omkring jorden .. . aren
1851-1853, Botanik, vol. II, p. 100, pl. 10, fig. 1 (m, n, 0), 1861.

Polygala galopagensis Cuopat, Mémoires Société de Physique et d’Histoire Naturelle, vol. 31,
p. 230, 1893.

Suffrutescent perennial, to about 1 m. high, herbage more or less glaucous;
stems spreading, glabrous, reddish below, yellowish above; leaves alternate,
numerous, linear to oblanceolate-elliptic, acute to cuspidate, glabrous, coriaceous,
5-15 mm. long, 1-1.5 mm. wide; petioles less than 1 mm. long; racemes be-
coming elongate, acute, 1.5-12 cm. long, about 1 cm. wide; flowers white or
whitish; pedicels less than 1 mm. long, glabrous; bracteoles caducous; outer
sepals ovate, acute, glabrous, green, margins white, more or less equal, about
1.5 mm. long; wings petaloid, obovate-elliptic, attenuate, glabrous, 3-nerved,
3.5—4 (or 5) mm. long, 2 mm. wide, more or less equaling capsule; corolla 2—3
mm. long; apical crest of keel inconspicuous, 4-parted and each part simple or
2-lobed, less than 1 mm. long; capsule oblong, glabrous, equally 2-loculed,
loculicidally dehiscent, 3—3.5 mm. long, 1.5 mm. wide; seeds 1 per locule, ob-
conical, apex obtuse and more or less apiculate, base attenuate, dark, shortly
lanate, trichomes more or less appressed apically, 2—2.5 mm. long; aril narrowly
elliptical, 2-lobed, white, about '% as long as seed.

SPECIMENS EXAMINED. FLOREANA: Darwin (CGE, type), Edmonston (K),
Andersson (S). IsaBELA: Macrae (CGE,K); Tagus Cove, Crossland 443 (BK),
Howell 9510 (stems yellow, leaves green, flowers whitish tinged lavender, CAS),
Snodgrass & Heller 142 (DS), Snow 307, 327 (DS), abundant from beach to
600 feet, Stewart 1774 (CAS). MARcHENA: Snodgrass & Heller 763 (DS).
SAN CRISTOBAL: Snodgrass & Heller 494 (DS); lower region, southwest end,

584 CALIFORNIA ACADEMY OF SCIENCES [Pcoc. 4TH SER.

Baur 9 (K); Puerto Chico, 20 m., Schimpff 172 (CAS); Wreck Bay, bushy
plant 2-3 feet high, Howell 8609 (CAS), abundant in sandy soil near shore,
Stewart 1777 (CAS).

The type specimens of P. galapageia and P. obovata Hooker fil. [P. galapageia
var. insularis (A. W. Bennett) Robinson] in the University of Cambridge
herbarium are very different in appearance, which would account for treating
them as distinct species. This difference is due largely to the shapes of the
leaves: in var. galapageia linear-oblanceolate or linear-oblong and pungently
acute; in var. insularis broader and obtuse. Although the wings in var. galapageia
are usually shorter than in var. inmsularis, the wings in flowers of the type of
P. galapageia are 5 mm. long, the longest measured in any specimen referable to
the typical variety.

2b. Polygala galapageia var. insularis (A. W. Bennett) Robinson.

Polygala obovata Hooxer Fit., Transactions Linnaean Society, Botany, vol. 20, p. 233, 1847.
ANDERSSON, Kongliga Vetenskaps-Akademiens Handlingar, 1853, p. 231, 1855. ANDERSSON,
Svenska Fregatten Eugenies Resa omkring jorden .. . aren 1851-1853, Botanik, vol. II,
p. 99, pl. 10, fig. 2 (c-i, k-m, p), 1861. Rogprnson AND GREENMAN, American Journal
Science, series 3, vol. 50, p. 146, 1895. Not P. obovata Sr. Hrvarre, Flora Brasiliae
Meridionalis, vol. 2, p. 37, 1835.

Polygala obovata {ma. angustifolia ANDERSSON, Kongliga Vetenskaps-Akademiens Handlingar,
1853, p. 232, 1855. ANpERsSON, Kongliga Svenska Fregatten Eugenies Resa omkring
jorden .. . aren 1851-1853, Botanik, vol. II, p. 99, 1861. P. obovata var. angustifolia
(ANDERSSON) RILEY as a synonym, Kew Bulletin, 1925, p. 219, 1925.

Probably also: P. obovata fma. latifolia ANDERSSON, Kongliga Vetenskaps-Akademiens
Handlingar, 1853, p. 231, 1855. ANDERSSON, Kongliga Svenska Fregatten Eugenies Resa
omkring jorden . . . aren 1851-1853, Botanik, vol. II, p. 99, 1861. P. obovata var.
latifolia (ANDERSSON) RILEY as a synonym of P. sancti-georgii, Kew Bulletin, 1925,
De 219, 1925;

Polygala chatamensis ANDERSSON, Kongliga Vetenskaps-Akademiens Handlingar, 1853, p. 232,
1855. ANDERSSON, Kongliga Svenska Fregatten Eugenies Resa omkring jorden . . . aren
1851-1853, Botanik, vol. II, p. 99, pl. 10, fig. 3 (p, q, r), 1861. P. galapageia [var. gala-
pageia| Ropinson in part as a synonym, Proceedings American Academy, vol. 38,
p. 160, 1902.

Polygala insularis A. W. BENNETT, Journal Botany, vol. 17, p. 204, 1879. ROBINSON AND
GREENMAN, American Journal Science, series 3, vol. 50, p. 146, 1895.

Polygala galapageia var. insularis (A. W. Bennett) Ropinson, Proceedings American Academy,
vol. 38, p. 161, 1902. Stewart, Proceedings California Academy Sciences, series 4,
vol. 1, p. 86, 1911.

Differing from var. galapageia in having stems 30-45 cm. high, reddish;
leaves oblanceolate to linear-lanceolate, obtuse to acute, more or less apiculate,
6-10 mm. long, 1-2 mm. wide; racemes oblong or rounded, obtuse to subacute,
1-4 cm. long, about 1 cm. wide; wings 4-5 mm. long.

SPECIMENS EXAMINED. Galapagos, Andersson (K), Andersson 181 (K).
FLOREANA: “Charles and Chatham,” Andersson (S). MARCHENA: occasional in

Vot. XXXII] HOWELL & PORTER: POLYGALA IN GALAPAGOS ISLANDS 585

tufaceous soil near the shore, Stewart 1776 (CAS,K). PINTA: fairly abundant
on lava beds in lower part, Stewart 1773 (CAS). SAN CRISTOBAL: Darwin
(CGE, type of P. obovata Hooker fil.), Andersson (S, type of P. chatamensis
Andersson, GH, isotype), Andersson 182 (K); northern Chatham, Baur 7 (K);
Cerro Brujo, north coast, Snow 350 (DS); Sappho Cove, abundant on sand
beaches, Stewart 1782 (CAS). Santa Cruz: Academy Bay, common on lower
parts, Stewart 1783 (CAS).

Riley (Kew Bull., 1925, p. 219) was probably in error when he referred
Andersson’s broad-leaved form of P. obovata to P. sancti-georgiu. Neither at
Stockholm nor at Kew was an Andersson specimen seen that would be referable
to P. sancti-georgii. It would seem probable that Andersson was distinguishing
by name the two leaf-forms of P. galapageia which are represented by his col-
lections. These formal names, however, were not seen on any Andersson collec-
tions, a fact that adds to the difficulty of placing them definitely.

The type of P. chatamensis Andersson would appear to represent a seedling
stage of P. galapageia var. insularis, although in general aspect it looks more
like var. galapageia. However, in the type of P. chatamensis, the leaves, although
linear-oblong or linear-oblanceolate, are obtuse, the inflorescence is short-
congested and rounded above, and the wings are over 4 mm. long. These details
do not correspond to Andersson’s figure of P. chatamensis which is more like
P. galapageia var. galapageia.

3. Polygala sancti-georgii Riley.

3a. Polygala sancti-georgii var. sancti-georgii.

Polygala sancti-georgii R1iLEY, Kew Bulletin, 1925, p. 218, 1925.

Polygala obovata var. latifolia Riley as a synonym, Kew Bulletin, 1925, p. 219, 1925.
Probably not P. obovata fma. latifolia ANDERSSON, Kongliga Vetenskaps-Akademiens
Handlingar, 1853, p. 231, 1855, basonym of Riley’s name.

Herbaceous annual, 15-50 cm. high, herbage glaucous; stems spreading,
glabrous, reddish; leaves alternate, usually numerous and crowded, broadly
spathulate to rotundate or rarely narrowly spathulate, obtuse, cuspidate, gla-
brous, coriaceous, blade decurrent into petiole, 6-13 mm. long, 3-8 mm. wide;
petioles less than 1 mm. long; racemes 1.5-3.5 (—6) cm. long, 1.5 cm. wide;
flowers purplish to white; pedicels 1 mm. long, glabrous; bracteoles caducous;
outer sepals ovate, acute, glabrous, green, margins white, more or less equal,
1.5-2 mm. long, 1 mm. wide; wings petaloid, ovate-elliptic, more or less obtuse,
glabrous, 5-nerved, 6—6.5 mm. long, 2-3 mm. wide, longer than capsule; corolla
about 3 mm. long; apical crest of keel conspicuous, petaloid, 14—16-lobed, about
1 mm. long; capsule oblong, glabrous, equally 2-loculed, loculicidally dehiscent,
3.5-4 mm. long, 2 mm. wide; seeds 1 per locule, obconical, apex obtuse and
more or less apiculate, base attenuate, dark, shortly lanate, trichomes more or
less appressed apically, 2.5-3 mm. long; aril narrowly elliptical, 2-lobed, white,
about % as long as seed.

586 CALIFORNIA ACADEMY OF SCIENCES [Pcoc. 47TH SER.

SPECIMENS EXAMINED. Galapagos, Mr. Brace (K). FLOREANA: Hicks 424
(K, type); Cormorant Bay, abundant on sand beaches, Stewart 1780, 1781
(CAS), sand dunes, Wiggins & Porter 564 (CAS); Post Office Bay, beaches
and environs at east end, Wiggins & Porter 531 (CAS).

As mentioned under P. galapageia var. insularis, no specimens of P. sancti-
georgii were seen among Andersson’s collections either in Stockholm or Kew.
Hence, it is believed that his P. obovata forma Jatifolia should be referred to
P. galapageia var. insularis, not to P. sancti-georgii. The epithet, sancti-georgit,
is taken from the name of the yacht, St. George, which in 1924 visited the
Galapagos Islands on the “St. George Pacific Expedition,’ under the auspices
of the Scientific Expeditionary Research Association of England.

3b. Polygala sancti-georgii var. oblanceolata Howell.

Polygala sancti-georgii var. oblanceolata Howe.1, Leaflets of Western Botany, vol. 10,
p. 351, 1966.

Differing from var. sancti-georgii in having stems 25-60 cm. high; leaves
narrowly to broadly oblanceolate, obtuse to acute, 8-25 mm. long, 1-4 mm.
wide: racemes 2—7 (—12) cm. long, 1—1.5 cm. wide; flowers light to deep laven-
der; wings oblanceolate-elliptic, 4-6 mm. long, 2—2.5 mm. wide; corolla 3.5 mm.
long; apical crest of keel 12—14-lobed, about 1.5 mm. long, lobes broad and
pink; seeds 2.5 mm. long.

SPECIMENS EXAMINED. RABIDA: slopes at north end, Howell 9737 (stems
and leaves glaucous, flowers light to deep lavender, CAS, type), DeRoy &
DeRoy 31 (DS), Baur 6, 8 (GH), north side, elevation 50 feet, Bowman, 30
July 1957 (CAS). SAN Satvapor: near the shore, Sulivan Bay, Howell 10021
(stems and leaves glaucous, flowers white tinged lavender, CAS). SANTA CRUZ:
Conway Bay, Baur 4 (GH, K), near the shore, Howell 9851 (CAS), sandy
patch near the coast, Cheesman 400 (K); 2 miles inland, Cheesman 401 (K).

PROCEEDINGS

OF THE

~ CALIFORNIA ACADEMY OF SCIENCES
FOURTH SERIES

G Dallas Hanna Anniversary Volume

Vol. XXXII, No. 20, pp. 587-677; 14 figs., 5 pls. November 29, 1968

A FLORA OF THE SAN BRUNO MOUNTAINS
SAN MATEO COUNTY, CALIFORNIA

By
Elizabeth McClintock
California Academy of Sciences

San Francisco, California

and

Walter Knight
Regional Parks Botanic Garden
Berkeley, California

In collaboration with

Neil Fahy

Standard Oil Company
San Francisco, California

INTRODUCTION

The San Bruno Mountains in northern San Mateo County and the hills in
San Francisco County, California, constitute the northern outliers of the Santa
Cruz Mountains. The San Bruno Mountains have remained relatively undis-
turbed, except for the City of Brisbane and today constitute an “island” entirely
surrounded by metropolitan areas. The covering of vegetation on the San Bruno
Mountains contains many of the plants which once were found to the north in
San Francisco where only fragmentary remnants of the natural vegetation re-
main. However, the pressure of metropolitan development is moving toward
the San Bruno Mountains and soon some of their sections will undergo changes.
This flora records the plants to be found there before these changes occur.

[587]

588 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER.

‘

=
San Bruno Mountain Block

Ficure 1. San Bruno Mountains looking north to the City of San Francisco and Mount
Tamalpais, showing locations of faults and fault blocks on the San Francisco Peninsula.

A Bir oF History (by Neil Fahy)

When members of the Portola Expedition discovered San Francisco Bay in
1769, they also were the first Europeans to see San Bruno Mountain, the main
ridge of the range. Five years later, 1774, Fernando Rivera and four soldiers
climbed the mountain and from its summit watched the sunrise across the bay.
The mountain was named the following year by Bruno Hecata for his patron
saint.

The San Bruno Mountain region included portions of the five Mexican land
grants indicated on figure 2. Rancho Buri Buri was granted to José Antonio
Sanchez, who as a child rode mule-back from Sonora, Mexico, to San Francisco
with Anza in 1776. Sanchez received the grant in 1827 and had it confirmed in
1835. The rancho extended from the salt flats on the Bay west to San Andreas
Valley and from Colma south to Burlingame.

The rancho with the longest name also included most of the mountain and
the Crocker Hills. Canada de Guadalupe, la Visitacién y Rodeo Viejo included
in its name three identifiable places, Guadalupe Valley, the site of Brisbane and
Crocker Industrial Park; the Visitacion Valley district of San Francisco; and
the old rodeo grounds along Islais Creek from Daly City to the intersection of
Alemany Boulevard and Mission Street. The rancho was requested in 1835 by
Miguel Sanchez but his petition was denied. However, it was given in the fol-
lowing year to Jacob P. Leese a naturalized Mexican citizen and a prominent

Vou. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 589

Laguna de la Merced

nine 2 tas
incon de las Salinas

Canada de Guadalupe, Visitacion y Rodeo Viejo Miameimas

Ficure 2. Mexican land grants of San Bruno Mountains and vicinity.

figure in early Bay Area history. The southern boundary of Leese’s rancho is
traced approximately by Hillside Boulevard. Leese later traded the rancho to
Robert Ridley for one in Sonoma County but Ridley had financial problems
and the rancho was divided and sold. Robert Eaton acquired 70 acres and
Alfred Wheeler acquired the rest. After American acquisition, the patent of
Henry R. Payson to 5473 acres was approved in 1865. The rancho was further
subdivided and in 1872 the Visitacion Land Company had the largest holdings.
In 1884 Charles Crocker acquired the holdings of the Visitacion Land Company
which were 3814 acres of the original land grant. In the following year he
acquired the Wheeler parcel comprising 183 acres lying on the shore of San
Francisco Bay. When Crocker’s estate was distributed in 1891, his lands passed
to the Crocker Estate Company, present owners of the San Bruno Mountains.

Three ranchos touched the northern portion of the San Bruno Mountains.
The rancho in Visitacion Valley known as Canada de Guadalupe y Rodeo Viejo
was included in Leese’s rancho of 1841 but was later separated from it. In 1865
it was granted to William Pierce. The rancho which contained Bernal Heights,
Hunters Point, and the area south to San Bruno Mountain was Rincon de las
Salinas y Potrero Viejo (which in translation refers to the corner of the salt
marsh and the old pasture). This was granted in 1839-1840 to José Bernal,
the son of Juan Bernal, a soldier in Anza’s expedition of 1776. The San Miguel
rancho, including Mt. Davidson and Twin Peaks, was granted to José de Jesus
Noé in 1845.

590 CALIFORNIA ACADEMY OF SCIENCES [PRoc. 4TH SER.

Several cities have grown up surrounding the mountains. On the north is
San Francisco. On the northeastern slope of the higher ridge is Brisbane. For-
merly known as Visitacion City, the name Brisbane was adopted when the post
office was established in 1931. The name was changed to avoid confusion with
the Visitacion Valley district of San Francisco. To the south is South San
Francisco, the name used from 1849 through the 1880’s for the area between
Hunters Point and Islais Creek. The present South San Francisco was founded
by a subsidiary of G. F. Swift as a meat packing center. The name was adopted
when the area was incorporated in 1908. To the west are Colma and Daly City.
Originally, Colma was a stop on the San Francisco-San Jose Railroad. It was
called Colma in the 1870’s and the name was given to the post office in 1891.
Daly City was named in 1911 for John Daly, a local property owner.

TopoGRAPHY (by Neil Fahy)

“The name ‘San Bruno Mountains’ |[is| given to the short range which ex-
tends in a direction diagonal to the peninsula from Sierra Point nearly across
to the Pacific.’ On the north it is ‘‘. .. separated by a low divide from the group
of hills on the San Miguel Ranch, to which the name of San Miguel Hills may
be given.” This is the extent of the San Bruno Mountains as recorded in 1865
by the Geological Survey of California under the leadership of J. D. Whitney.
The “hills on the San Miguel Ranch” culminate in Mount Davidson, 929 feet,
the highest point in San Francisco.

San Mateo County contains two distinct, northwest-trending mountain ranges
separated by the Merced Valley which lies to the southwest and south of the
San Bruno Mountains where part of Daly City, all of Colma, and part of South
San Francisco are located. The western range, the backbone of the county, is
the northern continuation of the Santa Cruz Mountains. The eastern range is
the San Bruno Mountain region, with the highest point, Radio Peak, 1314 feet
above the sea (fig. 3, no. 33).

The San Bruno Mountain region is itself composed of two parallel, northwest-
trending ranges, separated for over half their length by Guadalupe Valley (fig. 3,
no. 43). The ranges are united by the ridge or saddle at the northern end of
Colma Canyon (fig. 3, no. 36). The southern range, San Bruno Mountain
proper, extends from Sierra Point on San Francisco Bay to near San Diego
Avenue in San Francisco. It is the higher range and rises abruptly from Merced
Valley about 1000 feet in a horizontal distance of *% of a mile. The northern
range, the Crocker Hills (fig. 3, no. 47), extends from Visitacion Point on the
bay to the vicinity of the Guadalupe School in San Francisco. Its summit is
about 850 feet high.

The region is drained by two main streams and several smaller ones. They
are all intermittent. Guadalupe Creek, flowing through Guadalupe Valley, has
its headwaters in Wax Myrtle, Dairy, and Romanzoffia ravines (fig. 3, nos. 21,

Vor. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 591

Ficure 3. San Bruno Mountains. Key to numbered location on pages 602 and 603.

22, 23). Colma Creek, the other main stream, has its source below the Flower
Garden [in April Brook], and flows westward for a short distance, parallel to
Guadalupe Valley Parkway, and then enters Colma Canyon. The canyon of
Colma Creek makes a deep cleft in the mountain where it enters Merced Valley
(at Market Street in Daly City). The minor streams are mainly in ravines on
the southwest flank of the southern range. In the Crocker Hills, Islais Creek
which bears the Salinan Indian name for the Hollyleaf Cherry (Prunus ilicifolia),
had its headwaters near today’s Mission Drive-In Theater. Houses and freeways
have altered its course.

The eastern shoreline of the mountain group during the Mission Period had
the bay lapping against the sandstone cliffs. Today the shoreline is bay-fill.

GEOLOGY (by Neil Fahy)
Rock TyPEs
The great bulk of San Bruno Mountain and the Crocker Hills is composed
of late Cretaceous (100 million years old) dark greenish-gray graywacke of the
Franciscan Formation. This is a poorly sorted sandstone made up of angular
rock fragments, detrital chert, and with more than 10 percent feldspar. The
angular, unsorted content indicates it was derived from the rapid erosion of the

592 CALIFORNIA ACADEMY OF SCIENCES [PRoc. 4TH SER.

source area and rapid burial in a subsiding depositional basin. Fossils are rare.
The graywacke is exposed in the steep canyons and on the ridge crests but only
as isolated knobs on the gentle slopes. Excellent exposures are visible along the
Guadalupe Valley Parkway and Radio Road. The Franciscan Formation is the
name applied to a group of rock types first described from the San Francisco area.

Small lenses of a hard, brittle, siliceous rock called chert are exposed near
Telford and Diamond streets in South San Francisco and at Point San Bruno.
The marine origin of the chert is indicated by the abundance of microscopic
opaline skeletons of radiolaria, one-celled marine organisms.

Serpentine, the California State rock, outcrops on Point San Bruno, but in-
dustrialization has concealed much of it. Two very small serpentine areas are
present near Serbian Ravine.

There are areas of poorly consolidated sand at the head of Colma Canyon
in The Meadow and along the western base of the mountain. The sand was
originally thought to be dune sand but later work has questioned this.

The largest area of recent depositon is in the Guadalupe Valley which con-
tains alluvial deposits eroded from the surrounding hills.

SOIL

Because San Bruno Mountain is composed almost entirely of one rock type,
there is little variation in the type of soil. It is all derived from the Franciscan
graywacke. The varying factor is the soil depth. The graywacke produces a
relatively thin soil on the steep slopes and a thicker soil on the gentle slopes.

FAULTS

The northern portion of the San Francisco Peninsula is composed of a series
of northwest-trending earth blocks bounded by faults or zones of weakness.
These blocks are named and indicated on figure 1. They are from north to
south: the Telegraph Hill block, the San Miguel Hills block, the San Bruno
Mountain block, the Cemetery block, and the Merced Valley block. The two
which make up the San Bruno Mountain region are the San Bruno Mountain
and the Cemetery blocks.

The City College fault, which extends in an arc from Lands End through
the City College of San Francisco to the bay at Visitacion Valley, forms the
northern boundary of the San Bruno Mountain block. The Hillside fault forms
the southern boundary. At present (1967) no fault evidence has been found
to account for the abrupt termination of the San Bruno Mountains at Sierra
Point. There are, however, many small faults throughout the mountain. The
Cemetery bleck is bounded by the Hillside fault in the north and the Merced
Valley in the south.

STRUCTURE

San Bruno Mountain and the Crocker Hills are part of a great asymmetrical

Vot. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 593

arch or anticline whose axis roughly coincides with Ridge Road. The dipping
strata are visible in most of the earth cuts along Radio Road and the Guadalupe
Valley Parkway. It should be noted that the strata generally dip away from
the mountain.

HIsTORY

The San Bruno Mountain landscape began to develop about 130 million
years ago in the Cretaceous Period. At that time the western portion of Cali-
fornia was beneath a sea whose eastern shore was lapping against the base of
the ancestral Sierra Nevada. Great quantities of sediment were deposited in the
Cretaceous sea. Some of these sediments can be seen as the rocks of San Bruno
Mountain.

The present Coast Range landforms began to form about a million years ago.
A buckling of the earth’s crust produced fracture zones along which earth blocks
moved relative to one another. One of the elevated fault blocks was San Bruno
Mountain.

Economic GEOLOGY

Gold and zinc have been mined from the San Bruno Mountains in past years.
The gold occurred in quartz veins in the graywacke. Several uneconomic claims
were filed in the Crocker Hills just south of the county line, but little develop-
ment work was done.

Two veins of zinc ore were discovered near Diamond Avenue in South San
Francisco in 1919. A 75-foot shaft was sunk and a 6-foot vein was worked for
a few years.

The only mineral products from the mountain at present are crushed rock
and sand.

CLIMATE AND WEATHER

Weather data for the San Bruno Mountains which was supplied by the
Crocker Land Company! confirmed observations made on our plant collecting
forays. The San Bruno Mountains stand alone on the San Francisco Peninsula,
unprotected from the atmospheric conditions which bring about their weather
pattern (see figure 4).

Winp. Wind on the San Bruno Mountains displays the marked diurnal and
seasonal patterns which characterize the San Francisco Peninsula. However,
data for March to June 1967, taken from the Crocker Hills Survey (p. 11), show
that wind speeds are greater at comparable times at a site on the mountain area
than in the nearby City of San Francisco. Wind speeds are less in mornings

1Climate Survey of San Bruno Mountain. Phase 2: On-site Survey of Crocker Hills Climate. Aerosol
Laboratory, Metronics Associates, Inc., Stanford Industrial Park, Palo Alto, California. October 20, 1967.
Prepared for Crocker Land Company, San Francisco, California, and used with their permission.

594 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 4. Aerial view of the San Francisco Peninsula looking west and north with the
San Bruno Mountains in the middleground. The San Francisco Airport is in the immediate
foreground.

and increase from early to mid-afternoon. Speeds are greater in June than in
March. Matched data are given below:

TaBLe 1. Mean Wind Speeds—Crocker Hills vs. San Francisco City (page 11, Crocker
Hills Survey).

1967 TOPARVIES PES Hie 1 P.M. 4 P.M.
CH SF CH SF CH SF
March 10 8 13 10 13 12
April 10 9 12 WH 13 12
May 11 9 13 14 15 15
June 13 9 15 14 18 16

Visitors to the San Bruno Mountains will notice that some sites are more
windy than others owing to its irregularity of terrain and the distribution of the
several colonies of planted trees. Available data show that some of the windiest
sites, obviously those most exposed, are: (1) the 1314-foot summit, (2) the junc-
tion of Radio Road and the road to the Nike Base, and (3) the Nike Base. Kam-
chatka Point which always seemed to us to be very windy and bleak was actually
less windy than these three sites.

Data for wind speeds at night on the highest and most exposed points, such

Vor. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 59

as the 1314-foot summit and the long ridge to the east, are not available. How-
ever, on nights when winds were known to be strong at these sites, it was esti-
mated from known recorded speeds for Mount Tamalpais, in nearby Marin
County, that wind speeds on the San Bruno Mountains summit and ridge may
have reached to 50 to 80 miles per hour in gusts up until midnight.

Wind direction varies with the time of the day and with the season. During
winter the morning to afternoon pattern is frequently north becoming northwest,
and a change to a southerly direction indicates that stormy weather can be ex-
pected. During summer months wind flow is frequently southwest becoming west.

Foc. An extensive low fog cover is dominant over the San Bruno Mountains
during the months of July through August when the summer fog regime prevails
along the central California coast. The fog cover, however, does not usually
extend over the entire area of the mountains during an entire day, but tends to
cover only the western section while the eastern section, with Guadalupe Valley
and the City of Brisbane, frequently remain in the sunshine. Data from the
Crocker Hills Survey, page 17, comparing the number of foggy days between a
site in Guadalupe Valley and a site in the Crocker Hills, show that in the months
from March to May the Crocker Hills site was foggier on 30 percent of the days,
and in June to July on 79 percent of the days, than at the Guadalupe Valley
site. Frequently during afternoons the fog bank has been observed for several
hours in a practically stationary position over the western half of the mountain
area. During these periods the fog lies so close to the ground that a moisture
drip is produced from the trees of Eucalyptus globulus.

TEMPERATURE. The generally cool and mild temperatures characteristic of
the coastal part of the San Francisco Peninsula prevail on the San Bruno Moun-
tains. “The seasonal range of temperature is small, in keeping with the pre-
dominant flow of marine air over the site. Mean daily maximums range from
about 56° F. in winter to about 66° F. in fall. Mean daily minimums are above
freezing during all months. However, several daily minimums below freezing are
to be expected during the winter and early spring, particularly on clear, quiet
nights when extensive cold-air drift occurs in the gullies and canyons. The lowest
minimum temperature likely over the long term is 20° F.

“The elevation of the site tends to produce lower summer temperatures and
warmer winter temperatures on the average than found at nearby sea-level
stations. For example, summer fog on the site is associated with afternoon tem-
peratures near 52° F., while sea level temperatures are frequently 5 degrees
higher at the San Francisco airport and 20 degrees higher at Palo Alto (20 miles
to the south). Warmer winter site temperatures are likely on the slopes and knolls
in the morning hours whenever the general wind regime is weak (about ' of the
time). Under these conditions the exposed parts of the site are frequently in or
near the upper and warmer part of the radiation inversion layer that begins at
sea level.” (Crocker Hills Survey, Appendix A, page 3.)

RAINFALL. “Total yearly rainfall on the site is estimated to be somewhat

596 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

greater than at San Francisco or about 22 inches. ... The mean number of days
per year with measurable rain (snow is rare) is about 66, mostly during the
months November—April.” (Crocker Hills Survey, Appendix A, page 3.) Rain
is rare during the remainder of the year.

VEGETATION

The area of the San Bruno Mountains covered by this flora contains about
3000 acres and reaches 1314 feet at the highest point. Although a relatively
small area, the topography is sufficiently irregular to provide suitable habitats
for four of the plant communities which occur in the Santa Cruz Mountains.
These are grassland, coastal scrub, foothill woodland, and salt marsh. Grassland
covers the largest portion of the area, all of the Crocker Hills, the entire south-
eastern slopes, and most of lower western slopes of the taller southern range. A
mixture of native and introduced grasses and such associates as Brodiaea laxa,
B. pulchella, Sisyrinchium bellum, Ranunculus californicus, Lupinus nanus, L.
bicolor, Sidalcea malvaeflora, Sanicula arctopoides, Plantago erecta, and Baeria
chrysostoma may be found in this plant community.

Coastal scrub occurs in some of the ravines and canyons such as Colma
Canyon, the upper part of Buckeye Canyon, and Islay Ravine. Some of the
characteristic species are: Anaphalis margaritacea, Achillea millefolium, Artemisia
californica, Eriogonum latifolium, Baccharis pilularis var. consanguinea, Diplacus
aurantiacus, Rhus diversiloba, and Lupinus arboreus.

Coastal woodland is best developed in lower Buckeye Canyon. Aesculus
californica, Quercus agrifolia, and Umbellularia californica make a fairly dense
woodland.

The Salt Marsh community occurs at Point San Bruno and Sierra Point but
little of it remains today.

CoLMA CANYON WATERSHED

Colma Canyon is accessible from the eastern end of Market Street in Daly
City. In late 1964 a paved road called Guadalupe Parkway was completed along
the floor of the canyon and from there to the junction with Radio Road. The
thoroughfare now continues along the southeast edge of the Crocker Hills and
joins old Bayshore Highway north of Brisbane.

The construction of Guadalupe Parkway left the steep slopes of Colma
Canyon, with its floristically interesting plant cover, unchanged, but it did
change the course of Colma Creek. Originally only a foot trail wound along the
creek. Equestrians and hikers would meander through the vegetation along the
water’s edge. The creek ran all year around and occasional pools would invite
youthful swimmers who would be secluded by the overhanging boughs of
Coulter and arroyo willows, salmonberry, lady fern, twinberry, snowberry, and
creek dogwood which provided shelter from the brisk afternoon winds.

Vout. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 597

Along the present parkway on the lower slopes of the canyon can still be
seen elegant displays of lady ferns, salmonberries, and Coulter willow. Creek
dogwood, however, no longer remains due to construction of the parkway. Along
the canyon sides from the creek to the ridges, particularly in the vicinity of the
rocky outcroppings one can find coast barberry (Berberis pinnata), poison oak
(Rhus diversiloba), red berry (Rkamnus crocea), holly leaf cherry (Prunus
ilicifolia), California polypody (Polypodium californicum), spreading dudleya
(Dudleya cymosa), bluff lettuce (Dudleya farinosa), Pacific stonecrop (Sedum
spathulifolium), seaside daisy (Erigeron glaucus), rock cress (Arabis blepharo-
phylla), and serviceberry (Amelanchier pallida).

At the mouth of Colma Canyon, especially on the south side, is a sand de-
posit reminiscent of the former dune areas in the western part of San Francisco.
Here are found typical coastal dune plants such as yellow beach lupine (Lupinus
arboreus), blue beach lupine (Lupinus chamissonis), sand mat (Cardionema
ramosissimum), California whitlowwort (Paronychia franciscana), spine-flower
(Chorizanthe pungens var. hartwegii), and dune knotweed (Polygonum paro-
nychia). This dune area continues through the mountain to the southeast and
appears again at the mouth of Pig Ranch Ravine. Besides a partial representa-
tion of the Colma Canyon dune plants, this odoriferous ravine is the only locality
on the mountain for the California croton (Croton californicus).

As one proceeds up Colma Canyon in the areas where the soil has accumu-
lated humus in the decomposed sandstone, the vegetation cover consists of a
soft chaparral interspersed with herbs and grasses. This vegetation, which also
inhabits the little side ravines fingering out from the canyon, includes buckwheat
(Eriogonum latifolium), California sagebrush (Artemisia californica), sticky
monkeyflower (Diplacus aurantiacus), dwarf chaparral broom (Baccharis pilu-
laris), phacelia (Phacelia californica), pennyroyal (Monardella villosa var.
franciscana).

There are no trees in the canyon proper but introduced eucalyptus (/uca-
lyptus globulus) and cypress (Cupressus macrocarpa) trees are to be seen along
Radio Road. Many of the cypresses have their trunks adorned with the leather
fern (Polypodium scouleri) growing as an epiphyte as high as 20 feet. This
epiphytic habitat is probably subsidized by the extreme moisture from the heavy
fogs which roll through the entrance of Colma Canyon. For this reason, the
cypress grove has been called the Fog Forest.

The upper part of Colma Canyon opens into a large slightly sloped flat
area, which we call the Great Meadow, a grassland punctuated with an occasional
dark green prostrate mat of the dwarf chaparral broom (Baccharis pilularis).
Along the broad upper slopes draining into the Great Meadow are sheets of
dark yellow color in July consisting of lizard tail (Eriophyllum staechadifolium).
In August on adjacent slopes there are spreading white blankets of pearly ever-
lasting, California’s most beautiful cudweed (Anaphalis margaritacea).

598 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER.

To the south of the Great Meadow extending gradually uphill for about a
mile is April Brook Ravine. April Brook is damp during all the seasons of the
year and it empties into Colma Creek near a colony of Coulter willows (Salix
coulteri) just below the Great Meadow. The drainage areas of April Brook
Ravine are more gradual than the more steep slopes of Colma Canyon.

In the spring, on the eastern slopes of April Brook Ravine, there are sizeable
colonies of goldfields (Baeria chrysostoma), which well substantiate the com-
mon name. In colonies equally as large are white masses of johnny-tuck (Ortho-
carpus erianthus). On occasional slopes between Radio Road and April Brook
are colonies of both coast iris (Jris longipetala) and Douglas iris (Jris douglasiana).
Even though their flowering times partially overlap, there has never been evi-
dence of any interbreeding between the two irises. Also in this area the two
native wild strawberries, Fragaria californica and F. chiloensis, occur together.

An unusual distributional record in the April Brook drainage area is the
occurrence of the dune tansy (Tanacetum camphoratum) at the edge of Radio
Road near Summer Seep. In this same watershed are serviceberry (Amelanchier
pallida), coffeeberry (Rhamnus californica), and Indian hemp (A pocynum
pumilum).

The only other sizeable area draining ultimately into Colma Canyon is Cable
Ravine. It is accessible from Radio Road about a half mile south of Guadalupe
Parkway intersection at a eucalyptus grove. Up this little ravine is the only
locality hereabouts of salal (Gaultheria shallon) on a very prominent rocky out-
crop which overlooks the lower reaches of the ravine. Progressing up the slope
one can see in spring, colorful yellow patches of johnny-jump-up (Viola peduncu-
lata). Nearby is the best display of bearberry manzanita (Arctostaphylos uva-
ursi) on the entire mountain forming a ground-hugging drape as it clings to the
exposed grassy slope.

THE RIDGE AND ITS RAVINES

A gravel-surface road extends for two miles east of the parking lot along
the ridge. Both ends of the ridge are bisected by high voltage transmission lines.
In spring on the south-facing slopes at the western end of the mountain are
many plants of the bright red Indian paint brush (Castilleja franciscana), and
during late spring and early summer crimson sage (Salvia spathacea) displays
itself over a considerable acreage.

If one can endure the strenuous descent into the brushy ravines, thickets of
poison oak (Rhus diversiloba) can be found with some plants almost arboreal
in habit having trunks up to 11 inches in diameter. From the ravines, Ceanothus
thyrsiflorus extends in large colonies up to the ridge road and in April its large
clusters of blue flowers are a striking contrast against the dark green foliage.

The southeastern section of the mountain is almost exclusively grassland.

The north-facing slopes include ridges and ravines that are brushy in vary-

Vor. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 599

ing degrees with only a small amount of grassland. Starting near Brisbane and
progressing toward the quarry California buckeyes, California laurels, Cali-
fornia live oaks, and holly-leaf cherry are found in the ravines. Between the
quarry and westward toward the parking lot and Radio Road can be seen a
brushy mantle of vegetation consisting of evergreen huckleberry, manzanitas,
and other woody plants interspersed with herbaceous perennials such as coast
larkspur (Delphinium californicum), coast angelica (Angelica hendersonit),
several species of Lomatium and Sanicula, and in the spring Heracleum maxi-
mum conspicuously whitens the ravines.

To the west of Lipman School in Brisbane is the base of the densely wooded
ravine which we call Buckeye Canyon. California buckeyes occur along the
intermittent creek in the lower part of the ravine. Advancing up the slope from
the creek are shrubs of chaparral currant (Ribes malvaceum) and above these
California laurels (Umbellularia californica) dominate the plant cover of the
canyon. On the upper slopes of the canyon particularly around rocky out-
cropings are low shrubs of the gold-cup oak (Quercus chrysolepis var. nana).

On a windy promontory on the down-side of Radio Road as one approaches
the first radio station, there is an area which is frequently so cold and windy
that it has been named Kamchatka Point. Among the plants here are two un-
usual manzanitas, the evergreen huckleberry, and a deciduous huckleberry which
we consider to be Vaccinium arbuscula, and another rarity, false lily-of-the-
valley (Maianthemum dilatatum).

GUADALUPE VALLEY

Crocker Industrial Park now covers most of the valley. On the west end of
the drainage are the botanically interesting Fern Rock and Wax Myrtle Ravine.
Around Fern Rock are coastal wood fern (Drvyopteris arguta), leather fern
(Polypodium scouleri), western sword fern (Polystichum munitum), bracken
fern (Pteridium aquilinum), and chain fern (Woodwardia fimbriata). In the
ravine are fine specimens of wax myrtle (Mvyrica californica), and several
herbaceous annuals and perennials including bleeding heart (Dicentra formosa).
At the upper part of the ravine below the new Guadalupe Parkway are pitcher
sage (Lepechinia calycina), cascading, compact shrubs of redberry (RAamnus
crocea), and low dwarf shrubs of gold-cup oak (Quercus chrysolepis var. nana).

At the southwestern end of the valley is a large active quarry forming a
wide chasm where it penetrates the main mountain ridge. Recent enlargement
of the quarry eliminated a thicket of shrubs in which were creek dogwood
(Cornus californica) and coast red elderberry (Sambucus callicarpa).

CROCKER HILLS

This area constitutes the range of hills running east to west from the old
Bayshore Highway to the spot designated on the aerial photograph as Reservoir
Hill (fig. 3). It has a largely introduced and weedy collection of plants ac-

600 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

cented here and there by the spiny European gorse (Ulex europaeus). Despite
its obnoxious qualities, the prolificly flowering gorse makes a dark yellow ex-
panse of color in spring contrasting with adjacent sky lupine (Lupinus nanus)
and two adventive members of the mustard family, wild radish (Raphanus
sativus), and field mustard (Brassica campestris). An interesting concentration
of plants abounds at Reservoir Hill in the spring and the little area should be
visited at that time.

Point SAN BRUNO

Within the last 5 years much grading and filling has been done on this point
of land which extends into the bay. It is a continuation of San Bruno Moun-
tain and extends across the freeway. The interesting plants here were of a
coastal strand type with wind-shorn, low-growing buckeye trees and willows
along the top of the bluffs. Most of the bay below the bluffs has been filled.
Little salt flat coves originally supported pickleweed (Salicornia pacifica),
arrowgrass (7viglochin concinna), sea lavender (Limonium californicum), and
Jaumea carnosa, but these are disappearing in the path of development.

REGENERATION OF PLANTS AFTER FIRE

Most plants of chaparral and coastal scrub are characterized by their ability
to regenerate after fire by sprouting from root crowns. These basal structures
are usually noticeably enlarged at, or just below, the ground level. Other plants
not reproducing by crown sprouts sometimes produce abundant seedlings after
a fire. During our four years of plant collecting on the mountain there have
been fires in different areas. Listed here are the plants which we have observed
to regenerate after some of these fires:

Plants stump sprouting after fire:

Arctostaphylos pacifica Quercus wislizeni
Baccharis_ pilularis Rhamnus californica
Ceanothus thyrsiflorus Rosa gymnocarpa
Diplacus aurantiacus Rubus parviflorus
Heteromeles arbutifolia Salix lasiole pis
Holodiscus discolor Salvia spathacea
Monardella villosa var. franciscana Sambucus callicarpa
Osmaronia cerasiformis Symphoricarpos species
Prunus emarginata Vaccinium ovatum

Plants seeding abundantly after fire:

Arctostaphylos imbricata Grossularia californica
Arctostaphylos montaraensis Lepechinia calycina
Artemisia californica Ribes malvaceum

PLANTS OF SAN BRUNO MouNTAIN USED AS ORNAMENTALS

Many of the colorful and attractive California native plants which are used
as garden ornamentals may be seen in their native habitats on the San Bruno

Vout. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 601

Mountains. In the San Francisco Bay region are three botanical gardens where
many of the San Bruno plants may be seen growing under garden conditions
and where information regarding their culture may be obtained. Tilden Regional
Park Botanic Garden, Berkeley, devoted entirely to growing and displaying
California native plants, grows all of the plants which are listed here. The Uni-
versity of California Botanical Garden, Berkeley, and the Strybing Arboretum,
Golden Gate Park, San Francisco, have sections devoted to California natives
and some of the plants listed here may be seen at both gardens:

Allium dichlamydeum
Allium unifolium
Anaphalis margaritacea
Arabis blepharophylla
Arbutus menziesii
Arctostaphylos uva-ursi
Aristolochia californica
Baccharis pilularis
Berberis pinnata
Brodiaea laxa
Ceanothus thyrsiflorus
Clarkia rubicunda
Cornus californica
Delphinium californicum
Dicentra formosa
Diplacus aurantiacus
Dudleya cymosa
Dudleya farinosa
Erigeron glaucus
Erigeron philadelphicus
Erysimum franciscanum
Eschscholzia californica
Fragaria californica
Fragaria chiloensis
Garrya elliptica
Heteromeles arbutifolia
Heuchera micrantha

Holodiscus discolor

Iris douglasiana

Iris longipetala
Maianthemum dilatatum
M yrica californica
Osmaronia cerasiformis
Physocarpus capitatus

Potentilla egedii var. grandis

Prunus demissa
Prunus emarginata
Prunus ilicifolia

Ribes malvaceum
Romanzoffia suksdorfii
Rosa californica

Rosa gymnocarpa
Rubus parviflorus
Rubus spectabilis
Satureja douglasi
Sedum spathulifolium
Sisyrinchium bellum
Thalictrum polycarpum
Trillium chloropetalum
Vaccinium arbuscula
Vaccinium ovatum
Viola adunca

Viola pedunculata
Zauschneria californica

PLANT COLLECTIONS

The collections on which this flora is based were begun early in 1963. They
were made by Walter and Irja Knight (cited in the Catalogue of Plants as Kk),
E. McClintock (cited as McC), Philip Wheeler (cited as W), Javier Pefnalosa
(cited as P), Carlyn Halde, Virginia Ryder, J. H. Thomas, Peter Raven,
Katherine Toschi, Arthur Menzies, James B. Roof, and Lewis Rose. The cited
specimens are deposited in the Herbarium of the California Academy of Sciences
except those collected by J. H. Thomas which are in the Dudley Herbarium,
Stanford University.

602 CALIFORNIA ACADEMY OF SCIENCES

PLACE NAMES

[Proc. 4TH SER.

When we began our plant collecting on San Bruno Mountain in early 1963
we found that there were only a few place names in general use for parts of the
mountain. Since we needed to designate our collecting localities we assigned our
own names to these places. Most place names regardless of their origin are listed
here and are located approximately on the aerial photograph of the mountain

region (fig. 3).

April Brook

Bitter Cherry Ridge
Blue Blossom Hill
Brisbane

Brisbane Ravine
Buckeye Canyon
Cable Ravine
Colma Canyon
Cow Trough Ravine
Crocker Avenue
Crocker Hills
Crocker Industrial Park
Dairy Ravine
Devil’s Arroyo
Eucalyptus Grove
Fern Rock

Flower Garden

Fog Forest

Gladys Ravine

Glen Park Ravine in Brisbane
Guadalupe Valley
Islay Ravine

Juncus Ravine
Kamchatka Point
Manzanita Dike

Bitter Cherry Ridge

April Brook

Flower Garden

Kamchatka Point

Quarry watershed

Olivet Ravine

Pig Ranch Ravine

Serbian Ravine (serpentine)

ne wd

oOo CO ~I OO

Serbian Cemetery
10 Olivet Cemetery

11 Sage Ravine

12 Poison Oak Ravine
13. Tank Ravine

14 Juncus Ravine

15 Cow Trough Ravine

Monterey Pine

Nike Base

Olivet Cemetery

Olivet Ravine

Owl’s Canyon

Parking Lot at summit
Pig Ranch Ravine

Point San Bruno

Poison Oak Ravine
Powerlines, East and West
Quarry watershed

Radio Road

Randolph Ravine, end Randolph Avenue
Reservoir Hill

Ridge Road

Romanzoffia Ravine

Sage Ravine

Serbian Cemetery

Serbian Ravine (serpentine)
Sierra Point

Summer Seep

Tank Ravine

Trillium Gulch

Wax Myrtle Ravine

26 Owl’s Canyon
27 Islay Ravine

28 Buckeye Canyon
29 Brisbane Ravine
30 Gladys Ravine
31 Trillium Gulch
32 Nike Base

33 Parking Lot

34 Powerlines, East and West
35 Devil’s Arroyo
36 Colma Canyon
37 Radio Road

38 Ridge Road

39 Cable Ravine

40 Brisbane

Vou. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 603

16 Randolph Ravine, end Randolph Avenue 41 Reservoir Hill

17 Fog Forest 42 Summer Seep

18 Eucalyptus Grove 43 Crocker Industrial Park in Guadalupe
19 Fern Rock Valley

20 Monterey Pine 44 Point San Bruno

21 Wax Myrtle Ravine 45 Sierra Point

22 Dairy Ravine 46 Crocker Avenue

23 Romanzoffia Ravine 47 Crocker Hills

24 Manzanita Dike 48 Glen Park Ravine in Brisbane

25 Blue Blossom Hill

TasLe 2. Tabulation of the Vascular Plants.

Division
Class Species
Subclass Family Genus Native Introduced Total
Calamophyta 1 1 2 2
Pterophyta
Filicinae 1 8 9
Gymnospermae 2 2 2 2
Angiospermae
Monocotyledoneae 10 52 69 28 97
Dicotyledoneae 64 256 304 128 432
384 158 542
71% 29%

Taste 3. List of the 10 largest families of vascular plants.

Family Number of genera Number of species
Compositae 52 90
Gramineae aA 49
Cruciferae 18 25
Rosaceae 15 23
Leguminosae 13 42
Umbelliferae 13 18
Scrophulariaceae 9 19
Caryophyllaceae 9 17
Liliaceae 10 15
Labiatae 10 13

TaBLe 4. List of the largest genera.

Number of species in each genus

Lupinus, Trifolium 10
Juncus 8
Rumex, Senecio 7
Carex, Festuca, Gnaphalium, Orthocarpus 6
Bromus, Lotus 5

604 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

ACKNOWLEDGMENTS

We are grateful to those whose collections were used. In addition we wish
to thank John Thomas Howell who identified the grasses, sedges, and rushes,
helped with identifications in other groups, and gave freely of his time and sug-
gestions. We gratefully acknowledge his help and encouragement. Irja Knight
participated in all of the field work and helped with the project in numerous
ways. For her capable assistance we extend our special thanks. We thank
Lauramay Dempster, Rimo Bacigalupi, Alva Day, Peter Raven, and John
Hunter Thomas for identifications in several groups of plants. The Crocker
Estate Company allowed us to choose from their file the aerial photographs used
in figures 1 to 4. We acknowledge with thanks their cooperation and interest in
this project and for financial assistance given in 1963, making it possible to do
field work during that year. We are grateful to James B. Roof for his interest
in this project and for his knowledgeable comments when he accompanied us on
field trips.

ANNOTATED CATALOGUE OF VASCULAR PLANTS

In the Flora of the San Bruno Mountains we have, in most instances, used
the plant names in the Flora of the Santa Cruz Mountains of California by John
Hunter Thomas (in 1961). Keys to the nearly 550 different kinds of plants
of the San Bruno Mountains have not been included here since those in the
Flora of the Santa Cruz Mountains are applicable. For ease of use by both
amateur and professional botanists the sequence of families within their divisions,
classes, and subclasses is alphabetical.

DIVISION CALAMOPHYTA
EQUISETACEAE. Horsetail Family

Equisetum arvense Linnaeus. Common Horsetail. Rare, on wet ground. Colma Canyon,
K May 30, 1964; Radio Road, W May 23, 1965.

Equisetum telmateia J. F. Ehrhart var. braunii (Milde) Milde. Giant Horsetail.
Occasional, along streams and seepages. Colma Canyon, McC, Halde, K June 23, 1963; Cow
Trough Ravine, K 341; Devil’s Arroyo, McC March 28, 1965; Quarry, K July 14, 1963;
Radio Road, K August 5, 1963.

DIVISION PTEROPHYTA
CLASS FILICINAE
POLYPODIACEAE. Fern Family

Adiantum jordani K. Mueller. California Maidenhair Fern. Occasional, in damp
protected places. Quarry, K July 14, 1963; Romanzoffia Ravine, McC, Menzies February 16,
1963; Owl’s Canyon, K 1049; Colma Canyon, K March 15, 1964; Glen Park Canyon, McC,
W May 14, 1967.

Athyrium filix-femina (Linnaeus) Roth. Western Lady Fern. Rare, in wet gullies.
Base of Quarry, K July 14, 1963; Colma Canyon, along creek, McC, Halde, K June 23, 1963.

Vout. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 605

Dryopteris arguta (Kaulfuss) Watt. Coastal Wood Fern. Occasional, brushy, well
drained slopes. Buckeye Canyon, K August 5, 1963; Fern Rock, K July 14, 1963; Powerline
Ridge, McC & K September 2, 1965; 400 feet northeast of Parking Lot, K February 28, 1963;
Quarry, K July 14, 1963.

Pityrogramma triangularis (Kaulfuss) Maxon. Goldenback Fern. Rare, shaded well
drained slopes. Dies back during winter. Harold Road, Brisbane, W December 30, 1966;
Glen Park Canyon above Brisbane, McC April 16, 1967.

Polypodium californicum Kaulfuss. California Polypody. Occasional, rocky slopes.
Cable Ravine, McC March 14, 1965; ravine % mile east of 1314-foot summit, P 2700;
Romanzoffia Ravine, McC, Menzies February 16, 1963; Glen Park Canyon, McC April
16, 1967.

Polypodium scouleri Hooker & Greville. Leather Fern. Occasional, crevices and surfaces
of rocks and on tree trunks in Fog Forest. Fern Rock, K August 5, 1963; Fog Forest, on
trunk of Monterey cypress, K September 2, 1964; Fog Forest, on trunk of blue gum, McC,
Halde June 23, 1963; Cable Ravine, McC March 14, 1965.

Polystichum munitum (Kaulfuss) Presl. Western Sword-Fern. Common, in brushy
and rocky areas. April Brook, W July 7, 1965; Colma Canyon, K March 15, 1964; Fern
Rock, K August 5, 1963; east facing slope below Parking Lot, McC, Halde June 23, 1963;
West Powerline, McC March 28, 1965 (plants growing in an area burned during preceding
autumn, were sprouting from crown); Trillium Gulch, McC March 28, 1965.

Pteridium aquilinum (Linnaeus) Kuhn var. pubescens Underwood. Bracken Fern.
Widespread in grassland and on brushy slopes. Cable Ravine, McC March 14, 1965; Colma
Canyon, McC, Halde, K June 23, 1963; Fern Rock, K August 5, 1963; West Powerline,
McC March 28, 1965; slope below Parking Lot, McC June 9, 1963; east of Quarry, W June
19, 1965; Radio Road at Horseshoe Ridge, K August 5, 1965.

Woodwardia fimbriata J. E. Smith. Western Chain Fern. Rare, in moist places. Colma
Canyon, W May 18, 1965.

CLASS GYMNOSPERMAE
CUPRESSACEAE. Cypress Family

Cupressus macrocarpa Hartweg. Monterey Cypress. Large trees probably planted
many years ago; some young trees spontaneous. Fog Forest along Guadalupe Road, K
August 5, 1963.

PINACEAE. Pine Family

Pinus radiata D. Don. Monterey Pine. Single young tree, probably planted. Cable
Ravine, K September 30, 1964.

CLASS ANGIOSPERMAE
SUBCLASS DICOTYLEDONEAE

AIZOACEAE. Carpet-weed Family

Mesembryanthemum edule Linnaeus. Hottentot-Fig. Planted as a groundcover and
occasionally becomes naturalized as along Radio Road. Flowers spring and summer. Orna-
mental. Useful on banks and roadcuts. Native of South Africa.

606 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Tetragonia tetragonioides (Pallas) Kuntze [Tetragonia expansa Murray]. New
Zealand Spinach. Cultivated for food, it escapes and becomes naturalized in disturbed and
marshy areas. Point San Bruno, K, McC September 2, 1964; Brisbane Lagoon, W August
14, 1965; trash dumps at west end of mountain, K 1081. Native of southeastern Asia, Aus-
tralia, and New Zealand.

AMARANTHACEAE. Amaranth Family

Amaranthus powellii Watson. Amaranth. Pig-Weed. Weed in disturbed areas.
Abandoned quarry, near Hillside Boulevard and School Street, K October 7, 1963.

ANACARDIACEAE. Sumac Family

Rhus diversiloba Torrey & Gray. Poison-oak. Shrub with 3-foliolate leaves. It is
variable in habit from subarborescent to shrubby and climbing, and in leaf shape and size.
One of the common plants of the mountain, usually occurring with chaparral shrubs but in
one locality, on a south facing slope at the base of Blue Blossom Hill, it occurs alone. The
foliage turns reddish in autumn and is showy. Contact with any part of the plant may
cause a dermititis in those individuals who are susceptible.

Schinus molle Linnaeus. Pepper Tree. Cultivated as an ornamental tree, occasionally
becomes naturalized. Old Bayshore Highway, Brisbane, W October 16, 1966. Native of
South America.

ApocyNacEAE. Dogbane Family

Apocynum pumilum (Gray) Greene. Dogbane. Indian-Hemp. Rare, in grassy or rocky
areas. Radio Road at Summer Seep, K August 5, 1963; Radio Road, above Great Meadow,
W June 20, 1965.

Vinca major Linnaeus. Periwinkle. Cultivated as a groundcover, escapes and becomes
naturalized. Near Bayshore Highway, Brisbane, W June 8, 1965. Native of Mediterranean
region.

ARALIACEAE, Aralia Family

Hedera helix Linnaeus. English-Ivy. Cultivated as a vine or groundcover, escapes
and becomes naturalized. Radio Road, near radio towers, W June 13, 1965. Native of Europe.

ARISTOLOCHIACEAE. Aristolochia Family

Aristolochia californica Torrey. Dutchman’s Pipe. California Pipe Vine. Rare,
grassy slope with scattered shrubs, growing on the ground or scrambling over shrubs. Edge
of southern ravine of Glen Park Canyon, McC, W May 14, 1967. Deciduous scandant shrub
with characteristically U-shaped flowers, February to May. It is interesting because of the
unusual shape of its flowers which in some species are shaped like a pipe. Aristolochia cali-
fornica is a California endemic from the northern half of the state, San Bruno being its
southernmost known locality. Aristolochia, a large genus of about 450 species distributed
mainly in tropical and subtropical regions, has pendulous flowers of an extraordinary form,
often disagreeable odor, and complicated pollination mechanisms. Several, including A. cali-
fornica, are used as garden and greenhouse subjects for their unusual flowers.

VoL. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 607

BERBERIDACEAE. Barberry Family

Berberis pinnata Lagasca [Mahonia pinnata (Lagasca) Fedde]. Coast Barberry. Fre-
quent at rock outcroppings. In open windswept habitats it tends to be low-growing. Reser-
voir Hill, Menzies March 14, 1965; Radio Road at Horseshoe Ridge, K August 5, 1963; Radio
Road near radio towers, McC, Menzies February 16, 1963; Nike Base, K March 15, 1964;
Quarry, K982; Colma Canyon, McC, Halde, K June 23, 1963; Ridge Road near West Power-
line, McC, K September 2, 1963. Flowers March to May. Attractive evergreen shrub
throughout the year; the yellow flowers are showy in the spring and the new foliage which
precedes the flowers is glossy and reddish-green. Ornamental.

BETULACEAE. Birch Family

Corylus californica (A. DeCandolle) Rose. California Hazel. Occasional, in ravines or
along creeks. Colma Canyon, K March 15, 1964; upper Bitter Cherry Ravine in a thicket,
K 1110; Quarry, K June 24, 1963; Owl’s Canyon, W June 7, 1965. One of the mountain’s
interesting deciduous shrubs with flowers in pendant catkins in early spring before the
leaves, and small edible nuts in autumn. Ornamental.

BoRAGINACEAE. Borage Family

Allocarya chorisiana (Chamisso) Greene. Rare, in damp ground. Point San Bruno,
K April 24, 1965; Devil’s Arroyo, K 1100; Ridge Road at West Powerline, McC, K May
16, 1965; Upper Colma Canyon, McC, W June 11, 19067.

Amsinckia intermedia Fischer & Meyer. Fiddleneck. Frequent in grassland and dis-
turbed areas. Colma Canyon, McC, Halde, K June 23, 1963; Bitter Cherry Ridge, K May
4, 1964; Romanzoffia Ravine, K May 4, 1964; Devil’s Arroyo, K 1089; Owl’s Canyon, K
1088; Ridge Road at West Powerline, K April 13, 1964; Ridge Road at Brisbane Powerlines,
K March 21, 1965; Brisbane Ravine, W March 30, 1965.

Cryptantha hispidissima Greene. Rare, in grassland. Devil’s Arroyo, K 1124; Quarry,
K July 14, 1963. A stiff, bristly herbaceous plant, with small white flowers May to July.

Cryptantha micromeres (Gray) Greene. Occasional, edge of chaparral, sometimes in
burned over areas. In a ravine to west of Radio Road, K April 25, 1964; Trillium Gulch,
McC March 28, 1965; Devil’s Arroyo, McC March 28, 1965; West Powerline, K 1091. Plants
often form dense colonies, and have a profusion of small white flowers, April to June.

Cynoglossum grande Douglas ex Lehmann. Hound’s Tongue. Occasional, in grassland
and chaparral, sometimes in burned over areas. Ridge Road, in a ravine 12 mile east of
Parking Lot, P 2684; Romanzoffia Ravine, K March 21, 1965; Brisbane Ravine, W March
30, 1965; Trillium Gulch, K 1021; Glen Park Canyon, McC, W May 14, 1967. Sometimes
used as an ornamental for its attractive basal leaves and blue, forget-me-not-like flowers,
February to April.

Myosotis latifolia Poiret. Forget-me-not. An escape from cultivation, grows along
roadsides, stream sides and in shaded areas. Guadalupe Road, K August 5, 1963; April
Brook, W May 18, 1965; Brisbane Ravine, W May 25, 1965. Flowers in spring and summer.
Native of Europe and North Africa.

CALLITRICHACEAE. Water Starwort Family

Callitriche marginata Torrey. Water Starwort. Rare, floating in water of stream,
or on surface of mud at edge of stream. Randolph Ravine, P 2738. The tiny green plants
are usually numerous enough to be conspicuous, often forming mats.

608 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

CAMPANULACEAE. Bellflower Family

Triodanus biflora (Ruiz & Pavon) Greene. Venus’ Looking Glass. Rare, in grassland
and edge of chaparral. Crystal Cave Canyon, K 1125, and W June 19, 1965. The small blue
to lavender flowers are axillary and sessile, in June. May become weedy.

CAPRIFOLIACEAE. Honeysuckle Family

Lonicera hispidula Douglas ex Lindley. Honeysuckle. Scandant on adjacent plants or
sometimes on ground. Rare. Radio Road, K August 5, 1963.

Lonicera involucrata (Richardson) Banks ex Sprengel [Z. ledebourii Eschscholtz].
Twinberry. Rare, along streams and in shade of chaparral. Colma Creek, McC, Halde, K
June 23, 1963. (Much of Colma Creek was destroyed by road construction, therefore
Twinberry may no longer be growing at this locality.) Bitter Cherry Ravine, K April 4,
1965. Flowers April to June. Ornamental both in flower and fruit.

Lonicera japonica Thunberg. Japanese Honeysuckle. Escape from cultivation, along
roadsides and disturbed places. Crocker Avenue near intersection with Guadalupe Road,
K September 2, 1964. Native of Japan.

Sambucus callicarpa Greene. Red Elderberry. Rare, brushy ravines, edge of chaparral.
Near Olivet Cemetery, W May 18, 1965; base of Devil’s Arroyo, McC March 28, 1965;
Quarry, K June 24, 1963; Ridge Road near West Powerline, McC, K September 2, 1963, and
J. Roof March 27, 1966. Flowers March and April. Stump sprouts following fire. Orna-
mental particularly in fruit.

Sambucus mexicana Pres] ex DeCandolle. Blue Elderberry. Rare, brushy ravines, edge
of chaparral. Quarry, K June 24, 1963; Buckeye Canyon, K August 5, 1963; Point San
Bruno, K 1029; Sierra Point, W May 12, 1965. Flowers April to June. Ornamental for
foliage and blue-black glaucous fruits.

Symphoricarpos albus (Linnaeus) Blake var. laevigatus (Fernald) Blake [S. rivularis
Suksdorf]. Snowberry. Occasional, brushy slopes, edge of chaparral. Stump-sprouts after a
fire. East facing slope below Parking Lot, McC, Halde June 23, 1963; Radio Road, McC,
K September 2, 1963; Quarry, K July 14, 1963; Colma Canyon, K May 30, 1964. Flowers
May to August. White fruits in late summer and autumn are more conspicuous than flowers.
Ornamental, but spreads vigorously by underground rootstocks, therefore must be kept within
bounds.

Symphoricarpos mollis Nuttall. Trailing Snowberry. Rare, brushy slopes, edge of
chaparral. Quarry, K June 24, 1963; Live Oak Canyon, W April 27, 1965. Flowers April to
July followed by showy white berries.

CARYOPHYLLACEAE. Pink Family

Arenaria macrophylla Hooker. Sandwort. Rare, rocky outcrop. Kamchatka Point,
McC March 21, 1965; J. Roof April 24, 1965.

Cardionema ramosissimum (Weinmann) Nelson & Macbride. Sand-Mat. Occasional,
sandy soil. Lower Colma Canyon, McC, Halde, K June 23, 1963; Pig Ranch Ravine, K 1082;
near Olivet Cemetery, W May 18, 1965; Crocker Hills, W May 17, 1965. Flowers May to
June.

Vou. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 609

Cerastium arvense Linnaeus. Mouse-ear Chickweed. Rare, in grassland. Reservoir
Hill, McC March 14, 1965; Nike Base, K May 15, 1965. Flowers March to May. A showy
spring wildflower.

Cerastium viscosum Linnaeus [C. glomeratum Thuillier]. Mouse-ear Chickweed. Weed
of gardens, roadsides, and disturbed places. Ravine to south of Crocker Avenue, P 2653;
north slope of San Bruno Mountain above Crocker Avenue, Toschi 63:16; above the Quarry,
K July 14, 1963; West Powerline, K 948. Flowers March to July. Native of Europe.

Paronychia franciscana Eastwood. Rare, grassland or rocky slopes. Colma Canyon,
McC, Halde, K June 23, 1963; near Nike Base, McC, W May 23, 1965. Mat forming weedy
perennial, inconspicuous flowers May to June.

Polycarpon tetraphyllum (Linnaeus) Linnaeus. Weed of sandy soil and disturbed
places. Colma Canyon, McC, Halde, K June 23, 1963; Crocker Hills near Cow Palace, W
May 17, 1965. Native of Europe.

Sagina apetala Arduino var. barbata Fenzl ex Ledebour. Pearlwort. Inconspicuous
weed of disturbed places, often on hardpacked soil. Ridge Road near West Powerline, McC
April 25, 1965. Flowers April and May. Native of Europe and Asia.

Sagina occidentalis Watson. Western Pearlwort. Rare, grassland. Ridge Road, near
West Powerline, K 945; near Nike Base, McC, W May 23, 1965. Flowers March to May.

Silene gallica Linnaeus. Windmill Pink. Catchfly. Campion. Pinkish petals are twisted
simulating the blades of a windmill. Weed of roadsides and disturbed places. Near Nike
Base, K 697; Flower Garden near April Brook, K 976; Bitter Cherry Ridge, K April 4, 1965;
east facing slope below Parking Lot, McC June 9, 1963; Ridge Road about % mile east of

Parking Lot, K 870; ravine 12 mile east of Parking Lot, P 2696. Flowers spring and summer.
Native of Europe.

Silene scouleri Hooker subsp. grandis (Eastwood) Hitchcock & Maguire [S. pacifica
Eastwood]. Rare, grassland. East facing slope below Parking Lot, McC, Halde June 23,
1963; Quarry, K July 14, 1963. Perennial with long taproot, and pink flowers in June and
July.

Silene verecunda Watson. Rare, in grassland. Near summit of mountain below radio
towers, W May 11, 1965.

Spergula arvensis Linnaeus. Spurry. Weed of fields and disturbed areas. Crocker
Avenue, K August 5, 1963; Crocker Hills, K January 8, 1964; Colma Canyon, McC June
13, 1965; April Brook, McC March 28, 1965; Ridge Road at West Powerline, McC March
14, 1965; ridge above Brisbane, McC, K September 2, 1963. Flowers in spring and summer.
Native of Europe.

Spergularia macrotheca (Hornemann ex Chamisso & Schlechtendal) Heynhold. Rare,
salt-flat bordering San Francisco Bay. Point San Bruno, McC September 11, 1964; Sierra
Point, W May 8, 1965; Brisbane Lagoon, W June 19, 1965. Flowers May to September.

Spergularia marina (Linnaeus) Grisebach. Sand Spurry. Rare, salt flat bordering
San Francisco Bay or inland. Point San Bruno, K 984; Guadalupe Valley, P 2768. Flowers
spring and summer.

Spergularia rubra (Linnaeus) J. & C. Presl. Sand Spurry. Weed of disturbed places,
often in hard-packed soil. Quarry, K July 14, 1963; east facing slope below Parking Lot,

610 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER.

McC, Halde June 23, 1963; Crocker Hills near Cow Palace, W May 17, 1965; Ridge Road,
K April 25, 1964. Flowers spring and summer. Native of Europe.

Stellaria media (Linnaeus) Cyrillo. Chickweed. Weed of gardens, fields, and disturbed
areas. Guadalupe Road, K August 5, 1963; Quarry, K July 14, 1963; Crocker Hills, K May
4, 1964; Ridge Road, %4 mile east of Parking Lot, AK 875; Randolph Ravine, P 2746. Flowers
in spring and summer. Native of Europe and Asia.

Stellaria nitens Nuttall. Chickweed. Rare, in grassland. Ridge Road at West Powerline,
McC Apri! 25, 1965; Trillium Gulch, McC March 28, 1965.

CHENOPODIACEAE. Goosefoot Family

Atriplex patula Linnaeus var. hastata (Linnaeus) Gray [A. hastata Linnaeus]. Oc-
casional, salt marshes or salt flats, along San Francisco Bay shore or inland. Foothill Boule-
vard at School Street, K October 7, 1963; Crocker Hills, K 2041.

Atriplex rosea Linnaeus. Redscale. Weed, salt flat, Point San Bruno, McC, K September
11, 1964. Native of Eurasia.

Atriplex serenana A. Nelson. Weed, waste ground often forming tangled mats. Brisbane
Lagoon, W August 14, 1965.

Bassia hyssopifolia (Pallas) Kuntze [Echinopsilon hyssopifolium (Pallas) Moquin].
Weed of saline waste places. Brisbane Lagoon, McC, K September 11, 1964. Native of Asia.

Chenopodium album Linnaeus. Lamb’s-Quarters. Pigweed. Weed of waste ground.
Brisbane Lagoon, W June 19, 1965. Native of Europe. The edible young plants are often
cooked as greens.

Chenopodium berlandieri Moquin. Weed of waste ground. Brisbane Lagoon, W June
19, 1965. Introduced from southwestern United States and adjacent Mexico. Characterized
by an unpleasant odor which is lacking in C. album.

Chenopodium californicum (Watson) Watson. Rare, brushy slopes, becoming weedy
in disturbed areas. Lower Colma Canyon, McC, Halde, K June 23, 1963; Pig Ranch Ravine,
W May 11, 1965.

Chenopodium murale Linnaeus. Disturbed ground. Crocker Industrial Park, W June
19, 1965. A common weed throughout North America, unpleasantly scented. Native of
Europe.

Roubieva multifida (Linnaeus) Moquin [Chenopodium multifidum Linnaeus]. Weed
of waste places and roadsides. Lower Colma Canyon, McC, Halde, K June 23, 1963. Native
of Chile.

Salicornia pacifica Standley [S. virginica Linnaeus, in part]. Glasswort. Pickleweed.
Rare, alkaline and coastal salt flats. Point San Bruno, McC, K September 11, 1964. A
perennial forming large colonies, differing from the following which is an annual having
individual plants scattered among the perennial plants of S. pacifica.

Salicornia depressa Standley [S. rubra A. Nelson; S. europaea Linnaeus]. Glasswort.
Rare, alkaline and coastal salt flats. Point San Bruno, McC, K September 11, 1964.

Salsola kali Linnaeus var. tenuifolia Tausch. Russian Thistle. Weed of disturbed
ground. Brisbane Lagoon, W June 19, 1965. A common tumbleweed throughout most of
California. Native of Eurasia.

Vor. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 611

Ficure 5. Anaphalis margaritacea (Linnaeus) Gray.

ComposITar. Sunflower Family

Achillea millefolium Linnaeus var. californica (Pollard) Jepson [A. californica
Pollard]. Yarrow. Milfoil. Frequent, in open grassy or brushy areas. East facing slope
below Parking Lot, McC, Halde June 23, 1963; near Crocker Avenue, Toschi 63:15; Ran-
dolph Ravine, P 2750; Owl’s Canyon, K April 25, 1965; Colma Canyon, K August 5, 1963;
Romanzoffia Ravine, K May 4, 1964. Perennial with finely divided, fern-like leaves and
white flower heads April to July. Our plant is part of a variable species widespread in the
Northern Hemisphere.

Agoseris apargioides (Lessing) Greene. Coast Dandelion. Occasional, in grassland.
Colma Canyon, McC June 13, 1965; near Nike Base, McC, W May 23, 1965; West Power-
line, McC, K September 2, 1963. Flowers spring and summer.

Agoseris grandiflora (Nuttall) Greene. California Dandelion. Occasional, in grassland.
East facing slope below Parking Lot, McC June 9, 1963; near Nike Base, McC, W May 23,
1965; Pig Ranch Ravine, K May 15, 1965; Crocker Hills, K May 4, 1964. Flowers spring
and summer.

Anaphalis margaritacea (Linnaeus) Gray. Pearly Everlasting. Frequent, in grassland,
sometimes forming sizeable colonies. Lower Colma Canyon, McC, Halde, K June 23, 1963;
Flower Garden, McC March 28, 1965; near Nike Base, McC, W May 23, 1965; upper Radio
Road, W May 23, 1965; east facing slope below Parking Lot, McC, Halde June 23, 1963;
West Powerline, McC, K September 2, 1963. Flowers April to September. The pearly-white
flower-heads are showy. Ornamental.

612 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Anthemis cotula Linnaeus. Mayweed. Dog-Fennel. Chamomile. A showy weed, oc-
casionally in fields and roadsides. Lower Colma Canyon, McC, Halde, K June 23, 1963;
Guadalupe Road, K August 5, 1963; Crocker Avenue, W July 7, 1965; Brisbane Lagoon,
W June 19, 1965; Old County Road, Brisbane, W June 8, 1965. Flowers spring through
autumn. Native of Europe.

Artemisia californica Lessing. California Sagebrush. Frequent, on exposed slopes and
ridges. Colma Canyon, McC June 13, 1965; near Olivet Cemetery, W May 18, 1965; slope
immediately south of Parking Lot, McC, Halde June 23, 1963; West Powerline, McC, K
September 2, 1963; upper Buckeye Canyon, K July 14, 1963; Point San Bruno, McC, K
September 11, 1964; Owl’s Canyon, seedlings in burned area, McC March 28, 1965. One of
the shrubs of the coastal scrub, distinguished by its gray-green foliage and its aromatic sage-
like odor. Flowers inconspicuous in summer and early autumn. Regenerates by seeds
after fire.

Artemisia douglasiana Besser. California Mug-Wort. Wormwood. Occasional, on
grassy or brushy areas, sometimes on disturbed ground. Lower Colma Canyon, McC,
Halde, K June 23, 1963; summit of mountain, Raven 1796; Cow Trough Ravine, K October
7, 1963; Brisbane Lagoon, W August 14, 1965, Flowers summer and early autumn.

Aster chilensis Nees. California Aster. Occasional, in brushy areas Cow Trough Ravine,
K October 7, 1963; West Powerline, McC, K September 2, 1963; Point San Bruno, K 747.
Perennial with flowers July to December. Sometimes used as an ornamental for its late
season purplish flowers.

Aster exilis Elliott. Rare, marshy and often alkaline areas. Crocker Hills, K 2037.
Annual with small flower heads in autumn.

Aster radulinus Gray. Rough-leaved Aster. Rare, in brushy areas. Quarry, K July
14, 1963; Buckeye Canyon, K August 5, 1963. Perennial with upper surface of leaves rough
to the touch, flowers July to December.

Aster subspicatus Nees. Occasional, brushy areas. Lower Colma Canyon, McC, Halde,
K June 23, 1963; Radio Road, W May 23, 1965; April Brook, K June 20, 1965; Dairy
Ravine near radio towers, K August 5, 1963; Romanzoffia Ravine, K May 4, 1964; Quarry,
K July 14, 1963. Flowers May to October. Sometimes used as an ornamental.

Baccharis pilularis DeCandolle var. pilularis. Dwarf Chaparral-Broom. Occasional,
on exposed slopes and ridges. Lower Colma Canyon, McC September 29, 1966; Radio Road,
K August 5, 1963. In the low-growing form of the species plants are prostrate to decumbent
often forming mats scarcely more than 12 inches tall. Occasionally used as an ornamental.

Baccharis pilularis DeCandolle var. consanguinea (DeCandolle) Kuntze. Chaparral-
Broom, Coyote Brush. Frequent, on open slopes and in ravines, in chaparral. West Power-
line, McC, K September 2, 1963; Kamchatka Point, K July 27, 1963; Radio Road, McC,
K September 2, 1963; San Bruno Point, McC, K September 11, 1964. The erect form of the
species has plants 5 to 6 feet tall, occurring in the chaparral. Plants near the West Power-
line in an area burned in 1964 produced stump sprouts the following spring. The common
name, Fuzzy-Wuzzy, is sometimes given to these plants when in fruit because of the abundant
dry pappus which blows with the slightest movement of air.

Baeria chrysostoma Fischer & Meyer subsp. chrysostoma. Goldfields. Rare, open slopes
and ridges. Ravine % mile east of 1314-foot summit of mountain, P 2689.

Vou. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 613

Baeria chrysostoma Fischer & Meyer subsp. hirsutula (Greene) Ferris. Goldfields.
Frequent, exposed slopes and ridges. North of Randolph Drive and Hillside Boulevard,
P 2727; West Powerline, K 708; Brisbane Powerline, K 964; Devil’s Arroyo, McC March
28, 1963; top of mountain near radio station, McC, Menzies February 16, 1963; east facing
slope near top of mountain, McC April 17, 1964. Both subspecies are colorful spring wild-
flowers which form dense golden-yellow colonies. Flowers March to May.

Bellis perennis Linnaeus. English Daisy. Weed, along roadside and damp fields. Crocker
Avenue, K September 2, 1964. A common. and attractive, but sometimes troublesome, weed
of lawns, where it is in flower almost throughout the year. Native of Europe.

Blennosperma nanum (Hooker) Blake. Rare, in grassland. Owl’s Canyon, McC, K
April 25, 1965. A colorful spring wildflower with golden-yellow flowers in March and April.

Carduus pycnocephalus Linnaeus. Italian Thistle. Troublesome weed of disturbed
places. Brisbane Lagoon, W June 19, 1965. Flowers May and June. Native of Mediterranean
region.

Carduus tenuiflorus Curtis. Troublesome weed of disturbed places. Sierra Point, W
Apri! 27, 1965. Flowers May to July. Native of Europe.

Centaurea calcitrapa Linnaeus. Purple Star-Thistle. Troublesome weed of disturbed
areas. Crocker Hills, W May 17, 1965; Sierra Point, W June 21, 1965; Brisbane Lagoon,
W August 17, 1965; Cow Trough Ravine, K October 7, 1963. Flowers May to October.
Native of Eurasia.

Centaurea melitensis Linnaeus. Napa Thistle. Tocalote. Troublesome weed of dis-
turbed places. Quarry at end of School Street near Hillside Boulevard, K October 7, 1963;
Colma Canyon, K May 30, 1964. Flowers May to October. Native of Europe.

Centaurea solstitialis Linnaeus. Barneby’s Thistle. Troublesome weed of roadsides
and disturbed areas. Brisbane Lagoon, W June 19, 1965. Flowers June to December. Native
of Europe.

Chaetopappa alsinoides (Greene) Keck. Rare, in grassland. Above Harold Avenue,
Brisbane, McC, W May 14, 1967. Slender inconspicuous annual with few yellowish flowers
in head, April and May, ray flowers lacking.

Chrysanthemum coronarium Linnaeus. Crown Daisy. Garland Chrysanthemum. At-
tractive weed of roadsides and disturbed places. Base of Pig Ranch Ravine, K Apri] 24, 1965;
Crocker Avenue, K August 5, 1963; Sierra Point, W May 8, 1965. Showy yellow flowers,
April to August. Native of Eurasia and Northern Africa.

Chrysopsis villosa (Pursh) Nuttall var. bolanderi (Gray) Gray. Golden Aster.
Frequent, in grassland. Cow Trough Ravine, K October 7, 1963; eastern end of Ridge Road,
K July 14, 1963. Flowers April to November.

Cichorium intybus Linnaeus. Chicory. Weed with blue flowers particularly showy
in mornings, of roadsides and disturbed ground. Crocker Industrial Park, W October 16,
1966. Flowers in summer and autumn. The deep taproot is used as an adulterant or sub-
stitute for coffee. Native of Europe.

Cirsium brevistylum Cronquist. Indian Thistle. Occasional, on brushy slopes. Lower
Colma Canyon, McC, Halde, K June 23, 1963; top of mountain near Parking Lot, McC,
Halde June 23, 1963; Quarry, K July 14, 1963. Flowers April to July.

614 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Cirsium quercetorum (Gray) Jepson. Brownie Thistle. Occasional, in grassland. North
of Randolph Avenue at Hillside Boulevard, P 2703; Quarry, K July 14, 1963; near Nike
Base, K 588. Low-growing plants with pale flowers, April to July.

Cirsium vulgare (Savi) Tenore. Bull Thistle. Weed, in disturbed areas. Lower Colma
Canyon, McC, Halde, K June 23, 1963; Quarry, K July 14, 1963; Brisbane Lagoon, W
August 14, 1965. A coarse thistle with upper surface of leaves rough to touch and purplish
flowers June to October. Native to Eurasia.

Conyza bonariensis (Linnaeus) Cronquist. Weed, in waste ground and _ roadsides.
Crocker Hills in a marshy area between Geneva Avenue and Guadalupe Parkway, K 2042.
Flowers June to September, flower heads larger than those of C. canadensis. Native of
South America.

Conyza canadensis (Linnaeus) Cronquist. Horseweed. Weed of waste ground. Hillside
Boulevard near Serbian Cemetery, K 390; Radio Road, K August 5, 1963; Colma Canyon,
K May 30, 1964. Tall weedy annual with inconspicuous flowers July to October. Widely
distributed in North and South America.

Cotula australis (Sieber) J. D. Hooker. Weed, in disturbed places. Ridge Road, 4
mile east of Parking Lot, K 876; Ridge Road near West Powerline, K 957; Crystal Cave
Canyon, K 1253. Tiny inconspicuous but troublesome annual weed, often forming colonies,
common in gardens. Flowers March to October. Native of Australia.

Cotula coronopifolia Linnaeus. Brass-Buttons. Weed of moist places, sometimes in
salt flats. Point San Bruno, McC, K September 11, 1964; Sierra Point, W May 8, 1965;
ravine south of Crocker Avenue, P 2658; near Crocker Avenue, Toschi 63:33. Strong scented
perennial with yellow flowers in terminal heads, March to October. Native of South Africa.

Erechtites arguta (A. Richard) DeCandolle [Senecio glomeratus Desfontaines ex Poiret].
New Zealand Fireweed. Weed of disturbed places. Crocker Industrial Park, W June 19,
1965; near Nike Base, McC, W May 23, 1965. Coarse annual with pale flowers, June to
August. Native of New Zealand and Australia.

Erechtites prenanthoides DeCandolle [Senecio minimus Poiret]. Australian Fireweed.
Weed of disturbed places. Crocker Hills near Cow Palace, W May 17, 1965; Bitter Cherry
Ravine, K 1109. Coarse annual with lance-shaped, toothed leaves which distinguish it from
E. arguta in which leaves are deeply divided. Pale flowers, July to September. Native of
Australia and New Zealand.

Erigeron foliosus Nuttall. Rare, in grassland. Buckeye Canyon, W July 8, 1965. Flowers
June and July.

Erigeron glaucus Ker-Gawler. Seaside Daisy. Frequent, on wind-swept slopes and
ridges. East facing slope below Parking Lot, McC, Halde June 23, 1963; West Powerline,
McC, K September 2, 1963; Trillium Gulch, K 1016; Pig Ranch Ravine, W May 12, 1965.
Attractive low-growing perennial with somewhat thickish smooth leaves, and showy lavender
flowers, March to August. Ornamental.

Erigeron philadelphicus Linnaeus. Philadelphia Daisy. Rare, in wet ground. April
Brook, where creek joins Colma Canyon, W June 16, 1965; Radio Road, W May 23, 1965.
Slender perennial, attractive flower heads with over 150 slender lavender ray flowers, April
to June. Ornamental.

PLATE 1. Upper. Erigeron glaucus Ker-Gawler.
Lower. Sidalcea malvaeflora (DeCandolle) Gray ex Bentham.

( McCLINTOCK

AND PLATE 1

ACAD. SCI., 4TH SER., VOL.XXXII NO. 20

CALIF.

PROC.

KNIGHT)

Vor. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 615

Eriophyllum confertiflorum (DeCandolle) Gray. Yellow Yarrow. Occasional, on
brushy slopes. East facing slope below Parking Lot, McC, Halde June 23, 1963; Quarry,
K July 14, 1963; canyon southwest of radio towers, W May 12, 1965; above Olivet Cemetery,
McC June 13, 1965. Shrub, usually 1-foot tall, with narrowly linear leaf divisions and many
yellow flower heads in showy clusters, May to November. Ornamental, it stays within
bounds when planted and its gray-green finely divided foliage offers a pleasing contrast
with its golden-yellow flower heads.

Eriophyllum staechadifolium Lagasca. Lizard-Tail. Occasional, in brushland on
western part of mountain. Quarry, K July 14, 1963; Radio Road, McC, K September 2,
1963; slope below Parking Lot, McC June 9, 1963. Shrub usually 2 feet or more tall, with
lanceolate leaf divisions and many yellow flower heads in clusters, May to November and
often during remainder of year. Distinguished from preceding species by its leaves which
have broader divisions and a more coarse appearance than those of EF. confertiflorum.

Filago californica Nuttall. Occasional, in grassland and brushy areas. A ridge above
Brisbane, + 800-foot elevation, Raven 1803; South Powerline, McC March 14, 1965. Flowers
March and April.

Filago gallica (Linnaeus) Linnaeus. Occasional, in grassland, and disturbed areas. Lower
Colma Canyon, McC, Halde, K June 23, 1963; slope below Parking Lot, McC June 9, 1963.
Weed of gardens and other disturbed areas. Native of Europe.

Franseria chamissonis Lessing subsp. bipinnatisecta (Lessing) Wiggins & Stockwell.
Beach-Bur. Rare, beaches and salt flats. San Bruno Point, A September 2, 1964, Flowers
June to October.

Gnaphalium beneolens A. Davidson, Everlasting. Cudweed. Rare, brushy slopes.
Brisbane Ravine, K August 5, 1963. A perennial, flowers July to November.

Gnaphalium californicum DeCandolle. Green Everlasting. California Cudweed. Fre-
quent, on dry brushy slopes and ridges. East facing slope below Parking Lot, McC, Halde
June 23, 1963; slopes southeast of 1314-foot summit, P 2671; West Powerline, McC, K
September 2, 1963; Radio Road, K August 5, 1963; Sierra Point, W May 8, 1965, Strongly
scented biennial, leaves green not gray, flowers May to August.

Gnaphalium chilense Sprengel. Everlasting. Cudweed. Frequent, brushy areas at
margin of chaparral. Colma Canyon, K May 30, 1964; Brisbane Lagoon, W June 19, 1965;
Pig Ranch Ravine, W May 12, 1965; Crystal Cave Canyon, K June 20, 1965; Quarry, K July
14, 1963; Radio Road, K August 5, 1963. Gray woolly annual or biennial, flowers through-
out the year.

Gnaphalium luteo-album Linnaeus. Everlasting. Cudweed. Weed of disturbed areas.
Crocker Hills, K 2045; Radio Road, K August 5, 1963; Colma Canyon, McC, W June 13,
1965; Guadalupe Valley, P 2763; in coastal scrub, Thomas 10064. Gray woolly annual or
biennial, flowers throughout year. Native of Europe.

Gnaphalium purpureum Linnaeus. Purple Cudweed. Rare, dry disturbed areas.
Slope below Parking Lot, McC June 9, 1963; Pig Ranch Ravine, W May 12, 1965. White
woolly annual or biennial, flowers April to June.

Gnaphalium ramosissimum Nuttall. Pink Everlasting. Rare, open or brushy slopes.
Radio Road, McC, K September 2, 1963; Horseshoe Ridge, K August 5, 1963; Great Meadow

616 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

adjacent to upper Colma Canyon, W August 16, 1965. Sweet-scented, green-leaved biennial
with pink flowers, June to October. The most attractive of all of the Everlastings.

Grindelia camporum Greene. Gum-Weed. Occasional, on dry open slopes and ridges.
North city limits of South San Francisco, McC, K September 11, 1964; Brisbane Lagoon,
W August 14, 1965; Radio Road, McC, K September 2, 1963; Quarry, K July 14, 1963;
Buckeye Canyon, K July 14, 1963; Ridge Road, W July 8, 1965. Subshrub, leaves resinous,
yellow flower heads, June to December.

Grindelia hirsutula Hooker & Arnott. Occasional, on dry open or brushy slopes. Ravine
north of Randolph Drive at Hillside Boulevard, P 2736; Sierra Point, W April 27, 1965;
slope southeast of 1314-foot summit, P 2666; Quarry, K June 24, 1963; near radio stations,
McC, Menzies February 16, 1963. Flowers April to June.

Grindelia humilis Hooker & Arnott. Rare, in salt flats. Brisbane Lagoon, W June
19, 1965; near Bayshore Highway north of South San Francisco, K November 8, 1963.
Flowers throughout year. Sometimes used as an ornamental; fast growing and large yellow
flower heads are showy.

Grindelia maritima (Greene) Steyermark. Rare, in association with coastal scrub,
J. H. Thomas 9709. Perennial with yellow flowers August and September.

Haplopappus ericoides (Lessing) Hooker & Arnott. Mock-Heather. Rare, in sandy
soil on brushy slope. Lower Colma Canyon, McC September 29, 1966. Low shrub with
heather-like leaves and yellow flower heads September to November.

Helenium puberulum DeCandolle. Sneezeweed. Rosilla. Occasional, moist places or
along creeks. Wax Myrtle Ravine, K August 5, 1963; April Brook, K, J. Roof August 4,
1963; Great Meadow, W July 7, 1965. Biennial or shortlived perennial, yellow flower heads
with dome-shaped disk surrounded by short, reflexed rays, April to November.

Helianthella castanea Greene. Rare, in grassland. Sierra Point, W April 27, 1965.
Perennial herb with large leaves and sunflower-like flower heads, April and May. Superfi-
cially resembles Wyethia. Endemic to San Francisco Bay region.

Hemizonia corymbosa (DeCandolle) Torrey & Gray. Tarweed. Occasional, in grass-
land. Colma Canyon, McC June 13, 1965; Pig Ranch Ravine, K May 1965; abandoned
quarry at end of School Street near Hillside Boulevard, K October 7, 1963; Sierra Point,
W April 27, 1965. Annual with lower pinnatifid leaves in spring, leaves becoming entire
on older plants, pale yellow flower heads, May to November.

Hemizonia pungens (Hooker & Arnott) Torrey & Gray subsp. maritima (Greene)
Keck. Spikeweed. Rare. Crocker Avenue, McC September 24, 1967. Erect stiffly branched
annual to 12 inches tall, yellow flower heads, late summer to early autumn.

Heterotheca grandiflora Nuttall. Telegraph Weed. Rare, in disturbed places. Crocker
Industrial Park, W October 16, 1966. Tall weedy annual or biennial with grayish foliage
and yellow flower heads March to October. Native of interior central and southern Cali-
fornia, introduced as a weed in our area.

Hypochaeris glabra Linnaeus. Smooth Cat’s-Ear. Weed, in disturbed places. Colma
Canyon, McC June 13, 1965; north of Randolph Drive at Hillside Boulevard, P 2721. An-
nual weed with basal leaves and yellow flower heads, February to August. Native of
Eurasia and North Africa.

Vout. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 617

Hypochaeris radicata Linnaeus. Hairy Cat’s-Ear. Weed of disturbed places and grass-
lands. Lower Colma Canyon, McC, Halde, K June 23, 1963; east facing slope below Parking
Lot, McC, Halde June 23, 1963; Ridge Road at eastern end, K July 14, 1963; near Nike
Station, McC May 25, 1965. Perennial with taproot, basal leaves, and yellow flower heads,
March to September. Native of Eurasia and North Africa.

Jaumea carnosa (Lessing) Gray. Rare, in salt flat at edge of San Francisco Bay. Point
San Bruno, McC, K September 2, 1964. Fleshy perennial with yellow flowers, June to
November.

Lactuca saligna Linnaeus. Willow-Lettuce. Weed of disturbed ground. Cow Trough
Ravine, K October 7, 1963. Native of Eurasia.

Lactuca serriola Linnaeus. Prickly Lettuce. Weed of disturbed ground. Pig Ranch
Ravine, W May 11, 1965. Prickly-leaved annual, flowers May to November. Native of
Eurasia and North Africa.

Lactuca virosa Linnaeus. Wild Lettuce. Weed of disturbed ground. Ridge Road near
West Powerline, K August 5, 1963. Tall biennial to 6 to 8 feet, with stems hollow at least
below, flowers August and September. Native of Eurasia and North Africa.

Lagophylla ramosissima Nuttall. Hareleaf. Rare, known from a single collection in
“San Bruno Hills,” J. McMurphy June 13, 1909.

Layia chrysanthemoides (DeCandolle) Gray. Rare, known from a single collection in
“San Bruno Hills,’ J. McMurphy June 13, 1909.

Layia gaillardioides (Hooker & Arnott) DeCandolle. Rare, in coastal scrub, J. H.
Thomas 9307. Annual with showy golden-yellow flowers, April to June.

Layia hieracioides (DeCandolle) Hooker & Arnott. Frequent, in chaparral. Colma
Canyon, K August 5, 1963; Radio Road, McC, K September 2, 1963; Dairy Ravine, K
August 5, 1963; Quarry, K July 14, 1963; Devil’s Arroyo, seedlings in area burned over in
previous autumn, McC March 28, 1965. Pungent, hairy annual, with lobed leaves and rela-
tively small flowers, April to August.

Layia platyglossa (Fischer & Meyer) Gray subsp. campestris Keck. Tidy-Tips. Rare,
in grassland and chaparral. Colma Canyon, Raven 1780; Ridge Road at West Powerline,
K 636. Hairy annual with pinnately lobed leaves and showy pale yellow flower heads,
March to June.

Madia capitata Nuttall. Tarweed. Occasional, in grassland, and disturbed areas. Colma
Canyon, K 607; April Brook, K May 15, 1965; Pig Ranch Ravine, W May 11, 1965; Radio
Road, K August 5, 1963; east facing slope below Parking Lot, McC, Halde June 23, 1963;
summit of mountain, Raven 1794. Pungent, glandular-hairy annual with flower heads sessile
or shortly stalked, May to September.

Madia exigua (Smith) Gray. Small Tarweed. Rare, in grassland. Ridge Road at West
Powerline, K 625. Low growing, branched annual, flowers May and June. The small flower
heads with only one or two disk-flowers per head and the slender peduncles distinguish
this species.

Madia gracilis (Smith) Keck. Slender Tarweed. Rare, in grassland. Above Olivet
Cemetery, McC June 13, 1965. Glandular-viscid annual, flowers May to September.

618 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Madia sativa Molina. Coastal Tarweed. Frequent, in grassland, roadsides, and disturbed
areas. Becoming weedy. Lower Colma Canyon, McC, Halde, K June 23, 1963; Radio Road,
McC, K September 2, 1963; West Powerline, McC, K September 2, 1963; Quarry, K July 14,
1963; Devil’s Arroyo, K 1094, Stout, pungent, glandular-hairy annual with flower heads
clustered in upper leaves, June to November.

Matricaria matricarioides (Lessing) Porter. Pineapple-Weed. Weed of disturbed
areas, often in dry hard-packed soil. Crocker Hills, K May 4, 1964; Sierra Point, W May
8, 1965; Crocker Industrial Park, W June 19, 1965. Aromatic annual with finely pinnatifid
leaves and conical yellow flower heads, March to June.

Micropus californicus Fischer & Meyer. Slender Cottonweed. Rare, margin of chap-
arral. Canyon southwest of the National Broadcasting Company radio tower, W May 11,
1965. White woolly annual, inconspicuous flowers, April to June.

Microseris douglasii (DeCandolle) Schultz-Bipontinus subsp. douglasii. Rare, in grass-
land. Thomas 9259.

Microseris douglasii (DeCandolle) Schultz-Bipontinus subsp. tenella (Gray) Chambers.
Rare, in grassland. Buckeye Canyon, W April 27, 1965. Both subspecies are annuals lack-
ing a stem with leaves basal, yellowish to pale apricot-co'ored flowers on a stalk, April
and May.

Picris echioides Linnaeus. Bristly Ox-Tongue. Weed of disturbed places. Quarry, K
June 24, 1963; Colma Canyon, K August 5, 1963; Brisbane Lagoon, W June 19, 1965. Rough,
bristly annual or biennial, yellow flowers, April to December. Native of Mediterranean region.

Psilocarphus tenellus Nuttall. Slender Woolly-Heads. Occasional, in dry bare, often
hard packed, ground, forms colonies and mats, sometimes becomes weedy. Guadalupe Road,
McC April 11, 1965; Quarry, K June 24, 1963; Ridge Road, W June 19, 1965. Low-growing
grayish annual, flowers March to May.

Rafinesquia californica Nuttall. California Chicory. Occasional, on grassy or brushy
areas. Thomas 9294.

Senecio aronicoides DeCandolle. Butterweed. Frequent, in grassland and border of chap-
arral. Colma Canyon, K March 15, 1964; east slope of mountain below radio station,
Toschi 63:68; top of mountain, Raven 1375; 300 yards east of Parking Lot on area burned
in autumn 1964, K 869; Romanzoffia Ravine, K May 4, 1964; ravine 1% mile east of 1314-foot
summit, P 2690; South Powerline in area burned in autumn 1964, McC March 14, 1965. Tall
coarse perennial with yellow flower heads, April to June.

Senecio cruentus (L’Heritier) DeCandolle. Florist’s Cineraria. Garden Cineraria.
Escapes from cultivation in moist, shaded areas. Top of mountain near KRON radio station,
W June 20, 1965. Perennial with large purple ray flowers, summer. Native of Canary Islands.

Senecio elegans Linnaeus. Purple Ragwort. Escape from cultivation. Colma Canyon,
W June 16, 1965. Annual with lavender ray flowers, May and June. Native of South Africa.

Senecio mikanioides Otto. German Ivy. Weed of moist areas along roadsides. Crocker
Avenue, McC, K January 12, 1964. Evergreen perennial twining on other plants, bright
yellow flower heads, winter and spring. Native of South Africa.

Senecio sylvaticus Linnaeus. Weed in brushy areas. East facing slope below Parking
Lot, McC, Halde June 23, 1963; Ridge Road about 100 yards east of Parking Lot, K May
19, 1964. Coarse annual weed with pale flower heads, April to June. Native of Europe.

Vout. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 619

Senecio vulgaris Linnaeus. Common Groundsel. Weed of disturbed areas. East facing
slope below Parking Lot, McC April 17, 1964; South Powerline, McC March 14, 1965. An-
nual with pale flower heads, spring-summer. Black-tipped involucral bracts distinguish it
from S. sylvaticus. Native of Europe.

Silybum marianum (Linnaeus) Gaertner. Milk-Thistle. Weed of disturbed places and
roadsides. Lower Colma Canyon, McC, Halde, K June 23, 1963; Crocker Hills, K May
4, 1964. Coarse annual or biennial with spiny mottled leaves and purple flower heads. Native
of Mediterranean region.

Solidago californica Nuttall. California Golden-Rod. Occasional, dry open areas. West
Powerline, McC, K September 2, 1963; Cow Trough Ravine, K October 7, 1963; April
Brook, W August 16, 1965. Perennial with creeping rhizome and hairy leaves somewhat
rough to touch above, flowers July to November.

Solidago canadensis Linnaeus subsp. elongata (Nuttall) Keck. Meadow Goldenrod.
Occasional, on brushy areas. East facing slope below Parking Lot, McC, Halde June 23,
1963; Colma Canyon, K March 15, 1964. Differs from S. californica in its glabrous or only
slightly hairy leaves, flowers August to November.

Solidago spathulata DeCandolle. Dune Goldenrod. Frequent, on brushy areas. East
facing slope below Parking Lot, McC, Halde June 23, 1963; top of mountain near radio
towers, McC, Menzies February 16, 1963; West Powerline, McC September 2, 1963; lower
Colma Canyon, McC September 29, 1966; Crocker Hills, K August 5, 1963. Differs from
two preceding goldenrods in the shining, more or less glutinous inflorescence, flowers August
to October.

Soliva sessilis Ruiz & Pavon. Weed of roadsides and other disturbed areas. West Power-
line, K April 13, 1964; Crocker Hills near Cow Palace, W May 17, 1965; Randolph Avenue,
K May 11, 1967. Small depressed annual weed, flowers January to June. Native of Chile.

Sonchus asper (Linnaeus) Hill. Prickly Sow-Thistle. Weed of roadsides and other dis-
turbed areas. East facing slope below Parking Lot, McC, Halde June 23, 1963; lower Colma
Canyon, McC, Halde, K June 23, 1963; Radio Road, K August 5, 1963. Weedy annual,
flowers March to July. Native of Eurasia and North Africa.

Sonchus oleraceus Linnaeus. Sow Thistle. Weed of roadsides and other disturbed
areas. Brisbane Lagoon, W June 19, 1965; north of Randolph Drive at Hillside Boulevard,
P 2724; Horseshoe Ridge, K August 5, 1963. Weedy annual, flowers March to September.
Native of Europe.

Stephanomeria virgata Bentham. Rare, open rocky slope. South side of Ridge Road,
about 200 yards south of Parking Lot, McC, K August 1, 1967. Stiff annual with almost
leafless stems and inconspicuous pink or white flowers in summer.

Tanacetum camphoratum Lessing. Dune Tansy. Rare, known from only one locality
on the mountain. Distribution restricted to the San Francisco area. Radio Road, K August
5, 1963. Herbaceous perennial with gray fern-like leaves and showy yellow flowers, June
to October. Ornamental.

Taraxacum laevigatum (Willdenow) DeCandolle. Red-Seeded Dandelion. Weed of
disturbed areas. Flower garden adjacent to April Brook, McC March 28, 1965; West Power-
line, McC April 25, 1965. Flowers March to July. Native of Eurasia.

620 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Tragopogon porrifolius Linnaeus. Oyster-Root. Salsify. Weed of disturbed areas.
Crocker Industrial Park, W June 19, 1965. Perennial with long taproot and purple flowers,
March to October. Native of Mediterranean region. Sometimes grown for its edible root
called vegetable-oyster.

Wyethia angustifolia (DeCandolle) Nuttall. Mule-Ears. Occasional, in grassland.
Crocker Hills, K May 4, 1964; Point San Bruno, K April 24, 1965; Sierra Point, W April
27, 1965; above Harold Avenue, Brisbane, McC, W May 14, 1967; Glen Park Canyon, McC
April 16, 1967. Perennial herb with long narrow leaves and showy sunflower-like flower
heads April and May. Similar in appearance to Helianthella castanea from which it differs
in technical characters of flower and fruit.

Xanthium spinosum Linnaeus. Spiny Clotbur. Spanish Thistle. Weed of disturbed
places. Crocker Industrial Park, W June 19, 1965. Coarse spiny annual, inconspicuous
flowers, April to November. Nativity doubtful, has been reported from both North and
South America, as well as from Europe. Because of the aggressive weedy nature of this
plant it is difficult to determine where it may have originated.

CONVOLVULACEAE. Convolvulus Family

Convolvulus arvensis Linnaeus. Field Bindweed. Orchard Morning-Glory. Deep-rooted,
troublesome weed of fields and waste places. Lower Colma Canyon, W June 16, 1965; inter-
section of Radio Road with Guadalupe Parkway, K August 5, 1963; Quarry, W June 8,
1965. Flowers May to October. Introduced from Europe.

Convolvulus occidentalis Gray. Western Morning-Glory. Occasional, on brushy slopes,
often scrambling over other plants. Colma Canyon, K May 4, 1964; above Olivet Cemetery,
McC June 13, 1965; Randolph Drive at Hillside Boulevard, P 2729; east facing slope below
Parking Lot, McC, Halde June 23, 1963; upper Buckeye Canyon, K July 14, 1963. Flowers
April to July. May be distinguished from preceding by its larger flowers, about 1% to 14%
inches long, whereas, flowers in C. arvensis are about *4 inch long.

Convolvulus subacaulis (Hooker & Arnott) Greene. Occasional, in grassland. Crocker
Hills near Cow Palace, W May 17, 1965; Colma Canyon, W June 16, 1965; ravine east of
Quarry, W June 19, 1965; Wax Myrtle Ravine, K August 5, 1963; north facing slope south
of Guadalupe Parkway close to junction with Radio Road, McC, W June 11, 1967. Flowers
April to June. Differs from two preceding morning-glories by its short stem and its larger
flowers, from 14% to 2 inches long.

Dichondra donelliana Tharp & Johnston. Rare, known only from a slope above Olivet
Cemetery. Above Olivet Cemetery, W May 18, 1965. The gray-green foliage of this species
distinguishes it from D. micrantha Urban (usually referred to as D. repens Forster) which
is cultivated as a ground cover and occasionally escapes to become naturalized. Dichondra
donelliana is known from scattered localities in southern Oregon, northern and central Cali-
fornia, and Santa Cruz Island off the coast of southern California. Dichondra, a poorly
understood genus of about 14 species mostly in North and South America, New Zealand,
and Australia, is best known through D. micrantha (sometimes called D. repens) the much
used groundcover. D. donelliana has until recently been known as D. repens, a name which
applies to a plant in New Zealand and Australia. Its identity was established in 1961 when

it was described as a new species and named for the South American botanist, Professor
Carlos O’Donell.

Vou. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 621

CorNACEAE. Dogwood Family

Cornus californica C. A. Meyer. Creek Dogwood. Western Red Dogwood. Rare, in
moist canyons and ravines, sometimes forming thickets. Quarry, K July 14, 1963; lower
Colma Canyon along creek, McC, Halde, K June 23, 1963. However, at the present time
(1967) it may no longer exist at these localities. Clusters of white flowers, April to June.
Leaves turn red in autumn before falling and the slender bare reddish-brown branches are
showy in spring before the new leaves appear. Ornamental.

CRASSULACEAE. Stone Crop Family

Dudleya cymosa (Lemaire) Britton & Rose. Bluff Lettuce. Scattered on rocky slopes
and ledges. Flowers April to July.

Dudleya farinosa (Lindley) Britton & Rose. Scattered on rocky slopes and ledges.
East facing slope below Parking Lot, McC, Halde June 23, 1963. Flowers July and August.
Distinguished from preceding species by the leaves being farinose or white-mealy. Both
species of Dudleya are used as ornamental succulent plants.

Sedum spathulifolium Hooker. Stone Crop. Scattered on rocky and grassy slopes and
ledges. Ravine east of Quarry, W July 8, 1965; east facing slope below Parking Lot, McC,
Halde June 23, 1963. Flowers April to July. Useful as an ornamental succulent plant.
Sedum may be distinguished from Dudleya when in flower by its spreading and separate
petals. In Dudleya the petals are united to form a short tubular corolla.

Tillaea erecta Hooker & Arnott. Pigmyweed. Occasional, in open places in grassland
and edge of chaparral, often in pure colonies on exposed ground. Slope above Brisbane,
Raven 1371; Crocker Hills near Guadalupe Road, McC April 11, 1965; on ridge near West
Powerline, K 993. A tiny inconspicuous annual, only an inch or two tall. Close examination
with a hand-lens, however, reveals its minute but fleshy opposite leaves and tiny, sedum-like
flowers April and May.

CRUCIFERAE. Mustard Family

Arabis blepharophylla Hooker & Arnott. Coast Rock Cress. Frequent, grassy slopes
and rocky outcroppings. Near Crocker Avenue, P 2654; Cable Ravine, McC March 14, 1965;
Powerlines, K 967; Owl’s Canyon, K April 25, 1965; Colma Canyon, K 1226; slope southeast
of 1314 foot summit, P 2672; slope above Brisbane, Raven 1378. Early spring-flowering
perennial with attractive rose-purple flowers, March to May. Ornamental.

Arabis glabra (Linnaeus) Bernhardi. Tower Mustard. Occasional, grassy and brushy
slopes. Colma Canyon, K 1217; Flower Garden near April Brook, McC March 28, 1965;
West Powerline, Menzies March 14, 1965; summit of mountain, Raven 1799; Quarry, K
July 14, 1963; South Powerline Ridge, Menzies March 14, 1965; near radio station, McC,
Menzies February 16, 1963. Weedy in appearance with whitish to yellow flowers, February
to June.

Barbarea orthoceras Ledebour. American Winter Cress. Occasional, open or brushy
areas. Manzanita Dike, K 933; meadow between April Brook and Radio Road, McC March
28, 1965; Eucalyptus grove on Radio Road, K 861; east facing slope below radio station,
Toschi 63:63; Sierra Point, W April 27, 1965. Pale yellow flowers, March to July.

622 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Barbarea verna (Miller) Ascherson. Early Winter Cress. Weed, in disturbed areas.
Colma Canyon, K May 30, 1964; near Crocker Avenue, P 2651; east facing slope below
summit, McC April 17, 1964; ravine 42 mile east of summit, P 2697. Yellow flowers, March
to July. Native of Europe.

Brassica campestris Linnaeus. Field Mustard. Weed, disturbed areas. Near Crocker
Avenue, P 2662; Guadalupe Valley, P 2766; Colma Canyon, K 1236; Brisbane Lagoon, W
June 19, 1965. The bright yellow flowers are colorful in early spring, February to July.
Native of Europe.

Brassica geniculata (Desfontaines) J. Ball. Mediterranean Mustard. Summer Mustard.
Weed, in disturbed areas. Radio Road, K August 5, 1963; Quarry, K July 14, 1963; Guada-
lupe Valley, P 2770; Colma Canyon, McC, Halde, K June 23, 1963; Romanzoffia Ravine,
K May 4, 1964; base of Owl’s Canyon, W June 7, 1965; Point San Bruno, K, McC Septem-
ber 11, 1964. Yellow flowers mostly in summer, May to September. Native of Mediterranean
region.

Brassica kaber (DeCandolle) L. C. Wheeler. Charlock. Weed along roadsides and in
waste ground. Point San Bruno, K April 24, 1964. Flowers March to October. Native of
Europe.

Brassica oleracea Linnaeus. Cabbage. Escape from cultivation. Brisbane Lagoon, W
June 19, 1965. Sometimes persisting in maritime situations, flowers in spring and summer.
Native of western and southern Europe.

Cakile maritima Scopoli. Sea-Rocket. Occasional, as a weed on sand dunes and beaches.
Point San Bruno, McC, K September 11, 1964. Flowers lavender to purplish, March to
September. Native of Europe.

Capsella bursa-pastoris (Linnaeus) Medicus. Shepherd’s Purse. Weed of disturbed
areas and on open grassy slopes. West Powerline, K 734. Small whitish flowers throughout
year. Native of Europe.

Cardamine oligosperma Nuttall. Bitter Cress. Occasional, in grassland and disturbed
areas, sometimes in areas previously burned. Colma Canyon, K May 30, 1964; ravine 4% mile
east of summit, P 2686; Romanzoffia Ravine, K March 14, 1965; Trillium Gulch, K 1018.
Annual with small white flowers and seed pods which open explosively, often becoming
weedy, flowers February to June.

Cardaria draba (Linnaeus) Desvaux. Hoary Cress. Weed, in disturbed ground. Crocker
Hills near old Bayshore Highway, W May 7, 1967. Erect perennial with numerous tiny white
flowers in terminal panicles, March to December. Native of Europe.

Coronopus didymus (Linnaeus) J. E. Smith. Wart-Cress. Weed of waste and culti-
vated ground and along roadsides, sometimes in hard packed soil. Colma Canyon, K May
30, 1964; Crocker Hills near Cow Palace, W May 17, 1965; Guadalupe Road, McC April 11,
1965; east facing slope below Parking Lot, McC, Halde June 23, 1963; Ridge Road east
of Parking Lot, K 699. Much branched, strong-smelling annual with tiny flowers, March to
November. Native of Europe.

Dentaria californica Nuttall var. integrifolta (Nuttall) Detling. Toothwort. Rain-
Bells. Milk-Maids. Occasional, open grassy or brushy slopes. Slope above Brisbane, Raven
1380; Kamchatka Point, K January 8, 1964; Colma Canyon, K March 135, 1964; Radio

=
PLATE 2. Upper. Dudleya farinosa (Lindley) Britton & Rose.
Lower. Diplacus aurantiacus (Curtis) Jepson.

(McCLINTOCK
PROC. CALIF. ACAD. SCI., 4TH SER., VOL.XXXII NO. 20 AND PLATE 2
KNIGHT)

an toeee 7"

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fi cw steeaew 5 eco
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Bt as oe vial

Vou. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 623

Road, McC, V. Ryder February 16, 1964; near radio station close to summit, McC, Menzies
February 16, 1963; Cable Ravine, McC March 14, 1965; Trillium Gulch, McC March 28,
1965; ravine 4% mile east of 1314-foot summit, P 2698. Perennial with an underground
thickened stem or rhizome from which in late winter arise 3-foliolate leaves. The slender
erect stems have 3 to 5-foliolate leaves and pretty white to pink flowers, January to May.

Erysimum franciscanum Rossbach. Franciscan Wallflower. Occasional, open, rocky,
or grassy slopes. Quarry, K July 14, 1963, and W July 8, 1965; Buckeye Canyon, K July
14, 1963; below top of mountain, McC June 9, 1963; lower Colma Canyon, McC, Halde, K
June 23, 1963; Randolph Avenue at Hillside Boulevard, P 2735; South Powerline, McC
March 14, 1965. Attractive showy spring wildflower, a biennial or short-lived perennial,
leggy in habit, with terminal clusters of pale yellow or cream-white flowers, March to May.
Ornamental.

Lepidium nitidum Nuttall. Shining Pepper-Grass. Occasional, open rocky areas and
grassland. Side spur of San Bruno Mountain west of Brisbane, Raven 1367; Romanzoffia
Ravine, K March 14, 1965; West Powerline, K, McC March 14, 1965; Crocker Avenue, K
1201; Colma Canyon, K 1248; slope below top of mountain near radio station, McC, Menzies
February 16, 1963; Randolph Avenue, South San Francisco, K May 16, 1967. Erect annual
with tiny whitish flowers January to May, and small ovate fruits having a peppery taste.

Lepidium pinnatifidum Ledebour. Russian Pepper-Grass. Weed of disturbed ground.
Brisbane Lagoon, W June 19, 1965; Randolph Avenue, South San Francisco, K May 16,
1967. Annual or biennial with tiny flowers in spring. Native of southern Russia.

Lobularia maritima (Linnaeus) Desvaux [Alyssum maritimum (Linnaeus) Lamarck.
Koniga maritima (Linnaeus) R. Brown]. Sweet Alyssum. Weedy escape from cultivation
in disturbed ground. Crocker Avenue, K August 5, 1963; Reservoir Hill, McC March 14,
1965. Low much branched perennial with often a profusion of small white flowers through-
out the year. Native of Europe.

Lunaria annua Linnaeus. Moonwort. Honesty. Money-Plant. Weedy escape from
cultivation in moist shaded areas. Radio Road, Thomas 10632. Erect branched annual or
biennial with attractive purple flowers, March to June, followed by rounded to oval paper-
thin fruits resembling a silver coin and used in dry arrangements. Native of Europe.

Raphanus sativus Linnaeus. Wild Radish. Weed of disturbed areas. Guadalupe Road,
K August 5, 1963; ravine south of Crocker Avenue, P 2663; ridge above Brisbane, Mcc,
K September 2, 1963. Tall branched annual or biennial with showy clusters of flowers in
varying shades of white, yellowish, to pale and darker lavender, February to May. Native
of Europe. The edible radishes are selected forms of the species.

Rorippa nasturtium-aquaticum (Linnaeus) Schinz & Thellung [Nasturtium officinale
R. Brown]. Water Cress. Occasional, in shallow water or margins of streams. Cow Trough
Ravine, K October 7, 1963; Crocker Avenue, K August 5, 1963; Quarry, K July 14, 1963;
lower Colma Canyon, McC, Halde, K June 23, 1963; Devil’s Arroyo, McC March 28, 1965;
Sierra Point, W April 27, 1965. Leafy aquatic perennial with white flowers March to Novem-
ber. Native of the Old World. Cultivated for its edible leaves.

Sisymbrium officinale (Linnaeus) Scopoli. Hedge Mustard. Weed of waste ground.
Crocker Avenue, K August 5, 1963; Colma Canyon, K May 4, 1964; Romanzoffia Ravine,
K May 4, 1964; Crocker Hills, K May 4, 1964. Coarse stiffly branched annual with clusters
of small yellowish flowers, February to August, and elongate fruits closely appressed to stem.
Native of Europe.

624 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Ficure 6. Marah fabaceus (Naudin) Greene.

Sisymbrium orientale Linnaeus. Weed of waste ground. Guadalupe Road, K April
11, 1965. Coarse, branched annual or biennial, distinguished from Szsymbrium officinale by
its spreading fruits. Native of Europe.

Thelypodium lasiophyllum (Hooker & Arnott) Greene var. inalienum Robinson.
Occasional, open and brushy slopes in undisturbed areas. Crocker Hills, K May 4, 1964;
South Powerline, Menzies March 14, 1965; Devil’s Arroyo, K 1090, and K 1103; Guadalupe
Road, McC April 11, 1965. Tall, erect, annual which in this form of the species has the
slender fruits spreading away from the stem.

Thysanocarpus curvipes Hooker. Fringe-Pod. Rare, in grassland. Owl’s Canyon,
K May 1, 1967. Slender, branched annual with tiny whitish flowers, March to May, and a
small rounded fruit sometimes with perforate margin.

CUCURBITACEAE. Gourd Family

Marah fabaceus (Naudin) Greene. Manroot. Wild Cucumber. Occasional, in brushland
climbing on shrubs or trailing over the ground. Slope above Crocker Avenue, Toschi 63:27;
Colma Canyon, K May 30, 1964; lower Colma Canyon, McC, Halde, K June 9, 1963; above
the Quarry, K July 14, 1963; ravine east of West Powerline, McC March 28, 1965; slope
below summit of mountain, McC June 9, 1963. Flowers cream-white, corolla more or less
rotate, that is having a tube and spreading lobes, February to May, and mature fruits with
many stiff spines. This species and the following, Marah oregonus, are similar in appearance.
Both have herbaceous stems arising annually from a large perennial tuber, hence the common
name of Manroot. They may be distinguished by the shape of their corollas and the num-
ber and stiffness of the spines on the fruits.

Vou. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 625

Ficure 7. Arctostaphylos imbricata Eastwood.

Marah oreganus (Torrey & Gray) T. J. Howell. Manroot. Wild Cucumber. Occasional,
in brushland climbing on shrubs or trailing over the ground. Dairy Ravine, K August 5,
1963; west facing slope below summit of mountain, McC April 17, 1964; slopes southwest
of Brisbane, P 2760; Devil’s Arroyo, K 1060. Flowers whitish, corollas more or less campanu-
late, March to May, and mature fruits with few weak spines.

DrpsacacEaE, Teasel Family

Dipsacus fullonum Linnaeus [D. sativus Honckeny]. Fuller’s Teasel. Weed along
roadsides and in disturbed areas. Crocker Industrial Park, W March 27, 1966. Tall stout
biennial herb with terminal oblong flower clusters, of which the spine-like bracts are con-
spicuous and surpass the flowers, which appear in spring and summer. Has been cultivated
in Europe for fulling in textile mills, hence the common name. Native of Europe.

Scabiosa atropurpurea Linnaeus. Pincushion Plant. Escape from cultivation along
roadsides. Humboldt Road, Brisbane, K November 8, 1963. Attractive garden annual with
terminal flower heads, flowers in whites, pinks, to purples throughout most of the year.

ERICACEAE. Heather Family

Arbutus menziesii Pursh. Madrono. Madrone. Rare, on eastern end of mountain,
dry rocky ground. Brisbane Canyon (to east of Buckeye Canyon), W May 25, 1965; on
frontage road to south of Brisbane, McC, K September 11, 1964. One of the important broad-
leaved evergreen trees of the Pacific Coast region with clusters of small white bell-shaped
flowers in spring followed by red fruits looking much like those of its relative, the European
strawberry tree, Arbutus unedo. Ornamental.

626 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER.

Arctostaphylos imbricata Eastwood. Manzanita. Occasional, brushy slopes and ridges
forming dense low colonies. Summit of mountain, northwest of lowermost radio tower,
McC, Menzies February 16, 1963; West Powerline, McC, K September 2, 1963. Also re-
ported by W. Knight and J. Roof from Kamchatka Point, Blue Blossom Hill and Manzanita
Dike. Known only from San Bruno Mountain. The low-growing, much branched plants
form mat-like colonies, flowers February to May. Plants regenerate by seed following fire.

Arctostaphylos montaraensis J. Roof. Montara manzanita. Rare, adjacent to a rocky
outcrop about 300 yards east-northeast of Parking Lot, Menzies, February 5, 1963. Only
one shrub known prior to 1964 fire. The fire destroyed the shrub but since the fire ap-
proximately a hundred seedlings have appeared. Erect shrub to about 6 feet at maturity
with sessile leaves. Flowers in March. Known only from the San Bruno Mountain locality
and on Peak Mountain in Montara Range about 15 miles airline to south.

Arctostaphylos pacifica J. Roof. Rare, one rocky outcrop. Only in one locality, Ridge
Road about 300 yards northeast of Parking Lot on a prominent rocky outcrop, McC,
Menzies February 16, 1963. Known only from San Bruno Mountain. Plants have a basal
burl and stump-sprout after fire. Flowers in February. Ornamental.

Arctostaphylos uva-ursi (Linnaeus) Sprengel. Bear-berry. Kinnikinnick. Occasional,
on west end of mountain on uppermost ridges. West Powerline, McC, K September 2, 1963;
Cable Ravine, about 30 yards from the Monterey pine, largest colony on mountain, K 864;
Kamchatka Point, McC, Menzies February 16, 1963; northern end of West Powerline Ridge,
K May 4, 1964. San Bruno Mountain represents one of the southernmost stations known
in California for this manzanita which has a wide circumboreal distribution in North
America, Asia, and Europe. Low growing, mat-forming shrubs, flowers February and
March, Ornamental.

Although one of the smaller genera on San Bruno Mountain the genus Arctostaphylos
is one of the most interesting. Arctos!aphylos imbricata, an endemic, is a prostrate sessile-
leaved shrub which forms sizeable colonies and has been found at five locations. It is making
a good recovery from the fire of 1964 due to the large amounts of seed which lay on the
ground at the time of the fire. Arctostaphylos pacifica, a small-leaved burl-forming creep-
ing manzanita is also making a comeback after the fire due to regeneration from root sprouts.
It is found only at one rocky outcrop a couple of hundred yards below the parking lot and
is distinguished by its reticulate leaf surfaces and serrulate leaf margins. Just adjacent and
downhill a few yards was a six foot high shrub of A. montaraensis which was killed outright
by the fire in 1964 but is being perpetuated nicely by the appearance of numerous seedlings.
A cross section of this original shrub showed growth rings which indicated its age to have
been about thirty years. The largest and only other colony of A. montaraensis is at its type
locality on Montara Mountain Range, within sight, and about 15 airline miles to the south.

Arctostaphylos uva-ursi usually occurs on the north-facing slope of the mountain except
for the colony previously mentioned near Cable Ravine. On Kamchatka Point there is a
low-growing form with small suborbicular leaves borne on assurgent branchlets coming off
of cord-like branches which are absolutely prostrate. It has been introduced to cultivation
under the cultivar name of ‘Miniature’ and seems to be suitable as a ground cover. Arcto-
staphylos uva-ursi, in some of its localities along the Pacific Coast, hybridizes with adjacent
species. Some of its plants on San Bruno Mountain, however, are characteristic of the
species and do not appear to be of hybrid origin as they do farther to the south in Monterey
County.

Gaultheria shallon Pursh. Salal. Rare. Known from only one locality on San Bruno
Mountain. Brushy slope above Radio Road about 14 mile from Eucalyptus grove, Menzies

Vout. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 627

January 12, 1964, and K 863. Evergreen spreading shrub of medium height with small urn-
shaped flowers in spring followed by black berry-like fruits. Ornamental.

Vaccinium arbuscula (Gray) Merriam. Huckleberry. Rare, known only from two
exposed rocky outcrops. Kamchatka Point, McC, Menzies February 16, 1963; Huckleberry
Ridge, K September 2, 1964. Deciduous shrub to about 2 feet tall, flowers sparsely in ? March
and April. The occurrence of Vaccinium arbuscula if indeed our material belongs to this
species, is unexpected on San Bruno Mountain since it is a huckleberry of forested and
montane areas to the north as far as British Columbia and east to the Sierra Nevada and the
Rocky Mountains.

Vaccinium ovatum Pursh. Evergreen Huckleberry. Frequent, on exposed often rocky
slopes, ridges, and ravines on upper part of mountain. Shrubs stump sprout in spring fol-
lowing fire. In two of its localities it occurs close to Vaccinium arbuscula from which it is
distinguished by its evergreen, thickish and leathery leaves. Kamchatka Point, Menzies
May 17, 1964; Quarry, K June 24, 1963; ravine % mile east of 1314-foot summit, P 2693;
West Powerline, McC, K June 2, 1963; top of Dairy Ravine, K August 5, 1964; Huckleberry
Ridge, K May 4, 1964. Erect, much branched, evergreen shrub with small bellshaped white,
pink tinged flowers, March to May, followed by black edible huckleberries.

EUPHORBIACEAE. Euphorbia Family

Croton californicus Mueller-Argoviensis. Rare, in sandy soil. Pig Ranch Ravine, W
July 7, 1965, and K 1028. Silvery-leaved perennial, inconspicuous flowers, June to Novem-
ber. Used by California Indians for stupefying fish.

Euphorbia crenulata Engelmann. Chinese Caps. Wartweed. Occasional, on grassy
or brushy slopes, or disturbed areas. Near Crystal Cave Canyon, W June 20, 1965; Colma
Canyon, K March 15, 1964; Pig Ranch Ravine, W May 12, 1965, and K 1113; Olivet Ceme-
tery, W May 18, 1965. Green-leaved annual or biennial with milky juice, and inconspicuous
flowers February to November. Becomes weedy.

Euphorbia lathyris Linnaeus. Gopher Plant. Caper Spurge. Weed, escaping from
cultivation in waste places. Hillside Boulevard near Cypress Hill Golf Course, K October 7,
1963. Coarse, blue-green, annual or biennial, with milky juice, upper leaves clasping the
stem, and inconspicuous flowers, February to November. Native of Europe.

Euphorbia peplus Linnaeus. Petty Spurge. Weed, waste places. Crocker Avenue, McC
September 24, 1967. Erect usually branched annual, to about 10 to 12 inches tall, small
greenish flowers almost anytime. Native of Europe.

FAGACEAE. Oak or Beech Family

Quercus agrifolia Née. Coastal Live Oak. California Live Oak. Occasional, in woodland
of canyons and ravines. Quarry, K July 14, 1963; Buckeye Canyon, K August 5, 1963; Point
San Bruno on a bluff near shoreline, McC, K September 11, 1964. Also reported from the
ridge to west of lower part of Owl’s Canyon. Spreading, much branched evergreen oak,
flowers in catkins, February and March. The leaves of this oak tend to be convex, sharply
toothed, and to have tufts of hairs in the axils of the veins on the lower surface.

Quercus chysolepis Liebmann. Maul Oak. Canyon Oak. Gold-cup Oak. Occasional,
open rocky outcroppings on open slopes and in ravines. Crocker Hills, McC, K September
2, 1963; near Guadalupe Road, K August 5, 1963; Quarry, K July 14, 1963, and June 24,
1963; Buckeye Canyon, W May 25, 1965, and K 722; Bitter Cherry Ridge, K May 4, 1964.
On San Bruno Mountain this oak tends to be a low shrub or small tree 6 to 8 feet tall.

628 CALIFORNIA ACADEMY OF SCIENCES [| PRroc, 4TH SER.

The most dwarf of these low growing shrubs have been designated as Quercus chrysolepis
Liebmann var. nana Jepson. This is an evergreen oak in which the leaves are thick and
leathery, more or less brown hairy on lower sur‘ace, flowers in catkins in May and June.
The acorns have scales more or less covered by golden-brown felt-like hairs, hence the
common name, Gold-cup Oak.

Quercus moreha Kellogg. Oracle Oak. Rare, only on one ridge toward eastern end of
mountain. Brisbane Powerline, J. Roof April 25, 1965. Quercus moreha originated as a
hybrid between the deciduous Q. kelloggii and the evergreen Q. wislizenit and tends to be
semi-deciduous. At its one locality on the mountain it is low growing to about 18 inches tall
and spreading to several feet across. Has been called Quercus morehus.

Quercus wislizeni A. DeCandolle. Interior Live Oak. Occasional, in ravines and rocky
open slopes. Plants regenerate by stump sprouts after fire. Quarry, K July 14, 1963; Buck-
eye Canyon, A August 5, 1963; about 100 yards east of Parking Lot near Ridge Road,
K 1104. Quercus wislizent is an evergreen oak with thick, toothed, parallel-veined leaves.
Plants on San Bruno Mountain are low growing, a form which has been designated as Q.

wislizent A. DeCandolle var. frutescens Engelmann.

FRANKENIACEAE. Frankenia Family

Frankenia grandifolia Chamisso & Schlechtendal. Alkali Heath. Rare, salt flats and
alkaline areas. Beach at Point San Bruno, AK September 2, 1964; salt flat at Point San
Bruno, McC, K September 11, 1964; Brisbane Lagoon, W August 14, 1965. Perennial some-
times exuding crystals of salt, with heath-like leaves, and rose-purple flowers, May to
November.

FUMARIACEAE. Fumitory Family

Dicentra formosa (Haworth) Walpers. Pacific Bleeding Heart. Rare, in moist shaded
areas. Creek in Colma Canyon, K May 30, 1964; Fern Rock near Guadalupe Creek, K 1024.
In both localities it is sparse in occurrence. Although the creek in Colma Canyon was mostly
covered for the Guadalupe Parkway bleeding heart may still be found in one p'ace. Attrac-
tive perennial with much divided, fern-like leaves and attractive pale to deep lavender
flowers in March and April. Ornamental.

GARRYACEAE. Silk Tassel Family

Garrya elliptica Douglas. Coast Silk Tassel. Silk Tassel Bush. Quinine Bush. Oc-
casional, brushy slopes, in chaparral. Regenerates after fire by stump sprouting. Colma
Canyon, K May 4, 1964; Sierra Point, W April 27, 1965; Quarry, K June 24, 1963; Bitter
Cherry Ridge, K April 4, 1965; slope above Trillium Gulch, McC April 27, 1965; ravine just
below top of mountain, McC, Menzies February 16, 1963. Evergreen shrub with stiff,
leathery, undulate-margined leaves which are gray hairy on the lower surface, flowers are
in long tassels, January to March. Plants are either male or female and both staminate and
pistillate flewer tassels are showy although the staminate hang more gracefully. Crnamental.

GENTIANACEAE, Gentian Family

Centaurium davyi (Jepson) Abrams. Centaury. Rare, in grassy areas, occurring
sparsely. On fire trail west of Quarry, W June 19, 1965. Both this species and Centaurium
floribundum are rather inconspicuous anna's with pink flowers in spring and summer.

Centaurium floribundum (Bentham) Robinson. Centaury. Rare, in grassy and dis-
turbed areas, occurring sparsely. Crocker Industrial Park, W March 27, 1966.

Vou. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 629

GERANIACEAE. Geranium Family

Erodium botrys (Cavanilles) Bertoloni. Long-beaked Filaree. Common, weed in grass-
land and disturbed areas. Abandoned quarry at end of School Street, near Hillside Boulevard,
K October 7, 1963; Ridge Road near Brisbane Powerline, McC May 16, 1965, and K 958;
Crocker Avenue, P 2637, and Toschi 63:32; Hillside Boulevard, P 2709. Long-beaked filaree
has simple but pinnately lobed leaves with relatively large flowers (petals to nearly 14 inch
long) in spring. Native of Mediterranean region.

Erodium cicutarium (Linnaeus) L’Heritier. Red-stemmed Filaree. Common, weed in
grassland and disturbed areas. Colma Canyon, K March 15, 1964, and McC June 13, 1965;
abandoned quarry at end of School Street near Hillside Boulevard, K October 7, 1963;
Quarry, K July 14, 1963; Ridge Road, McC May 16, 1965, and K 970; Radio Road near
radio stations, McC, Menzies February 16, 1963. Red-stemmed filaree has pinnately com-
pound leaves with each leaflet finely divided, and relatively small flowers (petals about 4
inch long) late winter into autumn. Native of southern Europe.

Erodium moschatum (Linnaeus) L’Heritier ex Aiton. White-stemmed Filaree. Musk
Filaree. Weed in grassland and disturbed areas. Crocker Avenue, P 2636, and Toschi 63:40;
ravine north of Randolph Avenue, P 2747; Guadalupe Valley, P 2767; Romanzoffia Ravine,
K May 4, 1964. White-stemmed filaree has pinnately compound relatively long leaves (21%
to 12 inches long) with each leaflet coarsely serrate or cleft and relatively small flowers,
about the same size as the red-stemmed filaree. Flowers late winter into autumn. Native
of Mediterranean region.

Erodium obtusiplicatum (Maire, Weiler & Wilczek) J. T. Howell. Rare, weed in
grassland and disturbed places. Romanzoffia Ravine, K May 4, 1964; Ridge Road near
Brisbane Powerlines, McC May 16, 1965. This filaree which has no common name was
long confused with Erodium botrys which it resembles in its simple but lobed leaves, its
relatively large flowers and long beak of fruit. Flowers in spring and summer. Native of
North Africa.

Geranium carolinianum Linnaeus. Carolina Geranium. Rare, on brushy or grassy
slopes. Near Nike Base, McC, W May 23, 1965. Branched annual with pinkish flowers,
April to October. A widely distributed plant from the Pacific to the Atlantic states.

Geranium dissectum Linnaeus. Cut-leaved Geranium. Occasional, weed in grassland
and disturbed areas, often in partial shade. Colma Canyon, K May 4, 1964; Crocker Avenue,
P February 23, 1963. Annual with leaves more deeply divided than Geranium carolinanum
and small pink flowers, April to October. Native of Europe.

Geranium molle Linnaeus. Crane’s-bill Geranium. Occasional, weed in grassland and
disturbed areas, sometimes in shade of other plants. Colma Canyon, McC, Halde, K June
23, 1963; Nike Base, K 1199; Radio Road, W May 23, 1965. Annual or biennial with rose-
pink flowers, February to June.

Geranium retrorsum L’Heritier ex DeCandolle. New Zealand Geranium. Occasional,
weed in grassland and disturbed areas, sometimes in shaded areas. Radio Road, McC, K
September 2, 1963; Cable Ravine, K 860; April Brook, K 980. Perennial with a taproot,
plant covered with retrorse whitish hairs, flowers purple, March to September. Native of
Australia and New Zealand. The four species of Geranium on San Bruno Mountain are
inconspicuous annuals or perennials with small flowers and are not to be confused with the
garden geraniums which belong to the South African genus Pelargonium.

630 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

HALORAGIDACEAE. Water Milfoil Family

Myriophyllum brasiliense Cambessédes. Parrot Feather. Rare, growing in water.
Crocker Industrial Park, W March 27, 1966. Aquatic perennial, stems supported by water,
leaves finely dissected and appearing feathery, tiny flowers June to August. Cultivated as
an aquarium plant but escapes. Native of South America.

HrppocasTANaAceEAE. Horse Chestnut or Buckeye Family

Aesculus californica (Spach) Nuttall. California Buckeye. Horsechestnut. Rare, part
of woodland in ravines or slopes, often windswept and low in stature. Buckeye Canyon,
McC May 27, 1967; Point San Bruno, McC, K September 11, 1964. Deciduous tree with
round crown, leaves characteristically palmately 5-lobed and appear before the showy white
and pinkish flowers borne on thick spike-like clusters, April to June. The leaves fall early
even before the summer is over. The pear-shaped fruits usually have one glossy brown seed
which contains poisonous substances. The Indians used the seeds for food after leaching with
boiling water, and for stupefying fish.

HypROPHYLLACEAE. Waterleaf or Phacelia Family

Eriodictyon californicum (Hooker & Arnott) Torrey. Yerba Santa. California Moun-
tain Balm. Rare, in chaparral. Buckeye Canyon, K July 14, 1963; Owl’s Canyon, W June
17, 1965; West Powerline, K May 4, 1964. Evergreen shrub with small pale blue flowers,
April to July. Regenerates after fire both by root sprouts and by seeding itself.

Nemophila heterophylla Fischer & Meyer. Rare, moist brushy slope. Devil’s Arroyo,
McC March 28, 1965. Weak-stemmed annual with pale blue to white flowers, January to
April.

Nemophila menziesii Hooker & Arnott. Baby-Blue Eyes. Occasional, moist open
grassland or in chaparral, often coming up among other plants. Reservoir Hill, K 879, and
McC March 14, 1965; Bitter Cherry Ravine, K 1006. Weak-stemmed, diffusely branched
annual with showy pretty blue flowers, February to April. A favorite spring wild flower,
its name commemorates Archibald Menzies, the early Scottish explorer of the Pacific coast
who was with the Vancouver Expedition.

Nemophila parviflora Douglas ex Bentham. Rare, moist brushy slope. Devil’s Arroyo,
K April 26, 1965. Weak-stemmed annual with small inconspicuous flowers, March to June.

Phacelia californica Chamisso. One of the most common perennials on mountain, in
grassland and chaparral. Lower Colma Canyon, McC, Halde, K June 23, 1963; Colma
Canyon, K 1197, K1214, K 1222; ravine north of Randolph Avenue at Hillside Boulevard,
P 2739; east facing slope below radio station, Toschi 63:67; slope south of 1314-foot sum-
mit, P 2667; ravine % mile east of 1314-foot summit, P 2691; exposed slopes below sum-
mit, McC June 9, 1963; Quarry, K July 14, 1963; Romanzoffia Ravine, K May 4, 1964;
Huckleberry Ridge, K May 4, 1964; Cable Ravine, McC March 14, 1965; Bitter Cherry
Ravine, K 1010; Glen Park Canyon, McC April 16, 1967. Gray hairy perennial with blue
to purplish flowers, March to September.

Phacelia distans Bentham. Wild Heliotrope. Occasional, rocky, sandy, and brushy
slopes. South facing slope near West Powerline, McC March 14, 1965; Pig Ranch Ravine,
W May 11, 1965, K May 15, 1965. Weak-stemmed annual with blue flowers, March to June.

Phacelia malvaefolia Chamisso. Stinging Phacelia. Occasional, on brushy slopes. Colma
Canyon, McC, Halde, K June 23, 1963; Romanzoffia Ravine, K May 4, 1964, McC March

Vou. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 631

28, 1965. Weak stemmed annual with coarse conspicuous hairs, and dull white inconspicuous
flowers, April to July.

Pholistoma auritum (Lindley) Lilja ex Lindblom. Fiesta Flower. Climbing Nemophila.
Rare, under shade in wooded slopes. Sierra Point, W April 27, 1965; Gladys Canyon, W
June 16, 1965. Weak stemmed annual with coarse hairs, short recurved prickles, and blue
flowers, March to June.

Romanzoffia suksdorfii Greene. Mist-Maiden. Occasional, on shaded or open ‘moist
rocky areas. Romanzoffia Ravine, K May 4, 1964, McC March 28, 1965; Buckeye Canyon,
W May 8, 1965. Reported in Owl’s Canyon by James Roof. Attractive, dainty, tuberous-
rooted perennial with mostly basal leaves and small white flowers, March to May. Orna-
mental.

HYPERICACEAE. St. John’s Wort Family

Hypericum anagalloides Chamisso & Schlechtendal. Creeping St. John’s Wort. Tinkers
Penny. Rare, along seepages or stream sides. Cow Trough Ravine, K October 7, 1963;
stream in Colma Canyon, W June 16, 1965. Perennial with prostrate and a few erect stems
and small yellow flowers, March to July.

LABIATAE. Mint Family

Lepechinia calycina (Bentham) Epling [Sphacele calycina Bentham]. Pitcher Sage.
Occasional, in brushy areas. Regenerates by seedlings after a fire. Crocker Hills, K, McC
September 2, 1963; Guadalupe Road, K August 5, 1963; ravine west of Crystal Cave Canyon,
W June 20, 1965. Low-growing, aromatic shrub with somewhat woolly leaves and showy
white flowers, April to July. Ornamental.

Marrubium vulgare Linnaeus. White Hoarhound. Rare, weed of waste ground. Along
Bayshore Highway, South San Francisco, W March 20, 1966. White woolly perennial with
small white flowers in dense heads, February to July. Native of Europe.

Mentha spicata Linnaeus. Spearmint. Escape from cultivation in wet ground. Crocker
Avenue, W July 7, 1965. Pleasantly fragrant perennial with lavender flowers in a spike-like
inflorescence, July to October. Native of Europe.

Monardella villosa Bentham. Coyote Mint. Pennyroyal. Common, on brushy and
rocky areas. Regenerates after fire by root sprouts. Quarry, K July 14, 1963, and K June
4, 1963; lower Buckeye Canyon, McC September 29, 1966; lower Colma Canyon, McC,
Halde, K June 23, 1963; slope above April Brook, McC August 1, 1965; lower area of
Quarry, McC August 1, 1965; West Powerline, McC March 25, 1965; east facing slope below
Parking Lot, McC, Halde June 23, 1963; canyon east of Quarry, W June 19, 1965; Dairy
Ravine, K August 5, 1963. An erect perennial, 12 to 18 inches tall, with lavender flowers in
compact heads, June to August. Two forms are described which may sometimes be dis-
tinguished on San Bruno Mountain. Var. villosa has “thin” nearly glabrous leaves, and is
mostly on the eastern side of the mountain. Var. franciscana (Elmer) Jepson has “thickish”
usually densely hairy leaves, and is mostly on the western side of the mountain.

Pogogyne serpylloides (Torrey) Gray. Rare, moist grassy and brushy places. South
Powerline near summit, McC April 25, 1965; fire trail west of Quarry, W June 19, 1965.
Slender aromatic annual with small lavender flowers, April to June. Grows in shade of sur-
rounding plants and is easily overlooked.

632 CALIFORNIA ACADEMY OF SCIENCES [Proc, 4TH SER.

Prunella vulgaris Linnaeus subsp. lanceolata (Barton) Hultén. Rare, on grassy or
semi-shaded slopes. Upper Colma Canyon near Radio Road, W July 7, 1965; Colma Canyon,
K August 5, 1963. Perennial with lanceolate leaves, and light purple flowers, April to
December.

Salvia spathacea Greene. Crimson Sage. Pitcher Sage. Common, open grassy or rocky
slopes. Lower Colma Canyon, McC, Halde, K June 23, 1963; Colma Canyon, K 1218,
Raven 1787; slope southeast of 1314-foot summit, P 2681; West Powerline, McC, K Septem-
ber 2, 1963; Brisbane Powerline, K 953; Trillium Gulch, McC March 28, 1965. Coarse, gray
hairy, aromatic perennial with reddish flowers in showy heads, March to June.

Satureja douglasii (Bentham) Briquet [Micromeria douglasti Bentham]. Yerba Buena.
Frequent, in grassland and brushy areas, creeping on ground at base of surrounding plants.
Slope above April Brook, McC August 1, 1965; Ridge Road at Saddle Camp, K August 5,
1963; east facing slope below Parking Lot, McC, Halde June 23, 1963; West Powerline,
McC, K September 2, 1963; Point San Bruno, K March 13, 1964. Fragrant, evergreen trail-
ing perennial, rooting at the stem tips, small whitish flowers, May to August.

Scutellaria tuberosa Bentham. Skull-cap. Rare, on brushy slopes, under shade of
surrounding plants. Glen Park Canyon, McC April 30, 1967. Low growing, fragrant peren-
nial with tubers on the root-stocks and dainty blue flowers, March to May.

Stachys ajugoides Bentham. Hedge Nettle. Rare, low often wet ground and waste
places. Colma Canyon, K August 5, 1963; Radio Road, K August 5, 1963. Coarse perennial,
flowers May to October.

Stachys bullata Bentham. Hedge Nettle. Occasional, edge brushy areas and shaded
slopes. Lower Colma Canyon, McC, Halde, K June 23, 1963; Quarry, K July 14, 1963;
Cow Trough Ravine, October 7, 1963. Coarse perennial, flowers April to September.

Stachys chamissonis Bentham. Coast Hedge Nettle. Rare, in moist places and ravine
creek bottoms. Devil’s Arroyo, McC March 28, 1965, K 1101; near source of Guadalupe
Creek, K 988. Rather tall coarse perennial, flowers March to May.

Stachys rigida Nuttall ex Bentham subsp. quercetorum (Heller) Epling. Hedge Nettle.
Frequent, on open grassy slopes and brushy hillsides, often in colonies. East facing slope be-
low Parking Lot, McC, Ha'de June 23, 1963; Bitter Cherry Ridge, K 1012; near Nike Base,
McC, W May 23, 1965; Point San Bruno, K 1063; Crocker Hills, K May 4, 1964; Glen
Park Canyon, McC April 16, 1967. Coarse perennial, flowers March to July.

LAURACEAE. Laurel Family

Umbellularia californica (Hooker & Arnott) Nuttall. California Laurel. California
Bay. Oregon Myrtle. Woodland in Buckeye Canyon, McC September 29, 1966. Evergreen
tree with pungently aromatic, coriaceous leaves and clusters of small yellow-green flowers,
January to March. A relative of the Mediterranean laurel, Laurus nobilis, its leaves may be
used as a seasoning, but since its leaves are more pungent than the Mediterranean tree it
should be used with discretion. Its hard wood takes a fine polish and is used for making
small ornaments.

LEGUMINOSAE. Pea or Bean Family

Acacia decurrens Willdenow. Green Wattle. Escape from cultivation and occasionally
naturalized. Crocker Avenue, McC, K January 12, 1963. Tree with finely divided leaves
and yellow flowers, January to March. Native of Australia.

Vou. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 633

Acacia melanoxylon R. Brown. Blackwood Acacia. Black Acacia. Escape from culti-
vation and occasionally naturalized. Cable Ravine, McC March 14, 1965; Horseshoe Ridge,
K August 5, 1963. Tree with leaves reduced to 2 to 5 nerved phyllodia, yellow flowers,
February and March. Native of Australia.

Acacia retinodes Schlechtendal. Everblooming Acacia. Escape from cultivation and
occasionally naturalized. Crocker Avenue, McC, K January 12, 1964, W July 7, 1965. Tree
with leaves reduced to 1-nerved phyllodia, yellow flowers January to August. Native of
Australia. :

Albizia distachya (Ventenat) Macbride [A. lophantha (Willdenow) Bentham]. Stink
Bean. Plume Albizia. Escape from cultivation and occasionally naturalized. Bayshore High-
way, W June 8, 1965. Tree with finely divided leaves and yellow flowers in a spike-like
plume, late winter and spring. Native of Australia.

Astragalus nuttallii (Torrey & Gray) J. T. Howell var. virgatus (Gray) Barneby.
Locoweed. Occasional, in grassland, and in the open on sandy soil. Flower Garden near Radio
Road, W June 20, 1965, W August 16, 1965; south of Pig Ranch Ravine, W July 7, 1965;
near Olivet Cemetery, W May 18, 1954; April Brook, J. Roof July 17, 1963. Robust but
weak-stemmed perennial with greenish-white f'owers, April to October.

Cytisus maderensis Masferrer. Broom. Escape from cultivation and naturalized in
disturbed places. Radio Road, K August 5, 1963. Evergreen shrub with yellow flowers in
spring. Native of Madeira.

Cytisus monspessulanus Linnaeus. French Broom. Escape from cultivation and is
naturalized in disturbed places.

Cytisus scoparius (Linnaeus) Link. Scotch Broom. Escape from cultivation and
naturalized in disturbed places. Brisbane, W Apri! 27, 1965; Sierra Point, W April 27, 1965.
Nearly leafless shrub with large yellow or sometimes yellow and brown flowers, spring.

Lathyrus odoratus Linnaeus. Sweet Pea. Escape from cultivation, occasionally becoming
established in disturbed places. Point San Bruno, W May 8, 1965. Annual with fragrant
flowers in different colors, spring. Native of Italy.

Lathyrus vestitus Nuttall ex Torrey & Gray. Pacific Pea. Common, on brushy areas,
climbing on adjacent shrubs or spreading on the ground. Lower Colma Canyon, McC, Halde,
K June 23, 1963; Colma Canyon, W June 13, 1965, K March 15, 1964; slope southwest of
Brisbane, P 2759; South Powerline, McC March 14, 1965; Reservoir Hill, McC March 14,
1965; Devil’s Arroyo, McC March 28, 1965; West Powerline, McC March 10, 1965. Peren-
nial with well developed tendrils, pink to pale purplish flowers usually fading with age,
March to July. A variable species divided into two or more subspecies on the basis of
height of plant, internode length, and amount of pubescence.

Lotus corniculatus Linnaeus. Bird’s Foot Trefoil. Weed of roadsides and other dis-
turbed areas. Crocker Industrial Park, McC July 29, 1966; West Powerline, W May 18,
1966. Prostrate perennial with yellow flowers, June to August. Native of Europe.

Lotus micranthus Bentham. Bird’s Foot Trefoil. Frequent, in grassland and edge of
brushy areas. Colma Canyon, K May 30, 1964; April Brook, McC March 28, 1965; Bitter
Cherry Ridge, K April 4, 1965; Buckeye Canyon, W March 30, 1965; Crocker Hills, McC
April 11, 1965. Slender-stemmed annual with small salmon-colored flowers, March to June.

634 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Lotus purshianus (Bentham) Clements & Clements. Spanish Clover. Rare, in grassland.
Sierra Point, W May 8, 1965. Low-growing annual, sometimes forming mats, with small
pinkish-white flowers, April to October.

Lotus scoparius (Nuttall) Ottley. Deerweed. Frequent, in many habitats, in grassy
and brushy areas. Lower Colma Canyon, McC, Halde, K June 23, 1963; Radio Road, McC,
K September 2, 1963; Owl’s Canyon, W June 7, 1965; near Olivet Cemetery, W May 18,
1965; Crocker Hills, K May 4, 1964. Brushy perennial with erect to spreading green virgate
stems and flowers yellow tinged with red, May to October.

Lotus subpinnatus Lagasca. Frequent, in grassland and brushy slopes, sometimes in
disturbed places. Crystal Cave Canyon, K 1252; Devil’s Arroyo, K May 9, 1965; Pig Ranch
Ravine, K May 15, 1965; Crocker Industrial Park, McC March 28, 1965. Low growing,
weak-stemmed annual with small, yellowish, solitary, sessile flowers, March to June.

Lupinus affinis Agardh. Rare, in grassland. Colma Canyon, McC, W June 13, 1965.
Annual with blue flowers, April to June.

Lupinus albifrons Bentham ex Lindley. Silver Bush Lupine. Common, open rocky
slopes. Colma Canyon, K May 4, 1964; Randolph Drive at Hillside Boulevard, P 2725;
Sierra Point, W April 27, 1965; Motorcycle Ridge, K August 5, 1963; west facing slope be-
low top of mountain, McC April 17, 1964; ravine 4% mile east of summit, P 2685; Glen Park
Canyon, McC April 16, 1967. Attractive shrub or subshrub with silvery-gray foliage and
blue flowers April to June. This lupine is variable in habit and two forms may sometimes
be recognized. Plants erect and up to 3 to 4 feet tall may be called var. albifrons and those
lax and spreading with stems less than 18 inches may be called var. collinus Greene.

Lupinus arboreus Sims. Bush Lupine. Yellow Beach Lupine. Occasional, mostly in
sandy areas or disturbed places. Colma Canyon, McC June 13, 1965; Sierra Point, W May
8, 1965; top of mountain near radio station, McC, Halde June 23 1963. Attractive shrub
with yellow flowers, March to September. Occurring commonly in sandy soil along the
coast of northern and central California.

Lupinus bicolor Lindley. Frequent, in grassland. Slopes southwest of Brisbane, P 2757;
Bitter Cherry Ridge, K 1008; Ridge Road, K March 14, 1965; Guadalupe Road, McC April
11, 1965; Flower Garden adjacent to April Brook, K 975; Crocker Industrial Park, W June
19, 1965. Annual with small blue flowers, March to June. This common spring wildflower
is a variable species with several forms separated on the basis of technical characters of the
flowers.

Lupinus chamissonis Eschscholtz. Blue Beach Lupine. Rare, in sandy soil. Colma
Canyon, K 973. Shrubby, with blue flowers, March to August. Frequently seen in sandy
soil along the coast of central and southern California.

Lupinus formosus Greene. Rare, dry open often sandy areas. Sierra Point, W June
21, 1965. Perennial with lavender flowers, May and June.

Lupinus micranthus Douglas ex Lindley. Occasional, in grassland. Guadalupe Valley,
P 2764; West Powerline, K 709. Annual with small blue flowers, March to May. Resembles
Lupinus bicolor from which it usually may be distinguished by the glabrous or sparsely
hairy upper leaf surfaces.

Lupinus nanus Douglas ex Bentham. Frequent, in grassland, often forming showy
colonies. Colma Canyon, McC, W June 13, 1965; Crocker Industrial Park, W May hs

Vou. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 635

1966; Sierra Point, W April 27, 1965. Annual with dark blue or rarely white flowers, March
to June.

Lupinus succulentus Douglas ex Koch. Rare, in disturbed ground. Point San Bruno,
K March 30, 1964. Stout, hollow-stemmed annual with blue flowers.

Lupinus variicolor Steudel. Frequent, in grassland or open gravelly places. Colma
Canyon, K May 4, 1964; east facing slope below top of mountain, McC April 17, 1964; West
Powerline, K 1042. Perennial, often decumbent, with flowers of varying shades of blues,
pinks, yellows and whites.

Medicago polymorpha Linnaeus var. vulgaris (Bentham) Shinners [M. hispida
Gaertner]. Bur Clover. Weed of disturbed places. Randolph Drive at Hillside Boulevard,
P 2718; Colma Canyon, K May 30, 1964; Point San Bruno, K 1033. Weak-stemmed an-
nual with small yellow flowers, March to June, and a coiled spiny pod. Native of Europe.

Medicago sativa Linnaeus. Alfalfa. Lucerne. Weed of roadsides and waste ground, an
escape from cultivation. Frontage road at north city limits of South San Francisco, McC,
K September 11, 1964; Brisbane Lagoon, W June 19, 1965. Perennial with small blue flowers,
spring to autumn. An important forage plant, native of the Old World.

Melilotus albus Desrousseaux ex Lamarck. White Sweet Clover. Weed, waste places.
Crocker Avenue, McC September 24, 1967. Tall erect annual or biennial with tiny white
flowers, spring to autumn. Native of Eurasia.

Melilotus indicus (Linnaeus) Allioni. Indian Sweet Clover. Weed of waste ground and
disturbed places. Point San Bruno, McC, K September 11, 1964; Sierra Point, W May &,
1965. Erect annual with small yellow flowers, April to October. Native of Old World.

Melilotus officinalis (Linnaeus) Lamarck. Weed of roadsides and disturbed places.
Crocker Industrial Park, W June 8, 1965. Erect annual with small yellow flowers, May to
August. Native of Europe.

Pisum sativum Linnaeus. Garden Pea. Weed of disturbed places. Cow Palace area,
W May 17, 1965. Annual, flowers in spring. An edible plant which occasionally escapes
from cultivation. Native of the Old World.

Trifolium amplectens Torrey & Gray. Sack Clover. Occasional, in grassland. Point
San Bruno, K 1078; Bitter Cherry Ravine, K May 15, 1965; West Powerline, McC March
28, 1965. Annual with purple or sometimes pale flowers in small heads, March to June.
At maturity the flowers become inflated, like tiny sacks, hence the common name.

Trifolium bifidum Gray. Rare, in grassland. Radio Road, W May 23, 1965; April
Brook, K May 15, 1965. Annual with pale pink flowers, April and May, on slender recurved
pedicels.

Trifolium dubium Sibthorp. Shamrock. Weed of waste places, lawns, and other dis-
turbed areas. Near Cow Palace, W May 18, 1965. Annual with small yellow flowers, May
and June. Native of Europe.

Trifolium gracilentum Torrey & Gray. Occasional, in grassland and disturbed areas.
North of Randolph Drive at Hillside Boulevard, P 2719; Pig Ranch Ravine, K May 15,
1965; Crocker Hills near Guadalupe Road, McC April 11, 1965. Slender-stemmed annual
with small heads of little pink flowers, March to May.

636 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Trifolium microcephalum Pursh. Occasional, in open grassland. Slope above Quarry,
McC August 1, 1965; Devil’s Arroyo, K May 9, 1965; West Powerline, McC March 28, 1965.
Slender-stemmed annual with pinkish flowers in small heads, April to June.

Trifolium microdon Hooker & Arnott. Occasional, in open grassland or rocky slopes.
Pig Ranch Ravine, K May 15, 1965; West Powerline, McC March 28, 1965. Slender-
stemmed annual with pinkish flowers, March to July.

Trifolium repens Linnaeus. White Clover. Escapes from cultivation, sometimes used
in lawns, and naturalized. West Powerline, W May 18, 1965; Radio Road, K August 5,
1965. Perennial with whitish flowers in showy heads, April to October. Native of Europe.

Trifolium tridentatum Lindley. Tomcat Clover. Occasional, in grassland, sometimes
forms colonies. Pig Ranch Ravine, K May 15, 1965; Nike Base, K May 15, 1965; Devil’s
Arroyo, K May 9, 1965; Ridge Road, May 16, 1965. Annual with erect to decumbent stems
and red-purple flowers in showy heads, March to May.

Trifolium wormskjoldii Lehmann. Occasional, in wet or marshy places. April Brook,
W May 18, 1965; Crocker Industrial Park, W May 17, 1965. Creeping perennial with de-
cumbent stems and light purple flowers in shewy heads, April and May.

Ulex europaeus Linnaeus. Gorse. Furze. Escape from cultivation, becoming weedy in
disturbed areas. North of Crocker Avenue, Toschi 63:37; Crocker Hills, McC, K September
2, 1963. Spiny shrub with yellow flowers, February to July. Native of Europe.

Vicia americana Muhlenberg ex Willdenow. American Vetch. Frequent, in grassy and
brushy areas, often scrambling over other plants. Colma Canyon, K May 4, 1964; Nike
Base, K 1232; Kamchatka Point, K March 21 1965; South Powerline, McC March 14, 1965.
Trailing perennial with purplish flowers, March to May. Resembles Lathyrus vestitus.

Vicia angustifolia Linnaeus. Common Vetch. Weed of roadsides and waste places.
Crocker Hills, A May 4, 1964. Trailing perennial with fairly large purple flowers, April to
June. Native of Europe.

Vicia benghalensis Linnaeus. Vetch. Weed of roadsides and disturbed areas. Vicinity
of Cow Palace, W May 17, 1965. Pubescent annual with rose-purple flowers, April to June.
Native of Mediterranean region.

Vicia gigantea Hooker. Giant Vetch. Occasional, in moist open or brushy places,
rampant climber on adjacent shrubs. Colma Canyon, K May 30, 1964; base of Quarry,
K July 14, 1963; Radio Road, W May 23, 1965; April Brook, McC March 28, 1965; Wax
Myrtle Ravine, K August 5, 1963. Perennial with stout trailing stem and reddish-purple
flowers, March to August.

Vicia sativa Linnaeus. Spring Vetch. Weed along roadsides and disturbed areas. Upper
Colma Canyon, McC, W June 11, 1967. Annual with sessile, light and dark purple flowers
March to July. Native of Europe.

LINACEAE. Flax Family

Linum usitatissimum Linnaeus. Flax. Escape from cultivation and becoming weedy
along roadsides and disturbed places. San Bruno Hills, Vincent Jones April 1921. Slender,
branched annual with pretty blue flowers, March to July. Native of Europe. Not reported
from our area within recent years.

Vov. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 637

LYTHRACEAE. Loosestrife Family

Lythrum hyssopifolia Linnaeus. Loosestrife. Occasional, in moist places as damp
creek beds, and meadows. Crocker Hills, McC September 2, 1963; Radio Road, K August
5, 1963; fire trail west of Quarry, W June 19, 1965; Sierra Point, W August 8, 1965; Crocker
Industrial Park, W June 8, 1965. Slender annual with small linear leaves and pale purple
flowers, April to October. Widely distributed in the northern hemisphere from North America
to Europe.

MatvaceEaAE. Mallow Family

Lavatera arborea Linnaeus. Tree Mallow. Escape from cultivation along roadsides,
becoming spontaneous. Bayshore highway, Brisbane, W June 8, 1965. Shrubby with red-
purple flowers, May to July.

Lavatera assurgentiflora Kellogg. Malva-Rose. Escape from cultivation, used as a
windbreak, and occasionally becoming spontaneous. Hillside Boulevard near Cypress Hill
Golf Course, K October 7, 1963. Shrubby, sometimes a small tree, showy rose-purple flowers
almost throughout the year. Native of the Channel Islands.

Lavatera cretica Linnaeus. Weed of waste places, fields, and roadsides. Point San
Bruno, K 1032; lower Pig Ranch Ravine, K April 4, 1965; Crocker Hills, K 2043; Colma
Canyon, McC, Halde, K June 23, 1963. Annual with small pinkish flowers, spring through
autumn. Native of Mediterranean region.

Malva nicaeensis Allioni. Bull Mallow. Weed of roadsides, waste places, and around
habitations. Crocker Hills, W May 17, 1965. Annual with pale lavender flowers, spring
through summer. Native of Europe.

Malva parviflora Linnaeus. Cheeseweed. Weed of waste ground, fields, and around
habitations. Crocker Industrial Park, W October 16, 1966. Annual with pale pinkish flowers,
spring through summer. Native of Europe. The small disk-shaped fruits or “cheeses” are
edible.

Sidalcea malvaeflora (DeCandolle) Gray ex Bentham. Checker Bloom. Wild Holly-
hock. Common, on open grassy areas. Crocker Avenue, P 2633; Randolph Drive, P 2710;
Colma Canyon, McC, Halde, K June 23, 1963; Radio Road, K 1245; Flower Garden ad-
jacent to April Brook, McC March 28, 1965; top of mountain near radio station, McC,
Menzies February 16, 1963; slope southeast of summit, P 2674. Attractive perennial with
pink flowers, February to June.

MyricacEazE. Wax Myrtle or Bayberry Family

Myrica californica Chamisso & Schlechtendal. California Wax Myrtle. Rare, in moist
ravines. Wax Myrtle Ravine, K August 5, 1963; canyon south of Pig Ranch Ravine, W July
7, 1965. Evergreen tree with inconspicuous flowers, April to June. Ornamental.

Myrtaceak. Myrtle Family

Eucalyptus globulus Labillardiére. Blue Gum. Spontaneous with planted trees. Fog
Forest along old Guadalupe Road, K January 15, 1964; Horseshoe Ridge, K August 5, 1965.
Tall tree with peeling bark and white flowers, January to May. Native of Tasmania and
Victoria, Australia.

638 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER.

Ficure 8. Camissonia ovata (Nuttall ex Torrey & Gray) Raven.

NYCTAGINACEAE. Four-o’Clock Family

Mirabilis jalapa Linnaeus. Four-o’Clock. Marvel of Peru. Escape from cultivation
and occasionally spontaneous. Marshy area near Bayshore Freeway, K October 4, 1966.
Perennial with showy flowers in shades of rose-pink to purple and white, opening in late
afternoon, August to October. Native of tropical America.

ONAGRACEAE. Evening Primrose Family

Camissonia dentata (Cavanilles) Reiche [Oenothera contorta Douglas ex Hooker var.
strigulosa (Fischer & Meyer) Munz]. Rare, open areas in deep sandy soil. Mouth of Colma
Canyon, K April 15, 1965; Pig Ranch Ravine, K May 16, 1967. Annual with small yellowish
flowers, April to July.

Camissonia micrantha (Hornemann ex Sprengel) Raven [Oenothera micrantha Horne-
mann ex Sprengel]. Rare, in grassland or sandy areas. Pig Ranch Ravine, K April 24, 1965;
K May 16, 1967. Annual with small yellow flowers, April to July.

Camissonia ovata (Nuttall ex Torrey & Gray) Raven [Oenothera ovata Nuttall].
Suncup. Frequent, in grassland. Flower Garden near April Brook, McC March 28, 1965; near
Nike Base, McC May 23, 1965; eastern side of mountain about 4% mile east of Brisbane
Powerlines, K 952; Glen Park Canyon, McC April 16, 1967. Perennial with deep taproot,
basal rosette of leaves, and bright yellow flowers, February to June.

Clarkia davyi (Jepson) Lewis & Lewis. Occasional, grassy slopes. Crystal Cave Canyon,

=>
PLATE 3. Upper. Clarkia rubicunda (Lindley) Lewis & Lewis.
Lower. Castilleja wightii Elmer.

( McCLINTOCK

PLATE

AND
KNIGHT)

PROC. CALIF. ACAD. SCI., 4TH SER., VOL,XXXII NO.

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Vou. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 639

K June 20; 1965; Buckeye Canyon, W May 25, 1965. Annual, stem sometimes stout, leaves
ovate and more or less crowded, flowers pinkish, petals sometimes spotted, May and June.

Clarkia purpurea (Curtis) Nelson & Macbride. Occasional, in grassland... Above Harold
Avenue, Brisbane, McC, W May 14, 1967. Slender annual with dark red-purple flowers, the
petals lacking spots, April to July.

Clarkia rubicunda (Lindley) Lewis & Lewis. Farewell-to-Spring. Frequent, in grassland.
Sierra Point, W June 21, 1965; Colma Canyon, McC, Halde, K June 23, 1963; Crystal Cave
Canyon, K June 20, 1965; Quarry, K July 14, 1963; Buckeye Canyon, W June 7, 1965;
above Olivet Cemetery, McC June 13, 1965. Showy annual often branched above, with
pink flowers, petals often with red spot at base, April to August. Ornamental.

Epilobium adenocaulon Haussknecht. Northern Willow-Herb. Occasional, in moist
places. Quarry, McC August 1, 1965; Sierra Point, W May 8, 1965; Radio Road, McC, K
September 2, 1963; Cow Trough Ravine, K October 7, 1963; Crystal Cave Canyon, K June
20, 1965. Perennial with pale to dark pink flowers, June to October.

Epilobium franciscanum Barbey. San Francisco Willow-Herb. Occasional, in moist
places. Guadalupe Road, K August 5, 1963; Wax Myrtle Ravine, K August 5, 1963; slope
below top of mountain, McC June 9, 1963; Colma Canyon, K May 30, 1964; Radio Road,
K June 20, 1965; Crystal Cave Canyon, K June 20, 1965. Perennial with pink flowers,
April to June.

Epilobium paniculatum Nuttall. Willow-Herb. Occasional, in open usually dry dis-
turbed areas. Crocker Avenue, K August 5, 1963; Cow Trough Ravine, K October 7, 1963;
Radio Road, McC, K September 2, 1963; Great Meadow near Radio Road, W August 16,
1965; Quarry, K July 7, 1963. Annual, stem of mature plants almost woody and with ex-
foliating epidermis, small pinkish flowers, May to October.

Epilobium watsonii Barbey. Willow-Herb. Occasional, in moist areas. Radio Road,
McC, K September 2, 1963; Cow Trough Ravine, K October 7, 1963; Crystal Cave Canyon,
K June 20, 1965. Perennial with dark pink flowers, June and July.

Oenothera hookeri Torrey & Gray. Rare, ina seepage area. Upper Colma Canyon, McC,
W June 11, 1967. Coarse biennial, 3 to 4 feet tall with large yellow showy flowers, June
to October.

Zauschneria californica Presl. California Fuchsia. Occasional, dry rocky or brushy
areas. South Powerline, Menzies March 14, 1965; Pig Ranch Ravine, K 1112, W July 7,
1965. Suffrutescent, decumbent, grayish perennial with showy red flowers, July to Novem-
ber. Ornamental.

OROBANCHACEAE. Broomrape Family

Orobanche uniflora Linnaeus. Broomrape. Rare, on rocky ledges, parasitic on Sedum
spathulifolium. Owl’s Canyon, K, McC, W April 25, 1965. Stems fleshy, colorless, and leaf-
less, flowers purple, March and April.

OXALIDACEAE. Oxalis Family

Oxalis pes-caprae Linnaeus. Bermuda Buttercup. Cape Oxalis. Weedy escape from
cultivation in grassland and disturbed areas. Crocker Avenue, AK January 15, 1964; Reservoir
Hill, K February 28, 1965; South Powerline, Menzies March 14, 1965. Stemless perennial,
leaves all basal, bright yellow showy flowers February to June. Native of South Africa.

640 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER.

This attractive weed is undesirable because the scaly bulbs on its rootstocks make it difficult
to eradicate.

Oxalis pilosa Nuttall. Hairy wood sorrel. Occasional, grassland. Colma Canyon, McC,
Halde, K June 23, 1963; Guadalupe Road, McC, Ryder February 16, 1964; Nike Base, K
1239; Ridge Road near Powerlines, McC April 25, 1965; Glen Park Canyon, McC April 16,
1967. Low growing perennial with small yellow flowers, April to June.

PAPAVERACEAE. Poppy Family

Eschscholzia californica Chamisso. California Poppy. One of the mountain’s com-
monest spring wildflowers, in grassland and disturbed areas. Cable Ravine, McC March
14, 1965; rocky slope near Parking Lot, McC, Halde June 23, 1963; Randolph Avenue,
P 2706; Ridge Road on summit of mountain, K 632. Usually a biennial with gray-green
much divided leaves and showy orange colored flowers, March to October or occasionally
other times during the year.

Meconella californica Torrey. Rare, wet rocky slopes and grassland, sometimes in
small colonies. Ravine east of Brisbane Powerlines, McC March 21, 1965; Devil’s Arroyo, McC
March 28, 1965; Glen Park Canyon, McC April 16, 1967. Slender glabrous annual with
obovate to spathulate basal leaves, and small white flowers, February to April.

Meconella linearis (Bentham) Nelson & Macbride. Known only from a collection
made in “San Bruno Hills” by A. D. E. Elmer (no. 4594) in June, 1903. Differs from pre-
ceding species by its linear basal leaves and spreading hairs on flowering stems.

Papaver somniferum Linnaeus. Opium Poppy. Escape from cultivation. Pig Ranch
Ravine, K May 15, 1965. Coarse, gray-green annual with milky juice, upper leaves clasping
the stem and large, white, pink or purplish flowers. Native of Old World. Sometimes culti-
vated for its large flowers. Opium is obtained from milky juice of fruit.

Platystemon californicus Bentham. Cream-Cups. Occasional, grassland and brushy
areas. Colma Canyon, McC, Halde, K June 23, 1963; above Harold Avenue, Brisbane, W
March 30, 1965; Devil’s Arroyo, McC March 28, 1965; Quarry, W July 8, 1965. Annual with
spreading hairs, entire leaves, and cream-colored flowers, March to June.

Stylomecon heterophylla (Bentham) Taylor. Wind Poppy. Rare, moist shaded often
grassy slopes, sometimes appearing in abundance after a fire. Devil’s Arroyo, K 1017; Owl’s
Canyon, K 1072; lower slopes of West Powerline, W May 12, 1965. Glabrous annual with
divided leaves and scarlet flowers, March to May.

PLANTAGINACEAE. Plantago Family

Plantago coronopus Linnaeus. Weed, on salt flats. Sierra Point, W May 8, 1965;
Brisbane Lagoon, W June 19, 1965. Annual (sometimes a perennial) with coarsely toothed
leaves and flowers in spikes 2 to 3 inches long, March to August. Native of the Old World.

Plantago erecta Morris [P. hookeriana Fischer & Meyer var. californica (Greene)
Poe]. California Plantain. Frequent, open slopes at edge of chaparral and in grassland,
sometimes forming sparse to dense colonies. Randolph Avenue, P 2732; Colma Canyon,
K March 15, 1954; Glen Park Canyon, McC April 16, 1967; Devil’s Arroyo, K 1106; Crystal
Cave Canyon, W June 20, 1965; slope below Parking Lot, McC June 9, 1963. Slender an-

nual with entire linear leaves and flowers in spikes to as much as 1 inch long, January
to June.

Vout. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 641

Plantago hirtella Humboldt, Bonpland & Kunth var. galeottiana (Decaisne) Pilger.
Mexican Plantain. Rare, in damp areas and in places where water seeps. Radio Road ad-
jacent to radio station near top of mountain, McC, Halde, June 23, 1963, W June 20, 1965;
Colma Canyon, K August 5, 1963. Perennial with broadly ovate leaves and flowers in spikes
5 to 6 (or more) inches long.

Plantago lanceolata Linnaeus. Ribgrass. English Plantain. Buckhorn. Weed of
roadsides and other disturbed areas, and in grassland. South of Crocker Avenue, P 2644;
Quarry at end of School Street and Hillside Boulevard, K October 7, 1963; frontage road
at northern city limits of South San Francisco, McC, K September 11, 1964; Crocker In-
dustrial Park, W June 8, 1965; Glen Park Canyon, McC April 16, 1967. Perennial with
slender lanceolate leaves and flowers in spikes to 3 inches long, April to October. The
flowers of this species have much exerted anthers. Native of Europe.

PLUMBAGINACEAE. Plumbago Family

Limonium californicum (Boissier) Heller. Marsh Rosemary. Sea Lavender. Rare,
salt flats along San Francisco Bay. Point San Bruno, K 1047. Perennial with a basal tuft of
leaves and small purplish flowers at the ends of a branched inflorescence, July to November.

POLEMONIACEAE. Polemonium Family

Gilia achilleaefolia Bentham. California Gilia. Occasional, open rocky slopes and
brush covered areas. Pig Ranch Ravine, W May 12, 1965; Crystal Cave Canyon, K June
20, 1965. Erect branched annual with slender stems and blue-violet flowers, February to July.

Gilia clivorum (Jepson) V. Grant. Occasional, open rocky slopes and grassland. Crocker
Hills, K 1003; Ridge Road near West Powerline, K 1036; Owl’s Canyon, K 1067; Buckeye
Canyon, W April 2, 1967; Crystal Cave Canyon, K June 20, 1965. Erect branched annual,
blue-violet flowers, March to June; similar to preceding species from which it differs by
its generally shorter corollas and more numerous seeds in the capsules.

Linanthus androsaceus (Bentham) Greene. Occasional, open grassy slopes, often in
colonies. Colma Canyon, Raven 1793; Glen Park Canyon, McC, W May 14, 1967; Owl’s
Canyon, K 1068. Slender, attractive annual with clusters of pink to white flowers, April
to July.

Navarretia squarrosa (Eschscholtz) Hooker & Arnott. Skunkweed. Occasional, dry
open slopes in often hard-packed soil, sometimes in grassland. Colma Canyon, K August 5,
1963; Quarry, K July 14, 1963; Brisbane Lagoon, W June 19, 1965. Annual with skunk-
like odor and heads of small blue flowers, June to October.

Phlox gracilis (Hooker) Greene. [Gilia gracilis Hooker; Microsteris gracilis (Hooker)
Greene]. Occasional, in grassland, sometimes recurring after fire. Two ravines east of Brisbane
Powerlines, McC March 21, 1965; Owl’s Canyon, K 1044; Devil’s Arroyo, McC March 28,
1965; Glen Park Canyon, McC April 16, 1967; Quarry, W April 2, 1967. Diminutive annual
with small pink flowers, February to April.

POLYGONACEAE. Buckwheat Family

Chorizanthe pungens Bentham var. hartwegii (Bentham) Goodman. Spine-flower.
Rare, sandy areas. Lower Colma Canyon, McC June 13, 1965. Prostrate annual with in-
conspicuous flowers in spiny heads, April to June.

642 CALIFORNIA ACADEMY OF SCIENCES [PRoc. 4TH SER.

Eriogonum latifolium Smith. Wild Buckwheat. Frequent, open rocky slopes, grassy
and brushy areas. Quarry at end of School Street near Hillside Boulevard, K October 7,
1963; adjacent to Olivet Cemetery, W May 18, 1965; Randolph Avenue at Hillside Boule-
vard, P 2717; Guadalupe Road, McC April 11, 1965; Point San Bruno, K September 2,
1964; Quarry, K July 14, 1963; above Harold Avenue, Brisbane, McC, W May 14, 1967.
Perennial with mostly basal leaves densely hairy on lower surface and small flowers in dense
heads, June to October.

Eriogonum nudum Douglas ex Bentham. Rare, dry open or rocky areas. Sierra Point,
W June 21, 1965. Similar in appearance to preceding except that stem is glabrous, flowers
June to August.

Eriogonum vimineum Douglas ex Bentham. Rare, rocky slopes. Canyon south of
Pig Ranch Ravine, W July 7, 1965. Annual branched above with few flowers in small
clusters, July to September.

Polygonum aviculare Linnaeus. Dooryard Knotweed. Weed of dry packed soil in

disturbed areas. Crocker Avenue, K August 5, 1963. Prostrate annual, stems and leaves
gray-green, small whitish flowers along stems, May to October. Native of Eurasia.

Polygonum paronychia Chamisso & Schlechtendal. Dune Knotweed. Rare, in sandy
soil. Colma Canyon, McC June 13, 1965. Prostrate perennial, stems and leaves gray-green,
small whitish flowers in terminal clusters, May to July.

Pterostegia drymarioides Fischer & Meyer. Occasional, on open and rocky or brushy
slopes, scrambling under the adjacent shrubs. Rocky slopes below summit, McC June 9,
1963; Bitter Cherry Ridge, K May 4, 1964; South Powerline, Menzies March 14, 1965;
Crystal Cave Canyon, K June 20, 1965; Colma Canyon, K 1216. Delicate prostrate, twining
annual with tiny inconspicuous flowers, March to June.

Rumex acetosella Linnaeus. Sheep Sorrel. Weed, frequent on open and brushy areas
or grassland. Glen Park Canyon, McC April 16, 1967; Colma Canyon, McC, Halde, K June
23, 1963; meadow between April Brook and Radio Road, McC March 28, 1965; Sierra Point,
W May 8, 1965; south of Crocker Avenue, P 2642. Perennial with hastately lobed leaves
and small reddish flowers in terminal panicles, March to September. Native of Eurasia.

Rumex californicus Rechinger f. California Dock. Rare, in grassland. Lower Buckeye
Canyon, K September 2, 1964; near Bayshore Freeway, Brisbane, W June 8, 1965. Coarse
perennial with lanceolate leaves and small greenish flowers on terminal branches, May to
August.

Rumex conglomeratus Murray. Green Dock. Weed of low moist areas. Colma Canyon,
K August 5, 1963; April Brook, W August 16, 1965. Tall coarse perennial. Flowers April to
October. Native of Europe.

Rumex crispus Linnaeus. Curly Dock. Weed of low moist places. Ravine south of
Crocker Avenue, P 2656; Guadalupe Road, K August 5, 1963; Point San Bruno, W March
23, 1965. Perennial with somewhat undulate leaf margins and small flowers, April to July.
Native of Eurasia.

Rumex obtusifolius Linnaeus. Bitter Dock. Weed of low moist places. Guadalupe
Road, K August 5, 1963. Stout perennial with often large leaves (to 12 inches long), small
flowers, May to September. Native of Europe.

VoL. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 643

Rumex pulcher Linnaeus. Fiddle Dock. Weed of waste places. Colma Canyon, McC,
Halde, K June 23, 1963; above Randolph Avenue at Hillside Boulevard, P 2728; ridge above
Buckeye Canyon, K July 14, 1963. Perennial with divaricately branched stem, and small
flowers, May to July. Native of Mediterranean Region.

Rumex transitorius Rechinger f. [R. salicifolius Weinmann forma transitorius (Rech-
inger f.) J. T. Howell]. Rare, in moist disturbed areas. Crocker Hills, McC, K September 2,
1963.

PorTULACACEAE. Portulaca or Purslane Family

Calandrinia ciliata (Ruiz & Pavon) DeCandolle var. menziesii (Hooker) Macbride.
Red Maids. Occasional, open places and disturbed areas, sometimes bordering grassland.
Glen Park Canyon, McC April 16, 1967; lower Colma Canyon, McC, Halde, K June 23,
1963; Flower Garden adjacent to April Brook, McC March 28, 1965; Pig Ranch Ravine,
W May 12, 1965; slope above Crocker Avenue, Toschi 63:36. Somewhat fleshy annual with
small but conspicuous magenta flowers, February to May.

Montia perfoliata (Donn ex Willdenow) Howell. Miner’s Lettuce. Frequent, in
moist areas and grassland. Colma Canyon, K May 30, 1964; Glen Park Canyon, McC April
16, 1967; near Nike Base, McC, W April 23, 1965; Bitter Cherry Ridge, K May 4, 1964;
Ridge Road, K February 28, 1965. Glabrous annual with stem leaves opposite and connate,
small white flowers, February to May.

Portulaca oleracea Linnaeus. Purslane. Weed of disturbed areas. Hillside Boulevard,
K October 7, 1963. Prostrate, fleshy, glabrous annual with small yellow flowers, May to
October. Native of Europe.

PRIMULACEAE. Primrose Family

Anagallis arvensis Linnaeus. Scarlet Pimpernel. Weed of disturbed areas and in grass-
land. Lower Colma Canyon, McC, Halde, K June 23, 1963; Sierra Point, W April 27, 1965;
near Nike Base, McC, W May 23, 1965; West Powerline, McC, K April 13, 1964. Diffusely
branched, slender annual with dainty salmon-colored flowers at most times throughout the
year. Native of Europe.

Dodecatheon hendersonii Gray subsp. cruciatum (Greene) Thompson. Shooting Star.
Occasional, on grassy slopes, but sparse at all localities on San Bruno Mountain. Radio Road,
McC, V. Ryder, February 16, 1964; Cable Ravine, K 865; West Powerline, K 951; Glen
Park Canyon, McC April 16, 1967. Perennial with basal rosette of leaves and flower stalk
terminated by several magenta flowers, February to April. One of our most attractive early
spring wildflowers. The strongly reflexed corolla lobes indicate the relationship of this
genus to the European Cyclamen which we know as a florist’s and garden plant. Ornamental.

RANUNCULACEAE. Buttercup Family

Aquilegia formosa Fischer var. truncata (Fischer & Meyer) Baker. Columbine. Rare,
moist, partially shaded brushland, only a few plants in each locality. Bitter Cherry Ridge,
K May 4, 1964; Columbine Gulch, W March 28, 1965. Attractive perennial with divided
leaves and nodding salmon-red flowers, April to June. Ornamental.

Delphinium californicum Torrey & Gray. Coastal Larkspur. Occasional, moist, partially
shaded brushland. Colma Canyon, McC, W June 13, 1965; Owl’s Canyon, K 1048; Roman-
zoffia Ravine, K May 4, 1964. Perennial to 4 to 5 feet tall, racemes densely flowered, flowers
dull bluish or lavender, April to July.

644 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47H SER.

Delphinium decorum Fischer & Meyer. Blue Larkspur. Occasional, in grassland. Colma
Canyon, K March 15, 1964; Bitter Cherry Ridge, K May 4, 1964; Owl’s Canyon, McC April
25, 1965; grassy meadow between April Brook and Radio Road, McC March 28, 1965; above
Harold Avenue, Brisbane, McC, W May 14, 1967. Perennial to about 12 inches tall, with
deep blue flowers, March to May. A very attractive spring wildflower.

Ranunculus californicus Bentham. California Buttercup. Frequent, in moist grassland.
Slope above Parking Lot, McC February 16, 1964; Kamchatka Point, McC June 9, 1963;
above Harold Avenue, Brisbane, McC May 14, 1967; Randolph Drive north of Hillside
Boulevard, P March 30, 1963; near Nike Base, McC, W May 23, 1965. Perennial with
fibrous roots, stems more or less hairy, sparsely branched, usually 1 to 2 feet tall, bright
yellow flowers, February to June, smooth achenes.

Ranunculus muricatus Linnaeus. Prickle-fruited Buttercup. Weed of wet ground and
seepage areas. Near Olivet Cemetery, W May 18, 1965; Colma Canyon, W June 16, 1965.
Annual or perennial with fibrous roots, stems glabrous, to about 12 inches tall, yellow flowers
February to June, achenes roughened with sharp points. Native of Europe.

Thalictrum polycarpum (Torrey) Watson. Meadow Rue. Occasional, in moist places
on brush covered slopes. Lower Colma Canyon, McC, Halde, K June 23, 1963; Bitter Cherry
Ridge, K April 4, 1965; Romanzoffia Ravine, K May 4, 1964. Perennial with attractive fern-
like leaves, much branched panicles with dainty yellowish flowers, March to May. Orna-
mental.

RHAMNACEAE. Buckthorn Family

Ceanothus thyrsiflorus Eschscholtz. Blue Blossom. Frequent, on brushy slopes, seedlings
sometimes appear after fire. Near West Powerline, McC, K September 2, 1963; Quarry,
K June 24, 1963; Ridge Road, 4% mile east of summit, P 2695. Evergreen shrub to 15 feet
or more, with leaves 3-veined from base and blue flowers, March to May. Ornamental.

Rhamnus californica Eschscholtz. California Coffee Berry. Frequent, on brushy slopes.
Regenerates by crown sprouts after fire. Base of mountain adjacent to Olivet Cemetery,
W May 18, 1965; Pig Ranch Ravine, K 1108; above April Brook and east of Nike Base,
McC April 23, 1965; Huckleberry Ridge, K May 4, 1964; Buckeye Canyon, K July 14, 1963;
West Powerline, McC, K September 2, 1963; north-facing slope south of Guadalupe Parkway
and west of its junction with Radio Road, McC, W June 11, 1967. Evergreen shrub with
inconspicuous flowers, May to July, and black berries. Ornamental, especially the more
compact low-growing plants.

Rhamnus crocea Nuttall. Redberry. Occasional, on brushy slopes and rocky outcrops.
Ridge south of Parking Lot, K May 4, 1964; Guadalupe Road, K August 5, 1963; Colma
Canyon, McC June 13, 1965. Small leaved, prostrate, compact, evergreen shrub with in-
conspicuous flowers February to May, and red berries. Ornamental.

ROSACEAE. Rose Family

Acaena californica Bitter. Frequent, grassland and rocky slopes. Above Harold
Avenue, Brisbane, McC, W May 14, 1967; near Nike Base, K March 15, 1964; Flower
Garden, McC March 28, 1965; slope below Parking Lot, McC, Halde June 23, 1963; South

=>
Pirate 4. Left. Rhamnus californica Eschscholtz, stump sprouting after fire.
Right. Baerta chrysostoma Fischer & Meyer.

( McCLINTOCK
PROC. CALIF. ACAD. SCI., 4TH SER., VOL.XXXII NO. 20 AND PLATE 4

KNI GHT)

V4 ’

: No 4

Vou. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 645

Ficure 9. Fragaria chiloensis (Linnaeus) Duchesne.

Powerline, McC March 14, 1965. Perennial with basal, pinnately divided leaves and incon-
spicuous flowers in clusters on flower stalks usually 12 to 18 inches tall, March to June.

Alchemilla occidentalis Nuttall ex Torrey & Gray. Western Lady’s Mantle. Occasional,
on rocky and brushy slopes. Because of its small size easily overlooked. South Powerline,
McC March 14, 1965; West Powerline, K 949. Inconspicuous annual with tiny flowers lack-
ing petals, February to May.

Amelanchier pallida Greene. Service Berry. June Berry. Occasional, on rocky and
brushy slopes. Stump sprouts after fire. Slope just below summit, McC June 9, 1963; Horse-
shoe Ridge, K August 5, 1963; above Harold Avenue, Brisbane, McC, W May 14, 1967; April
Brook, W June 16, 1965. Deciduous shrub with white flowers April and May. Ornamental.

Cotoneaster pannosa Franchet. Rare, an escape from cultivation. Colma Canyon, W
June 16, 1965. Evergreen shrub, lower leaf surface gray hairy, small pinkish flowers, June,
reddish fruits later in season. Native of China.

Crataegus monogyna Jacquin. Hawthorn. Rare, an escape from cultivation. North of
Crocker Avenue, W July 7, 1965; in eucalyptus forest, Crocker Avenue near Guadalupe
Road, K September 2, 1964. Deciduous small tree, with lobed leaves, pink flowers in spring,
red fruits later in season. Native of Europe and western Asia.

Fragaria californica Chamisso & Schlechtendal. California Strawberry. Frequent, in
grassland and on brushy and rocky slopes. Growing with Fragaria chiloensis along Radio
Road. Colma Canyon, K March 15, 1964; below Nike Base, McC, W May 23, 1965; Radio
Road, McC, W May 23, 1965; April Brook, McC March 28, 1965; Cable Ravine, McC March

646 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER.

14, 1965; ravine west of Radio Road, K 700; Kamchatka Point, McC May 1, 1966; below
summit in vicinity of radio station, McC, Menzies February 16, 1963. Attractive perennial,
spreading by runners, leaves thin, dull green above and with silky hairs beneath, white
flowers % to *4 inch across, January to June, fruit edible. Ornamental.

Fragaria chiloensis (Linnaeus) Duchesne. Beach Strawberry. Occasional, in sandy
and rocky areas. Radio Road, K August 5, 1963; ravine west of Radic Road, K 688; April
Brook, McC March 28, 1965. Similar to Fragaria californica in habit, leaves leathery, shining
and silky hairy beneath, flowers usually *4 inch or more across, February to August, fruit
edible. Ornamental, probably superior to Fragaria californica because of its thicker and
more glossy leaves and larger flowers.

Heteromeles arbutifolia (Aiton) Roemer [Photinia arbutifolia (Aiton) Lindley].
Toyon. Christmas Berry. California Holly. Frequent, in chaparral and wooded ravines, or
sometimes occurring as single individuals in open areas. Stump sprouts after fire. Colma
Canyon, McC June 13, 1965; Quarry, K June 24, 1963; Bitter Cherry Ravine, K 1015;
Guadalupe Road, K 1001; Cable Ravine, McC March 14, 1965. Evergreen shrub or small
tree; leaves leathery, mostly glabrous, margins toothed; small white flowers in large clusters,
June and July, red holly-like fruits in winter. Ornamental.

Holodiscus discolor (Pursh) Maximowicz. Cream Bush. Ocean Spray. Frequent, in
chaparral and brushy areas. Stump sprouts after fire. Colma Canyon, McC, Halde, K June
23, 1963; Radio Road near summit of mountain, K September 2, 1964; Horseshoe Ridge,
K August 5, 1963; Quarry, K June 24, 1963; Romanzoffia Ravine, K 1019; Trillium Gulch,
McC March 28, 1965; Buckeye Canyon, McC, W May 14, 1967. Deciduous shrub with small
white flowers in showy panicles, April to July. Ornamental.

Horkelia californica Chamisso & Schlechtendal. Occasional, in grassland. Colma Canyon,
McC June 13, 1965; Quarry, K July 14, 1963; Radio Road, McC, K September 2, 1963; Pig
Ranch Ravine, W May 11, 1965; slope below Parking Lot, McC, Halde June 23, 1963.
Herbaceous perennial, leaves mostly basal, pinnately compound; stems to 2 feet or more
with small white flowers, June to October. Similar in appearance to Potentilla glandulosa
and often confused with it.

Osmaronia cerasiformis (Torrey & Gray ex Hooker & Arnott) Greene. Oso-Berry.
Frequent, in chaparral and in moist wooded ravines. Stump sprouts after fire. Colma
Canyon, K March 13, 1964; Cable Ravine near Eucalyptus grove, McC March 14, 1965;
Quarry, K July 14, 1963; Devil’s Arroyo, McC March 28, 1965; Bitter Cherry Ravine,
K 1011; ravine near Parking Lot, P 2701; Buckeye Canyon, W May 7, 1965, and McC April
16, 1967. Deciduous shrub or small tree with small white flowers in nodding racemes, Febru-
ary to April, and fruit berry-like, red at first and black when mature. The soft wood of
this shrub has the pith divided into transverse chambers. Ornamental.

Physocarpus capitatus (Pursh) Kuntze. Ninebark. Rare, on wooded slopes. Quarry,
K June 24, 1963. Deciduous shrub with exfoliating bark, small white flowers in capitate
clusters, May to June. The shrubs at this locality on San Bruno Mountain vary in height
from about 7 to 8 inches to 5 feet, the taller shrubs being toward the lower part of the

ravine in more protected places and the lower ones where it is more open and windswept.
Ornamental.

Potentilla egedii Wormskjéld var. grandis (Torrey & Gray) J. T. Howell. Cinquefoil.
Occasional, in seepages and other wet places. Colma Creek in Colma Canyon, McC April

11, 1965; Guadalupe Road, K August 5, 1963; Radio Road, W May 23, 1965. Low perennial

Vout. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 647

with runners, silvery pinnately compound leaves in basal rosettes, and yellow flowers, April
to June. Ornamental for its silvery foliage and yellow flowers.

Potentilla glandulosa Lindley. Sticky Cinquefoil. Occasional, on grassy or brushy
slopes. Crocker Avenue, P 2634; Colma Canyon, K 1224; Cow Trough Ravine, K October
7, 1963; Quarry, K July 7, 1963; Flower Garden, K 978; Devil’s Arroyo, McC March 28,
1965; above Harold Avenue, Brisbane, McC, W May 14, 1967; slopes below summit of
mountain, McC June 9, 1963. Herbaceous perennial with mostly basal pinnately compound
leaves, stems to 2 feet or more, and small yellowish flowers, March to July. Similar in: ap-
pearance to Horkelia californica from which it is distinguisheed by technical characters
of flowers.

Prunus demissa (Nuttall) Walpers. Western Choke Cherry. Occasional, in chaparral
and wooded ravines. Upper Colma Canyon, K April 11, 1965; lower Colma Canyon, McC,
Halde, K June 23, 1963; Devil’s Arroyo, Roof April 25, 1965; Sierra Point, W May 12, 1965.
Deciduous shrub with small white flowers in showy racemes, April to June. Ornamental.

Prunus emarginata (Douglas) Walpers. Bitter Cherry. Rare, on brushy slopes. Stump
sprouts after fire. Bitter Cherry Ridge, K, Roof May 15, 1965; Glen Park Canyon, McC, W
May 14, 1967. Deciduous shrubs with small white flowers in showy umbellate clusters, April
and May. Ornamental.

Prunus ilicifolia (Nuttall ex Hooker & Arnott) Walpers. Holly-leaved Cherry. Islay.
Occasional, brushy and wooded ravines. Crocker Hills, McC, K September 2, 1963; Dairy
Ravine, K August 5, 1963; Wax Myrtle Ravine, K August 5, 1963; between Devil’s Arroyo
and Buckeye Canyon, W June 7, 1963; Buckeye Canyon, K July 14, 1963; Sierra Point,
W May 12, 1965. Evergreen shrub with holly-like leaves and small white flowers in showy
racemes, March to June. Ornamental.

Rosa californica Chamisso & Schlechtendal. California Wild Rose. Occasional, on brushy
areas. Stump sprouts after fire. Point San Bruno, K September 2, 1964; Romanzoffia
Ravine, K May 4, 1964; Buckeye Canyon, W June 7, 1965. Deciduous shrub with stout flat-
tened usually recurved prickles, attractive pink flowers in corymbs, April to October, petals
usually % to 1 inch long.

Rosa gymnocarpa Nuttall ex Torrey & Gray. Wood Rose. Frequent, on brushy slopes.
Stump sprouts after fire. Colma Canyon, K May 30, 1964; Quarry, K June 24, 1963; Bitter
Cherry Ridge, K May 4, 1964; Owl’s Canyon, Roof April 25, 1965; Romanzoffia Ravine,
K May 4, 1964; Buckeye Canyon, W June 7, 1965; east facing slope below Parking Lot,
McC, Halde June 23, 1963. Slender-stemmed deciduous shrub with many terete mostly
straight prickles, dainty pink flowers, April to September, usually solitary, petals less than
Y% inch long. Ornamental.

Rubus parviflorus Nuttall var. velutinus (Hooker & Arnott) Greene. Thimble Berry.
Occasional, on moist brushy slopes. Stump sprouts after fire. Colma Canyon, K May 30,
1964; Romanzoffia Ravine, K May 4, 1964; Columbine Gulch, McC March 28, 1965; Quarry,
K June 24, 1964. Deciduous shrub lacking prickles, leaves 5-lobed, flowers white to pink,
March to August, fruit an edible red thimble-berry. Ornamental.

Rubus procerus P. J. Mueller ex Boulay. Himalaya Berry. Aggressive weedy shrub
forming roadside thickets. Colma Canyon, K August 5, 1963; Guadalupe Road, K September
2, 1964; Crocker Avenue, W July 7, 1965. Robust, more or less evergreen, sprawling, prickly
shrub, with 5 leaflets white hairy beneath, white or pink flowers, June to August, in large

648 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

terminal panicles, edible blackberries. Native of Europe. Its rampant growth makes this
shrub undesirable but its berries are delicious.

Rubus spectabilis Pursh var. franciscanus (Rydberg) J. T. Howell. Salmon Berry.
Occasional, on moist brushy slopes forming thickets. Colma Canyon, K April 11, 1965; ravine
to west of summit of mountain, K 692. Deciduous shrub, young stems with prickles and
3-{foliolate leaves, flowers single, red-purple, March to May, edible fruits reddish to salmon-
colored. Ornamental.

Rubus ursinus Chamisso & Schlechtendal. California Blackberry. Pacific Blackberry.
Frequent, forming brushy thickets by trailing on ground and over and among other shrubs.
Crocker Avenue, McC, K January 12, 1964; Colma Canyon, K May 30, 1964; ravine west
of Radio Road, K 696; near Nike Base, McC, W May 23, 1965; Owl’s Canyon, McC April
25, 1965. Scandent shrub with prickles, leaves white hairy at least when young, 3-foliolate on
young stems, mostly 3-lobed on flowering stems, flowers white in small clusters March to
August, edible blackberries. Cultivated for its excellent edible fruits and has entered into the
development of the Loganberry, the Youngberry, and the Boysenberry.

RuBIAcEAE. Madder Family

Coprosma repens J. D. Hooker. Mirror Shrub. Escapes from cultivation and persists
in waste places. Crocker Avenue, McC September 24, 1967. Evergreen shrub to 8 to 10 feet
tall with shining leaves, inconspicuous flowers and orange berry-like fruits, late summer to
autumn. Native of New Zealand.

Galium aparine Linnaeus. Bedstraw. Goose Grass. Cleavers. Frequent, scrambling
over other plants, mostly in brushy and shaded areas. Buckeye Canyon, McC April 16, 1967;
Colma Canyon, K May 30, 1964; near Nike Base, McC, W May 23, 1965; April Brook,
McC March 28, 1965; ravine west of Radio Road, K April 25, 1964; Quarry, K July 14,
1963; east facing slope below Parking Lot, McC, Halde June 23, 1963; Reservoir Hill, McC
March 14, 1965; Randolph Drive at Hillside Boulevard, P 2748; South Powerline, McC
March 14, 1965. Weak stemmed annual with usually linear leaves more than % inch long,
6 to 8 in a whorl, white flowers, March to August, dry fruits with hooked bristles. Nativity
is doubtful, may be introduced from Europe.

Galium californicum Hooker & Arnott. California Bedstraw. Occasional, shaded
slopes and brushy areas. East facing slope below Parking Lot, McC, Halde June 23, 1963;
canyon below radio towers, W May 12, 1965; East Powerline, K March 21, 1965. Perennial,
stems more or less erect but intertwining and clumping together, leaves mostly ovate and
longer than 4 inch, in 4’s, yellow-green flowers May to July, fruits fleshy, glabrous or hairy.

Galium nuttallii Gray. Climbing Bedstraw. Frequent, clambering over adjacent
plants, mostly on dry brushy slopes. Randolph Drive at Hillside Boulevard, P 2722; Reser-
voir Hill, McC March 14, 1964; lower Colma Canyon, McC, Halde, K June 23, 1963; near
Nike Base, K 1230; east of Brisbane Powerlines, K 965; above Harold Avenue, Brisbane,
McC, W May 14, 1967. Perennial, more or less woody at base, with stems to as much as
6 feet long, leaves mostly oval and less than 14 inch long, in 4’s, yellow-green flowers March
to August, fruit fleshy, glabrous.

SALICACEAE. Willow Family

Salix coulteri Andersson. Coulter Willow. Occasional, streams and seepage areas. Colma
Canyon, K August 5, 1963; junction of Colma Creek and April Brook, K March 28, 1965;
ravine south of Crocker Avenue, P 2640; Crocker Industrial Park, W October 16, 1966;

Vout. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 649

northern South San Francisco, McC, K September 11, 1964. Small tree with entire leaves
which are densely silvery hairy on lower surfaces, flowers in catkins, March to May.

Salix lasiolepis Bentham. Arroyo Willow. Common, streams and seepage areas. Crown
sprouts after fire. Lower Colma Canyon, McC March 30, 1963; Radio Road at Summer
Seep, K 862; above the Quarry, K July 14, 1963; Cow Trough Ravine, K October 7, 1963;
Devil’s Arroyo, McC March 28, 1965; northern South San Francisco, McC, K September 11,
1964; Guadalupe Valley along a creek bank, P March 30, 1963. Small tree with leaves nearly
entire and somewhat revolute on the margins, glaucous and more or less glabrous beneath,
flowers in catkins March to May.

Salix lasiolepis and S. coulteri hybridize occasionally and plants apparently of hybrid
origin with varying amounts of tomentum on the lower leaf surface are found in Colma
Canyon and at the junction between April Brook and Colma Creek.

SAXIFRAGACEAE. Saxifrage Family

Grossularia californica (Hooker & Arnott) Coville & Britton [Ribes californicum
Hooker & Arnott]. California Gooseberry. Hillside Gooseberry. Occasional, on shrubby or
wooded areas. Seedlings appear after fire. Guadalupe Road in Crocker Hills, K January
15, 1964; above the Quarry, K July 14, 1963; Devil’s Arroyo, McC March 28, 1965; Buck-
eye Canyon, W May 5, 1963. Deciduous shrub with nodal spines, reddish-green flowers
January to April, and prickly fruits.

Grossularia divaricata (Douglas) Coville & Britton [Ribes divaricatum Douglas]. Rare,
on shrubby slope. Colma Canyon, K May 4, 1964. Differs from Grossularia californica in
its mostly greenish and rather inconspicuous flowers February to May, and fruits lacking
prickles.

Grossularia leptosma Coville [Ribes menziesii Pursh var. leptosmum (Coville) Jepson].
Bay Gooseberry. Canyon Gooseberry. Buckeye Canyon, K August 5, 1963; above the
Quarry, K July 14, 1963. Deciduous shrub with nodal spines, leaves glandular hairy beneath,
reddish flowers March to June, and prickly fruits.

Heuchera micrantha Douglas ex Lindley. Alum Root. Occasional, shaded and brushy
slopes. Bitter Cherry Ravine, K May 15, 1965; Quarry, K July 14, 1963; Romanzoffia
Ravine, K May 4, 1964; Owl’s Canyon, K 1051; Dairy Ravine, K August 5, 1963. Perennial
herb with mostly basal leaves, small greenish-white flowers, May and June, having five
shortly exerted stamens. Ornamental.

Lithophragma affinis Gray. Woodland Star. Occasional, in moist wooded or open
rocky areas. Romanzoffia Ravine, K March 14, 1965; between Romanzoffia Ravine and
Trillium Gulch, K 969; Flower Garden, McC March 28, 1965; Bitter Cherry Ravine, K April
4, 1965. Slender perennial with basal leaves and white flowers, February to May, having
petals about 14 inch long and 10 stamens.

Ribes malvaceum Smith. California Black Currant. Chaparral Currant. Rare, wooded
ravine. Buckeye Canyon, K 944. Deciduous shrub with rose-pink flowers in racemes, Octo-
ber to March, fruits purple-black. The stems of the currants lack the nodal spines which
are present in the gooseberries, sometimes the two are united into a single genus.

Saxifraga californica Greene. California Saxifrage. Occasional, in moist grassland
or brushy or rocky areas. Colma Canyon, K 1247; Cable Ravine, Menzies March 14, 1965;
Devil’s Arroyo, McC March 28, 1965; ravine %4 mile east of Brisbane Powerlines, McC
March 21, 1965. Slender perennial with mostly basal leaves, flowering stems about 12 inches
tall, small flowers, February to April, petals white, 4% inch long, 10 stamens.

650 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Tellima grandiflora (Pursh) Douglas ex Lindley. Fringe Cups. Occasional, in moist
shrubby areas, seepage and stream banks. Creek bank in Colma Canyon, McC April 11,
1965; Romanzoffia Ravine, K April 25, 1964; north facing slope south of Guadalupe Park-
way near junction with Radio Road, McC, W June 11, 1967. Stout perennial with mostly
basal cordate leaves, cup-shaped flowers, March to June, petals whitish then turning red,
margins laciniate or fringed, hence the common name.

SCROPHULARIACEAE. Figwort Family

Bellardia trixago (Linnaeus) Allioni. Weed of roadsides and disturbed areas. Randolph
Avenue, K May 16, 1967; Crocker Industrial Park, Brisbane, W June 19, 1965; junction of
Radio Road and Crocker Avenue, W June 16, 1965; Sierra Point, W April 27, 1965. Coarse
annual, but attractive, weed with rose-pink and white flowers. Native of Mediterranean
region.

Castilleja affinis Hooker & Arnott. Indian Paint Brush. Occasional, in brushy or
open grassy and rocky areas. Randolph Drive at Hillside Boulevard, P 2740; near Nike
Base, McC, W May 23, 1965; Flower Garden near April Brook, McC February 28, 1965.
Perennial with red bracts, flowers March to September. Differs from Castilleja franciscana
by having the lower lip of the corolla not exerted.

Castilleja franciscana Pennell. Franciscan Paint Brush. Occasional, in grassland
and rocky areas. South side of Ridge Road near powerlines, K 921; Colma Canyon, Raven
1775; Guadalupe Road in Crocker Hills, K 1098; near Olivet Cemetery, McC June 13, 1965;
near Cow Palace in Crocker Hills, W May 17, 1965. Perennial with more or less hairy but
not glandular-hairy stems and leaves, and red bracts, flowers March to September, slender
yellow-green lower lip of corolla much exerted.

Castilleja wightii Elmer. Wight’s Paint Brush. Frequent, grassland, brushy and rocky
areas. Colma Canyon, Raven 1778; Quarry, K July 14, 1963; Flower Garden near April
Brook, McC March 28, 1965; near Nike Base, McC, W May 23, 1965; Trillium Gulch, McC
March 28, 1965; slope below Parking Lot, McC June 9, 1963; West Powerline, McC, K
September 2, 1963; South Powerline, K 947; Guadalupe Road in Crocker Hills, K 1013.
This is the most commonly seen of the paint-brushes on San Bruno Mountain. It differs
from the two preceding species in being glandular-hairy especially in the inflorescence and
somewhat viscid to the touch. Its bracts are either yellow or dull reddish, flowers March
to October.

Collinsia franciscana Bioletti. Chinese Houses. Occasional, in open grassy, or brushy
areas. Colma Canyon, K May 4, 1964; Bitter Cherry Ravine, K 1014; Devil’s Arroyo, McC
March 28, 1965; ravine east of Brisbane Powerlines, McC March 21, 1965; above Harold
Avenue, Brisbane, McC, W May 14, 1967. Annual with whorls of sessile, violet-blue and
whitish flowers, March to May.

Collinsia heterophylla Buist ex Graham. Chinese Houses. Rare, in shaded grassland.

Buckeye Canyon, W May 7, 1965. Annual with whorls of sessile lavender and white flowers,
April to July.

Collinsia sparsiflora Fischer & Meyer var. solitaria (Kellogg) Newsom. Chinese
Houses. Occasional, in grassland. Ravine east of Brisbane Powerlines, McC March 21, 1965;

>
PuaTeE 5. Upper. Artemisia californica Lessing, on slope in Owl’s Canyon.
Lower. Eschscholzia californica Chamisso.

( McCLINTOCK

PLATE 5

AND
KNIGHT)

PROG, “CATIF. ACAD. SC!;, 41TH SER., VOL.XxxII No. 20

a sae ls am

Vor. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 651

Owl’s Canyon, K 1069; Buckeye Canyon, W April 2, 1967. Annual with mostly single small
blue flowers, March to May, borne on pedicels.

Diplacus aurantiacus (Curtis) Jepson. Sticky Monkey Flower. Bush Monkey Flower.
Common, in dry brushland and open areas. Regenerates by crown sprouts after fire. Cow
Trough Ravine, K October 7, 1963; Dairy Ravine below radio tower, K August 8, 1963; Pig
Ranch Ravine, W May 12, 1965; ridge above Owl’s Canyon, W June 7, 1965; Quarry,
K June 14, 1963; east facing slope below Parking Lot, McC, Halde June 23, 1963; Gladys
Canyon, Brisbane, W June 14, 1965; above Harold Avenue, Brisbane, McC, W May. 14,
1967. Evergreen shrub with large and showy yellow-orange flowers March to July. Orna-
mental.

Linaria texana Scheele [L. canadensis (Linnaeus) Dumont de Courset var. texana
(Scheele) Pennell]. Blue Toad Flax. Occasional, sandy soil in open areas. Colma Canyon,
K April 11, 1965; west of Pig Ranch Ravine, W May 11, 1965; Manzanita Dike, K 971.
Slender annual with violet-blue flowers, March to May, spur on the corolla. Linaria texana
has a wide distribution outside of California, it goes northward to British Columbia and
eastward to the southeastern states.

Mimulus guttatus DeCandolle. Monkey Flower. Occasional, in seepage and wet areas.
Lower Colma Canyon, McC, Halde, K June 23, 1963; Radio Road at Summer Seep, K 1105;
Quarry, W June 19, 1965; Devil’s Arroyo, McC March 28, 1967; Glen Park Canyon, McC
April 16, 1967; ravine north of Randolph Drive and Hillside Boulevard, P 2744. Perennial
herb with showy bright yellow flowers, March to August.

Orthocarpus densiflorus Bentham. Owl’s Clover. Frequent, grassy and brushy areas.
Colma Canyon, Raven 1776; Flower Garden, K March 28, 1965; April Brook, K May 15,
1965; near Nike Base, W May 23, 1965; Pig Ranch Ravine, W May 11, 1965; summit above
Owl’s Canyon, K 1045; Romanzoffia Ravine, K March 13, 1965; above Harold Avenue,
Brisbane, McC, W May 14, 1967. Annual with purple flowers in terminal clusters, March
to May.

Orthocarpus erianthus Bentham var. erianthus. Butter-and-eggs. Johnny Tuck.
Rare, in grassland. Colma Canyon, W June 13, 1965. Annual often forming colonies, with
yellow flowers, March to May.

Orthocarpus erianthus Bentham var. roseus Gray. Frequent, in grassland, usually
forming colonies. Ravine south of Crocker Avenue, P 2657; Pig Ranch Ravine, K 1030;
Flower Garden, K 977; ravine west of Radio Road, K 1244; Crocker Industrial Park, W
March 27, 1966. Annual with whitish flowers aging rose-pink, March to May.

Orthocarpus faucibarbatus Gray var. albidus (Keck) Howell. Rare, in sandy soil.
Sierra Point, W May 2, 1965. Annual with whitish flowers, April to June.

Orthocarpus floribundus Bentham. Rare, in moist grassland. Ravine north of junc-
tion of Randolph Drive and Hillside Boulevard, P 2734; Point San Bruno, K April 25, 1965.
Annual with whitish flowers, April and May. Orthocarpus floribundus occurs only in Marin,
San Francisco and San Mateo counties.

Orthocarpus purpurascens Bentham var. latifolius Watson. Escobita. Frequent, in
grassland and open fields. Quarry, K June 24, 1963; Colma Canyon, McC June 13, 1965;
near Nike Base, K 1234; Radio Road, K August 5, 1963; slope below Parking Lot, McC
June 9, 1963; Kamchatka Point, K May 15, 1965; Ridge Road near West Powerline, K 968;
above Harold Avenue, Brisbane, McC, W May 14, 1967. Annual, stems and leaves hairy,
stems usually purplish and branched, flowers purplish March to May.

652 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Orthocarpus pusillus Bentham. Occasional, in open and grassy areas, sometimes in
hard packed soil and then somewhat depauperate. West Powerline, K 876; Glen Park
Canyon, McC April 16, 1967. Small slender annual with foliage brownish tinged, tiny in-
conspicuous flowers, April and May.

Scrophularia californica Chamisso & Schlechtendal. California Bee Plant. Frequent,
in brushland and bordering it. Lower Colma Canyon, McC July 18, 1967; Quarry, McC
August 1, 1965; east facing slope below Parking Lot, McC April 17, 1964; West Powerline,
McC March 28, 1965. Coarse perennial to as much as 5 to 6 feet tall with small insignificant
reddish-brown flowers February to July. Tends to become weedy.

Veronica americana (Rafinesque) Schweinitz ex Bentham. American Brooklime. Oc-
casional, in seepages and other wet areas. Colma Canyon, W June 16, 1965; north facing
slope south of Guadalupe Parkway, McC, W June 11, 1967; west of Radio Road, K June
20, 1965; canyon east of Quarry, W June 8, 1965; April Brook, W May 18, 1965. Erect
perennial with small bright blue flowers, May to October.

Veronica persica Poiret. Rare, in disturbed sandy soil. Lower Colma Canyon, McC,
K, Kasapligil July 18, 1967. Procumbent annual with small blue flowers, March to September.
Native of Eurasia.

SOLANACEAE. Nightshade Family

Solanum furcatum Dunal. Nightshade. Weed in disturbed areas. Crocker Hills, McC,
K September 2, 1963; Pig Ranch Ravine, W May 11, 1965; adjacent to Olivet Cemetery,
W May 18, 1965; on summit near Parking Lot, McC, Halde June 23, 1963. Weak-stemmed,
straggling perennial, with insignificant white to pale lavender flowers April to October,

corolla about °& inch or more across. Native of South America.

Solanum nodiflorum Jacquin [has been called S. nigrum by some authors but it is
not S. nigrum Linnaeus]. Nightshade. Weed in disturbed areas. Guadalupe Road, K August
5, 1963; Wax Myrtle Ravine, K August 5, 1963; near Olivet Cemetery, W August 18, 1965.
Weak stemmed straggling annual or perennial, with small insignificant pale lavender or
whitish flowers, March to December, corolla about ™% inch across. Nativity uncertain, but
it is probably South America.

Solanum umbelliferum Eschscholtz. Blue Witch. Frequent, on brushy slopes. Slopes
southeast of 1314-foot summit, P 2678; Quarry, K June 24, 1963; South Powerline, Menzies
March 14, 1965. Subshrub, more or less rounded in habit, hairy, with blue flowers January
to September.

TROPAEOLACEAE. Tropaeolum Family

Tropaeolum majus Linnaeus. Garden Nasturtium. Escape from cultivation and per-
sisting in waste places. Crocker Avenue, McC September 24, 1967. Trailing annual or peren-
nial with peltate leaves and attractive yellow-red flowers during most of the year. Native
of South America

UMBELLIFERAE. Carrot or Parsley Family

Angelica hendersonii Coulter & Rose. Occasional, in brushland. Regenerates after fire
by sprouting from rootstocks. Quarry, K July 14, 1963; Romanzofiia Ravine, McC March
28, 1965. Also in Colma Canyon. Stout perennial to 3 to 4 feet tall with large pinnately
compound leaves, each leaflet more or less ovate and pinnately veined, white flowers April

Vout. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 653

Ficure 10. Heracleum maximum Bartram.

to July. Superficially resembles Heracleum maximum but may be distinguished by the
shape of the leaves.

Anthriscus caucalis Bieberstein [A. scandicina (Weber) Mansfeld]. Bur-chervil. Weed
of disturbed areas. Guadalupe Road, K August 5, 1963; Sierra Point, W May 12, 1965.
Annual or biennial with finely divided leaves, small white flowers April to June, and fruits
with little hooks. Native of Europe.

Apiastrum angustifolium Nuttall. Wild Celery. Rare, in sandy soil. Pig Ranch Ravine,
K 1200. Annual with finely divided leaves, small white flowers February to May, and fruits
with inconspicuous ribs.

Bowlesia incana Ruiz & Pavon. Occasional, in grassy and brushy areas. Devil’s Arroyo,
K 1099; Trillium Gulch, K 1107. Delicate, weakly trailing, prostrate annual with tiny
inconspicuous flowers, March to May. Superficially resembles Pterostegia drymarioides,
another weakly trailing annual.

Caucalis microcarpa Hooker & Arnott. California Hedge Parsley. Rare, on open
grassy slopes. South Powerline, K 1215. Hispid annual, leaves finely divided, flowers small,
white, April to June, and fruits with numerous hooks.

Conium maculatum Linnaeus. Poison Hemlock. Spotted Hemlock. Weed of disturbed
areas. Crocker Hills, McC, K September 2, 1963. Also in upper Colma Canyon. Tall bien-
nial, stems purple spotted, leaves pinnately divided, flowers white, small, April to August,
fruits with prominent ribs.

Native of Europe. The plant is poisonous, more so when fresh than when dry, and by
it “criminals and philosophers were put to death at Athens.” It was said to have been
used to execute Socrates.

654 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47H SER.

Daucus pusillus Michaux. Rattlesnake Weed. Common, in grassland. Quarry, K July
14, 1963; Devil’s Arroyo, K April 25, 1965; Guadalupe Road, McC April 11, 1965; Radio
Road in eucalyptus area, McC, W June 11, 1967; Nike Base, K April 25, 1964; Colma
Canyon, McC June 13, 1965. Annual, leaves finely divided, flowers small white or pinkish,
April to June, fruits with stout hooks.

Foeniculum vulgare Miller. Sweet Fennel. Weed in disturbed areas. Guadalupe Road,
K August 5, 1963; Brisbane Lagoon, W August 14, 1965. Biennial or perennial with anise-
like odor, leaf divisions linear, flowers yellow, May to October, fruits smooth, angled. Native
of Europe where cultivated edible forms have been selected and used as a vegetable especially
in Italy.

Heracleum maximum Bartram [H. lanatum Michaux]. Cow Parsnip. Occasional, on
open slopes and in bases of ravines. Romanzoffia Ravine, K May 4, 1964; top of mountain
near radio station, McC, Menzies February 16, 1963; Crocker Avenue, P 2660; southwest
of Brisbane, P 2761; Colma Canyon, K May 30, 1964. Stout coarse perennial, to 5 to 6
feet tall, leaves ternately lobed with broad leaflets palmately veined and lobed, flowers white
in large flat-topped clusters April to July, fruits with narrow wings. See Angelica.

Ligusticum apiifolium (Nuttall) Gray. Lovage. Occasional, on open and exposed
grassy or brushy areas. Sometimes root sprouts after fire. Quarry, K June 24, 1963; Kam-
chatka Point, McC March 21, 1965; Nike Base, K 690; east facing slope below Parking Lot,
McC, Halde June 23, 1963; West Powerline, McC March 28, 1965. Perennial, leaves pin-
nately divided, flowers whitish, April to June, turning rusty in age, fruits smooth but ribbed.

Lomatium caruifolium (Hooker & Arnott) Coulter & Rose. Alkali Parsnip. Occasional,
in grassy areas. Radio Road, McC, Ryder February 16, 1964; Randolph Drive at Hillside
Boulevard, P 2731; above Harold Avenue, Brisbane, McC, W May 14, 1967. Acaulescent
perennial with ternately divided leaves, yellow flowers, March and April, fruits broadly
winged.

Lomatium dasycarpum (Torrey & Gray) Coulter & Rose. Lace Parsnip. Frequent, in
grassy and rocky areas. Colma Canyon, Raven 1789; ravine north of Randolph Avenue at
Hillside Boulevard, P 2745; ravine south of Crocker Avenue, P 2647; Quarry, K July 14,
1963; top of mountain on slope above Parking Lot, McC, Ryder February 16, 1964; Radio
Road near radio station, McC, Menzies February 16, 1963; Guadalupe Road, McC April 11,
1965; above Harold Avenue, Brisbane, McC, W May 14, 1967. Perennial differing from L.
caruifolium by its woolly fruits.

Lomatium utriculatum (Nuttall) Coulter & Rose. Bladder Parsnip. Occasional, on
open grassy slopes and ridges. Radio Road near radio station, McC, Menzies February 16,
1963; Reservoir Hill north of Colma Canyon, K 878; Cable Ravine, McC March 14, 1965;
Brisbane Powerlines, K 954; Guadalupe Road in Crocker Hills, K April 11, 1965; meadow
between April Brook and Radio Road, McC March 28, 1965. Perennial, differs from two
previous species by the thin wings of the fruit being broader than the main body of the fruit.

Oenanthe sarmentosa Presl. Occasional, in sluggish streams and marshy areas. Colma
Canyon, K August 5, 1963; Pig Ranch Ravine, K May 30, 1964; Guadalupe Creek, K April
11, 1965; April Brook, K 982. Perennial with pinnately compound leaves, each leaflet toothed
or lobed, white flowers May to October, fruits with corky ribs.

Sanicula arctopoides Hooker & Arnott. Footsteps-of-spring. Yellow mats. Snake-root.
Common, on open grassy slopes and windswept ridges. Above Harold Avenue, Brisbane,
McC, W May 14, 1967; Radio Road, McC, Ryder February 16, 1964; Cable Ravine, McC

Vor. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 655

March 28, 1965; meadow between April Brook and Radio Road, McC March 28, 1965;
Radio Road near radio station, McC, Menzies February 16, 1963; Kamchatka Point, McC
May 1, 1966; Crocker Avenue, P 2646. One of the mountain’s colorful spring wildflowers.
Prostrate perennial, entire plant a chartreuse-yellow color when in flower, flowers small, in
rounded clusters, February to May, surrounded by conspicuous bracts, seeds with few hooks.

Sanicula bipinnatifida Douglas ex Hooker. Purple Sanicle. Occasional, in grassland
and open slopes. Above Harold Avenue, Brisbane, McC, W May 14, 1967; Owl’s Canyon,
K 1065; Randolph Avenue, P 2726. Stout perennial with pinnately compound leaves, usually
purple flowers in a tight head without bracts, February to May, seeds with hooks.

Sanicula crassicaulis Poeppig ex DeCandolle. Pacific Sanicle. Common, on open or
shaded and wooded slopes. Colma Canyon, A1220; east facing slope below Parking Lot,
McC, Halde June 23, 1963; west facing slope below mountain summit, K 694; Quarry, K
July 14, 1963; West Powerline, K 707; Trillium Gulch, K 1020; Owl’s Canyon, K 1050;
Bitter Cherry Ridge, K April 15, 1965. Perennial with palmately lobed leaves, yellow flowers
in tight heads with tiny bracts, February to June, seeds with hooks.

Sanicula laciniata Hooker & Arnott. Coast Sanicle. Rare, on open exposed slope. Below
top of mountain near Parking Lot, McC June 9, 1963. Perennial with palmately lobed leaves,
leaf divisions laciniate, flowers yellow in tight heads with small bracts, April to June, seeds
with hooks.

UrTICACEAE. Nettle Family

Soleirolia soleirolii (Requien) Dandy. [Helxine soleirolii Requien]. Baby’s Tears.
Weed in shaded disturbed areas. Crocker Avenue under Monterey cypress, K September 2,
1964. Delicate creeping mat-forming perennial with inconspicuous flowers during most of
the year. Native of Corsica and Sardinia.

Hesperocnide tenella Torrey. Rare, in moist shaded areas. Colma Canyon, McC,
Halde, K June 23, 1963. Weak-stemmed but erect annual with stinging hairs and incon-
spicuous flowers, March to May. Nettle-like in appearance.

Urtica holosericea Nuttall. Nettle. Rare, along banks of creeks. Buckeye Canyon,
K 725; Guadalupe Creek, McC, K September 11, 1964. Stout coarse perennial with stinging
hairs and inconspicuous flowers, May to October.

VALERIANACEAE, Valerian Family

Centranthus ruber (Linnaeus) DeCandolle. Jupiter’s Beard. Red Valerian. Weed in
disturbed ground. Brisbane Lagoon, W August 14, 1965. Attractive perennial with white,
pink, and rose-pink flowers from late winter into autumn. A garden plant which escapes
from cultivation. Native of the Mediterranean region.

Plectritis congesta (Lindley) DeCandolle. Occasional, in grassland and moist brushy
areas, sometimes forms small colonies. Meadow between Radio Road and April Brook,
McC March 28, 1965; Romanzoffia Ravine, K April 24, 1964; Devil’s Arroyo, McC March
28, 1965; Owl’s Canyon, K 1095; Glen Park Canyon, McC April 16, 1967. Annual with
small pink flowers, February to May, in spike-like inflorescences.

VIOLACEAE. Violet Family

Viola adunca Smith. Blue Violet. Frequent, in grassland. Colma Canyon, K March
15, 1964; Flower Garden adjacent to April Brook, K 978; Cable Ravine, McC March 14,

656 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER.

1965; west facing slope below top of mountain, McC April 17, 1964. Low growing perennial
with violet flowers, February to April. This attractive violet resembles the cultivated
English violet of gardens.

Viola pedunculata Torrey & Gray. Johnny-Jump-Up. California Golden Violet.
Frequent, in grassland. Point San Bruno, K 1026; Randolph Drive at Hillside Boulevard,
P 2716; Glen Park Canyon, McC April 16, 1967; above Harold Avenue, Brisbane, McC, W
May 14, 1967; Cable Ravine, McC March 14, 1965; Flower Garden, McC March 28, 1965;
ridge above Great Meadow, W June 20, 1965. Perennial with yellow flowers, March and
April, the two upper petals are marked with red-brown on the back. An attractive spring
wildflower.

SUBCLASS MONOCOTYLEDONEAE
ARACEAE. Calla Family

Zantedeschia aethiopica (Linnaeus) Sprengel. Calla. Calla-Lily. An escape from
cultivation in damp sandy soil. Pig Ranch Ravine, K April 25, 1965; W May 14, 1965.
Native of South Africa.

CYPERACEAE. Sedge Family

Carex barbarae Dewey. Occasional, wet grassy ground. Colma Canyon, McC, K June
23, 1963; Radio Road, W May 23, 1965; north city limits of South San Francisco in seepage
area with Salix coulteri and S. lasiolepis, McC, K September 11, 1964.

Carex brevicaulis Mackenzie. Rare, on exposed slopes. J. T. Howell 13774.
Carex densa (Bailey) Bailey. Rare, in seepages. Crocker Hills, K May 4, 1964.

Carex obnupta Bailey. Rare, wet ravines. Crystal Cave Canyon, W June 19, 1965; Cow
Trough Ravine, K October 7, 1963.

Carex subbracteata Mackenzie. Rare, in wet ravine. Crystal Cave Canyon, K June 20,
1965.

Carex tumulicola Mackenzie. Rare, on grassy slope. Olivet Cemetery, W May 18, 1965.

Cyperus eragrostis Lamarck. Occasional, in marshy places and roadside ditches. Buckeye
Canyon, K September 2, 1964; Cow Trough Ravine, K October 7, 1963; Crocker Avenue,
K August 5, 1963; Crocker Hills, K October 4, 1966; Crocker Industrial Park, Brisbane,
W June 19, 1965; Sierra Point, W May 8, 1965.

Scirpus californicus (C. A. Meyer) Steudel. California Tule. Rare, in marsh, growing
with cattails. Near Bayshore Freeway, K October 4, 1966.

Scirpus cernuus Vahl var. californicus (Torrey) Beetle. Rare. Edge of creek in Colma
Canyon, K May 30, 1964.

Scirpus microcarpus Presl. Rare, in wet and marshy places. Meadow adjacent to
April Brook, W May 18, 1965; Colma Canyon, K May 30, 1964.

Scirpus robustus Pursh. Rare. Moist roadside ditch. Crocker Hills, K 2047.

GRAMINEAE. Grass Family

Agrostis diegoensis Vasey. Rare, open grassy slope. Near Quarry, W June 8, 1965.

Vou. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 657

Agrostis exarata Trinius. Rare, in low moist places. Pig Ranch Ravine, W May 11,
1965.

Agrostis hallii Vasey. Occasional, on brushy slopes. East facing slope below Parking
Lot, McC, Halde June 23, 1963; Quarry, K July 14, 1963; Crystal Cave Canyon, W June
19, 1965.

Agrostis semiverticillata (Forskal) C. Christensen. Moist disturbed ground. Crocker
Industrial Park, Brisbane, W June 8, 1965. Introduced from Europe and Asia.

Aira caryophyllea Linnaeus. Silvery Hair Grass. Delicate graceful annual. Well dis-
tributed on grassy or brushy slopes. Crocker Hills, near Cow Palace, W May 17, 1965;
Devil’s Arroyo, K May 9, 1965; near Nike Station, McC, W May 23, 1965; Owl’s Canyon,
K April 25, 1965; Pig Ranch Ravine, W May 11, 1965; east facing slope below Parking Lot,
McC, Halde June 9, 1963; south of West Powerline, K March 14, 1965. Introduced from
Europe.

Avena barbata Brotero. Slender Wild Oat. Weed of waste places and grassy slopes.
Colma Canyon, McC, Halde, K June 23, 1963; Crocker Avenue, Toschi February 23, 1963;
Quarry, K July 14, 1963. Native to Mediterranean region.

Briza maxima Linnaeus. Rattlesnake Grass. Big Quaking Grass. Attractive weedy grass
of waste ground. Sierra Point, W April 27, 1965. Introduced from Europe.

Briza minor Linnaeus. Little Quaking Grass. Occasional, on grassy areas and waste
ground. Guadalupe Valley, P March 30, 1963; near Olivet Cemetery, McC June 13, 1965;
Colma Canyon, McC, Halde, K June 23, 1963; Pig Ranch Ravine, W May 11, 1965; Roman-
zoffia Ravine, K May 4, 1964; Point San Bruno, K 1027. Introduced from Europe.

Bromus carinatus Hooker & Arnott. Common, on exposed or shaded habitats. Plants
mostly biennial. Randolph Ravine, P 2711; Colma Canyon, AK March 15, 1964; near Nike
Base, K April 25, 1964; McC, W May 23, 1965; Radio Road, below radio stations, Toschi
March 3, 1963; Owl’s Canyon, K April 25, 1965.

Bromus diandrus Roth [B. rigidus sensu auct. Amer.]. Rare, in disturbed soil. Guadalupe
Valley, P 2765a. Introduced from Europe.

Bromus marginatus Nees von Esenbeck. Occasional, open grassy slopes. North facing
slope above Crocker Avenue, Toschi 63:29; West Powerline, K April 13, 1964; Owl’s Canyon,
Ket0 52.

Bromus mollis Linnaeus. Soft Chess. Occasional, waste places and in grassland. West
Powerline, K April 13, 1964; Pig Ranch Ravine, K May 15, 1965; Colma Canyon, W July 7,
1965; Sierra Point, W April 27, 1965. Introduced from Europe.

Bromus rubens Linnaeus. Foxtail Chess. Rare, in disturbed dry soil. Pig Ranch Ravine,
K May 15, 1965; Sierra Point, W May 8, 1965. Introduced from Europe.

Calamagrostis nutkaensis (Presl) Steudel. Large clumping perennial, occasional, near
summit of mountain. Colma Canyon, McC, Halde, K June 23,1963; Dairy Ravine, K August
5, 1963; east facing slope below Parking Lot, McC June 9, 1963.

Cortaderia selloana (Schultes) Ascherson & Graebner. Pampas Grass. Tall perennial
planted as an ornamental but occasionally naturalized. Brisbane Lagoon, W August 14, 1965.
Native of South America.

658 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Danthonia californica Bolander. Rare, open brushy or grassy slopes. East facing
slope below Parking Lot, McC June 9, 1963; Crocker Hills, K May 4, 1964.

Deschampsia caespitosa (Linnaeus) Beauvois subsp. holciformis (Presl) Lawrence.
California Hair Grass. Rare, clumps on grassy slopes, a tufted perennial. Crocker Hills,
K May 4, 1964.

Distichlis spicata (Linnaeus) Greene. Salt Grass. Rare. Salt marsh near edge of bay.
Brisbane Lagoon, W August 14, 1965.

Elymus glaucus Buckley. Blue Wild Rye. Western Rye Grass. Common, in grassland
and on brushy hillsides. East facing slope below Parking Lot, McC, Halde June 23, 1963;
upper Buckeye Canyon, A July 14, 1963; Bitter Cherry Ridge, K May 15, 1965; Quarry,
W June 17, 1965.

Elymus triticoides Buckley. Rare, disturbed areas. Brisbane Lagoon, W August 14,
1965.

Festuca californica Vasey. Tall clumping perennial. Frequent, on somewhat shaded
brushy hillsides. Colma Canyon, K May 4, 1964; Bitter Cherry Ridge, K May 4, 1964; near
Parking Lot, McC April 17, 1964; Romanzoffia Ravine, K May 4, 1964; Owl’s Canyon,
K April 25, 1965.

Festuca dertonensis (Allioni) Ascherson & Graebner. Common, on brushy and grassy
slopes or flats. Near Crocker Avenue, P 2648; Randolph Ravine, P 2712; slope above Park-
ing Lot, McC April 17, 1964; Ridge Road, 42 mile east of Parking Lot, P 2694. Introduced
from Europe.

Festuca idahoensis Elmer. Blue Bunch Grass. Rare, rocky grassy slope at summit of
mountain. East facing slope below Parking Lot, McC, Halde June 23, 1964.

Festuca megalura Nuttall. Occasional, mostly in disturbed areas, sometimes on grassy
slopes. Crystal Cave Canyon, K June 20, 1965; ridge southwest of Brisbane, P 2756; near
Nike Station, McC, W May 23, 1965; Sierra Point, W April 27, 1965.

Festuca myuros Linnaeus. Weed, in disturbed areas and grassy slopes. Ravine % mile
east of Parking Lot, P March 9, 1963; Crystal Cave Canyon, K June 20, 1965. Introduced
from Europe.

Festuca rubra Linnaeus. Rare, on brushy slope below summit. Near Parking Lot, McC
June 9, 1963.

Holcus lanatus Linnaeus. Velvet Grass. Occasional, moist places and grassy slopes.
Lower part of Colma Canyon, McC, Halde, K June 23, 1963; Crocker Avenue, K September
2, 1964; Crocker Industrial Park, Brisbane, W June 8, 1965. Introduced from Europe.

Hordeum brachyantherum Nevski. Meadow Barley. Rare, in grassland and brushy
areas. Crocker Hills, W May 17, 1965; Randolph Avenue, at Hillside Boulevard, P 2733.

Hordeum geniculatum Allioni [H. hystrix Roth]. Mediterranean Barley. Rare, in
grassland, Crocker Hills near Cow Palace, W May 17, 1965. Introduced from Europe.

Hordeum leporinum Link. Farmer’s Foxtail. Disturbed or grassy areas. Guadalupe
Valley, P 2765; Randolph Avenue at Hillside Boulevard, P 2741; West Powerline, K April
13, 1964; Colma Canyon, W August 16, 1965. Introduced from southern Europe.

Vout. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 659

Koeleria macrantha (Ledebour) Sprengel [K. gracilis Persoon; K. cristata (Linnaeus)
Persoon]. June Grass. Frequent, on brushy and often rocky slopes. Sierra Point, W April
27, 1965; near Nike Station, McC, W May 23, 1965; upper Buckeye Canyon, K July 14,
1963; Owl’s Canyon, K 1053; east facing slope below Parking Lot, McC, Halde June 23, 1963.

Lolium multiflorum Lamarck. Italian Rye Grass. Occasional, in waste places and
grassy slopes and flats. Lower part of Colma Canyon, McC, Halde, K June 23, 1963; Guada-
lupe Road, K August 5, 1963. Introduced from Europe.

Melica californica Scribner. Occasional, in grassland. Ridge southwest of Brisbane,
P 2754; Randolph Avenue at Hillside Boulevard, P 2720; near Olivet Cemetery, W May 18,
1965; Owl’s Canyon, K April 25, 1965; Pig Ranch Ravine, K May 15, 1965.

Mclica imperfecta Trinius. Delicate perennial. Rare. Brushy slope, Colma Canyon,
K May 30, 1964.

Melica torreyana Scribner. Frequent, on brushy slopes and grassland. Near Olivet
Cemetery, W May 18, 1965; Huckleberry Ridge, K May 4, 1964; Owl’s Canyon, K April
25, 1965; Buckeye Canyon, K May 24, 1966; slope southeast of summit, P 2676; West
Powerline, McC March 14, 1965.

Panicum capillare Linnaeus. Witch Grass. Rare. Disturbed area, Brisbane Lagoon,
W June 19, 1965. Introduced from eastern North America.

Parapholis incurva (Linnaeus) C. E. Hubbard. Sickle Grass. Rare. Salt flat, Sierra
Point, W May 8, 1965. Native of Europe.

Phalaris californica Hooker & Arnott. California Canary Grass. Common, in moist
areas and on grassy ridges. Randolph Avenue at Hillside Boulevard, P 2737; Colma Canyon,
K May 30, 1964; Quarry, K July 14, 1963; Bitter Cherry Ridge, K May 4, 1964; Crystal
Cave Canyon, K June 20, 1965; Pig Ranch Ravine, K May 15, 1965.

Phalaris canariensis Linnaeus. Canary Grass. Rare. Disturbed ground. Brisbane
Lagoon, W June 19, 1965. Introduced from Europe. Used in bird-seed mixtures.

Poa annua Linnaeus. Annual Bluegrass. Occasional, in grassland. Randolph Avenue,
K May 16, 1967; Crocker Avenue between Templeton Avenue and Scenic Way, J. H. Thomas
May 2, 1967.

Poa pratensis Linnaeus. Kentucky Blue-Grass. Disturbed area along Radio Road, W
May 23, 1965. Probably introduced from Europe and often used in lawn grass mixtures.

Poa unilateralis Scribner. Occasional, open grassy or rocky slopes. Ridge southwest
of Brisbane, P 2755; east facing slope below Parking Lot, McC April 17, 1964; Owl’s Canyon,
K April 25, 1965; Brisbane Powerlines, K March 21, 1965.

Polypogon monspeliensis (Linnaeus) Desfontaines. Rabbit-foot Grass. Occasional, in
wet or damp areas. Point San Bruno, McC, K June 8, 1965; Sierra Point, W May 8, 1965;
Crocker Industrial Park, W June 15, 1965; lower Colma Canyon, McC, Halde, K June 23,
1963. Introduced from Europe.

Secale cereale Linnaeus. Cereal Rye. Rare. Disturbed ground, Point San Bruno,
K 1038. Tufted annual, frequently cultivated and becoming naturalized. Native of south-
west Asia.

Sitanion jubatum J. G. Smith. Big Squirrel-tail Grass. Rare, dry open often rocky
places or waste places. Sierra Point, W June 20, 1965; Brisbane Ravine, W May 25, 19065.

660 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Sorghum halepense (Linnaeus) Persoon. Johnson Grass. Rare. Disturbed ground,
Point San Bruno, W June 19, 1965. Native of Mediterranean region.

Stipa cernua Stebbins & Love. Rare. Rocky grassland. Pig Ranch Ravine, K May 15,
1965.

Stipa lepida Hitchcock. Rare. Rocky open grassland, Raven 1783; near Olivet Cemetery,
W May 18, 1965.

Stipa pulchra Hitchcock. Rare, grassland and rocky slopes. Randolph Avenue, at
Hillside Boulevard, P 2613; slope southeast of summit, P 2670.

Triticum aestivum Linnaeus. Wheat. Spontaneous in waste places, may not persist.
Point San Bruno, K 1037; Brisbane Lagoon, W June 19, 1965. Widely cultivated, introduced
from Old World.

TRIDACEAE. Iris Family

Chasmanthe aethiopica (Linnaeus) N. E. Brown [Antholyza aethiopica Linnaeus].
Head of Buckeye Canyon, persisting. Planted in 1965 by Philip Wheeler. Native of South
Africa.

Freesia refracta Klatt. Freesia. Slope below radio station on Radio Road. Toschi
63:64. Native of South Africa.

Iris douglasiana Herbert. Douglas Iris. Frequent, open grassland and meadows. Point
San Bruno, K March 16, 1964; Colma Canyon, McC, Halde, K June 23, 1963; Nike Base,
K March 15, 1964; Meadow between April Brook and Radio Road, McC March 28, 1965;
Cable Ravine, McC March 14, 1965; Horseshoe Ridge, K August 5, 1963; Pig Ranch Ravine,
W August 11, 1965. Showy flowers in various shades of cream-white to blue, March to
May. Ornamental.

Tris longipetala Herbert. Frequent, open grassland and meadows. Occurs in same
habitat as preceding iris and in at least one locality (the Flower Garden) the two are grow-
ing together. Colma Canyon, K 508; near Crocker Avenue, P 2655; north of Randolph Drive
at Hillside Boulevard, P 2707; Point San Bruno, K March 16, 1964; east end of ridge near
Brisbane Powerlines, K 956. Flowers in various shades of blue, April and May. Petals to
3 inches long, often longer than petals of Douglas Iris. In the absence of flowers the two
irises may be distinguished by their leaves which in Jris douglasiana are dark green and
shining on upper surface and dull grayish-green on lower surface while in J. longipetala the
leaves have the same color on both surfaces.

Sisyrinchium bellum Watson. California Blue-eyed Grass. Widely distributed, open
grassland. Colma Canyon, McC, Halde, K June 23, 1963; east facing slope below Parking
Lot, McC April 17, 1964; Cable Ravine, McC March 14, 1965; Manzanita Dike, K 932;
Olivet Cemetery, McC June 13, 1965; Sierra Point, W May 21, 1965; north of Randolph
Drive at Hillside Boulevard, P 2705; Glen Park Canyon, McC April 16, 1967.

Sisyrinchium californicum (Ker) Dryander. Yellow-eyed Grass. Rare, in boggy places
or edge of water. Colma Creek, in water, W June 15, 1965.

Tritonia crocosmaeflora Lemoine. Montbretia. Rare, an escape from cultivation. In
shade of eucalyptus trees, Radio Road, McC September 24, 1967. Perennial, growing from
a corm, 3 to 4 feet tall, sword-shaped leaves, showy orange flowers, August and September.
A garden hybrid between two South African plants, 7. pottsii Baker and 7. aurea Planchon.

Vout. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 661

JUNCACEAE. Rush Family

Juncus balticus Willdenow. Baltic Rush. Rare. Salt flat, Point San Bruno, K, McC,
September 11, 1964.

Juncus bufonius Linnaeus. Toad Rush. Occasional, in wet places or seepage areas.
Quarry, K July 7, 1963; Colma Canyon, K May 30, 1964; between Nike Base and April
Brook, McC, W May 23, 1965; Point San Bruno, K 1040.

Juncus effusus Linnaeus var. brunneus Engelmann. Bog Rush. Occasional, in’ wet
places. Lower Colma Canyon, McC, Halde, K June 23, 1963; Quarry, K July 14, 1963;
Buckeye Canyon, K September 2, 1964; Crocker Avenue, K August 5, 1963.

Juncus leseurii Bolander. Salt Rush. Rare, wet places, in both fresh water and salt
flats. Creek in lower Buckeye Canyon, K September 2, 1964; Point San Bruno, K Septem-
ber 11, 1964.

Juncus mexicanus Willdenow. Mexican Rush. Rare. Dry ground, Point San Bruno,
K September 11, 1964.

Juncus occidentalis (Coville) Wiegand [J. tenuis Willdenow var. congestus Engelmann].
Western Rush. Rare. Sierra Point, W May 8, 1965.

Juncus phaeocephalus Engelmann. Rare. Sierra Point, W May 8, 1965; north facing
brushy slope south of Guadalupe Parkway and near its junction with Radio Road, McC,
W June 11, 1967.

Juncus xiphioides E. Meyer. Rare. Sierra Point, W May 8, 1965.

Luzula multiflora (Retzius) Lejeune [L. subsessilis (Watson) Buchenau]. Wood Rush.
Common, in grassland and seepage areas. Bitter Cherry Ridge, K April 4, 1965; Radio Road
below transmitting towers, McC, Menzies February 16, 1963; east facing slope below Park-
ing Lot, McC March 17, 1964; Crocker Hills, K January 8, 1964; Crystal Cave Canyon,
K June 20, 1965; Randolph Drive at Hillside Boulevard, P 2677; Glen Park Canyon, McC
April 16, 1967; Sierra Point, W May 8, 1965.

JUNCAGINACEAE. Arrowgrass Family

Triglochin concinna Davy. Slender Arrowgrass. Rare. Salt flat, Point San Bruno,
K 1084.

LEMNACEAE. Duckweed Family

Lemna minima Philippi. Duckweed. Rare, floating in water. Cow Trough Ravine,
K October 23, 1963; April Brook, K June 20, 1965; Crocker Hills, in a marsh between
Geneva Avenue and Guadalupe Parkway, K October 4, 1966.

LintacesE. Lily Family

Allium dichlamydeum Greene. Coastal Onion. Occasional, grassland and rocky slopes.
Colma Canyon, McC, Halde, K June 23, 1963; Bitter Cherry Ridge, K 1094; near Nike
Base, McC May 23, 1965; Flower Garden, W June 20, 1965; east facing slope below Parking
Lot, McC, Halde June 23, 1963; Sierra Point, W May 8, 1965. Attractive rose-purple flowers,
May and June. Ornamental.

Allium triquetrum Linnaeus. A weed in gardens and disturbed places. Crocker In-
dustrial Park, W March 27, 1966. Native of southern Europe.

662 CALIFORNIA ACADEMY OF SCIENCES [PRoc. 47TH SER.

Ficure 11. Allium dichlamydeum Greene.

VoL. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 663

Allium unifolium Kellogg. Rare, in moist areas. Colma Creek, W May 18, 1965; April
Brook, W May 24, 1965; north facing slope south of Guadalupe Parkway near its junction
with Radio Road, McC, W June 11, 1967. Attractive rose-pink flowers, May.

Brodiaea elegans Hoover. Harvest Brodiaea. Rare, in grassland. Sierra Point, W June
21, 1965.

Brodiaea laxa (Bentham) Watson. Grass-Nut. Ithuriel’s Spear. Frequent, in grass-
land. Colma Canyon, McC, Halde, K June 23, 1963; Crocker Hills, W June 8, 1965; grass-
land along Radio Road, W June 16, 1965; east facing slope below Parking Lot, McC, Halde
June 23, 1963; Sierra Point, W May 8, 1965. Attractive robust plant with blue-purple flowers
in umbels which look like a miniature agapanthus, May and June. Ornamental.

Brodiaea pulchella (Salisbury) Greene [B. capitata Bentham]. Blue-Dicks. Wild-
Hyacinth. Frequent, in grassland or in brush. Colma Canyon, K May 30, 1964; Devil’s
Arroyo, K 1023; Bitter Cherry Ravine, K April 4, 1965; Crystal Cave Canyon, W June 19,
1965; east facing slope below Parking Lot, McC April 17, 1964; Randolph Avenue at Hillside
Boulevard, South San Francisco, P 2730. The congested heads of flowers on weak stems
distinguish this species from the other three brodiaeas in our area.

Brodiaea terrestris Kellogg [B. coronaria (Salisbury) Engler var. macropoda (Torrey)
Hoover]. Rare, on western section of mountain. Near Quarry, W June 19, 1965; Radio Road
in Eucalyptus area, McC, W June 11, 1967. The corms of the species of Brodiaea are edible
and were used as food by California Indians.

Chlorogalum pomeridianum (DeCandolle) Kunth. Amole. Soap-Plant. Occasional,
in grassland. Colma Canyon, K May 4, 1964; Trillium Gulch (where there had been a fire
during previous autumn), McC March 28, 1965; east facing slope below Parking Lot, McC
June 9, 1963; Sierra Point, W June 23, 1965. The wavy-margined leaves appear in winter
and spring before the tall flowering branches. The star-like flowers open in the evening or
on gray foggy or overcast days. The large underground bulb is covered with old fibrous
coats from previous seasons. The bulb which contains a saponin was used by Indians and
early settlers for its saponifying effect. The Indians roasted the bulb and used it for food.

Disporum hookeri (Torrey) Nicholson. Rare, in damp ground. Devil’s Arroyo, McC
March 28, 1965. Ornamental.

Fritillaria lanceolata Pursh. Checker-Lily. Mission Bells. Occasional, grassland and
rocky or brushy slopes. Sierra Point, W June 21, 1965; Radio Road, east facing slope below
radio station, Toschi March 17, 1963; near Parking Lot at summit, K 630; 1 mile east of
Parking Lot at summit, P 2702; Cable Ravine, McC March 14, 1965; Trillium Gulch, K 990.
Brownish-purple flowers are nodding. Ornamental.

Maianthemum dilatatum (Wood) Nelson & Macbride. False Lily-of-the-Valley. Rare.
Rocky outcrop. Kamchatka Point, McC June 9, 1963; Thomas 9288. Maianthemum dilata-
tum occurs in eastern Asia and western North America and reaches its southernmost known
American distribution in San Mateo County. It belongs to a small genus of three species
known from the north temperate regions of Eurasia and North America.

Muilla maritima (Torrey) Watson. Rare, in grassland. Guadalupe Road, L. Rose
March 25, 1965. Muilla is an anagram of Allium. This yellow-flowered species has an onion-
like appearance but its plants lack the odor or taste of onion and the underground part is a
corm and not a bulb as in Allium.

664

CALIFORNIA ACADEMY OF SCIENCES

Ficure 12. Fritillaria lanceolata Pursh.

[Proc. 4TH SER.

Vout. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO 665

~ a a

Ficure 13. Trillium chloropetalum (Torrey) T. J. Howell.

Smilacina racemosa (Linnaeus) Desfontaines var. amplexicaulis (Nuttall) Watson.
Fat Solomon. Occasional, in grassy and brushy areas. Colma Canyon, W June 13, 1965;
Devil’s Arroyo, K 1055; Romanzoffia Ravine, K May 4, 1964; Trillium Gulch, McC March
28, 1965. Ornamental, with numerous white flowers in a panicle.

Smilacina stellata (Linnaeus) Desfontaines var. sessilifolia (Baker) Henderson. Slim
Solomon. Occasional, on grassy or brushy areas, sometimes with Fat Solomon, from which
it is distinguished by its fewer flowers in a raceme. Colma Canyon, McC, Halde, K June 23,
1963; Trillium Gulch, McC March 28, 1965; Bitter Cherry Ridge, K May 4, 1964; Devil’s
Arroyo, K 1056; Crystal Cave Canyon, W June 20, 1965. Ornamental.

Trillium chloropetalum (Torrey) T. J. Howell. Wake-Robin. Rare, in brushy areas
under Baccharis and Heracleum. Devil’s Arroyo, McC March 28, 1965; Trillium Gulch,
K March 21, 1965; Colma Canyon, McC April 11, 1965. Flowers March and April. The
pale purple flowers are sessile and surrounded by the mottled whorled leaves. Ornamental.

Zigadenus fremontii (Torrey) Torrey ex Watson var. minor (Hooker & Arnott)
Jepson. Zygadene. Rare, in grassland. Point San Bruno, K March 16, 1964. Attractive white
flowers, February to April. In some species of Zigadenus, especially Z. venenosus, the death-
camas, all parts of the plants are poisonous. Bulbs of all species are suspect.

OrcHIDACEAE. Orchid Family

Habenaria unalascensis (Sprengel) Watson var. maritima (Greene) Correll [H.
elegans (Lindley) Bolander var. maritima (Greene) Ames]. Rein Orchis. Rare, grassland
or brushy slopes. April Brook, W August 16, 1965; Horseshoe Ridge along Radio Road,

666 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

FicureE 14. Fog over top of ridge above the quarry on north side.

K August 5, 1963. Flowers small, greenish in a spike, August and September, each flower
with a basal spur. Neither the small flowers of this orchid nor of the following orchids have
any ornamental value.

Habenaria unalascensis (Sprengel) Watson var. elata (Jepson) Correll. Rein Orchis.
Rare. Grassy slope above April Brook, McC August 1, 1965.

Habenaria unalascensis (Sprengel) Watson var. unalascensis. Rein Orchis. Rare.
Quarry, K July 14, 1963.

Spiranthes romanzoffiana Chamisso & Schlechtendal. Ladies-Tresses. Rare, in grass-
land. Radio Road near radio towers, Marsh Pitman 482. Small greenish-white flowers in
June occur in a spike and lack the spur of the rein-orchis.

TYypHACEAE. Cattail Family

Typha domingensis Persoon. Rare, marshy area in Crocker Hills between Geneva
Avenue and Guadalupe Road, K October 4, 1966.

Typha latifolia Linnaeus. Broad-leaved Cattail. Soft Flag. Rare, marshy area in
Crocker Hills between Geneva Avenue and Guadalupe Road, K October 4, 1966. Both of
these cattails grew together in the same area which has now been filled for industrial de-
velopment. With phasing out of this habitat these two species probably no longer exist
in our area.

INDEX TO FAMILIES, GENERA, AND CoMMON NAMES

Acacia, 632, 633
Black, 633
Blackwood, 633
Everblooming, 633

Acaena, 644

Achillea, 611

Adiantum, 604

Aesculus, 630

Agoseris, 611

Agrostis, 656, 657

Aira, 657

Aizoaceae, 605

Albizia, 633

Alchemilla, 645

Alfalfa, 635

Alkali Heath, 628

Alkali Parsnip, 654

Allium, 661, 663

Allocarya, 607

Alum Root, 649

Alyssum, 623

Amaranth, 606

Amaranth Family, 606

Amaranthaceae, 606

Amaranthus, 606

Amelanchier, 645

Amole, 663

Amsinckia, 607

Anacardiaceae, 606

Anagallis, 643

Anaphalis, 611

Angelica, 652

Anthemis, 612

Anthriscus, 653

Apiastrum, 653

Apocynaceae, 606

Apocynum, 606

Aquilegia, 643

Arabis, 621

Araceae, 656

Aralia Family, 606

Araliaceae, 606

Arbutus, 625

Arctostaphylos, 626

Arenaria, 608

Aristolochia, 606

667

Aristolochia Family, 606
Aristolochiaceae, 606
Arrowgrass Family, 661
Arrowgrass, Slender, 661
Artemisia, 612
Aster, 612
California, 612
Golden, 613
Rough-leaved, 612
Astragalus, 633
Athyrium, 604
Atriplex, 610
Avena, 657

Baby-Blue Eyes, 630
Baby’s Tears, 655
Baccharis, 612
Baerila, 612, 613
Barbarea, 621, 622
Barberry, Coast, 607
Barberry Family, 607
Barley, Meadow, 658
Mediterranean, 658
Bassia, 610
Bay, California, 632
Bayberry Family, 637
Beach-Bur, 615
Bean Family, 632
Bean, Stink, 633
Bear-berry, 626
Bedstraw, 648
Bee Plant, 652
Beech Family, 627
Bellardia, 650
Bellflower Family, 608
Bellis, 613
Berberidaceae, 607
Berberis, 607
Bermuda Buttercup, 639
Betulaceae, 607
Bindweed, 620
Birch Family, 607
Bird’s Foot Trefoil, 633
Bitter Dock, 642
Black Berry, Pacific, 648
Bleeding Heart, 628

668 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

Blennosperma, 613
Blue Blossom, 644
Blue Dicks, 663
Blue-eyed Grass, California, 660
Blue Gum, 637
Blue Witch, 652
Bluff Lettuce, 621
Borage Family, 607
Boraginaceae, 607
Bowlesia, 653
Brass-Buttons, 614
Brassica, 622
Bristly Ox-Tongue, 618
Briza, 657
Brodiaea, 663
Harvest, 663
Bromus, 657
Brooklime, American, 652
Broom, Chaparral, 612
French, 633
Scotch, 633
Broomrape, 639
Broomrape Family, 639
Buckeye, 630
Buckeye Family, 630
Buckhorn, 641
Buckthorn Family, 644
Buckwheat Family, 641
Buckwheat, Wild, 642
Bur-chervil, 653
Butter-and-eggs, 651
Buttercup, Bermuda, 639
California, 644
Prickle-fruited, 644
Buttercup Family, 643
Butterweed, 618

Cabbage, 622
Cakile, 622
Calamagrostis, 657
Calandrinia, 643
Calla, 656

Lily, 656
Calla Family, 656
Callitrichaceae, 607
Callitriche, 607
Camissonia, 638
Campanulaceae, 608
Campion, 609
Caprifoliaceae, 608

Capsella, 622
Cardamine, 622
Cardaria, 622
Cardionema, 608
Carduus, 613
Carex, 656
Carpet-Weed Family, 605
Carrot Family, 652
Caryophyllaceae, 608
Castilleja, 650
Catchfly, 609
Cat’s Ear, Hairy, 617
Smooth, 616
Cat-tail, 666
Cat-tail Family, 666
Caucalis, 653
Ceanothus, 644
Celery, Wild, 653
Centaurea, 613
Centaurium, 628
Centaury, 628
Centranthus, 655
Cerastium, 609
Cereal Rye, 659
Chaetopappa, 613
Chamomile, 612
Chaparral-Broom, 612
Dwarf, 612
Charlock, 622
Chasmanthe, 660
Checker Bloom, 637
Lily, 663
Cheeseweed, 637
Cherry, Bitter, 647
Holly-leaved, 647
Western Choke, 647
Chess, Foxtail, 657
Soft, 657
Chenopodiaceae, 610
Chenopodium, 610
Chickweed, 609, 610
Mouse-ear, 609
Chicory, 613
California, 618
Chinese Caps, 627
Houses, 650
Chorizanthe, 641
Chorogalum, 663
Christmas Berry, 646

Vor. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO

Chrysanthemum, 613
Garland, 613
Chrysopsis, 613
Cichorium, 613
Cineraria, 618
Garden, 618
Cinquefoil, 646, 647
Cirsium, 613, 614
Clarkia, 638, 639
Cleavers, 648
Clotbur, Spiny, 620
Clover, Bur, 635
Indian Sweet, 635
Owl’s, 651
Sack, 635
Spanish, 634
Tomcat, 636
White, 636
Coffee Berry, 644
Collinsia, 650
Columbine, 643
Compositae, 611
Conium, 653
Convolvulaceae, 620
Convolvulus, 620
Convolvulus Family, 620
Conyza, 614
Coprosma, 648
Cornaceae, 621
Cornus, 621
Coronopus, 622
Cortaderia, 657
Corylus, 607
Cotoneaster, 645
Cottonweed, 618
Cotula, 614
Cow Parsnip, 654
Coyote Brush, 614
Mint, 631
Crane’s Bill Geranium, 629
Crassulaceae, 621
Crataegus, 645
Cream Bush, 646
Cream-Cups, 640
Cress, American Winter, 621
Bitter, 622
Coast Rock, 621
Early Winter, 622
Hoary, 622
Wart, 622

Water, 623
Croton, 627
Cruciferae, 621
Cryptantha, 607
Cucumber, Wild, 624, 625
Cucurbitaceae, 624
Cudweed, 615

California, 615

Everlasting, 615

Purple, 615
Cupressaceae, 605
Cupressus, 605
Curly Dock, 642

Currant, California Black, 649

Chaparral, 649
Cynoglossum, 607
Cyperaceae, 656
Cyperus, 656
Cypress Family, 605
Cypress, Monterey, 605
Cytisus, 633

Daisy, Crown, 613
English, 613
Philadelphia, 614
Seaside, 614

Dandelion, California, 611
Coast, 611
Red-Seeded, 619

Danthonia, 658

Daucus, 654

Deerweed, 634

Delphinium, 643

Dentaria, 622

Deschampsia, 658

Dicentra, 628

Dichondra, 620

Diplacus, 651

Dipsacaceae, 625

Dipsacus, 625

Disporum, 663

Distichlis, 658

Dock, Bitter, 642
California, 642
Curly, 642
Fiddle, 643
Green, 642

Dodecatheon, 643

Dogbane, 606

Dogbane Family, 606

669

670 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER.

Dog-Fennel, 612
Dogwood, Creek, 621
Western, 621
Dogwood Family, 621
Dryopteris, 605
Duckweed, 661
Duckweed Family, 661
Dudleya, 621
Dutchman’s Pipe, 606

Elderberry, Blue, 608

Red, 608
Elymus, 658
Epilobium, 639
Equisetaceae, 604
Equisetum, 604
Erechtites, 614
Ericaceae, 625
Erigeron, 614
Eriodictyon, 630
Eriogonum, 642
Eriophyllum, 615
Erysimum, 623
Erodium, 629
Eschscholzia, 640
Escobita, 651
Eucalyptus, 637
Euphorbia, 627
Euphorbia Family, 627
Euphorbiaceae, 627
Evening Primrose Family, 638
Everlasting, 615

Green, 615

Pearly, 611

Pink, 615

Fagaceae, 627

False Lily-of-the-Valley, 663

Farewell-to-Spring, 639

Fat Solomon, 665

Fennel, Sweet, 654

Fern, Bracken, 605
California Maidenhair, 604
Coastal Wood, 605
Golden-back, 605
Leather, 605
Western Chain, 605
Western Lady, 604
Western Sword, 605

Fern Family, 604

Festuca, 658
Fiddleneck, 607
Fiesta Flower, 631 |
Figwort Family, 650
Filago, 615
Filaree, Long-beaked, 629
Musk, 629
Red-stemmed, 629
White-stemmed, 629
Fireweed, Australian, 614
New Zealand, 614
Flax, 636
Blue Toad, 651
Flax Family, 636
Foeniculum, 654
Footsteps-of-Spring, 654
Forget-me-not, 607
Four-o’clock, 638
Four-o’clock Family, 638
Foxtail, Chess, 657
Farmer’s, 658
Fragaria, 645, 646
Frankenia, 628
Frankenia Family, 628
Frankeniaceae, 628
Franseria, 615
Freesia, 660
Fringe Cups, 650
Fringe-Pod, 624
Fritillaria, 663
Fuchsia, California, 639
Fuller’s Teasel, 625
Fumariaceae, 628
Fumitory Family, 628
Furze, 636

Galium, 648
Garland Chrysanthemum, 613
Garrya, 628
Garryaceae, 628
Gaultheria, 626
Gentian Family, 628
Gentianaceae, 628
Geraniaceae, 629
Geranium, 629
Carolina, 629
Crane’s Bill, 629
Cut-leaved, 629
New Zealand, 629
Geranium Family, 629

VoL. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO

Gilia, 641

California, 641
Glasswort, 610
Gnaphalium, 615
Golden-Rod, California, 619

Dune, 619

Meadow, 619
Goldfields, 612, 613
Gooseberry, 649

Bay, 649

California, 649

Canyon, 649

Hillside, 649
Goosefoot Family, 610
Gopher Plant, 627
Gorse, 636
Gourd Family, 624
Gramineae, 656
Grass, Annual Blue, 659

Big Quaking, 657

Big Squirrel-tail, 659

Blue Bunch, 658

Blue Wild Rye, 658

California Blue-eyed, 660

California Canary, 659

California Hair, 658

Canary, 659

Cereal Rye, 659

Goose, 648

Italian Rye, 659

Johnson, 660

June, 659

Kentucky Blue, 659

Little Quaking, 657

Pampas, 657

Rabbit-foot, 659

Rattlesnake, 657

Russian Pepper, 623

Salt, 658

Shining Pepper, 623

Sickle, 659

Silvery Hair, 658

Velvet, 658

Western Rye, 658

Witch, 659

Yellow-eyed, 660
Grass Family, 656
Grass-Nut, 663
Grindelia, 616
Groundsel, 619

Grossularia, 649
Gum-Weed, 616

Habenaria, 665, 666
Hair-Grass, California, 658
Silvery, 657
Haloragidaceae, 630
Haplopappus, 616
Hareleaf, 617
Hawthorn, 645
Hazel, California, 607
Heath, Alkali, 628
Heather Family, 625
Hedera, 606
Hedge Nettle, 632
Helenium, 616
Helianthella, 616
Heliotrope, Wild, 630
Helxine, 655
Hemizonia, 616
Hemlock, Poison, 653
Spotted, 653
Heracleum, 654
Hesperocnide, 655
Heteromeles, 646
Heterotheca, 616
Heuchera, 649
Himalaya Berry, 647
Hippocastanaceae, 630
Hoarhound, 631
Holcus, 658
Holly, California, 646
Hollyhock, Wild, 637
Holodiscus, 646
Honesty, 623
Honeysuckle, 608
Honeysuckle Family, 608
Hordeum, 658
Horkelia, 646
Horse Chestnut, 630
Horse Chestnut Family, 630
Horsetail, Common, 604
Giant, 604
Horsetail Family, 604
Horseweed, 614
Hottentot-Fig, 605
Hound’s Tongue, 607
Huckleberry, 627
Evergreen, 627
Hydrophyllaceae, 630

672

Aypericaceae, 631
Hypericum, 631
Hypochaeris, 616, 617

Indian-Hemp, 606

Indian Paint Brush, 650

Tridaceae, 660

Tris, 660

Iris Family, 660

Islay, 647

Ithuriel’s Spear, 663

Ivy, English, 606
German, 618

Jaumea, 617
Johnny-Jump-Up, 656
Johnny Tuck, 651
Juncaceae, 661
Juncaginaceae, 661
Juncus, 661

June Berry, 645
Jupiter’s Beard, 655

Kinnikinnick, 626

Knotweed, Dooryard, 642

Dune, 642
Koeleria, 659
Koniga, 623

Labiatae, 631
Lactuca, 617
Ladies-Tresses, 666

Lady’s Mantle, Western, 645

Lagophylla, 617
Lamb’s Quarters, 610
Larkspur, Blue, 644
Coastal, 643
Lathyrus, 633
Lauraceae, 632
Laurel, California, 632
Laurel Family, 632
Lavatera, 637
Layia, 617
Leguminosae, 632
Lemna, 661
Lemnaceae, 661
Lepechinia, 631
Lepidium, 623
Lettuce, Bluff, 621
Miner’s, 643

CALIFORNIA ACADEMY OF SCIENCES

Prickly, 617

Wild, 617

Willow, 617
Ligusticum, 654
Liliaceae, 661
Lily Family, 661

Lily-of-the-Valley, False, 663

Limonium, 641
Linaceae, 636
Linanthus, 641
Linaria, 651
Linum, 636
Lithophragma, 649
Lizard-tail, 615
Lobularia, 623
Locoweed, 633
Lolium, 659
Lomatium, 654
Lonicera, 608
Loosestrife, 637
Loosestrife Family, 637
Lotus, 633, 634
Lovage, 654
Lucerne, 635
Lunaria, 623
Lupine, 634, 635
Blue Beach, 634
Bush, 634
Silver Bush, 634
Yellow Beach, 634
Lupinus, 634, 635
Luzula, 661
Lythraceae, 637
Lythrum, 637

Madder Family, 648
Madia, 617, 618
Madrone, 625
Madrono, 625
Mahonia, 607
Maianthemum, 663
Mallow, Bull, 637
Tree, 637
Mallow Family, 637
Malva, 637
Malvaceae, 637
Malva-Rose, 637
Manroot, 624, 625
Manzanita, 626
Marah, 624, 625

[Proc. 4TH SER.

Vou. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO

Marrubium, 631
Marsh Rosemary, 641
Marvel-of-Peru, 638
Matricaria, 618
Mayweed, 612
Meadow Rue, 644
Meconella, 640
Medicago, 635
Melica, 659
Melilotus, 635
Mentha, 631
Mesembryanthemum, 605
Micropus, 618
Microseris, 618
Microsteris, 641
Milfoil, 611
Milk-Maids, 622
Mimulus, 651
Miner’s Lettuce, 643
Mint, Coyote, 631
Mint Family, 631
Mirabilis, 638
Mirror Shrub, 648
Mission-Bells, 663
Mist Maiden, 631
Mock-Heather, 616
Monardella, 631
Money-Plant, 623
Monkey Flower, 651
Bush, 651
Sticky, 651
Montbretia, 661
Montia, 643
Moonwort, 623
Morning-Glory, Orchard, 620
Western, 620
Mountain Balm, 630
Mug-Wort, 612
Muilla, 663
Mule-Ears, 620
Mustard, Field, 622
Hedge, 623
Mediterranean, 622
Summer, 622
Tower, 621
Mustard Family, 621
Myosotis, 607
Myrica, 637
Myricaceae, 637
Myriophyllum, 630

Myrtaceae, 637
Myrtle Family, 637
Myrtle, Oregon, 632

Nasturtium, 652
Navarretia, 641
Nemophila, 630
Nettle, 655
Coast Hedge, 632
Hedge, 632
Nettle Family, 655
New Zealand Spinach, 606
Nightshade, 652
Nightshade Family, 652
Ninebark, 646
Nyctaginaceae, 638

Oak, California Live, 627
Canyon, 627
Coastal Live, 627
Gold-cup, 627
Interior Live, 628
Maul, 627
Oracle, 628
Poison, 606

Oak Family, 627

Ocean Spray, 646

Oenanthe, 654

Oenothera, 638, 639

Onagraceae, 638

Onion, Coastal, 661

Orchid Family, 665

Orchidaceae, 665

Orchis, Rein, 665

Orobanchaceae, 639

Orobanche, 639

Orthocarpus, 651

Osmaronia, 646

Oso Berry, 646

Owl’s Clover, 651

Oxalidaceae, 639

Oxalis, 639, 640
Cape, 639

Oxalis Family, 639

Ox-Tongue, 618

Oyster-Root, 620

Paint Brush, 650
Franciscan, 650
Indian, 650
Wight’s, 650

673

674 CALIFORNIA ACADEMY OF SCIENCES

Pampas Grass, 657
Panicum, 659
Papaver, 640
Papaveraceae, 640
Parapholis, 659
Paronychia, 609
Parrot Feather, 630
Parsley, California Hedge, 653
Parsley Family, 652
Parsnip, Alkali, 654
Bladder, 654
Cow, 654
Lace, 654
Pea, Garden, 635
Pacific, 633
Sweet, 633
Pea Family, 632
Pearlwort, 609
Western, 609
Pearly Everlasting, 611
Pennyroyal, 631
Pepper-Grass, Russian, 623
Shining, 623
Pepper Tree, 606
Periwinkle, 606
Phacelia, 630
Phacelia Family, 630
Phalaris, 659
Phlox, 641
Pholistoma, 631
Photinia, 646
Physocarpus, 646
Pickleweed, 610
Picris, 618
Pigmyweed, 621
Pigweed, 606, 610
Pimpernel, Scarlet, 643
Pinaceae, 605
Pine, Monterey, 605
Pine Family, 605
Pineapple-Weed, 618
Pincushion Plant, 625
Pink, Windmill, 609
Pink Family, 608
Pinus, 605
Pipe Vine, California, 606
Pisum, 635
Pitcher Sage, 631, 632
Pityrogramma, 605
Plantaginaceae, 640

Plantago, 640, 641
Plantago Family, 640
Plantain, California, 640
English, 641
Mexican, 640
Platystemon, 640
Plectritis, 655
Plumbaginaceae, 641
Plumbago Family, 641
Poa, 659
Pogogyne, 631
Poison Hemlock, 653
Poison Oak, 606
Polemoniaceae, 641
Polemonium Family, 641
Polycarpon, 609
Polygonaceae, 641
Polygonum, 642
Polypodiaceae, 604
Polypodium, 605
Polypody, 605
Polypogon, 659
Polystichum, 605
Poppy, California, 640
Opium, 640
Wind, 640
Poppy Family, 640
Portulaca, 643
Portulaca Family, 643
Portulacaceae, 643
Potentilla, 646, 647
Primrose Family, 643
Primulaceae, 643
Prunella, 632
Prunus, 647
Psilocarphus, 618
Pteridium, 605
Pterostegia, 642
Purslane, 643
Purslane Family, 643

Quercus, 627, 628
Quinine Bush, 628

Radish, Wild, 623
Rafinesquia, 618
Ragwort, 618
Rain-Bells, 622
Ranunculaceae, 643
Ranunculus, 644

[Proc. 4TH SER.

Vor. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO

Raphanus, 623

Rattlesnake Weed, 654

Red Maids, 643

Redberry, 644

Redscale, 610

Rein Orchis, 665, 666

Rhamnaceae, 644

Rhamnus, 644

Rhus, 606

Ribes, 649

Ribgrass, 641

Romanzoffia, 631

Rorippa, 623

Rosa, 647

Rosaceae, 644

Rose, California, 647
Wood, 647

Rose Family, 644

Rosilla, 616

Roubieva, 610

Rubiaceae, 648

Rubus, 647, 648

Rumex, 642, 643

Rush, Baltic, 661
Bog, 661
Mexican, 661
Salt, 661
Toad, 661
Western, 661
Wood, 661

Rush Family, 661

Russian Thistle, 610

Sage, Crimson, 632
Pitcher, 631, 632
Sagebrush, 612
Sagina, 609
St. John’s Wort, 631
St. John’s Wort Family, 631
Salal, 626
Salicaceae, 648
Salicornia, 610
Salix, 648, 649
Salmon Berry, 648
Salsify, 620
Salsola, 610
Salvia, 632
Sambucus, 608
Sand-Mat, 608
Sand Spurry, 609

Sandwort, 608
Sanicle, Coast, 655
Pacific, 655
Purple, 655
Sanicula, 654, 655
Satureja, 632
Saxifraga, 649
Saxifragaceae, 649
Saxifrage, 649
Saxifrage Family, 649
Scabiosa, 625
Scarlet Pimpernel, 643
Schinus, 606
Scirpus, 656
Scrophularia, 652
Scrophulariaceae, 650
Scutellaria, 632
Sea Lavender, 641
Sea-Rocket, 622
Seaside Daisy, 614
Secale, 659
Sedge Family, 656
Sedum, 621
Senecio, 618, 619
Service Berry, 645
Shamrock, 635
Shepherd’s Purse, 622
Shooting Star, 643
Sidalcea, 637
Silene, 609
Silk Tassel Bush, 628
Silk Tassel Family, 628
Silybum, 619
Sisymbrium, 623, 624
Sisyrinchium, 660
Sitanion, 659
Skull-cap, 632
Skunkweed, 641
Slim Solomon, 665
Smilacina, 665
Snake-root, 654
Sneezeweed, 616
Snowberry, 608
Soap-Plant, 663
Soft Flag, 666
Solanaceae, 652
Solanum, 652
Soleirolia, 655
Solidago, 619
Soliva, 619

675

676 CALIFORNIA ACADEMY OF SCIENCES

Solomon, Fat, 665
Slim, 665
Sonchus, 619
Sorghum, 660
Sorrel, Hairy Wood, 640
Sheep, 642
Sow-Thistle, 619
Prickly, 619
Spearmint, 631
Spergula, 609
Spergularia, 609
Sphacele, 631
Spikeweed, 616
Spinach, New Zealand, 606
Spine-Flower, 641
Spiranthes, 666
Spotted Hemlock, 653
Spurge, Caper, 627
Petty, 627
Spurry, 609
Stachys, 632
Star-Thistle, 613
Starwort, Water, 607
Stellaria, 610
Stephanomeria, 619
Stipa, 660
Stone Crop, 621
Stone Crop Family, 621
Strawberry, Beach, 646
California, 645
Stylomecon, 640
Sumac Family, 606
Suncup, 638
Sunflower Family, 611
Sweet Alyssum, 623
Symphoricarpos, 608

Tanacetum, 619
Tansy, Dune, 619
Taraxacum, 619
Tarweed, 616, 617
Coastal, 618
Slender, 617
Small, 617
Teasel, Fuller’s, 625
Teasel Family, 625
Telegraph Weed, 616
Tellima, 650
Tetragonia, 606
Thalictrum, 644

Thelypodium, 624
Thimble Berry, 647
Thistle, Barneby’s, 613

Brownie, 614

Bull, 614

Indian, 613

Italian, 613

Milk, 619

Napa, 613

Russian, 610

Spanish, 620

Star, 613
Thysanocarpus, 624
Tidy-Tips, 617
Tillaea, 621
Tinkers Penny, 631
Toad Flax, Blue, 651
Tocalote, 613
Toothwort, 622
Toyon, 646
Tragopogon, 620
Trefoil, Bird’s Foot, 633
Trifolium, 635, 636
Triglochin, 661
Trillium, 665
Triodanus, 608
Triticum, 660
Tritonia, 660
Tropaeolaceae, 652
Tropaeolum, 652
Tropaeolum Family, 652
Tule, California, 656
Twinberry, 608
Typha, 666
Typhaceae, 666

Ulex, 636
Umbelliferae, 652
Umbellularia, 632
Urtica, 655
Urticaceae, 655

Vaccinium, 627
Valerian, Red, 655
Valerian Family, 655
Valerianaceae, 655
Venus’ Looking Glass, 608
Veronica, 652
Vetch, 636

American, 636

[Proc. 47H SER.

VoL. XXXII] McCLINTOCK & KNIGHT: FLORA OF SAN BRUNO

Common, 636
Giant, 636
Spring, 636
Vicia, 636
Vinca, 606
Viola, 655, 656
Violaceae, 655
Violet, Blue, 655
California Golden, 656
Violet Family, 655

Wake-Robin, 665
Wallflower, 623
Wartweed, 627

Water Cress, 623

Water Milfoil Family, 630
Water Starwort, 607
Water Starwort Family, 607
Waterleaf Family, 630
Wattle, Green, 632

Wax Myrtle, 637

Wax Myrtle Family, 637
Wheat, 660

Wild Oat, Slender, 657

Willow, Arroyo, 649
Coulter, 648
Willow Family, 648
Willow-Herb, 639
Woodland Star, 649
Woodwardia, 605
Woolly-Heads, 618
Wormwood, 612
Wyethia, 620

Xanthium, 620

Yarrow, 611

Yellow, 615
Yellow-eyed grass, 660
Yellow mats, 654
Yerba Buena, 632
Yerba Santa, 630

Zantedeschia, 656
Zauschneria, 639
Zigadenus, 665
Zygadene, 665

677

INDEX TO VOLUME XXXII

FOURTH SERIES

New names and principal reference in boldface type

Zantedeschia, 656
aethiops, Ophiocoma, 235
affinis, Castilleja, 650
Lithophragma, 649
Lupinus, 634.
Syngathus, 370
Syngathus fuscus, 363, 370
afghanus, Staurois, 137
A Flora of the San Bruno Mountains, San
retinodes, 633 Mateo County, California, by Eliza-
Acaena californica, 644. beth McClintock, and Walter Knight,
Acanthina brevidentata, 238 In collaboration with Neil Fahy,

abdominalis, Silis, 459
Silis (Ditemnus), 452, 455, 459
stercorarius, Camponotus (Myrmo-
thrix), 250
Abraliopsis hoylei, 243
abrupta, Silis (Silis), 465, 503, 504
abstrusa, Silis (Silis), 462, 482, 485
Acacia decurrens, 632
melanoxylon, 633

Acanthochitona discrepans, 434
hirudiniformis, 242

Acer spicatum, 192, 205

acetosella, Rumex, 642

(Achelous) brevimanus, Portunus, 246

Achillea californica, 611
millefolium, 596
millefolium californica, 611
achilleaefolia, Gilia, 641
Achirus inscriptus, 366
Acmaea (Collisella) aeruginosa, 238
(Nomaeopelta) mesoleuca, 238
striata, 238
strigatella, 238
Acropora, 45
Acroteuthis winslowensis, 524, 526
acuminata, Citharina, 550, 556, 567
acuta, Pocillopora, 47
acuta, Silis (Silis), 463, 489, 491
acutangulum, Tristix, 562, 563
Adiantum jordani, 604:
adenocaulon, Epilobium, 639
adunca, Viola, 601, 655
adustum, Cerithium, 239
advena, Eggerella, 230
aenigma, Pauliella, 234
aequale, Haplophragmium, 536, 542
aeruginosa, Acmaea (Collisella) 238
Aesculus californica, 596, 630
aestivum, Triticum, 660
aethiopica, Antholyza, 660
Chasmanthe, 660

587-677

Agama, 129

himalayana, 138

tuberculata, 110, 129, 130, 138
Agamidae, 110, 138
agarici, Clythia, 64, 74

Clythia (Platypeza), 61
agassizi, Cocornis, 253

Sphenocarcinus, 246
agassizli, Ozius, 247
Agathomyia lucifuga, 64

nemophila, 64

sylvania, 64
agglutinatus, Reophax, 230

Agkistrodon himalayanus, 116, 130,

Agoseris apargioides, 611
grandiflora, 611
Agraulis vanillae, 250
agrifolia, Quercus, 596, 627
Agrostis diegoensis, 656
exarata, 657
halli, 657
semiverticillata, 657
Ahaetulla fronticincta, 139
nasuta, 139
prasina, 139
Aira caryophallea, 657
Aizoaceae, 605
ajugoides, Stachys, 632
Alabamanidae, 566
alascanus, Sebastolobus, 316
albertensis, Quadrimorphina, 566

[679]

680 CALIFORNIA ACADEMY OF SCIENCES

albidus, Orthocarpus faucibarbatus, 651

albifrons, Lupinus, 634
albirostris, Corythoichthys, 363
Albizia distachya, 633
lophantha, 633
albocintus, Oliogodon, 114, 141
albolabris, Trimeresurus, 116, 142
albopunctata, Riopa, 139
album, Chenopodium, 610
albus laevigatus, Symphoricarpos, 608
Melilotus, 635
Alchemilla occidentalis, 645
alexandri, Ophiocoma, 235
Allium, 663
dichlamydeum, 601, 661, 662
unifolium, 601, 663
triquetrum, 661
Allocarya chorisiana, 607
almgreni, Gaudryinella, 538, 540
alsinoides, Chaetopappa, 613
Alsophylax, 129
tibetanus, 138
alticola, Scutiger, 127, 136
Altirana parkeri, 127, 137
Allysum maritimum, 623
Amalthea grayana, 239
Amaranthaceae, 606
Amaranthus powellii, 606
Amelanchier, 195
pallida, 597, 598, 645
americana, Veronica, 652
Vicia, 636
americanum, Macrobranchium, 248
americanus americanus, Coccyzus, 405
occidentalis, Coccyzus, 405
Ammobaculoides romaensis, 535
Ammodiscacea, 532
Ammodiscidae, 532
Ammodiscinae, 532
Ammodiscus, 532, 533, 570
glabratus, 533, 568
pacificus, 229
Amphelikturus, 366
dentriticus, 364
Amphibia, 107, 117, 136, 143, 144
Amphineura, 242
Amphiodia violacea, 234
amphioxys, Nodosaria, 543
Amphipoda, 245
amplectens, Trijolium, 635

[Proc. 4TH Serr.

amplexicaulis, Smilacina racemosa, 665
ampulla, Globulina, 565, 572
Amsinckia intermedia, 607
spectabilis, 386
Anabus, 429
(Anadara) reinharti, Arca, 237
Anadara (Scapharca) reinharti, 237
anagallis arvensis, 64.3
anagalloides, Hypericum, 631
Anaphalis margaritacea, 596, 597, 601, 611
Anarcardiaceae, 606
Anarchopterus, 372
(Anarchopterus) crinigerus, Micrognathus,
364, 371
crinitus, Micrognathus, 364, 371
andersoni, Bufo, 107, 123, 136
andersoni, Planulina, 567, 572
andersonii, Polygala, 582
androsaceus, Linanthus, 641
Angelica, 654
hendersonii, 599, 652
angelica, Silis (Silis), 464, 495, 497
Angiospermae, 603, 605
Anguidae, 138
Anguilliformes, 88
angulata, Silis (Silis), 463, 483, 488
angustifolia, Vicia, 636
angustifolium, Apiastrum, 653
angustirostris, Mirounga, 377, 378
annandali, Philautus, 137
annandalii, Rana, 137
Annelida, 244
annua, Lunaria, 623
Poa, 659
Anolis townsendi, 252
Anomalina popenoel, 567
Anomalinidae, 566
Anomalininae, 566
Anomura, 248
Anoplodera nitens, 189
Anoplopoma fimbria, 316
Anosia plexipus, 419
Anthemis cotula, 612
Antholyza aethiopica, 660
Anthophilax, 193
Anthopleura elegantissima, 515
Anthozoa, 231
Anthriscus caucalis, 653
scandicina, 653
Antigonia capros, 366

VoL. XXXII]

antiquissima, Bigenerina, 539, 542
Anura, 136, 137
apargioides, Agoseris, 611
aparine, Galium, 648
Apatophsis, 154, 155, 196
barbara, 155, 192, 195
caspica, 155
kamarowl, 155
kashmiriana, 155
modica, 155
montana, 155
serricornis, 155
sinica, 155
toxotoides, 155
aper, Capros, 330
apetala barbata, Sagina, 609
Apiastrum angustifolium, 653
apiculata, Chaetopleura, 434
apiifolium, Ligusticum, 654
Aplousina filum, 232
Apocynaceae, 606
Apocynum pumilum, 598, 606
Apoda, 136
Aquilegia formosa truncata, 643
aquilinum, Pteridium, 599
pubescens, Pteridium, 605
Arabis blepharophylla, 597, 601, 621
glabra, 621
Araceae, 656
Arachnida, 252
Araliaceae, 606
arborea, Lavatera, 637
arboreus, Lupinus, 386, 596, 597, 634:
arbuscula, Vaccinium, 599, 601, 627
arbutifolia, Heteromeles, 600, 601, 646
Photinia, 646
Arbutus menziesii, 195, 202, 601, 625
unedo, 625
Arca (Anadara) reinharti, 237
arctopoides, Sanicula, 596, 654
_ Arctostaphylos, 626
imbricata, 600, 626
montaraensis, 600, 626
pacifica, 600, 626
uva-ursi, 598, 601, 626
Arenaria macrophylla, 608
arenicola, Brandtothuria, 236
areolata, Retevirgula, 233
Argonauta cornuta, 243
cornutus, 243

INDEX

expansa, 243
arguta, Dryopteris, 599, 605
Erechtites, 614.
Argyroepeira nigriventris, 252
arida, Silis (Silis), 463, 482, 485
aristata, Lithophaga (Myoforceps), 238
Aristolochia, 606
californica, 601, 606
Aristolochiaceae, 606

arizonica, Silis (Silis), 455, 464, 489, 493

armitagel, Silis, 458

arnensis, Oligodon, 114, 117, 141
aronicoides, Senecio, 618
Arrhinotermes oceanicus, 250

Artemisia californica, 596, 597, 600, 612,

650
douglasiana, 612
Arthropoda, 245
articulata, Textularia, 230
arvense, Cerastium, 609
Equisetum, 604.
arvensis, Anagallis, 643
Convolvulus, 620
Spergula, 609
asellus, Lepidopleurus, 434, 438, 441
asper, Sonchus, 619
aspersa, Littorina, 239
asperula, Madracis, 44, 46
assamensis, Rana, 137
assurgentiflora, Lavatera, 637
Astacolus grata, 555, 558
incurvata, 555
perobliqua, 550, 556
Aster chilensis, 612
exilis, 612
radulinus, 612
subspicatus, 612
Asterias, 328
stellata, 328
Asteroidea, 234
asthenes, Lepidozona, 438, 439, 441, 442
Astragalus nuttallii virgatus, 633
Astrangia, 44, 52
dentata, 45, 46, 52, 231
equatorialis, 44, 45, 46, 52
gardnerensis, 44, 45, 46, 52
hondaensis, 44, 46, 231
Astrorhizidae, 539
Ataxophragmiidae, 536
Athyrium filix-femina, 604

682 CALIFORNIA ACADEMY OF SCIENCES

atra, Ludwigothuria, 237
Silis, 402
Silis (Silis), 445, 462, 477, 480
Atriplex hastata, 610
patula hastata, 610
rosea, 610
serenana, 610
atropurpurea, Scabiosa, 625
Atta cephalotes, 250
attenuata, Lithophaga, 238
Aucella, 527
crassicollis, 527
inflata, 525, 527
solida, 525, 527
uncitoides, 525, 527
aulicus, Lycodon, 113, 140
aurantiacus, Diplacus, 596,
651
aurea, Tritonia, 660
aureola, Dendroica, 253
Dendroica petechia, 253
Auricula stagnalis, 239
auritum, Pholistoma, 631
aurivilliusi, Lissa, 245
auropunctata rugosa, Wasmannia, 251
Wasmannia, 224, 251
australis, Cotula, 614
autumnata, Centrodera, 149, 159, 160, 162,
163,164, 166, 168.1725 0-73eu 74 mite
17S 182 1185 setevies, 11s 7 atsomtene
197, 200, 201, 202, 203, 204, 205, 206,
207
Avena barbata, 657
Aves, 253
aviculare, Polygonum, 642
avis, Kionotrochus (?), 44, 46
Azteca emmae, 250

597, 600, 601,

Baccharis pilularis, 597, 600, 601
pilularis consanguinea, 596, 612
pilularis pilularis, 612

Baeria chrysostoma, 596, 598, 644
chrysostoma chrysostoma, 612
chrysostoma hirsutula, 613

baileyi, Thermophis, 129

Balanophyllia, 44, 52
galapagensis, 44, 46
osburni, 44, 45, 46, 52, 53
scheeri, 42, 45, 46, 53, 54, 231

balticus, Juncus, 661

[Proc. 4TH SER.

ee ee en

barbara, Apatophysis, 155, 192, 195
barbarae, Carex, 656 |
Barbarea orthoceras, 621

verna, 622
barbata, Avena, 657

Sagina apetala, 609
barbatus, Hipponix, 239
barnesi, Chiton, 442
barremiana, Gavelinella, 566, 572
barri, Silis (Silis), 467, 503, 507
Bassia hyssopifolia, 610
beebei, Helicocranchia, 243

Melanoteuthis, 243
Bellardia trixago, 650
Belemnopsis, 524

gerardi, 524

sulcatus, 524
Bellis perennis, 613
belti, Pseudomyrma, 251
bellum, Sisyrinchium, 596, 601, 660
beneolens, Gnaphalium, 615
benghalensis, Vicia, 636
Berberidaceae, 607
Berberis pinnata, 597, 601, 607
beolandieri, Chenopodium, 610
bernhardi, Nerita, 240

Nerita fulgurans, 240
berryi, Menkenina, 550, 560, 561
Betula lutea, 192
Betulaceae, 607
Bhanotichthys, 365
biauriculata, Silis, 453
bicolor, Centrodera, 191

Lupinus, 596, 634.

Pheidole biconstricta, 251
biconstricta bicolor, Pheidole, 251
bidentata, Cantharis, 452

Ditemnus, 451

Silis (Ditemnus), 451, 452, 455
bifasciatum, Rhagium, 207
bifida, Euprognatha, 245
bifidum, Trifolium, 635
biflora, Triodanus, 608
Bigenerina antiquissima, 539, 542

deciusi, 539, 540, 452
Bimonilina variana, 540

biolleyi, Camponotus (Myrmobrachys), 250
Leptinaria, 242
Ochrodermella, 242
Rhinocricus (Eurhinocricus), 249

VoLt. XXXII]

bipinnatifida, Franseria, 386
Sanicula, 655
bipinnatisecta, Franseria chamissonis, 615
biserialis, Thais haemastoma, 241
biunguiculatus, Brachycarpus, 248
blainvilleanus, Leptonotus, 364
blainvillei, Placiphorella, 243
blandus, Camponotus (Myrmobrachys), 250
Camponotus (Myrmocamelus), 250
blanfordi, Rana, 125, 130, 137
Blennosperma nanum, 613
blepharophylla, Arabis, 597, 601, 621
Boidae, 112, 139
Boiga ceylonensis, 112, 140
cyanea, 140
cynodon, 140
forsteni, 131, 140
gokool, 140
multifasciata, 112, 140
ochracea, 140
trigonata, 113, 140
bolanderi, Chrysopsis villosa, 613
Boraginaceae, 607
bothriorynchus, Typhlops, 142
botrys, Erodium, 629
boulengeri, Rana, 130
Bowlesia incana, 653
bowringi, Hemidactylus, 139
Brachiopoda, 234
brachyantherum, Hordeum, 658
Brachycarpus biunguiculatus, 248
brachycephalus, Corythoichthys, 363
Brachymyrmex longicornis, 250
Brachyura, 245
bramius, Typhlops, 142
branchialis, Microphrys, 246
brandti, Schizoplax, 435, 437, 442
Brandtothuria arenicola, 236
impatiens, 236
brasiliense, Myriophyllum, 630
.Brassica campestris, 600, 622
geniculata, 622
kaber, 622
oleracea, 622
braunii, Equisetum telmateia, 604.
Breeding Characteristics of Yellow-Billed
Cuckoos in Arizona, by William J.
Hamilton, III, and Marian E. Ham-
ilton, 405-432
brevicaulis, Carex, 656

INDEX 683

breviceps, Rana, 108, 137
Brevicipitidae, 136
brevidentata, Acanthina, 238
Monoceros, 238
brevidentatum, Monoceros, 238
brevior, “Conger”, 87, 90, 92, 95, 96, 98, 99
brevistylum, Cirsium, 613
brighami, Psammocora, 45
Briza maxima, 657
minor, 657
Brodiaea, 663
capitata, 663
coronaria macropoda, 663
laxa, 596, 601, 663
pulchella, 596, 663
terrestris, 663
Bromus, 603
carinatus, 657
diandrus, 657
marginatus, 657
mollis, 657
rubens, 657
brooki, Hemidactylus, 139
(Brotulidarum), Otolithus, 92
brunnea, Holoporella, 233
brunneus, Conus, 239
Juncus effusus, 661
Bryozoa, 232
Buchia crassicollis, 527
inflata, 527
solida, 527
Bufo andersoni, 107, 123, 136
himalayanus, 107, 136
melanostictus, 107, 136
Bufonidae, 107, 136
bufonius, Juncus, 661
Bulbophragmium, 535, 542
Bulimina, 570
ovata, 359
Buliminacea, 570
Bulimulus, 242
bullata, Marginulina, 548
Stachys, 632
Bungaris caeruleus, 141
lividus, 141
niger, 142
walli, 142
bungaroides, Bungarus, 141
Bungarus bungaroides, 141
fasciatus, 141

684 CALIFORNIA ACADEMY OF SCIENCES

Bursa caelata, 238
bursa-pastoris, Capsella, 662
Butler, C. P., The Goose Lake Fragments,
by, 291-313
Butorides virescens, 253
virescens maculatus, 253

Cactornis inornata, 253
Caducifer cinis, 238
thalia, 238
caelata, Bursa, 238
Ranella, 238
caeruleus, Bungaris, 141
caespitosa holciformis, Deschampsia, 658
Cakile maritima, 386, 622
Calamagrostis nutkaensis, 657
calamaria, Liopeltis, 140
Calamphyta, 603, 604
Calandrinia ciliata menziesii, 643
Calappa convexa, 248
Calappidae, 248
calceata, Calotarsa, 57, 60, 61
Calcinus explorator, 248
calcitrapa, Centaurea, 613
californianus californianus, Zalophus, 379
japonicus, Zalophus, 379
wollenbaeki, Zalophus, 379
Zalophus, 377, 378, 379
californica, Acaena, 644.
Achillea millefolium, 611
Aesculus, 596, 630
Aristolochia, 601, 606
Artemisia, 596, 597, 600, 612, 650
Chenopodium, 386
Cornus, 599, 601, 621
Corylus, 607
Danthonia, 658
Eschscholzia, 601, 64.0, 650
Festuca, 658
Filago, 615
Fragaria, 598, 601, 64.5, 646
Grossularia, 600, 649
californica, Gubkinella, 568, 573
Horkelia, 64.6, 647
integrifolia, Dentaria, 622
Meconella, 640
Melica, 659
Myrica, 599, 601, 637
Phacelia, 597, 630
Phalaris, 659

[Proc. 4TH Serr.

Plantago hookeriana, 640
Pleurobranchaea, 515, 516, 517, 519
Porites, 51
Protoclythia, 58, 59, 60, 61, 62, 63, 64,
66, 67, 68, 75
Rafinesquia, 618
Rhamnus, 598, 600, 644.
Rosa, 601, 647
Saxifraga, 649
Scrophularia, 652
Silis, 494
Silis (Silis) , 463, 464, 489, 493
Solidago, 619
Umbellularia, 596, 599, 632
Zauschneria, 601, 639
californicum, Chenopodium, 610
Delphinium, 599, 601, 643
Eriodictyon, 630
Galium, 648
Gnaphalium, 615
Limonium, 600, 64.1
Polypodium, 597, 605
Ribes, 649
Sisyrinchium, 660
californicus, Corylus, 195
Croton, 597, 627
Micropus, 618
Platystemon, 640
Ranunculus, 596, 644.
Rumex, 64.2
Scirpus, 656
californiensis, Lingulina, 560, 564
Pagurus, 248
calliati, Murex, 366
callicarpa, Sambucus,
callichroa, Ostrea, 238
Calliophis macclellandi, 142
macclellandii, 116
Callistochiton viviparus, 435, 436, 441, 442
Callitrichacae, 607
Callitriche marginata, 607
Callomyia, 64
Callorhinus ursinus, 378, 401
Calotarsa calceata, 57, 60, 61
insignis, 60, 61, 64, 74
Calotes versicolor, 105, 110, 123, 138
calycina, Lepechinia, 599, 600, 631
Sphacele, 631
Camissonia dentata, 638
micrantha, 638

204, 599, 600, 608

Voi. XXXII]

ovata, 638
Campanulaceae, 608
campestris, Brassica, 600, 622
Layia platyglossa, 617
camphoratum, Tanacentum, 598,
Camponotus (Myrmaphaenus)
251
(Myrmobrachys) biolleyi, 250
(Myrmobrachys) blandus, 250
(Myrmocanelus) blandus, 250
(Myrmothrix) abdominalis
ius, 250
camporum, Grindelia, 616
canadensis, Conyza, 614:
elongata, Solidago, 619
Linaria, 651
Shepherdia, 195
canariensis, Phalaris, 659
Cantharis bidentata, 452
Cantharus gemmatus, 238
(Gemmophos) gemmatus, 238
sanguinolentus, 238
cantoris, Elaphe, 113, 140
capensis, Pentaceros, 321, 322, 324, 325, 326,
327, 331
Quinquarius, 324, 325
capillare, Panicum, 659
capitata, Brodiaea, 663
Madia, 617
capitatus, Physocarpus, 601, 646
Caprifoliaceae, 608
Capros, 330
capros, Antigonia, 366
Capros aper, 330
Capsella bursa-pastoris, 622
caracolla caracolla, Epistomina,
Epistomina caracolla, 569, 572
carbo, Silis (Silis), 461, 475, 477
Cardamine oligosperma, 622
Cardaria draba, 622
Cardionema ramosissimum, 597, 608
Cardiosoma crassum, 247
Carduus pycnocephalus, 613
tenuiflorus, 613
Carex, 603
barbarae, 656
brevicaulis, 656
densa, 656
obnupta, 656
subbracteata, 656

619

cocosensis,

stercorar-

DOG), Siz

INDEX 685

tumulicola, 656
carinata, Mabuya, 111, 139
carinatus, Bromus, 657
carmelita, Silis (Silis), 461, 467, 469
carnosa, Jaumea, 600, 617
carolinianum, Geranium, 629
Carpilodes cinctimanus, 246
caruifolium, Lomatium, 654.
Caryophyllaceae, 603, 608
caryophyllea, Aira, 657
Caryophyllia, 44
diomedae, 44, 46
caspica, Apatophysis, 155
Cassidulinacea, 566, 569
castanea, Helianthella, 616, 620
Castanopsis chrysophylla, 204
casteneus, Neotermes, 250
Castilleja affinis, 650
franciscana, 598, 650
wightii, 650
catenula, Dentalina, 544
caucalis, Anthriscus, 653
caucilis microcarpa, 653
cava, Silis, 449, 492
Silis (Silis), 455, 460, 463, 489, 492
Cavolina longirostris constricta, 242
Ceanothus thyrsiflorus, 598, 600, 601, 644
Cellaria veleronis, 232
Celtis, 407
reticulata, 407
Centaurea calcitrapa, 613
melitensis, 613
solsitialis, 613
Centaurium davyi, 628
floribundum, 628
Centranthus ruber, 655
Centrechinus (Diadema) mexicanus, 235
Centrocidaris doederleini, 236
Centrodera, 149, 154, 155, 156, 157, 184, 190,
191, 196, 201, 204, 205
autumnata, 149, 159, 160, 162, 163, 164,
166. 168 172, 17S, W7As17 7 7s eee
183, 184, 185, 187, 190, 192, 197, 200,
201, 202, 203, 204, 205, 206, 207
bicolor, 191
dayi, 149, 159, 162, 163, 164, 166, 168,
VAs, 172) AW TLIO U9 Sy 18385:
187, 190, 192, 200, 201, 202, 204, 205,
207

686 CALIFORNIA ACADEMY OF SCIENCES

decolorata, 154, 157, 158, 184, 185, 186,
191, 202
hirsuta, 178
nevadica, 158, 186
picta, 154, 156, 186, 192, 202, 205
Souci, WA), WSO) iWsal USA. ash ISS, ISO.
157, ie, ICO, soil, Wee, ies, leek
LG GHG OS On lyre aemely Samal.
L757, LIS 180. Weow ese aied:
185, 187, 188, 189, 190, 191, 192, 193,
194, 195, 196, 197, 198, 201, 202, 203,
204, 205, 206, 207, 208
Centrodera spurca (LeConte) and Two New
Species Resembling it, with Biolog-
ical and other notes (Coleoptera:
Cerambycidae), by Hugh B. Leech,
149-218
(Centrodera) spurcus, Evodinus, 160
Centrodera sublineata, 185, 201
Centroderus, 154, 162
Cephalopoda, 243
cephalotes, Atta, 250
cerasiformis, Osmaronia, 600, 601, 64.6
cerasogaster, Xenochropis, 166, 141
Cerastium arvense, 609
glomeratum, 609
viscosum, 609
Ceratobuliminidae, 569, 570
Ceratobulimininae, 570
cereale, Secale, 659
Cerithium adustum, 239
maculosum, 239
cernua, Stipa, 660
cernuus californicus, Scirpus, 656
cervinus, Typocerus, 160, 161, 168
cespitosa, Pocillopora damicornis, 42, 44, 45,
48
Cetoconcha scapha, 237
ceylonensis, Boiga, 112, 140
Chaetopappa alsinoides, 613
Chaetopleura apiculata, 434

chamissonis bipinnatisecta, Franseria, 615
Lupinus, 597, 634.

Chaperiella condylata, 232

characteristica, Siphonaria gigas, 241, 243

charlottensis, Trochammina, 230
Chasmanthe aethiopica, 660
chatamensis, Polygala, 585
cheiranthifolia, Denothera, 386
Chelonia, 137, 143, 144

[Proc. 4TH Ser.

chemnitziana, Melina, 237
chemnitzianum, Isognomon (Melina), 237
Pedalion 237
Perna, 237
Chenopodiaceae, 610
Chenopodium album, 610
berlandieri, 610
californica, 386
californicum, 610
multifidum, 610
murale, 610
chera, Rachodrassus, 79, 83
Chesterman, Charles W., Contributions to
the Petrography of the Galapagos,
Cocos, Malpelo, Cedros, San Benito,
Tres Marias, and White Friars Is-
lands, 339-362
chilense, Gnaphalium, 615
chilensis, Aster, 612
chiloensis, Fragaria, 598, 601, 645, 646
Chilostomellinae, 566
Chiton barnesi, 442
(Chiton) goodallii, 243
(Chiton) stokesii, 242
goodallii, 243
(Chiton) goodallii, Chiton, 243
Chiton marmoratus, 434
polii, 434, 436, 441
(Radsia) goodallii, 243
(Radsia) stokesi, 242
squamosus, 434
stokesi, 242
stokesii, 243
tuberculatus, 434
Chitra indica, 105, 110, 138
Chivers, Dustin D., Observations on Pleuro-

branchaea californica MacFarland,
1966 (Opisthobranchia, Notaspidea),
515-521

Chloeia entypa, 244
euglochis, 244
viridis, 244
Chlorogalum pomeridianum, 663
chloropetalum, Trillium, 601, 665
Chlorostoma gallinum multifilosum, 241
maculostriatum, 241
Chordata, 252
chorisiana, Allocarya, 607
Chorizanthe pungens hartwegii,
chrysanthemoides, Layia, 617

597, 641

Vout. XXXII]

Chrysanthemum coronarium, 613
chrysolepis nana, Quercus, 599, 628
Quercus, 627
Chrysopelea ornata, 140
chrysophylla, Castanopsis, 204
Chrysopsis villosa bolanderi, 613
chrysostoma, Baeria, 596, 598, 644
Baeria chrysostoma, 612
chrysostoma, Baeria, 612
hirsutula, Baeria, 613
Church, Clifford C., Lower Cretaceous
Foraminifera of the Orchard Peak-
Devils Den Area, California, 523-580
Cibicididae, 567
Cichorium intybus, 613
cicutarium, Erodium, 629
ciliata menziesii, Calandrinia, 643
Microporella, 233
Mopalia, 434
cinctimanus, Carpilodes, 246
cinerea, Clythia, 64, 68
Juglans, 192
Nuttallina, 436, 442
Nuttallina (Middendorffia), 434, 441
cinereus, Ischnochiton, 433, 434, 441, 442
cingulata, Herse, 250
Phleganthontius, 250
cinis, Caducifer, 238
Pollia, 238
Tritonidea, 238
cirratulus hirsutus, Nerine, 244
Cirripedia, 245
Cirsium brevistylum, 613
quercetorum, 614
vulgare, 614.
cisovnicensis, Pseudoreophax, 537, 542
Citharina, 526, 550, 556, 557
acuminata, 550, 556, 557
kernensis, 556, 557, 558
Cladocora, 44
debilis, 44, 46, 231
Clarkia davyi, 638
purpurea, 639
rubicunda, 601, 639
Cleidochasma contracta, 232
clivorum, Gilia, 641
clivosa, Pavona, 42, 44, 45, 46
Pavona (Pavona), 49
Clypeaster ochrus, 236
rotundus, 236

INDEX 687

speciosus, 236
Clythia, 67
agarici, 64, 74
cinerea, 64, 68
coraxa, 64
cylindrica, 230
dymka, 64
hunteri, 64
(Platypeza) agarici, 61
polypori, 64
coarctata, Orthomorpha, 249
Coccyzus americanus americanus, 405
americanus occidentalis, 405
ferrugineus, 253
(Nesococcyx), 253
cocoénsis, Cotylopus, 252
Petrolisthes, 248
Prenolepis (Nylanderia) guatemalensis,
251
Prenolepis guatemalensis, 251
(Cocopupa) cocosensis, Nesopupa, 242
Cocornis agassizi, 253
Cocos Island, Including a_ Bibliography,
Contributions to the Biogeography of,
by Leo George Hertlein, 219-289
cocosana, Liomera, 246
(Cocosconus) hopkinsi, Habroconus, 242
cocosensis, Camponotus (Myrmaphaenus),
251
Nesopupa (Cocopupa), 242
Olivella (Olivella) , 240
Pecten (Cyclopecten), 238
Phos, 240
cocosi, Encope, 236
Encope micropora, 236
(Cocoslens) pallidus, Habroconus, 242
Coeciliidae, 136
Coelenterata, 230
Coenobita compressus, 248
Coenobitidae, 248
coerulea, Sambucus, 204
collaris, Sibynophis, 115, 141
Colletosia radiata, 233
collinsi, Marginulinopsis, 551
Collinsia franciscana, 650
heterophylla, 650
sparsiflora solitaria, 650
(Collisella) aeruginosa, Acmaea, 238
Collisella mesoleuca, 238

688 CALIFORNIA ACADEMY OF SCIENCES

Coluber diadema, 140
fasciolatus, 113, 140
monticolus, 117
ventromaculatus, 140

Colubridae, 112, 117, 139, 140, 141

Columbella cribraria, 240
labrosa, 241

columbiae, Linckia, 234

columellaris, Purpura, 241
Thais, 241

commissuralis, Obelia, 230

communis, Dentalina, 544

Compositae, 603

compressus, Coenobita, 248

Conchoderma, 245
virgatum, 245

concinna, Frondicularia, 559
Triglochin, 600, 661

Condalia, 425, 426
lycioides, 407

condanarus, Psammophis, 141

condylata, Chaperiella, 232

confertiflorum, Eriophyllum, 615

conica, Textularia, 230

conicus, Eryx, 139

Conium maculatum, 653

consanguinea, Baccharis pilularis, 596, 612
Conger, 88, 90, 92
“Conger” brevior, 87, 90, 92, 95, 96, 98, 99

Conger conger, 101
dissimilis, 92, 94, 95, 96, 98
fallax, 99
fornicatus, 92, 95, 96, 97
meridies, 92, 95, 96, 97, 99
sanctus, 92, 93, 95, 96, 98, 99
vetustus, 88, 90, 92, 95, 100, 101
(Congeris) , Otolithus, 87, 92
“Congermuraena” sector, 87
congesta, Plectritis, 655
congestus, Juncus tenuis, 661
conglomeratus, Rumex, 64.2
Congridae, 88
(Congridarum), Otolithus, 92
Conorboides, 570
umiatensis, 570
conspersa, Littorina, 239
constanti, Ischnochiton, 436, 442
constricta, Cavolina longirostris, 242
constricta, Silis (Silis), 467, 503, 506

[Proc. 4TH SER.

contorta murrayana, Pinus, 204
strigulosa, Oenothera, 638
contracta, Cleidochasma, 232
Contribution of the Biogeography of Cocos
Island, Including a Bibliography, by
Leo George Hertlein, 219-289
Contributions to the Petrography of the
Galapagos, Cocos, Malpelo, Cedros,
San Benito, Tres Marias, and White
Friars Islands, by Charles W. Ches-
terman, 339-362
controversa mexicana, Cypraea, 239
Conus brunneus, 239
dalli, 239
gladiator, 239
recurvus, 239
tiaratus, 239
convexa, Calappa, 248
Convolvulaceae, 620
convolvulus arvensis, 620
occidentalis, 620
subacaulis, 620
Conyza canadensis, 614
cooksoni, Leptodius, 247
Tegula, 241
cooperl, Lepidozona, 434
Copepoda, 245
Copidozoum tenuirostre, 233
Coprosma repens, 648
Corals from the Galapagos and Cocos Is-
lands, by J. Wyatt Durham, 41-56
coraxa, Clythia, 64
cordiformis, Lovenia, 236
coriacea inarmata, Micopora, 233
Cornaceae, 621
corniculatus, Lotus, 633
Cornus californica, 599, 601, 621
cornuta, Argonauta, 243
cornutus, Argonauta, 243
coronaria macropoda, Brodiaea, 663
coronarium, Chrysanthemum, 613
Coronaster marchenus, 234
coronopifolia, Cotula, 614:
Coronopus didymus, 622
coronopus, Plantago, 640
corrugata, Textularia, 230
Cortaderia selloana, 657
Corylus californica, 607
californicus, 195
corymbosa, Hemizonia, 616

VoL. XXXII] INDEX 689

Corythoichthys, 365 Croton californicus, 597, 627
albirostris, 363 erucialis, Silis (Silis), 463, 489, 490
brachycephalus, 363 cruciatum, Dodecatheon hendersonii, 643
ensenadae, 372 Cruciferae, 603
profundus, 363, 364, 365 cruentus, Senecio, 618
vittatus, 372 Crustacea, 245

Cosymbotus platyurus, 111, 123, 138 Cryptantha hispidissima, 607

Cotoneaster pannosa, 645 micromeres, 607

cotula, Anthemis, 612 Cryptocerus cristatus, 251

Cotula australis, 614 Cryptochiton stelleri, 434
coronopifolia, 614 cubana, Kathetostoma, 366

Cotylopus cocoensis, 252 Cuckoos in Arizona, Breeding character-

coulteri, Salix, 598, 648, 649, 656 istics of Yellow-Billed, by William J.

covensis, Drassylus, 79, 83, 84 Hamilton, IIJ, and Marian E. Ham-

craneana, Hemithyris, 234 ilton, 405-432
Hispanirhynchia ?, 234 Cucurbitaceae, 624

crassa, Thais, 241 cumingiana, Ochrodermella, 242

crassicaulis, Sanicula, 655 Cupressaceae, 605

crassicollis, Aucella, 527 Cupressus macrocarpa, 597, 605
Buchia, 527 curvipes, Thysanocarpus, 624.

crassicornis, Eciton (Labidus), 251 curvirostris, Parellisina, 233

crassipes, Pachygrapsus, 247 curvitura, Marginulina, 548

Crassulaceae, 621 cyanea, Boiga, 140

crassum, Cardiosoma, 247 Planes, 247

Crataegus monogyna, 645 cyanochloris, Dendrelaphis, 140

crenata, Harpa, 239 cyanophlyctis, Rana, 108, 137

Crenella divaricata, 237 Canoplax, 434

crenulata, Euphorbia, 627 dentiens, 434, 436, 442

Crepidacantha poissoni, 233 Cyclamen, 643
setigera, 233 (Cyclopecten) cocosensis, Pecten, 238

Creseus virgula, 242 exquisitus, Pecten, 238

cretica, Lavatera, 637 Cycloseris, 44, 51

cribraria, Columbella, 240 elegans, 45, 46, 52

Cribrostomoides, 535 mexicana, 44, 46, 231

crinigerus, Micrognathus, 371 cyclurus, Oligodon, 141
Micrognathus (Anarchopterus), 364, 371 cylindrica, Clytia, 230

crinita, Notopygos, 244 Cymatium vestitum, 239

crinitus, Micrognathus, 371, 372 cymodoce ferruginea, Trapezia, 247
Micrognathus (Anarchopterus), 364, cymosa, Dudleya, 597, 601, 621

371 cynodon, Boiga, 140
Syngnathus, 372 Cynoglossum grande, 607

crispus, Rumex, 64.2 Cyperaceae, 656

cristata, Koelaria, 659 Cyperus eragrostis, 656

cristatus, Cryptocerus, 251 Cyphastrea, 45

Cristellaria hamata, 551 Cyphomyrmex rimosus slavini, 251

cristulipes, Teleophrys, 246 Cypraea, 244

crocea, Rhamnus, 597, 599, 644: controversa mexicana, 239

Crocodilidae, 109, 138 isabella, 239

Crocodilus palustris, 138 isabella mexicana, 239

crocosmaeflora, Tritonia, 660 (Luria) isabella, 239

690 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Srp.

moneta, 239 dendriticus, Amphelikturus, 364

(Monetaria) moneta, 239 Dendroeca petechia, 253

rashleighana, 239 Dendroica aureola, 253
Cyrtodactylus fasciolatus, 138 petechia aureola, 253

gubernatoris, 139 densa, Carex, 656

khasiensis, 139 densiflorus, Lithocarpus, 204

lawderanus, 139 Orthocarpus, 651
Cytisus maderensis, 633 Dentalina, 546, 547

monspessulanus, 633 catenula, 544

scoparius, 633 communis, 544.

debilis, 547
dalli, Conus, 239 jugosa, 230
damicornis cespitosa, Pocillopora, 42, 44, 45, filiformis, 230
46, 48 grahami, 545, 550

Pocillopora, 42, 45, 46, 47, 54, 231 Dentaria californica integrifolia, 622
danianum, Diopederma, 234 dentata, Astrangia, 45, 46, 52, 231
Danthonia californica, 658 Camissonia, 638
daphnense, Flabellum, 44, 46 dentelifer; Drassylus, 84
Darbyella, 555, 568 denticulata gracilis, Reteporellina, 233
dasycarpum, Lomatium, 654. denticulatifrons, Hippa, 248
Dasypeltidae, 141 denticulatus, Mithrax (Mithraculus), 246
Daucus pusillus, 654 dentiens, Cyanoplax, 434, 436, 442
davyi, Centaurium, 628 dentigera, Silis, 505

Clarkia, 638 dentigera, Silis (Silis), 465, 497, 499

dayi, Centrodera, 149, 159, 162, 163, 164, 166, | depressa, Salicornia, 610
168, 171, 172, 177, 178, 179, 182, 183, depsarius, Tragosoma, 195
185, 187, 190, 192, 200, 201, 202, 204, depressus, Paradasygius, 246

205, 207 dertonensis, Festuca, 658
debilis, Cladocora, 44, 46, 231 Deschampsia caespitosa holciformis, 658
Dentalina, 547 deserticola, Silis, 462
Stenorhynchus, 246 Silis (Silis), 455, 460, 463, 483, 485
Vaginulina, 546, 564 Desmophyllum, 44
decacanthus, Pentaceros, 326, 327, 331 galapagense, 44, 46
Decapoda, 245 dhongoka, Kachuga, 137
deciusi, Bigenerina, 539, 540, 542 Diacria quadridentata parva, 242
decolorata, Centrodera, 154, 157, 158, 184, diadema, Coluber, 140
185, 186, 191, 202 (Diadema) mexicana, Centrechinus, 235
decoloratum, (Rhagium), 154 diandrus, Bromus, 657
Rhamnusium ?, 154 Dicentra formosa, 599, 601, 628
decorum, Delphinium, 644. dichlamydeum, Allium, 601, 661, 662
decurrens, Acacia, 632 Dichondra, 620
Delphinium californicum, 599, 601, 643 donelliana, 620
decorum, 644. micranthra, 620
demissa, Prunus, 601, 64.7 repens, 620
Prunus virginiana, 201 dictyota, Ophionereis, 235
Dempster, Lillian J., see Follett, W. I. didymus, Coronopus, 622
Dendrelaphis cyanochloris, 140 diegoensis, Agrostis, 656
gorei, 140 difficilis, Microthele, 237
pictus, 140 occidens, Silis (Silis), 462, 479

tristis, 140 Silis, 449, 475, 477, 479, 480

Vor. XXXII]

Silis (Silis), 455, 460, 461, 462, 477, 478
digitata, Leptoseris, 44, 46, 231
dilacerata, Silis, 455
dilatatum, Maianthemum, 599, 601, 663
Dinodon gammei, 140
septentrionalis, 140
diomedae, Caryophyllia, 44, 46
diomedeae, Psolus, 237
Diopederma danianum, 234
Diplacus aurantiacus, 596, 597, 600, 601, 651
Dipsaceae, 625
Dipsacus fullonum, 625
sativus, 625
Discodon, 447, 451, 457
discolor, Holodiscus, 600, 601, 646
Discoporella umbellata, 233
discrepans, Acanthochitona, 434
disjuncta, Silis (Silis), 465, 497, 500
Disporum hookeri, 663
dissectum, Geranium, 629
dissimilis, Conger, 92, 94, 95, 96, 98
distachya, Albizia, 633
distans, Phacelia, 630
Distichlis spicata, 386, 658
Distinctive “Congrid Type” Fish Otoliths
from the Lower Tertiary of the Gulf
Coast (Pisces: Anquilliformes), by
Don L. Frizzell, and C Kurt Lamber,
87-101
Ditemnus, 447, 448, 450, 451, 457
(Ditemnus) abdominalis, Silis, 452, 455, 459
Ditemnus bidentata, 451
(Ditemnus) bidentata, Silis, 451, 452, 455
fossiger, Silis, 451, 453, 455
freemani, Silis, 451, 453, 455
howdeni, Silis, 452, 455, 456
knulli, Silis, 452, 455, 458
latiloba, Silis, 451, 452, 455
Ditemnus marginella, 451
(Ditemnus) nigerrima, Silis, 452, 455, 458
obtusa, Silis, 451, 455, 456
perforata, Silis, 452, 455, 458
tricornis, Silis, 451, 455, 456
vandykei, Silis, 452, 455, 457
divaricata, Crenella, 237
Grossularia, 649
divaricata, Silis (Silis), 464, 497, 498
divaricatum, Ribes, 649
diversa, Platypezina, 64
diversiloba, Rhus, 596, 597, 598, 606

INDEX 691

Dodecatheon hendersonii cruciatum, 643
doederleini, Centrocidaris, 236
domingensis, Typha, 666
donelliana, Dichondra, 620
Dorothia, 536
oxycona, 536, 538, 542
douglasiana, Artemesia, 612
Tris, 598, 601, 660
douglassii douglassii,
Micromeria, 632
Microseris douglasii, 618
Satureja, 601, 632
tenella, Microseris, 618
draba, Cardaria, 622
Drassylus covensis, 79, 80, 83, 84
dentelifer, 84
drymarioides, Pterostegia, 642, 653
Dryopteris arguta, 599, 605
dubia, Podoceropsis, 245
dubium, Trifolium, 635
dubius, Philautus, 137
Dudleya, 621
cymosa, 597, 601, 621
farinosa, 597, 601, 621
dunckeri, Syngnathus, 364, 366, 375
Durham, J. Wyatt, Corals from the Gala-
pagos and Cocos Islands, 41-56
duvergieri, Otolithus (Congeris) , 92
Dycotyledoneae, 603, 605
dymka, Clythia, 64

618

Microseris,

Echeneis remora, 245
Echeveria farinosa, 386
Echinodermata, 234
Echinoidea, 235
Echinometra van-brunti, 236
Echinopsilon hyssopifolium, 610
echinus, Rachodrassus, 79, 80, 83
echioides, Picris, 618
Eciton (Labidus) crassicornis, 251
ecuadoriana, Solenastrea, 45
edule, Mesembryanthemum, 605
edwardsii, Petrolisthes, 248
effusus brunneus, Juncus, 661
egedii grandis, Potentilla, 601, 646
Eggerella, 537, 538
advena, 230
egregia, Silis (Silis), 462, 477, 481
eichenbergi, Lenticulina, 554
Lenticulina (Lenticulina) , 554:

692 CALIFORNIA ACADEMY OF SCIENCES

Elachistodon westermanni, 131, 141
Elaphe cantoris, 113, 140
helena, 140
hodgsoni, 127, 129, 130
hodgsonii, 113, 140
porphyracea, 140
prasina, 140
radiata, 113, 124, 140
taeniura, 140
Elapidae, 116, 118, 141, 142
elata, Habenaria unalascensis, 666
elegans, Cycloseris, 45, 46, 52
Pocillopora, 42, 44, 45, 46, 48, 231
Senecio, 618
elegantia, Nodosaria, 541, 562
elegantissima, Anthopleura, 515
elliptica, Garrya, 601, 628
Ellobium stagnalis, 239
elongata, Hyperammina, 539
Solidago canadensis, 619
Testudo, 110, 132, 138
elucens, Syngnathus, 364, 366, 374
Elymus glaucus, 658
triticoides, 658
emarginata, Prunus, 600, 601, 647
emarginata, Silis (Silis), 455, 463, 485
emmae, Azteca, 250
Empimorpha, 58, 61
geneatis, 62
Empis, 58, 61
Emydidae, 109, 137, 138
Enantiosula manica, 233
Encope cocosi, 236
micropora cocosi, 236
Endopachys vaughani, 44, 46, 231
Enhydris enhydris, 140
enhydris, Enhydris, 140
Enhydris sieboldi, 140
ensenadae, Corythoichthys, 372
Micrognathus, 372, 373
Micrognathus vittatus, 373
entypa, Chloeia, 244
Eopsetta jordani, 315
Epilobium adenocaulon, 639
franciscanum, 639
paniculatum, 639
watsonii, 639
Epinannolene pittieri, 249
Epistomina, 569
caracolla caracolla, 569, 572

Eponididae, 567
equatorialis, Astrangia, 44, 45, 46, 52
Equisetaceae, 604
Equisetum arvense, 604
telmateia braunii, 604
eragrostis, Cyperus, 656
Erechtites arguta, 614.
prenanthoides, 614:
erecta, Plantago, 596, 64.0
Tillaea, 621
erectus, Hippocampus, 367
Ergates spiculatus, 170
erianthus erianthus, Orthocarpus, 651
Orthocarpus, 598
Orthocarpus erianthus, 651
roseus, Orthocarpus, 651
Ericaceae, 625
Epistomininae, 569
Eponides, 567
ericoides, Haplopappus, 616
Erigeron foliosus, 614
glaucus, 597, 601, 614
philadelphicus, 601, 614:
Eriodictyon californicum, 630
Eriogonum latifolium 596, 597, 642
nudum, 642
vimineum, 642
Eriophyllum confertiflorum, 615
staechadifolium, 386, 597, 615
Eriphia granulosa, 246
squamata, 246
Erodium botrys, 629
cicutarium, 629
moschatum, 629
obtusiplicatum, 629
erosus, Thyrolambus, 246
Erysimum franciscanum, 601, 623
erythrogaster, Hirundo rustica, 253
Oligodon, 115, 118, 125, 141
erythrurus, Trimeresurus, 142
Eryx conicus, 139
Eschscholzia californica, 601, 64.0, 650
Eucalyptus globulus, 595, 597, 637
Eucidaris thouarsli, 236
euglochis, Chloeia, 244
Eumetopias, 378, 380
jubata, 377, 378, 390
Euphorbia crenulata, 627
lathyris, 627
peplus, 627

[Proc. 4TH SER.

Vor. XXXIi]

Euphorbiaceae, 627
Euponera (Trachymesopus) stigma, 251
Euprognatha bifida, 245
granulata, 245, 246
(Eurhinocricus) biolleyi, Rhinocricus, 249
europaea, Salicornia, 610
europaeus, Ulex, 600, 636
Eusigalion spinosum, 244
Eutermes ripperti, 250
Evodinus, 184
(Centrodera) spurcus, 160
spurcus, 160
exarata, Agrostis, 657
excavata, Porites, 42, 45, 46, 51, 232
excentricitas, Lagena striatopunctata, 230
excentricus, Reophax, 230
exigua, Madia, 617
exilis, Aster, 612
Vaginulina, 230
eximia, Silis (Silis), 462, 477, 481
Exline, Harriet, Two Gnaphosid Spiders
from Arkansas, 79-85
expansa, Argonauta, 243
Teteragonia, 606
explanulata, Pavona, 44, 46
Pavona (Pavona), 231
explorator, Calcinus, 248
exquisitus, Pecten (Cyclopecten), 238

fabaceus, Marah, 624

fabulosa, Silis (Silis), 464, 497, 498

Fagaceae, 627

Fagus grandifolia, 192

Fahy, Neil, see McClintock,

fallax, Conger, 99

farinosa, Dudleya, 597, 601, 621
Echeveria, 386

fasciatus, Bungarus, 141
Lycodon, 140

fasciolatus, Coluber, 113, 140
Cyrtodactylus, 138

faucibarbatus albidus, Orthocarpus, 651

Favia, 45

fenderi, Silis, 487

fenderi, Silis (Silis), 463, 485, 486

fenestrata, Silis (Silis), 455, 464, 489, 495

ferruginea, Trapezia cymodoce, 247

Elizabeth

ferrugineus, Coccyzus, 253
Nesococcyx, 253

INDEX | 693

Festuca, 603
californica, 658
dertonensis, 658
idahoensis, 658
megalura, 658
myuros, 658
rubra, 658
Filago californica, 615
gallica, 615
Filicinae, 603, 604
filicornis, Silis (Silis), 455, 460, 467, 471
filiformis, Dentalina, 230
filigera, Silis, 470
Silis (Silis) , 461, 467, 470
filix-femina, Athyrium, 604.
filum, Aplousina, 232
fimbria, Anoplopoma, 316
fimbriata, Woodwardia, 599, 605
Fissurella virescens, 239
fistulatus, Syngnathus, 364
Flabellum daphnense, 44, 46
flavescens, Varanus, 112, 139
flavida, Silis, 472, 473, 475, 477
Silis (Silis), 461, 467, 472
flaviviridis, Hemidactylus, 139
flavus, Rachodrassus, 83
flexuosa, Tubulipora, 233
Flora of the San Bruno Mountains, San
Mateo County, California, by Eliza-
beth McClintock and Walter Knight,
In Collaboration with Neil Fahy,
587-677
floribundum, Centaurium, 628
floribundus, Orthocarpus, 651
floridae floridae, Syngnathus, 364, 367, 369
hubbsi, Syngnathus, 637, 368, 369
mckayi, Syngnathus, 364, 367, 370
nesiotes, Syngnathus, 364, 368
Siphostoma, 369
Syngnathus, 367, 368, 369
Foeniculum vulgare, 654
foliosus, Erigeron, 614
folium, Leptodesmus, 249

Follett, W. I. and Lillian J. Dempster,
Relationships of the Percoid fish
Pentaceros richardsoni Smith, with

description of a specimen from the
coast of California, 315-338

folletti, Syngnathus, 364

Foraminifera, 229

694 CALIFORNIA ACADEMY OF SCIENCES

Foraminifera of the Orchard Peak-Devils
Den Area, California, by Clifford C.
Church, 523-580

Foraminiferida, 532

formosa, Dicentra, 599, 601, 628

Rana, 108, 137
truncata, Aquilegia, 643

formosus, Lupinus, 634

fornicatus, Conger, 92, 95, 96, 97

forsteni, Boiga, 140

fossiger, Silis, 453

Silis (Ditemnus), 451, 453, 455
Fragaria californica, 598, 601, 64.5, 646
chiloensis, 598, 601, 645, 646

franciscana, Castilleja, 598, 650

collinsia, 650

Monardella villosa, 597, 600

Paronychia, 597, 609
franciscanum, Epilobium, 639

Erysionum, 601, 623

franciscanus, Rubus spectabilis, 64.8

frankei, Frondicularia, 550, 557, 564

Frankenia grandifolia, 628

Frankeniaceae, 628

frankliniana, Rooseveltia, 254

Franseria bipinnatifida, 386

chamissonis bipinnatisecta, 615

Fraxinus, 407

pennsylvanica, 407
freemani, Silis, 453
Silis (Ditemnus), 451, 4.53, 455
Freesia refracta, 660
fremontii minor, Zigadenus, 665
Populus, 406

frenatus, Hemidactylus, 111, 139

Fritillaria lanceolata, 663, 664

fritzi, Talorchestia, 245

Frizzell, Don L., and C Kurt Lamber, Dis-
tinctive “Congrid Type” Fish Otoliths
from the Lower Tertiary of the Gulf
Coast
101

Frondicularia, 557, 560

concinna, 559
frankei, 550, 557, 564

fronticincta, Ahaetulla, 139

frutescens, Quercus wislizeni, 193, 628

fulgurans bernhardi, Nerita, 240

fullonum, Dipsacus, 625

fultoni, Mitra, 240

(Pisces: Anguilliformis), 87—

[Proc. 47H SER.

Fumariaceae, 628
funiculata, Nerita, 240
furcata, Porites, 373
furcatum, Solanum, 652
furcillatus, Lepidonotus, 244
fusca, Sthenelais, 244
fuscum, Siphostoma, 369
Trachischium, 115, 141
fuscus affinis, Syngnathus, 363, 370
fuscus, Syngnathus, 363
Syngnathus, 369, 370

gaillardioides, Layia, 617
galapageia galapageia, Polygala, 582, 583
584, 585
insularis, Polygala, 582, 584, 585, 586
Polygala, 583, 584, 585
Polygala galapageia, 582, 583, 585
galapagense, Desmophyllum, 44, 46
galapagensis, Balanophyllia, 44, 46
Madrepora, 44, 46
Uca, 247, 248
Galium aparine, 648
californicum, 648
nuttallii, 648
galleottiana, Plantago hirtella, 64.1
gallica, Filago, 615
Silene, 609
gallina multifilosa, Tegula, 241
gallinum multifilosum, Chlorostoma, 241
gammei, Dinodon, 140
Rana, 137
gangeticus, Gavialis, 109, 138
Trionyx, 110, 138
gardnerensis, Astrangia, 44, 45, 46, 52
garnoti, Hemidactylus, 139
Garrya elliptica, 601, 628
Garryaceae, 628
Gastropoda, 238
Gaudryinella almgreni, 538, 540
Gaultheria shallon, 598, 626
Gavelinella, 566
barremiana, 566, 572
Gavialis gangeticus, 109, 138
G Dallas Hanna, by Robert C. Miller, 1-
40
Gecarcinidae, 247
Gecko gecko, 139
gecko, Gecko, 139
Gekkonidae, 111, 138, 139

Vor. XXXII]

geminata, Solenopsis, 251
gemmatus, Cantharus, 238
Cantharus (Gemmophos), 238
(Gemmophos) gemmatus, Cantharus, 238
geneatis, Empimorpha, 62
geniculata, Brassica, 622
geniculatum, Hordeum, 658
Gentianaceae, 628
Geoclemys hamiltoni, 137
Geoemyda tricarinata, 132, 137
Geoemyda trijuga, 132, 137
Geraniaceae, 629
Geranium, 629
carolinianum, 629
dissectum, 629
molle, 629
retrorsum, 629
gerardi, Belemnopsis, 524
gerbillus, Rana, 137
giardi, Pyroteuthis, 243
gibbus, Pentaceros, 328
gigantea, Pavona, 42, 45, 46
Pavona (Pavona), 50
Vicia, 636
gigas characteristica, Siphonaria, 241, 243
Siphonaria, 241, 243
Gilchristia, 317
Gilia achilleaefolia, 641
clivorum, 641
gracilis, 641
glabella, Polinices, 240
glabra, Arabis, 621
Hypochaeris, 616
glabratus, Ammodiscus, 533, 568, 616
gladiator, Conus, 239
Glandulina, 541, 563
Glandulinidae, 563
Glandulininae, 563
glandulosa, Potentilla, 646, 647
glasselli, Thyrolambrus, 246
glauca, Sambucus, 407
glaucus, Elymus, 658
Erigeron, 597, 601, 614
Globanomalina, 571
hexensis, 568, 571
Globigerina infracretacea, 571
Globigerinacea, 571
globispira, Succinea, 242
globosa, Gyroidina, 566, 572
Neritina, latissima, 240

INDEX 695

Globotextulariinae, 536
Globulina, 565

ampulla, 565, 572

prisca, 565
globulus, Eucalyptus, 595, 597, 637
Gloiopotes ornatus, 245
glomeratus, Senecio, 614
Glomospira, 534

gordialis, 534, 538
glutinosis, Ichthyophis, 136
Glyptocephalus zachirus, 315
Gnaphalium, 603

beneolens, 615

californicum, 615

chilense, 615

luteo-album, 615

purpureum, 615

ramosissimum, 615
Gnaphosidae, 80
gokool, Boiga, 140
Gonothyraea, gracilis, 230
roodallii, Chiton, 243

Chiton (Chiton), 243

Chiton (Radsia), 243
gooddingil, Salix, 406
goodei, Sebastodes, 316
Goose Lake Fragments, The, by C. P. But-

ler, 291-313

gordialis, Glomospira, 534
gorei, Dendrelaphis, 140
gracile, Opeas, 242
eracilentum, Trifolium, 635
gracilis, Gilia, 641

Gonothyraea, 230

Koelaria, 659

Madia, 617

Microsteris, 641

Narcissia, 234

Ophisaurus, 119, 138

Panulirus, 249

Phlox, 641

Reteporellina denticulata, 233
gracillissima, Marginulinopsis, 551, 564
grahami, Dentalina, 545, 550
Gramineae, 603, 656
gramineus, Trimeresurus, 118
grande, Cynoglossum, 607
grandiflora, Agoseris, 611

Heterotheca, 616

Tellima, 650

696 CALIFORNIA ACADEMY OF SCIENCES

grandifolia, Fagus, 192
Frankenia, 628
grandis, Meoma, 236
Potentilla egedii, 601, 646
Silene scouleri, 609
granulata, Euprognatha, 246
granulosa, Eriphia, 246
Grapsidae, 247
Grapsus grapsus, 247
grapsus, Grapsus, 247
grata, Astacolus, 555, 558
grayana, Amalthea, 239
Hipponix, 239
Green, John Wagener, Revision of the
Nearctic Species of Silis (Canthar-
idae: Coleoptera), 447-513
Griffineta nelsonensis, 318
Grindelia camporum, 616
hirsutula, 616
humilis, 616
maritima, 616
Grossularia californica, 600, 649
divaricata, 649
leptosma, 649
guatemalensis cocoénsis, Prenolepis, 251
cocoénsis, Prenolepis (Nylanderia), 251
gubernatoris, Cyrtodactylus, 139
Gubkinella, 573
asiatica, 573
californica, 568, 573
Guembelitriinae, 573
guentheri, Trachischium, 115, 127, 141
guineénse, Tetramorium, 251
Gumbelitria harrisi, 573
gurneyi, Ptilosarcus, 231
guttatus, Mimulus, 651
Guttulina, 563, 568
guyonianum, Limoniastrum, 192
gymnocarpa, Rosa, 600, 601, 64.7
Gymnospermae, 603, 605
Gyroidina, 533, 566
globosa, 566, 572

Habenaria unalascensis elata, 666
unalascensis maritima, 665
unalascensis unalascensis, 666

Habroconus, 242
(Cocosconus) hopkinsi, 242
(Cocoslens) pallidus, 242
(Pseudoguppya) pacificus, 242

[Proc. 4TH SER.

haemastoma biserialis, Thais, 241
haematodes insularis, Odontomachus, 251
Odontomachus, 251
Haeuslarella, 540
Halecium washingtoni, 230
halepense, Sorghum, 660
haliaétus, Pandion 253
hallii, Agrostis, 657
Haloragidaceae, 630
hamata, Cristellaria, 551
Hamilton, Marian E., see Hamilton, Wil-
liam J., III
Hamilton, William J., III, and Marian E.
Hamilton, Breeding Characteristics of
Yellow-Billed Cuckoos in Arizona, by,
405-432
hamiltoni, Geoclemys, 137
hancocki, Haplophragmoides, 230
Macrobranchium, 248
Pecten (Oppenheimopecten), 238
Sphenotrochus, 44, 46
Hanna, G Dallas, by Robert C. Miller, 1-40
hannah, Ophiophagus, 142
Hantkeninidae, 571
Haplopappus ericoides, 616
Haplophragmium, 536
aequale, 536, 542
Haplophragmoides, 536, 538
hancocki, 230
Haplophragmoidinae, 535
Hardella thurjii, 137
Harpa crenata, 239
hartwegii, Chorizanthe pungens, 597, 64.1
harrisi, Giimbelitria, 573
hastata, Atriplex, 610
Atriplex patula, 610
heathiana, Stenoplax, 434, 439, 441
Hedbergella, 571
planispira, 568, 571
Hedbergellinae, 571
Hedera helix, 606
helena, Elaphe, 140
Helenium puberulum, 616
Helianthella castanea, 616, 620
Helicocranchia beebei, 243
helicoides, Polinices, 240
helix, Hedera, 606
helleri, Perinereis, 244
Helxine soleirolii, 655
Hemiarthrum setulosum, 435, 436, 442

Vor. XXXII]

Hemidactylus bowringi, 139
brooki, 139
flaviviridis, 139
frenatus, 111, 139
garnoti, 139
Hemithyris craneana, 234
Hemizonia corymbosa, 616
pungens, 616
hemphilli, Podochela, 246
hendecacanthus, Pentaceros, 326, 327, 331
hendersonii, Angelica, 599, 652
cruciatum, Dodecatheon, 643
Heracleum lanatum, 654
maximum, 599, 653, 654.
Herald, Earl S., Studies on the Atlantic
American Pipefishes with Descriptions
of New Species, 363-375
Herbstia tumida, 245
Herpetology of Nepal: A History, Check
List, and Zoogeographical Analysis of
the Herpetofauna, The, by Lawrence
W. Swan and Alan E. Leviton, 103-
147
Herse cingulata, 250
Hertlein, Leo George, Contribution to the
Biogeography of Cocos Island, In-
cluding a Bibliography, 219-289
Hesperocidaris panamensis, 236
Hesperocnide tenella, 655
Heterenchelys, 92
Heterohelicidae, 573
Heteromeles arbutifolia, 600, 601,
heterophylla, Collinsia, 650
Nemophila, 630
Stylomecon, 640
Heterotheca grandiflora, 616
Heterozona subviridis, 436
Heuchera micrantha, 601, 649
hexensis, Globanomalina, 568, 571
Nodosaria, 544, 564
hieracioides, Layia, 617
Hilara, 58, 61
hildebrandi, Syngnathus, 364, 366
himalayana, Agama, 138
Natrix, 113, 141
himalayanum, Leiolopisma, 111, 127,
139
himalayanus, Agkistrodon, 116, 130, 142
Bufo, 107, 136
Hippa denticulatifrons, 248

646

130,

INDEX 697

Hippidae, 248
Hippocampus erectus, 367
hudsonius, 367
Hippocastanaceae, 630
Hippocrepininae, 539
Hipponix barbatus, 239
grayanus, 239
pilosus, 239
hirsuta, Centrodera, 178
Spio, 244
hirsutula, Baeria chrysostoma, 613
Grindelia, 616
hirsutus, Nerine cirratulus, 244
hirtella galleottiana, Plantago, 64.1
hirtimanus, Pylopagurus, 248
hirudiniformis, Acanthochitona, 242
Hirundo rustica erythrogaster, 253
Hispanirhynchia ?craneana, 234
hispida, Medicago, 635
hispidissima, Cryptantha, 607
hispidula, Lonicera, 608
Histiopterus richardsoni, 317
Historis odius, 250
Histrio odius, 250
orion, 250
hodgsoni, Elaphe, 127, 129, 130
hodgsonii, Elaphe, 113, 140
holciformis, Deschampsia caespitosa, 658
Holcus lanatus, 658
Holodiscus discolor, 600, 601, 646
Holoporella brunnea, 233
holosericea, Urtica, 655
Holothurioidea, 236
hondaensis, Astrangia, 44, 46, 231
Kionotrochus (?), 44, 46
hookeri, Disporum, 663
Oenothera, 639
hookeriana californica, Plantago, 640
hopkinsi, Habroconus (Cocosconus), 242
Hordeum brachyantherum, 658
geniculatum, 658
hystrix, 658
leporinum, 658
Horkelia californica, 646, 647
howdeni, Silis (Ditemnus), 451, 452,
456
Howell, John Thomas, and Duncan M.
Porter, The Plant Genus Polygala in
the Galapagos Islands, 581-586
hoylei, Abraliopsis, 243

455,

698 CALIFORNIA ACADEMY OF SCIENCES

hubbsi, Syngnathus floridae, 367, 368, 369
hudsonius, Hippocampus, 367
humilis, Grindelia, 616
Murex, 240
Nodosaria, 541, 568
hunteri, Clythia, 64
Hurriah sanguiniventer, 117
hurum, Trionyx, 138
hyadesi, Nuttallochiton, 434
Hydrobius fuscipes, 156
Hydrophyllaceae, 630
Hydrophilus, 156
Hydrozoa, 230
Hymenoptera, 250
Hyperammina, 539
elongata, 539
Hypericaceae, 631
Hypericum anagalloides, 631
Hypochaeris glabra, 616
radicata, 617
hyssopifolia, Bassia, 610
Lythrum, 637
hyssopifolium, Echinopsilon, 610
hystrix, Hordeum, 658

Ichthyocampus pawneei, 364
Ichthyophis glutinosis, 136
Icteria virens, 407
idahoensis, Festuca, 658
ilicifolia, Prunus, 591, 597, 601, 647
imbricata, Arctostaphylos, 600, 626
imitans, Semperothuria, 237
imitator, Ischnochiton, 434, 436, 442
immaculata, Plagusia, 247
impatiens, Brandtothuria, 236
imperfecta, Melica, 659
Inachoides laevis, 245
inarmata, Micopora coriacea, 233
incana, Bowlesia, 653
incongrua, Silis (Silis), 463, 483, 485
incurva, Parapholis, 659
incurvata, Astacolus, 555
indica, Chitra, 105, 110, 138
indicum, Sphenomorphus, 139
indicus, Melilotus, 635
inflata, Aucella, 525, 527

Buchia, 527

Limacina, 242
infractacea, Globigerina, 571
inhabilis, Jaegerothuria, 237

[Proc. 4TH Ser.

inornata, Cactornis, 253
Pinaroloxias, 253
inscriptus, Achirus, 366
Insecta, 250
insignis, Calotarsa, 60, 61, 64, 74
insolita, Silis (Silis), 467, 503, 507
insperata, Silis (Silis), 467, 503, 509
instructa, Pennella, 245
insularis, Leucotermes, 250
Odontomachus haematodes, 251
Polygala galapageia, 582, 584, 585, 586
integrifolia, Dentaria californica, 622
intermedia, Amsincka, 607
introversa, Silis (Silis), 465, 503
intybus, Cichorium, 613
Involutina, 533, 569
stinemeyeri, 568, 569
involucoata, Lonicera, 608
Involutinidae, 569
Tridaceae, 660
Tris douglasiana, 598, 601, 660
longipetala, 598, 601, 660
isabella, Cypraea, 239
Cypraea (Luria), 239
mexicana, Cypraea, 239
Ischnochiton, 441
cinereus, 433, 434, 441, 442
constanti, 436, 442
imitator, 434, 436, 442
magdalenensis, 434
subviridis, 442
Isognomon (Melina) chemnitzianum, 237
quadrangularis, 238
italica, Saracenaria, 552, 558
Ithaginis, 127

Jaegerothuria inhabilis, 237
jalapa, Mirabilis, 638
jamaicense, Macrobranchium, 248
januarii, Polypus, 243
Japalura, 127
kumaonensis, 138
major, 111, 127, 131, 138
tricarinata, LIT 1275131 188
variegata, 138
japonica, Lonicera, 608
japonicus, Pentaceros, 321, 326, 327, 331
Quinquarius (Pentaceros), 319
Zalophus californianus, 379
jara, Lycodon, 117, 140

Vout. XXXII]

Jaumea carnosa, 600, 617
jerdoni, Ophisops, 139
Rhacophorus, 137
Typhlops, 142
jordani, Adiantum, 604.
Eopsetta, 315
jubata, Eumetopias, 377, 378, 390
jubatum, Sitanion, 659
juglandifer, Oligodon, 141
Juglans, 407
cinerea, 192
major, 407
jugosa, Dentalina, 230
juliflora, Prosopis, 407
Juncaceae, 661
Juncaginaceae, 661
Juncus, 603
balticus, 661
bufonius, 661
effusus brunneus, 661
leseurli, 661
mexicanus, 661
occidentalis, 661
phaeocephalus, 661
tenuis congestus, 661
xiphioides, 661

kaber, Brassica, 622
Kachuga dhongoka, 109, 137
kachuga, 109, 138
kachuga, Kachuga, 138
Kachuga smithi, 138
tecta, 138
kali tenuifolia, Salsola, 610
Kalotermes (Neotermes) larseni, 250
kamarowl, Apatophysis, 155
kaouthia, Naja naja, 118
Katharina tunicata, 434
Kathetostoma cubana, 366
kefersteini, Ludwigothuria, 237
kellogii, Quercus, 628
kernensis, Citharina, 556, 557, 558
Kessel, Berta B., see Kessel, Edward L.
Kessel, Edward L., and Berta B. Kessel,
Mating Behavior and Activity-Rest
in Protoclythia califor-
nica (Diptera: Platypezidae), 57-78
khasiensis, Cyrtodactylus, 139
Kionotrochus (?)avis, 44, 46
(?) hoodensis, 44, 46

Periodicity

INDEX 699

Knight, Walter, see McClintock, Elizabeth
knulli, Silis (Ditemnus), 452, 455, 458
kochii, Vaginulina, 547, 548, 558
Koelaria cristata, 659
gracilis, 659
macrantha, 659
Koniga maritima, 623
kugleri, Lenticulina, 553
kumaonensis, Japalura, 138

Labiatae, 603, 631
(Labidus) crassicornis, Eciton, 251
labiosa, Pyrene, 241
labrosa, Columbella, 241
laciniata, Sanicula, 655
Lactuca saligna, 617
serriola, 617
virosa, 617
ladacensis, Leiolopisma, 111, 129, 139
laeve, Trachischium, 141
laevigatum, Taraxacum, 619
laevigatus, Symphoricarpos albus, 608
laevis, Inachoides, 245
Lagena striatopunctata excentricitas, 230
Lagophylla ramosissima, 617
Lamber, C Kurt, see Frizzell, Don L.
Lampyridae, 448
lanatum, Heracleum, 654
lanatus, Holcus, 658
lanceolata, Fritillaria, 663, 664
Plantago, 64.1
Prunella vulgaris, 631
larseni, Kalotermes (Neotermes), 250
Larval Development of Chitons (Amphi-
neura), The, by Allyn G. Smith, 433-
446
lasiolepis, Salix, 600, 649, 656
lasiophyllum, Thelypodium, 624.
lasseni, Silis, 472, 473
lasseni, Silis (Silis), 461, 467, 472
latestyla, Silis (Silis), 464, 497
lathyris, Euphorbia, 627
Lathyrus odoratus, 633
vestitus, 633, 636
latifolia, Myosotis, 607
Polygala, 586
Typha, 666
latifolium, Eriogonum, 596, 597, 642
latifolius, Orthocarpus purpurascens, 651

700 CALIFORNIA ACADEMY OF SCIENCES

latiloba, Silis, 452
Silis (Ditemnus), 451, 452, 455
Latirus tuberculatus, 239
latissima globosa, Neritina, 240
Lauraceae, 632
Laurus nobilis, 632
Lavatera arborea, 637
assurgentiflora, 637
cretica, 637
lawderanus, Cyrtodactylus, 139
Layia chrysanthemoides, 617
caillardioides, 617
hieracioides, 617
platyglossa campestris, 617
laxa, Brodiaea, 596, 601, 663
lecontei, Silis (Silis), 464, 497, 499
ledebourii, Lonicera, 608
Leech, Hugh B., Centrodera spurca (Le-
Conte) and Two New Species Re-
sembling it, with Biological and Other
Notes (Coleoptera: Cerambycidae),
149-218
Leguminosae, 603, 632
Leiolopisma himalayanum, I11, 127, 130,
139
ladacensis, 111, 129, 139
sikkimense, 111, 132, 139
Leioptilus undulatus, 231
leithi, Psammophis, 141
Trionyx, 138
Lemna minima, 661
Lemnaceae, 661
Jens, Mitra, 240
Lenticulina, 552, 553, 555, 558
eichenbergi, 554
(Lenticulina) eichenbergi, Lenticulina, 554
Lenticulina kugleri, 553
(Lenticulina) eichenbergi, 554.
(Lenticulina) miinsteri, 552, 562
(Lenticulina)
552, 562

Lenticulina ouachensis,

munsteri, Lenticulina,

ro)

saxonica saxonica, 552, 562

valanginiana, 551
lenzi, Popilius, 250
leopolitana, Pseudopolymorphina, 565
Lepechinia calycina, 599, 600, 631
lepida, Silis, 452

Stipa, 660

[Proc. 47TH Serr.

Lepidium nitidum, 623
pinnatifidum, 623
Lepidonotus furcillatus, 244
Lepidopleurus asellus, 434, 438, 441
Lepidozona asthenes, 438, 439, 441, 442
cooperi, 434
mertensi, 434
leporinum, Hordeum, 658
Leptinaria, 242
biolleyi, 242
Leptodesmus folium, 249
Leptodius cooksoni, 247
Leptonotus blainvilleanus, 364
Leptorhabdium, 156
Leptoseris digitata, 44, 46, 231
panamensis, 45
leptosma, Grossularia, 64.9
leptosmum, Ribes menziesii, 649
Leptura serricornis, 155
leseurii, Juncus, 661
Lessonothuria pardalis, 237
Leucomelina pica, 224, 250
Leucotermes insularis, 250
Leviton, Alan E., see Swan, Lawrence W.
liebigii, Rana, 108, 130, 131, 137
Ligusticum apiifolium, 654
Liliaceae, 603, 661
Limacina inflata, 242
limnocharis, Rana, 108, 137
Limoniastrum guyonianum, 192
Limonium californicum, 600, 641
Linaceae, 636
Linanthus androsaceus, 64.1
Linaria canadensis, 651
texana, 651
Linckia columbiae, 234
linearis, Meconella, 64.0
lineata, Sigsbeia, 235
lineatus, Oostethus, 363
lineolatus, Sphaerodactylus, 252
Lingulina californiensis, 560, 564
tenera, 550, 560
Lingulininae, 560
Linum usitatissimum, 636
Liocranchia reinhardti, 243
Liomera cocosana, 246
Liopeltis calamaria, 140
rappii, 113, 140
stoliczkae, 140

- im meas

Voi. XXXII]

Liriodendron tulipifera, 191
Lissa aurivilliusi, 245
tuberosa, 245
Lissemys punctata, 138
Lithocarpus densiflorus, 204
Lithophaga attenuata, 238
(Myoforceps) aristata, 238
Lithophragma affinis, 649
Littorina aspersa, 239
conspersa, 239
modesta, 239
Lituolacea, 533
Lituolidae, 535
Lituolinae, 535
livida, Rana, 137
lividus, Bungarus, 131, 141
lobata, Porites, 42, 45, 46, 51, 54
Silis (Silis), 465, 503, 505
lobatus, Xanthodius, 247
Lobularia maritima, 623
Lolium multiflorum, 659
Lomatium, 599
caruifolium, 654.
dasycarpum, 654.
utriculatum, 654:
longicornis, Brachymyrmex, 250
longimanus, Pylopagurus, 248
longipetala, Iris, 598, 601, 660
longirostris constricta, Cavolina, 242
Lonicera hispidula 608
involucrata, 608
japonica, 608
ledebourii, 608
lophantha, Albizia, 633
Lophophorus, 127
Lophosmilia wellsi, 44, 46
Lotus, 603
corniculatus, 633
micranthus, 633
purshianus, 634.
scoparius, 634.
subpinnatus, 634
louisianae, Siphostoma, 369
Syngnathus, 364, 369, 370, 371
Lovenia cordiformis, 236
Lower Cretaceous Foraminifera of the Or-
chard Peak-Devils Den Area, Cali-
fornia, by Clifford C. Church, 523-
580
lucifuga, Agathomyia, 64

INDEX 701

Ludwigothuria atra, 237
kefersteini, 237
Lunaria annua, 623
Lupinus, 603
affinis, 634.
albifrons, 634.
arboreus, 386, 596, 634
bicolor, 596, 634
chamissonis, 597, 634.
formosus, 634.
micranthus, 634
nanus, 596, 600, 634.
ramosissimum, 597
succulentus, 635
varlicolor, 635
(Luria) isabella, Cypraea, 239
lutea, Silis, 470, 471
Silis (Silis), 455, 460, 461, 467, 470
luteo-album, Gnaphalium, 615
Luzula multiflora, 661
subsessilis, 661
Lycidae, 448
lycioides, Condalia, 407
Lycodon, 117
aulicus, 113, 140
fasciatus, 140
jara, 117, 140
mackinnoni, 140
striatus, 140
Lytechinus pictus, 236
Lythraceae, 637
Lythrum hyssopifolia, 637
Lytoceras, 525
saturnale, 525, 526, 527, 531

Mabuya carinata, 111, 139
macularia, 112, 139
multifasciata, 139

macelayi, Silis (Silis), 463, 485, 489

macclellandii, Calliophis, 116, 142

mackinnoni, Lycodon, 140

macrantha, Koeleria, 659

Macrobranchium americanum, 248
hancocki, 248
jamaicense, 248
olfersii, 249

macrocarpa, Cupressus, 597, 605

macrodactyla, Uca, 248

macrophylla, Arenaria, 608

macropoda, Brodiaea coronaria, 663

702 CALIFORNIA ACADEMY OF SCIENCES

macrops, Pseudoxenodon, 115, 123, 141
macrotheca, Spergularia, 386, 609
macularia, Mabuya, 112, 139
macularius, Pareas, 141
maculatum, Conium, 653
Sphenomorphus, 132, 139
maculatus, Butorides virescens, 253
Rhacophorus, 109, 137
maculostriata, Tegula, 241
maculostriatum, Chlorostoma, 241
maculosum, Cerithium, 239
Madia capitata, 617
exigua, 617
gracilis, 617
sativa, 618
maderensis, Cytisus, 633
Madracis asperula, 44, 46
Madrepora galapagensis, 44, 46
magdalenensis, Ischnochiton, 434
Mahonia pinnata, 607
Maianthemum dilatatum, 599, 601, 663
Majidae, 245, 246
major, Japalura, 111, 127, 131, 138
Juglans, 407
Megophrys, 136
Vinca, 606
majus, Tropaeolum, 652
Malacostraca, 245
Malva nicaeensis, 637
parviflora, 637
Malvaceae, 637
malvaceum, Ribes, 599, 600, 601, 649
malvae flora, Sidalcea, 596, 637
Phacelia, 630
mammata, Scutiger, 136
manica, Enantiosula, 233
Marah fabaceus, 624.
oregonus, 624, 625
marchenus, Coronaster, 234
margaritacea, Anaphalis, 596, 597, 601, 611
marginata, Callitriche, 607
marginatus, Bromus, 657
Ophiophragmus, 235
Petrolisthes, 248
marginella, Ditemnus, 451
Marginella minor, 240
Marginulina 548
bullata, 548
curvitura, 548
parkeri, 548, 550

[Proc. 4TH SER.

pyramidalis, 549, 564
robusta, 549, 558
sigali, 54.9, 564
texasensis, 548
Marginulinopsis, 551, 558
collinsi, 551
gracillissima, 551, 564
marianum, Silybum, 619
marina, Spergularia, 609
maritima, Cakile, 386, 622
Grindelia, 616
Habenaria unalascensis, 665
Koniga, 623
Lobularia, 623
muilla, 663
Silis (Silis), 467, 511
maritimum, Alyssum, 623
marmoratus, Chiton, 434
Marrubium vulgare, 631
marsupiata, Microporella, 233
Mating Behavior and Activity-Rest Perio-
dicity in Protoclythia californica (Dip-
tera: Platypezidae), by Edward L.
Kessel, and Berta B. Kessel, 57-78
Matricaria matricarioides, 618
matricarioides, Matricaria, 618
maxima, Briza, 657
maximum, Heracleum, 599, 653, 654:
maximus, Rhacophorus, 109, 125, 137
McClintock, Elizabeth, and Walter Knight,
In Collaboration with Neil Fahy,
A Flora of the San Bruno Mountains,
San Mateo County, California, 587—
677
mckayi, Siphostoma, 370
Syngnathus, 367
Syngnathus floridae, 364, 367, 370
meandrina, Pocillopora, 42, 45, 46, 48, 232
Meconella californica, 640
linearis, 640
media, Stellaria, 610
Medicago hispida, 635
polymorpha vulgaris, 635
sativa, 635
megalura, Festuca, 658
Megophrys major, 136
Melampus panamensis, 240
tabogensis, 240
trilineatus, 240
melaneus, Oligodon, 141

Vor. XXXII]

melanostictus, Bufo, 107, 136
melanostoma, Otocryptops, 249
Melanoteuthis beebei, 243
melanoxylon, Acacia, 633
Melica californica, 659
imperfecta, 659
torreyana, 659
Melilotus albus, 635
indicus, 635
officinalis, 635
Melina chemnitziana, 237
(Melina) chemnitzianum, Isognomon, 237
melitensis, Centareua, 613
melo, Purpura, 241
melones, Purpura, 241
Thais (Vasula), 241
Menkenina, 526, 560
berryi, 550, 560, 561
Menkeninae, 560
menziesii, Arbutus, 195, 202, 601, 625
Calandrina ciliata, 643
leptosmum, Ribes, 649
Nemophila, 630
Pseudotsuga, 201
Meoma grandis, 236
Menthe spicata, 631
meridies, Conger, 92, 95, 96, 97, 98, 99
mertensi, Lepidozona, 434
Mesembryanthemum edule, 605
mesoleuca, Acmaea (Nomaeopelta), 238
Collisella, 238
SSGabhia 258
Metaclythia, 64
mexicana, Centrechinus (Diadema),
Cycloseris, 44, 46, 231
Cypraea controversa, 239
Cypraea isabella, 239
Sambucus, 608
mexicanus, Juncus, 661
Micopora coriacea inarmata, 233
micrantha, Camissonia, 638
Dichondra, 620
Heuchera, 601, 649
Oenothera, 638
microcanthus, Lotus, 633
Lupinus, 634.
microcarpa, Caucalis, 653
microcarpus, Scirpus, 656
microcephalum, Trifolium, 636
microdon, Trifolium, 636

iss)
WwW
UL

INDEX

Micrognathus, 366, 372
(Anarchopterus) crinigerus, 364, 371
(Anarchopterus) crinitus, 364, 371
crinigerus, 371
crinitus, 371, 372
ensenadae, 372, 373
nitidus, 372
vittatus, 364, 372, 373
vittatus-ensenadae, 373

Microhyla ornata, 108, 124, 136

Microhylidae, 108

micromeres, Cryptantha, 607

Micromeria douglasii, 632

Micropanope polita, 247

Microphrys branchialis, 246
triangulatus, 246

microphylla, Morus, 407

micropora cocosi, Encope, 236

Microporella ciliata, 233
marsupiata, 233

Micropus californicus, 618

Microseris douglasii douglasii, 618
douglasii tenella, 618
gracilis, 641

Microstomus pacificus, 315

Microthele difficilis, 237

703

(Middendorffia) cinerea, Nuttallina, 434, 441

mikanioides, Senecio, 618
millefolium, Achillea, 596
californica, Achillea, 611
milleporosa, Tetraclita squamosa, 245
Miller, Robert C., G Dallas Hanna, 1-40
Mimulus guttatus, 651
mindii, Pseudophallus, 364
minima, Lemna, 661
minimus, Senecio, 614
minor, Briza, 657
Marginella, 240
Zigadenus fremontii, 665
minutus, Planes, 247
Mirabilis jalapa, 638
mirandus, Notochiton, 434
Mirounga angustirostris, 377, 378
(Mithraculus) denticulatus, Mithrax, 246
Mithrax (Mithraculus) denticulatus, 246
(Mithrax) spinipes, 246
(Mithrax) spinipes, Mithrax, 246
Mitra fultoni, 240
lens, 240
tristis, 240

704 CALIFORNIA ACADEMY OF SCIENCES

Mitrella ocellata, 240
modesta, Littorina, 239
modica, Apatophysis, 155
molle, Geranium, 629

Schinus, 606
mollis, Bromus, 657

Symphoricarpos, 608
molurus, Python, 112, 139
Mollusca, 237
Monardella villosa, 631

villosa franciscana, 597, 600
moneta, Cypraea, 239

Cypraea (Monetaria), 239
(Monetaria) moneta, Cypraea, 239
monitor, Varanus, 112, 139
Monoceros brevidentata, 238

brevidentatum, 238
Monocotyledoneae, 603, 656
monogyna, Crataegus, 645
monspeliensis, Polypogon, 659
monspessulanus, Cytisus, 633
montana, Apatophysis, 155
montanica, Silis (Silis), 464, 497, 498
montaraensis, Arctostaphylos, 600, 626
Montia perfoliata, 643
monticola, Pareas, 141

Rana, 108, 137

Trimeresurus, 116, 125, 142
monticolus, Coluber, 117
Montipora, 45
Mopalia ciliata, 434

muscosa, 434
mordax, Rhagium, 207
moreha, Quercus, 628
Morus, 407

microphylla, 407
moschatum, Erodium, 629
mucosus, Ptyas, 115, 123, 141
Muilla, 663

maritima, 663
multifasciata, Boiga, 112, 140

Mabuya, 139
multifida, Roubieva, 610
multifidum, Chenopodium, 610
multifilosa, Tegula gallina, 241
multifilosum, Chlorostoma gallinum, 241
multiflora, Luzula, 661
multiflorum, Lolium, 659
muntuim, Polystichum, 599, 605
miunsterl,

562

Lenticulina 552,

(Lenticulina),

[Proc. 4TH Serr.

murale, Chenopodium, 610
Murex calliati, 366
humilis, 240

muricatus, Ranunculus, 644
mutrayana, Pinus contorta, 204
muscosa, Mopalia, 434
mutabilis, Nodosaria, 543
(Myoforceps) aristata, Lithophaga, 238
Myosotis latifolia, 607
Myrica californica, 599, 601, 637
Myricaceae, 637
Myriophyllum brasiliense, 630
Myriopoda, 249
(Myrmaphaenus)

251
(Myrmobrachys) biolleyi, Camponotus, 250

blandus, Camponotus, 250

(Myrmocamelus) blandus, Camponotus, 250
(Myrmothrix) abdominalis

Camponotus, 250
Myrtaceae, 637
myuros, Festuca, 658

cocosensis, Camponotus,

stercorarius,

naja kaouthia, Naja, 118
Naja naja, 142
naja, Naja, 142
Naja naja kaouthia, 118
naja naja, 118
tripudians, 118
nana, Quercus chrysolepis, 599, 628
nanum, Blennosperma, 613
nanus, Lupinus, 596, 600, 634
Penetopteryx, 364
Narcissia gracilis, 234
nasidens, Odynerus (Pachodynerus), 250
nasturtium-aquaticum, Rorippa, 623
Nasturtium, officinale, 623
nasuta, Ahaetulla, 139
Natrix himalayana, 113, 141
parallela, 114, 123, 141
piscator, 107, 114, 141
platyceps, 114, 130, 141
stolata, 114, 141
subminiata, 117, 141
Navarretia squarrosa, 641
Neanthes obscura, 244
nelsonensis, Griffineta, 318
nemophila, Agathomyia, 64
Nemophila heterophylla, 630
menziesii, 630

Won. xoxox Ll

parviflora, 630
Neotermes casteneus, 250
(Neotermes) larseni, Kalotermes, 250
Nerine cirratulus hirsutus, 244
Nerita bernhardi, 240
fulgurans bernhardi, 240
funiculata, 240
ornata, 240
scabricosta, 240
scabricosta ornata, 240
Neritina latissima globosa, 240
pilsbryi, 240
nesiotes, Syngnathus floridae, 364, 368
(Nesococcyx), Coccyzus, 253
Nesococcyx ferrugineus, 253
Nesopupa, 242
(Cocopupa) cocosensis, 242
Nesotriccus ridgwayl, 253
nevadica, Centrodera, 158, 186
Silis, 461
nevadiea, Silis (Silis), 461, 467, 474
Newportia rogersi, 249
nicaeensis, Malva, 637
nielseni, Octopodoteuthis, 243
niger, Bungaris, 142
nigerrima, Silis (Ditemnus), 452, 455, 458
nigriventris, Argyroepeira, 252
nigromarginatus, Zaocys, 116, 141
nigrovirens, Sypharochiton, 437, 438, 441,
442, 443
nigrum, Solanum, 652
nitens, Anoplodera, 189
Stellaria, 610
nitida, Trochammina, 230
Nitidella sertularium, 240
nitidula, Silis, 459, 462
nitidum, Lepidium, 623
nitidus, Micrognathus, 372
nobilis, Laurus, 632
nodiflorum, Solanum, 652
Nodosaria, 541, 563
amphioxys, 543
elegantia, 541, 562
hexensis, 544, 564
humilis, 541, 568
mutabilis, 543
sceptrum, 543
tenuicosta, 543, 564
(Nomaeopelta) mesoleuca, Acmaea, 238
Nonionidae, 566

INDEX 705

norvegicus, Rattus, 226

Notochiton mirandus, 434

Notopygos cirnita, 244

Nouria polymorphinoides, 230

nuda, Ophionereis, 235

nudum, Eriogonum, 642

nutkaensis, Calamagrostis, 657

nutkana, Rosa, 193

nuttallii, Galium, 648
virgatus, Astragalus, 633

Nuttallina cinerea, 436, 442
(Middendorffia) cinerea, 434, 441
thomasi, 434, 436, 438, 442

Nuttallochiton hyadesi, 434

Nyctaginaceae, 638

(Nylanderia) guatemalensis cocoénsis, Pre-

nolepis, 251

Obelia commissuralis, 230

oblanceolata, Polygala sancti-georgii, 582
586

obnupta, Carex, 656

obovata, Polygala, 584, 585, 586

obscura, Neanthes, 244

Observations on Pleurobranchaea califor-
nica MacFarland, 1966 (Opistho-
branchia, Notaspidea), by Dustin

D. Chivers, 515-521

obstipa, Tamaria, 234
obtusa, Silis (Ditemnus), 451, 455, 456
obtusifolius, Rumex, 642
obtusiplicatum, Erodium, 629
occidens, Silis (Silis) difficilis, 462, 479
occidentalis, Alchemilla, 645

Coccyzus americanus, 405

Convolvulus, 620

Juncus, 661

Oreaster, 234

Sagina, 609
oceanicus, Arrhinotermes, 250
ocellata, Mitrella, 240
ochracea, Boiga, 140

Ostrea, 238
Ochrodermella, 242

biolleyi, 242

cumingiana, 242
ochrus, Clypeaster, 236
octocostata, Vaginulina, 546, 562, 564
octolineatus, Simotes, 117
Octopodoteuthis nielseni, 243

706 CALIFORNIA ACADEMY OF SCIENCES

Octopus (Octopus) pusillus, 243
(Octopus) pusillus, Octopus, 243
Ocypodidae, 247
odius, Historis, 250
Histrio, 250
Odontomachus haematodes, 2
haematodes insularis, 251
odoratus, Lathyrus, 633

Odynerus (Pachodynerus) nasidens, 250

Oenanthe sarmentasa, 654
Oenothera cheiranthifolia, 386
contorta strigulosa, 638
hookeri, 639
micrantha, 638
ovata, 638
officinale, Sisymbrium, 623, 624
officinalis, Melilotus, 635
oleracea, Brassica, 622
Portulaca, 643
oleraceus, Sonchus, 619
offersii, Macrobranchium, 249
Oligodon albocintus, 114, 141
arnensis, 114, 117, 141
cyclurus, 141
erythrogaster, 115, 118, 125, 141
juglandifer, 141
melaneus, 141
oligolepis, Typhlops, 142
oligosperma, Cardamine, 622
(Olivella) cocosensis, Olivella, 240
Olivella (Olivella) cocosensis, 240
Onagraceae, 638
Oostethus lineatus, 363
Opeas gracile, 242
Ophiacantha phragma, 235
Ophiactis savignyi, 235
simplex, 235
Ophiocoma aethiops, 235
alexandri, 235
Ophioderma panamense, 235
variegatum, 235
Ophiomyxa panamensis, 235
Ophionereis dictyota, 235
nuda, 235
Ophiophagus hannah, 142

wat

Ophiophragmus marginatus, 23
paucispinus, 235
Ophisaurus gracilis, 119, 138
Ophisops jerdoni, 139
Ophiurioidea, 234

[Proc. 4TH SER.

(Oppenheimopecten) hancocki, Pecten, 238
Opuntia, 407
Orbitoidacea, 567
orchardensis, Trochammina, 534, 542
Orchidaceae, 665
Oreaster occidentalis, 234
oregonensis, Silis (Silis), 461, 475, 477
oregonus, Marah, 624, 625
orientale, Sisymbrium, 624:
orion, Histrio, 250
orisioides, Thuriaria, 230
ornata, Chrysopelea, 140
Microhyla, 108, 124, 136
Nerita, 240
Nerita scabricosta, 240
ornatus, Gloiopotes, 245
Xylosteus, 156
Orobanchaceae, 639
Orobanche uniflora, 639
Orr, Robert T., and Thomas C. Poulter,
The Pinniped Population of Ano
Nuevo Island, California, 377-404
Orthocarpus, 603
densiflorus, 651
erianthus, 598
erianthus erianthus, 651
erianthus roseus, 651
faucibarbatus albidus, 651
floribundus, 651
purpurascens latifolius, 651
pusillus, 652
orthoceras, Barbarea, 621
Ortholeptura, 151
insignis, 150
valida, 150, 151
Orthomorpha coarctata, 249
ortmanni, Petrolisthes, 248
osburni, Balanophyllia, 44, 45, 46, 52, 53
Osmaronia cerasiformis, 600, 601, 646
Ostrea callichroa, 238
ochracea, 238
palmula, 238
Otocryptops melanostoma, 249
Otolithus, 99
(Brotulidarum), 92
(Congeris) , 87, 92
(Congeria) duvergieri, 92
(Congeris) papointi, 92
(Congridarum), 92
(Platessae), 87

VoL. XXXII]

Otostigmus scabricauda, 249
ouachensis, Lenticulina, 555
oulaniensis, Symplectoteuthis, 243
ovata, Camissonia, 638
Oenothera, 638
Stenocionops, 246
ovatum, Vaccinium, 600, 601, 627
oweniana, Plectis, 254
Oxalidaceae, 639
Oxalis pes-caprae, 639
pilosa, 640
oxycona, Dorothia, 536, 538, 542
Ozius agassizii, 247
tenuidactylos, 247
tenuidactylus, 247
verreauxii, 247

(Pachodynerus) nasidens, Odynerus, 250
Pachygrapsus crassipes, 247
transversus, 247
pachynota, Vaginulinopsis, 548, 558
Pachyta, 161
(Parapachyta) spurca, 160
spurca, 154, 160, 168, 193, 205
spurcata, 169
pacifica, Arctostaphylos, 600, 626
Platypezina, 58, 59, 60, 61, 64, 75
Salicornia, 600, 610
Silene, 609
Trivia, 241
pacificum, Taonidium, 243
pacificus, Ammodiscus, 229
Habroconus (Pseudoguppya), 242
Microstomus, 315
Plagiobrissus, 236
Sphaerodactylus, 252
Paguridae, 248
Pagurus californiensis, 248
Palaemon (Palaemon) ritteri, 249
(Palaemon) ritteri, Palaemon, 249
Palaemonidae, 248
Palinuridae, 249
pallida, Amelanchier, 597, 598, 645
Silis, 493
Silis (Silis), 460, 467, 503, 508
pallidus, Habroconus (Cocoslens) , 242
palmula, Ostrea, 238
paludosa, Polygala, 581
palustris, Crocodilus, 138
panamense, Ophioderma, 235
panamensis, Hesperocidaris, 236

INDEX | 707

Leptoseris, 45

Melampus, 240

Ophiomyxa, 235

Porites, 45

Textularia, 230

Tralia, 240

Uca, 247
Pandion haliaétus, 253
paniculata, Polygala, 581
paniculatum, Epilobium, 639
Panicum capillare, 659
pannosa, Cotoneaster, 645
pansa, Purpura patula, 240
Panulirus gracilis, 249

penicillatus, 249
Papauer somniferum, 640
Papaveraceae, 640
Papilio turnus, 419
papointi, Otolithus (Congeris) , 92
Paradasygius depressus, 246
parallela, Natrix, 114, 123, 141
parallela, Silis (Silis), 467, 503, 506
Parapachyta, 154

spurca, 154, 160, 192
(Parapachyta) spurca, Pachyta, 160
Parapholis incurva, 659
paraprinceps, Theelothuria, 237
pardalis, Lessonthuria, 237
Pareas macularius, 141

monticola, 141
Parellisina curvirostris, 233
parkeri, Altirana, 137

Marginulina, 548, 550
Paronychia franciscana, 597, 609
paronychia, Polygonum, 597, 642
Parophrys vetulus, 315
Parthenope (Platylambrus) exilipes, 246
Parthenopidae, 246
parva, Diacria quadridentata, 242
parviflora, Malva,, 637

Nemophila, 630
parviflorus, Rubus, 600, 601

velutinus, Rubus, 647
Pasimachus punctulatus, 206
patula hastata, Atriplex, 610

pansa, Purpura, 240

Purpura, 240

Thais, 240, 241
paucispinus, Ophiophragmus, 235

Sebastodes, 316

708 CALIFORNIA ACADEMY OF SCIENCES

Pauliella aenigma, 234
Pavona, 42, 49
clivosa, 42, 44, 45, 46
(Pavona) clivosa, Pavona, 49
Pavona explanulata, 44, 46, 231
gigantea, 42, 45, 46
(Pavona) gigantea, Pavona, 50
Pavona (Pavona) clivosa, 49
(Pavona) gigantea, 50
(Pavona) varians, 50, 231
Pavona (Polyastra) ponderosa, 42, 45, 46,
Zoi
(Polyastrea) ponderosa, 50
ponderosa, 54
varians, 42, 45, 46, 54
(Pavona) varians, Pavona, 50, 231
pawneel, Ichthyocampus, 364
Pecten (Cyclopecten) cocosensis, 238
(Cyclopecten) exquisitus, 238
(Oppenheimopecten) hancocki, 238
(Pecten) sericeus, 238
(Pecten) sericeus, Pecten, 238
Pedalion chemnitzianum, 237
pedunculata, Viola, 598, 601, 656
pelagicus, Syngnathus, 364
Pelargonium, 629
Pelecypoda, 237
Pelobatidae, 108, 117, 136
Penetopteryx nanus, 364
penicillatus, Panulirus, 249
Pennatulacea, 231
Pennatulidae, 231
Pennella instructa, 245
pennsylvanica, Fraxinus, 407
Pentaceros, 324, 325, 326, 328, 329, 331
capensis, 321, 322, 324, 325, 326, 32
331
decacanthus, 326, 327, 331
gibbus, 328
hendecacanthus, 326, 327, 331
japonicus, 321, 326, 327, 331
(Pentaceros) japonicus, Quinquarius, 319
Pentaceros richardsoni, 315, 316, 317, 3
SI) Sa) Si Say SO, 2)
330, 3315 332
richardsonii, 317

323,

Pentacerotos, 329
pentandra, Tamarix, 407
peplus, Euphorbia, 627
percomis, Silis, 469

[Proc. 4TH SER.

Silis (Silis), 455, 460, 467, 468
perennis, Bellis, 613
perfoliata, Montia, 643
Silis, 504
perfoliata, Silis (Silis), 465, 503
perforata, Silis, 459
Silis (Ditemnus), 452, 455, 458
Perinereis helleri, 244
Perna chemnitzianum, 237
quadrangularis, 238
perobliqua, Astacolus, 550, 556
persica, Veronica, 652
pes-caprae, Oxalis, 639
petechia aureola, Dendroica, 253
Dendroeca, 253
Petrography of the Galapagos, Cocos, Mal-
pelo, Cedros, San Benito, Tres Mar-
jas, and White Friars Islands, contri-
butions to the, by Charles W. Chester-
man, 339-362
Petrolisthes cocoensis, 248
edwardsii, 248
marginatus, 248
ortmanni, 248
tonsorius, 248
Phacelia californica, 597, 630
distans, 630
malvaefolia, 630
phaeocephalus, Juncus, 661
Phalaris californica, 659
canariensis, 659
Pheidole biconstricta bicolor, 251
punctatissima, 251
subarmata, 251
philadelphicus, Erigeron, 601, 614
Philautus annandali, 137
dubius, 137
Phlegathontius cingulata, 250
Phlox gracilis, 641
Phoca vitulina, 377, 378
phoebe, Prionodes, 366
Pholistoma auritum, 631
Phos cocosensis, 240
Photinia arbutifolia, 646
phragma, Ophiacantha, 234
Phrynocephalus, 129
theobaldi, 138
phrynoides, Rana, 130
Physocarpus capitatus, 601, 646
pica, Leucomelina, 224, 250

Vor. XXXII]

Picris echioides, 618
picta, Centrodera, 154, 156, 186, 192, 202,
205

pictus, Dendrelaphis, 140
Lytechinus, 236
Toxotus, 154

pilosa, Oxalis, 640

pilosus, Hipponix, 239

pilsbryi, Neritina, 240

pilularis, Baccharis, 597, 600, 601
Baccharis pilularis, 612
consanguinea, Baccharis, 596, 612
pilularis, Baccharis, 612

Pinaceae, 605

Pinaroloxias inornata, 253

pinnata, Berberis, 597, 601, 607
Mahonia, 607

pinnatifidum, Lepidium, 623

Pinniped Population of Afio Nuevo Island,

California, The, by Robert T. Orr
and Thomas C. Poulter, 377-404
Pinus contorta murrayana, 204
radiata, 605
piscator, Natrix, 107, 114, 141
Pisces, 252
Pisum sativum, 635
pittieri, Epinannolene, 249
Pityrogramma triangularis, 605
Placiphorella blainvillei, 243
Plagiobrissus pacificus, 236
Plagusia immaculata, 247
Planaxis planaxis, 240
planaxis, Planaxis, 240
Planaxis planicostatum, 240
planicostatus, 240
Planes cyanea, 247
minutus, 247
planicostatum, Planaxis, 240
planispira, Hedbergella, 568, 571
Plant Genus Polygala in the Galapagos Is-
lands, The, by John Thomas Howell
and Duncan M. Porter, 581-586
Plantaginaceae, 640
Plantago coronopus, 64.0
erecta, 596, 640
hirtella galeottiana, 641
hookeriana californica, 640
lanceolata, 641
Planulina, 567
andersoni, 567, 572

INDEX 709

spissocostata, 567
Planulininae, 567
parkeri, Altirana, 127
(Platessae), Otolithus, 87
platyceps, Natrix, 114, 130, 141
platyglossa campestris, Layia, 617
(Platypeza) agarici, Clythia, 61
Platypezina, 60

diversa, 64

pacifica, 58, 59, 60, 61, 64, 75
Platystemon californicus, 64.0
platyurus, Cosymbotus, 111, 123, 138
Plectis oweniana, 254
Plectonotum, 447
Plectritis congesta, 655
Plesiastrea, 45
pleuraden, Rana, 130
Pleurobranchaea, 517

californica, 515, 516, 517, 519
Pleurobranchaea california MacFarland, 1966

(Opisthobranchia, Notaspidea), Ob-
servations on, by Dustin D. Chivers,
515-521

plexipus, Anosia, 419
plicata, Thuja, 201
Plumbaginaceae, 641
Poa annua, 659
pratensis, 659
unilateralis, 659
Pocillopora, 42, 47
acuta, 47
damicornis, 42, 45, 46, 47, 54, 231
damicornis cespitosa, 42, 44, 45, 46, 48
elegans, 42, 44, 45, 46, 48, 231
meandrina, 42, 45, 46, 48, 232
verrucosa, 42, 45, 46, 48, 232
Pocilloporidae, 47
Podoceropsis dubia, 245
Podochela hemphilli, 246
Pogogyne serpylloides, 631
poissoni, Crepidacantha, 233
Polemius, 447, 451, 457
Polemoniaceae, 641
polii, Chiton, 434, 436, 441
Polinices glabella, 240
helicoides, 240
polita, Micropanope, 247
Pollia cinis, 238
sanguinolenta, 238
polunini, Rana, 109, 125, 130, 131, 137

710 CALIFORNIA ACADEMY OF SCIENCES

Polyastra, 50
(Polyastra) , 42
ponderosa, Pavona, 42, 45, 50, 231
Polycarpon tetraphyllum, 609
polycarpum, Thalictrum, 601, 644
Polygala, 581, 582
anderssonii, 582
chatamensis, 585
galapageia, 583, 584, 585
galapageia galapageia, 582, 583, 585
galapageia insularis, 582, 584, 585, 586
obovata, 584, 585, 586
paludosa, 581
paniculata, 581
puberula, 583
sancti-georgii, 585, 586
sancti-georgii oblanceolata, 582, 586
sancti-georgil 582,
Polygonaceae, 641
Polygonum aviculare, 642
paronychia, 597, 642
polymorpha vulgaris, Medicago, 635
Polymorphinidae, 563
Polymorphininae, 563
polymorphinoides, Nouria, 230
Polypodiaceae, 604
Polypodium californicum, 597, 605
scouleri, 597, 599, 605
Polypogon monspeliensis, 659
polypori, Clythia, 64
Polypus januarii, 243
pusillus, 243
Polystichum munitum, 599, 605
pomeridianum, Chlorogalum, 663
ponderosa, Pavona, 54
Pavona (Polyastra), 42, 45, 46, 50, 231
ponticeriana, Sitana, 138
popenoei, Anomalina, 567
popeorum, Trimeresurus, 142
Popilius lenzi, 250
Populus fremontii, 406
tremuloides, 201

585

sancti-georgil,

Porcellanidae, 248

Porites, 42, 51
californica, 51
excavata, 42, 45, 46, 51, 232
furcata, 373
lobata, 42, 45, 46, 51, 54
panamensis, 45

porosa, Tetraclita, 245

[Proc. 4TH Ser.

porphyracea, Elaphe, 140
porrectus, Typhlops, 142
porrifolius, Tragopogon, 620
Porter, Duncan M., see
Thomas
Portunidae, 246
Portunus (Achelous) brevimanus, 246
Potentilla egedii grandis, 601, 646
glandulosa, 646, 647
pottsii, Tritonia, 660
Portulaca oleracea, 643
Portulacaceae, 643
Poulter, Thomas C., see Orr,
pourtalesi, Thecopsammia, 44, 46
powellii, Amaranthus, 606
Praebulimina, 570
seabeensis, 570
prasina, Ahaetulla, 139
Elaphe, 140
pratensis, Poa, 659
prenanthoides, Erechtites, 614:
Prenolepis guatemalensis coco€énsis, 251
(Nylanderia) guatemalensis coco€nsis,
251
Primula, 127
Primulaceae, 643
Prionodes phoebe, 366
prisca, Globulina, 565
procerus, Rubus, 647
profundacella, Psammocora, 42, 45, 46, 54,
232
Psammocora (Psammocora), 49
profundus, Corythoichthys, 363, 364, 365
Prosopis juliflora, 407
Protoclythia californica, 58, 59, 60, 61, 62,
63, 64, 66, 67, 68, 75
Protozoa, 229
protracta, Silis, 504
protracta, Silis (Silis), 465, 497, 501
proxima, Silis (Silis), 465, 503, 504
Prunella vulgaris lanceolata, 631
Prunus demissa, 601, 647
emarginata, 600, 601, 647
ilicifolia, 591, 597, 601, 647
virginiana demissa, 201

Howell, John

Robert T.

Psammocora, 49

brighami, 45

profundacella, 42, 45, 46, 54, 232
(Psammocora) profundacella, Psammocora,

VoL. XXXII] INDEX alo
Psammocora (Psammocora) profundacella, melones, 241
49 patula, 240
(Stephanaria) stellata, 42, 44, 45, 46, patula pansa, 240
49, 232 purpurascens latifolius, Orthocarpus, 651

Psammodynastes pulverulentes, 141
pulverulentus, 115
Psammolyce spinosa, 244
Psammophis condanarus, 141
leithi, 141
Pseudoglandulina, 541
tenuis, 544
(Pseudoguppya) pacificus, Habroconus, 242
Pseudomyrma belti, 251
Pseudopentaceros, 317, 324, 325, 326, 331
richardsoni, 317, 318, 324
richardsonii, 317
Pseudophallus mindii, 364
Pseudopolymorphina, 565
leopolitana, 565
Pseudoreophax, 537
cisovnicensis, 537, 542
Pseudotsuga menziesii, 201
taxifolia, 192
Pseudoxenodon macrops, 115, 123, 141
Psilocarphus tenellus, 618
Psolus diomedeae, 237
Pteridium aquilinum, 599
aquilinum pubescens, 605
Pterophyta, 603, 604
Pteropoda, 242
Pterostegia drymarioides, 642, 653
Ptilosarcus gurneyl, 231
Ptyas mucosus, 115, 123, 141
puberula, Polygala, 583
puberulum, Helenium, 616
pubescens, Pteridium aquilinum, 605
pulchella, Brodiaea, 596, 663
pulcher, Rumex, 643
pulchra, Stipa, 660
pulverulentes, Psammodynastes, 141
pulverulentus, Psammodynastes, 115
pumilum, Apocynum, 598, 606
punctata, Lissemys, 138
Riopa, 112, 139
punctatissima, Pheidole, 251
punctulatus, Pasimachus, 206
pungens hartwegii, Chorizanthe, 597, 64.1
Hemizonia, 616
Purpura columellaris, 241
melo, 241

purpurea, Clarkia, 639
purpureum, Gnaphalium, 615
purshianus, Lotus, 634.
pusillus, Daucus, 654.
Octopus (Octopus), 243
Orthocarpus, 652
Polypus, 243
pycnocephalus, Carduus, 613
Pylopagurus hirtimanus, 248
longimanus, 248
pyramidalis, Marginulina, 54.9, 564
Pyrene labiosa, 241
Pyroteuthis giardi, 243
Pyrulina, 565
Python molurus, 112, 139

quadrangularis, Isognomon, 238
Perna, 238
Quadratina, 563
strombecki, 562, 563
quadridentata parva, Diacria, 242
Quadrimorphina albertensis, 566
(Quadrimorphina) ruckeri, 566, 572
(Quadrimorphina) ruckeri, Quadrimorphina,
566, 572
quercetorum, Cirsium, 614.
Stachys rigida, 632
Quercus, 204
agrifolia, 596, 627
chrysolepis, 627
chrysolepis nana, 599, 628
kellogii, 628
moreha, 628
wislizeni, 600, 628
wislizeni frutescens, 193, 628
Quinquarius, 317, 325, 326, 329, 331
capensis, 324, 325
(Pentaceros) japonicus, 319

racemosa amplexicaulis, Smilacina, 665
Rachodrassus, 80

chera, 79, 83

echinus, 79, 80, 83

flavus, 83
radiata, Colletosia, 233

Elaphe, 113, 124, 140

Pinus, 605

712 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER.

radicata, Hypochaeris, 617
(Radsia) goodallii, Chiton, 243
stokesi, Chiton, 242
radulinus, Aster, 612
Rafinesquia californica, 618
ramosissima, Lagophylla, 617
ramosissimum, Cardionema, 597, 608
Gnaphalium, 615
Lupinus, 597
Ramulina, 565
spandeli, 565
Ramulininae, 565
Rana annandalii, 137
assamensis, 137
blanfordi, 125, 130, 137
boulengeri, 130
breviceps, 108, 137
cyanophlyctis, 108, 137
formosa, 108, 137
gammel, 137
gerbillus, 137
liebigii, 108, 130, 131, 137
limnocharis, 108, 137
livida, 137
monticola, 108, 137
phrynoides, 130
pleuraden, 130
polunini, 109, 125, 130, 131, 137
swani, 109, 117, 125, 137
tigrina, 109, 137
vicina, 137
randalli, Syngnathus, 364, 373, 374, 375
Ranella caelata, 238
Ranidae, 108, 117, 137
Ranunculaceae, 643
Ranunculus californicus, 596, 644.
muricatus, 644.
Raphanus sativus, 600, 623
rappii, Liopeltis, 113, 140
raristriata, Vaginulina, 557
rashleighana, Cypraea, 239
Rattus norvegicus, 226
raymondi, Trachydermon, 434
recta, Silis (Silis), 455, 464, 489, 495
Vaginulina, 54.5, 562
recurvus, Conus, 239
refracta, Freesia, 660
reinhardti, Liocranchia, 243
reinharti, Anadara (Scapharca), 237
Arca (Anadara), 237

Relationships of the Percoid Fish Penta-
ceros richardsoni Smith, with De-
scription of a specimen from the
Coast of California, by W. I. Fol-
lett, and Lillian J. Dempster, 315-
338

remora, Echeneis, 245

Reophax agglutinatus, 230

excentricus, 230
repens, Coprosma, 648
Dichondra, 620
Trifolium, 636

Reptadeonella violacea, 233

Reptilia, 109, 137, 138, 252

Reteporellina denticulata gracilis, 233

Retevirgula areolata, 233

reticulata, Celtis, 407

retinodes, Acacia, 633

retrorsum, Geranium, 629

reversa, Silis, 494

reversa, Silis (Silis), 464, 489, 494.

Revision of the Nearctic species of Silis
(Cantharidae: Coleoptera) by John
Wagener Green, 447-513

Rhacophoridae, 109, 137

Rhacophorus jerdoni, 137

leucomystax, 137
maculatus, 109, 137
maximus, 109, 125, 137
taeniatus, 131
tuberculatus, 131

Rhagium, 184

bifasciatum, 207

(Rhagium) decoloratum, 154

Rhagium mordax, 207

Rhamnaceae, 644

Rhamnus californica, 598, 600, 644

crocea, 597, 599, 644

Rhamnusium ?decoloratum, 154

Rhinocricus (Eurhinocricus) biolleyi, 249

Rhizangiidae, 52

Rhododendron, 127

Rhus diversiloba, 596, 597, 598, 606

Rhynchonellidae, 234

Rhynchoteuthis, 243

Ribes californicum, 649

divaricatum, 649
malvaceum, 599, 600, 601, 649
menziesii leptosmum, 649

Vou. XX XI1]

richardsoni, Histiopterus, 317

Pentaceros, 315, 316, 317, 318, 319, 322,
323, SWB, SS, BAD, SPT5 SHO) Soul, 2872

Pseudopentaceros, 317, 318, 324

richardsonii, Pentaceros, 317, 330, 332

Pseudopentaceros, 317
riedeli riedeli, Vaginulina, 547, 558
Vaginulina riedeli, 547, 558
rigida quercetorum, Stachys, 632
Riopa albopunctata, 139
punctata, 112, 139
ripperti, Eutermes, 250
ritteri, Palaemon (Palaemon), 249
rivularis, Symphoricarpos, 608
Robertiacea, 569, 570
robusta, Marginulina, 549, 558
robustus, Scirpus, 656
rogersi, Newportia, 249
romaensis, Ammobaculoides, 535
Romanzoffia suksdorfii, 601, 631
romanzoffiana, Spiranthes, 666
Rooseveltia frankliniana, 254
Rorippa nasturtium-aquaticum, 623
Rosa californica, 601, 647
gymnocarpa, 600, 601, 647
nutkana, 193
Rosaceae, 603, 644
rosea, Atriplex, 610
roseus, Orthocarpus erianthus, 651
Rotaliporidae, 571
rotundus, Clypeaster, 236
Roubieva multifida, 610
rousseau, Syngnathus, 364, 368
rubens, Bromus, 657
ruber, Centranthus, 655
Rubiaceae, 648
rubicunda, Clarkia, 601, 639
rubidus, Toxotus, 154
rubra, Festuca, 658
Salicornia, 610
Spergularia, 609
Rubus parviflorus, 600, 601
parviflorus velutinus, 647
procerus, 647
spectabilis, 601
spectabilis franciscanus, 648
ursinus, 648
ruckeri, Quadrimorphina, 566, 572
ridgwayi, Nesotriccus, 253
ruficollis, Silis, 451

INDEX

rugosa, Silis (Silis), 455, 460, 467, 468
Wasmannia auropunctata, 251
Rumex, 603
acetosella, 642
californicus, 642
conglomeratus, 64.2
crispus, 642
obtusifolius, 642
pulcher, 643
salicifolius transitorius, 643
transitorius, 643
russelli, Vipera, 142
rustica erythrogaster, Hirundo, 253

saggittarius, Sibynophis, 115, 141
Sagina apetala barbata, 609
occidentalis, 609
Salamandridae, 136
Salicaceae, 648
salicifolius transitorius, Rumex, 643
Salicornia depressa, 610
europaea, 610
pacifica, 610
rubra, 610
virginica, 610
Salientia, 107
saligna, Lactuca, 617
Salix coulteri, 598, 64:8, 649, 656
gooddingii, 406
lasiolepis, 600, 649, 656
Salsola kali tenuifolia, 610
salvator, Varanus, 139
Salvia spathacea, 598, 600, 632
salvini, Sicydium, 252
Sambucus, 407
callicarpa, 204, 599, 600, 608
coerulea, 204
glauca, 407
mexicana, 608
sancti-georgii oblanceolata, Polygala,
586
Polygala, 585, 586
polygala sancti-georgii, 582, 585
sancti-georgii, Polygala, 582, 585
Sanctus, 100
sanctus, Conger, 92, 93, 95, 98, 99
sanguiniventer, Hurriah, Ik
sanguinolenta, Pollia, 238
Tritonidea, 238
sanguinolentus, Cantharus, 238

713

iS)

714 CALIFORNIA ACADEMY OF SCIENCES

Sanicula, 599
arctopoides, 596, 654.
bipinnatifida, 655
crassicaulis, 655

Polypodium, 597, 599, 605

scovelli, Syngnathus, 364
Scrophylaria californica, 652
Scrophulariaceae, 603, 650

[Proc. 4rH Srp.

laciniata, 655 Scutellaria tuberosa, 632
Saracenaria, 551, 552 Scutiger, 129
italica, 552, 558 alticola, 127, 136
spinosa, 550, 552 mammata, 127, 136
Sarcodina, 229 sikkimmensis, 108, 117, 127, 131, 133,
sarmentosa, Oenanthe, 654 136
sativa, Madia, 618 seabeensis, Praebulimina, 570
Medicago, 635 Sebastodes goodei, 316
Vicia, 636 paucispinis, 316
sativum, Pisum, 635 Sebastolobus alascanus, 316
sativus, Dipsacus, 625 Secale cereale, 659
Raphanus, 600, 623 sector, “Congermuraena”’, 87
Satureja douglasii, 601, 632 Sedum, 621

saturnale, Lytoceras, 525, 526, 527, 531
Sauria, 110, 143, 144

savignyi, Ophiactis, 235

Saxifraga californica, 649

spathulifolium, 597, 601, 621, 639
Selenkothuria theeli, 237
selloana, Cortaderia, 657
semiverticillata, Agrostis, 657

Saxifragaceae, 649 Semperothuria imitans, 237
saxonica, Lenticulina saxonica, 552, 562 sempervirens, Sequoia, 60, 195
saxonica, Lenticulina, 552, 562 Senecio, 603
Scabiosa atropurpurea, 625 aronicoides, 618
scabricauda, Otostigmus, 249 cruentus, 618
scabricosta, Nerita, 240 elegans, 618
ornata, Nerita, 240 glomeratus, 614
Scalesia, 581 mikanioides, 618
scandicina, Anthriscus, 653 minimus, 614
scapha, Cetoconcha, 237 sylvaticus, 618, 169
(Scapharca) reinharti, Anadara, 237 vulgaris, 619

“Scarria”’ mesoleuca, 238 septentrionalis, Dinodon, 140
Sceloporus, 419 Sequoia sempervirens, 60, 195
sceptrum, Nodosaria, 543 serenana, Atriplex, 610

scheeri, Balanophyllia, 42, 45, 46, 53, 54 sericeus, Pecten (Pecten), 238

’

231 Serpentes, 112, 143, 144

schencki, Textularia, 230 serpylloides, Pogogyne, 631
Schinus molle, 606 serricornis, Apatophysis, 155
Schizoplax, 437 Leptura, 155

brandti, 435, 437, 442 serriola, Lactuca, 617
Scincidae, 111, 139 sertularium, Nitidella, 240
Scirpus californicus, 656 sessilifolia, Smilacina stellata, 665

cernuus californicus, 656 sessilis, Soliva, 619

microcarpus, 656 setigera, Crepidacantha, 233

robustus, 656 setulosum, Hemiarthrum, 435, 436, 442
Scleractinia, 231 shallon, Gaultheria, 598, 626
scoparius, Cytisus, 633 Sheperdia canadensis, 195

Lotus, 634 Sibynophis collaris, 115, 151

scouleri grandis, Silene, 609 saggittarius, 115, 141

VoL. XXXII]

Sicydium salvini, 252
Sidalcea malvaeflora, 596, 614, 637
sieboldi, Enhydris, 140
sigali, Marginulina, 54.9, 564
Sigsbeia lineata, 235
sikkimense, Leiolopisma, 132, 139
sikkimmensis, Scutiger, 108, 117, 127, 131,
133, 136
Silene gallica, 609
pacifica, 609
scouleri grandis, 609
verecunda, 609
Silis, 447, 448, 450, 451, 455, 457, 459,
460, 462
abdominalis, 459
(Silis) abrupta, Silis, 465, 503, 504.
abstrusa, Silis, 462, 482, 485
acuta, Silis, 463, 489, 491
angelica, Silis, 464, 495, 497
angulata, Silis, 463, 483, 485
arida, Silis, 463, 482, 485
arizonica, Silis, 455, 464, 489,493
Silis armitagei, 458
atra, 462
(Silis) atra, Silis, 455, 462, 477, 480
barri, Silis, 467, 503, 507
Silis biauriculata, 453
californica, 494
(Silis) californica, Silis, 463, 464, 489, 493
carbo, Silis, 461, 475, 477
carmelita, Silis, 461, 467, 469
Silis cava, 449, 492
(Silis) cava, Silis, 455, 460, 463, 489, 492
constricta, Silis, 467, 503, 506
crucialis, Silis, 463, 489, 490
Silis dentigera, 505
(Silis) dentigera, Silis, 465, 497, 499
Silis deserticola, 462
(Silis) deserticola, Silis, 455, 460, 463, 483
485
Silis difficilis, 449, 475, 477, 479, 480
(Silis) difficilis, Silis, 455, 460, 461, 462,
477,478
difficilis oeeidens, Silis, 462, 479
Silis dilacerata, 455
(Silis) disjuneta, Silis, 465, 497, 500
Silis (Ditemnus) abdominalis, 452, 455, 459
(Ditemnus) bidentata, 451, 452, 455
(Ditemnus) fossiger, 451, 453, 455
(Ditemnus) freemani, 451, 453, 455

INDEX 715

(Ditemnus) howdeni, 452, 455, 456
(Ditemnus) knulli, 452, 455, 458
(Ditemnus) latiloba, 451, 452, 455
(Ditemnus) nigerrima, 452, 455, 458
(Ditemnus) obtusa, 451, 455, 456
(Ditemnus) perforata, 452, 455, 458
(Ditemnus) tricornis, 451, 455, 456
(Ditemnus) vandykei, 452, 455, 457

(Silis) divaricata, Silis, 464, 497, 498
egregia, Silis, 462, 477, 481
emarginata, Silis, 455, 463, 485
eximia, Silis, 462, 477, 481
fabulosa, Silis, 464, 497, 498

Silis fenderi, 487

(Silis) fenderi, Silis, 463, 485, 486
fenestrata, Silis, 455, 464, 489, 493
filicornis, Silis, 455, 460, 467, 471

Silis filigera, 470

(Silis) filigera, Silis, 461, 467, 470

Silis flavida, 472, 473, 475, 477

(Silis) flavida, Silis, 461, 467, 472

Silis fossiger, 453
freemani, 453
howdeni, 451

(Silis) inecongrua, Silis, 463, 483, 485
insolita, Silis, 467, 503, 507
insperata, Silis, 467, 503, 509
introversa, Silis, 465, 503

Silis lasseni, 472, 473

(Silis) lasseni, Silis, 461, 467, 472
latestyla, Silis, 464, 497

Silis latiloba, 452

(Silis) leeontei, Silis, 464, 497, 499

Silis lepida, 452

(Silis) lobata, Silis, 465, 503, 505

Silis lutea, 470, 471

(Silis) lutea, Silis, 455, 460, 461, 467, 470
macelayi, Silis, 463, 485, 489
maritima, Silis, 467, 510, 511
montanica, Silis, 464, 497, 498
Silis nevadica, 461
nitidula, 459, 462

(Silis) oregonensis, Silis, 461, 475, 477

Silis pallida, 493

(Silis) pallida, Silis, 460, 467, 503, 508
parallela, Silis, 467, 503, 506

Silis percomis, 469

(Silis) percomis, Silis, 455, 460, 467, 468

Silis perfoliata, 504

(Silis) perfoliata, Silis, 465, 503

716 CALIFORNIA ACADEMY OF SCIENCES

Silis perforata, 459
protracta, 504

(Silis) protracta, Silis, 465, 497, 501
proxima, Silis, 465, 503, 504
recta, Silis, 455, 464, 489, 495

Silis reversa, 494

(Silis) reversa, Silis, 464, 489, 494.

Silis ruficollis, 451

(Silis) rugosa, Silis, 455, 460, 467, 468

Silis (Silis) abrupta, 465, 503, 504
(Silis) abstrusa, 462, 482, 485
(Silis) aeuta, 463, 489, 491
(Silis) angeliea, 464, 495, 497
(Silis) angulata, 463, 483, 485
(Silis) arida, 463, 482, 485
(Silis) arizonica, 455, 464, 489, 493
(Silis) atra, 455, 462, 477, 480
(Silis) barri, 467, 503, 507
(Silis) californica, 463, 464, 489, 493
(Silis) carbo, 461, 475, 477
(Silis) earmelita, 461, 467, 469
(Silis) cava, 455, 460, 463, 489, 492
(Silis) constricta, 467, 503, 506
(Silis) erueialis, 463, 489, 490
(Silis) dentigera, 465, 497, 499

(Silis) deserticola, 455, 460, 462, 463,

483, 485

(Silis) difficilis, 455, 460, 461, 462, 477,

478
(Silis) difficilis oecidens, 462,479
(Silis) disjuncta, 465, 497, 500
(Silis) divarieata, 464, 497, 498
(Silis) egregia, 462, 477, 481
(Silis) emarginata, 455, 463, 485
(Silis) eximia, 462, 477, 481
(Silis) fabulosa, 464, 497, 498
(Silis) fenderi, 463, 485, 486
(Silis) fenestrata, 455, 464, 489, 493
(Silis) filicornis, 455, 460, 467, 471
s) filigera, 461, 467, 470
(Silis) flavida, 461, 467, 472
)
)

incongrua, 463, 483, 485
insolita, 467, 503, 507
(Silis) insperata, 467, 503, 509
(Silis) introversa, 465, 503

(Silis) lasseni, 461, 467, 472

(Silis) latestyla, 464, 497

(Silis) lecontei, 464, 497, 499
(Silis) lobata, 465, 503, 505

(Silis) lutea, 455, 460, 461, 467, 470

(Silis) maeelayi, 463, 485, 489
(Silis) maritima, 467, 510, 511
(Silis) montaniea, 464, 497, 498
(Silis) nevadiea, 461, 467, 474
(Silis) oregonensis, 461, 475, 477
(Silis) pallida, 460, 467, 503, 508
(Silis) parallela, 467, 503, 506
(Silis) percomis, 455, 460, 467, 468
(Silis) perfoliata, 465, 503

(Silis) protracta, 465, 497, 501
(Silis) proxima, 465, 503, 504
(Silis) reeta, 455, 464, 489, 495
(Silis) reversa, 464, 489, 494
(Silis) rugosa, 455, 460, 467, 468
(Silis) simulata, 465, 497, 501
(Silis) singularis, 462, 477, 481
(Silis) solitaria, 464, 489, 494.
(Silis) spathulata, 455, 460, 467, 469
(Silis) spinigera, 455, 460, 467
(Silis) striatella, 462, 477, 478
(Silis) subtruneata, 463, 485, 487
(Silis) tardella, 455, 460, 467, 471
(Silis) tenuata, 461, 474, 477
(Silis) thermalis, 463, 489, 491
(Silis) tripliecata, 465, 497, 500
(Silis) ursina, 463, 482, 485

(Silis) vulnerata, 455, 460, 464, 467, 503,

505
(Silis) simulata, Silis, 465, 497, 501
singularis, Silis, 462, 477, 481
solitaria, Silis, 464, 489, 494
Silis spathulata, 469

(Silis) spathulata, Silis, 455, 460, 467, 469

Silis spinicollis, 459

(Silis) spinigera, Silis, 455, 460, 467
striatella, Silis, 462, 477, 478
subtruneata, Silis, 463, 485, 487
tardella, Silis, 455, 460, 467, 471
tenuata, Silis, 461, 474, 477
thermalis, Silis, 463, 489, 491
triplicata, Silis, 465, 497, 500
ursina, Silis, 463, 482, 485

Silis vulnerata, 462, 499

(Silis) vulnerata, Silis, 455, 460, 464, 467,

503, 505
Silybum marianum, 619
Simotes octolineatus, 118
simplex, Ophiactis, 235
simulata, Silis (Silis), 465, 497, 501
singularis, Silis (Silis), 462, 477, 481

[Proc. 4TH Serr.

VoL. XXXII]

sinica, Apatophysis, 155
Siphonaria gigas, 241, 243
gigas characteristica, 241, 243
Siphostoma floridae, 369
fuscum, 369
louisianae, 369
mckayi, 370
Sisymbrium officinale, 623, 624
orientale, 624
bellum, 596, 601, 660
californicum, 660
Sitana ponticeriana, 138
Sitanion jubatum, 659

INDEX

Smilacina racemosa amplexicaulis, 665

stellata sessilifolia, 665

Smith, Allyn G., The Larval Development

of Chitons
smithi, Kachuga, 138
Solanaceae, 652
Solanum, 58
furcatum, 652
nigrum, 652
nodiflorum, 652
umbelliferum, 652
Soleirolia soleirolii, 655
soleirolii, Helxine, 655
Soleirolia, 655
Solenastrea ecuadoriana, 45
Solenopsis geminata, 251
succinea, 251
solida, Aucella, 525, 527
Buchia, 527
Solidago californica 619
canadensis elongata, 619
spathulata, 619
solitaria, Collinsia sparsiflora, 650
solitaria, Silis (Silis) , 464, 489, 494.
Soliva sessilis, 619
solstitialis, Centaurea, 613
somniferum, Papavera, 640
Sonchus asper, 619
oleraceus, 619
Sorghum halepense, 660
spandeli, Ramulina, 565
sparsiflora solitaria, Collinsia, 650
spathacea, Salvia, 598, 600, 632
spathulata, Silis, 469
Silis (Silis) 455, 460, 467, 469
Solidago, 619

(Amphineura),

433-446

spathulifolum, Sedum, 597, 601, 621, 639

speciosa, Thais, 241

speciosus, Clypeaster, 236

spectabilis, Amsinckia, 386
franciscanus, Rubus, 648
Rubus, 601

Spergula arvensis, 609

Spergularia macrotheca, 386, 609
marina, 609
rubra, 609

Sphacele calycina, 631

Sphaerodactylus lineolatus, 252
pacificus, 252

Sphenocarcinus agassizi, 246

Sphenomorphus indicum, 139
maculatum, 132, 139

Sphenotrochus hancocki, 44, 46

spicata, Distichlis, 386, 658
Menthe, 631

spicatum, Acer, 192, 205

spiculatus, Ergates, 170

Spilotes melanurus, 118

spinicollis, Silis, 459

spinigera, Silis (Silis), 455, 460, 467

spinipes, Mithrax (Mithrax), 246

spinosa, Psammolyce, 244
Saracenaria, 550, 552

spinosum, Eusigalion, 244
Xanthium, 620

Spio hirsuta, 244

Spiranthes romanzoffiana, 666

spissocostata, Planulina, 567

springeri, Syngnathus, 364, 370, 371

spurca, Centrodera, 149, 150, 151, 152,
158, 160, 161,
163, 164, 166, 167, 168, 170, 171,
DS eels Se lie Sees Onn s2e
184, 185, 187, 188, 189, 190, 191,
193, 194, 195, 196, 198, 201, 202,

155; 1565 1576

204, 205, 206, 207, 208
Pachyta, 154, 160, 168, 193, 205
Pachyta (Parapachyta), 160
Parapachyta, 154, 160, 192

spurcata, Pachyta, 169

spurcus, Evodinus, 160
Evodinus (Centrodera), 160
Toxotus, 160, 161

Squamata, 110

squamata, Eriphia, 246

squamosa, Tetraclita, 245

squamosus, Chiton 434

717

154,
162,
2h
183,
192,
203,

718 CALIFORNIA ACADEMY OF SCIENCES

squarrosa, Navarretia, 641
Stachys ajugoides, 632
bullata, 632
chamissonis, 632
rigida quercetorum, 632
staechadifolium, Eriophyllum, 386, 597, 615
stagnalis, Auricula, 239
Ellobium, 239
Staurois afghanus, 137
himalayana, 137
stejnegeri, Trimeresurus, 116, 118, 142
Stellaria media, 610
nitens, 610
stellata, Asterias, 328
Psammocora (Stephanaria), 23, 42, 44,
49
sessilifolia, Smilacina, 665
stelleri, Cryptochiton, 434
Stenocionops ovata, 246
Stenoplax, 441
heathiana, 434, 439, 441
Stenorhynchus debilis, 246
Stephanaria, 49
(Stephanaria) stellata, Psammocora, 42, 44,
45, 46, 49, 232
Stephanomeria virgata, 619
stercorarius, Camponotus (Myrmothrix) ab-
dominalis, 250
Sthenelais fusca, 244
stigma, Euponera (Trachymesopus), 251
stinemeyeri, Involutina, 568, 569
Stipa cernua, 660
lepida, 660
pulchra, 660
stokesii, Chiton, 242, 243
Chiton (Radsia), 242
stolata, Natrix, 114, 141
stoliczkae, Liopeltis, 140
striata, Acmaea, 238
striatella, Silis (Silis), 462, 477, 478
striatopunctata, Lagena, 230
striatus, Lycodon, 140
strigatella, Acmaea, 238
strigulosa, Oenothera contorta, 638
striolata, Vaginulina, 54.7, 558, 564
strombecki, Quadratina, 562, 563
Studies on the Atlantic
fishes with Descriptions of New Spe-
cies, by Earl S. Herald, 363-375
Stylomecon heterophylla, 640

American Pipe-

[Proc. 4TH SER.

subacaulis, Convolvulus, 620
subarmata, Pheidole, 251
subbracteata, Carex, 656
sublienata, Centrodera, 185, 201
subminiata, Natrix, 117, 141
subpinnatus, Lotus, 634.
subsessilis, Luzula, 661
subspicatus, Aster, 612
subtrunceata, Silis (Silis), 463, 485, 487
subviridis, Heterozona, 436
Ischnochiton, 442
Succinea, 242
globispira, 242
succulentus, Lupinus, 635
suksdorfii, Romanzoffia, 601, 631
sulcatus, Belemnopsis, 524
Swan, Lawrence W., and Alan E. Leviton,
The Herpetology of Nepal: A His-
tory, Check List, and Zoogeographical
Analysis of the Herpetofauna, 103-—
147
swanl, Rana, 109, 117, 125, 137
sylvania, Agathomyia, 64
sylvaticus, Senecio, 618, 619
Symphoricarpos, 608
albus laevigatus, 608
mollia, 608
rivularis, 608
Symplectoteuthis oulaniensis, 243
Syngnathus, 366, 373
affinis, 370
crinitus, 372
dunckeri, 364, 366, 375
elucens, 364, 366, 374
fistulatus, 364
floridae, 367, 368, 369
floridae floridae, 364, 367, 369
floridae hubbsi, 367, 368, 369
floridae mckayl, 364, 367, 370
floridae nesiotes, 364, 368
folletti, 364
fuscus, 369, 370
fuscus affinis, 363, 370
fuscus fuscus, 363
hildebrandi, 364, 366
louisianae, 364, 369, 370, 371
mckayl, 367
pelagicus, 364
randalli, 364, 373, 374, 375
rousseau, 364, 367, 368

———

Won xoxox II]

scovelli, 364
springeri, 364, 370, 371

Sypharochiton nigrovirens, 437, 438, 441,

442, 443

tabogensis, Melampus, 240
taeniatus Rhacophorus, 131
taeniura, Elaphe, 140
Talorchestia fritzi, 245
Tamaria obstipa, 234
Tamarix pentandra, 407
Tanacetum camphoratum, 598, 619
Taonidium pacificum, 243
Taraxacum laevigatum, 619
tardella, Silis (Silis) , 455, 460, 467, 471
taxifolia, Pseudotsuga, 192
tecta, Kachuga, 138
Tegula cooksoni, 241
gallina multifilosa, 241
maculostriata, 241
Teleophrys cristulipes, 246
(Teleoteuthis) , Onykia, 243
Tellima grandiflora, 650
telmateia braunii, Equisetum, 604.
tenella, Hesperocnide, 655
Microseris douglasii, 618
tenellus, Psilocarphus, 618
tenera, Lingulina, 550, 560
tenuata, Silis (Silis), 461, 474, 477
Tenues, 581
tenuiceps, Trachischium, 115, 151
tenuicosta, Nodosaria, 543, 564
tenuidactylos, Ozius, 247
tenuidactylus, Ozius, 247
tenuiflorus, Carduus, 613
tenuifolia, Salsola kali, 610

tenuilamellosa, Tubastrea, 42, 44, 45, 46, 54,

Zoe
tenuirostre, Copidozoum, 233
tenius congestus, Juncus, 661
Pseudoglandulina, 544
terrestris, Brodiaea, 663
Testudinata, 109
Testudinidae, 110, 138
Testudo elongata, 110, 132, 138
Tetraclita squamosa, 245
squamosa milleporosa, 245
Tetraelita porosa, 245
Tetragonia expansa, 606
tetragonioides, 606

INDEX 719

tetragoniodes, Tetragonia, 606
Tetramorium guineénse, 251
tetraphyllum, Polycarpon, 609
texana, Linaria, 651
texasensis, Marginulina, 548
Textularia articulata, 230
conica, 230
corrugata, 230
panamensis, 230
schencki, 230
Textulariina, 532
Thais columellaris, 241
crassa, 241
haemastoma biserialis, 241
patula, 240, 241
speciosa, 241
triangularis, 241
(Vasula) melones, 241
thalia, Caducifer, 238
Thalictrum polycarpum, 601, 644:
Thamnasteriidae, 49
The Goose Lake Fragments, by C. P. But-
ler, 291-313
The Herpetology of Nepal: A _ History,
Check List, and Zoogeographical Anal-
ysis of the Herpetofauna, by Law-
rence W. Swan and Alan E. Leviton,
103-147
The Larval Development of Chitons (Am-
phineura), by Allyn G. Smith, 433-
446
The Pinniped Population of Ano Nuevo Is-
land, California, by Robert T. Orr,
and Thomas C. Poulter, 377-404
The Plant Genus Polygala in the Gala-
pagos Islands, by John Thomas How-
ell, and Duncan M. Porter, 581-586
Thecopsammia pourtalesi, 44, 46
theeli, Selenkothuria, 237
Theelothuria paraprinceps, 237
Thelypodium lasiophyllum, 624.
theobaldi, Phrynocephalus, 138
Thermophis baileyi, 129
thomasi, Nuttallina, 434, 436, 438, 442
thouarsii, Eucidaris, 236
Thuja plicata, 201
Thuriaria orisioides, 230
thurjii, Hardella, 137
Thyrolambrus erosus, 246
glasselli, 246

720 CALIFORNIA ACADEMY OF SCIENCES

thyrsiflorus, Ceanothus, 598, 600, 601, 644
Thysanocarpus curvipes, 624
tiaratus, Conus, 239
tibetanus, Alsophylax, 138
tigrina, Rana, 109, 137
Tillaea erecta, 621
Tomopterna, 125
tonsorius, Petrolisthes, 248
torreyana, Melica, 659
townsendi, Anolis, 252
toxotoides, Apatophysis, 155
Toxotus pictus, 154
rubidus, 154
spurcus, 160, 161
Trachischium fuscum, 115, 141
guentheri, 115, 127, 141
laeve, 141
tenuiceps, 115, 141
Trachydermon, 434
(Trachymesopus) stigma, Euponera, 251
raymondi, 434
Tragopan, 127
Tragopogon porrifolius, 620
Tragosoma depsarius, 195
Tralia panamensis, 240
transitorius, Rumex, 643
Rumex salicifolius, 643
transversus, Pachygrapsus, 247
Trapezia cymodoce ferruginea, 247
tremuloides, Populus, 201
triangularis, Pityrogramma, 605
Thais, 241
triangulatus, Microphrys, 246
tricarinata, Geoemyda, 132, 137
Japalura, 111, 127, 131, 138
Trichopsetta ventralis, 366
tricornis, Silis (Ditemnus), 451, 455, 456
tridentatum, Trifolium, 636
Trifolium, 603
amplectens, 635
bifidum, 635
dubium, 635
gracilentum, 635
microcephalum, 636
microdon, 636
repens, 636
tridentatum, 636
wormskjoldii, 636
Triglochin concinna, 600, 661
trigonata, Boiga, 113, 140

[Proc. 4TH SER.

trijuga, Geoemyda, 132, 137
trilineatus, Melampus, 240
Trillium chloropetalum, 601, 665
Trimeresurus albolabris, 116, 142
erythrurus, 142
gramineus, 118
monticola, 116, 125, 142
popeorum, 142
stejnegeri, 116, 118, 142
viridis, 118
Triodanus biflora, 608
Trionychidae, 110, 138
Trionyx gangeticus, 110, 138
hurum, 138
leithi, 138
Triplasia, 535
triplicata, Silis (Silis), 465, 497, 500
tripudians, Naja, 118
triquetrum, Allium, 661
tristis, Dendrelaphis, 140
Mitra, 240
Tristix, 563
acutangulum, 562, 563
triticoides, Elymus, 658
Triticum aestivum, 660
Tritonia aurea, 660
crocosmaeflora, 660
pottsii, 660
Tritonidea cinis, 238
sanguinolenta, 238
Trivia pacifica, 241
trixago, Bellardia, 650
Trochammina, 533
charlottensis, 230
nitida, 230
orchardensis, 534, 542
Trochamminidae, 533
Trochammininae, 533
Tropaeolaceae, 652
Tropaeolum majus, 652
truncata, Aquilegia formosa, 643
Vaginulina, 548
Trypostega venusta, 233
Tubastrea, 54
tenuilamellosa, 42, 44, 45, 46, 54, 232
tuberculata, Agama, 110, 129, 130, 138
tuberculatus, Chiton, 434
Latirus, 239
tuberosa, Lissa, 245
Scutellaria, 632

Vor. XXXII]

Tubulipora flexuosa, 233
tulipifera, Liriodendron, 191
tumida, Herbstia, 245
tumulicola, Carex, 656
tunicata, Katharina, 434
turnus, Papilio, 419
Turrilinidae, 570
Turrilininae, 570
Two Gnaphosid Spiders from Arkansas, by
Harriet Exline, 79-85
Tylototriton verrucosa, 136
Typha domingensis, 666
latifolia, 666
Typhaceae, 666
Typhlopidae, 142
Typhlops bothriorynchus, 142
braminus, 142
jerdoni, 142
oligolepis, 142
porrectus, 142
Typocerus cervinus, 160, 161, 168

Uca galapagensis, 247, 248
macrodactyla, 248
panamensis, 247

Ulex europaeus, 600, 636

umbellata, Discoporella, 233

Umbelliferae, 603, 652

umbelliferum, Solanum, 652

Umbellularia californica, 596, 599, 632

umiatensis, Conorboides, 570

unalascensis elata, Habenaria, 666
Habenaria unalascensis, 666
maritima, Habenaria, 665
unalascensis, Habenaria, 666

uncitoides, Aucella, 525, 527

undulatus, Leioptilus, 231

unedo, Arbutus, 625

uniflora, Orobanche, 639

unifolium, Allium, 601, 663

unilateralis, Poa, 659

Uroconger, 92

Urodela, 136

ursina, Silis (Silis), 463, 482, 485

ursinus, Callorhinus, 378, 401
Rubus, 648

Urtica holosericea, 655

Urticaceae, 655

usitatissimum, Linum, 636

utriculatum, Lomatium, 654

uva-ursi, Arctostaphylos, 598, 601, 626

INDEX

Vaccinium arbuscula, 599, 601, 627

ovatum, 600, 601, 627
Vaginulina, 545, 560, 561

debilis, 546, 564

exilis, 230

kochii, 547, 548, 558

octocostata, 546, 562, 564

raristriata, 557

recta, 54.5, 562

riedeli riedeli, 54.7, 558

striolata, 54.7, 558, 564

truncata, 548
Vaginulinopsis, 548

pachynota, 548, 558
valanginiana, Lenticulina, 551
Valerianaceae, 655
van-brunti, Echinometra, 236

vandykei, Silis (Ditemnus) , 452, 455, 457

vanillae, Agroaulis, 250

Varanidae, 112, 139

Varanus flavescens, 112, 139
monitor, 112, 139
salvator, 139

variana, Bimonilina, 540

varians, Pavona, 42, 45, 46, 54
Pavona (Pavona), 50, 231

variegata, Japalura, 138

variegatum, Ophioderma, 235

variicolor, Lupinus, 635

(Vasula) melones, Thais, 241

vaughani, Endopachys, 44, 46, 231

veleronis Cellaria, 232

velutinus, Rubus parviflorus, 647

venenosus, Zigadenus, 665

ventralis, Trichopsetta, 366

ventromaculatus, Coluber, 140

venusta, Trypostega, 233

verecunda, Silene, 609

verna, Barbarea, 622

Verneuilininae, 537, 540

Veronica americana, 652
persica, 652

verreauxii, Ozius, 247

verrucosa, Pocillopora, 42, 45, 46, 48, 232
versicolor, Calotes, 105, 110, 123, 138

vestitum, Cymatium, 239
vestitus, Lathyrus, 633, 636
vetulus, Parophrys, 315

vetustus, Conger, 88, 90, 92, 95, 100, 101

Vicia americana, 636

~
Do
bo

angustifolia, 636
benghalensis, 636
gigantea, 636
sativa, 636

vicina, Rana, 137

villosa bolanderi, Chrysopsis, 613
franciscana, Monardella, 597, 600
Monardella, 631

vimineum, Eriogonum, 642

Vinca major, 606

Viola adunca, 601, 655
pedunculata, 598, 601, 656

violacea, Amphiodia, 234
Reptadeonella, 233

Violaceae, 655

Vipera russelli, 142

Viperidae, 116, 117, 142

virens, Icteria, 407

virescens, Fissurella, 239
maculatus, Butorides, 253

virgata, Stephanomeria, 619

virgatum, Conchoderma, 245

virgatus, Astragalus nuttallii, 633

virginiana demissa, Prunus, 201

virginica, Salicornia, 610

virgula, Creseus, 242

viridis, Chloeia, 244
Trimeresurus, 118

virosa, Lactuca, 617

viscosum, Cerastium, 609

vittatus, Corythoichthys, 372
ensenadae, Micrognathus, 373
Micrognathus, 364, 372, 373

vitulina, Phoca, 377, 378

viviparus, Callistochiton, 435, 436, 441, 442

vulgare, Cirsium, 614.
Foeniculum, 654.
Marrubium, 631

vulgaris lanceolata, Prunella, 631
Medicago polymorpha, 635
Senecio, 619

CALIFORNIA ACADEMY OF SCIENCES

[Proc. 4TH SER.

vulnerata, Silis, 462, 499
Silis (Silis), 455, 460, 464, 467, 503,
505

walli, Bungaris, 142
washingtoni, Halecium, 230
Wasmannia auropunctata, 224, 251
auropunctata rugosa, 251
watsonii, Epilobium, 639
wellsi, Lophosmilia, 44, 46
westermanni, Elachistodon, 131, 141
wightii, Castilleja, 650
winslowensis, Acroteuthis, 524, 526
wislizeni frutescens, Quercus, 193, 628
Quercus, 600, 628
wollebaeki, Zalophus californianus, 379
Woodwardia fimbriata, 599, 605
wormskjoldii, Trifolium, 636
Wyethia, 616
angustifolia, 620

Xanthias politus, 247
Xanthidae, 246
Xanthium spinosus, 620
Xanthodius lobatus, 247
Xenochropis cerasogaster, 116, 141
xiphioides, Juncus, 661
Xylosteus, 156

ornatus, 156

zachirus, Glyptocephalus, 315
Zalophus, 378, 380

californianus, 377, 378, 379

californianus californianus, 379

californianus japonicus, 379

californianus wollebaeki, 379
Zantedeschia aethiopica, 656
Zaocys nigromarginatus, 116, 141
Zauschneria californica, 601, 639
Zigadenus, 665

fremontii minor, 665

venenosus, 665

ERRATA

Page 50. Line 10 from bottom: for (Polyastrea) read (Polyastra).

Page 132. Line 22 from top: for sikkimensis read sikkimense.

Page 231. Line 20 from bottom: for Cladocera read Cladocora.

Page 235. Line 3 from bottom: for mexicana read mexicanus.

Page 245. Line 13 fromtop: for Tetraelita read Tetraclita.

Page 443. After line 11 from top, in center of page add: LITERATURE CITED.
Page 451. Line 18 from top: for Silas read Silis.

Page 627. Line 5 from bottom:

for chysolepis read chrysolepis.

PROCEEDINGS

OF THE

California Academy of Sciences

FOURTH SERIES

G Dallas Hanna Anniversary Volume

Vol. XXXII, No. 1, pp. 1-40; frontispiece; 2 figs. April 24, 1962

G DALLAS HANNA
by

Robert C. Miller

California Academy of Sciences

Marine Biological Laboratory)
i Sa i =f 5 4’ SS eS :

APR 1 8 1963
WOODS HOLE, MASS.

SAN FRANCISCO
PUBLISHED BY THE ACADEMY
1962

; >
LG UAT
COMMITTEE ON PUBLICATION
Dr. Roperr C. Minter, Chairman — > |
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Dr. Epwarp L. Kesseu, Editor Dr. Leo G/HERTLEIN Fo
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