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TRANSACTIONS HARVARD 
UNIVERSITY 


OF THE 


SAN DIEGO SOCIETY OF NATURAL HISlvORyY 


2 
VOLUME 13 


PRINTED FROM THE 


W. W. WuHitNeY PUBLICATION ENDOWMENT 


SAN DIEGO, CALIFORNIA 
PRINTED FOR THE SOCIETY 


1962-1964 


COMMITTEE ON PUBLICATION 


BAYLorR Brooks 
JoHN Apams Comstock 
Cart L. Husss 


LAURENCE M. KLAUBER 


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CONTENTS 


. The Belvedere Expedition to the Gulf of California. By George 


Be bindsay-INoverber OIG xe ace: ee ee te se 1-44 


. The early stages of Stenaspilates apapinaria Dyar (Lepidoptera: 


Geometridae). By John Adams Comstock. April 19, 1963... 45-48 


. Birds of the Belvedere Expedition to the Gulf of California. By 


Richard’@.bankss June) 1963" eee Bert er sa oe 49-60 


. A second new species of megathymid from Baja California, 


Mexico (Lepidoptera: Megathymidae). By Charles F. Harbison. 
TPG (ay (1s Sembee emcee, War cakes, OR 61.72 


. A new insular subspecies of the speckled rattlesnake. By L. M. 


KiauberSeptember-27, 19603.) " ey: ea ee ee ee ee 73-80 


. Two new opisthobranch mollusks from Baja California. By Wes- 


leyalVie Harmer. September. 27, 01963: seers ee een ee 81-84 


. The mammals of Baja California, Mexico. By Laurence M. 


Inluey Sanwaty- 15 OG Mie re Set ten 85-168 


. The larva and pupa of A gathymus dawsoni (Lepidoptera: Mega- 


thymidae). By John Adams Comstock. February 14, 1964... 169-172 


. A new marine beetle from the Gulf of California (Coleoptera: 


Staphylinidae). By Ian Moore. February 14, 1964.02... 173-176 


Birds and mammals of the voyage of the “Gringa.” By Richard 
Gr Banike: Bebruatyol 4 1964 2th sate ee ole eo eee 177-184 


Evolution of the pit vipers. By Bayard H. Brattstrom. May 4, 
De oi bk ue tly Nee 4) enema ts SRE ran tye A eee ee Ee Loe & 185-268 


The Staphylinidae of the marine mud flats of southern Cali- 
fornia and northwestern Baja California (Coleoptera). By Ian 


Dooce: Vey F Gace ees eee eh ee ene ee 269-284 


The structure and reproduction of the red alga Chondria nidifica 
Harvey. By E. Yale Dawson and Bilgin Toztin. August 28, 1964 285-300 


PiA FR 
UNIVE 


18. 


19. 


20. 


. Notes on variability and range in the elk kelp Pelagophycus. By 


Bruce C. Parker and E. Yale Dawson. August 28, 1964................ 


. The larva of Hadrotes crassus (Mannerheim) (Coleoptera: Sta- 


phylinidae). By Ian Moore. August 28, 1964.........-..-...-------------- 


. Terrestrial mollusks of the Belvedere Expedition to the Gulf of 


California. By William K. Emerson and Morris K. Jacobson. 
ANUGUSEEZO, PLO Gite ee 2a eee ieee Se se eee 


. Invertebrate megafossils of the Belvedere Expedition to the Gulf 


of California. By William K. Emerson and Leo George Hertlein. 
December 0. A19G4 sen Ses a eee 4 Ee eee 


The races of Haliotis fulgens Philippi (Mollusca: Gastropoda). 
By Robert Ro Malmage: December 501064. = ee 


Additions to the nudibranch fauna of the east Pacific and the 
Gulf of California. By Clinton L. Collier and Wesley M. Farmer. 
Wecember BO 1964 yo ee as 2S were. 1s acl ne SR ee 


The mammals of Cerralvo Island, Baja California. By Richard C. 
Banks December 3041964: <a Sees es a ee 


ERRATA 


Page 52, paragraph 3, line 1 — read aethereus for aetherus. 


Running headings, odd-numbered pages 89-165 — read 1964 for 1963. 


Page 181, paragraph 4, line 1 — read nigripes for nigrepes. 


Page 218, paragraph 1, line 2— read mokeson for mokasen. 


Page 372, paragraph 2, line 5 — read Vokes for Voke. 


Page 379, paragraph 3, line 3 — read Harald for Harold. 


Page 381, paragraph 8, line 3 — read CASIZ 26 for CASIZ 27. 


301-308 


309-312 


313-332 


333-368 


369-376 


377-396 


397-404 


405-415 


TRANSACTIONS 
OF THE 


SAN DIEGO "SOCIETY OF NATURAL HISTORY 


VoLuME 13, No. 1, pp. 1-44, figs. 1-28 


THE BELVEDERE EXPEDITION 
TO THE GULF OF CALIFORNIA 


BY 


GerorGE E. LINDSAY 


Director, San Diego Natural History Museum 


SAN DIEGO, CALIFORNIA 
PRINTED FOR THE SOCIETY 


NovEMBER 9, 1962 


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Fic. 1. Route of the expedition. For detail of area A, see page 4; for area B, see 
page 17; for area C, see page 28. 


MUS. COMP. z00 
DEC 31 1969 
naRyARn 

UNIVERSITY 


THE BELVEDERE EXPEDITION 
TO THE GULF OF CALIFORNIA 


BY 
GeorGE E. LINDSAY 


CONTENTS 
LbaeRo (eke ata c(i Rb Reet RAE UUs, 2S RS ea ae Soe PER EGY AN Dual cote n'a 3 
Mopmatetheyespecitone ta enue ee Re = eee ae ee 5 
Preliminary report of the scientific collections —..0...-..--...---0ec-sceseeeseeceeeceeeeeeeeees 40 
ENcknOw edo inedist: Yr sei mals ett, WAR, | Soe BC laa Meer ile es ei 43 
INTRODUCTION 


The Belvedere Expedition was a voyage for the biological exploration of 
the islands in the Gulf of California. It was a function of the San Diego Natural 
History Museum, supported by a grant from the Belvedere Scientific Fund of 
San Francisco. The “San Agustin IT”, an eighty-five foot motor vessel with a 
crew of eight, was chartered for the trip. The owner, Mr. Antero Diaz, accom- 


panied the boat. 


Twelve biologists participated, seven for the entire voyage. George Lindsay, 
as Director of the Museum, was in charge. Ira L. Wiggins, Professor of Biology 
at Stanford University and Scientific Director of the Belvedere Scientific Fund, 
made general botanical collections, as did Reid Moran, Curator of Botany of 
the Museum. Richard C. Banks, Curator of Birds and Mammals, collected birds 
and mammals. Charles F. Harbison, Curator of Entomology, collected land 
arthropods. William K. Emerson, of the American Museum of Natural History, 
collected fossil and recent invertebrates. Charles Shaw, Assistant Director and 
Curator of Reptiles of the San Diego Zoo, collected reptiles with the assistance 
of Michael Soulé, a graduate student at Stanford University. Don Hunsaker I], 
Assistant Professor of Zoology at San Diego State, Dennis Bostic, his graduate 
student, and Chris Parrish, Museum Assistant and student at San Diego State 
College, joined the expedition at La Paz for eight days during their spring holiday. 
Ambrosio Gonzalez C., Jefe de Investigadores of the Instituto Mexicano de 
Recursos Naturales Renovables, was with the expedition from March 14th through 
April 3rd. 

The “San Agustin II” sailed from Bahia de los Angeles March 15th, 1962, 
and returned April 26th. During the voyage, 32 islands were visited, with several 
stations on each of the larger ones. A stop was made at Punta Sargento, Sonora, 
and there were seven stations along the Baja California coast. 


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1962 | LINDSAY: BELVEDERE EXPEDITION 5 


Fig. 3. The “San Agustin IT”, the eighty-five foot motor vessel chartered for 
the expedition. 


LoG OF THE EXPEDITION 


Wiggins and Soulé were the first to leave San Diego, taking a load of 
expedition gear in the International Travelall of the Belvedere Scientific Fund. 
They left March 11th and arrived at Bahia de los Angeles March 13th. Gonzalez, 
Emerson, Moran, Banks, Harbison, Shaw, and Lindsay flew from Tijuana to 
Bahia de los Angeles the morning of March 14th, arriving at 1105. All hands 
spent the afternoon getting the equipment and supplies aboard the “San Agustin 
HES 


March 15: Bahia de los Angeles to Puerto Refugio, Isla Angel de la Guarda. 


We breakfasted ashore, got the last gear aboard, and departed from Bahia 
de los Angeles at 0900, en route to Isla Angel de la Guarda. The crossing was 
choppy, and there was a cold wind. The first stop was at Arroyo Estaton, at 1200. 

Angel de la Guarda is 47 miles long — the longest island in the Gulf — and 
12 miles wide in the south-central part. It has a peak 4315 feet high in the northern 
section. The upper part of the island is composed of volcanic rocks, resting on 
a basement of granitic and metamorphic rocks, which are exposed at several 
localities. The flora includes the elephant tree (Pachycormus discolor), cardon 
cactus (Pachycereus pringlei), and other plants of the central desert flora of Baja 
California. Angel de la Guarda is uninhabited. 


6 SAN Dteco Society oF NAtTurRAL History { VoL. 13 


il . i i 
3 ~ ae rs “4 P ‘ & 2 “SQ ; 


Fic. 4. Biologists of the expedition. Upper row, left to right: Richard C. Banks, 
Michael Soulé, Don Hunsaker II, Chris Parrish, Ira L. Wiggins, Charles Shaw, 
Reid Moran. Lower row: William K. Emerson, Dennis Bostic, Charles F. Har- 
bison. Behind the camera: George Lindsay. Back in Mexico City: Ambrosio 
Gonzalez C. 


We were pleased, when we went ashore at Arroyo Estaton, to find the island 
relatively moist, and many plants in flower. Lindsay climbed a ridge to where a 
Museum party had collected two giant rattlesnakes (Crotalus mitchelli ssp.) in 
1960; he found no more snakes but did recover a geologist’s hammer left at that 
time. All were aboard at 1500, and we went to Puerto Refugio, where we found a 
good anchorage between Isla Mejia and the main island. Harbison spent the 
night ashore to collect insects with gasoline lanterns; Banks set small mammal 
traps. 


March 16: Puerto Refugio, Isla Angel de la Guarda. 


Shore parties worked Isla Mejia and Angel de la Guarda. Shaw, Soule, 
Gonzalez, Emerson and Lindsay checked reptile traps left by John Sloan in 
August 1961. These were five-gallon cans sunk level with the soil surface and 
covered with masonite tops but with space for reptiles to enter. The first trap, 
in a dune area, had filled with sand; but the others, placed in rocky situations, 
contained many lizards, rodents, scorpions, and insects. These were removed 
for shipment to Sloan in San Diego. 


1962 | LINDSAY: BELVEDERE EXPEDITION 7 


# ee . Se ac 


Fic. 5. Michael Soulé and Charles Shaw removing specimens from a reptile trap 


at Puerto Refugio, Isla Angel de la Guarda. 


ta: 


Fic. 6. Charles Shaw and a friend, a large black chuckwalla (Sauromalus his- 
pidus), at Puerto Refugio, Isla Angel de la Guarda. 


8 San Disco Society oF Naturat History {Vox 13 


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Fic. 7. North end of Isla Angel de la Guarda. 


The botanists found collecting good, and there was a fine display of wild- 
flowers. Among several new records was Phacelia pauciflora, a rare plant known 
only from a few collections from the vicinity of Bahia de los Angeles but not 
known from any of the islands. 


We made a special effort to collect scorpions on each of the islands and 
did so without exception. At Puerto Refugio we were surprised to find many 
small scorpions under rocks in the upper littoral zone: five were collected under 
a single beach boulder. They were in sand moist with sea water, in an area 
subject to inundation at extreme high tide. Apparently they are an undescribed 
V ejovis known also from Bahia de los Angeles. 


Large black chuckwallas (Sauromalus hispidus) are abundant on Isla Angel 
de la Guarda, and a series of living ones was taken for the San Diego Zoo. 
One specimen of speckled rattlesnake (Crotalus mitchelli) was collected. 


March 17: Puerto Refugio to Pond Island, Isla Angel de la Guarda. 


We were under way at 0700, coasting down the east side of Angel de la 
Guarda to Pond Island. Because it is supposed to be crawling with rattlesnakes, 
Pond Island is known locally as Isla Vibora; and it did yield one specimen of 
red rattlesnake (Crotalus ruber) , for a new record. 


Pond Island is only about one mile long and 400 feet high. It is connected 
to Angel de la Guarda by a rocky reef which drys in places at low water. Wiggins, 
Banks, Soulé, Gonzalez, Shaw, and Lindsay spent the afternoon on Pond; Moran 
and Harbison went to the main island near Pond Island Bay; and Emerson took 


1962 } LINDSAY: BELVEDERE EXPEDITION 2 


a small boat down the coast about two miles to sample some fossil deposits. Banks 
collected a nesting Craveri’s murrelet (Endomychura craver1) for a new breeding 
station. An attractive lagoon at Pond Island has a prolific fauna. Emerson 
returned to collect there; and Diaz turned malacologist to the extent of collecting 
two sacks of large rock oysters, which we had steamed at cocktail time and fried 
for dinner. We enjoyed a comfortable anchorage south of Pond Island. 


March 18: Pond Island to Punta Sargento, Sonora, and Bahia Agua Dulce, 
Isla Tiburon. 


We were under way at 0330 on the fifty-mile run to Punta Sargento, through 
flat calm water. We changed course several times to photograph finback whales, 
about 25 of which were observed off the west side of Isla Tiburon. Near Isla 
Patos the sea was covered by feeding Bonaparte’s gulls, dainty birds which pick 
macroplankton from the water as they swim or hover. There were also many 
terns, loons, cormorants, and pelicans. A long line of flying pelicans had three 
blue-footed boobies in formation, a situation we were to observe often. 


Shoal water near Punta Sargento made it necessary to anchor a mile off- 
shore. We landed at a campsite, apparently deserted, but a canoe with three young 
Seri Indians, two men and a woman, approached evidently expecting to land. 
Seeing us, they changed course and went around the point. We found a tremen- 
dous midden just above the beach, a yard thick and hundreds of yards long. 


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Fic. 8. Reid Moran collecting plants at Puerto Refugio, Isla Angel de la Guarda. 


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10 San Disco Society oF Natrurat History {Vor./13 


Fic. 9. Ambrosio Gonzalez C. examining a sour pitahaya plant (Machaerocereus 
gummosus) at Punta Sargento, Sonora. This cactus had not been known to occur 
on the Mexican mainland. 


One reason for visiting Punta Sargento was to verify the occurrence there 
of the sour pitahaya (Machaerocereus gummosus) , seen from the air in December 
1960. The cactus was thought to be endemic to Baja California and the Gulf 
islands and not known to occur on the Sonora mainland. We found the sour 
pitahaya in abundance, as well as a tuberous-rooted cactus (Wilcoxia digueti1) . 


The shore party returned to the ship at 1200; and we went to Isla Tiburon. 
Tiburon is the largest island in the Gulf of California, 30 miles long and 20 miles 
wide, the only large island on the eastern side of the Gulf. The highest range in 
the eastern margin rises to nearly 4000 feet. The island is separated from Sonora 
by a shallow, narrow strait, 2 to 4 miles wide and in places less than 20 feet deep. 


We spent the afternoon collecting at Bahia Agua Dulce, at the north end 
of Tiburon. Crucifixion thorn (Koeberlinia spinosa) was found here for the first 
insular record. Four canoe loads of fishermen were camped on shore, their camp 
surrounded by drying shark meat. Harbison went ashore for the night, to collect 
insects. 


1962 | LINDSAY: BELVEDERE EXPEDITION 11 


March 19: Bahia Agua Dulce to Punta Willard, Isla Tiburon. 


A brisk offshore breeze held us sideways to the swell, causing a heavy roll 
and a confusion of fallen books, boxes, and gear in the cabin. The plant drier 
secured to the cabin roof, used heat lamps run by the main generator. The drier 
was covered with a tarp at night after the generator was shut down, to protect 
the plant presses from dew. When the engineer started the main generator at 
0430, no one thought to remove the cover from the plant presses, and the whole 
thing burst into flame about 0600. There was much excitement as crewmen 
threw burning tarpaulins, corrugates, and specimens into the sea. The previous 
day’s plant collections were lost. 


We were under way down the west coast of Tiburén at 0800. Many birds 
and several whales were observed. Anchoring at Punta Willard at 1100, the 
shore party worked until 1430, when a sudden shower sent everybody aboard. 
Diaz and a crewman, José Toledo, went hunting and returned with a nice little 
mule deer (Odocoileus hemionus sheldoni), which is endemic to Tiburén. 
Banks supervised skinning and kept the skin and skull for a specimen. Three 
shrimp trawlers passed and anchored under the point in the evening. 


March 20: Punta Willard to Sauzal, Isla Tiburon. 


We were under way soon after 0400 and anchored off Arroyo Sauzal, on 
the south side of Tiburén, at 0600. Shrimp boats anchored nearby supplied 
us with fresh shrimp and agreed to take mail to Guaymas. José Toledo brought 
aboard some antelope jack rabbits (Lepus alleni tiburonensis), one of which 
was alive and looked strange indeed, hopping about the deck. 


Two new reptile records were obtained on Isla Tiburon: Harbison found 
a Sonora boa (Lichanura trivirgata); and Soulé collected a Sonoran gopher 
snake (Pituophis catenifer affinis). 


March 21: Isla Tiburon to Isla San Pedro Martir and Isla San Esteban. 


The trip from Isla Tiburén to Isla San Pedro Martir was calm. San Pedro 
Martir is a triangular rock mass 1052 feet high and less than a mile wide, lying 
in mid-Gulf. It apparently is rhyolite. Being the nesting place of thousands of 
blue-footed and brown booby birds, it is almost entirely covered with guano. 
Brown pelicans also nest on the island, and red-billed tropic birds were observed 
flying above. 

We anchored in a small cove on the northeast side. Guano was collected 
commercially in the past, and many stone terraces and platforms for its ac- 
cumulation remained on the steep slopes. Large iron rings attached to rocky cliffs 
apparently were used to hold lighters at the small shingle beach where we put 
ashore. California sea lions were tame and played about the shoreboats. 


Two large dark endemic lizards (Uta palmeri and Cnemidophorus tigris 
martyris) were very abundant over the whole island, even foraging on the algae- 
covered rocks of the intertidal zone. They were not timid, and many were taken 


12 San Disco Society oF Natura History { VoL. 13 


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Fic. 10. Blue-footed boobies (Sula nebouxii) on Isla San Pedro Martir. 


alive without difficulty. The only snake known from Isla San Pedro Martir is 
a large rattlesnake (Crotalus atrox), a single specimen of which was taken by 
Reid Moran in 1952. Our party collected seven additional examples of this 
large rattlesnake, which will make possible a more critical taxonomic study. 


The upper part of Isla San Pedro Martir was covered with a bright globe- 
mallow (Sphaeralcea hainesit) in full bloom, forming a colorful understory of 
a fine forest of Pachycereus pringlei. The biologists returned to the beach at 
noon, rather reluctant to leave this interesting island. Collections were processed 
on the three-hour run to Isla San Esteban, where we anchored off the mouth of 
a broad arroyo on the southeast side, just north of an offlying rock. 


Isla San Esteban is about 4 miles long north and south, 3 miles wide, and 
1772 feet high. Tidal currents are very fast, and tide rips cause broken water 
during the most rapid flow. A low shingle spit projecting about 3/4, mile from 
the southwest end of the island, is the nesting place of western gulls. Osprey 
nests occupied the tops of most pinnacles and cliffs. 


1962] LinpsAy: BELVEDERE EXPEDITION 13 


Fic. 11. Ira L. Wiggins with a large rattlesnake (Crotalus atrox) which he 
collected on Isla San Pedro Martir. 


14 San Dreco Society oF NaturAL History { Vo. 13 


March 22: Isla San Esteban. 


The whole day was spent at the same anchorage. Moran climbed to the 
top of the island and found particularly good collecting on north slopes, adding 
many species to the known flora. Interesting finds included a crucifer (Lyrocarpa 
linearifolia) previously known only from one collection from Angel de la Guarda 
and an ironwood (Sideroxylon leucophyllum) known from no other island but 
Angel de la Guarda. Harbison and Moran saw racers which they were unable 
to collect. A large pink-and-brown blotched chuckwalla (Sauromalus varius) 
and another large lizard (Ctenosaura hemilopha conspicuosa) inhabit the rocky 
atroyos and slopes. A series of living specimens was taken for the San Diego 
Zoo. An endemic blacktailed rattlesnake, only three specimens of which have 
been taken, was not found by our party. 


A large hedgehog cactus (Echinocereus grandis) was in flower, as were an 
as-yet-undescribed species of Mammillaria and a cholla cactus. Emerson found 
a Pliocene outcrop just north of the anchorage, from which he collected echino- 
derm spines, oysters, and corals. Diaz, while fishing for the galley, caught a 
large garoupa. Scorpions (Hadrurus sp. and Centruroides exilicauda) were 
fairly abundant but small. 


March 23: Isla San Esteban to Isla Salsipuedes. 


A strong west wind came up during the night, but our anchorage was pro- 
tected. Banks and Wiggins went ashore with the boat that picked up Harbison, 
who had collected insects during the night. We were under way about 0700. The 
wind continued fresh, and the crew took unusual precautions, stowing all gear 
below and even removing the plant presses from the cabin top. We learned why 
when we cleared the protection of Isla San Esteban, en route to Isla San Lorenzo. 
The Gulf was rough, and spray flew the length of the boat. We changed course 
for Isla Salsipuedes and reduced speed. Arriving at a good anchorage in a cove 
in the southwest side of that island about 1045, we had an early lunch, and were 
ashore before noon. 


Isla Salsipuedes, Isla San Lorenzo Norte (called Isla Partida by the Mexi- 
cans), and Isla San Lorenzo Sur form the eastern side of the Canal de Salsi- 
puedes. They are long narrow islands, the southeastern end of the same ridge 
whose northwestern end is Isla Angel de la Guarda. The shoreline of the islands 
is bold, and there are few places to land. Isla Salsipuedes is about 114 miles long, 
VY, mile wide, and 376 feet high, with several offlying rocks. The island is barren 


and monotonous, and the continuing strong wind made collecting difficult. 


March 24: Isla Salsipuedes, Isla San Lorenzo Norte and Isla San Lorenzo Sur. 
The wind abated during the night. We went to the south end of Isla San 


Lorenzo Norte and landed at the mouth of a steep arroyo. Emerson worked a 
small lower Pliocene sandstone deposit, while the others climbed to the top of 
the island. Banks caught a deer mouse (Peromyscus guardia) by hand, for the 
first record for the island. Pelicans were nesting in the arroyo bottom. Some of 
the eggs made noises, and in other nests the ugly little plum-colored, hiccuping 


1962 } LinpsAY: BELVEDERE EXPEDITION 15 


Fic. 12. Isla San Lorenzo Sur, as seen from Isla San Lorenzo Norte. 


chicks had hatched. The giant hedgehog cactus, endemic to San Esteban and 
the two San Lorenzos, had golden colored spines and rather pink flowers. A 
finback whale passed while we were ashore. Unusually beautiful specimens of 
chalcedony scattered over the top of the island inspired some of the biologists 
to become rockhounds. Vegetation was sparse. 


Returning to the ship at midday, we sailed to the mouth of an arroyo about 
one-third the way down the west side of Isla San Lorenzo Sur. The vegetation 
was much more interesting and included the wild fig tree (Ficus palmeri), 
Galvezia, Bursera, and Pachycereus. There were tremendous deposits of gyp- 
sum, forming cliffs and vertical walls of the narrow canyon in which we worked. 


Returning to the ship at 1630, we sailed to the Salsipuedes anchorage for 
the night. Americans in a sportfishing boat, who had come from Bahia de los 
Angeles, were unable to return because of rising wind and seas and so stayed 
aboard our ship. 


March 25: Isla Salsipuedes to Bahia de los Angeles. 


Our schedule called for a stop at Isla Raza; but since the rough seas pre- 
cluded landing there, the small boats were put aboard and we headed for Bahia 
de los Angeles, where we arrived at 1245. An inspector from the Mexican Fish- 
eries Department was there to see the Vermilion Sea Field Station, and we 


16 SAN Disco Society oF Naturat History { VoL, 13 


needed water and fresh supplies. Lindsay left the expedition for six days, return- 
ing to San Diego by air and rejoining the boat at Mulejé on March 31st. The 
biologists spent the afternoon ashore, boarding after dinner for an early de- 
parture the following morning. 


March 26: Bahia de los Angeles to Isla Raza and Isla Partida. 


The crew were aboard at 0330, the engines started at 0415, and we were 
under way at 0500. We visited Isla Raza first to take advantage of the calm sea, 
because landings are difficult in rough weather. It is a small island, only 34 mile 
long east and west, 1/4, mile wide, and 100 feet high, and is the nesting place of 
thousands of Heermann’s gulls. These trim dark gray birds with white heads and 
red bills occupied the whole island but had not yet started to lay. Commercial 
eggers had come from Santa Rosalia by canoe and were waiting for the birds to 
produce. Royal terns chattered and scolded at the water’s edge, where they were 
confined by the Heermann’s gulls; they would later establish nesting colonies 
near the north end of the island. 


Raza is another island where guano was formerly harvested commercially. 
The flat areas are cleared of rocks, which are stacked in great piles, to facilitate 
guano collecting. 


We departed at 1350 on the one-hour run to Isla Partida. Partida is 114 
miles long and 14 mile wide and consists of two peaks about 400 feet high joined 
by a low narrow isthmus. Anchorage was made in a little cove on the east side. 

Botanical collecting was poor. Emerson worked on a Pliocene deposit on 
the south side of the isthmus, collecting pectens, barnacles, and oysters. Banks 
caught five fish-eating bats (Pizonyx vivesi) under the loose rocks of a talus 
slope. Two of them bit him. These curious mammals apparently feed almost 
exclusively on small fish, which they catch with their long feet and long com- 
pressed claws. Black petrels, too, nest under the talus rocks, and one flew aboard 
at night while we were at anchor. 


March 27: Isla Partida to Santa Rosalia. 


Banks and Harbison were picked up at 0500. Harbison amused and amazed 
the crew by letting a large tarantula, which he found on Isla Partida, walk over 
his arm. The whole day was spent on the run to Santa Rosalia, where the ship 
was moored to the dock at 1930. 


March 28: Santa Rosalia to Isla San Marcos. 


We stopped at Santa Rosalia because of a radio message from San Diego 
that a biologist from the University of Mexico would join the expedition there. 
We stayed in Santa Rosalia until the arrival of the Aeronaves plane; but the 
scientist was not aboard, so we sailed to Isla San Marcos. San Marcos is almost 
6 miles long and 21/4 miles wide; its highest peak is 891 feet. Most of the island 
is volcanic, but there are large lower Pliocene deposits of fossiliferous marine 
sediments and gypsum at the south end. The gypsum beds are about 400 feet 
thick where they are quarried on the southwest side of the island. The gypsum 
apparently accumulated in an ancient saline lake or lagoon. 


11° 


Bir 
FROM |. PARTIDA 
ISLA TORTUGA 


ie 


/ 
4 
/ 


| 
1 
fo 
i! MARCOS 
| 
| 


~~--~--- 


-_ 


Sans Arroyo de los 
Rosalia yChiv 

SS Pe 
ra 
4 


zalene! 


4 
San Lucas 4 


1 
! Cos 
Punta Santa /nes 


San Bruno 
yf ra 
WS /SLAS SANTA INES 


unta Concepcion 


oe 


Puerto Ballandra 


Bahia Marquer 


1Zs 
Fig. 13. Central Gulf islands, the route of the expedition shown by a dash line. 


18 SAN DrieGo Society oF NaAturAL History { VoL. 13 


We landed at 1430 on the northwest coast for 31/4 hours’ collecting. Leaving 
Harbison to camp on the beach, we moved south to the village to anchor for 


the night. 


March 29: Isla San Marcos to San Lucas and return. 


Six biologists were left on the east side of Isla San Marcos, about three 
miles from its northern end, while the ship took Wiggins to the San Lucas air- 
strip to meet a plane from Guaymas. The biologist was not aboard, and the ship 
returned to Isla San Marcos and picked up the shore party in the evening, pro- 
ceeding to the village to anchor for the night. 


March 30: Isla San Marcos to Isla Tortuga and Mulejé. 


We were under way at 0415, landing on the middle of the south side of 
Isla Tortuga at 0800. Tortuga is a barren and precipitous island about two miles 
long and one mile wide, the top of a huge volcanic mountain, composed of 
recent-appearing dark lava, with a crater about 200 feet deep and half a mile 
across. 


Tortuga was very dry, and the botanists had rather poor collecting; but on 
one parched slope Moran was amazed to see lush green moss growing under a 
rock. When he reached to collect a sample, he felt warm moist air, from a small 
fumarole. The island still has central heating. 


A small endemic rattlesnake (Crotalus tortugensis), was fairly abundant, 
and the herpetologists had good collecting. Harbison found many insects, in 
spite of the dry condition. 


Departing from Isla Tortuga at 1430, we went toward the Islas Santa 
Inez, three low barren islands surrounded by shoals and sunken rocks. The sea 
was uncomfortable, the islands offered little protection, the time was late, and 
the biologists were not particularly interested in going ashore. It was decided 
to proceed to Mulejé through the shallow passage between the islands and Punta 
Santa Inez. We anchored at 1945. 


March 31: Mulejé to Puerto Santispaquis, Bahia Concepcion. 


The “San Agustin II” rolled heavily during the night in the exposed 
anchorage at Mulejé. We moved closer to the beach after daylight, and the 
biologists worked ashore from 0800 until 1230. Mr. and Mrs. Lawrence Kuebler 
flew over the ship at 1245, bringing Lindsay from San Diego. Diaz met them 
at the Rancho Loma Linda airstrip when they landed, and brought them aboard 
about 1300. The Kueblers visited the ship for a short time, then went ashore 
loaded with boxes of specimens and mail, which they took back to San Diego 
next day. We cruised to the beautiful Santispaquis Cove on the western shore 
of Bahia Concepcion, anchored, and went ashore in the evening. Harbison spent 
the night ashore, and Soulé hunted reptiles with the gasoline lantern. 


1962 } LINDSAY: BELVEDERE EXPEDITION 19 


Fic. 14. Baby pelicans (Pelecanus occidentalis) on Isla Blanca, Bahia Concepcién. 


April 1: Puerto Santis paquis. 


Wiggins, Banks, and Lindsay worked some of the islets near the anchorage, 
the most interesting of which was Isla Blanca. Pelicans were nesting on the 
ground and in palo-verde trees, some of which bore white stalactites of guano. 
Blotched solitary eggs of oystercatchers lay in very slight depressions in the 
beach gravel. Western gulls also were nesting along the beach, but they did use 
a few twigs and bits of seaweed to house their eggs. 


Everybody but Moran returned to the ship at noon. Two American couples 
who were camped ashore, joined us in a big turtle barbecue. A turtle had been 
split and the halves of the shell propped on either side of a small fire to cook. 
Whole yellowtail and cabrilla were roasted on coals. Clams from a beautiful 
little lagoon were also toasted on coals. The campers brought cocktails of hacha, 
and sabichi, a raw fish cocktail made of sierra mackerel. Cold beer was from our 
cold room. It was a beautiful day and a pleasant party. Moran returned to the 
beach and was picked up just at dark. Many man-of-war birds circled and spiralled 
at great altitude in the evening sky. 


Fic. 15. William K. Emerson collecting fossils from a large Pliocene bed at El 
Pulpito, Baja California. 


1962 } LINDSAY: BELVEDERE EXPEDITION 21 


April 2: Bahia Concepcion to Isla Ildefonso and El Pilpito. 


The engines started at 0330, and an outboard stuttered toward shore to 
pick up Harbison, who had started his lantern to guide it in. The anchor was 
up at 0400, and we had a very smooth passage to Isla Ildefonso, arriving at 0800. 
The island is a little more than one mile long, perhaps half a mile wide, and 
387 feet high. The top is a rather flat tableland. We anchored and landed in a 
cove at the south end. Pelicans had finished their nesting, but there were many 
blue-footed and brown boobies with eggs or chicks. Two large finback whales 
cruised along the island in fairly shallow water. 


Boarding at 1230, we lunched during the six-mile run to Punta Pulpito. 
Anchoring about half a mile south of that great volcanic plug, which is 500 feet 
high, we went ashore for the afternoon. Emerson, Wiggins, and Lindsay col- 
lected fossils from a large Pliocene bed just above the water line. Soulé and 
Banks found two rattlesnakes (Crotalus ruber). Harbison again spent the night 
ashore. 


April 3: El Pulpito to Isla Coronados, Loreto, and Coronados. 


Juan Largo took Banks ashore to pick up his traps, and they brought 
Harbison aboard. We departed at 0630 on the three-hour run to Isla Coronados, 
another volcanic island. It is about 134 miles long, north and south, 11/4 miles 
wide, and 928 feet high. The island is covered with boulders of volcanic rock, 
which make walking difficult; and the crater is a Pleistocene andesite volcano. 
The biologists were put ashore near the base of a sand and rock spit, at the south 
end of Los Coronados, and Wiggins and Moran went to the top. The ship pro- 
ceeded to Loreto. There Mr. Gonzalez left the expedition, to go to the United 
States and visit museums and herbaria in connection with his current study of 
the platyopuntia cacti. We checked with the Captain of the Port for our mail 
and papers and found no word from the biologist. A canoe brought several 
drums of fresh water to the ship. We left at 1445, reaching Los Coronados at 
1545, and anchored for the night. 


Several large scorpions including an undescribed Hadrurus 14 am. long, 
were taken under rocks and rotting nets at deserted shark fishing camps. 


April 4: Isla Coronados to Isla Carmen. 


Isla Carmen is about 18 miles long and 2 miles wide except at the north- 
ern end, where it widens to 5 miles. The taller mountains west of Salinas Bay 
rise to an elevation of 1572 feet. A permanent spring supplies water to a large 
salt works at Salinas Bay. The salt deposit is about 11/4 miles long and 1 mile 
wide. The vegetation of Carmen is heavier than that of the more northern 
islands. 


We anchored in Puerto Ballandra, on the northwest side of Carmen, at 
0800. This is a beautiful circular cove, with a narrow entrance between high 
headlands. There is a small lagoon with mangroves behind the bay, and a wide 
arroyo gives access to the central part of the island. In an interior valley on the 
trail to the salina are a dry well and a stone pumphouse, said to have supplied 


22 SAN Dteco Socrety oF Naturav History Mott 


Fig. 17. Well preserved coral (Porites californica) in the fossil reef shown above. 


1962 | LINDSAY: BELVEDERE EXPEDITION 23 


water for mules when salt was packed to Puerto Ballandra for shipment. Moran 
climbed the peak overlooking the salinas. One of the interesting finds here was 
the dormant bulbs of a zephyr lily (Zephyranthes arenicola), which later 
flowered in San Diego. Moran got it again on Isla Catalina. 


Emerson found a fossil reef with beautifully preserved coral heads at the 
south end of the bay. D’az and one of the crewmen, José Toledo, drove the 
Pak-Jak, a motor-driven field scooter, across the island to the village at the salt 
works, no doubt the first trans-island trip by a wheeled vehicle. In the evening 
a gasoline lantern was suspended over the water, attracting larval fish, marine 
worms, and other organisms. The engineer, impressed with the results, made an 
electrical extension light to hang over the side. 


A pril 5: Puerto Ballandra to Bahia Marquer and return. 


Banks got nine mammals in his 43 traps during the night. Harbison’s insect 
collection was good. After the two were picked up, we cruised to Bahia Marquer. 
This is the type locality of the Marquer formation of an upper Pliocene calcare- 
ous conglomerate containing volcanic pebbles, sandstones, algal limestone, and 
coral reef material. Fossils were abundant throughout the high cliffs but were 


badly leached. 


South of Bahia Marquer at about 50 meters elevation Moran was surprised 
to find a small dry lake with two partially aquatic plants (Eryngium nasturtit- 
folium, Marsilea fournieri) not reported before from any of the islands. 

Shaw and Lindsay went up a narrow arroyo in whose limestone bottom 
were several tinajas with damp sand and some water. Turning over several hun- 
dred rocks they got only two scorpions (Centruroides exilicauda) but collected 
a worm snake (Leptotyphlops sp.) and a black headed snake (Tantilla sp.), 
both new records for the Gulf islands. We returned to Puerto Ballandra for the 
night. 

The new night light was put over, and soon a great quantity of fish, at- 
tracted by the swimming organisms, were darting through. Then a large green 
turtle swam up and was promptly speared by Juan Largo. 


April 6: Puerto Ballandra to Loreto and Puerto Escondido. 


Early in the morning we went to Loreto, where Diaz had arranged for 
diesel fuel to be brought out with small boats. Rough water in the open road- 
stead made loading a tricky operation. Various supplies, including fresh toma- 
toes, a goat, live chickens, and a live turtle, were loaded the same way. Shaw, 
Banks, Harbison, and Soulé took a car to San Telmo, a pool on the road to 
Parras, to collect tree frogs (Hyla regilla) for David Jameson of San Diego 
State College. They were unable to find the frogs but did collect one tadpole. 
The rest had lunch and showers at the Garayzars, mailed letters, and picked up 
mail and ship’s papers from the Captain of the Port; and then we all departed 
for Puerto Escondido. 

Puerto Escondido is a beautiful little harbor, perfectly landlocked. For- 
merly, the bay was a strait between the peninsula and two small islands, but a 


24 SAN Disco Society oF NAtrurRAL History { VoL. 13 


low tombolo has connected the islands and the peninsula at the northern end, 
The very narrow entrance to an inner harbor is kept open by rushing tidal cur- 
rents. Tremendous rugged cliffs of the Sierra Giganta form a spectacular back- 
We anchored south of the entrance of the inner harbor, and Wiggins, 
drop for the bay. 
Moran, and Harbison went ashore in a mangrove area to the east. Moran found 
a specimen of Crotalus ruber lucasensis. Emerson. Soulé, and Lindsay surveyed 
the inner harbor and at its northern end found an unusual chocolate-brown 
scorpion with cream-colored legs. 


April 7: Puerto Escondido to Isla Danzante and return. 


We breakfasted on the short run to Danzante, where we anchored in a 
protected cove on the western side near the north end. Danzante is a rough 
island about 31/4, miles long and less than a mile wide, composed of Comondu 
volcanics. From the air it looks like a dinosaur. There were pelicans nesting at 
the north end of the island. A dead yellow-bellied sea snake (Pelamis platurus) , 
was found on the beach. Moran and Wiggins more than tripled the known 
flora of the island. 


Moran needed the full day for botanical collecting on Danzante, while the 
rest wanted to work the peninsula at Puerto Escondido. Arrangements were 
made for a small boat to pick Moran up at 1630, and the ship returned to Puerto 
Escondido at noon. A native named Ramon Villalejo reported frogs in a tinaja 
called Carrizal, which he stated was but two kilometers distant. He agreed to 
guide Shaw, Soulé, Wiggins, Harbison, Banks, and Lindsay to the tinaja. We 
started in boats which deposited us at a trail to the north end of the inner harbor, 
then we walked over a low pass and down a beach to a grove of date palms called 
Chenque. There Banks shot a red fox which was eating a kid, and Shaw found 
a banded burrowing snake (Chilomeniscus sp.) under a rock. Ramon thought 
it was dangerous and was very concerned when Shaw picked it up. We had al- 
ready hiked much more than two kilometers when we turned inland for the 
longer walk to the base of the Sierra Giganta. We finally reached a beautiful 
canyon and the tinajas, where we found nine specimens of Hyla regilla for 
Jameson. They sang in the plastic bags as we hiked back to the boats in the dusk. 

Diaz had arranged for goats to be barbecued for a party on shore. Mr. 
Pedro Mayu, owner of Puerto Escondido, and his son Pierre had arrived from 
Hermosillo with several Mexican guests. Captain and Mrs. West of the yacht 
“Monsoon IT”, with their guests Mr. and Mrs. Joe Kelly, also joined the party, 


which was very pleasant. 


April 8: Puerto Escondido to Isla Monserrate. 


Departing from Puerto Escondido at 0600, we reached Isla Monserrate an 
hour and a half later and anchored near its southwest corner. Monserrate is 4 
miles long and 2 miles wide, with a highest elevation of 734 feet. Though most 
of the island is composed of volcanic rock, there are Middle Pliocene fossiliferous 
limestone deposits at the beach, and more grey limestone caps the mesa at about 
600 feet elevation. 


1962 | LINDSAY: BELVEDERE EXPEDITION 25 


Fic. 18. George Lindsay with a giant barrel cactus (Ferocactus diguetiu) on Isla 
Santa Catalina. The giant cacti in the background are cardons (Pachycereus 


pringlei). 


Fic. 19. Michael Soulé with one of the rattleless rattlesnakes (Crotalus catalin- 
ensis) of Isla Santa Catalina. 


1962 } LINDSAY: BELVEDERE EXPEDITION 27 


Moran and Harbison were on the island all day, while the rest came aboard 
at noon for lunch. Moran went to the top of the island and to the northern 
edge of the limestone mesa. Wiggins found a spicy-scented succulent composite 
with only one flower per head (Coulterella capitata) ; it was previously known 
only from a limited area near La Paz and from the four southernmost islands. 
Emerson, Wiggins, and Diaz went about halfway up the west coast of the island 
in a small boat and found some fine specimens of Pleistocene sand dollars. 


Vegetation was rather dense in some arroyos and made travel inland dif- 
ficult. Many specimens of Peniocereus johnstonii, a thin-stemmed cactus with 
huge tuberous roots, were found, one with stems nearly ten feet long and a 
tuber that would weigh more than fifty pounds. Another cactus seen was Wil- 
coxia striata, called jaramatraca by Baja Californians, which has dahlia-like tubers 
and stems thinner than a pencil. 


A south wind blew all afternoon, so we anchored off the north end of the 
island for the night. Harbison stayed ashore. Shaw and Soulé worked some sand 
dunes with lanterns after dark. Soulé caught a banded burrowing snake and 
had another in his hand, as well as several joints of cholla cactus. He had tracked 
the snake over the dunes, made a grab for it, and got snake and cholla at the 
same time — but the snake hit the sand, submerged, and “swam” away. 


It was windy during the night. Wiggins and Lindsay were up from time 
to time, securing things on deck so they would not blow overboard. 


April 9: Isla Monserrate to Isla Santa Catalina. 


We arrived at Catalina at 0830 and anchored in a cove about midway on 
the west side of the island. Catalina is a rugged island, about 71 miles long, 2 
miles wide, and 1500 feet high, surrounded by deep water. The bottom at the 
anchorage was deep and steep, so we arranged for everyone to be aboard by 
1600 in order to find a more secure place for the night. Catalina is the type 
locality of a ponderous barrel cactus (Ferocactus digueti1), often more than ten 
feet tall. These yellow-spined giants, mixed with clean-looking cardon cacti 
(Pachycereus pringlei), give the island an unusual appearance. In the arroyos 
near the beach the Burseras, Machaerocereus, and other plants formed dense 
thickets. One could often see half a dozen iguanas (Dipsosaurus catalinensis) 
under a single bush, and the utas (Uta squamata) were blue-green. Moran 
found a rattlesnake and complained that it did not rattle, although it shook its 
tail. It had no rattle and was thought to be an anomaly, but in the afternoon 
Emerson found another, also without a rattle. The snake was Crotalus cata- 
linensis which was described from a single specimen in 1954. All three of the 
known specimens of C. catalinensis lack rattles. A spotted night snake (Hypsi- 
glena) taken under a rock was a new record for the island. 


The biologists returned to the ship at 1615, and we went to the north end 
of the island, where the chart showed a bight and sand beach between two head- 
lands. It was very windy, and there was no bight or protection. Catalina has 
not been surveyed, and the chart is inadequate. We returned to a small cove on 


FROM PUERTO ESCONDIDO 
| “Wy, fl 
Vt ISLA SANTA 
\. CATALINA SAN JOSE (4 


R San Jose 


Apia 
- 


ISLA SAN 
. \ FRANCISCO 
* 


os ee Sor 


LAS AN/MAS , i SEAL 
ROCKS 


Ostiones 


/SLA \ 
BALLENA aah 


Fic. 20. South Gulf islands, the route of the expedition shown by a dash line. 


1962 | LinpsAY: BELVEDERE EXPEDITION 29 


the northwest side where we had fair protection. Banks, Harbison, and Moran 
went ashore. Wiggins and Lindsay fished with Juan Largo in a small boat, 
catching several large yellowtail. Then a fish struck Lindsay’s outfit, taking it 
out of his hands and overboard. The night was uncomfortable, with heavy roll. 


April 10: Isla Catalina 


The ship was moved to a rock-and-sand spit on the northeast side of 
Catalina. A canyon south of the spit contains a small spring and a grove of date 
palms, which were visited by Moran, Harbison, and Soulé. Moran went south 
on the main ridge to the highest point on the island, about 500 meters by alti- 
meter. The east side of the island was sterile compared with the west side, and 
most of the biologists spent the afternoon aboard, working on their collections. 
A cold wind came up at dusk. The small boats were put aboard, and the ship 
moved as close to shore as was deemed safe, in order to secure as much protec- 
tion as possible in the lee of the spit. There was considerable motion during 


the night. 


April 11: Isla Catalina to Isla San José. 


The engines started at 0410, but there was some difficulty in getting the 
anchor unfouled. We departed for Isla San José about 0500. San Jose is 161, 
miles long, 2 to 6 miles wide, and 2000 feet high. It is also composed of Com- 
ondu volcanics overlaid by deposits of marine Pliocene. We anchored on the 
northeast side, at Arroyo de Aguada, at 0930. Emerson found tremendous de- 
posits of middle or early Pliocene pectens and oysters in a remarkable state of 
preservation. The beds were more than 150 feet thick. The gastropods were 
casts, the shells having leached away. 


There was a single date palm in the arroyo bed at Aguada, and a deserted 
goat ranch. Antero Diaz and José Toledo went hunting and saw two deer but 
did not shoot. Emerson worked the fossil deposit and was joined by Wiggins 
in the afternoon. Harbison found a speckled rattlesnake (Crotalus m. mitchellt) , 
which Soulé collected. Scorpions were small and of at least two species. The 
biologists were aboard at 1600 and the ship moved to Los Ostiones, a lagoon en- 
closed by a sand spit on the northwest side of San José, where there is protection 
from north winds. Banks set mammal traps after dinner, using a flashlight, and 
Harbison spent the night on shore. 


April 12: Los Ostiones, Isla San José, to Isla Las Animas and Isla San José. 


Soulé was anxious to sample the herpetofauna on the Baja California penin- 
sula adjacent to San José, so Juan Largo took him to Puerto Nopalo, where he 
found a goat ranch and a Mexican family with many children. The other bi- 
ologists worked the mountains behind Los Ostiones. The country was largely 
granitic but also had fossil deposits. Two rattlesnakes were taken (Crotalus 
ruber lucasensis and C. e. enyo), which with the C. mitchelli collected the day 
before make a total of three species for San José. A spotted night snake was 
collected under a fallen cardon. 


30 San Disco Society oF Naturav History { VoL. 13 


Fic. 21. Whitewashed Isla Las Animas, where cacti predominate. In the fore- 
ground, thick-stemmed Machaerocereus gummosus and smaller, creeping Coche- 
miea poselgeri. Also, Opuntia cholla and A triplex barclayana. 


Antero Diaz and José Toledo went hunting early, returning with a nice 
buck deer at 0900. It was very dark but probably the same subspecies (Odocoi- 
leus hemionus peninsulae) found on the peninsula. The skull and hide were 
preserved by Banks for a specimen. 


We departed for Isla Las Animas at 1015. The sea was calm, and we saw 
many mantas, sharks, yellowtail, trumpetfish, and sierra in the clear water as 
we crossed a shoal area off the north point of San José. 


Las Animas is a barren granite rock a few hundred yards long and perhaps 
150 feet high, with a few offlying rocks and surrounded by very deep water. The 
ship stood off while the biologists went ashore, probably the first “biological 
survey” the islet has experienced. 


We were surprised to find two kinds of lizards, one possibly Urosaurus 
microscutatus and the other Phyllodactylus tuberculosus, and several scorpions 
(Vejovis sp. and Centruroides exilicauda) , were also taken, under rocks. Pelicans 
were nesting; and man-of-war birds wheeled overhead, but none of their nests 
were seen. Only ten kinds of flowering plants were observed, six of which were 
cacti, Pachycereus pringlet, Machaerocereus gummosus, Lemaireocereus thur- 


beri, Cochemiea poselgeri, Opuntia cholla, and Mammillaria albicans. The 


1962 | LINDSAY: BELVEDERE EXPEDITION 31 


others were Atriplex barclayana, Perityle robusta, Portulaca pilosa, and Trian- 
thema portulacastrum. California sea lions inhabited rocks just east of the 
island, and a dead one was floating in the water, surrounded by an oil slick 
fifty yards across, presumably from the decomposing animal. 


Aboard again at 1345. we had lunch en route to the south end of Isla San 
José. There we anchored off the base of the rock spit enclosing the southernmost 
lagoon. Diaz came ashore and shot a ring-tailed cat (Bassariscus astutus) in a 
mangrove thicket. The lagoon had many oysters and large puffer fish. 


April 13: Isla San José to Isla San Francisco and La Paz. 


Wiggins, Moran, Shaw, Soulé, and Lindsay went to Isla San Francisco in 
the 15-foot boat, while the ship waited for Banks to pick up his traps and to 
take Harbison aboard. San Francisco, lying about 414 miles south of San José, 
is an irregular island with an area of about 11/4 square miles and an altitude of 
689 feet. It was very dry, and even the cacti showed drought distress. We found 
emaciated specimens of the southern chuckwalla (Sauromalus ater) and a few 
other reptiles. 

The ship arrived at 0900, and we boarded and proceeded to La Paz, arriving 
at 1600. It was Good Friday afternoon and rather difficult to locate the port 
officials, but in due time we had permits to take fuel and water on Saturday 
morning and received accumulated mail from the Captain of the Port. Dinner 
was at the Perla Hotel to give Rosario, our cook, a well-deserved rest. 


April 14: La Paz 


The ship was watered, fueled, and provisioned. Don Hunsaker II, an Assis- 
tant Professor of Zoology at San Diego State College, Dennis Bostic, a graduate 
student, and Chris Parrish, a museum assistant and student at San Diego State 
College, arrived by plane at 1800, to join the expedition during their Easter 
holiday. 


April 15: La Paz to Isla Cerralvo. 


The engines started at 0400, and we were under way soon after. The yachts 
in the quiet bay and the lights in the backdrop of the town were beautiful. We 
passed several ships in the La Paz and San Lorenzo Channels and arrived at 


Arroyo Aguaje, on the northwest side of Cerralvo, at 0900. 


Isla Cerralvo — the name is also spelled “Ceralbo” — is 16 miles long and 
41/, miles wide. It has several high peaks, one of which, about 7 miles from the 
north end of the island, is 2518 feet high. The island is a single mountain range, 
composed of coarse greenish-white granitic rock, with some metamorphosed 
sediments. Its vegetation is heavier than that of the other Gulf Islands and 
resembles that of the lower areas of the Cape region of Baja California. 

We found two small springs in the arroyo, shaded by giant fig trees (Ficus 
palmeri). The spring water was potable but had a sweet taste. Damsel flies were 
mating and depositing eggs. There were many cardinals and woodpeckers. 


32 SAN Disco Society oF Naturat History {Vornls 


Fic. 22. Richard C. Banks with a live fish-eating bat (Pizonyx vivesi) collected 


on Isla Cayo. 


1962 | LinpsAY: BELVEDERE ExPEDITION 33 


Boarding at 1500, we cruised to the southwest corner of the island and 
anchored just north of a sandspit which gives protection. All went ashore and 
worked sand dunes until dark. 


Soule, Hunsaker, and Bostic were particularly interested in Sator, a genus 
of primitive lizards found only on Cerralvo, Santa Cruz, and San Diego Islands. 
The itinerary of the expedition was adjusted so that Hunsaker and Bostic could 
visit these three islands during their limited time in the field during their spring 
holidays. Several living specimens of Sator grandaevus, which occurs only on 
Cerralvo, were collected for subsequent behavior studies in the laboratory and 
for the San Diego Zoo. Hunsaker and Bostic spent the night on shore, as did 
Harbison. Soule and Parrish also collected by lantern light, returning to the ship 
about 2000. 


April 16: Isla Cerralvo. 


Moran, Wiggins, and Emerson wanted a full day at the arroyo at Rancho 
Ruffo and were taken there in a small boat while the rest of the biologists worked 
the sand dunes and arroyo at the anchorage. Hunsaker and Bostic did not come 
aboard for breakfast but spent the morning collecting sators. The animals showed 
four distinct color phases. One female had a bright orange head. The ship moved 
to Rancho Ruffo at noon. Emerson had worked a fossil deposit in the morning 
and continued in the afternoon, accompanied by Wiggins. Moran came to the 
beach at dark, after an all-day hike to the top of the island. The vegetation was 
dry, but the giant barrel cacti, Ferocactus diguetii, were in bloom. Parrish collected 
scorpions, six species of which were taken, collected lizards, and shot a canyon 
wren for Banks. 


April 17: Isla Cerralvo to Islas San Francisco and San José. 


The crew was up before 0400 and picked up Harbison from the shore, where 
he had spent the night, and we were under way north at 0430. We cruised past 
the east side of Isla Espiritu Santo and Isla Partida. The Gulf was flat and glassy. 
At one place we changed course to follow a pod of about 12 pilot whales. A 
Ridley sea turtle was speared by Juan Largo and the specimen saved because it 
has only rarely been taken in the Gulf. We observed many dolphin fish, manta 
rays, and marlin. 


The short stop at the isthmus of Isla San Francisco on April 13th did not 
provide sufficient time to work that island, and in passing we had observed a 
mangrove lagoon at the north end. Moran suggested that an additional stop be 
made there, so we went ashore from noon until 1500. The vegetation, though 
dry, was in better condition than that at the isthmus. Also the scorpion collecting 
was much better. On the previous stop only one kind was found, but on this 
stop the known fauna was increased to 6 species. Proceeding toward the west 
side of Isla San José, we passed Isla Coyote, with a picturesque group of fisher- 
men’s houses snuggled at the base of rock bluffs, and approached Isla Cayo. This 


34 SAN Disco Society oF NAatrurAL History | VoL. 13 


Fic. 23. Isla Santa Cruz as seen from Isla San Diego. 


is a small rocky ridge which we decided to visit. Moran, Wiggins, Banks, Hun- 
saker, Parrish, Bostic, and Lindsay landed. Four fish-eating bats were taken from 
crevices in the rocks, Hunsaker smoking them out with cigar smoke. Great blue 
herons, black-crowned night herons, brown pelicans, cormorants, and boobies 
were on the islet and offlying rocks. Bostic and Lindsay collected several scorpions 
(Broteas alleni) and Urosaurus and Phyllodactylus lizards. The very short stop 


was interesting and productive. 


We anchored for the night at the salt works on the southwest side of Isla 
San José. Several went ashore and visited the small village and the ponds, where 
the salt is harvested by hand. About 25 laborers work there. Wild goats and deer 
are a principal food in their rather primitive mess. A compact raft of eared grebes 
passed near the boat — perhaps 500 birds in the mass, all diving in unison, then 
popping to the surface. 


April 18: Isla San José to Islas Santa Cruz and San Diego. 


We made another 0430 departure, for Isla Santa Cruz, where we anchored 
off the southwest corner at 0800. Santa Cruz is a steep rugged granitic island, 33/4 
miles long, about 114 miles wide, and 1500 feet high. Bold bluffs 300 to 1000 
feet high make it inaccessible except on the west side, where there are steep arroyos 
and the slopes are about 45 degrees. 


1962 } LinpsAY: BELVEDERE EXPEDITION 35 


The biologists were soon ashore, and Moran found a rattlesnake (Crotalus 
atrox) a few minutes after landing. Parrish collected another almost immediately. 
Three species of lizards were the only reptiles recorded from the islands, so the 
rattlers were of particular interest, but no others were found. The second known 
species of sator lizard (Sator angustus) was very common and tame. One cardon 
cactus was in flower, with a sator stationed beside each blossom, eating the insects 
that it attracted. Using a trout pole and noose, Shaw captured all of the lizards 
on the plant. Then while resting for a few minutes, he observed another crop of 
sators climbing the same cactus to take stations beside the open flowers. They, 
too, were collected. 


A group of approximately 60 harbor porpoises swam north past the anchored 
ship and returned, in scattered bunches, later in the day. One blackfish went 
through express, traveling at about six knots and swimming about 150 feet 
between breaths. 

Moran went up a steep arroyo to the summit of the island. A few plants of 
Ferocactus diguetti were observed. 

We moved to tiny Isla San Diego at 1600. This, too, is a granitic island, 
about one mile long and 722 feet high. We anchored off the southwest shore, 
and had an hour’s collecting before dark. 


it. > ma 
ke a a, » on 


Fic. 24. Don Hunsaker and Chris Parrish with some of the 270 scorpions col- 
lected on the expedition. 


36 SAN Disco Society oF NaturAL History { VoL. 13 


Fic. 25. Music at Banks’s birthday party, off Isla San Francisco. 


April 19: Isla San Diego to Islas Coyote and San Francisco. 


Banks had traps ashore and caught 11 deer mice (Peromyscus sejugis). 
Moran was ashore before breakfast, to have all available time on the island. The 
other biologists were ashore at 0630 and collected until noon. Emerson got a fine 
series of a land snail (Bulimulus chamberlini) previously known only from one 
imperfect specimen. A white globular cactus (Mammillaria albicans) was pat- 
ticularly abundant. Hunsaker and Parrish dove for fish and invertebrates. When 
they were picked up, a group of harbor porpoises accompanied the small boat. 


We departed at noon, cruising down the west side of Isla San José and 
stopping at the fishing village on Isla Coyote to inspect a mechanical device 
which the fishermen had found floating near Las Animas. This was a deep-sea 
free vehicle, with a fish trap attached, which had been released from the California 
Fish and Game vessel “Alaska” in March and not recovered. Its radar reflector, 
buoyancy floats, and trap had puzzled the fishermen. 


Isla Coyote, known locally as Partida, is inhabited by one large family. 
The head of the house has 24 children. The barren islet also produces two kinds 
of lizards, Uta and Phyllodactylus, and two species of scorpions (Broteas alleni 
and Centruroides exilicauda) . 


1962 } LINDSAY: BELVEDERE EXPEDITION 37 


We anchored for the night at the north end of Isla San Francisco. Banks 
went ashore to set mammal traps; and while he was gone, Antero Diaz prepared 
a birthday party for him. Impromptu gifts, doggerel, and songs made an enter- 
taining evening. Antero had sent a boat to Coyote for a guitar, and the crew 
serenaded Banks with appropriate birthday songs. 


April 20: Isla San Francisco to Islas Partida and Espiritu Santo. 


We cruised to the first bay on the northwest side of Isla Partida. Soulé took 
a small boat to Los Islotes, a group of rocks off the north end of the island, and 
found a Urosaurus lizard and a scorpion (Broteas alleni). The other biologists 
went ashore on Isla Partida. 


Isla Partida and Isla Espiritu Santo are separated by a narrow strait which 
can sometimes be navigated with a canoe. Partida is about 31/, miles long, and 
Espiritu Santo is 714 miles long, 2 to 5 miles wide, and nearly 2000 feet high. 
The islands are composed of lavas, mudflows, tuffs, and other eruptive rocks 
of the Comondu age, dipping 8 to 10 degrees westward. There are several deep 
indentations along the west sides of the islands, and there are three offlying islets. 
The eastern sides of the islands are great bluffs of banded rocks. 


We departed at 1500 and cruised south to anchor in a cove at Isla Espiritu 
Santo, just opposite Isla Ballena. All were ashore from 1600 until dark. José 
Toledo shot two of the black jackrabbits (Lepus insularis) that are known only 
from Espiritu Santo. These are large hares with black backs and cinnamon under- 
parts. Soulé shot a Uta and when he retrieved it from a bush he found he had 
also killed a rattlesnake! Wiggins also took a rattler while tearing a fallen cardon 
apart. Both were Crotalus mitchelli mitchelli. Parrish collected many scorpions, 
and surprised himself by shooting a flying bat (Pipistrellus hesperus australis) 
with his 22-caliber dust-shot pistol. Hunsaker and Bostic made an industrious 
assault on the island, tearing up dead cardons, turning boulders, and otherwise 
altering the landscape in their search for reptiles. 


Espiritu Santo was much more interesting than Isla Partida. There was 
heavy vegetation in the arroyo. Harbison stayed ashore all night. Hunsaker 
joined him about 2100, after preparing one of the rabbits for Banks. 


April 21: Isla Espiritu Santo, Isla Ballena, and La Paz. 


Banks found two woodrats (Neotoma lepida vicina) in his traps when he 
went ashore. Moran also went ashore without breakfast, in order to have as much 
time as possible on the island, and climbed to the cliffs at the head of the main 
canyon. Wiggins, Emerson, Hunsaker, Bostic, Parrish, and Lindsay spent two 
hours on Isla Ballena, a small island about 34 mile long, 1/4 mile wide, and 228 
feet high. Agustin Toledo picked the group up at 0930, dropped Hunsaker, 
Bostic, and Parrish at the boat, and took Wiggins, Emerson, and Lindsay to the 
main island for further collecting. All were back aboard at 1145, and we went to 
La Paz, arriving at 1500. 


38 SAN Disco Society oF NaturaL History { Vot. 13 


Hunsaker, Bostic, and Parrish had reservations to fly to Tijuana on April 
23rd. The collecting activity of the expedition was completed, and we tried to 
make reservations for several of the other biologists to return on the same plane, 
but there was no space available. Therefore we radio-phoned Francisco Munoz 
and arranged for his Cessna 195 to meet the ship at Loreto on Wednesday, 
April 25th. Most of the scientific crew spent the night ashore. 


A pril 22: La Paz, Easter Sunday. 


This was a quiet day of rest and recuperation. Mr. Albert E. Schwabacher 
of San Francisco kindly allowed us to use his radio for some necessary traffic 
with the United States and invited several of the scientists aboard his yacht 
“Pez Espada V” for cocktails in the late afternoon. 


A dinner for the biologists at La Perla Hotel celebrated Chris Parrish’s 
birthday and the successful completion of the collecting activities of the expedition. 


April 23: La Paz and north. 


Hunsaker, Bostic, and Parrish departed by plane for Tijuana. It was impos- 
sible to get water and fuel, but we had sufficient supplies of both to reach Bahia 
de los Angeles so departed for the north at 1400. We ran through the night and 
had calm seas. The bioluminescence was spectacular. Flying fish flashed light, 
then great balls of small fish spread like an exploding rocket. Occasional glows 
deep down probably indicated sharks. 


A pril 24: Enroute, La Paz to Bahia de los Angeles. 


Daybreak light at 0430 silhouetted Isla Monserrate and played on Isla 
Danzante and the Sierra Giganta behind Puerto Escondido. We stopped at 
Puerto Chenque for an hour, while Diaz went ashore, and then at Loreto. There 
a telegram was sent to Munoz revising our rendezvous to Santa Rosalia. We 
had a short visit with Miss Annetta Carter and departed for Santa Rosalia 
at 0930. 


The biologists enjoyed a day of relaxation, caught up on their reading and 
specimen preparation, and made preliminary reports of their field results. After 
dark, the bioluminescence was particularly beautiful. We passed to the east of 
Isla San Marcos, picked up the lights of Santa Rosalia, and anchored in its 
harbor at 2200. 


April 25: Santa Rosalia, enroute to Bahia de los Angeles . 


The “San Agustin II” was tied up to the dock for water at daylight. The 
Cessna 195 buzzed the ship at 0600; and Emerson, Soulé, and Banks met the 
plane at the hillside airport and departed for Tijuana. The crew did considerable 
personal shopping at Santa Rosalia, and we finally left for Bahia de los Angeles 
at 1130. There was more chop than we had experienced for several days, and it 
became rough after dinner. Jackets were broken out of sea bags for the first time 
in a month. 


1962 | LinpsAY: BELVEDERE EXPEDITION 39 


Fig. 26. Antero Diaz, owner of the “San Agustin II” and our genial and efficient 
host on the expedition. 


April 26: Bahia de los Angeles. 


We arrived at Bahia de los Angeles at sunrise, 0500, in heavy wind, and 
breakfasted aboard because the chop made it too wet to go ashore. The ship was 
unloaded in a surprisingly short time: all of the collections and gear were on 
the beach by 0800. Carl Hubbs and a party from Scripps Institution of Oceanog- 
raphy were at the Vermilion Sea Field Station, making a survey of the fish fauna 
of the bay. The research vessel “Traveler”, with J. Laurens Barnhard and party 
from the Beaudette Foundation, were engaged in a benthic survey of the same 
area. There was much scientific activity connected with the field station. 


Collections were sorted. Wiggins and Moran loaded the International 
Travelall of the Belvedere Scientific Fund with the dried plant specimens, Banks’s 
birds and mammals, and Harbison’s insects, and departed for the north at 1300. 
The fossils, preserved reptiles, living cacti, and other heavy specimens were 
packed for shipment to Tijuana by truck. Harbison, who had left a jeep at the 
station some months earlier, had it prepared for the return trip. Shaw and 
Lindsay left by plane with Munoz at 1450, arriving at Tijuana at 1700. 


40 SAN Dreco Society oF NaAtuRAL History { Vor 


PRELIMINARY REPORT OF THE SCIENTIFIC COLLECTIONS 


The Belvedere Expedition was a success; and I am confident that, when 
its scientific results are known, it will compare favorably with the “Albatross” 
expedition of 1911 and the “Silvergate” expedition of the California Academy 
of Sciences in 1921. This can be only a preliminary sketch of what was accom- 


plished. 


BoTANY 


Dr. Ira L. Wiggins, Professor of Botany at Stanford University, and Dr. 
Reid Moran, Curator of Botany at the San Diego Natural History Museum, 
were the botanical collectors. Altogether, they made nearly 2000 numbered col- 
lections of vascular plants. Since they collected independently, there were duplica- 
tions; however, each found many plants not seen by the other, and the total 
collection is thus greater. 


Although half a dozen botanists have at one time or another visited one 
or more islands in the Gulf, the only one to visit all of the principal islands 
and to report on the flora of the area was Ivan M. Johnston, who was with the 
California Academy expedition of 1921. Dr. Johnston was an extremely able 
field man and taxonomist and wrote an excellent report. However, he simply 
did not have time to cover some of the larger islands; and furthermore, the trip 
was too early for the summer annuals and too late for many of the spring annuals. 
Therefore, many plants remained to be found. 


Johnston observed that, whereas various of the islands have endemic reptiles 
and mammals, they have few endemic plants. Although several species were first 
described from insular material, most of them have turned up later on the adja- 
cent peninsula. Johnston explained the much lower degree of endemism among 
plants as probably resulting from their greater powers of overwater dispersal, 
with consequent greater likelihood of swamping out of incipient endemism. 
Botanically, one might say that the Gulf islands are not so much islands as 
detached bits of the mainland and peninsula: the flora of each island seems to 
include just about every plant of the adjacent land for which there is a suitable 
habitat. Perhaps for this reason, Johnston, who collected also at many stations 
on the peninsula and mainland and wrote of the flora of the Gulf region as a 
whole, did not emphasize the floras of the individual islands. For various of the 
widespread species, he made no attempt to list each station at which he saw 
each plant. 


On this trip, Drs. Moran and Wiggins placed more emphasis on the floras 
of the individual islands. They carried ashore on each island a list of the plants 
previously reported, and made a special effort to collect those plants not listed. 
As a result, many widespread plants were collected which otherwise might have 
been overlooked on the assumption that they must have been reported before. 
Various unexpected plants were also found, some of them apparently repre- 
senting considerable range extensions. On some islands many species were added 


1962 | LinpsAY: BELVEDERE EXPEDITION 41 


Fic. 27. Charles F. Harbison camped ashore for insect collecting by lantern light. 


to the poorly known flora; an extreme example is Danzante Island, where only 
about 30 species were definitely reported and about 80 additional ones were col- 
lected on our trip. 


The specimens collected on this trip not only amplify the lists of species 
known from the various islands but also give valuable information about the total 
range of species in the Sonoran Desert, whose detailed distribution has been 
poorly known. It is possible that some of the specimens that could not be identi- 
fied in the field represent species as yet undescribed. 


Dr. George Lindsay investigated the cactus flora of the islands in connection 
with a current study of the Baja California representatives of that family. He 
collected many living plants, of which some were later prepared as herbarium 
specimens and others were grown for information about their flowers. 


ENTOMOLOGY 


Mr. Charles F. Harbison, Curator of Entomology of the San Diego Nat- 
ural History Museum, collected about 10,000 arthropods, including insects, 
spiders, ticks, scorpions, pseudoscorpions, and centipedes. Harbison was an 
enthusiastic and energetic collector, turning innumerable rocks, wielding his 
net wherever the spiny vegetation would permit, and camping ashore at night 
to collect by lantern light. 


42 San Dieco Society oF Natura History { VoL. 13 


Many insects known from the Baja California peninsula can now be recorded 
from the Gulf islands for the first time. The insect material, when worked up 
by specialists, will be a valuable contribution to the knowledge of the entomo- 
fauna of the area. 


Chris Parrish, a young research assistant with the San Diego Natural His- 
tory Museum and a student at San Diego State College, was with the expedition 
for eight days. Parrish has a large collection of scorpions from the Baja California 
peninsula. During the expedition some 270 specimens of about 14 species in 7 
genera were collected by or for him on the Gulf islands. These may well prove 
to be of particular interest. 


Brrps AND MAMMALS 


Dr. Richard C. Banks, Curator of Birds and Mammals at the Natural His- 
tory Museum, was in charge of collections of these animals. He prepared 77 bird 
and 98 mammal specimens, ranging in size from hummingbirds to deer. Skulls 
of marine mammals picked up on beaches and saved as locality records are not 
included in the above count. 


There is a high rate of endemism in the mammals of the Gulf islands, 
particularly among the mice and rats. Of the forms taken on this trip, 29 are 
new to the Museum’s collection. Range extensions were established for two 
species. Little information about breeding time, litter size, or habitat selection is 
published for the insular mammals, and many data were accumulated on this 
expedition. 

The distribution of birds on the islands is not well known, so the specimens 
and sight records obtained add much distributional information. Most of the 
species recorded, however, are of widespread distribution. Few insular populations 
of birds are recognized as taxonomically distinct; but four new forms were added 
to the Museum’s collection, all from Tiburon Island. Many data on breeding 
were accumulated, especially for the colonial sea birds. 


The information upon bird distribution obtained on this trip complements 


Dr. Banks’ current detailed study of the birds of Cerralvo Island. 


REPTILES 


There are many species of reptiles on the Gulf islands. The diversity of 
reptiles and the large number of islands combine to furnish the biogeographer 
and evolutionist with examples of the effects of isolation, restriction of area and 
habitat, and presence or absence of predators and of other species of reptiles, 
as well as offering many clues to the history of the herpetofaunas of Baja Cali- 
fornia and western North America. 


Mr. Charles Shaw, Curator of Reptiles of the San Diego Zoo, and Mr. 
Michael Soulé, of Stanford University, were responsible for the reptile collecting, 
but all of the biologists participated. Dr. Hunsaker and Mr. Bostic, of San Diego 


State College, made behavioral studies, in addition to collections for the Museum. 


1962 | LinpsAY: BELVEDERE EXPEDITION 43 


Mr. Soulé took more than 100 serum samples for electrophoretic and immuno- 
logical analysis. 

The large number of specimens from the islands is an important addition 
to the Museum’s reptile collection. Approximately 15 new records for the islands 
were discovered, some of which doubtless represent undescribed taxa. The herpe- 
tologists also collected many live lizards for display in the San Diego Zoo and 
for behavioral studies. 

Dr. Hunsaker and Mr. Bostic were particularly interested in the behavioral 
differentiation of two species of the primitive lizard genus Sator, which occupy 
separate islands and have been isolated probably since Pliocene and possibly 
since Miocene times. They made observations and took motion pictures to show 
the behavior of the two. Also, they took a large sample of living sator lizards 
for laboratory studies. 


INVERTEBRATES 


Dr. William K. Emerson of the American Museum of Natural History 
collected fossil and Recent invertebrates, mostly mollusks. He took fossils, ranging 
in age from early Pliocene to late Pleistocene, at fourteen different localities, 
several of which were previously unknown. The collections include critical mate- 
rial that will establish the precise age of several poorly known deposits. 

Dr. Emerson collected Recent mollusks from the intertidal zone at thirty-five 
localities. This material will greatly aid in establishing the distribution of the 
molluscan fauna within the Gulf. He also collected invertebrates from kitchen 
middens on some of the islands. 

An extensive series of land shells was taken from the islands by Emerson 
and other members of the scientific party. This material is expected to be of 
considerable zoogeographical interest. 


ACKNOWLEDGMENTS 


The Belvedere Scientific Fund of San Francisco provided the principal finan- 
cial support of the Belvedere Expedition to the Gulf of California. Its grant 
paid for ship charter, air transportation, and many incidental expenses. 

The salaries of the scientific personnel were paid by their respective institu- 
tions. Some equipment supplied to the San Diego Natural History Museum by 
the National Science Foundation was used on the expedition. 

The scientific collections were made with the permission of various officers 
and officials of Mexico. The generous cooperation and help of the following 
gentlemen is particularly appreciated. Marine collections were made under permits 
granted by C. Almirante Antonio Vazquez del Mercado, Director General de 
Pesca de Industrias Conexas, de la Secretaria de Industria y Comercio. C. Dr. 
Enrique Beltran, Subsecretario de Agricultura y Ganaderia, personally arranged 
for permits to collect plants and for additional courtesies from his offices. C. Ing. 
Luis Macias Arellano, Director General de Caza de la Secretaria de Agricultura 
y Ganaderia, issued the permits for taking land animals. 


{Wor 13 


San Dreco Soctety oF NATURAL History 


44 


(Pachcereus pringle:) on 


, 


. Emerson with a dwarf card6n 


Fic. 28. William K 


Isla Santa C 


atalina. 


TRANSACTIONS | 


OF THE 


SAN DIEGO SOGIETY OF NATURAL HISTORY 


Vo . 13, No. 2, pp. 45-48, figs. 1-2. Apri 19, 1963 


THE EARLY STAGES OF 
STENASPILATES APAPINARIA DYAR 
(Lepidoptera: Geometridae ) 


BY 


JoHN Apams CoMSTOCK 
Del Mar, California 


The geometrid moth Stenaspilates apapinaria was described by Dyar (1908) 
from four examples taken in San Diego, California. It has been collected spar- 
ingly at light in Del Mar for several years. I have found in the literature no 
reference to the life history or food habits of any member of this genus. 


Eggs obtained from a gravid female on April 30, 1962, hatched on May 8. 
A second batch of eggs hatched on July 4. The young larvae were offered oak, 
Eriogonum, Ceanothus, willow, and honeysuckle. They preferred willow and 
honeysuckle and were successfully reared on these, even though in nature they 
might choose some other plant. The first larva pupated July 23, in a fragile 
cocoon. 


Eacs (fig. 1A) laid on their sides, singly or in lines, oval, 0.6 mm long, 
0.51 mm wide, light gray tinged with blue, the surface appearing finely granular 
but at higher magnification seen to have minute pits surrounded by hexagonal 
walls. 


First INSTAR LARVA (fig. 1B) 3.5 mm long. Head weakly bilobed, nearly 
twice as wide as the body, 0.4 mm wide, white tinged with yellow and spotted on 
the upper part of the cheeks with darker yellow, with a few black dots close to 
the thoracic juncture; ocelli black; setae short, black. Thorax and abdomen white 
with black stripes; thoracic segments tapering towards the abdomen; middorsal 
region with a broad longitudinal band, sometimes discontinuous or lacking on 
the thorax but well defined along the other segments, narrowing at the junctures, 
and with a narrow black stripe lateral to this band, interrupted at segment junc- 
tures; side with a prominent black subspiracular double stripe not visible from 
above; venter white. Legs, anal prolegs, and single additional pair of prolegs all 
concolorous with the body; setae short, inconspicuous. 


46 San Drieco Society oF Natura History {Vor 43 


he 
oi 
HE 
Be 


Fic. 1. Egg and larval instars of Stenaspilates apapinaria Dyar. 


_ A, Egg, lateral aspect, x 40; B, first instar larva, dorsal aspect, X 30; C, intermediate 
instar larva, dorsal aspect, X 10; D, penultimate instar larva, lateral aspect, on leaf of 
honeysuckle, X 3, From a water-color drawing by the author. 


1963 } Comstock: STENASPILATES APAPINARIA 47 


INTERMEDIATE INSTAR LARVA (fig. 1C) sticklike, slender, cylindrical, 13 
mm long. Head strongly bilobed, slightly wider than the body, 1.1 mm wide, with 
dark and narrow but clearly defined sutural lines, cream, heavily spotted with 
brown; sides of cheeks dark gray spotted and streaked with black; ocelli hyaline; 
setae black, arising from small black nodules. Body cream, heavily overlaid with 
longitudinal bands and stripes of brown and black; dorsal surface with a fine 
black midline in a wide band of mottled brown and on each side a narrow stripe 
of cream; side with a suprastigmatal band of mottled brown, then another longi- 
tudinal band of cream gradually merging with the striped and mottled venter; 
ventral surface lighter than the lateral and dorsal; setae numerous, scattered, 
black. Legs dark brown to black on the distal two-thirds, mottled cream and 
brown proximally; prolegs concolorous with the body. 


PENULTIMATE INSTAR LARVA (fig. 1D) similar to the prior instar but with 
very distinct longitudinal lines, the middorsal accented with a black dash at the 
front half of each segment, and with four longitudinal stripes laterally, the lower- 
most stripe in line with the spiracles and extending cnto the upper portion of 


the head. 


MAarTovre LARVA cylindrical, without nodules or excrescences, 43 mm long, 
the head 2.2 mm wide, the average body segment 1.9 mm wide; similar to the 
penultimate instar larva but with most lines and markings somewhat blurred and 
reduced in intensity, thus more uniformly mottled tan; spiracles small, with light 
circlets surrounding the black margins and with minute light central points; setae 
short, concolorous with the body, indistinct. 


Pupa (fig. 2) 16 mm long, 4 mm wide through the center; head well 
rounded; maxillae and antennae extending to the wing margins; surface smooth 
and glistening, the abdominal segments and head brownish yellow, the thorax, 
wing cases, and appendages at first greenish brown, soon becoming darker brown 
than the abdomen, the green disappearing; prothoracic spiracle conspicuous, black, 
protruding, the other spiracles small, narrow, darker than the body; cremaster 
a dark pyramidal protrusion with two minute recurved hooklets at the tip and 
a cluster of smaller hooklets around the base. 


LiTERATURE CITED 


Dyar, Harrison G. 


1908 Descriptions of some new moths from southern California: Stena- 


spilates apapinaria n. sp. Proc. Ent. Soc. Wash. 10:55. 


48 San Dreco Society oF NATURAL History { VoL. 13 


Fic. 2. Pupa of Stenaspilates apapinaria Dyar, x 5. 


A, Dorsal aspect; B, ventral aspect; C, lateral aspect. From a water-color drawing by 
the author. 


wwii a I | 


HARVARD 


UNIVERSIT 
TRANSACTIONS 5 a 


OF THE 


SAN DIEGO SOCIETY OF NATURAL HISTORY 


VoLuME 13, No. 3, pp. 49-60 


BIRDS OF THE BELVEDERE EXPEDITION 
TO THE GULF OF CALIFORNIA 


BY 


RicHarp C. BANKS 


Curator of Birds and Mammals 
San Diego Natural History Museum 


SAN DIEGO, CALIFORNIA 
PRINTED FOR THE SOCIETY 


JUNE 10, 1963 


as 


MUS. COMP. 2001 
LIBRAR} 


JUN 21 1963 


HARVARD 
GRIVERSITY 


BIRDS OF THE BELVEDERE EXPEDITION 
TO THE GULF OF CALIFORNIA 


BY 


RicHarp C. BANKS 


Few of the islands in the Gulf of California have been thoroughly investigated by ornitholo- 
gists. Early collectors in the Cape region of Baja California visited some of the southernmost, 
especially Espiritu Santo and San José islands, and representative series from these localities 
are available. Most islands in the Gulf were visited by the “Albatross” expedition of 1911 
(Townsend, 1923) and by the California Academy of Sciences’ expedition of 1921 (Mailliard, 
1923), but on neither of these voyages was there a major ornithological emphasis. Bird records 
from these trips, as well as from earlier work on the islands, are included in Grinnell’s (1928) 
summary of the ornithology of Baja California. More recently A. J. van Rossem collected on 
several of the islands, but a complete list of birds is available only for Tiburén Island, Sonora 
(van Rossem, 1932). Recent work on Cerralvo Island, Baja California, (Banks, 1963 and in 
press) has emphasized that these insular avifaunas are, in general, very poorly known. 


Further study of the birds of the Gulf of California was made possible by the Belvedere 
Expedition of the San Diego Natural History Museum in March and April, 1962. This voyage, 
sponsored by the Belvedere Scientific Fund, was not primarily ornithological, but there was 
opportunity for considerable collecting and observation of birds. The expedition began at Bahia 
de los Angeles, Baja California, March 15, 1962, and ended at La Paz, April 21. Thirty-two 
islands in the Gulf of California were visited, with several landings on some of the larger ones. 
The expedition also visited seven localities on the peninsula of Baja California and one on the 
coast of Sonora. Lindsay (1962) gave a general account and a detailed itinerary of the expedi- 
tion, with maps of the route followed. For convenience, a list of the islands with the dates of 
our visits is presented here as table 1. 


The present report concerns itself only with observations made and specimens collected on 
the islands and on the waters of the Gulf of California. Little effort has been made to correlate 
the findings of this expedition with the results of previous surveys. Thus, this paper is not a 
complete list of the avifaunas of the islands. Such a list would be premature and could include 
little but an indication of presence or absence of species. On this trip considerable emphasis was 
placed on other aspects of the biology of the insular populations. 


With the exception of representatives presented to the Direccidn General de Caza, Mexico 
City, specimens mentioned in this paper are in the San Diego Natural History Museum. For 
their assistance and contributed observations, I wish to express my appreciation to my com- 
panions on the voyage: Dennis Bostic, William K. Emerson, Ambrosio Gonzalez C., Charles F. 
Harbison, Don Hunsacker II, George E. Lindsay, Reid Moran, Chris Parrish, Charles Shaw, 
Michael Soulé, and Ira L. Wiggins. Similar gratitude is expressed to Antero Diaz and the crew 
of the San Agustin II for their interest and assistance. Comparison of some specimens with those 
in the Dickey collection was made possible through the courtesy of Thomas R. Howell. 


Species ACCOUNTS 
Podiceps caspicus. Eared Grebe—Flocks of Eared Grebes, ranging in number from nine to 


approximately 400, were common throughout the Gulf. Seldom was a single individual seen. 
Specific localities for sightings are Angel de la Guarda, Tiburon, San Pedro Martir, Salsi- 
puedes, San Marcos, Tortuga, San Ildefonso, Monserrate, and San José islands, the dates 
ranging from March 16 to April 17. 


52 San Drieco Society oF NaturAL History { VoL. 13 


TABLE | 
Brief itinerary of the Belvedere Expedition. 

Island Date Island Date 
Angel de la Guarda Mar. 15-17 Coronado Apr. 3-4 
Mejia Mar. 16 Carmen Apr. 4-5 
Pond Mar. 17 Danzante Apr. 7 
Tiburon Mar. 18-20 Monserrate Apr. 8-9 
San Pedro Martir Mar. 21 Santa Catalina Apr. 9-10 
San Esteban Mar. 21-23 San Jose Apr. 11-12 
Salsipuedes Mar. 23 Las Animas Apr. 12 
North San Lorenzo Mar. 24 San Francisco Apr. 13517, 19 
South San Lorenzo Mar. 24 Cayo Apr. 17 
Raza Mar. 26 Coyote Apr. 19 
Partida Mar. 26 Santa Cruz Apr. 18 
San Marcos Mar. 28-29 San Diego Apr. 18-19 
Tortuga Mar. 30 Espiritu Santo, incl. Partida Apr. 20-21 
Islands in Bahia Concepci6n Apr. 1 Cerralvo Apr. 15-17 
San Ildefonso Apr. 2 


Loomelania melania. Black Petrel—This species was observed from San Esteban Island to 
San Marcos Island in the northern part of the Gulf, and near Cerralvo and Espiritu Santo 
islands. It was not noted in the approximately 200 miles between San Marcos Island and the 
southern end of Espiritu Santo. 


The single known breeding locality in the area visited by this expedition is Partida Island 
(Grinnell, 1928). Here, on March 26, three birds were taken from roosting sites under the rocks. 
One bird flew aboard the boat at night, when many could be heard calling and flying about the 
island. There was no direct evidence of breeding although the specimens taken, all males, had 
testes 7 to 9 mm. in length. 


Phaéthon aetherus. Red-billed Tropicbird—Several birds were seen at San Pedro Martir 
Island on March 20. The only other individual seen was near San Francisco Island on April 17; 
this bird, sitting on the water, seemed to be struggling as the boat approached. Just before we 
reached it, the bird swallowed what appeared to be a large fish, and flew. 


Pelecanus occidentalis californicus. Brown Pelican—Pelicans were evident in all parts of 
the Gulf visited except for the vicinity of Tiburén Island. Counts of clutch size were possible 
in three colonies, and are recorded in table 2. 


On San Pedro Martir Island most nests contained hatching eggs or newly hatched young; 
only one half-grown bird was noted. There were small dead fish around the edges of many nests, 
and lizards on the nest rims were feeding both on the fish and on the flies attracted by them. 
Breeding was in a similar stage on North San Lorenzo Island, where the colony consisted of an 
estimated 3000 nests. About one-third of the nests examined contained newly hatched young or 
pipping eggs. 

Asynchrony in nesting activity was first noted on Isla Blanca, a small island in Bahia Con- 
cepcion, on April 1. Large numbers of young nearly ready to fly were clustered around the 
southwest part of the island. A cactus forest on the northwest part of the island held nests in 
all stages of development from eggs to nearly grown young. Clutch counts here were made only 
in nests with eggs. Lindsay reports that pelicans were nesting on this island in January, 1962. 

Nesting appeared to have been completed in a colony on San Ildefonso Island, visited on 
April 2. No active nests were found, but a number of immature birds was flying near the island. 
In a colony of several hundred nests on Danzante Island, on April 7, no eggs or newly hatched 
young were seen. All young observed had the flight feathers well developed, with down still 


covering most of the body. There were two colonies of pelican nests on Las Animas, a small 


1963 | BANKS: BirDS OF BELVEDERE EXPEDITION 53 


TABLE 2 
Clutch sizes of pelicans and boobies. 
Island Date No. of Clutch size 
nests 
examined ] 2 3 

Pelecanus 

occidentalis 
San Pedro Martir Mar. 21 106 6 18 82 
North San Lorenzo Mar. 24 398 24 106 268 
Isla Blanca Apr. 1 42 0 7 35 
Sula nebouxi 
San Pedro Martir Mar. 21 32 8 19 5 
San Ildefonso Apr. 2 4 0 4 0 
Sula leucogaster 
San Ildefonso Apr. 2 6 2 4 0 


islet near San José Island, on April 12. The one colony examined consisted of approximately 
100 nests in stages from eggs to nearly grown young. 

A juvenile bird taken on San Pedro Martir Island on March 21 was preserved in alcohol. 

Sula nebouxti. Blue-footed Booby.—A colony of at least 1000 nests on San Pedro Martir 
Island was visited on March 21. Nesting stages ranged from scrapes without eggs, but to which 
the birds were “attached”, to young nearly ready to fly. Because the half-grown young leave the 
nest at the approach of an intruder, clutch counts were possible only where eggs or newly hatched 
young were present (table 2). On San Ildefonso Island, on April 2, a few Blue-footed Boobies 
were nesting with a large colony of Brown Boobies. Of four nests found, three contained eggs or 
newly hatched young, and one held larger juveniles. 

Two unsexed juveniles were taken on San Pedro Martir Island; one was preserved in alcohol. 

Sula leucogaster. Brown Booby.—Several Brown Boobies were seen near Tiburén Island 
on March 18; three were flying in a line of 25 pelicans. A few were noted at San Pedro Martir 
Island, on March 20, and Wiggins reported a nest of this species there. 

Large numbers of Brown Boobies had nests with eggs on San Ildefonso Island in January, 
1962 (Lindsay, pers. comm.). A few nests were active here on April 2. Only six of the widely 
scattered nests observed held eggs or nearly hatched young (table 2), whereas 18 had half- 
grown birds and five had nearly full-grown young. 

Birds of this species were also seen in the vicinity of Partida, Santa Cruz, San Francisco, 
and Las Animas islands in mid-April. 

Phalacrocorax auritus. Double-crested Cormorant.—This bird was noted near San Marcos 
Island on March 29, and in the southern part of the Gulf near San Diego, Cayo, San Francisco, 
Espiritu Santo, and Cerralvo islands in April. 

Phalacrocorax penicillatus. Brandt Cormorant.—A colony of twenty nests was found on 
the southeast side of Salsipuedes Island on March 23, 1962. Approximately sixty birds were 
observed. There were 13 very young birds, half-grown or less, and 15 larger juveniles, nearly 
ready for flight. 

Van Rossem (1945) reported a breeding colony of Brandt Cormorants on San Pedro 
Martir Island in 1925, but the species is apparently a recent addition to the Gulf waters of 
Baja California. Grinnell (1928) listed only a single record. The only previously reported 
breeding colony on islands associated with Baja California was on Roca Blanca, between Partida 
and Angel de la Guarda islands (Osorio Tafall and del Toro Aviles, 1945). Dr. Carl L. Hubbs 
(pers. comm.) has indicated that the species also breeds on islands in Bahia de los Angeles. 

Fregata magnificens. Magnificent Frigatebird—This bird was seen commonly from San 
Marcos Island to the south. 


54 San Disco Society oF Naturav History | Voiz13 


Ardea herodias. Great Blue Heron.—This species was recorded on Raza, San Marcos, 
Coronado, Carmen, Danzante, Santa Catalina, San José, and Cayo islands. 

Casmerodius albus. Common Egret.—This bird was seen around the mangroves on the 
southern side of San José Island on April 13. 


Leucophoyx thula. Snowy Egret—The Snowy Egret was observed on Coronado Island, 
April 3. 

Aythya affinis. Lesser Scaup.—A group of six males and one female was seen in a bay 
at the northern end of Angel de la Guarda Island on March 16. On the following day a lone 


scaup was seen in the lagoon at Pond Island. 


Melanitta perspicillata. Surf Scoter—Eight birds, four of each sex, were seen at the northern 
end of Angel de la Guarda Island on March 16. 


Mergus serrator. Red-breasted Merganser.—This species was observed in small numbers 


at Angel de la Guarda, Pond, San Esteban, and San Marcos islands. 


Cathartes aura. Turkey Vulture.—In the northern part of the Gulf vultures were observed 
only on Tiburén Island, but they were more common farther south. On San Marcos Island, on 
March 28, over 50 vultures were seen sitting on cliffs near the beach or flying over the island. 
The species was also noted on Coronado, Carmen, Danzante, Monserrate, San José, and Espiritu 
Santo islands. 

Buteo jamaicensis. Red-tailed Hawk.—A lack of large animals to serve as prey is probably 
a limiting factor for large hawks on the Gulf islands. Red-tails were observed only on San 
Esteban, South San Lorenzo, San Marcos, Tortuga, Coronado, and Cerralvo islands. They 
should be expected on the other larger islands as well. 


Pandion haliaetus. Osprey—The Osprey nests commonly on the Gulf islands. On March 
22 a nest was under construction on San Esteban Island; little was done but the base. Three 
nests were found on Salsipuedes Island, within a half-mile of one another. One nest contained 
three young birds, about one-fourth grown. Another had two young about that size but the 
two birds in the third nest were only about a week old. A nest on North San Lorenzo Island 
was not accessible, but one on South San Lorenzo Island held two young about two-thirds grown. 


A nest on a pile of rocks on Raza Island contained two young, approximately one-third 
grown. On San Marcos Island, on March 28, a nest with two quarter-grown young and an 
unhatched egg was attended by parents carrying a fish. A nest with young similarly developed, 
and also with an unhatched egg, was seen on a small island in Bahia Concepcion. Three nests 
were found on San Ildefonso Island. One of these contained two partly grown young and an 
egg, one was unoccupied although adult birds were present, and one which was apparently in 
use could not be examined closely. One nest on Danzante Island was empty, but another con- 
tained three eggs. 

Ospreys were also seen on Angel de la Guarda, Pond, Partida, Tortuga, Coronado, Carmen, 
Monserrate, Santa Catalina, Santa Cruz, San Francisco, and Espiritu Santo islands. Nesting 
probably occurs on most of these islands. 

Falco peregrinus. Peregrine Falcon—Adult peregrines were giving alarm notes near an 
empty nest on Salsipuedes Island on March 23. Bones of at least three Craveri Murrelets (Endo- 
mychura craveri) were picked up at the base of the nest cliff. Peregrines were also observed on 
Tortuga, Danzante, and Santa Cruz islands. 

Lophortyx gambelii pembertoni. Gambel Quail.—Quail were seen only on Tiburén Island, 
where, on March 20, a small covey was found in dense cover. A single male specimen was col- 
lected; its testes measured 7 mm. Quail are not known to occur on any island associated with 
Baja California (Grinnell, 1928). 

Haematopus palliatus. American Oystercatcher.—Oystercatchers are common in the Gulf 
of California; Grinnell (1928) listed records for several islands. Additional localities recorded 
during the period covered by this report are Pond, San Esteban, Raza, and Santa Cruz islands. 
A nest containing a single egg was found on a small island in Bahia Concepcion on April 1. 


A phriza virgata. Surfbird—One bird was seen on Raza Island on March 26. 


1963 } Banks: Birps OF BELVEDERE ExPEDITION aD 


Actitis macularia. Spotted Sandpiper —This is a fairly common bird on the peninsula and 
on the Pacific islands, but records for the islands in the Gulf of California are few. Grinnell 
(1928:95) listed the species from Carmen Island, and it is known from Cerralvo Island (Banks, 
in press). Other localities where Spotted Sandpipers were observed are Pond, San Esteban, 
Santa Catalina, and San Francisco islands. 

Catoptrophorus semipalmatus. Willet—Willets were seen on Pond, San Esteban, and 
San Marcos islands. 

Erolia minutilla. Least Sandpiper—A flock of approximately 10 birds was observed on 
the tidal sand flats on Raza Island on March 26. Least Sandpipers were also seen at a lagoon 
on the west side of San José Island on April 12. A male specimen from Raza Island was slightly 
fat and molting. 


Larus occidentalis. Western Gull—These birds were seen throughout the Gulf and around 
nearly all the islands, but nowhere was there a large concentration. Nests were found on several 
islands, mostly by the crew members. One bird observed at San Ildefonso Island had symmetrical 
white wing patches as described by Hubbs (1954). 

Larus delawarensis. Ring-billed Gull—This gull was common near Tiburén and San 
Esteban islands, and a few were noted near Angel de la Guarda. It was not seen in the southern 
part of the Gulf. 

Larus philadelphia. Bonaparte Gull—Bonaparte Gulls were abundant near Tiburén, San 
Esteban, and San Pedro Martir islands, but were not seen in Baja California waters in this part 
of the Gulf. Farther south they were found at San Marcos Island, and in the late afternoon 
of April 5 large flocks gathered to feed near the west side of Carmen Island; at least 2000 birds 
were congregated there. 

Larus heermanni. Heermann Gull.—Fairly common in the northern part of the Gulf, the 
Heermann Gull was seen near Angel de la Guarda, Pond, Tiburén, San Esteban, Salsipuedes, 
Raza, Partida, San Marcos, Tortuga, San Ildefonso, and Santa Catalina islands. The largest 
concentration was at Raza Island, on March 26, where thousands of adults were on territories 
preparatory to breeding. No eggs were found on this date. Dr. Raymond J. Ryckman informed 
me that hatching had just begun on that island on May 20. Only a few immature birds were 
noted on Raza Island, whereas in flocks of approximately 200 at San Marcos Island, March 
28, and 150 at Mulegé, March 31, most birds were in immature plumage. 

Thalasseus elegans. Elegant Tern—An estimated 1000 terns were noted on Raza Island 
on March 26, but no nesting was yet in progress. Two were seen at San Marcos Island on 
March 28. 

Endomychura craveri. Craveri Murrelet—A male bird incubating two fresh eggs was found 
in a cleft under vine-covered rocks on the steep slope of Pond Island on March 17. The bird, 
which was extremely fat, had paired brood patches. Its left testis measured 12 x 6 mm. but the 
right testis was only about two-thirds that size. Nesting of the Craveri Murrelet on Pond Island 
has not previously been reported. Twenty or more murrelets were seen in the waters around San 
Marcos Island on March 28, and a few birds were seen between Puerto Escondido and Mon- 
serrate Island on April 8. Indirect evidence associates this species with Salsipuedes and San 
Francisco islands (see pp. 54 and 56). 

Zenaida asiatica. White-winged Dove.—A nest was found on Santa Catalina Island on 
April 9, about three feet above the ground in an inner fork of a large, multi-branched cardon. 
The nest held two young birds, estimated to be four or five days old. Nesting on this island 
was thus as much as two months earlier than on Cerralvo Island, 100 miles to the south, in the 
same year (Banks, in press). White-winged Doves were seen also on Tiburén, Carmen, Mon- 
serrate, Santa Cruz, San José, and Cerralvo islands during April. 

Zenaidura macroura. Mourning Dove.—This dove was seen in small numbers on San 
Esteban, Carmen, Santa Catalina, and San Francisco islands. 

Tyto alba. Barn Owl.—Wiggins reported what was probably a Barn Owl on Tortuga Island 
on March 30. Hunsacker and Bostic flushed one from a roost in jagged sea cliffs near the north 
end of San Francisco Island on April 17. The perching area was directly over the water, and 


56 San Dteco Society oF NaturaL History { Vot. 13 


the presence of bones in tide pools indicates that most pellets probably fall into the water and 
are washed away or disintegrate. Some pellet material and several bird wings were found, how- 
ever. Remains of at least six Craveri Murrelets and at least four wood rats (Neotoma lepida, 
presumably of the endemic race abbreviata) were identified. 

Bubo virginianus. Great Horned Owl.—This owl was heard on Carmen Island on April 4. 
The species was not noted elsewhere on this voyage, although it is known to occur on several of 
the larger islands (Grinnell, 1928; Banks, in press) . 

Speotyto cunicularia. Burrowing Owl.—A single bird flushed several times on Coronado 
Island on April 3. On March 16, Lindsay found two pellets presumed to be from this species on 
Angel de la Guarda Island, the only Gulf island for which Grinnell (op.cit.) had records of 
the Burrowing Owl. One of these pellets contained parts of three pocket-mice, Perognathus 
spinatus. 

Phalaenoptilus nuttalli. Poor-will—Poor-wills were flushed on Cerralvo Island on April 16. 

Calypte costae. Costa Hummingbird—This is the most common hummingbird on the 
islands in the Gulf of California, as it apparently is throughout the peninsula. It was recorded, 
during this trip, on the following islands: Angel de la Guarda, Tiburén, San Esteban, South 
San Lorenzo, San Marcos, Tortuga, Coronado, Carmen, Danzante, Monserrate, Santa Catalina, 
Espiritu Santo, and Cerralvo. 

The display of a male Costa Hummingbird was closely observed on Angel de la Guarda 
on March 15. A female was sitting in a bush over which the display occurred. The male dove 
nearly straight down from a height of approximately 40 feet and rose again nearly vertically, 
forming a broad, deep U. At the top of the dive it closed the U and repeated the dive on the 
same course. This display was accompanied by a high-pitched whine which seemed louder near 
the bottom of the descent. This individual performed over 30 dives without a stop. The sound 
accompanying this performance was heard on most of the islands, and was often the first indi- 
cation of the presence of the species. 

Another type of display was noted on Coronado Island on April 3. A female humming- 
bird, apparently attracted to my “squeaking”, hovered within two feet of me, then rested in a 
small tree about six feet away. A male appeared and hovered approximately two feet in front 
of the female, facing her with his gorget fully expanded. His wings made a noise like a loud 
bumblebee; superimposed on this was a whine identical to the sound heard in the diving display. 
After half a minute, during which time the male hovered or moved in small circles, both flew, 
with bills clashing, to another tree, where the female landed. The male once again displayed 
in the same manner. He appeared to be jabbing at her with his bill, but I could not be certain 
of this. After nearly a minute he flew; the female remained on her perch and preened. 

On San Esteban Island a female Costa Hummingbird fluttered and hovered in front of 
me, about 10 feet feet high, with her tail fully spread, for nearly a minute before she perched 
nearby. 

Few data on the breeding of this species were obtained. Females with young birds were 
noted on Tiburén Island on March 20 and on San Esteban Island on March 22. Shaw found 
a nest with two eggs, presumably of this species, on Coronado Island on April 3. A male taken 
on San Marcos Island on March 29 had testes 2 mm. long. 

Female Costa Hummingbirds were noted feeding on Asclepias sp., Ruellia californica, 
and Passiflora sp. The mummy of a bird (sex not determined; not an adult male) was found 
tangled among the sticky branches of a dead plant of the latter on Coronado Island on April 4. 

| Ree xantusi. Xantus Hummingbird. — This bird was noted only on Cerralvo 
Island. 

Megaceryle alcyon. Belted Kingfisher. — Kingfishers were seen at Angel de la Guarda, 
San Ildefonso, and San Francisco islands. 

Centurus uropygialis ssp. Gila Woodpecker. — The Gila Woodpecker was recorded on 
Tiburon, Monserrate, San José, Espiritu Santo, and Cerralvo islands, and on a small island 


in Bahia Concepcién. Its absence from many of the islands probably reflects the lack of suitable 
vegetation. 


1963 } BANKS: BrirDS OF BELVEDERE EXPEDITION 57 


Males from Monserrate and San José islands, taken April 8 and 11, respectively, each 
had 5 mm. testes. These are of the race C. u. brewsteri. A male taken at the north end of 
Tiburon Island on March 18 had testes 7 mm. in length. The tentative referral of this specimen 
to C. u. tiburonensis is purely on geographic grounds. That race was described by van Rossem 
(1942) on the basis of a supposed similarity in paleness to the disjunct C. u. albescens of the 
lower Colorado River Valley. I was not impressed with this similarity when I examined a series 
of tiburonensis in the Dickey collection. The birds from Tiburén are darker around the head 
than are those from the Colorado River region, and show considerable variation in the paleness 
of the back. The specimen presently under consideration agrees well in back color with the 
darkest of the tiburonensis examined. In color of the ear coverts and nape, however, the best 
agreement was with an example of C. u. sulfuriventer from Rosario, Sinaloa. The bird does 
not particularly resemble C. u. uropygialis in any of these respects. The impressions recorded 
here lead me to doubt the validity of C. u. tiburonensts, or at least the validity of its character- 
ization. A more detailed and comprehensive analysis of this population is, I believe, necessary. 


Dendrocopos scalaris ssp. Ladder-backed Woodpecker—This is by far the more common 
of the two woodpeckers found on islands in the Gulf of California. The species was seen on 
Tiburon, San Esteban, South San Lorenzo, San Ildefonso, Carmen, Danzante, Monserrate, 
Santa Catalina, San José, Espiritu Santo, and Cerralvo islands. On April 5, at Carmen Islana, 
Soulé reported a pair mating. A female taken at a nesting hole on Santa Catalina Island, April 
9, had an egg in the oviduct and a brood patch. The birds here were highly territorial. There 
was no sign of breeding, however, in four females secured on Cerralvo Island on April 15. The 
birds on Cerralvo Island had young in the nest in late May of that year (Banks, in press). 


A female taken on Tiburon Island on March 20 is of the race Dendrocopos s. sinaloensis. 
This is the first report of this form of the Ladder-backed Woodpecker on Tiburén, where van 
Rossem (1945) recorded D. s. cactophilus. The females from Cerralvo Island, April 15, are 
referrable to D. s. soulei. A male from Carmen Island, April 4, and a female from Santa Catalina, 
April 9, are both of the southern peninsular form, D. s. lucasanus. 


Myiarchus cinerascens ssp. Ash-throated Flycatcher.—This species, one of the most abundant 
on the islands as well as on the peninsula, was noted on most of the islands, as follows: Angel 
de la Guarda, Tiburén, San Esteban, South San Lorenzo, San Marcos, Tortuga, Coronado, 
Carmen, Danzante, Monserrate, Santa Catalina, San José, Las Animas, Espiritu Santo, and 
Cerralvo. The islands on which Ash-throated Flycatchers were not recorded are the smaller 
ones and those with little or no sizeable vegetation. 


A male specimen taken on Angel de la Guarda Island on March 15 is Myiarchus c. cin- 
erascens; a male from San José Island, April 11, and a female from Cerralvo Island, April 15, 
are M. c. pertinax. 


Corvus corax. Common Raven.—Ravens were seen on most of the islands in the Gulf, the 
exceptions being Angel de la Guarda, Mejia, Raza, Tortuga, Carmen, and Cayo. The lack of 
records for these islands is probably due to oversight. On North San Lorenzo Island I saw a 
raven take an egg from a pelican nest, while pelicans sat quietly on other nests a few feet away. 
The raven poked a hole in the egg and carried it across an arroyo in its bill. 


A nest was found on San IIdefonso Island on April 2. It contained two newly hatched 
young and two eggs. A bird on Danzante Island was carrying something in its bill, but whether 
it was food or nesting material could not be determined. 


Auriparus flaviceps ssp. Verdin. — The Verdin is a common resident on most of the 
islands. It was noted on Angel de la Guarda, Tiburon, San Esteban, South San Lorenzo, San 
Marcos, Tortuga, Coronado, Carmen, Danzante, Monserrate, Santa Catalina, San José, San 
Francisco, Santa Cruz, Espiritu Santo, and Cerralvo islands. Nesting activities were noted on 
San Esteban and Coronado islands. A nest at the former locality held three eggs on March 22. 
On Coronado Island, on April 3, one nest contained three eggs and another was occupied by 
two young birds which were ready to leave. 


58 San Disco Society oF NATuRAL History { VoL. 13 


A female taken on Angel de la Guarda on March 15 and a male from Cerralvo Island, 
April 15, are both Auriparus f. flaviceps. The latter had testes 5 mm. in length, although nesting 
on that island apparently did not begin until late May (Banks, in press). 

Campylorhynchus brunneicapillus. Cactus Wren.—The Cactus Wren was noted on Tiburén 
Island and on all mainland areas visited, but it was not observed on any of the islands associated 
with Baja California, except within Bahia Concepcién. On Tiburon Island, March 20, birds 
were observed in the process of nest building. 

Catherpes mexicanus conspersus. Cahon Wren.—This bird was noted only on Espiritu 
Santo and Cerralvo islands. A female taken on the latter island by Parrish on April 15 was not 
in breeding condition. 

Salpinctes obsoletus obsoletus. Rock Wren—The Rock Wren is a common bird on most 
of the islands, inhabiting even many of the smaller, sparsely vegetated rocks. It was noted during 
the course of this expedition on the following islands: Angel de la Guarda, Pond, Tiburon, San 
Pedro Martir, Salsipuedes, North San Lorenzo, Partida, San Marcos, Tortuga, San Ildefonso, 
Coronado, Danzante, Monserrate, Santa Catalina, San José, and Las Animas. Its apparent 
absence from the other islands visited is probably fortuitous; there are definite records, for 
example, for Espiritu Santo and Cerralvo islands (Grinnell, 1928; Banks, in press) although 


we did not record them there at this time. 


A male taken March 21 on San Pedro Martir Island had testes 7 mm. in length. A female 


and a juvenile male were taken on Salsipuedes Island on March 23. 


Mimus polyglottos. Mockingbird—Mockingbirds were noted on Angel de la Guarda, 
Tiburén, San Esteban, San Pedro Martir, Santa Catalina, San José, and Espiritu Santo islands. 


Toxostoma cinereum. Gray Thrasher.—This species was observed only on San José and 
Cerralvo islands. 

Oreoscoptes montanus. Sage Thrasher—Wiggins and Shaw reported Sage Thrashers on 
Mejia Island on March 16, and one was secured by Shaw. 


Polioptila caerulea obscura. Blue-Gray Gnatcatcher.—Gnatcatchers of this species were 
observed definitely on Coronado, Carmen, Danzante, Monserrate, San José, Espiritu Santo, and 
Cerralvo islands. Females taken on Danzante and Monserrate islands, April 7 and 8, were 
molting dorsally. 


Polioptila melanura ssp. Black-tailed Gnatcatcher. This species was noted on Angel de la 
Guarda, Tiburon, and San José islands. All gnatcatchers seen could not be identified to species 
with certainty. A male taken on Angel de la Guarda on March 16 had testes 4 mm. long; this 
specimen is referrable to P. m. lucida. A male with 3 mm. testes from Tiburén Island, March 
20, is P. m. curtata. 


Anthus spinoletta pacificus. Water Pipit—Wiggins secured a female pipit on Tiburén 
Island on March 18; this was the only record of the species on the islands. 


Phainopepla nitens. Phainopepla—This bird was fairly common on Tiburon Island, but 
was not noted elsewhere on the trip. 


Lanius ludovicianus ssp. Loggerhead Shrike —Shrikes were recorded on Pond, Mejia, North 
San Lorenzo, and Santa Cruz islands. On North San Lorenzo Island a shrike flew into a large 
clump of sour pitahaya (Machaerocereus) with a large grasshopper which it impaled on a spine. 
It flew away immediately but returned soon with another grasshopper; although the shrike 
landed momentarily, it carried the second grasshopper over a nearby ridge. 


An unsexed juvenile, recently out of the nest, was taken by Wiggins on Mejia Island on 
March 16; it is of the race L. |. grinnelli. 


Vermivora celata lutescens. Orange-ctowned Warbler—This is the only species of warbler 
noted on the trip. It was seen on South San Lorenzo, San Marcos, Tortuga, and San IIdefonso 
islands. A bird was seen feeding on Fouquieria on San Marcos Island. An unsexed, slightly 
fat bird was taken on San Ildefonso Island on April 2. 


1963 } BANKs: BrrpsS OF BELVEDERE EXPEDITION 59 


Icterus cucullatus trochiloides. Hooded Oriole.—This species was noted only on San José 
and Cerralvo islands. A female was taken on the former island on April 11, and a male, with 
moderate fat and 3 mm. testes, was secured on Cerralvo Island on April 15. 

Piranga ludoviciana. Western Tanager.—A male tanager collected by Soulé on Santa Cruz 
Island on April 18 was the only one recorded on the trip. This bird was molting and was very 
fat; the testes were 3 mm. in length. 

Richmondena cardinalis ssp. Cardinal.—Cardinals were observed on Tiburon, Coronado, 
Monserrate, Santa Catalina, San José, Espiritu Santo, and Cerralvo islands. A female obtained 
on Tiburon Island on March 20 is of the race Richmondena c. townsendi. Three other speci- 
mens taken are referrable to R. c. ignea; these are a male and a female from Coronado Island, 
April 3, and a female from Santa Catalina Island, April 9. The single male obtained had 
testes 8 mm. long, suggesting that the breeding season was approaching. 

Carpodacus mexicanus ssp. House Finch.—This species was found on most of the islands, 
and must rank as one of the most widely distributed birds in the Gulf of California. Records 
obtained on this expedition are for the following islands: Angel de la Guarda, Tiburon, San 
Pedro Martir, North San Lorenzo, South San Lorenzo, Partida, San Marcos, Tortuga, San 
Ildefonso, Coronado, Carmen, Danzante, Monserrate, Santa Catalina, Santa Cruz, San Diego, 
Espiritu Santo, and Cerralvo. 

On several islands House Finches were observed feeding on the flowers of the cardon. 
Lindsay reported both members of a pair carrying nesting material on Carmen Island on April 


5. Wiggins found a nest on Santa Catalina Island on April 10. 


A female of the race Carpodacus m. frontalis was taken on San Pedro Martir Island on 
March 21. A male with testes 5 mm. long, referrable to C. m. ruberrimus, was obtained on San 
Marcos Island on March 29. 

Chlorura chlorura. Green-tailed Towhee—This bird was noted on Tiburon, San Esteban, 
San Marcos, Coronado, and Carmen islands. 

Calamospiza melanocorys. Lark Bunting —A flock of approximately 25 Lark Buntings was 
seen on San Ildefonso Island on April 2. The single specimen obtained was a molting female 
with moderate fat. 

Passerculus sandwichensis. Savannah Sparrow.—This species was noted only on San IIde- 
fonso Island. 

Amphispiza bilineata ssp. Black-throated Sparrow.—One of the more widely distributed 
species on the islands, the Black-throated Sparrow was noted on Tiburén, San Esteban, San 
Marcos, Tortuga, Coronado, Carmen, Danzante, Monserrate, Santa Catalina, Santa Cruz, 
San José, San Francisco, and Cerralvo islands. Wherever it occurred it was abundant. 

Black-throated Sparrows were observed feeding on cacti of the genera Mammillaria and 
Pachycereus. Birds noted on Fouquieria on Coronado Island were apparently taking small aphids 
from the leaves. 

The pattern of breeding activities in this species is complex. Specimens are available from 
eight islands; at least five subspecies, including four insular endemics, are involved, but breeding 
activity is apparently not correlated with morphologic variation. Evidence from five islands 
indicates breeding, in various stages, whereas the evidence from the other three indicates a lack 
of breeding activity. Extreme differences in gonadal development were noted in populations on 
neighboring islands, such as Monserrate and Santa Catalina. The factors responsible for the 
timing of breeding are difficult to determine in such a situation. There was no evidence that fall 
or winter breeding had taken place on any of the islands visited. 

Two male Black-throated Sparrows from San Esteban Island had testes 8 and 9 mm. in 
length; a female had a brood patch. All three birds, of the endemic race A. 6. cana, were taken 
on March 22. All birds seen here appeared to be paired, and one pair was observed carrying 
nesting material. 

The single specimen available from San Marcos Island, taken March 29, is a female whose 
ova ranged to 5 mm. in diameter. This suggests readiness to breed, although no brood patch was 
noted in this bird. The specimen is referrable to A. 6. bangst. 


60 San Dreco Society oF NATURAL History { VoL. 13 


Testis length of three males of A. 6. tortugae from Tortuga Island varied from 3 to 5 mm. 
on March 30. Three females showed no evidence of breeding readiness. On nearby Coronado 
Island, on April 3, a population of A. b. bangsi was well into breeding. Several young birds, 
recently out of the nest, were seen; the tail of one such bird was only 25 mm. long. The testes 
of one of the two males obtained were 6 mm. long. 

Two males taken on Carmen Island offer conflicting evidence of breeding. One bird, April 
4, had testes 2.5 mm. in length, whereas another, April 5, had testes 7 mm. long and a cloacal 
protuberance. A female taken on April 4 did not show evidence of breeding. These specimens 
are of the race A. b. carmenae. 

Two males from Monserrate Island, April 8, each had testes 3 mm. long. Two females 
taken on the same date were not in breeding condition. This population is referrable to A. b. 
bangsi. A male of the same race from Santa Catalina Island had 10 mm. testes on April 10. 
These islands are less than 15 miles apart. 

No evidence of breeding was found in four specimens of A. b. belvederei taken on Cerralvo 
Island on April 15, although one male had testes 4 mm. in length. Nesting began on this island 
in late May (Banks, in press). 

Spizella passerina. Chipping Sparrow.—A Chipping Sparrow was seen on North San 
Lorenzo Island on March 24. 

Lonotrichia leucophrys. White-crowned Sparrow.—White-crowned Sparrows were seen on 
Angel de la Guarda, Partida, San Ildefonso, and Coronado islands. Large flocks were noted in 


each instance. 


LITERATURE CITED 


Banks, R. C. 
1963. Three new birds from Cerralvo Island, Baja California, Mexico. Occ. Pap. Calif. 
Acad. Sci. No. 37, 5 pp. 


GRINNELL, J. 
1928. A distributional summation of the ornithology of Lower California. Univ. Calif. 
Publ. Zool. 32:1-300. 


Husss, C. L. 
1954. Western gull, with symmetrical wing patches, resembling aberrant Heermann gulls. 
Condor 56:228. 


Linpsay, G. E. 
1962. The Belvedere expedition to the Gulf of California. Trans. San Diego Soc. Nat. 
Hist. 13: 1-44. 


MAILLIARD, J. 
1923. Expedition of the California Academy of Sciences to the Gulf of California in 1921. 
The birds. Proc. Calif. Acad. Sci., 4th ser., 12:443-456. 


Osorio TAFALL, B. F., and M. pEL Toro AvILEs 
1945. Notas sobre la distribucién de Phalacrocorax penicillatus (Brandt) en el Golfo de 
Cortes y la costa occidental de Baja California. Rev. Soc. Mex. Hist. Nat. 6:85-93. 
TOWNSEND, C. H. 
1923. Birds collected in Lower California. Bull. Amer. Mus. Nat. Hist. 48:1-26. 
VAN Rossen, A. J. 
1932. The avifauna of Tiburon Island, Sonora, Mexico, with descriptions of four new 
races. Trans. San Diego Soc. Nat. Hist. 7:110-150. 
1942. Four new woodpeckers from the western United States and Mexico. Condor 
44-22-26. 
1945. A distributional survey of the birds of Sonora, Mexico. Occ. Pap. Mus. Zool., La. 
State Univ. No. 21, 379 pp. 


YUL LE 150 


HARVARD 
UNIVERSITY 


TRANSACTIONS 


OF THE 


SAN DIEGO SOCIETY OF NATURAL HISTORY 


VoLuME 13, No. 4, pp. 61-72 


A SECOND NEW SPECIES OF MEGATHYMID FROM 
BAJA CALIFORNIA, MEXICO 


(Lepidoptera: Megathymidae) 


BY 


CHARLES F. HarBISON 


Curator of Entomology 
San Diego Society of Natural History 


SAN DIEGO, CALIFORNIA 
PRINTED FOR THE SOCIETY 


JUNE 30, 1963 


“Te are 


ad “ ae : oe See s ~~ © 
— g 9 es he ig a elk: =" 
a te a ec ES 
Fig. 1. Agave goldmaniana Trelease as it grows 17.7 miles north of Punta Prieta, Baja 
California, Mexico. 


_ 


ce, 


, 


7 _. 
r 2 
~ 


ae. ween eh. OE eK 
Serene — ~ im fas 
fF ee a ea ee 
Fig. 2. Agave goldmaniana 29 miles north of Punta Prieta. A smaller species of agave grows 


on the hillside in the background. Mr. Fred T. Thorne is standing by the plant. 


A SECOND NEW SPECIES OF MEGATHYMID FROM 
BAJA CALIFORNIA, MEXICO 
(Lepidoptera: Megathymidae) 


lane 


CHARLES F. HARBISON 


On October 8, 1946, while collecting plants north of Punta Prieta (28°58’N, 114°17/W), 
Baja California, Mexico, Dr. E. Yale Dawson observed a flight of megathymids associated with 
Agave goldmaniana Trelease (Dawson, 1948). Dr. Dawson succeeded in netting eight of these 
rapidly flying insects, seven males and one female, for the collection of the Allan Hancock 
Foundation of the University of Southern California. In 1956 Mr. Lloyd M. Martin of the 
Los Angeles County Museum rediscovered the material collected by Dr. Dawson and thought 
that these specimens were possibly additional examples of the new species of the Baja California 
megathymid that I was describing at that time (Harbison, 1957). Therefore, through Dr. John 
Adams Comstock, arrangements were made with Dr. John S. Garth of the University of Southern 
California for the loan of the Dawson material to me for study. Examination of the macula- 
tions of the insects and of prepared microscope slides of the genitalia proved that they were not 
Agathymus comstocki (Harbison) but were probably another new species. 


Because the eight specimens of this megathymid were not a long enough series on which 
to base the description of a new species, a trip was made back to the original Dawson locality 
to obtain more material in October, 1956, with Arthur A. Lee. Despite diligent work for two 
days only eight additional specimens were captured — still a very short series. In October, 1961, 
I succeeded in netting forty-five megathymids during three days of collecting near that locality. 
When the entire series of sixty-one specimens was studied, a great amount of variation was 
found in maculations and genitalia. A male holotype was chosen to represent the new species, 
but when an allotype had to be selected there was no way of knowing which female could most 
appropriately be chosen. On October 7, 8, and 9, 1962, Fred T. Thorne and I revisited the 
region. We were especially anxious to obtain mating pairs, and Mr. Thorne was successful in 
capturing three pairs in coitus. 


Food plants in which larvae are found and reared are often important in descriptions of 
megathymids (dos Passos, 1960). Because the one live pupa that I collected in October, 1961, 
did not emerge, it was not possible to be certain that these insects were living in Agave gold- 
maniana Trelease. In order to answer the host plant problem a late summer trip was made to 
the vicinity of Punta Prieta, August 30 to September 8, 1962 (fig. 1). No adult megathymids 
were seen during the nine days in the vicinity. Living larvae and pupae were found in the leaves 
of the large species of agave, Agave goldmaniana (fig. 2). A smaller gray-leaved agave grow- 
ing on the hillsides above the area occupied by the Goldman Century Plant was also examined 
for infestation. No living immature megathymids were found in these plants, but a few leaves 
showed that the smaller agave is at times infested by a megathymid. Nineteen leaves of A. 
goldmaniana infested with living larva or pupa were collected. Four of the immature insects 
had died in their cells before the material was turned over to Dr. John A. Comstock, who had 
agreed to try to raise to adult stage any megathymid material brought back. Of 15 immature 
insects submitted to Dr. Comstock, only three males and two females emerged. 


For a short time on the morning of October 9, 1962, I searched for infested leaves. One 
insect emerged immediately after the leaf was removed from the host plant. It was allowed to 
expand and develop rigidity before being killed. A second emerging adult was accidentally 


killed as it tried to come out of its cell in a leaf. The specimen was preserved and sent to Dr. 


64 SAN Disco Society oF Naturav History {VoL. 13 


Charles L. Remmington of Yale University for a possible chromosome count. A third specimen 
successfully emerged next morning in our camp. Three infested leaves were brought back to 
Dr. Comstock; from these, two adult megathymids emerged on October 14 and 15. One pupa 
was parasitized. 


COMMENTS ON PROCEDURE 


An illustration of the typical megathymid venation was given in a previous paper (Harbi- 
son, 1957). The veins were named there as on Plate III in the revision of the Megathymidae 
by Barnes and McDunnough (1912). The first vein of the secondary is the subcosta and the 
second vein is the radius. To bring this plate up to date and into agreement with the Comstock- 
Needham system as revised by Snodgrass, change Sc to Sc + R, and change R to Rs (radial 
sector) . 


Although several recent authors have chosen females as holotypes (Freeman, 1960; Stall- 
ings, Turner and Stallings, 1961), I prefer to use a male for four reasons: 1. Males are more 
uniform than females in maculations and genitalia; 2. Male genitalia are less subject to distor- 
tion; 3. More males than females are usually collected and reared; 4. Evans (1955) illustrated 
only male genitalia, making comparisons easier. I do believe that both sexes should be equally 
well described and illustrated so that any specimen coming to hand can be definitely determined. 


Evans (op. cit.) believed that the genitalia should be studied in place on the insect or re- 
moved and mounted dry on cards. Many megathymid experts keep the genitalia in vials of ethyl 
alcohol to which a few drops of glycerine have been added. My laboratory procedure has been 
described previously (Harbison, 1957). I believe that the concavity slides which I use for male 
genitalia do not distort the genitalia and are much more safe to handle than cards or vials. The 
completed slide should be placed flat in a small unit tray beside the unit containing the rest of 
the insect; the same data are on each. 


I believe, contrary to the opinion of most megathymid experts, that a study of the genitalia 
of the holotype should be made. The holotype is not as beautiful a specimen after the removal 
of the genitalia, but, how is a student to know what the genitalia of that specimen is like by 
studying the genitalia of another individual? Genitalia do vary from specimen to specimen. I 
believe that as the study of the Megathymidae expands, and more species and subspecies are 
found, it will be necessary for students to see the genitalia of all holotypes. 


Agathymus dawsoni, new species 


The type series consisted of the male holotype, the female allotype and 87 male and 29 ad- 
ditional female paratypes for a total of 118 specimens (table 1). Slides of the genitalia of 111 
specimens were prepared. 


The holotype and allotype are in the type collection of the San Diego Society of Natural 
History. Paratypes will be placed in the following collections: Allan Hancock Foundation, 
University of Southern California, ¢ ¢ 1 to 7, 2 2 (original Dawson material) and ¢ 4 
14, 46; American Museum of Natural History, ¢ 4 15, 48; British Museum (Natural His- 
tory), ¢ $ 16, 49; California Academy of Sciences, $ ¢ 17, 50; Carnegie Museum, ¢ ¢ 
98, 59; Los Angeles County Museum, ¢ ¢ 18, 52; Museum of Comparative Zoology, Har- 
vard University, ¢ ¢ 20, 53; National University of Mexico, Institute of Biology, ¢ ¢ 21, 
94; Philadelphia Academy of Sciences, ¢ ¢ 22, 55; United States National Museum, ¢ ¢ 
24, 56; Yale University, ¢ ¢ 25,57; Burns, John M., 3 $ 28, 68; Comstock, John A., 


3 6 84, 85, 86 and 2 2 28, 29, 30 (raised through from agave leaf); Freeman, H. Avery, 
6 4 30,69 and 2 24 (latter courtesy of Thorne); Nicolay, S.S., ¢ $ 31, 70; Roever, Kilian, 
8 6 36,71 and 2 19 (latter courtesy of Thorne); Stallings, Don B. and J. R. Turner, 
6 4 38,72 and 2 20 (latter courtesy of Thorne); Thorne, Fred T., $ 3 76, 77, 78 and 
2 @ 21, 22, 23 (collected by Thorne); Tilden, -j..W.. “6 6 40) 735 Tinkham, (EWR: 
38 6 45, 74. The remainder of the paratypes are in the collection of San Diego Society of 
Natural History. Female topotypes will be distributed as they become available. 


1963] 


Locality 


17.7 mi. N 


Punta Prieta 


20 mi. N 


Punta Prieta 


7 atowl-9 
mi. N 


Punta Prieta 


24 mi. N 


Punta Prieta 


20 mi. N 


Punta Prieta 


17.7 to 20 mi. N 


Punta Prieta 


Harsison: A New MEGATHYMID 


Data on the Type Series of A gathymus dawsoni 


Collector 


Harbison 


Dawson 


Lee 


Harbison 


Harbison 


Thorne 
Harbison 


Harbison 


Thorne 


Harbison 


*Pairs collected in copulation. 


TABLE 1 


Date 


Oct. 4, 1961 


Oct. 8, 1946 


@Oet..15,, 1956 
Oct. 16, 1956 


Oct. 16, 1956 


Oct. 3, 1961 
Oct. 4, 1961 


Oct. 5,196! 


Oct. 7, 1962 
Oct. 7, 1962 


Oct. 8, 1962 


Oct. 8, 1962 


Sept. 2-5 and 
Oct. 9, 1962 


Type Designation 
or Paratype No. 


Holotype 


Allotype (= 2 1) 


O 10 &8 O3 4O O 40 40 40 O3 
oOo Noo on 


+0 


Oo 8 O&O +10 BF +40 O +0 


40 O3 


$ 1-7 


Os +0 
Co A 
oO wa 


29,10 


$ 37-58 
2 11-15 
$ 
2 


59-78 
16-24 
79, 2 253 
80, 2 261 
81, e378 


$ 82-87* 
2 28-30° 


*Collected as larvae or pupae and raised to adult stage; see table 2. 


Males 


12 


11 


22 


20 


65 


Females 


66 San Dreco Society oF NATURAL History {Vot. 13 


TABLE 2 


Larvae and Pupae of A gathymus dawsoni Raised to Adult Stages in 1962 


Paratype No. Collected Emerged Raised by 
$ 82 Oct. 9 Ort Harbison 
g 83 Oct9 Oct. 10 2? 
é 84 Sept. 3 Oct AZ Comstock 
& 85 Sept. 4 Oct. 20 ? 
3 86 Oct-93 Oct. 15 a 
3 87 Sept. Nov. 11 i 
O27 Sept. 2 Sept. 24 "5 
2 28 @Oces? Oct. 14 2 
2 29 Sept. 3 Oct. 30 4 


HototyPeE MALE 
Upper Surface 


Primaries. — Most of this surface is covered with dark grayish-brown appressed scales, be- 
coming more brownish toward the outer margin of the wing (fig. 3a). The light maculations 
are tan to yellowish white. From the base of the wing in the area between A, and Cuz to ap- 
proximately half way to the lower spot of the discal band there are numerous tan scales. Admixed 
with these are long hair-like scales of the same color. The cell spot is small and oval. The sub- 
apical spots are four in number. The upper one has its inner margin much nearer to the base of 
the wing than the other three. Subapical spot 2, between R, and R,, and spot 4 have their inner 
edges about the same distance from the base of the wing; the edge of spot 3 is slightly nearer the 
base. The extra-discal (submarginal of Stallings and Turner) spots vary as to shape and size in 
the fore wings of the holotype. In the left wing the upper one, between M, and Ms, is about three 
times larger than the one below. In the other primary the spots are of about equal size. The 
discal band in the holotype is composed of two aligned spots and the usual middle spot between 
Cu, and Cu, is lacking completely. Spot 1 is roughly rectangular, being about twice as long as 
broad. Its inner side lines up with the inner margin of subapical spot 4. Spot 3 of the discal band 
is small and roundish. There seems to be no tendency for this to be double. The marginal fringe 
is made up of long gray and white scales. The gray scales extend out from the margins of the 
wings where the veins reach the edge thus making a dark and light fringe. 


Secondaries. — The grayish-brown, flattened, saw-toothed scales of the secondaries are 
almost hidden under long, brownish-white hair-like scales similar to those covering the dorsum 
of the thorax but not of the same degree of whiteness. These extend out to the band of light 
maculations but not to the edge of the wing. Basally, in the area between A, and A,, the hair- 
like scales are very long and of the same light color as those covering the upper surface of the 
thorax. There is no spot in the area between Sc + R, and Rs. The outer edge of the maculation 
between Rs and M, is almost connected to the inner edge of the spot below. The outer edges of 
the next four spots are in a straight line. The spots between M, and M;, C, and A, are the 
larger. The scales forming the fringes of the secondaries are mostly white, but are dark where 
the veins reach the edge of the wing, as in the primaries. 


Under Surface 


Primaries. — The base of the wing is covered with dark brownish scales to the band of 
maculations. Beyond the maculations there is an admixture of whitish and darkish scales to the 
edge of the wing. The discal band shows faint indications of the middle discal spot. 


1963 | HarsBison: A New MescaATHYMID 67 


Fig. 3. Upper (left) and lower (right) surfaces of adults of A gathymus dawsoni. a. Holo- 
type male; b. Allotype female; c. Paratype male no. 80; d. Paratype female no. 26. These para- 
types were collected in copulation. 


68 San Dieco Society oF NAtTurAL History {VoL. 13 


Secondaries. — The same spots appear as are on the upper surface of the wing but they are 
less distinct and have a frosted appearance because the dark areas have an admixture of flattened 
whitish scales. A spot between Sc + R, and Rs which did not show on the upper surface can be 
seen basad to the spot between Rs and M,. Also, a light indistinct spot is basad to this spot by 
about its own width. Several indistinct spots between A, and M, basad to the outer band by its 
width, almost form a paralleling light band. 


Genitalia 


The terminology used for the genitalia is that of Barnes and McDunnough (1912) and 
Harbison (1957). Nothing distinctive was noted in segment IX of the abdomen. Viewed 
from beneath, the uncus is relatively short, extremely broad and heavily chitinized at the tip. 
Its tip is more than half the breadth between the wide part of the uncus, about two-thirds of the 
way to its base. The gnathos are roundly and evenly curved upward. The valva is three times 
longer than broad. Three characteristics are distinctive in the new species (fig. 4b): 1. The 
cucullus of the blade of the valva tends to be truncate with numerous sharp teeth along the up- 
turned edge; 2. Sharp teeth also extend back along the cucullus to a position occupied by the 
proharpe; 3. The latter structure is heavily spined and slightly bent rostad, with little tendency 
to a branch at the base. 


Other Characters 


The antenna segments are covered with blackish scales for the basal five-sixths of each 
segment. The club is light below and dark at the tip. The labial palpi are densely covered with 
whitish, up-right scales. The legs are brown covered with whitish scales. The total expanse of 
the wings is 44 mm.; male paratypes vary from 37 to 48 mm. 


ALLOTYPE FEMALE 


Upper Surface 


Primaries. — The general scalation is much as in the holotype, but the light maculations 
are much more extensive (fig. 3b). The cell spot is much larger than in the male and tends to 
be squarish with slight emarginations on the outer and inner sides. It completely crosses the 
cell from the front at R, to the upper inner angle of maculation 1 of the discal band. There are 
four subapical spots which are arranged as in the holotype. The extra-discal spots are unequal in 
size, the second being the larger and nearer to the base of the wing. The inner lower angle of the 
latter is removed from the upper outer angle of the upper spot of the discal band (discal spot 1) 
by about the width of the extradiscal spot. The discal band is of three spots. Spot 1 is slightly 
wider than spot 2 and its inner lower angle forms an obtuse angle with Cu,. The outer edge of 
the spot is angled outward for about one-quarter of its length, then inward to vein Cu,. The 
front edge is wider than the rear. Spot 2 of the discal band along vein Cu, is the same width as 
the contiguous rear edge of spot 1. This spot is slightly longer than spot 1 and reaches Cus. Spot 
3 is roughly an isosceles triangle but the apex, which would point inward, is replaced by a notch. 
This spot does not extend completely from vein Cu, to A». 


Secondaries. — The maculations are much as in the holotype and are not markedly more 
extensive. 
Under Surface 

Primaries. — Discal spots 1 and 2 are about the same width basad to distad and are wider 


than spot 3. Spot 1 touches vein M, and Cu, for its entire width and its inner margin is at right 
angles with the latter vein. The outer margin of this spot is bowed outward. The inner edge of spot 
2 of the discal band is bowed inward; the outer edge is almost perpendicular to vein Cuy. This 
spot touches both veins Cu, and Cup. Spot 3 of the discal band extends almost to vein A.; the 
front edge almost touches Cu, and is shorter than the rear edge. 


1963 } Harsison: A New MEGATHYMID 69 


Fig. 4. Genitalia and pupal cremasters. a. Valva of A. dawsoni male paratype no. 1, show- 
ing extreme of truncateness; b. Valva of holotype of A. dawsoni; c. Valva of holotype of A. 
comstocki; d. Uncus and segment IX of holotype of A. dawsoni; e. Genitalia of allotype of 
A. dawsoni; £. Genitalia of a paratype of A. comstocki: g. Pupal cremasters (left to right) of 
A. dawsoni paratypes 75, from above, ¢ 74, from below, 2 20 and 2 21, both from 
above. 


70 San Disco Society oF NATURAL History {Vot. 13 


Secondaries. — The maculations are much more distinct than in the holotype. The con- 
trast between light and dark areas is very plain, with the light areas appearing as spots rather 
than bands. The spot between Sc+ R, and Rs, basad to the spot between Rs and M,, is smaller 
than in the male and is much more noticeable because of the predominance of dark scales in the 
areas basad and distad. It is removed from the outer spot by twice its own width. 


Genitalia 


The lateral fold on the left edge of the vaginal plate (genital plate of some authors) is 
quite unusual in that the pointed posterior end makes a 90 degree turn and ends beside a fold 
that extends downward from the ala (fig. 4e). It is quite different from the lateral fold on the 
other edge, which is normal. Because of this abnormality in the allotype an accurate measure- 
ment cannot be made transversely across the plate to compare with the medial measurement 
from just below the ostium to the anterior margin in the emargination between the convergent 
inner ends of the lateral fold. 


Other Characters 


The antenna is very similar to that of the holotype male. It tends to be less ringed with 
white apical scales, the black basad scaling covering most of each segment. The club is more 
uniform in color and is dark. The legs are similar to those of the male. The total expanse of 
the wings of the allotype is 48 mm.; female paratypes vary from 44 to 53 mm. 


PARATYPE SERIES 


A male and female paratype collected in copulation by Mr. Thorne in October, 1962, are 
shown in figure 3, c and d. The male has the middle spot of the discal band present, smaller than 
the spot above and about the size of the spot below; the first spot of the discal band is much 
larger than the other two spots. In the holotype, spot 1 is about equal to spot 3, and spot 2 is 
lacking. The maculations on the secondaries are not as clearly defined as in the holotype. The 
genitalia shows the extreme truncateness of the cucullus of the valva that is shown by one of 
Dawson’s paratypes, illustrated as figure 4a. Also, the cucullus of the valva is shorter and 
thicker than in the holotype or illustrated paratype. The proharpe varies considerably in the 
paratype series, but has no tendency to fork at the base. 


The maculations of the female taken in copulation (fig. 3d) are very much like those of the 
allotype. The genitalia are similar although the emargination at the anterior edge of the vaginal 
plate is not so distinct. There is considerable variation in the type series. 


COMPARISON WITH CONGENERIC SPECIES 


In males of Agathymus dawsoni all spots of the discal band are round. Spot 2 tends to be 
absent, but if present it is smaller than 1 and equal to 3. Spot 3 is single. Spot 3 is double in 
comstocki. In comstocki, stephensi, remingtoni, and fieldi, spots 1 and 2 are equal in size. Spot 1 
of remingtoni is triangular rather than round, and all spots of aryxna are squarish. Spots of 
estelleae and fieldi are all larger than in dawsoni. 


In the female of A. dawsoni, spot 3 of the discal band is roughly triangular with the inner 
apex notched, and it is smaller than spots 1 and 2. This spot is Z-shaped in comstocki, nearly 
thomboidal in stephensi, irregular in remingtoni and pentagonal in fieldi. Spot 3 is larger than 
spots 1 and 2 in remingtoni, estelleae, and mariae, and equal in size to 1 and 2 in field. 


The male genitalia of dawsoni differ from all species for which comparative material is 
available in the truncateness of the cucullus of the valva. The female genitalia have an emargi- 
nation at the anterior edge of the plate which separates this species from all species but mariae. 


1963 } HarsBison: A New MEGATHYMID 71 


ACKNOWLEDGEMENTS 


Thanks are due to Dr. E. Yale Dawson for taking time out from his botanical collecting 
to net the first eight specimens; the new species is named in honor of this outstanding botanist 
and algologist. Thanks are also extended to Mr. Lloyd M. Martin of the Los Angeles County 
Museum, who called my attention to Dawson’s material in the Allan Hancock Foundation’s 
collection. I am grateful to Dr. John S. Garth of the Foundation who loaned me this valuable 
series and to Dr. John A. Comstock who brought the specimens to my laboratory and who 
raised some additional material from immature stages. On two of the trips to the type locality 
I had the pleasurable companionship of two of my best friends, who collected a vital part of 
the type series of the new insect. I deeply appreciate their help on these field trips; Mr. Arthur 
A. Lee and Mr. Fred T. Thorne have my lasting gratitude. 


Mr. T. W. Pace took the photographs of the insects and of the genitalia. Dr. Reid Moran 
read the first draft of this paper and made many valuable suggestions and Dr. Richard C. Banks 
read the finished manuscript. Dr. George E. Lindsay, Director of the San Diego Natural History 
Museum, made it possible for me to be away from the museum on the numerous field trips. 
To Dr. John A. Comstock and to Mr. Kilian Roever goes my special appreciation for their as- 
sistance and continued encouragement. The latter gentleman supplied me with much needed 
comparative material from Texas, New Mexico and Arizona. 


LITERATURE CITED 


Barnes, WILLIAM, AND JAMES H. McDUNNOUGH 
1912. Revision of the Megathymidae. Cont. Nat. Hist. Lepid. North Amer. Vol 1, no. 
3. 43 pp., 6 pls. 
Dawson, E. YALE 
1948. A naturalist’s diary on the Mexican west coast. Cact. Succ. Jour. 20:161-165. 


pos Passos, Cyrit F. 
1960. Taxonomic notes on some nearctic Rhopalocera. 1. Hesperioidea. Jour. Lepid. Soc. 


14:24-36. 


Evans, W. H. 
1955. A catalogue of the American Hesperiidae. Part IV. London. 500 pp., pl. 54-88. 


FREEMAN, H. A. 
1960. Notes on Agathymus in Texas, and the description of a new species from Mexico 
(Megathymidae) . Jour. Lepid. Soc. 14:58-62. 
Harpsison, C. F. 


1957. A new species of Megathymus from Baja California, Mexico (Lepidoptera: Mega- 
thymidae). Trans. San Diego Soc. Nat. Hist. 12:231-262. 


STALLINGS, Don B., J. R. TURNER, AND VIOLA N. STALLINGS 


1961. A new subspecies of Agathymus mariae from Mexico (Megathymidae). Jour. 
Lepid. Soc. 15:19-22. 


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TRANSACTIONS 
OF THE 


SAN DIEGO SOCIETY OF NATURAL HISTORY 


VoLuME 13, No. 5, pp. 73-80, figs. 1-2 


A NEW INSULAR SUBSPECIES OF THE 
SPECKLED RATTLESNAKE 


BY 


L. M. KLAUBER 


Honorary Curator of Reptiles, 
San Diego Society of Natural History 
8 


SAN DIEGO, CALIFORNIA 
PRINTED FOR THE SOCIETY 


SEPTEMBER 27, 1963 


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A NEW INSULAR SUBSPECIES OF THE | | badd LCRS 
SPECKLED RATTLESNAKE 


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BY 
L. M. KLAuBER 


The recent establishment by the San Diego Society of Natural History of a field station 
at Bahia de los Angeles, on the west shore of the Gulf of California, has greatly facilitated 
collecting in central Baja California and on adjacent islands. This has been particularly true of 
Isla Angel de la Guarda, one of the largest of the Gulf islands, which lies about 20 miles offshore 
from the field station. 


It has long been known that the speckled rattlesnakes, Crotalus mitchelli, found on the 
island reach a large size— larger than in any mainland section of the extensive range of the 
species in Baja California and the southwestern United States. Although the presence of this 
snake on the island has been known for more than 80 years (Streets 1877), until recently only 
six specimens had been available — not enough to judge the validity of any differences from the 
mainland rattlers. Now an additional 15 specimens are at hand, besides an unborn brood of 
eight in which some characters may be ascertained. Not only is the impressive size difference 
still in evidence. but divergences of squamation and pattern are also apparent. Although no 
invariable key character has been detected, the differences from the mainland subspecies are 
sufhciently consistent and significant to warrant the description of the island form. My reasons 
for placing such insular forms in subspecific categories, despite the virtual impossibility of gene 


flow, have already been presented (Klauber 1956, p. 28). Therefore I name the form 


Crotalus mitchelli angelensis subsp. nov. 
ANGEL DE LA GUARDA ISLAND SPECKLED RATTLESNAKE. 


1877. Crotalus pyrrhus, Streets, Bull. U.S. Nat. Mus. 7:39. 

1882. Crotalus pyrrhus (part), Yarrow, Bull. U.S. Nat. Mus. 24:73. 

1883. Crotalus confluentus mitchellii (part) , Garman, Mem. Mus. Comp. Zool. 8 (3) :173. 

1887. Crotalus mitcheli (part), Belding, West Am. Sci. 3 (24) :98. 

1889. Crotalus pyrrhus, Townsend, Proc. U.S. Nat. Mus. 13 (800) :143. 

1891. Crotalus pyrrhus (part), Stejneger, West Am. Sci. 7 (59) :165. 

1891. Crotalus pyrrhus (part) , Cope, Proc. U.S. Nat. Mus. 14 (882) :694. 

1894. Crotalus mitchellii (part) , Van Denburgh, Proc. Calif. Acad. Sci. ser. 2, 4:450. 

1896. Crotalus mitchelli (part) , Boulenger, Cat. Snakes Brit. Mus. 3:580. 

1929. Crotalus tigris mitchellii (part) , Amaral, Bull. Antivenin Inst. Am. 2:82. 

1930. Crotalus confluentus mitchellii (part), Klauber, Trans. San Diego Soc. Nat. Hist. 6 
(3) :108, 128. 

1936. Crotalus mitchellii pyrrhus (part), Klauber, Trans. San Diego Soc. Nat. Hist. 8 
(19) :157; (20) :191. 

1952. Crotalus mitchelli pyrrhus (part), Klauber, Bull. Zool. Soc. San Diego 26:117. 

1956. Crotalus mitchelli pyrrhus (part) , Klauber, Rattlesnakes 1:38. 


Type SpEcIMEN. — No. 51994 in the collection of the San Diego Society of Natural 
History. Collected about 4 miles southeast of Refugio Bay, at 1500 feet elevation, Isla Angel 
de la Guarda, Gulf of California, Mexico (near 29° 2914’ N, 113° 33’ W), March 22, 1963, 
by Dr. Reid Moran. 


DIFFERENTIAL DraGnosis.— A subspecies of Crotalus mitchelli differing from the other 
subspecies in its larger size, proportionately larger head in adults, smaller initial rattles, greater 
number of dorsal scale rows, higher number of ventral scutes, greater frequency of supraoculars 
with sutures, larger and fewer scales between the prenasals and rostral, and higher number of 
middorsal blotches on the body. In nearly all of these characters there is some overlapping with 
the mainland subspecies, but the differences are statistically significant. 


76 San Disco Society oF Natura History {VoL.73 


DEscrIPTION OF THE Horotype. — An adult male. Length overall, as measured before 
shrinkage in preservative, 1331 mm.; tail length 100 mm. or 714 per cent; length of head (some- 
what damaged in killing the snake) 62 mm., contained 211/ times in the body length overall. 


The scale rows number 25-27-21, with 15 at the middle of the tail. All dorsals are keeled, 
except the row next the ventrals on each side. Paired apical scale pits are evident on the posterior 
dorsals. The ventrals number 185 and the subcaudals 27. The anal is entire. Ten irregular scales 
border the rattle, which comprises an incomplete string of seven segments; the dorso-ventral 
width of the proximal lobe is 15 mm. 


The supralabials number 14-12, and the infralabials 14-15. The last supralabials are some- 
what longer than the others. The first infralabials are undivided; there are neither intergenials 
nor submentals. The rostral is triangular and is wider than high. As is usual in C. mitchelli sub- 
species, the scales on the snout and crown are so subdivided as to make it difficult to determine 
to which categories some should be assigned. There is no line of demarcation between the scales 
of the prefrontal and frontal areas; there are about 26 scales in the prefrontal section, and five 
scales comprise the minimum intersupraocular bridge. 


As is generally the case in C. mitchelli subspecies (except stephensi), the prenasals do not 
touch the rostral; however, the separation is not produced by granules but by a single scale on 
each side having approximately the same height as the prenasal. Besides these scales, the first 
supralabials and a pair of small internasals contact the rostral. The prefoveals number 8-7. 


Each supraocular is sutured by short seams at the outer edge. The scales anterior to the 
orbit on each side cannot be as readily classified as in most rattlesnakes; it may be said either 
that the normal upper preocular is divided both horizontally and vertically, and that there are 
two loreals on each side, or that there are three preoculars and four loreals. The usual crescentic 
lower preoculars bordering the pit above almost fail to reach the orbit. There are three rows of 
scales between the supralabials and orbit. The anterior scales on the crown are somewhat knobby; 
the posterior ones are flat and keeled. 


The head is pink above, with ill-defined and irregular fawn blotches. No lateral head stripes 
are evident. The underside of the head is tan. Dorsally, the body pattern comprises a series of 
36 somewhat ill-defined hexagonal, fawn or vinaceous blotches on a pink ground color, with 
a secondary series of blotches on either side. Posteriorly, the dorsal blotches engage the secondaries 
and become cross stripes. The dorsum is liberally sprinkled with darker dots, especially within 
the blotches; these punctations are characteristic of C. mitchelli subspecies (except stephens?) 
and give these snakes the name of speckled rattlesnakes. The scales comprising the anterior row 
bounding the dorsal blotches are often tipped on their posterior ends with dark brown or black. 


The ventrum is pinkish-cinnamon, especially centrally, with extensive irregular dark-brown 
blotching along the outer edges of the ventrals. There are six tail rings, the first four fawn, the 
last black; all are wider and darker dorsally. The rattle matrix is black. The tail is mottled below. 


DESCRIPTION OF THE SUBSPECIES. — The paratypic series includes the following specimens: 
SDSNH_ 19717-8, 19990-5, 44358, 51991-3, 51995-6; CAS 50869-70, 50904; USNM 8562, 
15978, 64588.* Together with the holotype, whose statistics are included in the following sum- 
mary, there are 12 males and 9 females. There is also an unborn brood of 8 (SDSNH 44401-8), 
which are fully formed (except one defective) and with patterns already clearly in evidence. To 
avoid the effects of familial uniformity, I have not used the statistics of these juveniles except 
in blotch counts. 


*The USNM catalogue entry indicates uncertainty as to the origin of this specimen with the statement “doubt- 
less from Cedros Island’. But Townsend’s field notes indicate that 64588 came from Angel de la Guarda. 
Furthermore, extensive collecting on Cedros during the past 50 years has failed to produce another specimen 
of C. mitchelli, although many specimens of C. exsul have been found there. 


1963 } KLAuBER: New INSULAR RATTLESNAKE 77 


In presenting statistics and data on the paratypic series, and therefore of the new subspecies 
as now known, I shall at the same time point out the nature and extent of the differences that 
have been observed between C. m. angelensis and the other subspecies of C. mitchelli. 


The new island form is certainly the largest of the C. mitchelli subspecies. Of the 21 speci- 
mens now available (excluding the brood), 5 reach or exceed 1240 mm. The longest is a male 
of 1367 mm. Of the two largest mainland subspecies, m. mitchelli and m. pyrrhus, only 9 out of 
250 adults that I have examined measure 1000 mm. or more; the longest of these, a male from 
Escondido, San Diego County, is 1114 mm. In appearance, the new subspecies is even more 
striking than indicated by length alone: a 1367 mm. snake is nearly twice as bulky or heavy as 
one 1114 mm. long. 


In C. m. angelensis the adult tail proportion of males averages 7.7 per cent, of females 5.7 
per cent. The male proportion falls between those of m. mitchelli and m. pyrrhus; the female 
seems shorter than either (see Klauber 1956, table 4:3, p. 158). The new subspecies has a 
considerably larger head, proportionately, than m. mitchelli and slightly larger than that of 
m. pyrrbus (Klauber 1956, table 4:2, p. 154). The following figures represent the number of 
times the head is contained in the overall body length of a large male: m. mitchelli 27.0, pyrrhus, 
23.0, angelensis 22.6. But at a body length of 1000 mm., which would be a fully grown pyrrhus 
but a still growing angelensis, the respective figures would be 23.0 and 21.0. This fact tends to 
validate a prior conclusion concerning head proportionalities in related groups (Klauber 1938, 


p- 20). 


The rattle of C. m. angelensis in its early segments is peculiarly small. Crotalus m. mitchelli, 
although a rattlesnake of only moderate size, has the largest rattle button of any rattlesnake, 
even exceeding the buttons of such giants as C. adamanteus and C. atrox. Fortunately, I have 
available 5 specimens of angelensis with buttons; of these the dorso-ventral widths are 5.1 to 
5.3 mm., mean 5.2 mm. The corresponding means of other C. mitchelli subspecies are: mitchelli 
8.2, pyrrhus 6.7, muertensis 5.8, stephensi 5.1 (Klauber 1956, table 5:6, p. 289). The suc- 
ceeding average increment in rattle width for each of the next 3 or 4 segments in angelensis is 
1.1 mm., compared with 1.5 in m. mitchelli, 1.1 in pyrrhus, and 0.9 in muertensis. Thus angelensis 
never attains the rattle size of either mitchelli or pyrrhus until the snake has reached a body length 
which the mainland subspecies never attain. 


The fangs of angelensis are relatively shorter than in the other subspecies; their length is 
contained in the head length about 7.6 times and in the length overall about 170 times (for the 
other subspecies see Klauber 1956, table 11:1, p. 736). 


Data on the meristic characters of the paratypic series of angelensis are as follows: midbody 
scale rows 25 or 27 (6 out of 21 have 25 rows); ventrals (males) 180 to 187, mean 184.8 
(females) 186 to 190, mean 188.2; subcaudals (males) 23 to 28, mean 25.4 (females) 19 to 21, 
mean 20.1; supralabials 12 to 15, mean 13.8; infralabials 13 to 17, mean 14.9; body blotches 
36 to 46, mean 41.0; tail rings (males) 6 to 8, mean 6.9 (females) 4 to 6, mean 5.0. Comparisons 
with the other C. mitchelli subspecies may readily be made by consulting Klauber 1956, table 
2:7, p. 124. Crotalus m. angelensis differs from the others to an important degree in four of 
these characters, as indicated in the following table: 


Scale rows Ventrals (mean) Body blotches 
(mode) Males Females (mean) 
C. m. angelensis 27 184.3 188.2 41.0 
C. m. mitchelli 25 176.3 178.9 32.3 
C. m. pyrrbus 25 178.0 178.7 33.3 
C. m. muertensis 23 17. 178.3 59.7. 
C. m. stephensi 23 174.3 179.0 36.8 


78 San Disco Society oF NATurRAL History { VoL. 13 


Crotalus m. angelensis has the peculiarity in head scales that distinguishes C. mitchelli 
subspecies (except stephens:) from all other rattlesnakes, namely the almost universal separation 
of the rostral from the prenasals by other scales. This feature is found to an appreciable extent 
(about 17 per cent) in only one other form — Crotalus viridis cerberus, the black rattlesnake of 
Arizona. But C. m. angelensis differs from m. mitchelli and m. pyrrhus in usually having the 
interposed scale between each prenasal and the rostral in the form of a single scale of substantially 
the same height as the prenasal, whereas in the others the interposed scales are usually smaller. 
Prefoveals are more often in contact with the rostral in the two mainland forms, and the inter- 
nasals are generally smaller. These differences are best indicated by the ranges and means of the 
total scales in contact with the rostral, as shown in the following statistics: 


Scales in contact with the rostral 


Range Mean 
C. m. angelensis 6— 10 6.9 
C. m. mitchelli 7—11 9.5 
C. m. pyrrbus >—11 7.7 
C. m. muertensis 6— 10 8.4 


Probably the most consistent key character wherewith to segregate angelensis from either 
m. mitchelli or pyrrhus is the presence of sutures or other blemishes in the supraoculars. These 
are extensively present in stephensi and muertensis, but we need not rely on the supraocular 
sutures to key out these two: the prenasal-rostral contact in stephensi and its absence in an gelensis, 
will segregate stephensi; and scale rows and body size will differentiate muertensis. But supra- 
ocular sutures are important in distinguishing the other two subspecies; for they occur in only 
3 per cent of m. mitchelli, about 10 per cent of pyrrhus, and 95 per cent of angelensis. They are 
not so obvious in angelensis as in stephensi; nevertheless they are nearly always present, particu- 
larly as seams indenting or parallel with the outer edges of the supraoculars. 


In color and pattern, C. mitchelli is the most variable of all rattlesnakes, with the single 
exception of C. viridis, which is found throughout a much larger and ecologically more variable 
area. Some local color phases of C. m. pyrrhus are particularly striking: the burnt-orange colored 
specimens of the Santa Ana mountains of southern California, the coral pink or red snakes of 
some parts of central Arizona, and the cream-colored snakes of the Tinajas Altas range, a 
southerly continuation of the Gila Mountains in extreme southern Arizona. 


Crotalus m. angelensis shows considerable uniformity in color but no special brilliance. It 
begins life as a gray snake with dark gray hexagonal blotches, and then changes gradually through 
tan or buff with brown blotches to an eventual pink with russet blotches when fully adult. The 
blotches do not have the even outlines or light borders characteristic of so many rattlers; they 
are produced by a darker ground color with increased stippling. There is a smaller, secondary 
series of blotches on either side; with these the posterior dorsal blotches merge to form cross- 
bands. In almost all specimens, some anterior scales of the blotches are tipped posteriorly with 
dark brown or black; these are more prevalent in angelensis than in other mitchelli subspecies. 


The ventrum in adults is pinkish-cinnamon, especially midventrally, with irregular brown 
blotching on the outer edges of the ventral scutes. 


The head is colored similarly to the body. The darker blotches are variable and indistinct. 
Ventrally, the head is cream colored. The side stripes characteristic of the heads of most rattle- 
snakes, if present at all, are quite inconspicuous. Very young snakes have a light supraocular 
crossdash widening inwardly, but this disappears at an early age. 


The anterior tail rings are of the same color as the body and are largely composed of puncta- 
tions. The last two or three rings are black. The rings narrow laterally and become less distinct 
ventrally. The rattle matrix is black. 


1963 | KLAUBER: New INSULAR RATTLESNAKE 79 


Fic. 2. Allan Kuebler and Chris Parrish with Angel de la Guarda Island Speckled Rattlesnakes 
which they collected at Arroyo Estatén, Angel de la Guarda Island, April 22, 1960. Photograph 
by George Lindsay. 


80 San Disco Society oF NATURAL History {VoL. 13 


In the more important characters that distinguish pyrrhus from m. mitchelli, such as head 
size, rattle width, and the nature of the separation between the rostral and prenasal, angelensis 
is shown to be more closely allied to pyrrhus than to m. mitchelli, just as would be expected from 
territorial considerations, for Angel de la Guarda lies off the part of the peninsula occupied by 
pyrrhus. A rather frequent occurrence of elongated final supralabials in angelensis is the only 
character suggesting a closer relationship with m. mitchelli than with pyrrhus. 


The occurrence of the largest subspecies of C. mitchelli on an island is contrary to the usual 
relationships between insular forms of Crotalus and their nearest mainland relatives; for the 
insular forms generally are smaller. In each of the following pairs, for example, the insular form 
(named first) is smaller: C. tortugensis and C. atrox, C. exsul and C. ruber ruber, C. catalinensis 
and C. scutulatus scutulatus, C. molossus estebanensis and C. m. molossus, C. unicolor and 
C. durissus terrificus, C. viridis caliginis and C. v. helleri, and C. mitchelli muertensis and C. m. 
pyrrhus. (Not enough specimens of C. enyo cerralvensis are available to determine its size rela- 
tionship with C. e. enyo.) Of all the pairs mentioned above, the size relationship of C. m. muer- 
tensis to C. m. pyrrhus is most in contrast with that of C. m. angelensis to C. m. pyrrhus: whereas 
angelensis is larger than pyrrhus, muertensis is a stunted form about 60 per cent of the length of 
pyrrhus. Yet muertensis also occurs on an island (El Muerto) off Baja California, and only 
about 75 miles from Angel de la Guarda. 


Why C. m. angelensis should reverse the usual trend of island forms in rattlesnakes (and 
other genera as well — Klauber 1956, p: 304) is not known. Angel de la Guarda is a large island, 
with a length of 48 miles and a maximum width of 12 miles. Ecologically, it is favorable to 
rattlesnakes, with good cover and a plentiful food supply, of both small mammals and lizards. 
Yet C. ruber ruber, also found on the island, though not so plentiful as angelensis, does not 
reach so large a size as ruber on the adjacent peninsula, although it is not conspicuously stunted. 


It should not be assumed, from the number of island subspecies mentioned, that subspecific 
differentiation is a normal or invariable result of island colonization by rattlesnakes. On the 
contrary, in the Gulf of California alone, besides the pairs already mentioned, no less than 7 
species of rattlers are found on 14 other islands without subspecific differentiation having become 
evident, at least so far as can be determined from the available material. Among these there are 


5 other islands inhabited by C. m. mitchell. 


LITERATURE CITED 
KLAUBER, LAURENCE M. 


1938 A statistical study of the rattlesnakes: V. Head dimensions. Occ. Pap. San Diego 
Soc. Nat. Hist. 4: 1-53. 


1956 Rattlesnakes: their habits, life histories, and influence on mankind. 1: i-xxix, 1-708; 
2: i-xvii, 709-1476. Univ. Calif. Press. 


STREETS, THOMAS H. 


1877 Contributions to the natural history of the Hawaiian and Fanning Islands and 
Lower California. Bull. U. S. Nat. Mus. 7: 1-172. 


TRANSACTIONS 
OF THE UNIVER 


SAN, DIEGO SOCIETY OF NATURAL HISTORY 


VoLuME 13, No. 6, pp. 81-84 SEPTEMBER 27, 1963 


TWO NEW OPISTHOBRANCH MOLLUSKS 
FROM BAJA CALIFORNIA 


BY 


WesLey M. FarMerR 


Curator of Exhibits 
San Diego Natural History Museum 


Very little work has been done on nudibranchs in waters associated with Baja California, 
a fact mentioned by Marcus (1961:57). Recent work has shown that the waters of the Gulf of 
California and of the Pacific coast of Baja California support a high population of these animals. 
Among the most interesting finds are two very similar appearing members of the family Dorididae 
which are newly described below. 


Collections have been made possible through a variety of circumstances. Dr. Kenneth Norris, 
whom I accompanied on a trip to the Sebastian Vizcaino Bay area in 1953, provided time to 
collect nudibranchs. I accompanied Dr. Richard C. Banks to Cerralvo Island in June, 1962, 
and collected marine organisms in that area; this was made possible with the aid of his grant 
from the National Science Foundation and the cooperation of the San Diego Natural History 
Museum. John Sloan provided specimens from Cedros Island (WMF 129) and Angel de la 
Guardia Island (WMF 250). Mrs. Fay Wolfson donated a specimen from Gonzaga Bay 
(WMEF 112). Clinton Collier gave me a specimen from Gonzaga Bay (\WMF 303). 


I wish to thank Mrs. Rosemarie Fiebig for translation of several German references. 


I follow Odhner (1957) in using the name Chromodoris Alder and Hancock 1855 for 
glossodorid species with hamate teeth. 


Chromodoris norrisi, spec. nov. 


Type. — The type was collected by the author (WME 49) at Cerralvo Island (24° 10’ N, 
109° 52’ W), Baja California, Mexico, on May 26, 1962. It is deposited in the Invertebrate 
Type Series of the California Academy of Sciences, San Francisco, California, where it is 
registered as IZ No. 4. With two of the paratypes (IZ Nos. 5 and 6), it will be associated 
with the Frank Mace MacFarland Memorial Collection of Opisthobranchs. 


Description. — The body is white with a faint tinge of cobalt violet on the notum between 
the rhinophores and gills (plate -la). This band of violet, apparently due to the violet liver, 
is present only on some specimens. The notum is edged with cadmium orange dashes of varying 
lengths. The surface of the notum in larger animals is covered with as many as 200 deep cobalt 
violet dots, and a few light red ones, of varying size. Approximately 100 cadmium yellow and 
light yellow dots of greater and slightly varying diameter are intermingled with the violet dots. 
The number of dots is considerably less in smaller animals. The side of the body has a few 
violet dots; the underside of the notum is plain, except around the head in some specimens. The 


82 SAN DieGo Soctety OF NATURAL HIstTory { VoL. 13 


PLATE 1b. Chromodoris banksi from Puertecitos, Baja California, Mexico. 


1963 } FARMER: Two New OPpiIstTHOBRANCH MOLLUSKS 83 


f 


Fic. 1. a-e, Chromodoris norrisi: a, ventral view; b, far lateral tooth; c, 30th radular tooth; 
d, 2nd radular tooth; e, rachis. f-k, Chromodoris banksi: f, ventral view; g, 29th tooth; h, 25th 
tooth; i, 7th tooth; j, Ist tooth; k, rachis. 


exposed posterior edge of the foot is cadmium orange. The rhinophore is largely white; the 
distal part is sometimes deep cadmium orange-red, except for a white tip. The lower portion of 
the clavi contain very small black specks. The branchae are white, with deep cadmium orange 
tips in some specimens. There are very small eye spots beneath two clear areas which are just 
posterior and medial to the rhinophores. There are numerous opaque white subepidermal glands 
visible under the inner edge of the notum. 


The body shape is very similar to other chromodorids. Ten specimens collected at Cerralvo 
Island varied in size from 17-61 mm. long, 10-32 mm. broad, and 6-13 mm. high. The notum 
covers the entire animal except for the foot, which extends for a short distance beyond the pos- 
terior rim. The notum is generally smooth. The foot is slightly expanded and rounded anteriorly, 
and slightly bilabiate (fig. la). The lower lip is thick and the upper lip thinner. The tail is 
slightly pointed. 

The pronounced rhinophores, with 20-25 deeply perfoliate clavi, are borne on short stalks 
and are completely retractile into low, smooth sheaths. 

There are 11 simply pinnate retractile branchial filaments. Some animals have the filaments 
branched at the distal end up to five times. The branchae are joined at their bases in a circlet. 
The most anterior filament is the largest. The tubular anal papilla is slightly elevated and fleshy, 
and all but the posterior part is surrounded by the branchial base. 


The penis is unarmed. 


The radulae (fig. 1b-e) from 6 individuals had dental formulae ranging from 41 to 65-1-64 
to 48, with 63 to 111 rows. The type has the formula 68 x 45-1-48. There is a lip plate present 
composed of small hooks. 


84 San Disco Society oF Natura History [Vor 


Range. — Animals have been taken at Cedros Island (28° 7’ N, 115° 11’ W) and Puerto 
Rompiente (27° 43’ N, 115° 00’ W) on the Pacific coast of Baja California and Cerralvo 
Island (24° 10’ N, 109° 52’ W), Bahia de los Angeles (29° 05’ N, 113° 36’ W) and San 
Luis Gonzaga Bay (29° 49’ N, 114° 24’ W) in the Gulf of California. Animals have been 
collected in May, June, December and January. 


Comments. — Most animals were taken intertidally in rocky areas; some at Cerralvo Island 
were seen moving about on the rocks during the day but most were taken from under rocks. 
Some were taken in 10 feet of water. 


Chromodoris banksi, spec. nov. 


Type. — The type was taken by the author (WMEF 101) 2.3 miles south of Puertecitos 
(30° 24’ N, 114° 40’ W), Baja California, Mexico, on December 14, 1962. The type is 
deposited in the Invertebrate Type Series of the California Academy of Sciences, where it is 
registered as IZ No. 1. With two paratypes (IZ Nos. 2 and 3), it will be associated with 
the Frank Mace MacFarland Memorial Collection of Opisthobranchs. 


Description. — The body is white with brown-black spots of varying size on the notum, 
the side of the body, and under the mantle (plate 1b). Mixed randomly with the dark spots 
on the notum are cream colored spots and occasionally an orange one. The edge of the notum 
is translucent white with an uninterrupted orange line next to it. The edge of the foot is ringed 
with light yellow in one animal, the color intensifying to a bright cadmium orange at the exposed 
part of the tail; this area is colorless in the other specimens. The tip of the tail has a few cream 
colored spots. The tips of the rhinophores and gills are rimmed with orange. The remainder of 
the rhinophores and gills is white. 


The body shape is very similar to that of other chromodorids. The three specimens meas- 
ured 24, 33, 15 mm. long, 13, 8, 5 mm. broad and 7, 5, 3.5 mm. high. The notum covers the 
eritire animal except for the foot, which extends for a short distance beyond the posterior rim. 
The notum is generally smooth. The foot is slightly expanded and rounded anteriorly, and 


slightly bilabiate (fig. 1f). The lower lip is thick and the upper lip thinner. The rhinophores . . 


have 11 perfoliate clavi and low, smooth sheaths. There are nine simply pinnate retractile bran- 
chial filaments. The branchae are joined at their bases in a crescent. The most anterior filament 
is the largest. The tubular anal papilla is slightly elevated, fleshy, and all but the posterior part 


is surrounded by the crescentic branchial base. 
The penis is unarmed. 


The radula (fig. 1g-k) from the type has the dental formula 124 x 32-1-32. A lip plate 


composed of small hooks is present. 


Range. — Animals have been taken at 2.3 miles south of Puertecitos (30° 24’ N, 114° 
40’ W), San Luis Gonzaga Bay (29° 49’ N, 114° 24’ W) and Puerto Refugio, Angel de la 
Guardia Island (29° 33’ N, 113° 35’ W), Baja California. Animals have been collected in 
December, January and March. 


LITERATURE CITED 


OpHNER, N. H. 


1957. Chromodoris contra Glossodoris. A systematic nomenclatorial controversy. Proc. 


Malacol. Soc. London 32:250-253. 


Marcus, E. 
1961. Opisthobranch mollusks from California. Veliger 3 (Suppl., Pt. 1) : 1-84, pls. 1-10. 


JAN 6 IU 


HARVARD 
UNIVERSIT 


TRANSACTIONS 
OF THE 


SAN DIEGO SOCIETY OF NATURAL HISTORY 


Zz 
VoLuME 13, No. 7, pp. 85-168 


THE MAMMALS OF BAJA CALIFORNIA, MEXICO 


BY 


LAURENCE M. HueEy 


Former Curator of Birds and Mammals 


San Diego Natural History Museum 


SAN DIEGO, CALIFORNIA 
PRINTED FOR THE SOCIETY 


JANuaARY 15, 1964 


TRANSACTIONS 


OF THE 


SAN DIEGO SOCIETY OF NATURAL HISTORY 


VoLuME 13, No. 7, pp. 85-168 


THE MAMMALS OF BAJA CALIFORNIA, MEXICO 


BY 


LAURENCE M. Huey 


Former Curator of Birds and Mammals 
San Diego Natural History Museum 


SAN DIEGO, CALIFORNIA 
PRINTED FOR THE SOCIETY 


JANUARY 15, 1964 


EDITOR'S NOTE 


When Laurence M. Huey retired from the San Diego Natural History Museum in 
April, 1962, after 40 years of service as Curator of Birds and Mammals, he left the manuscript 
of the present report in essentially finished form. Early versions of the manuscript had been 
read by two persons, and many of their editorial comments had been incorporated. The present 
editor saw the manuscript through a final retyping and the proof stages. Editing done at this 
time consisted only of checking punctuation and grammar and verifying and making consistent 
the citations and references. The editor was also responsible for preparing the table of contents 
and the index. In only a few instances was Huey’s wording or phraseology changed. 


Laurence Huey died on June 10, 1963, before having the opportunity of reviewing the 
final edited typescript, and only a few weeks before the paper went to the printer. Whereas 
the format, content, and statements of fact in this paper are those set forth by Mr. Huey, the 
editor takes responsibility for typographical errors and inconsistencies which may occur. 


This paper is, in effect, the culmination of Laurence Huey’s work on the mammals of 
Baja California. It summarizes not only the published literature but also, and more importantly, 
the knowledge obtained from nearly 50 years of intimate experience, in the field and in the 
museum, with the mammals of that area. 


MUS. COMP. ZOOL 
LIBRARY 


JAN 28 1364 


HARVARD 
UNIVERSITY. 


THE MAMMALS OF BAJA CALIFORNIA, MEXICO 


BY 


LAURENCE M. Hugy 


TABLE,.OF GCONTENES 


AnteOcticre ee eee te) eres. ee Ree play. ANU gr, alle a Bg EONS el ee ee &8 
12) aie Meat. te aR et nck OO Ate ey gt OL ce OR RS OT RR SN ea Mele AD 90 
PSECOUMES 401 (SPECIES. 5 ates tne chest eee SRT ae St) Pci ey 8s 91 
Widelphidae.-3 3. 30a rh ct oe A Wellna ts Gee bee Bie EN ethene eee 91 
SOMCIgaGagt eek Mel a eet Rae Re a OL Oi Se EAP ll 91 
flea piclaien. eee rere ees, See eis 2h hab ee 1S ee at eh eee oe 92 
Emballonutidae: less. 325 ne ee Bee ee PE Be EE heed 92 
Phwllostomudae; 398000 lei ha aes at oo ee Tk eee eS 93 
Die lite PVs ghee ok webs telo NT siherades Ate AeA i Peet IMR SHULL Ls EAR EBs gh ab Pie aD A Os 93 
Wes perciionicaeye treat anne Seen Me een ean see ee meee 93 
Dytolossicl tes mers Bled rey et eins ene ee Ne ae ne ie ae eed ce te ee 98 
[Ee poridac tes alia fA seu) Soh et UE ae 2 era Nich PAK 99 
Serie vette ae Wee tae ae Deal Uo an rane Anat ink ip Ae et aad 102 
(GDS Tan STS YS? hese sales ate, a2 AO Se a rE 105 
Pele Ger cumy 1G AC 8, 2i eR ce Oa ee tee A Bt re ee 110 
Gastoridae: a: a. DD Pi ovgteiamis eines pat Dn oe 1429) wnedsree: We baieaieas 124 
Griceticae Peete ORs ile 64. AAT IG, hod dy Ane a BYE ON ED 124 
I Arieccheve ees es Bestia ee ee Ace A ee eer ae Pe een iS 
PESTON SUT SEV Me Rect ONE, SICA coat mare fe Sem Feces, Skee PLgME oe OEE are 138 
Bhiyseceridac, oeet mu tate FAO nt ee 1) AGA AS he a See 138 
| JCA EIG Ver Bete Dies ae aay ohne 8 eae ee Va ee se ee 138 
[Delphinidaes cet ons ke te Ba eas Ee Ak eee eine 138 
Ste grad avete ete age cp 5) aeeea ae RnS  RN EN Pemreey wate t Cee Pye eee AT. 140 
Balaenoptericae ys! ese te Ae es, Soe i Poieeee, Sgn) sei card RS 140 
Baltenicae-g2-2 a 2.8 Wee 28 Ate ee Dee ee eee hs ee 141 
(BETTY 3 BERR, eee ee Ee DONO: LO Sah UME Ene teat Oar aaNeS Seme ee Ona 142 
Whresidse Aes re ges Sah MES IA ol be hie | nee ese kL 143 
Peocy orice ee tare en renre: WEN CBs: Nr noes. deters Ee ee 144 
IVE ias elidel Mee eI Nate ee ate a eee te ee 145 
Beligaces ts cee ove sey ee de eh SN Seen eee ee. 147 
OEIC eM es Ss ee ene oO, tae tle? la dee 5 Skt ia A ta a HO 148 
PD austere aeeaeer beer he Bet eres Be. ewe Se eh Rey ee ie Co ee 149 
Geyer Ws er ash ARG lla le Fen sd res oe en EN ahd SPE aah 150 
Antilocapridae .............. peered eR a Sete a eae eee a 151 
[BYSSUATS 1S (Se eae, SU 8 EE ot gl! RCN ie: nee Ber AEN ates? SERINE a ME 151 
Dessbstea cet nak ee rs cc BN ee ae neice ea rr 153 


ciclo come mine al 6 pleas ov weve) tyke’ oie ores fan al eer ae 159 


88 SAN Dreco Society oF Natura History { VoL. 13 


INTRODUCTION 


This paper is the first attempt to catalog the kinds of mammals known to occur in the long 
peninsula of Baja California and on the surrounding coastal islands. 


The early Spanish missionaries and explorers in their accounts of this remote land offered 
singular and vague descriptions of mountain sheep as early as 1757, but the first definite account 
of mammalian species in Baja California is by Spencer F. Baird (1859) in a paper on a collection 
of birds made by John Xantus at Cape San Lucas. 


From then until 1902, many naturalists collected in this interesting region, but they were 
mainly seeking minerals or botanical or ornithological specimens. Mammals were collected only 
incidentally or by only a few persons. Among the most prominent were Anthony, Belding, 
Bryant, and Price, who made small collections of mammals from scattered localities along the 
peninsula and the Pacific islands. The novelties of these collections were reported by J. A. Allen, 
C. Hart Merriam, Gerrit S. Miller, Jr., and others. Captains C. M. Scammon and Charles H. 
Townsend, while in pursuit of their separate interests, made notable captures or observations 
along the coasts and recorded them in their reports. 


In 1902, Edmund Heller, in the interests of the Field Columbian Museum, now the 
Chicago Natural History Museum, made an extensive collecting trip by means of pack animals 
across the northern part of the peninsula from San Quintin on the Pacific coast, over the Sierra 
San Pedro Martir and Sierra Juarez to San Felipe on the Gulf of California. This trip was one 
of the early major efforts to collect mammals on the peninsula. The results were published by 


D. G. Elliot (1903b). 


In 1905 and 1906, E. W. Nelson and E. A. Goldman, using pack animals, traversed the 
entire length of the peninsula, as part of the field program of the Bureau of Biological Survey 
of the United States Department of Agriculture. From their carefully made collections of plants, 
birds, and mammals, numerous novelties were described by the collectors and others. Later, 
Nelson (1921) wrote the best descriptive book on the natural history of this arid land so far 
written, still a necessary reference for any worker. A general list of the mammals was not 
made, but references to various species are to be found throughout the text. 


Burt (1932) reported on his own magnificent collection and on two smaller collections 
made by D. R. Dickey and H. H. Sheldon. On a cruise with J. R. Pemberton and others, W. H. 
Burt trapped on all of the important islands in the Gulf of California north to Mejia Island 
(29° 33’ N). This outstanding paper described twenty species and subspecies of three genera — 
Perognathus, Peromyscus, and Neotoma—the largest number of mammals described from 
Baja California in one paper by a single author. A summary lists all known species and races 
of the three genera found on islands in the Gulf. 


My mammal-trapping experience in Baja California started in May of 1914, when I camped 
on North Los Coronados Island. The chief objective was to collect bird eggs, but a series of 
Peromyscus maniculatus, then swarming on the island, was preserved. But it was not until 
March, 1923, when I was employed by the San Diego Society of Natural History as Curator 


of Birds and Mammals, that I commenced serious mammal collecting in Baja California. 


As a guest of Griffing Bancroft in April, 1923, I made my first trip by automobile into 
this enchanting land. We reached El Marmol and Santa Catarina Landing, then the end of a 
rough and tortuous road. The purpose of the trip was oological collecting —a complete day- 
light occupation. The considerable number of mammals collected was eviscerated and prepared 
by candlelight in the evening and often far into the early morning. 


That summer, with several other California collectors, I was the guest of Prof. José M. 
Gallegos, who was working as a scientific explorer in northern Baja California for the Mexican 
government. We visited with pack animals the forested heights of the Sierra San Pedro Martir. 


1963 | Huey: MAmMats oF BajA CALIFORNIA 89 


Later, as the Professor’s guest, I went on the Mexican fisheries’ patrol boat “Tecate” to Guadalupe 
Island, to census the elephant seals, then in the dawn of their population resurgence. In the 
following three years, Professor Gallegos joined Mrs. May Canfield and me on trips into the Sierra 
Juarez and other remote areas in northern Baja California. We used two Model T Fords as 
transportation. A close friendship with Professor Gallegos was ended by his untimely death on 
an official trip in British Honduras. During these years the Museum’s mammal collection grew 


rapidly. 
March 22 to April 25, 1926, was profitably spent by Mrs. Canfield and myself collecting 


birds and mammals at San Felipe, a remote fishing village located on the shore of the Gulf of 
California. This was a hazardous trip at that date and we had the pleasure of the company of 
Chester C. Lamb and Raymond Gilmore, representing the Museum of Vertebrate Zoology, 
University of California, who, while working independently, were road companions and camp 
neighbors during our stay and provided the pleasant association of fellow workers. 


Fisherman friends gave me the opportunity of visiting all of the Pacific coastal islands north 
of San Ignacio Lagoon except the San Benitos, and Griffing Bancroft invited me to accompany 
him on a boat trip into Scammon’s Lagoon. 


In 1926-27 the Mexican government appropriated 3,500 pesos to extend the automobile 
trail south from El Marmol to San Ignacio. It was completed in 1927 as a contour road without 
fills or bridges, rough with rocks and chuck holes. Over this primitive trail, from February to 
May, 1928, Mrs. Canfield and I in our two Model T’s, reached the end of automobile travel 
on the peninsula — then at Mulegé on the Gulf of California. Enroute, at Punta Prieta, Mrs. 
Canfield was overtaken by a recurrence of a very serious illness and almost lost her life; but, 
after a two weeks’ delay and without any medical help, she revived and insisted that the expedi- 
tion continue. She was the first woman to drive a car over this road. Upon our return home she 
was again stricken in September and was thereafter confined to a wheel chair until her death 
twenty-two years later. 


Several productive short trips with different companions were made into the central and 
arid desert sections of Baja California in the 1930’s. In this time the final link in the automobile 
road between Mulegé and Cape San Lucas had been completed. 


In 1941 the Museum organized an expedition by truck the length of the peninsula, financed 
by Max Miller, a local author seeking material for a book on this primitive region. With him 
and Frank F. Gander of the Museum staff, I collected mammals vigorously southward to La Paz, 
where Mr. Miller crossed the Gulf to Guaymas enroute home and left the two collectors on their 
own. On the return trip we stopped where mammal trapping would yield the most desirable 
specimens. December 7, 1941, the day of the Japanese attack on Pearl Harbor, found us at Santo 
Domingo Landing on the remote Vizcaino Desert. We heard the news on our small portable 
radio that evening. This place is an important type locality, but we stayed only two days longer 
and then started north, arriving at the international boundary a week later to find great changes 
in a homeland now at war. 


As a result of the collecting described above, the mammals of the mainland were well enough 
represented at San Diego to permit several reports to be made in the next five years. These 
reports revealed the need for collecting at several additional localities and so in the spring of 
1947, Charles Harbison of the Museum staff and I, in a truck, made a productive trip to the 
Gulf of California midway down the peninsula. In June a companion and I revisited El Marmol 
and Santa Catarina Landing. This general area was again visited in the early spring of 1958 by 
A. B. Howell, myself, and our wives to augment series of several species that were then being 
studied. This was the last collecting trip made into Baja California by the writer. That there are 
yet some novelties to be found is shown by the description of a relict kangaroo rat trapped on 
a high mesa in central Baja California (Huey, 1962). 


90 SAN Dtieco Society oF Naturav History | Vov. 13 


In February, 1960, I participated in the annual gray whale census made by airplane along 
the Pacific coast and in the lagoons of Baja California. This survey has been made each February 
for several years by Dr. Carl L. Hubbs and associates. From airplanes at 4000 to 5000 feet 
elevation, along the coastline and over the lagoons where these cetaceans gather each year to 
calve and breed, observers were able to make satisfactory population counts. This four-day expedi- 
tion was most interesting; but the highlight of the trip was in the criss-cross flights over the 
peninsula to and from Bahia de los Angeles on the Gulf side, which was our headquarters. To 
view from the comfortable seat of an airplane the tortuous, rough, and rocky roads winding 
ribbon-like through familiar country and to peek into the deep, labyrinthian, palm-filled canyons 
among lava-covered mesas, where the field naturalist longs to explore but may never have the 
chance, is an unmeasured thrill difficult to express. It does, however, offer to the observant 
naturalist a limited understanding of the problems involved in the ecology of terrestrial mammal 
populations. 


PLAN 


The name of each subspecies or monotypic species is followed by a vernacular name. The 
synonymy includes the following names, each with the place of first publication: (1) the first 
available name in the form originally used; (2) the present combination, if different; (3) syn- 
nonyms which apply to Baja California populations, each with the type locality in parentheses. 
Then follows a statement of the type locality for the accepted name and a statement of the 
range in Baja California, based on the literature and on specimens, mostly in the collection of 
the San Diego Society of Natural History. Any specimen that extends the known range is cited, 
with date of capture, and brief commentary is made on a few species in regard to either their 
habits, habitats, or ranges. 


The names of the species and subspecies listed herein are arranged geographically, from 
north to south with the international boundary as the starting point. This will facilitate their 
location down this long peninsula, as neither a map of Baja California nor a gazetteer of localities 
is offered. 


The following abbreviations are used in the species accounts: S. D. N. H. M., San Diego 
Natural History Museum; M. V. Z., Museum of Vertebrate Zoology; U. S. N. M., United 
States National Museum. 


Three new subspecies in the genus Perognathus are described. 


1963 } Huey: MammMats oF BajA CALIFORNIA 9] 
ACCOUNTS OF SPECIES 


Class MAMMALIA — Mammals 
Order MARSUPIALIA — Pouched Mammals 
Family DIDELPHIDAE — Opossums 
Genus Didelphis Linnaeus 
Didelphis marsupialis virginiana Kerr 
Virginia Opossum 
1792. Didelphis virginiana Kerr, Animal Kingdom .. . :193. 
1951. [Didelphis marsupialis} virginiana Hershkovitz, Fieldiana-Zool. 31:550. July 10. 

Type locality. — Virginia. 

Range. — Opossums were introduced into the San Gabriel Valley near Los Angeles, Cali- 
fornia, in 1906 (Little, 1916) and have extended their range southward to the Mexican boundary 
where they are common residents along the Tijuana River bottom. 

A caged specimen has been reported from a ranch six miles south of Tijuana, where it 


was captured in May, 1962. 


Order INSECTIVORA — Insectivores 
Family SORICIDAE — Shrews 
Genus Sorex Linnaeus 
Sorex ornatus ornatus Merriam 


Adorned Shrew 


1895. Sorex ornatus Merriam, North Amer. Fauna 10:79. December 31. 
1903. Sorex oreinus Elliot, Field Columb. Mus. Publ. 74, Zool. Ser. 3:172. May 7. (Aguaje 

de las Fresas, Sierra San Pedro Martir, Baja California. Alt. 6000 feet.) 

Type locality. — Head of San Emigdio Canyon, Mount Pinos, Kern Co., California. Alt. 
about 5500 feet. 

Range. — Pacific slope, sea level to mountain tops, of northern sections of the peninsula 
from latitude 30°N northward to international boundary. 

There are specimens in the S. D. N. H. M. from El Rosario and the Sierra San Pedro 
Martir. 

Sorex ornatus lagunae Nelson and Goldman 


San Lucas Adorned Shrew 


1909. Sorex lagunae Nelson and Goldman, Proc. Biol. Soc. Wash. 22:27. March 10. 
1928. Sorex ornatus lagunae Jackson, North Amer. Fauna 51:169. July 24. 
Type locality. — La Laguna, Sierra Laguna, Baja California. Alt. 5500 feet. 
Range. — Known only from the type locality. 


Sorex juncensis Nelson and Goldman 
Tule Shrew 


1909. Sorex californicus juncensis Nelson and Goldman, Proc. Biol. Soc. Wash. 22:27. March 
10. 
1928. Sorex juncensis Jackson, North Amer. Fauna 51:172. July 24. ; 
Type locality. — Socorro, a small coastal Salicornia marsh, 15 miles south of San 
Quintin, Baja California. 
Range. — Known only from the type locality. . 
Nelson and Goldman caught 2 specimens at this locality in September, 1905. The writer 
has set several trap lines through this small salt marsh without success. 


92 San Dreco Society oF Natura History { VoL. 13 


Genus Notiosorex Coues 


Notiosorex crawfordi crawfordi (Coues) 
Desert Shrew 


1877. Sorex (Notiosorex) crawfordi Coues, Bull. U. S. Geol. Geogr. Surv. Terrs. 3:651. 
May 15. 

1884. Sorex crawfordi True, Proc. U. S. Nat. Mus. 7:606. November 29. 

1895. Notiosorex crawfordi Merriam, North Amer. Fauna 10:32. December 31. 
Type locality — Near old Fort Bliss, about 2 miles above El Paso, El Paso County, Texas. 
Range. — Probably occurs in suitable habitat over the length of the peninsula. 


A mummy specimen in the S. D. N. H. M., first preserved in tequila and then dried, was 
collected at Cape San Lucas, Baja California, by T. S. Brandegee in November of 1902. 


Family TALPIDAE — Moles 
Genus Scapanus Pomel 


Scapanus latimanus occultus Grinnell and Swarth 
Southern California Broad-footed Mole 


1912. Scapanus latimanus occultus Grinnell and Swarth, Univ. Calif. Publ. Zool. 10:131. 
April 13. 


Type locality. — Santa Ana Canyon, 12 miles northeast of Santa Ana, Orange County, 
California. Alt. 400 feet. 


Range. — Known to occur over the higher parts of the Sierra Juarez in northern Baja 
California. 


A single male specimen from Laguna Hanson, taken July 22, 1924, is in the S. D. 
NE He M: 


Scapanus latimanus anthonyi J. A. Allen 
Sierra San Pedro Martir Mole 


1893. Scapanus anthonyi J. A. Allen, Bull. Amer. Mus. Nat. Hist. 5:200. August 18. 
1937. Scapanus latimanus anthonyi F. G. Palmer, Jour. Mamm. 18:312. August 14. 
Type locality. — Sierra San Pedro Martir, Baja California. Alt. 7000 feet. 
Range. — Higher levels of the Sierra San Pedro Martir in northern Baja California. 


Order CHIROPTERA — Bats 
Family EMBALLONURIDAE — Sac-winged Bats 
Genus Balantiopteryx Peters 


Balantiopteryx plicata pallida Burt 
Peters’ Pale Bat 
1948. Balantiopteryx plicata pallida Burt, Occ. Pap. Mus. Zool. Univ. Michigan 515:1. October 
30. 


Type locality. — San Bernardino, Rio Mayo, Sonora. 
Range. — Known in Baja California from the Cape District. 
Four male and two female specimens in the collection of Luther Little were collected 


March 21 and 22, 1923, by Chester Lamb at San José del Cabo (fide letter, 1958). This sub- 


species is also known from Santa Anita. 


1963 } Hurty: MammMats oF BayA CALIFORNIA 93 


Family PHYLLOSTOMIDAE — American Leaf-nosed Bats 
Genus Macrotus Gray 
Macrotus californicus Baird 


California Leaf-nosed Bat 


1858. Macrotus californicus Baird, Proc. Acad. Nat. Sci. Philadelphia 10:116. May. 
Type locality. — Old Fort Yuma, Imperial County, California, opposite Yuma, Arizona. 


Range. — Found chiefly on the desert slopes, from the international boundary south to 


the Cape region at La Paz (H. Allen, 1893). 


Genus Choeronycteris Tschudi 
Choeronycteris mexicana Tschudi 


Mexican Long-tongued Bat 


1844. Ch{oeronycteris } mexicana Tschudi, Untersuchungen uber die Fauna Peruana. . . :72. 

Type locality. — Mexico. 

Range. — Found coastwise from the international boundary southward to Rancho Ra- 
mona, latitude 30° N (Huey, 1954). 

This Mexican species seems to be of irregular occurrence in the northern section of its 
Baja California range. Specimens from Aguaito, 20 miles east of El Rosario, and from Rancho 


Ramona are in the S. D. N. H. M. 


Family NATALIDAE — Funnel-eared Bats 
Genus Natalus Gray 
Natalus stramineus mexicanus Miller 


Funnel-eared Bat 


1902. Natalus mexicanus Miller, Proc. Acad. Nat. Sci. Philadelphia 54:399. September 12. 
1959. Natalus stramineus mexicanus Goodwin, Amer. Mus. Novit. 1977:6. December 22. 
Type locality. — Santa Anita, Baja California. 
Range. — Known from two nearby localities on the eastern side of the Cape region of 


the peninsula (Dalquest and Hall, 1949). 


Family VESPERTILIONIDAE — Vespertilionid Bats 
Genus Myotis Kaup 
Myotis yumanensis yumanensis (H. Allen) 


Desert Yuma Bat 


1864. Vespertilio yumanensis H. Allen, Smiths. Misc. Coll. 7 (165) :58. June. 
1897. Myotis yumanensis Miller, North Amer. Fauna 13:66. October 16. 
Type locality. — Old Fort Yuma, Imperial County, California, opposite Yuma, Arizona. 
Range. — Over the Colorado River delta area and adjoining desert foothills in extreme 
northeastern Baja California, thence south to Cape San Lucas except the area about San Ignacio. 


Myotis yumanensis sociabilis H. W. Grinnell 
Tejon Yuma Bat 


1914. Myotis yumanensis sociabilis H. W. Grinnell, Univ. Calif. Publ. Zool. 12:318. Decem- 
ber 4. 
Type locality. —Old Fort Tejon, Tehachapi Mountains, Kern Co., California. Alt. 
3200 feet. ; 
Range. — Along the Pacific slope of the coast range mountains, Sierra Juarez and Sierra 
San Pedro Martir, west and south to the vicinity of San Quintin, and probably farther to the 
south. 
A specimen in the S. D. N. H. M. from Santo Domingo, near latitude 31° N, was 
collected June 2, 1925. 


94 SAN Deco Society oF Natura History {Vot. 13 


Myotis yumanensis lambi Benson 
San Ignacio Yuma Bat 


1947. Myotis yumanensis lambi Benson, Proc. Biol. Soc. Wash. 60:45. May 19. 

Type locality. — San Ignacio (27°17’N), Baja California. 

Range. — Known only from the type locality, but probably present in other suitable areas 
in the central section of the peninsula. 


Myotis velifer peninsularis Miller 
Cave Bat 


1898. Myotis peninsularis Miller, Ann. Mag. Nat. Hist., ser. 7. 2:124. August. 

1928. Myotis velifer peninsularis Miller and G. M. Allen, U. S. Nat. Mus. Bull. 144:93. 
May 25. 
Type locality. — San José del Cabo, Baja California. 
Range. — Found in the southern end of the peninsula. 


Myotis evotis evotis (H. Allen) 
Little Big-eared Bat 


1864. Vespertilio evotis H. Allen, Smiths. Misc. Coll. 7 (165) :48. June. 
1897. Myotis evotis Miller, North Amer. Fauna 13:77. October 16. 
1909. Myotis micronyx Nelson and Goldman, Proc. Biol. Soc. Wash. 22:28. March 10. 
(Comondu, Baja California.) 
Type locality. — Monterey, Monterey County, California. 
Range. — From the international boundary as far south as Comondu (latitude 26°5’N) 
in the southern section of the peninsula. 


Myotis milleri Elliot 
Sierra San Pedro Martir Bat 


1903. Myotis milleri Elliot, Field Columb. Mus. Publ. 74, Zool. Ser. 3:172. May 7. 
Type locality. —La Grulla, Sierra San Pedro Martir, Baja California. Alt. 8000 feet. 
Range. — Higher mountains in northern Baja California. 
There are specimens in the S. D. N. H. M. from El Valle de la Trinidad and the Sierra 
San Pedro Martir. 


Myotis thysanodes thysanodes Miller 
Fringed Bat 


1897. Myotis thysanodes Miller, North Amer. Fauna 13:80. October 16. 

Type locality. — Old Fort Tejon, Tehachapi Mountains, Kern County, California. Alt. 
3200 feet. 

Range. — The hilly and mountainous parts of the northern section of Baja California. 

A male taken from a tunnel at the 50 foot level in the El Fendmeno Mine in the Sierra 
Juarez, January 31, 1954, is in the S. D. N. H. M. 


Myotis volans volans (H. Allen) 
Long-legged Bat 


1866. V{espertilio} volans H. Allen, Proc. Acad. Nat. Sci. Philadelphia 18:282. 
1909. Myotis capitaneus Nelson and Goldman, Proc. Biol. Soc. Wash. 22:28. March 10. 
(San Jorge, 30 miles southwest of Comondu, Baja California.) 
1914. Myotis volans Goldman, Proc. Biol. Soc. Wash. 27:102. May 11. 
Type locality. — Cape San Lucas, Baja California. 
Range. So far as now known, this form is found only in the southern part of the 
peninsula. 


1963 } Hurty: MAmmats of BAJA CALIFORNIA 95 


Myotis volans interior Miller 
Interior Long-legged Bat 


1914. Myotis longicrus interior Miller, Proc. Biol. Soc. Wash. 27:211. October 31. 
1928. Myotis volans interior Miller and G. M. Allen, U.S. Nat. Mus. Bull. 144:142. May 25. 
Type locality. — Five miles south of Twining, Taos County, New Mexico. Alt. 11,300 
feet. 
Range. — So far as known, over northeastern parts of the peninsula. 
A male attributable to this race in the S. D. N. H. M. was taken on the western side 
of Laguna Salada at Demara’s Well, on November 25, 1936. 


Myotis californicus californicus (Audubon and Bachman) 
Little California Bat 


1842. Vespertilio californicus Audubon and Bachman, Jour. Acad. Nat. Sci. Philadelphia, 
ser. I, pt. 2, 6:28). ; 
1897. Myotis californicus Miller, North Amer. Fauna 13:69. October 16. 
Type locality. — California. By subsequent restriction Monterey, Monterey County, 
California (Miller and Allen, 1928). 
Range. — Along the Pacific coast from the United States boundary south to Cape San 
Lucas. 


Myotis californicus stephensi Dalquest 
Stephens Little California Bat 


1900. Myotis californicus pallidus Stephens, Proc. Biol. Soc. Wash. 13:153. June 13. (Name 
pre-occupied by Kf erivoula} pallida. Blyth, Cat. Mamm. Mus. Asiat. Soc. Bengal, 1863.) 
1946. Myotis californicus stephensi Dalquest, Proc. Biol. Soc. Wash. 59:67. March 11. 
Type locality. — Vallecito, San Diego County, California. 
Range. — Along desert slopes and over the Colorado River delta area in northeastern 


Baja California. 
Myotis subulatus melanorhinus (Merriam) 
Black-nosed Bat 


1890. Vespertilio melanorhinus Merriam, North Amer. Fauna 3:46. September 11. 
1903. Myotis orinomus Elliot, Field Columb. Mus. Publ. 79, Zool. Ser. 3:228. August 1). 
(La Grulla, Sierra San Pedro Martir, Baja California.) 
1928. Myotis subulatus melanorhinus Miller and G. M. Allen, U. S. Nat. Mus. Bull. 144:169. 
May 25. 
Type locality. — Little Spring, north base of San Francisco Mountain, Coconino County, 
Arizona. 
Range. — In the forest areas of the higher mountains of northern Baja California. 


Genus Pizonyx Miller 
Pizonyx vivesi (Menegaux) 


Fish-eating Bat 


1901. Myotis vivesi Menegaux, Bull. Mus. Hist. Nat. Paris 7:323. 

1906. Pizonyx vivesi Miller, Proc. Biol. Soc. Wash. 19:85. June 4. 
Type locality. —“Ilot du Cardonal au Islo, parti d I’Archipel Salsi puedes”, Gulf of 

California, Baja California. Now understood to be Isla Partida (28°53’N, 113°04’W), Gulf 

of California, Baja California (Reeder and Norris, 1954). ; 
Range. — On both Sonora and Baja California shores of the upper half of the Gulf of 

California and on the Pacific side of the peninsula in the vicinity of Punta Eugenia. 


96 SAN Drseco Society oF NATURAL History { Vot. 13 


Genus Pipistrellus Kaup 
Pipistrellus hesperus hesperus (H. Allen) 


Western Pipistrelle 


1864. Scotophilus hesperus H. Allen, Smiths. Misc. Coll. 7 (165) :43. June. 
1897. Pipistrellus hesperus Miller, North Amer. Fauna 13:88. October 16. 

Type locality. — Old Fort Yuma, Imperial County, California, opposite Yuma, Arizona. 

Range. — Over the delta of the Colorado River and through El Valle de la Trinidad 
into the arid western slopes of Sierra Juarez and south through the higher Sierra San Pedro 
Matrtir at least to Calmalli (latitude 28°15’N). 

Specimens are in the S. D. N. H. M. from San Felipe (April), Laguna Hanson 
(October), Sangre de Cristo (June), El Valle de la Trinidad (March, June, July), San Fer- 
nando (April), Catavifa (June), and Calmalli (April). 


Pipistrellus hesperus merriami (Dobson) 
Merriam Western Pipistrelle 


1886. Vesperugo merriami Dobson, Ann. Mag. Nat. Hist., ser. 5, 18:124. August. 
1913. Pipistrellus hesperus merriami Grinnell, Proc. Calif. Acad. Sci., ser. 4, 3:279. August 28. 
Type locality. — Red Bluff, Tehama County, California. 
Range. — This subspecies occurs in summer over the coastal plain and through the live 
oak belt from the international boundary south to Ensenada. 


Pipistrellus hesperus australis Miller 
Southern Western Pipistrelle 


1897. Pipistrellus hesperus australis Miller, North Amer. Fauna 13:90. October 16. 

Type locality. — Barranca Ibarra, Jalisco. 

Range. — This small bat is found in suitable localities over the southern third of the 
peninsula from San Ignacio southward. 

There are specimens in the S. D. N. H. M. from Los Barriles (November, 1941). 


Genus Eptesicus Rafinesque 
Eptesicus fuscus pallidus Young 


Pallid Brown Bat 


1908. Eptesicus pallidus Young, Proc. Acad. Nat. Sci. Philadelphia 60:408. October 14. 
1912. Eptesicus fuscus pallidus Miller, U. S. Nat. Mus. Bull. 79:62. December 31. 

Type locality. — Boulder, Boulder County, Colorado. 

Range. — Northeastern section of Baja California including, during summertime, the 
higher parts of the Sierra San Pedro Martir, thence seasonally over lower sections of the northern 
parts of the mid-peninsula. 

Specimens in the S. D. N. H. M. taken in the Sierra San Pedro Martir in summer are 
assignable to this race. 


Eptesicus fuscus bernardinus Rhoads 
Pacific Brown Bat 


1902. Eptesicus fuscus bernardinus Rhoads, Proc. Acad. Nat. Sci. Philadelphia 53:619. Feb- 
ruary 6. 
Type locality. — Near San Bernardino, San Bernardino County, California. 
Range. — Along the coastal region from the international boundary south to San Quintin. 
Specimens of this race in the S. D. N. H. M. were collected from a small colony in an 
attic at the Hamilton Ranch, Santo Domingo, near latitude 31°N, in June. At this time young 
of the year were on the wing. 


1963 | Hury: Mammats oF BAJA CALIFORNIA 97 


Eptesicus fuscus peninsulae (Thomas) 
Cape San Lucas Brown Bat 


1898. Vespertilio fuscus peninsulae Thomas, Ann. Mag. Nat. Hist., ser. 7, 1:43. January. 
1912. Eptesicus fuscus peninsulae Miller, U. S. Nat. Mus. Bull. 79:63. December 31. 
Type locality. — Sierra Laguna, Baja California. 
Range. — Tip of the peninsula of Baja California. 


Genus Lasiurus Gray 
Lasiurus borealis teliotis (H. Allen) 


Western Red Bat 


1891. Atalapha teliotis H. Allen, Proc. Amer. Philos. Soc. 29:5. April 10. 
1897. Lasiurus borealis teliotus Miller, North Amer. Fauna 13:110. October 16. 
Type locality. — Unknown, probably some part of California. 
Range. — From the international boundary south to Comondu, southern Baja California 


(Anthony, 1928). 


Lasiurus cinereus cinereus (Palisot de Beauvois) 
Hoary Bat 


1796. Vespertilio cinereus (misspelled linereus) Palisot de Beauvois, Catal. Raisonne Mus. 

Peale Philadelphia, p. 18 (page 15 of English edition by Peale and Palisot de Beauvois) . 
1864. Lasiurus cinereus H. Allen, Smiths. Misc. Coll. 7 (165) :21. June. 

Type locality. — Philadelphia, Pennsylvania. 

Range. — Extreme northern humid section of the peninsula. 

One specimen in the S. D. N. H. M. was shot flying amid pine trees at 9:30 p.m. on 
October 11, 1926, at Laguna Hanson. This is a late flying species. 


Lasiurus ega xanthinus (Thomas) 
Sierra Laguna Bat 


1897. Dasypterus ega xanthinus Thomas, Ann. Mag. Nat. Hist., ser. 6, 20:544. December. 
1953. Lasiurus ega xanthinus Dalquest, Louisiana State Univ. Studies, Biol. Ser. 1:61. Decem- 

ber 28. 

Type locality. — Sierra Laguna, Baja California. 

Range. — A specimen recorded from Palm Springs on the desert side of the coast range 
mountains in southern California (Constantine, 1946) offers evidence that this tropical species 
sporadically wanders into the northern desert sections of Baja California as has Choeronycteris 
mexicana on occasion. The species seems to be associated with Fan Palms, in which it finds 
shelter. 


Genus Plecotus E. Geoffroy Saint-Hilaire 


Plecotus townsendii pallescens (Miller) 
Pale Big-eared Lump-nosed Bat 


1897. Corynorhinus macrotis pallescens Miller, North Amer. Fauna 13:52. October 16. 
1959. Plecotus townsendii pallescens Handley, Proc. U. S. Nat. Mus. 110:190. 

Type locality. — Kearn Canyon, Navajo County, Arizona. 

Range. — Probably in acceptable places over the northern section of the peninsula south 
at least to Calmalli. 

A female collected April 10, 1928, at the latter locality, and a male taken from a mine 
tunnel 25 miles north of Punta Prieta, November 15, 1947, are in the S. D. N. H. M. 


98 SAN Disco Society oF Natura History [VoLs 13 


Genus Antrozous H. Allen 
Antrozous pallidus pallidus (Le Conte) 


Desert Pallid Bat 


1856. V{espertilio} pallidus Le Conte, Proc. Acad. Nat. Sci. Philadelphia 7:437. 
1864. Antrozous pallidus H. Allen, Smiths. Misc. Coll. 7 (165) :68. June. 

Type locality. — El Paso, El Paso County, Texas. 

Range. — This subspecies occurs along the international boundary from the Colorado 
River westward to the vicinity of Jacumba, San Diego County, California. 


Antrozous pallidus pacificus Merriam 
Pacific Pallid Bat 


1897. Antrozous pallidus pacificus Merriam, Proc. Biol. Soc. Wash. 11:180. July 1. 

Type locality. — Old Fort Tejon, Tehachapi Mountains, Kern County, California. 

Range. — This bat ranges from the international boundary on the Pacific slope south 
into the Ilano area below the Sierra San Pedro Martir. 

There are specimens in the S. D. N. H. M. from 10 miles northwest of San Fernando, 
San Fernando, and 7 miles north of Santa Catarina. 


Antrozous pallidus minor Miller 
Cape San Lucas Pallid Bat 


1902. Antrozous minor Miller, Proc. Acad. Nat. Sci. Philadelphia 54:389. September 3. 
1951. Antrozous pallidus minor Goldman, Smiths. Misc. Coll. 115:356. July 31. 

Type locality. — Comondu, Baja California. 

Range. — Southern section of the peninsula. 

There are specimens in the S. D. N. H. M. from Los Barriles, on the southeastern tip 
of the peninsula, where a small colony of about 20 individuals was found roosting in a slight 
cave-like overhang on the shady side of a boulder beside the main road on November 4, 1951. 
A single clap of a butterfly net against the rock captured four females and one male. Another 
female was netted at 10:00 p.m. in an old abandoned adobe house at Santo Domingo (25°30’N) 
on the Magdalena Bay side of the peninsula on November 19, 1941. 


Family MOLOSSIDAE — Free-tailed Bats 
Genus Tadarida Rafinesque 


Tadarida brasiliensis mexicana (Saussure) 
Mexican Free-tailed Bat 


1860. Molossus mexicanus Saussure, Rev. Mag. Zool. Paris, ser. 2, 12:283. July. 
1955. Tadarida brasiliensis mexicana Schwartz, Jour. Mamm. 36:108. February 28. 

Type locality. — Cofre de Perote, Vera Cruz. Alt. 13,000 feet. 

Range. — The length of the peninsula. 

There are specimens in the S. D. N. H. M. from El Valle de la Trinidad, where they 
were shot in flight in the early evening of March 18, 1936, and from a small colony found roosting 
behind large framed oil paintings hanging on the wall of the San Ignacio Mission, March 12, 


1928. Both sexes were taken in the same colony. 


Tadarida femorosacca (Merriam) 
Pocketed Bat 


1889. Nyctinomus femorosaccus Merriam, North Amer. Fauna 2:23. October 30. 
1924. Tadarida femorosacca Miller, U. S. Nat. Mus. Bull. 128:86. April 29. 

Type locality. — Agua Caliente, now Palm Springs, Riverside County, California. 

Range. — The length of the peninsula; recorded from the Sierra San Pedro Martir and 
from Santa Anita in the Cape District (Shamel, 1931). 


1963 | Huey: MAMMALS oF BAJA CALIFORNIA 99 


Tadarida molossa (Pallas) 
Large-eared Free-tailed Bat 


1766. V{espertilio} molossus Pallas, ... Misc. Zool. .. .; Spicilegia Zool... . fasc. 3:8, 1767. 
1931. Tadarida macrotis Shamel, Proc. U.S. Nat. Mus. 78 (19) :15. May 6. (Part.) 
1949. Tadarida molossa Hershkovitz, Proc. U. S. Nat. Mus. 99:452. 

Type locality. — “America’’; “not improbably from Surinam” (Miller, 1913). 

Range. — This rare species has been recorded from Santa Anita (Shamel, 1931) on 


the basis of four U. S. N. M. specimens preserved in alcohol. 


Order LAGOMORPHA — Hares, Rabbits and Pikas 
Family LEPORIDAE — Rabbits and Hares 
Genus Sylvilagus Gray 
Sylvilagus bachmani cinerascens (J. A. Allen) 


California Brush Rabbit 


1890. Lepus cinerascens J. A. Allen, Bull. Amer. Mus. Nat. Hist. 3:159. October 8. 
1907. Sylvilagus bachmani cinerascens Nelson, Proc. Biol. Soc. Wash. 20:84. July 22. 

Type locality. — San Fernando, Los Angeles County, California. 

Range. — From the international boundary south to Ensenada, and eastward through 
the chaparral region to the northwestern slopes of the Sierra Juarez. 


Sylvilagus bachmani rosaphagus Huey 
San Quintin Brush Rabbit 


1940. Sylvilagus bachmani rosaphagus Huey, Trans. San Diego Soc. Nat. Hist. 9:221. July 31. 
Type locality. — Two miles west of Santo Domingo Mission (30°45’N, 115°58’W), 
Baja California. 
Range. — Coastal benches from Santo Tomas south to El Rosario and inland to the 


higher foothills in the mid-Sierra San Pedro Martir. 


Sylvilagus bachmani howelli Huey 
Inland Brush Rabbit 


1927. Sylvilagus bachmani howelli Huey, Trans. San Diego Soc. Nat. Hist. 5:67. July 6. 
Type locality. — 10 miles southeast of Alamo (31°35’N, 116°03’W), Baja California. 
Range. — Brushy western slopes of the southern Sierra Juarez south along the western 

chaparral mesas and higher hills bordering the northern Sierra San Pedro Martir. 


Sylvilagus bachmani exiguus Nelson 
Lower California Brush Rabbit 


1907. Sylvilagus bachmani exiguus Nelson, Proc. Biol. Soc. Wash. 20:84. July 22. 

Type locality. — Yubay, central Baja California. 

Range. — South from the vicinity of San Fernando Mission and El Marmol to the 
vicinity of Comondu. 


Sylvilagus bachmani cerrosensis (J. A. Allen) 


Cedros Island Brush Rabbit 


1898. Lepus cerrosensis J. A. Allen, Bull. Amer. Mus. Nat. Hist. 10:145. April 12. 
1909. Sylvilagus bachmani cerrosensis Nelson, North Amer. Fauna 29:255. August 31. 
Type locality. — Cerros [Cedros } Island, Baja California. 
Range. — Cedros Island. 


100 San Dreco Society oF NATURAL History { Vor. 13 


Sylvilagus bachmani peninsularis (J. A. Allen) 
Cape San Lucas Brush Rabbit 


1898. Lepus peninsularis J. A. Allen, Bull. Amer. Mus. Nat. Hist. 10:144. April 12. 
1909. Sylvilagus bachmani peninsularis Nelson, North Amer. Fauna 29:255. August 31. 
Type locality. — Santa Anita, Baja California. 
Range. — Southern part of Baja California on both coasts from below Comondu to the 
Cape. 
Sylvilagus mansuetus Nelson 


San José Island Brush Rabbit 


1907. Sylvilagus mansuetus Nelson, Proc. Biol. Soc. Wash. 20:83. July 22. 
Type locality. — San José Island, Gulf of California, Baja California. 
Range. — San José Island. 


Sylvilagus audubonii sanctidiegi (Miller) 
San Diego Cottontail 


1899. Lepus floridanus sanctidiegi Miller, Proc. Acad. Nat. Sci. Philadelphia 51:389. Octo- 
ber 5. 
1909.  Sylvilagus auduboni sanctidiegi Nelson, North Amer. Fauna 29:218. August 31. 
Type locality. — Mexican boundary monument 258, shore of the Pacific Ocean, San 
Diego County, California. 
Range. — Northwestern section of Baja California from the international boundary south 
to the El Rosario River and from the sea coast to the lower slopes of the higher mountains. 


Sylvilagus audubonii arizonae (J. A. Allen) 
Arizona Cottontail 


1877. {Lepus sylvaticus} var. arizonae J. A. Allen, in Coues and Allen, Monographs of North 
American Rodentia (U. S. Geol. Surv. Terr. Rep. Wash. 11:332). August. 
1909. Sylvilagus auduboni arizonae Nelson, North Amer. Fauna 29:222. August 31. 
Type locality. — Beal Spring, 2 miles from Kingman, Mohave County, Arizona. 
Range. — Desert section of northeastern Baja California from the international boundary 
south to the latitude of the head waters of the Gulf of California, and to the lower slopes of the 
western mountains. 


Sylvilagus audubonii confinis (J. A. Allen) 
Lower California Cottontail 


1898. Lepus arizonae confinis J. A. Allen, Bull. Amer. Mus. Nat. Hist. 10:146. April 12. 
1909. Sylvilagus auduboni confinis Nelson, North Amer. Fauna 29:220. August 31. 

Type locality. — Playa Maria, Baja California (Pacific side, 28°54’N). 

Range. — South from the El Rosario River on the west coast and Santa Rosalia on the 
east coast to the Cape. 


Genus Lepus Linnaeus 
Lepus californicus bennettii Gray 


San Diego Black-tailed Jack Rabbit 


1843. Lepus bennettii Gray, Zoology of the Voyage of H. M. S. Sulphur, under the Command 
of Capt. Sir E. Belcher, vol. 1, no. 1, Mammalia, pt. 1, p. 35, pl. 14. April. 
1909. Lepus californicus bennettii Nelson, North Amer. Fauna 29:136. August 31. 
Type locality. — San Diego, San Diego County, California. 
Range. — West of the summit of the Sierra Juarez along the international boundary 
to the coast, and southward over the coastal mesas to San Quintin. 


1963 } Hury: MAmMaALs oF BAJA CALIFORNIA 101 


Lepus californicus deserticola Mearns 
Colorado Desert Black-tailed Jack Rabbit 


1896. Lepus texianus deserticola Mearns, Proc. U. S. Nat. Mus. 18:564. June 24. 
1909. Lepus californicus deserticola Nelson, North Amer. Fauna 29: 137. August 31. 


Type locality. — Western edge of Colorado Desert, at east base of Coast Range (near 
Mexican boundary) , Imperial County, California. 


Range. — Deserts of northeastern Baja California, east of the Sierra Juarez and Sierra 


San Pedro Martir south to Calamajué Bay (Nelson, 1909). 


Lepus californicus martirensis Stowell 
San Pedro Martir Black-tailed Jack Rabbit 


1895. Lepus martirensis Stowell, Proc. Calif. Acad. Sci., ser. 2, 5:51. May 28. 
1909. Lepus californicus martirensis Nelson, North Amer. Fauna 29:152. August 31. 

Type locality. — Sierra San Pedro Martir, Baja California. 

Range. — From the southern end of the Sierra Juarez through El Valle de la Trinidad, 
southward over the Sierra San Pedro Martir to the vicinity of San Fernando Mission; from that 
latitude southward over the breadth of the peninsula to the latitude of Scammon’s Lagoon. 


Lepus californicus sheldoni Burt 
Carmen Island Black-tailed Jack Rabbit 


1933. Lepus californicus sheldoni Burt, Proc. Biol. Soc. Wash. 46:37. February 20. 
Type locality. — Carmen Island (26°N, 111°12’W), Gulf of California, Baja Cali- 


fornia. 
Range. — Carmen Island. 


Lepus californicus magdalenae Nelson 
Magdalena Island Black-tailed Jack Rabbit 


1907. Lepus californicus magdalenae Nelson, Proc. Biol. Soc. Wash. 20:81. July 22. 
Type locality. — Magdalena Island, Baja California. 


Range. — Magdalena and Margarita islands, on the Pacific side of the southern part of 
Baja California. 


Lepus californicus xanti Thomas 
Cape San Lucas Black-tailed Jack Rabbit 


1898. Lepus californicus xanti Thomas, Ann. Mag. Nat. Hist., ser. 7, 1:45. January. 

Type locality. — Santa Anita, Baja California. 

Range. — Southern part of the peninsula, from Scammon’s Lagoon on the west coast 
and Loreto on the east coast southward to the Cape. 


Lepus insularis Bryant 
Espiritu Santo Island Black Jack Rabbit 


1891. Lepus insularis Bryant, Proc. Calif. Acad. Sci., ser. 2, 3:92. April 23. He 

1895. Lepus edwardsi Saint-Loup, Bull. Mus. Hist. Nat. Paris 1:5. (Espiritu Santo Island.) 
Type locality. — Espiritu Santo Island, Gulf of California, Baja California. 
Range. — Espiritu Santo Island. 


102 San Drieco Socrety oF Natura History [Vot. 13 


Order RODENTIA — Rodents 
Family SCIURIDAE — Squirrels and Relatives 
Genus Eutamias Trouessart 
Eutamias merriami merriami (J. A. Allen) 


Merriam’s Chipmunk 


1889. Tamias asiaticus merriami J. A. Allen, Bull. Amer. Mus. Nat. Hist. 2:176. October 21. 
1897. Ej utamias | merriami Merriam, Proc. Biol. Soc. Wash. 11:191. July 1. 

Type locality. —San Bernardino Mountains, due north of San Bernardino, San Ber- 
nardino County, California. Alt. 4500 feet. 

Range. — Barely within the bounds of Baja California. 

There are specimens in M. V. Z. from the north end of Nachogiiero Valley (Johnson, 
1943133). 


Eutamias merriami obscurus (J. A. Allen) 
San Pedro Martir Chipmunk 


1890. Tamias obscurus J. A. Allen, Bull. Amer. Mus. Nat. Hist. 3:70. June. 
1909. Ef utamias} m{erriami} obscurus Nelson and Goldman, Proc. Biol. Soc. Wash. 22:23. 
March 10. 
Type locality. — Sierra San Pedro Martir, near Vallecitos, Baja California. 
Range. — The higher chaparrel and forested areas of the Sierra Juarez and Sierra San 
Pedro Martir of northern Baja California. 


Eutamias merriami meridionalis Nelson and Goldman 
Peninsula Chipmunk 


1909. Eutamias merriami meridionalis Nelson and Goldman, Proc. Biol. Soc. Wash. 22:23. 

March 10. 

Type locality. — Aguaje de San Esteban, about 25 miles northwest of San Ignacio, Baja 
California. 

Range. — Known only from the type locality and San Pablo, Baja California. 

This relict chipmunk is one of the rarest mammals living on the peninsula, where its 
range is, as far as is now known, an area less than 25 miles in diameter. It stands in strange 
contrast with its northern relatives by living in lava-bound palm-cactus associations. 


Genus Ammospermophilus Merriam 
Ammospermophilus leucurus leucurus (Merriam) 


White-tailed Antelope Squirrel; Desert Chipmunk 


1889. Tamias leucurus Merriam, North Amer. Fauna 2:20. October 30. 
1907. Ammospermophilus leucurus Mearns, U.S. Nat. Mus. Bull. 56:299. April 13. 
Type locality. — San Gorgonio Pass (east of Banning), Riverside County, California. 
Range. — Desert section of northeastern Baja California from the international boundary 
south to San Felipe on the Gulf of California. 


Ammospermophilus leucurus peninsulae (J. A. Allen) 
Western Peninsula White-tailed Antelope Squirrel 


1893. Tamias leucurus peninsulae J. A. Allen, Bull. Amer. Mus. Nat. Hist. 5:197. August 18. 
1907. Ammospermophilus leucurus peninsulae Mearns, U. S. Nat. Mus. Bull. 56:299. April 
13 


Type locality. — San Telmo, Baja California. 
Range. — Dryer western slopes of the mountains of northern Baja California from Santo 
Tomas southward to San Fernando. 


1963 | Huey: MamMMats oF BajA CALIFORNIA 103 


Ammospermophilus leucurus canfieldae Huey 
Mid-peninsula White-tailed Antelope Squirrel 


1929. Ammospermophilus leucurus canfieldae Huey, Trans. San Diego Soc. Nat. Hist. 5:243. 
February 27. 
Type locality. — Punta Prieta (28°56’N, 114° 12’W), Baja California. 
Range. — Desert region of central Baja California from Catavifia south to the vicinity 
of San Ignacio. 


Ammospermophilus leucurus extimus Nelson and Goldman 
Southern Peninsula White-tailed Antelope Squirrel 


1929. Ammospermophilus leucurus extimus Nelson and Goldman, Jour. Wash. Acad. Sci. 
19:281. July 19. 
Type locality. — Saccaton (15 miles north of Cape San Lucas) , Baja California. 
Range. — Along the eastern coast of the peninsula from Santa Rosalia south and from 
San Jorge on the west coast south to the Cape. 


Ammospermophilus insularis Nelson and Goldman 
Espiritu Santo Island Antelope Squirrel 


1909. Ammospermophilus leucurus insularis Nelson and Goldman, Proc. Biol. Soc. Wash. 
22:24. March 10. 

1959. Ammospermophilus insularis Hall and Kelson, Mamm. North Amer., p. 334. March 31. 
Type locality. — Espiritu Santo Island, Gulf of California, Baja California. 
Range. — Espiritu Santo Island. 


Genus Spermophilus Cuvier 
Spermophilus beecheyi nudipes (Huey) 
Sierra Juarez Ground Squirrel 


1931. Citellus beecheyi nudipes Huey, Trans. San Diego Soc. Nat. Hist. 7:18. October 6 
1959. Spermophilus beecheyi nudipes Hall and Kelson, Mamm. North Amer., p. 355. March 
31 


Type locality. — Laguna Hanson (31°58’N, 115°53’W), Sierra Juarez, Baja California. 
Range. — From the international boundary south to the vicinity of San Quintin, includ- 
ing the higher parts of the Sierra Juarez and the Sierra San Pedro Martir. 


Spermophilus beecheyi rupinarum (Huey) 
Catavina Ground Squirrel 

1931. Citellus beecheyi rupinarum Huey, Trans. San Diego Soc. Nat. Hist. 7:17. October 6. 
1959. Spermophilus beecheyi rupinarum Hall and Kelson, Mamm. North Amer., p. 355. 

March 31. 

Type locality. — Catavifia (29°54’N, 114°57’W/), Baja California. ie 

Range. — Deserts of central Baja California, south of the Sierra San Pedro Martir and 
San Fernando Mission; southern extent yet unknown. 


Spermophilus atricapillus W. E. Bryant 
Lower California Rock Squirrel 


1889. Spermophilus grammurus atricapillus W. E. Bryant, Proc. Calif. Acad. Sci., ser. 2, 2:26. 
June 20. 

1959. Spermophilus atricapillus Hall and Kelson, Mamm. North Amer., p. 355. March 31. 
Type locality. — Comondu, Baja California. ; 
Range. — South from latitude 28° N through the mountains to areas below Comondu; 

southern limits not now known. 


104 San Dreco Society or NAtTurRAL History { Vot. 13 


Spermophilus tereticaudus tereticaudus Baird 
Round-tailed Ground Squirrel 


1858. Spermophilus tereticaudus Baird, Mammals, in Rep. Expl. Surv. Railr. to Pacific, 
8(1) :315. July 14. 

1926. Citellus tereticaudus vociferans Huey, Proc. Biol. Soc. Wash. 39:29. July 30. (San 
Felipe, Baja California. ) 
Type locality. — Old Fort Yuma, Imperial County, California, opposite Yuma, Arizona. 
Range. — Desert region of northeastern Baja California from the international boundary 

south to San Felipe. 


Spermophilus tereticaudus apricus (Huey) 
Trinidad Valley Round-tailed Ground Squirrel 


1927. Citellus tereticaudus apricus Huey, Trans. San Diego Soc. Nat. Hist. 5:85. October 10. 
1959. Spermophilus tereticaudus apricus Hall and Kelson, Mamm. North Amer., p. 358. 
March 31. 
Type locality. — EI Valle de la Trinidad (31°20’N, 115°40’W), Baja California. 
Range. — Known only from the type locality. 


Genus Sciurus Linnaeus 
Sciurus carolinensis carolinensis Gmelin 


Eastern Gray Squirrel 


1788. {| Sciurus } carolinensis Gmelin, Caroli a Linné Systema Naturae, ed. 13, 1:148. 
1884. Sciurus carolinensis carolinensis True, Proc. U. S. Nat. Mus. 7:595. November 29. 

Type locality.— “Carolina.” 

Range. — This species was introduced on the western slopes of the Sierra San Pedro 
Martir in 1946 by Mr. E. E. Utt. 

Liberations were made in two localities. Nine squirrels were set free at La Sanja, and 
later in the same year eight more were released in Arroyo San Rafael near the head of the old 
Soccoro ditch. A letter dated December 8, 1958, regarding this introduction, was received from 
Mrs. Alberta Meling of San José, Baja California, the ranch from which the liberating expedition 
started. Mrs. Meling stated that her son Andrew accompanied Mr. Utt when the squirrels were 
turned loose. Further information in the same letter stated that squirrels had been seen one or 
two years ago at La Sanja by a now-forgotten observer and that this year (1958) Mr. Meling 
had observed squirrels in Arroyo San Rafael. 

The success of this enterprise, though contrary to the concepts of mammalogists regarding 
alien species, seems to be positive, for twelve years have passed and the species is still to be found 
in at least one area where liberated. 

The problem with this introduced species is: should it succeed in populating the forested 
area of the higher Sierra, what would happen to the native Chickaree, Tamiasciurus d. mearnst, 
that now occupies the small mountaintop arboreal habitat? Could or would they live side by 
side, or would the alien destroy and replace the native species? 


Sciurus griseus anthonyi Mearns 
Anthony’s Gray Squirrel 


1897. Sciurus fossor anthonyi Mearns, Preliminary diagnoses of new mammals of the genera 
Sciurus, Castor, Neotoma, and Sigmodon, from the Mexican border of the United 
States, p. 1. March 5. (Preprint of Proc. U.S. Nat. Mus. 20:501. January 19, 1898.) 

1907. Sciurus griseus anthonyi Mearns, U. S. Nat. Mus. Bull. 56:264. April 13. 

Type locality. — Campbell’s Ranch, Laguna Mountains, San Diego County, California. 
Range. — A single adult male specimen collected April 2, 1936, between El Rayo and 

Laguna Hanson in the Sierra Juarez by the writer marks the only known occurrence of the 

species in Baja California. 


1963 } Huey: MAMMA ts oF Baja CALIFORNIA 105 


Genus Tamiasciurus Trouessart 
Tamiasciurus douglasii mearnsi (Townsend) 
Mearns’ Chickaree 


1897. Sciurus hudsonius mearnsi Townsend, Proc. Biol. Soc. Wash. 11:146. June 9. 
1940. Tamiasciurus douglasii mearnsi Hayman and Holt in Ellerman, The families and genera 
of living rodents, British Mus. 1:348. June 8. 
Type locality. — Sierra San Pedro Martir, Baja California. Altitude about 7000 feet. 
Range. — This rather rare relict subspecies is to be found only in limited numbers in 
the higher coniferous forests of the Sierra San Pedro Martir. 
The introduction of Sciurus carolinensis into the area in 1946 may exterminate Mearns’ 
Chicakaree. 
Family GEOMYIDAE — Pocket Gophers 
Genus Thomomys Weid-Neuwied 
Thomomys umbrinus albatus Grinnell 
Imperial Valley Pocket Gopher 


1912. Thomomys albatus Grinnell, Univ. Calif. Publ. Zool. 10:172. June 7. 
1959. Thomomys umbrinus albatus Hall and Kelson, Mamm. North Amer., p. 419. March 31. 

Type locality. — West side of Colorado River at old Hanlon Ranch near Pilot Knob, 
Imperial County, California. 

Range. — Irrigated section of the region south of the international boundary to El 
Major, east to the Colorado River and west to the limits of gravity water on the east base of 
Sierra Cocopah. 

Thomomys umbrinus affinis Huey 


Jacumba Pocket Gopher 


1945. Thomomys bottae affinis Huey, Trans. San Diego Soc. Nat. Hist. 10:254. August 31. 
1959. Thomomys umbrinus affinis Hall and Kelson, Mamm. North Amer., p. 419. March 31. 
Type locality. — Jacumba, San Diego County, California. 
Range. — Jacumba Valley on both sides of the international boundary. 


Thomomys umbrinus nigricans Rhoads 
Tawny Pocket Gopher 


1895. Thomomys fulvus nigricans Rhoads, Proc. Acad. Nat. Sci. Philadelphia 47:36. Feb- 
ruary 21. 
1959. Thomomys umbrinus nigricans Hall and Kelson, Mamm. North Amer., p. 429. March 
Sie 
Type locality. — Witch Creek, 7 miles west of Julian, San Diego County, California. 
Alt. 2755 feet. 
Range. — Along the international boundary from Nachogiero Valley westward to El 
Valle de las Palmas, thence south to Las Cruces, some 15 miles inland from Ensenada. This 
subspecies does not at any point reach the sea coast. 


Thomomys umbrinus sanctidiegi Huey 
San Diego Pocket Gopher 


1945. Thomomys bottae sanctidiegi Huey, Trans. San Diego Soc. Nat. Hist. 10:258. August 
SA: 
1959. Thomomys umbrinus sanctidiegi Hall and Kelson, Mamm. North Amer., p. 432. March 
31. 
Type locality. — Balboa Park, San Diego, San Diego County, California. 
Range. — Coastal strip from the international boundary south to the vicinity of Ensenada. 
The type specimen of this race was captured alive within the Natural History Museum 
building in Balboa Park where it had wandered through an open door during daylight hours! 


106 SAN Deco Society or Natura History | Vous 


Thomomys umbrinus lucidus Hall 
Laguna Salada Pocket Gopher 
1932. Thomomys bottae lucidus Hall, Proc. Biol. Soc. Wash. 45:67. April 2. 
1959. Thomomys umbrinus lucidus Hall and Kelson, Mamm. North Amer., p. 427. March 31. 
Type locality. — Las Palmas Canyon, mesquite association, 2 miles east of Gaskill’s 
Tank, west side of Laguna Salada (north of latitude 32°N), Baja California. 
Range. — Known only from the type locality. 


Thomomys umbrinus juarezensis Huey 
Sierra Juarez Pocket Gopher 
1945. Thomomys bottae juarezensis Huey, Trans. San Diego Soc. Nat. Hist. 10:255. August 
ae 
1959. Thomomys umbrinus juarezensis Hall and Kelson, Mamm. North Amer., p. 426. March 
ile 
Type locality. — Laguna Hanson, Sierra Juarez, Baja California. 
Range. — Known from the forested area of the higher parts of the Sierra Juarez. 


Thomomys umbrinus cunicularius Huey 
Pattie Basin Pocket Gopher 

1945. Thomomys bottae cunicularius Huey, Trans. San Diego Soc. Nat. Hist. 10:252. August 

Sil: 
1959. Thomomys umbrinus cunicularius Hall and Kelson, Mamm. North Amer., p. 423. 

March 31. 

Type locality. — Los Palmitos, western end of Pattie Basin (31°44’N, 115°36’W), on 
southeastern slope of Sierra Juarez, Baja California. 

Range. — Known only from the type locality. 


Thomomys umbrinus jojobae Huey 
Sangre de Cristo Pocket Gopher 
1945. Thomomys bottae jojobae Huey, Trans. San Diego Soc. Nat. Hist. 10:256. August 31. 
1959. Thomomys umbrinus jojobae Hall and Kelson, Mamm. North Amer., p. 426. March 31. 
Type locality. — Sangre de Cristo (31°52’N, 116°06’W) , Baja California. 
Range. — Western foothills of the Sierra Juarez, in Valle de San Rafael. 


Thomomys umbrinus proximarinus Huey 
Coastal Pocket Gopher 
1945. Thomomys bottae proximarinus Huey, Trans. San Diego Soc. Nat. Hist. 10:261. August 
Bile 


1959. Thomomys umbrinus proximarinus Hall and Kelson, Mamm. North Amer., p. 431. 
March 31. 
Type locality.— Boca la Playa (31°32’N, 116°38’W), mesa bordering the sea, 16 
miles west of Santo Tomas, Baja California. 
Range. — Known only from the type locality. 


Thomomys umbrinus aphrastus Elliot 
Perplexing Pocket Gopher 
1903. Thomomys aphrastus Elliot, Field Columb. Mus. Publ. 79, Zool. Ser. 3:219. August 15. 
1959. Thomomys umbrinus aphrastus Hall and Kelson, Mamm. North Amer., p. 420. March 

31: 

Type locality. — San [| =Santo] Tomas, 18 miles south of Ensenada, Baja California. 
Alt. 50-100 feet. 

Range. — From Santo Tomas in Santo Tomas Valley eastward to extreme western end 
of El Valle de la Trinidad, thence southward along the foothills of the Sierra San Pedro Martir 
at least to Las Cabras. Southward from Santo Tomas, the range of this race reaches the coast at 
Johnson’s Ranch (Rancho San Antonio) ; thence over the coastal plain to or below San Quintin. 


1963 } Hury: MAmMMALs oF BAjJA CALIFORNIA 107 


Thomomys umbrinus xerophilus Huey 
San Matias Pass Pocket Gopher 


1945. Thomomys bottae xerophilus Huey, Trans. San Diego Soc. Nat. Hist. 10:257. August 
311. 
1959. Thomomys umbrinus xerophilus Hall and Kelson, Mamm. North Amer., p- 434. March 
Bile 
Type locality. — Near Diablito Spring, summit of San Matias Pass, between Sierra 
Juarez and Sierra San Pedro Martir, Baja California. 
Range. — San Matias Pass and eastern section of El Valle de la Trinidad at least to 
Aguaita Spring. 
Specimens from the western end of El Valle de la Trinidad are intergrades and referable 


to T. b. aphrastus. 


Thomomys umbrinus martirensis J. A. Allen 
San Pedro Martir Pocket Gopher 
1898. Thomomys fulvus martirensis J. A. Allen, Bull. Amer. Mus. Nat. Hist. 10:147. April 
12 


1959. Tharctiye umbrinus martirensis Hall and Kelson, Mamm. North Amer., p. 427. March 
31. 
Type locality. —La Grulla Meadow, Sierra San Pedro Martir, Baja California. Alt. 
7400 feet. 
Range. — Higher levels of the Sierra San Pedro Martir. 


Thomomys umbrinus siccovallis Huey 
El Cajén Cafion Pocket Gopher 


1945. Thomomys bottae siccovallis Huey, Trans. San Diego Soc. Nat. Hist. 10:258. August 
31. 

1959. Thomomys umbrinus siccovallis Hall and Kelson, Mamm. North Amer., p. 433. March 
31 


Type locality. — El Cajén Caton (30°54’N, 115°10’W), east base of Sierra San 
Pedro Martir, Baja California. Alt. 3200 feet. 

Range. — From the type locality on the low east base of the Sierra San Pedro Martir 
south to Mattomi (Hall and Kelson, 1952b). 


Thomomys umbrinus abbotti Huey 
El Rosario Pocket Gopher 


1928. Thomomys bottae abbotti Huey, Trans. San Diego Soc. Nat. Hist. 5:89. January 18. 
1959. Thomomys umbrinus abbotti Hall and Kelson, Mamm. North Amer., p. 416. March 31. 
Type locality. — One mile east of El Rosario (30°03’N, 115°48’W), Baja California. 
Range. — Known only from the environs of the river bottom association at the type 
locality. 


Thomomys umbrinus brazierhowelli Huey 
San Fernando Pocket Gopher 


1960. Thomomys umbrinus brazierhowelli Huey, Trans. San Diego Soc. Nat. Hist. 12:407. 
February 1. ; 
Type locality — San Fernando Mission, latitude 30° N, Baja California. 
Range. — Known from the type locality and eastward over the alluvial silt soils of Llano 


de San Agustin. 


108 San Dreco Society oF NaATuRAL HIstTory { VoL. 13 


Thomomys umbrinus catavinensis Huey 
Catavina Pocket Gopher 


1931. Thomomys bottae catavinensis Huey, Trans. San Diego Soc. Nat. Hist. 7:45. December 
19. 

1959. Thomomys umbrinus catavinensis Hall and Kelson, Mamm. North Amer., p. 422. 
March 31. 
Type locality. — Catavifia (29°54’N, 114°57’W), Baja California. 
Range. — Known only from the palm-filled valley at the type locality. 


Thomomys umbrinus ruricola Huey 
Santa Catarina Pocket Gopher 


1949. Thomomys bottae ruricola Huey, Trans. San Diego Soc. Nat. Hist. 11:53. January 31. 
1959. Thomomys umbrinus ruricola Hall and Kelson, Mamm. North Amer., p. 432. March 31. 

Type locality. — Four miles north of Santa Catarina Landing (29°35’N, 115°17’W), 
Baja California. 

Range. — Known from the type locality and Rancho Ramona, a locality some four miles 
northeast of Santa Catarina. 

The type specimen was caught in a mouse trap while away from its burrow at night. 


Thomomys umbrinus cactophilus Huey 
Punta Prieta Pocket Gopher 


1929. Thomomys bottae cactophilus Huey, Trans. San Diego Soc. Nat. Hist. 5:241. February 
De 

1959. Thomomys umbrinus cactophilus Hall and Kelson, Mamm. North Amer., p. 421. March 
31 


Tare locality. — Punta Prieta (28°56’N, 114°12’W), Baja California. 
Range. — Known from the type locality and an area near the Pacific coast at Santa 
Rosalia Bay. 


Thomomys umbrinus borjasensis Huey 
San Borjas Pocket Gopher 


1945. Thomomys bottae borjasensis Huey, Trans. San Diego Soc. Nat. Hist. 10:262. August 
a 

1959. Thomomys umbrinus borjasensis Hall and Kelson, Mamm. North Amer., p. 421. March 
31. 
Type locality. — San Borjas Mission (28°52’N, 113°53’W), Baja California. 
Range. — Known only from the type locality. 


Thomomys umbrinus rhizophagus Huey 
Los Angeles Bay Pocket Gopher 


1949. Thomomys bottae rhizophagus Huey, Trans. San Diego Soc. Nat. Hist. 11:54. January 
a1, 
1959. Thomomys umbrinus rhizophagus Hall and Kelson, Mamm. North Amer., p. 432. 
March 31. 
Type locality. — Los Flores, 7 miles south of Bahia de los Angeles (28°50’N, 113° 
32’W) , Baja California. 
Range. — Known only from the type locality. 


1963 } Huey: Mammats oF Baja CALIFORNIA 109 


Thomomys umbrinus russeolus Nelson and Goldman 
San Angel Pocket Gopher 
1909. Thomomys bottae russeolus Nelson and Goldman, Proc. Biol. Soc. Wash. 22:25. 
March 10. 


1959. Thomomys umbrinus russeolus Hall and Kelson, Mamm. North Amer., p. 432. March 
Bile 


Type locality. —San Angel, 30 miles west of San Ignacio, Baja California. 


Range. — Found on the eastern side of the Vizcaino Desert. 


Thomomys umbrinus homorus Huey 
Rancho Lagunitas Pocket Gopher 


1949. Thomomys bottae homorus Huey, Trans. San Diego Soc. Nat. Hist. 11:55. January 31. 
1959. Thomomys umbrinus homorus Hall and Kelson, Mamm. North Amer., p. 425. March 
31 


Type locality.— One mile east of Rancho Lagunitas (28°20’N, 113°15’W), Baja 
California. 

Range. —Local populations are found in suitable localities in hilly sections of the 
extreme northeastern Vizcaino Desert from the vicinity of Calmalli (1200 feet) eastward to 
the summit of the peninsular ridge near the type locality. 


Thomomys umbrinus incomptus Goldman 
Magdalena Plain Pocket Gopher 
1939. Thomomys bottae incomptus Goldman, Proc. Biol. Soc. Wash. 52:29. March 11. 


1959. Thomomys umbrinus incomptus Hall and Kelson, Mamm. North Amer., p. 426. March 

Bile 

Type locality. — San Jorge, near Pacific coast, west of Pozo Grande and about 25 miles 
southwest of Comondiu, Baja California. Alt. 50 feet. 

Range. — Northern half of the vast Magdalena Plain with the exception of the very 
restricted coastal habitat of T.u. litoris (Huey, 1945a). 


Thomomys umbrinus litoris Burt 
Magdalena Bay Pocket Gopher 
1940. Thomomys bottae litoris Burt, Occ. Pap. Mus. Zool. Univ. Michigan 424:1. November 
29 


1959. Thomomys umbrinus litoris Hall and Kelson, Mamm. North Amer., p. 427. March 31. 
Type locality. — Stearns Point, west side of Magdalena Bay, Baja California. 
Range. — Known only from the type locality. 


Thomomys umbrinus magdalenae Nelson and Goldman 
Magdalena Island Pocket Gopher 
1909. Thomomys magdalenae Nelson and Goldman, Proc. Biol. Soc. Wash. 22:24. March 10. 
1959. Thomomys umbrinus magdalenae Hall and Kelson, Mamm. North Amer., p. 427. March 
Sie 
Type locality. — Magdalena Island, Baja California. 
Range. — Known only from the type locality. 


Thomomys umbrinus imitabilis Goldman 
La Paz Pocket Gopher 


1939. Thomomys bottae imitabilis Goldman, Proc. Biol. Soc. Wash. 52:30. March 11. 
1959. Thomomys umbrinus imitabilis Hall and Kelson, Mamm. North Amer., p. 426. March 
34 


Type locality. — La Paz, Baja California. 
Range. — Known only from the type locality. 


110 San Dteco Society oF Naturat History { Vot. 13 


Thomomys umbrinus alticolus J. A. Allen 


Sierra Laguna Pocket Gopher 


1899. Thomomys fulvus alticolus J. A. Allen, Bull. Amer. Mus Nat. Hist. 12:13. March 4. 


1959. Thomomys umbrinus alticolus Hall and Kelson, Mamm. North Amer., p. 419. March 
2 il, 


Type locality. — Sierra Laguna, Baja California. 
Range. — Higher sections of the Victoria Mountains in the Cape District. 


Thomomys umbrinus anitae J. A. Allen 
Cape San Lucas Pocket Gopher 


1898. Thomomys fulvus anitae J. A. Allen, Bull. Amer. Mus. Nat. Hist. 10:146. April 12. 
1959. Thomomys umbrinus anitae Hall and Kelson, Mamm. North Amer., p. 420. March 31. 
Type locality. — Santa Anita, Baja California. 


Range. — Lower levels of the Cape District, south of the Magdalena Plain, except the 
section about La Paz, which is occupied by T.u. imitabilis (Huey, 1945a). 


Family HETEROMYIDAE — Heteromyid Rodents 
Genus Perognathus Wied-Neuwied 
longimembris Group 
Perognathus longimembris internationalis Huey 


Jacumba Valley Silky Pocket Mouse 


1939. Perognathus longimembris internationalis Huey, Trans. San Diego Soc. Nat. Hist. 9:47. 
August 31. 


Type locality. — Baja California side of international boundary at Jacumba, San Diego 
County, California. 
Range. — Known only from both sides of the international boundary in Jacumba Valley. 


Perognathus longimembris aestivus Huey 
San Rafael Valley Silky Pocket Mouse 


1928. Perognathus longimembris aestivus Huey, Trans. San Diego Soc. Nat. Hist. 5:87. 
January 18. 
Type locality. — Sangre de Cristo (31°52’N, 116°06’W), Valle de San Rafael, western 
base of the Sierra Juarez, Baja California. 


Range. — Known from the type locality in Valle de San Rafael and from the eastern 
end of El Valle de la Trinidad, some 20 miles south (Huey, 1939:49). 


Perognathus longimembris venustus Huey 
San Agustin Silky Pocket Mouse 


1930. Perognathus longimembris venustus Huey, Trans. San Diego Soc. Nat. Hist. 6:233. 
December 24. 
Type locality. — San Agustin (30°N, 115°W), Baja California. 
Range. — Known only from the type locality. 
Collecting in suitable areas and at an auspicious season will, without doubt, extend the 
range southward. 


1963 } Hurry: MAmmats oF Baja CALIFORNIA 111 


Perognathus longimembris bombycinus Osgood 
Yuma Silky Pocket Mouse 


1907. Perognathus bombycinus Osgood, Proc. Biol. Soc. Wash. 20:19. February 23. 
1929. Perognathus longimembris bombycinus Nelson and Goldman, Proc. Biol. Soc. Wash. 
42:104. March 25. 
Type locality. — Yuma, Yuma County, Arizona. 
Range. —In suitable sandy areas from the international boundary at Algodones, on 
the Colorado River, south to San Felipe, on the Gulf of California. 
Specimens from the latter locality are in the S. D. N. H. M. 


formosus Group 
Perognathus formosus mesembrinus Elliot 


Southwestern Long-tailed Pocket Mouse 


1904. Perognathus mesembrinus Elliot, Field Columb. Mus. Publ. 87, Zool. Ser. 3:251. 
January 7. 


1929. Perognathus formosus mesembrinus Nelson and Goldman, Proc. Biol. Soc. Wash. 
42:106. March 25. 

Type locality. — Palm Springs, Riverside County, California. 

Range. — Desert area from the international boundary south to the end of Laguna 


Salada. 


Perognathus formosus cinerascens Nelson and Goldman 
Ashy Pocket Mouse 


1929. Perognathus formosus cinerascens Nelson and Goldman, Proc. Biol. Soc. Wash. 42:105. 
March 25. 
Type locality. — San Felipe, northeastern Baja California. 
Range. — Arid rocky hills in desert area from 35 miles north of San Felipe to San Felipe. 
Without doubt this subspecies lives in the arid area bordering the Gulf for an as yet 
undetermined distance southward. 


Perognathus formosus infolatus Huey 


San Francisquito Pocket Mouse 


1954. Perognathus formosus infolatus Huey, Trans. San Diego Soc. Nat. Hist. 12:1. March 1. 
Type locality. —7 miles west of San Francisquito Bay (28°30’N), Gulf of California, 
Baja California. 
Range. — The desert areas bordering the Gulf of California from El Marmol south to 
Barril (28°06’N). 
baileyi Group 
Perognathus baileyi hueyi Nelson and Goldman 


Huey Pocket Mouse 


1929. Perognathus baileyi hueyi Nelson and Goldman, Proc. Biol. Soc. Wash. 42:106. March 

25. 

Type locality. — San Felipe, northeastern Baja California. 

Range. — Desert slopes of the coast range mountains from the international boundary 
southward to the region east of the southern extent of the Sierra San Pedro Martir. Southern 
limits are unknown. 


112 San Drieco Society of Naturav History [ VoL. 13 


Perognathus baileyi rudinoris Elliot 
Western Bailey Pocket Mouse 


1903. Perognathus baileyi rudinoris Elliot, Field Columb. Mus. Publ. 74, Zool. Ser. 3:167. 
May 7. 
1903. Pe aibae knekus Elliot, Field Columb. Mus. Publ. 74, Zool. Ser. 3:169. May 7. 
(Rosarito, Sierra San Pedro Martir, Baja California. Regarded as an extra large specimen 
of P. b. rudinoris by Nelson and Goldman, 1930.) 
Type locality. — San Quintin, Baja California. 
Range. — The coastal plain about San Quintin and foothill slopes of the southern end 
of the Sierra San Pedro Martir south to the region of El Marmol and San Fernando Mission. 


Perognathus baileyi mesidios' subsp. nov. 
Midpeninsula Bailey Pocket Mouse 


Type. — Adult male; from San Borja Mission, near latitude 28°45’N, Baja California, 
Mexico; collected by Laurence M. Huey, October 13, 1941; no. 14470, San Diego Natural 
History Museum. 

Diagnosis. — Perognathus baileyi mesidios is recognizable by its silky, grizzled dorsal 
pelage with a darker buff median line and long tail thickly haired over the outer half of its 
length. Its skull is broad and flat across the parietals with swollen audital bullae. 

Comparisons. — Compared with Perognathus baileyi extimus from the Cape District, 
P. b. mesidios has more grizzled dorsal pelage with darker colored sides and brighter, broader 
median buff stripes. The skull is more nearly flat and broader and has slightly more swollen, less 
truncated audital bullae. Compared with Perognathus baileyi hueyi from the San Felipe district, 
P. b. mesidios has darker and more grizzled pelage dorsally. The skull is not so broad and is 
slightly more rounded and not so flattened in profile. Compared with Perognathus baileyi rudi- 
noris from the El Marmol-San Quintin district, P. 6. mesidios is lighter in dorsal pelage color. 
The skull has less inflated bullae with slightly more slender nasals. 

Measurements of type.— Total length, 212; tail, 121; hind foot, 25; ear, 6; greatest 
length of skull, 29.7; width across bullae, 15.6; interorbital constriction, 7.0; nasals, 11.7; tooth- 
row, 4.1. : 

Range. — South from Catavifia to Conception Bay. 


Perognathus baileyi fornicatus Burt 
Monserrate Island Pocket Mouse 
1932. Perognathus baileyi fornicatus Burt, Trans. San Diego Soc. Nat. Hist. 7:164. October 
ele 
Type locality. — Monserrate Island (25°38’N, 111°02’W), Gulf of California, Baja 


California. 
Range. — Monserrate Island. 


Perognathus baileyi extimus Nelson and Goldman 
San Lucas Pocket Mouse 


1930. Perognathus baileyi extimus Nelson and Goldman, Jour. Wash. Acad. Sci. 20:223. 
June 19. 
Type locality. — Tres Pachitas, 36 miles south of La Paz, Baja California. 
Range. — Cape District of the peninsula. 


' From the Greek, mesidios, middle. 


1963} Huey: MAMMALS oF Baja CALIFORNIA 113 


penicillatus Group 
Perognathus penicillatus angustirostris Osgood 


Colorado Desert Pocket Mouse 


1900. Perognathus penicillatus angustirostris Osgood, North Amer. Fauna 18:47. September 

20. 

Type locality. — Carrizo Creek, western edge of Colorado Desert, Imperial County, 
California. 

Range. — Suitable areas of soft sandy soil along the international boundary from Algo- 
dones west to the slopes of the coastal mountains and south to San Felipe. Southern limits not 
known. 

Perognathus arenarius mexicalis Huey 
Laguna Salada Sand Pocket Mouse 
1939. Perognathus arenarius mexicalis Huey, Trans. San Diego Soc. Nat. Hist. 9:57. August 
ale 

Type locality.— Los Muertos Canyon fan (32°27’N, 115°53’W), in Palo Verde- 
Ironwood association at Gaskill’s Tank, near Laguna Salada, northern Baja California. 

Range. — Found on sandy ground on the western side of Laguna Salada. 


Perognathus arenarius albescens Huey 
San Felipe Sand Pocket Mouse 


1926. Perognathus arenarius albescens Huey, Proc. Biol. Soc. Wash. 39:67. July 30. 

Type locality. — San Felipe, Baja California. 

Range. — Sandy ground from about 40 miles north of San Felipe southward along the 
Gulf slope for an undetermined distance, possibly to Los Angeles Bay. 


Perognathus arenarius helleri Elliot 
San Quintin Sand Pocket Mouse 


1903. Perognathus helleri Elliot, Field Columb. Mus. Publ. 74, Zool. Ser. 3: 166. May 7. 
1926. Perognathus arenarius helleri Huey, Proc. Biol. Soc. Wash. 39:68. July 30. 

Type locality. — San Quintin, northwestern Baja California. 

Range. — Sandy areas over the San Quintin plain, northwestern Baja California. 


Perognathus arenarius ambiguus Nelson and Goldman 
Yubay Sand Pocket Mouse 


1929. Perognathus arenarius ambiguus Nelson and Goldman, Proc. Biol. Soc. Wash. 42:108. 
March 25. 
Type locality. — Yubay, 30 miles southeast of Calamahué, Baja California. 
Range. — Central section of the peninsula from Chapala southward to the middle of 
the Vizcaino Desert. 


Perognathus arenarius paralios' subsp. nov. 
Barril Sand Pocket Mouse 


Type. — Adult male; from Barril (28°20’N), on the Gulf of California, Baja Cali- 
fornia, Mexico; collected by Laurence M. Huey, March 23, 1947; no. 15542, San Diego Natural 
History Museum. 

Diagnosis. — Perognathus arenarius paralios is extremely pale, almost white with a tinge 
of brown over the lightly grizzled dorsal area. The side stripes are obscure or missing on some 
specimens and the tail stripe is notably pale. The skull of P. a. paralios is large and is flat across 
the brain case. 


1 From the Greek, paralios, by or near the sea. In reference to the coastal range of this race. 


114 San Dieco Society oF NAtTuRAL History { Vor. 13 


Comparisons. — Perognathus arenarius paralios differs from its northern coastal relative, 
Perognathus arenarius albescens in being slightly larger and in having a brownish cast to its dorsal 
pelage. The tail stripe is notably pale and the brownish side stripes found on the darker subspecies 
of this species are obscure or missing on most specimens of P. a. paralios. From the subspecies 
Perognathus arenarius ambiguus, which lives in higher altitudes westward on the Pacific side 
of the peninsula, this newly named subspecies differs in being decidedly paler, and but little 
comparison is required to separate specimens. Compared with Perognathus arenarius sublucidus 
from La Paz, in the Cape District, P. a. paralios differs even more; P.a. paralios is pale and 
appears to be nearly white. Cranially, P. a. paralios differs from all three of the geographically 
adjoining subspecies in minor characters but is, in the main, larger and the braincase is more 
nearly flat. 

Measurements of type. — Total length, 150; tail, 81; hind foot, 22; ear, 5; greatest length 
of skull, 24.0; mastoid breadth, 13.0; interorbital constriction, 6.3; nasals, 8.9; toothrow, 2.9. 

Range. — Fairly abundant, over sandy stretches on the Gulf slope, mainly near the 
shore, from the type locality northward to the vicinity of Los Angeles Bay, where intergradation 
takes place with the more northern subspecies, P. a. albescens. 

Series of specimens in the S. D. N. H. M. are from Barril, San Francisquito Bay, Los 
Flores and Los Angeles Bay; the latter are not typical. 


Perognathus arenarius sabulosus' subsp. nov. 
Scammon Lagoon Sand Pocket Mouse 


Type. — Adult male; from mainland on south side of Scammon’s Lagoon, Baja Cali- 
fornia, Mexico; collected by Laurence M. Huey, May 22, 1926; no. 5300, San Diego Natural 
History Museum. 

Diagnosis. — Perognathus arenarius sabulosus is pallid in color on the face, sides of 
head, and body. Dorsally the pelage is slightly grizzled. The median side stripe is light brown 
and very faintly marked, as is the upper side of the tail. The skull of this subspecies is dome-like 
across the brain case and has a compressed appearance when viewed dorsally. 

Comparisons. — Compared with Perognathus arenarius ambiguus, whose range lies north 
and east at higher altitudes, P. a. sabulosus is paler in color and has a narrower, dome-shaped 
skull. Compared with Perognathus arenarius arenarius, whose range lies south of San Ignacio 
Lagoon and over the Magdalena Plain, P. a. sabulosus is paler in overall color, is less grizzled 
dorsally and the side stripes and stripe on top of the tail are paler and less heavily marked. The 
skull is more dome-shaped and not as broad across the bullae. 

Measurements of type. — Total length, 165; tail, 91; hind foot, 21; ear, 5; greatest 
length of skull, 23.5; mastoid breadth, 12.3; interorbital constriction, 6.1; nasals, 8.8; toothrow, 2.9. 

Range. — Sparsely distributed over the bleak sandy areas about the lagoons on the 
western Vizcaino Desert, southward to the northern part of San Ignacio Lagoon and westward 
over the desolate wind-blown reaches of the triangular shaped area to Point Eugenia. Specimens 
from the vicinity of the type locality and non-typical specimens coastwise from north of the 


type locality at Santo Domingo Landing and Santa Rosalia Bay are in the S. D. N. H. M. 


Perognathus arenarius arenarius Merriam 
San Jorge Sand Pocket Mouse 


1894. Perognathus arenarius Merriam, Proc. Calif. Acad. Sci., ser. 2, 4:461. September 25. 
1926. Perognathus arenarius arenarius Huey, Proc. Biol. Soc. Wash. 39:68. July 30. 

Type locality. — San Jorge, near Comondu, Baja California. 

Range. — Northern section of Magdalena Plain and southern part of the Vizcaino Desert, 
on the Pacific side of the peninsula. 


' From the Greek sabulosus, sandy, in reference to the sandy habitat of this subspecies. 


1963 } Hury: MAammMats oF BajJA CALIFORNIA 115 


Perognathus arenarius sublucidus Nelson and Goldman 
La Paz Sand Pocket Mouse 


1929. Perognathus arenarius sublucidus Nelson and Goldman, Proc. Biol. Soc. Wash. 41:109. 
March 25. 
Type locality. — La Paz, Baja California. 
Range. — Sandy sections of the desert basin in the La Paz region. 


Perognathus arenarius albulus Nelson and Goldman 
Magdalena Island Sand Pocket Mouse 


1923. Perognathus penicillatus albulus Nelson and Goldman, Proc. Biol. Soc. Wash. 36:159. 
May 1. 

1926. Perognathus arenarius albulus Huey, Proc. Biol. Soc. Wash. 39:68. July 30. 
Type locality. — Magdalena Island, off west coast of Baja California. 
Range. — Magdalena Island. 


Perognathus arenarius ammophilus Osgood 
Santa Margarita Island Sand Pocket Mouse 


1907. Perognathus penicillatus ammophilus Osgood, Proc. Biol. Soc. Wash. 20:20. February 
23: 

1926. Perognathus arenarius ammophilus Huey, Proc. Biol. Soc. Wash. 39:68. July 30. 
Type locality. —Santa Margarita Island, off the west coast of Baja California. 
Range. — Santa Margarita Island. 


Perognathus arenarius siccus Osgood 
Cerralvo Island Sand Pocket Mouse 


1907. Perognathus penicillatus siccus Osgood, Proc. Biol. Soc. Wash. 20:20. February 23. 
1929. Perognathus arenarius siccus Nelson and Goldman, Proc. Biol. Soc. Wash. 42:108. 
March 25. 
Type locality. — Cerralvo Island, Gulf of California, Baja California. 
Range. — Cerralvo Island. 


intermedius Group 
Perognathus fallax fallax Merriam 


San Diego Short-eared Pocket Mouse 


1889. Perognathus fallax Merriam, North Amer. Fauna 1:19. October 25. 

Type locality. — Reche Canyon, 3 miles southeast of Colton, San Bernardino County, 
California. Alt. 1250 feet. 

Range. — Northern Baja California along the international boundary from Jacumba 
west to the Pacific Ocean, south to El Valle de la Trinidad and to Ensenada, mainly on the 
western slopes of the mountains. 


Perognathus fallax pallidus Mearns 
Pallid Short-eared Pocket Mouse 


1901. Perognathus fallax pallidus Mearns, Proc. Biol. Soc. Wash. 14:135. August 9. 

Type locality. — Mountain Spring, east slope of Coast Range near Mexican boundary, 
in extreme western Imperial County, California. Alt. 2500 feet. 

Range. — The type locality of this subspecies is within one mile of the Mexican boundary, 
where it is fairly abundant. Continuous habitat extends southward over the international line, 
where the subspecies is to be found in Baja California. 

Trapping in likely habitats at two localities 25 miles and 60 miles southward failed to 
capture P. fallax. 


116 SAN Dieco Society oF NaturAL History { Vo. 13 


Perognathus fallax majusculus Huey 
San Quintin Short-eared Pocket Mouse 
1960. Perognathus fallax majusculus Huey, Trans. San Diego Soc. Nat. Hist. 12:418. February 
1 


Type locality. — San Quintin, Baja California. 
Range. — Coastal area from below Santo Tomas south to El Rosario, eastward to the 


base of the Sierra San Pedro Martir. 


Perognathus fallax xerotrophicus Huey 
Chapala Short-eared Pocket Mouse 


1960. Perognathus fallax xerotrophicus Huey, Trans. San Diego Soc. Nat. Hist. 12:419. 
February 1. 
Type locality. — Two miles northwest of Chapala, Baja California. 
Range. — Inland plains on the western slope from San Fernando south to the eastern 
Vizcaino Desert area. 


Perognathus fallax inopinus Nelson and Goldman 
Turtle Bay Short-eared Pocket Mouse 


1929. Perognathus fallax inopinus Nelson and Goldman, Proc. Biol. Soc. Wash. 42:110. 
March 25. 
Type locality. — Turtle (San Bartolomé) Bay, west coast of Baja California. 
Range. — Coastal strip from Santa Catarina Landing south to the type locality. 


Perognathus anthonyi Osgood 
Cedros Island Pocket Mouse 


1900. Perognathus anthonyi Osgood, North Amer. Fauna 18:56. September 20. 
Type locality. — South Bay, Cerros (Cedros) Island, Baja California, Mexico. 
Range. — Known only from Cedros Island. 


californicus Group 
Perognathus californicus femoralis J. A. Allen 


Dulzura Pocket Mouse 


1891. Perognathus (Chaetodipus) femoralis J. A. Allen, Bull. Amer. Mus. Nat. Hist. 3:281. 
June 30. 
1913. Perognathus californicus femoralis Grinnell, Proc. Calif. Acad. Sci., ser. 4, 3:335. 
August 28. 
Type locality. — Dulzura, San Diego County, California. 
Range. — Brushy foothills area on the western slopes of the mountains from the boundary 
at Jacumba Valley westward to Tijuana, thence south to the vicinity of Ensenada and the north- 
western slopes of the Sierra San Pedro Martir. 


Perognathus californicus mesopolius Elliot 
San Pedro Martir Pocket Mouse 


1903. Perognathus femoralis mesopolius Elliot, Field Columb. Mus. Publ. 74, Zool. Ser. 3:168. 
May 7. 

1955. Perognathus californicus mesopolius Miller and Kellogg, U. S. Nat. Mus. Bull. 205:280. 
March 3. 
Type locality. — Pifién, Sierra San Pedro Martir, Baja California. 
Range. — Sierra San Pedro Martir. 


1963 } Huey: MAammMats oF Baja CALIFORNIA 117 


spinatus Group 
Perognathus spinatus spinatus Merriam 
Needles Spiny Pocket Mouse 


1889. Perognathus spinatus Merriam, North. Amer. Fauna 1:21. October 25. 
Type locality. —25 miles below Needles, Colorado River, San Bernardino County, 


California. 
Range. — This subspecies extends from the international boundary on the south slopes 
of Pilot Knob, in the extreme northeastern corner of Baja California, south through the Sierra 


Cocopah to San Felipe. 


Perognathus spinatus rufescens Huey 
Western Spiny Pocket Mouse 
1930. Perognathus spinatus rufescens Huey, Trans. San Diego Soc. Nat. Hist. 6:231. Decem- 
ber 24. 


Type locality.— Mouth of Palm Canyon, Borrego Valley, northeastern San Diego 
County, California. 


Range. — The desert slopes of the Sierra Juarez from the international boundary south- 
ward; the limits of this subspecies are as yet unknown. 


Perognathus spinatus oribates Huey 
San Fernando Spiny Pocket Mouse 
1960. Perognathus spinatus oribates Huey, Trans. San Diego Soc. Nat. Hist. 12:409. February 
1 


Type locality. — San Fernando Mission (30°N), Baja California. 

Range. — Rocky terrain over the western foothills bordering Llano de San Agustin at 
latitude 30°N, northward along the arid rocky foothills of the western slopes of the Sierra San 
Pedro Martir to Las Cabras, inland and east of San Telmo near latitude 31°N. 


Perognathus spinatus evermanni Nelson and Goldman 
Mejia Island Spiny Pocket Mouse 


1929. Perognathus evermanni Nelson and Goldman, Proc. Biol. Soc. Wash. 42:111. March 25. 
1932. Perognathus spinatus evermanni Burt, Trans. San Diego Soc. Nat. Hist. 7:165. October 
Sih 


Type locality. — Mejia Island, north of Angel de la Guardia Island, Gulf of California, 
Baja California. 


Range. — Mejia Island. 


Perognathus spinatus guardiae Burt 
Guardian Island Spiny Pocket Mouse 


1932. Perognathus spinatus guardiae Burt, Trans. San Diego Soc. Nat. Hist. 7:165. October 
i) 
Type locality. — Puerto Refugio, north end of Angel de la Guardia Island, Gulf of 
California, Baja California. Alt. 30 feet. 
Range. — Angel de la Guardia Island. 


Perognathus spinatus prietae Huey 


Mid-peninsula Spiny Pocket Mouse 
1930. Perognathus spinatus prietae Huey, Trans. San Diego Soc. Nat. Hist. 6:232. December 
24 


Type locality. — 25 miles north of Punta Prieta (29°24’N, 114°24’W), Baja California. 


Range. — From Catavifia south over the rocky parts of the peninsula to the vicinity of 
Santa Gertrudis Mission and Barril on the Gulf. 


118 SAN Drieco Society oF Naturat History {Vot. 13 


Perognathus spinatus broccus Huey 
San Ignacio Spiny Pocket Mouse 
1960. Perognathus spinatus broccus Huey, Trans. San Diego Soc. Nat. Hist. 12:410. February 


IF 
Type locality. — San Ignacio (27°17’N), Baja California. 
Range. — Over the lava-covered slopes of the Sierra de la Giganta. 


Perognathus spinatus marcosensis Burt 
San Marcos Island Spiny Pocket Mouse 


1932. Perognathus spinatus marcosensis Burt, Trans. San Diego Soc. Nat. Hist. 7:166. 


October 31. 
Type locality. —San Marcos Island (27°13’N, 112°05’W), Gulf of California, Baja 


California. 
Range. — San Marcos Island. 
Perognathus spinatus pullus Burt 
Coronados Island Spiny Pocket Mouse 


1932. Perognathus spinatus pullus Burt, Trans. San Diego Soc. Nat. Hist. 7:166. October 31. 
Type locality. —Coronados Island (26°06’N, 111°18’W), Gulf of California, Baja 


California. 
Range. — Coronados Island. 
Perognathus spinatus occultus Nelson 
Carmen Island Spiny Pocket Mouse 


1912. Perognathus spinatus nelsoni Townsend, Bull. Amer. Mus. Nat. Hist. 31:122. June 14. 
(Not of Merriam, 1894.) 

1912. Perognathus spinatus occultus Nelson, Proc. Biol. Soc. Wash. 25:116. June 29. (Substi- 
tute for P. s. nelsoni Townsend, preoccupied. ) 
Type locality. — Carmen Island, Gulf of California, Baja California. 
Range. — Carmen Island. 


Perognathus spinatus seorsus Burt 
Danzante Island Spiny Pocket Mouse 


1932. Perognathus spinatus seorsus Burt, Trans. San Diego Soc. Nat. Hist. 7:167. October 31. 
Type locality. —Danzante Island (25°47’N, 111°11’W), Gulf of California, Baja 


California. 
Range. — Danzante Island. 
Perognathus spinatus bryanti Merriam 
San José Island Spiny Pocket Mouse 
1894. Perognathus bryanti Merriam, Proc. Calif. Acad. Sci., ser. 2, 4:458. September 25. 
1932. Perognathus spinatus bryanti Burt, Trans. San Diego Soc. Nat. Hist. 7:167. October 31. 
Type locality. — San José Island, Gulf of California, Baja California. 
Range. — San José Island. 
Perognathus spinatus latijugularis Burt 
San Francisco Island Spiny Pocket Mouse 
1932. Perognathus spinatus latijugularis Burt, Trans. San Diego Soc. Nat. Hist. 7: 168. October 
31 


Type locality. — San Francisco Island (24°50’N, 110°34’W), Gulf of California, Baja 


California. 
Range. — San Francisco Island. 


1963 } Hury: Mammats or Baya CALIFORNIA 119 


Perognathus spinatus magdalenae Osgood 
Magdalena Island Spiny Pocket Mouse 


1907. Perognathus spinatus magdalenae Osgood, Proc. Biol. Soc. Wash. 20:21. February 23. 
Type locality. — Magdalena Island, off the west coast of Baja California. 
Range. — Magdalena Island. 


Perognathus spinatus margaritae Merriam 
Santa Margarita Island Spiny Pocket Mouse 


1894. Perognathus margaritae Merriam, Proc. Calif. Acad. Sci., ser. 2, 4:459. September 25. 
1930. Pferognathus} s{ pinatus | margaritae Benson, Univ. Calif. Publ. Zool. 32:452. Septem- 
ber 6. 
Type locality. — Santa Margarita Island, off the west coast of Baja California . 
Range. — Santa Margarita Island. 


Perognathus spinatus lambi Benson 
Espiritu Santo Island Spiny Pocket Mouse 
1930. Perognathus spinatus lambi Benson, Univ. Calif. Publ. Zool. 32:452. September 6. 


Type locality. — San Gabriel, Espiritu Santo Island, Gulf of California, Baja California. 
Range. — Espiritu Santo Island. 


Perognathus spinatus peninsulae Merriam 
Southern Peninsula Spiny Pocket Mouse 


1894. Perognathus spinatus peninsulae Merriam, Proc. Calif. Acad. Sci., ser. 2, 4:460. Septem- 
ber 25. 
Type locality. — San José del Cabo, Baja California. 
Range. — Cape region of Baja California. 


Genus Dipodomys Gray 
agilis Group 
Dipodomys agilis cabezonae (Merriam) 


Cabezon Kangaroo Rat 


1904. Perodipus cabezonae Merriam, Proc. Biol. Soc. Wash. 17:144. July 14. 
1921. Dipodomys agilis cabezonae Grinnell, Jour. Mamm. 2:96. May 2. 

Type locality. — Cabezon, San Gorgonio Pass, Riverside County, California. 

Range. — Along the international boundary, from Jacumba Valley to Campo, thence 
south over the northwestern chaparral-covered slopes of the Sierra Juarez. 


Dipodomys agilis simulans (Merriam) 
Dulzura Kangaroo Rat 


1904. Perodipus streatori simulans Merriam, Proc. Biol. Soc. Wash. 17:144. July 14. 

1921. Dipodomys agilis simulans Grinnell, Jour. Mamm. 2:96. May 2. 

1925. Dipodomys agilis latimaxillaris Huey, Proc. Biol. Soc. Wash. 38:84. May 26. (Two 
miles west of Santo Domingo Mission (30°45’N, 115°58’W), Baja California. Regarded 
as identical with simulans by Huey, 1951.) 
Type locality. — Dulzura, San Diego County, California. . 
Range. — Over the great coastal foothill areas from the western base of the Sierra Juarez 

to the Pacific Ocean, and from the international boundary directly south of Campo, California, 

as far south as Rancho San Pablo, 10 miles south of Alamo, thence diagonally southwest to San 


Quintin and the Pacific Ocean. 


120 SAN Disco Society or NAtuRAL History [Vor 13 


Dipodomys agilis martirensis Huey 
Sierra Kangaroo Rat 


1927. Dipodomys agilis martirensis Huey, Trans. San Diego Soc. Nat. Hist. 5:7. February 20. 

Type locality.—La Grulla (east side of valley), Sierra San Pedro Martir, Baja 
California. Alt. 7500 feet. 

Range. — Higher parts of the Sierra Juarez, southward to La Grulla in the Sierra San 
Pedro Martir and westward along the higher foothill slopes to the vicinity of San José (Huey, 
1951). 

Dipodomys agilis plectilis Huey 
EI Rosario Kangaroo Rat 


1951. Dipodomys agilis plectilis Fuey, Trans. San Diego Soc. Nat. Hist. 11:240. April 30. 
Type locality. — Mouth of Canon San Juan de Dios (30°7’N), Baja California. 
Range. — El Rosario Valley eastward to the type locality, thence south to Aguaito and 
E] Marmol and west to the coast at Santa Catarina Landing. 


Dipodomys paralius Huey 
Santa Catarina Kangaroo Rat 


1951. Dipodomys paralius Huey, Trans. San Diego Soc. Nat. Hist. 11:241. April 30. 
Type locality. — Santa Catarina Landing (29°31’N), Baja California. 
Range. — So far as now known, near Santa Catarina and around Santa Catarina 
Landing. 
peninsularis Group 
Dipodomys peninsularis pedionomus Huey 


Chapala Kangaroo Rat 


1951. Dipodomys peninsularis pedionomus Huey, Trans. San Diego Soc. Nat. Hist. 11:247. 

April 30. 

Type locality. —2 miles north of Chapala Dry Lake (29°30’N, 114°35’W), on Llano 
de Santa Ana, Baja California. 

Range. — Inland Ilanos from southeast of San Fernando Mission south to the valley 
region below Punta Prieta and eastward to Valle de Agua Amarga and San Borja Mission. 


Dipodomys peninsularis peninsularis (Merriam) 
Vizcaino Desert Kangaroo Rat 


1907. Perodipus simulans peninsularis Merriam, Proc. Biol. Soc. Wash. 20:79. July 22. 
1921. Dipodomys agilis peninsularis Grinnell, Jour. Mamm. 2:96. May 2. 
1951. Dipodomys peninsularis peninsularis Huey, Trans. San Diego Soc. Nat. Hist. 11:246. 
April 30. 
iT pe locality. — Santo Domingo Landing (28°51’N, 114°W), Baja California. 
Range. — Vizcaino Desert from below Punta Prieta (about 28°40’N) south to vicinity 
of San Ignacio (27°20’N), thence over the peninsular divide to El Valle de Yaqui between 
San Ignacio and Santa Rosalia on the Gulf slope. 


Dipodomys peninsularis eremoecus Huey 
Gulf Coast Kangaroo Rat 


1951. Dipodomys peninsularis eremoecus Huey, Trans. San Diego Soc. Nat. Hist. 11:248. 
April 30. 
Type locality. — Seven miles west of San Francisquito Bay (28°30’N), Baja California. 
Range. — Wide Ilano-like region bordering Santa Teresa and San Francisquito bays 
on the Gulf side of Baja California. 


1963 } Huey: MAmMMALs oF BAJA CALIFORNIA 121 


Dipodomys peninsularis australis Huey 
Southern Peninsula Kangaroo Rat 
1951. Dipodomys peninsularis australis Huey, Trans. San Diego Soc. Nat. Hist. 11:249. 
April 30. 
ie locality. — Santo Domingo (25°30’N), Magdalena Plain, Baja California. 
Range. — Magdalena Plain from San Jorge in the northernmost section south to Matan- 
cita, a ranch inland from the mouth of Magdalena Bay at about latitude 24°40’N. 


heermanni Group 
Dipodomys gravipes Huey 
San Quintin Broad-faced Kangaroo Rat 
1925. Dipodomys gravipes Huey, Proc. Biol. Soc. Wash. 38:83. May 26. , 

Type locality.— Two miles west of Santo Domingo Mission (30°45’N, 115°58’W), 
Baja California, or precisely, on cactus covered slope south of huge red cliff that marks the 
entrance of Santo Domingo River Canyon from the coastal plain. 

Range. — So far as known, from the vicinity of the type locality south over the Ilano 
and foothills east of San Quintin, a distance of about 20 miles, and north over Llano de Camalu 
to the San Telmo River. 

Dipodomys antiquarius Huey 
San Borjas Kangaroo Rat 
1962. Dipodomys antiquarius Huey, Trans. San Diego Soc. Nat. Hist. 12:477. August 30. 

Type locality. —San Juan Mine, Sierra San Borjas (28°41’N, 113°37’W), Baja Cali- 
fornia. Alt. 4000 feet. 

Range. — Known only from the type locality. 


merriami Group 
Dipodomys merriami merriami Mearns 
Merriam Kangaroo Rat 

1890. Dipodomys merriami Mearns, Bull. Amer. Mus. Nat. Hist. 2:290. February 21. 
1914. Dipodomys merriami merriami Grinnell, Univ. Calif. Publ. Zool. 12:241. March 20. 

Type locality. — New River, between Phoenix and Prescott, Maricopa County, Arizona. 

Range. — A narrow fringe along the international boundary eastward from Mexicali to 
the Colorado River and a short distance southward along the eastern base of the Sierra Cocopah. 


Dipodomys merriami arenivagus Elliot 
San Felipe Kangaroo Rat 
1904. Dipodomys merriami arenivagus Elliot, Field Columb. Mus. Publ. 87, Zool. Ser. 3:249. 
January 7. 
Type locality. — San Felipe, Baja California. 
Range. — Arid deserts east of the Sierras Juarez and San Pedro Martir and west of the 
Sierra Cocopah, from the international boundary south to an undetermined area between San 


Felipe and Los Angeles Bay. 


Dipodomys merriami trinidadensis Huey 
Trinidad Kangaroo Rat 


1951. Dipidomys merriami trinidadensis Huey, Trans. San Diego Soc. Nat. Hist. 11:220. 
April 30. 
Type locality. — Aguaito Spring, El Valle de la Trinidad, Baja California. 
Range. — El Valle de la Trinidad and more arid parts of El Valle de San Rafael, at 
least to Sangre de Cristo along the western base of the Sierra Juarez. 
Lidicker (1960), in his analysis of the merriami group, extended the range of this race 
to include the population living in a limited area on the international boundary at Jacumba. 


This region is divided from El Valle de San Rafael by a section of pine- and chaparral-clad, 


122 SAN Dteco Society oF NATuRAL History {Vov. 13 


rock-bound hill country of more than 30 miles in breadth in which this species could never have 
lived. This Jacumba Valley population is one of a number of variants or incipient subspecies 
in isolated or elevated localities which border the hotter deserts that constitute the principal range 
of D. merriami. 

Dipodomys merriami quintinensis Huey 


San Quintin Kangaroo Rat 


1951. Dipodomys merriami quintinensis Huey, Trans. San Diego Soc. Nat. Hist. 11:222. 

April 30. 

Type locality. — Five miles east of San Quintin, Baja California. 

Range. — Coastal plain about San Quintin Bay as far north as the vicinity of Santo 
Domingo, some 25 miles north of San Quintin, and south to the vicinity of the El Rosario 
River bottom. 

All specimens so far known have been collected from low elevations similar to the San 
Quintin-Santo Domingo coastal terraces. 


Dipodomys merriami semipallidus Huey 
Mid-peninsula Kangaroo Rat 


1927. Dipodomys merriami semipallidus Huey, Trans. San Diego Soc. Nat. Hist. 5:65. July 6. 

Type locality.— Seven miles north of Santa Catarina (29°45’N, 115°10’W), Baja 
California; now (1958) known as Rancho Ramona. 

Range. — In its most typical form this race is found in a limited region over the Llano 
de San Agustin from Mission San Fernando southward to Santa Catarina and eastward to 
El Marmol, all between elevations of 1500 and 2000 feet. 

As has been recorded (Huey, 1951), this race differs from its more southern relative 
D. m. platycephalus by darker pelage dorsally. The two subspecies are easily differentiated by 
this character. A recently collected series of specimens from San Fernando and another series 
from the type locality have amply demonstrated the correctness of the preceding statement. A 
wide area uninhabited by the species separates D. m. semipallidus from the darker race, D. m. 
quintinensis, found to the north; the latter lives at a much lower elevation and under cool, humid, 
coastal conditions. A wide area of intergradation extends south from the limited range of D. m. 
semipallidus. Lidicker (1960) has emphasized this race’s affinities with the more southern race, 
D. m. platycephalus and, indeed, arranged semipallidus as a synonym of platycephalus. Because 
the color of the pelage permits separation of the two, I feel that D. m. semipallidus is worthy 
of recognition. 


Dipodomys merriami platycephalus Merriam 
Calmalli Kangaroo Rat 


1907. Dipodomys platycephalus Merriam, Proc. Biol. Soc. Wash. 20:76. July 22. 
1927. Dipodomys merriami platycephalus Huey, Trans. San Diego Soc. Nat. Hist. 5:66. 
July 6. 
Type locality. — Calmalli, Baja California. 
Range. — From near latitude 29° on the Gulf side diagonally southwestward to Santa 
Rosalia Bay on the Pacific side and over the great Vizcaino Desert region. 


Dipodomys merriami annulus Huey 
San Francisquito Kangaroo Rat 


1951. Dipodomys merriami annulus Huey, Trans. San Diego Soc. Nat. Hist. 11:224. April 
30. 
Type locality. —Barril (28°20’N, 112°50’W), Gulf of California, Baja California. 
Range. — Gulf of California side of the peninsula over the coastal plains bordering San 
Francisquito and Santa Teresa bays and thence northward into a non-typical population at Los 
Flores near Los Angeles Bay. 


1963 } Huery: Mammats oF Baja CALIFORNIA 12 


W 


Dipodomys merriami brunensis Huey 
San Bruno Kangaroo Rat 
1951. Dipodomys merriami brunensis Huey, Trans. San Diego Soc. Nat. Hist. 11:225. April 
30. 
Type locality. — Llano de San Bruno, Baja California. 
Range. — El Valle de Yaqui, about 8 or 10 miles northwest of Santa Rosalia, Llano 


de San Bruno south of Santa Rosalia, and the gradually rising region from the south end of 
Conception Bay south to Canipolé. 


Dipodomys merriami llanoensis Huey 
Magdalena Plain Kangaroo Rat 
1951. Dipodomys merriami llanoensis Huey, Trans. San Diego Soc. Nat. Hist. 11:226. April 
30 


Type locality. — Buena Vista (24°50’N, 111°50’W) , Magdalena Plain, Baja California. 

Range. — From San Jorge, on the northern end of the Magdalena Plain, south to the 
area south of El Refugio, and probably southward to Arroyo Seco below Magdalena Bay. 

Lidicker (1960:203) questioned the validity of this subspecies. Recognition of the sub- 
species seems to me desirable as a means of cataloging the geographic variation shown by the 
specimens. 

Dipodomys merriami melanurus Merriam 
Cape San Lucas Kangaroo Rat 


1893. Dipodomys merriami melanurus Merriam, Proc. Calif. Acad. Sci., ser. 2, 3:345. January 
5 


Type locality. — San José del Cabo, Baja California. 

Range. — In its typical form this race occupies a crescentic area, having an arid tropical 
climate, fringing the southern base of the Sierra Victoria from Cape San Lucas eastward to 
San José del Cabo and Miraflores, thence northward to Agua Caliente. 

Between Agua Caliente and La Paz a mountainous area interrupts the range of D. mer- 
riam1; the specimens from La Paz, while nearer those of the Cape District, show some characters 
of D. m. llanoensis from the Magdalena Plain District, as would be expected. 


Dipodomys margaritae Merriam 
Margarita Island Kangaroo Rat 


1907. Dipodomys margaritae Merriam, Proc. Biol. Soc. Wash. 20:76. July 22. 

Type locality. — Santa Margarita Island, off the west coast of southern Baja California. 

Range. — Santa Margarita Island. 

Lidicker (1960) regarded this insular population as a subspecies of D. merriami. He had 
only the same material on which to base his conclusions that was used by Huey (1951), who 
accorded specific rank to the population. At that time Huey recorded only specimens from the 
Dickey collection, but had before him also those from the U.S. N. M. The latter were so faded 
that they were of little use. Huey found a low proportion of intergrading characters, as did 
Lidicker. To me, specific instead of subspecific rank seems preferable. 


Dipodomys insularis Merriam 
San José Island Kangaroo Rat 


1907. Dipodomys insularis Merriam, Proc. Biol. Soc. Wash. 20:77. July 22. 
Type locality. — San José Island, Gulf of California, Baja California. 
Range. — San José Island. 


124 SAN Dieco Society ofr Naturav History pViorels 


deserti Group 
Dipodomys deserti deserti Stephens 
Big Desert Kangaroo Rat 
1887. Dipodomys deserti Stephens, Amer. Nat. 21:42. January. 

Type locality. — Mojave River (3 or 4 miles from and opposite Hesperia), San Ber- 
nardino County, California. 

Range. — Colorado Desert district of northeastern Baja California, from the vicinity 
of Pilot Knob westward along the international boundary to the base of the coast range moun- 
tains, southward along the eastern edge of Pattie Basin, skirting the desert base of the Sierra 
San Pedro Martir, at least to the latitude of San Felipe and probably considerably farther south 
in suitable sandy habitats. 

Family CASTORIDAE — Beavers 
Genus Castor Linnaeus 
Castor canadensis repentinus Goldman 
Colorado River Beaver 
1932. Castor canadensis repentinus Goldman, Jour. Mamm. 13:266. August 9. 

Type locality. — Bright Angel Creek, 4000 feet, Grand Canyon, Arizona. 

Range. — Beavers are to be found in extreme northeastern Baja California, oftentimes 
in abundance, where they follow the Colorado River and larger canals through the delta region. 

This beaver is by habit a “bank digger” for its shelter and has been the cause of consid- 
erable trouble in canals where regular waterflow is carried and where food for sustenance, such 
as willow and cottonwood saplings, is growing. 


Family CRICETIDAE — Cricetid Rodents 
Genus Oryzomys Baird 
Oryzomys peninsulae Thomas 
Lower California Rice Rat 
1897. Oryzomys peninsulae Thomas, Ann. Mag. Nat. Hist., ser. 6, 20:548. December. 
Type locality. — Santa Anita, Baja California. 
Range. — Known only from limited marshy areas near sea level in extreme southern Baja 


California. 
Genus Reithrodontomys Giglioli 
Reithrodontomys megalotis megalotis (Baird) 
Desert Harvest Mouse 

1858. Reithrodon megalotis Baird, Mammals, in Rep. Expl. Surv. Railr. to Pacific 8(1) :451. 

July 14. 
1893. Reithrodontomys megalotis J. A. Allen, Bull. Amer. Mus. Nat. Hist. 5:79. April 28. 

Type locality. — Between Janos, Chihuahua, and San Luis Springs, Grant County, New 
Mexico. 

Range. — The riparian section of the Colorado River Delta in the extreme northeastern 
part of Baja California. 


Reithrodontomys megalotis longicaudus (Baird) 
Long-tailed Harvest Mouse 
1858. Reithrodon longicauda Baird, Mammals, in Rep. Expl. Surv. Railr. to Pacific 8(1) :451. 


July 14. 
1913. Reithrodontomys megalotis longicaudus Grinnell, Proc. Calif. Acad. Sci., ser. 4, 3:303. 
August 28. 
Type locality. — Petaluma, Sonoma County, California. 
Range. — In marshlike areas south from the international boundary to Jacumba along 


the lower slopes of the Sierra Juarez to El Valle de la Trinidad, and west to the coast from 
Tijuana to Ensenada. 


1963 } Huey: Mammats oF Baja CALIFORNIA 12 


Ww 


Reithrodontomys megalotis peninsulae (Elliot) 
San Quintin Harvest Mouse 

1903. Rhithrodontomys peninsulae Elliot, Field Columb. Mus. Publ. 74, Zool. Ser. 3:164. 

May 7. 
1914. Reithrodontomys megalotis peninsulae A. H. Howell, North Amer. Fauna 36:35. June 5. 

Type locality. — San Quintin, Baja California. 

Range. — Coastal plain, south from San Antonio del Mar to San Fernando Mission 
and inland to the base of the coast range mountains, mainly the Sierra San Pedro Martir, or 
wherever marshy conditions forming suitable habitat for the species are to be found. 


Genus Peromyscus Gloger 
Peromyscus crinitus stephensi Mearns 
Stephens Canyon Mouse 

1897. Peromyscus stephensi Mearns, Proc. U. S. Nat. Mus. 19:721. July 30. 
1909. Peromyscus crinitus stephensi Osgood, North Amer. Fauna 28:232. April 17. 

Type locality. — Lowest water on wagon road in canyon at eastern base of Coast Range 
near Mexican boundary (=3 miles east of Mountain Spring), Imperial County, California. 

Range. — Along the lower slopes on the desert side of the Coast Range mountain chain 
south from the international boundary to the latitude of San Felipe. Southern extent of range 
unknown. 

Peromyscus crinitus pallidissimus Huey 
Pallid Canyon Mouse 

1931. Peromyscus crinitus pallidissimus Huey, Trans. San Diego Soc. Nat. Hist. 6:389. 

August 28. 

Type locality. — Small island in Gonzaga Bay (29°50’N, 114°20’W), Gulf of Cali- 
fornia, Baja California. 

Range. — Known from the type locality only. 


Peromyscus pseudocrinitus Burt 
False Canyon Mouse 
1932. Peromyscus pseudocrinitus Burt, Trans. San Diego Soc. Nat. Hist. 7:173. October 31. 
Type locality. — Coronados Island (26°60’N, 111°18’W), Gulf of California, Baja 
California. 


Range. — Coronados Island. 


Peromyscus californicus insignis Rhoads 
Southern Parasitic Mouse 
1895. Peromyscus insignis Rhoads, Proc. Acad. Nat. Sci. Philadelphia 47:33. February 21. 
1907. Peromyscus californicus insignis Mearns, U. S. Nat. Mus. Bull. 56:429. April 13. 
Type locality. — Dulzura, San Diego County, California. 
Range. — The brush-covered slopes west of the higher mountains to the coast, and from 
the international boundary south to or immediately below San Quintin. 


Peromyscus eremicus eremicus (Baird) 
Desert White-footed Mouse 

1858. Hesperomys eremicus Baird, Mammals, in Rep. Expl. Surv. Railr. to Pacific 8(1) :479. 

uly 14. 
1895. eee eremicus J. A. Allen, Bull. Amer. Mus. Nat. Hist. 7:226. June 29. 

Type locality. — Old Fort Yuma, Imperial County, California, on Colorado River 
opposite Yuma, Arizona. ; 

Range. — Northeastern desert section of Baja California from the international boundary 
south to San Luis Bay, and from the Colorado River and Gulf of California west to the foothills 
of the eastern slopes of the higher mountains. 


126 


1892. 
1898. 


1903. 


SAN Drieco Society oF Naturav History { Vo. 13 


Peromyscus eremicus fraterculus (Miller) 
Dulzura White-footed Mouse 


V esperimus fraterculus Miller, Amer. Nat. 26:261. March. 

Pi eromyscus} eremicus fraterculus J. A. Allen, Bull. Amer. Mus. Nat. Hist. 10:154. 
April 12. 

Peromyscus homochroia Elliot, Field Columb. Mus. Publ. 74, Zool. Ser. 3:158. May 7. 
(San Quintin, Baja California.) 

Type locality. — Dulzura, San Diego County, California. 


Range. — Along the international boundary from Jacumba Valley westward to the 


Pacific Ocean and south over the brushy western slopes of the higher mountains to El Rosario 
and the San Fernando Mission. 


1898. 
1909. 


1S 28 


19311. 


1909. 


1909. 


1909. 


Peromyscus eremicus cedrosensis J. A. Allen 
Cedros Island White-footed Mouse 


Peromyscus cedrosensis J. A. Allen, Bull. Amer. Mus. Nat. Hist. 10:154. April 12. 
Peromyscus eremicus cedrosensis Osgood, North Amer. Fauna 28:244. April 17. 
Type locality. —Cerros (=Cedros) Island, Baja California. 

Range. — Cedros Island. 


Peromyscus eremicus carmeni Townsend 


Carmen Island White-footed Mouse 


Peromyscus eremicus carmeni Townsend, Bull. Amer. Mus. Nat. Hist. 31:126. June 14. 
Type locality. — Carmen Island, Gulf of California, Baja California. 
Range. — Carmen Island. 


Peromyscus eremicus cinereus Hall 
San José Island White-footed Mouse 


Peromyscus eremicus cinereus Hall, Proc. Biol. Soc. Wash. 44:87. June 29. 
Type locality. — San José Island (25°N), Gulf of California, Baja California. 
Range. — San José Island. 


Peromyscus eremicus insulicola Osgood 
Espiritu Santo Island White-footed Mouse 


Peromyscus eremicus insulicola Osgood, North Amer. Fauna 28:246. April 17. 
Type locality. — Espiritu Santo Island, Gulf of California, Baja California. 
Range. — Espiritu Santo Island. 


Peromyscus eremicus polypolius Osgood 
Margarita Island White-footed Mouse 
Peromyscus eremicus polypolius Osgood, North Amer. Fauna 28:248. April 17. 


Type locality. — {Santa} Margarita Island, off west coast of southern Baja California. 
Range. — Santa Margarita Island. 


Peromyscus eremicus avius Osgood 
Cerralvo Island White-footed Mouse 


Peromyscus eremicus avius Osgood, North Amer. Fauna 28:247. April 17. 
Type locality. — Cerralvo Island, Gulf of California, Baja California. 
Range. — Cerralvo Island. 


1963 } Huey: MamMMa ts oF BAajA CALIFORNIA 12 


N 


Peromyscus eremicus eva Thomas 


Lower California White-footed Mouse 


1898. Peromyscus eva Thomas, Ann. Mag. Nat. Hist., ser. 7, 1:44. January. 
1898. Peromyscus eremicus propinquus J. A. Allen, Bull. Amer. Mus. Nat. Hist. 10:154. 
April 12. (San Pablo Point, Baja California.) 
1909. Peromyscus eremicus eva Osgood, North Amer. Fauna 28:245. April 17. 
Type locality. — San José del Cabo, Baja California. 
Range. — On the mainland of the southern part of the peninsula from Cape San Lucas 
northward to about latitude 29°N, where it meets the southern extent of the ranges of P. e. 
eremicus and P. e. fraterculus. A broad band of intergradation exists. 


Peromyscus caniceps Burt 
Monserrate Island Canyon Mouse 
1932. Peromyscus caniceps Burt, Trans. San Diego Soc. Nat. Hist. 7:174. October 31. 
Type locality. — Monserrate Island (25°38’N, 111°02’W), Gulf of California, Baja 


California. 
Range. — Monserrate Island. 


Peromyscus guardia mejiae Burt 

Mejia Island White-footed Mouse 
1932. Peromyscus guardia mejiae Burt, Trans. San Diego Soc. Nat. Hist. 7:174. October 31. 
Type locality. — Mejia Island (29°33’N, 113°35’W), Gulf of California, Baja Calli- 


fornia. 


Range. — Mejia Island. 


Peromyscus guardia guardia Townsend 
Angel de la Guardia White-footed Mouse 
1912. Peromyscus guardia Townsend, Bull. Amer. Mus. Nat. Hist. 31:126. June 14. 


Type locality. — Angel de la Guardia Island, Gulf of California, Baja California. 
Range. — Angel de la Guardia Island. 


Peromyscus guardia interparietalis Burt 
San Lorenzo Island White-footed Mouse 


1932. Peromyscus guardia interparietalis Burt, Trans. San Diego Soc. Nat. Hist. 7:175. 
October 31. 
Type locality. — South San Lorenzo Island (28°36’N, 112°51’W), Gulf of California, 
Baja California. 
Range. — South San Lorenzo Island. 


Peromyscus dickeyi Burt 
Tortuga Island White-footed Mouse 


1932. Peromyscus dickeyi Burt, Trans. San Diego Soc. Nat. Hist. 7:176. October 31. 
Type locality.— Tortuga Island (27°21’N, 111°54’W), Gulf of California, Baja 
California. 


Range. — Tortuga Island. 


Peromyscus maniculatus gambelii (Baird) 
Gambel White-footed Mouse 


1858. Hesperomys gambelii Baird, Mammals, in Rep. Expl. Surv. Railr. to Pacific, 8(1) :464. 
July 14. 

1893. Sitomys americanus thurberi J. A. Allen, Bull. Amer. Mus. Nat. Hist. 5:185. August 
18. (Sierra San Pedro Martir, Baja California. ) 


128 SAN Deco Society oF Naturav History { VoL. 13 


1896. Peromyscus texanus medius Mearns, Preliminary diagnoses of new mammals from the 
Mexican border of the United States, p. 4. March 25. (Preprint of Proc. U. S. Nat. 
Mus. 18:446. May 23. Nachoguero Valley, Baja California.) 
1909. Peromyscus maniculatus gambeli Osgood, North Amer. Fauna 28:67. April 17. 
Type locality. — Monterey, Monterey County, California. 
Range. — Westward along the international boundary from Nachogiiero Valley and the 
western slopes of the northern Sierra Juarez to the Pacific Ocean, south to Ensenada. 
Specimens from El Valle de la Trinidad, Sierra San Pedro Martir and westward to San 
Quintin are mainly non-typical and often strongly approach either P. m. sonoriensis or P. m. 
coolid gei. 
Peromyscus maniculatus sonoriensis (Le Conte) 


Sonora White-footed Mouse 


1853. Hesp{eromys }sonoriensis Le Conte, Proc. Acad. Nat. Sci. Philadelphia 6:413. 
1903. Peromyscus oresterus Elliot, Field Columb. Mus. Publ. 74, Zool. Ser. 3:159. May 7. 
(Vallecitos, Sierra San Pedro Martir, Baja California.) 
1909. Peromyscus maniculatus sonoriensis Osgood, North Amer. Fauna 28:89. April 17. 
Type locality. — Santa Cruz, Sonora. 
Range. — Northeastern Baja California east of the higher parts of the Sierra Juarez 
and the Sierra San Pedro Martir, south along the northern Gulf coast to an undetermined 
section below San Felipe; thence around the southern slopes of the Sierra San Pedro Martir. 


In the latter region a broad band of intergradation exists where this subspecies blends 
with the north coastal P. m. gambelii and the southern peninsular subspecies P. m. coolidgei. 


Peromyscus maniculatus assimilis Nelson and Goldman 
Los Coronados Islands White-footed Mouse 
1931. Peromyscus maniculatus assimilis Nelson and Goldman, Jour. Mamm. 12:305. August 
we 
Type locality.—Coronados Island (=Los Coronados Islands) off the west coast of 
northern Baja California. 


Range. — Los Coronados Islands. 


Peromyscus maniculatus dubius J. A. Allen 
Todos Santos Islands White-footed Mouse 


1898. Peromyscus dubius J. A. Allen, Bull. Amer. Mus. Nat. Hist. 10:157. April 12. 

1909. Peromyscus maniculatus dubius Osgood, North Amer. Fauna 28:98. April 17. 
Type locality. —Todos Santos Islands, off the west coast of northern Baja California. 
Range. — Todos Santos Islands. 


Peromyscus maniculatus exiguus J. A. Allen 
San Martin Island White-footed Mouse 


1898. Peromyscus exiguus J. A. Allen, Bull. Amer. Mus. Nat. Hist. 10:157. April 12. 

1931. Peromyscus maniculatus martinensis Nelson and Goldman, Jour. Wash. Acad. Sci. 
21:534. December 19. (San Martin Island, Baja California.) 

1932. Peromyscus maniculatus exiguus Nelson and Goldman, Jour. Mamm. 13:371. Novem- 
ber 2. 
Type locality. — San Martin Island, off the west coast of northern Baja California. 
Range. — San Martin Island. 


1963] 


1932. 


Huey: MAmMMALs oF BAJA CALIFORNIA 129 


Peromyscus maniculatus hueyi Nelson and Goldman 
Gonzaga White-footed Mouse 


Peromyscus maniculatus hueyi Nelson and Goldman, Trans. San Diego Soc. Nat. Hist. 
7:51. April 15. 
Type locality. — Small island in Gonzaga Bay (29°50’N, 114°20’W/), Gulf of Cali- 


fornia, Baja California. 


1898. 
1909. 


1931: 


1898. 
1909. 


1909. 


1909. 


1898. 
1909. 


Range. — Known only from the type locality. 


Peromyscus maniculatus geronimensis J. A. Allen 
San Gerénimo Island White-footed Mouse 


Peromyscus geronimensis J. A. Allen, Bull. Amer. Mus. Nat. Hist. 10:156. April 12. 
Peromyscus maniculatus geronimensis Osgood, North Amer. Fauna 28:99. April 17. 
Type locality. — San Geronimo Island, off the west coast of northern Baja California. 
Range. — San Gerénimo Island. 


Peromyscus maniculatus dorsalis Nelson and Goldman 
Natividad Island White-footed Mouse 


Peromyscus maniculatus dorsalis Nelson and Goldman, Jour. Wash. Acad. Sci. 21:535. 
December 19. 

Type locality. — Natividad Island, off the west coast of central Baja California. 
Range. — Known only from Natividad Island. 


Peromyscus maniculatus cineritius J. A. Allen 
San Roque Island White-footed Mouse 


Peromyscus cineritius J. A. Allen, Bull. Amer. Mus. Nat. Hist. 10:155. April 12. 
Peromyscus maniculatus cineritius Osgood, North Amer. Fauna 28:100. April 17. 
Type locality. — San Roque Island, off the west coast of central Baja California. 
Range. — San Roque Island. 


Peromyscus maniculatus magdalenae Osgood 
Magdalena Island White-footed Mouse 


Peromyscus maniculatus magdalenae Osgood, North Amer. Fauna 28:101. April 17. 
Type locality. — Magdalena Island, off the west coast of southern Baja California. 
Range. — Magdalena Island and a narrow strip of adjacent mainland. 


Peromyscus maniculatus margaritae Osgood 
Santa Margarita Island White-footed Mouse 


Peromyscus maniculatus margaritae Osgood, North Amer. Fauna 28:95. April 17. 
Type locality. — (Santa) Margarita Island, off the west coast of southern Baja California. 
Range. — Santa Margarita Island. 


Peromyscus maniculatus coolidgei Thomas 
San Lucas White-footed Mouse 


Peromyscus leucopus coolidgei Thomas, Ann. Mag. Nat. Hist., ser. 7, 1:45. January. 
Peromyscus maniculatus coolidgei Osgood, North Amer. Fauna 28:94. April 17. 
Type locality. — Santa Anita, Cape region of Baja California. 


Range. — Mainland of the peninsula south from the area below El Marmol to the Cape. 


130 San Dreco Society oF NaturaAL HIstTory [Vot. 13 


Peromyscus sejugis Burt 
Santa Cruz Island White-footed Mouse 


1932. Peromyscus sejugis Burt, Trans. San Diego Soc. Nat. Hist. 7:171. October 31. 
Type locality. —Santa Cruz Island (25°17’N, 110°43’W), Gulf of California, Baja 
California. 


Range. — Santa Cruz and San Diego islands, Gulf of California. 


Peromyscus slevini Mailliard 
Santa Catalina Island White-footed Mouse 


1924. Peromyscus slevini Mailliard, Proc. Calif. Acad. Sci., ser. 4, 14:1221. July 22. (See 
Burt, 1934.) 
Type locality. — Santa Catalina Island, 17 miles northeast of Punta San Marcial, Gulf 
of California. 


Range. — Santa Catalina Island, Gulf of California. 


Peromyscus boylii rowleyi (J. A. Allen) 
Rowley White-footed Mouse 


1893. Sitomys rowleyi J. A. Allen, Bull. Amer. Mus. Nat. Hist. 5:76. April 28. 
1896. P{eromyscus} bl oyli} rowleyi Mearns, Preliminary diagnoses of new mammals from 
the Mexican border of the United States, p. 3. May 25. (Preprint of Proc. U. S. Nat. 
Mus. 19:139. December 21.) 
1903. Peromyscus gaurus Elliot, Field Columb. Mus. Publ. 74, Zool. Ser. 3:157. May 7. (San 
Antonio, Sierra San Pedro Martir, Baja California. ) 
Type locality.— Noland Ranch, on north side of San Juan River, San Juan County, 
Utah. (Hall, 1931.) 
Range. — Brush covered slopes of the higher mountains of northern Baja California, 
especially in weed patches near water courses. 


Peromyscus truei martirensis (J. A. Allen) 
San Pedro Martir White-footed Mouse 


1893. Sitomys martirensis J. A. Allen, Bull. Amer. Mus. Nat. Hist. 5:187. August 18. 
1903. Peromyscus hemionotis Elliot, Field Columb. Mus. Publ. 74, Zool. Ser. 3:157. May 7. 
(Rosarito Divide, Sierra San Pedro Martir, Baja California.) 
1909. Peromyscus truei martirensis Osgood, North Amer. Fauna 28:171. April 17. 
Type locality. — Sierra San Pedro Martir, Baja California. Alt. 7000 feet. 
Range. — Higher mountains of northern Baja California, mainly in the pifon belts of 


the Sierra Juarez and the Sierra San Pedro Martir (Hoffmeister, 1951). 


Peromyscus truei lagunae Osgood 
Sierra Laguna White-footed Mouse 


1909. Peromyscus truei lagunae Osgood, North Amer. Fauna 28:172. April 17. 
Type locality. — La Laguna, Sierra Laguna, Baja California. 
Range. — Wooded slopes, above 4200 feet, on the higher mountains of the Cape District 
of Baja California. 
Genus Onychomys Baird 


Onychomys torridus pulcher Elliot 
Desert Grasshopper Mouse 


1904. Onychomys pulcher Elliot, Field Columb. Mus. Publ. 87, Zool. Ser. 3:243. January 7. 
1913. Ofnychomys} t{orridus} pulcher Hollister, Proc. Biol. Soc. Wash. 26:215. December 
20. 


1963 | Huey: MAMMALS oF BAJA CALIFORNIA 131] 


Type locality. — Morongo Pass, east end of San Bernardino Mountains, San Bernardino 
County, California. 

Range. — This subspecies ranges into Baja California in the extreme northeastern part 
along the international boundary, where there is but one record, a specimen collected at Seven 


Wells (Hollister, 1914). 


Onychomys torridus ramona Rhoads 
Ramona Grasshopper Mouse 


1893. Onychomys ramona Rhoads, Amer. Nat. 27:833. September. 
1904. Onychomys torridus ramona Merriam, Proc. Biol. Soc. Wash. 17:124. June 9. 
Type locality. — San Bernardino Valley (more exactly, Reche Canyon, 4 miles southeast 
of Colton), San Bernardino County, California. Alt. 1250 feet. (Grinnell, 1933.) 
Range. — In northwestern Baja California from Jacumba Valley westward to the Pacific 
Ocean and south a short distance to El Canon Misién and El Valle de las Palmas. 
Specimens of O. ¢. ramona in the S. D. N. H. M. are from Jacumba Valley, El Valle 


de las Palmas, and the south side of Cafién Misidn near La Misién. 


Onychomys torridus macrotis Elliot 
Lower California Grasshopper Mouse 


1903. Onychomys macrotis Elliot, Field Columb. Mus. Publ. 74, Zool. Ser. 3:155. May 7. 
1914. Onychomys torridus macrotis Hollister, Proc. U. S. Nat. Mus. 47:469. October 29. 

Type locality. — Head of San Antonio River, west slope of Sierra San Pedro Martir, 
Baja California. 

Range. — Along the western base of the Sierra Juarez in El Valle de San Rafael and 
from the summit of San Matias Pass through El Valle de la Trinidad to the coast at San 

uintin. 
: The southernmost point of capture to date is at Santa Maria, immediately south of 
San Quintin. 
Genus Sigmodon Say and Ord 
Sigmodon hispidus eremicus Mearns 


Western Cotton Rat 


1897. Sigmodon hispidus eremicus Mearns, Preliminary diagnoses of new mammals of the 
genera Sciurus, Castor, Neotoma, and Sigmodon, from the Mexican border of the United 
States, p. 4. March 5. (Preprint of Proc. U. S. Nat. Mus. 20:504. January 19, 1898.) 
Type locality. — Cienega Well, 30 miles south of Mexican boundary monument No. 204 
on the east bank of the Colorado River, Sonora. 
Range. — Found in the riparian association on the Colorado River Delta and in the agri- 
cultural section of extreme northeastern Baja California. 


Genus Neotoma Say and Ord 
albigula Group 
Neotoma albigula venusta True 


Colorado Valley White-throated Wood Rat 


1894. Neotoma venusta True, Diagnoses of some undescribed wood rats (genus Neotoma) 
in the National Museum, p. 2. June 27. (Preprint of Proc. U.S. Nat. Mus. 17:354. 
November 15.) 

1910. Neotoma albigula venusta Goldman, North Amer. Fauna 31:33. October 19. 

Type locality. — Carrizo Creek, Imperial County, California. 

Range. — Through the dryer mesquite-dune association in the extreme northeastern 
section of Baja California. From the Colorado River it extends westward to the foothills of the 
great coast range and south along these slopes to the southern end of Laguna Salada. 


132 SAN Dreco Society or NAtTurRAL History { Vot. 13 


lepida Group 
Neotoma lepida gilva Rhoads 


Banning White-footed Wood Rat 


1894. Neotoma intermedia gilva Rhoads, Amer. Nat. 28:70. January. 
1932. Neotoma lepida gilya Goldman, Jour. Mamm. 13:63. February 9. 

Type locality. — Banning, Riverside County, California. 

Range. — A rather narrow strip on the eastern slopes of the Sierra Juarez from the inter- 
national boundary south to, and probably a short distance beyond, Gaskill’s Tank. 

The preferred habitat of this race is amid large outcropping boulders on hillsides cr 
mountain crests. 

Neotoma lepida intermedia Rhoads 


Intermediate White-footed Wood Rat 


1894. Neotoma intermedia Rhoads, Amer. Nat. 28:69. January. 
1932. Neotoma lepida intermedia Goldman, Jour. Mamm. 13:64. February 9. 

Type locality. — Dulzura, San Diego County, California. 

Range. — Northwestern Baja California, over the chaparral-covered area from the Sierra 
Juarez westward to the Pacific Ocean, and from the northern Sierra San Pedro Martir obliquely 
northwestward to the coast in the vicinity of the mouth of the Santo Tomas River. 

The habitat of this subspecies is chiefly large clumps of flat jointed cactus, but it occupies 
also rocky outcroppings and boulder-covered hillsides amid open chaparral. 


Neotoma lepida felipensis Elliot 
San Felipe Desert White-footed Wood Rat 
1903. Neotoma bella felipensis Elliot, Field Columb. Mus. Publ. 79, Zool. Ser. 3:217. August 
15 


1932. Neotoma lepida felipensis Goldman, Jour. Mamm. 13:64. February 9. 

Type locality. — San Felipe, Baja California. 

Range. — The arid desert section of eastern Baja California from the southern base of 
Sierra Cocopah south along the lower eastern base of the Sierra San Pedro Martir to an undeter- 
mined area south of San Felipe. 

This wood rat prefers cactus clumps but it also is found commonly under large prostrate 
mesquites or along steep banks of dry arroyos, where it lives in underground crevices and burrows 
guarded by cut cactus thorns. 


Neotoma lepida egressa Orr 
El Rosario White-footed Wood Rat 


1934. Neotoma lepida egressa Orr, Proc. Biol. Soc. Wash. 47:109. June 13. 

Type locality. — One mile east of El Rosario, Baja California. Alt. 200 feet. 

Range. — Coastal region south from San Antonio del Mar to El Rosario, and probably 
coastwise some distance below. Inland toward the base of the Sierra San Pedro Martir as far as 
San José and Socorro. 

The greater numbers of this subspecies are to be found living in the thorny cactus-wild 
rose chaparral of the lower foothills. 


Neotoma lepida insularis Townsend 
Angel de la Guardia Island White-footed Wood Rat 


1912. Neotoma insularis Towsend, Bull. Amer. Mus. Nat. Hist. 31:125. June 14. 

1932. Neotoma lepida insularis Burt, Trans. San Diego Soc. Nat. Hist. 7:182. October 31. 
(see Burt and Barkalow, 1942.) 
Type locality. — Angel de la Guardia Island, Gulf of California, Baja California. 
Range. — Angel de la Guardia Island. 


1963 | Hury: Mammats or Baya CALIFORNIA 133 


Neotoma lepida molagrandis Huey 
Vizcaino Desert White-footed Wood Rat 
1945. Neotoma lepida molagrandis Huey, Trans. San Diego Soc. Nat. Hist. 10:307. August 
31. 


Type locality. — Santo Domingo Landing, more precisely, at the site of the old well near 
the edge of a mesa-like shelf, alt. 50 feet, some 3 miles inland from the landing beach (28°15’N), 
Baja California. 

Range. — So far as now known, the northern and coastal section of the Vizcaino Desert 
region, from Punta Prieta to San Ignacio. 


This subspecies lives in partly subterranean nests built under any large prostrate growth 
and always guarded by beds of cactus thorns about the entrances. It seems to have a habit of 
wandering farther from its shelter and runways than any other form of N. lepida, according to 
the writer’s experience. 


Neotoma lepida aridicola Huey 
San Francisquito White-footed Wood Rat 
1957. Neotoma lepida aridicola Huey, Trans. San Diego Soc. Nat. Hist. 12:287. September 
25. 
Type locality. — El Barril (near 28°20’N), on the Gulf of California, Baja California. 


Range. — So far as known, the desert slopes on the Gulf side of the peninsula in the 
region from San Francisquito Bay to Barril. Almost certainly this subspecies will be found 
farther north and south of the above range when further explorations are made. 


This light colored wood rat has a predilection for arroyo banks where deep washes are 
present in the soil. Nests are located in fissures and guarded by beds of shredded cactus thorns. 


Neotoma lepida marcosensis Burt 
San Marcos Island White-footed Wood Rat 


1932. Neotoma lepida marcosensis Burt, Trans. San Diego Soc. Nat. Hist. 7:179. October 31. 
Type locality. — San Marcos Island (27°13’N, 112°05’W), Gulf of California, Baja 


California. 
Range. — San Marcos Island. 


Neotoma lepida ravida Nelson and Goldman 
Comondt White-footed Wood Rat 
1931. Neotoma intermedia ravida Nelson and Goldman, Proc. Biol. Soc. Wash. 44:107. 
October 17. 
1932. Neotoma lepida ravida Goldman, Jour. Mamm. 13:64. February 9. 
Type locality. — Comondu, southern Baja California. 


Range. — Through the volcanic region, including extensive lava beds, along the back- 
bone of the peninsula from about latitude 28°N south to the Sierra de la Giganta. 


Neotoma lepida nudicauda Goldman 
Carmen Island White-footed Wood Rat 
1905. Neotoma nudicauda Goldman, Proc. Biol. Soc. Wash. 18:28. February 2. 
1932. Neotoma lepida nudicauda Burt, Trans. San Diego Soc. Nat. Hist. 7:182. October 31. 
Type locality. — Carmen Island, Gulf of California, Baja California. 
Range. — Carmen Island. 


134 San Deco Society oF Naturat History | Vorwl3 


Neotoma lepida latirostra Burt 
Danzante Island White-footed Wood Rat 


1932. Neotoma lepida latirostra Burt, Trans. San Diego Soc. Nat. Hist. 7:180. October 31. 
Type locality. —Danzante Island (25°47’N, 111°11’W), Gulf of California, Baja 


California. 
Range. — Danzante Island. 


Neotoma lepida perpallida Goldman 
San José Island White-footed Wood Rat 


1909. Neotoma intermedia perpallida Goldman, Proc. Biol. Soc. Wash. 22:139. June 25. 
1932. Neotoma lepida perpallida Goldman, Jour. Mamm. 13:65. February 9. 

Type locality. — San José Island, Gulf of California, Baja California. 

Range. — San José Island. 


Neotoma lepida abbreviata Goldman 
San Francisco Island White-footed Wood Rat 


1909. Neotoma abbreviata Goldman, Proc. Biol. Soc. Wash. 22:140. June 25. 
1932. Neotoma lepida abbreviata Burt, Trans. San Diego Soc. Nat. Hist. 7:182. October 31. 
Type locality. — San Francisco Island (near southern end of San José Island) , Gulf of 
California, Baja California. 
Range. — San Francisco Island. 


Neotoma lepida pretiosa Goldman 
Magdalena Plain White-footed Wood Rat 


1909. Neotoma intermedia pretiosa Goldman, Proc. Biol. Soc. Wash. 22:139. June 25. 
1932. Neotoma lepida pretiosa Goldman, Jour. Mamm. 13:64. February 9. 

Type locality. — Matancita (also called Soledad), 50 miles north of Magdalena Bay, 
Baja California. Alt. 100 feet. 

Range. — The vast Magdalena Plain from San Jorge (southwest of Comondt) south- 
ward to Santa Margarita Island. 


Neotoma lepida vicina Goldman 
Espiritu Santo Island White-footed Wood Rat 


1909. Neotoma intermedia vicina Goldman, Proc. Biol. Soc. Wash. 22:140. June 25. 
1932. Neotoma lepida vicina Goldman, Jour. Mamm. 13:65. February 9. 
Type locality. — Espiritu Santo Island, Gulf of California, Baja California. 
Range. — Espiritu Santo Island. 


Neotoma lepida notia Nelson and Goldman 
Sierra Laguna White-footed Wood Rat 


1931. Neotoma intermedia notia Nelson and Goldman, Proc. Biol. Soc. Wash. 44:108. Octo- 
ber 17. 
1932. Neotoma lepida notia Goldman, Jour. Mamm. 13:65. February 9. 
Type locality. —La Laguna, Sierra de la Victoria, southern Baja California. Alt. 5500 
feet. 
Range. — Known only from the mountains of the Cape region. 


1963 } Huey: Mammats oF BajA CALIFORNIA 135 


Neotoma lepida arenacea J. A. Allen 
Cape San Lucas White-footed Wood Rat 
1898. Neotoma arenacea J. A. Allen, Bull. Amer. Mus. Nat. Hist. 10:150. April 12. 
1932. Neotoma lepida arenacea Goldman, Jour. Mamm. 13:65. February 9. 
Type locality. — San José del Cabo, Baja California. 
Range. — Coastal plains and lower hills in the Cape District. 
This form is found living abundantly in dry rock walls which surround small farms. 


Neotoma anthonyi J. A. Allen 
Todos Santos Island Wood Rat 


1898. Neotoma anthony J. A. Allen, Bull. Amer. Mus. Nat. Hist. 10:151. April 12. 
Type locality. — Todos Santos Island, off the coast of northwestern Baja California. 
Range. — Todos Santos Island. 


Neotoma martinensis Goldman 


San Martin Island Wood Rat 


1905. Neotoma martinensis Goldman, Proc. Biol. Soc. Wash. 18:28. February 2. 
Type locality. — San Martin Island, off the west coast of northern Baja California. 
Range. — San Martin Island. 


Neotoma bryanti Merriam 
Cedros Island Wood Rat 


1887. Neotoma bryanti Merriam, Amer. Nat. 21:191. February. 
Type locality. — Cerros (=Cedros) Island, Baja California. 
Range. — Cedros Island, off the west coast of Baja California. 


Neotoma bunkeri Burt 
Coronados Island Wood Rat 
1932. Neotoma bunkeri Burt, Trans. San Diego Soc. Nat. Hist. 7:181. October 31. 
Type locality. —Coronados Island (26°06’N, 111°18’W), Gulf of California, Baja 


California. 


Range. — Coronados Island. 
fuscipes Group 
Neotoma fuscipes macrotis Thomas 
San Diego Dusky-footed Wood Rat 


1893. Neotoma macrotis Thomas, Ann. Mag. Nat. Hist., ser. 6, 12:234. September. 
1894. Neotoma fuscipes macrotis Merriam, Proc. Acad. Nat. Sci. Philadelphia 14:246. 

September 25. 

Type locality. — San Diego, California. 

Range. — A narrow strip along the international boundary, from the northern slopes of 
the Sierra Juarez below Jacumba westward to the Pacific Ocean, south to the vicinity of Ensenada. 

Dusky-footed wood rats are found chiefly in heavy chaparral and live oak associations 
where they build large nests of sticks and leaves. These nests are also occasionally placed up in 
the branches of live oak trees, and are often used by several generations, each tenant adding more 
material to the nest. 

Neotoma fuscipes martirensis Orr 
San Pedro Martir Dusky-footed Wood Rat 

1934. Neotoma fuscipes martirensis Orr, Proc. Biol. Soc. Wash. 47:110. June 13. 

Type locality. — Valladares, Sierra San Pedro Martir, Baja California. Alt. 2700 feet. 

Range. — South from latitude 32° to El Rosario, and from the mid-section of the Sierra 
Juarez and the higher reaches of the Sierra San Pedro Martir west through the chaparral belt to 
the coast. 


136 San Dreco Society of NATuRAL HIstTory { VoL. 13 


Genus Microtus Schrank 
Microtus californicus sanctidiegi R. Kellogg 


San Diego Meadow Mouse 


1918. Microtus californicus neglectus R. Kellogg, Univ. Calif. Publ. Zool. 21:31. December 
28. (Not of Jenyns, 1941.) 
1922. Microtus californicus sanctidiegi R. Kellogg, Proc. Biol. Soc. Wash. 35:78. March 20. 
(Substitute for neglectus R. Kellogg.) 
Type locality. — Escondido, San Diego County, California. Alt. 640 feet. 
Range. — Moist sections of the lower Tijuana River drainage in extreme northwestern 
Baja California, from Tecate west to the ocean and coastwise south to and slightly below Ensenada. 


Microtus californicus grinnelli Huey 
Sangre de Cristo Meadow Mouse 


1931. Microtus californicus grinnelli Huey, Trans. San Diego Soc. Nat. Hist. 7:47. December 
19. 
Type locality. — Sangre de Cristo (31°52’N, 116°06’W), in El Valle de San Rafael, 
on the western base of the Sierra Juarez, Baja California. 
Range. — So far as known, in the region immediately west of the Sierra Juarez. 


Microtus californicus hyperythrus. Elliot 
San Pedro Martir Meadow Mouse 


1903. Microtus californicus huperuthrus Elliot, Field. Columb. Mus. Publ. 74, Zool. Ser. 3: 161. 
May 7. 

1903. Wicca: californicus hyperythrus Elliot, Field Columb. Mus. Publ. 79, Zool. Ser. 
3:218. August 15. (A change of spelling permitted by International Code of Zoological 
Nomenclature; Proc. Biol. Soc. Wash. 39:89. July 7, 1926.) 

1926. Microtus californicus perplexabilis Grinnell, Jour. Mamm. 7:223. August 9. (La Grulla, 
7000 feet, Sierra San Pedro Martir, Baja California.) 

Type locality. —La Grulla, Sierra San Pedro Martir, Baja California. (In the original 
account the type locality was said to be San Quintin, Baja California, but this designation was 
later declared to have been an “inexplicable inadvertence”; see Elliot, 1907:292.) 

Range. — Sierra San Pedro Martir, at altitudes from 6000 to 8000 feet. 


Microtus californicus aequivocatus Osgood 
San Quintin Meadow Mouse 
1928. Microtus californicus aequivocatus Osgood, Jour. Mamm. 9:56. February 9. 
Type locality. — San Quintin, Baja California. 
Range. — Coast region from marshy areas near Arroyo Seco southward over the San 
Quintin Plains area to El Rosario, and inland to elevations below 3000 feet. 


Genus Ondatra Link 
Ondatra zibethicus bernardi Goldman 


Lower Colorado River Muskrat 


1932. Ondatra zibethicus bernardi Goldman, Proc. Biol. Soc. Wash. 45:93. June 21. 

Type locality. — Four miles south of Gadsden, Yuma County, Arizona. 

Range. — Colorado River, including seepage ponds, tributary sloughs and irrigation 
canals, in extreme northeastern Baja California. 


1963 } Huey: MAmMMALs oF BajA CALIFORNIA 137 


Family MURIDAE Gray 
Genus Rattus Fischer 
Rattus norvegicus (Berkenhout) 


Norway Rat 
1769. Mus norvegicus Berkenhout, Outlines of the Natural History of Great Britain and Ire- 
land, 1:5. 
1932. Rattus norvegicus Cabrera, Trab. Mus. Nac. Cien. Nat., Madrid Zool. 57:264. Decem- 
ber 30. 


Type locality. — England. 


Range. — Found in sea port towns such as La Paz, San José del Cabo, Santa Rosalia 
and Ensenada, where the species has probably escaped from calling cargo vessels, and at Mexicali 
where it has been introduced by railroad or by invasion from nearby border towns. 


Rattus rattus rattus (Linnaeus) 


Black Rat 


1758. {Mus} rattus Linnaeus, Systema Naturae, ed. 10, 1:61. 
1916. Rattus rattus Hollister, Proc. Biol. Soc. Wash. 29:126. June 6. 
Type locality. — Uppsala, Sweden. 


Range. — Found under similar range restrictions as Rattus norvegicus, but will spread 
into residential areas where dwellings are not so closely located. The surrounding farm regions 
are not acceptable to either of the Rattus and they have remained in limited numbers within the 
towns. 


Rattus rattus alexandrinus (E. Geoffroy Saint-Hilaire) 
Gray-bellied Rat 


1803. Mus alexandrinus E. Geoffroy Saint-Hilaire, Catalogue des Mammiféres du Muséum 
National d’Histoire Naturelle, Paris, p. 192. 


1918. R{attus } rattus alexandrinus Hinton, Jour. Bombay Nat. Hist. Soc. 26:63. December 20. 
Type locality. — Alexandria, Egypt. 


Range. — Similar to that of two preceding Rattus. Fresh stock from overseas vessels 
calling at seaports probably has been introduced at various times. Recognition of races is difficult 


and they are here listed arbitrarily. (See Hall and Kelson, 1959:768.) 


Genus Mus Linnaeus 
Mus musculus domesticus Rutty 


House Mouse 


1772. Mus domesticus Rutty, Essay Nat. Hist. County Dublin, 1:281. 
1943. Mus musculus domesticus Schwarz and Schwarz, Jour. Mamm. 24:65. February 20. 

Type locality. — Dublin, Ireland. 

Range. — House mice are to be found in and around human habitations practically 
throughout the peninsula and on occasion have spread into nearby fields, as at San Ignacio. 

This is the only rodent to be found on oceanic Guadalupe Island where, after being 
introduced during the past century by human occupation, it is now firmly established over most 
of the island. It subsists on the seeds and annual growth of feral weeds and grasses, likewise 
introduced by humans. Specimens of this mouse from Guadalupe Island are in the S. D. N. 


H. M. 


138 San Dteco Society oF NATuRAL History { VoL. 13 


Order CETACEA — Whales and Porpoises 
Family ZIPHIIDAE — Beaked Whales 
Genus Ziphius G. Cuvier 


Ziphius cavirostris G. Cuvier 
Cuvier’s Beaked Whale 


1823. Ziphius cavirostris G. Cuvier, Recherches sur les ossemens fossiles . . . ed. 2, 5:352. 
Type locality. — Near Fos, Bouches-du-Rhone, France. 


Range. — This rare cetacean has been recorded once from the shores of Baja California 


(Hubbs, 1951a). 
Family PHYSETERIDAE — Sperm Whales 


Genus Physeter Linnaeus 
Physeter catodon Linnaeus 


Sperm Whale 


1758. {Physeter} catodon Linnaeus, Systema Naturae, ed. 10, 1:76. 


Type locality. — Kairston, Orkney Island, Scotland (Thomas, Proc. Zool. Soc. London, 
1911, p. 157. March 22.). 


Range. — From the Pribilof Islands south to the Gulf of Panama. The species occurs 
along the entire Pacific coast of Mexico and can thus be given as ranging within the bounds 


of Baja California. 


A school of about 25 Sperm Whales was observed by Hubbs and Moran in January, 
1960, near Guadalupe Island. 


Family KOGIIDAE — Pigmy Sperm Whales 
Genus Kogia Gray 
Kogia breviceps (Blainville) 


Pigmy Sperm Whale 


1833. Physeter breviceps Blainville, Ann. Anat. Phys. 2:337. 


1846. Kogia breviceps Gray, in The Zoology of the Voyage of H. M. S. Erebus and Terror 
eel (Mana, 222. 


Type locality. — Region of Cape of Good Hope, Union of South Africa. 


Range. — The occurrence of this cosmopolitan species at Mazatlan, Mexico, and the 
capture of a living specimen, storm-bound and washed ashore a short distance north of the 
United States-Mexican boundary at Imperial Beach, California (Hubbs, 1951b) offer ample 


evidence that Baja Californian waters are within the range of this rare species. 


Family DELPHINIDAE — Porpoises 
Genus Delphinus Linnaeus 


Delphinus bairdii Dall 
Baird’s Dolphin 
1873. Delphinus bairdii Dall, Proc. Calif. Acad. Sci. 5:12. January 29. (See Miller, 1936.) 


Type locality. — Cape (now Point) Arguello, Santa Barbara County, California. 


Range. — The Pacific coast of Baja California south from the United States boundary, 
and into the Gulf of California as far north as Escondido Bay, latitude 25°35’N. 


1963 | Hury: Mammats oF Baya CALIFORNIA 139 


Genus Tursiops Gervais 
Tursiops gillii Dall 
Gill’s Bottle-nosed Dolphin 
1873. Tursiops gilli Dall, Proc. Calif. Acad. Sci. 5:13. January 29. 
Type locality. — Monterey, California. 


Range. — Both coasts of Baja California; recorded from San Bartolomé Bay and the 
head of the Gulf of California at Puerto Pefasco, Sonora. 


Tursiops nuuanu Andrews 
Pacific Bottle-nosed Dolphin 


1911. Tursiops nuuanu Andrews, Bull. Amer. Mus. Nat. Hist. 30:233. August 26. 

Type locality. — North Pacific Ocean, approximately 12°N, 120°W. 

Range. — Two specimens of beached skulls from Santa Catalina Island, Gulf of Cali- 
fornia, constitute the known occurrence of this species in Baja California (Andrews, 1911). 


Genus Lagenorhynchus Gray __ 
Lagenorhynchus obliquidens Gill 


Striped Dolphin 
1865. Lagenorhynchus obliquidens Gill, Proc. Acad. Nat. Sci. Philadelphia 17:177. 


Type locality. — Pacific Ocean, near San Francisco, California. 
Range. — Along the entire Pacific coast of Baja California (Scheffer, 1950). 


Genus Orcinus Fitzinger 
Orcinus rectipinna (Cope) 
Pacific Killer Whale 
1869. Orca rectipinna Cope, Proc. Acad. Nat. Sci. Philadelphia 21:22. 
1961. Orcinus {rectipinna} Herschkovitz, Fieldiana-Zoology 39:549. July 25. 


Type locality. — Coast of California. 
Range. — Both Gulf and Pacific coasts of Baja California. 


Genus Pseudorca Reinhardt 
Pseudorca crassidens (Owen) 


False Killer Whale 


1846. Phocaena crassidens Owen, A History of British Fossil Mammals and Birds, p. 516. 
1862. Pseudorca crassidens Reinhardt (on title page to reprint of article from Overs. Danske 

Vid. Selsk. Forh., pp. 103-152). 

Type locality. — Lincolnshire Fens, England (subfossil) . 

Range. — Found in eastern Pacific Ocean from Puget Sound, Washington, to Acapulco, 
Guerrero. One record for Pichilinque Bay, near La Paz, places this species on the Baja California 
list (Miller, 1920). 

Genus Globicephala Lesson 
Globicephala scammonii Cope 


Scammon’s Blackfish 


1869. Globicephalus scammonii Cope, Proc. Acad. Nat. Sci. Philadelphia 21:21. 
1955. Globicephala scammonii Miller and Kellogg, Bull. U. S. Nat. Mus. 205 :664. 

Type locality. — Ten miles off Pacific coast in latitude 31° N, Baja California. 

Range. — Both coasts of Baja California, south from the United States-Mexican boundary 
on the Pacific side and well into the Gulf of California on the eastern side (Scammon, 1874:87). 


140 San Dteco Society of NaAturAL History { Vo. 13 


Genus Phocoena G. Cuvier 
Phocoena vomerina Gill 


Harbor Porpoise 


1865. Phocaena vomerina Gill, Proc. Acad. Nat. Sci. Philadelphia 17:178. 
1942. Phocoena vomerina Scheffer, Murrelet 23:45. August 14. 
Type locality. — Puget Sound, Washington. 


Range. — Along the Pacific coast of the peninsula, and into the Gulf for an unknown 
distance; ranges as far south as the Tres Marias Islands off the Mexican mainland. 


Phocoena sinus Norris and McFarland 
Gulf of California Harbor Porpoise 


1958. Phocoena sinus Norris and McFarland, Jour. Mamm. 39:24. February 20. 

Type locality. — Northeast shore of Punta San Felipe, Baja California Norte, Gulf of 
California, Mexico. 

Range. — To date, with certainty, only from the northern waters of the Gulf of California. 


Family ESCHRICHTIDAE — Gray Whale 
Genus Eschrichtius Gray 
Eschrichtius glaucus (Cope) 


Gray Whale 


1777. {Balaena} gibbosa Erxleben, Systema regni animalis . . . p. 610. (Based on “scrag whale” 
of Dudley, Phil. Trans. Roy. Soc. London 33:258, 1725. Coast of New England.) 


1868. Agaphelus glaucus Cope, Proc. Acad. Nat. Sci. Philadelphia 20:160. (Monterey Bay, 
California.) 

1869. Rhachianectes glaucus Cope, Proc. Acad. Nat. Sci. Philadelphia 21:15. 

1937. Eschrichtius gibbosus Van Deisne and Junge, Temminckia 2:181. 

1952. Eschrichtius glaucus Schevill, Breviora, Mus. Comp. Zool. 7:3. September 29. 


[Specific identity of gibbosus and glaucus regarded as not demonstrated (Miller and 
Kellogg, 1955). For further comments on the status of the name for the Gray Whale, see recom- 
mendation by Hershkovitz, 1961. } 


Type locality. — Monterey Bay, California. 


Range. — Regular visitor from January to April each year in the large lagoons and bays 
on the west coast of middle Baja California, where the calves are born and the species breeds. 
Occurs sparingly in the Gulf of California, where it has been observed as far north as the vicinity 
of Guaymas on the coast of Sonora. 


Family BALAENOPTERIDAE — Fin-backed Whales 
Genus Balaenoptera Lacépéde 
Balaenoptera physalus (Linnaeus) 
Common Fin-backed Whale 


1758. {Balaena} physalus Linnaeus, Systema Naturae, ed. 10, 1:75. 

1862. Balaenoptera physalus Schlegel, De Dieren van Nederland . . . Zoogdieren, p. 101. 
Type locality. — Spitzbergen seas. 
Range. — Found in deeper water off both Gulf and Pacific coasts of Baja California. 


1963 | Hury: MAmMaAts oF Baya CALIFORNIA i41 


Balaenoptera borealis Lesson 
Sei or Pollack Whale 


1828. Balaenoptera borealis Lesson, Histoire naturelle générale et particuliere des mammiféres 
et des oiseaux découverts depuis 1788 jusqu’a nos jours, { Half-title: } Complément des 
oeuvres de Buffon . . . complément 2, vol. 1, Cétacés, p. 342. 


Type locality. — Gromitz, Lubeck Bay, Schleswig-Holstein, Germany. 


Range. — Western North Pacific from Bering Sea south at least to waters off Baja Cali- 
fornia (Miller and Kellogg, 1955). 


Balaenoptera acutorostrata Lacépéde 
Sharp-headed Finner Whale 


1804. Balaenoptera acuto-rostrata Lacépéde, Histoire naturelle des Cétacées . . . Pp. XXXVil. 
Type locality. — European seas. 
Range. — Most abundant in North Pacific waters, and reported from Antarctica. 


A specimen was recorded for La Jolla, California (Abbott, 1930); the range is given by 
Miller and Kellogg (1955) as south at least to waters off Baja California. 


Genus Sibbaldus Gray 


Sibbaldus musculus (Linnaeus) 
Blue or Sulphur-bottom Whale 


1758. | Balaena} musculus Linnaeus, Systema naturae, ed. 10, 1:76. 
1923. Sibbaldus musculus Miller, Smiths. Misc. Coll. 76(5) :20. August 31. 
Type locality. — Firth of Forth, Scotland. 


Range. — Found in deeper waters off the west coast of Baja California (Scammon, 


1874:72). 


Genus Megaptera Gray 
Megaptera novaeangliae (Borowski) 
Hump-backed Whale 
1781. Balaena novae angliae Borowski, Gemeinniizzige Naturgeschichte des Thierreichs . . . 
2( 1221. 
1932. Megaptera novaeangliae Kellogg, Proc. Biol. Soc. Wash. 45:148. September 9. 
Type locality. — Coast of New England. 


Range. — This cosmopolitan species is found in fair numbers along the coasts of Baja 
California, ranging well up into the Gulf. 


Family BALAENIDAE — Right Whales 
Genus Balaena Linnaeus 
Balaena japonica Lacépéde 


Pacific Right Whale 


1818. Balaena japonica Lacépéde, Mém. Mus. Hist. Nat., Paris, 4:469, 472. 

1961. Balaena japonica Hershkovitz, Fieldiana-Zoology 39:561. July 25. 
Type locality. — Coast of Japan and northwest coast of North America. 
Range. — West coast of Baja California (Scammon, 1869:38). 


142 San Dreco Society of Naturav History { Vov. 13 


Order CARNIVORA — Carnivores 
Family CANIDAE — Wolves, Coyotes, Dogs and Foxes 
Genus Canis Linnaeus 
Canis latrans mearnsi Merriam 


Mearns Coyote 


1897. Canis mearnsi Merriam, Proc. Biol. Soc. Wash. 11:30. March 15. 
1932. Canis latrans mearnsi Nelson, Proc. Biol. Soc. Wash. 45:224. November 26. 

Type locality. — Quitobaquito, Pima County, Arizona. 

Range. — Over the lower deserts in extreme northeastern Baja California, south to El 
Major and around Laguna Salada; ranges into higher sections during warmer seasons. 


Canis latrans clepticus Elliot 
San Pedro Martir Coyote 


1903. Canis clepticus Elliot, Field Columb. Mus. Publ. 79, Zool. Ser. 3:225. August 15. 
1932. Canis latrans clepticus Nelson, Proc. Biol. Soc. Wash. 45:224. November 26. 
Type locality. — Vallecitos, Sierra San Pedro Martir, Baja California. Alt. 8500 feet. 
Range. — Over the Pacific slope of northwestern Baja California from the sierras west- 
ward to the Pacific Ocean and from the international boundary south through the high mountains 
to Llano de San Agustin and El Rosario. Ranges in wintertime over the lower desert areas 
including San Felipe, on the shores of the Gulf of California (Jackson in Young and Jackson, 
1951). 


Canis latrans peninsulae Merriam 


Peninsula Coyote 


1897. Canis peninsulae Merriam, Proc. Biol. Soc. Wash. 11:28. March 15. 
1932. Canis latrans peninsulae Nelson, Proc. Biol. Soc. Wash. 45:224. November 26. 

Type locality. — Santa Anita, Cape San Lucas, Baja California. 

Range. — Peninsula of Baja California south of latitude 30°N (Jackson in Young and 
Jackson, 1951). 


Genus Vulpes Bowdich 


Vulpes macrotis arsipus Elliot 
Desert Kit Fox 


1904. Vulpes arsipus Elliot, Field Columb. Mus. Publ. 87, Zool. Ser. 3:256. January 7. 
1913. Vulpes macrotis arsipus Grinnell, Proc. Calif. Acad. Sci., ser. 4, 3:287. August 28. 
Type locality. — Daggett, Mohave Desert, San Bernardino County, California. 
Range. — Found in small numbers in the sandy, arid desert sections of extreme north- 
eastern Baja California from the Colorado River west to Laguna Salada and south to San Felipe. 
Specimens from the latter locality are in the S. D. N. H. M. 


Vulpes macrotis tenuirostris Nelson and Goldman 
El Valle de la Trinidad Kit Fox 


1931. Vulpes macrotis tenuirostris Nelson and Goldman, Jour. Mamm. 12:302. August 24. 
Type locality. — Trinidad Valley, northwest base of Sierra San Pedro Martir, Baja 
California. Alt 2600 feet. 
Range. — Known only from the desert-like valley floor of El Valle de la Trinidad. 
There are specimens from this locality in the S. D. N. H. M. 


1963 | Huey: MAMMALS oF BajA CALIFORNIA 143 


Vulpes macrotis devia Nelson and Goldman 
Magdalena Plain Kit Fox 
1909. Vulpes macrotis devius Nelson and Goldman, Proc. Biol. Soc. Wash. 22:25. March 10. 
1955. Vulpes macrotis devia Miller and Kellogg, Bull. U. S. Nat. Mus. 205:636. March 3. 


Type locality. — Llano de Yrais, opposite Magdalena Island, Baja California. 
Range. — Sandy desert areas over the vast Magdalena Plain. 


Genus Urocyon Baird 
Urocyon cinereoargenteus scottii Mearns 


Arizona Gray Fox 


1891. Urocyon virginianus scottti Mearns, Bull. Amer. Mus. Nat. Hist. 3:236. June 5. 

1895. Urocyon cinereo-argenteus scottu J. A. Allen, Bull. Amer. Mus. Nat. Hist. 7:253. June 
29. 
Type locality. — Pinal County, Arizona. 
Range. — Through the lower desert section of northeastern Baja California south at least 


to San Felipe. 
Specimens from the latter locality are in the S. D. N. H. M. 


Urocyon cinereoargenteus californicus Mearns 
California Gray Fox 


1897. Urocyon cinereoargenteus californicus Mearns, Preliminary diagnoses of new mammals 
of the genera Lynx, Urocyon, Spilogale, and Mephitis, from the Mexican boundary 
line, p. 3. January 12. (Preprint of Proc. U. S. Nat. Mus. 20:459. December 24.) 
Type locality. — Tahquitz Valley, San Jacinto Mountains, Riverside County, California. 


Alt. 8000 feet. 

Range. — This subspecies is found over the Pacific drainage through the boulder- 
chaparral association of the higher hills and mountains from the international boundary south 
to the southern parts of the Sierra San Pedro Martir. 


Urocyon cinereoargenteus peninsularis Huey 
Lower California Gray Fox 


1928. Urocyon cinereoargenteus peninsularis Huey, Trans. San Diego Soc. Nat. Hist. 5:203. 
September 1. 
Type locality. — San Ignacio (27°24’N, 112°59’W), Baja California. 
Range. — Southern half of the peninsula of Baja California. 


Family URSIDAE — Bears 
Genus Ursus Linnaeus 
Ursus magister Merriam 


California Grizzly Bear 


1914. Ursus magister Merriam, Proc. Biol. Soc. Wash. 27:189. August 13. 

Type locality. — Los Biacitos, head of San Onofre Canyon, Santa Ana Mountains, San 
Diego County, California. 

Range. — The only record for bears in Baja California is to be found in trapper Pattie’s 
diary where he states that he had seen grizzly bears near Santa Catarina Mission in the Sierra 
Juarez in 1825 (Nelson, 1921:110). 

Ursus magister is the species that occurred in San Diego County and is presumably the 
species that formerly occupied the Sierra Juarez — thus the use of this name. There are no known 
specimens from Baja California. 


144 SAN Dreco Society of Naturav History { Vor. 13 


Family PROCYONIDAE — Raccoons and Allies 
Genus Bassariscus Coues 
Bassariscus astutus octavus Hall 


San Diego Ring-tailed Cat 


1926. Bassariscus astutus octavus Hall, Univ. Calif. Publ. Zool. 30:39. September 8. 
Type locality. — San Luis Rey River, near Escondido, San Diego County, California. 
Range. — Slopes and higher parts of the Sierra Juarez and Sierra San Pedro Martir 
south to an as yet undetermined locality. 
This species generally inhabits areas where large boulders and chaparral form the usual 
habitat of small rodents, on which the ring-tailed cat preys. There are specimens of B. a. octavus 


from the Sierra Juarez in the S. D. N. H. M. 


Bassariscus astutus palmarius Nelson and Goldman 
Palm Grove Ring-tailed Cat 


1909. Bassariscus astutus palmarius Nelson and Goldman, Proc. Biol. Soc. Wash. 22:26. 

March 10. 

Type locality. — Comondu, Baja California. 

Range. — Known from the rocky areas in central Baja California. 

This subspecies is especially abundant in the date-palm filled canyons of San Ignacio 
and Comondu, where rocky canyon rims offer ideal habitat and in which a large population of 
rodents exists. Specimens in the S. D. N. H. M. are from San Ignacio. 


Bassariscus astutus insulicola Nelson and Goldman 
San José Island Ring-tailed Cat 


1909. Bassariscus astutus insulicola Nelson and Goldman, Proc. Biol. Soc. Wash. 22:26. 
March 10. 
Type locality. — San José Island, Gulf of California, Baja California. 
Range. — San José Island. 


Bassariscus astutus saxicola Merriam 
Espiritu Santo Island Ring-tailed Cat 


1897. Bassariscus saxicola Merriam, Proc. Biol. Soc. Wash. 11:185. July 1. 

1926. Bassariscus astutus saxicola Hall, Univ. Calif. Publ. Zool. 30:46. September 8. 
Type locality. — Espiritu Santo Island, Gulf of California, Baja California. 
Range. — Espiritu Santo Island. 


Genus Procyon Storr 
Procyon lotor pallidus Merriam 


Pallid Raccoon 


1900. Procyon pallidus Merriam, Proc. Biol. Soc. Wash. 13:151. June 13. 
1923. Procyon lotor pallidus Grinnell, Univ. Calif. Publ. Zool. 21:316. January 27. 

Type locality. — New River (about 6 miles west of Imperial) , Colorado Desert, Imperial 
County, California. 

Range. — This subspecies is found throughout the delta of the Colorado River in extreme 
northeastern Baja California, from the international boundary south to the tidal estuaries of the 
Gulf and in cultivated areas wherever gravity water is carried by canals. 


1963 | Huey: MAmMMALs oF BAJA CALIFORNIA 145 


Procyon lotor psora Gray 
California Raccoon 


1842. Procyon psora Gray, Ann. Mag. Nat. Hist., ser. 1, 10:261. December. 
1914. Procyon lotor californicus Mearns, Proc. Biol. Soc. Wash. 27:66. March 20. (Ocean 
beach near last Mexican boundary monument { No. 258}, San Diego County, California.) 
1923. Procyon lotor psora Grinnell, Univ. Calif. Publ. Zool. 21:316. January 27. 
Type locality. — Sacramento, Sacramento County, California. 
Range. — Along the watercourses of the Pacific drainage from the higher mountains 
westward to the Pacific Ocean and south from the international boundary to El Rosario. 


Procyon lotor grinnelli Nelson and Goldman 
Lower California Raccoon 


1930. Procyon lotor grinnelli Nelson and Goldman, Jour. Wash. Acad. Sci. 20:82. March 4. 

Type locality. — La Paz, Baja California. 

Range. — This subspecies is found over the southern part of the peninsula from the mid- 
Vizcaino area southward. 

The greatest number inhabits the tidal lagoons where shellfish and crustaceans are the 
chief items of their food. 


Family MUSTELIDAE — Mustelids 
Genus Mustela Linnaeus 
Mustela frenata latirostra Hall 


Long-tailed Weasel 


1936. Mustela frenata latirostra Hall, Carnegie Inst. Wash. Publ. 473:96. November 20. 
Type locality. — San Diego, San Diego County, California. 
Range. — Extreme northwestern Baja California. 
A crushed specimen was found on the highway near Rosarito, ten miles south of Tijuana, 


June 1, 1962. 


Genus Taxidea Waterhouse 
Taxidea taxus berlandieri Baird 


Mexican Badger 


1858. Taxidea berlandieri Baird, Mammals, in Rep. Expl. Surv. Railr. to Pacific 8(1) :205. 
July 14. 
1895. Taxidea taxus berlandieri J. A. Allen, Bull. Amer. Mus. Nat. Hist. 7:256. June 29. 
Type locality. — Llano Estacado, Texas, near the border of New Mexico. 
Range. — Over the desert area from the Colorado River westward to the base of the 
coast range mountains and south to San Felipe. 
The writer observed a badger and found numerous workings at the latter locality in 


January, 1931. 


Taxidea taxus neglecta Mearns 
California Badger 


1891. Taxidea americana neglecta Mearns, Bull. Amer. Mus. Nat. Hist. 3:250. June 5. 
1901. Taxidea taxus neglecta Miller and Rehn, Proc. Boston Soc. Nat. Hist. 30:218. December 

27. 

Type locality. — Fort Crook, Shasta County, California. 

Range. — The coastal benches and dryer valleys from the Sierra Juarez and Sierra San 
Pedro Martir westward to the Pacific Ocean, and from the international boundary south to San 
Quintin. 


146 SAN Dieco Society or NaturAL History { VoL. 13 


Taxidea taxus infusca Thomas 
Lower California Badger 


1898. Taxidea taxus infusca Thomas, Proc. Zool. Soc. London for 1897, p. 899. April 1. 
Type locality. — Santa Anita, Baja California. 
Range. — Over the Ilanos and dryer plains of the southern parts of the peninsula from 
Punta Prieta south to Cape San Lucas. 


Genus Spilogale Gray 
Spilogale putorius martirensis Elliot 
San Pedro Martir Spotted Skunk 


1903. Spilogale arizonae martirensis Elliot, Field Columb. Mus. Publ. 74, Zool. Ser. 3:170. 

May 7. 

1906. S§ talaga microdon A. H. Howell, North Amer. Fauna 26:34. November 24. (Co- 
mondu, Baja California.) 
1959. Spilogale putorius martirensis Van Gelder, Bull. Amer. Mus. Nat. Hist. 117:369. 

June 15. 

Type locality. — Vallecitos, Sierra San Pedro Martir, Baja California. 

Range. — From the international boundary southward over the peninsula to Comondu 
(26°N). 

According to the latest reviser of this genus (Van Gelder, 1959), S. p. martirensis inter- 
grades with S. p. phenax along the international boundary and specimens from this region, while 
not typical of martirensis, can best be referred to the Baja California race. Specimens from the 
region about Comondu, the southern extent of the range of S. p. martirensis, bear the same 
relative position towards S. p. lucasana found in the Cape District as the population along the 
international boundary does towards S. p. phenax. 

However, there are great stretches down the peninsula between the southern San Pedro 
Martir and San Ignacio from which there are no specimens available and with the accession of 
a good series the status of the Spilogale of Baja California may well be re-examined. It is possible 
that the species is non-existant in much of this region. There are four specimens from Laguna 
Hanson, Sierra Juarez, which show their relationship both to S. p. phenax and to S. p. martirensis. 
One from the chaparral belt just north of El Valle de la Trinidad, 10 miles southeast of Alamo, 
is characteristic of S. p. martirensis. Four specimens from San Ignacio show intergradation with 
the southern form, S. p. lucasana. In this series the whole clinal gamut is shown. All of these 


specimens are in the S. D. N. H. M. 


Spilogale putorius lucasana Merriam 
Cape San Lucas Spotted Skunk 
1890. Spilogale lucasana Merriam, North Amer. Fauna 4:11. October 8. 
1959. Spilogale putorius lucasana Van Gelder, Bull. Amer. Mus. Nat. Hist. 117:374. June 15. 
Type locality. — Cape San Lucas, Baja California. 
Range. — Cape District, including the vast Magdalena Plain of the southern section of 
the peninsula. 


One specimen collected at Santo Domingo, Magdalena Plain, is in the S. D. N. H. M. 


Genus Mephitis E. Geoffroy Saint-Hilaire and G. Cuvier 
Mephitis mephitis estor Merriam 
Arizona Striped Skunk 

1890. Mephitus estor Merriam, North Amer. Fauna 3:81. September 11. 
1933. Mephitis mephitis estor Grinnell, Univ. Calif. Publ. Zool. 40: 108. September 26. 

Type locality. — Little Spring, north base San Francisco Mountain, Coconino County, 
Arizona. Alt. 8200 feet. 

Range. — Along the Colorado River Delta and the irrigated districts of northeastern 
Baja California. 


1963 | Hury: MAMMALS oF Baja CALIFORNIA 147 


Mephitis mephitis holzneri Mearns 
Southern California Striped Skunk 


1897. Mephitis occidentalis holzneri Mearns, Preliminary diagnoses of new mammals of the 
genera Lynx, Urocyon, Spilogale, and Mephitis, from the Mexican boundary line, p. 4. 
January 12. (Preprint of Proc. U. S. Nat. Mus. 20:461. December 24.) 

1933. Mephitis mephitis holzneri Grinnell, Univ. Calif. Publ. Zool. 40: 107. September 26. 
Type locality. —San Isidro Ranch, within 2 miles of the United States boundary and 

19 miles east of the Pacific sea coast, Baja California. 

Range. — Found over the chaparral covered slopes from the higher mountains westward 
to the coast and south to the Santo Domingo River and the northern end of the San Quintin 

Plain. The latter localities are the southernmost points known for striped skunks, Mephitis, in 


Baja California. 
Genus Enhydra Fleming 


Enhydra lutris nereis (Merriam) 
Southern Sea Otter 


1904. Latax lutris nereis Merriam, Proc. Biol. Soc. Wash. 17:159. October 6. 
1923. Enhydra lutris nereis Grinnell, Univ. Calif. Publ. Zool. 21:316. January 27. 

Type locality. — San Miguel Island, Santa Barbara Islands, California. 

Range. — Formerly, along the sea coast from the international boundary south to 
Sebastian Vizcaino Bay (Scammon, 1874:169). Now extinct in Baja California. 

Sea otter bones have been exhumed from kitchen middens near Rosarito Beach and 
Descanso Bay, between Ensenada and Tijuana. 


Family FELIDAE — Cats 
Genus Felis Linnaeus 


Felis concolor californica May 
California Mountain Lion 


1896. Felis californica May, California Game “marked down” p. 22. 
1929. Felis concolor californicus Nelson and Goldman, Jour. Mamm. 10:347. November 11. 
Type locality. — Kern County, California. 
Range. — Through the higher mountains and chaparral slopes from the international 
boundary south to the El Rosario River. 
This large cat preys on deer and is to be found in the general range of deer wherever 
they occur. 


Felis concolor browni Merriam 
Yuma Mountain Lion 


1903. Felis aztecus browni Merriam, Proc. Biol. Soc. Wash. 16:73. May 29. 
1929. Felis concolor browni Nelson and Goldman, Jour. Mamm. 10:347. November 11. 

Type locality. — Lower Colorado River, 12 miles south of Yuma, Yuma County, 
Arizona. 

Range. — Over the delta region of the Colorado River, and southward around the 
southern Sierra San Pedro Martir to the coast south of El Rosario. The southern limits of this 
race down the peninsula are unknown. 

A mountain lion was observed at close range by Gordon Marsh in mid-morning on 
April 5, 1950, on a cactus-covered mesa 25 miles south of El Rosario. At that time he was on 
an expedition in company of Charles F. Harbison, scouting for rare cacti. 


148 SAN Dieco Society oF Naturat History { VoL. 13 


Felis concolor improcera Phillips 
Lower California Mountain Lion 
1912. Felis improcera Phillips, Proc. Biol. Soc. Wash. 25:85. May 4. 
1929. Felis concolor improcera Nelson and Goldman, Jour. Mamm. 10:347. November 11. 
Type locality. — Calmalli, Baja California. 
Range. — Vizcaino Desert region and south to the mountains of Cape San Lucas. 
A specimen without skull from near Calmalli, of this pallid colored race, is in the 
S' Dy NeEM. 
Genus Lynx Kerr 
Lynx rufus californicus Mearns 


California Wildcat 

1897. Lynx rufus californicus Mearns, Preliminary diagnoses of new mammals of the genera 

Lynx, Urocyon, Spilogale, and Mephitis, from the Mexican boundary line, p. 2. January 

12. (Preprint of Proc. U. S. Nat. Mus. 20:458. December 24.) 

Type locality. — San Diego, San Diego County, California. 

Range. — Over the higher mountains westward to the Pacific Ocean and from the inter- 
national boundary southward to El Rosario and possibly beyond. 

This feline follows the range of California Quail, and wherever quail are abundant there 
also will be found wildcats. 

Lynx rufus baileyi Merriam 


Desert Wildcat 


1890. Lynx baileyi Merriam, North Amer. Fauna 3:79. September 11. 
1932. Lynx rufus baileyi V. Bailey, North Amer. Fauna 53:291. March 1. 

Type locality. — Mocassin Spring, north of Colorado River, Coconino County, Arizona. 

Range. — Over the Colorado River Delta in northeastern Baja California to the base of 
the western mountains, and from the international boundary southward over the deserts around 
the southern base of the Sierra San Pedro Martir to the Pacific coast, and south to or below 
Comondu. 

Specimens in the S. D. N. H. M. from San Ignacio, while not typical of L. r. baileyi, 
more closely resemble that subspecies than any other. A good series of this species from the 
central section of the peninsula might reveal characters that would set them apart from the other 
races. 

Lynx rufus peninsularis Thomas 
Cape San Lucas Wildcat 
1898. Lynx rufus peninsularis Thomas, Ann. Mag. Nat. Hist., ser. 7, 1:42. January. 
Type locality. — Santa Anita, Baja California. 
Range. — Cape region of Baja California. 


Order PINNIPEDIA — Seals and Sea Lions 
Family OTARIIDAE — Eared Seals 
Genus Callorhinus Gray 
Callorhinus ursinus cynocephalus (Walbaum) 


Pribilof Fur Seal 


1792. Siren cynocephala Walbaum, in Artedi, Petr. Artedi . . . ichthyologica, part 3, Genera 
piscium, p. 560. (Based on the sea ape of Steller; see Stejneger, Georg Wilhelm Steller 
... p. 285, August, 1936.) 
1940. Callorhinus ursina cynocephala Hall, Calif. Fish and Game 26:76. January. 
Type locality. — North Pacific Ocean south of the Alaska Peninsula at approximately 
53°N, 155° W (see Stejneger, op. cit., pl. 12, facing p. 278). 
Range. — A straggler has been observed off the coast of extreme northwestern Baja 
California, south of Los Coronados Islands. 


1963 | Huey: MAmMMALs oF BAJA CALIFORNIA 149 


Genus Arctocephalus E. Geoffroy Saint-Hilaire and F. Cuvier 
Arctocephalus philippii townsendi Merriam 


Guadalupe Fur Seal 


1897. Arctocephalus townsendi Merriam, Proc. Biol. Soc. Wash. 11:178. July 1. 
1958. Arctocephalus philippti townsendi Scheffer, Seals, Sea Lions and Walruses, A Review 
of the Pinnipedia, p. 80. April 24. 
Type locality. — Guadalupe Island, Baja California. 
Range. — Formerly abundant on Guadalupe Island but brought almost to the vanishing 
point by sealers in the latter part of the last century. A few escaped, and after almost three- 
fourths of a century the number has increased to between 200 and 500 seals (Scheffer, 1958:81). 


Genus Zalophus Gill 


Zalophus californianus californianus (Lesson) 
California Sea Lion 


1828. Otaria californiana Lesson, Dictionnaire Classique d’Histoire Naturelle 13:420. 

1858. Otaria gillespi (lapsus for gilliespii) MacBain, Proc. Royal Soc. Edinburgh 1:422. 
(Probably the northern end of Gulf of California. Type skull now in University of 
Edinburgh Anatomical Museum no. P.EU. ca 1; Schefter, 1958.) 

1958. Zalophus californianus californianus Scheffer, Seals, Sea Lions and Walruses, a Review 
of the Pinnipedia, p. 60. April 24. 

Type locality. — Rocks in vicinity of San Francisco Bay, California. 
Range. — Reefs and rocks, including near coast islands and outlying islands such as 

Guadalupe, along the entire coast of Baja California from the international boundary and Los 

Coronados Islands to near the head of the Gulf of California at Consag Rock. 


Family PHOCIDAE — Earless Seals 
Genus Phoca Linnaeus 
Phoca vitulina geronimensis J. A. Allen 


California Harbor Seal 


1902. Phoca richardii geronimensis J. A. Allen, Bull. Amer. Mus. Nat. Hist. 16:495. December 

12: 

1942. Phoca vitulina geronimensis Doutt, Ann. Carnegie Mus. 29:116-117. May 12. 

Type locality. — San Gerénimo Island, Baja California. 

Range. — Mainly coastwise, from San Ignacio Lagoon, San Gerénimo and San Martin 
islands north to the international boundary. 

The latest reviser of this group (Scheffer, 1958) has treated P. r. geronimensis as a 
synonym of P. r. richardi. After closely scrutinizing the painstaking review of the group by 
Doutt (1942), who considered all aspects and compared numerous specimens, the writer feels 
that P. v. geronimensis is a valid subspecies and should be recognized. 


Genus Mirounga Gray 
Mirounga angustirostris (Gill) 


Northern Elephant Seal 


1866. Macrorhinus angustirostris Gill, Comm. Essex Inst. 5(1):13. April 7. Proc. Chicago 
Acad. Sci. 1:33. April, 1866. 

1904. [Mirounga} angustirostris Elliot, Field Columb. Mus. Publ. 95, Zool. Ser. 4:545. 
August 2. 
Type locality. — San Bartolomé Bay, Baja California. 


150 SAN Drteco Society or Natura History { Vo. 13 


Range. — Pacific coast from international boundary south to the San Benitos Islands. 

During the heyday of Pacific coast whaling and sealing, elephant seals were sought for 
their oil and were reduced to near extinction. In the year 1892 only nine individuals were found 
on Guadalupe Island and the greater number of these were collected for museum specimens 
(Huey, 1930). Since this population low, the species, through isolation and lack of further pur- 
suit, has made a prodigious increase in numbers. At present the number is estimated at from 
8 to 10 thousand (Scheffer, 1958). This increase in numbers has caused the animals to occupy, 
again, some sections of their ancestral range. Assembled groups have been found from the Santa 
Barbara Islands south to the San Benitos Islands from latitude 36°N south to 26°N. Further 
population pressure probably will cause further range expansion; “‘scouts” have been noted as 
far north as the coasts of Washington and British Columbia. 


Order ARTIODACTYLA — Even-toed Ungulates 
Family CERVIDAE — Deer 
Genus Odocoileus Rafinesque 
Odocoileus hemionus fuliginatus Cowan 


Southern Black-tailed Deer 


1933. Odocoileus hemionus fuliginatus Cowan, Jour. Mamm. 14:326. November 13. 

Type locality. —Barona Ranch, 30 miles east of San Diego, San Diego County, Cali- 
fornia. 

Range. — Over the higher brush and forested areas of the Pacific slopes from the inter- 
national boundary south to the southern Sierra San Pedro Martir and the San Quintin-El 
Rosario region. South of this area, over the cactus-covered mesas, it intergrades with the penin- 
sular form. 

The name Odocoileus is here used in place of Dama in conformance with Opinion 58 
(September 16, 1960) of The International Zoological Commission. 


Odocoileus hemionus cerrosensis Merriam 
Cedros Island Black-tailed Deer 


1898. Odocoileus cerrosensis Merriam, Proc. Biol. Soc. Wash. 12:101. April 30. 
1915. Odocoileus hemionus cerrosensis Lydekker, Catalogue of the Ungulate Mammals in 
the. . . British Museum, 4: 180. 
Type locality. — Cerros { =Cedros } Island, off the west coast of Baja California. 
Range. — Cedros Island, Baja California. 
Due to over-hunting by residents of the island, this subspecies is fast approaching extinc- 


tion. There are three specimens in the S. D. N. H. M. 


Odocoileus hemionus peninsulae (Lydekker) 
Cape San Lucas Black-tailed Deer 


1898. Mazama (Dorcelaphus) hemionus peninsulae Lydekker, Proc. Zool. Soc. London for 
1897, pp. 899-900. April. 
1901. Odocoileus hemionus peninsulae Miller and Rehn, Proc. Boston Soc. Nat. Hist. 30:16. 
December 27. 
Type locality. — Between La Laguna and Victoria Mountain, Sierra Laguna, Baja Calli- 
fornia. Alt. about 6000 feet. 
Range. — From the southern peninsula district northward in an irregular range over 
the cactus-covered mesas to Aguaita, below El Rosario. 
A deer, the status of which is unknown, lives on San José Island (latitude 25°N), Gulf 
of California. There is one specimen of this deer in the S. D. N. H. M. 


1963 } Huery: MammMats oF Baja CALIFORNIA 151 


Family ANTILOCAPRIDAE — Prong-horn 
Genus Antilocapra Ord 
Antilocapra americana americana (Ord) 


American Prong-horn Antelope 


1815. Antilope americana Ord in Guthrie, A new Geographical, Historical and Commercial 

Grammar . . . Philadelphia, ed. 2, 2:292 (described on p. 308). 

1818. Antilocapra americana Ord, Journ. Phys. Chim. Hist. Nat. et Arts, 87:149. 

Type locality. — Plains and highlands of the Missouri River. 

Range. — American Prong-horn Antelope once ranged over the deserts of northeastern, 
and the coastal mesas of northwestern, Baja California but have long since extirpated. 

The last account of the species for the Pacific side was in 1882-83 (Van Dyke, 1888:73) 
where the record reveals antelope to have ranged over the mesas between El Cajon and Otay 
Mesa, California, localities very close to the international boundary. 

Another record of extreme interest was found in the translation of the log of Juan 
Rodriguez Cabrillo (Ferrel, 1879:304) when he discovered and landed at what is now known 
as San Diego Bay, September 17, 1542. He records hundreds of animals observed, the descrip- 
tions of which can only be applied to Prong-horns. Thus, the Prong-horn Antelope is the first 
species of mammal to be recorded for California. 

The writer, as an apprentice working in a local taxidermist shop, vividly recalls freshly- 
killed antelope heads, taken in El Valle de la Trinidad by Mr. Walter Dupee, a local wealthy 
big game hunter, in the years 1908 and 1909. Subsequent trips by the writer in later years have 
failed to record the species from that locality. 


Antilocapra americana peninsularis Nelson 
Lower California Prong-horn Antelope 


1912. Antilocapra americana peninsularis Nelson, Proc. Biol. Soc. Wash. 25:107. June 29. 

Type locality. — Forty-five miles south of Calmalli, Baja California. 

Range. — Formerly, antelopes of this subspecies ranged from below San Felipe on the 
eastern coast and from San Quintin on the western coast south over the Ilanos and plains of the 
peninsula to the vicinity of Magdalena Bay. 

Today the subspecies totters on the verge of extinction with a few reputedly living on 
the arid remote desert Ilanos just north and south of Bahia de Los Angeles on the east coast 
and a larger group in the Vizcaino Desert east of Scammon’s Lagoon on the western side of 
the peninsula. It is safe to predict that within the next 20 years this race will have disappeared 
from Baja California. 


Family BOVIDAE — Bovids 
Genus Ovis Linnaeus 
Ovis canadensis cremnobates Elliot 


Sierra San Pedro Martir Bighorn 


1904. Orvis cervina cremnobates Elliot, Field Columb. Mus. Publ. 87, Zool. Ser. 3:239. January 
ip 
1912. Ovis canadensis cremnobates Miller, U. S. Nat. Mus. Bull. 79:396. December 31. 

Type locality. — Mattomi, Sierra San Pedro Martir, Baja California. 

Range. — Bighorn sheep are found in scattered small numbers on the steeper remote 
desert slopes from the international boundary southward through the Sierra Juarez and the 
Sierra San Pedro Martir to the vicinity of Los Angeles Bay. , 

Formerly a few lived on the Cocopah range, southwest of Mexicali, but according to 
late accounts this small herd has been exterminated. Some live on the San Carlos Mesa, an 
elevated plateau on the Pacific side, which lies coastwise southwest of the San Fernando Mission. 


152 SAN Dreso Society oF Naturat History {Vot. 13 


Bighorn Sheep from this population are often found on the immediate coast where fishermen, 
landing through the surf, have killed them near the beach. A female from San Carlos Mesa 
is in the S. D. N. H. M. 

The Bighorn Sheep of Baja California are doubtfully holding their own. As the human 
population increases, sheep are killed at every opportunity. The method of hunting is to locate 
remote springs or watering places, lie in wait, and kill as many as possible when the sheep 
come to drink. This method is especially successful in the dry summer. Only remote desert 
isolation leaves any hope of long survival and such regions will not support large populations. 


Ovis canadensis weemsi Goldman 
Sierra Giganta Bighorn 


1937. Ovis canadensis weemsi Goldman, Proc. Biol. Soc. Wash. 50:30. April 2. 

Type locality. — Cajon de Tecomaja, Sierra de la Giganta, about 30 miles south of 
Cerro de la Giganta, southern Baja California. 

Range. — This race of Bighorn Sheep lives in very limited numbers along the moun- 
tainous backbone of the peninsula from the vicinity of Los Angeles Bay southward into and 
through the Sierra de la Giganta. 

The mountainous region that comprises the greater part of the habitat of O. c. weemsi 
is chiefly lava; hence the darker pelage color. 

Ovis canadensis weemsi, like its more northern relative, is being reduced in numbers by 
hunters and it is only a matter of time until the population is reduced beyond the point of 
possible recovery. 


1963 | Hury: Mammats oF BajJA CALIFORNIA 153 


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1963 | Huey: Mammats oF Baya CALIFORNIA 


abbreviata, Neotoma, 134 
acutorostrata, Balaenoptera, 141 
Agaphelus, 140 
glaucus, 140 
agilis, Dipodomys, 119-120 
albatus, Thomomys, 105 
albigula, Neotoma, 131 
alexandrinus, Mus, 137 
americana, Antilocapra, 151 
Antilope, 151 
Taxidea, 145 
americanus, Sitomys, 127 
Ammospermophilus, 102-103 
insularis, 103 
leucurus leucurus, 102 
leucurus peninsulae, 102 
leucurus canfieldae, 103 
leucurus extimus, 103 
leucurus insularis, 103 
angustirostris, Mirounga, 149 
anthonyi, Neotoma, 135 
Perognathus, 116 
Scapanus, 92 
Antiocapridae, 151 
Antilocapra, 151 
americana americana, 151 
americana peninsularis, 151 
Antilope, 151 
americana, 151 
antiquarius, Dipodomys, 121 
Antrozous, 98 
minor, 98 
pallidus pacificus, 98 
aphrastus, Thomomys, 106 
Arctocephalus, 149 
philippii townsendi, 149 
townsendi, 149 
arenacea, Neotoma, 135 
arenarius, Perognathus, 113-115 
arizonae, Lepus, 100 
Spilogale, 146 
arsipus, Vulpes, 142 
asiaticus, Tamias, 102 
astutus, Bassariscus, 144 
Atalapha, 97 
teliotis, 97 
atricapillus, Spermophilus, 103 
auduboni, Sylvilagus, 100 
aztecus, Felis, 147 
bachmani, Sylvilagus, 99-100 
baileyi, Lynx, 148 
Perognathus, 111-112 
bairdii, Delphinus, 138 
Balaena, 140-141 
gibbosa, 140 
japonica, 141 
musculus, 141 
novae angliae, 141 
physalus, 140 
Balaenidae, 141 
Balaenoptera, 140 
acuto-rostrata, 14] 
borealis, 141 
physalus, 140 


Balaenopteridae, 140-141 
Balantiopteryx, 92 
plicata pallida, 92 
Bassariscus, 144 
astutus insulicola, 144 
astutus octavus, 144 
astutus palmarius, 144 
astutus saxicola, 144 
saxicola, 144 
beecheyi, Citellus, 103 
Spermophilus, 103 
bella, Neotoma, 132 
bennettii, Lepus, 100 
berlandieri, Taxidea, 145 
bombycinus, Perognathus, 111 
borealis, Balaenoptera, 141 
Lasiurus, 97 
bottae, Thomomys, 105-109 
Bovidae, 151-152 
boylii, Peromyscus, 130 
brasiliensis, Tadarida, 98 
breviceps, Kogia, 138 
Physeter, 138 
bryanti, Neotoma, 135 
Perognathus, 118 
bunkeri, Neotoma, 135 
cabezonae, Perodipus, 119 
californiana, Otaria, 149 
californianus, Zalophus, 149 
californica, Felis, 147 
californicus, Lepus, 100-101 
Macrotus, 93 
Microtus, 136 
Myotis, 95 
Perognathus, 116 
Peromyscus, 125 
Sorex, 91 
Callorhinus, 148 
ursinus cynocephalus, 148 
canadensis, Ovis, 151-152 
caniceps, Peromyscus, 127 
Canidae, 142-143 
Canis, 142 
clepticus, 142 
latrans clepticus, 142 
latrans mearnsi, 142 
latrans peninsulae, 142 
mearnsi, 142 
peninsulae, 142 
capitaneus, Myotis, 94 
carolinensis, Sciurus, 104 
Castor, 124 
canadensis repentinus, 124 
Castoridae, 124 
catodon, Physeter, 138 
cavirostris, Ziphius, 138 
cedrosensis, Peromyscus, 126 
cerrosensis, Lepus, 99 
Odocoileus, 150 
cervina, Ovis, 151 
Cervidae, 150 
Choeronycteris, 93 
mexicana, 93 
cinerascens, Lepus, 99 


159 


160 San DigEGo Society or NATURAL History 


cinereoargenteus, Urocyon, 143 
cinereus, Lasiurus, 97 
Vespertilio, 97 
cineritius, Peromyscus, 129 
Citellus, 103 
beecheyi nudipes, 103 
beecheyi rupinarum, 103 
tereticaudus apricus, 104 
tereticaudus vociferans, 104 
clepticus, Canis, 142 
concolor, Felis, 147-148 
Corynorhinus, 97 
macrotis pallescens, 97 
Cricetidae, 124-136 
cynocephala, Siren, 148 
Dasypterus, 97 
ega xanthinus, 97 
Delphinidae, 138-140 
Delphinus, 138 
bairdii, 138 
deserti, Dipodomys, 124 
dickeyi, Peromyscus, 127 
Didelphidae, 91 
Didelphis, 91 
marsupialis virginiana, 91 
virginiana, 91 
Dipodomys, 119-124 
agilis cabezonae, 119 
agilis latimaxillaris, 119 
agilis martirensis, 120 
agilis peninsularis, 120 
agilis plectilis, 120 
agilis simulans, 119 
antiquarius, 121 
deserti deserti, 124 
gravipes, 121 
insularis, 123 
margaritae, 123 
merriami annulus, 122 
merriami arenivagus, 121 
merriami brunensis, 122 
merriami llanoensis, 122 
merriami melanurus, 123 
merriami merriami, 121 
merriami platycephalus, 122 
merriami quintinensis, 122 
merriami semipallidus, 122 
merriami trinidadensis, 121 
paralius, 120 
peninsularis australis, 120 
peninsularis eremoecus, 120 
peninsularis pedionomus, 120 
peninsularis peninsularis, 120 
platycephalus, 122 
simulans peninsularis, 120 
domesticus, Mus, 137 
douglasii, Tamiasciurus, 105 
dubius, Peromyscus, 128 
edwardsi, Lepus, 101 
ega, Dasypterus, 97 
Emballonuridae, 92 
Enhydra, 147 
lutris nereis, 147 
Eptesicus, 96 
fuscus bernardinus, 96 


fuscus pallidus, 96 


fuscus peninsulae, 97 

pallidus, 96 
eremicus, Hesperomys, 125 

Peromyscus, 125-127 
Eschrichtidae, 140 
Eschrichtius, 140 

gibbosus, 140 

glaucus, 140 
estor, Mephitis, 146 
Eutamias, 102 

merriami meridionalis, 102 

merriami merriami, 102 

merriami obscurus, 102 
eva, Peromyscus, 127 
evotis, Myotis, 94 
exiguus, Peromyscus, 128 
fallax, Perognathus, 115-116 
Felidae, 147-148 
Felis, 147 

aztecus browni, 147 

californica, 147 

concolor browni, 147 

concolor californicus, 147 

concolor improcera, 148 

improcera, 148 
femoralis, Perognathus, 116 
femorosacca, Tadarida, 98 
femorosaccus, Nyctinomus, 98 
floridanus, Lepus, 100 
formosus, Perognathus, 111 
fossor, Sciurus, 104 
fraterculus, Vesperimus, 126 
frenata, Mustela, 145 
fulvus, Thomomys, 105, 107, 110 
fuscipes, Neotoma, 135 
fuscus, Eptesicus, 96-97 

Vespertilio, 97 
gambelii, Hesperomys, 127 
gaurus, Peromyscus, 130 
Geomyidae, 105-110 
geronimensis, Peromyscus, 129 
gibbosa, Balaena, 140 
gibbosus, Eschrichtius, 140 
gillespii, Otaria, 149 
gillii, Tursiops, 139 
glaucus, Agaphelus, 140 

Eschrichtius, 140 

Rhachianectes, 140 
Globicephala, 139 

scammonii, 139 
Globicephalus, 139 

scammonu, 139 
grammurus, Spermophilus, 103 
gravipes, Dipodomys, 121 
griseus, Sciurus, 104 
guardia, Peromyscus, 127 
helleri, Perognathus, 113 
hemionotis, Peromyscus, 130 
hemionus, Mazama, 150 

Odocoileus, 150 
Hesperomys, 125 

eremicus, 125 

gambelii, 127 

sonoriensis, 128 
hesperus, Pipistrellus, 96 

Scotophilus, 96 


1963 } Huey: Mammats or Baja CALIFORNIA 


Heteromyidae, 110-124 
hispidus, Sigmodon, 131 
homochroia, Peromyscus, 126 
hudsonius, Sciurus, 105 
improcera, Felis, 148 
insignis, Peromyscus, 125 
insularis, Ammospermophilus, 103 
Dipodomys, 123 
Lepus, 101 
Neotoma, 132 
intermedia, Neotoma, 132, 134 
juncensis, Sorex, 91 
japonica, Balaena, 141 
Kerivoula, 95 
pallida, 95 
Kogia, 138 
breviceps, 138 
Kogiidae, 138 
Lagenorhynchus, 139 
obliquidens, 139 
lagunae, Sorex, 91 
Lasiurus, 97 
borealis teliotis, 97 
cinereus cinereus, 97 
ega xanthinus, 97 
Latax, 147 
lutris nereis, 147 
latimanus, Scapanus, 92 
latrans, Canis, 142 
lepida, Neotoma, 132-135 
Leporidae, 99-101 
Lepus, 99, 100, 101 
arizonae confinis, 100 
bennettii, 100 
californicus bennettii, 100 
californicus deserticola, 101 
californicus magdalenae, 101 
californicus martirensis, 101 
californicus sheldoni, 101 
californicus xanti, 101 
cerrosensis, 99 
cinerascens, 99 
floridanus sanctidiegi, 100 
insularis, 101 
martirensis, 101 
peninsularis, 100 
sylvaticus, 100 
texianus deserticola, 101 
leucopus, Peromyscus, 129 
leucurus, Ammospermophilus, 102-103 
Tamias, 102 
longicauda, Reithrodon, 124 
longicrus, Myotis, 95 
longimembris, Perognathus, 110 
lotor, Procyon, 144-145 
lucasana, Spilogale, 146 
lutris, Enhydra, 147 
Latax, 147 
Lynx, 148 
baileyi, 148 
rufus baileyi, 148 
rufus californicus, 148 
rufus peninsularis, 148 
macrotis, Corynorhinus, 97 
Neotoma, 135 
Onychomys, 131 


Tadarida, 99 
Vulpes, 142-143 
Macrotus, 93 
californicus, 93 
magdalenae, Thomomys, 109 
magister, Ursus, 143 
maniculatus, Peromyscus, 127-129 
mansuetus, Sylvilagus, 100 
margaritae, Dipodomys, 123 
marsupialis, Didelphis, 91 
martinensis, Neotoma, 135 
Sitomys, 130 
martirensis, Lepus, 101 
Mazama, 150 
hemionus peninsulae, 150 
mearnsi, Canis, 142 
megalotis, Reithrodon, 124 
Reithrodontomys, 124-125 
Megaptera, 141 
novaeangliae, 141 
melanorhinus, Vespertilio, 95 
Mephitis, 146 
estor, 146 
mephitis estor, 146 
mephitis holzneri, 147 
occidentalis holzneri, 147 
merriami, Dipodomys, 121-123 
Eutamias, 102 
Vesperugo, 96 
mexicana, Choeronycteris, 93 
Tadarida, 98 
mexicanus, Molossus, 98 
Natalus, 93 
microdon, Spilogale, 146 
micronyx, Myotis, 94 
Microtus, 136 
californicus aequivocatus, 136 
californicus grinnelli, 136 
californicus huperuthrus, 136 
californicus hyperythrus, 136 
californicus neglectus, 136 
californicus sanctidiegi, 136 
milleri, Myotis, 94 
minor, Antrozous, 98 
Mirounga, 149 
angustirostris, 149 
molossa, Tadarida, 99 
Molossidae, 98 
Molossus, 98 
mexicanus, 98 
molossus, Vespertilio, 99 
Muridae, 137 
musculus, Mus, 137 
Sibbaldus, 141 
Mustela, 145 
frenata latirostra, 145 
Mustelidae, 145 
Mus, 137 
alexandrinus, 137 
domesticus, 137 
musculus domesticus, 137 
norvegicus, 137 
rattus, 137 
Myotis, 93-95 
californicus californicus, 95 
californicus pallidus, 95 


161 


162 SAN Disco Society oF NATURAL History 


californicus stephensi, 95 
evotis evotis, 94 
longicrus interior, 95 
micronyx, 94 
milleri, 94 
orinomus, 95 
peninsularis, 94 
subulatus melanorhinus, 95 
thysanodes thysanodes, 94 
velifer peninsularis, 94 
vivesi, 95 
volans volans, 94 
volans interior, 95 
yumanensis lambi, 94 
yumanensis sociabilis, 93 
yumanensis yumanensis, 93 
Natalidae, 93 
Natalus, 93 
mexicanus, 93 
stramineus mexicanus, 93 
Neotoma, 131-135 
abbreviata, 134 
albigula venusta, 131 
anthonyi, 135 
arenacea, 135 
bella felipensis, 132 
bryanti, 135 
bunkeri, 135 
fuscipes macrotis, 135 
fuscipes martirensis, 135 
insularis, 132 
intermedia, 132 
intermedia gilva, 132 
intermedia notia, 134 
intermedia perpallida, 134 
intermedia pretiosa, 134 
intermedia ravida, 133 
intermedia vicina, 134 
lepida abbreviata, 134 
lepida arenacea, 135 
lepida aridicola, 133 
lepida egressa, 132 
lepida felipensis, 132 
lepida intermedia, 132 
lepida insularis, 132 
lepida gilva, 132 
lepida latirostra, 134 
lepida marcosensis, 133 
lepida molagrandis, 133 
lepida notia, 134 
lepida nudicauda, 133 
lepida perpallida, 134 
lepida pretiosa, 134 
lepida ravida, 133 
lepida vicina, 134 
macrotis, 135 
martinensis, 135 
nudicauda, 133 
venusta, 131 
norvegicus, Mus, 137 
Rattus, 137 
Notiosorex, 92 
crawfordi crawfordi, 92 
novae angliae, Balaena, 141 
Megaptera, 141 
nudicauda, Neotoma, 133 


nuuanu, Tursiops, 139 
Nyctinomus, 98 
femorosaccus, 98 
obliquidens, Lagenorhynchus, 139 
obscurus, Tamias, 102 
occidentalis, Mephitis, 147 
Odocoileus, 150 
cerrosensis, 150 
hemionus cerrosensis, 150 
hemionus fuliginatus, 150 
hemionus peninsulae, 150 
Ondatra, 136 
zibethicus bernardi, 136 
Onychomys, 130-131 
macrotis, 131 
pulcher, 130 
ramona, 131 
torridus macrotis, 131 
torridus pulcher, 130 
torridus ramona, 131 
Orca, 139 
rectipinna, 139 
Orcinus, 139 
rectipinna, 139 
oreinus, Sorex, 91 
oresterus, Peromyscus, 128 
orinomus, Myotis, 95 
ornatus, Sorex, 91 
Oryzomys, 124 
peninsulae, 124 
Otaria, 149 
californiana, 149 
gillespii, 149 
gilliespii, 149 
Otariidae, 148-149 
Ovis, 151-152 
canadensis cremnobates, 151 
canadensis weemsi, 152 
cervina cremnobates, 151 
pallida, Kerivoula, 95 
pallidus, Antrozous, 98 
Eptesicus, 96 
Procyon, 144 
paralius, Dipodomys, 120 
penicillatus, Perognathus, 113, 115 
peninsulae, Canis, 142 
Reithrodontomys, 125 
peninsularis, Dipodomys, 120-121 
Lepus, 100 
Myotis, 94 
Perodipus, 119-120 
cabezonae, 119 
simulans peninsularis, 120 
streatori simulans, 119 
Perognathus, 110-119 
anthonyi, 116 
arenarius albescens, 113 
arenarius albulus, 115 
arenarius ambiguus, 113 
arenarius ammophilus, 115 
arenarius arenarius, 114 
arenarius helleri, 113 
arenarius mexicals, 113 
arenarius paralios, 113 
arenarius sabulosus, 114 
arenarius siccus, 115 


{Vot. 13 


1963 } Huey: Mammats oF Baya CALIFORNIA 


arenarius sublucidus, 114 
baileyi extimus, 112 
baileyi fornicatus, 112 
baileyi hueyi, 111 
baileyi mesidios, 112 
baileyi rudinoris, 112 
bombycinus, 111 
bryanti, 118 
californicus femoralis, 116 
californicus mesopolius, 116 
evermanni, 117 
fallax fallax, 115 
fallax inopinus, 116 
fallax majusculus, 116 
fallax pallidus, 115 
fallax xerotrophicus, 116 
femoralis mesopolius, 116 
formosus cinerascens, 111 
formosus infolatus, 111 
formosus mesembrinus, 111 
helleri, 113 
knekus, 112 
longimembris aestivus, 110 
longimembris bombycinus, 111 
longimembris internationalis, 110 
longimembris venustus, 110 
margaritae, 119 
mesembrinus, 111 
penicillatus albulus, 115 
penicillatus ammophilus, 115 
pencillatus angustirostris, 113 
penicillatus siccus, 115 
spinatus broccus, 118 
spinatus bryanti, 118 
spinatus evermanni, 117 
spinatus guardiae, 117 
spinatus lambi, 119 
spinatus latijugularis, 118 
spinatus magdalenae, 119 
spinatus marcosensis, 118 
spinatus margaritae, 119 
spinatus nelsoni, 118 
spinatus peninsulae, 119 
spinatus prietae, 117 
spinatus pullus, 118 
spinatus occultus, 118 
spinatus oribates, 117 
spinatus rufescens, 117 
spinatus seorsus, 118 
spinatus spinatus, 117 
Peromyscus, 125-130 
boylii rowleyi, 130 
californicus insignis, 125 
caniceps, 127 
cedrosensis, 126 
cineritius, 129 
crinitus pallidissimus, 125 
crinitus stephensi, 125 
dickeyi, 127 
dubius, 128 
eremicus avius, 126 
eremicus carmeni, 126 
eremicus cedrosensis, 126 
eremicus cinereus, 126 
eremicus eremicus, 125 
eremicus eva, 127 


eremicus fraterculus, 126 
eremicus insulicola, 126 
eremicus polypolius, 126 
eremicus propinquus, 127 
eva, 127 
exiguus, 128 
gaurus, 130 
geronimensis, 129 
guardia guardia, 127 
guardia interparietalis, 127 
guardia mejiae, 127 
hemionotis, 130 
insignis, 125 
leucopus coolidgei, 129 
maniculatus assimilis, 128 
maniculatus cineritius, 129 
maniculatus coolidgei, 128 
maniculatus dorsalis, 129 
maniculatus dubius, 128 
maniculatus exiguus, 128 
maniculatus gambelii, 127 
maniculatus geronimensis, 129 
maniculatus hueyi, 129 
maniculatus magdalenae, 129 
maniculatus margaritae, 129 
maniculatus martinensis, 128 
maniculatus sonoriensis, 128 
oresterus, 128 
pseudocrinitus, 125 
sejugis, 130 
slevini, 130 
stephensi, 125 
texanus medius, 128 
truei lagunae, 130 
truei martirensis, 130 
philippii, Arctocephalus, 149 
Phoca, 149 
richardii geronimensis, 149 
vitulina geronimensis, 149 
Phocidae, 149 
Phocoena, 140 
sinus, 140 
vomerina, 140 
Phyllostomidae, 93 
physalus, Balaena, 140 
Balaenoptera, 140 
Physeter, 138 
breviceps, 138 
catodon, 138 
Physeteridae, 138 
Pipistrellus, 96 
hesperus australis, 96 
hesperus hesperus, 96 
hesperus merriami, 96 
Pizonyx, 95 
vivesi, 95 
platycephalus, Dipodomys, 122 
Plecotus, 97 
townsendii pallescens, 97 
plicata, Balantiopteryx, 92 
Procyon, 144-145 
lotor californicus, 145 
lotor grinnelli, 145 
lotor pallidus, 144 
lotor psora, 145 
pallidus, 144 


163 


164 San Deco Society oF NATURAL History 


psora, 145 
Procyonidae, 144-145 
psora, Procyon, 145 
pulcher, Onychomys, 130 
ramona, Onychomys, 131 
Rattus, 137 
norvegicus, 137 
rattus alexandrinus, 137 
rattus rattus, 137 
rattus, Mus, 137 
Rattus, 137 
rectipinna, Orca, 139 
Orcinus, 139 
Reithrodon, 124 
longicauda, 124 
megalotis, 124 
Reithrodontomys, 124-125 
megalotis longicaudus, 124 
megalotis megalotis, 124 
megalotis peninsulae, 125 
peninsulae, 125 
Rhachianectes, 140 
glaucus, 140 
richardii, Phoca, 149 
rowleyi, Sitomys, 130 
rufus, Lynx, 148 
saxicola, Bassariscus, 144 
scammonii, Globicephala, 139 
Globicephalus, 139 
Scapanus, 92 
anthonyi, 92 
latimanus anthonyi, 92 
latimanus occultus, 92 
Sciuridae, 102-105 
Sciurus, 104 
carolinensis carolinensis, 104 
fossor anthonyi, 104 
griseus anthonyi, 104 
hudsonius mearnsi, 105 
Scotophilus, 96 
hesperus, 96 
sejugis, Peromyscus, 130 
Sibbaldus, 141 
musculus, 141 
Sigmodon, 131 
hispidus eremicus, 131 
simulans, Dipodomys, 120 
Perodipus, 120 
sinus, Phocoena, 140 
Siren, 148 
cynocephala, 148 
Sitomys, 127, 130 
americanus thurberi, 127 
martirensis, 130 
rowleyi, 130 
slevini, Peromyscus, 130 
sonoriensis, Hesperomys, 128 
Sorex, 91 
californicus juncensis, 91 
crawfordi, 92 
juncensis, 91 
lagunae, 91 
oreinus, 91 
ornatus lagunae, 91 
ornatus ornatus, 91 


Soricidae, 91 


Spermophilus, 103-104 
atricapillus, 103 
beecheyi nudipes, 103 
beecheyi rupinarum, 103 
grammutus atricapillus, 103 
tereticaudus apricus, 104 
tereticaudus tereticaudus, 104 

Spilogale, 146 
arizonae martirensis, 146 
lucasana, 146 
microdon, 146 
putorius lucasana, 146 
putorius martirensis, 146 

spinatus, Perognathus, 117-119 

stephensi, Peromyscus, 125 

stramineus, Natalus, 93 

streatori, Perodipus, 119 

subulatus, Myotis, 95 

sylvaticus, Lepus, 100 

Sylvilagus, 99 
auduboni arizonae, 100 
auduboni confinis, 100 
auduboni sanctidiegi, 100 
bachmani cerrosensis, 99 
bachmani cinerascens, 99 
bachmani exiguus, 99 
bachmani howelli, 99 
bachmani peninsularis, 100 
bachmani rosaphagus, 99 
mansuetus, 100 

Tadarida, 98 
brasiliensis mexicana, 98 
femorosacca, 98 
macrotis, 99 
mexicana, 98 
molossa, 99 

Talpidae, 92 

Tamias, 102 
asiaticus merriami, 102 
leucurus, 102 
leucurus peninsulae, 102 
obscurus, 102 

Tamiasciurus, 105 
douglasii mearnsi, 105 

Taxidea, 145 
americana neglecta, 145 
berlandieri, 145 
taxus berlandieri, 145 
taxus infusca, 146 
taxus neglecta, 145 

taxus, Taxidea, 145-146 

teliotis, Atalapha, 97 

tereticaudus, Citellus, 104 
Spermophilus, 104 

texanus, Peromyscus, 128 

texianus, Lepus, 101 

Thomomys, 105-110 
albatus, 105 
aphrastus, 106 
bottae abbotti, 107 
bottae affinis, 105 
bottae borjasensis, 108 
bottae cactophilus, 108 
bottae catavinensis, 108 
bottae cunicularius, 106 
bottae homorus, 109 


Acre 15: 


1963 | 


Huey: MAMMALS oF BajJA CALIFORNIA 


bottae imitabilis, 109 
bottae incomptus, 109 
bottae jojobae, 106 
bottae juarezensis, 106 
bottae litoris, 109 
bottae lucidus, 106 
bottae proximarinus, 106 
bottae rhizophagus, 108 
bottae ruricola, 108 
bottae russeolus, 109 
bottae sanctidiegi, 105 
bottae siccovallis, 107 
bottae xerophilus, 107 
fulvus alticolus, 110 
fulvus anitae, 110 
fulvus martirensis, 107 
fulvus nigricans, 105 
magdalenae, 109 


umbrinus 


abbotti, 107 


umbinus afhnis, 105 


umbrinus 
umbrinus 
umbrinus 
umbrinus 
umbrinus 
umbrinus 
umbrinus 
umbrinus 
umbrinus 
umbrinus 
umbrinus 
umbrinus 
umbrinus 
umbrinus 
umbrinus 
umbrinus 
umbrinus 
umbrinus 
umbrinus 
umbrinus 
umbrinus 
umbrinus 
umbrinus 
umbrinus 
umbrinus 
umbrinus 


albatus, 105 
alticolus, 110 
anitae, 110 
aphrastus, 106 
brazierhowelli, 107 
borjasensis, 108 
cactophilus, 108 
catavinensis, 108 
cunicularius, 106 
homorus, 109 
imitabilis, 109 
incomptus, 109 
jojobae, 106 
juarezensis, 106 
litoris, 109 
lucidus, 106 
magdalenae, 109 
martirensis, 107 
nigricans, 105 
proximarinus, 106 
rhizophagus, 108 
ruricola, 108 
russeolus, 109 
sanctidiegi, 105 
siccovallis, 107 
xerophilus, 107 


thysanodes, Myotis, 94 
torridus, Onychomys, 130-131 
townsendi, Arctocephalus, 149 
townsendii, Plecotus, 97 
truei, Peromyscus, 130 


Tursiops, 139 
gillii, 139 


nuuanu, 139 


umbrinus, Thomomys, 105-110 


Urocyon, 143 


cinereoargenteus californicus, 143 
cinereoargenteus peninsularis, 143 
cinereoargenteus scottii, 143 
virginianus scottii, 143 


Ursidae, 143 


ursinus, Callorhinus, 148 


Ursus, 143 
magister, 143 
velifer, Myotis, 94 


venusta, Neotoma, 131 


Vesperimus, 126 
fraterculus, 126 


Vespertilio, 94-95, 97-99 


californicus, 95 
cinereus, 97 
evotis, 94 


fuscus peninsulae, 97 


melanorhinus, 95 
molossus, 99 
pallidus, 98 
volans, 94 
yumanensis, 93 


Vespertilionidae, 93-98 


Vesperugo, 96 
merriami, 96 


virginiana, Didelphis, 91 
virginianus, Urocyon, 143 


vitulina, Phoca, 149 

vivesi, Myotis, 95 
Pizonyx, 95 

volans, Myotis, 94 
Vespertilio, 94 


vomerina, Phocoena, 140 


Vulpes, 142 
arsipus, 142 


macrotis arsipus, 142 
macrotis devia, 143 
macrotis devius, 143 
macrotis tenuirostris, 142 
yumanensis, Myotis, 93 


Vespertilio, 93 
Zalophus, 149 


californianus californianus, 149 


zibethicus, Ondatra, 136 


Ziphiidae, 138 
Ziphius, 138 


cavirostris, 138 


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ee Pe A eee 


LIBRARY 

-B 27 1964 
TRANSACTIONS PER et is 
HARVARD 

OF THE UNIVERSITY. 


SAIN DIEG@ SOCIETY OF, NATURAL HISTORY 


Vol. 13, No. 8, pp. 169-172 February 14, 1964 


THE LARVA AND PUPA OF AGATHY MUS DAWSONI 
(Lepidoptera: Megathymidae) 
BY 


JOHN ADAMS COMSTOCK 
Del Mar, California 


During the collecting of megathymids, September 1 to 7, 1962, to augment a series 
for describing an additional species (see Comstock, 1957), Charles F. Harbison 
obtained fifteen leaves of the Goldman Century Plant, Agave goldmaniana Trelease, 
which were infested with the Agave borer. These were collected at a point 17 miles 
north of Punta Prieta, Baja California. They were turned over to me for observation, 
and were placed individually in separate rearing cages, numbered consecutively 1 to 15. 


The leaves had been trimmed in the field to save weight and to facilitate transport 
over difficult terrain. Unfortunately, this trimming dried out and altered the character 
of the succulent pulp, resulting in the death of nine larvae by September 24, 1962. Five 
examples remained, however, and from these, drawings and notes were made in a short 
period of time. 


The species was described by Harbison (1963) as Agathymus dawsoni, named 
for Dr. E. Yale Dawson, now Director of the San Diego Natural History Museum. 


No observations on the habit of ovulation, or on the egg, were made in the field, 
but it was noted that the exit of the burrows in the leaves was always on the underside, 
and that the silken ‘window’ of exit was not spun until the larva had reached its final 
instar. Frass was not observed in the burrows, and its absence below the openings 
indicated that the larvae were sap feeders. 


In the case of larvae which had died in their earlier instars, the walls of the 
burrows were brown, with no indication of the flaky white powder which is so 
characteristic of the burrows made by Agathymus stephensi larvae (Comstock and 
Dammers, 1934). This waxy secretion is apparently formed only in the later period of 
the final instar, shortly before pupation, and in the case of A. dawsoni, it is scanty. 


The larval chambers vary in length from 24 to 50 mm., and in width from 12 to 
24 mm. The exit window is round in the majority of cases, but a few are oval. The 
maximum diameter of the round exits was 7.5 mm., and the minimum 3.5 mm. As 
some of the larvae were starved, the minimum measurements are probably abnormal. 


Emergence of the imagos occurred from September 24 to October 30, 1962. 


170 San Dieco Society or Narurat History [VoL. 13 


Fig. 1. Larva and pupa of Agathymus dawsoni. 


A. Pupa, lateral aspect, 314; B. Mature larva, dorsal aspect, X 314; C. Pupa, ventral 
aspect, 314. Reproduced from a water color drawing by J. A. Comstock. 


1964] Comstock: AGATHYMUS DAWSONI 171 


Mature larva (fig. 1B).—Length, 32 mm.; greatest width through 7th segment, 


9.5 mm.; head width, 3.6 mm. 


In healthy larvae the body surfaces are thinly obscured, at least in part, by the 
flaky white powder. This covering was brushed off before the following details were 
recorded: 


The head is light brown, well rounded, with the sutures almost indistinguishable. 
The mouth parts shade to black, and the ocelli are concolorous with the cheeks. The 
surface appears granular, due to a thick covering of short yellow-brown setae. 


The body ground color is whitish-green, with a darker shading on the cauda. Each 
segment has numerous transverse ridges. The first segment has a conspicuous black 
cervical shield, slightly wider middorsally, but divided in the center by a narrow light 
stripe (fig. 2A). There is a middorsal longitudinal dark green stripe, absent on the 
thoracic and caudal segments, and narrowing at each segmental juncture. 


The legs are dull yellow with brown tips, and the prolegs are concolorous with the 
body. The crochets are light brown, arranged in a complete oval, with those arising 
from the inner margin arching outwardly. They are placed in triordinal rows (fig. 2B). 


The spiracles are black-rimmed, with centers slightly darker than the body ground 
color, but usually appearing white because of the whitish flakes covering them. 


The entire body surface is thickly covered with very short yellow-brown setae, and 
a few slightly longer ones, which arise from small yellow papillae. There is some 
variation in larval coloration and intensity of markings. 


Pupa (fig. 1A and C).—Length, 29 mm.; greatest width through 5th abdominal 
segment, 7.5 mm.; head width, 4 mm., comparatively small in relation to body width. 
The eyes are red-brown, widely separated, and protruding. 


The body ground color is yellow-brown, the wing veins yellow, and the cremaster 
shades to brownish-black at the tip. 


The antennae terminate two-thirds the distance toward the wing margins, and 
the maxillae extend slightly beyond that point. The spiracles are concolorous with the 


body. 


The cauda ends in a spatulate cremaster which inclines ventrally. Figure 2, C and 
D, illustrates both this and the caudal segments in ventral and dorsal aspects. 


LITERATURE CITED 


Comstock, John A. 
1957. Notes on the metamorphosis of an Agave-boring butterfly from Baja 
California, Mexico. Trans. San Diego Soc. Nat. Hist. 12:263-276. 


Comstock, John A., and Charles M. Dammers 
1934. The metamorphoses of three California diurnals. Bull. So. Calif. Acad. 
Sci. 33:79-92. 


Harbison, Charles F. ay Af 
1963. A second new species of megathymid from Baja California, Mexico 
(Lepidoptera: Megathymidae). Trans. San Diego Soc. Nat. Hist. 15:61-72. 


172 San Dieco Society of Naturat History [VoL. 13 


yyy Nh yD 


ear ees 


Fig. 2. Agathymus dawsoni. 


A. Cervical shield of larva; B. Crochets on typical proleg of larva; C and D. Caudal 
segments of pupa, including cremaster, in ventral and dorsal aspects, respectively. All figures 
highly magnified. Reproduced from a water color drawing by J. A. Comstock. 


MUS. COMP. ZOOL’ 


LIBRARY 
FEB 27 1964 
TRANSACTIO 
; NS HARVARD 
OF THE UNIVERSITY. 


SAN DIEGO SOCIETY OF NATURAL HISTORY 


VoLuME 13, No. 9, pp. 173-176 


A NEW MARINE BEETLE FROM THE GULF OF CALIFORNIA 
(Coleoptera: Staphylinidae) 


BY 
IAN MoorRE 


Research Associate in Entomology 
San Diego Natural History Museum 


SAN DIEGO, CALIFORNIA 
PRINTED FOR THE SOCIETY 


FEBRUARY 14, 1964 


" 


Fig. 1. Cameronium sonorensis, new species. 


FEB 27 (304 


HARVARD 
A NEW MARINE BEETLE FROM THE GULF OF CADFORNIXY 


(Coleoptera: Staphylinidae) 


BY 
IAN MOORE 


Species of the genus Cameronium have hitherto been known only from the shores 
of the warm arid regions of North Africa and the Red Sea and from Zanzibar. The 
discovery of a new species in cracks in the rocks in the intertidal zone of the desert 
coast of Sonora, Mexico, discloses a very interesting example of insect distribution. 
We can anticipate the discovery of other species of this genus in similar situations in 
other parts of the world. 

Members of this genus differ from other marine bolitocharinids by the combina- 
tion of their unarmed tibiae and bifid ligulae. A more detailed discussion of the genus 


may be found in Moore (1956). 


Cameronium sonorensis, new species 


Holotype. — From cracks in the rocks in the intertidal zone at Punta de los 
Cuervos, San Carlos Bay, Sonora, Mexico; collected by Ian Moore, November 7, 1962; 
in the California Academy of Sciences. 


Paratypes. — Three specimens, same data as holotype, in the California Academy 
of Sciences, the San Diego Natural History Museum, and the collection of lan Moore. 


Description of holotype. — Form: Small, elongate, subparallel (fig. 1). Color: 
Head piceous, becoming paler anteriorly, pronotum rufo-testaceous, mesosternum 
piceous, elytra rufo-testaceous, with the scutellar region clouded, abdomen piceous, the 
tergites paler apically, antennae with the first two segments pale, the others progres- 
sively darker, legs and mouth parts testaceous. 


Head: Orbicular; surface evenly convex at the sides, shallowly impressed centrally 
between the eyes, densely reticulate, finely and rather densely punctured throughout. 
Eyes at about their length from base of head. Antennae: First two segments more than 
twice as long as wide, third a little narrower and a little shorter than second, fourth 
through sixth about as wide as second, not longer than wide, seventh through tenth 
distinctly transverse, eleventh longer than wide, gradually narrowed to apex. 


Pronotum: About as wide as head, transverse, widest near apical third; sides 
arcuate in apical half, sinuate before the hind angles; apical margin straight; base 
arcuate. Surface convex except for a broad, shallow central impression which is deepest 
near the base. Integument sculptured very much like the head. Elytra: Distinctly 
wider and a little longer than the pronotum. Sides gently arcuate, apices straight. 
Surface sculpture similar to that of pronotum. 


Abdomen: The first three segments nearly parallel, thence gradually tapered to 
apex. Surface with a sparse network of fine lines; finely, sparsely, somewhat asperately 
punctured; interspaces very shining. Length: 2 mm. Sex: unknown. 


The paratypes differ in no noticeable way from the holotype. 


This species is very similar in structure to C. obockianum but has the head black, 
whereas it is pale in the latter. 


176 San Dieco Society oF NATURAL History [VoL. 13 


KEY TO THE SPECIES OF Cameronium 


lirigg TRIER VG) TORS. Geese cet seeeere Ar cber hats nar oaant oconeak backs uecheboososonccc subd liebmanni, obockianum 
lb. Head dark 

Dae Ron Obi LACK: -c<.<.05 desk cos esdadetds omen on teee reer one eee flavipenne 

Phe “Pronotumi Pale: ...62..c0<.-cosnteeceess00edee nedonsnessespesesuc wosccsed sss eioees sonorensis 


BIBLIOGRAPHY OF THE GENUS Cameronium 


Cameron, Malcolm 
1944. New oriental Staphylinidae (Col.). Ann. Mag. Nat. Hist., ser. 11, 
11:312-322. 
Fauvel, Albert 
1905. Staphylinides exotique nouveaux, pt. 3, Revue d’Ent. 24:113-147. 


Koch, Carlos 


1936. Wissenshaftliche Ergebnisse der entomologischen Expeditionen Seiner 
Durchlaucht des Fursten Alessandro C. Dello Torre e Tasso nach Aegypten 
und auf die Halbinsel Sinae, XIII: Staphylinidae. Publ. Mus. Ent. 
Pietro Rossi 1:115-232. 


Moore, lan 
1956. A revision of the Pacific coast Phytosi with a review of the foreign 
genera (Coleoptera: Staphylinidae). Trans. San Diego Soc. Nat. Hist. 
12:103-151. 
Scheerpeltz, Otto 


1957. Ein neue Art der Gattung Cameronium (Col. Staphylinidae) (75. Beitrag 
zur Kenntnis der Palaarctischen Staphyliniden). Ent. Blatter 53:147-150. 


“LIBRARY 
FEB 27 1964 
TRANSACTIONS HARVARD 


UNIVERSITY. 
OF THE 


SAN DIEGO SOGIELY OF NATURAL HISTORY 


VoLuME 13, No. 10, pp. 177-184 


BIRDS AND MAMMALS OF THE VOYAGE OF THE “GRINGA”’ 


BY 


RICHARD C. BANKS 


Curator of Birds and Mammals 


San Diego Natural History Museum 


SAN DIEGO, CALIFORNIA 
PRINTED FOR THE SOCIETY 


FEBRUARY 14, 1964 


Publication of this paper was sponsored by 
The American Foundation for Oceanography. 


FEB 27 1964 


HARVARD 


UNIVERSITY 
BIRDS AND MAMMALS OF THE VOYAGE OF THE “GRINGA” 


BY 


RICHARD C. BANKS 


Biologists have been making sporadic visits to the islands of Baja California for 
approximately 70 years. As a result of this activity the taxonomy and distribution of 
the vertebrate fauna has become fairly well known. However, little has been recorded 
about other aspects of the biology of these insular populations. This paper records 
some additional data obtained on a voyage south along the Pacific coast of Baja 
California and into the southern part of the Gulf of California during April and May. 
1963. It is not a complete report of all species observed or collected, but consists mainly 
of selected information, from field notes and from specimens, bearing particularly on 
the abundance and reproductive biology of the animals concerned. 


The “Gringa” (fig. 1) is a 65-foot yacht of Norwegian design and construction, 
powered by twin diesel engines. Ample space and comfortable quarters make it ideal 
either for pleasure cruising or for scientific field work. The crew of the “Gringa” 
included Mr. and Mrs. Richard F. Dwyer, owners of the vessel, Harton B. Smith, 
engineer, and Jaime Eulogio Pacheco, steward. Mr. John Emmel and Mr. and Mrs. 
Thomas Crawford were guests of the Dwyers for portions of the trip. The scientific 
party was composed of three staff members of the San Diego Natural History Museum 
— Reid Moran, Curator of Botany, Boyd Seavey, Assistant in Birds and Mammals, 
and the author. 


Moran and Seavey join me in expressing heartfelt appreciation to Mr. and Mrs. 
Dwyer for the opportunity to make this trip. Their accommodation to the needs of the 
biologists was both gracious and generous. To the other companions of the voyage. 
as well, go our thanks for their interest and assistance. 


Both Moran and Seavey have read the manuscript and have offered comments on 
the material presented. Dr. Richard G. Van Gelder provided information concerning 
his visit to Cedros Island in 1957. Comparisons of several specimens were made in the 
Museum of Vertebrate Zoology. 


ITINERARY 


Departed San Diego in the late afternoon of April 9, 1963. 


Isla San Martin. — Anchored near Hassler’s Cove in mid-morning, April 10; 
departed after picking up traps, early April 12. 


Isla San Ger6nimo. — Arrived early afternoon, April 12, and spent the remainder 
of the day ashore: left early April 13, after picking up traps. 

Isla Cedros. — Anchored at the village, on the southeast part of the island, in the 
evening, April 13; worked in the vicinity of the village. April 14 and 15; moved north 
along the east side of the island, April 16, and worked two arroyos, April 16 and 17: 
left temporarily, morning of April 18. 


Islas San Benito. — Anchored among the islands near noon, April 18, spending 
that afternoon on West San Benito Island; visited Middle and East San Benito Islands 
briefly, April 19, leaving after noon. 


180 San Dieco Society oF NATURAL HiIsToRY (Wor<l3 


Fig. 1. Mr. and Mrs. Dwyer aboard the “Gringa” at Mulege. 


Isla Cedros. — Anchored at South Bay in mid-afternoon, April 19; remained until 
early morning, April 21. 


Isla Natividad. — Arrived in mid-morning, April 21, working ashore until dusk; 
departed about midnight. 
Isla Magdalena. —- Landed on the west side of the island, 2.5 miles north of 


Punta Hughes, on morning of April 23; proceeded after two hours to Puerto 
Magdalena, in Magdalena Bay, arriving shortly after noon; worked on this island until 
departure, morning of April 25. 


Isla Santa Margarita. — Reached Puerto Cortés early in afternoon, April 25; 
departed in mid-morning, April 20. 

La Paz. — Arrived at mid-day, April 29, after stops at Los Frailes and Bahia de 
los Muertos; Moran and Seavey returned to San Diego, April 30; proceeded to 
Pichilinque Harbor, May 4. 

Isla Partida (Espiritu Santo). — Anchored in Bahia Cardenal about noon, May 
5; departed in morning, May 6. 

Isla San José. — Ashore on the south end of the island for three hours in 
afternoon, May 6; left anchorage in Bahia Amortajada in morning, May 7. 

Puerto Escondido. — Arrived in late afternoon, May 7, spending some time 
ashore: remained here until afternoon, May 10. 

Isla Danzante. — Visited by small boat from Puerto Escondido in afternoon, 
May 8, and morning, May 9, remaining approximately two hours each time. 

Isla Carmen. — Visited the south end of this island by small boat from Puerto 
Escondido in afternoon, May 9, and in morning, May 10. 


Isla Monserrate. — Arrived in late afternoon. May 10; left in mid-afternoon, 
May 11, after spending the morning ashore. 


1964] BANKS: VOYAGE OF THE “GRINGA” 181 


Isla Carmen. — Arrived at Bahia Ballandra late, May 12; left about noon, May 
13: 
Isla Coronado. — Anchored in mid-afternoon, May 13; left in mid-afternoon. 
May 14. 
Mulegé. — Arrived about noon, May 15; departed for San Diego, May 16. 
BIRDS 
Diomedea nigrepes. Black-footed Albatross. — One was seen off Abreojos Point 


(approximately 26° 42’ N) on the morning of April 22. 


Puffinus puffinus opisthomelas. Manx Shearwater. —- Thousands of shearwaters 
were seen south of Cedros Island on April 21, where hey were feeding in a large, 
loose flock. Nesting burrows were abundant on the flatter parts of Natividad Island. 
and most were occupied by this species. Each burrow investigated contained a single 
white egg. Skeletal remains of shearwaters were much in evidence on the island, 
probably reflecting the abundance of semi-feral house cats. 


Pandion haliaetus. Osprey. — This species was observed on most of the islands 
visited. Active nests were found on San Martin, Cedros, Natividad, Santa Margarita, 
and all three of the San Benito islands. 


Lophortyx californicus. California Quail. — Quail were heard or seen in two 
places at Ballandra Bay, Carmen Island, on May 13. The female obtained was ready 
to lay. The subspecific status of this bird is uncertain. This is the first report of quail 
on the islands of Baja California. 


Larus spp. Gulls. — Adult Western Gulls, L. occidentalis, were abundant on the 
Pacific coastal islands south to Natividad. The birds seemed to have selected nest 
sites, although nesting had not yet begun. The largest concentration was on Natividad 
Island, where we estimated conservatively 5000 birds on the low, flat parts of the island. 


In Magdalena Bay a mixed flock of well over 5000 gulls, mostly immature, was 
feeding on pelagic red crabs which were teeming in the bay. Species represented. in 
decreasing order of abundance, were the California Gull, L. californicus, Ring-billed 
Gull. L. delawarensis, Heermann Gull, L. heermanni and Bonaparte Gull, L. philadel- 


phia. Immature Western Gulls, also present in the bay, stayed apart from the flock. 


Ptychoramphus aleutica. Cassin Auklet. — The flat, sandy southern end of San 
Geronimo Island was honeycombed with the nesting burrows of this auklet. Some ne wly 
hatched young were found, as were also nearly fresh eggs. A burrow on West San 
Benito Island held an adult and a nearly full grown young bird. Van Rossem (1939: 
443) noted that the birds on the San Benito Islands (P. a. australe) nest some two 
months earlier than the species does farther to the north (P. a. aleutica). My specimens 
from San Geronimo Island are intermediate in measurements between these two 
races (cf. van Rossem, loc. cit.). 

Colaptes sp. Flicker. — A partial skeleton of a flicker was found on San Martin 
Island by Seavey on April 10. Although specific allotment of this specimen is not 
possible, size suggests that it is C. cafer “rather than C. chrysoides. Records ue flickers 
on the islands a northwestern Baja California are very few (Grinnell, 1928). There 
is no habitat suitable for the species on San Martin Island. 


Thryomanes bewickii cerroensis. Bewick Wren. — This was one of the more 
abundant species on Cedros Island. One bird which flushed, as though from a nest. 
was carrying food in its bill. Males taken had testes 5 to 7 mm. long. 

Salpinctes obsoletus. Rock Wren. — Several family groups of S. 0. tenuirostris 
were seen on West San Benito Island; the young were fully grown and nearly indepen- 
dent. Young of S. 0. obsoletus on Cedros Island were at the same stage of development. 
Rock Wrens seem to me to be less abundant on the Pacific coastal islands of Baja 
California than on the islands in the Gulf of California. 


182 San Dieco Society oF NATURAL HISTORY [VoL. 13 


Polioptila melanura margaritae. Black-tailed Gnatcatcher. — Males in molt were 
taken on Santa Margarita and Partida (Espiritu Santo) islands. The Blue-gray 
Gnatcatcher, P. caerulea, has previously been recorded on Espiritu Santo Island 
(Banks, 1963a), and there is a specimen in the Museum of Vertebrate Zoology from 
that locality. 


Icterus spurius. Orchard Oriole. — Seavey found the mummy of an Orchard 
Oriole on West San Benito Island. The plumage, although badly abraded, indicates 
that the bird was a female. There is no previous record of the Orchard Oriole for Baja 
California. 


Carpodacus mcgregort. McGregor House Finch. — No house finches were seen 
on the San Benito Islands despite a thorough search (least extensive on the east island). 
The extinction of this form has been forecast since early in the century (Grinnell, 1928, 
and references therein). The last report of which | am aware is by Bancroft (1932:89), 
who estimated the population to be 25 individuals. It seems unlikely that the number 
is currently that high, and the species may now be extinct. 


Passerculus sandwichensis sanctorum. Savannah Sparrow. — This was by far the 
most abundant land bird on all three of the San Benito Islands. Young were out of the 
nest and nearly full grown at the time of our visit. 


Amphispiza belli belli. Bell Sparrow. — This was the only common land bird on 
San Martin Island. Breeding had begun, but no young were seen. 


Amphispiza bilineata ssp. Black-throated Sparrow. — This species was one of the 
most common on Cedros Island. Many pairs were accompanied by young birds; feather 
growth had been completed in the two juveniles taken. Reproduction appeared to be at 
the same stage on Natividad Island. These two populations are referred to A. b. 
deserticola. 


Black-throated Sparrows of the race A. b. bangsi were common on Magdalena 
Island, and were seen in pairs or in small flocks. Two males taken from pairs had 
completely ossified skulls and testes 6 and 8 mm. in length. A male taken from a flock 
had an incompletely ossified skull and testes only 3 mm. long. A female from a flock 
also had the skull incompletely ossified, and was molting dorsally. Both the immatures 
(as determined by the skulls) were in the adult plumage. 


Data from Cerralvo Island, Baja California (Banks, 1963b), indicate that young 
Black-throated Sparrows join into flocks at the end of the breeding season, at about 
the time of their post-juvenal molt, and remain in flocks until the next breeding 
season. Adults remain in pairs during this time. On Cerralvo Island, adults were not 
found to be in breeding condition when young birds were in flocks. Testis size of the 
adults taken on Magdalena Island in April, 1963, was equivalent to that of nesting 
birds taken on Cerralvo Island; this suggests that breeding was in progress or was 
about to begin. The fact that young birds in! adult plumage were in flocks, however, 
suggests that breeding had occurred several months previously, perhaps in the fall. 
None of these data suggests a February breeding season, as stated for Magdalena and 
Santa Margarita islands by van Rossem (1945). 


Black-throated Sparrows of the race A. b. bangsi were in breeding condition on 
Partida (Espiritu Santo), Monserrate, and Danzante islands, in early May. 


Representatives of A. b. carmenae were also breeding in early May; the season 
had not advanced far enough for young to be out of the nest. Evidence from both 
1962 (Banks, 1963a) and 1963 indicate that the March date given by van Rossem 
(1945) for the breeding season of Black-throated Sparrows on ‘Carmen Island is too 
early. 


1964] BaNnKs: VOYAGE OF THE “GRINGA” 18: 


hy 
ww 


MAMMALS 


Pipistrellus hesperus. — Two specimens obtained on Monserrate Island are 
referrable to P. A. australis on geographic grounds (Hatfield, 1936). There seem to be 
no previous records for bats on this island. 


Sylvilagus bachmani cerrosensis. — None was seen during six days at four 
different areas on Cedros Island. Old droppings, possibly from a rabbit, were seen, 
but no definite sign was found. Nelson (1909) knew of only four specimens, and | 
have been unable to find a reference to any taken since that study. This rabbit perhaps 
was never very common, and its present status is unknown. It certainly is not as 
abundant as other insular rabbits in Baja California. 


Perognathus baileyi fornicatus. — A single specimen was obtained on Monserrate 
Island; it was a female with three 9 mm. embryos. 


Perognathus spinatus. — Representatives of six subspecies of this pocket mouse 
were obtained, from six islands. The only certain evidence of breeding was for P. s 
seorsus on Danzante Island, where a female held three 8 mm. embryos. 


Peromyscus pseudocrinitus. — A single gray-pelaged male, with testes 8 mm. long, 
was taken on Coronado Island. 


Peromyscus eremicus. — Eleven specimens of P. e. cedrosensis were taken on 
Cedros Island, where the population of rodents seemed to be low. Six males had testes 
8 to 14 mm. long. The single pregnant female obtained held five 22 mm. embryos. 
Four females had no embryos; three of these were lactating. 


A single P. e. pullus was obtained on Santa Margarita Island; it was a male with 
testes 8 mm. in length. 


A male P. e. carmeni from the south end of Carmen Island had testes 3 mm. long. 
A female of this race had no embryos. 


Peromyscus maniculatus. — White-footed mice of the race P. m. exiguus were 
abundant on San Martin Island; the trap yield was nearly 50 per cent. Most of the 
series of 37 mice obtained were caught on the heavily vegetated lava slopes, but one 
individual was taken in low growth on sand dunes near the beach. Approximately 
one-third of the mice had parasites imbedded at the base of one or both ears. These 
have been identified as chigger fleas, Tunga caecata, by Dr. Frank J. Radovsky. Eight 
of 11 females were pregnant, embryos ranging in number from three to six and in 
length from 6 to 18 mm. Two females were lactating and one had no embryos. Testes 
of adult males were 9 to 12 mm. in Jength; juvenile and sub-adult males had testes 5 
to 6 mm, long. 


On San Geronimo Island, mice were even more abundant than on San Martin; 
the trap yield of P. m. geronomoensis was over 60 per cent. Many were taken in and 
around the nearly deserted village, but the population in the low, sparse vegetation of 
the sand flats seemed to be equally dense. One female had five 15 mm. embry os; Ll 
were not pregnant and none was recorded as lactating. The single juvenile male taken 
had testes 3 mm. long; testis length of adult males ranged from 5 to 10 mm. The 
average testis length of adult males on San Geronimo Island was 3.5 mm. less than 
that of adults on San Martin Island. 


Neotoma lepida. —- Wood rats, N. 1. pretiosa, were extremely abundant on Mag- 
dalena Island. Several elevated houses were found in a mangrove swamp, and drop- 
pings were seen commonly on fallen or prostrate tree trunks. An immature male had 
testes 7 mm. in length; testes of adult males measured 14 to 18 mm. None of the 
females taken was pregnant. 


164, San Dieco Society or NATuRAL History [VoL. 13 


Houses of NV. l. latirostra were fairly common on Danzante Island. None of the 
four females obtained was pregnant; two males had testes 15 and 17 mm. long. A 
male taken April 2, 1962, had testes measuring 17 mm. Only a single specimen of this 
subspecies has been reported previously (Burt, 1932). 


Neotoma bryanti. — Sign of this wood-rat was scarce on Cedros Island. Only 
two females were obtained, both at South Bay. Neither was pregnant. 


LITERATURE CITED 


Bancroft. G. 
1932. Lower California: A Cruise. The Flight of the Least Petrel. G. P. 
Putnam’s Sons, New York. 403 pp. 


Banks, R. C. 
1963a. Birds of the Belvedere Expedition to the Gulf of California. Trans. San 
Diego Soc. Nat. Hist. 13:49-60. 
1963b. The birds of Cerralvo Island, Baja California, Mexico. Condor 65:300-312. 
Burt, W. H. 


1932. Descriptions of heretofore unknown mammals from islands in the Gulf of 


California, Mexico. Trans. San Diego Soc. Nat. Hist. 7:161-182. 
Grinnell, J. 


1928. Distributional summation of the ornithology of Lower California. Univ. 


Calif; Publ) Zool. 3221-300: 
Hatfield, D. M. 


1936. A revision of the Pipistrellus hesperus group of bats. Jour. Mamm. 


17:257-262. 


Nelson, E. W. 
1909. The rabbits of North America. North American Fauna 29. 314 pp. 


van Rossem, A. J. 
1939. Some new races of birds from Mexico. Ann. Mag. Nat. Hist., ser. 11, 
4.:439-443, 
1945. Preliminary studies on the black-throated sparrows of Baja California, 
Mexico. Trans. San Diego Soc. Nat. Hist. 10:237-244. 


TRANSACTIONS 
OF THE 


SAN DIEGO SOCIETY OF NATURAL HISTORY 
Pos 


YA 
Jf 


VotuME 13, No. 11, pp. 185-268 


EVOLUTION OF THE PIT VIPERS 


BY 


BayYARD H. BRaTTsSTROM 


California State College at Fullerton 
Fullerton, California 


SAN DIEGO, CALIFORNIA 
PRINTED FOR THE SOCIETY 
May 4, 1964 


TRANSACTIONS 
OF THE 
SAN DIEGO SOCIETY OF NATURAL HISTORY 


VoLuME 13, No. 11, pp. 185-268 


EVOLUTION OF THE PIT VIPERS 


BY 


BayarD H. BrRatTTsTROM 


California State College at Fullerton 
Fullerton, California 


SAN DIEGO, CALIFORNIA 
PRINTED FOR THE SOCIETY 


May 4, 1964 


23 2 3 21 10 413 6 20 Il 14 
‘ tom 1 ' ‘inet ’ ' ’ 
: ‘ H 7 org fe 


: Pa, a? ee 


Fig. 1. Skull of Crotalus ruber ruber. From Klauber, 1956. 


Key to bones: 1, Premaxilla; 2, Prefrontal; 3, Frontal; 4, Parietal; 5, Basisphenoid; 6, Squamosal; 
7, Maxilla; 8, Palatine; 9, Pterygoid; 10, Ectopterygoid; 11, Quadrate; 12, Mandible (or lower jaw); 
12A, Dentary; 12B, Articular; 13, Pro-otic; 14, Exoccipital; 15, Poison fang; 16, Mandibular teeth; 
17, Pterygoid teeth; 18, Palatine teeth; 19, Supraoccipital; 20, Stapes; 21, Postfrontal; 22, Basioccipital; 
23, Nasal; 24, Turbinal; 25, Vomer. 


EVOLUTION OF THE PIT VIPERS 


BY 
Bayarp H. BratTtrsTROM 


CONTENTS 

Introduction Feed accede Ae et 187 
Materials and Methods .............. 632 SEP My Optom IY 188 
Acknowledgments aR Rae SRS er St ABC ERE ea eR Ne 188 
Comparative Ostelogy of the Crotalids ao va) Sn ee ee bee ea RO 
General Osteological Observations eee See eee RAS LOO. 
Anomalies, diseases, and injuries Sestrih ath sd 190 
Ontogenetic variation . See Siero eect en ee Es 190 
Geographic variation sie Rtas tetany Scand ee ee ee 192 
Descriptive Osteology ee ee ee ee ee eos 194 
Teeth. ...... sre Ray hve alle Bem Hereetniyy bd ce aan amie boom WP tel DD 

Bonesee te at. see, ee eee ee ee Se een See a Oe ee We 195 
Diylopenyar emt te ea eee eer ee oe ee i econ ae ee DAZ 
| (Cd aoe a= aca de ee aoc ie dei th 2 aM iy, coolly! nk dates Wie 219 
Iintercenenceelationships : 2 ¢.d..cs6; gots janet esate ee eek eee. 
Phylogeny of A gkistrodon 5 WORE TARE pc eT EMER 25 ote MUTE, 2a Oe eee 2d!) 
Phylogeny of Crotalus and Sistrurts ......0..cc.0cc00cccscseeeeeeees f 238 
Phylogeny of Bothrops ..... peas! A) 5 fe 3 Me Mees Le AN A ee 246 
Phylogetyat linimeresuras 5.0 ce 247 
Rasommomniygrc ee xsi tn. eRe ok ae Sie A ee A eet 248 
Zoogeography ......... PS RAIS EERE et So oc Rete Paste ote Mee Ree eke 251 
Zoogeography of the Crotalidae Eta SR Maia: Po Ma Roce! 255 
Summary ...... ee ee Ap caghs..:cfecacest todte cm sane ae are 261 
Literature Cited ........ 2A ee le tees aot 262 

INTRODUCTION 


The skeleton is one of the most important structures available for the analysis of relation- 
ships of vertebrates. This paper assesses relationships within one group of vertebrates, the 
Crotalidae, by a study of their comparative osteology. 

The Crotalidae, or pit vipers, of America and Eurasia, is a family of poisonous snakes 
with movable fangs. The Crotalidae shows remarkable parallel evolution with the Old World 
family of true vipers, the Viperidae. The Crotalidae differs from the Viperidae in many 
characters, the most conspicuous of which is the presence of heat-sensitive facial pits. 

The Crotalidae, as currently defined, includes six genera: Crotalus (rattlesnakes; North, 
Central, and South America), Sistrurus (pigmy rattlesnakes; North America), Lachesis (bush- 
master; Central and northern South America, Trinidad), Bothrops (fer-de-lance, hog-nosed pit 
vipers, palm pit vipers; Mexico, Central and South America), Trimeresurus (Asiatic pit 
vipers; southeast Asia), and A gkistrodon (copperheads, moccasins; North America, Asia, and 
extreme southeastern Europe). 


188 SAN Dreco Society oF NaturaAL History {Vor 13 


TABLE 1 
Summary of material examined. 
Genus Species in genus Species examined Skeletons Examined 

A gkistrodon REE I es. 14 91 50 
BCE DTOP See ree 30? 13 40 
(Cra G1 tere Poe ee een eae eRe 24 232 281 
LACE SIS Maes oN ae: 1 1 yi 
WSiishrtirtica@e. ne are te st 3 3 2 
Wir inter ESUTUS hv cisstes TE es 33 8 15 

SIO Eal Simeteertees eee 105 7 414 


Not examined: A. annamensis, himalayanus, millardi, monticola, nepa. 
2Not examined: C. transversus. 


MATERIALS AND METHODS 


In the course of this study, skeletons of crotalids were borrowed from most of the major 
museums in the United States. A report on the fossil material has been given elsewhere 
(Brattstrom, 1954). The recent material examined is listed in tables 1 to 3. Skeletons of 
several species of the Viperidae were also examined. Through the courtesy of curators at several 
museums, I was allowed to remove the skulls of rare crotalids from preserved specimens. This 
can be done without damage to any of the scales, and the skull can be replaced with cotton. 
References to individual specimens use standard abbreviations for names of institutions. 


Measurements were made to the nearest tenth of a millimeter with a vernier caliper. 
Paired bones were measured consistently on the more convenient side (e.g., the frontal and 
ectopterygoid on the right side, the pterygoid on the left) except when one member of a pair 
was absent. No significant difference was noted in measurements of paired bones. Klauber 


(1939) found the same with regard to fangs. 


The skeletons examined had been prepared in many ways. Bones that had remained too 
long in bleach had rough edges; these bones were not measured. Specimens that had been in 
formaldehyde for many years often had soft bones, and these also were not measured. Except 
for these two easily detected conditions, the skeletons examined showed no differences resulting 
from method of preparation. 


Drawings were made of most bones of each skeleton examined. The mode or the most 
typical example of each skull bone of each species is illustrated herein. Measurements of length 
and width were graphed as were ratios of measurements of one bone to measurements of an- 
other (e.g., quadrate length to pterygoid length). Graphs were made for all bones measured 
and for almost all possible combinations of them. Only a few of these graphs are reproduced 
here, though most of them are discussed. 


The terms and names used are those of recent authors on each group or geographic area: 
Cochran, 1943; Gloyd, 1940; Gloyd and Conant, 1943; Klauber, 1936a, 1949, 1952, 1956; 
Maslin, 1942; Pope, 1935; Pope and Pope, 1933; Rooij, 1917; Schmidt, 1953; M. A. 
Smith, 1931. 


ACKNOWLEDGMENTS 


The following people and institutions have my thanks for the loan of recent and fossil 
osteological material: Mr. Charles M. Bogert, American Museum of Natural History; Dr. 
John C. Brown, Los Gatos, California; Dr. Charles L. Camp, Museum of Paleontology, 
University of California, Berkeley; Dr. Doris M. Cochran, Department of Herpetology, 


1964 } BRATTSTROM: EVOLUTION OF Pit VIPERS 189 


TABLE 2 

Specimens examined: Agkistrodon, Bothrops, Lachesis, Sistrurus, and Trimeresurus. 
Animal Number Animal Number Animal Number 
A gkistrodon EPS!) Bothrops 40 Sistrurus 21 
acutus .... . 1 alternata 2 catenatus ssp. 6 
bilineatus 5 atrox . 18 catenatus 4 
blomhoffi_. 2 bilineatus 1 tergeminus I 
contortrix ssp. 8 cotiara 1 miliarius =P 4 
Ae g streckeri | 

laticinctis 1 dunni 1 

: : ravus a) 
contortrix 1 godmani 2 Te opiate 15 
mokeson e...  3 jararaca .. 4 Plo labyic 1 
12:71 GS Ni cee me aes eRe 2 Jararacussu 4 flaviviridis 2 
hypnale .. i lancealatus, 7. ..-. 1 gramineus 1 
DISCIVOTUS SSDig.ct ac (O nasuta 1 mucrosquamatus 1 
leucostoma ............ 9 neuwiedi 2 puniceus 1 
piscivorus ne NITE nummifer .. 1 purpureomaculatu 72 
Gut Sean ee or Renee | schlegeli 1 stejnegert 1 
rhodostoma .... .... 1 Lachesis muta .. 7 wagleri 6 


United States National Museum; Dr. Edwin H. Colbert, American Museum of Natural His- 
tory; Dr. D. Dwight Davis, Chicago Natural History Museum; Mr. William E. Duellman, 
Museum of Zoology, University of Michigan; Dr. D. H. Dunkle, Department of Paleon- 
tology, United States National Museum; Dr. Herman Gunter, Florida Geological Survey; 
Dr. Claude W. Hibbard, Museum of Paleontology, University of Michigan; Dr. Hildegarde 
Howard, Los Angeles County Museum; Dr. Robert F. Inger, Chicago Natural History 
Museum; Dr. L. M. Klauber, Zoological Society of San Diego and San Diego Society of 
Natural History; Dr. Arthur Loveridge, Museum of Comparative Zoology, Harvard Univer- 
sity; Dr. James E. Moismann, Museum of Zoology, University of Michigan; Dr. Wilfred T. 
Neill, Ross Allen Reptile Institute; Mr. William Otto, then of the Department of Geology, 
California Institute of Technology; the late Dr. Karl P. Schmidt, Chicago Natural History 
Museum; the late Mr. Joseph R. Slevin, California Academy of Sciences; Dr. Hobart M. 
Smith, Department of Zoology, University of Illinois; Dr. Robert C. Stebbins, Museum of 
Vertebrate Zoology, University of California, Berkeley; Dr. Ernest Williams, Museum of 
Comparative Zoology, Harvard University. 

In addition, I wish to thank the many people who have given me specimens or skeletons, 
especially Mr. Charles E. Shaw and the late C. B. Perkins of the San Diego Zoo. 

The work was carried out under the guidance of Dr. Raymond B. Cowles. 

Various aspects of the paper have been discussed in person or through correspondence 
with many people. I would especially like to thank the following: Mr. Charles M. Bogert, 
Mr. W. Leslie Burger, Dr. Walter Auffenberg, Dr. Max Hecht, Dr. Daniel I. Axelrod, Dr. 
George A. Bartholomew, Dr. Raymond B. Cowles, Dr. Carl Epling, the late Dr. Frank Pea- 
body, Dr. Jay M. Savage, Dr. Howard K. Gloyd, and Dr. Laurence M. Klauber. 


COMPARATIVE OSTEOLOGY OF THE CROTALIDAE 


GENERAL OSTEOLOGICAL OBSERVATIONS 
The osteology of various crotalids has been described by Tyson (1683), Owen (1853), 
Giebel (1866), Hoffman (1890), W. E. Taylor (1895), Boulenger (1896), Gadow (1920), 
Phisalix (1922), Klauber (1936a, 1939, 1956), Radovanovic (1937), M. A. Smith (1943), 
and Ruiz (1951). Most of these descriptions are concerned primarily with the fang mechanism, 
poison apparatus, or family characterization. Skulls of crotalids were figured by Ditmars, 1931 


190 SAN Disco Society oF Natrurav History [Vot. 13 


TABLE 3 
Specimens of Crotalus examined. 
Animal Number Animal Number Animal Number 
adamanteus  ................ yen GUICAUGAINS one) pusilla ee eee 5 
BET OK RR Ps ei 38 INLEFMEAIUS ......:...2000000..0 1 TUDET ee ee ee 20 
DaASHISCUS TR ee ee nee 5 lepiaastsspse rece re van 2 SCULUIGLUS. ee ee 8 
COVASEES SSDS ooh n ceed. 53h 6 lepidus® eve ccG. 2 stejne gente his eth: 2 
COVdslesnees ont ee 6 klauberi .... Westie 3 (OTIS: pone Gein A 2 
laterorepens® ...2..;...... 17 mitchelli LOTEUGENSIS: “aie eee al 
durissus Witcbellia eee hye 2 EviSeVidttUSa-. 0 eee 
AUTISS USHERS hone ceca. 4 PYIMQUS oo eee 13 ViTIGIS SSP. eee 1 
terrificus .... cass D Shephensi pee ogee decolor (=concolor) .. 1 
ECAR | ee ae 5 molossus hellert. 25 ee pa ene 31 
EMICOIOT eco cce hens. i gh? THOLOSSILSER ee ONE RANI Sie ee ree te 12 
CRIN O Bey eer Rete se D) IIT ESCONS ee oon eee 1 lutosus 2 aoe eee 4 
horridus ssp. .. Me) DOyStIChUS” =a ae ee 1 VITIDIS ees Be es ee 17 
OTTIOUC Bee ee oes es 11 DICED A ete Mette Bee 4 willandpscohs em 3 


(Lachesis) and 1933 (Lachesis, B. atrox, C. adamanteus); Gadow, 1920 (C. “durissus”!); 
Phisalix, 1912, 1914, 1922 (Lachesis) ; Radovanovic, 1937 (C. horridus, A. contortrix, A. blom- 
hoffi, T. wagleri, B. atrox); H. M. Smith, 1946 (C. intermedius, C. gloydi, C. triseriatus) ; 
M. A. Smith, 1943 (A. rhodostoma); and Steindachner, 1913 (T. mucrosquamatus). The 
vertebrae have been discussed from the viewpoints of myology (Mosauer, 1935), evolution 
(Johnson, 1955a, 1955b, 1956), osteology (de Rochebrune, 1881; Goette, 1897), and paleontol- 
ogy (Simpson, 1933; Gilmore, 1938; Brattstrom, 1953, 1954, 1955, 1958). The shaker (or style) 
of rattlesnakes has been described in some detail by Zimmermann and Pope (1948). The 
general anatomy of the skull and vertebrae of a rattlesnake were fully described and illustrated 
by Klauber (1956). Some of Klauber’s illustrations are reproduced here (figs. 1, 2). 
Anomalies, Diseases, and Injuries 

Injured bones were relatively rare among the skeletons examined in this study. The most 
common abnormality seen was the fusion of several vertebrae. Another abnormality or injury 
found was the enlargement of the bone at the junction of the dentary with the angular and 
splenial. The lower jaw is probably the weakest group of bones in the snake’s body, and the 
dentary junction seems to be especially fragile. It is possible that fracture of the bones of the 
lower jaw occurs during striking or during the eating of oversized prey. A double squamosal 
bone was found on each side of the head of one specimen of Crotalus horridus. 

Ontogenetic Variation 

Small variations were noted in the shapes of certain bones and processes. Some of these 
variations appeared to be ontogenetic, but most appeared to be individual or geographic varia- 
tions. Study of variation in species represented by large samples (such as A. contortrix, A. 
piscivorus, B. atrox, C. atrox, C. cerastes, C. ruber, C. viridis, T. wagleri) helped evaluate 
differences found in species of which few specimens were available. 

There is no major change in the shape of the various bones of the skull of crotalids after 
birth. Cranial bones become strongly ossified during the first or second year. After ossification, 
the bones increase in size with the general growth of the snake. In a series of 11 new-born 
Bothrops atrox from a single brood from Panama, there was very little variation in the skulls. 
These individuals did not differ greatly from the adults except in three characters: the dorsal 
surface of the parietal had not yet become completely ossified and was triangular and arched; 
the ectopterygoid was not greatly forked; and the ventral process of the basisphenoid had not | 
yet developed. 


1The figure looks like C. horridus, however. In one of the drawings there are four teeth in the left 
pterygoid, in two other drawings, supposedly of the same individual, there are five. 


1964 } BRATTSTROM: EVOLUTION OF Pir VIPERS 19] 


oe eee 5 
lone 
pps 7 
se dee 
een 
2 SS ee 
a a 2 
a 
6~ <---4 
ANTERIOR 
POSTERIOR 


Fig. 2. Mid-thoracic vertebra of Crotalus ruber ruber. From Klauber, 1956. a, front 
view; b, rear view; c, side view. 

Key to parts: 1, Condyle or ball of centrum; 2, Diapophysis; 3, Cup of centrum; 4, Hypapophysis; 
>, Neural spine; 6, Parapophysis; 7, Postzygapophysis; 8, Prezygapophysis; 9, Zygantrum; 10, Zygosphene. 


Most crotalids show little ontogenetic change; however, Lachesis muta apparently is an 
exception. If the lines of Lachesis measurements are extrapolated backward on the various 
graphs, they do not conform with the size changes shown in the other genera of the family. 
It is therefore apparent that either there is ontogenetic change in relative lengths of bones in 
this species, or the young hatch extremely large. 

Because bone dimensions increase with age, it is useful to have some standard reference 
with which to compare measurements. Since fang length and head length were measurable on 
most skulls examined, these were used as a basic index of age. The “head length” measured 
on preserved specimens by Klauber (1937, 1938, 1939) differs by only a few millimeters 
(the thickness of the skin) from the “lower jaw length” used here. 


192 SAN Dieco SociETY OF NATURAL History | VoL. 13 


TABLE 4 
Counts of tooth sockets in Agkistrodon, showing number of specimens', (range), and mean. 
Species Palatine Prerygoid Dentary 
acutus # 2(4)4 23)13 2 (17-18) 17.5 
Prentas Se 106 5)3. 8 10 (14-17) 15.1 9 (16-18) 16.2 
blomhoffi 2(4)4 2 (12) 12 2 (11-12) 11.5 
eentorent _ 22(45)4.4 24 (12-21) 16.1 19 (12-17) 15.1 
aly: a ee eee 2 (3-5)3.5 3 (9-13) 113 4 (12-13) 12.3 
hypnale Re ee 2(4)4 2(15-16) 15.5 ava nvs 
piscivorus ssp. ice eer LOO) 15 (13-16) 14.9 15 (16-20) 18.3 
piscivorus RR OL ME iat yea iy 14(4-5)4.9 14(13-17) 15.1 14 (18-21) 18.6 
leucostoma Bie Pee ce trees AG ie) Ee 16(14-18) 15.4 12 (16-18) 17.8 
8 ae te ae A eee em 41(4-5)4.9 45 (13-18) 15.1 41 (16-21) 18.2 
strauchi OS 3, Wir oo Pence 1Gs)32 2(7-8)7.5 2 (10-11) 10.5 
Tegel Wa cteshe: 82 (3-5) 4.3 90(7-21) 13.6 81(10-21) 15.5 


1JTn tables 4-7, the number of specimens is the number of bones with countable sockets, not the number 
of skulls. Occasionally the paired bones differed by one or two sockets, and occasionally a tooth-bearing 
bone was lost or broken on one side of the skull. 


At the suggestion of L. M. Klauber, I tested the methods used by Bryuzgin (1939), 
Petter-Rousseaux (1953) and Senning (1940) for determining the age of reptiles and amphi- 
bians by counting growth rings in bones. Most of the bones of crotalids show a series of light 
and dark rings. These are most evident and most easily counted on the neural spines and the 
ectopterygoid and articular bones and particularly on the frontal bone. Counts of the number 
of rings on each of these bones from one individual were the same. 

The southern California population of Crotalus viridis helleri is relatively homogeneous 
(Klauber, 1936b, 1937, 1945, 1949). When ring counts on skulls from this population were 
plotted on a log-log scale against head length and against fang length, S-shaped growth curves 
resulted. Similar data from another relatively homogeneous series (C. viridis oreganus from 
Washington and Idaho; Klauber, supra cit.) resulted in very similar curves. Similar data for 
other species of Crotalus show the same type of growth curves as for C. viridis, though the 
slopes of the curves, and hence probably the rates of growth, are different for the different 
species examined. 

The greatest number of rings counted in a rattlesnake born in captivity with the skull 
available was 16, though C. viridis has been kept in captivity for 19 years (Shaw, 1957). 

These data plus the conclusions reached by Peabody (1958, 1961) and by Senning, 
Bryuzgin, and Petter-Rousseaux (supra cit.) suggest that these rings are useful for the deter- 
mination of age and may, in fact, be growth rings, one added per year. 


Geographic Variation 


Small geographic variations were noted in size and shape of certain bones and processes. 
In general, the variation noted was minor; the skulls of any one species were usually very 
similar. The osteological similarities within species and differences between species were sufh- 
ciently constant, not only for characterization of species and determination of relationships, but 
also for construction of artificial keys to skulls (see Klauber, 1956). Although the variation 
found among individuals of local populations was small, a greater amount of variation was 
evident between widely separated populations of the same species. Some examples of geographic 
variation follow. 


1964 } BrRATTSTROM: EVOLUTION OF Pit VIPERS 


TABLE 5 


193 


Counts of tooth sockets in Bothrops, showing number of specimens, (range), and mean. 


See footnote in table 4. 


Species Palatine 
LUG TITREN Bo, SARA ee PSOE DS sR 


Pterygoid 
2 (10-11) 10.5 


Dentary 
3 (10-12) 11 


atrox: total ...... an WG>)41 13 (13-18) 15.5 18 (14-19) 16.7 
Bnleehe 4 oe Ae ht! 10 (3-5) 4.2 7 (13-17) 15.7 11 (16-19) 16.8 
at + | <n eee 6 (14-18) 15.2 7 (14-18) 16.6 

UTA AELT Cet See tenn ete <1 A Seek) Ree 2(5)5 2 (17-18) 17.5 2(17) 17 

Con ANNO © iene SIN Rais on 1 (32) 3? 2(11-12) 11.5 2(12-13) 12.5 

ROC anOR Veh Aa By ol Abedin  2(4)4 4(12) 12 4(14-17) 15.3 

Faria net ene oe ee oe 245)45 4(15-17) 16.3 4(6-17) 10.8 

HANETACUASIE Go AA. te: eee we SR oe as ccces 8 (3-5) 3.8 8 (14-17) 16.0 8 (13-18) 15.4 

Lanceolalisme, Seek EMA T ees. 2(2)2 2 (10-11) 10.5 2 (12-13) 12.5 

TESTLTEN a> AS ene SPR ea He ee eee — 2 (10-11) 10.5 2 (12) 12 

NOTE Tas | eae eae eee Bek a Sahar Tee eoe 4 (3-4) 3.5 4 (10-12) 11.5 4(10-13) 11.5 

ARG) ee eee 2(4)4 2 (13-14) 13.5 2(14-15) 14.5 

schlegelt| <...2... ae en 2O)e 2 (16) 16 2 (15-16) 15.5 

Tact eek 5 RAEN id. 44(2-5)3.8 49 (10-18) 13.5 55 (6-19) 14 


1New-born brood. 


The nasal bone is variable in many crotalids, but some constancy apparently has developed 
in Crotalus cerastes. About 90 per cent of the skeletons of two subspecies can be distinguished. 
In C. c. cerastes, from the Mohave Desert, the nasal is square, whereas in C. c. laterorepens, 
from the Colorado Desert, it is elongate. No specimens were available of the Arizona form, 
C. c. cercobombus. 


In C. durissus terrificus and C. d. unicolor the nasal bones are squarish, with anterolateral 
processes; in C. d. tzabcan they are squarish or rectangular, usually with posterolateral processes. 
The medial edge of the squamosal is usually straight in the former two and curved in the 
latter. The anterior hump of the process of the basisphenoid is usually low and rounded in 


C. d. tzabcan and well developed and pointed in C. d. terrificus. 


To judge from the few skulls with subspecific or locality data, in C. h. horridus the basis- 
phenoid process has a pointed anterior hump, while in C. h. atricaudatus the hump is absent 
or low. 


Crotalus viridis varies in the width of the parietal, the curvature of the squamosal, and the 
shape of the dorsal process of the premaxilla. In C. v. decolor (=concolor), the parietal is a 
relatively elongate triangle, and the squamosal has straight sides except anteriorly, where it is 
curved. In C. y. lutosus, the parietal is almost an equilateral triangle, the squamosal has 
straight sides which taper anteriorly, and the dorsal process of the premaxilla is an elongate 
triangle. In C. v. helleri the parietal is an elongate triangle, the squamosal has curved sides, 
and the dorsal process of the premaxilla is thin and expands into a small triangle at its base. 
In C. v. oreganus from Washington and Idaho, the parietal is a broad triangle, the squamosal 
has straight sides, though somewhat curved and blunt anteriorly, and the dorsal process of the 
premaxilla is thin, with a broad triangle at its base. In one specimen of C. v. oreganus from the 
Sierra Nevada of California, the parietal is broader than usual, and the dorsal process of the 
premaxilla is a broad triangle. In C. v. viridis the parietal is a broad triangle, the squamosal 
has straight sides and is blunt anteriorly, and the dorsal process of the premaxilla is thin. 


194 


San Disco Society oF NATURAL History 


TABLE 6 


{Vot. 13 


Counts of tooth sockets in Crotalus, showing number of specimens, (range), and mean. 


See footnote in table 4. 


Species Palatine Prerygoid Dentary 
adamanteus 2 39\(123))227 40 (7-11) 8.1 39 (9-10) 9.5 
Ga ee EE Ce RE Ot 67 (3)3 70(7-9)8 70(7-11) 9.9 
BPECHECEU SS Vd occy secs Pa ee ees: 4(0-2) 1 4 (7-8) 7.5 4(9-11) 9.8 
eee 44 (2-4)3 46 (7-9) 8.4 45 (8-11) 9.8 
ET ERSTLG a eee et eee 18(0-2) 1.1 12(7-9)7.8 14(8-11)9.8 
Sag en eee ee 6(1-3)2.7 2(8)8 4 (9-11) 10 
horridus 28 (2-3) 2.9 34 (8-11) 8.7 34 (10-13) 10.9 
TRLTITCOUUS 280s 2 5 cca cae Renee: 1(3)3 ING) )3: 1(8)8 
[Sa Li Ta Seas aOR RO oP SE Casas 7 (2-3) 2.7 8 (6-8) 6.9 10(8-9) 8.1 
HEPCICLUE hho oot traE e Be: 1d(223)) 29 24 (7-10) 8.2 23 (7-10) 8.9 
111 GATOS Ne an ae 3 Re ce rer 12 (1-3) 221 12 (6-10) 8.8 9(9-11) 10 
PIAUYSEICEMS © Oooo: esc ee ewe tse 2(0)0 2(7)7 2(8)8 
Pes tllriG Take BE ee nt hi OO Ree ee, 9(3)3 9 (7-9) 7.7 10(9-11) 10.1 
PUREN A hiae se 8 op chat gk te oe se 3(3)3 5 (6-7) 6.6 3 (9-10) 9.3 
PUDET PRs CL 27 (3)3 26(6-9)8 31 (8-11) 9.5 
SST ET ET en A EE LTE 16(3)3 16(6-8) 7.3 14(9-10) 9.6 
BRC CEI ages. Corn mmdetea aetna: 4(0)0 31(4-5) 4-7, 4 (6-7) 6.5 
PEATE Mai ana aactne so tana sas 4(3)3 4(7-9)8 3 (9-10) 9.3 
ESI CEIISES, icin, gore eased oe Sunes tos «ease 2(8)3 2(9)9 2 (10) 10 
PF ISERIGEUG. cree Meena Bu React 4(3)3 9 (6-7) 6.2 3 (7-8) 7.3 
a ate hcg 0 ey PR Oe ee ane Re aE ae 103 (2-4) 2.9 108 (6-10) 7.4 107 (6-11) 9.3 

ALECOLUT eee tees nG)s 2 (6-7) 6.5 2(9)9 
PEO SIS sree Meee’. :_ 09 te Ade: Sel. Serpe | 8 (2-3) 2.5 6(7-8) 7.2 6(9-10) 9.3 
Preller tat Le) ACO Nee Tn 47(3)3 52 (7-10) 7.8 50 (8-11) 9.4 
oreganus 17 (2-4) 3.1 16(6-9) 7.8 18 (9-10) 9.6 
NELLA Ue RR ee tae) Ee AMR Pec Sie yeas 30(2-3)3 32 (7-9) 7.7 31(6-11)9.1 
MALIA TRE Stee he ee ae NS ee 4(1-2) 1.8 6 (5-7) 6.2 5(8)8 
Tico ieee pe 417 (0-4) 2.4 440(4-11)7.4 437 (6-13) 8.7 


In one specimen of Lachesis muta from Colombia (AMNH 63419), the basisphenoid 


process is triangular, whereas in three specimens from Panama it is rectangular. In the Colom- 
bian specimen, the lower two processes of the prefrontal are not strongly fused, and the dorsal 
process is about equal to the middle process in length; the lower process is the shortest. In the 
Panamanian specimens, the lower two processes of the prefrontal are fused and are longer than 


the upper. 
DESCRIPTIVE OSTEOLOGY 


The osteology of each species is not described in detail, but rather is illustrated (figs. 15 
to 32) and summarized (tables 10 to 12). In the tables, the various characters are listed for 
each species. The detailed descriptions of the bones, the variations noted, and the specific 
characters used, are given below. A summary of the tables, in terms of relationships, will be 
given beyond. 


1964 } BRATTSTROM: EVOLUTION OF Pit VIPERS 195 


Teeth 


In the Crotalidae, teeth are found on the maxilla, pterygoid, palatine, and dentary bones, 
with the fang in the maxilla. The fang has been fully discussed by Klauber (1939, 1956) and 
by H. M. Smith (1952). The fangs of the various crotalids are similar but differ slightly in 
length and curvature, and in the size and shape of the lumen (Klauber, 1939). Trimeresurus 
wagleri has the most atypical fang of any crotalid, for the lower lumen usually is on the inner 
or medial side, not on the anterior surface as in the rest of the crotalids. In one specimen of 
T. wagleri, however, the lower lumen is anteromedial. 


In one specimen of Lachesis muta (BHB 1635) from Panama, one of the functional 
fangs has two lower lumen openings, one above the other, with a groove between them and 
another groove above the upper of the two. There are two functional fangs in each maxilla of 


one specimen of Bothrops atrox examined (USNM-NZP 4072). 


The number of teeth was determined for each crotalid skull examined. As shown by 
Bogert (1943), accurate tooth counts can be obtained only by counting the sockets and not 
by just counting the teeth present. Counts of tooth sockets can be made on preserved speci- 
mens as well as on skeletons. Tooth socket counts for the crotalids examined are given in 
tables 4 to 7. The number of tooth sockets plotted against size of snake indicate that in crota- 
lids there is no change in the number of teeth with age. 


It is a general rule that more primitive vertebrates have a greater number of teeth. How- 
ever, a large number of teeth can sometimes be a specialization, as in mosasaurs and porpoises. 
Since it is difficult to determine whether a large number of teeth is specialized or primitive 
among crotalids, little value was given to this character in determining relationships. It is 
interesting, however, that several crotalids have a small number of palatine teeth: C. durissus, 


C. polystictus, C. stejnegeri, C. willardi, T. jerdoni, and T. flaviviridis (tables 4-7). 


Among species of crotalids that are relatively homogeneous in external characters, there 
is little variation in tooth counts, but in heterogeneous species, such as C. viridis, there is wide 
variation. Some species have extremely elongate, straight teeth, and others have short, stubby 
teeth. Crotalus polystictus is unusual in having extremely elongate, narrow dentary and ptery- 
goid teeth, with the basal one-fourth of each tooth enlarged. 


Bones 


Premaxilla (Premaxillary).— The premaxilla (figs. 15, 16) of crotalids is T-shaped, 
with the cross-bar of the T forming the anterior ventral edge of the bone. The vertical part 
of the T forms a dorsal projection. This projection has a thin posterior process that acts as a 
septum between the nares. The anterior edge of the basal portion of the bone may be pointed, 
curved, or flat. The dorsal process may be thin, thick, or thick at the base and thin dorsally 
(see C. atrox, fig. 15). The premaxilla supports the rostral scale, and there appears to be some 
relationship between the shape of the premaxilla and that of the rostral scale. A gkistrodon 
acutus, for example, has a pointed rostral scale and a pointed premaxilla (fig. 16). 


The shape of the premaxilla is variable within some species. In the various subspecies of 
C. viridis (fig. 15), the basal portion is usually pointed, but the dorsal process varies in thick- 
ness and shape. 


Prefrontal. — The prefrontal (figs. 15, 16) of crotalids does not vary greatly within a 
species. In Bothrops, Trimeresurus, and Lachesis, it extends medially more than in A gkistrodon, 
Crotalus, and Sistrurus. The three posterior processes of the prefrontal are quite characteristic 
for each species with regard to relative size and fusion of processes (figs. 15, 16). 


Vomer, turbinal, ethmoid. —These bones were not examined critically since their shapes 
and positions seemed not to vary greatly. The turbinals are relatively small in most species; 
however, they are large and have a high dorsal projection in the larger snakes, such as 
C. ruber, C. atrox, C. adamanteus, C. durissus, B. atrox, and Lachesis muta. 


196 San Dieco Society oF Natrurat History (Vor. 13 


TABLE 7 
Counts of tooth sockets in Lachesis, Sistrurus, and Trimeresurus, showing number of specimens, 


(range), and mean. See footnote in table 4. 


Species Palatine Pterygoid Dentary 
L. muta ee eee oer 8(3)3 8 (9-10) 9.6 8(9-12) 11 
De AP ORALUS. cites hohe oa OER Re 17 (1-3) 2.6 19 (5-7) 6.5 20(9-10) 9.1 
S. miliarius A eee ae Faerie #{(6)))3) 4(7-9)7.8 7 (10-11) 10.4 
SAVE ecg: ” RPE Soh A 10(3)3 10(5-7)7.9 10(8-10) 8.8 
Sistrurus totals Se pk Pies lag 29 (1-3) 3 33 (5-9) 6.4 37 (8-11) 9.4 
De Dolabrig. ss 2) een Sn eee 2 (4)4 Det) 2GM-1S) AZ 
T.. flaviviridis heen = AO) Os 4 (11-14) 12.5 4 (10-17) 13.3 
T. mucrosquamatus .. dee steer eens —— — 2 (10-11) 10.5 
DD PUTIC CUS tert eee Sea ee sSesee Zi) > 212) 2 2 (15-17) 16 
T. purpureomaculatus ER Re A 2(4)4 4 (8-12) 10.8 4 (LEZ) 8 
TEMS MEG ETE \ccutte eie ace RBar ous eee Bee —. 2 (12-13) 12.5 2 (12-14) 13 
Mire pidplent rar nae oe ents oder VO)2 10 (13-14) 13.5 8 (11-16) 14.5 
Trimeresurus totals: 5.20000 ncsen<- 19 (0-5) 3.6 24 (8-14) 12.1 24 (10-17) 11.6 


1M. A. Smith (1943), quoting Walls, said that T. jerdoni also lacks palatine teeth. 


Nasal. — The nasals of crotalids (figs. 17, 18) are paired flat bones, occasionally curved 
laterally and anteriorly, with ventrally projecting medial processes which meet, but do not fuse, 
at the midline. These processes articulate independently on the ventral surface of each frontal. 
There is usually a cartilage-like tissue connecting the anterior and lateral edges of the nasals 
with the premaxillae and prefrontals. The nasals are usually longer than wide. 

The shape of the flat dorsal surface of the nasal (fig. 17) is variable within some species, 
such as Crotalus durissus, C. horridus, C. mitchelli, C. molossus, C. viridis, and C. cerastes. 
As noted above, some of this variation is geographic. Nasal variability in most species is so 
great as to eliminate the use of this bone in determining relationships. 


Many species of crotalids have accessory hooks and processes on the lateral and posterior- 
lateral corners of the nasals (e.g., C. horridus and C. c. cerastes, fig. 17). These processes are 
variable among the individuals within a species; for example, they were present on the nasals 
of only 3 of the 25 C. cerastes skulls examined. Possibly the variation results from differences 
2 pone during development. These processes are often destroyed or lost in preparation 
of skeletons. 


One of the two skulls examined of the ridge-nosed rattlesnake, C. willardi, had the nasals 
slightly raised anteriorly. 


Palatine. — The palatine (figs. 17, 18) is a thin, vertical tooth-bearing bone articulating 
with the anterior end of the pterygoid. It lies between, but does not articulate with, the medial 
wall of the maxilla and the lateral edge of the vomer. It is deeply imbedded in connective 
tissue and is dorsal to the ventral level of the maxilla. This position, plus its weak articulation 
2 ‘e pterygoid, probably reduces its effectiveness in food-holding during the swallowing of 
ood. 

In the supposedly primitive forms of the Crotalidae and in the Viperidae, the palatine is 
small and narrow. In each genus of the Crotalidae, it seems to have undergone similar evolu- 
tionary changes, becoming thin and high. It is high and triangular in most species of Crotalus 
and A gkistrodon (figs. 17, 18). There is often a small anterior process or knob on the palatine 
in advanced forms of these two genera. The palatine has a small dorsal process in Lachesis. 


1964 | BRATTSTROM: EVOLUTION OF Pit VIPERS 197 


FRONTAL PARIETAL 


L H 
WwW : 
Hz-| << 
SIDE y 


VIEW OF 
| VERTEBRA 
M 


Soe 


LOWER JAW 


HV 


I W,Z | agwWe PRE 


ee : 
SS @ 
= H WV 
L L Ww 
Ss ) _we post 
KH 


DORSAL VIEW 


a Ee OF VERTEBRA 


SQUAMOSAL ECTOPTERYGOID PTERYGOID QUADRATE W POST 
Fig. 3. Methods of measurement of crotalid bones. 
Abbreviations: A, articular; C, centrum; H, height; L, length; LH, length of hypapophysis; M, 
Meckelian foramen lateral border; NS, neural spine; P, process; POST, postzygapophysis; PRE, prezygapo- 
physis; V, vertebra; W, width; Z, zygosphene. 


One of the more important characters that distinguish most species of Bothrops from Trimere- 
surus is that the palatine is forked in most of the former and in none of the latter (Ruiz, 
1951). It is not forked in B. nummifer and B. schlegeli, two species that Ruiz did not examine. 

Frontal. — The frontal bones (figs. 19, 20) are flat and elongate, square, or, occasionally, 
wider than long. They articulate posteriorly with the parietal and sometimes with the post- 
frontal, anteroventrally with the nasals, and anterolaterally with the prefrontals. The antero- 
lateral process has many shapes. 

The lateral edge of the frontal lies above the eye socket and often is curved inward, 
conforming to the shape of the eye. Probably for the same reason, the frontal is often turned 
up laterally. In the sidewinder or horned rattlesnake, Crotalus cerastes, this raised portion sup- 
ports the basal tissue of the horn. The horn itself is not, however, supported by bone, nor is it 
hard like that of the viperid Cerastes cerastes of the Sahara Desert. Crotalus enyo also has a 
raised frontal, where a small fleshy horn appears in life. The frontal of Bothrops schlegeli is 
flat and does not contribute to, nor seem to aid in, the support of the thin flap-like scale 
above the eye in that species. A raised frontal may contribute to the general shape of the 
head in other crotalids, aiding in support of the canthal ridge or supraocular scale. 

The shape of the frontal is relatively constant in each species. The width of the bone was 
measured at its narrowest part, and the length was measured midway between the sides (fig. 3). 
The frontal is longer than wide in all species of Bothrops examined except B. schlegeli and 
B. nummifer. It is longer than wide in all species of Sistrurus and Crotalus except C. ada- 
manteus and sometimes C. atrox. In C. adamanteus it is wider than long except in some very 
old specimens, in which it may measure longer than wide though it looks wider than long. 
The frontal of C. atrox is elongate in young individuals and in most adults, but a few large 
adults have frontals wider than long. The frontal is wider than long in Lachesis muta and 
Trimeresurus wagleri. The latter is the only crotalid with the center of each frontal depressed. 
All the other species of Trimeresurus have elongate frontals. The frontals of most species of 
Agkistrodon are elongate; those of A. acutus, A. bilineatus, and A. piscivorus are squarish 
and occasionally wider than long. 


198 SAN Disco Society oF NATURAL History | Vorms 


IN MM. 


LENGTH 


QUADRATE 


10 20 30 40 
ECTOPTERYGOID LENGTH IN MM. 


Fig. 4. Relationship of quadrate length to ectopterygoid length in various crotalids. Solid 
line, observed range; dashed line, extrapolation; solid squares, Lachesis. 


Do) 
° 


IN MM. 


LENGTH 


3 


QUADRATE 


10 20 30 40 50 
LOWER JAW LENGTH IN MM. 


Fig. 5. Relationship of quadrate length to lower jaw length in a few species of Crotalus. 


Solid line, observed range; dashed line, extrapolation; solid circles, durissus group; open circles, 
viridis group; crosses, C. cerastes. 


1964 } BRATTSTROM: EVOLUTION OF Pit VIPERS 199 


Maxilla (Maxillary).— The maxilla of crotalids (figs. 21, 22) is a heavy bone holding 
the functional fang. The maxilla has a large lateral opening which contains the heat-sensitive 
facial pit. Bullock and Cowles (1952) and Bullock and Fox (1957) discuss the anatomy and 
physiology of the facial pit and its relation to the shape of the cavity of the maxilla. 


As described by Klauber (1939), the forward rotation of the fang is due to a change in 
position of the quadrate, forcing the ectopterygoid to push on the maxilla and to rotate it 
against the prefrontal. In Crotalus and Sistrurus, most of this forward rotation of the maxilla 
is afforded by the medial anterior process of the ectopterygoid, which lies flat against the 
posterior part of the maxilla. In Bothrops and Agkistrodon, the movement is due to the lateral 
process of the anterior end of the ectopterygoid pushing on the posterolateral corner of the 
maxilla. This articulation differs greatly in the various species of Trimeresurus (fig. 22). 


The shape of the anterior edge of the pit cavity is of some taxonomic importance, as 
Ruiz (1951) has shown. This edge is a smooth open curve in Agkistrodon and Bothrops 
(fig. 22). In Trimeresurus it is irregular and may have a posteriorly pointing median process. 
This process is barely discernible in Bothrops schlegeli; it is absent in the related B. nummifer. 
The curvature of the anterior edge of the pit cavity of Crotalus, Sistrurus, and Lachesis has a 
definite, prominent posterior or dorsal process. In Lachesis this process has a small depression 
on its outer surface (fig. 22 and Ruiz, 1951). The prominence of this process is emphasized 
in Crotalus and Sistrurus (fig. 21) because of a depression posterior to the process. In some 
species (C. viridis, tigris, mitchelli, cerastes, enyo, and scutulatus) this depression extends 
anteriorly; this extension is one of the characters that suggest a close relationship among these 
species. The maxilla of Trimeresurus resembles the hypothetical primitive form of the family 
because of the relatively small size of the pit cavity. 


Parietal. — The parietal (figs. 23, 24) is the largest and heaviest bone of the crotalid 
skull. It is a single bone that protects the brain and gives support and attachment for many 
of the head and jaw muscles. 


The definitive shape of the parietal appears to be acquired during the first and second years 
of growth. In many species, however, there are changes throughout life in the shape and posi- 
tion of the lateral processes. The lateral processes are flat and thin and are parallel to the 
dorsal surface of the parietal, but more ventrally placed. The muscles of the lower jaw, the 
muscles of the squamosal-quadrate complex, and the muscles that force venom out of the 
poison gland are all attached to these processes. One, or part of one, of these processes usually 
forms a curved ridge at the anteromedial end of the squamosal. The medial limit of these 
processes is difficult to determine on young pit-vipers and on large adults of a few species. 
Because of the variability of shape and position of these processes, they were given little value 
in determining relationships. The term “dorsal surface of the parietal” is used in this paper 
for the shape of the dorsal surface excluding these lower lateral processes. 

The shape of the dorsal surface of the parietal is characteristic for each species. It usually 
expands laterally just posterior to its anterior edge. The postfrontal bone articulates with the 
parietal on these lateral extensions. The greatest width of the parietal and of the skull is at 
this lateral extension; this width may be increased by the lateral extent of the postfrontal. The 
head width, as measured just posterior to the eye on preserved snakes, is equal to the greatest 
width of the parietal (or parietal plus postfrontal where the postfrontal extends beyond the 
parietal) plus the thickness of the skin. The anterolateral extensions of the parietal are often 
raised slightly, especially in those species in which the frontal is also raised. 

Posterior to this lateral extension, the dorsal surface of the parietal narrows to a point or 
ridge, which is usually notched posteriorly. In some species there may be a second lateral 
expansion of the dorsal surface of the parietal (fig. 23, C. atrox, horridus, and molossus). The 
parietal extends posteriorly and laterally beyond the dorsal surface of the parietal and meets 
the occipital, squamosal, and otic areas. 

In all crotalids the dorsal surface of the parietal is wider than long. It is T-shaped or 
triangular in Agkistrodon. It is squarish, trapezoidal, or triangular in Bothrops, except for 


200 SAN Dteco Society oF NAtTurRAL History {Vour. 13 


LENGTH IN MM. 


QUADRATE 
fo) 


10 30 50 70 
LOWER JAW LENGTH IN MM. 


30 


QUADRATE LENGTH IN MM. 
fo) 


10 re) 50 70 
LOWER JAW LENGTH IN MM. 

Fig. 6 (above). Relationship of quadrate length to lower jaw length in Agkistrodon. 
Solid line, observed range; dashed line, extrapolation; solid circles, A. piscivorus; crosses, other 
species of A gkistrodon. 

Fig. 7 (below). Relationship of quadrate length to lower jaw length in a few genera of 


crotalids. Solid line, observed range; dashed line, extrapolation; solid squares, Lachesis. 


B. nummifer and B. schlegeli, in which it is T-shaped. In Lachesis it is T-shaped, and in 
Trimeresurus it is elongately triangular or T-shaped. In Crotalus and in Sistrurus ravus the 
parietal is triangular. It is trapezoidal in S. catenatus and S. miliarius. In a small specimen of 
S. catenatus tergeminus (USNM 29132, Fort Davis, Texas) there is a small groove down 
the middle of the parietal. A similar groove was found on one specimen of Agkistrodon 
contortrix, a medium-sized adult from Kansas (UIMNH, no number). 

The length of the dorsal surface of the parietal was measured at the mid-line, and the 
width was measured across the anterolateral extensions. On those species in which the post- 
frontal extends laterally beyond the parietal, two measurements were taken — parietal width 
and parietal width plus postfrontal. Because of the shape of the parietal under the postfrontal, 
the difference between the two measurements was not great except in Lachesis, in which the 
postfrontals extend laterally more than in any other crotalid. 


1964 } BRATTSTROM: EVOLUTION OF Pit VIPERS 201 


A graph (not reproduced here) of length of the parietal plotted against width shows 
little variation for all crotalids. The curve for Trimeresurus wagleri and those for the “arboreal” 
group of Bothrops fall close together. The curves for Trimeresurus (other than T. wagleri) 
and those for the large terrestrial species of Bothrops also are close together. The curve for 
Agkistrodon contortrix is close to that for the Old World Agkistrodon, and the curve for 
A. bilineatus is closest to that for A. piscivorus. The curves for Crotalus and Sistrurus are 
closer to those for A gkistrodon than to those for Bothrops and Trimeresurus. 


Postfrontal.— The postfrontal, though large and wide in the Viperidae, is very small in 
the Crotalidae. It appears to be undergoing a reduction in size within the family and is 
extremely small in Bothrops nasuta. Apparently, however, it is present in all crotalids though 
often lost in the preparation of the skull. Because of apparent loss, it is not shown on some 


of the drawings (figs. 23, 24). 


The postfrontal is in a depression on the anterolateral expansion of the parietal and 
may articulate on the top, side, or end of this parietal extension. The postfrontal either touches 
the frontal or it does not. Which condition obtains is usually characeristic for a species, and 
in some groups it is useful in determining relationships. In some species, however, the con- 
dition is variable. In these species, the number of skeletons in which the postfrontal touches 
the frontal and the number in which it does not are as follows: C. horridus, 5-9; C. mitchelli, 
1-9; C. molossus, 3-2; C. ruber, 10-4; C. atrox, 21-10; B. neuwiedi, 1-1; A. bilineatus, 5-1. In 
C. ruber, C. atrox, and A. bilineatus, the distance separating the two bones is always less than 
1 mm. 


Otic. —No great differences were noted in the otic areas of the skulls examined. 


Occipitals. —The pit-viper skull contains a single supraoccipital, which is often fused 
with the parietal. The two exoccipitals (=lateral occipitals) meet on the mid-line dorsally 
and articulate with the basioccipital ventrally. The exoccipitals, together with the basioccipitals, 
form the foramen magnum and contribute to the occipital condyle. The basioccipital, which has 
a large ventral process, articulates with the basisphenoid and otic bones. The occipitals seem 
not to vary significantly in crotalids except for the basioccipital, which will be discussed below 
with the basisphenoid. 

Stapes (Columella auris).— The stapes is a thin bone extending posteriorly and laterally 
from the oval window to the ventral side of the quadrate bone. Because the stapes is often 
lost in preparation, no special note was made of its variation. 

It is often said that snakes “hear” by feeling vibrations from the ground (Tumarkin, 
1955). Though no physiological or behavioral work has been done, it is suggested here that 
the stapes and its fusion to the quadrate might well act as a mechanism or route for transfer 
of sound to the inner ear. A vibration detected by the snake from the substrate might be trans- 
mitted by the lower jaw to the quadrate and then to the stapes, or by the ribs to the vertebrae, 
to the skull, to the quadrate, and to the stapes. 

Basisphenoid. — The basisphenoid (figs. 25, 26) is a flat bone with a thin ventral process. 
This process fuses posteriorly with the ventral process of the basioccipital and separates the 
two muscle masses of the ventral surface of the skull. The basisphenoid and basioccipital form 
most of the ventral surface of the skull. The shapes of the anterior and anterolateral processes 
of the basisphenoid show individual variation that precludes the use of this character in deter- 
mining relationships. 

The ventral processes of the basioccipital and basisphenoid are shown in side view in 
figures 25 and 26. The ventral process of the basioccipital generally is heavy and curves pos- 
teriorly; it varies in curvature, height, and width. The ventral process of the basisphenoid is 
thinnest anteriorly. It becomes thick just before it joins the process of the basioccipital. 

Members of the atrox group of Crotalus (adamanteus, atrox, ruber, and tortugensis) 
have a high ventral process on the basisphenoid. Crotalus durissus and basiliscus have a small 
anterior hump on this process. The members of the viridis group, except C. mitchelli, all have 
low ventral processes. Most of the small Mexican rattlesnakes (C. lepidus, pricei, and tri- 


202 SAN Disco Society oF NaAtTurRAL History EVor) 13 


30 


IN MM. 


LENGTH 
nm 
oO 


QUADRATE 


lo 20 30 
PARIETAL LENGTH IN MM. 


Fig. 8. Relationship of quadrate length to parietal length in a few genera of crotalids. 
Solid line, observed range; dashed line, extrapolation; solid squares, Lachesis. 


seriatus) have low processes, as does Sistrurus miliarius. An anterior hump is characteristic of 
all Agkistrodon skulls examined. It is especially high and well developed in A. acutus, bilinea- 
tus, and piscivorus, and is small in all others, including contortrix. The anterior hump is 
present in the species of Bothrops except in schlegeli. Lachesis muta has a well developed 
rectangular process. The process is triangular in Trimeresurus wagleri. It is low in T. albolabris, 
purpureomaculatus, and stejnegeri, in the last having an anterior hump. Trimeresurus puniceus 
and flavomaculatus have well developed processes and anterior humps. 


Squamosal. — The squamosal is a thin, flat bone lying on the posterolateral corner of the 
parietal. The anterior end of the quadrate lies on the dorsal surface of the posterior end of 
the squamosal. This junction is movable, but the motion is largely due to the action of the 
quadrate. 


The squamosal varies in length and shape among the various pit vipers (figs. 27, 28). In 
Crotalus and Sistrurus, it has a lateral process or hook at its posterior end. The quadrate lies 
on the entire posterior end of the squamosal except for this process, which is round or pointed 
and forms an angle with the main part of the bone. This angle is 90° or less in Crotalus 


1964 } BRATTSTROM: EVOLUTION OF Pit VIPERS 203 


basiliscus, horridus, lepidus, tortugensis, triseriatus, viridis, willardi, pusillus, and Sistrurus cate- 
natus, and usually (10 out of 11 specimens) in S. muliarius. The angle is so acute that it 
forms a hook in C. pusillus, S. catenatus, and S. miliarius. The hooked squamosal was used 
by W. E. Taylor (1895) to distinguish skulls of Sistrurus from those of Crotalus. However, 
the hook is lacking in S. ravus and is present in C. pusillus; thus this distinction does not hold. 


Crotalus price: has a squamosal unique among crotalids, with a peculiar posterior process. 
Furthermore, in one of the two specimens examined, the anterior end of the bone is forked, 
with several small processes or serrations between the arms of the fork (fig. 27), though in 
the other specimen the anterior end is flat. 


In one skull examined (Crotalus horridus, LMK R-51), there are two squamosal bones 
on each side, one posterior and lateral to the other. The normal quadrate lies on the posterior 
squamosal, which lies on the anterior squamosal. The squamosals are abnormal in shape for 
this species, and it is probable that this condition developed during the healing of some injury. 


Squamosal length was plotted against quadrate length for all species. The resulting graphs 
(not shown here) are a straight line for each species, with very little scatter. Lines of the 
species groups of Crotalus are close together. Crotalus tortugensis has a squamosal relatively 
shorter than that of C. atrox. This is one of the few characters in which this island species, 
related to atrox, extends beyond the scatter of atrox. Sistrurus ravus has a relatively short 
squamosal as compared to other species of Sistrurus and thus is similar to C. lepidus and C. 
triseriatus. The slopes of the curves are distinct for Bothrops, Trimeresurus, Lachesis, and 
A gkistrodon. The greatest divergence from the family norm is found in Lachesis, which has a 
squamosal relatively longer than that of the other genera. Trimeresurus wagleri and T. puniceus 
differ greatly from most species of Trimeresurus in having relatively small squamosals and 
large quadrates. This condition is found also in Bothrops schlegeli, but not in B. nummifer. 


Quadrate. — The quadrate does not vary greatly in shape among the crotalids examined. 
It is usually wide anteriorly, narrowing posteriorly until just before the posterior articulation, 
where it expands. This articulation is wide, smooth, and often forked. No generic or major 
specific differences in shape were noted. 


The quadrate bone is of different relative length in different crotalids. In was hoped that 
the differences in length could be associated with a possible increase in efficiency in fang rotation. 
From a physical model and from mathematical calculations, Dr. Blaine H. Levedahl has sug- 
gested (personal communication) that any increase in relative length of any of the segments 
in the fang rotation sequence (skull, quadrate, pterygoid, ectopterygoid) would increase the 
mechanical efficiency of the system. An equal proportional increase in all parts would not 
increase the efficiency, but an increase in length of one part would. If this is the case, the 
relatively large ratio of squamosal to quadrate in Lachesis would make the bushmaster the most 
efficient in this regard. The primitive Trimeresurus wagleri, with its relatively short squamosal, 
would probably be the most inefficient. The quadrate is relatively short in Lachesis and rela- 
tively long in T. wagleri, however. 


Graphs of squamosal length plotted against quadrate length have already been discussed. 
Pterygoid length plotted against quadrate length also resulted in straight-line graphs with 
almost no scatter. Specific and generic differences can be noted but are not great. 


Graphs of ectopterygoid length versus quadrate length are similar — they are straight lines 
with slight scatter. Generic and specific differences are noticeable (fig. +). In Agkistrodon 
piscivorus, the quadrate is longer than the ectopterygoid. In other crotalids, it is usually 
shorter; the only exceptions seen are one specimen of Crotalus scutulatus and a few very large 
individuals of C. atrox and C. adamanteus. Measurements for A. contortrix fall on the same 
general line as those of the Old World species of Agkistrodon. The line for A. bilineatus is 
intermediate between those for the Old World Agkistrodon and A. piscivorus (fig. 4). The 


latter two lines have very little scatter. 


204 SAN D1geGo Socrety oF NATURAL HIstory {Vot. 13 


Quadrate length plotted against lower jaw length also gives straight lines, with almost 
no scatter. Specific differences in ratios are difficult to discern, but generic and subgeneric lines 
are distinct. A few of these graphs are shown in figures 5 to 7. 

Graphs of quadrate length against parietal length also are straight lines, though the scatter 
is great. The quadrate is always longer than the parietal. In T. wagleri the quadrate is rela- 
tively longer than that in most species of Trimeresurus. A relatively elongate quadrate is found 
also in B. schlegeli and B. nummifer. Except for these three species, the lines for Bothrops are 
different from those for Trimeresurus (fig. 8). No other major differences are noticeable. 


Pterygoid. — The pterygoid (figs. 29, 30) is a toothed bone that is narrow anteriorly, 
where the teeth are present, and thin and flat posteriorly. Anteriorly, the pterygoid articulates 
with the palatine, and posteriorly it joins with the articular bone. The pterygoid is curved 
laterally in its middle third. The curvature on the lateral edge of the bone is more anterior 
than that on the medial edge. The ectopterygoid usually fits into a groove on the lateral edge 
of the pterygoid and may articulate anterior to, in the middle of, or posterior to the lateral 


curvature (figs. 29, 30). 


In Crotalus, Sistrurus, and Lachesis, the pterygoid teeth extend at most only to the middle 
of the junction with the ectopterygoid and often not so far. In the other genera of crotalids 
and in the viperids, the teeth extend beyond this junction. 


The posterior part of the pterygoid usually has a ridge on its ventrolateral surface. This 
ridge may be bordered by deep lateral grooves. Occasionally the top of this ridge is bent over 
laterally and thus encloses a small cavity. Many muscles attach on the ventral surface of the 
pterygoid, and the shape and position of the ridge are often related to the position of attach- 
ment of these muscles. An additional ventral process is present on the pterygoid of A gkistrodon 
acutus at the ectopterygoid junction. 

The shape of the posterior portion of the pterygoid, the curvature of the medial and 
lateral edges, the position of the ectopterygoid junction, and the size and shape of the ridge 
on the ventral surface of the pterygoid, all are quite characteristic of each species (figs. 29 
and 30). These characters are, however, difficult to classify or describe and thus are difficult 
to use in determining relationships. 


Pterygoid-quadrate graphs have been discussed. A graph of pterygoid length against 
ectopterygoid length is shown in figure 9. Generic differences are apparent. Tvrimeresurus 
wagleri conforms in this character with the rest of the genus. The lines for Sistrurus and 
Crotalus appear different when extrapolated. The range (solid line) of Sistrurus, however, 
conforms closely with the range (solid line) of Crotalus and falls within its scatter. The 
measurements for A gkistrodon contortrix fall within the scatter of the Old World A gkistrodon 
line. The line for A. piscivorus is distinct but close to that for A. bilineatus. The number of 
specimens available of Lachesis and arboreal species of Bothrops was too small to permit draw- 
ing any curves. 


Graphs of pterygoid length potted against lower jaw length show no specific or generic 
differences. All measurements of all species fall very close to a single line. 

Ectopterygoid (External pterygoid, transpalatine, transverse, transverse pterygoid) .— The 
ectopterygoid is a flat, thin bone extending from about the middle of the pterygoid, passing 
under the eye, and articulating with one or more points on the posterior wall of the maxilla. 
The ectopterygoid is usually forked anteriorly. In a few species the shape of this fork shows 
individual and ontogenetic variation, but in others the shape is quite diagnostic and is useful 
in determining relationships (figs. 29, 30). 

The lateral process of the fork of the ectopterygoid is quite round in Crotalus ruber, and 
this process turns ventrally in C. tortugensis. Crotalus triseriatus has a thin lateral flap on its 
ectopterygoid. The primitive shape of this bone is probably broad and flat, with a wide anterior 
end. This condition is seen in some Viperidae and in Trimeresurus wagleri, purpureomaculatus, 


albolabris, stejnegeri, and Bothrops schlegeli (figs. 29, 30). 


1964] 


IN MM. 


ECTOPTERYGOID LENGTH 


40 


30 


20 


BRATTSTROM: EVOLUTION OF Pir VIPERS 205 


20 30 


40 50 


PTERYGOID LENGTH IN MM. 


Fig. 9. Relationship of ectopterygoid length to pterygoid length in the crotalids. Solid 
line, observed range; dashed line, extrapolation; dots, arboreal group of Bothrops; solid squares, 
Lachesis. 


IN MM. 


FANG LENGTH 


30 


20 


30 


60 90 
LOWER JAW LENGTH IN MM. 


Fig. 10. Relationship of fang length to lower jaw length in the crotalids. Solid line, 
observed range; dashed line, extrapolation; dot, arboreal group of Bothrops; solid squares, 
Lachesis. 


206 SAN Disco Society oF NATURAL History {Vor13 


The posterior part of the ectopterygoid generally is narrower than the anterior part and 
usually is flat or twisted. The junction of the ectopterygoid with the pterygoid is either flat, 
in a groove in the pterygoid, or supported by lateral flaps of the ectopterygoid. The junction 
of the ectopterygoid with the pterygoid is usually characteristic for each species. The curvature, 
twisting, and posterior articulations of the ectopterygoid are difficult to interpret in terms of 
relationships. The shape of the anterior end of the bone is more useful in this regard. 


The ectopterygoid is shorter than the basal portion of the pterygoid in all species of 
Agkistrodon except acutus and strauchi; thus this character cannot be used to distinguish all 
species of Agkistrodon from other crotalids as Ruiz (1951) and W. E. Taylor (1895) 
proposed. 


Graphs of ectopterygoid length against lower jaw length show few distinctions except on 
the generic and subgeneric levels. 


Lower Jaw. — The lower jaw of crotalids (figs. 31, 32) contains four bones, the articular, 
angular(e), dentary, and splenial. The coronoid bone is absent in crotalids, as in most other 
snakes. Both angular and splenial are small and lie on the medial side of the lower jaw. The 
shape and relationship of these two bones and their position in regard to the internal Meckelian 
foramen are of some taxonomic importance. 


The tooth-bearing dentary bone is usually blunt anteriorly and is joined to the dentary of 
the opposite side by a ligament at the mid-line. The dentary has two posterior processes, a 
dorsal and a ventral, which fit above and below an elongate process of the articular. The 
splenial and angular lie medial to this junction. This dentary-articular junction is probably 
the weakest point in the crotalid skull, to judge from the frequency with which it is broken. 
The teeth on the dentary extend almost to the posterior tip of the dorsal surface of the dorsal 
process. The lower process of the dentary extends farther posteriorly than does the dorsal, 
except in a few species (A. hypnale, A. piscivorus, B. bilineatus, B. jararaca, B. schlegeli, 
figs. 31, 32; tables 10 and 12). The two processes are equal in length in A. acutus, in B. atrox, 
and in T. wagleri. 


The dentary has a small mental foramen on its external surface, the position and shape 
of which do not appear to be taxonomically significant. 


The articular is the longest bone in the lower jaw and probably represents the fused arti- 
cular, prearticular, and surangular (Gilmore, 1938; Williston, 1925). The posterior portion 
of the articular meets the quadrate in a dorsal saddle-shaped articulating surface. Posterior to 
this junction there is a heavy process which curves inward. The pterygoid articulates with the 
U-shaped area formed by this process. The ventral curvature of the articular appears to be a 
character of no taxonomic value, changing somewhat with growth. The articular may have a 
thin flap or process on its ventrolateral edge just below the point of articulation of the quad- 
rate, as in Trimeresurus wagleri and Crotalus atrox, mitchelli, molossus, ruber, and viridis 
(figs. 31, 32). This appears to be a specific characteristic in some species and a variable 
character in others (e.g., C. ruber and viridis). 


The posterior part of the articular has a thin high medial hump probably representing 
the embryonic surangular. This hump lies just anterior to the junction of the quadrate, and it 
forms the medial wall of the large posterior Meckelian foramen. The shape of this hump is 
characteristic for each species, but the differences are those of slight changes in curvature and 
hence are difficult to interpret for use in determining relationships. 


The posterior Meckelian foramen is open dorsally and laterally. Ventrolaterally, it is 
bordered by another part of the articular. The extent and shape of the opening of the foramen 
are specific characters. There seem to be trends in several phyletic lines of crotalids towards 
increase in the size of this opening and trends in other groups toward a reduction in its size. 


1964 | BRATTSTROM: EVOLUTION OF Pit VIPERS 207 


There is a small foramen in the lateral wall of the articular. This foramen is usually on 
the long, narrow part of the articular, but in some species (e.g., most species of Trimeresurus, 
B. schlegeli, C. intermedius; figs. 31, 32) it occurs within or below the area of the posterior 


dorsal hump. 


NEURAL SPINE W/H RATIO 


e) 50 100 150 200 
VERTEBRAL NUMBER 


0 50 100 ISO 200 
VERTEBRAL NUMBER 


Fig. 11. Relationship of size of neural spine to position within the vertebral column in 
two specimens of Crotalus durissus tzabcan (MVZ 45726 and 45727) from Yucatan. 


208 SAN Dieco Society oF NaTurAL History {Vot. 13 


Figure 10 is a graph of lower jaw length plotted against fang length. The lines for 
Bothrops and Trimeresurus are separate. The lines for T. wagleri and B. schlegeli differ from 
those for the other members of their respective genera. This is one of the few characters in 
which the Old World and New World species of A gkistrodon are different. With very little 
scatter, the measurements for A. contortrix, A. bilineatus, and A. piscivorus all approximate 
a single line. The Crotalus and Sistrurus lines are indistinguishable. The few measurements 
of S. ravus suggest a possible ontogenetic allomorphic change for these characters during 
growth. Nothing significant is revealed in graphs of measurements of several other characters 
of the lower jaw. 


V ertebrae. — A typical crotalid vertebra was described and figured by Klauber (1956) and 
is shown here in figure 2. The thoracic vertebrae of the Crotalidae each have a long hypapo- 
physis projecting ventrally from the centrum. The diapophyses are small tubercles. A small 
process extends anterior and ventral to the diapophysis. It is probably part of the diapophysis 
and is termed the parapophysis (Simpson, 1933; Brattstrom, 1954). The first two vertebrae 
are relatively simple, with the first lacking a neural spine. The hypapophysis divides into two 
distinct and separated parts posterior to the sacrum. These parts are called lymphapophyses. 


In general, the vertebrae of all the crotalids are similar, differing only in minor shapes 
and proportions. For example, the midthoracic vertebrae of Bothrops and Trimeresurus each 
have a well developed ridge extending from the posterolateral corner of the zygosphene to the 
posterior edge of the articulating surface of the prezygapophysis. This ridge is absent in the 
other genera. In Bothrops and Trimeresurus and in a few species of Agkistrodon, the lateral 
process of the prezygapophysis usually does not extend more than 1 millimeter beyond the 
articulating surface. There is a triangular depression on the anterior face of the prezygapophysis 
in Agkistrodon. This depression is lacking in Crotalus and Sistrurus except for the area imme- 
diately around the foramen and lateral to the centrum cup. There is a groove in the posterior 
edge of the neural spine in most species of Crotalus. This groove is usually absent in Sistrurus 
catenatus and miliarius or, if present, extends only halfway up the neural spine; vertebrae of 
S. ravus were not seen. 


The above-mentioned generic differences are relatively constant in most species, but they 
are not always valid in specialized or primitive species or in young individuals. It is important, 
however, to be able to identify the various species of crotalids by the vertebrae since they are 
the most common elements found in fossil deposits. Numerous subtle differences can be used 
to distinguish between the vertebrae of the different species, but these are often difficult to 
describe. The situation is complicated further by the fact that there are relative and propor- 
tional size differences among the vertebrae of a single individual. For example, the neural spine 
is tall and narrow on the anterior vertebrae of most crotalids. It increases in height in the first 
50 to 80 vertebrae and then decreases (fig. 11). At the point of maximum height, the neural 
spine is still increasing in width. Similar changes within the vertebral column can be noted for 
other characters (fig. 12). 


Several methods could be used for a quantitative differentiation between the vertebrae of 
different species of crotalids. One specific vertebra could be used as a standard for a species 
(e.g., vertebra number 80 or number 100). In fossil deposits, however, the vertebrae are 
usually separated, and it is impossible to determine which vertebra is number 80 or 100. A 
more useful standard would be to use the vertebra which has, for example, the neural spine 
height equal to the neural spine width. Fragmentation and weathering in fossils would require 
that several criteria be used. 

Where large series of vertebrae are available, it is sometimes useful to use ratio diagrams 
on a log-difference scale of a series of analogous measurements, comparing a sample with a 
known standard. This method, developed by Simpson (1941), was used by Brattstrom (1954) 
in distinguishing Pleistocene and Recent crotalids of Florida. Vertebrae can also be distin- 
guished by the use of simple proportions and ratios (Johnson, 1955b); see table 8. 


1964} BRATTSTROM: EVOLUTION OF Pit VIPERS 209 


TABLE 8 
Ratios of measurements of mid-thoracic vertebrae. 

W Post! Hv? HV3 

Species H NS H NS W Post 

C. adamanteus Fre eee Le 7 82 2.76 1.52 
C. atrox FO eee ie ree 1.69 pNeg 1.60 
CO ART IG IG restpese oer ent ARAN a A a 12 2.31 BA 1.61 
CRCETASLOS RE ere eee We eee 2.84 1.31 
COMA UTISSIEGEZAP CAN! pon is ee eee A a 1.41 2.42 eae 
C. enyo SPA te See BR a en ES Se 1.26 2.26 1.79 
(Ct 011 Gad 2 or cee Ne ARE 237, 3.00 1.26 
CPE TIED CLUE ee ete cet ear are Gains See ee Reece 2.38 2.46 1.03 
COLOSSUS eee ce ee acer — = 
C. pricet Ne Ne eetok Re Gar ae Oy EY _ 3.05 3.95 1.29 
(Ce Fag pate nish a tea ae cad ae eee 223 2.99 1.34 
TET (C1a Thief eR a” REOPEN OY ROE 217 3.10 1.43 
COROT aa eee a a oe OR ne eek 2 aa Se 1.68 2.81 1.67 
CR MCONCOLO Te oan ee ic AE Ee 2.85 3.39 1.19 
CMa Ids te re es ea etn ee pe ere 2.90 3.50 17 
Ceapallardita te Nat ee Ace Mee RECN BD) 3.66 0.97 
MCE LETIALILG ie St Me Ne Inthe sa RA EAE Sa hea ae 1.93 2.96 1.53 
SEM TPLIAT IC Seu at MO eee ys ee 2.70 3.80 1.41 
SIR GOTT TON Ags Clean Gch eRe RR Re mee oe 1.85 2.90 ey, 
AM PESCIV ONLESCMR i fe ict chp ARM GPE cy SRE seer 1.54 SD. jes 


1Width across postzygapophyses/height of neural spine. 
*Height of vertebra/height of neural spine. 
3Height of vertebra/width across postzygapophyses. 


Since differences occur along the vertebral column of a single snake, it was decided to 
graph measurements of every 10th or 20th vertebra for the length of the vertebral column. 
This was first done with two specimens of Crotalus durissus, using two characters (fig. 11), 
and then with a skeleton of Crotalus ruber, using several characters (fig. 12). The two graphs 
show the kinds of changes within the vertebral column of one snake but do not permit species 
comparisons because of size differences. 


To compare species, ratios of the various characters to height of the vertebrae were com- 
puted and graphed for C. ruber and durissus (figs. 13, 14) and for some 15 other species of 
crotalids. The position and crossing of the various lines seem to be characteristic for each 
species. These graphs also can be used for identifying vertebrae. For an unidentified vertebra, 
first the ratios of the measurements to the height of the vertebra are plotted on a card with 
the same scale as the graph. The card is then moved across each graph until a point is found 
where all the lines on the card conform with the lines on the graph. Conformity should occur 
at only one position on only one graph, thus indicating the serial position and the species of 
the unknown vertebra. This technique has proved successful, but, of course, it works only with 
species for which graphs are available. Furthermore, these graphs fail to show such important 
details as the curvature and shape of parts, which are characters of some importance in species 
determination. 


210 SAN Disco Society oF NAtuRAL History {Vot. 13 


fe) 20 40 60 80 100 120 140 160 180 


VERTEBRAL NUMBER 


Fig. 12. Relationship of several characters to position within the vertebral column in one 


specimen of Crotalus ruber ruber (BHB 1628). 

Characters: 1, width of vertebra; 2, length of vertebra; 3, height of neural spine; 4, length of neural 
spine; 5, width across prezygapophyses; 6, width across postzygapophyses; 7, hypapophysis length including 
centrum; 8, hypapophysis length; 9, height of zygosphene; 10, width of zygosphene; 11, height of vertebra. 


1964 } KRATTSTROM: EVOLUTION OF Pit VIPERS 211 


) 20 40 60 80 1 120 140 
VERTEBRAL NUMBER 


160 180 
Fig. 13. Relationship of characters to position within the vertebral column (expressed as 


ratios of vertebral height) in one specimen of Crotalus ruber ruber (BHB 1628). Characters 
as in figure 12. 


H/X RATIOS 


20 80 


VERTEBRAL NUMBER 


Fig. 14. Relationship of characters to position within the vertebral column (expressed as 


ratios of vertebral height) in one specimen of Crotalus durissus tzabcan (MVZ 45726). 
Characters as in figure 12. 


212 San Disco Society oF NATURAL History [Vot. 13 
TABLE 9 
Length /width! ratios of midthoracic ribs. 

Species Ratio Species Ratio Species Ratio 
A. bilineatus .....5....% 3.33 Gemcenasles: ea o40! IC. scutulatus.c ee OO 
A. contortrix .. speay a8.) Gacerastes ant ee 31 C. viridis helleri .... 3.98 
Ae COMLOTITIX. oo ocsea ns 3.61 C. durissus tzabcan...... 3.52 CC. viridis helleri ........ 4.65 
A. piscivorus One SHOT, C. durissus tzabcan...... 3.200 ~=C.._ viridis lutosus ........ 5.08 
GR GETONE oc 00 Crihormiaise cee 3.68  S. catenatus dake O78 
Cribadiscus esc: 2418 C. mitchelli pyrrbus .... 3.19 


1Width is depth of curvature, i.e., the maximum distance from the central part of the rib to the line 
connecting the ends. 


Measurements of crotalid vertebrae are taken in essentially the same manner as described 
by Johnson (1955b) or as shown in figure 3. A few measursements, however, need further 
elaboration. 


Height of vertebra. — Greatest height of the vertebra (HV), measured from the top of the neural 
spine to the tip of the hypapophysis, with the calipers flat along the top of the neural spine. 

Width of vertebra. — Width of the vertebra (WV) at its narrowest point between the prezygapo- 
physes and the postzygapophyses. 

Length of vertebra. — Length of the centrum, from the center of the cup anteriorly to the center of 
the ball posteriorly (LV). 

Width across prezygapophyses. — Greatest width across the prezygapophyses (W Pre. Z), including 
the tip of the process. 

Width of process of prezygapophysis. — Anteroposterior width of the articulating surface (= process) 
of the prezygapophysis (W P Pre). (Not the width of the small lateral process measured in “width 
across prezygapophyses.” ) 

Width across postzygapophyses. — Greatest width across the postzygapophyses (W Post). 


Width of process of postzygapophysis. — Anteroposterior width of the articulating surface of the 
postzygapophysis (W P Post). 

Centrum length. — See length of vertebra. 

Centrum height and width. — Height and width of the ball of the centrum (HC and WC). 

Neural spine height.— Height of the neural spine measured from the base of the neural spine 
(= zygosphene-zygantrum ridge) to the top of the spine at a point about half-way between its anterior 
and posterior edges (H NS). The height of the neural spine as measured at the zygosphene or at the 
zygantrum is often unreliable because of the variation in thicknesses of the zygantrum and zygosphene. 

Neural spine length. — Anteroposterior length of the neural spine at the top of the spine (L NS). 
(Erroneously referred to as “neural spine width” by Brattstrom, 1953, 1954.) 

Zygosphene height. — Height measured from the dorsal edge of the cup of the centrum to the top 
or shelf part of the zygosphene (HZ). 

Width of process of zygosphene. — Anteroposterior width of one side (= process) of the zygosphene 
WZ). 

Hypapophysis length. — Measured in two ways: first, from the top of the ball of the centrum to 
the ventral tip of the hypapophysis (LH); second, from the ventral base of the ball of the centrum to the 
tip of the hypapophysis (LH + C). The second measurement is distinguished by the term “including 
centrum.” 

Hypapophysis width. — Anteroposterior width of the middle of the hypapophysis (unless mentioned 
otherwise, e.g., Crotalus giganteus, Brattstrom, 1954). 

Parapophysis length. — Length of parapophysis (LP) from the base (junction with the anterior face 
of the prezygapophysis) io the tip. 

Parapophysis width. — Width (from side to side) at the middle of the parapophysis (WP). 


Ribs. — Ribs are found on all the presacral vertebrae of crotalids except the first two 
(atlas and axis). The first sacral rib is forked distally, with the ventral process longer than 
the dorsal. The second sacral rib is forked to the base. The third sacral rib has the dorsal part 


completely separate from the ventral. The dorsal part is apparently immovably articulated and 


1964 } BRATTSTROM: EVOLUTION OF Pir VIPERS 218 


ee, 


Qe Wea wae. OE 
_ ¢ a vy 


ADAMANTEUS ATROX BASILISCUS CERASTES DURISSUS 


ii 


<(ennoGi ae res 
aol geal £2 ea 
ag Seer: = i 
A < eS 


PRICEI RUBER SCUTULATUS TIGRIS 


cals 
27 


tS 


a 


=e 
va pal 


TORTUGENSIS TRISERIATUS V.HELLERI V.LUTOSUS 


U4 


V.OREGANUS V.OREGANUS SIERRA V.VIRIDIS WILLARD! 


“al an 
ga ga (as 


CATENATUS MILIARIUS RAVUS PUSILLUS 


14 


Fig. 15. Premaxillae and prefrontals of species of Crotalus and Sistrurus. 


the ventral movably articulated to the vertebra. In each of the next two vertebrae, evidently 
the first and second caudal vertebrae, both the dorsal and ventral processes are fused to the 
vertebra. In the third and fourth caudal vertebra, these processes are fused to each other. On 
each succeeding caudal vertebra they remain as thin, narrow, immovable lateral processes. These 
processes may serve as attachments for cloacal and scent-gland muscles. 


A typical crotalid rib is thin and elongate, with two dorsal heads. No major differences 
were noted in the articulating surfaces of the ribs in any of the crotalids. Some individuals 
seemed to have heavier heads on the ribs than others, but this condition is usually associated 
with the large size of the snake. 


214 San Dreco Society oF Natura History {Vor. 13 
ee Sar ae 
ya PP P 2 oe 


A.ACUTUS  BILINEATUS BLOMHOFF | CONTORTRIX HALYS HYPNALE 
V A Jn age 
a DP ge & Fe 
A. PISCIVORUS RHODOSTOMA B.ALTERNATA ATROX BILINEATUS COTIARA 
re A a ene ays 
ee eee vv ED aa fA 
B.GODMANI JARARACA JARARACUSSU LANCEOLATUS NASUTA NEUWIEDI 


pai 9 ea 


B. NUMMIFER SCHLEGEL! L.MUTA T. ALBOLABRIS 


T. FLAVIVIRIDIS PURPUREOMACULATUS WAGLERI 


Fig. 16. Premaxillae and prefrontals of species of Agkistrodon, Bothrops, Lachesis, and 


Trimeresurus. 


Ribs vary in length and curvature within one snake. The curvature of the mid-thoracic 
ribs often reveals (or results from) the general habitus of the snake; for example, Crotalus 
durissus, with a high body, has long ribs with slight curvature, whereas C. basiliscus, with a 
low, flat body, has more curvature to the ribs. Length-width ratios of midthoracic ribs of a few 
crotalids are presented in table 9. Width in this case means curvature (= depth of arc) and 
is measured from the lateral curvature of the rib to a line connecting the ends. Some differ- 
ences between the ribs of several species (e.g., C. basiliscus and durissus, as just mentioned) 
can be seen from table 9, but also note the variation in two specimens of durissus and in two 
specimens of C. y. helleri. Ribs were not used as a taxonomic character in determining relation- 
ships because of their ontogenetic, serial, and individual variation. 


Shaker. — The shaker or style of the rattlesnakes, Crotalus and Sistrurus, represents the 
fused terminal 9th to 11th caudal vertebrae. The muscles that vibrate the rattle are inserted on 
the base of the style. The anatomy of the shaker was described by Klauber (1940, 1956) and 
by Zimmermann and Pope (1948). 


No major differences could be found in the shaker of the various species of rattlesnakes 
examined, and no differences could be found to distinguish the shakers of Crotalus and 


1964} BRATTSTROM: EVOLUTION OF Pit VIPERS 215 


) a Poo , 
ey NS 
Ooo p b OD 
ae Ze ee 


DURISSUS PUSILLUS ENYO HORRIDUS INTERMEDIUS 


LEPIDUS MITCHELLI MOLOSSUS PRICE! RUBER 


SCUTULATUS TIGRIS TORTUGENSIS TRISERIATUS WILLARDI 


pyar ee al iy 
Ds : 


VIRIDIS HELLER! VLUTOSUS VOREGANUS  V. VIRIDIS 


‘e 0 D 
= eae. 


CATENATUS MILIARIUS RAVUS 
Fig. 17. Nasals and palatines of species of Crotalus and Sistrurus. 


Sistrurus except for the temporal difference noted by Zimmerman and Pope (1948). Length 
of the shaker was plotted against width and against height. There were no consistent differences 
or similarities among the various species. The variation in one species was often as great as 
the entire range for the two genera. There was a general increase in size of the shaker with 
increase in size of the snake, but with no consistency in ratio. The variability found is probably 
related to the fact that the shaker represents many centers of ossification plus extravertebral 
additions. There may be differences in the number of vertebrae in the shakers of various 
species, but because of the method of development and the fusion of parts, the number is 
almost impossible to determine. 


216 San Disco Society oF NATURAL History [ VoL. 13 


! : ED) a 
A.ACUTUS BILINEATUS BLOMHOFF! CONTORTRIX HALYS 
A. HYPNALE pisciVORUS RHODOSTOMA STRAUCHI  B. ATROX 
B. BILINEATUS COTIARA GODMANI JARARACA JARARACUSSU 

B.LANCEOLATUS NEUWIEDI NUMMIFER SCHLEGEL] L.MUTA 


T. ALBOLABRIS FLAVIVIRIDIS PUNICEUS 


T. PURPUREOMACULATUS WAGLERI 


Fig. 18. Nasals and palatines of species of Agkistrodon, Bothrops, Lachesis, and 


Trimeresurus. 


1964 } BRATTSTROM: EVOLUTION OF Pit VIPERS 217 


Po, ie a Sh 


ADAMANTEUS — ATROX BASILISCUS CERASTES DURISSUS 
ENYO HORRIDUS LEPIDUS MITCHELLI MOLOSSUS 
PRICE! RUBER SCUTULATUS TIGRIS TORTUGENSIS 


Flea. (ego icn eile we 


TRISERIATUS VIRIDIS DECOLOR VIRIDIS HELLERI  VIRIDIS LUTOSUS 


im me [f 


VIRIDIS OREGANUS VIRIDIS OREGANUS SIERRA VIRIDIS VIRIDIS 
WILLARDI CATENATUS MILIARIUS RAVUS PUSILLUS 


Fig. 19. Frontals of species of Crotalus and Sistrurus. 


PHYLOGENY 


Several workers have suggested phylogenies for the Crotalidae. Among the first of these 
was Garman (1889). Recent contributions to knowledge of relationships within the family 
were made by Amaral (1929), Githens and Butz (1929), Githens and George (1931), 
Githens (1935), Mosauer (1932, 1935), Maslin (1942), and Minton (1956). Phylogenetic 
trees were presented for Crotalus and Sistrurus by Amaral (1929), Githens and George 
(1931), Gloyd (1940), H. M. Smith (1946), and Klauber (1956). 


Fossil crotalids contribute little to our knowledge of relationships within the family. A 
detailed report on the fossil crotalids was made by Brattstrom (1954) and summarized by 
Klauber (1956). 


Except for scutellation, non-osteological data are lacking or contribute little to our knowl- 
edge of relationships within the family. Data concerning the biochemistry and physiological 
action of crotalid venoms (Minton, 1956) contribute supportive evidence for some relation- 
ships, however. Minton (1956 and personal communication) cautioned against using venom 
data for determining definite relationships and suggested that they be used in conjunction with 
other data. 


218 SAN Disco Society of NaturAL History { VoL. 13 


Paar oe 


A.ACUTUS  BILINEATUS BLOMHOFF! CONTORTRIX HALYS HYPNALE 


ee irri 


A. PISCIVORUS RHODOSTOMA  STRAUCHI B.ALTERNATA ATROX BILINEATUS 


em meet ieee igeleen 2c 


B. COTIARA GODMANI JARARACA JARARACUSSU LANCEOLATUS NASUTA 


P(E tee eae ay 


B.NEUWIEDI NUMMIFER SCHLEGELI L.MUTA  T. ALBOLABRIS FLAVIVIRIDIS 


(a ies Cs ees een 


T. MUCROSQUAMATUS  PUNICEUS PURPUREOMACUL ATUS STEJNEGERI WAGLERI 


Fig. 20. Frontals of species of A gkistrodon, Bothrops, Lachesis, and Trimeresurus. 


Mosauer (1932, 1935) studied the trunk muscles of several snakes, including Crotalus 
horridus, Agkistrodon mokasen, A. piscivorus, Lachesis muta, and Sistrurus catenatus. He 
found no major differences in trunk musculature among the crotalids examined. No one has 
studied the head muscles of crotalids as Haas (1952) did for viperids. 

Scale characters in reptiles are one of the most important criteria used by modern 
herpetologists to determine relationships. In addition, Picado (1931) showed the usefulness 
of epidermal micro-ornaments in determining relationships in snakes, but no one has carried 
his investigations to other species and genera. On the basis of scale characters, relationships of 
crotalids were suggested by Amaral (1929), Klauber (1936a, 1952, 1956), Gloyd (1940), 
Gloyd and Conant (1943), and H. M. Smith (1946). 

Though vibration of the tail occurs in many snakes, the rattle is unique in Sistrurus and 
Crotalus. A structure resembling an incipient rattle is found in the caudal spine and four rows 
of subcaudal scales in Lachesis muta. The rattle was discussed in great detail by Klauber 
(1940) and by Zimmermann and Pope (1948) and was used by Klauber (1956) in deter- 
mining relationships. Probably the rattle was evolved only once (in a proto-Crotalus-Sistrurus) 
primarily as a defense against being stepped upon by large hoofed mammals, as suggested by 
Klauber (1940, 1956). 

Color and color pattern are occasionally useful in determining relationships between 
species. This is especially true with many rattlesnakes. For example, the evolution of color 
pattern in the Sistrurus ravus-Crotalus triseriatus complex was discussed in detail by Gloyd 
(1940:244). Klauber (1956) also took color and pattern into account in the construction 
of his phylogenetic chart. 


1964 } BRATTSTROM: EVOLUTION OF Pit VIPERS 219 


IMG SAS 


ADAMANTEUS ATROX BASILISCUS CERASTES  purRissus  ENYO 


Jd 


HORRIDUS LEPIDUS MITCHELLI — MOLOSSUS PRICE! 
RUBER PUSILLUS SCUTULATUS TIGRIS. = TORTUGENSIS 
VIRIDIS WILLARD! CATENATUS RAVUS 


Fig. 21. Maxillae of species of Crotalus and Sistrurus. 


The few interspecific crotalid hybrids (Crotalus ruber x C. viridis hellert, C. durissus 
unicolor x C. scutulatus, C. horridus atricaudatus x C. adamanteus, C. viridis oreganus x C. 
scutulatus; Klauber, 1956) do not add significantly to knowledge of relationships within the 
family. They indicate only that under rare circumstances in nature and in captivity isolating 
mechanisms may break down (or may never have developed, as in the case of C. d. unicolor 
and C. scutulatus) and hybridization will occur. One intergeneric hybrid (C. h. horridus x S. c. 
catenatus) reported by Bailey (1942) is of interest in further indicating the close relationship 
between Crotalus and Sistrurus. 

METHODS 

All phylogenies are somewhat subjective, as they indicate the worker’s concept of relation- 
ship within a group. In this paper, the various characteristics of each species of crotalid are 
compared in order to determine relationships as accurately as possible. This comparison is 
made with drawings of bones (figs. 15 to 32) and a tabulation of characters (tables 10 to 12) 
for the species available. In order to determine the relationships among members of each 
genus, these tables are summarized in a series of additional tables (13 to 16), which give the 
number and percentage of characters that each species has in common with each other member 
of its genus. The following discussion of the phylogeny of the pit vipers is based primarily on 
the osteology and hence on the similarities and differences shown in tables 13 to 16. This basis 
is taken for two reasons: first, osteological characters are generally regarded as the most basic 
or least changeable and hence as more likely to indicate relationships than are features of the 
external morphology; and second, extensive data for other characters are not available except 
in Crotalus and Sistrurus. Data concerning venoms, hemipenes, scutellation, etc., have been 
used when available. 


220 SAN Dreco Society oF NaturaAt History { Vox. 13 


33 dD FB OSA 


A.ACUTUS BILINEATUS BLOMHOFF! CONTORTRIX HYPNALE PISCIVORUS 


SYS IAS 


B.ALTERNATA ATROX  BILINEATUS COTIARA GODMAN! JARARACA 


mE 


B. JARARACUSSU LANCEOLATUS NEUWIED! NUMMIFER SCHLEGEL! L.MUTA 


y 3 


T. ALBOLABRIS FLAVIVIRIDIS T. PUNICEUS PURPUREOMACUL ATUS 
STEJNEGERI WAGLERI 


Fig. 22. Maxillae of species of Agkistrodon, Bothrops, Lachesis, and Trimeresurus. 


At one time in the analysis of the phylogeny of crotalids, I considered weighting charac- 
ters, giving more weight to those that are more stable intraspecifically. In approximate order 
of decreasing consistency and hence of decreasing usefulness in determining relationships, the 
various bones are: frontal, parietal, postfrontal, maxillary (especially the pit cavity), basis- 
phenoid-basioccipital complex, prefrontals, squamosals, palatine, pterygoid, ectopterygoid, lower 
jaw, premaxilla, nasals, vertebrae, shaker, and ribs. But the weighting of characters is in itself 
subjective unless an analysis of variability is made for each character. I have therefore followed 
the procedure outlined above, comparing percentages of unweighted characters. 


The terms “group” and “subgroup” denote only groups of related species, and the terms 
are not to be taken as equal in the different genera. 


Most of the characters listed in tables 10 to 12 are sufficiently consistent to be useful in 
a key to skulls. Several keys to crotalids and crotalid skulls were made at the request of L. M. 
Klauber for his book (Klauber, 1956). These keys are used most successfully on adult skulls 


and in conjunction with the figures and tables presented here. 


1964 } BRATTSTROM: EVOLUTION OF Pit VIPERS 


i) 
i) 
—_ 


ADAMANTEUS ATROX BASILISCUS 


CERASTES 


HORRIDUS LEPIDUS 

DURISSUS 
MITCHELLI MOLOSSUS PRICEI RUBER 
SCUTULATUS TIGRIS TOR TUGENSIS TRISERIATUS 

V. DECOLOR V. HELLERI V. HELLERI Vv. LUTOSUS 

Old 
V. OREGANUS Vv. VIRIDIS WILLARDI CATENATUS 
MILIARIUS RAVUS PUSILLUS 


Fig. 23. Parietals and postfrontals of species of Crotalus and Sistrurus. 


222 SAN Disco Society oF NAturAL History [Vor 13 


INTERGENERIC RELATIONSHIPS 


The Solenoglypha, comprising the Crotalidae and the Viperidae, is characterized by the 
presence of movable fangs. Various osteological and myological studies (Haas, 1952; Johnson, 
1955b, 1956) indicate that the Viperidae is more primitive than the Crotalidae. The Crotalidae 
also seems to be more specialized than the Viperidae in having the heat-sensitive loreal pit. The 
problems of the origin of the Solenoglypha (Boulenger, 1896; Hewitt, 1911; Mehely, 1911; 
Radovanovic, 1935; Haas, 1938, 1952) and of the differentiation of the two families, however, 
are not within the scope of this paper. 


The most primitive genus of crotalids supposedly would have more characters in common 
with the more primitive Viperidae than would other crotalids. This genus would be the least 
divergent from its ancestor. Trimeresurus and Agkistrodon have more viperid characters than 
do Bothrops, Lachesis, Sistrurus and Crotalus. The pterygoid teeth extend beyond the junction 
of the ectopterygoid in A gkistrodon, Bothrops, and Trimeresurus, as they do in the Viperidae. 
The palatine in T. wagleri and in some species of Agkistrodon is short and stubby and similar 
to that in the Viperidae. The anterior end of the ectopterygoid is viperid-like in most species 
cf Trimeresurus. The species of Trimeresurus also have relatively small loreal pit cavities — 
seemingly a primitive character. The members of the genus A gkistrodon all have large head 
plates, as do the primitive members of the Viperidae. Sistrurus also has large head plates, but 
most of its osteological characters and the presence of a rattle indicate that it is a more 
specialized genus. Minton (1956) noted that the venom of Agkistrodon is that of a relatively 
primitive crotalid stock. These characters all suggest that Agkistrodon and Trimeresurus are 
the most primitive genera in the Crotalidae. To judge from these and from other characters 
discussed below, T. wagleri and the small species of A gkistrodon (contortrix, halys, hypnale, 
blomhoffi, etc.) are probably the most primitive members of their genera and hence the most 
primitive members of the family. 


Figure 33 is my concept of the relationships of the genera within the Crotalidae. This 
chart is based primarily on osteology. Superimposed on this phylogenetic tree are two major 
external characters: type of head scales and presence or absence of a rattle. There are large 
head plates in Agkistrodon and Sistrurus and small scales on the top of the head in Bothrops, 
Trimeresurus, Lachesis, and Crotalus. The head scales are keeled in Bothrops, Lachesis, and 
T. wagleri. The head scales anterior to the parietal region are not keeled in the remaining 
species of Trimeresurus. 


Lachesis. — The relationship of the bushmaster, Lachesis muta, has been uncertain. It has 
been included in the same genus (Lachesis) with all of the species currently in Bothrops 
(Brazil, 1914). The species of Trimeresurus, Lachesis, and Bothrops have all been included 
in the same genus, or the species have been variously separated into the three genera. Most 
modern herpetologists have considered Lachesis a distinct genus but have still thought its rela- 
tionships to be with Bothrops. As Garman (1889) first pointed out and as Ruiz (1951) also 
hinted, however, Lachesis muta is much closer to the rattlesnakes than to Bothrops. This 
similarity is especially evident in the osteology, wherein the majority of the characteristics of 
Lachesis are Crotalus-like. Notably, the pterygoid teeth do not extend beyond the middle of 
the junction of the ectopterygoid in Lachesis, Crotalus, and Sistrurus, and these forms are 
similar in having a knob-like process in the curvature of the pit-cavity. Other similarities are 
in the shapes of the prefrontals, parietal, squamosals, basioccipital, palatines, pterygoids, and 
ectopterygoids (tables 10, 11). The subcaudal scutes of Lachesis are in four rows posteriorly, 
in contrast to the one or two rows in all other crotalids. These scutes and the horny terminal 
spine probably resemble those of the Sistrurus-Crotalus ancestor before the rattle was perfected. 
The only major similarity of Lachesis and Bothrops is in their large size, but the subtropical 
Crotalus durissus is also large. The large size of these forms is in accordance with Bergmann’s 
rule for ectotherms in the tropics, as discussed by Cowles (1945). The large size of the scales 


1964 } BRATTSTROM: EvoLUTION OF Pit VIPERS 223 


WY 


J 


BILINEATUS CONTORTRIX 
A. ACUTUS BLOMHOFFI 
ee HYPNALE . 
PISCIVORUS 
LS es RHODOSTOMA 
B. ALTERNATA 
BILIN re) 
anne EATUS COTIARA 
INES | 
JARARACA LANCEOLATUS 
B. GOOMANI JARARACUSSU 
B. NASUTA NEUWIEDI NUMMIFER SCHLEGELI 
| | | MUCROSQUAMATUS 
L. MUTA IRIDIS 
T. ALBOLABRIS Sa) 
T. PUNICEUS ; i WAGLERI 
PURPUREOMACUL ATUS STE JUNEGERI 


Fig. 24. Parietals and postfrontals of species of A gkistrodon, Bothrops, Lachesis, and 


Trimeresurus. 


on the body of Lachesis, C. durissus, and the large species of Bothrops, may well be an example 
of convergence; it may be an adaptation of scale size in tropical regions for a primitive insulat- 
ing mechanism for trapping and retaining non-movable air close to the body, as suggested by 


observations by Cowles (1958 and MS). 


[Vot. 13 


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Ol ATaV.L 


225 


EvoLuTION OF Pir VIPERS 


BRATTSTROM 


1964} 


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226 San Dteco Society of NaturAL History [Vot. 13 


ADAMANTEUS ATROX BASILISCUS CERASTES 
DURISSUS ENYO HORRIDUS LEPIDUS 
il aera ee 
MITCHELL! MOLOSSUS PRICE| RUBER 
SGUGUIEATUS TIGRIS TORTUGENSIS TRISERIATUS 


VIRIDIS DECOLOR VIRIDIS HELLERI VIRIDIS OREGANUS WASH.-IDA. 


VIRIDIS OREGANUS SIERRA VIRIDIS VIRIDIS WILLARDI 
CATENATUS MILIARIUS RAVUS PUSIIEEUS 


Fig. 25. Basioccipitals and basisphenoids of species of Crotalus and Sistrurus. The bones 
are drawn upside down and as viewed from the side. 


Crotalus and Sistrurus.— As has just been stated, Crotalus, Sistrurus, and Lachesis are 
probably closely related, to judge by their osteology and caudal scutellation. A gkistrodon seems 
a more likely ancestor of a Crotalus-Sistrurus-Lachesis stock than does either Bothrops or 
Trimeresurus. In Agkistrodon, Lachesis, Crotalus, and Sistrurus, the greatest breadth of the 
head is due to the width of the parietal plus the poison glands. In Trimeresurus and Bothrops, 


1964 } BRATTSTROM: EVOLUTION OF Pit VIPERS 227 


A. ACUTUS BILINEATUS BLOMHOFFI CONTORTRIX 
A.HALYS HYPNALE PISCIVORUS B. ALTERNATA 
B. ATROX COTIARA GODMANI JARARACA 
B. JARARACUSSU LANCEOLATUS NASUTA NEUWIEDI 
B.NUMMIFER SCHLEGEL L.MUTA T. ALBOLABRIS 
T.FLAVIVIRIDIS PUNICEUS PURPUREOMACULATUS 
T, WAGLERI STEJNEGERI 


Fig. 26. Basioccipitals and basisphenoids of species of Agkistrodon, Bothrops, Lachesis, 


and Trimeresurus. The bones are drawn upside down and as viewed from the side. 


however, the greatest breadth of the head is due to the extremely elongate quadrate and squa- 
mosal. This gives a pronounced triangular shape to the head. A gkistrodon has large head plates, 
as does Sistrurus. If Sistrurus and the triseriatus group of rattlesnakes are considered the most 
primitive members of the Crotalus-Sistrurus stock (see below), then one of the small species 
of Agkistrodon (contortrix, hypnale, halys, blomhoffi) seems the most likely ancestor of this 
stock. This is suggested by the shape of the frontal, parietal, and basisphenoid (tables 10 
to 12). The durissus group might instead be considered the most primitive member of the 
Crotalus-Sistrurus stock; then the Agkistrodon acutus-bilineatus-piscivorus stock would be the 
most likely ancestor of the Crotalus-Sistrurus-Lachesis line. The latter relationship is suggested 
by similarity in the shape of the parietal, the prefrontal processes, the frontal, and the basis- 
phenoid process (tables 10 to 12) in Lachesis, Crotalus durissus, and the Agkistrodon acutus- 
bilineatus-piscivorus group. The first relationship seems more likely, though neither can be 
proven as yet. It is apparent from the above and from figure 33 that, regardless of the rela- 
tionships, small head scales have evolved several times in crotalids —in a proto-Bothrops- 
Trimeresurus, in Lachesis, and in Crotalus. 


228 San Dteco Society oF Naturat History { Vox. 13 


A 


ADAMANTEUS ATROX BASILISCUS CERASTES LATEROREPENS 
OLD YOUNG i ¢ 
G GERASHTES DURISSUS ENYO HORRIDUS 
LEPIDUS MITCHELL! MOLOSSUS PRICE! RUBER 


ae ile: ae aa 


SCUTULATUS TIGRIS TORTUGENSIS TRISERIATUS  VIRIDIS DECOLOR 


fo BE Li 


WASH.- IDA. SIERRA 
V. HELLERI V EUTOSUS V. OREGANUS V. VIRIDIS 
WILLARD! CATENATUS MILIARIUS RAVUS PUSILLUS 


Fig. 27. Squamosals of species of Crotalus and Sistrurus. 


Bothrops and Trimeresurus. — A major problem in the taxonomy of the Crotalidae is the 
question of the distinctness of the genera Bothrops and Trimeresurus. Both genera are tropical, 
the former in the New World, the latter in the Old, and many species in the two genera have 
similar colorations. No extensive work has been done on the taxonomy of the two genera 
except that of Maslin (1942) and Burger (MS). The two genera are quite distinct osteologi- 
cally, though this distinctness is based on a combination of several characters. There are two 
relatively exclusive characters. First, the curvature of the pit cavity bears a small process in 
Trimeresurus, whereas it is smooth in Bothrops, except in B. schlegeli, in which a process is 
slightly developed. Second, the palatine is entire in Trimeresurus and forked in Bothrops, 
except in B. schlegeli and B. nummifer. The major problem in distinguishing the two genera 
lies in certain similarities of these two species of Bothrops and Trimeresurus wagleri in the 


1964 } BRATTSTROM: EVOLUTION OF Pit VIPERS 229 


ee 


A.ACUTUS  BILINEATUS BLOMHOFF| CONTORTRIX HALYS HYPNALE 


na Sg ae ei 


A. PISCIVORUS RHODOSTOMA STRAUCHI B. ALTERNATA ATROX 
B. BILINEATUS COTIARA GODMANI JARARACA JARARACUSSU 
B.LANCEOLATUS NEUWIEDI NUMMIFER SCHLEGEL | L.MUTA 


ROWSE ESE 


T. ALBOLABRIS FLAVIVIRIDIS PUNICEUS PURPUREOMACULATUS WAGLERI 


Fig. 28. Squamosals of species of A gkistrodon, Bothrops, Lachesis, and Trimeresurus. 


shapes of the prefrontal, frontal, parietal, and postfrontal, and, according to Maslin (1942), 
in hemipenes and in scutellation. Trimeresurus wagleri is distinct from most species of Trimere- 
surus in its wide and depressed frontals, in its scutellation (Maslin, 1942), and in its neuro- 
toxic venom (M.A. Smith, 1931; Maslin, 1942). It differs from B. schlegeli and B. nummifer 
in its elongate prefrontal, its low palatine, and its triangular basisphenoid. It differs from all 
other species of Bothrops in having a process in the curvature of the pit cavity. Perhaps most 
importantly, it differs from all other crotalids in having the lower lumen of the fang medial 
rather than anterior and in having depressed frontals. Trimeresurus wagleri thus can be dis- 
tinguished from the small species of Bothrops as well as from the other species of Bothrops 
and Trimeresurus. The other species of the two genera are easily separable on the basis of 
the two major characters mentioned above. As will be discussed below, the distinctness of 
T. wagleri and of the small Bothrops will be recognized by placing them in separate subgenera 
within their respective genera. 


PHYLOGENY OF AGKISTRODON 


Little has been said in the literature concerning the relationships of the species of A gkis- 
trodon. Nevertheless, many workers have been concerned over the occurrence of the same 
genus in North America and in Asia. Some have supposed that the two groups were not in 
the same genus, although no evidence supporting this supposition has been presented. The 
present study shows that there are no major osteological characters to distinguish all Old 


World from all New World species of Agkistrodon. Most of the characters shared by Old 


230 SAN Dreco Society oF NaAturRAL History { Vou. 13 


5000099 
ADAMANTEUS ATROX BASILISCUS 


ke 


CERASTES DURISSUS ENYO 


HORRIDUS INTERMEDIUS LEPIDUS MITCHELL 

“= 609000@) 

Zz L Vi me) LT eae ( 

MOLOSSUS PRICE! RUBER SCUTULATUS 

eooe 0o00cSg @o00° 

fx LTA: = 

TIGRIS TORTUGENSIS = TRISERIATUS VIRIDIS 

WILLARDI CATENATUS RAVUS PUSILLUS 


Fig. 29. Pterygoids and ectopterygoids of species of Crotalus and Sistrurus. 


The pterygoids are shown in ventral view, the ectopterygoids in dorsal view (anterior ends). The 
junction of the two bones is indicated in the drawing of the pterygoid. Teeth are indicated by circles, the 
mean number for each species being shown. 


World forms or by New World forms are also shared by other members of the genus. 
A gkistrodon contortrix and the piscivorus-bilineatus group show closer osteological relationships 
to various Asiatic forms than they do to each other (table 13). 


1964 } BRATTSTROM: EVOLUTION OF Pir VIPERS 231 


SETS SED 


A.ACUTUS BILINE AT US BLOMHOFFI 


A.CONTORTRIX HALYS HYPNALE 


—S © 606006 9 8009 


ao 2° S0CS0SGc0 


PISCIVORUS 


Via 
ATROX = 


Sedo000028 


A.RHODOS TOMA B. ALTERNATA 


coo 
—soooool OF S© 3 
° 
o 


000000005 
img ae 
= aS 
B.BILINEATUS COTIARA GODMANI 


VS 


LANCEOLATUS 
B.JARARACA JARARACUSSU 
<a 060808 lo 
B.NEUWIEDI NUMMIFER 


FLAVIVIRIDIS 


f- coo Cor: ) 
4 


PUNICEUS PURPURE OMACULATUS STEJNEGERI WAGLERI 


Fig. 30. Pterygoids and ectopterygoids of species of A gkistrodon, Bothrops, Lachesis, and 


Trimeresurus. See caption of figure 29. 


SAN Dirco Society of NATURAL History | Vor. 13 


232 


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Il ATaVvL 


233 


EvoLUTION OF Pit VIPERS 


BRATTSTROM 


1964] 


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| winipaut 
| 
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| | 
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(penunucy) 1 ATAV.L 


234 SAN Drieco Society oF NaturaAt History {[Vorsd3 


ADAMANTEUS AT ROX | BASILISCUS 


CERASTES : DURISSUS ENYO 


HORRIDUS INTERMEDIUS LEPIDUS 


MITCHELL! MOLOSSUS PRICE] 


= Wid 
RUBER SCUTULATUS STEJNEGERI 
TIGRIS Sy: 


TORTUGENSIS TRISERIATUS 
VIRIDIS ene 
CATENATUS 


eee is th EF pass 


Fig. 31. Lower jaws of species of Crotalus and Sistrurus. 
Teeth are indicated by simple lines, the mean number for the species being shown. The posterior 
extents of the dorsal and ventral posterior processes of the dentary are indicated by heavy lines. 


Osteologically, the genus Agkistrodon is divisable into two main groups (fig. 34). One 
of these groups includes the Asian acutus and the North American bilineatus and piscivorus. 
The species in this group have 13 of 26 characters in common, including square frontals, 
T-shaped parietals, broader anterior ends to the ectopterygoids, and high basisphenoid processes 
(table 10). Within this group, bilineatus and piscivorus are closely related, being similar in 
most of the characters studied (tables 10, 13). Agkistrodon bilineatus is similar to contortrix 
in a few minor characters such as the ratio of the lower jaw length to quadrate length. Accord- 
ing to Minton (1956), bilineatus is intermediate in several venom characters between contortrix 
and piscivorus. Gloyd and Conant (1943) suggested that bilineatus might be closely related 
to mokeson (=contortrix). From their Table I, it is apparent that bilineatus is similar to 
mokeson in scale rows and caudal scales. However, bilineatus is similar to piscivorus in the 
number of ventrals and infralabials. All osteological evidence indicates a close relationship of 
bilineatus and piscivorus. Agkistrodon acutus has several peculiar characteristics, such as a 
pointed premaxilla, an ectopterygoid process, and an elongate prefrontal. It still has, however, 
several characters that indicate relationship to bilineatus and piscivorus, such as a square frontal 


and a T-shaped parietal (table 10). A gkistrodon acutus is one of the few ovoviviparous 
crotalids (Pope, 1935). 


1964 } BRATTSTROM: EVOLUTION OF Pir VIPERS 235 


———— A. ACUTUS BILINEATUS BLOMHOFFI 
ee ae ts ST 
aa HALYS HYPNALE 
A. CONTORTRIX 


Ce ZF T “strauchi 
RHODOSTOMA 
A. PISCIVORUS 

= BILINEATUS 


ATROX 


ie ALTERNATA ; 
JARARACUSSU 
JARARACA 
B. COTIARA 
Kea, NEUWIEDI 
NASUTA 
B. LANCEOLATUS 


es 
B. NUMMIFER SCHLEGEL! L. MUTA 
PURPUREOMACULATUS 
FLAVIVIRIDIS 
T. ALBOL ABRIS 
T. STEJNEGERI WAGLERI 


Fig. 32. Lower jaws of species of Agkistrodon, Bothrops, Lachesis, and Trimeresurus. See 
explanation under figure 31. 


The second group of closely related species includes the New World contortrix and the 
remaining Asiatic species (hypnale, halys, blomhoffi, rhodostoma, etc.; fig. 34). The forms 
within this group have 8 of 14 characters in common (57.1 per cent). The most important of 
these characters are the elongate frontal, the triangular parietal, and the low basisphenoid. 
A gkistrodon hypnale, with its trapezoidal nasal and elongate parietal, seems the most divergent 
member of this group. Agkistrodon blomhoffi is similar in many respects to halys. The 
esteology of rhodostoma is known only from a drawing in M. A. Smith (1943: 498, fig. 158); 
rhodostoma appears to be quite specialized, but certain of its characteristics point to a relation- 
ship with the hypnale stock. It has a large number of characters in common with hypnale and 
strauchi. Agkistrodon contortrix is more closely related to the Old World forms such as halys 
and hypnale (fig. 34, tables 10, 13) than it is to either bilineatus or piscivorus of the New 
World. The relationship of strauchi to hypnale, of nepa to hypnale, and of monticola and 
himalayanus to strauchi (fig. 34) is suggested by comments on the external morphology of 
these forms by Pope (1935) and by M. A. Smith (1943). The relationship of strauchi to 
hypnale is also supported by osteological evidence. The phylogenetic positions of the rare 
annamensis and millardi are at present unknown. They may belong in the hypnale-halys sub- 


group (fig. 34). 


Regardless of where the center of origin of the genus Agkistrodon was (though it was 
probably Asia), there are members of both major groups of the genus in the Old and New 
Worlds (fig. 34). 


{Vot. 13 


SAN Disco Society oF NAturRAL History 


236 


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238 San Disco Society oF NATURAL History { Vou. 13 


CROTALUS 


ele 
- 
- 
a= 
=o5 
-- 


SISTRURUS 


TERRESTRIAL 
BOTHROPS 


ARBORE AL 
BOTHROPS 


-- 
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-- 


RATT 
l 
~ Es 


PRES 
E 
ne NT 


AGKISTRODON Rarry @ —— ae 
Ss 


T. (TRIMERESURUS) 
A8Sey7 


~e-ene 


T. (TROPIDOLAEMUS) 
TO VIPERIDAE 


¢ 
got* ¢ Z 
yer se" sf 
\o o ee 
yah - oh 
Bom er? 


Fig. 33. Suggested phylogeny of the Crotalidae. 


PHYLOGENY OF CROTALUS AND SISTRURUS 


The genus Crotalus, as here defined (fig. 36), includes five main groups: durissus, atrox, 
viridis, triseriatus, and Sistrurus. These groups and the species within them have been recog- 
nized on the basis of external morphology (Gloyd, 1940; Klauber, 1956; H. M. Smith, 1946; 
Amaral, 1929). The relationships of the forms in the friseriatus group have been the least 
known. One of the groups, Sistrurus, is usually considered a separate genus. The Sistrurus 
group is similar in many of its characters (Amaral, 1929; Gloyd, 1940) to the friseriatus 
group. In my opinion, the characters that distinguish Crotalus and Sistrurus break down within 
S. ravus, C. pusillus, and C. intermedius. In general, the differences between Crotalus and 
Sistrurus are of the same type and order of magnitude as those between any of the groups of 


Crotalus (table 14). 


The nine large plates on the top of the head, supposedly characteristic of Sistrurus, are 
broken up into smaller scales in a fair number of the specimens of S. ravus (Gloyd, 1940: 
241-242). Klauber (1956:175) noted that the parietal scale often splits transversely in 
S. ravus. Fragmentation of the head plates also occurs as a rarity in S. catenatus and S. 
miliarius streckeri (Gloyd, op. cit.). This fragmentation of the head plates in some species of 
Sistrurus resembles the head scutellation of certain species of Crotalus (intermedius, polystictus, 
pricet, pusillus). The occurrence of fragmented head plates in rare individuals of Sistrurus 
does not invalidate the character of 9 head scales as diagnostic of Sistrurus, but it suggests 
that this character may not have as complex a genetic basis as thought by some. 


In the hemipenes of Sistrurus there is a gradual transition from spines to calyces, whereas 
in those of Crotalus there is an abrupt change from spines to fringes (Gloyd, 1940). Unfor- 
tunately, descriptions are not available of the hemipenes of the crucial species, S. ravus and 
C. pusillus. Klauber (1956) pointed out that in C. stejnegeri and C. lepidus the transition 
from spines to calyces is not so sharp as in most species of Crotalus, and in fact is similar to 
that found in Sistrurus. He considered this condition unique with stejnegeri and lepidus rather 


than primitive or intermediate. 


1964 | BRATTSTROM: EvoLUTION OF Pit VIPERS 239 


RHODOSTOMA 


HYPNALE MONTICOLA 


NEPA | HIMALAYANUS 7 


/~ straucH | + / 
HALYS ’ ) 
Boer a / PISCIVORUS 


BLOMHOFFI 


CONTORTRIX BILINEATUS 


ACUTUS \ 


ANNAME 
NSIC oer Si, \ NORTH 
MILLARD); ___ : \ AMERICA 
je x 
\ 
ASIA 
YS 
\ 


Fig. 34. Suggested phylogeny of A gkistrodon. 


On the basis of its osteology, Sistrurus is similar to the triseriatus group of Crotalus, with 
12 of 29 characters (41.4 per cent) in common. One of the supposed diagnostic features of 
Sistrurus is the acute angle formed by the posterolateral process of the squamosal. This angle 
is 80° or less in S. catenatus and miliarius. This is not a distinguishing character, however, for 
the angle is not acute in S. ravus, whereas it is acute in some of the small species of Crotalus, 
such as pusillus (table 11). Though the species of Sistrurus have many characters (72.7 per 
cent) in common, the group shares no character that distinguishes it from Crotalus. The dis- 
tinctness of the two genera breaks down in the similarity of S. ravus to several members of the 
triseriatus group, especially C. pusillus. I therefore consider Sistrurus as a subgenus of Crotalus. 


Sistrurus ravus is usually considered the most primitive member of the Sistrurus group 
(Gloyd, 1940; Klauber, 1956). This position is supported by osteological evidence, such as 
the shapes of the palatine, frontal, parietal, and prefrontal and the presence of a basisphenoid 
process. Sistrurus ravus is similar to C. pusillus in scutellation and osteology. Sistrurus ravus 
is the most Crotalus-like species in its group. 


The osteology of S. catenatus and miliarius is very similar (87 per cent of characters in 
common, table 14). Minton (1956) noted that miliarius venom seems more primitive than 
catenatus venom. It is most probable that catenatus and miliarius represent separate branches 
from a primitive ravus-like form (fig. 36). 


Two rattlesnakes, C. polystictus and stejnegeri, are unusual in head length, tail length, 
teeth, rattles, etc. (Klauber, 1952, 1956). The osteology of the rare stejnegeri (only 5 speci- 
mens known) is known only from counts of tooth sockets (table 6) and from a dentary bone. 
The latter has the dorsal process shorter than the ventral process. There are no palatine teeth. 
The osteology of the equally rare polystictus is unknown except that one specimen has 8 den- 
tary, 7 pterygoid, and no palatine teeth. The dentary and pterygoid teeth are extremely thin 
and elongate but are wide at the base. The reduction of the palatine teeth and the unusual 
proportional and scutellation differences of these two species (Klauber, 1937, 1938, 1939, 1952, 
1956) suggest that probably they are closely related and diverged from the main stock of 
rattlesnakes soon after one of the more primitive groups, Sistrurus. 


240 SAN Disco Society oF Natura History {Vou13 


wABEE 6 
Numerical summary of relationships? in A gkistrodon, based on table 10. 
S = os oe o = .. Number of 
3 5 2 = = 8 £ x characters 
Species 3 5S = & x & 8 shared by 
= = s 8 ss 2 = groups? 
Wiesel Ni7bq iazemis sou 0m ISON oLIG§  etiG 
acutus 61.5 654, 153080 S77 Sl 577 Soe 500 
a 4 1826) 13-26) 1426 ENIIL20" IS 26Nn NGO MSON 116-26 
bilineatus 6912, 2500) 9538" 550) 577 ozo 500 500 
_— oe? = 13-26 15-26 820 1626 7-16 10-16 
pisctvorus 50.0 Df, 40.0 61.5 43.8 62.5 
ea 172260) 1620 G@DGUNMLOIG NN ILIG 
blomhoffi 65.4 80.0 61.5 625 68.8 
; 15208 1O-26mm TONG mn IacIG 
contortrix 75.0 73.1 62.5 81.3 
14.20 9-14 11-20 8-14 
halys 70.0) (6A3) 55.0) Al 
1016 el 
hypnale 62.5 64.7 
ds 9-13 
rhodostoma 69.2 


lNumber of characters in common-number of characters used 


per cent of characters in common. 


“The New World forms of Agkistrodon share only 38.5 per cent (10 of 26) of characters; 
all members of the genus share 25.0 per cent (4 of 16) of characters. 


On the basis of osteology, the friseriatus group of rattlesnakes is divided here into three 
subgroups. The most primitive of these seems to be the intermedius subgroup, which includes 
intermedius and transversus. Closely related is the triseriatus subgroup, which includes pricet, 
triseriatus, and pusillus (fig. 36). The third is the lepidus subgroup, which includes lepidus 
and willardi. The distinctness of these species and groups is supported by external morphology 
(H. M. Smith, 1946; Klauber, 1952, 1956). H. M. Smith (1946) separated triseriatus from 
price. H. M. Smith (1946) and Klauber (1952) separated intermedius (or its synonyms) 
and transversus from the other small crotalids and placed them in the intermedius group 
(omiltemanus subgroup of H. M. Smith, 1946). Unfortunately, the osteology of intermedius 
is known only from the figures and description in H. M. Smith (1946; listed as C. gloydi 
lautus), which show the pterygoid with a medial angle in the posterior third, the palatine low, 
and the anterior border of the splenial passing straight ventrally, posterior to the anterior 
Meckelian foramen. Little is known about transversus, as the species is known from only three 
preserved specimens. On the basis of color and scutellation, Klauber (1952) believed it to be 
most closely related to intermedius. 

The shapes of the parietal, frontal, nasal, maxilla, and pterygoid suggest a close relation- 
ship of pusillus to pricei and triseriatus. It also has many characters in common with C. willardi 
and S. ravus. Crotalus pusillus is probably most closely related to triseriatus. Crotalus pusillus 
differs from price: and triseriatus in that the basisphenoid has a well developed process with 
an anterior hump. This character may indicate a primitive relationship with C. durissus or with 
the primitive Crotalus stock. 


1964 } BRATTSTROM: EVOLUTION OF Pit Vipers 241 


GODMANI 


ATROX ALTERNATUS 


NEUWIEDI 


JARARACUSSU LATERALIS 
JARARACA 
BRACHYSTOMA 
BILINEATUS BICOLOR DUNNI 
ITAPE TININGAE 
LANCEOLATUS 
NIGROVIRIDIS 
; Y 
COTIARA / LANSBERGI / 
NASUTA / , 
i OPHRYOMEGAS J 
/ 
NUM 
INSUL ARIS = / J UMMIFER 
CASTELNAUDI - i ips 
PICADO! : 


Fig. 35. Suggested phylogeny of Bothrops. 


Crotalus lepidus is probably most closely related to willardi. This is suggested by the shape 
of the parietal, frontals, prefrontals, squamosals, pterygoids, and process of the basisphenoid. 
Both species have many characters in common with C. triseriatus, C. pricei, S. catenatus, and 
S. miliarius (fig. 36, tables 11, 14). Crotalus lepidus and willardi seem to form a subgroup 
related to the ¢triseriatus subgroup. 


The relationships of the forms within the durissus group (horridus, molossus, basiliscus, 
durissus, and unicolor) were first pointed out by Gloyd (1940). These relationships are sup- 
ported by osteological evidence. Crotalus horridus and molossus are more closely related to 
each other than either is to the basiliscus-durissus-unicolor branch of the group. 


Crotalus basiliscus is very closely related to durissus. The two differ in minor ways in 
the shapes of the prefrontal, palatines, etc. (table 11). They differ also in general body shape 
and in the height of the neural spines on the anterior vertebrae. 


Crotalus d. unicolor van Lidth de Jeude, from Aruba Island, Dutch West Indies, has 
been considered both as a distinct species and as a subspecies of durissus (see Klauber, 1956: 
44). The relationships are very close. Even though the material of unicolor examined consisted 
of young specimens, it was apparent that unicolor is more closely related to C. d. terrificus than 
to C. d. tzabcan on the basis of nasal and squamosal shape. For various reasons, Klauber 
(1956) and Gloyd (1940) kept unicolor, tortugensis, exsul, and catalinensis as distinct island 
species, even though their relationships to certain mainland species are apparent and close. 


Gloyd (1940) and Klauber (1956) pointed out the distinctness of the atrox group of 
Crotalus. Both workers presented phylogenetic trees of this group. The osteological data sup- 
port these suggested relationships. The atrox group includes adamanteus (with its Pleistocene 
clinal ancestor, C. a. pleistofloridensis) and the extinct giganteus. The group also includes atrox 
(with its island derivitive, tortugensis) and ruber (with its island derivitives, exsul and cata- 
linensis). Crotalus atrox is intermediate in many osteological characters between ruber to the 
west and adamanteus to the east. Crotalus atrox is especially similar to adamanteus in frontal 
and parietal shape and to ruber in the shape of the basisphenoid, palatine, prefrontal, and 
ectopterygoid. Minton (1956) suggested, on the basis of venom, that ruber and atrox are 
closely related, showing less relationship to adamanteus than to each other, but within the 


{Vor. 13 


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243 


Pir VIPERS 


BRATTSTROM: EVOLUTION OF 


1964] 


05 
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gnssi8® %, 2) 
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Fig. 36. Suggested phylogeny of Crotalus, including Sistrurus. 


244 SAN Disco Society oF NAturRAL History [Vot. 13 


OKINAVENSIS 
CONVICTUS CHASENI /’ ELEGANS JERDONI 
i / (MONTICOLA || / 
\L id A B ' ff 
il 77 |MUGROSQUAMATUS‘ 
x Ke ‘ / FLAVOVIRIDIS 
Sas M, STRIATUS 
i Wet 
ACUTIMENTALUS i 
f STEJNEGERI 
( PURPUREOMACULATUS 
FLAVOMAGUL ATUS f GANTORIS GRAMINEUS 
\ \ 
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SUMATRANUS | i! — ! 


/ 
' 


' ScHULTZI \-~ 
; ‘ 


\_-~” MGGREGORI 
y 


ALBOLABRIS 


FASCIATUS 
MAGCROLEPIS; 
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CORNUTUS 


LABIALIS 


4 
“ MUTABILIS 


4 
7’ /POPIORUM ¢ PUNIGEUS GRACILIS 


BORNEENISIS _-~ 


/ - ‘pk, =< 
SS \ --~ _-~ANAMALLENSIS 


— --~~ TRIGONOCEPHALUS 


BOER OPS sae ee PHILIPPINENSIS 


Fig. 37. Suggested phylogeny of Trimeresurus. Solid lines, species examined osteologi- 
cally; dashed lines, species known only from external characters, largely from Maslin, 1942. 


same venom group. Crotalus adamanteus has the most specialized venom of the three species. 
On the basis of both osteology and external morphology, tortugensis appears to be a small 
form of atrox. Crotalus tortugensis resembles ruber in one peculiar character, having a ventrally- 
turned anterolateral process on the ectopterygoid. 


Crotalus exsul from Cedros Island, Baja California, is so closely related to ruber that if it 
were not for complicated nomenclatorial changes the two forms would probably be considered 
subspecific by most people. No skeletons were seen of exsul or of the recently described 
catalinensis Cliff from Catalina Island, Gulf of California, Mexico, but the relationships of 
exsul to C. r. ruber and of catalinensis to C. r. lucasensis ate so close on the basis of external 
characters that the osteology probably does not differ greatly. 


The relationship of scutulatus has been assumed to be with the atrox group, but scutulatus 
differs from that group in many respects, as noted by Klauber (1930). The venom and certain 
head scales suggest relationships with durissus. Klauber (1930, 1956) and Gloyd (1940) con- 
sidered scutulatus intermediate between the atrox and viridis groups. On the basis of its 
osteology, especially the shapes of the parietal, frontal, maxilla, and basisphenoid (table 11), 
scutulatus appears to be very closely related to the viridis group. It retains some of the primi- 
tive scale characters of the genus Crotalus and of the atrox group. The species probably 
represents an early offshoot of the viridis group, soon after the differentiation of the viridis 


and atrox groups (fig. 36). 


The viridis group, as here defined, includes mitchelli, tigris, viridis, cerastes, enyo, and 
potterensis. These forms have 14 of 29 characters (48.3 per cent) in common. They are 
especially similar in frontal and parietal shape. They each have an anteriorly directed depres- 
sion posterior to the process in the curvature of the pit-cavity of the maxilla. They all have 
relatively simple prefrontals and (except for mitchelli) low basisphenoids. The relationships of 
these species were first noted by Amaral (1929). Gloyd (1940) considered the relationship of 


1964 } BRATTSTROM: EVOLUTION OF Pir VIPERS 245 


enyo, cerastes, and tigris unknown. Klauber (1956) placed mitchelli, tigris, and viridis close 
together, but considered the relationship of cerastes and enyo to be with durissus (see discus- 
sion below). 


The most divergent member of the viridis group is mitchelli. In the shape of the processes 
of the prefrontal and in the well-developed process of the basisphenoid, it resembles the atrox 
and durissus groups. Crotalus mitchelli is probably the most primitive member of the viridis 
group, 1.e., it has changed least from the main Crotalus stock. This view is suggested by the 
shape of the basisphenoid and prefrontal and by the configuration of certain head scales. 


Osteologically, viridis seems to be closely related to potterensis, tigris, and enyo (table 14). 
On the basis of osteology, intraspecific relationships in viridis are difficult to determine. 
Crotalus y. concolor and C. v. lutosus seem closely related, as do C. yv. helleri and C. y. 
oreganus. In many characters, however, C. v. oreganus is more like C. y. viridis than like either 
C. v. lutosus or C. v. helleri. 

Crotalus potterensis is an extinct fossil species of the viridis group, described by Brattstrom 
(1953) from the Pleistocene of Potter Creek Cave, Shasta County, California, on the basis 
of two vertebrae, a part of a pterygoid, ribs, and other fragments. Subsequently, many other 
vertebrae from the same locality have been found in the paleontological collections of the 
University of California at Berkeley. This new material shows a change through time from 
typical viridis to typical potterensis within the cave deposits (Brattstrom, 1958). Crotalus 
potterensis differs from viridis in several characters, such as larger size and greater centrum 
length and diameter; they are similar in shape of the pterygoid and number of pterygoid teeth. 


As noted above, the relationships of cerastes and enyo have been uncertain (Amaral, 
1929; Gloyd, 1940; Klauber, 1956). It is suggested here that cerastes and enyo are members 
of the viridis group in view of the deep anteriorly-directed depression in the pit-cavity curvature, 
the low basisphenoid, the elongate frontals, etc. (tables 11, 14). On the basis of some minor 
hemipenal characters, the prominent scale tuberculations, and a high neural spine, Klauber 
(1956) suggested that cerastes and enyo are related to durissus. The hemipenal characters are 
also shared with such species as adamanteus and pricei. Klauber (1936b) had previously 
noted the scale and color similarities of cerastes and mitchelli. 


A high neural spine is found on the anterior vertebrae of durissus (fig. 11), cerastes, and 
enyo, as pointed out by Klauber (1956), but I believe that the similarity results from con- 
vergence. In durissus the high neural spine accounts for the high dorsal ridge evident externally. 
This ridge may be associated with the method of forming the resting coil in which each upper 
body loop is placed partly on the lower body loop rather than flat on the ground as in other 
species, or it may be associated with the peculiar striking position taken by this species. A 
very low neural spine is found in basiliscus (table 8), which is extremely similar to durissus 
in scutellation and in most osteological characters examined. Crotalus basiliscus differs from 
durissus externally primarily in being an extremely heavy flat snake; as far as known it does 
not coil or strike like durissus. 

In cerastes and enyo the tall neural spine is only slightly evident as a ridge externally. 
Here the tall neural spine may be primarily associated with increased musculature in the 
anterior thoracic region, used in the lateral undulations of enyo and the sidewinding locomotion 
of cerastes. Both species occur typically in open desert areas. The high neural spine in cerastes 
and enyo might represent an increased area for muscle attachment permitting fast movements 
across open areas (Cowles, 1956), with cerastes using an additional, more elaborate, mechanism 
—sidewinding. It is thus probable that the high neural spine has evolved twice in Crotalus 
for two separate functions and does not indicate, as Klauber (1956) believed, a relationship 
of cerastes and enyo to durissus. 

Crotalus cerastes and enyo are similar to the viridis group in external characters (Amaral, 
1929), in venom characters (Minton, 1956), and in skeletal characters (tables 11, 14). 
Crotalus cerastes and enyo have 70 to 86 per cent of their skeletal characters in common with 
members of the viridis group, and only 41 to 45 per cent in common with durissus. 


246 San Disco Society oF Natura History {Vot. 13 


TABLE Wb 


Numerical summary of relationships! in Bothrops, based on table 12. 


Species 


atrox 
bilineatus 
godmani 
jararaca 
jararacussu 
lanceolatus 
neuwtiedi 
cotiara 
nasuta 
nummifer 
schlegeli 


p24. 1621 13-20) 18:24 18:24" 16245 1624 916-22 S213") 1103224 99223 
alternatus 75.0 76.2 65.0 75.0 66.7 T27 61.5 } 


14-22 11-18 17-25 18-25 1825 16-25 15-23 7-13 10-23 8-24 


NI 
wr 
(>) 
loa) 
nN 
N 
aS 
Nn 
wn 
wn 
i) 
i) 


atrox GG 66. 680 20) 720) (GON Osea ose) ido aoe 
Pt [1e19) 1622 Nano) ENS 22 e140 MS IGEN o> OMITEON 
bilineatus 579) 727. | SAA 50) 9500 7010.) 385) tno od 
a z 1470) Silom NI Z0NdG 20) Nt2sIS elton ni re meMeTo 
godmani 7010) 5550) (55.0... 80.0, 9667 7510 uGll | 65729 
or [725) M6 2501625 15230 ONG IO een IB 

jararaca 68i0"7 64.0 16410-1" 165.2109 69D) w4psumioa2 
ji ; 2025 M1525) 1G 230) eeLIs ETO Sunes 
jararacussu S0L0F "6010 56.5 615 9435" 333 
oe 17-25) ElI523 0 elias ose 
lanceolatus 68.0 65.2 61.5 34.8 33.3 
os 17.23) leulge nia-pse nog 
neuwiedi 73.9 61.5 56.5 45.8 
Gill, MOD IIODS 
cotidara 54.5 47.6 43.5 
7Al3n ease 

nasuta 53.8 33.3 
14.22 
nummuifer 63.6 


INumber of characters in common-number of characters used 


per cent of characters in common. 


PHYLOGENY OF BOTHROPS 

The relationships of the species within the genus Bothrops are poorly known. About 30 
or 40 species have been described in the genus, of which probably two-thirds are valid. Many 
species are known from only one or two specimens each. Because of the lack of material, only 
a few species have been examined in this study (table 2). It is hoped that the separation of 
the major groups and subgroups of Bothrops on the basis of osteology will aid further sys- 
tematic work. 

As noted above and in figure 35, the genus Bothrops is divisable into two major groups 
based on osteology and scutellation. One of the groups is here termed the “arboreal” group, 
though not all members are arboreal; it includes at least nummifer and schlegeli. These two 
species are very much alike in the shapes of the frontal, parietal, lower jaw, palatine, ectoptery- 
goid, and premaxilla. Both appear to be quite primitive, with schlegeli probably the most 
primitive member of the genus. As noted above, schlegeli has several characters in common 
with the unique Trimeresurus wagleri. Bothrops schlegeli and T. wagleri probably have changed 
very little since their differentiation from the main lines of evolution of their respective genera. 
To judge from scutellation and habit, B. bicolor, lateralis, brachystoma, dunni, and nigroviridis 


may be related to nummifer and schlegeli (fig. 3D). 


1964 } BRATTSTROM: EVOLUTION OF Pit VIPERS 247 


TABLE 16 
Numerical summary of relationships’ in Trimeresurus and Bothrops, based on table 12. 


os 3 3 sea ~~ 

3 iS S 8 5 ar ws 

obs 4 x 00 2 * = 

Species ay 5 a 5 = ts i = 

: = ee = . Ww aS) : 

w : = By Q 

& eS 

13-260 We dcIBe (9515) Ie 2G S140 610 26u e626 S25 
ke gramineus 50.0 30.8 60.0 50.0 Diol 38.5 231 34.8 
i: Hew Gy MuceL yAC NE Ly QnpLaa us 
T. flaviviridis 385 935 50.0 50.0 42.3 46.2 34.8 
ar ar 15 po eee) WNnRO NScIa | ne Is cane 
T. muscrosquamatus 28.6. 9 53:8) 88.9. 385) 2 Ol5. Glo 
Bina hecs. as Oli5a) ALO Mm TdeG oNeeeIG 
T. puniceus 60.0 444 50.0 20.0 50.0 
- N46 OG s 
T. purpureomaculatus 64.3 46.2 38.5 65.2 
Be Auction 6-14 5:14 8-14 
T. stejnegeri 42.9 35.7 57.1 
— 9.26 14-23 
Ties wagleri 34.6 60.9 
ohio . 10-23 


B. atrox 43.5 


Number of characters in common-number of characters used 


per cent of characters in common. 


The second group is here called the “terrestrial” group, although a few members are 
somewhat arboreal. This group includes two subgroups, the small “hog-nosed pit vipers” and 
the large terrestrial forms. The first subgroup includes cotiara and nasuta, which have several 
characters in common (tables 12, 15). To judge from external characters, this group may also 


include lansbergi, itapetiningae, and ophryomegas (fig. 35). 


The subgroup of large terrestrial forms of Bothrops includes jararaca, jararacussu, atrox, 
neuwiedi, godmani, alternatus, bilineatus, and lanceolatus. Bothrops insularis, castelnaudi, and 
picadoi also may belong here, to judge from external characters. These species all seem closely 
related. The number of characters each has in common with other members of its genus can 
be seen in table 15. As most of the species have not been examined osteologically, the relation- 


ships diagrammed (fig. 35) are highly tentative. 
PHYLOGENY OF TRIMERESURUS 


Only seven species of this large genus have been examined osteologically. Contributions 
to our knowledge of relationships within this genus have been made by Pope and Pope (1933), 
Pope (1935), and Maslin (1942). The taxonomic status of many of the species of 
Trimeresurus is still, however, in doubt. This is due to the absence of adequate distributional 
and ecological data as well as to the paucity of specimens. Fortunately, specimens were 
examined of species representative of most of the groups of Trimeresurus of Maslin (1942). 


248 SAN Draco Society oF NATURAL History {Vot. 13 


As has been mentioned several times above, T. wagleri is the most distinct species in the 
Crotalidae. It has many characters that distinguish it not only from other species of Trimere- 
surus but also from most other crotalids; these include the medial location of the lower lumen 
of the fang, the depressed centers of the frontals, the low palatine, and the shape of the post- 
frontal, lower jaw, and ectopterygoid. In skull type, wagleri probably approaches the Viperidae 
more closely than does any other crotalid, and in this way it is probably the most primitive 
member of the family. Even though it may have retained many of the primitive characters of 
the family, however, it is highly specialized in other characters, such as its depressed frontals 
and medial lower Jumen of the fang. Because of its differences, wagleri is here separated from 
the other species of Trimeresurus into a distinct subgenus, for which the name Tropidolaemus 
Wagler is available. The very similar and probably subspecific (see E. H. Taylor, 1922) form, 
T. philippinensis, is included with it in this subgenus. No skulls of T. philippinensis were 


examined. 


Table 16 shows the number of characters each member of the genus has in common with 
each other member. Trimeresurus stejnegeri appears similar in many respects to gramineus and 
puniceus. The most striking osteological similarity is in parietal shape. Maslin (1942) placed 
stejnegeri in the same group as gramineus. The puniceus group of Maslin (1942) includes 
puniceus, gracilis, borneensis, cornutus, anamallensis, and trigonocephalus. Osteologically, T 
puniceus appears to be related to a primitive gramineus-stejnegeri as evidenced by the es 
of its parietal, frontal, and maxilla. The presence of the anterior hump on the basisphenoid in 
flaviviridis and puniceus may be due to convergence or may indicate some relationship. 


Trimeresurus mucrosquamatus is similar in many respects to flaviviridis, stejnegeri, and 
gramineus. Maslin (1942) placed it in a group with elegans. Trimeresurus purpureomaculatus 
is similar to stejnegeri in the shapes of frontal and basisphenoid and to gramineus in the shapes 
of pterygoid, maxilla, and parietal. Maslin (1942) included purpureomaculatus in a group with 
acutimentalis and erythrurus. The reduction of palatine and pterygoid teeth indicates, along 
with scutellation and characters of the basisphenoid and ectopterygoid, that flaviviridis is 
quite specialized. This specialization is probably related to its isolation on the Ryukyu Islands. 
Maslin (1942) placed flaviviridis in a group with jerdoni, which also lacks palatine teeth. 
Osteologically, flaviviridis seems to be related to a primitive mucrosquamatus-stejnegeri- 
gramineus stock, as indicated by the shapes of the frontal, parietal, postfrontal, and_basi- 
sphenoid. 


Figure 37 presents a very tentative suggestion of relationships within the genus Trimere- 
surus. The probable phylogenetic positions of members of the genus that were not studied 
osteologically are included from the information presented by Maslin (1942). The position 
of wagleri and its related form is well documented. On the basis of both osteological and 
external characters, the relationship among the monticola, mucrosquamatus, jerdoni, and 
flaviviridis groups and their position relative to stejnegeri and gramineus seem fairly well estab- 
lished. These groups also show a similarity in that the hemipenes bear true spines, lacking in 
other members of the genus (Maslin, 1942). The position of the other forms in the genus 
is quite tentative. 


TAXONOMY 


In view of the relationships suggested above on the basis of osteology and supported by 
evidence from the study of scutellation, venoms, hemipenes, and hybridization, a few taxonomic 
changes seem necessary. These changes are primarily associated with the use of the subgeneric 
concept. Though the subgenus is seldom used in modern herpetology, it is my opinion that it 
is useful in indicating relationships within a genus. For this reason, the two basic groups of 
Trimeresurus are separated as subgenera: wagleri and philippinensis are placed in Tropidolae- 
mus Wagler, and the remaining species in the nominate subgenus. It is also suggested that the 
two basic groups in the genus Bothrops should be treated as subgenera: Bothrops and Bothrie- 
chis, the latter to include the so-called “arboreal” group of species. 


1964 } BRATTSTROM: EVOLUTION OF Pit VIPERS 249 


It has been noted that the characters of the genera Crotalus and Sistrurus merge or break 
down. This breakdown is especially true in the Sistrurus-C. triseriatus complex, as seen in the 
osteology, hemipenes, squamation, venoms, and color. Also, hybridization occurs. It is my 
belief that the differences between Crotalus and Sistrurus are best indicated by placing all the 
rattlesnakes in one genus and considering Sistrurus as a subgenus. More than likely, for pur- 
poses of stability, most modern herpetologists will not readily take up Sistrurus as a subgeneric 
name. This is not really important as long as it is realized that the difference between Sistrurus 
and Crotalus is almost of the same order of magnitude as that between other species groups 
within Crotalus. 


Because of the seemingly different directions of evolution, the development of the loreal 
pit in the Crotalidae, and their geographic distribution, it is desirable that the Viperidae and 
Crotalidae be given full family rank, as has been done by various authors (Gilmore, 1938; 
Klauber, 1936a; H. M. Smith, 1950; Brattstrom, 1954). 


In the past, when the Crotalidae has been given family rank, two subfamilies have often 
been designated: the Lachesinae, including Lachesis, Bothrops, Trimeresurus, and sometimes 
A gkistrodon (Amaral, 1938), and the Crotalinae, including Crotalus and Sistrurus. This sub- 
family arrangement is untenable in view of the relationships of Lachesis to Crotalus suggested 
above and in view of the supposedly primitive condition of Agkistrodon. The only logical 
suprageneric grouping would place Bothrops and Trimeresurus in one group and A gkistrodon, 
Crotalus, and Lachesis in another. However, in my opinion, the division of the Crotalidae into 
subfamilies does not clarify any relationships and would be highly artificial. 

A summary of the generic and subgeneric taxonomy of the Crotalidae is given below. The 
characteristics of each genus and subgenus are listed, as well as the forms included in the taxa 
as here defined. No attempt has been made to give complete synonymies of the taxa, as these 
can be found in Gloyd (1940), Gloyd and Conant (1943), Klauber (1936b, 1952, 1956), 
Maslin (1942), Smith and Taylor (1945), and M. A. Smith (1943). 


Family CROTALIDAE Gray 
Genus Agkistrodon Beauvois 

Genotype. — Cenchris mokeson (Daudin). 

Characteristics. —Top of head with large plates anteriorly, usually nine in number. Sub- 
caudals in one or two rows. Pterygoid teeth extending posteriorly past middle of junction of 
ectopterygoid. Ectopterygoid usually shorter than basal portion of pterygoid. Frontals flat or 
turned up laterally. Palatine with a dorsal hump. Curvature of pit cavity a simple, open curve. 

Range. — United States, Mexico, Central America, southeastern Asia to southeastern 
Europe. 

Referred species. —acutus, annamensis, bilineatus, blomhoffi, contortrix, halys, hima- 
layanus, hypnale, millardi, mokeson, monticola, nepa, piscivorus, rhodostoma, strauchi. 


Genus Bothrops Wagler 
Subgenus Bothrops Wagler 

Genotype. — Coluber lanceolatus Lacépede. 

Characteristics. —Top of head with numerous small scales of varying size, including 
several scales in frontal area. Head scales usually keeled; gular scales not keeled. Subcaudals 
in one or two rows. Pterygoid teeth extending posteriorly past middle of articulation of ecto- 
pterygoid with pterygoid. Ectopterygoid longer than basal portion of pterygoid. Palatine 
forked anteriorly. Postnasal pore usually absent. Frontals flat or turned up laterally, longer 
than wide. Edge of pit cavity with a smooth curve. Primarily terrestrial. 

Range. — Mexico, Central and South America. 

Referred species. — alternatus, atrox, bilineatus, godmani, insularis, jararaca, jararacussu, 
lanceolatus, neuwiedi. The species cotiara, lansbergi, nasuta, ophryomegas, picadoi, and other 
little-known species of Bothrops may belong here, or they may represent additional subgenera. 


250 SAN Dirco Society oF NaturaAt History {Vot. 13 


Subgenus Bothriechis Peters 

Genotype. — Bothriechis nigroviridis Peters. 

Characteristics. — Like the above except palatine (roundish) not forked anteriorly; frontal 
wider than long; curvature of pit cavity sometimes with a median hump. Usually arboreal. 

Range. — Mexico, Central and South America. 

Referred species. — bicolor, brachystoma, dunni, lateralis, nigroviridis, nummifer, and 
schlegeli. Probably some of the other little-known arboreal species of Bothrops belong here, 
but they are not assigned to either subgenus for lack of information. The name Teleuraspis 
Cope, (Proc. Acad. Nat. Sci. Philadelphia, 1859:338, Type Trigonocephalus schlegeli 
Berthold) is available if the arboreal group of Bothrops is further divided in the future. 


Genus Crotalus Linnaeus 
Subgenus Crotalus Linnaeus 

Genotype. — Crotalus horridus Linnaeus. 

Characteristics. —Top of head with scales of varying size; more than one scale in frontal 
area. Parietal scales, if enlarged, not in contact nor symmetrical. Rattles present. Subcaudals 
in one or two rows. Pterygoid teeth not extending posteriorly past middle of articulation of 
ectopterygoid with pterygoid. Ectopterygoid longer than basal portion of pterygoid. Frontals 
flat or turned up laterally. Curvature of pit cavity interrupted by a dorsal hump or process. 
Lateral process of squamosal blunt or pointed, but if forming a fork with the main bone, angle 
between them 90° or greater. 

Range. — North, Central, and South America. 

Referred species. — adamanteus, atrox, basiliscus, catalinensis, cerastes, durissus, enyo, 
exsul, horridus, intermedius, lepidus, mitchelli, molossus, polystictus, pricei, pusillus, ruber, 
scutulatus, stejnegeri, tigris, tortugensis, transversus, unicolor, viridis, willardi, + potterensis, 
Tgiganteus. 

Subgenus Sistrurus Garman 

Genotype. — Crotalus miliarius Linnaeus. 

Characteristics. —Top of head with large plates anteriorly, usually nine in number, in- 
cluding a single frontal and a pair of large symmetrical parietals in contact. Rattles present. 
Subcaudals in one or two rows. Pterygoid teeth not extending past middle of articulation of 
ectopterygoid with pterygoid. Ectopterygoid longer than basal portion of pterygoid. Frontals 
flat or turned up laterally. Curvature of pit cavity interrupted by a dorsal hump or process. 
Lateral process of squamosal forked at an angle of 80° or less with the main bone, except in 
C. ravus. 

Range. — Eastern Canada, eastern United States, and central Mexico. 

Referred species. — catenatus, miliarius, ravus. 


Genus Lachesis Daudin 


Genotype. — Crotalus mutus Linnaeus. 

Characteristics. —Top of head with small, keeled scales. Posterior subcaudals in four 
rows. Pterygoid teeth not extending posteriorly past middle of junction of ectopterygoid with 
pterygoid. Ectopterygoid longer than basal portion of pterygoid. Palatine with high dorsal 
hump. Top of frontals flat. Curvature of pit cavity interrupted by a well-developed process 
with a small depression on its outer surface. 

Range. — Central and northern South America, Trinidad Island. 

Referred species. — muta. 


Genus Trimeresurus Lacépede 
Subgenus Trimeresurus Lacépéde 
Genotype. — Trimeresurus viridis Lacépede (= T. gramineus (Shaw) ). 


Characteristics. —Top of head with numerous small scales of varying size, including sev- 
eral or many scales in the frontal area. Nasal pore usually present on posterior wall of nostril. 


1964 } BRATTSTROM: EVOLUTION OF Pit VIPERS 251 


Scales on top of head anterior to parietal region smooth; gular scales not keeled. Subcaudals 
in one or two rows. Pterygoid teeth extending posteriorly beyond middle of articulation of 
ectopterygoid with pterygoid. Ectopterygoid longer than basal portion of pterygoid. Palatine 
not forked anteriorly. Frontals flat or at least without depressed centers. Lower lumen of fang 
anterior. Prefrontal laterally longer than wide. Curvature of pit cavity having a small hump 
or process. Gular scales not keeled. Venom, as far as known, hemotoxic. Terrestrial or arboreal. 


Range. — Southeastern Asia to India and Ceylon. 


Referred species. —acutimentalis, albolabris, anamallensis, borneensis, cantoris, chaseni, 
convictus, cornutus, elegans, erythrurus, fasciatus, flavomaculatus, flavoviridis, gracilis, gramt- 
neus, halieus, jerdoni, kanburiensis, kaulbacki, labialis, macrolepis, malabaricus, mcegregori, 
monticola, mucrosquamatus, mutabilis, okinavensis, popiorum, puniceus, purpureomaculatus, 
schultzi, stejnegeri, strigatus, sumatranus, trigonocephalus. 

Subgenus Tropidolaemus Wagler 

Genotype. —Cophias wagleri Boie. 

Characteristics. — Top of head with numerous small scales of varying size, including many 
or several in the frontal area. All scales on top of head usually keeled; gular scales keeled. 
Nasal pore absent. Subcaudals in one or two rows. Pterygoid teeth extending posteriorly past 
middle of junction of ectopterygoid with pterygoid. Ectopterygoid longer than basal portion 
of pterygoid. Palatine low, not humped or forked. Frontals with centers depressed. Lower 
lumen of fang medial. Prefrontals laterally wider than long. Curvature of pit cavity interrupted 
by a small process or knob. Venom neurotoxic. Color primarily green. Arboreal. 

Range. — Malaya, Sumatra, Philippine Islands. 

Referred species. —wagleri and probably philippinensis. The latter was not seen, but was 
considered by Taylor (1922) to be very closely related to wagleri, if not subspecific. 


ZOOGEOGRAPHY 


After the basic relationships for a group have been established, it is interesting and enjoy- 
able to speculate on the evolution and zoogeography of the group. 

Darlington (1957) recently summarized the zoogeography of the pit vipers as follows: 
“Crotalines are unknown fossil before the Pliocene (when rattlesnakes were in North America). 
Other clues suggest that the subfamily originated in Asia and that both Agkistrodon and 
Trimeresurus (Bothrops) moved from Asia to America, one of these general [ sic} (or an earlier 
stock) presumably giving rise to the rattlesnakes” (p. 228), and “. . . pit-vipers prob- 
ably arose (from true vipers) in southern Eurasia, and two or three stocks reached America, 
one producing the rattlesnakes” (p. 203-204). 

As will be seen below, I do not disagree with this interpretation but wish only to expand 
the discussion. Someone has said that all zoogeography is speculation. I do not agree com- 
pletely, but it must be realized that the following comments are largely that. It is my inter- 
pretation of events which may have taken place if the data on the fossils and phylogeny are 
accurate and the basic assumptions are correct. 

For an evolutionary and zoogeographic analysis, something should be known of (1) the 
relationships of the species and genera, (2) the fossil history, (3) the ecology, behavior, and 
physiology of recent forms, (4) the distribution of land areas and bridges at various times 
in the past, and (5) past climatic conditions. 

Phylogeny and fossils. —The relationships and phylogeny of the pit vipers are sum- 
marized in figures 33 to 37 and tables 13 to 16. Fossil crotalids do not aid greatly in inter- 
preting relationships or in constructing past distributions. Many recent species have changed 
little since the Pliocene (Brattstrom, 1954; Klauber, 1956). 

Land areas and bridges. — The distribution of the continents during the history of the 
Crotalidae has probably been similar to that of today. Theories for Cenozoic and Mesozoic 
continental drift and trans-oceanic land bridges (between South America and Africa and be- 
tween South America and Asia) have been discarded by most workers (Mayr et. al., 1952; 


252 SAN Disco Society oF Naturat History {Vot. 13 


Press in Poldervaart et al., 1955). The presence and absence of certain island chains are of 
critical importance in interpreting zoogeographic distribution. In regard to the history of the 
Crotalidae, the North and South American connections, the Bering land bridge, and the 
changes in the Sundaland islands are extensively documented and discussed in the literature 
(Underwood, 1954; Inger, 1947, 1954; Hesse, Allee, and Schmidt, 1951; Darlington, 1948, 
1957; Simpson, 1947, 1953, 1956; Stirton, 1950; Durham, Arellano, and Peck, 1955). These 
data will not be reviewed here but will be taken into consideration in the speculations to be 
presented. For example, a group is not stated as having dispersed to a certain area until a land 
bridge is inferred to have been present. 

The possibility of chance dispersal on rafts, etc., (Simpson, 1952, 1953, 1956; Axelrod, 
1952b) is probably not of major importance in crotalid zoogeography. No crotalids occur in 
the West Indies, the Galapagos or Revillagigedos islands, New Guinea, Australia, Celebes, or 
the islands of Oceania. The possibility of rafting is not entirely ruled out, however, and there 
are even records of Crotalus adamanteus swimming 22 miles out to sea between the Florida 
Keys (Klauber, 1956: 464, 470). The swimming ability of other crotalids is also documented 
(Klauber, 1956: 464-472). Though the distribution of crotalids in the Malaya-Borneo-Philip- 
pine area is primarily associated with Sundaland (Darlington, 1948) or other known land 
bridges (see Inger, 1947, 1954, for discussion and review), there remains a possibility of 
crotalids having rafted to some of these islands, or across the Bering and Panama sea straits, 
cr to the islands of Trinidad, Martinique, or Formosa. 

Chance dispersal by crotalids on rafts would be facilitated by their being ectothermic. In 
a cool area they would be able to withstand long periods without food. Rafting in cold areas, 
however, would probably be lethal to the snakes (Brattstrom, MS; Klauber, 1956: 387-391). 


Ecology of recent forms. — The temperature requirements of a few crotalids are rela- 
tively well known (Cowles and Bogert, 1944; Cowles, 1945; Fitch, 1960; Brattstrom, MS). 
As ectothermic reptiles, their body heat comes entirely from external sources (Cowles and 
Bogert, 1944; Brattstrom, MS). As noted by Cowles (1945) and correlated with Bergmann’s 
rule for ectotherms, large ectotherms do not live in more northern latitudes today. This is 
largely because in more northern latitudes there is not enough solar radiation (the major source 
of heat to ectotherms) to heat a large reptile to body temperatures under which it can carry 
out its normal physiological and ecological functions. Small lizards and snakes (Eumeces, 
Thamnophis) and frogs, toads, and salamanders (with their lower thermal preferences) can 
and do occur in northern latitudes today. Cold or cool northern climates apparently inhibit 
the northern distribution of crotalids. This is of importance in interpreting the zoogeography 
of the crotalids, especially in regard to the times of dispersal across the Bering land bridge. 
For example, though such warm-blooded forms as horses, mammoths, and mastodons could 
cross the Bering land bridge during the cold periods of the Pleistocene (Simpson, 1947; Stir- 
ton, 1950), probably the cold Pleistocene glacial climates and probably even the cool inter- 
glacial climates, if they were similar to that of today, inhibited dispersal of crotalids across 


the Bering land bridge. 


The present distribution of the Crotalids is shown in figure 38, and the present distribu- 
tion of the various genera in figures 40C and 41. Except for Crotalus and Sistrurus (Klauber, 
1956) and the copperheads (Fitch, 1960), the ecology of most crotalids is not well docu- 
mented. Trimeresurus and Bothrops are tropical today, Trimeresurus in southeastern Asia and 
the Sundaland, Philippine, and Ryukyu islands, Bothrops in Central and South America. 
A gkistrodon is primarily warm temperate. A gkistrodon contortrix is primarily associated with 
the Eastern Deciduous Forest, occurring rarely into semiarid areas in Texas. The semiaquatic 
A. piscivorus occurs in the warm temperate to subtropical areas of the southeastern United 
States. A gkistrodon bilineatus, though occurring within the tropical region, is apparently pri- 
marily associated with temperate climates. The ecology of the Asiatic forms of A gkistrodon 
is not well known, but apparently most occur in temperate climates, often in relatively cool 
montane regions within the tropics; hence the apparent overlap in range of Trimeresurus and 


1964 } BRATTSTROM: EVOLUTION OF Pir VIPERS 253 


A gkistrodon (fig. 40C). Lachesis muta occurs today in the tropical regions of southern Central 
and northern South America and in Trinidad. 


The genus Sistrurus occurs today in eastern North America and central Mexico. Sistrurus 
cdtenatus occurs primarily in the prairie and the deciduous hardwood forest, muliarius occurs 
primarily in the warm temperate forests of the southeastern United States, and ravus occurs in 
the arid temperate regions of central Mexico. Most of the species of Crotalus occur today in 
arid or subarid open areas (deserts, prairies, etc.) or in dry deciduous or coniferous forests in 
western North America. The major exceptions to this distribution are adamanteus, in warm 
temperate to subtropic parts of the southeastern United States, and the members of the durissus 
group: horridus in warm temperate to subtropic parts of the eastern United States, molossus 
on the dry plateau of Mexico and the southwestern United States, basiliscus in the arid sub- 
tropics of the west coast of Mexico and Central America, and durissus in the arid subtropics 
to wet tropics of Central and South America (fig. 41). 


Past climates. — A wealth of information has been accumulated concerning the Cenozoic 
climates of the world, and especially of North America, based on the distributions of fossil 
plants and the climatic requirements and morphological adaptations of their modern close rela- 
tives. The basic assumptions, the floras, and the inferred climates have been described and 
discussed by Chaney (1940, 1947, 1948), Chaney et. al. (1944), Borek and Ryan (1952), 
Axelrod (1938, 1949, 1950a, 1950b, 1952a, 1952b, 1956, 1958), and MacGinitie (1941, 1953). 
Of primary interest in terms of crotalid zoogeography are the changes in the Madro- Tertiary, 
Arcto-Tertiary, and Neotropical Tertiary Geofloras and the evolution of the desert vegetation 
in western North America as outlined by Chaney, Axelrod, and others (supra cit.). The 
details of these changes are elaborated by these authors, but the essential features are as fol- 
lows: Subtropical and tropical conditions were more widespread in the early Tertiary than at 
present and occurred farther north, with subtropical conditions occurring in southern Alaska. 
It is possible that in the Paleocene and Eocene subtropical climates may have occurred in the 
area of the Bering land bridge. The tropics and subtropics were restricted southward during 
the remainder of the Tertiary and were essentially in their present position by the Pliocene. 
With a southward retreat of the tropics and with topographic changes, the more northern 
Arcto-Tertiary and Madro-Tertiary Geofloras developed and differentiated into several basic 
types. This differentiation occurred primarily in late Oligocene, Miocene, and Early Pliocene 
and was largely related to an expansion of the Madro-Tertiary Geoflora in the southwestern 
parts of North America (Axelrod, 1949:159; 1958); the latter geoflora differentiated into 
grasslands, woodlands, chaparral, and semidesert (Axelrod, 1949:159, 161; 1950b:287). The 
grasslands in the present prairie states apparently had their origin in Late Oligocene and their 
major expansion in the Miocene (Chaney and Elias, 1938). The present deserts did not come 
into existence until relatively recently — post mid-Pliocene, according to Axelrod (1950b: 


296-298). 


The general assumption in the interpretation of past climates from fossil floras involves 
the use of the ecological requirements of modern plants. Past climates are inferred on the basis 
of needs of modern plants. The assumption made is that the plants have not greatly altered 
their physiological or ecological requirements, or, if they have, that physiological changes in the 
plant might be reflected in some change in the morphology of the plant (1.e., changes in leaf 
size, shape, thickness, etc.; Axelrod, Chaney, etc., supra cit.). Most of the paleoclimates that 
have been inferred on the basis of fossil plants are supported by the few paleotemperatures 
determined by the use of oxygen isotopes of marine environments (Epstein and Lowenstam, 
1953; Lowenstam and Epstein, 1954; Emiliani, 1954, 1955). A method of suggesting past 
climates on the basis of fossil reptiles and amphibians (especially snakes, lizards, frogs, toads, 
and salamanders), based on the same sort of assumptions used by the paleobotanists, is being 


proposed by Brattstrom (MS). 


Fig. 38. Present distribution of the Viperidae (horizontal lines) and of the Crotalidae (vertical lines) . 


1964 | BRATTSTROM: EVOLUTION OF Pit VIPERS 25 


ZOOGEOGRAPHY OF THE CROTALIDAE 


In the absence of critical fossils, the speculations on the evolution and zoogeography of 
the crotalids as presented below are based largely on the past distribution of climates as deter- 
mined from paleobotanical data. The first assumption made is that if a group or species occurs 
in a certain climatic area today (ji.e., tropics, subtropics), it probably has been adapted to and 
remained in this climatic area for most of its evolutionary history, and that the past distribu- 
tion of this form can be suggested by the past distribution of this climate or flora. It is further 
assumed that snakes have not altered their physiological and ecological requirements in the 
past. If this is true, it follows that each form must have continued to live in the same environ- 
ment. Therefore, the potential area of each form at any time in the past is shown by the 
distribution of suitable climate and vegetation at that time. As with plants, the basic assump- 
tions would appear to be safe for faunas that remained intact but less safe for individual 
species. Recent studies by Brattstrom (1961) on fossil tortoises associated with fossil floras 
suggest that in this group, at least, there has been no great change in ecological preference 
through the Cenozoic; a group of tortoises which is tropical today was associated in the past 
with tropical floras. 


With regard to crotalids, it is known that the related Bothrops and Trimeresurus occur 
in the tropics today (fig. 40C), the former in the New World, the latter in the Old. If we 
assume that Bothrops and Trimeresurus were always associated with tropical or subtropical 
floras, then the maximum possible past distribution of the two genera is suggested by the dis- 
tribution of tropical and subtropical floras for the various epochs. As the tropics and subtropics 
have been contracted for the latter part of the Cenozoic (Chaney, Axelrod, etc., supra cit.), if 
Bothrops and Trimeresurus were in tropical or subtropical areas they could not have crossed 
the Bering land bridge (fig. 40) during that time. In the Paleocene and Eocene, however, 
subtropical conditions occurred in the area of the Bering land bridge (fig. 40A), and at that 
time dispersal of these forms could have occurred across that area. In post-Eocene times, the 
tropics became restricted more to the south (fig. 40B) and thus, according to the two assump- 
tions, Bothrops or Trimeresurus could not have crossed the Bering land bridge. The predomi- 
nately warm-temperate A gkistrodon, however, apparently could occur at these latitudes until 


later times (fig. 40). 


Using the phylogenetic and geological data available, the two stated assumptions, and the 
inferred past climates based on paleobotanical studies, the following are my speculations on the 
evolution and zoogeography of the Crotalidae. A summary of these speculations is diagrammed 


in figure 39. 


Crotalidae.— The various criteria of Matthew (1915) for determining the center of 
origin or center of dispersal on the basis of the number of species, distribution of primitive 
forms, etc., are not always convincing. Some of these criteria suggest, but do not prove, an 
Old World origin for the Crotalidae. For example, the most primitive members of the 
Crotalidae (Trimeresurus wagleri and the hypnale-contortrix group of Agkistrodon) occur 
primarily in the Old World today (except A. contortrix). The most specialized members of 
the family (Crotalus and Sistrurus) occur in the New World. Probably the greatest con- 
centration of species occurs in the Old World Tropics (species of Trimeresurus), but nearly 
as great a concentration occurs in the New World Tropics (species of Bothrops). 


Snakes probably had their origin in the Lower Cretaceous (Bellairs and Underwood, 
1951). The time of origin of the Crotalidae is unknown, but it may have been in the Upper 
Cretaceous, when several families of snakes originated (Gilmore, 1938; Bellairs and Under- 
wood, 1951). The fossil history of the Viperidae is better known than that of the Crotalidae, 
and suggests that the Viperidae had an Old World center of origin and dispersal. The present 
distribution of the Viperidae (fig. 38) would also suggest this. It is probable that the Crotali- 
dae originated from the Viperidae in the Old World, as Darlington (1957) suggested. 


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Fig. 40. Hypothetical past distribution of Agkistrodon (dots) and of Bothrops and 
Trimeresurus (lines). A, Cretaceous-Eocene; B, Miocene; C, Recent. Solid line, presumed 
boundary between Tropical and Subtropical vs. Warm Temperate floras. 


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tween Tropical and Subtropical vs. Warm Temperate floras. 


1964 } BRATTSTROM: EVOLUTION OF Pit VIPERS 259 


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260 SAN Disco Society oF NaturaAt History {Vot. 13 


Soon after the origin of the Crotalidae, apparently two major groups arose, a proto- 
Trimeresurus-Bothrops stock, occurring in or becoming adapted to tropical or subtropical con- 
ditions, and A gkistrodon, occurring in or becoming adapted to temperate climates. When times 
were equable in the Paleocene and Eocene, both these basic stocks apparently dispersed into the 
New World over the Bering land bridge (figs. 39, 40A). With the subsequent restriction of 
the tropics and subtropics southward through the remainder of the Cenozoic, the differentiat- 
ing Bothrops and Trimeresurus stocks presumably retreated southward to their present posi- 
tions. One of the temperate members of the Agkistrodon stock probably gave rise to a 
Lachesis-Crotalus stock soon after reaching the New World. 


A gkistrodon. — Because of some of its primitive characters and its present temperate 
distribution, the basic A gkistrodon stock is assumed to be an early (perhaps Eocene or Paleo- 
cene) branch of the crotalid line occurring in or becoming adapted to temperate climates 
and floras. The two major groups of Agkistrodon perhaps differentiated in Late Eocene times. 
basic stocks of both groups invading the New World across the Bering land bridge, associated 
with warm temperate climates (figs. 39, 40A). 


One of these groups (A. acutus-bilineatus-piscivorus) may have differentiated in the New 
World into the present-day piscivorus and bilineatus, which perhaps moved southward with 
the climatic zones in late Tertiary. The Old World relic of this group is acutus, of South 
China and Formosa. 


The other group of A gkistrodon survives today mainly in the Old World (hypnale, halys, 
etc.), with only contortrix in the New World. A contortrix-like stock of this group soon after 
its arrival in the New World probably gave rise to the Lachesis-Crotalus-Sistrurus lineage. 
The Old World members of this Agkistrodon group perhaps differentiated along with Old 
World temperate floras or with isolation on various mountain ranges. The southeasternmost 
member of this Asiatic group, A. rhodostoma, occurs in isolated warm temperate mountain 
regions in Malaya, Sumatra, and Java. 


Bothrops-Trimeresurus. — It is suggested above that in the Paleocene or Eocene the proto- 
type of Bothrops and Trimeresurus, occurring in tropical or subtropical regions, crossed the 
Bering area and that with the restriction of the tropics southward through the rest of the 
Cenozoic, the ranges of the two genera also became restricted southward. The similarity noted 
between Bothrops and Trimeresurus today is probably due to their retention of characters, 
many of them primitive in the family, associated with tropical or subtropical conditions — 
conditions where they are assumed to have lived for most or all of their evolutionary history 


(figs. 39,40). 
In the Old World, the wagleri group of Trimeresurus may have diverged early from the 


main stock, possibly in Eocene times, and perhaps even before the Trimeresurus-Bothrops 
prototype reached the New World. The relationships, distribution, and ecology of the remain- 
ing species of Trimeresurus are too little known to allow further comment. 


The genus Bothrops consists of an arboreal group, mainly in Central America, and a 
terrestrial group, mainly in South America but with B. atrox reaching southern Mexico. In 
view of the diversity of the numerous species, it seems unlikely that the two groups could have 
diverged since the reestablishment of the Panama land bridge in Upper Pliocene to Recent. 
Rather, they probably developed in isolation on the two sides of the Panama Sea Straits, 
present from Eocene to Pliocene. It is uncertain whether Bothrops originally reached South 
America by rafting over this strait as certain mammals and tortoises appear to have done 


(Simpson, 1943, 1947, 1953a; Stirton, 1950; Brattstrom, 1961) or by land during the Eocene. 


Lachesis-Crotalus-Sistrurus.— One of the primitive members of A gkistrodon probably 
soon after arriving in the New World gave rise to the Crotalus-Lachesis line. This line prob- 
ably soon diverged into Lachesis and Crotalus (figs. 39, 41). Assuming that the Lachesis stock 


was associated with more subtropical floras, the basic Lachesis stock probably retreated south- 


1964 | BRATTSTROM: EVOLUTION OF Pit VIPERS 261 


ward with the retreating flora. The southward dispersal of Lachesis may have been stopped by 
the Panama Sea Straits, or Lachesis may have rafted into South America. Its present distri- 
bution is Central and northern South America and Trinidad. Its large size, the homogeneity 
within the species, and its ovoviviparity may be related to a long time of isolation, possibly 
since Oligocene times. It is possible that the small range of this species is related to competition 
with the large terrestrial species of Bothrops. Very few of the large species of Bothrops occur 
within or north of the range of Lachesis. 


Probably after the differentiation of the Lachesis stock, a rattle developed in the remain- 
ing Crotalus line. Within this line, two groups differentiated, Sistrurus retaining large head 
plates, and Crotalus developing small scales on top of the head. The primitive C. polystictus 
and stejnegeri probably soon diverged from the primitive Crotalus stock. The durissus, viridis, 
atrox, and triseriatus groups of Crotalus perhaps differentiated in Late Eocene or Early Oligo- 
cene. The subsequent history of each group is suggested in figure 41. The Pliocene and 
Pleistocene fossil crotalids (Brattstrom, 1954, 1958, MS) suggest that most modern species of 
rattlesnakes had differentiated and attained their present ranges by Middle or Late Pliocene. 
At least two forms, potterensis and giganteus, became extinct near the end of the Pleistocene. 


SUMMARY 


As a means of determining relationships among members of the family Crotalidae, some 
414 skeletons of 105 species were examined, described, and figured. Effects on the skeleton of 
injury, disease, form of preservation, and method of preparation, were negligible or were 
sufficiently obvious to be taken into consideration. Some bones such as nasals and ribs are 
quite variable within species and hence of little value in study; others, such as frontals and 
parietals, are relatively constant and distinctive for each species and hence useful in determin- 
ing relationships. A tentative phylogenetic plan of crotalid evolution is presented, based 
primarily on the osteology of the forms studied but supported by data from studies of fossils, 
venoms, hemipenes, scutellation, muscles, and hybridization. 


For the family Crotalidae, six genera are recognized: Bothrops, in the New World, with 
the subgenera Bothrops and Bothriechis; Trimeresurus, in the Old World, with the subgenera 
Trimeresurus and Tropidolaemus; Lachesis, in the New World, monotypic; Agkistrodon, in 
the Old and New World; and Crotalus, in the New World, with the subgenera Crotalus and 


Sistrurus. 


Trimeresurus wagleri and some of the Asiatic members of the genus A gkistrodon are con- 
sidered the most primitive members of the family. The former, while retaining several primitive 
characters, has also evolved some divergent osteological specializations (depressed centers of the 
frontals, medial position of the lower lumen of the fang). The rattlesnakes, Crotalus, are 
probably the most specialized members of the family. Lachesis is most closely related to them, 
and both groups probably were derived from a primitive Agkistrodon stock. Trimeresurus and 
Bothrops are closely related but distinct. 


The age of crotalids can be determined by counting the annual growth rings in the bones. 
These rings are best counted in the frontal bone. 


Because of the lack of adequate fossils, little can be said about the zoogeography of the 
family at present. A few of the writer’s speculations are presented. 


262 SAN Disco Society of NATuRAL History { Vor. 13 


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1964 | BRATTSTROM: EVOLUTION OF Pit VIPERS 265 


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chez les serpentes. Ann. Sci. Nat. Zool. 16:161-205. 


266 San Disco Society of NATURAL History {Vot. 13 


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Rapovanovic, M. 
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TRANSACTIONS 
OF THE 


SAN DIEGO SOCIETY OF NATURAL HISTORY 


VoLuME 13, No. 12, pp. 269-284 


THE STAPHYLINIDAE OF THE MARINE MUD FLATS 
OF SOUTHERN CALIFORNIA AND 
NORTHWESTERN BAJA CALIFORNIA 
(COLEOPTERA) 


BY 
IAN Moore 


Research Associate in Entomology 


San Diego Natural History Museum 


SAN DIEGO, CALIFORNIA 
PRINTED FOR THE SOCIETY 


May 4, 1964 


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THE STAPHYLINIDAE OF THE MARINE MUD FLATS 
OF SOUTHERN CALIFORNIA AND 


NORTHWESTERN BAJA CALIFORNIA ; 
MUS, COMP ZOOL 


(COLEOPTERA) LIBRARY 
BY 1 9 13x04 
IaN Moore mAnKVARD 
UNIVE RSILY, 
INTRODUCTION 


In southern California and in northern Baja California, Mexico, Coleoptera occupy three 
rather distinct seashore habitats — rocky areas, sandy beaches, and mudflats — with compara- 
tively little overlap in species from one to another. I have dealt previously with the Coleoptera 
of rocky shores in southern California and with certain of those of the sandy beaches (Moore, 
1956a, 1956b). The present paper deals with the Coleoptera, and especially the Staphylinidae, 
of estuaries and their associated mudflats — the habitat for maritime Coleoptera that has been 
least studied in southern California. 

LeConte (1877) and Casey (1889) reported several species of Staphylinidae from salt 
marshes at San Diego, California, but conditions have so changed in San Diego County that 
it is now difficult to find even a small undisturbed area where the salt marsh fauna can be 
investigated adequately. In northern Baja California, however, changes are occurring more 
slowly, and undisturbed areas may still be found. 

The bays and estuaries of southern California and northern Baja California are of two 
distinct types, the first an open bay with a direct connection with the ocean, the other a lagoon 
which is usually separated from the ocean by a sand bar. Along the shore of either may be 
a salt marsh. The few open bays of southern California have been greatly modified in the past 
fifty years by the activities of man. Most of their once extensive mud flats have been covered 
with fill-dirt and utilized for industry or commerce, and their waters are often heavily polluted 
by sewage efHuent, oils, and industrial wastes. In some small areas in the southern part of 
San Diego Bay it is still possible to find mud flats covered with sea lettuce. Relatively larger 
areas in Baja California, as at El Estero, seven miles south of Ensenada, are unmodified by 
man, but the situation there is changing rapidly. 

The small lagoons of southern California are more numerous than the open bays, occurring 
at the mouths of major canyons which drain the mesa areas to the east. Their protective bars 
are broken only occasionally during rare periods of heavy rain, or possibly sometimes by the 
surf of severe winter storms. The activities of man have not yet altered these habitats as greatly 
as they have those of the open bays, but the changes are noticeable. It therefore seems worth- 
while to describe in detail the relatively undisturbed salt marsh in northwestern Baja California 
where this study was made. 


A Sact MarsH IN NorTHERN BAJA CALIFORNIA 


During the summer of 1961 I spent several weekends near a small salt marsh called La 
Salina, which proved to have a fauna of several species of Staphylinidae, with individuals in 
great abundance. La Salina is on the Guerrero Ranch about four miles south of La Mision de 
San Miguel. Being approximately fifty miles south of San Diego, it has a similar climate. The 
salt marsh is small, approximately one-half mile long and one-quarter mile wide. According to 
Sefior Angel Covarubias, who has lived on the adjacent ranch to the south for about twenty 
years, this marsh last had a direct connection with the ocean following the major storm of 
1941. Since that time, it has been separated from the ocean by a sand bar over which the tides 
flow only eight to twelve times a year during the winter months. The marsh is at the mouth 
of a large canyon which drains an extensive high inland mesa. One of the driest years in 
recent history of this area was 1961, a fact which may have some bearing on the abundance of 
insects I found there at that time. 


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1964] Moore: STAPHYLINIDAE OF MARINE Mup Friars 273 


TABLE 1 
Zonation of the salt marsh at La Salina. 
Predominant 
Zone Surface Plants Coleoptera 
Water Algae Berosus metaliceps Sharp 
Tropisternis salsamentus Fall 
I Muddy sand Sparse Salicornia Ochthebius interruptus LeConte 
Ochthebius sp. near rectus LeConte 
Carpelimus confinis Casey 
Carpelimus debilis Casey 
Bledius punctatissimus LeConte 
Bledius ornatus LeConte 
Thinobius frizzelli Hatch 
II Salt crust underlaid Dense Salicornia Bembidion ephippigerum LeConte 
with one inch of Tachys vittiger LeConte 
muddy sand Carpelimus confinis Casey 


Bledius ferratus LeConte 
Bledius punctatissimus LeConte 
Thinobius frizzelli Hatch 
Actidium sp. 


III Salt crust underlaid Distichlis, Salicornia, — Bledius ferratus LeConte 


with muddy sand numerous others 
IV Not investigated Juncus Not investigated 


The marsh area may conveniently be divided into several zones, as shown in figure 1. A 
surface profile of Zones I and II also is shown in figure 1. Table 1 lists the more noticeable 
characteristics of each of the zones. 


Zone I was a narrow strip of the immediate shore of the lagoon, extending eastward along 
its northern shore from the extreme western terminus. It was composed of dark, muddy sand 
without a salt crust. It supported a sparse growth of Salicornia, which in some places over- 
hung the bank into the water. It contained a very rich fauna of staphylinids. In 1959, I found 
there only Thinobius frizzelli, which was, however, present in great numbers. In 1961, Bledius 
punctatissimus was the predominant species, with some Thinobius frizzelli, Carpelimus debilis, 
and Carpelimus confinis. Specimens were readily collected in this area by pouring water on 
the sand, a technique which brought the individuals rapidly to the surface. They were then 
easily picked up with tweezers and dropped into alcohol. The north shore of the lagoon to the 
east of this area gradually changed to sticky mud, in which no insects were found. The south- 
ern shore of the lagoon, which was used as the local trash dump, was not investigated. 


Zone II was an extensive salt-encrusted flat which covered most of the central part of the 
marsh. It became quite narrow at the western section of the marsh, adjacent to Zone I. It 
supported there a lush growth of Salicornia but no other plants. In this area the surface con- 
sisted of a hard salt crust from one thirty-second to one sixteenth of an inch thick, underlying 
which was a one-inch layer of muddy sand, and below that a base of fine sand of undetermined 
depth, but at least twelve inches deep. The predominant insect fauna was found in the top 


274 San Dreco Society oF Natura History {Vor13 


inch of muddy sand, but some specimens were found as much as four inches below the surface. 
The most abundant insect here was Bledius ferratus (adults and larvae), with numerous Bem- 
bidion ephippigerum and Tachys vittiger. Also present in lesser numbers were Thinobius friz- 
zelli and Carpelimus confinis. Pouring water on the surface, so effective for collecting in Zone 
I, produced no results in Zone II, as the surface was nearly impenetrable. Placing a shovel-full 
of mud and sand in a bucket of salt water (the only water convenient) and agitating slightly 
caused the insects to come to the surface of the water in a minute or two. They were then 
lifted from the surface with a small strip of window screen, which was dipped into a jar of 
alcohol to dislodge them. Later, when the specimens were removed from the alcohol, they were 
coated with a sticky, white precipitate, probably caused by combination of the salt water and 
alcohol. This was easily removed by transferring them to a vial of benzine for a few minutes 
and then for a short time back to alcohol before mounting. In the broad central expanse of 
Zone II, the Salicornia was sparse and less vigorous. Although the small piles of castings which 
indicate the presence of Bledius occurred there, I was unable to find specimens. The indications 
were that Bledius ferratus had inhabited the entire area earlier in the year. 


Zone III was a foot or two higher than Zone II. Its plant cover was much denser and 
more varied than that of Zones I or II, being a mixture of Distichlis, Salicornia, and other 
plants. Indications of Bledius occurred here in the few open spaces, but they all appeared dry 
and old. I was unable to find any staphylinids in this zone in the summer of 1961. 


Zone IV, in which the predominent plants were dense clumps of Juncus acutus, was not 
investigated. 


The highly saline water area of the lagoon at La Salina occupied, in the summer of 1961, 
a narrow winding channel in the western half of the area. 


I visited La Salina again early in March of 1962, after a month of heavy rainfall, and 
found Zones I and II completely submerged. Zone III was muddy in places, and almost all 
of its plant cover was brown and dormant. At the outer edges of Zone III were a few damp 
open spaces without a salt crust, in the black sandy mud of which I found numerous adults 
of Bledius ferratus. 


Early in 1963 a causeway was under construction across the center of the marsh to accom- 
modate a new four-lane toll road from Tijuana to Ensenada. The water in the lagoon was being 
pumped out for use in the highway construction. 


COLEOPTERA OF THE MARINE SALT MARSHES 


The following list includes only those species of beetles which are known to occur in the 
salt water (the Hydrophilidae) or in the adjacent saturated sand and mud (all the others). 


Carabidae Limnebiidae 
Bembidion ephippigerum LeConte Ochthebius interruptus LeConte 
Tachys vittiger LeConte Ochthebius sp. near rectus LeConte 
Hydrophilidae Staphylinidae 
Berosus metalliceps Sharp Carpelimus confinis Casey 
Tropisternis salsamentus Fall Carpelimus debilis Casey 
Enochrus hamiltoni Horn ssp. Carpelimus salinus n. sp. 
pacificus Leech Bledius diagonalis LeConte 
Limnichidae Bledius ferratus LeConte 
Throscinus crotchi LeConte Bledius ornatus LeConte 
Ptiliidae Bledius punctatissimus LeConte 


Actidium sp. Thinobius frizzelli Hatch 


1964} Moore: STAPHYLINIDAE OF MARINE Mup FLats 27D 


Many other species of Coleoptera are restricted to the outer drier areas of this environ- 
ment. Some, such as Cicindela gabbi, are found on the open dry spaces at the edges of bays 
and estuaries; others, such as Amara jacobina, are found under cover in these areas, and still 
others are associated with plants which are limited to this region. Many other Coleoptera which 
are normally found outside this area invade the outer regions when conditions are favorable. 


STAPHYLINIDAE OF MarINE MupFLatTs 


The known staphylinid fauna of marine reefs of southern California is composed entirely 
of members of the subfamily Aleocharinae. That of sandy beaches includes members of four 
subfamilies: Aleocharinae, Staphylininae, Omaliinae, and Oxytelinae. That of marine mudflats 
is composed entirely of members of the subfamily Oxytelinae. The subfamily Oxytelinae is 
distinguished from all other Staphylinidae by the presence of a complete second sternite, this 
sclerite being absent or rudimentary in members of the other subfamilies. The three genera 
found in the salt marshes of this area all belong to the tribe Oxytelini, characterized by tarsi 
of fewer than five segments. The tarsi of the Oxytelini are usually said to be three-segmented, 
but some species of Bledius have four segments and Thinobius has only two. 


Genus Bledius Leach 


Members of this large genus are found throughout the world, being particularly abundant 
in temperate regions. The larvae and adults inhabit burrows in the sandy or muddy margins 
of streams and ponds. Many species are known from salt marshes in various parts of the 
world. Four species are recorded from the marine salt marshes of southern California. Bledius 
jacobinus LeConte (1877:220) is closely related to one of these, B. ferratus LeConte, and is 
probably also a halobiont. Its type locality, San Diego, California, may be in error. It is 
unknown to me. Other species of Bledius occur in the decaying seaweed on the sandy beaches 
of the Pacific coast of North America. The adults are often attracted to lights. This genus, 
much in need of revision, is being studied by William Steel of Maidenhead, England. 

Bledius ferratus LeConte (fig. 2a).— Color variable, from dark to light brown, with the 
legs and usually the antennae and elytra paler. Head and pronotum sparsely, elytra moderately, 
and abdomen very sparsely, punctured. Integuments reticulate throughout, the elytra more 
feebly than the other parts. Surface of head with a central tubercle, longitudinally impressed 
from the tubercle to the base; mentum concave, its basal margin straight; gula deeply impressed. 
Sides of pronotum with a lateral margin; longitudinal central line well-impressed; lateral pro- 
sternal suture absent; hypomera concave. Length 5 to 7 mm. 

LeConte Secrbed this species from San Diego. I have found it at La Salina and Laguna 
Santa Maria, Baja California, Mexico. At La Salina, larvae and adults were common in Zones 
II and III, whexe they probably feed on the larvae and pupae of a fly that was abundant in 
the mud. 

This species belongs to the mandibularis group of LeConte, of which he wrote (1877:218) : 
“The species of this group resemble each other very closely and might be readily confounded, 
if it were not for the differences in the gular process, and sexual characters. In the neighbor- 
hood of the ocean they live in salt marshes, and their occurrence on both sides of the continent, 
with a closely allied form in the interior, indicates an origin and distribution similar to that of 
species of various genera already mentioned by me. I have interpreted such cases to be unmodi- 
fied, or (in this instance), slightly modified survivals of ancient forms and probably from the 
Cretaceous period.” To suggest such an ancient ancestry as the Cretaceous (more than sixty 
million years) for species so closely related appears unnecessary. LeConte probably speculated 
that the Cretaceous period, the time of the last submergence of the continent, was the final 
opportunity for these species to become widely disseminated. However, these beetles are active 
fliers, often being taken at lights, and could readily be carried long distances by strong winds. 
During the late Cenozoic period, the continent was dotted with undrained saline basins which 
would have provided excellent stepping stones for halobionts. It is probable that the winged 
halobionts were widely distributed during the arid Miocene and Pliocene epochs and then 


276 SAN D1EGo SocreTy OF NATURAL HIstory 


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1964] Moore: STAPHYLINIDAE OF MARINE Mup EFiats 277 


were reduced in number and contracted into isolated pockets during the Pleistocene for a long 
enough time to produce separate but closely related species. 

Bledius ornatus LeConte (fig. 2b) .— Head, thorax, and abdomen black; elytra black with 
an irregular testaceous maculation at the outer apical nacle epipleura pale; antennae piceous, 
with the basal and apical segments darker; legs testaceous. Head and pronotum moderately 
punctured, with the interspaces reticulate. Elytra more closely and somewhat more coarsely 
punctured, shining. Abdomen sparsely punctured, feebly reticulate. Surface of head evenly 
convex, the mentum flat. Pronotum with a lateral margin and a central longitudinal groove, 
the hind angles distinct. Fissure of anterior coxal cavity long, open, widely exposing the 
trochantin. Lateral posternal suture distinct. Hypomera concave. Length 4 mm. 

The color pattern of the elytra varies greatly in extent, the elytra in some specimens being 
almost entirely dark and in others having only a basal dark band. Many intergrades occur. 

The species has been reported from British Columbia to southern California. I have taken 
it also at Rosarito Beach and La Salina in Baja California. 

In my experience, this species is always restricted to a very narrow habitat, where, how- 
ever, adults and larvae can often be found in large numbers. I have found it nowhere but in 
the sand of the landward shore of a sandbar which separates an estuary fromthe ocean (Zone 
I at La Salina). Splashing or pouring water within a few inches of the water’s edge often 
brings myriads of these insects to the surface. Fall and Cockerell (1907:168) reported this 
species from Santa Fe Canyon, New Mexico, but considering the specialized habitat, this record 
is probably based on a misidentification. 

LeConte stated that this species may be related to B. albonatus Maklin, but the description 
of the latter mentions that the legs are dark. 

Bledius punctatissimus LeConte (fig. 2c). — Body black, with the legs and antennae pale. 
Head and pronotum densely, very finely, punctured and densely reticulate. Elytra densely, 
more coarsely punctured, shining. Abdomen moderately punctured, feebly reticulate. Surface 
of head evenly convex; mentum flat. Pronotum with rounded hind angles, lateral margins dis- 
tinct, and a very fine central longitudinal line; fissure of anterior coxal cavity long and closed, 
the trochantin concealed. Lateral prosternal suture distinct. Hypomera rectangular, concave. 
Length 3 mm. 

This species was described from one specimen from “Southern California.” LeConte 
(1877:226) stated, “Another specimen . . . from Florida. This locality is perhaps doubtful; 
if correct it indicates a very unusual distribution.” I have seen it from Carlsbad, California, 
and La Salina and El Estero, Baja California. At La Salina it was very abundant in Zone I 
and was taken occasionlly in Zone II. 

Bledius diagonalis LeConte.— This species is unknown to me. LeConte placed it near 
ornatus, but stated (1877:230) that the elytra were “dark, dirty yellow, with a large common 
triangular spot extending from the humeri nearly to the tip of the suture blackish; epipleura 
fuscous.” Lengh 4.7 mm. 

This species was based on a single specimen collected by LeConte (1877:230) at “San 
Diego, Cala., in salt marsh.” To my knowledge, it has not been found since. 

Genus Carpelimus Leach 

Members of this moderately large genus are found throughout the world, and, like 
Bledius, are most common in temperate regions. The species are mostly of monotonously small 
size and somber color, the relatively minor specific differences between many of them rendering 
identifications tedious and difficult. The habitat is similar to that of Bledius, the two genera 
often being found together. 

Carpelimus debilis Casey (fig. 3a).— This species is very similar to C. confinis. Color 
dark, with the legs a little paler. Integuments finely and densely punctured throughout. Tenth 
antennal segment about as long as wide. 

Carpelimus debilis was described from “California (San Diego) . . . abundant on banks 
of wet ditches near the town” (Casey, 1889:375). The ditches mentioned may have been 
brackish connections with the bay. I have taken this species under partly dried mats of Ulva 


278 San Dieco Society oF Natura History { VoL, 13 


on the shores of San Diego Bay and Mission Bay. I have also collected it from the muddy 
sand of the shores of Buena Vista Lagoon, San Diego County, California, and La Salina and 
Estero Beach, Baja California. At La Salina, I found only two specimens, in Zone I, whereas 
C. confinis was present there in great numbers. 

Carpelimus confinis Casey (fig. 3c).— Very similar to C. debilis. Dark, with tarsi paler. 
Integuments finely, densely punctured except for a small, narrow area at the center of the 
pronotum. Tenth antennal segment transverse. 

This species and C. debilis are so similar that they might easily be confused. Examination 
of a long series of each has convinced me that there are no intergrades between the two, the 
few specific differences being constant. It was described from “California (San Diego)” (Casey, 
1889:369). I have found it at Buena Vista Lagoon, San Diego County, California, and at 
La Salina, El Estero, and Laguna Santa Maria, Baja California. 


Carpelimus salinus, new species 

Holotype. — From El Estero, seven miles south of Ensenada, Baja California, Mexico, 
April 12, 1959; Ian Moore, collector. In the California Academy of Sciences (Entomology). 

Paratypes. — Seven specimens from El Estero, Baja California, Mexico; one taken on 
April 12, 1959, one on April 22, 1959, and five on May 1, 1959, by Ian Moore. In the Cali- 
fornia Academy of Sciences, the San Diego Natural History Museum, and the collections of 
Melville Hatch and Ian Moore. 

Form. — Rather slender, subcylindrical (fig. 3b). 

Color. — Largely black, with the femora piceous, the tibiae, tarsi, and posterior coxae pale. 
Pubescence fulvous, much longer and more conspicuous on the abdomen than elsewhere. 

Head. — Slightly narrower than pronotum, minutely densely punctured, the neck trans- 
versely strigulose. Eyes moderately prominent, coarsely faceted; tempora shorter than the 
eyes, parallel, not narrowed behind. Antennae shorter than the head and pronotum, the second 
segment shorter than the first, the third a little shorter than the second, longer than wide, the 
fourth as wide as the third, one-half as long, not longer than wide, the fifth to tenth progres- 
sively increasing in width but not in length, the eighth to tenth distinctly transverse, the eleventh 
as long as the preceding two together. 

Pronotum. — A little wider than long, widest near the middle, only very slightly narrowed 
to the rounded apical angles, gradually narrowed from behind the middle to the base, which 
is distinctly narrower than the apex, the sides distinctly but not strongly sinuate before the 
hind angles. Surface evenly convex except for a very feeble impression near the base on each 
side. Evenly, very densely, finely punctured throughout except for a narrow, elongate, central 
smooth area, the punctures separated by less than their diameters. Interspaces finely, densely 
microreticulate. 

Elytra. — Wider than long, not longer, but distinctly wider, than the pronotum, convex 
except for a distinct but feeble sutural impression in the basal half. Evenly and densely punc- 
tured throughout, the punctures twice as large as those of the pronotum, separated by less than 
their diameters. Interspaces shining. 

Abdomen. — About as wide as the elytra, widest at about the fourth visible tergite. Very 
finely and rather sparsely punctured, the punctures smaller than those of the pronotum, sepa- 
rated by three to four times their diameters. Interspaces finely, evenly microreticulate. 

Length 2 mm. Sex unknown. 

Remarks. — The specimens were taken from beneath pieces of moist paper in a path 
through dense Salicornia, just above the reach of the normal daily tides. This would correspond 
with Zone II at La Salina. 

This species is quite distinct from the other two salt marsh species of this area in its 
relatively shorter and wider elytra. In Casey’s (1889) key to the Nearctic species it will run 
to C. providus, from which it differs by the presence of the median impunctate line of the 
pronotum, by the sides of the pronotum being sinuate before the hind angles, and in other 
details. 


1964} Moore: STAPHYLINIDAE OF MARINE Mup FLats 279 


2a, Gea % # 
cy A 
= we 
<i) | an 


=| 


~ b 
0000 0000. 50000 
Q /) 
LAE 
U\ “N\ 
d 


Fig. 3. Outline drawings of imagos. a, Carpelimus debilis; b, C. salinus; c, C. confinis; 
d, Thinobius frizzelli. Bar represents 1 mm. 


280 San Dreco Society oF NATuRAL History {VoL. 13 


Genus Thinobius Keissenwetter 


The members of this genus are usually minute slender beetles found in the same type 
of habitat as Bledius and Carpelimus. As with Carpelimus, many of the species so closely 
resemble each other as to require very careful study to distinguish them. They are of world- 
wide distribution. Kincaid (1961) called attention to the fact that the two-segmented condition 
of the tarsi has generally been overlooked by American students. 


Thinobius frizzelli Hatch (fig. 3d).— This species was described from Washington and 
British Columbia. An excellent illustration was given by Hatch (1957). Kincaid (1961) gave 
good figures of the adult and larva, with ecological notes. I found a few specimens on the salt 
marsh at Torrey Pines, San Diego County, California, and great numbers in Zone I at La 
Salina, Baja California. 


Key TO THE STAPHYLINIDS 


The foregoing eight species can be distinguished by the following key. 
la. Elytra dehiscent. 


2a. Anterior tibiae not spinose externally (Thinobius) 200.000.000.000 T. frizzelli 
2b. Anterior tibiae spinose externally (Bledius). 
3a. Hypomera not separated from the pronotal disc by a carina.................. B. ferratus 
3b. Hypomera separated from the pronotal disc by a carina. 
fae gelytralvepipleura yellow: .cc nce ..ccuctn chee eee ee ONAL 
4b. Elytral epipleura dark. 
Namely era CO Ore dr san. sin de tp iher Bac ata ae ae eer B. diagonalis 
Sbswblyragdarksunicolored o.c.c6) ae Meee B. densissimus 
Ib. Elytra not dehiscent (Carpelimus). 
6a. Disc of pronotum evenly punctured throughout... ees C. debilis 
6b. Disc of pronotum with a smooth central space. 
7a. Elytra one and one-half times as long as pronotum............................... C. confinis 
7p elytcanvery Wetlevlonger: than) promotumier. 2... sec ae eee C. salinus 
LARVAE 


The larvae of possibly less than one per cent of the known species of Staphylinidae have 
been described, most of these from Europe. The number of described larvae of the Nearctic 
species would be very small except that the larvae of some of the Holarctic species have been 
made known in the European literature. This neglect is due in part to a general lack of interest 
in this large and difficult family, but more to the difficulty of associating the larvae and imagos 
for proper identification. Difficulties in rearing members of this family have been known for 
a long time. A unique opportunity to associate larvae with known adults occurs in certain 
restricted habitats such as with the inquilines, parasites, cave-dwellers, and marine species. It 
has been my good fortune to obtain larvae of all but three of the known Staphylinidae of the 
southern California salt marshes under circumstances permitting almost positive identification. 
Parts of these larvae are illustrated in figure 4. 


The key which follows will readily distinguish the larvae of the species of these salt 
marshes from one another, but, in view of our limited knowledge of other species of these 
large genera, it may not suffice to separate them from other larvae when these become known. 
The characters used to distinguish the three genera in the key will probably be found to be 
unsatisfactory as the knowledge of these genera increases. An attempt to define the subfamily 
Oxytelinae on the basis of larval characters would be fruitless at the present time. 


1964} Moore: STAPHYLINIDAE OF MariNnE Mup FLats 281 


Qa 
a 
ass” 
@ 


Se >e/L- \WV 


Fig. 4. Larval parts of staphylinids. a-c, Bledius ferratus: a, mandible; b, antenna; c, 
urogomphus. d-f, Bledius ornatus: d, mandible; e, antenna; f, urogomphus. g-i, Bledius puncta- 
tissimus: g, mandible; h, antenna; i, urogomphus. j-l, Carpelimus confinis: j, mandible; k, an- 


tenna; I, urogomphus. m, urogomphus of C. debilis. 


282 SAN Digeco Society oF NaturaAL History | Vou. 13 


Key TO SOME STAPHYLINID LARVAE 
la. Modified seta of apex of second antennal segment slender, several times as long as wide 
(Carpelimus). 
Zac eogonp USystralGlitte. ...4 ede.) sacutaisen ty. soak ice Pena ee eo enact tke ee C. confinis 
2b. Wrogomphus abruptly hooked: at apex c.0 2.1020. 00006 eae ee Joineeeeandebilas 
lb. Modified seta of apex of second antennal segment stout, almost as wide as long. 


3a. Apex of mandible with seven subequal denticles internally (Thinobius) ......T. frizzelli 


3b. Apex of mandible with two prominent teeth, sometimes with one to three short inter- 
mediate denticles (Bledius). 


4a. Mandible without denticles between apical teeth, 
thestecth serrulatemternally eo Cee ee ee  B. ferratus 


4b. Mandible with denticles between apical teeth, the teeth entire internally. 


5a. Apex of urogomphus abruptly narrowed to a parallel-sided process bearing 
two setae at the (St Nise eat te eer e Roney reer: ae Mens 2 eRe B. punctatissimus 


5b. Apex of urogomphus gradually conical to a tip bearing a single seta ....B. ornatus 
ACKNOWLEDGMENTS 


I am indebted particularly to the following people for assistance with this paper: Hugh 
B. Leech, of the California Academy of Sciences, for identification of the water beetles and for 
numerous other favors; Trevor Kincaid, for specimens of Thinobius frizzelli and other marine 
species; Melville Hatch for a paratype of Thinobius frizzelli and other useful material; Henry 
Dybas, for identification of the ptiliid; and Charles Harbison, for help with the plant names. 


1964 } Moore: STAPHYLINIDAE OF MARINE Mup FLats 283 


LITERATURE CITED 


CaseY, THOMAS L. 
1889. A preliminary monograph of the North American species of Trogophloeus. Ann. 
New York Acad. Sci. 4:322-383. 


Ariat, 
1901. List of the Coleoptera of southern California, with notes cn habits and distribu- 
tion and description of new species. Occas. Pap. Calif. Acad. Sci. no. 8. 282 pp. 


Fatt, H. C., AND T. D. A. COCKERELL 
1907. The Coleoptera of New Mexico. Trans. Amer. Ent. Soc. 33:145-272. 


Hatcu, MELVILLE 
1957. The beetles of the Pacific Northwest. Part II. Staphyliniformia. Univ. Wash. 
Publ. Biol. 16(pt. 2). xii + 384 pp. 


KINCAID, TREVOR 
1961. The ecology and morphology of Thinobius frizzelli Hatch, an intertidal beetle. 
15 pp., 6 pls. Seattle. 


LeConreE, JOHN L. 
1877. On certain genera of Staphylinidae Oxytelini, Piestidae and Micropeplidae as 
represented in the fauna of the United States. Trans. Amer. Ent. Soc. 6:213-252. 


Moore, IAN 
1956a. A revision of the Pacific coast Phytosi with a review of the foreign genera (Coleop- 
tera: Staphylinidae). Trans. San Diego Soc. Nat. Hist. 12: 103-152. 


1956b. Notes on some intertidal Coleoptera with descriptions of the early stages (Cara- 
bidae, Staphylinidae, Malachiidae). Trans. San Diego Soc. Nat. Hist. 12:207-230. 


: Z ~~ 
fel} AP, 
Srikiids 


se en ae 


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| yeaa: 4 Bais re A, =, { v7 4. 
we , y ; 
e. , nie Pe: f a 5 hay ae : . i Le 
i ee 
Weal a flap a aT diy Ai hy ¥ ere a i Ly ¥ ei? ee 


+ 
' ‘ (ie? Gy tg hie i) Wig ’ Ypo tse es x 


UO a alion alte iene a, He Wt oa} 
’ 


PAS A Sete cuabrie. Gel Wsterisgoebead te “goth! 
a9, lela! eh fee ack ten Me 


° § ‘ 4 j : vty 
plates obi Et eo Sey 
A Fi er - ; a} j “1 ph Z : 
Byer Leia: amma STA Ay, we (3 F i nok . ue ei te ES hate ‘itt a wD 
a i es hd > , - Cm ea — > tre % ;: ; : Hi ; 
a i ae any 4 ( *) pel ‘ oe Rite ; p 
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v . TA iv) - “! ? we ¥) it 
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Ae é i er E eee be Mi 
ae : ; “f ) x i 
iv wi fo y/ “<i - ‘ M4 ‘< 
re! Dea. ree PUG e fehl Ped MRS OSs wit kes pte 
a i } A € , 
= Ade w2P 
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’ ev iy} % ‘ 
a | wk > ,. ( bad") : j wis X 4, - song a ; 
ao a “aA f . ‘ wy ay 
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haa te 4 say - Awa ast be leegingeen Siu Jyelgy? 4 eis “ ri sla Th) wababs aD 
OP te dS 6th eS ese oe ONG. ove ae! ai sa bares Nhe rt BONE, * 
ere bs ¢ ' | 


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at ee ' es 


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FFB 8 1965 


HARVARD 
UNIVERSITY 


TRANSACTIONS 
OF THE 


SAN DIEGO SOCIETY “OF NATURAL-HISTORY. 


VoLuME 13, No. 13, pp. 285-300 


THE STRUCTURE AND REPRODUCTION OF THE 
RED ALGA CHONDRIA NIDIFICA HARVEY 


BY 


E. YaLE Dawson 
Director, San Diego Natural History Museum 
AND 


BiLtciIn TozuNn 


Department of Pharmaceutical Botany and Genetics 
University of Istanbul, Turkey 


SAN DIEGO, CALIFORNIA 
PRINTED FOR THE SOCIETY 


AucGustT 28, 1964 


th 


me: | Z 
‘ ‘ f 
A a ee es = 
Fig. 1. Chondria nidifica Harvey. 


A. Habit of a male plant. X 1. B. A branch of the same, showing terminal spermatangial tufts. 
% 3. C. Upper part of a tetrasporangial plant to show clustered determinate fertile branchlets. * 1. 


LIBRARY 
FFB 8 1965 


THE STRUCTURE AND REPRODUCTION) OF THE 
RED ALGA CHONDRIA NIDIFICA HARVEY’ 
BY | 
E. YaLE Dawson AND BiLcGIN TOZUN 


Chondria nidifica was described by Harvey in 1858 in a supplement to his final part of 
Nereis Boreali-Americana. The specimen had been collected by Dr. Schott during the Mexican 
Boundary Survey, but no data regarding the habitat accompanied the specimen. The type 
locality, thus, is unknown, although it is presumed now to be in the vicinity of San Diego, 
California, near the Mexican boundary. Harvey’s illustration, which is the very last in his 
book of North American algae, shows a mature tetrasporangial axis with its characteristic tufts 
of sporangial branchlets. This figure has served for recognition of the plant for a century. 


Chondria nidifica does not appear in the list of marine algae of San Diego, California, 
prepared by Daniel Cleveland in 1885 for Orcutt’s Flora of Southern and Lower California, 
but it was listed by C. L. Anderson (1891) as of southern California occurrence. Farlow’s 
earlier references (1875, 1876) merely followed Harvey, for Farlow apparently had not seen 
any specimens. J. G. Agardh, too (1863:801), reiterated Harvey and discussed the original 
description in connection with his transfer of the species to Chondriopsis. Later, Agardh 
(1892:161) discussed Harvey’s plant again, remarking on the peculiarly tufted tetrasporangial 
stichidia, but mentioned no new materials. 


Apparently, the second collections of the plant were made by Mrs. E. Snyder at Pacific 
Beach, near San Diego, California, in the late 1890’s. McClatchie (1897:359) indicated her 
collaboration in field work, and mentioned that in southern California the plant is “frequent on 
rocks and in sand between tide marks.” Snyder was also a collaborator of Collins, Holden and 
Setchell in the preparation of Phycotheca Boreali-Americana, and her Pacific Beach materials 
appear as specimen number 841 in volume 17 of 1901. The examples we have seen in various 
sets of this publication are all tetrasporic. The first sexual materials to be mentioned in the 
literature are those numbered 1646 in volume 33 of Phycotheca Boreali-Americana of 1910, 
for which Snyder contributed cystocarpic material from Pacific Beach. 


Subsequent to the collections issued in P.B.A. the plant is scarcely mentioned in the 
literature except as the host of the parasitic Janczewskia lappacea Setchell (1914:14). Collec- 
tions had been made repeatedly by N. L. Gardner in the vicinity of San Pedro, California, 
and a set of these was issued in 1916 as P.B.A. number 2146, representing the parasitic plant. 


So far as we are aware, no formal description of Chondria nidifica, apart from Harvey’s 
original account of a tetrasporic individual, appeared until Dawson (1963) reported male 
plants for the first time and pointed out the dimorphism of sexual and asexual plants (fig. 1). 


Since 1945, Dawson has given several collection records for the plant in Mexico and in 
southern California (Dawson, 1945, 1948, 1951, 1954, 1959; Dawson, Neushul, and Wildman, 
1960). The most recent report of its range (Dawson, 1961) is from Government Point, Santa 
Barbara County, California, to Bahia Asuncion, Baja California. 

The species is characteristically an inhabitant of the lower intertidal zone on surfy shores 
provided with small cobbles and stones embedded in sand. It is commonly associated with 
Gracilariopsis andersonii and Gracilariopsis sjoestedtii, which also are commonly found on 
stones partially embedded in sand. In such habitats Chondria nidifica is often exceedingly 
abundant. Indeed, in southern California, from La Jolla to Point Conception, the species 
was found by Dawson (1959) to be frequent to abundant at a majority of 42 intertidal survey 


1This study was supported in part by a contract Nonr 228(25) between the Office of Naval Research and 
the University of Southern California. The junior author was supported by the Committee on International 
Exchange of Persons. 


288 SAN Dieco Society oF Naturat History [Voras 


stations, and the twelfth most frequently encountered benthic alga of the region. In the south- 
ern part of its range it becomes less conspicuous, and south of Punta Baja it is confined to 
areas affected by cool upwelling water. 

Detailed examination of the structure and reproduction of a species of Chondria was first 
carried out by Falkenberg (1901), although Phillips (1896) had made some comparative 
studies of procarp and early cystocarp development in Chondria tenuissima. Falkenberg dealt 
principally with Chondria dasyphylla and included the first account of spermatangial develop- 
ment for the genus. Kylin (1928) followed with a thorough review of the same species insofar 
as tetrasporangial and carposporic plants were concerned, but he gave no further account of 
spermatangial development. The principal knowledge, thus, of the structure and reproduction 
of Chondria is based upon study of a single species in the subgenus Coelochondria. The 
available data have recently been incorporated by Hommersand (1963) in his study of the 
classification and evolution of the Ceramiaceae and Rhodomelaceae. 

Gross morphological treatments of various species of Chondria have been presented, mainly 
incidental to taxonomic and floristic accounts such as those of Bérgesen (1918) and Dawson 
(1963), but structural details are generally lacking. Chondria nidifica has been treated in this 
manner by Dawson (1963) in comparison with six other Chondria species along California 
and Pacific Mexico, and it is an expansion of that descriptive study that is here undertaken. 

Material.— The specimens used in this study were collected October 7, 1963, at 
the seaward end of Sunset Boulevard on Santa Monica Bay, Los Angeles County, California. 
The species was growing abundantly at about the + 0.5 foot tide level in sandy pool areas 
among small cobbles. All reproductive phases were present in approximately equal proportions, 
and the plants generally exhibited active growth. 

Most of the studies were made using sections prepared with the freezing microtome, 
stained with aqueous aniline blue, and illustrations were drawn with the camera lucida and 
microprojector. 


VEGETATIVE DEVELOPMENT 


Chondria nidifica is a laxly branched, coarse species. It reaches a height of 15 cm. under 
average conditions, but occasional or exceptional plants 20 to 40 cm. long may be found. The 
species is perennial and regenerates for several years from the bases of old axes above the 
fleshy holdfast, so that quite old plants may have a considerable basal tuft of branch stubs 
and their regenerative branches. The plant is subject to extensive grazing by inshore animals 
and commonly exhibits much proliferous branching from broken and torn ends. 

Well-developed young plants consist of a few to many erect, cylindrical axes from the 
basal disc. These are 1.0 to 1.5 mm. in diameter, rather remotely, irregularly, multifariously 
branched at intervals of 2 to 6 cm. The branches are terminally acuminate when in active 
growth (fig. 2, D). Trichoblasts are ordinarily too small and few and too early deciduous 
to be conspicuous. Older axes become increasingly coarse and may have a semi-rigid texture, 
sometimes reaching 2 mm. in diameter. 

The acuminate apex terminates in a prominent, capitate apical cell about 18 y tall and 
nearly as broad. The apical cell cuts off, usually by a concave wall, the first vegetative initial 
(fig. 2, A). This initial may first undergo a transverse division to form two broad, flat cells, 
or it may form directly a lateral protuberance after which an oblique wall cuts off the first 
trichoblast initial. These initials are cut off successively from each segment of the apical cell 
in an approximate 2/7 left hand spiral divergence. The trichoblast initial usually divides once 
before the first pericentral cell is cut off from the axial cell immediately at its base. The re- 
maining four pericentral cells are cut off within about 7 segments of the apex. At this point 
the trichoblasts have developed into 7- to 8-celled filaments and have acquired branches 
(fig. 2, A). The pericentral cells continue to divide to build up the cortical tissue. Mean- 
while, the axial cells enlarge, assuming first a somewhat quadrate form (as seen in longisection) 
followed by progressive axial elongation. During this cortical enlargement the basal cell of the 
trichoblast, with its prominent connection to the axial cell, becomes elongated in keeping with 
the cortical expansion, so that the second trichoblast cell always remains at the surface. The 


1964} Dawson AND TO0zUN: Chondria nidifica 289 


trichoblasts are deciduous and break away usually within less than 1 mm. of the apex, leaving 
the first trichoblast cell as a slender scar-cell which extends through the cortex and is provided 
near the base with several secondary pit connections to surrounding cells. 

The five pericentral cells in part retain their identity in younger parts of the axis. In 
transections at some levels near branch tips they may be recognized by their large size and 
prominent pit connections to the axial cell. At other levels only smaller progeny cells may be 
evident. Prominent intercellular spaces are present in these juvenile parts of the axis (fig. 2, B). 
Continued division of the pericentral cells and their derivatives gives rise to a parenchymatous 
inner cortex (medulla) and a palisade-like outermost cortical layer (fig. 2, A). In older 
parts the axial cell row remains prominent (to 100 j: in diameter) and is surrounded by first 
an irregular layer of smaller parenchymatous cells, then by a medullary tissue composed mainly 
of large, thin-walled cells interspersed with a few smaller ones (fig. 3, A). The outer cortex 
loses its palisade-like character by tangential division of the cells into small, angular, deeply- 
pigmented cells 15-20 « in diameter. These form a compact, smooth surface layer (fig. 3, B). 

In fully mature areas the walls of the medullary cells are about 7 1 thick and there 
are thicker points of juncture between the cells (fig. 3, A). There is some stratification evident. 
Some of the intercellular spaces noted in young parts become filled with a clear matrix at 
maturity. Others are filled by the smaller progeny cells of late development. 


ORIGIN OF BRANCHES 


After the falling away of the trichoblast just back of the apex, the scar-cell remains as a 
prominent circular structure, sometimes with a clear indication of the former pit connection 
and pit membrane (fig. 3, D). In younger or slender secondary branches these scar-cells may 
remain unmodified throughout the length of the branch (fig. 3, C), whereas in larger, primary 
axes subject to heavy cortical development the scar-cell may soon lose its identity by division 
and overgrowth of the outer cortical cells that at first usually form a rosette arrangement 
around it (fig. 3, E). 

The relatively distant and infrequent branches in Chondria nidifica seem not to arise 
ordinarily near the apex of an axis, but well below it. The branch initial may arise directly 
from the distal side of the scar-cell a little below its end and before cortical overgrowth has 
been completed (fig. 2, C). In other instances the branch initial may arise later, after cortical 
overgrowth has obscured the scar-cell, and the young branch then may push out between the 
cells of the superficial layer. In still other instances, the cells immediately overlying the end 
of the trichoblast scar-cell seem to be capable of initiating one or more accessory branches 
and, remaining meristematic, of producing additional branches a second season after those 
of the first have eroded away in whole or in part. This is especially true of tetrasporangial 
plants, in which most of the branching is accessory and exogenous from the base of old or 
eroded primary branches. These regenerative branches are small, slender, determinate, mainly 
fertile ones which form the characteristic tufts on the axis of tetrasporic plants (fig. 1, C). 
Similar regenerative tufts develop from renewed meristematic activity of cells from broken or 
grazed ends of main branches. The determinate and proliferous character of the tetrasporangial 
branchlets is contrasted with the branches of sexual plants which are all of essentially the same, 
indeterminate kind. This accounts for the dimorphism commonly noted between sexual and 
asexual specimens (fig. 1). 


DEVELOPMENT OF THE PROCARP 


The branch tips of fertile female plants are generally more blunt than those of sterile 
and tetrasporic plants, and good developmental stages for procarps are best found on more 
slender, proliferous branches. Developing procarps are found immediately behind the dividing 
apical cell. They arise from trichoblast initials and are intermingled with sterile members in 
the spiral succession of trichoblasts. The sterile trichoblasts are distinct from the fertile ones 
at the youngest stages (fig. 4, A) and very early develop into the characteristic slender, 
branched, deciduous processes. The fertile trichoblast initial cuts off its first cell in the usual 
manner, but this cell becomes much enlarged and undergoes several successive divisions to 


290 San Dieco Society oF NAtTurAL History { VoL. 13 


form an adaxial enlargement at the apex of the short, bearing trichoblast (fig. 4, B-E). The 
basal cell by this time has divided to form the beginning of a pedicel for the growing cystocarp. 

At a later stage, but still within a fraction of a millimeter of the apex, the developing 
cystocarp is seen as a nearly sessile, irregularly hemispherical protuberance 60 to 70 y in 
diameter and exhibiting a prominent, projecting trichogyne about 50 « or more long. A 
median optical view of the cystocarp at this time reveals the procarp lying in a shallow groove 
on the adaxial side and surrounded by developing pericarp tissue except on the free side 
(fig. 4, F-G). The two axial cells within the protuberance are large and relatively little pig- 
mented. The supporting cell is situated just adaxial to the middle of the cystocarp and bears 
basally a sharply angled row of 3 to 5 sterile cells, and adaxially, but parallel to its longer 
dimension, the 4-celled carpogonial branch. The trichogyne is relatively massive on the small 
terminal carpogonium, and during development it shows a median swelling which later dis- 
appears. During its elongation the trichogyne membrane is thick in the lower part and thin 
at its lengthening tip. 

After fertilization, the trichogyne may remain intact for some time while the development 
of the pericarp continues adaxially. The result often shows a cystocarp advanced to incipient 
ostiole formation, but with the trichogyne remaining far displaced as a projection from the 
base of the pericarp. 

Early post-fertilization stages seem to follow closely those described by Kylin (1928) for 
Chondria dasyphylla, in which the supporting cell cuts off an auxiliary cell while the pericarp 
enlarges rapidly and the sterile cells multiply. The stage at which the auxiliary cell is clearly 
evident must be brief, for one sees in nearly all post-fertilization cystocarps a developing fusion 
cell (fig. 4, H) which involves the auxiliary cell, the supporting cell, the axial cell, and a 
number of the developing sterile cells. These sterile cells appear in branched series scarcely 
distinguishable from the similarly developing gonimoblast. Fusion and enlargement continues 
(fig. 5, B) in the developing gonimoblast until a large irregular mass is evident, from the 
lobes of which carpospores are produced successively (fig. 5, C). 

The appearance of maturing cystocarps at successive stages of enlargement is shown in 


figure 5, A. 
SPERMATANGIAL DEVELOPMENT 


The male reproductive structures of Chondria are so unusual in their fan-like form that 
they have attracted the attention of marine botanists for over a century. The first good illus- 
tration of these was given by Thuret (1851) for Chondria tenuissima; subsequently the male 
structures of various species have been described or illustrated by Buffham (1890), Falkenberg 
(1901), Borgesen (1918), and others. In all of these, the spermatangial sori are produced 
on the trichoblasts and take the form of rounded or elongate, thick, pedicellate fans. Similar 
structures occur in the genus Pterochondria. The fans of Chondria nidifica show close similarity 
to those of C. dasyphylla and C. tenuissima. Their development is as follows. 

Fertile spermatangial trichoblasts arise from the apex in the same manner as trichoblasts 
on sterile plants (fig. 7, C). The juvenile trichoblast at the 3-celled stage (at which time the 
pericentral cells are not yet all cut off) shows an adaxial bulge or protuberance of the second 
cell (fig. 6, A). This bulge cuts off the initial of a spermatangial branch while the sterile 
portion of the trichoblast continues to extend and to produce sterile branches. A second sper- 
matangial branch initial may be cut off slightly later from the second cell above the first, so 
that two spermatangial sori are produced from the same trichoblast (fig. 6). The first division 
of the male initial produces four cells oriented in a plane perpendicular to the main branch 
axis (fig. 6, B). Further development produces a gradual flattening of the structure in the 
same plane, and the appearance of a marginal row of initials (fig. 6, C-H). 

In these very early stages, the flattened spermatangial branch remains only 1 cell thick. 
However, soon after the stage shown in figure 6, G, some of the lower, primary cells of the 
branch cut off cells (usually obliquely) toward the outer flat surfaces. At the same time these 
ptimary cells begin to show an elongation in the direction of the length of the expanding, flat- 


1964} DAWSON AND T6zUN: Chondria nidifica 291 


tened spermatangial branch (fig. 6, E-F). These soon cease cutting off new cells, gradually 
become much elongated with the expansion of the spermatangial branch, and are transformed 
into the veins shown in figure 6, H. Meanwhile, the cells cutting off to either face of the blade 
divide repeatedly to produce its expansion, while the marginal initials add to the primary cells 
until they cease dividing, become enlarged, and form the marginal row of border cells (fig. 7, A). 

In the mature sorus the vein cells extend all the way to the border cells, and the super- 
ficial cells on either side become divided up into chains of small, angular, widely separated 
cells which cut off to the surface numerous ovoid spermatia (fig. 7, B). 


DEVELOPMENT OF TETRASPORANGIA 


Sporangial development in Chondria was first investigated by Falkenberg (1901), and 
his observations on C. dasyphylla were in part disputed by Kylin (1928) working on the same 
species. Chondria nidifica shows a somewhat different development than that worked out by 
Kylin, especially in later stages. 

Sporangial development is initiated very near the fertile apex by the successive cutting off 
of three cells from a very young pericentral cell. The first two are cut off more distally than 
the third, which comes to lie somewhat adaxial on the upper or inner side of the pericentral 
cell. The first two cells become the cover cells and the third becomes the sporangial initial. 
At the earliest stages at which these are readily distinguished in the fertile apex, the pericentral 
cell has already elongated somewhat and the cover cells are much enlarged compared to ordinary 
superficial cells (fig. 7, D). The cover cells, in fact, enlarge very rapidly, become somewhat 
inflated and falcately curved. The surrounding cortical cells arising from sterile pericentral cells 
do not develop a complete outer cover, and the developing sporangium and cover cells 
(fig. 7, E) lie in a kind of cavity seen as through a small window from the outside. The entire 
developing tetrasporangial branch retains extensive intercellular spaces during this enlargement, 
in the same manner as Kylin (1928) observed in C. dasyphylla. These spaces are filled later, 
in part, by the rapidly enlarging tetrasporangium which matures within one or a few mm. of 
the apex, where it is discharged (fig. 8, B). The discharge of the tetrasporangium occurs by 
an extrusion through the circular opening in the cortex, and the cover cells are left in the 
remaining cavity with a residue of the sporangial membrane lying on the now shrunken peri- 
central cell (fig. 8, C). 

The pericentral cell never becomes as markedly elongated as in Chondria dasyphylla, but 
it does consistently lack the development of secondary pit connections with surrounding cells 
(fig. 7, D). Furthermore, and unlike the condition reported by Kylin (op. cit.) in C. dasy- 
phylla, no additional cover cells or daughter cells are cut off, and the entire sporangial com- 
plex consists only of the tetrasporangium and the two conspicuously inflated cover cells at- 
tached by large pit connections to the pericentral cell. Neighboring pericentral cells, from 
which the vegetative tissue of the branch arises, show conspicuous secondary pit connections as 
well as a progeny of extensive overarching, branched filaments making up the cortex. 

The appearance of the mature, determinate tetrasporangial stichidia in their characteristic 


tufts is shown in figures 1, C, and 8, A. 


LITERATURE CITED 
AGarpH, J. G. 
1863. Species genera et ordines algarum. Vol. 2. pp. 701-1291. Lund. 
1892. Analecta algologica. Actis Soc. Physiographicae Lundensis 28. 182 pp., 3 pl. 
ANDERSON, C. L. 
1891. List of California marine algae, with notes. Zoe 2:217-225. 
BOrGESEN, F. 
1918. The marine algae of the Danish West Indies. 2. Rhodophyceae. Dansk. Bot. Ark. 
3 :241-304. 
BuFFHaAM, T. H. 
1890. On the reproductive organs, especially the antheridia, of some of the Florideae. 
Journ. Quekett Micr. Club II 4:246-253. 


292 San Disco Society oF Natura History {Vors 13 


CLEVELAND, D. 
1885. Marine algae of San Diego, California. In C. R. Orcutt, Flora of Southern and 
Lower California. San Diego. 
Cotuins, F. S., I. HoLpEN, AND W. A. SETCHELL 
1895-1919. Phycotheca Boreali-Americana. Fasc. 1-46, A-E. Malden, Mass. 


Dawson, E. Y. 

1945. Marine algae associated with upwelling along the northwestern coast of Baja 
California, Mexico. Bull. So. Calif. Acad. Sci. 44:57-71. 

1948. Resultados preliminares de un reconocimiento de las algas marinas de la costa 
Pacifica de Mexico. Rev. Soc. Mex. Hist. Nat. 9:215-255. 

1951. A further study of upwelling and associated vegetation along Pacific Baja Cali- 
fornia, Mexico. Jour. Mar. Research 10:39-58. 

1954. Resumen de las investigaciones recientes sobre algas marinas de la costa Pacifica 
de Mexico, con una sinopsis de la literatura, sinonimia y distribucion de las 
especies descritas. Rev. Soc. Mex. Hist. Nat. 13:97-197. 

1959. Oceanographic survey of the continental shelf area of southern California, pp. 
169-218. State Water Pollution Control Board, Sacramento, California. 

1961. A guide to the literature and distributions of Pacific benthic algae from Alaska 
to the Galapagos Islands. Pac. Sci. 15:370-461. 

1963. Marine red algae of Pacific Mexico. Part 8. Ceramiales: Dasyaceae, Rhodo- 
melaceae. Nova Hedwigia 6:401-481, 46 pl. 


Dawson, E. Y., M. NeusHuL, AND R. D. WiLpMAN 
1960. Seaweeds associated with kelp beds along southern California and northwestern 
Mexico. Pacific Naturalist 14:1-81. 


FALKENBERG, P. 

1901. Die Rhodomelaceen des Golfes von Neapel und der angrenzenden Meeres- 
Abschnitte. Fauna und Flora Des Golfes von Neapel und der angrenzenden 
Meeres-Abschnitte. Monographie 26. xvi + 754 pp., 24 pl. 

Fartow, W. G. 

1875. List of the marine algae of the United States, with notes on new and imperfectly 
known species. Proc. Amer. Acad. Arts & Sci. 10 [N. S. 2}: 351-380. 

1876. List of the marine algae of the United States. U. S. Comm. of Fish and Fisheries. 
Report of Commissioner for 1873-74 and 1874-75, pp. 691-718. 

Harvey, W. H. 

1858. Nereis Boreali-Americana. Part III. Chlorospermeae. Smithsonian Contr. to 

Knowl. Vol. 10, Art. 2. 140 pp., pl. 37-50. 
HomMErRSAND, M. H. 

1963. The morphology and classification of some Ceramiaceae and Rhodomelaceae. 

Univ. Calif. Publ. Bot. 35:165-366. 
KYLin, H. 
1928. Entwicklungsgeschichtliche Florideenstudien. Lunds Univ. Arsskr., N. F. 24(4) : 
T1275 

McCratcuig, A. J. 

1897. Seedless plants of southern California. Proc. So. Calif. Acad. Sci. 1:337-395. 
PHILuips, R. W. 

1896. On the development of the cystocarp in Rhodomelaceae. II. Ann. Bot. 10:185-204. 
SETCHELL, W. A. 

1914. Parasitic Florideae, I. Univ. Calif. Publ. Bot. 6:1-35. 
TuHureT, G. M. 

1851. Recherches sur les zoospores des algues et les antéridies des Cryptogames. Part 


II. Ann. Sci. Nat. Bot. 16(3¢ Ser. 2) : 53-93. 15 pl. 


1964} Dawson AND T6zUN: Chondria nidifica 293 


Fig. 2. Vegetative development of Chondria nidifica. 


A. Apex of a primary axis (the lower part shown in median longitudinal sectional view) showing: 
a, apical cell; b, first trichoblast initial; c, first pericentral cell; d, two of the developing ring of five peri- 
central cells; e, trichoblast branch; f, cortical cells developing from pericentral cells; g, axial cell row; 
h, basal cell of trichoblast which becomes the scar-cell; i, outer palisade cortex. X 466. B. A slightly oblique 
transection near the apex of a sterile axis, showing the axial cells, intercellular spaces, early development 
of the cortex from pericentral cells (still partly recognizable), and part of the basal cell of a trichoblast 
which has formed a secondary pit connection. X 110. C. Diagrammatic representation of the origin of an 


ordinary lateral branch from the distal, inner side of a scar-cell, D. Upper portion cf a sterile axis with 
branches. X 2.5. 


294 San Drieco Society oF NaturaAL History iAoies lB) 


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Fig. 3. Vegetative development and origin of branches in Chondria nidifica. 


A. Transection of a mature sterile axis, showing prominent axial cell, the parenchymatous medulla 
with smaller cells representing late-developed cells that have filled the earlier intercellular spaces, and the 
small-celled outer cortex which has lost its earlier palisade character. X 80. B. Surface aspect of the outer 
cortical layer of a mature axis. 466. C. Superficial view of a scar-cell at an early stage after the fall 
of the trichoblast, showing the rosette form of the surrounding cortical cells. X 466. D. Superficial view 
(somewhat lateral) showing a scar-cell with still prominent pit-connection evident at the broken surface. 
* 466. E. Scar-cell overgrown by cortical cell encroachment. 466. The two central cells may have been 
derived from the scar-cell below. 


1964] Dawson AND TO0zUN: Chondria nidifica 295 


f 


Sor Da 


Ah O 


Fig. 4. Procarp and cystocarp development in Chondria nidifica. 


A. Apex of a fertile female plant, showing the distinction between the early stages of a sterile (right) 
and a fertile (left) trichoblast. 466. B-E. Progressive development of a fertile trichoblast before maturity 
of the procarp. 466. F. Transectional view of adaxial side of the fertile branch, showing position of the 
procarp (p) in a shallow groove. G. Median optical view of a young cystocarp ready for fertilization, show- 
ing the two enlarged axial cells, the supporting cell bearing a sharply angled row of five sterile cells, and 
the four-celled carpogonial branch with exserted trichogyne. H. Cystocarp in early post-fertilization stage, 
showing rapid development of the pericarp and the beginning of fusion of the auxiliary cell, supporting 
cell, axial cell, and sterile cells. X 466. 


7 dig 


296 San Dreco Society oF Naturat History { VoL. 13 


Fig. 5. The cystocarp of Chondria nidifica. 


A. Upper part of a cystocarpic axis with branches and developing cystocarps. X 2.5. B. Fusion cell 
with developing gonimoblast branches as seen squeezed out of a young cystocarp. X 466. C. Mature 
cystocarp with ostiole, massive gonimoblast fusion cell, and young carpospores being protruded from its 
lobes. X 111. 


1964} Dawson AND T0zUN: Chondria nidifica 297 


Fig. 6. Spermatangial development in Chondria nidifica. 


A. Apex of a spermatangial axis, showing an early stage in the development of a male branch on 
the fertile trichoblast. B. A young spermatangial trichoblast, showing a well-developed sterile part and a 
fertile branch in the four-celled stage. C-D. Young spermatangial branches in the unistratose stage, one in 
face view, one in oblique view. E-F. Median optical lateral views of young spermatangial branches showing 
the first divisions of primary cells, producing the pluristratose condition, and the beginning of elongation of 
the primary cells. G. A spermatangial trichoblast bearing two juvenile sori, in oblique view. H. A young 
spermatangial sorus showing development from marginal initials and the elongation of primary cells to form 


the veins. All X 466. 


298 


San Dreco Socrety oF NATuRAL History { VoL. 13 


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Fig. 7. Spermatangial and tetrasporangial development in Chondria nidifica. 


A. Diagrammatic representation of a maturing spermatangial sorus showing border cells and veins 
and their relationship to the sterile branches of the trichoblast. B. Median optical view of the marginal 
part of a mature spermatangial branch showing border cells, vein cells, and ultimate spermatangial filaments. 
X 466. C. Upper part of a male axis showing branches and spermatangiophores. X 2.5. D. Median optical 
view of a tetrasporangial branch near the tip, showing central axial filament, a fertile pericentral cell bear- 
ing two cover cells, and a sterile pericentral cell bearing many cortical cells and with numerous secondary 
pit connections. Note that the fertile pericentral cell does not develop these secondary pit connections. 
Note the incomplete cortex over and around the cover cells. E. Detail of a young tetrasporangium develop- 


ing from a fertile pericentral cell, showing the prominent cover cells. 466. 


1964} Dawson AND TOzUN: Chondria nidifica 299 


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Fig. 8. Tetrasporangial development in Chondria nidifica. 


A. Upper part of a tetrasporangial axis, showing tufts of fertile branchlets. 2.5. B. Mature tetra- 
sporangium still attached to fertile pericentral cell which bears two cover cells. 466. C. Superficial view 
of a tetrasporangial branch at the place of discharge of a tetrasporangium, showing the two large cover 
cells filling much of the cavity and a shrunken fertile pericentral cell in the center below the remains of 
the sporangial membrane. X 466. 


w 


FEB 8 1965 


HARVARD 
UNIVERSITY 


TRANSACTIONS 
OF THE 


SAIN DIEGO SOGIETY OF NATURAL HISTORY 


VoLuME 13, No. 14, pp. 301-308 


NOTES ON VARIABILITY AND RANGE 
IN THE ELK KELP PELAGOPHYCUS 


BY 


Bruce C. PARKER 


Department of Botany and Plant Biochemistry 
University of California, Los Angeles 


AND 


E. YaLE Dawson 
San Diego Natural History Museum 


SAN DIEGO, CALIFORNIA 
PRINTED FOR THE SOCIETY 


AuGusT 28, 1964 


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FEB 8 1965 


NOTES ON VARIABILITY AND RAN Eee 
IN THE ELK KELP PELAGOPHYCUS  ~*=!TY 
BY 


BrucE C. PARKER AND E. YALE Dawson 


The elk kelp, Pelagophycus, is one of the giant seaweeds of Pacific North America be- 
longing to the family Lessoniaceae. The genus now contains two species, P. porra (Leman) 
Setchell and P. giganteus (Areschoug) Areschoug, the second only recently recognized as 
distinct and redescribed by Dawson (1962). Pelagophycus porra has been known for well 
over a century and has been reported from the vicinity of San Pedro, California, south to 
Punta San José, Baja California. P. giganteus has been known positively only from Santa 
Catalina Island, California, from where it was first described in 1876. Dawson pointed out the 
principal distinctions between the two species — P. porra has a smaller holdfast, a longer stipe, 
more numerous blades, and a larger, more spherical pneumatocyst. P. giganteus was known 
to him only from specimens with four blades, flattened antlers, elliptical pneumatocysts, and 
stipes less than 3 meters long. 


In May, 1963, the senior author found a portion of a plant of Pelagophycus which had 
washed ashore near Avalon, Santa Catalina Island. The specimen had six “petioles” (antler 
branches) indicating six blades instead of the maximum of four previously described for 
P. giganteus (Dawson, 1962). This discovery, in consideration of the very limited collections 
of this genus existing in herbaria, suggested that a more thorough search for specimens was 
needed to determine the variability and the systematic relationships within the genus. A pro- 
gram was initiated by the senior author using the Scuba (self-contained underwater breathing 
apparatus) diving technique, to collect specimens of these deep-growing kelps and to observe 


the characteristics of several natural populations of P. giganteus at the type locality, Santa 
Catalina Island. 


More than 200 plants of P. giganteus from Catalina Island were examined. Measurements 
were taken of a select number of plants from each collecting site. At least 30 specimens were 
preserved in glycerine:water:phenol (50:50:trace) solution for about one week, then removed 
from the preservative and stored in polyethylene bags for future reference. These specimens 
and accompanying data are deposited in the Herbarium of the University of California, 
Los Angeles. 


COLLECTIONS AND OBSERVATIONS AT SANTA CATALINA ISLAND 


The first collections were made at Avalon Bay on August 7 and September 6, 1963, 
where observations of more than 50 plants disclosed the following facts: (1) More than 50 
per cent of the population possesses six blades, and specimens with four blades are next in 
abundance; (2) Plants with more than six blades (i.e. seven, eight, or nine) occur occasionally 
(fig. 1); (3) Although the antlers of many specimens were more or less flattened as described 
previously for P. giganteus (Dawson, 1962), the variability is considerable and at least a few 
plants have essentially cylindrical antlers. 


The second collection site was Long Point, four nautical miles north of Avalon, and also 
on the east side of the island. This site is close to White Cove, where Dawson dredged the 
first collections of intact P. giganteus in 1948-49. Here the following observations were made 
on October 27, 1963: (1) About 50 per cent of the 30 plants observed had six blades (fig. 2) 
and most of the others had four; (2) A complete range in antler form from flattened to nearly 
cylindrical occurred; (3) Stipes averaged about 50 per cent longer than those of Avalon plants. 
In this connection it is noteworthy that a floating specimen collected some weeks later one 
mile north of Long Point had one of the longest stipes yet recorded (2.75 meters). This 
plant also possessed the largest pneumatocyst yet measured (11.4 * 13.2 cm.). 


304 San Deco Society oF Natura History {Vot. 13 


Fig. 1. Pelagophycus giganteus with nine blades, collected from Avalon, Santa Catalina 
Island, September 6, 1963. Approx. 1/15 life size. 


The third major collecting site was Emerald Bay, 13 nautical miles north of Avalon and 
four miles south of the northernmost tip of Santa Catalina Island. This area was chosen 
because Dawson’s characterization of P. giganteus had resulted mainly from collections made 
there. The divers reported that about 20 plants in a population of about 30 had six blades, 
while the remaining 10 plants possessed four blades. There were no juvenile plants at this 
location during the visit on December 8, 1963. Most of the plants observed had longer stipes 
than the Avalon specimens, and at least one plant had a nearly spherical pneumatocyst 


(S'16-X79°7 em): 

Among all of these specimens from Catalina Island there is marked variation with respect 
to the distance between the top of the pneumatocyst and the first dichotomy leading to the 
two antlers. Sometimes the distance is very small (ca. 1 cm.), as is the case with P. porra, 
but in other specimens it approaches 25 cm. 


COLLECTIONS AND OBSERVATIONS AT SAN CLEMENTE ISLAND 


Dawson (1962) reported evidence that P. giganteus might occur also at San Clemente 
Island. North had recalled seeing plants of this type while diving at the northeast end of 
that island on January 28, 1957. On the same date, Neushul had observed Pelagophycus at 
Northwest Harbor and on the west side of the island. A fragmentary specimen lacking stipe 
and pneumatocyst was collected from Wilson Cove (Herb. U.C.: Silva 4715, February 22, 
1949, pulled up on anchor from 12 meters). 


During the studies at Santa Catalina Island, two of the divers also visited Wilson Cove, 
San Clemente Island, and located there a large population of Pelagophycus from which they 
collected two specimens. The plants were growing at the base of a cliff beginning at about 25 
meters depth and continuing beyond 37 meters depth as far as visibility permitted observation. 
This appears to be the greatest depth yet recorded for P. giganteus. The plants lay along the 


1964 } PARKER AND Dawson: THE ELK KE Lp Pelagophycus 305 


Fig. 2. A six-bladed example of Pelagophycus giganteus in the natural habitat at ap- 
proximately 25 meters depth at Long Point, Santa Catalina Island, October 27, 1963. Approx. 
1/15 life size. 


bottom in a strong current. So that the divers could return quickly to the surface without 
exhausting their air supply, it was necessary to remove all but the bases of the large blades 
while collecting. Of the two specimens collected, one had six blades and the other had four. 
The divers reported that the proportion of 4- and 6-bladed specimens was about 1:1 as in the 
Catalina Island populations. The holdfasts, stipe lengths, and approximate dimensions of the 
blades removed by the divers resemble those of the Catalina specimens of P. giganteus. How- 
ever, in marked contrast to the Catalina Island plants, the two San Clemente Island speci- 
mens possess perfectly spherical pneumatocysts, 10.7 cm. in diameter, and perfectly cylindrical 
antlers. In these characters they correspond with P. porra. These incomplete data indicate that 
more collecting will be necessary at San Clemente Island before the range of morphological 
variation can be ascertained and its ecologic and phylogenetic implications assessed. 


COLLECTIONS AT Point Loma, SAN D1EGo, CALIFORNIA 


Pelagophycus porra collected at Point Loma for comparative purposes provided the fol- 
lowing major results: (1) Plants with 12 or more blades were found to be common; (2) One 
specimen had a holdfast measuring 33 cm. in diameter, well in excess of the maximum of 
20 cm. previously known for the species (Dawson, 1962), possibly due to its growth to some 
extent in sand and shell which covered the rock to which the holdfast was primarily attached; 
(3) Plants with 12 blades had pneumatocysts as small as 9 to 10 cm. in diameter and stipes 
as short as 3.51 meters. 


DISCUSSION 


The new data collected has permitted a re-evalution of the characteristics used to separate 
the two species of Pelagophycus. We now recognize the holdfast and stipe length as two of 


306 San Dieco Society oF NaTuRAL History {VoL. 13 


the more stable criteria for separating them. The number and size of blades also seem distinc- 
tive, for there is little overlap between the two species. The occurrence of 12 (to 20) blades 
now appears to be more characteristic of fully grown specimens of P. porra than eight blades 
as previously indicated by Dawson (1962). This agrees with an opinion expressed earlier by 
Setchell (1896) who examined Pelagophycus in the vicinity of San Pedro, California. The 
plants found at Santa Catalina and San Clemente islands have a maximum of six (rarely 
seven, eight, or nine) blades at maturity. These blades are always much longer (up to 10 or 
more meters) and wider (0.4 or more meters) than are those of P. porra. 


There is a considerably less consistent distinction between the species when one examines 
the pneumatocysts and antlers. The maximum size of P. giganteus at Catalina Island, 
11.4 X 13.4 cm., exceeds that earlier reported for this species, 9 X 12 cm. (Dawson, 1962). The 
range of pneumatocyst dimensions for P. giganteus also overlaps that for P. porra (minimum 
of 9 to 10 cm. in diameter for a 12-bladed specimen). The character of spherical versus ellipti- 
cal pneumatocysts in the respective species now appears less distinct. For example, a specimen 
of P. giganteus from Catalina had a pneumatocyst measuring 8.6 X 9.7 cm., while a specimen 
of P. porra from Point Loma measured 11.1 X 12.2 cm. Furthermore, the specimens from 
San Clemente Island, which resemble P. giganteus from Catalina in their holdfast, stipe, and 
blade characteristics, are more like P. porra in their possession of spherical pneumatocysts and 
cylindrical antlers. The antlers of Catalina Island specimens of P. giganteus vary considerably 
in the degree of flattening and sometimes are also nearly cylindrical like those of P. porra 
and of the San Clemente Island plants. Although little tendency for antler flattening was 
observed in P. porra from Point Loma, it is noteworthy that Setchell (1896), in describing 
plants from San Pedro, remarks that the antlers “are very decidedly flattened.” 


In view of the apparent stability of the holdfast, stipe, and blade characteristics, we may 
continue to recognize two distinct species of Pelagophycus: P. giganteus, from Santa Catalina 
and San Clemente islands, and P. porra, primarily located along the coast from Los Angeles 
County, California, south at least to Punta San José, Baja California. They may be distin- 
guished thusly: 


P. giganteus 


P. porra 


Holdfast: Haptera developing from progres- Haptera confined to a short length 
sively higher levels on base of stipe, of lowermost stipe, not extensive, 
becoming extensive, to 60 or more usually 10-20 cm. broad (rarely 
cm. broad, spreading into sand more), usually attached to a rock. 
and shell. 

Stipe: Short, reaching a maximum of Long, 7 to 27 meters. 

2.5-3 meters in adult plants. 

Blades: Usually 6 at maturity, rarely 7 Usually 12-20 in adult plants, to 
8 or 9; 6 to 10 or more meters 5.5 meters long and up to 0.4 
long and 0.4 or more meters wide. meters wide. 

Pneumatocyst: ae elliptical (from Catalina) Spherical to subspherical, 10-20 cm. 

6.4 X 7.6 to 11.4 X 13.4 cm. or in diameter. 
spherical (from Clemente) to 11 
cm. diameter. 
Antlers: Pronouncedly flattened to nearly Cylindrical (from Point Loma). 


cylindrical (from Catalina) or per- 
fectly cylindrical (from Clemente). 


The differences observed between specimens of P. giganteus from Santa Catalina and from 
San Clemente islands are viewed as representing subspecific variation in well-separated popu- 


1964} PARKER AND Dawson: THE ELK KELp Pelagophycus 307 


lations. Further collections from San Clemente Island may make possible more definite con- 
clusions regarding the systematic relationships of these two island populations. 

We would like to stress that subsequent collections from other locations, such as the 
several submarine banks in the area, may provide evidence for additional changes in the 
systematics of this genus. No collections have been made from the other Channel Islands, al- 
though it is probable that certain of these have populations of Pelagophycus. Santa Cruz Island 
is believed to support plants of this genus, allegedly of the P. porra type (Dawson, 1962), 
but we have made no observations in that area. Professor Orda Plunkett, of the Department 
of Botany and Plant Biochemistry, University of California, Los Angeles, recalls seeing plants 
possibly of this genus while fishing at Santa Rosa Island some years ago; the plants were 
visible from the boat and were associated with a bed of Nereocystis luetkeana. 

A single specimen of Pelagophycus was collected by Taylor Gabbard while free-diving 
on the north side of Anacapa Island, May 26, 1963. The genus has not been reported pre- 
viously from that island, and the specimen may have been detached. The holdfast was 20 cm. 
broad, but may have been larger, for it possessed a few severed main hapteral branches. The 
stipe measured 4.1 meters in length and the pneumatocyst 15.3 X 17.3 cm. (elliptical). Antlers 
were essentially cylindrical, and there were eight blades. Although the antlers, blades and, to 
a lesser extent, the holdfast, suggest P. porra, the short stipe accompanied by the large, elliptical 
pneumatocyst is not in accord with the usual developmental sequence of P. porra at Point 
Loma, where the pneumatocyst is spherical and normally only half as large when the stipe has 
reached four meters. It seems, therefore, that this single specimen from Anacapa Island points 
to a need for continued explorations for Pelagophycus there and for further tests of the 
stability of the characters now used to distinguish the species. 


ACKNOWLEDGEMENTS 


We are grateful to the Scuba divers, John Bleck, Larry Cushman, Edmund Hobson, 
David Joss, and Harold Warner, for volunteering their services for this study. The investiga- 
tions were supported by funds from University Grant No. 1827, University of California, 
Los Angeles. 


LITERATURE CITED 
Dawson, E. YALE 


1962. On the recognition of a second species of the genus Pelagophycus. Bull. So. 
Calif. Acad. Sci. 61:153-160. 


SETCHELL, WILLIAM A. 
1896. The elk-kelp. Erythea 4:179-184. 


RS 


—_ 
@ 


bae 


FFB 8 1965 
TRANSACTIONS HARVARD 


OF THE UNIVERSITY 


SaNE DIEGO SOCIETAGOESNATURAL HISTORY 
VotuME 13, No. 15, pp. 309-312 Aucust 28, 1964 


THE LARVA OF HADROTES CRASSUS (MANNERHEIM) 
(COLEOPTERA: STAPHYLINIDAE). 


BY 


IAN Moore 


Research Associate in Entomology 
San Diego Natural History Museum 


Hadrotes crassus (Mannerheim), a member of the tribe Xanthopygini of the subfamily 
Staphilininae, is the only known species of the genus. It occurs in decaying seaweed from 
Alaska to Baja California. It is usually not common, but occasionally is encountered in 
numbers. Its larva has not hitherto been described. 


The tribe Xanthopygini is moderately large, embracing thirty-three known genera, most 
of which contain only one or a few species. However, some genera include as many as ten to 
thirty species. Most of the species inhabit the tropics, with very few in temperate regions. 


Larvae of only two species of the tribe have been described. Both of these are known 
from the Nearctic region. They are Creophilus maxillosus (Linné) and Thinopinus pictus 
LeConte. The former is of holarctic distribution and the latter is confined to the shores of the 
Pacific coast of North America. The larva of Creophilus maxillosus has been described by 
several European students, including Paulian (1941:251); that of Thinopinus pictus was 
illustrated by Boving and Craighead (1930:118, 123). 


A key to the known larvae of the Xanthopygini follows: 


1. Last segment of antenna very short, not longer than the 


larcemnodihed seta at apexiOl thitdsegiie nit: <2. a cea ee Thinopinus 

Last segment of antenna several times as long as 

modified seta at.apexof third segmetit..0 5.500 oR eee eee eee 2 
2. Ligula about one-half as long as first segment 

of Jabiall palpi snot densely pubbescemts: 211.0522 hee ae ee ee eee Creophilus 

Ligula longer than first segment of labial palpi, 

densely, ‘pubescent :at-apex ste: te sec 202s ietace teeta ake ae see Hadrotes 


Larva OF Hadrotes crassus (MANNERHEIM) 


Color.— Head and thorax reddish-brown above; abdomen dark testaceous above; under 
surface, palpi and legs pale testaceous. 


Head. — Wider than long, widest near base of mandibles; sides nearly straight to the 
middle, thence curved into the broadly rounded posterior angles; base straight. Ocelli four, 
large, located in a compact group posterior to the base of the mandibles. Frontal suture form- 
ing an obtuse angle with the epicranial suture, which is about one-half the length of the head. 
Clypeolabral margin with five large, evenly spaced blunt teeth of equal size, and an additional 


310 San Dreco Society oF Natura History {Vor 13 


=o 


5 


%, 
fool 


& 
y CA 


GE 


ky 
| 


Fig. 1. Larva of Hadrotes crassus. a, prosternum; b, clypeolabral margin; c, labial palpi 
and ligula; d, apex of urogomphus; e, antenna; f, dorsal view of larva; g, anterior leg, front 
view; h, middle leg, front view; i, maxilla; j, claw of posterior leg. 


LIBRARY 


1964 } Moore: Larva OF Hadrotes crassus FFR 8 36 5 
Ee & 


smaller tooth at each outer angle. Antennae four-segmented, first segment shorter than” wide 
second narrower at base than the first, widened gradually to apex, which is about¥AY Wid&ad Y 
the first segment, about twice as long as wide; third three-fourths as wide as the second, narrow 
at base, as wide at apex as the third segment; fourth little longer than width of the third, 
about twice as long as wide; modified seta at apex of the third segment, hardly one-fifth as 
long as the fourth segment, about as long as wide. Maxilla with the mala cylindrical, nearly 
twice as long as wide; palpi three-segmented, first two segments about equal in length, about 
twice as wide as long; third shorter than the second, little more than half as wide as apex of 
the second, cylindrical in basal two-thirds, thence narrowed to the pointed apex. Ligula about 
four times as long as wide, shorter than the labial palpi, constricted before the apex, which 
is densely pubescent. Labial palpi two-segmented, first segment four times as long as wide; 
second of equal width but only one-half as long, gradually narrowed to the rounded apex. 
Gular sutures united. 


Pronotum. — Subrectangular, transverse, about as long as the mesonotum and metanotum 
together. Legs stout; lower side of femora with a double row of stout setae; tibiae with irregu- 
larly placed stout setae throughout. 


Abdomen. — A little wider than the head, longer than the head and thorax together. 
Pseudopode twice as long as first two segments of the urogomphus. Urogomphus three-seg- 
mented, first segment widest, twice as long as wide; second a little narrower, longer, about 
four times as long as wide; third very slender, about one-half as wide and as long as the second. 


Integument. — Finely, densely reticulate, the legs obscurely so. 
Length. — 16 mm. 


One specimen was taken from decaying seaweed at Sunset Cliffs, San Diego, California, 
October 5, 1953, in company with eight adults. 


LITERATURE CITED 


Bovine, A. G., and F. C. CrAIGHEAD 


1930-1931. An illustrated synopsis of the principal larval forms of the order Coleoptera. 
Ent. Americana 11:1-351. 


MANNERHEIM, Car G. 


1846. Nachtrag zur Kaefer-Fauna der Aleutischen Inseln und der Inseln Sitkha. Bull. 
Soc. Imp. Moscou 19:501-516. 


PAULIAN, RENAUD 


1941. Les premiers états des Staphylinoidea. Etude de Morphologie comparée. Mem. 
Hist. Nat. Paris, nov. ser., 15:1-361, 3 pls. 


cFR 8 1960 


HARVARD 
UNIVERSITY 


TRANSACTIONS 
OF THE 


SAN DIEGO SOGIETY OF NATURAL HISTORY 


VoLuME 13, No. 16, pp. 313-332 


TERRESTRIAL MOLLUSKS 
OF THE BELVEDERE EXPEDITION 
TO THE GULF OF CALIFORNIA 


BY 


WILLIAM K. EMERSON 
AND 


Morris K. JACOBSON 


American Museum of Natural History 


SAN DIEGO, CALIFORNIA 
PRINTED FOR THE SOCIETY 


AuGustT 28, 1964 


Pp ey 


TERRESTRIAL MOLLUSKS OF THE BELYE ERE? 
EXPEDITION TO THE GULF OF CALIFOR RE 


BY 


Wi_tiaM K. EMERSON AND Morris K. JAcoBson 


INTRODUCTION 


This paper records the terrestrial mollusks that were collected by the senior author and 
other members of the scientific party from the islands in the Gulf of California and from 
the east coast of Baja California during the course of the expedition. This collection was made 
incidental to other collecting interests and every member of the scientific party assisted in the 
field work. Therefore, we are pleased to acknowledge the assistance of Richard C. Banks, 
Dennis Bostic, Ambrosio Gonzalez C., Charles F. Harbison, Don Hunsacker II, George E. 
Lindsay, Reid Moran, Chris Parrish, Charles E. Shaw, Michael Soulé, and Ira L. Wiggins. 
We are also indebted to Antero Diaz and the crew of the M/V “San Agustin II” for their 
interest and co-operation. 

The expedition, sponsored by the San Diego Natural History Museum and the Belvedere 
Scientific Fund of San Francisco, started from Bahia de los Angeles, Baja California, on 
March 15, 1962, and terminated at La Paz, Baja California, on April 21, 1962. During the 
cruise, 32 islands were visited, seven localities on the peninsula of Baja California were investi- 
gated, and one brief stop was made on the Sonoran coast. A general account and log of the 
expedition, including maps of the route, was published by Lindsay (1962). 

Our knowledge of the terrestrial molluscan fauna of the islands in the Gulf of California 
is based largely on the study by G Dallas Hanna (1923), who reported on collections made 
by the California Academy of Sciences on expeditions to the Gulf in 1919 and 1921. Hanna 
described 12 new species, nine of which were insular populations, and redescribed many of 
the previously known species. More recently, Pilsbry and Lowe (1932) described two insular 
species and Jacobson (1958) recorded the land mollusks collected by the “Puritan”-American 
Museum expedition to western Mexico, including two insular species from the Gulf of Cali- 
fornia. Dr. Hanna and Allyn G. Smith currently are working on a large collection that has 
accumulated in recent years as a result of numerous collecting trips to the Gulf of California 
by the California Academy of Sciences. When their study is completed, the insular faunas 
will be much better known. At the present time, the terrestrial mollusks of most of the islands 
are too poorly known for us to undertake a regional analysis of the origin and evolution of 
these insular faunas. 

The present collection is composed largely of specimens of the family Bulimulidae. All 
of the known insular species of this family in the Gulf of California are represented in the 
collection and are referred to eight nominal species of the genus Rabdotus (sensu lato). The 
smaller and less conspicuous members of the insular molluscan faunas are poorly represented 
in the collection. The remainder of the collection consists of one new subspecies of the family 
Helminthoglyptidae and of two previously known species for each of the families Urocoptidae 
and Pupillidae. The collecting localities are indicated on the maps that accompany the general 
account of the expedition (Lindsay, 1962: figs. 1, 2, 7, 13, 20). The present collection was 
divided between the San Diego Natural History Museum and the American Museum of 
Natural History. 

We wish to thank G Dallas Hanna and Allyn G. Smith of the California Academy of 
Sciences, Emery P. Chase of the San Diego Natural History Museum, R. Tucker Abbott 
and H. Burrington Baker of the Academy of Natural Sciences of Philadelphia, and Harald A. 
Rehder and Joseph P. E. Morrison of the United States National Museum for their kindness 
in loaning pertinent specimens from the collections of their respective institutions. Meredith L. 
Jones of the American Museum of Natural History assisted us in certain of our statistical 
studies and William E. Old, Jr., of the American Museum of Natural History provided 


technical aid of various kinds. 


316 San Dieco Society oF Natura History { VoL. 13 


The senior author is very much indebted to George E. Lindsay for inviting him to join 
the expedition and to his companions on the trip for their assistance in the field collecting. 
We are most grateful to the Belvedere Scientific Fund for making the expedition possible. 


SYSTEMATIC TREATMENT 
Family BULIMULIDAE 


Genus Bulimulus Leach, 1814 


Bulimulus Leach, 1814, p. 41. 
Type species. — Helix exilis Gmelin, 1791 {= Bulimulus guadalupensis Bruguiére, 1792] 
by subsequent designation of Herrmannsen (1846:136). 


Remarks. — Pilsbry (1897b:127), in his monograph of the Bulimulidae, recognized three 
major divisions within the nominate genus. These were: (I) the subgenus Bostryx Troschel, 
having shells with “Apical whorls smooth, not wrinkled, costate or granular”, represented by 
species from Ecuador, Peru, Chile, Bolivia, and Argentina; (II) the subgenera Plectostylus 
Beck, Scutalus Albers, Bulimulus s.s., and Rhinus Albers, having shells with “Apical whorls 
irregularly wrinkled subvertically, or with the wrinkles wavy, dislocated or variously broken 
into granules”, represented by species from tropical America, West Indies, and the Galapagos 
Islands; (III) the subgenera Protoglyptus Pilsbry, Naesiotus Albers, and Orthotomium Crosse 
and Fischer {= Rabdotus Albers}, having shells with “Apical whorls with regular, straight 
vertical riblets”, represented by species from northern Mexico, Baja California and the southern 
United States. The species of bulimuli treated in the present paper are referable to Pilsbry’s 
Division III. 

Although Pilsbry’s ultraconservative arrangement was generally accepted for many years, 
recently it has undergone revision. Weyrauch (1956:1-4) noted that the nepionic sculpture 
of the subgenera of Pilsbry’s Division III differed decidedly from that of the type species of 
Bulimulus, B. exilis (Gmelin) {= guadalupensis (Bruguiére) }. As a result of this and other 
observations, he gave full generic standing to Naesiotus Albers, 1850, and he indicated that 
Protoglyptus Pilsbry, 1897b, also was worthy of that rank. Parodiz (1962:445) subsequently 
elevated Protoglyptus to generic status. 

Weyrauch (1956:3) also expressed doubt that Rabdotus Albers, 1850 (=Orthotomium 
Crosse and Fischer, 1875), the third of the subgenera that Pilsbry recognized, should be 
considered a subgenus of Naesiotus. Among other things he noted a difference in the nepionic 
sculpture, stating: “The nepionic shell of Rabdotus differs mainly from that of Naesiotus in 
the initial half whorl, its surface appearing shriveled by a very low, ill defined sculpture, often 
without traces of riblets, or if present, they are often broken or diversely curved. The remain- 
ing part of the embryonic shell is sculptured with narrowly spaced, straight, vertical or slightly 
slanting riblets, which are in most specimens continuous from suture to suture, but in other 
specimens in places interrupted at the middle. The embryonic sculpture of Naestotus is rela- 
tively stronger, in most species more widely spaced, at least on the first whorl, than in any 
species of Rabdotus. The vertical or slightly slanting riblets of Naesiotus are thin, high, al- 
ways continuous and on the initial half whorl equally strong as on the remaining part of the 
embryonic shell.” 

In addition to Weyrauch’s observations, we should like to call attention to the following 
significant points. 1. The distribution of Rabdotus is confined to central and northern Mexico, 
Baja California, and central and southern United States. In this area no species of Protoglyptus 
or Naesiotus are known to occur. 2. Rabdotus may or may not have an internal axial denticle 
or other axial structures (subgenus Leptobyrsus). 3. Rabdotus is generally unicolored or is 
less commonly streaked with axial stripes. The colors are generally subdued, never vivid. In 
addition, as was noted by Pilsbry (1946:4), spiral markings are wanting. 4. Rabdotus, in the 
wide sense used in this report, frequently has a strong, well-developed peristome, which is 
commonly strongly reflected, unlike the simple lip of the type species of Bulimulus. 5. Rabdotus 
has that indefinable characteristic that Pilsbry has described as “the general appearance.” After 


1964} EMERSON AND JACOBSON: TERRESTRIAL MoLiusks 317 


working for a while with Rabdotus, one finds that the group as a whole becomes readily dis- 
tinguishable. 

Therefore, we feel justified, in the absence of anatomical data to the contrary, to raise 
Rabdotus to full generic standing and to recognize the following subgenera: 

Rabdotus (sensu stricto) Albers, 1850, type species Bulimulus dealbatus (Say); 

Leptobyrsus Crosse and Fischer, 1875, type species Bulimulus spirifer (Gabb) ; 

Hannarabdotus new subgenus, type species Bulimulus slevini Hanna; 

Puritanina Jacobson, 1958, type species Bulimulus montezuma Dall. 

It is also of interest to note in passing that Pilsbry himself, who in 1897 was pessimistic 
about the possibility that his subgenera would eventually be raised indiscriminately to full 
genera, felt differently in later years. In connection with Bostryx, Pilsbry (1944:120) stated: 
“When I revised the classification of the Bulimulidae nearly fifty years ago the genus Buli- 
mulus was left with rather wide limits . .. As genera are somewhat more narrowly limited now, 
it seems desirable to allow some groups of Bulimuli the generic status.” 


Genus Rabdotus Albers, 1850 


Rabdotus Albers, 1850, p. 164. 

Ortkotomium Crosse and Fischer in Fischer and Crosse, 1875, p. 473; type species Bulimulus sufflatus 
Gould, 1859, by original designation. 

Globulinus Crosse and Fischer in Fischer and Crosse, 1875, p. 475; type species Bulimulus sufflatus 
Gould, 1859, by original designation. 

Type species. — Bulimulus dealbatus Say, 1821, by subsequent designation of Kobelt 
(1880:267), for Rhabdotus {sic}. 

Remarks. — Zilch (1960:490) incorrectly cited Bulimulus (R.) sufflatus Gould, 1859, 
as the type species of Rabdotus. This species, however, was not included by Albers (1850: 164- 
165) in the original list of species referred to Rabdotus. The earliest valid type designation 
appears to be Bulimulus dealbatus Say, 1821, made by Kobelt (1880:267). 


KEY TO THE SUBGENERA OF RABDOTUS 
1. Surface not pebbled 


1.1. No internal axial ornamentation 


1.1.1. Lip simple, sometimes internally thickened............2.2.0.0..020202.2-.-... Rabdotus (s.s.) 

ihe Liptexpanded*and/onretiected# aos Ui a SS Hannarabdotus 

1.2. With internal axial ornamentation commonly present..............-.-.--.-.--.-------- Leptobyrsus 

Pen surtace. closely pebbled i(ecanulate i. 2 ee ee eee Puritanina 


Subgenus Rabdotus 
Rabdotus (Rabdotus) sufflatus (Gould), 1859 


Bulimulus vesicalis Gould, 1853 [October], p. 375, pl. 14, fig. 1, “Lower California.” Not Pfeiffer, 
1853. [March]. 

Bulimulus sufflatus Gould, in Binney, 1859, p. 25. 

Bulimulus (Orthotomium) sufflatus Gould, Dall, 1893, p. 646, pl. 72, fig. 9. 

Bulimulus sufflatus var. insularis Cooper, 1893a, pp. 212; Cooper, 1893b, 340, pl. 14, fig. 6, Isla 
Espiritu Santo; Cooper, 1894, p. 140. 

Bulimulus sufflatus var. chinchensis Cooper, 1894, p. 140, pl. 5, fig. 10, “El Chinché Mountains”, 
Baja California. 

Bulimulus (Orthotomium) sufflatus (Gould), Pilsbry, 1897b, pp. 136, 137, pl. 18, figs. 38-44, pl. 25, 
fig. 63, San José del Cabo to La Paz, Baja California. 

Bulimulus (Orthotomium) sufflatus var. insularis Cooper, Pilsbry, 1897b, p. 137. 

Bulimulus (Orthotomium) sufflatus var. chinchensis Cooper, Pilsbry, 1897b, p. 137, pl. 25, fig. 63. 

Bulimulus juarezi Pfeiffer, 1865, p. 832; Pilsbry, 1897b, pp. 136, 137, pl. 18, figs. 40, 41. 

Bulimulus sufflatus Gould, Lowe, 1933, p. 75, “aestivating, buried under dead leaves of the yucca 
and other desert vegetation”, sand hills at Ensenada de los Muertos, Baja California. 

Records. —Isla San José, east side, 1 dead, fresh specimen (Banks); Isla San José, west 
side, 3 mature, 2 immature, dead specimens (Harbison and Wiggins). 

Remarks. — Gould (1853:375) stated that this species “Inhabits Lower California”, and 
that the specimens were collected by Major William Rich along the lower east coast of Baja 


California, possibly at La Paz or San Juan. Subsequent writers have cited it from numerous 


318 San Dreco Socrety oF NaTurAL History [Vors13 


localities in the Cape region of the peninsula. La Paz is here designated the type locality. 
An extensive synonymy of the earlier literature is given by Pilsbry (1897b:136). 

Cooper (1893a:212; 1893b:340, pl. 14, fig. 6; 1894:140) described specimens from 
Isla Espiritu Santo, off La Paz, as a “variety” of R. sufflatus. The insular race was considered 
by Cooper to be more slender than the nominate subspecies. After examining the original 
description and discussions of the subspecies, however, we find ourselves in agreement with 
Hanna (1923:485) who stated that “the differentiating characters . . . are not well described.” 
Cooper gave a confusing account of the supposed size differences. In one place (Cooper, 
1893a:213), the length is given as 1.20 inches, as compared to 1.35 inches for the peninsular 
form, whereas later Cooper (1894:140) nliaw to the insular specimens as a “much larger 
form.” The figure of msularis given by Cooper (1893b: pl. 14, fig. 6) is 1.25 inches, which 
is well within the range of variation for sufflatus as given ip Pilsbey (1897b: 136). No speci- 
mens of the R. sufflatus group were reported by Hanna (1923) on Isla Espiritu Santo from 
the collections made in 1921, nor did the present expedition obtain any specimens there. 

The type specimens of Cooper’s insularis were apparently lost in the disastrous fire of 
1906 that destroyed the collections of the California Academy of Sciences. Until additional 
specimens are taken on Isla Espiritu Santo, this taxon must be considered a nomen inquirendum. 

Hanna (1923:485) reported a single juvenile specimen of the sufflatus group from Isla 
Cerralvo, off La Paz, but concluded that it could not be differentiated from typical sufflatus. 

The present specimens are the first record of this species from Isla San José. The four 
larger specimens range from 24.0 to 25.0 mm. in altitude, with an average altitude of 24.1 
mm., and they range from 15.4 to 16.4 mm. in diameter, with an average diameter of 15.7 mm. 
Although these specimens are smaller than most specimens with the same number of whorls 
from peninsular populations of this species, we hesitate to describe an insular race on the basis 
of our small sample. Additional material from Isla San José, however, may demonstrate con- 
clusively that this insular population merits subspecific recognition. 


Subgenus Leptobyrsus Crosse and Fischer, 1875 

Leptobyrsus Crosse and Fischer in Fischer and Crosse, 1875, p. 475. 

Sonorina Pilsbry, 1896, p. 114; type species Bulimulus spirifer (Gabb), 1867, by original designation. 

Type species. — Bulimulus spirifer (Gabb), 1867, by original designation. 

Remarks. — Pilsbry (1896:114) proposed the name Sonorina in the belief that Leptobyrsus 
Crosse and Fischer, 1875, was preoccupied by Leptobyrsa Stal, 1873. However, under the 
present Zoological Code the two names are not homonyms. 

This group is characterized by the common presence of a lamina on the columella within 
the body whorl. The denticle-like lamina generally is only partly visible in the aperture. In rare 
instances, specimens occur in which the denticle is not present, but, as we indicate below, this 
may be due to calcium absorption. 

The junior author recently had the opportunity of discussing the nature of the internal 
denticle with Dr. H. Burrington Baker of the Academy of Natural Sciences of Philadelphia. 
He was kind enough to point out that there may be a very simple explanation for the varying 
size of the denticle and for the apparent lack of correlation between size of the denticle and 
maturity of the shell (see p. 323). These phenomena become more understandable if the 
denticle is considered to be a storehouse of lime that the snail secretes copiously when lime 
conditions are favorable and reabsorbs when lime is difficult to obtain. Reabsorption of shell 
material is not unknown in many species of terrestrial mollusks. If this interpretation is correct, 
there is even less reason for considering the size and shape of the denticle to be a significant 
specific character (see discussion beyond) as utilized by Hanna (1923) and Pilsbry and Lowe 
(1932). However, much additional information, including ecological and physiological studies, 
will be needed to determine why this lime preserving characteristic is found only in certain 
species of Rabdotus. 


Rabdotus (Leptobyrsus) chamberlini (Hanna), 1923 
Figure 1 
Bulimulus chamberlini Hanna, 1923, p. 494, pl. 8, fig. 21, Isla San Diego. 


1964} EMERSON AND JACOBSON: TERRESTRIAL MOLLUSKS 319 


a 


Fig. 1. Rabdotus (Leptobyrsus) chamberlini (Hanna). a-b, apertural view, showing 
epiphragm in first specimen; c-f, dorsal view, showing variation in the development of the 
internal denticles. * 1.25. 


Records. — Isla San Diego, 132 dead specimens, 50 feet to crest of island, locally common 
under blocks of granite which were overgrown by vegetation (Emerson, Shaw, and Soulé). 

Remarks. — This distinctive species was based on a single, incomplete specimen found on 
tiny, granitic Isla San Diego. The present specimens agree very well with the original descrip- 
tion and corroborate the conclusion of Hanna (1923:494), based on the unique type, that 
this species is closely related to R. (L.) dentifer, a larger form occurring on Isla Tortuga. 

The present species is smaller than any of its congeners and there seems to be very little 
variation in size. Of ten specimens chosen at random for measuring, the altitude ranges from 
17.0 to 19.7 mm., with an average of 18.6 mm., and the width ranges from 7.5 to 9.5 mm., 
with an average of 8.7 mm. (table 1). 

The textnre of the shell is thin and fragile, and the peristome is weakly expanded and 
unreflected (fig. 1, a-b). 

The axial denticle is commonly rounded at the summit, but in some specimens it tends 
to show the squarish outline of R. (L.) lamellifer (cf. figs. 1f and 2h). The size of the 
denticle varies considerably, there being no apparent correlation between the maturity of the 
shell and the size and thickness of the denticle. 

An immature, well-preserved, specimen of four whorls shows the following characters of 
the nuclear whorls—a series of approximately nine irregularly spaced, weakly developed, 
whitish spiral lines on a surface weakly ornamented by shallow, irregular, axial wrinkles. The 
wrinkles become more regular and stronger on the first postnuclear whorl, where they are 
most pronounced just below the suture. 


Rabdotus (Leptobyrsus) dentifer (Mabille), 1895 

Bulimulus (Leptobyrsus) dentifer Mabille, 1895, p. 67, “Ile volcanique de la Tortuga.” 

Bulimulus dentifer Mabille, Pilsbry, 1898, pp. 161, 162. Hanna, 1923, pp. 492-494, pl. 8, figs. 16-20, 
Isla Tortuga. Lowe, 1933, p. 110, Isla Tortuga, “under blocks of black lava not far from the crater’s rim.” 

Records. —Isla Tortuga, above 200 feet elevation, 10 living and 71 dead specimens 
(Banks, Moran, and Soulé). 

Remarks. — Dead specimens of this small species were seen in abundance near and in 
the crater of this volcanic island, and the living specimens were found adhering to the under- 
sides of rocks in talus deposits. Some of the present specimens are larger than the maximum 
size reported by Hanna (1923:493). One specimen measures 26.4 mm. in height, whereas 
the largest specimen reported by Hanna is 23.7 mm. Next to R. (L.) chamberlini (Hanna), 
this is the smallest species in the subgenus. 

Mabille (1895:67) gave a largely undiagnostic description of this species, but cited the 
“Volcanic Island of La Tortuga” as the habitat. As Hanna (1923:494) has stated, the geo- 


{ VoL. 13 


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1964 | EMERSON AND JACOBSON: TERRESTRIAL MOLLUSKS 


Fig. 2. Kabdotus (Leptobyrsus) lamellifer lamellifer (Pilsbry). a-c, paratypes of Buli- 
mulus sanmarcosensis Pilsbry and Lowe from Isla San Marcos; d-f, topotypic specimens; g-k, 
topotypic specimens showing variation in the development of the denticles. * 1. 


graphic evidence seems conclusive, despite the inadequate original description. Also, the measure- 
ments given in the original description, height 24 mm. and diameter 9 mm., fall within the 
range of variation of specimens known from this locality. Table 1 gives a summary of the 
measurements of the altitude and diameter of seven specimens from the present collection. 

As Hanna (1923:492) pointed out, contrary to the original description, microscopical 
spiral striae are present only on the uppermost two or three whorls. Hanna (1923:493) also 
described and figured the genitalia and jaw of this species. 


Rabdotus (Leptobyrsus) lamellifer lamellifer (Pilsbry) , 1897 
Figure 2 

Bulimulus lamellifer Pilsbry, 1897a, p. 103, “Lower California”? (W. M. Gabb). Pilsbry, 1898, 
p. 160, pl. 21, figs. 94-99. Hanna, 1923, pp. 495-497, Coyote Bay, Agua Verde Bay, and west of Puerto 
Escondido, Baja California and Isla Danzante. Lowe, 1933, p. 110, Isla Danzante, living specimens. 

Bulimulus sanmarcosensis Pilsbry and Lowe, 1932, pp. 49, 50 {not pl. 5, fig. 1}, Isla San Marcos. 

Bulimulus (Leptobyrsus) sanmarcosensis Pilsbry and Lowe, Pilsbry, 1935, p. 2, pl. 1, fig. 8 {holotype}. 

Records. — Mulegé estuary, Baja California, south entrance at an elevation of 165 to 330 
feet, 5 dead specimens (Moran). 

Puerto Santispaquis, Bahia de Concepcidn, Baja California, elevation 985 to 1300 feet, 
18 dead, mature specimens (Moran). 

Tinajas Carriza, northwest of Puerto Escondido, Baja California, 5 specimens (Banks). 

About 2 miles northwest of Puerto Escondido, Baja California, 2 dead specimens (Banks). 

Puerto Escondido, Baja California, elevation 50 to 100 feet, 18 mature, 11 immature, 
dead specimens (Moran, Soulé). 


Isla Danzante, elevation 32 to 165 feet, 26 dead specimens (Moran). 


322 San Dreco Society oF NaturAL History { VoL. 13 


TABLE 2 


Measurements (in mm.) of the internal denticles in samples of Rabdotus (L.) lamellifer. 


Number of HEIGHT WIDTH 
Species specimens Range Average Median Range Average Median 

R. (L.) lamellifer lamellifer (Pilsbry) 

Santispaquis, Baja Calif. (type locality) ll 2.9-4.5 3.4 33 4.5-7.7 6.5 6.3 

Isla Danzante 231 1.8-3.2 2.6 DD 4.9-6.4 5.8 De) 
R. (L.) lamellifer ximenez (Hanna) 

Marquer Bay, Isla Carmen (type locality) 22? 2.0-5.3 35 3/2 4.4-8.5 63 6.4 

Puerto Ballandra, Isla Carmen 10 3} D5).3) 4.3 4.3 6.0-8.0 72 UL») 


1Only 7 possess measurable denticles. 
“Only 17 possess measurable denticles. 


Isla San Marcos, northeast side, elevation 150 to 700 feet, 3 dead specimens and 1 living 
juvenile specimen (Banks, Moran, Soulé); Isla San Marcos, southwest side, elevation about 
300 feet, 1 dead specimen (Moran). 

Remarks. — Bulimulus lamellifer was based by Pilsbry (1897a) on specimens collected by 
William M. Gabb from an unknown locality in Baja California. Later, Pilsbry (1898) sug- 
gested that Gabb had collected the type specimens “probably on the Gulf side between San 
Antonio {in the Cape Region} and San Borja [west of Bahia de los Angeles}.” As Hanna 
(1923) pointed out, subsequent collecting has failed to find this species in these widely sepa- 
rated areas of the peninsula, and it seems likely that Gabb collected his specimens somewhere 
between these localities. We here designate Puerto Santispaquis, Bahia de Concepcion, as the 
type locality, since this is near the center of the known geographic range of this species, Santa 
Rosalia (Pilsbry, 1898) to Isla Danzante (Hanna, 1923). A summary of the measurements 
of the altitude and diameter of eleven specimens from the type locality is given in table 1. 

Bulimulid populations occurring on some of the islands appear to be closely related, if not 
identical, to those of this species occurring on the peninsula. The population from Isla San 
Marcos was named Bulimulus sanmarcosensis by Pilsbry and Lowe (1932), whereas the 
one on Isla Carmen has received two names, Bulimulus ximenez Hanna (1923) and B. carmen 
Pilsbry and Lowe (1932). The population occurring on Isla Danzante was considered by 
Hanna (1923) to be referable to the peninsular species, R. lamellifer. In order to determine 
the biological significance of these insular populations, we undertook a statistical analysis of 
samples available to us. These data are summarized in table 1. 

The measurements of altitude and diameter of the shells show that the ranges of variation 
for the population from the peninsula (Rabdotus lamellifer lamellifer) and the populations 
from Isla San Marcos and Isla Danzante are essentially the same, whereas the population on 
Carmen Island is significantly larger in these dimensions. Moreover, the difference in size 
between the shells from Isla Carmen and the shells of other populations of this complex is 
strikingly apparent, making this a convenient, easily observable distinction. Therefore, we pro- 
pose to recognize the population from Carmen Island as an insular subspecies, R. (L.) lamel- 
lifer ximenez. 

Pilsbry and Lowe (1932) believed that the much smaller internal denticle of R. san- 
marcosensis served to distinguish the shells occurring on Isla San Marcos from R. lamellifer 
and R. ximenez. However, measurements of the denticles of samples representing these taxa 
show great variation in the size of this structure (table 2). Hence, we must conclude that this 
character is too variable to serve as a criteron for separating these populations. It is of interest 
to note that in a series of ten specimens from Isla San Marcos, the denticle ranged from 5.0 
mm. in height and 3.1 mm. in width to the slightest indication of a fold in most of the 
specimens. 


1964 } EMERSON AND JACOBSON: TERRESTRIAL MoLLusKs 323 


Fig. 3. Topotypic specimens of Rabdotus (Leptobyrsus) lamellifer ximenez (Hanna) 
showing variation in the development of the denticles. a-b, specimens from the present collec- 
tion; c-e, specimens from the “Puritan” collection (Jacobson, 1958:6). * 1. 


In the population occurring on Isla Danzante, the denticles appear to be poorly developed 
(table 2). The size of the specimens (table 1), however, suggests affinity with the nominate 
subspecies rather than with insular populations occurring on Islas Carmen and Coronados. 

Future studies of the insular populations of the R. lamellifer complex occurring on Islas 
San Marcos, Danzante and Coronados may show that some of these populations represent 
valid subspecies. However, we are taking a conservative point of view on the basis of the 
material previously studied and presently at hand, believing that these populations represent a 
single plastic species with only two recognizable subspecies. 


Rabdotus (Leptobyrsus) lamellifer ximenez (Hanna), 1923 
Figure 3 
Bulimulus ximenez Hanna, 1923, pp. 497-499, pl. 8, figs. 4-9, Isla Carmen and Isla Coronados. 
Lowe, 1933, p. 110, Isla Carmen, under rock slides. 
Bulimulus (Leptobyrsus) ximenez Hanna, Jacobson, 1958, p. 6, Marquer Bay, Isla Carmen, in an 
arroyo, east of the bay. 


Bulimulus carmen Pilsbry and Lowe, 1932, pp. 50, 51 [Not pl. 5, fig. 2], Salinas Bay, Isla Carmen. 

Bulimulus (Leptobyrsus) carmen Pilsbry and Lowe, Pilsbry, 1935, p. 2, pl. 1, fig. 7 {holotype}. 

Records. — Marquer Bay, Isla Carmen, 100 mature, dead specimens (Emerson et al.). 

Puerto Ballandra, Isla Carmen, at elevations of 100 to 300 feet, 17 dead specimens 
(Banks, Moran, Soulé). 

South end of Isla Coronados, at all elevations, 14 dead specimens (Moran). 

Remarks. — Bulimulus ximenez was described by Hanna (1923:497-499) from samples 
collected at Marquer Bay (type locality) and several other localities on Isla Carmen and from 
Isla Coronados. He described the internal denticle as follows: “square in shape, slightly emar- 
ginate or sinuous on its outer edge.” However, this is by no means a fixed character, as was 
noted previously (Jacobson, 1958:6). A series of mature specimens, those which possess a 
strong reflection of the peristome and a well-developed parietal callus, were opened and the 
denticles were measured. This examination showed that this character is quite variable in the 
two samples studied (see table 2 and discussion below). We must conclude that “Bulimulus” 
ximenez is merely an insular population of the wide ranging species Rabdotus lamellifer, one 
which attains a noticeably larger size and which deserves only subspecific recognition. 

Pilsbry and Lowe (1932:50-51) believed “Bulimulus” carmen, based on specimens from 
Salinas Bay, Isla Carmen, could be distinguished from “Bulimulus” ximenez, by having the 


324 San Dieco Society oF Naturav History { VoL. 13 


shell less solid and the outline rather stout, and by possessing an internal denticle that is 
“much weaker.” These characters, however, are quite variable in this group, as Jacobson 
(1958:6) has already stated, and the type specimens of Bulimulus carmen show no other 
characters that serve to separate them from topotypic specimens of Rabdotus lamellifer ximenez. 

The small sample in the present collection from Isla Coronados is composed of specimens 
having a large, well-developed denticle. On the basis of size, height and width, this insular 
population is referred to R. |. ximenez. Hanna (1923:498) recorded 77 specimens from Isla 
Coronados under the name of Bulimulus ximenez. 

The denticles of this species are extremely variable in shape, no two being alike. In this 
they resemble the internal protuberances in such species as Jeanneretia subtussulcata (Pfeiffer) 
from Cuba and Cepolis cepa (Miller) from Hispaniola. Although the shell in life is, in the 
words of Hanna (1923:497), “light vinaceous flesh color”, the denticle of two fresh 
specimens in the present collection is white at the summit, gradually becoming flesh color at 
the base. The outer edge is either strongly emarginate, weakly so, or in some specimens, no 
thicker than the rest of the structure. The denticle varies in thickness and is either developed 
on one plane or moderately to strongly twisted horizontally as well as vertically. The strength 
of the denticle seems to bear no relationship to maturity of the shell; a shell with a weak 
parietal callus and a scarcely reflected peristome, both of which are signs of immaturity, has 
a denticle 6.3 mm. long and 4.0 mm. high, whereas a shell with a strongly raised callus and a 
strongly reflected peristome, signs of maturity, has the denticle reduced to a mere sinuous 
lamella, 1.8 mm. high, along the axis. The ventral side of the denticle is either straight or 
variously indented at one or more points. In some specimens it is so high that it practically 
closes, at its peak, the passage of the whorl. 


Rabdotus (Leptobyrsus) veseyianus (Dall), 1893 
Bulimulus (Leptobyrsus) veseyianus Dall, 1893, p. 645, pl. 71, figs. 4, 5, Isla Espiritu Santo 
(Belding). 
Bulimulus veseyianus Dall, Pilsbry, 1898, pp. 160, 161, pl. 21, figs. 92, 93 [enlarged copies of Dall’s 
figs.}. Hanna, 1923, pp. 499, 500, pl. 8, figs. 1-3, Isla Espiritu Santo and Isla Partida [“North Island” }. 


Records. — Ballena Cove, west side of Isla Espiritu Santo, elevation about 100 feet, 88 
mature, 2 immature, dead specimens, along the side of an arroyo, among rocks (Emerson 
et al.). 


West side of Isla Partida in an arroyo, 3 complete specimens, many fragmental specimens 
perhaps eaten by rodents (Emerson et al.). 

Remarks. — The axial internal ornamentation in this species differs decidedly from the 
internal denticle of R. lamellifer. The denticle does not rise abruptly, but rather is a sinuous 
enlargement of the axis itself, terminating rather sharply at the summit of the body whorl. It 
does not extend beyond this whorl. An examination of 11 specimens picked at random, how- 
ever, does not seem to substantiate Hanna’s (1923:500) statement that “the development of 
this denticle is somewhat dependent upon age, younger individuals having the lamella projected 
no more than in B. spirifer.” The more mature shells, judging this by the thickness of the 
parietal wash and the degree of reflection of the peristome, do not generally have a higher 
and thicker lamella than less mature examples. 

Our measurements indicate a greater range of size than appears in the measurements given 
by Hanna (1923:500), 41.7 to 36.5 mm.; our specimens range from 44.7 to 34.5 mm. in 
altitude (table 1). 

This species is easily distinguished by its large size, strongly reflected and wide peristome, 
and characteristic internal lamella. The peristome in a fresh specimen is flesh colored, becoming 
white along the reflected margin. The aperture of the shell, beyond the slight extension of the 
flesh color of the lip, is white, as are the axis and the lamella. 

Isla Partida, where some recognizable fragments and three mature individuals were 
found, is a smaller island directly north of Isla Espiritu Santo. The two islands are very close 
to one another, being separated by a narrow channel. 


1964 } EMERSON AND JACOBSON: TERRESTRIAL MoLiusks 325 


Fig. 4. Rabdotus (Hannarabdotus) slevini (Hanna). a-c, specimens from Isla Santa 
Cruz (= Bulimulus santacruzensis Hanna); d-f, topotypic specimens. ™ 1. 


Subgenus Hannarabdotus, new subgenus 


Type species. — Bulimulus slevini Hanna, 1923. 

Description. — Shell is similar to Leptobyrsus, but lacks internal ornamentation on the 
axis. Columellar fold, which is visible at the aperture, may or may not be present. Peristome 
is more or less expanded or may be revolute, but is not simple as in Rabdotus (sensu stricto). 

Remarks. — The type species of Rabdotus, Bulimulus dealbatus Say, 1821, differs in hav- 
ing a simple peristome, sometimes thickened within. As was suggested previously (Jacobson, 
1958), the many bulimuli of Baja California that have the lip strongly expanded and frequently 
revolute appear to form a natural group of species that deserve subgeneric recognition. 

Species included in Hannarabdotus are: baileyi Dall, beldingi Cooper, ceralboensis Hanna, 
excelsus Gould (= elatus Gould), hannai Pilsbry, inscendens W. G. Binney, johnstoni Hanna, 
pallidior Sowerby, slevini Hanna [= santacruzensis Hanna}, and vegetus Gould {= ?pallidior}. 

We take extreme pleasure in naming this taxon in honor of Dr. G Dallas Hanna, who 
has made numerous contributions to the knowledge of the west American terrestrial mollusks. 


Rabdotus (Hannarabdotus) slevini (Hanna), 1923 
Figure 4 

Bulimulus santacruzensis Hanna, 1923, pp. 487, 488, pl. 7, figs. 12-15, Isla Santa Cruz. Lowe, 1933, 
p. 109, Isla Santa Cruz. 

Bulimulus slevini Hanna, 1923, pp. 488, 489, pl. 7, figs. 16-19, Isla Monserrate. Lowe, 1933, p. 109, 
Isla Monserrate. 

Bulimulus (Orthotomium) santacruzensis Hanna, Jacobson, 1958, p. 6, Isla Santa Cruz, southwest side. 

Records. — Isla Monserrate, 34 dead specimens and 2 live specimens, under rocks (Harbi- 
son, Moran); southeast side of Isla Monserrate, 87 dead and 3 live specimens (Emerson). 

Isla Santa Cruz, 114 dead specimens, at all elevations (Moran, Wiggins); southwest side 
of Isla Santa Cruz, at elevations greater than 50 feet, 72 dead and 1 living specimen 
(Emerson). 

Remarks. — Hanna (1923) considered the bulimulid populations occurring on Islas Mon- 
serrate and Santa Cruz to represent two closely related, but distinct species, which he named 
Bulimulus slevini and B. santacruzensis, respectively. He believed that the shells of the latter 
could be separated from those of the former by being larger on the average and by having a 
broader peristome and a more strongly developed columellar fold. However, a visual examina- 
tion of the large samples in the present collection indicates that these characters can not be 
used to separate most of the specimens from the two islands. Even the columellar fold was 
found to be a variable character in both populations. 


326 San Disco Society of NaturaL History {Wort 


TABLE 3 
Measurements (in mm.) of specimens of Rabdotus (Hannarabdotus) slevini (Hanna) 
in the present collection. 
Number of Altitude Diameter 
specimens Range Average Median Range Average Median 
Isla Monserrate, 85 30.0-38.9 34.15 34.2 12.0-15.1 Sal 13.7 
southeast side 
Isla Monserrate, 33 30.3-39.7 33.86 D307, 12.0-16.0 13.40 13.4 
all elevations 
Isla Santa Cruz,! 62 32.7-42.3 37.01 36.9 12.5-15.8 13.96 13.9 
all elevations 
Isla Santa Cruz,! 29 32.5-40.0 36.44 36.5 125-15:0 13.98 14.1 
arroyos 
Isla Santa Cruz,! 68 29.9.43.4 35.42 3) IPD=UWDH 13.68 1327, 


southwest side 
1Type locality of Bulimulus santacruzensis Hanna. 


Inasmuch as these two taxa were based on insular populations, we undertook, with the 
generous assistance of Dr. Meredith L. Jones of the American Museum of Natural History, 
an analysis of the two populations to determine whether statistically valid differences were 
observable among the various samples at our disposal. These samples consisted of 85 specimens 
from the southeast side of Isla Monserrate, 33 specimens from the southwest side of Isla Santa 
Cruz, 29 specimens from arroyos of Isla Santa Cruz, 62 specimens from Isla Santa Cruz, all 
elevations, and 68 specimens from the southwest side of Isla Santa Cruz( see table 3). 

Altitudes and diameters of all specimens were measured and altitude-diameter ratios of 
all specimens were calculated. Statistical comparisons among the five lots were made by the 
use of Student’s “t” test and the modification of the 75% rule presented by Mayr, Linsley, 
and Usinger (1953:145-148). 

According to Dr. Jones, no consistent valid differences were noted among all of the com- 
parisons made. Therefore, we feel that this justifies our synonymizing Rabdotus santacruzgensis 
with R. slevini and explains our difficulty in separating collections from the two islands in 
question. This finding is in harmony with the conclusion of Mayr et al. (1953:31) who 
stated: “The difference [between two populations} must be sufficiently great so that it is pos- 
sible to identify the great majority of specimens without knowledge of their provenience.” 

Although the name Bulimulus santacruzensis has page priority, we have elected to use 
the first reviser’s prerogative in selecting “Bulimulus” slevini, inasmuch as Hanna (1923) de- 
scribed that taxon thoroughly but limited his description of R. santacruzensis to the apparent 
differences between it and R. slevini. 

A fresh, immature specimen of 414 whorls from Isla Santa Cruz shows the following 
characteristics: nuclear whorls with numerous, vertical, regularly spaced, axial riblets, interstices 
about double the width of the ribs; riblets progressively more oblique on the first postnuclear 
whorl, gradually growing obsolete and being replaced by retractive, oblique, irregular growth 
lines. 

Rabdotus (Hannarabdotus) johnstoni (Hanna), 1923 
Bulimulus joknstoni Hanna, 1923, p. 491, pl. 7, figs. 1-6; pl. 11, fig. 3, Isla Santa Catalina. 


Bulimulus johnsoni {sic} Hanna, Lowe, 1933, p. 109, Isla Santa Catalina, “living under rock slides 
on the north side of the main canyon.” 


Records. —Isla Santa Catalina, west side, under rocks and dead trees, dead shells num- 
erous, 95 mature, 10 immature, dead specimens (Emerson et al.) ; Isla Santa Catalina, common 
on backside of rocks and under dead trees, 83 mature, many immature, live specimens 
(Emerson) ; Isla Santa Catalina, east side, 23 dead, 6 live specimens (Emerson et adl.). 

Remarks. — This species, the type lot of which consisted of 2000 individuals, is appar- 
ently one of the most common and easily collected species on the islands. It is the smallest of 
the known insular representatives of this subgenus. 


1964 | EMERSON AND JACOBSON: TERRESTRIAL MoL_usks 527 


Fig. 5. Holotype of Micrarionta (Eremarionta) rowelli bechteli. X 1.5. 


Rabdotus (Hannarabdotus) ceralboensis (Hanna), 1923 

Bulimulus ceralboensis Hanna, 1923, p. 490, pl. 7, fig. 11; pl. 11, figs. 2, 4, west of Ruffo’s ranch 
house, Isla Ceralbo. 

Records. — El Mostrador, north of Ruffo’s ranch, west side of Isla Cerralvo, 88 dead 
specimens, fragments, uncommon at 75 to 150 feet among boulders (Wiggins and Emerson). 

Remarks. — This species is most readily determined, aside from its locality, by the strongly 
expanded, but not reflected, peristome, the large size, the absence of a columellar fold and the 
generally smooth surface. 

In the present collection are specimens that have greater dimension than those of the 
type lot (Hanna, 1923:490), which apparently consisted of only four specimens. Our largest 
specimen measures 42.8 mm. in altitude and 22.5 mm. in diameter, whereas the largest in the 


type lot is 36.8 mm. by 19.7 mm. 


Family HELMINTHOGLYPTIDAE 
Genus Micrarionta Ancey, 1880 

Micrarionta Ancey, 1880, p. 334. 
Type species. — Helix facta Newcomb, 1864, by monotypy. 

Subgenus Eremarionta Pilsbry, 1913 
Eremarionta Pilsbry, 1913, p. 382. 
Type species. — Micrarionta desertorum Pilsbry and Ferriss, 1908, by monotypy. 

Micrarionta (Eremarionta) rowelli bechteli, new subspecies 
Figure 5 

Type locality. —Isla San Esteban, Sonora, Mexico, in rock slide, G D. Hanna leg., April 
2, 1953, C.A.S. no. 36377. Another lot in the present collection was taken on the same island 
by Reid Moran at 300 meters, March 22, 1962. 

Type depository. — Holotype and four paratypes in the California Academy of Sciences, 
San Francisco; one paratype in the San Diego Natural History Museum; three paratypes in 
the American Museum of Natural History. 

Diagnosis. —A Micrarionta of the group of M. rowelli (Newcomb) distinguished from 
M. rowelli mexicana Pilsbry and Lowe (1934) by its somewhat more depressed shape, smaller 
umbilicus and less ample aperture, and from M. rowelli mearnsi Bartsch (1904), another sub- 
species occurring in the general region, by its larger size and much larger umbilicus. 

Description.— The depressed shape and postnuclear retractive sculpture are typical of 
the group of M. rowelli. The papilose nuclear whorl is eremariontid. The umbilicus is deep, 
showing most of the whorls on the inner side, contained about five times in the diameter. The 
number of whorls is a little less than five, the last one descending sharply at the aperture. The 
lip is barely expanded and non-reflected, widening at the columellar insertion. The aperture 
is circular, oblique. 

The dead and somewhat bleached shell is shiny, light brown cn the spire and the upper 
half of the body whorl, where this area is sharply limited just above the periphery by a narrow 
brown band, set off above by a somewhat wider, whitish area. The subperipheral and basic area 
is much paler, approaching cream color. 


328 SAN D1seGo Society oF NATURAL HIstTory {VoL. 13 


Measurements. — Measurements in millimeters are as follows: 


Diameter Height Aperture 

Holotype 14.8 To 4.5 Hanna leg. 
Paratypes 14.4 6.5 4.6 se 

137 7.0 4.4 s 

14.2 Ip? 4.4 -. 

14.7 7-4 5.0 2 

13.7 8.2 5.0 Moran leg. 

27 6.8 Dell Hy 

12.6 6.8 4.8 “a 


Remarks. — This is the only insular subspecies known for the widely distributed group 
of M. rowelli. It is easily distinguished from the two geographically related subspecies — 
from mearnsi Bartsch (San José Mountains, Sonora) by its decidely larger umbilicus and from 
mexicana Pilsbry and Lowe (near Puerto Penasco, Sonora) by its obviously less capacious 
aperture. This feature, which is apparent to the eye, is supported by measurements. The lateral 
diameter of the aperture of paratypic specimens of M. mexicana in the collection of the 
Academy of Natural Sciences of Philadelphia measure from 5.9 to 6.2 mm., whereas the new 
subspecies has measurements from 4.4 to 5.1 mm. for specimens of about the same size. 

We take pleasure in naming this new subspecies in honor of Mr. K. K. Bechtel of the 
Belvedere Scientific Fund. 

Family UROCOPTIDAE 


Genus Coelocentrum Crosse and Fischer, 1872 
Coelocentrum Crosse and Fischer, 1872, p. 302. 
Type species. —Cylindrella turris Pfeiffer, 1856, by original designation. 
Subgenus Spartocentrum Dall, 1895 
Spartocentrum Dall, 1895, p. 3. 
Teneritia Mabille, 1897, p. 79; type species Berendtia digueti Mabille, 1895 [= Coelocentrum (Sparto- 
centrum) digueti (Mabille) }, here designated. 


Type species. —Cylindrella (Urocoptis) irregularis Gabb, by original designation. 
Remarks. — Mabille (1897:79) designated “Berendtia digueti J. Mabille et B. minorina 
J. Mab.” the cotypes of Teneritia. We have restricted the type species to the former. 
Coelocentrum (Spartocentrum) vanduzeei Hanna, 1923 
Coelocentrum vanduzeei Hanna, 1923, pp. 508, 509, pl. 9, figs. 31-34; pl. 11, fig. 7, Puerto Escon- 


dido, Baja California. Lowe, 1933, p. 110, “Puerto Escondido in company with . . . Berendtia taylori 
(Pfr.).” 


Records. — Puerto Escondido, Baja California, 1 complete specimen, 2 fragments (Moran, 
Soulé). 

Remarks. — The single complete specimen in the present collection has almost 18 whorls; 
the aperture is on a very small extension. In other respects, the specimen agrees well with 
the original description. 

Coelocentrum (Spartocentrum) oweni Hanna, 1923 

Coelocentrum oweni Hanna, 1923, pp. 511-512, pl. 9, figs. 3-8; pl. 11, fig. 8, Agua Grande, Isla 
Carmen (Virgil Owen). 

Records. — Marquer Bay, Isla Carmen, 88 complete specimens, many fragments, all eleva- 
tions (Emerson et al.). 

Isla Danzante, 14 complete, but not mature, specimens, many fragments (Emerson et al.). 

Remarks. — Hanna (1923:510) refers a series of 23 specimens collected at Marquer Bay 
to C. insulare Hanna, which was described from Puerto Ballandra, Isla Carmen. However, if 
his shells present the same range of dimensions as the present lot from Marquer Bay, Isla 
Carmen, it is apparent that they should have been referred instead to C. oweni from Agua 
Grande, Isla Carmen. Agua Grande is nearer to Marquer Bay than to Puerto Ballandra, but 
is situated on the opposite side of the island. The measurements for altitude and diameter of 
the present specimens are definitely larger than the measurements given by Hanna for C. in- 
sulare. They are even larger than the measurements of C. oweni, but we prefer to assign them 
to the latter taxon rather than to establish still a third species on the basis of this character alone. 


1964 } EMERSON AND JACOBSON: TERRESTRIAL MoLLusks 329 


TABLE 4 
Measurements (in mm.) of Coelocentrum oweni. 
Number of 
Altitude Diameter Whorls 
19.7 4.4 15 
17.9 Be/, 14 
17.4 4.8 141, 
We, 4.7 15 
19.4 4.3 161, 
17.0 4.3 13Y, 
16.5 4.4 14, 
16.3! 4.9 13, 
172. 4.6 16 
17.8 4.5 14/, 
14.6 4.6 14 
16.6 4.7 14 


1Probably not completely mature, but included to show extreme in size of diameter. 


In measuring the altitude of these shells, one cannot always easily determine what speci- 
mens are fully mature. The number of whorls alone, as can be seen from table 4, is not a 
sufficiently accurate index. Nor is the degree of soluteness of the aperture definitive, as was 
noted by Hanna (1923:507). The peristome of a mature shell does not seem to differ 
materially from that of an immature one. Nevertheless, to insure some sort of uniformity, 
the shells measured were limited to specimens with a noticeably solute aperture, on the sup- 
position that whatever can be said about specimens with adnate apertures, certainly those with 
free apertures must be mature or nearly so. 

A strong possibility exists, as Hanna noted (1923:510), that there is only a single 
species of Spartocentrum on Isla Carmen, the differences in the populations probably being 
due to local ecological differences. Careful field studies are needed to settle this question. 

Judging by the number of whorls, the specimens collected on Isla Danzante are not 
mature. However, they closely resemble specimens of C. oweni from Isla Carmen and there 
seems to be no reason to question Hanna’s statement (1923:511) that only one species is 
involved. 

Family PUPILLIDAE 
Genus Pupoides Pfeiffer, 1854 

Pupoides Pfeiffer, 1854, p. 192. 

Leucochila von Martens, 1860, p. 296; type species Pupa fallax [= -Pupoides albilabris (C. B. Adams, 
1841) }, by original designation. 

Leucochiloides Pfeiffer, 1879, p. 292; type species Bulimus lardeus Pfeiffer, 1852, by subsequent 
designation of Connolly (1912:176). 

Type species. — Bulimus nitidulus Pfeiffer, 1839, by subsequent designation of Kobelt 
(1880:267). 

Remarks. — Although Pilsbry (1948:920) refers Themapupa Iredale, 1930 (type species 
by montotypy Pupa beltiana Tate, 1894) to the synonymy of Pupoides, Zilch (1959:168) 


gives subgeneric rank to Iredale’s taxon. 


Pupoides (Pupoides) albilabris (C. B. Adams), 1841 


P. {upa} albilabris C. B. Adams, 1841, p. 271. 

Pupoides marginatus (Say), Hanna, 1923, p. 514. Pilsbry, 1926, p. 249. Not Cyclostoma marginatum 
Fischer von Waldheim, 1807, p. 219. 

Pupoides albilabris (C. B. Adams), Pilsbry, 1948, p. 921, fig. 499:1-7. 

Records. — Marquer Bay, Isla Carmen, 1 specimen (Emerson). 

Remarks. — Hanna (1923:514) records this species from the following islands in the 
Gulf of California: Islas Angel de la Guarda, Tortuga, San Lorenzo, San Esteban, and 
Monserrate. The present record should be added to this list. 


330 SAN Drieco Society oF Naturat History { VoL. 13 


Pilsbry (1948:923) sets forth the reason for the change of name of P. marginatus (Say) 
to P. albilabris (C. B. Adams). He also gives an exhaustive synonymy for the species. 


Pupoides (Pupoides) catalinensis Hanna, 1923 
Pupoides catalinensis Hanna, 1923, pp. 514-515, pl. 10, figs. 1-4, Isla Santa Catalina. Pilsbry, 1926, 
p. 249, text fig. 17. 
Records. — East side of Isla Santa Catalina, 1 dead specimen (Emerson). 
Remarks. — The height of the shell and its pronounced cylindrical outline sets this species 
off from the related P. albilabris that occurs on other islands in the vicinity. 


SpreciEs LISTED BY COLLECTING LOCALITIES 


Isla San Esteban, March 21, 22: Micrarionta (Eremarionta) rowelli bechteli, new subspecies. 

Isla San Marcos, March 28, 29: Rabdotus (Leptobyrsus) lamellifer lamellifer (Pilsbry). 

Isla Tortuga, March 30: Rabdotus (Leptobyrsus) dentifer (Mabille). 

Mulegé estuary, March 31: Rabdotus (Leptobyrsus) lamellifer lamellifer (Pilsbry). 

Puerto Santispaquis, April 1: Rabdotus (Leptobyrsus) lamellifer lamellifer (Pilsbry). 

Isla Coronados, April 3, 4: Rabdotus (Leptobyrsus) lamellifer ximenez (Hanna). 

Isla Carmen, April 4, 5: Rabdotus (Leptobyrsus) lamellifer ximenez (Hanna). 
Coelocentrum (Spartocentrum) oweni Hanna. 
Pupoides (Pupoides) albilabris (C. B. Adams). 

Puerto Escondido, April 6, 7: Rabdotus (Leptobyrsus) lamellifer lamellifer (Pilsbry) . 

Coelocentrum (Spartocentrum) vanduzeei Hanna. 

Isla Danzante, April 7: Rabdotus (Leptobyrsus) lamellifer lamellifer (Pilsbry). 
Coelocentrum (Spartocentrum) oweni Hanna. 

Isla Monserrate, April 9, 10: Rabdotus (Hannarabdotus) slevini (Hanna). 

Isla Santa Catalina, April 10: Rabdotus (Hannarabdotus) johnstoni (Hanna). 

Pupoides (Pupoides) catalinensis Hanna. 

Isla San José, April 11, 12: Rabdotus (Rabdotus) sufflatus (Gould). 

Isla Cerralvo, April 15-17: Rabdotus (Hannarabdotus) ceralboensis (Hanna). 

Isla Santa Cruz, April 18: Rabdotus (Hannarabdotus) slevini (Hanna). 

Isla San Diego, April 18, 19: Rabdotus (Leptobyrsus) chamberlini (Hanna). 

Isla Espiritu Santo, April 20: Rabdotus (Leptobyrsus) veseyianus (Dall). 

Isla Partida, April 20: Rabdotus (Leptobyrsus) veseyianus (Dall). 


LITERATURE CITED 
Apams, C. B. 
1841. Catalogue of the Mollusca of Middlebury, Vt., and vicinity, with observations. 
Amer. Jour. Sci. Arts 40:266-277. 
ALBERS, J. C. 
1850. Die Heliceen, nach natiirlicher Verwandtschaft systematisch geordnet. Ist ed. 
Berlin. 262 pp. 
AncEY, C. F. 
1880. Descriptions de mollusques nouveaux. Le Naturaliste, 2me Année 1:334-335. 
BarTSCH, PAUL 
1904. Notes on the genus Sonorella, with descriptions of new species. Smiths. Misc. 
Coll. 47:187-200, pls. 28-33. 
Binney, W. G. 


1859. The terrestrial air-breathing mollusks of the United States. Boston. Vol. 4. 
viii + 207 pp., pls. 75-80. 


1964 | EMERSON AND JACOBSON: TERRESTRIAL MoLiusks 331 


ConNoLLy, M. 
1912. A revised reference list of South African non-marine Mollusca: with descriptions 


of new species in the South African Museum. Ann. South African Mus. 11:59- 
306, pl. 2. 


Cooper, J. G. 
1893a. On land and fresh water shells of Lower California. Proc. Calif. Acad. Sci.. 
ser. 2, No. 2. 3:207-217. 
1893b. On land and fresh water shells of Lower California. Proc. Calif. Acad. Sci. 
set. 2, No. 3.3':336-344, pls. 13, 14: 
1894. On land and fresh water shells of Lower California. Proc. Calif. Acad. Sci., 
set.) 25) Nowa. 4: 130-1435 pls, 5,6: 


Crossk, H., AND P. FiscHER 
1872. Diagnoses molluscorum novorum, Reipublicae Mexicanae incolarum. Jour. 


Conchyl. 20:301-303. 


DATE. We Fi: 
1893. Land shells of the genus Bulimulus in Lower California, with description of 
several new species. Proc. U.S. Nat. Mus. 16:639-647, pls. 71, 72. 


1895. Diagnoses of new mollusks from the survey of the Mexican boundary. Proc. 
U.S. Nat. Mus. 18:1-6. 


FiscHER, P., AND H. Crosse 
1870-1878. Etudes sur les mollusques terrestres et fluviatiles du Mexique et du Guate- 
mala. In Recherches zoologiques pour servir a l’histoire de la faune de |’Amérique 
Centrale et du Mexique, publiées sous la direction de M. Milne Edwards. Paris. 
1-702 pp., pls. 1-31. 


FISCHER VON WALDHEIM, G. 
1806-1807. Muséum Demidoff, ou catalogue systématique et raisonné des curiosités de 
la nature et de I’art . . . Moscow. {not seen}. 


GouLp, A. A. 
1853. Descriptions of shells from the Gulf of California and the Pacific coasts of 
Mexico and California. Boston Jour. Nat. Hist. 6:374-408, pls. 14-16. 


Hanna, G D. 
1923. Expedition of the California Academy of Sciences to the Gulf of California in 
1921. Land and freshwater mollusks. Proc. Calif. Acad. Sci., ser. 4, 12:483-527. 


HERRMANNSEN, A. N. 
1846-1847. Indicis Generum Malacozoorum Primordia. Cassellis. Vol. 1. xlii +> 637 pp. 


Jacosson, M. K. 
1958. Results of the Puritan-American Museum of Natural History Expedition to 
western Mexico. 3. The terrestrial mollusks. Amer. Mus. Novitates 1899. 14 pp. 


KosBELT, W. 
1876-1881. Illustriertes Conchylienbuch. Nurnberg [2 vols.}, xvi + 392 pp., 112 pls. 


Leacu, W. L. 
1814. Zoological miscellany; being descriptions of new, or interesting animals. London. 


1:1-144, pls. 1-60. 


Linpsay, G. E. 
1962. The Belvedere Expedition to the Gulf of California. Trans. San Diego Soc. Nat. 
Hist. 13:1-44. 


332 San Drieco Society oF Natura History {Vou 13 


Lowe, H. N. 
1933. The cruise of the ‘“Petrel.”’ Nautilus 46:73-76, 109-115. 


MABILLE, JULES 
1895. Mollusques de la Basse Californie. Bull. Soc. Philomat. Paris, ser. 8, 7:54-76. 
1897. Notitiae malacologicae. Bull. Soc. Philomat. Paris, ser. 8, 9:78-102. 


Martens, E. von, In J. C. ALBERS 
1860. Die Heliceen nach naturlicher Verwandtschaft systematisch geordnet. 2nd ed. 
Leipzig. xviii + 359 pp. 
Mayr, E., E. G. LinsLey AND R. L. USINGER 
1953. Methods and Principles of Systematic Zoology. McGraw-Hill Book Co., Inc. 
New York. ix + 328 pp. 
Paropiz, J. J. 


1962. New and little-known species of South and Central American land snails (Buli- 
mulidae). Proc. U.S. Nat. Mus. 113:429-456. 


PFEIFFER, LOUIS 
1853. Diagnosen neuer Heliceen. Zeits. f£. Malak. 10:51-58. 
1854. Zur Molluskenfauna der Insel Cuba. Malak. Blatt. 1:170-213. 
1865. Descriptions of five new species of land-shells, from the collection of the late 
Hugh Cuming. Proc. Zool. Soc. London for 1865:831-832. 
1879-1881. Nomenclator Heliceorum viventium qui continentur nomina omnium hujus 
familiae generum et specierum hodie cognitorum diposita ex afhnitate naturali. 


Kassel. 606 pp. 


Pitspry, H. A. 

1896. Sculpture of the apical whorls, a new character for distinguishing groups of 
Bulimuli. Nautilus 9:112-115. 

1897a. New Lower Californian Bulimuli. Nautilus 10:102-103. 

1897b-1898. American Bulimulidae: Bulimulus, Neopetraeus, Oxychona, and South 
American Drymaeus. Manual of Conchology, ser. 2, vol. 11. Philadelphia. 339 
pp., 51 pls. 

1913. Notes upon some Lower Californian helices. Proc. Acad. Nat. Sci. Philadelphia 
65 :380-393, pls. 15, 16. 

1922-1926. Pupillidae (Orculinae, Pagodulinae, Acanthinulinae, etc.). Manual of Con- 
chology, ser. 2, vol. 27. Philadelphia. v + 369 pp., 32 pls. 

1935. Description of middle American land and fresh water Mollusca. Proc. Acad. 
Nat. Sci. Philadelphia 87:1-6, pl. 1. 

1944. Peruvian land Mollusca — II. Nautilus 57:118-127. 

1946-1948. Land Mollusca of North America (North of Mexico). Acad. Nat. Sci. 
Philadelphia, Monog. 3. Vol. 2, part 1: vi + 520 pp., part 2: pp. i-xlvii, 521- 
1113, figs. 282-585. 


Picssry, H. A., AND H. N. Lowk 


1932. New west American species of Bulimulus and Nassa. Nautilus 46:49-52. 
1934. Some desert helices of the genus Micrarionta. Nautilus 48:67-68. 


WEYRAUCH, WOLFGANG 
1956. The genus Naesiotus with descriptions of new species and notes on other Peruvian 
Bulimulidae. Proc. Acad. Nat. Sci. Philadelphia 108:1-17, pl. 1. 
ZILCH, ADOLF 
1959-1960. Gastropoda, Euthyneura. In O. H. Schindewolf, Handbuch der Palao- 
zoologie. Berlin. Vol. 6. xii + 834 pp., 2515 figs. 


TRANSACTIONS 
OF THE 


SAN DIEGO SOCIETY OF NATURAL HISTORY 


VoLuME 13, No. 17, pp. 333-368 


INVERTEBRATE MEGAFOSSILS 
OF THE BELVEDERE EXPEDITION 
TO THE GULF OF CALIFORNIA 


BY 


Wittiam K. EMERSON 


Chairman, Department of Living Invertebrates 
American Museum of Natural History, 
and Research Associate 
San Diego Natural History Museum 


AND 


LEO GEORGE HERTLEIN 


Curator of Invertebrate Paleontology 
Department of Geology 
California Academy of Sciences 


SAN DIEGO, CALIFORNIA 
PRINTED FOR THE SOCIETY 


DECEMBER 30, 1964 


Penasco 


Guoymas 
y 28° 


4 


Punta Santa 
Antonita 


lOO Miles 


100 Kilometers 
W2° 


Fig. 1. Map showing the areas from which invertebrate megafossils were collected by the 
Belvedere Expedition. 
Localities indicated by numbered circles are as follows: 1, Isla Angel de la Guarda; 2, Isla Partida; 


3, Isla Salsipuedes; 4, Isla San Esteban; 5, El Pulpito; 6, Isla Coronados; 7, Isla Carmen; 8, Isla Mon- 
serrate; 9, Isla San Diego; 10, Isla San José; 11, Isla Cerralvo. 


JAN 15 1965 


HARVARD 


INVERTEBRATE MEGAFOSSILS | ).)))2e6)ry 
OF THE BELVEDERE EXPEDITION 
TO THE GULF OF CALIFORNIA 


BY 
WILLIAM K. EMERSON AND LEO GEORGE HERTLEIN 


CONTENTS 
RETO CICeOt. meester ee eye Po PSP A See Se ON, ie Bian ee ene ee 335 
Descriptions of Collecting Localities, Faunal Lists, and Age Determinations... 336 
List of Species by Taxonomic Groups, with Age and Collecting Localities... 348 
Sys age ate Bd BS oat wo 01 eee eee oe ki nc ore eek an SEP I ee ee ete Phys, eee LN 352 
Pcetactite® ited meee. se ae Gee G Med, Jf EA 2 ae Ce aR ge eo Tl Road 5 Lee ee 367 
INTRODUCTION 


This paper records the fossil invertebrates, mostly mollusks, echinoids and corals, collected 
by members of the scientific party during the course of the expedition. Fossils ranging in age 
from early Pliocene to late Pleistocene were obtained from localities on ten islands in the Gulf 
of California and at El Pulpito on the peninsula of Baja California. Some of the localities 
were previously unknown, and the collections include guide fossils that have aided in the 
determination of more precise ages for several poorly known deposits. As a result of our 
taxonomic studies of the collections, we have proposed herein two new subgeneric names, 
Bechtelia, in the gastropod family Bursidae, and Morunella, in the gastropod family Thai- 
didae. We also have described a new species of gastropod, Cancellaria (Aphera) wigginsi, 


from Pleistocene deposits on Isla Monserrate. 


Most of the fossils were collected by the senior author, who divided his field work 
between the- Recent and fossil invertebrates, and by Ira L. Wiggins, who collected fossils 
incidental to his botantical studies. Other members of the scientific party, including Richard 
C. Banks, Charles F. Harbison, George E. Lindsay, Reid Moran, Charles E. Shaw, and 


Michael Soulé, also assisted in the collection of fossils when time permitted. 


The expedition, sponsored by the San Diego Natural History Museum and the Belvedere 
Scientific Fund, started from Bahia de los Angeles, Baja California, on March 15, 1962, and 
terminated at La Paz, Baja California, on April 21, 1962. A general account and log of the 
expedition, including charts showing the route and anchorage stops of the expedition’s ship, 
M/V “San Agustin IT”, was published by Lindsay (1962). A report on the terrestrial 
mollusks obtained by the expedition has been published (Emerson and Jacobson, 1964). 


Apparently the first geologist to record the presence of fossil invertebrates in the area of 
the Gulf of California was the German scientist Grewingk (1848), who described the geology 
of parts of this region. Although a considerable number of studies on the geology of this area 
have appeared subsequently, our understanding of the general stratigraphy and knowledge of 
the Tertiary paleontology of many parts of this region is far from complete. For summaries 
and bibliographies of the previous reports pertaining to late Cenozoic fossils from the Gulf 
of California area, the reader is referred to Hanna and Hertlein (1927), Beal (1948; see 
also Anon., 1924), Durham (1950), and Hertlein (1957). Of particular interest to the 


336 San Deco Society oF NaturaL History {Vor 13 


present study are the reports concerning the Pliocene and Pleistocene faunas from the islands 
in the Gulf and from El Pulpito on the peninsula. Hanna and Hertlein (1927) described 
the fossil invertebrates collected by the late Dr. Fred Baker on the expedition of the California 
Academy of Sciences to the Gulf of California in 1921. Durham (1950) authored a report 
on the megascopic paleontology and marine stratigraphy resulting from a cruise of the schooner 
“E. W. Scripps” of the Scripps Institution of Oceanography to the Gulf of California in 
1940. Hertlein (1957) discussed the Pliocene and Pleistocene fossils collected by Dr. Edwin 
T. Hammond of the University of California in the region about La Paz in 1949, and those 
obtained by Dr. G Dallas Hanna and J. R. Slevin of the California Academy of Sciences 
while on an expedition of the M/V “Orca” to the Gulf of California, including stops at 
El Mostrador on Isla Cerralvo and at El Pulpito, in 1953. Emerson (1960a) reported 
Pleistocene invertebrates from Isla Cerralvo obtained by the “Puritan”-American Museum of 
Natural History expedition to western Mexico in 1957. 


A number of people have kindly assisted us in the completion of this study. Dr. A. 
Myra Keen provided us with information on the W. W. Valentine collection of fossils in 
the Department of Geology of Stanford University. Mr. E. P. Chace of the San Diego 
Natural History Museum, Mr. and Mrs. John Q. Burch of Los Angeles, California, and 
Mr. Gordon Usticke of St. Croix, Virgin Islands, provided specimens on loan. Dr. Donald 
F, Squires of the U. S. National Museum and Dr. J. Wyatt Durham of the University of 
California Museum of Paleontology examined some of the stony corals in the present col- 
lections. Dr. John D. Soule of the Allan Hancock Foundation, University of Southern 
California, and Mr. Arnold Ross of the Department of Geology of the University of Florida 
identified the bryozoa and barnacles, respectively. Dr. G Dallas Hanna and Mr. Allyn G. 
Smith of the California Academy of Sciences, Mrs. Dina Lee Hernandez and Mr. William 
E. Old, Jr., of the American Museum of Natural History aided us in various ways. The 
senior author is indebted to Dr. George E. Lindsay for inviting him to join the expedition 
and to his companions on the trip for their assistance in the field collecting. We are grateful 


to Mr. K. K. Bechtel and the Belvedere Scientific Fund for making the expedition possible. 
The following abbreviations are used to designate institutions listed in this paper: 


AMNH, American Museum of Natural History, New York, New York; CAS, Department 
of Geology, California Academy of Sciences, San Francisco, California; SD-BE, San Diego 
Natural History Museum, Belvedere Expedition, Natural History Museum, San Diego, Cali- 
fornia; SU, Department of Geology, Stanford University, Stanford, California; UCMP, 
University of California Museum of Paleontology, Berkeley, California. 


DESCRIPTIONS OF COLLECTING LOCALITIES, FAUNAL LISTS, AND 
AGE DETERMINATIONS 


In this section, the localities are briefly described, the faunas are listed using a convenient 
system of nomenclature that should enable the non-specialist to recognize the taxa without 
reference to monographic studies, and the age of the deposits is given with as much certainty 
as our present knowledge permits. Determinations of age assignments for the Pliocene and 
Pleistocene faunas described in this report generally follow the geochronology that was recog- 
nized by Anderson (1950) and Durham (1950) for the marine strata containing metazoan 
invertebrates on the islands in the Gulf of California. Three divisions were recognized for 
the Pliocene epoch, namely the Marquer, Carmen and San Marcos formations. These were 
considered to represent strata deposited during late, middle and early periods of Pliocene time, 
respectively. Deposits of Pleistocene age were referred to either upper or lower beds. As 
Durham (op. cit.) pointed out, this is a tentative classification that will undoubtedly require 
refinement as the chronology of these faunas becomes better understood (see fig. 1). 

Regional correlations of the Cenozoic formations of the Gulf trough, northern and 
southern Baja California, and coastal California were presented by Durham and Allison 
(1960:59-64) in a review of the geology of Baja California. The following correlations 
were suggested for the Gulf trough sediments of Pliocene age: the Marquer (Anderson, 1950; 


1964 } EMERSON AND HERTLEIN: INVERTEBRATE MEGAEOSSILS 337 


Durham, 1950), Inferno (Wilson, 1948, 1955) and Borrego (Tarbet and Holman, 1944) 
formations, late Pliocene; Carmen (Anderson, 1950; Durham, 1950), Gloria (Wilson, 1948, 
1955), and Palm Spring formations, mid-Pliocene; and Imperial (Hanna, 1926), San Marcos 
(Anderson, 1950; Durham, 1950), and Boleo (Wilson, 1948, 1955) formations, early 
Pliocene. The Santa Rosalia formation (Wilson, 1948, 1955) was applied to Pleistocene sedi- 
ments overlying the Infierno formation in the vicinity of Santa Rosalia, Baja California. 


Collections from localities F-1 to F-14 are deposited in the San Diego Natural History 
Museum (SD-BE) and were made by Lindsay, Moran, Banks, Harbison and Emerson. 
Collections from localities 38539 to 38562 (CAS) were made by Wiggins and are deposited 
in the Department of Geology of the California Academy of Sciences. 


Locality F-1 (SD-BE), Isla Angel de la Guarda, ?Pliocene 


Fossils were collected from exposures in the sea cliff on the southeastern end of Isla 
Angel de la Guarda, about 2 miles south of Isla Pond, March 17, 1962, by Emerson. Sedi- 
ments consist of fine, poorly consolidated sands filling a terrace-like depression between a low 
hill and a higher volcanic flow. The fossiliferous bed is a maximum of 50 feet above the 
present beach and is locally covered with 1 to 5 feet of volcanic rock and alluvium. Oyster 
shells are common; other fossils are uncommon. 


Chione californiensis Broderip 

Chione undatella Sowerby 

Chlamys (Argopecten) species indeterminate 
Ostrea angelica Rochebrune 


Unfortunately, the age of this deposit cannot be determined on the basis of its meager 
fauna. The two worn, imperfect valves of a species of Chlamys (Argopecten) are poorly pre- 
served, and we are uncertain whether the specimens represent the Pliocene species, C. (A.) 
abietis Jordan and Hertlein. The other species are not of temporal significance. 


Although the surface truncation of the fossiliferous bed suggests a possible Pleistocene age 
for the deposit, the absence of coarse sediments and the general geologic setting suggest a 
Pliocene age. Similar deposits that are exposed in a large coastal arroyo south of this locality 
were considered by Durham (1950:21) to be correlative with the Marquer formation of 
late Pliocene age. 


Locality F-2 (SD-BE), Isla San Esteban, Pliocene 


Fossils were collected from exposures in the sea cliff on the east side of Isla San Esteban, 
opposite our anchorage, March 22, 1962, by Wiggins and Emerson. The sediments are of 
coarse, well-cemented sand, with an exposed thickness of approximately 100 feet (base not 
exposed), and the beds are overlain with 1 to 3 feet of soil; the beds are tilted to the north. 

Chlamys abietis Jordan and Hertlein 

Ostrea species 

Eucidaris thouarsit Valenciennes (spines) 

coral 

This appears to be the first record of the occurrence of fossiliferous sediments on this 
island. The presence of Chlamys (Argopecten) abietis in the fauna indicates a Pliocene age 
for this deposit. 

Locality F-2a (SD-BE), Isla Salsipuedes, ?Pliocene 

Fossils were found as float on the northwest side of Isla Salsipuedes at an elevation of 
about 100 feet, March 24, 1962, by Moran and Emerson. 

Ostrea cf. O. californica Marcou 


On the afternoon of March 23, Moran discovered, in the course of his botanical collect- 
ing, a number of large valves of this fossil oyster. The next morning Moran and the senior 
author visited the area and found several of the large fossils scattered on the surface of a 
kitchen midden that was composed largely of Recent mollusks. As the source beds for the 


338 San Dieco Society or Natura History { VoL. 13 


Fig. 2. Above. View of sea cliff, south side of El Pulpito, April 2, 1962. Locality F-4 (SD-BE), 


?Pleistocene; fossiliferous sandstone overlies volcanic rocks and is overlain by conglomerates and soil cover. 


Below. View of sea cliff at Arroyo de la Aguada, Isla San José, April 11, 1962. Locality F-11 
(SD-BE), Pliocene; fossiliferous beds are near top of exposure. Large blocks of fossiliferous sandstone have 
fallen to the beach (senior author is standing on one). Photographs courtesy of G. E. Lindsay. 


1964 } EMERSON AND HERTLEIN: INVERTEBRATE MEGAFOSSILS 339 


fossil oysters could not be located in the vicinity of the midden, we must conclude that the 
oysters had been transported by Indians or by some other non-geological agent to this site. 


It is interesting to note that Ostrea californica has been reported previously only from 
early Pliocene deposits of the Imperial formation of Imperial County, California; see the 
discussion of this species in the systematic section of this paper. 


Localities F-3 (SD-BE) and 38562 (CAS), Isla Partida, ?Pliocene 


Fossils were collected from exposures in the cliff on the southwest side of the isthmus, 
March 26, 1962, by Wiggins and Emerson. The top of the cliff is truncated and is covered 
with 3 to 4 feet of boulder conglomerate of probable Pleistocene age, with 1 to 2 feet of 
loose boulders resting on the terrace. The lower bed is composed of 65 to 70 feet of locally 
fossiliferous mudstone and fine sandstone, with pebbles, that weathers a light gray. It is 
overlain by the conglomerate and is covered at the base by a cobble beach. The terraced cliff 
is about 75 feet in total thickness. Casts and molds of Chlamys are extremely common, with 
preserved specimens uncommon. Other fossils are rare. 


In the following list, taxa marked ' are from locality F-3 (SD-BE); those marked * are 
from locality 38562 (CAS) = 38441 (CAS). 


Barbatia reeveana form lasperlensis Sheldon and Maury” 
Chlamys tumbezensis d’Orbigny’ * 

Plicatula species indeterminate! 

Balanus species! 


Although none of the faunal elements from this locality is of temporal significance, the 
geologic setting and the composition of the sediments suggest a probable Pliocene age for the 
fossiliferous sediments. This collection appears to be the first record of the occurrence of fossils 
on this island. 


Localities F-4 (SD-BE) and 38553 (CAS), El Pulpito, Baja California, ?Pleistocene 


Fossils were collected from exposures in the sea cliff, on the south side of El Pulpito, 
Baja California, April 2, 1962, by Lindsay, Wiggins, and Emerson (fig. 2). The richly 
fossiliferous bed is 4 to 18 feet thick, is composed of well-cemented sandstone weathering 
pinkish-gray, and rests on volcanic rocks. Locally the base of the exposure is covered by the 
cobble beach. The deposit is overlain by 20 to 30 feet of boulder conglomerate with coarse 
sand, pebbles and rocks, and is capped locally by a thin layer of soil and wind-blown sand. 
Fossils are well preserved; many clams have paired valves, and complete echinoid tests are 


common (see Lindsay, 1962:20, fig. 15). 


The taxa preceded by an asterisk (**) in the following list were not previously reported 
from the area of El Pulpito. Taxa marked ! are from locality F-4 (SD-BE); those marked 
? are from locality 38553 (CAS). 

Pecten subnodosus Sowerby! 

Pecten vogdesi Arnold! 

Spondylus princeps Broderip! 

Clypeaster speciosus Verrill* 

Encope californica Verrill'; 2 

Encope cf. E. grandis inezana Durham}: ? (extinct) 

Encope species indeterminate! 

Eucidaris thouarsti Valenciennes (spines) 1; ? 

*Meoma cf. M. grandis Gray? 
*bryozoal 

calcareous algae! 

The age of the fossiliferous deposits in the El Pulpito area has yet to be conclusively 
demonstrated. Grant and Hertlein (1938:47, 98) inferred a Pliocene age for the fauna from 
El Pulpito Point (W. W. Valentine, MS) with reference to the occurrence of Encope 
micropora Agassiz and Clypeaster speciosus Verrill. Durham (1950:47, 50) subsequently 
indicated that Valentine’s specimen of Encope was not referable to E. micropora, but sug- 
gested that it might be his “Encope n. sp. B”, which he recorded from the Pleistocene of 


340 San Deco Society or NaturaL History { VoL. 13 


Isla Coronados. Unfortunately, the manuscript describing the collection from El Pulpito 
made by the late W. W. Valentine in 1928 has not been published. 


At our request, Dr. A. Myra Keen provided us with a list of fossils in the Valentine 
collection from El Pulpito (SU locality no. 807). Unfortunately, none of the ten species 
appears to be of temporal significance, although some of the fossils were not identified to 
species. However, a large collection of fossils recently was recorded by Hertlein (1957:62-65; 
CAS localities 34036 and 34164) from deposits in an arroyo about one mile northwest of 
the coast near El Pulpito. None of the species listed by Hertlein is geochronologically re- 
stricted to pre-Pleistocene time, but one subspecies, Encope grandis inezana Durham, may 
be restricted to the Pleistocene, and all the other species but one, Haliotis fulgens Philippi, 
are now living in the Gulf of California. 


No species in the present collection from this locality are restricted to strata of known 
Pliocene age. All the species are extant, with the possible exception of the Encope identified 
as E. cf. E. grandis inezana Durham, and are known to occur in the Gulf of California at 
the present time. Many of the species are known to range from early Pliocene to the Present. 
The field evidence suggests a possible Pliocene age for the fossiliferous sediments. These 
sediments are well compacted, the sandstone is well sorted, and the deposit is cemented directly 
on the uncomformable surface of the underlying volcanic rocks. In most Pleistocene deposits 
in this region, the sediments are poorly compacted and sorted, and they generally overlie 
basal conglomerates which are cemented to the surface of the terrace platform. This fossiliferous 
deposit is covered by 20 to 30 feet of poorly sorted, largely uncompacted sediments that 
probably are at least in part of Pleistocene age (fig. 2). Additional field work in this area 


will have to be undertaken before a more precise age for this fauna can be determined. 


Locality F-5 (SD-BE), Isla Coronados, Pleistocene 


Fossils were collected from exposures in the sea cliff that forms a low terrace about 20 
feet above the high tide zone on the southern end of Isla Coronados, April 3, 1962, by 
Emerson. Sediments are locally very fossiliferous, and are composed of coarse, poorly sorted 
sand that is in places cemented with calcareous algae. Heads of the coral Porites are common. 
The taxa in the following list that are preceded by an asterisk (*) were not previously 
reported from this locality. 


Anadara multicostata Sowerby 


*Anomalocardia subimbricata tumens Verrill 


*A polymetis cognata clarki Durham 
Barbatia reeveana d’Orbigny 
Cardita megastropha Gray 
Cardium consors Sowerby 


Cardium biangulatum Broderip and Sowerby 


Cardium elenense Sowerby 
Chama frondosa Broderip 
*Chione californiensis Broderip 
Codakia distinguenda Tryon 
*Ctena mexicana Dall 
Diplodonta subquadrata Carpenter 
“Glycymeris gigantea Reeve 
Glycymeris multicostata Sowerby 
*Tsognomon janus Carpenter 
Lima tetrica Gould 

Lucina lampra Dall 

Lucina nuttalli Conrad 
Megapitaria squalida Sowerby 
Miltha xantusi Dall 
*Ostrea megodon Hanley 
*Ostrea palmula Carpenter 
Pecten subnodosus Sowerby 
Pecten vogdesi Arnold 

Petricola robusta Sowerby 
Pinctada mazatlanica Hanley 
* Semele cf. S. flavescens Gould (juvenile) 


Me 


*Semele verrucosa Morch 

*Spondylus calcifer Carpenter 
Spondylus princeps Broderip 
Tellina cumingii Hanley 
Teilina viridotincta Carpenter 
Acanthina muricata Broderip 
Aletes centiquadrus Valenciennes 
Bulla cf. B. punctulata A. Adams (fragment) 
Bursa californica Hinds 
Cancellaria obesa coronadoensis Durham 

(extinct ) 

*Cancellaria pulchra Sowerby 
Cantkarus anomalus Reeve 
Cerithium gemmatum Hinds 
Cerithium maculosum Kiener 

*Cerithium sculptum Sowerby 

*Conus diadema Sowerby 

*Conus gladiator Broderip 
Conus princeps Linné 
Conus purpurascens Broderip 
Conus regularis Sowerby 
Conus scalaris Valenciennes 
Conus ximenes Gray 

*“Crepidula arenata Broderip 
Crepidula onyx Sowerby 
Cypraea annettae Dall 

*Cypraea arabicula Lamarck 
Cypraecassis coarctata Sowerby 


1964] 


Diodora inaequalis Sowerby 
*“Enaeta cumingn form pedersenii Verrill 
*Fasciolaria princeps Sowerby 

Fusinus cinereus Reeve 

Gyrineum strongi Jordan (extinct) 
*Hipponix antiquatus Linné 

Knefastia funiculata Kiener 

Modulus cerodes A. Adams 

Morula lugubris C. B. Adams 
“Murex cf. M. elenensis Dall (fragment) 
*“Nassarius tiarula Kiener 
“Oliva porphyria Linné 

Olivella dama Wood 

Olivella species indeterminate (2 fragments) 

Parametaria dupontii Kiener 


EMERSON AND HERTLEIN: INVERTEBRATE MEGAFOSSILS 341 


Polinices bifasciata Gray 
*Polinices uber Valenciennes 
Strombus granulatus Swainson, including 
form cortezianus Durham 
Tegula mariana Dall 
*“Terebra cf. T. specillata Hinds 
Terebra variegata Gray 
*Trima californica Gray 
Trivia solandri Sowerby 
Turbo fluctuosus Wood 
Turbo squamiger Reeve 
*Turritella gonostoma Valenciennes 
Encope californica Verrill 
Porites californica Verrill 
crab claw 
calcareous algae 


The fauna of these extremely fossiliferous terrace deposits was enumerated by Durham 
(1950: tables 2 and 3, localities UCMP A-3548 and A-3550). The field evidence and the 


composition of the fauna suggest that these deposits are late Pleistocene in age. 


Localities F-6a, b (SD-BE) and 38539, 38540 (CAS), Puerto Ballandra, 


Isla Carmen, Pliocene 


Fossils were collected from a bioherm and sediments at the base of the bioherm that 
outcrop in the sea cliff on the southeast shore of the bay, April 4, 1962, by Wiggins and 
Emerson. The cliff is 20 to 25 feet in elevation, with its surface truncated by terraces and 
with 1 to 2 feet of soil cover. The reef is largely composed of branching heads of Porites 
californica (see Lindsay, 1962, figs. 16, 17), mollusks, echinoid spines, and coarse sand. 
Sediments of the basal bed are composed of fine sand (mudstone), with well-preserved 
pelecypods common and poorly preserved gastropods uncommon. These deposits abut against 
volcanic rocks that are exposed in an arroyo and on the adjacent beach. 


Coral reef facies {localities F-6a (SD-BE) and 38539 (CAS) ]. Fossils are locally 
common in coarse sand. The taxa preceded by an asterisk (*) in the following lists were 
not previously reported from this locality. Taxa marked ' are from locality F-6a (SD-BE); 
those marked ” are from locality 38539 (CAS). 


Anadara multicostata Sowerby 

* A nomalocardia subimbricata tumens Verrill'; 7 
Apolymetis cognata clarki Durham? 
Barbatia reeveana d’Orbigny!; 2 

* Basterotia hertleint Durham! (extinct) 
Cardita affinis californica Deshayes! 

*“Cardita megastropha Gray 
Cardium biangulatum Broderip and Sowerby? 
Cardium elenense Sowerby}, 2 
Chione californiensis Broderip!: 2 
Codakia distinguenda Tryon}; 2 

*Corbula bicarinata Sowerby! 
Ctena mexicana Dall; 2 
Divaricella eburnea Reeve! 
Glycymeris gigantea Reevel, 2 
Glycymeris multicostata Sowerby!; 2 
Isognomon janus Carpenter!, = 
Megapitaria squalida Sowerby}, 7 
Ostrea angelica Rochebrune!; 
Pinctada mazatlanica Hanley!: ? 
*Semele cf. S. bicolor C. B. Adams? 
Semele flavescens Gould! 
Solecardia eburnea Conrad! 

*Tellina meropsis Dall, 2 

*Tellina viridotincta Carpenter! 


*Astraea unguis Wood! 

Cerithium gemmatum Hinds}, 2 

Cerithium maculosum Kiener” 
*Conus princeps Linneé! 

*Crassispira cf. C. nymphia Pilsbry and 
Lowel 

Crucibulum scutellatum Wood}, 

Cypraea annettae Dall!, 2 

Diodora alta C. B. Adams!; 7 

Diodora inaequalis Sowerby}, 

Fusinus cinereus Reeve! 

Heliacus robertsae Durham! (extinct) 
*Hipponix antiquatus Linne!; 2 
*Modulus cerodes A. Adams! 

Morula ferruginosa Reevet 

Nassarius species! (juvenile) 

Oliva spicata (Bolten) Roding! 

Olivella dama Wood!; 7 
**Polinices uber Valenciennes!, = 
*Strombus galeatus Swainson}, 7 

Strombus granulatus Swainson}, 2 

Tegula mariana Dall!, 2 

Turbo fluctuosus Wood!; * 

calcareous algae!: 7 

Porites californica Verrill! 2 
*Eucidaris thouarsiti Valenciennes!, 2 


342 San Disco Society oF NaturaAL History {Vot. 13 


Sand and mudstone facies {locality 6b (SD-BE), marked ', and 38540 (CAS), marked 7}. 


Paired valves of pelecypods common. 


Apolymetis cognata clarki Durham1 Megapitaria squalida Sowerby? 
Basterotia hertleint Durham1; 2 (extinct) Ostrea angelica Rochebrune? 
Cardium biangulatum Broderip and Sowerby? Tagelus politus Carpenter!, 2 
Cardium elenense form apicinum Tellidora burneti Broderip and Sowerby! 
Carpenter!, 2 *Tellina amianta Dall? 

*Chione californiensis Broderip!, 2 *Tellina reclusa Dall 
Cyathodonta undulata Conrad}; 2 Tellina simulans C. B. Adams}; 2 

*Diplodonta sericata Reevel, 2 *Nassarius tiarula Kiener! 
Divaricella eburnea Reevel, 2 *Natica unifasciata Lamarck? 
Lucina lampra Dall}, 2 * Oliva polpasta Duclos? 

*Lucina nuttalli Conrad? *Olivella dama Wood1 


These deposits were described and the fossils listed by Durham (1950:39, tables 2, 3, 
localities UCMP A-3534, reef facies; UCMP A-3670, sand and mudstone facies). Durham 
(1950:23) considered the deposits referable to the Marquer formation of late Pliocene. 


Pleistocene faunas were reported from low terraces at the south end of the bay by 
Hertlein (1957, CAS locality 34165). 


Locality F-7 (SD-BE), Marquer Bay, Isla Carmen, Pliocene 


Fossils were collected from exposures in sea cliffs and arroyos adjacent to the beach of 
the bay on April 4, 1962, by Emerson. The sea cliffs are about 40 feet in elevation, with the 
surface truncated. The sandstone weathers a light buff and contains well-preserved oysters, 
pectens and echinoids; other fossils are poorly preserved. Volcanic pebbles and calcareous 
sand are common. 

Anomalocardia subimbricata tumens Verrill 

Chione cf. C. californiensis Broderip 

Chlamys abietis Jordan and Hertlein 

Ostrea angelica Rochebrune 

Pecten marquerensis Durham 

Pecten vogdesi Arnold 

Tagelus species indeterminate (mold) 

Turritella species indeterminate (mold) 

Encope species indeterminate (mold) 

Balanus trigonus Darwin 

The field evidence and the composition of the fauna suggest that these beds are near the 
base of the Marquer formation. 


Locality 38544 (CAS) [= 38544a}, Marquer Bay, Isla Carmen, Pliocene 


Fossils were collected by Wiggins from exposures in arroyos about 14 mile from Marquer 
Bay, April 5, 1962. 

Anadara multicostata Sowerby 

Barbatia cf. B. reeveana d’Orbigny 

Chlamys abietis Jordan and Hertlein 

Ostrea angelica Rochebrune 

Pecten vogdesi Arnold 

The field evidence and the composition of the fauna suggest that this collection was 
made from beds near the base of the Marquer formation. 


Locality 38545 (CAS), Marquer Bay, Isla Carmen, Pliocene 
Fossils were collected from exposures in arroyos about 1/, mile inland from Marquer Bay, 
April 5, 1962, by Wiggins. 
Chlamys abietis Jordan and Hertlein 


Field evidence suggests that this collection was made from beds near the middle of the 
Marquer formation. 


1964 } EMERSON AND HERTLEIN: INVERTEBRATE MEGAFOSSILS 343 


Locality 38546 (CAS) {= 38546a}, Marquer Bay, Isla Carmen, Pliocene 


Fossils were collected from exposures near the top of a mesa, about 14 mile inland from 


Marquer Bay, April 5, 1962, by Wiggins. 


Antigona isocardia Verrill Cypraea annettae Dall 

Barbatia bramkampi Durham Conus dispar Sowerby 

Cardium biangulatum Broderip and Sowerby Conus gladiator Broderip 
Chlamys circularis Sowerby Oliva spicata (Bolten) Roding 
Codakia distinguenda Tryon Polinices cf. P. uber Valenciennes 
Megapitaria squalida Sowerby Strombus galeatus Swainson 
Ostrea angelica Rochebrune Strombus gracilior Sowerby 
Pecten vogdesi Arnold Strombus granulatus Swainson 
Spondylus princeps Broderip Turbo species (operculum) 
Cerithium uncinatum Gmelin coral 


The field evidence and the composition of the fauna suggest that this collection was 
from near the top of the Marquer formation. 


Locality F-8 (SD-BE) Isla Monserrate, Pliocene 


Fossils were collected from exposures in terraced, light-colored limestone bluffs along 
the beach near the southwest corner of the island, April 8, 1962, by Emerson. The fossils 
are poorly preserved, mostly as casts and molds. 

Chlamys cf. C. abietis Jordan and Hertlein 

Pecten cf. P. stearnsu Dall 

The two species in the present collection indicate a Pliocene age for the deposit. Durham 
(1950:26) referred similar deposits on the southeastern side of the island to the Carmen 
formation, which he believed to be mid-Pliocene in age. 

Wiggins also collected on April 8, 1962, specimens of a pecten that has been identified 
as Chlamys (Argopecten) species indeterminate from deposits of probable Pliocene age on 


the southwest side of Isla Monserrate (CAS locality 38550). 
Localities F-9 (SD-BE) and 38551 (CAS), Isla Monserrate, Pliocene 


Fossils were collected from exposures in limestone bluffs that form a small headland 
north of F-8 (SD-BE), on April 8, 1962, by Wiggins and Emerson. Pectens and oysters 
are well preserved. 

Taxa marked ! are from locality F-9 (SD-BE); those marked * are from locality 
38991 (CAS): 

Chlamys abietis Jordan and Hertlein’: * 

Ostrea angelica Rochebrune! 

Pecten subnodosus Sowerby’ 

Clypeaster species indeterminate” (fragments) 


These deposits are correlative with the beds comprising Locality F-8 (SD-BE). 
Locality F-9a (SD-BE), Isla Monserrate, ?Pliocene 


Fossils were collected from terraces at an elevation of about 600 feet, April 8, 1962, by 
Moran. 


Chlamys aff. C. abietis Jordan and Hertlein 
The field evidence suggests that the beds at this elevation are Pliocene. 


Localities F-10 (SD-BE) and 38554 (CAS), Isla Monserrate, Pleistocene 
Fossils were collected from exposures in terrace deposits along the shore on the west side 
of the island, south of F-9 (SD-BE), April 8, 1962, by Wiggins and Emerson. One to 18 
feet of fossiliferous, fine grained sand rests on the terrace platform, which was cut into 
volcanic rocks a few feet above the present beach. 


344 


SAN Dreco Society OF NATURAL HISTORY 


i Vor. 13 


Taxa marked ! are from locality F-10 (SD-BE); those marked * are from locality 
38554 (CAS). 


Anadara multicostata Sowerby!; 2 

Apolymetis cognata clarki Durham}, 2 

Barbatia reeveana d’Orbigny? 

Cardita megastropha Gray! 

Cardium biangulatum Broderip and 
Sowerby!, 2 

Cardium consors Sowerby!; 2 

Cardium elenense Sowerby!; 2 

Chione californiensis Broderip!, 2 

Codakia distinguenda Tryon 

Divaricella eburnea Reeve}, 2 

Dosinia ponderosa Gray; 

Glycymeris gigantea Reeve 

Glycymeris multicostata Sowerby! 

Lucina nuttalli Conrad! 

Lucina lampra Dall; 2 

Megapitaria squalida Sowerby}: 2 

Pecten subnodosus Sowerby!, 2 

Pecten vogdesi Arnold}; 2 

Periglypta multicostata Sowerby! 

Pinctada mazatlanica Hanley! 

Polymesoda species” 

Pseudochama saavedrai Hertlein and Strong? 

Solecardia eburnea Conrad? 

Tagelus californianus Conrad! 

Tellina cumingii Hanley! 

Tellina reclusa Dall! 

Tellina simulans C. B. Adams! 

Astraea unguis Wood? 

Calliostoma bonita Strong, Hanna, and 
Hertlein1 

Calliostoma eximium Reeve? 

Cancellaria wigginsi, new species? (extinct) 

Cantharus pallidus Broderip and Sowerby? 


Cerithium gemmatum Hinds}, 2 


Cerithium maculosum Kienert 

Crucibulum scutellatum Wood! 

Conus brunneus Wood 

Conus dispar Sowerby” 

Conus nux Broderip! 

Conus patricius Hinds! 

Conus perplexus Sowerby! 

Conus princeps Linné! 

Conus purpurascens Broderip!: = 

Conus regularis Sowerby! 

Cypraea annettae Dall! 

Gyrineum strongi Jordan! (extinct) 

Knefastia funiculata Kiener!, * 

Nassarius tiarula Kiener!, 7 

Nerita scabricosta Lamarck!; 2 

Oliva incrassata Solander!, * 

Oliva spicata (Bolten) Roding! 2 

Olivella dama Wood!, ? 

Polinices bifasciata Gray! ; 

Polinices uber Valenciennes!; * 

Strombina maculosa Sowerby!; 7 

Strombus galeatus Swainson!; 

Strombus gracilior Sowerby! 

Strombus granulatus Swainson!, = 

Terebra strigata Sowerby! 

Terebra specillata Hinds! 

Terebra variegata Gray}, * 

Turbo fluctuosus Wood!, 2 

Dentalium semipolitum Broderip and 
Sowerby! 

Pocillopora robusta Verrill* 

Porites species! 

Encope californica Verrill*, 2 

Encope grandis Agassiz! = 

Eucidaris thouarsii Valenciennes! 

calcareous algae! 


» 


The presence of late Pleistocene fossils in deposits on a 20-foot terrace at the south end 
of the island was reported by Durham (1950:26, 37, 138), but the fossils, which were said 
to include only species that are still living, were not listed by him. The present collection was 
made from deposits that are apparently correlative with those mentioned by Durham (UCMP 
locality A-3567). The fauna is composed of species that are now living in the Gulf of Cali- 
fornia, with the exception of Gyrineum strong: Jordan and Cancellaria wigginsi, new species, 
both of which are apparently extinct, and Conus patricius Hinds. 


Locality 38555 (CAS), Isla Monserrate, Pleistocene 


Fossils exposed in deposits on the northwest end of the island were collected on April 
8, 1962, by Wiggins. 


Basterotia peninsularis Jordan 


Cardium biangulatum Broderip and Sowerby 


Cardium elenense Sowerby 
Divaricella eburnea Reeve 
Lucina lampra Dall 

Pitar newcombianus Gabb 
Tagelus subteres Conrad 
Olivella dama Wood 
Polinices uber Valenciennes 
Terebra variegata Gray 


These deposits may be correlative with those occurring at localities F-10 (SD-BE) and 
38554 (CAS). All the species in this collection are modern constituents of the fauna of the 
Gulf of California except Basterotia peninsularis Jordan, which is reported to range from 
Port Guatulco, Oaxaca, Mexico, south to the Galapagos Islands. 


1964 } EMERSON AND HERTLEIN: INVERTEBRATE MEGAFOSSIIS 345 


Localities F-11 (SD-BE) and 38549 (CAS), Isla San José, Pliocene 

Fossils were collected from exposures on the northeast side of the island along the shore 
north and south of Arroyo de la Aguada, April 11, 1962, by Wiggins and Emerson. The 
sea cliffs locally are from 100 to 150 feet in elevation. Their basal part is composed of 
apparently non-fossiliferous sandstone weathering a buff color, and is overlain by 0 to 25 feet 
of lighter colored sandstone that is extremely fossiliferous. In the upper beds, pectens, oysters 
and echinoids are common, but gastropods are preserved only as casts or molds (see fig. 2). 

In this area, fossiliferous deposits occur inland at least two miles, as fossils were noted 
at elevations of more than 500 feet (CAS locality 38548) in beds with an estimated maximum 
thickness of 40 feet that rest on unfossiliferous sandstones and granite. Although the field 
evidence suggests that the Pliocene beds have been subsequently truncated by terraces along 
the adjacent coast and in the arroyo, no Pleistocene fossils were found in the terrace deposits. 

Taxa marked ! are from locality F-11 (SD-BE) ; those marked ” are from 38549 (CAS). 

Ostrea angelica Rochebrune! 

Ostrea megodon Hanley” 

Chlamys abietis Jordan and Hertlein!, 2 

Pecten cf. P. carrizoensis Arnold 

Pecten marquerensis Durham1; 2 

Pecten subnodosus Sowerby! 

Spondylus cf. S. princeps Broderip! 

Conus species indeterminate 

Dendraster casseli Grant and Hertlein4 

Dendraster cf. D. granti Durham! 

Balanus tintinnabulum californicus Pilsbry (on pectens)1 

Although Hanna and Hertlein (1927) reported the presence of Pliocene fossils on this 
island, the fauna remains poorly known. On the basis of the seven species recorded by Hanna 
and Hertlein (1927:149) and the echinoid cited by Grant and Hertlein (1938:82), a 
mid-Pliocene age is suggested for at least part of the extensive fossil deposits on the island. 

The composition of the present collection, however, suggests that beds representing 
lower, middle, and upper Pliocene deposits may occur on this island. Valves of a large species 
of Pecten (sensu stricto), some 90 mm. high, are not uncommon in these deposits. Un- 
fortunately, only left valves were encountered, and these do not permit a definite identification. 
These specimens may represent P. (P.) carrizoensis Arnold, a species believed to be re- 
stricted to the early Pliocene (Durham, 1950:19, 61). The echinoids indicate a mid-Pliocene 
age (see systematic section of this paper), whereas the presence of Pecten (Patinopecten) 
marquerensis Durham indicates a late Pliocene age (Durham, op. cit.:19, 67). Detailed 
field studies will have to be undertaken before a more precise age can be determined for the 
strata at this locality. 


Localities F-1la (SD-BE) and 38548 (CAS), Isla San José, Pliocene 


Fossils were collected from exposures in Arroyo de la Aguada on the northeast side of 
the island at an elevation of about 530 feet, April 11, 1962, by Banks and Wiggins. 

Taxa marked ! are from locality F-1la (SD-BE); those marked * are from locality 38548 
(GAS). 

Ostrea angelica Rochebrune? 

Ostrea megodon Hanely? 

Chlamys abietis Jordan and Hertlein!. * 

Chlamys revellei Durham? ‘ 

Pecten (Patinopecten) cf. P. (P.) marquerensis Durham” 

Pecten subnodosus Sowerby 

Balanus species! 

The presence of Chlamys (Argopecten) revellei Durham suggests a mid-Pliocene age 
for these deposits. This species was previously known only from the Carmen formation at 
Salinas Bay, Isla Carmen, and from middle Pliocene deposits on Isla Monserrate (Durham, 
1950:19, 64). There is, however, a Patinopecten in this collection that is close to Pecten 
(Patinopecten) marquerensis Durham, a species supposedly restricted to the late Pliocene 


(Durham, 1950:19, 67). 


346 San Dieco Society oF NATuRAL History [ VoL. 13 


Locality 38547 (CAS), Isla San José, ?Pliocene 


Fossils were collected from exposures inland about 14 mile along the south side of the 
main arroyo leading eastward from Bahia de Ostiones, on the west side of the island, at an 


altitude of about 100 feet, on April 12, 1962, by Wiggins. 


Anadara cf. A. multicostata Sowerby 

Ostrea angelica Rochebrune 

Pecten species 

Placunanomia cumingi Broderip 

Although this meager fauna is not geochronologically diagnostic, it should be noted that 
fossiliferous sediments of Pliocene age were previously known from the west side of the island 


(Hanna and Hertlein, 1927, CAS locality 839). 


Extraordinarily large shell middens of apparent Recent age were reported by Emerson 
(1960b) from near Amortajada Bay, on the southwestern end of the island. 


Locality F-12 (SD-BE), Isla Cerralvo, Pleistocene 


Fossils were collected from deposits on a narrow terrace platform that is cut into granitic 
rocks, at an elevation of 15 to 20 feet, on the west side of the island, April 12, 1962, by 
Emerson. The terrace deposits are composed of 1 to 4 feet of pebble to boulder conglomerates 
that are locally fossiliferous. Fossils other than coral fragments and occasional coral heads 
are uncommon. 

Anadara multicostata Sowerby 

Pinctada mazatlanica Hanley 

Conus purpurascens Broderip 

Cerithium maculosum Kiener 

Muricanthus princeps Broderip 

Strombus galeatus Swainson 

Pocillopora robusta Verrill 


These terrace deposits are apparently correlative with similar deposits reported by Emer- 
son (1960a) on the south end of the island. All the species in the present collection are 
living at the present time in the Gulf of California. A late Pleistocene age is indicated for 
this and other low, emergent, terrace deposits in the Gulf of California region. 


Localities F-13 (SD-BE) and 38541, 38542, and 38543 (CAS), Isla Cerralvo, Pliocene 


Fossils were collected from exposures along the beach and in arroyos on the west side of 
the island near El Mostrador, April 16, 1962, by Wiggins and Emerson. The fossiliferous 
sandstones of the basal beds along the coast weather to a light buff color, carry fragments of 
calcareous algae, echinoid spines and mollusks, are 10 to 20 feet in thickness, are tilted, 
warped and faulted, and locally rest on conglomerates and mudstones. The overlying sand- 
stones and conglomerates are locally extremely fossiliferous, with pectens and echinoids com- 
mon. Other fossils, especially the gastropods, are badly leached. These beds are at least 60 
feet thick along the coast and exposures of 100 to 150 feet in thickness were noted in the 
arroyos. The upper fossiliferous sandstones weather dark buff to gray and are well compacted 
and cemented. 


The taxa in the following list that are preceded by an asterisk (*) were not previously 
known from this locality. The mark 1 indicates taxa from locality F-13 (SD-BE), ” indicates 
38541 (CAS), * indicates 38542 (CAS), and #4 indicates locality 38543 (CAS). 


*Ostrea aff. O. heermanni Conrad}, 2 Balanus tintinnabulum californicus Pilsbry1 
Chlamys abietis Jordan and (on pectens) 

Hertlein!, 3, 4 Balanus trigonus Darwin! (on pectens) 
Chlamys revellei Durham4 *colonial stony coral of undetermined generic 
Pecten subnodosus Sowerby}; 4 afhnity! (internal molds) 

Spondylus cf. S. princeps Broderip1, 4 *Conopeum commensale Kirkpatrick and 
*Clypeaster bowersi Weaver!, 4 Metzelaar1 


*Clypeaster cf. C. speciosus Verrill4 *calcareous algae! 


1964 } EMERSON AND HERTLEIN: INVERTEBRATE MEGAFOSSILS 347 


The occurrence of Pliocene faunas in deposits near El Mostrador was previously noted 
by Hanna and Hertlein (1927) and Hertlein (1957) at CAS localities 840 and 34033 
respectively. In addition to the field evidence, the Pliocene age assignment of these deposits 
was based largely on the presence in one of the collections (Hertlein, 1957) of Chlamys 
(Argopecten) abietis Jordan and Hertlein, a Pliocene species that is not known to occur in 
beds younger than late Pliocene (Durham, 1950). Only seven other species of megafossils 
were previously reported from this area. Of these, all are species comprising the modern 
Panamic fauna, and all but one also occur in the Pleistocene fauna of the lowest emergent 
terrace of Isla Cerralvo (Emerson, 1960a). 

Although the present collection is also small in number of species, the presence of the 
echinoid, Clypeaster bowersi Weaver, suggests an early- or mid-Pliocene age for these deposits. 
This echinoid is known to occur in strata of early Pliocene age of the Imperial and the San 
Marcos formations as well as in beds of apparently mid-Pliocene age at Santa Antonita Point, 
Baja California (Durham, 1950:41). Ostrea heermanni Conrad also is known from lower 
and middle Pliocene deposits (see systematic section of this report). Chlamys (Argopecten) 
revellei Durham, however, was previously known only from beds of mid-Pliocene age of the 
Carmen formation at Salinas Bay, Isla Carmen, and beds of probable mid-Pliocene age at 
Isla Monserrate (Durham, 1950:64), and its presence in the collection suggests a mid-Pliocene 
age for these deposits. 

Dr. D. F. Squires considered the internal molds of the colonial coral in the present col- 
lection to represent the genus Solenastrea, species indeterminate, but Dr. J. Wyatt Durham 
questioned that assignment. The occurrence of a hermatypic coral of the genus Solenastrea 
in these deposits would be paleogeographically significant. At the present time, Solenastrea 
fairbanksi Vaughan is known in the west Americas only from the early Pliocene beds of the 
Imperial formation of California and from beds believed to be mid-Pliocene in age on Maria 
Madre Island of the Tres Marias Islands, off the west Mexican state of Nayarit (Durham, 
1950). Representatives of the genus Solenastrea are known elsewhere from the Oligocene and 
Miocene of Europe and Asia Minor and from the Miocene to the present time in the West 
Indies and Florida (Hertlein and Emerson, 1959; Squires, 1959). It should be noted that 
Durham and Barnard (1952) described a Recent species of Solenastrea, S. ecuadoriana, on 
the basis of a single beach-worn specimen thought to have been collected on the shore at La 
Plata Island, Ecuador. Other species of coral having West Indian affinities may be living at 
present in the southern part of the Panamic faunal province. As pointed out by Squires 
(1959), the coral fauna of that zoogeographically critical area is still poorly known. 


Locality F-13a (SD-BE), Isla Cerralvo, Pliocene 
Fossils were collected from float found along the north side of the arroyo near the site 
of the old Ruffo Ranch on the west side of the island, April 16, 1962, by Emerson. Fossili- 
ferous sediments were noted for a distance of about a quarter of a mile inland from the 
mouth of the arroyo. The folded and faulted Pliocene deposits here appear to be several hun- 
dred feet thick, and the fossils are embedded in an extremely hard matrix. See remarks for the 
previous locality. 


Chlamys abietis Jordan and Hertlein 
Pecten vogdesi Arnold 


Locality F-14 (SD-BE), Isla San Diego, Pleistocene 

Fossils were collected from isolated deposits on the platform of a small terrace on the 
southwestern end of this granitic island, on April 19, 1962, by Soulé. The truncated face of 
the terrace forms a cliff, 12 to 22 feet in elevation, above a cobble beach. The fossils are 
exposed in coarse sands and conglomerates that rest on the platform about 6 feet above sea 
level, and the fossiliferous sediments are overlain by 1 to 12 feet of unfossiliferous talus that 
is composed largely of granitic boulders. Heads of the coral, Porites, some of which are six 
inches in diameter, were found attached to boulders in the basal conglomerates, which are 
locally preserved in the sea cliff. 


348 SAN Dteco Society oF NATURAL History 


Cardita megastropha Gray 
Chama buddiana Adams 
Chione californiensis Broderip 
Chione picta Willett 

Chione undatella Sowerby 
Codakia distinguenda Tryon 


Glycymeris multicostata Sowerby 


Pinctada mazatlanica Hanley 
Spondylus princeps Broderip 
Astraea unguis Wood 
Cerithium maculosum Kiener 
Conus perplexus Sowerby 
Cypraea annettae Dall 


Cypraea albuginosa Gray 

Diodora inaequalis Sowerby 

Gyrineum strongi Jordan (extinct) 

Oliva spicata (Bolten) Roding 

Parametaria duponti Kiener 

Strombus granulatus Swainson 

Turbo fluctuosus Wood 

Eckinometra vanbrunti Agassiz 

Pocillopora robusta Verrill 

Porites californica Verrill (heads and 
fragments ) 

calcareous algae (large clumps and 
fragments ) 


{Vor 13 


Squires (1959:424) mentioned this locality in a report on the stony corals collected by 
the “Puritan”-American Museum of Natural History Expedition to western Mexico in 1957. 
The fossils, other than Porites californica, were not listed. 

This terrace is typical of the low emergent terraces throughout the Gulf of California 
which represent an apparent higher sea level of 12 to 30 feet. Such terraces are known from 
near the head of the Gulf (Hertlein and Emerson, 1956) to the Cape region of the peninsula 
(Hertlein, 1957). A late Pleistocene age is suggested for the faunas from these terrace 
deposits based upon the physiographic and faunal evidence (Emerson, 1960a:4). 


List OF SPECIES BY TAXONOMIC GROUPS, WITH AGE AND COLLECTING LOCALITIES 
The available data do not permit definite age assignments for the faunas from the fol- 


lowing localites: SD-BE F-1, F-2a, F-3, F-4, and F-9a; CAS 38547, 38550, 38553, and 38562. 
See the locality descriptions in the previous section of this paper. Extinct Pleistocene species 
are noted. 
Mo .iusca 
PELECYPODA 

Anadara multicostata Sowerby, Pliocene: Isla Carmen (CAS 38539 and 38544). Pleistocene: Isla 
Coronados (SD-BE F-5); Isla Monserrate (SD-BE F-10, CAS 38554); Isla Cerralvo (SD-BE F-12). 

Anadara cf. A. multicostata Sowerby, Pliocene: Isla San José (CAS 38547). 

Anomalocardia subimbricata tumens Verrill, Pliocene: Isla Carmen (SD-BE F-6a and F-7, CAS 38539). 
Pleistocene: Isla Coronados (SD-BE F-5). 

Antigona isocardia Verrill, Pliocene: Isla Carmen (CAS 38546). 

Apolymetis cognata clarki Durham, Pliocene: Isla Carmen (SD-BE F-6b, CAS 38539). Pleistocene: 
Isla Coronados (SD-BE F-5); Isla Monserrate (SD-BE F-10, CAS 38554). 

Barbatia bramkampi Durham, Pliocene: Isla Carmen (CAS 38546). 

Barbatia reeveana d’Orbigny, Pliocene: Isla Carmen (SD-BE F-6a, CAS 38539). Pleistocene: Isla 
Coronados (SD-BE F-5); Isla Monserrate (CAS 38554). 

Barbatia cf. B. reeveana d’Orbigny, Pliocene: Isla Carmen (CAS 38544). 

Barbatia reeveana form lasperlensis Sheldon and Maury, Pliocene: Isla Partida (CAS 38562). 

Basterotia hertleini Durham (extinct), Pliocene: Isla Carmen (SD-BE F-6a and F-6b, CAS 38540). 

Basterotia peninsularis Jordan, Pleistocene: Isla Monserrate (CAS 38555). 

Cardita affinis californica Deshayes, Pliocene: Isla Carmen (SD-BE F-6a). 

Cardita megastropha Gray, Pliocene: Isla Carmen (SD-BE F-6a). Pleistocene: Isla Coronados (SD-BE 
F-5); Isla Monserrate (SD-BE F-10); Isla San Diego (SD-BE F-14). 

Cardium biangulatum Broderip and Sowerby, Pliocene: Isla Carmen (CAS 38539, 38540, and 38546). 
Pleistocene: Isla Coronados (SD-BE F-5); Isla Monserrate (SD-BE F-10, CAS 38554 and 38555). 

Cardium consors Sowerby, Pleistocene: Isla Coronados (SD-BE F-5); Isla Monserrate (SD-BE F-10, 
CAS 38554). 

Cardium elenense Sowerby, Pliocene: Isla Carmen, (SD-BE F-6a, CAS 38539). Pleistocene: Isla 
Coronados (SD-BE F-5); Isla Monserrate (SD-BE F-10, CAS 38554 and 38555). 

Cardium elenense form apicinum Carpenter, Pliocene: Isla Carmen (SD-BE F-6b, CAS 38540). 

Chama buddiana Adams, Pleistocene: Isla San Diego (SD-BE F-14). 

Chama frondosa Broderip, Pleistocene: Isla Coronados (SD-BE F-5). 

Chione californiensis Broderip, Pliocene: Isla Angel de la Guarda (SD-BE F-1); Isla Carmen (SD-BE 
F-6a and F-6b, CAS 38539 and 38540). Pleistocene: Isla Coronados (SD-BE F-5); Isla Monserrate 
(SD-BE F-10, CAS 38554); Isla San Diego (SD-BE F-14). 

Chione cf. C. californiensis Broderip, Pliocene: Isla Carmen (SD-BE F-7). 

Chione picta Willett, Pleistocene: Isla San Diego (SD-BE F-14). 

Chione undatella Sowerby, Pliocene: Isla Angel de la Guarda (SD-BE F-1). Pleistocene: Isla San 
Diego (SD-BE F-14). 


1964] EMERSON AND HErTLEIN: INVERTEBRATE MEGAFOSSILS 349 


Chlamys (Argopecten) species indeterminate, Pliocene: Isla Angel de la Guarda (SD BE F-1): Isla 
Monserrate (CAS 38550). 

Chlamys abietis Jordan and Hertlein, Pliocene: Isla San Esteban (SD-BE F-2): Isla Carmen (SD-BE 
F-7, CAS 38544 and 38545); Isla Monserrate (SD-BE F-9, CAS 38551); Isla San José (SD-BE F-1] 
and F-lla, CAS 38548 and 38549); Isla Cerralvo (SD-BE F-13 and F-13a, CAS 38542 and 38543). 

Chlamys cf. C. abietis Jordan and Hertlein, Pliocene: Isla Monserrate (SD-BE F-8). 

Chlamys aff. C. abietis Jordan and Hertlein, Pliocene: Isla Monserrate (SD-BE F-9a). 

Chalmys circularis Sowerby, Pliocene: Isla Carmen (CAS 38546). 

Chlamys revellei: Durham, Pliocene: Isla San José (CAS 38548); Isla Cerralvo (CAS 38543). 

Chlamys tumbezensis d’Orbigny, Pliocene: Isla Partida (SD-BE F-3, CAS 38562). 

Codakia distinguenda Tryon, Pliocene: Isla Carmen (SD-BE F-6a, CAS 38539 and 38546). Pleisto- 
cene: Isla Coronados (SD-BE F-5); Isla Monserrate (SD-BE F-10); Isla San Diego (SD-BE F-14). 

Corbula bicarinata Sowerby, Pliocene: Isla Carmen (SD-BE F-6a). ‘ 

Ctena mexicana Dall, Pliocene: Isla Carmen (SD-BE F-6a, CAS 38539). Pleistocene: Isla Coronados 
(SD-BE F-5). 

Cyathodonta undulata Conrad, Pliocene: Isla Carmen (SD-BE F-6b, CAS 38540). 

Diplodonta sericata Reeve, Pliocene: Isla Carmen (SD-BE F-6b, CAS 38540). 

Diplodonta subquadrata Carpenter, Pleistocene: Isla Coronados (SD-BE F-5). 

Divaricella eburnea Reeve, Pliocene: Isla Carmen (SD-BE F-6a and F-6b, CAS 38540). Pleistocene: 
Isla Monserrate (SD-BE F-10, CAS 38554 and 38555). 

Dosinia ponderosa Gray, Pleistocene: Isla Monserrate (SD-BE F-10, CAS 38554). 

Glycymeris gigantea Reeve, Pliocene: Isla Carmen (SD-BE F-6a, CAS 38539). Pleistocene: Isla 
Coronados (SD-BE F-5); Isla Monserrate (CAS 38554). 

Glycymeris multicostata Sowerby, Pliocene: Isla Carmen (SD-BE F-6a, CAS 38539). Pleistocene: 
Isla Coronados (SD-BE F-5); Isla Monserrate (SD-BE F-10); Isla San Diego (SD-BE F-14). 

Isognomon janus Carpenter, Pliocene: Isla Carmen (SD-BE F-6a, CAS 38539). Pleistocene: Isla 
Coronados (SD-BE F-5). 

Lima tetrica Gould, Pleistocene: Isla Coronados (SD-BE F-5). 

Lucina lampra Dall, Pliocene: Isla Carmen (SD-BE F-6b, CAS 38540). Pleistocene: Isla Coronados 
(SD-BE F-5); Isla Monserrate (SD-BE F-10, CAS 38554 and 38555). 

Lucina nuttalli Conrad, Pliocene: Isla Carmen (CAS 38540). Pleistocene: Isla Coronados (SD-BE 
F-5); Isla Monserrate (SD-BE F-10). 

Megapitaria squalida Sowerby, Pliocene: Isla Carmen (SD-BE F-6a, CAS 38539, 38540, and 
38546). Pleistocene: Isla Coronados (SD-BE F-5); Isla Monserrate (SD-BE F-10, CAS 38554). 

Miltha xantusi Dall, Pleistocene: Isla Coronados (SD-BE F-5). 

Ostrea angelica Rochebrune, Pliocene: Isla Angel de la Guarda (SD-BE F-1); Isla Carmen (SD-BE 
F-6a and F-7, CAS 38539, 38540, 38544, and 38546); Isla Monserrate (SD-BE F-9); Isla San José 
(SD-BE F-11, CAS 38548); Isla San José (CAS 38547). 

Ostrea cf. O. californica Marcou, Pliocene: Isla Salsipuedes (SD-BE F-2a). 

Ostrea aff. O. heermanni Conrad, Pliocene: Isla Cerralvo (SD-BE F-13, CAS 38541). 

Ostrea megodon Hanley, Pliocene: Isla San José (CAS 38548 and 38549). Pleistocene: Isla Coro- 
nados (SD-BE F-5). 

Ostrea palmula Carpenter, Pleistocene: Isla Coronados (SD-BE F-5). 

Ostrea sp., Pliocene: Isla San Esteban (SD-BE F-2). 

Pecten cf. P. carrizoensis Arnold, Pliocene: Isla San José (SD-BE F-11). 

Pecten marquerensis Durham, Pliocene: Isla Carmen (SD-BE F-7); Isla San José (SD-BE F-11, 
CAS 38549). 

Pecten (Patinopecten) cf. P. (P.) marquerensis Durham, Pliocene: Isla San José (CAS 38548). 

Pecten cf. P. stearnsii Dall, Pliocene: Isla Monserrate (SD-BE F-8). 

Pecten subnodosus Sowerby, Pliocene: Isla Monserrate (SD-BE F-9); Isla San Jose (SD-BE F-11, 
CAS 38548); Isla Cerralvo (SD-BE F-13, CAS 38543). Pleistocene: El Pulpito (SD-BE F-4); Isla Coro- 
nados (SD-BE F-5); Isla Monserrate (SD-BE F-10, CAS 38554). 

Pecten vogdesi Arnold, Pliocene: Isla Carmen (SD-BE F-7, CAS 38544 and 38546); Isla Cerralvo 
(SD-BE F-13a). Pleistocene: El Pulpito (SD BE F-4); Isla Coronados (SD-BE F-5); Isla Monserrate 
(SD-BE F-10, CAS 38554). 

Pecten sp., Pliocene: Isla San José (CAS 38547). 

Periglypta multicostata Sowerby, Pleistocene: Isla Monserrate (SD-BE F-10). 

Petricola robusta Sowerby, Pleistocene: Isla Coronados (SD-BE F-5). 

Pinctada mazatlanica Hanley, Pliocene: Isla Carmen (SD-BE F-6a, CAS 38539). Pleistocene: Isla 
Coronados (SD-BE F-5); Isla Monserrate (SD-BE F-10); Isla Cerralvo (SD-BE F-12); Isla San Diego 
(SD-BE F-14). 

Pitar newcombianus Gabb, Pleistocene: Isla Monserrate (CAS 38555). 

Placunanomia cumingii Broderip, Pliocene: Isla San José (CAS 38547). 

Plicatula sp., Pliocene: Isla Partida (SD-BE F-3). 

Polymesoda sp., Pleistocene: Isla Monserrate (CAS 38554). 

Pseudochama saavedrai Hertlein and Strong, Pleistocene: Isla Monserrate (CAS 38554). 


350 San Dieco Society oF NaAturAL History {VoL. 13 


Semele cf. S. bicolor C. B. Adams, Pliocene: Isla Carmen (CAS 38539). 

Semele flavescens Gould, Pliocene: Isla Carmen (SD-BE F-6a). 

Semele cf. S. flavescens Gould (juvenile), Pleistocene: Isla Coronados (SD-BE F-5). 

Semele verrucosa Morch, Pleistocene: Isla Coronados (SD-BE F-5). 

Solecardia eburnea Conrad, Pliocene: Isla Carmen (SD-BE F-6a). Pleistocene: Isla Monserrate 
(CAS 38554). 

Spondylus calcifer Carpenter, Pleistocene: Isla Coronados (SD-BE F-5). 

Spondylus princeps Broderip, Pliocene: Isla Carmen (CAS 38546). Pleistocene: El Pulpito (SD-BE 
F-4); Isla Coronados (SD-BE F-5); Isla San Diego (SD-BE F-14). 

Spondylus cf. S. princeps Broderip, Pliocene: Isla San José (SD-BE F-11); Isla Cerralvo (SD-BE 
F-13, CAS 38543). 

Tagelus californianus Conrad, Pleistocene: Isla Monserrate (SD-BE F-10). 

Tagelus politus Carpenter, Pliocene: Isla Carmen (SD-BE F-6b, CAS 38540). 

Tagelus subteres Conrad, Pleistocene: Isla Monserrate (CAS 38555). 

Tagelus sp. (mold), Pliocene: Isla Carmen (SD-BE F-7). 

Tellidora burneti Broderip and Sowerby, Pliocene: Isla Carmen (SD-BE F-6b). 

Tellina amianta Dall, Pliocene: Isla Carmen (CAS 38540). 

Tellina cumingit Hanley, Pleistocene: Isla Coronados (SD-BE F-5); Isla Monserrate (SD-BE F-10). 

Tellina meropsis Dall, Pliocene: Isla Carmen (SD-BE F-6a, CAS 38539). 

Tellina reclusa Dall, Pliocene: Isla Carmen (SD-BE F-6b). Pleistocene: Isla Monserrate (SD-BE 
F-10). 

Tellina simulans C. B. Adams, Pliocene: Isla Carmen (SD-BE F-6b, CAS 38540). Pleistocene: Isla 
Monserrate (SD-BE F-10). 

Tellina viridotincta Carpenter, Pliocene: Isla Carmen (SD-BE F-6a). Pleistocene: Isla Coronados 
(SD-BE F-5). 


GASTROPODA 


Acanthina muricata Broderip, Pleistocene: Isla Coronados (SD-BE F-5). 

Aletes centiquadrus Valenciennes, Pleistocene: Isla Coronados (SD-BE F-5). 

Astraea unguis Wood, Pliocene: Isla Carmen (SD-BE F-6a). Pleistocene: Isla Monserrate (CAS 
38554); Isla San Diego (SD-BE F-14). 

Bulla cf. B. punctulata A. Adams (fragment), Pleistocene: Isla Coronados (SD-BE F-5). 

Bursa californica Hinds, Pleistocene: Isla Coronados (SD-BE F-5). 

Calliostoma bonita Strong, Hanna and Hertlein, Pleistocene: Isla Monserrate (SD-BE F-10). 

Calliostoma eximium Reeve, Pleistocene: Isla Monserrate (CAS 38554). 

Cancellaria obesa coronadoensis Durham (extinct; type locality), Pleistocene: Isla Coronados (SD-BE 
F-5). 

Cancellaria pulchra Sowerby, Pleistocene: Isla Coronados (SD-BE F-5). 

Cancellaria wigginsi, new species (extinct), Pleistocene: Isla Monserrate (CAS 38554). 

Cantharus anomalus Reeve, Pleistocene: Isla Coronados (SD-BE F-5). 

Cantharus pallidus Broderip and Sowerby, Pleistocene: Isla Monserrate (CAS 38554). 

Cerithium gemmatum Hinds, Pliocene: Isla Carmen (SD-BE F-6a, CAS 38539). Pleistocene: Isla 
Coronados (SD-BE F-5); Isla Monserrate (SD-BE F-10, CAS 38554). 

Cerithium maculosum Kiener, Pliocene: Isla Carmen (CAS 38539). Pleistocene: Isla Coronados 
(SD-BE F-5); Isla San Diego (SD-BE F-14); Isla Monserrate (SD-BE F-10); Isla Cerralvo (SD-BE 
Buz )e 

Cerithium sculptum Sowerby, Pleistocene: Isla Coronados (SD-BE F-5). 

Cerithium uncinatum Gmelin, Pliocene: Isla Carmen (CAS 38546). 

Conus brunneus Wood, Pleistocene: Isla Monserrate (SD-BE F-10). 

Conus diadema Sowerby, Pleistocene: Isla Coronados (SD-BE F-5). 

Conus dispar Sowerby, Pliocene: Isla Carmen (CAS 38546). Pleistocene: Isla Monserrate (CAS 
38554), 

Conus gladiator Broderip, Pliocene: Isla Carmen (CAS 38546). Pleistocene: Isla Coronados 
SD-BE F-5). 

Conus nux Broderip, Pleistocene: Isla Monserrate (SD-BE F-10). 

Conus patricius Hinds, Pleistocene: Isla Monserrate (SD-BE F-10). 

Conus perplexus Sowerby, Pleistocene: Isla Monserrate (SD-BE F-10); Isla San Diego (SD-BE F-14). 

Conus princeps Linné, Pliocene: Isla Carmen (SD-BE F-6a). Pleistocene: Isla Coronados (SD-BE 
F-5); Isla Monserrate (SD-BE F-10). 

Conus purpurascens Broderip, Pleistocene: Isla Coronados (SD-BE F-5); Isla Monserrate (SD-BE 
F-10, CAS 38554); Isla Cerralvo (SD-BE F-12). 

Conus regularis Sowerby, Pleistocene: Isla Coronados (SD-BE F-5); Isla Monserrate (SD-BE F-10). 

Conus scalaris Valenciennes, Pleistocene: Isla Coronados (SD-BE F-5). 

Conus ximenes Gray, Pleistocene: Isla Coronados (SD-BE F-5). 

Conus sp., Pliocene: Isla San José (CAS 38549). 

Crassispira cf. C. nymphia Pilsbry and Lowe, Pliocene: Isla Carmen (SD-BE F-6a). 

Crepidula arenata Broderip, Pleistocene: Isla Coronados (SD-BE F-5). 


1964} EMERSON AND HERTLEIN: INVERTEBRATE MEGAFOSSILS 351 


Crepidula onyx Sowerby, Pleistocene: Isla Coronados (SD-BE F-5). 

Crucibulum scutellatum Wood, Pliocene: Isla Carmen (SD-BE F-6a, CAS 38539). Pleistocene: Isla 
Monserrate (SD-BE F-10). 

Cypraea albuginosa Gray, Pleistocene: Isla San Diego (SD-BE F-14). 

Cypraea annettae Dall, Pliocene: Isla Carmen (SD-BE F-6a, CAS 38539 and 38546). Pleistocene: 
Isla Coronados (SD-BE F-5); Isla Monserrate (SD-BE F-10); Isla San Diego (SD-BE F-14). 

Cypraea arabicula Lamarck, Pleistocene: Isla Coronados (SD-BE F-5). 

Cypraecassis coarctata Sowerby, Pleistocene: Isla Coronados (SD-BE F-5). 

Diodora alta C. B. Adams, Pliocene: Isla Carmen (SD-BE F-6a, CAS 38539). 

Diodora inaequalis Sowerby, Pliocene: Isla Carmen (SD-BE F-6a, CAS 38539). Pleistocene: Isla 
Coronados (SD-BE F-5); Isla San Diego (SD-BE F-14). 

Enaeta cumingit form pedersenti Verrill, Pleistocene: Isla Coronados (SD-BE F-5). 

Fasciolaria princeps Sowerby, Pleistocene: Isla Coronados (SD-BE F-5). 

Fusinus cinereus Reeve, Pliocene: Isla Carmen (SD-BE F-6a). Pleistocene: Isla Coronados (SD-BE 
F-5). 

Gyrineum strongi Jordan (extinct), Pleistocene: Isla Coronados (SD-BE F-5); Isla Monserrate 
(SD-BE F-10); Isla San Diego (SD-BE F-14). 

Heliacus robertsae Durham (extinct), Pliocene: Isla Carmen (SD-BE F-6a). 

Hipponix antiquatus Linné, Pliocene: Isla Carmen (SD-BE F-6a, CAS 38539). Pleistocene: Isla 
Coronados (SD-BE F-5). 

Knefastia funiculata Kiener, Pleistocene: Isla Coronados (SD-BE F-5); Isla Monserrate (SD-BE F-10, 
CAS 38554). 

Modulus cerodes A. Adams, Pliocene: Isla Carmen (SD-BE F-6a). Pleistocene: Isla Coronados 
(SD-BE F-5). 

Morula ferruginosa Reeve, Pliocene: Isla Carmen (SD-BE F-6a). 

Morula lugubris C. B. Adams, Pleistocene: Isla Coronados (SD-BE F-5). 

Murex cf. M. elenensis Dall (fragment), Pleistocene: Isla Coronados (SD-BE F-5). 

Muricanthkus princeps Broderip, Pleistocene: Isla Cerralvo (SD-BE F-12). 

Nassarius tiarula Kiener, Pliocene: Isla Carmen (SD-BE F-6b). Pleistocene: Isla Coronados (SD-BE 
F-5); Isla Monserrate (SD-BE F-10, CAS 38554). 

Nassarius sp. (juvenile), Pliocene: Isla Carmen (SD-BE F-6a). 

Natica unifasciata Lamarck, Pliocene: Isla Carmen (CAS 38540). 

Nerita scabricosta Lamarck, Pleistocene: Isla Monserrate (SD-BE F-10, CAS 38554). 

Oliva incrassata Solander, Pleistocene: Isla Monserrate (SD-BE F-10, CAS 38554). 

Oliva polpasta Duclos, Pliocene: Isla Carmen (CAS 38540). 

Oliva porphyria Linné, Pleistocene: Isla Coronados (SD-BE F-5). 

Oliva spicata (Bolten) Réding, Pliocene: Isla Carmen (SD-BE F-6a, CAS 38546). Pleistocene: 
Isla Monserrate (SD-BE F-10, CAS 38554); Isla San Diego (SD-BE F-14). 

Olivella dama Wood, Pliocene: Isla Carmen (SD-BE F-6a and F-6b, CAS 38539). Pleistocene: 
Isla Coronados (SD-BE F-5); Isla Monserrate (SD-BE F-10, CAS 38554 and 38555). 

Olivella sp. (2 fragments), Pleistocene: Isla Coronados (SD-BE F-5). 

Parametaria dupontii Kiener, Pleistocene: Isla Coronados (SD-BE F-5); Isla San Diego (SD-BE 
F-14). 

Polinices bifasciata Gray, Pleistocene: Isla Coronados (SD-BE F-5); Isla Monserrate (SD-BE F-10). 

Polinices uber Valenciennes, Pliocene: Isla Carmen (SD-BE F-6a, CAS 38539). Pleistocene: Isla 
Coronados (SD-BE F-5); Isla Monserrate (SD-BE F-10, CAS 38554 and 38555). 

Polinices cf. P. uber Valenciennes, Pliocene: Isla Carmen (CAS 38546). 

Strombina maculosa Sowerby, Pleistocene: Isla Monserrate (SD-BE F-10, CAS 38554). 

Strombus galeatus Swainson, Pliocene: Isla Carmen (SD-BE F-6a, CAS 38539 and 38546). Pleisto- 
cene: Isla Monserrate (SD-BE F-10, CAS 38554); Isla Cerralvo (SD-BE F-12). 

Strombus gracilior Sowerby, Pliocene: Isla Carmen (CAS 38546). Pleistccene: Isla Monserrate 
(SD-BE F-10, CAS 38554). 

Strombus granulatus Swainson, Pliocene: Isla Carmen (SD-BE F-6a, CAS 38539 and 38546). Pleis- 
tocene: Isla Monserrate (SD-BE F-10, CAS 38554); Isla San Diego (SD-BE F-14). 

Strombus granulatus Swainson, including form cortezianuvs Durham, Pleistocene: Isla Coronados 
(SD-BE F-5). 

Tegula mariana Dall, Pliocene: Isla Carmen (SD-BE F-6a, CAS 38539). Pleistocene: Isla Coronados 
(SD-BE F-5). 

Terebra specillata Hinds, Pleistocene: Isla Monserrate (SD-BE F-10). 

Terebra cf. T. specillata Hinds, Pleistocene: Isla Coronados (SD-BE F-5). 

Terebra strigata Sowerby, Pleistocene: Isla Monserrate (SD-BE F-10). nist 

Terebra variegata Gray, Pleistocene: Isla Coronados (SD-BE F-5); Isla Monserrate (SD-BE F-10, 
CAS 38554 and 38555). 

Trivia californica Gray, Pleistocene: Isla Coronados (SD-BE F-5). 

Trivia solandri Sowerby, Pleistocene: Isla Coronados (SD-BE F-5). 


352 San Disco Society OF NATURAL HIsTory {Vou. 13 


Turbo fluctuosus Wood, Pliocene: Isla Carmen (SD-BE F-6a, CAS 38539). Pleistocene: Isla Coro- 
nados (SD-BE F-5); Isla Monserrate (SD-BE F-10, CAS 38554); Isla San Diego (SD-BE F-14). 

Turbo squamiger Reeve, Pleistocene: Isla Coronados (SD-BE F-5). 

Turbo species (operculum), Pliocene: Isla Carmen (CAS 38546). 

Turritella gonostoma Valenciennes, Pleistocene: Isla Coronados (SD-BE F-5). 

Turritella species indeterminate (mold), Pliocene: Isla Carmen (SD-BE F-7). 


ScAPHOPODA 
Dentalium semipolitum Broderip and Sowerby, Pleistocene: Isla Monserrate (SD-BE F-10). 


ECHIN ODERMATA 
ECHIN OIDEA 

Clypeaster bowersi Weaver, Pliocene: Isla Cerralvo (SD-BE F-13, CAS 38543). 

Clypeaster speciosus Verrill, Pleistocene: El Pulpito (SD-BE F-4). 

Clypeaster cf. C. speciosus Verrill, Pliocene: Isla Cerralvo (CAS 38543). 

Clypeaster sp. (fragments), Pliocene: Isla Monserrate (CAS 38551). 

Dendraster casseli Grant and Hertlein, Pliocene: Isla San José (SD-BE F-11). 

Dendraster cf. D. granti Durham, Pliocene: Isla San José (SD-BE F-11). 

Echinometra vanbrunti Agassiz, Pleistocene: Isla San Diego (SD-BE F-14). 

Encope californica Verrill, Pleistocene: El Pulpito (SD-BE F-4, CAS 38553); Isla Coronados 
(SD-BE F-5); Isla Monserrate (SD-BE F-10, CAS 38554). 

Encope grandis Agassiz, Pleistocene: Isla Monserrate (SD-BE F-10, CAS 38554). 

Encope cf. E. grandis inezana Durham, Pleistocene: El Pulpito (SD-BE F-4, CAS 38553). 

Encope sp., Pliocene: Isla Carmen (SD-BE F-7). Pleistocene: El Pulpito (SD-BE F-4). 

Eucidaris thouarsii Valenciennes, Pliocene: Isla San Esteban (SD-BE F-2); El Pulpito (SD-BE F-4, 
CAS 38553); Isla Carmen (SD-BE F-6a, CAS 38539). Pleistocene: Isla Monserrate (SD-BE F-10). 

Meoma cf. M. grandis Gray, Pleistocene: El Pulpito (CAS 38553). 


COELENTERATA 
SCLERACTINIA 
Colonial stony coral of undetermined generic afhnity (internal molds), Pliocene: Isla Cerralvo 
(SD-BE F-13). 
Coral, Pliccene: Isla San Esteban (SD-BE F-2); Isla Carmen (CAS 38546). 
Pocillopora robusta Verrill, Pleistocene: Isla Monserrate (SD-BE F-10); Isla Cerralvo (SD-BE F-12); 
Isla San Diego (SD-BE F-14). 
Porites californica Verrill, Pliocene: Isla Carmen (SD-BE F-6a, CAS 38539). Pleistocene: Isla Coro- 
nados (SD-BE F-5); Isla San Diego (SD-BE F-14). 
Porites species, Pleistocene: Isla Monserrate (SD-BE F-10). 


BrYOZOA 
EcToPROCcTA 
Bryozoa, Pleistocene: El Pulpito (SD-BE F-4). 
Conopeum commensale Kirkpatrick and Metzelaar, Pliocene: Isla Cerralvo (SD-BE F-13). 
ARTHROPODA 
CRUSTACEA 
Crab claw, Pleistocene: Isla Coronados (SD-BE F-5). 


CIRRIPEDIA 
Balanus tintinnabulum californicus Pilsbry (on pectens), Pliocene: Isla San José (SD-BE F-11); Isla 
Cerralvo (SD-BE F-13). 
Balanus trigonus Darwin, Pliocene: Isla Carmen (SD-BE F-7); Isla Cerralvo (SD-BE F-13). 
Balanus sp., Pliocene: Isla Partida (SD-BE F-3); Isla San José (SD-BE F-11a). 


ALGAE 
CALCAREOUS ALGAE 

Calcareous algae, Pliocene: Isla Carmen (SD-BE F-6a, CAS 38539); Isla Cerralvo (SD-BE F-13). 
Pleistocene: El Pulpito (SD-BE F-4); Isla Coronados (SD-BE F-5); Isla Monserrate (SD-BE F-10); 
Isla San Diego (SD-BF. F-14). 

SYSTEMATIC TREATMENT 

In this section, synonymies and notes are presented for mollusks and echinoids in the 
present collection that are of particular interest. The synonymies of these species are restricted, 
with a few exceptions, to records reported from the Gulf trough. The collecting stations for 
each species are listed in the previous section of this paper. References in the synonymies and 
the discussions in this section are not repeated in the “Literature Cited” of this report. 

A ae species of gastropod and two new subgeneric names in the Class Gastropoda are 
proposed. 


1964 } EMERSON AND HERTLEIN: INVERTEBRATE MEGAFOSSILS 


Ww 
Ww 
Ww 


Phylum MOLLUSCA 
Class PELECYPODA 
Family OSTREIDAE 


Ostrea angelica Rochebrune, 1895 
Figures 3a-c 


Ostrea cumingiana “Dunker’” [of west American authors}.— Vokes in Wilson, Bull. Amer. Assoc. 
Petrol. Geol., vol. 32, no. 9, p. 1784, 1948. “Santa Rosalia Formation (Pleistocene)”.— Vokes in Wilson, 
Univ. Nac. Auton. de Mexico, Inst. Geol. Geofis., and Geodesia, Bol. no. 53, p. 41, 1948. Same record 
as preceding reference. — Vokes in Wilson and Veytia, U.S. Geol. Surv., Bull. 960-F, p. 199, 1949. 
South side of Arroyo del Infierno, uppermost Pliocene or Pleistocene. Range, uppermost Pliocene to Recent. 
— Durham, Geol. Soc. America, Mem. 43, pt. 2, p. 58, pl. 5, fig. 6, and probably fig. 7 (as O. vespertina 
Conrad), 1950. Late Pliocene to Recent in the Gulf of California region. — Vokes in Wilson and Rocha, 
U.S. Geol. Surv., Prof. Paper 273, p. 40, 1955. Same as earlier records by Vokes. — Durham and Hert- 
lein in Arnold, Univ. Calif. Publ. Geogr., vol. 10, no. 4, p. 242, 1957. In small arroyo in the 200(?)-foot 
terrace, near the coast at Santa Rosalillita, Baja California. Probably uppermost Pliocene or lowermost 
Pleistocene. 

Not Ostrea cumingiana Dunker, Zeitschr. f. Malakozool., Jahrg. 3, p. 48, 1846. “Patria ignota.” 
{Mow believed to be a Japanese species}. 

Ostrea angelica Rochebrune, Bull. Mus. Nat. d’Hist. Nat. Paris, vol. 1, p. 241, 1895. “Baie de Los 
Angeles,” east coast of Baja California, Recent. — Contreras, An. Inst. Biol. (Mexico), vol. 3, no. 3, p. 
207, figs. 18 and 19, 1932. “Bahia de Los Angeles, Golfo de California.” —lHertlein and Emerson, Trans. 
San Diego Soc. Nat. Hist., vol. 12, no. 8, p. 165, 1956. Puerto Pefiasco, Sonora, Mexico; Pleistocene. — 
Hertlein, Bull. South. Calif. Acad. Sci., vol. 56, pt. 2, pp. 59, 63, 1957. Sea cliff about 1/ mile east of 
mouth of Arroyo del Coyote, 21/4 miles southeast of Punta Coyote, 11 miles northeast of La Paz, Baja 
California; Pleistocene. Also, Pleistocene elevated beach at Santa Inez Bay, Baja California; and Carmen 
Island; Pleistocene. Probably also “Ostrea cf. angelica” from low hills back of Santa Inez Bay, Baja Cali- 
fornia; Pleistocene. — Emerson and Chace, Trans. San Diego Soc. Nat. Hist., vol. 12, no. 2, pp. 338, 341, 
May 27, 1959. Tecolote Creek, San Diego, California; Pleistocene. ?Spanish Bight, San Diego, California; 
Pleistocene. Punta Pefasco, Sonora, Mexico; Recent. — Hertlein and Emerson, Amer. Mus. Novitates, no. 
1940, p. 8, June 5, 1959. Maria Madre Island, Tres Marias Islands, Mexico; Pliocene. — Emerson, Amer. Mus. 
Novitates, no. 1995, p. 5, March 29, 1960. “South side of Ceralvo Island,” Pleistocene. — Emerson, Amer. 
Mus. Novitates, no. 2013, p. 6, August 18, 1960. Middens of San José Island, Gulf of California. 

{?]} Ostrea cf. angelica Rochebrune, Vokes in Wilson, Bull. Amer. Assoc. Petrol. Geol., vol. 32, no. 9, 
pp. 1782 and 1783, 1948. “Gloria Formation (Middle Pliocene)” and “Infierno Formation (Upper Plio- 
cene)”’. — Vokes in Wilson, Univ. Nac. Auton. Mexico, Inst. Geol. Geofis. y Geodesia, Bol., no. 53, pp. 
37, 38, 1948. Same records as in preceding reference. — Vokes in Wilson and Veytia, U.S. Geol. Surv., 
Bull. 960-F, p. 197, 1949. Same record as Vokes, 1948. — Vokes in Wilson and Rocha, U.S. Geol. Surv., 
Prof. Paper 273, pp. 36, 39, 1955. Same records as in preceding reference. 


Range. — Mid-Pliocene to Recent. Recent from San Ignacio Lagoon to San Felipe, Baja 
California, and south to Mazatlan, Sinaloa, Mexico. 

Remarks. — Many specimens of this species were collected from beds of Pliocene and 
of Pleistocene age. The valves are identical with Recent specimens living in the same region. 

The valves of Ostrea angelica are rather thick, strongly plicated, and interlock along the 
margins. Denticles are present along the margin just below the hinge. The interior of Recent 
specimens is tinged with green. Large specimens attain a length (beak to base) of 123 mm. 

The valves of Ostrea angelica are much larger and the area of attachment is larger than 
that of O. vespertina Conrad from the type locality, Imperial County, California (compare 
figures 3a-c with 3d-h). 


Ostrea cf. O. californica Marcou, 1858 


The following references are to typical O. californica. : 

Ostrea virginica var. californica Marcou, Geol. North America (Zurich), p. 32, pl. 5, figs. 2, 2a, 
1858. “In the Colorado desert, California; near Carrizo Creek, between San Diego and Fort Yuma. 
Miocene or Pliocene. 

Ostrea iridescens Gray, G D. Hanna, Proc. Calif. Acad. Sci., Fourth Ser., vol. 14, no. 18, p. 468, 
pl. 26, figs. 4, 5, 6, 7, March 23, 1926. Coyote Mountain, Imperial County, California, Pliocene. 

Not Ostrea iridescens Gray, in Hanley, 1854. a: 

Ostrea californica Marcou, Hanna and Hertlein, Nautilus, vol. 41, no. 2, pp. 45, 46, 1927. Earlier 
records cited. eux 

Geologic Record. — Previously known only from the Imperial formation. 


354 SAN Drseco Society OF NATURAL HISTORY pVornl3 


Remarks. — A large, thick, elongated lower valve of an oyster’ was collected at Loc. F-2A, 
on the south slope on the northwest side of Isla Salsipuedes at an altitude of about 100 feet. 
It is 271 mm. long, 105 mm. wide, and 60 mm. thick. This valve was found with Recent 
shells in kitchen middens and its original source is unknown. 

The shape, ligamental groove and general shell characters of this valve are similar to 
those of Ostrea californica from strata of Pliocene age in Imperial County, California, except 
that it is much larger and thicker. Likewise, it is much larger and thicker than O. corteziensis 
Hertlein, a species that lives at the present time in the Gulf of California. Ostrea corteziensis 
was cited as a fossil from the Gulf region by Durham (1950:58, pl. 4, fig. 1), as Ostrea 
chilensis Philippi (see Hertlein, L. G., Bull. South. Calif. Acad. Sci., vol. 50, pt. 2, p. 72, 


1951). 
The general appearance of the exterior of this fossil, except for greater size and thick- 


ness, resembles O. englekeyi Hertlein, a species described from mid-Miocene strata on Santa 
Rosa Island, California. The characters of the interior of the type specimen of O. englekeyi 


are not known. 
Positive identification of this oyster, based on this somewhat worn valve, is doubtful; 


certainly it is a fossil form. 
Ostrea heermanni Conrad, 1855 


Ostrea heermanni Conrad, Proc. Acad. Nat. Sci. Philadelphia, vol. 7, p. 267, February, 1855 [under 
heading of Miocene Species}. “Colorado Desert. Dr. Heermann. Shell silicified.”” — Conrad, House Exec. 
Decument 129, 33rd Congress, Ist Session, p. 15, July, 1855. “Locality. — Carrizo creek, Colorado desert. 
Dr. Heermann.” — Conrad, U.S. Pac. Railroad Repts., vol. 5, pt. 2, p. 326, 1857. “Locality. — Carrizo 
creek, Colorado desert. Dr. Heermann.” — Gabb, Geol. Surv. Calif., Palaeo., vol. 2, p. 107, 1869. “Pliocene, 
Colorado Desert.”’ — Heilprin, U.S. Geol. Sury., Fourth Ann. Rept., p. 314, 1884. Earlier records cited. 
— Orcutt, West Amer. Sci., vol. 12, no. 1 (102), p. 12, 1901 (as Ostrea hermanni). Miocene: Carrizo 
Creek. — Hanna, Proc. Calif. Acad. Sci., Fourth Ser., vol. 14, no. 18, p. 467, pl. 22, figs. 7, 8; pl. 23, 
figs. 1, 2, 1926. Loc. 693 (CAS), Yuba Buttes, Imperial County, California; Pliocene. — Woodring, 
Carnegie Inst. Washington, Publ. no. 418, p. 11, 1931. Painted Hill, Miocene. — Hertlein, Jour. Paleo., 
vol. 5, no. 4, p. 366, 367, 1931. Float five kilometers north of Santa Rosalia, Baja California; late Pliocene 
or Pleistocene; Arroyo Cadejé above the Rancho Cadejé, Baja California [Pliocene}.— Woodring, U. S. 
Geol. Surv., Prof. Paper, 190, p. 46, 1938. Imperial formation, Colorado Desert. — Hertlein, Bull. South. 
Calif. Acad. Sci., vol. 56, pt. 2, p. 58, 1957. Two miles southeast of La Rivera and about one mile from 
the coast of Baja California; also in arroyo wall 414 miles northeast of Sierra de la Trinidad, 9 miles 
southeast of Santiago, Baja California; Pliocene. 

Range. — Pliocene; early and mid-Pliocene of Durham (1950). 

Remarks. — Several fragments of this large, thick-shelled, rounded oyster, one 126 mm. 
long, were collected on Isla Cerralvo. These reveal the presence of the large ligamental groove 
and the rounded, sunken muscle impression, comparable to typical Ostrea heermanni from 
strata of Pliocene age in Imperial County, California, as well as other specimens from south- 
eastern Baja California. The present specimens closely resemble those illustrated by Hanna 
(1926). 

Family PECTINIDAE 


Chlamys (Argopecten) abietis (E. K. Jordan and Hertlein, 1926) 
Figures 4a-e 

Pecten (Plagioctenium) abietis E. K. Jordan and Hertlein, Proc. Calif. Acad. Sci., Fourth Ser., vol. 
15, no. 4, p. 214, pl. 23, figs. 1, 3, 7, April 26, 1926. “Arroyo Hondo, Maria Madre Island, Mexico; 
upper Pliocene.” — Touade, Econ. Geol., vol. 25, no. 2, p. 122, 1930. Upper Salada or calcareous series, 
Santa Rosalia, Baja California; Pliocene, probably upper Pliocene. — Hertlein, Bull. South. Calif. Acad. 
Sci., vol. 56, pt. 2, p. 60, 1957. “Santa Inez Bay, east coast of Lower California. From hard, brownish, 
limy, sandy strata in low hills back of the bay which, toward the beach, are overlain by Pleistocene terrace 
material”; Pliocene. — Foose, Bull. Amer. Assoc. Petrol. Geol., vol. 46, no. 9, p. 1743, 1962. Maria Cleofa 
Island, Tres Marias Islands, Pliocene. 

Aequipecten abietis Jordan and Hertlein, Vokes in Wilson, Bull. Amer. Assoc. Petrol. Geol., vol. 32, 


'Since the present paper was submitted for publication, oysters similar to the present species were collected 
by Dr. C. C. McFall from strata on the southwest end of Concepcion Peninsula, Concepcion Bay, Baja 
California. Species of early or middle Pliocene age accompanying the oysters include, Ostrea heermanni 
Conrad, Chlamys abietis (Jordan and Hertlein), Pecten subnodosus Sowerby, Cantharus cf. C. pallidus 
Broderip and Sowerby and Turritella cf. T. imperialis Hanna. 


1964 } EMERSON AND HERTLEIN: INVERTEBRATE MEGAFOSSIIS 355 


no. 9, pp. 1780, 1782, 1783, 1948. Gloria formation, middle Pliocene and Inferno fort 


Pliocene. — Vokes in Wilson, Univ. Nac. Auton. Mexico, Inst. Geol., Geofis. y Geodesia. Bol. no. 53. a 
37, 39, 1948. Same records as in preceding reference. — Vokes in Wilson and Veytia. U.S. Geol. Seve 
Bull. 960-F, pp. 194, 197, 198, 1949. Cited from Lower, middle, and upper Pliocene. — Durham, Geol. 
Soc. America, Mem. 43, pt. 2, p. 62, pl. 10, figs. 4, 7; pl. 11, fig. 4, August 10, 1950. Various localities 
cited in Gulf of California region, lower, middle and upper Pliocene. — Vokes in Wilson and Rocha. U.S 


Geol. Surv., Prof. Paper 273, pp. 32, 36, 39, 1955. Early, middle and late Pliocene, 

Pecten (Argopecten) abietis E. K. Jordan and Hertlein, Hertlein and Emerson, Amer. Mus. Novitates. 
no. 1940, p. 8, 1959. Maria Cleofa and Maria Madre Islands, Tres Marias Islands, Pliocene. 

Range. — Pliocene; early to mid-Pliocene of Durham (1950). 

Remarks. — This species is one of the most common fossils characteristic of strata of 
Pliocene age in the Gulf of California region. 

The radial ribs on shells of this species, especially in the earlier stage, are triangular to 
subtriangular and the interspaces are very marrow. A dense fringe of concentric lamellae 
cover the sides of the ribs and interspaces. These characters serve to separate it from Chlamys 
(Argopecten) circularis (Sowerby), in which the ribs are more rounded and narrower in 
cross-section, the interspaces are wider, and the concentric lamellae are only weakly developed 
(see fig. 4a-e). 

Chlamys (Pacifipecten) tumbezensis (d’Orbigny, 1835) 

Pecten aspersus Sowerby, Proc. Zool. Soc. London for 1835, p. 110, October 9, 1835. “Hab. ad 
Tumbez, Peruviae. Dredged in soft mud at a depth of five fathoms.” 

Not Pecten aspersus Lamarck, 1819. 

Pecten tumbezensis d’Orbigny, Voy. dans |’Amérique Méridionale, vol. 5, p. 663, 1846. Tumbez, 
Peru. New name for Pecten aspersus Sowerby, 1835, not Pecten aspersus Sowerby, 1825. — Peile in Bos- 
worth, Geol. northwest Peru, p. 178, pl. 25, fig. 8, 1922. “Talara Tablazo” and “Lobitos Tablazo”’, 
Quaternary, Peru. — Olsson, Bull. Amer. Paleo., vol. 27, no. 106, p. 171 (19), 1942. Rio La Vaca, Costa 
Rica, Pliocene. — Lemon and Churcher, Amer. Jour. Sci., vol. 259, no. 6, p. 415, 1961. Talara Tablazo, 
Peru, Pleistocene. 

Pecten (Leptopecten) tumbezensis d’Orbigny, Hertlein, Proc. Calif. Acad. Sci., Fourth ser., vol. 21, 
no. 25, p. 314, pl. 19, figs. 11, 12, 1935. Gulf of California to Tumbez and Paita, Peru. Also Quaternary 
of Peru. {With synonymy}. — Hoffstetter, Bol. Inform. Cienc. Nat., vol. 2, nos. 13-14, p. 73, 1948. Tercer 
Tablazo [youngest], Santa Elena Peninsula, Ecuador. — Hertlein and Strong, Bull. Amer. Mus. Nat. Hist., 
vol. 107, art. 2, p. 180, 1955. Gulf of California and east of San José del Cabo, Baja California to Paita, 
Peru. [With synonymy]. — Grau, Allan Hancock Pac. Exped., vol. 23, p. 118, pl. 40, 1959. Gulf of Cali- 
fornia to Paita, Peru. [With synonymy }. 

Pecten (Plagioctenium) tumbezensis d’Orbigny, Pilsbry and Olsson, Proc. Acad. Nat. Sci. Philadelphia, 
vol. 93, p. 55, 1941. Canoa formation, Punta Blanca, Ecuador. . 

Aequipecten (Pacifipecten) tumbezensis d’Orbigny, Olsson, Mollusks of the Tropical Eastern Pacific 
(Paleo. Res. Inst.: Ithaca, New York), p. 164, pl. 21, figs. 2-2c, 1961. Gulf of California to Paita, Peru. 

Range. — Pliocene to Recent. Recent from south of Consag Rock (31° N.), Gulf of 
California, to Paita, Peru, in 1.8-110 meters (1-61 fathoms). 

Remarks. — About 25 valves, in various stages of preservation, were collected in grayish- 
brown sediment on Isla Partida. Many of these are bent and deformed. This feature, as 
well as the character of the strata in which the shells occur, leads us to assign questionably a 
Pliocene age to these fossils. 

The shell of this species is rather small, rarely more than 35 mm. high, and comparatively 
thick for its size. It is sculptured with 13 to 15 radial ribs; those on the right valve are 
nearly square, whereas those on the left are narrowly triangular and are separated by wider 
interspaces. 

Family SPORTELLIDAE 
Basterotia (Basterotella) hertleinij Durham, 1950 
Figures 4g-j 

Basterotia kertleini Durham, Geol. Soc. Amer. Mem. 43, pt. 2, pp. 94, 95, pl. 25, figs. 4, 11, August 
10, 1950. Puerto Ballandra, Carmen Island (type locality), late Pliocene; Marquer Bay, Carmen Island, 
late Pliocene. ; 

Range. — Late Pliocene (Marquer formation). . 

Remarks. — Paired valves of this species were common in the siltstone bed that forms 
the basal part of the large coral reef of Pliocene age at Puerto Ballandra, Isla Carmen; see 
photographs in Lindsay (1962: figs. 16, 17). Only one single valve was found in the coarser 
sediments of the reef. 


356 San Disco Society oF Naturat History {[ VoL. 13 


1964} EMERSON AND HErTLEIN: INVERTEBRATE MEGAFOSSIIS 357 


This species seems to be most closely related to Basterotia (Basterotella) ecuadoriana 
Olsson, which was described from Recent specimens taken at Manta and Santa Elena. Ecuador. 
However, the present specimens appear to be smaller and to have finer surface sculpture 
than those described by Olsson. Basterotia (B.) peninsularis Jordan, known from the Pleisto- 
cene of Magdalena Bay, Baja California, and Islas Monserrate and Coronados, and reported to 
be living at present from Port Guatulco, Oaxaca, Mexico, to the Galapagos Islands, is also a 
similar species, but has a less elongate and more inflated shell. 


Family VENERIDAE 


Chione (Nioche) picta Willett, 1944 


{?] Tapes squamosa Carpenter, Catalogue of the Collection of Mazatlan Shells, . . . (British Museum: 
London), p. 78, 1857. “Mazatlan; from washings of Chamae and Spondyli; extremely rare.” Recent. 

Chione picta Dall, Manger, Johns Hopkins Univ., Stud. Geol., no. 11, p. 289, 1934. San Quintin 
Bay, Baja California, Mexico. Pleistocene. [Nomen nudum] 

Chione picta Willett, ex Dall MS., Bull. So. Calif. Acad. Sci., vol. 43, pp. 21, 22, pl. 8, May 31, 
1944. Living in Magdalena Bay, Baja California, Mexico (type locality) and at other west Mexican 
localities, and from Pleistocene deposits in the Los Angeles basin of southern California. — Woodring, U. S. 
Geol. Surv. Prof. Paper no. 207, p. 84, pl. 37, figs. 3, 4, 1946. Magdalena Bay, Lower California to 
Panama. Recent. San Pedro district, Los Angeles Co., California, Pleistocene. — DuShane, Veliger (Berkeley, 
California), vol. 5, p. 43, July 1, 1962. Puertecitos, Baja California, Mexico. Recent. 

Chione (Nioche) picta “Dall” Willett, Hertlein and Strong, Zoologica (New York, Zool. Soc.), vol. 
33, pp. 187, 188, December 31, 1948. “Port Guatulco, Mexico, in 6-7 fathoms.” 

“Chione” picta “Dall” Willett, Valentine, Trans. San Diego Soc. Nat. Hist., vol. 12, p. 195, July 2, 
1956. Potrero Canyon, Los Angeles Co., California. Pleistocene. — Valentine, Trans. San Diego Soc. Nat. 
Hist. vol. 12, p. 296, September 25, 1957. Near Punta Descanso, Baja California, Mexico. Pleistocene. — 
Kanakoff and Emerson, Contrib. Sci., Los Angeles Co. Museum, no. 13, pp. 22, 34 [table 1], October 
14, 1959. Newport Bay, Orange Co., California. Pleistocene. 

Chione (Timoclea) picta Willett, Keen, Sea Shells of Tropical West America (Stanford Univ. 
Press: Stanford, California), p. 146, fig. 335, 1958. Magdalena Bay, Lower California and southward to 
Panama, mostly offshore in depths to 7 fathoms. — McLean, Trans. San Diego Soc. Nat. Hist., vol. 12, 
p. 461, August 15, 1961. Los Angeles Bay, Baja California, Mexico. Recent. 

Nioche (Nioche) squamosa (Carpenter), Olsson, Mollusks of the Tropical Eastern Pacific (Paleo. 
Res. Inst.: Ithaca, New York), pp. 309, 310, March 10, 1961, pl. 49, fig. 10. Puerto Armuelles, Panama; 
Puerto Callo and Santa Elena, Ecuador; Zorritos and Bayovar, Peru. Recent. 

{?} Not Tapes squamosa Carpenter, 1857. 

Range. — Pleistocene to Recent. Recent from Magdalena Bay, Baja California, to Punta 
Penasco, Sonora, Mexico, in the Gulf of California, and south to Bayovar, Peru. 

Remarks. — Although this distinctive venerid is known from deposits of Pleistocene age 
in the Los Angeles basin of southern California and along the west coast of Baja California, 
Mexico, the present record, based on a single specimen, appears to be the first reported oc- 
currence of this species as a fossil in the Gulf of California. 

According to Keen (1958, p. 622), the type lot of Tapes squamosa Carpenter, which 
she examined in the British Museum (Natural History), “comprises two juvenile shells of 
which the larger may be a Protothaca sp. rather than an Irus” [= ?Irus (Paphonotia) 
ellipticus (Sowerby) }. Olsson (1961, p. 310), on the other hand, considered the unpublished 
drawings of Tapes squamosa for Carpenter’s (1857) Mazatlan Catalogue to represent the 
species later described by Willett (1944), from a manuscript name of the late W. H. Dall, 


Fig. 3. a-c, Ostrea angelica Rochebrune, 1895. a, b, F-1 (SD-BE), Isla Angel de la Guarda, ?Plio- 
cene, height (beak to base) 90 mm., width 65 mm.; outside and inside views, respectively, of upper valve. 
c, 38539 (CAS), Isla Carmen, Pliocene, ventral view of both valves in place, length 55.5 mm., height 
60.8 mm. 

d-h, Ostrea vespertina Conrad, 1854, topotypes. d, e, 691 (CAS), east of Coyote Mountain, Imperial 
County, California, Pliocene; d, inside view of upper valve, length 23.4 mm., height 29.5 mm.; e, inside 
view of lower valve, length 26.2 mm., height 31 mm. f-h, 35010 (CAS), northeast side of Coyote Moun- 
tain, Imperial County, California, Pliocene; f, outside view of upper valve, length 34 mm., height 36 mm.; 
g, h, outside and inside views, respectively, of lower valve, length 48.5 mm., height 43 mm. 

ints Semele verrucosa Morch, 1860. F-5 (SD-BE), Isla Coronados, Pleistocene, length 59.5 mm., 
height 47 mm. (specimen incomplete). Outside and inside views, respectively, of right valve. 

k, Morula (Morunella) lugubris (C. B. Adams), 1852. F-5 (SD-BE), Isla Coronados, Pleistocene, 
height 10 mm., diameter 4.8 mm., apertural view. Type species of Morunella, new subgenus. 


IVOrE213 


SAN Deco Society OF NATURAL HIsTory 


358 


1964 } EMERSON AND HErTLEIN: INVERTEBRATE MEGAFOSSILS 359 


as “Chione picta Dall.” The identity of Carpenter’s taxon, however, can not be determined 
until one of the syntypes is designated the lectotype of Tapes squamosa Carpenter, 1857. 


Family PETRICOLIDAE 
Petricola (Petricola) robusta Sowerby, 1834 


Petricola robusta Sowerby, Proc. Zool. Soc. London for 1834, p. 47, September 26, 1834. “Hal 
Panaman et ad Insulam Muerte dictam.” “Found in rocks at the depth of from six to eleven fathoms. 
Sowerby, Thes. Conch., vol. 2, p. 775, pl. 166, figs. 16, 17, 1854. “In rocks, six to eleven fathoms, 
Panama.” — Sowerby, Conch. Icon., vol. 19, Petricola, species 20, pl. 3, figs. 20a, 20b, 1874. Panama. 
Durham, Geol. Soc. America, Mem. 43, p. 87, pl. 23, fig. 15, 1950. Coronado Island and San Marcos 
Island; Pleistocene. 

Petricola sinuosa Conrad, Proc. Acad. Nat. Sci. Philadelphia, vol. 4, p. 155, 1849. [No exact locality 
cited, title states “Shells from the coasts of Lower California and Peru.” } 

Petricola bulbosa Gould, Proc. Boston Soc. Nat. Hist., vol. 4, p. 88, November, 1851. “Inhabits 
Guaymas.”’ — Gould, Otia Conch. (Gould and Lincoln: Boston), p. 210, 1862. “Petricola bulbosa {is 
P. robusta, Sowb.}.” 

Petricola venusta De Folin, Les Meéléagrinicoles, Soc. Havraise d’Etudes Diverses (Havre), p. 20, pl. 3, 
figs. 5-7, 1867. [Two localities cited (p. 9)} “les unes ont été péchées aux environs des Negritos . . . les 
autres ont été prises autour des iles aux Perles, dans la Baie de Panama.” 

Petricola (Petricola) robusta Sowerby, Hertlein and Strong, Zoologica, New York, Zool. Soc., vol. 
33, pt. 4, no. 13, p. 194, 1948. Gulf of California to Guayaquil, Ecuador. — Olsson, Mollusks of the 
Tropical Eastern Pacific (Paleo. Res. Inst.: Ithaca, New York), p. 315, pl. 54, figs. 2, 2a, 1961. Gulf of 
California to Caleta Sal, Peru. 

Petricola (Rupellaria) robusta Sowerby, Keen, Sea Shells of Tropical West America (Stanford Univ. 
Press: Stanford, California), p. 152, fig. 348, 1958. Puerto Pefasco and the Gulf of California to Guaya- 
quil, Ecuador, boring in hard clay. 

Range. — Pleistocene to Recent. Recent from Punta Peftasco, Gulf of California, to Boca 
Pan and Caleto Sal, Peru, intertidal to 24 meters (13 fathoms). 

Remarks. — Specimens of this species were collected from burrows in coral heads from 
Pleistocene deposits on Isla Coronados. 

Petricola robusta was originally described from a tropical west American locality. It has 
been cited as occurring in South Africa by E. A. Smith (Ann. Natal Mus., vol. 1, pt. 1, 
p- 65, 1906), by Turton (Mar. Shells of Port Alfred, South Africa, p. 246, no. 1733, 1946) 
and more recently by Barnard (Beginner’s Guide to South African Shells, p. 176, pl. 27, 
fig. 2, 1951), but Bartsch (U.S. Nat. Mus. Bull. 91, p. 247, 1915), who reported on the 


Turton Collection, pointed out that this is a Panamanian species and does not occur in 


South Africa. 
Family SEMELIDAE 
Semele verrucosa Morch, 1860 
Figures 3i, | 

Semele (Amphidesma) verrucosa Mérch, Malakozool. Blatter, Bd. 7, p. 190, December, 1860. “Los 
Bocorones” Islands, Costa Rica. 

Semele verrucosa Morch, Hertlein and Strong, Zoologica, New York Zool. Soc., vol. 34, pt. 4, no. 
19, p. 249, pl. 1, figs. 21, 24, December 30, 1949. Los Bocorones Islands, Costa Rica, to Hannibal Bank, 
Panama. — Keen, Sea Shells of Tropical West America (Stanford Univ. Press: Stanford, California), 
p. 202, fig. 504, 1958. Costa Rica to Panama, possibly intertidally, and to depths of 40 fathoms. — Olsson, 
Mollusks of the Tropical Eastern Pacific (Paleo. Res. Inst.: Ithaca, New York), p. 366, pl. 65, figs. 1-1b, 
March 10, 1961. Gulf of California to the Pearl Islands, Panama. 


Fig. 4 a-e, Chlamys (Argopecten) abietis (E. K. Jordan and Hertlein, 1926). a, F-11 (SD-BE), 
Isla San José, Pliocene, right valve, length 46 mm., height 43.6 mm.; b, F-9 (SD-BE), Isla Monserrate, 
Pliocene, left valve, length 49.6 mm., height 46.5 mm., convexity 13.6 mm.; c, F-9 (SD-BE), Isla Monserrate, 
Pliocene, right valve, length 53.9 mm., height 50.9 mm., convexity 14.5 mm.; d, F-11 (SD-BE), Isla San 
José, Pliocene, left valve, length 66 mm., height 63.5 mm.; e, F-11 (SD-BE), Isla San Jose, right 
valve, length 128 mm., height 118 mm. ' 

f, Pecten (Patinopecten) marquerensis (Durham), 1950. F-11, Isla San Jose, Pliocene, right valve, 
length 134 mm., height 120.5 mm. ‘ Lc 

g-j, Basterotia (Basterotella) hertleini Durham, 1950. F-6b (SD-BE), Puerto Ballandra, Isla_ varmen, 
Pliocene. g, h, exterior and interior views, respectively, of left valve, length 12.2 mm., height 7.8 mm.; 
i, interior view of right valve, length 12.6 mm., height 7 mm.; j, exterior view of right valve, 
mm., height 7 mm. 


length 11.4 


360 SAN Disco SocrETY OF NATURAL HISTORY { VoL. 13 


Range. — Pleistocene to Recent. Recent from Los Bocorones Islands, Costa Rica, to the 
Pearl Islands, Panama, in 64-73 meters (35-40 fathoms). ?Gulf of California. 

Remarks. —One right valve of this species, 60 mm. long and 46.5 mm. high, was col- 
lected from strata of Pleistocene age on Isla Coronados. The specimen retains much of the 
scaly, verrucose projections on the concentric ribs that are typical of this species. 

This is the first fossil record for this species. We have not been able to verify the presence 
of living populations of this species in the Gulf of California (see Olsson, 1961:366). 


Class GASTROPODA 
Family BURSIDAE 
Genus Gyrineum Link, 1807 

Gyrineum Link, Beschreibung der Naturalien-Sammlung der Universitat zu Rostock, p. 123, 1807. — 
Dall, 1904, Smithsonian Miscell. Colls., vol. 47, p. 131. 

Type species. — Murex gyrinus Linné, 1758, by subsequent designation of Dall (1904). 

Diagnosis. —Cymatiid species having a small to medium size, subovate shell, with two 
varices that are generally continuous and opposite and with a short, interiorly reflected, 
anterior canal; posterior canal obsolete, closed. 

Remarks. — The type species is a common constituent of the modern Indo-Pacific 
faunal province. 

The type designation by Rovereto (Atti Soc. Ligustica, vol. 10, pp. 101-110, 1899) of 
“G.[yrineum} spinosum Dillw.{yn]}? is invalid, because that taxon was not included in the 
list of species originally assigned to Gyrineum by Link (1807). 


Subgenus Bechtelia, new subgenus 


Type species. —Gyrineum strongi Jordan, 1936, Pleistocene, west Mexico. 

Diagnosis. — Shell similar to Gyrineum, but with the two varices interrupted by rounded 
nodes and with a short, nearly straight, open posterior canal. 

Remarks. — Several Gyrineum-like species are known from west North American deposits, 
ranging in age from Eocene to Pleistocene. The type species of this new taxon appears to 
represent the evolutionary climax of this group in the Cenozoic deposits of this region, as no 
congeneric species are known at the present time from eastern Pacific waters. 

We take pleasure in naming this subgenus in honor of Mr. K. K. Bechtel of San 
Francisco, California. 


Gyrineum (Bechtelia) strongi Jordan, 1936 
Figure 5g 

Gyrineum strongi E. K. Jordan, Contrib. Dept. Geol. Stanford Univ., vol. 1, no. 4, p. 160, pl. 18, 
fig. 10, November 13, 1936. Loc. 982 (CAS), along beach, south of village, Magdalena Bay, Baja Cali- 
fornia, Mexico. Pleistocene. — Durham, Mem. Geol. Soc. Amer., no. 43, pp. 28, 115, pl. 29, figs. 19, 22, 
August 10, 1950. Loc. A-3550 (UCMP), Coronados Island. Pleistocene. Loc. A-3598 (UCMP), Con- 
cepcion Bay, Baja California. Pleistocene. Santa Rosalia area, Baja California. Pleistocene. 

Range. — Pleistocene. 

Remarks. — This distinctive species, which is not known to be living, was not previously 
reported from Islas Monserrate and San Diego. All records are from terrace deposits of ap- 
parently late Pleistocene age. 


Family THAIDIDAE 
Genus Morula Schumacher, 1817 


Morula Schumacher, Essai d’une Nouveau Systeme des Habitations des vers Testacés, p. 227, 1817. 

Type species. — Morula papillosa Schumacher, 1817 {=Drupa uva (Bolten) Roding, 
1798}, by monotypy. 

Diagnosis. — Shell small, thick, sub-fusiform, spire elevated; aperture sub-linear, generally 
constricted by several singularly arranged labial teeth; columella typically with a prominent 
axial fold, this fold less commonly weakly developed or completely lacking; radula similar to 
Drupa (sensu stricto) but base of central tooth modified, margins generally smooth or 
wrinkled, without true denticles; operculum similar to that of Drupa. 


1964} EMERSON AND HErTLEIN: INVERTEBRATE | SILS 361 


Remarks. — On shell characters alone, some members of this group at 


lificult to separate 
from Drupa (sensu lato). In Morula the shell tends to be more fusiform in ‘ae daa do 
somewhat oval shell of Drupa (sensu stricto). There are also several major differences in 
the radular structure of the two groups. The base of the central tooth in Morula is medially 
convex instead of being straight or concave as in Drupa (s.s.). The medial bulge is actually 
the base of the central cusp, which, unlike Drupa (s.s.), extends the entire width of the 


central tooth as a heavy, raised rod. The margins of the central teeth are never denticulate. 
but have denticle-like wrinkles that indent the margin and terminate marginally as small 
serrations. The radula in species of the subgenus Drupella Thiele is greatly modified. 


Subgenus Morunella, new subgenus 
Type species. — Buccinum lugubre Adams, 1852. 


Diagnosis. — Surface of shell finely nodose; edge of labrum sharp, thickened within by 
a ridge possessing several lirae, each of which terminates in minute, labial denticles. 


Remarks. — Although the type species was originally placed in the Buccinidae and was 
subsequently referred to the Fasciolariidae, Fusinidae and Muricidae, it appears to be refer- 
able to the Muricacea and to be most closely allied with the genus Morula of the subfamily 
Drupinae. 


Morula (Morunella) lugubris (C. B. Adams), 1852 


Figure 3k 


Buccinum lugubre Adams, Ann. Lyceum Nat. Hist. New York, vol. 5, p. 293, June, 1852. “Panama, 
and Taboga.” Recent.— Adams, Catalogue of Shells collected at Panama with Notes on Synonymy, 
Station and Geographic Distribution (New York), p. 293 (separate p. 69), 1852. — Turner, Occas. 
Papers Moll., vol. 2, p. 61, pl. 5, fig. 7, {type specimen] September 22, 1956. — Hertlein, Veliger (Ber- 
keley, California), vol. 3, p. 8, July 1, 1960. 

Latirus lugubris C. B. Adams, Pilsbry and Lowe, Proc. Acad. Nat. Sci. Philadelphia, vol. 83, p. 114, 
May 21, 1932. Guaymas, Mazatlan, and Acapulco, Mexico and San Juan del Sur, Nicaragua. Recent. — 
Lowe, Trans. San Diego Soc. Nat. Hist., vol. 8, p. 30. Punta Penasco, Mexico. Recent. 

Fusinus luteopictus (Dall) Willett {in part, fide Burch, 1940, and others}, Trans. San Diego Soc. 
Nat. Hist., vol. 8, p. 396, December 15, 1937. Baldwin Hills, near Playa del Rey, California. Pleistocene. 

Not Fusus luteopictus Dall, 1877. 

Cantharus lugubris C. B. Adams, Burch, T., Nautilus, vol. 54, p. 47, pl. 2, figs. 5-7, October, 1940. 
Off Catalina Island and Redondo Beach, California; and Playa del Rey, California, Pleistocene. 

Drupa lugubris C. B. Adams, Burch, T., Minutes Conchol. Club South. Calif., no. 17, p. 7, Novem- 
ber, 1942. Redondo Beach, California. — Burch, J. Q. [editor}, Ibid., no. 52, p. 10, September, 1945. 
Redondo Beach, California to Panama. Recent. Baldwin Hills, Los Angeles Co., California. Pleistocene 
{specimens confused with Fusinus}. 

Morula lugubris (C. B. Adams), Keen, Sea Shells of Tropical West America, p. 376, fig. 412, 1958. 
San Diego, California, to Panama. Recent. — Emerson and Chace, Trans. San Diego Soc. Nat. Hist., vol. 
12, pp. 338, 342, May 27, 1959. Tecoiote Creek, San Diego, California. Pleistocene. —DuShane, Veliger 
(Berkeley, California) vol. 5, p. 47, July 1, 1962. Puertecitos, Baja California, Mexico. Recent. 


Range. — Pleistocene to Recent. Recent from off Catalina Island and Redondo Beach, 
California; Punta Pefiasco, Sonora, Mexico, in the Gulf of California, and south to Panama. 


Remarks. — The present specimen apparently is the first fossil record for this species in 
the Gulf of California. This species was previously reported from Pleistocene deposits in the 
Los Angeles and San Diego basins of southern California, and the species is known to range 
at the present time from southern California to Panama. 


A very similar species, Morula (Morunella) didyma (Schwengel) was described from 
200 feet, off Palm Beach, Florida (Schwengel, Nautilus, vol. 56, pp. 76, 77, pl. 7, fig. 7, 
1943) and is also reported from St. Croix, Virgin Islands (Nowell-Usticke, A Check List 
of the Marine Shells of St. Croix, p. 63, 1959), from off the coast of Texas in 2431 fathoms 
(Parker and Curray, Bull. Amer. Assoc. Petrol. Geol., vol. 40, p. 2433, pl. 1, fig. 7, 1996), 
and from Bocas Island, east Panama (Olsson and McGinty, Bull. Amer. Paleont., vol. 39, 
p. 15, 1958). The western Atlantic form may prove to be merely a subspecies of M. lugubris. 


362 SAN Deco Society oF Naturat History {Vot. 13 


Family CANCELLARIIDAE 
Cancellaria (Aphera) wigginsi, new species 


Figures 5d, e 


Holotype. — No. 12607, California Academy of Sciences Department of Geology type 
collection, from Loc. 38554 (CAS), west side of Isla Monserrate, Baja California, Mexico, 
in the Gulf of California; Pleistocene; Ira L. Wiggins, collector, April 8, 1962. 


Description. — Shell ovate, six rounded whorls, the body whorl much larger than the 
spire; the first two and one half nuclear whorls smooth, the third, fourth and fifth whorls 
sculptured with seven or eight fine concentric riblets gradually increasing in coarseness, the 
body whorl bearing 25 or 26 spirals which are crossed by longitudinal, raised lines of growth 
which lend a pitted appearance to the sculpture; aperture ovate, inner margin of outer lip bear- 
ing about 12 denticles, the third posterior one the largest, which extend inward as short lirae; 
inner lip covered with a broad, thick callus, the upper portion of which is very faintly longi- 
tudinally grooved; columella with two well developed plications, the upper one the larger. 


Measurements of type.— Length, 26.7 mm.; maximum diameter, 14.2 mm.; length of 
body whorl, 19.5 mm. 


Remarks. — This new species bears a resemblance to species described from strata of 
Miocene age in the Caribbean region and in Peru. Cancellaria (Aphera) wigginsi differs 
from C. (A.) islacolonis Maury (Bull. Amer. Paleont., vol. 5, p. 229 [65], pl. 36 [10], 
figs. 12, 12a, 12b, 1917) described from Miocene beds in Santo Domingo, in the more 
slender outline, the thicker callus on the body whorl, the lack of grooving on the columellar 
plications, and in the absence of a denticle on the upper portion of the columellar wall. 
Cancellaria (A.) ellipsis Pilsbry (Proc. Acad. Nat. Sci. Phila., vol. 73, p. 333, pl. 22, figs. 
8, 9, 1922), a small form 12 mm. in length, also described from Miocene beds in Santo 
Domingo, may be a juvenile form of the species described by Maury, as mentioned by Pilsbry. 


The shell characters separating C. (A.) wigginsi from C. (A.) islacolonis also serve to 
separate it from C. (A.) peruana Nelson (Trans. Conn. Acad. Sci., vol. 2, no. 1, p. 190, 
pl. 6, fig. 3, 1870; Spieker, Johns Hopkins Univ. Stud. Geol., no. 3, p. 42, pl. 4, fig. 13, 


1922), which also has a more globose form and coarser sculpture. 


Cancellaria (Aphera) tessellata Sowerby (Proc. Zool. Soc. London, p. 51, June, 1832; 
Conch. Illustr., Cat. p. 3, January 11, 1833, pl. 11, figs. 20, 20*, December 21, 1832), 
living in tropical west American waters, is very much more slender in outline in comparison 
to C. (A.) wigginsi and to its Miocene relatives. 


A second Recent species of the genus Cancellaria and the subgenus A phera occurs in 
the Panamic faunal province, namely C. (A.) oblonga Sowerby, 1825. 


We take pleasure in naming this species in honor of Dr. Ira L. Wiggins, Professor of 
Biology, Emeritus, Stanford University. 


Fig. 5. ac, Dendraster casseli Grant and Hertlein, 1938. F-11 (SD-BE), Isla San José, Pliocene. 
a, greater diameter 53.5 mm., lesser diameter (anterior-posterior) 52.1 mm., abactinal view; b, greater 
diameter (anterior-posterior) 44.6 mm., lesser diameter 43 mm., actinal view; c, greater diameter (anterior- 
posterior) 36 mm., lesser diameter 35 mm., abactinal view. 

d, e, Cancellaria (Aphera) wigginsi, new species. 38554 (CAS), Isla Monserrate, Pleistocene, holo- 
See 26.7 mm., maximum diameter 14.2 mm. d, dorsal view; e, apertural view. (Photos by A. G. 
mith ). 

f, Dendraster cf. D. granti Durham, 1950, Isla San José, lesser diameter 68.5 mm., greater diameter 
72.5 mm., abactinal view. 
ae! Gyrineum (Bechtelia) strongi Jordan, 1936. F-10 (SD-BE), Isla Monserrate, Pleistocene, height 
ay mm. (spire incomplete), maximum diameter 29 mm., apertural view. Type species of Bechtelia, new 
subgenus. 


1964 } EMERSON AND HERTLEIN: INVERTEBRATE MEGAFOSSILS 363 


364 San Dreco Society oF NaturaL History [ VoL. 13 


Family CONIDAE 
Conus (Chelyconus) patricius Hinds, 1843 


Conus patricius Hinds, Ann. Mag. Nat. Hist., new ser., vol. 11, p. 256, April, 1843. “Gulf of 
Nicoya. Central America . . . in 7 fathoms”.— Hertlein and Strong, Bull. Amer. Mus. Nat. Hist., vol. 
107, p. 218, November 28, 1955. Pifias Bay, Panama; Ardita Bay and Bahia Cuevita, Colombia. — Hertlein 
in Brandt, Coastal Study of southwest Mexico, pt. 2, p. 248, Dept. Geogr. Univ. Texas, 1958. Coast 
of Michoacan. — Shasky, Minutes Conchol. Club, South. Calif., no. 186, March, 1959. West Mexico. — 
Hanna, Occas. Papers, Calif. Acad. Sci., no. 35, pp. 49, 51, pl. 6, figs. 1-10; pl. 8, fig. 1; pl. 10, figs. 
4, 5, January 28, 1963. Michoacan, Mexico, south to Punta Carnero, Ecuador. Recent. 

Conus pyriformis Reeve, Conch. Icon., vol. 1, Conus pl. 13, fig. 70, May, 1843. “Bays of Caraccas 
and Montija, West Columbia.” 

Conus (Dendroconus) patricius Hinds, Keen, Sea Shells of Tropical West America (Stanford Univ. 
Press: Stanford, California), p. 484, fig. 934, 1958. Nicaragua south to Ecuador. Recent. 


Range. — Pleistocene to Recent. Recent from Michoacan, Mexico, to Punta Carnero, 
Ecuador. 

Remarks. — One beautifully preserved specimen, measuring nearly 90 mm. in length, was 
collected in the richly fossiliferous Pleistocene sands at Isla Monserrate. Although this 
species is known from the Pleistocene of Ecuador (Rivera, 1953, Minutes Conchol. Club 
So. Calif., no. 129, p. 14), this appears to be the first fossil record from Mexico for this 
species, which is not known to be living at the present time in the Gulf of California. 

Recently cited by Keen (1958) to range southward from Nicaragua, this species is now 
known to occur in Mexican waters. Hertlein (1958) and Hanna (1963) report it to be 
living as far north as the Mexican State of Michoacan, and Shasky (1959) records a speci- 
men from an unknown locality along the west Mexican coast. In 1960, Mr. Frederick D. 
Sibley collected living specimens at Revolcadero Beach, near Acapulco, Guerrero, Mexico 


(AMNH collection). 


Phylum ECHINODERMATA 
Class ECHINOIDEA 
Family CLYPEASTRIDAE 
Clypeaster bowersi Weaver, 1908 


Clypeaster bowersi Weaver, Univ. Calif. Publ. Bull. Dept. Geol., vol. 5, no. 17, p. 271, pl. 21, fig. 1; 
pl. 22, fig. 1, 1908. “Colorado Desert.” Presumed to be of Miocene age. — Kew, Univ. Calif. Publ. Bull. 
Dept. Geol., vol. 8, no. 5, p. 50, pls. 4 and 5, April 16, 1914. “Lower Division of the Carrizo formation 
at Coyote Mountain.” — Clark and Twitchell, U.S. Geol. Surv., Monogr. 54, p. 209, pl. 96, figs. la and 
lb, 1915. “Colorado Desert (Weaver). — Kew, Univ. Calif. Publ. Bull. Dept. Geol., vol. 12, no. 2, p. 58, 
pl. 5, figs. la and 1b; pl. 6, fig. 1, 1920. “Coyote Mountain, Imperial County, California.” “Lower Divi- 
sion of the Carrizo Creek beds, Pliocene.” —G D. Hanna, Proc. Calif. Acad. Sci., Fourth Ser., vol. 14, 
no. 18, p. 479, 1926. Several localities cited from Coyote Mountain, Imperial County, California and 
neighboring region, Pliocene. — Grant and Hertlein, Univ. Calif. Los Angeles Math. Phys. Sci., vol. 2, 
p. 43, 1938. Earlier records cited. Pliocene. — Shimer and Shrock, Index Fossils of North America (John 
Wiley & Sons, Inc.: New York), p. 221, pl. 84, figs. 17, 18, 1944. (Copies of Kew’s illustrations, 1920). 
“Plioc.: Cal. (Carrizo); L. Cal., Mex.” —Durham, Geol. Soc. America, Mem. 43, pt. 2, p. 40, pl. 42, 
fig. 9; pl. 46, figs. 1, 2, 1950. “Lower? Pliocene from mainland southeast of San Marcos Island, some 
distance west of Chivato Point.” Also Santa Antonita Point, Baja California. Pliocene. 

Clypeaster testudinalis Gray, Hanna and Hertlein, Proc. Calif. Acad. Sci., Fourth ser., vol. 16, no. 6, 
pp- 140, 145, 153 (as Clypeaster testudinarius), April 22, 1927. One half mile back from shore at Santa 
Antonita, Baja California; Pliocene. 

Not Echinanthus testudinarius Gray, Proc. Zool. Soc. London for 1851, p. 35. “Hab. Indian Ocean; 
Borneo.” (See also Hertlein and Grant, 1938, p. 43, footnote 47). 

Range. — Early (Durham) and Mid-Pliocene of the Gulf of California region and 
Imperial County, California. 

Remarks. —Two specimens of a huge Clypeaster were collected by Ira Wiggins from 
beds of Pliocene age on Isla Cerralvo. One of these is 168.2 mm. long. The petals on one 
side of the abactinal surface are visible, and these as well as other observable shell characters 
agree with those of specimens of Clypeaster bowersi Weaver from Imperial County, California, 
and others from Baja California. Another large, but crushed, specimen from the same locality 


is imperfectly preserved; it is, however, apparently referable to C. bowersi. 


1964 } EMERSON AND HERTLEIN: INVERTEBRATE M: S 36 


5 

One specimen in the present collection, collected by Emerson at E] Mostrador on Isla 

Cetralvo F-13 (SD-BE), is partly covered with matrix, but the shape and observable shell 
characters agree well with those of C. bowersi. 

The record of Clypeaster bowersi Weaver by Royo Gomez (Comp. Estud. Geol 


Colombia, Tomo 5, p. 469, 1942) from beds of late Tertiary age in Colombia is probably 
referable to a similar but different species. 


Family SCUTELLIDAE 
Dendraster casseli Grant and Hertlein, 1938 


Figures 5a-c 


Dendraster casseli Grant & Hertlein, Univ. Calif. Publ. Math. and Phys. Sci., vol. 2, p. 81, pl. 1, 
figs. 1, 2; pl. 30, fig. 3, April 19, 1938. From “Loc. 356 (UCLA), at head of ravine in SE. corner of 
Sec. 10, T. 4N., R. 17W., S.B.B. & M., about 0.2 mi. N. of corner, in the Dendraster bed, elevation 
1300 feet by contour (U.S. Geol. Surv. Topog. Map of Santa Susana Quadrangle).” —Hertlein and 
Grant, Mem. San Diego Soc. Nat. Hist., vol. 2, pt. 2a, p. 120, pl. 27, fig. 7 (San Diego formation, 
middle Pliocene), July 7, 1960. Earlier record (1938) from San José Island cited. 

Range. — Mid-Pliocene, southern California, and Isla San José, Gulf of California. 

Remarks. — More than 40 specimens ranging from 11 to 53 mm. in greatest diameter 
were collected on Isla San José. This species was previously reported from strata of Pliocene 
age on this island by Grant and Hertlein (1938:82). 

The apical system of this species is only slightly excentric and the ambulacral petals are 
long, nearly straight, narrow, open at the ends, and raised above the general surface of the test. 


Dendraster cf. D. granti Durham, 1950 


Figure 5f 


{?}] Dendraster granti Durham, Geol. Soc. America, Mem. 43, pt. 2, p. 41, pl. 47, figs. 3-12, August 
10, 1950. From Loc. A 3559 (Univ. Calif.), “Middle (?) Pliocene, Arroyo de Arce, north of Arroyo de 
Gua, Lower California. In cliff along north side of arroyo. Estimated to be about three quarters of a mile 
from beach. Stratigraphically higher than unconformity in section. In low-dipping alternating gravels and 
leached calcareous beds.” 


Range. — Mid-Pliocene (Durham), Baja California, Mexico and ?Isla San José, Gulf 
of California. 

Remarks. — The present record is based upon a Dendraster which is 68 mm. long (an- 
terior-posterior) and 72 mm. wide. The general characters of the test agree with those de- 
scribed for D. granti, and it occurs in beds of about the same age. 

The type specimen of Dendraster granti is only 35.7 mm. long and 37 mm. wide. It was 
compared with D. vizcainoensis Grant and Hertlein (1938, p. 90, pl. 8, figs. 1, 2, and 3), 
a species described from a terrace deposit of Quaternary age at Punta Santa Rosalia in Bahia 
de Sebastian Vizcaino, Baja California. The test of the present specimen is raised centrally, 
and the petals are elevated above the surface of the test, which bears large, rather widely 
spaced tubercles. 

The petals of D. granti were described as “ovate, almost parallel-sided.” The illustrations 
of it reveal variation in the curvature of the sides of the petals. The sides of the petals of 
the bivium in the present fossil are decidedly curved (fig. 5f). It seems probable that this 
specimen represents a large form of D. grant. is 

The present specimen also bears a decided resemblance to Dendraster vizcainoensis 
similaris Grant and Hertlein (1938, p. 90, pl. 27, figs. 1, 2; pl. 28, fig. 8), described from 
beds of late Pleistocene age in the vicinity of Los Cerritos Station, near Signal Hill, Long 
Beach, California. The type specimen is 88 mm. long and 93 mm. wide. The petals of the 
bivium of that subspecies are a little more ovate and the test is more elevated centrally than 
those of present specimen from Isla San José, but otherwise the resemblance is remarkable. 


Encope californica Verrill, 1870 


Figures 6a-e 


Encope californica Verrill, Amer. Jour. Sci., Ser. 2, vol. 49, p. 97, January, 1870. bag ied os 
“Cape St. Lucas”, “Gulf of California.” Recent. — Grant and Hertlein, Univ. Calif. Publ. Math. an 


366 San Drieco Society oF Natura History { VoL. 13 


Fig. 6. a-e, Encope californica Verrill, 1870. F-10 (SD-BE), Isla Monserrate, Pleistocene. a-c, height 
(anterior-posterior) 97 mm., width 98.5 mm.; a, abactinal view; b, side view; c, actinal view. d, e, height 
95 mm., width 93 mm.; d, abactinal view; e, side view. 

f-i, Encope grandis L. Agassiz, 1841. F-10 (SD-BE), Isla Monserrate, Pleistocene. f, g, height 


(anterior-posterier ) 95 mm., width 93 mm.; f, abactinal view; g, actinal view; h, i, height (anterior-posterior ) 
99.5 mm., width 96.3 mm.; h, side view; i, abactinal view. 


1964 } EMERSON AND HERrTLEIN: INVERTEBRATE MEGAFOSSILS 367 


Phys. Sci., vol. 2, p. 96, pl. 11, fig. 4, pl. 30, fig. 2, April 19, 1938. | about Ye mile southwest 
of the eastern tip of} Punta Santa Ynez, Gulf coast of Lower California, Mexico: Pleistocene. — Durham. 
Geol. Soc. America, Mem. 43, pt. 2, p. 44, pl. 37, fig. 2; pl. 38, figs. 1, 5 st 10, 1950. Range 
lower Pliocene to Recent. 

Range. — Early Pliocene (Durham) to Recent. Recent from San Luis Gonzaga Bay, 


Baja California, to La Paz, Baja California, Mexico. 


Remarks. — This echinoid occurs frequently in beds of Pliocene and Pleistocene age in 
the Gulf of California region. The test is highest posteriorly at the anterior end of the inter- 
ambulacral lunule. The margin of this species is much thicker and the lunules are closed in 
contrast to the similar Encope carrizoensis Kew, which was originally described from strata 
of Pliocene age in Imperial County, California. The test of Encope grandis Agassiz is highest 
posteriorly, like that of E. californica, but the margin is much thicker, the marginal lunules 
are open, and the interambulacral lunule is much larger (see fig. 6). 


A. H. Clark (Smithson. Miscell. Coll., vol. 106, no. 5, p. 6, 1946) believed Encope 
californica to be “a localized race confined to the Gulf of California” and H. L. Clark (Allan 
Hancock Pac. Exped., vol. 8, no. 5, p. 329, 1948), placed it in the synonymy of Encope 
micropora Agassiz, which was originally described without a type locality. Durham (1950) 
considered E. californica to be a valid species, and Mortensen (Mon. Echin., IV. 2, Clypeas- 
troidea, p. 445, 1948) recognized it as a subspecies or variety of E. micropora. 


Literature Cited! 


ANONYMOUS 
1924. Informe sobre la exploracién geoldgica de la Baja California, por la Marland 
Oil Company of Mexico. Bol. del Petroleo, vol. 17, no. 6, pp. 417-453, pls. 1, 
4-12, 14-22, 26-46, June; vol. 18, no. 1, pp. 14-53, pls. 77-92, 1 map, July. 


ANDERSON, C. A. 
1950. 1940 E. W. Scripps cruise to the Gulf of California. Pt. 1. Geology of islands 
and neighboring land areas. Mem. Geol. Soc. Amer., no. 43. 53 pp. 


Bear, Goi. 
1948. Reconnaissance of the geology and oil possibilities of Baja California, Mexico. 
Mem. Geol. Soc. Amer., no. 31. x + 138 pp., 11 pls. 


DuruaM, J. W. 
1950. 1940 E. W. Scripps cruise to the Gulf of California. Pt. 2. Megascopic paleon- 
tology and marine stratigraphy. Mem. Geol. Soc. Amer., no. 43. viti 4- 216 pp. 
DuruaM, J. W., AND E. C. ALLISON . 
1960. The geologic history of Baja California and its marine faunas. in The biogeo- 
graphy of Baja California and adjacent seas. Pt. 1. Geologic history. Syst. Zool. 
9:47-91. 


DuruaM, 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 16:1-110. 


Emerson, W. K. . . S 
1960a. Results of the Puritan-American Museum of Natural History expedition to 
western Mexico. 11. Pleistocene invertebrates from Ceralvo Island. Amer. Mus. 


Novitates no. 1995. 6 pp. . ee 
1960b. Results of the Puritan-American Museum of Natural History Expedition to 
western Mexico. 12. Shell middens of San José Island. Amer. Mus. Novitates 


no. 2013. 9 pp. 


1Excluding references cited in the Systematic Treatment section 


368 San Dieco Society oF Natura History [Vors13 


Emerson, W. K., AND M. K. JAcoBsoN 
1964. Terrestrial mollusks of the Belvedere expedition to the Gulf of California. Trans. 
San Diego Soc. Nat. Hist. 13:313-332. 


GREWINGK, C. 
1848. Beitrag zur Kenntniss der Geognostischen Beschaffenheit Californiens. Ver- 
handlungen der Russisch-Kaiserlichen Mineralog. Gesellschaft zu St. Petersburg, 
“1847”, pp. 143-162. 


Grant, U. S., IV, AND L. G. HERTLEIN 
1938. The west American Cenozoic Echinoidea. Publs. Univ. Calif. Los Angeles in 
Math. and Phys. Sci., Vol. 2. vi ++ 226 pp. 


Hanna, G D. 
1926. Paleontology of Coyote Mountain, Imperial County, California. Proc. Calif. 
Acad. Sci., ser. 4, 14:427-503. 


Hanna, G D., ANp L. G. HERTLEIN 
1927. Expedition of the California Academy of Sciences to the Gulf of California in 
1921. Geology and Paleontology. Proc. Calif. Acad. Sci., ser. 4, 16:137-157, pl. 5. 


HErTLEIN, L. G. 
1957. Pliocene and Pleistocene fossils from the southern portion of the Gulf of Cali- 
fornia. Bull. So. Calif. Acad. Sci. 56:57-75, pl. 13. 


Her TLEIN, L. G., AND W. K. EMERSON 
1956. Marine Pleistocene invertebrates from near Puerto Penasco, Sonora, Mexico. 
Trans. San Diego Soc. Nat. Hist. 12:154-176. 
1959. Results of the Puritan-American Museum of Natural History expedition to 
western Mexico. 5. Pliocene and Pleistocene megafossils from the Tres Marias 


Islands. Amer. Mus. Novitates no. 1940. 15 pp. 


Linpsay, G. E. 
1962. The Belvedere expedition to the Gulf of California. Trans. San Diego Soc. Nat. 
Hist. 13:1-44. 


Saumes, D. F. 
1959. Results of the Puritan-American Museum of Natural History expedition to 
western Mexico. 7. Corals and coral reefs in the Gulf of California. Bull. Amer. 
Mus. Nat. Hist. 118:367-432, pls. 28-34. 


TARBET, L. A., AND W. H. HoLMAN 
1944. Stratigraphy and micropaleontology of the west side of Imperial Valley, Cali- 
fornia. Bull. Amer. Assoc. Petrol. Geol. 28:1781-1782 { Abstract}. 


Witson, I. F. 
1948. Buried topography, initial structures, and sedimentation in Santa Rosalia area, 


Baja California, Mexico. Bull. Amer. Assoc. Petrol. Geol. 32:1762-1807. 


Wison, I. F. {in collaboration with V. S. RocHa} 
1955. Geology and mineral deposits of the Boleo Copper District, Baja California, 
Mexico. U. S. Geol. Surv. Prof. Paper 273. vi + 134 pp., 11 pls. 


TRANSACTIONS 
OF THE 
SAN DIEGO SOCIETY OF NATURAL HISTORY 


VoLuME 13, No. 18, pp. 369-376 


THE RACES OF HALIOTIS FULGENS PHILIPPI 
(MOLLUSCA: GASTROPODA) 


BY 


Ropert R. TALMADGE 
Willow Creek, California 


SAN DIEGO, CALIFORNIA 
PRINTED FOR THE SOCIETY 


DECEMBER 30, 1964 


IAN 1.5 196! 
THE RACES OF HALIOTIS FULGENS PHILIPPY 2” 


(MOLLUSCA: GASTROPODA ) HARVARD 
UNIVERSITY 
BY 


RoBerT R. TaLMADGE 


For a number of years there has been some confusion among malacologists as to the status 
of certain populations of Haliotis fulgens Philippi 1845. Relatively few names, each rather well 
defined, have been applied to this species, and there is no question concerning the synonymy 
and priority of these. However, some malacologists declare that there is a single form with 
only minor local variations, while others recognize two geographical races, one of which appears 
to be based upon features that can be found in almost any population of H. fulgens. It is 
probable that much of this difference of opinion is based on the examination of a rather small 
number of specimens, as few collections have large series of shells from various localities. Be- 
cause of limitations of space, the species is usually represented only by a growth series, with 
perhaps a few odd items. 


Perhaps the largest and most complete collection of Haliotis fulgens is in the San Diego 
Natural History Museum. Mr. Emery P. Chace, Curator of Conchology at that institution, 
kindly made available to me their comprehensive collection of this species, taken in many 
localities, some of which are represented in few collections. From a study of these, augmented 
by examination of most of the major malacological collections in the United States, I think 
a clearer understanding of this species and its variations may be obtained. 


VARIATION AND DISTRIBUTION 


Like other haliotids, this species exhibits definite variation related to age. Reeve (1846) 
figured and described a juvenile shell as Haliotis planilirata, the type of which is in the British 
Museum (Natural History). Juvenile specimens are usually more elongate than adults; they 
lack the muscle scar, and have a more or less pure silvery nacre. The muscle scar, as well as 
the brilliant green, pink, and blue coloration, starts to develop in shells of approximately 100 
to 125 mm. in length, and this development continues until the animals reach maturity. As 
the animal becomes older the shell deepens and becomes more rounded, with a dark blue colora- 
tion becoming predominant. Dark stains, probably of organic origin, cover portions of the 
columellar plate and interior, and the muscle scar becomes larger and rugose. Such old speci- 
mens are usually so eroded, by the action of borers and by wear, that most of the sculpturing 
is lost. The number of open siphonal pores usually decreases; one specimen examined is totally 
imperforate. 


At present the species appears to be restricted to the warmer waters of the Pacific coast 
of North America, from the vicinity of Magdelena Bay, Baja California (approximately 
24° N), north to Point Conception, California (approximately 34° N), including the offshore 
islands. There are some references to the species at Cape San Lucas, Baja California, but | 
have not been able to verify these occurrences. North of Point Conception, the species is 
known from several specimen records. There is a set of juvenile shells in the collection of the 
California Academy of Sciences taken from near Monterey, California. Mr. Keith Cox (pers. 
comm.) has also noted juveniles in that area. I have one in my collection from near Carmel, 


California. Keep (1935) refers to Haliotis fulgens from Monterey. 


There is a story told among the abalone divers that in past years the Japanese divers 
operating out of Monterey, who often worked as far south as the Channel Islands, attempted 
to establish this species in the north by planting the animals as they returned home. If this is 
true, it could explain the northern specimens; perhaps the plantings did not prosper, but a few 
individuals were able to survive in isolated spots, producing the juveniles that we find today. 


372 San Disco Society oF Natura History {Vor13 


Fossit RECORD 
The family Haliotidae is poorly represented by fossil specimens. Perhaps one of the 
earliest species is Haliotis lomaensis Anderson, late Cretaceous, from San Diego, California. 
This single specimen, in the California Academy of Sciences, is a well developed haliotid 
comparable to Recent species. 


Based upon shape and sculpturing there are two fossil examples which could be considered 
to pertain to Haliotis fulgens. Woodring described Haliotis lasia from the Miocene of San 
Luis Obispo County, California. The site is not far north of the main part of the present 
range. With at least one paratype, the type is in the United States National Museum 
(no. 371767). Another similar, if not identical, species, Haliotis elsmerensis Voke, is repre- 
sented by no. 32465 in the Museum of Invertebrate Paleontology, University of California, 
Berkeley. This is referred to the Pliocene of Los Angeles County, California. The locality is 
well within the range of the Recent species. 

After examination of both of these type specimens, I am of the opinion that they repre- 
sent the same species; both are either Haliotis fulgens or they are an early form of the complex 
which contains H. fulgens, H. walallensis and the European H. tuberculata. 


Hertlein (1957) recorded Haliotis fulgens at Punta Pulpito in the Gulf of California, 
dating the deposits as Pleistocene. As far as is known, the species does not inhabit that area 
today. Other paleontologists working with the elevated marine terraces of Pleistocene age have 
found the species along the coast of the Californias well within the range of the Recent form. 


METHOD 


In an attempt to evaluate the living populations, I had available for study, over a period 
of time, several hundred examples. The locality of each of these was plotted on a large scale 
map. A work sheet was prepared covering the nacre coloration, number and type of open 
siphonal pores, measurements (length, width, height, height of spire) and ratios, and variation 
in the sculpturing. Soft parts were compared, especially the formation of the epipodium, the 
fleshy girdle which encircles the muscular foot. 


Comparison of the soft parts indicated that there was a definite, persistent similarity, 
enough to establish that a single species was present. A study of the shells revealed non-clinal 
variation that could be used to separate populations from certain regions. 


The coloration of the nacre and the number of open pores were so similar in all age 
groups, and in shells from all localities, that separation on this basis was impractical. It was 
found that the ratios of certain measurements of the shell from at least two areas indicated a 
definite separation, and when combined with the sculpturing furnished evidence of two distinct 
geographical races. When sculpturing alone was considered, it was noted that there were three 
definite types, two of which merged. As the forms of sculpturing were plotted on a map, it 
was noted that they fell into distinct geographical areas, and the merging type of sculpturing 
was found to be at the point of contact of two such regions. Talmadge (1962, 1963) 
indicated that this was as good a diagnostic feature as was available while working on popula- 
tions of Haliotis varia Linnaeus, and these same characteristics proved useful in an attempt to 
evaluate the populations of H. fulgens. 


TAXONOMY 
At the present time, the variations in the sculpturing indicate two distinct populations on 
the mainland coast of the Californias, and the nearby offshore islands, with a third separate 
population, differing in both sculpturing and ratios, on Guadalupe Island. Based on this data, 
I recognize the following races of Haliotis fulgens. 


Haliotis fulgens Philippi 
Range. — Magdelena Bay, Baja California, north to Monterey, California, including the 


offshore islands. 


1964 } TALMADGE: Races oF Haliotis fulgen: 373 


Aaliotis fulgens fulgens Philippi 
Haliotis fulgens Philippi, 1845, Zeits. f£. Malak. 2, p. 150 
Haliotis splendens Reeve, 1846, Conch. Icon. 3, sp. 9 
Haliotis fulgens Philippi, 1847, Abbil. u. Besch., Tab. VIII, fig. Ia 
Aaliotis fulgens Philippi, Cox, 1962, Fish. Bull. 118, p. 32 

Type. — Lost ? Oldroyd (1927:234) gives the location of the type as “Philippi Collec- 
tion, Chile.” 

Type locality. — Philippi (1845) gave only “Patria”, but in 1847 used “Patria Califor 
nia.” Reeve (1846) used “Hab. California” for H. splendens and “Hab.?” for H. planilirata. 

Remarks. — This is the form which has been best figured and of which specimens are 
found in most collections. It is one of the commercially taken abalone in southern California 
and northern Baja California, where it is known as either Green or Blue Abalone. It is a 
medium elevated, ovate shell when adult, with a rather large, brilliant muscle scar, either 
greenish or bluish. The interior nacre is shaded with hues of pink, green, and blue, which 
become dark and brilliant with age. Very old specimens show dark stains and a rather dark 
nacreous interior. The sculpturing consists of numerous, rather irregular, rounded small cords. 
The siphonal angle is rounded and the five to seven open pores are on small, slightly elevated 
projections. 

This form is found from Point Conception south to Cedros Island. There are some color 
phases, usually in Mexican waters. In the vicinity of Punta Banda, Ensenada, and the Todos 
Santos Islands, Baja California, there is a strong tendency for specimens to have a yellowish 
or golden wash over the interior nacre. Near Corona del Mar, California, specimens are a 
rather pale green, with white or silvery nacre. In these areas the shape, sculpturing, and size 
of the shell are similar to those features in other regions within the range of the race; the 
differences noted can only be referred to as color phases. 


Haliotis fulgens turveri Bartsch 


Haliotis fulgens turveri Bartsch, 1942, Nautilus 56 (2), p. 57 

Type. — No. 508764, United States National Museum. 

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

Remarks. — Unfortunately, Bartsch used a badly worn, senile shell as his type. A desig- 
nated paratype in the Turver Collection is extremely similar. Both of these shells are deep 
and rounded, have dark blue nacre, and have five open pores. The cording is eroded away 
except for a slight area on the right or growing edge of the shell, but sufficient remains to 
establish the sculpturing form. 

Twenty examples were examined from Magdelena Bay; it was found that 18 had extremely 
wide, flat, irregular cording, which could be separated easily from that of the nominate race 
without detailed comparison. The other two shells had wide irregular cording which was still 
separable, but not as distinctly so. From Turtle Bay, Baja California, 18 of 25 examples were 
identical to Magdelena Bay specimens. The remaining 7 were similar, but had a more uniform 
and narrower cording, which probably indicates genetic contact with the more northern 
populations of the nominate race. 

Thirty specimens were examined from Cedros Island, and it was noted that 20 were 
sculptured with the narrow, rounded, irregular cording of H. fulgens fulgens, five were like 
H. f. turveri, and the remaining five were intermediate, fitting neither race yet similar to both. 

Bartsch’s description, without a figure, appears to be based on a worn senile example 
(examination confirms this), which could fit almost any population of Haliotis fulgens. How- 
ever, the sculpturing noted and discussed above is consistent, Is noticeable without undue — 
parison of specimens, and is restricted to a distinct geographical region. Hence, I believe t = 
there is a poorly defined geographic race of H. fulgens inhabiting the southern portion of the 
range of the species, from Magdelena Bay north to Turtle Bay, Baja California, Mexico. 


37/4 


San Deco Society oF Natura History 


[ VOL. 


Fig. 1. Holotype of Haliotis fulgens guadalupensis. Photos by Mead French. 


13 


1964 } TALMADGE: RAcEs OF Haliotis fulgens 375 


Haliotis fulgens guadalupensis, subsp. nov. 

Type. — No. H-1-30-60, San Diego Natural History Museum (fig. 1). 

Type locality. — Collected at South Tip, Morro Sur, Guadalupe Island, Baja California, 
Mexico. All specimens examined are from this one island. They were collected by divers 
Robert Clutter, Pete Taylor, and Carl Boyd, all of Scripps Institution of Oceanography, 
January 30, 1960. 


Description. — The shell is auriform; the apex of the spire is inset approximately one 
sixth of the major diameter of the shell and the spire is not elevated. Both the apex of the 
shell and the dorsal surface are on about the same plane. The siphonal angle is not acute; 
rather, the shell tends to bend gracefully over this region. There is a shallow, nearly obsolete 
groove below the siphonal angle. There are seven small open pores on low projections on the 
type, but in the series examined the number of open pores varies from five to seven. The shell 
is proportionately deeper and wider than in the other two races. The dorsal sculpturing is dis- 
tinct; the cording is remarkably uniform and more or less equal in width and height, separated 
by interspaces of equal width and depth. Close observation shows that the interspaces contain 
a second set of rather uniform cords, well below the general surface of the shell. 


The exterior coloration is dull reddish-brown, but in most specimens this is somewhat 
obscured by a growth of hard, red sponge. The interior nacre is pinkish, with green and blue 
iridescent tints. The rather large muscle scar is swirled in pinks and greens. The type is a 
nearly adult shell, chosen to best illustrate the very distinctive cording. 


Remarks. — Although some specimens taken in deeper water reach well into the range of 
size of the nominate race, the average shell is much smaller. Specimens as small as 115 mm. 
in length exhibit well developed muscle scars, whereas mainland specimens of this size would 
have only a trace of this scar. 


Based upon a formula of length equals 1000, the following ratios are noted: mainland 
specimens, length 1000, width 744, elevation 219; Guadalupe Island specimens, length 1000, 
width 810, elevation 293. A comparison of the Guadalupe Island population shows that 87.5% 
were virtually identical to the type. The remaining 12.5% had variations in the cording, but 
could be separated easily from either of the two mainland races. Haliotis fulgens guadalupensis 
appears to be an isolated, dwarfed, endemic race of the species. Another abalone, Haliotis 
cracherodii californiensis Swainson, also appears to be endemic to this island. 


ACKNOWLEDGMENTS 


I wish to express my appreciation to Mr. Emery P. Chace and the San Diego Natural 
History Museum for making the comparative material available. The scientific library of the 
California Academy of Sciences was opened to me, and Dr. Leo G. Hertlein of that institu- 
tion made special efforts to assist in checking certain aspects of the literature. Mr. S. P. Dance, 
British Museum (Natural History) examined and compared the type of Haliotis planilirata 
for me. The opportunity to examine material in the U.S. National Museum was made pos- 


sible by a Grant-in-Aid, No. P2958, from the American Philosophical Society of Philadelphia. 


LITERATURE CITED 
BartscH, Paul 
1942. A new subspecies of Haliotis (H. fulgens turver:). Nautilus 56:57. 
Cox, Keith W. 
1962. California abalones, family Haliotidae. Fish Bull. 118, Calif. Dept. Fish and 
Game. 133 pp. Sacramento. 
HErTLEIN, Leo G. ; er ee 
1957. Pliocene and Pleistocene fossils from the southern portion of the Gulf of Cali- 
fornia. Bull. So. Calif. Acad. Sci. 56:57-75. 
Keep, Josiah (Revised by Joshua Bailey, Jr.) 
1935. West Coast Shells. Stanford Univ. Press. xii + 350 pp. 


376 San DreGo Society OF NATURAL HIstTory [Vornls 


Ovproyp, Ida Shepard 
1927. The Marine Shells of the West Coast of North America. Vol. 2, part 3, pp. 
605-941, pl. 73-108. Stanford Univ. Press. 


Puiiprp!, R. A. 
1845. Diagnoses Testaceorum quorundam novarum. Zeits. fur Malak. 2:147-152. 


1847 (1845-1851). Abbildungen und Beschreibungen neuer oder wenig gekannter Con- 
chylien. Cassel. [Usually three volumes and about 144 color plates. Copy in 
the library of the California Academy of Sciences has been rebound into six 
volumes, four of text and two of plates. The separates covering H. fulgens are 


dated April, 1847. ] 


REEVE, Lovell A. 
1846. Monograph of the genus Haliotis. Conchologia Iconica 3. 17 color plates. London. 


TALMADGE, Robert R. 
1962. The Linnaean Haliotis varia in Australia. Mem. { Aust.} Nat. Mus., Melbourne, 
29:233-241e 
1963. Insular haliotids in the western Pacific (Mollusca:Gastropoda). Veliger 5:129- 


139, pl. 14. 


TRANSACTIONS 
OF THE 


SAN DIEGO SOCIETY OF NATURAL HISTORY 


VoLuME 13, No. 19, pp. 377-396 


ADDITIONS TO THE NUDIBRANCH FAUNA OF THE 
EAST PACIFIC AND THE GULF OF CALIFORNIA 


BY 


CLINTON L. COLLIER 


San Diego State College 
(Mailing address: 4374 Wilson Avenue, 
San Diego, California, 92104) 


AND 


WesLey M. FARMER 


San Diego Natural History Museum 


SAN DIEGO, CALIFORNIA 
PRINTED FOR THE SOCIETY 


DECEMBER 30, 1964 


ADDITIONS TO THE NUDIBRANCH FAUNA OF THE: 


EAST PACIFIC AND THE GULF OF CALIFORNIA | 5 j 


CLINTON L. CoLLIER AND WESLEY M. FARMER 


The coasts of the peninsula of Baja California represent an area where the nudibranchs 
have been little studied. Unlike the coast of California, most of Baja California is rugged, 
with difficult access. Consequently, when early workers were making extensive collections all 
along the Pacific coast they stopped when they reached San Diego and did not continue 
despite an equally rich nudibranch fauna to the south. When Marcus (1961) worked on a 
collection of California opisthobranchs he drew the only conclusion possible: new additions to 
the warm-temperate and tropical water fauna could be expected south of San Diego as soon 
as systematic collecting was done there. 


During the past few years the authors have made many trips along both coasts of Baja 
California to observe and collect the opisthobranch fauna of the region. We have found areas 
in which the animals are likely to occur and in which we may find more of the unusual 
forms. More importantly, we are able to obtain accurate color descriptions and measurements 
from the living animals and are not dependent on badly contracted specimens for descriptions. 
We have also made an effort to return living animals to San Diego so that they could be 
photographed. Such a procedure is of the utmost importance in the case of animals whose 
form and color are of such taxonomic significance. On extended trips, we took photographs in 


the field. 


We would like to express our deep gratitude to those who have helped us in the prepara- 
tion of this paper: Dr. Ernst Marcus and Miss Joan Steinberg answered many questions, Drs. 
Harold Rehder and Ronald P. Kenny sent specimens from the United States National Museum 
and from Australia, respectively, and Mr. James Lance gave suggestions and locality records. 
Mr. John Sloan of the San Diego Natural History Museum collected some of the specimens 


and Mrs. Rosemarie Fiebig, also of the museum, kindly translated some of the literature for us. 


Special thanks are due to Mrs. Mildred Le Compte, San Diego State College inter- 
library loan librarian for her untiring efforts in locating and obtaining literature. Without her 
assistance this paper would not have been possible. 


All types have been deposited in the California Academy of Sciences Invertebrate Type 
Collection. Some paratypes remain in the authors’ collections. We have followed the classi- 


fication of Odhner (1939) as modified by the works of Marcus. 


Collections were made at the following locations: 


California 

Newport! Bay,” Bos sAngeles iC ovtiaty nace care ce tee ete 33° 36’N_ 117° ~=54’°W 
Baja California, Pacific Side 

Jig Evie lx Srere | ee A ee EP thn, 28°. “O7N Eee ee 

Bala Sebastian ISCainO <2 oe ack epee Pas ecbeeene nace .27° 50’N 114° 51’W 
Baja California, Gulf Side 

Isla Angel de fa Guarda... ..----.nc--n-:neeoeeon--neeserenennennennaee 29° 33’/N_ 113° 35°W 


29° 49°N_ 114° 25’°W 


Se leven, Setar AHL m @foy oy 21: ee aeons oe 
30 25’N 114 40’W 


DD ITeLCECIEO Smee EI oe oe OE Suir an eased 3 


96 


BY HARVAR 


380 San Dreco Society oF NaturAL History { VoL. 13 


Order NUDIBRANCHIA 
Suborder DORIDACEA 
Section EUDORIDACEA 
Tribe CRYPTOBRANCHIA 
Family DORIDIDAE 
Subfamily GLOSSODORIDINAE 
Cadlina evelinae Marcus, 1958 
(Plate 1; fig. 1, A-B) 


Distribution. — The type of Cadlina evelinae is from the upper littoral of Ilhabela and 
Guaruja, Brazil (Marcus, 1958). In the present collection seven specimens are from a rocky 
sand flat at Coloradito, 15 miles north of Puertecitos, Baja California. One specimen was col- 
lected by John Sloan at Isla Angel de la Guarda. The species has also been recorded photo- 


graphically from Bahia Sebastian Viscaino, but specimens from that area have been lost. 


Description. — The entire animal is white to light cream except for the dorsal surface of 
the notum, which is speckled with numerous light red to orange spots. The spots are more 
numerous around the edge of the notum than in the middle. As reported by Marcus (1958:18) 
these spots are glandular and fade in alcohol, leaving a smooth, slightly raised area. Animals 
fixed in an FAA mixture retained the color of the glands. 


The body form is typical of animals of the subfamily Glossodoridinae (MacFarland, 
1905). The notum is uniformly broad along its entire length. In two specimens, however, it 
narrowed posteriorly. Of the seven animals collected at Coloradita the largest, while actively 
crawling, is 18 mm. long by 7 mm. wide. In two of the animals examined closely, the 
thinophores have 18 and 19 leaves on the clavi. Each animal has six bipinnate gills. The foot 
is bilabiate anteriorly. The oral tentacles are stout with an external fold. 


There is a strong labial cuticle which is heavily armed over most of its surface. The 
spines are 10 to 25 yu long and bifrucate at their distal ends into two short denticles (fig. 1, A). 
The radular formula for the largest animal is 78 by 65.1.65 while a smaller one has the formula 
of the type, 75 by 65.1.65. The rachidian tooth (fig. 1, B) has either four or five cusps. There 
seems to be more of the five-cuspidate teeth in the older section of the radula, whereas the 
area that has just been formed has rachidian teeth with only four cusps. The first lateral has 
three denticles on each side of the large middle cusp; some have only two denticles on the inner 
side. The next two laterals have four small outer denticles. The rest of the teeth out to the 
marginals have either two or, more commonly, three denticles on the outer side of the cusp. 
We noted that some were without any denticles, but this condition could be due to wear. The 
marginals are small and have five or six inner denticles; these outer teeth are often poorly 
formed. 


The highly variable radulae of the specimens at hand cast doubt on the validity of the 
idea that the radula can be separated within the genus. The number of denticles on both the 
rachidian and the first lateral varied from that described by Marcus, although teeth from 
different rows, if put together in one row, would fit his description. This also points to the 
variation of teeth within the one radula. Another discrepancy was in the size of the denticles. 
Marcus described the denticles of the laterals as fairly large, while in the present animals they 
were quite small and could only be seen by using a phase microscope. The same is true with 
the denticles of the first lateral. These variations in the teeth indicate that the radulae of other 
species of Cadlina should be re-examined before they are accepted as a valid taxonomic feature. 
The teeth with the most denticles are in the older part of the radula and indicate that the 
presence of fewer denticles is not necessarily a product of wear. The reproductive system is 


described by Marcus (1958:20-21). 


Discussion. — Marcus (1958) differeniated Cadlina evelinae from the other members of 
the genus listed by him in 1955. 


1964 } COLLIER AND Farmer: ADDITIONS TO Nupipr \NCH FAUNA 381 


Much emphasis has been placed on the radula of the members 
separation of the species. The species C. evelinae is undoubtedly valid 
from the other members of the genus by features other than the radula 
thinophores, and labial cuticles of our specimens agree completely 
the type. 

This animal brings the number of Cadlina recorded from the Pacific coast of North 
America to six. Cadlina evelinae can be differentiated from each of five listed by Lance 
(1962:157) by coloration and by other features. Cadlina evelinae is the only member of the 
genus recorded from the Gulf of California. 


Subfamily CONUALEVINAE, subfam. nov. 


The body is somewhat soft, lacking noticeable spicules, with the notum depressed and 
either minutely papillose or smooth. The rhinophores are retractile and have neither clavi nor 
perfoliations. The branchial aperture is round and without a ridge; gills vary from unipinnate 
to tripinnate. The anterior edge of the foot is bilabiate; the oral tentacles are short and stout. 
The teeth are edentulate and simply hooked, and the rachidian is absent. Two seminal re- 
cepticles are present. 

The type genus of the subfamily is Conualevia, gen. nov. 


of this genus for the 
and can be separated 
The coloration, gills, 
with the description of 


Genus Conualevia, gen. nov. 


The primary characteristic of the genus is the presence of smooth rhinophores. The notum 
may be smooth or minutely papillose. The oral tentacles are short, thick, and stout. The 
radula lacks a rachidian tooth or plate. The teeth on each side of the radula are offset at the 
rachidian space to form an interlocking pattern along the center line. There is either no labial 
cuticle or only a very light one. The retractile gills vary from unipinnate to tripinnate. The 
penis is unarmed. 

The female reproductive system is characterized by an X pattern at the end of a 
long vaginal duct. One leg of the X is the vaginal duct, one connects to the spermatocyst, one 
to the spermatheca, and the other is a short fertilization duct. This gives the female genital 
system a compact appearance. 

The type species of the genus is Conualevia marcusi, sp. nov. 


Conualevia marcusi, sp. nov. 
Plate 2; fig. 1, C-H 

Type. — The holotype (CASIZ 25), a whole animal, was collected 3.8 miles south of 
Puertecitos, Baja California, by the senior author in June, 1963. One paratype from the same 
locality is CASIZ 27 and one from Puerto Refugio on Isla Angel de la Guarda is CASIZ 33. 
A total of 9 animals was collected; others remain in the authors’ collections. 

Distribution. — The animals are known from one mile north of Puertecitos and from 
Puerto Refugio on Isla Angel de la Guarda, Baja California. They have been collected in 
March, June, and November. 

Description. — The animals are very light orange to white when living; all are white after 
preservation. The dorsal surface of the notum is closely papillose, with papillae a fraction of 
a millimeter long, giving the animal’s surface a fine textured appearance. The gills are con- 
colorous with the rest of the animal but appear slightly darker in the colored forms because 
they lack the softening effect which the papillae impart to the dorsal surface. 

The body form is like that of a typical dorid. The notum is high and extends over the 
foot (Plate 2). The preserved holotype measures 14.5 mm. long and 9 mm. wide. The 
retractile rhinophores are smooth (fig. 1, H) in the living animal although they appear annu- 
lated, due to contraction, in the preserved material. When only partly extended, the thinophores 
may also appear annulated in living specimens. The 16 unipinnate gills arranged in a circle 
around the anus are completely retractile into a branchial chamber which can be closed leaving 
only a small dot to mark the position of the anus and gill chamber. The foot is moderately 
large, but does not extend beyond the notum. The anterior margin of the foot is bilabiate 
(fig. 1, C). The oral tentacles are short and stout. 


382 San Dieco Society oF NaTurAL History { VoL. 13 


soace SOP 


Fig. 1. A-B, Cadlina evelinae. A, labial spines; B, partial row of radula. C-H, Conualevia 
marcusi. C, ventral view of preserved animal; D, diagram of the reproductive system, dorsal 
view; E, diagrammatic dorsal view of internal anatomy; F, radular tooth; G, Ist through 6th 
radular teeth; H, rhinophore. 


1, prostate; 2, vas deferens; 3, genital orifice; 4, nedimental opening; 5, vagina; 6, spermatocyst; 7, 
oviduct; 8, fertilization duct; 9, female gland mass; 10, ampulla; 11, spermatheca; 12, buccal area; 13, eye 
spot; 14, ganglion; 15, nerve ring; 16, esophagus; 17, reproductive system; 18, stomach; 19, hermaphrodite 
gland; 20, digestive gland; 21, intestine; 22, anus. 


1964} COLLIER AND FARMER: ADDITIONS TO Nupyp: iu | 


One of the specimens has the radular formula 59 by 9° 
hooks 40 to 60 long (fig. 1, F-G). There is no labial cuticle. 

The buccal mass (fig. 1, E 12) is large and muscular as is suggested by the well developed 
polyodont radula. The esophagus (16) is surrounded by the central nervous system (14) 
immediately adjacent to the buccal area. It then twists slightly before passing along the left 
ventral side of the stomach (18). The stomach is slightly almond shaped with the small end 
pointing anteriorly. The intestine (21) originates on the right side of a small knob at the 
narrow anterior end of the stomach and runs back along the right dorsolateral side of the 
digestive gland. The intestine is thin walled and straight, and fits into a slight groove in the 
digestive gland. Just anterior to the gill cavity it dips down the side of the digestive gland to 
the floor of the body cavity and then rises to open to the outside by means of the anal papillae 
(22). The large digestive gland (20) occupies the posterior half of the body cavity. In the 
preserved animals it is yellow and appears granular. It partly surrounds the stomach, extending 
to its anterior margin along the ventral side but only covering the posterior half on the dorsal 
surface. It communicates with the stomach where the esophagus enters. 


\I] the teeth are simple 


The heart is immediately anterior to the gill cavity. The auricle is thin walled and its 
boundaries are not well defined. The two ventricles are better developed and are visible along 
the top of the digestive gland. There is an aortic blood space which runs along the middorsal 
line of the digestive gland and stomach before emptying into sinal areas anterior to the 
stomach. 


The central nervous system was not examined closely. It is composed of two prominent 
cerebral ganglia. There is a black eyespot (13) on the anterior dorsolateral surface of each 
ganglion. Many small nerve fibers extend anteriorly from this area to innervate the buccal area 
and rhinophores. There is a nerve ring (15) connecting the cerebral ganglia and passing under 
the esophagus. 

The reproductive system (fig. 1, D) is compact, lying in contact with the anterior edge 
of the digestive gland and the right cerebral ganglion. The ampulla (10) is large, thin walled, 
and flattened dorso-ventrally; it lies along the back distal end of the female gland mass. A thin 
sinuous duct connects with the hermaphrodite gland (19), which is completely embedded in 
the digestive gland along the right ventral lateral line of the stomach. As it leaves the ampulla, 
the hermaphrodite duct bifurcates into a short oviduct (7) and the vas deferens (2). The 
prostate (1) is moderately large but not extremely glandular, and although twisting it is not 
sinuous. It lies along the left margin of the female gland mass, partly embedded in it. The vas 
deferens (2) proceeds directly to the genital orifice (3) after leaving the prostate. The penis 
is not muscular and is unarmed. 

The vagina (5) is thin and long. The spermatheca (11) communicates with the vagina 
through a short duct of the same diameter as the vagina. The spermatheca is large and globular 
and lies with its lower hemisphere embedded in the female gland mass. The hemisphere above 
the gland is jet black while that embedded is a light yellow. The spermatocyst (6) is conical 
with its narrow end embedded in the gland mass. The top is rounded giving the organ a 
spherical appearance before it is removed from the gland. The fertilization duct (8) is short 
and moderately sinuous. The female gland mass (9) is dull yellow and communicates to the 
outside through a pore posterior to the genital oriface (4). The ducts in the female reproductive 
system form an X (fig. 1, D). 

Conualevia alba, sp. nov. 
(Plate 3; fig. 2, A-D) 
Type. — The holotype (CASIZ 29) was taken on December 27, 1963, at Newport Bay, 


California, by the senior author. It is an entire animal. Paratypes from the same locality bear 
the numbers CASIZ 30 and 31. Some paratypes are retained in the authors’ collections. 
Distribution. — This animal occurs abundantly on a rocky mud flat at Newport Bay, 


California, where it is numerous at certain times of the year. In November, 1963, more than 
30 specimens were collected in a matter of minutes; many others were seen. In December of 


384 San Dteco Society oF Naturat History [Vot. 13 


Fig. 2. A-D, Conualevia alba. A, radular teeth at rachidian space; B, C, radular teeth; 
D, diagram of reproductive system. E-F, Polycera alabe. E, ventral view of living animal; F, 
half row of radula. G-H, Dendrodoris atropos. G, diagram of reproductive system; H, penis 
spine and hook. 


1, ampulla; 2, oviduct; 3, fertilization duct; 4, female gland mass; 5, prostate; 6, spermatheca; 7, 
vaginal duct; 8, genital vestibule; 9, spermatocyst; 10, vas deferens; 11, penis sheath. 


1964} COLLIER AND FARMER: AppITIONs to Nw IBRANCH FAUNA 385 


the same year about ten more were collected, but there were fewer present than before. Mr 
James Lance has found them at Point Loma in San Diego, California a? 

Description. — The living animals are white. Around the edge of the notum are embedded 
small opaque, white glands which form an unorganized row of spots (Plate 3) 
also appear white to the unaided eye, but on close observation it is seen that 
small black dots, hundreds of dots comprising each ring. 

The body form is that of a typical dorid. The notum varies in height, tending to be high 
when the animal is crawling and flattened when at rest. One of the largest specimens measured 
24 mm. long and 14 mm. wide when actively crawling. The notum covers the entire foot. The 
surface of the notum is also quite variable. In many animals it is entirely smooth (Plate 3), 
with no papillae, whereas in others, it is thickly covered with uniform papillae closely set and 
much less than a millimeter high. Animals of the latter phase resemble C. marcusi but are of 
less height. The anterior edge of the foot is bilabiate. The mouth is very noticeable with two 
small oral tentacles flanking it. These tentacles are quite short but very stout, looking almost 
like part of the oral area. The retractile rhinophores are smooth and, relative to other west 
coast dorids, are quite long and slender. There is no ridge around the opening of the 
thinophoral or branchial chambers. There are eight tripinnate gills in a circle around the anus. 
These are retractile and when pulled in it is impossible to determine the site of the opening 
on a living animal because the branchial chamber is so tightly closed. The gills, like the rest of 
the animal, are white. 

The radula of one specimen has the formula 34 by 56.0.56. The teeth (fig. 2, A-C) are 
simple hooks. The rows of teeth on each side of the radula are staggered (fig. 2, A) at the 
rachidian space (a-a’). The teeth are approximately the same size in a radular row. There is 
some variance between the rows, especially in the newly formed area of the radula. There is a 
moderately large labial cuticle but it is very thin. 

The esophagus is fairly large and appears to be glandular along much of its length. It 
enters a small, round, muscular stomach from the ventral side. The stomach is almost com- 
pletely embedded in the digestive gland; only the anterior end protrudes. The intestine leaves 
the stomach anteriorly. 

There are two prominent black eyespots on the ganglia. 

The reproductive system (fig. 2, D) is compact. A thin duct leads from the hermaphrodite 
gland to the thin, flat ampulla (1) which lies along the back of the female gland mass (4). 
The hermaphrodite duct bifurcates, after leaving the ampulla, into a thin, flattened oviduct (2) 
and a long, flattened vas deferens and prostate. The prostatic part of the vas deferens (5) is 
poorly differentiated and is almost nonglandular. The whole tract is embedded in the female 
gland mass but can be seen from the outside without dissecting the mass. The penis (11) is 
unarmed. The vagina (7) leads into a long vaginal duct which ends where a duct from the 
spermatheca (6) joins it. The spermatheca is round but flattened, and lies along the top of 
the female gland mass, not embedded in it. The spermatocyst (9) is a small round organ 
embedded in the female gland and connected by a short duct to a point just distal from the 
connection of the vaginal duct and the duct of the spermatheca. A small fertilization duct (3) 
bifurcates from this point. An X is formed at the junction of these ducts, as can be seen in 
figure 2, D. 

Discussion. —Conualevia marcusi and C. alba can be separated by several important 
anatomical and morphological features. The most noticeable is the body form — Conualevia 
alba is a much thinner animal than C. marcusi, and is more delicate in appearance. Although 
both species have glands around the notal rim they are much more evident in C. alba. The 
thinophores of C. alba are much longer relative to their width than are those of C. marcusi. 
Conualevia marcusi has a large number of unipinnate gills while C. alba has only about half 
as many tripinnate gills; this is an important external feature. 

The dental formulae of the two species are quite different, but this is of unknown signifi. 
cance as an insufficient number of radulae were examined. The prostate of C. marcusi is 
moderately developed, that of C. alba is poorly so. The actual size and shape of the reproduc- 


. the rhinophores 
they are ringed by 


386 SAN Disco Society oF NATURAL History { VoL. 13 


Plate 1. Cadlina evelinae 


Plate 2. Conualevia marcusi, sp. nov. 


<> 3 TER és . 


, te] j 2 
“Y > FY , Ms 
pee % EL Sil O86 ome a, 
* be Br iat aye en at hed a . 
Loe : 3S ae ae | vate oy 
a or” ahaa’ < 


yh . 
: wnt . aa 
xy oy , ras } ‘ 
* x “ay zt €) , 


La Meh ge ce Pe? saa 
AG od Sama a arti ae 


Plate 3. Conualevia alba, sp. nov. 


388 San Dreco Society oF Natura History { VoL. 13 


tive organs are probably of little importance, as these features, as well as the coloration of these 
organs, are related to the stage of the breeding cycle. The reproductive systems can be separated 
by the relative position of the spermatheca and spermatocyst. In C. marcusi they are on op- 
posite sides of a line drawn from the vaginal duct to the fertilization duct; that is they are on 
opposing sides of the X. In C. alba they are on the same side. 

The genus Conualevia possesses strong affinities to the members of the family Dorididae. 
It is much closer to that family than to any other group within the Eudoridacea. It has gills 
and rhinophores that are completely retractile, conforming to the tribe Cryptobranchia as 
established by Odhner (1939). The body shape, reproductive system, and general form of the 
digestive system conform to the family Dorididae within this tribe. The radula is strikingly 
similar to most species of the Dorididae. The feature of the smooth rhinophores, however, is 
very distinct; no other cryptobranch is recorded in the literature as possessing nonperfoliate 
rhinophores. 

Because of these affinities to the Dorididae, we have placed the genus Conualevia in that 
family. We feel, however, that the presence of smooth rhinophores indicates an evolutionary 
line distinct from that of the other members of the family, and have thus set the genus apart 
in the new subfamily Conualevinae. 

It is difficult to determine how much importance should be placed on the smooth 
rhinophores in attempting to relate these animals to the other cryptobranchs and in placing 
them in the presently accepted classification. This is partly because the exact function of the 
thinophores is unknown. Various workers (see Agersborg, 1922; Arey, 1917, 1918) have 
shown that these organs are sensitive to chemical and tactile stimulation, but to what extent 
and precisely how the animals use this ability is not adequately known. If the rhinophores are 
used to some degree as sensory receptors, it would seem that perfoliation would be an advan- 
tage tending to make the organ more sensitive simply by the increase in area. 

The name Conualevia means “smooth horns” and was chosen to call attention to the 
smooth rhinophores. In recognition of the contributions that Dr. Ernst Marcus has made to 
the field of opisthobranch taxonomy, we take pleasure in naming the type species of the genus 
in his honor. 


Tribe PHANEROBRANCHIA 
Superfamily NONSUCTORIA 
Family POLYCERIDAE 
Polycera alabe, sp. nov. 
(Plate 4; fig. 2, E-F) 

Type. — The holotype (CASIZ 27; original number WMF 130) was taken on the 
southeast side of Isla de Cedros, Baja California, by John Sloan in January, 1963. The radula 
and jaws have been removed from the preserved animal and are mounted on a slide (Box 1, 
Slide 69). One of two specimens from Puerto Refugio on Isla Angel de la Guarda in the 
Gulf of California bears the number CASIZ 32. It consists of a serial section of the entire 
animal mounted on eleven slides (Box 1, Slides 71-81) and was collected in March, 1963. 
Plate 4 illustrates the holotype. 

Distribution. — This species has been taken only in the localities mentioned above. 

Description. — The living animal is blue-black and, with the exception of the bottom of 
the foot, is covered with orange spots arranged more or less in rows posterior to the rhinophores. 
The black gills and rhinophores also have orange markings on them. Two of the four processes 
on the velum of the holotype are black; the two lateral ones are white. All processes are black 
in the two animals from the Gulf of California. The edge of the foot is white. The area above 
the eye spots is translucent. There are 20 colorless translucent projections on the body, some 
with yellow tips. There are 11 close-set black projections on the tail. 

The body form is typical of Polycera, with four processes on the velum. When actively 
crawling the holotype measured 25 mm. long, 5 mm. wide, and 5 mm. high in the heart region. 
Other individuals measured 12 and 15 mm. long, 4 mm. wide, and 5 mm. high. The foot is 
slightly expanded anteriorly (fig. 2, E) and is transversely grooved. 


1964 } COLLIER AND FARMER: AppITIONS To NupiprANCH FAUNA 389 


There are six centrally located gills, all unipinnate; the four anterior gills are 5} mm. high, 
the posterior ones smaller. The nonretractile rhinophores are perfoliate, with 11 leaves. 


The radula has the formula 9 by 3.2.0.2.3. Figure 2, F shows a half row from the radula. 
From the center, the first two teeth each have a strong denticle at the distal end, at right angles 
to the main axis of the tooth. The second tooth is slightly larger than the first. The third, 
fourth, and fifth teeth each lack a denticle. The third tooth is about half the size of the second. 
and the teeth get progressively smaller laterally. The axis of all the teeth is slightly curved. 
Jaw plates are present. 


The penis is armed. 


Discussion. — Three species of Polycera are known from the east Pacific — P. atra Mac- 
Farland, 1905, P. zosterae O’Donoghue, 1924a, and P. hedgpethi Marcus, 1964. Polycera alabe 
differs from them by its color pattern and radular characteristics. The radula of P. alabe lacks 
the basal spine on the second tooth that is found on the other three species. 


The name alabe refers to the way the bluish-black color of the animal diffused into the 
preservative (formalin); it is Greek for a type of ink. 


Section POROSTOMATA 
Family DENDRODORIDIDAE 
Dendrodoris atropos (Bergh, 1879) 
(Plate 5; fig. 2, G-H) 

Distribution. — Dendrodoris atropos has been reported from Brazil by Marcus (1957). 
It is one of the most common nudibranchs in the Gulf of California. We have taken this ani- 
mal at Coloradito, Puertecitos, and Isla Angel de la Guarda. In addition it has been found 
at Bahia Loreto (Gulf), Bahia de La Paz (Gulf), Bahia de los Angeles (Gulf), Bahia Se- 
bastian Viscaino, and at Tenacatita, Jalisco, on the mainland of Mexico. Thus, it is common 
throughout the southern end of the peninsula on the Pacific side and the length of the Gulf. 
We have found it around rocks in early summer and in late November. It is apparently most 

abundant from late winter through late spring or early summer. 


Description. — The living animal is black except for the tips of the rhinophores and gills, 
which are white, and the edge of the mantle ruffle, which has a red line around it. In the liv- 
ing animal the viscera can sometimes be seen through the notum. The foot and underside of 
the notum are sooty gray. The black color of the animal is moderately well retained in preser- 
vative; the red line fades but is still noticeable in most specimens. 


The body is plump. The notum is smooth or slightly undulate. There is a 6 mm. broad 
undulating ruffle which extends around the edge of the notum. The distal edge of this ruffle 
is dull red (Plate 5). The clavi of the rhinophores are set on stout stalks, are retractile into 
simple chambers, and have 20 to 24 leaves. In one of the animals dissected there were eight 
retractile bipinnate to tripinnate gills. The foot is very muscular, allowing the animal to cling 
tenaciously to rocks; we tore several animals apart trying to remove them, leaving the foot still 
clinging to the substrate. The largest animal we collected measured 45 mm. long, 15 mm. wide, 
and 7 mm. high when living; this measurement included the notal ruffle. 

The hermaphrodite duct leads into a large thick ampulla (fig. 2, G 1) which is bean 
shaped, bent in the middle so that it recurves on itself. The oviduct (2) is narrow and short. 
The vas deferens winds anteriorly and gradually widens as it becomes glandular. The prostatic 
part of the vas deferens (5) is highly convoluted. The vas deferens gradually narrows and 
loses its glandular texture to become a winding, flattened duct (10) running latero-venterally 
along the penial sheath. As soon as the vas deferens enters the penis it becomes enclosed in 
the large, muscular penial sheath, which gradually tapers and, just before entering the genital 
vestibule (8), bends sharply to the outside to connect with the vestibule at right angles. The 
penis is armed with both short hooks and long spines (fig. 2, H). | 

The vagina is short but muscular, and leads into a small winding duct (7) leading to 
the small spherical spermatheca (6). The fertilization duct (3) debouches from the vaginal 


390 SAN DigEGo Socrety oF NATURAL History [Vor 


' 


Plate 5. Dendrodoris atropos 


Plate 6. Cerberilla pungoarena, sp. nov. 


1964} CoLLIER AND FARMER: ADDITIONS TO NupIBRANCH FAUNA 39] 


duct and spermatheca at this point. This duct is very long and convoluted, although not 
winding upon itself. It enters the female gland mass (4) just after connecting with a duct 
leading from the small pear-shaped spermatocyst (9). The female gland mass is not excep- 
tionally large; it opens to the outside posterior to the genital vestibule. No accessory glands in 
the area of the genital vestibule were seen. 

The digestive and nervous systems are as described for D. atropos by Marcus (1957) 
and Marcus and Marcus (1962). There is a large buccal bulb with a large ptyaline gland lying 
to the right of it, anterior to the genital area. The pharynx leaves the buccal bulb and passes 
through the nerve ring, doubling back to a position just posterior and ventral of the nerve 
ring where the buccal ganglia connect to it. There are two small glands on the esophagus at 
this point. The esophagus is thick and glandular with the mosaic pattern described by Marcus 
and Marcus (1962:474). The esophagus ends in a prominent sphincter. 


Discussion. — For some time we thought that these animals were Dendrodoris nigra. The 
color seemed to agree perfectly with one of the phases of D. nigra and although there was a 
striking similarity between its digestive system and that of D. atropos there were enough minor 
differences elsewhere to allow for its being D. nigra. Among these is the muscular penial 
sheath which is so noticeable in the specimens at hand but which Marcus (1957) does not 
mention. Not until we were able to dissect specimens of D. nigra from two different locations, 
through the courtesy of Drs. Rehder and Kenny, were we sure that our specimens were not 
D. nigra and were indeed Dendrodoris atropos. 


Many authors (Marcus, 1957; O’Donoghue, 1924b; Eliot, 1906) have discussed the 
differences in classification and the inherent difficulties encountered in this group of animals 
and it is not necessary to repeat them here. Color may be one of the most important factors 
as Marcus (1957) states, but it alone is not enough, as these animals indicate. Classification 
in this group must still take into account all of the organ systems together and not single out 
any one as “the most important.” 


Suborder EOLIDACEA 
Tribe CLEIOPROCTA 
Family AEOLIDIIDAE 
Cerberilla pungoarena, sp. nov. 
(Plate 6; fig. 3, A-D) 

Type. — The holotype (CASIZ 28) and only specimen was collected by John Sloan at 
Puerto Refugio on the north end of Isla Angel de la Guarda in March, 1963. It consists of 
the preserved animal and a slide of the radula and jaw element (Box 1, Slide 70). 

Description. — The body and foot of the living animal are translucent white with light 
brown or tan on the dorsal surface. The posterior edges of the oral tentacles are lined with 
black pigment which makes them appear dark gray. The distal end of each cerata is opaque 
white but has a small translucent cap at the tip. The major portion of each cerata is translucent 
with a thin, speckled-appearing thread of liver diverticula running up the middle. 

The foot is wide (fig. 3, B) and rounded behind. It does not extend posteriorly beyond 
the cerata. The anterior edge of the foot is rounded and produced into lateral extensions at 
the corners. When viewed dorsally (fig. 3, A), the foot is very broad, extending laterally 
beyond the outer margins of the cerata. The animal measured 20 mm. long, 7 mm. wide and 
2.5 mm. high when actively crawling. The anterior cerata are short, whereas the posterior ones 
are very long and extend beyond the tail when the animal is actively crawling. The cerata are 
dorso-venterally flattened and fit next to each other like a stack of books. The cerata trail 
straight behind when the animal is crawling, but when the animal stops the cerata become dis- 
oriented and thrash around. When the animal is touched or disturbed the cerata move about 
wildly, extending and shortening in length (Plate 6). | | 

The rhinophores are relatively short and are perfoliate. The terminal half is white, the 
base, tan. There is a small black eye spot at the base of each rhinophore. T he thin, pointed 
oral tentacles are slightly contractile and can extend to almost half the length of the animal. 


392 SAN Disco SociETY OF NATURAL History [VoL. 13 


Fig. 3. Cerberilla pungoarena. A, dorsal view of living animal; B, ventral view of living 
animal; C, radular teeth; D, jaw plate. 


1964} COLLIER AND FARMER: ADDITIONS TO NupIBRANCH FAUNA 393 


The radula has the formula 21 by 0.1.0. A tooth from the center of the radula contained 
28 denticles (fig. 3, C). There are two large lateral denticles of unequal size on each side of 
the tooth with many small denticles between the large ones. In some teeth there is a large cen- 
tral denticle. Older teeth of the radula showed considerable wear with most of the denticles 
worn off and rounded. The thin, smooth, oval jaw plates (fig. 3, D) are pale yellow and 
slightly convex, with a smooth masculatory process. 


Discussion. — This animal was collected crawling on top of sandy mud, and was later 
observed to have definite burrowing abilities. It completely submerged into the sand in an 
aquarium, the sand collapsing behind the animal as it passed through. As far as we can deter- 
mine, this is the first recorded observance of an aeolid burrowing in sand. We did not deter- 
mine what the animal eats in this peculiar habitat. The broad foot is particularly adapted for 
this kind of existence and the cerata are aligned for easy passage through the sand. 


It has been pointed out by Marcus and Marcus (1959:260) that Cerberilla is a repre- 
sentative genus of the Indo-West Pacific, occurring principally, but not exclusively, in warm 
areas. They described C. tanna from the coast of Texas, the first known Atlantic Cerberilla. 
The specimen at hand is the first known representative of this genus from the east Pacific. 


In a discussion of Cerberilla, Bergh (1905) pointed out that the most important charac- 
teristic for separating the species of this genus from one another is the relationship of their 
colors. Marcus and Marcus later (1959) stated that the radula is the best distinguishing 
characteristic. 


Cerberilla ambonensis Bergh 1905 (see his plate XIX) differs from C. pungoarena by 
having black tipped cerata and rhinophores. Although Marcus and Marcus (1959) did not 
take color notes from the living animal, they described C. tanna as having an orange-brown 
spot on the outer surface of many but not all cerata, just under the cnidosac. Cerberilla pungo- 
arena can be distinguished from C. tanna by the presence of a large central denticle on some 
of the teeth. Baba (1940:110) described C. asamusiensis as having smooth rhinophores, a 
broad yellow area on the head between the oral tentacles and rhinophores, and teeth with only 
6 to 8 large denticles with smaller accessory denticles, characteristics not common to C. pun- 
goarena. 


394 San Disco Society OF NaTurRAL History [VoL. 13 


LITERATURE CITED 


AcErSBORG, H. P. KJERSCHOV 
1922. Some observations on qualatative chemical and physical stimulations in nudi- 
branchiate mollusks with special reference to the role of the ‘rhinophore.’ Jour. 
Exper. Zool. 36:423-444. 


Arey, LESLIE B. 
1917. The sensory potentials of the nudibranch “rhinophore.” Anat. Rec. 11:514-516. 
1918. The multiple sensory activities of the so-called rhinophores of nudibranchs. Amer. 
Jour. Physiol. 46:526-532. 


Basa, K. 
1940. Some additions to the nudibranch fauna of the northern part of Japan. Bull. 
Biogeog. Soc. Japan 10:103-111. 


BerGH, R. 
1879. Die Doriopsen des Atlantischen Meeres. Jb. Dtsch. Malakozool. Ges. Jahrg. 
6:42-6). 
1905. Die Opisthobranchiata der Siboga Expedition. Siboga Expedite 50:1-248, pls. 
1-20. 
EriomnGxNae: 


1906. Report upon a collection of Nudibranchiata from the Cape Verd Islands, with 
notes by C. Crossland. Proc. Malacol. Soc. London 7:131-159, pl. 14. 


LANCE, JAMES R. 
1962. Two new opisthobranch mollusks from southern California. Veliger 4:155-159, 
pl. 38. 


MacFar.anp, F. M. 
1905. A preliminary account of the Dorididae of Monterey Bay, California. Proc. Biol. 
Soc. Wash. 18:35-54. 


Marcus, ERNST 

1955. Opisthobranchia from Brazil. Bol. Fac. Fil. Univ. S. Paulo, Zool. 20:89-262, 
30 pls. 

1957. On Opisthobranchia from Brazil (2). Jour. Linnean Soc. London 43 :390-486. 

1958. On western Atlantic opisthobranchiate gastropods. Amer. Mus. Novitates 1906. 
82pp. 

1961. Opisthobranch mollusks from California. Veliger 3, Supplement. 85pp. 

1964. A new species of Polycera (Nudibranchia) from California. Nautilus 77: 128-131. 


Marcus, EVELINE and Ernst Marcus 
1959. Some opisthobranchs from the northwestern Gulf of Mexico. Inst. Marine Sci. 
6:251-261, 19 figs. 
1962. Opisthobranchs from Florida and the Virgin Islands. Bull. Marine Sci. Gulf and 
Carib. 12:450-488. 


Opuner, Nis H. 
1939. QOpisthobranchiate Mollusca from the western and northern coasts of Norway. 
K. Norske Vidensk. Selsk. Skr. 1939. 1. 93pp. 


O’DoNoGHUE, CHARLEs H. 
1924a. Notes on the nudibranchiate Mollusca from the Vancouver Island region. IV. 
Trans. Roy. Canadian Inst. 15:1-33, pls. 1-2. 
1924b. Report on Opisthobranchiata from the Abrolhos Islands, western Australia, with 
description of a new parasitic copepod. Jour. Linnean Soc. London 35:521-579, 
pl. 27-30. 


Tee j 


ns 
*. = 7 


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rie 


TRANSACTIONS 
OF THE 
SAN DIEGO SOCIETY OF NATURAL HISTORY 


VoLuME 13, No. 20, pp. 397-404 


THE MAMMALS OF CERRALVO ISLAND, 
BAJA CALIFORNIA 


BY 


RicHaRD C. BANKS 


Curator of Birds and Mammals 
San Diego Natural History Museum 


SAN DIEGO, CALIFORNIA 
PRINTED FOR THE SOCIETY 


DECEMBER 30, 1964 


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THE MAMMALS OF CERRALVO ISLAND ! 5 1955 


BAJA CALIFORNIA HARVARD 
UNIVERSITY 
BY 


RicHARD C. Banks 


Cerralvo Island, the southernmost of the islands in the Gulf of California, Mexico. has 
received but slight mention in the literature of mammalogy. This is not unexpected, as only 
four native and two feral species of mammals are known to occur there. Two bats have been 
recorded, but only in the nature of locality records. Two mice are considered to be endemic 
subspecies, but the original descriptions are somewhat incomplete and no subsequent studies 
have been made. Nothing has been reported of the population size of the feral species or of 
any results of their introduction. A summary of work on Cerralvo Island prior to the present 
investigation (Banks, 1962) omitted mention of mammal specimens taken by Donald R. 


Dickey in 1928. 
Between October, 1960, and June, 1962, I spent a total of six full weeks on Cerralvo 


Island. Several other visits, lasting from a few hours to a few days, were made to the island 
in this period. A complete schedule of this investigation has been presented previously (Banks, 
1963b), as have results of other parts of the study (Banks, 1963a, 1963b; Banks and Farmer, 
1962). 


ACKNOWLEDGEMENTS 


Work on Cerralvo Island was begun under the sponsorship of the Belvedere Scientific 
Fund of San Francisco, and continued under a grant from the National Science Foundation 
(G-19333). Facilities of the California Academy of Sciences and the San Diego Natural His- 
tory Museum were used, and the latter institution sponsored one visit to the island. Specimens 
were borrowed from Dr. Robert T. Orr, California Academy of Sciences; Dr. Richard H. 
Manville, Bird and Mammal Laboratories, United States Fish and Wildlife Service; and Dr. 
Seth B. Benson, Museum of Vertebrate Zoology. Dr. Thomas R. Howell made Donald 
Dickey’s field notes available to me. Cat scat was dissected by Mr. Boyd K. Seavey, and the 
contents were identified by Dr. Richard Etheridge. Dr. Richard G. Van Gelder provided 
information from his notes taken on the island. My field companions on the island, Robert G. 
Crippen, Michael Soulé, and Wesley Farmer, were of great assistance. Mr. and Mrs. Richard 
Adcock, of La Paz, Baja California, provided transportation to the island, and their friendly 
assistance in many ways is appreciated. 


List oF MAMMALS 


Big Brown Bat. Eptesicus fuscus peninsulae.— A single specimen of this bat was taken 


on May 27, 1962. Details have been reported elsewhere (Banks, 1964). 


Western Pipistrelle. Pipistrellus hesperus australis. — This bat was first reported on Cer- 
ralvo Island by Townsend (1912), who mentioned a specimen taken by H. E. Anthony on 
April 19, 1911. Specimens taken during this study are as follows: October 25, 1960; November 
9 and 17, 1961; April 16, 1962; May 24, 1962; June 2, 1962. These bats were common at 
dawn and dusk on all parts of the island. 


Males taken on October 25, 1960, and November 9, 1961, had testes 6 and 3 mm. long, 
respectively. A female taken on June 2, 1962, had two embryos 7 mm. in length. 


Desert Pocket Mouse. Perognathus arenarius siccus. — The population of pocket mice on 
Cerralvo Island was originally described by Osgood (1907) as Perognathus penicillatus sICCUS, 
but it was assigned to Perognathus arenarius by Nelson and Goldman (1929) when the dis. 
tinctness of these two species became evident. In the original description of this subspecies, 
Osgood did not consider it to be an endemic form. Although the type was selected from 


PR : sa 
~~. Vii ZU 


400 San Deco Society oF Natura History iVor. 1s 


Cerralvo (“Ceralbo”) Island, Osgood (1907:20) stated: “This form was found not only on 
Ceralbo Island but also at several localities on the neighboring end of the Peninsula. Speci- 
mens from the peninsular localities Tres Pachitas and Pescadero seem referable to it, while 
others from slightly farther north show intergradation with [ P. penicillatus | arenarius.” Nelson 
(1921:91) first stated that the race was endemic to the island. 


The specimens from Tres Pachitas and Pescadero were not mentioned by Hall and Kelson 
(1959:499) , whose range map does not extend far enough south to include them. Information 
on the labels of those specimens indicates that Tres Pachitas is 36 miles south of La Paz, or 
about 18 miles north of Todos Santos. Pescadero is indicated as being ten miles south of 
Todos Santos. Todos Santos is on the west side of the peninsula of Baja California, approxi- 
mately 54 miles south of La Paz and about the same distance southwest of the southern tip 
of Cerralvo Island. 


Nelson and Goldman (1929) described two additional subspecies of Perognathus arenarius 
from Baja California, including P. a. sublucidus from La Paz. They did not compare material 
from La Paz to specimens from Cerralvo Island, nor did they mention specifically those from 
the other localities in the Cape region. A very localized range was ascribed to sublucidus by 
the authors, as follows: “This subspecies has an unusually limited but well defined range, 
covering the very arid desert of the small sloping basin a few miles in extent, lying about 
the southern and southwestern part of La Paz Bay. To the north and south its range is limited 


by mountainous areas, and to the west by the divide between the drainage to the Gulf and to 
the Pacific.” 


However, with no additional information and apparently with no specimens from localities 
other than La Paz, Hall and Kelson (1959) mapped the range of P. a. sublucidus as extending 
across the peninsula in the latitude of La Paz and north along the Pacific coast to the latitude 
of Santa Margarita Island. This implies that the specimens from Tres Pachitas and Pescadero 
would be referred to sublucidus, although they actually have been overlooked. 


Samples of the three populations from this part of Baja California were examined in an 
attempt to determine their affinities. The population from Cerralvo Island included the speci- 
mens taken by Nelson and Goldman in 1906 and those collected by me in 1960-1962. The 
small series taken by Nelson and Goldman at Tres Pachitas and Pescadero in 1906 was aug- 
mented by a series from Todos Santos, taken by Chester C. Lamb in 1928. The sample from 
the La Paz population was collected by Laurence M. Huey in 1941. 


In the original descriptions, both P. a. siccus and P. a. sublucidus were compared to P. a. 
arenarius from the Magdalena Plain and Vizcaino Desert region of western Baja California. 
Measurements of several characters from adult topotypes were given in each description, but 
no mention was made of sexual variation. Sexes were segregated in the present study, and 
considerable variation was noted. Males averaged larger than females in virtually every charac- 
ter examined. The differences were slight in many instances, but in some features, as the 
greatest length of the skull, were as great as three per cent. This suggests that future studies 
of pocket mice might yield more definite results if the possibility of sexual variation 1s 
considered. 


Although there is considerable overlap in the ranges of measurements in these three popula- 
tions (fig. 1), there are rather clear differences between the means of the populations (table 1). 
The small number of specimens available, especially from the Todos Santos population, pre- 
cluded a more sophisticated statistical analysis. 

Males from Cerralvo Island average larger than those from the Todos Santos region in 
greatest length of the skull, basilar length, and greatest width of the skull, as well as in total 
length and tail length. The males from La Paz are smaller than those from Cerralvo Island 
in all these characters, in head and body length, and in frontonasal length as well. Males from 
La Paz have slightly longer tails than those from the Todos Santos area, but in all other char- 
acters mentioned the La Paz population averages smallest, despite one exceptionally large male 
from La Paz that tended to raise the average considerably. 


1964} Banks: MAMMALS OF CERRALVO ISLAND 40] 


TABLE 1 


Selected measurements (in mm.) of Perognathus arenarius from southern Baja California. 


Males 
Cerralvo Island La Paz Todos Santos area 
N Range Mean N Range Mean N Range Mean 
Miotal@lencthyeese =e 17: 168-188 176.5 12 160-184 170.0 7 168-180 173.7 
Earle ensth eee 17. —- 92-107 O7e7, 12. 86-100 94.1 7 81-102 93.7 
Head and body length... 18 71-86 77.8 13 71-84 75.9 8 73-92 79.2 
Length of skull ............. 17 24.9-26.9 25.97 12 23.5-26.95 24.82 6 25.05-26.05 25.56 
Bastlarslengthyee ss 16 —-18.8-20.4 19.51 12 17.9-20.1 18.77 6 = 18.5-19.7 19.07 
Frontonasal length -_.. 19 16.9-18.6 17.68 13 15.95-18.45 16.82 6 = 17.2-18.2 17.68 
Mastoid width ............. 17 1279-1829 13233 1a. 125-138 13.14 6 13:0-15,55..) W323 
Females 
Weal Weagin soe 16 = 161-186 169.8 12. 160-185 170.6 11 155-180 168.3 
Marl @lencthweeee se 16 87-98 O22 12. 88-100 94.0 11 80-96 90.5 
Head and body length... 17. 73-94 77.3 13-72-85 76.6 12 70-85 77.7 
Length of skull 16 24.4-26.2 25.13 12. = 23.3-25.85 24.64 8 23.7-25.5 24.89 
Basilar length -............ 16 =:18.2-20.15 18.96 13 17.5-19.6 18.60 7  17.4-19.4 18.61 
Frontonasal length -..... 17 ~—-16.35-17.8 Va 2, 9-17.28 16.80 8  16.5-17.4 16.96 
Mastoid width .............. 17 1226-133 13.00 13 12.4-13.7 13.01 9 = 12.7-13.6 13.2] 


Females from the La Paz population average slightly greater in tail length and in total 
length than females from the other populations, but the differences between the means are very 
slight. In other characters, the females from La Paz average smallest of the populations. Females 
from Cerralvo Island are larger than those from the Todos Santos region except in head and 
body length. One small female included in the population from Todos Santos tended to reduce 
the means for that population. 


There is but little color difference in the three series. The population from Cerralvo Island 
averages slightly grayer; it is less brownish dorsally, and somewhat lighter overall, than that 
from the Todos Santos area. The specimens from La Paz are very similar in color to those 
from Cerralvo Island. 


The following conclusions can be drawn concerning the status of these three populations. 
Perognathus arenarius siccus is distinguishable from the mainland populations and is endemic 
to Cerralvo Island; it does not include populations from the vicinity of Todos Santos, as 
originally stated (Osgood, 1907). Perognathus a. sublucidus is still known only from La Paz 
and the immediate vicinity, as stated by Nelson and Goldman (1929). The population from 
Pescadero, Todos Santos, and Tres Pachitas, on the west side of the Cape region of Baja 
California, is probably best referred to P. a. arenarius. This seems to be essentially the same 
concept of the relationship of these forms held by Nelson, who (1921) listed Perognathus 
penicillatus arenarius (now P. arenarius arenarius) as characteristic of both the Cape and 
Vizcaino Desert districts, presumably regarding specimens from Pescadero and Tres Pachitas 
as representative of that form. 

This concept differs from that mapped by Hall and Kelson (1959), however, in extend 
ing the range of P. a. arenarius southward through the Magdalena Plain to Pescadero on the 
Pacific side of the peninsular divide. This arrangement must be considered tentative, for I have 
not examined examples of more northern P. a. arenarius in this study. It is comp! ated by 
the report by Alvarez (1958) of specimens of P. a. albulus, supposedly endemic to Santa 


; : . te th | . 
Margarita Island, from Estero Salinas, on the peninsular mainland opposite that ts! ind. How 


402 San Disco Society oF NATuRAL History {VOL 13 


ise | ee La Paz mal 
é Tel ee ce Todos Santos 12 | 
dS PEE oS ee Cerralvo A el Mahe at Nae 
ype a PES . LaPaz [SOL 
(ee Sn Todos Santos Saale 
16 | Cerralvo (Bia ee 
\7 18 RS) 20 70 80 90 
Basilar length Head and body length 
| Vea eee all 2 Sear, La Paz Zeiten ote eee 
8 | Todos Santos 28a 
oR Meee ea ee Cerralvo Nie mel ae ees 
[ana be Oe an é La Paz [i ae ee te 
(sieeee EAs Todos Santos Gas eee 
Nias ao | NO aia Cerralvo [Dea So 
23 24 25 26 em 16 Wig 18 
Length of skull Frontonasal length 


Fig. 1. Range and means of some measurements in three populations of Perognathus 
arenarius in southern Baja California. 


The upper half of each section represents females, the lower half, males. Dots mark measurements of 
an extremely small female in the Todos Santos population and an especially large male in the La Paz sample. 


ever, in their remarks on P. a. albulus, Nelson and Goldman (1923) compared the insular 
population to P. a. arenarius and mentioned “the darker form which occurs on the adjacent 
mainland.” Even if albulus occurs on the eastern edges of Magdalena Bay, opposite Santa 
Margarita Island, arenarius could extend south beyond it, in upland situations, to the vicinity 
of Todos Santos and Pescadero. 

These animals were not common in the narrow, rocky arroyo on the east side of the 
island, where we camped in 1960. In the larger, flatter arroyo near Ruffo’s Ranch on the west 
side of Cerralvo Island, burrows and mounds indicated fairly widespread activity. Sign of this 
nature was found far up the arroyos where the substrate was sandy and soft enough to permit 
digging. Pocket mice seem to be much less abundant on the slopes and ridges. They are quite 
abundant on the flat, sandy southwestern portion of the island. 

The breeding season of pocket mice on Cerralvo Island is little known. A female with 
two 5 mm. embryos on May 25, 1962, was the only pregnant animal obtained. 

Weights of adults taken on Cerralvo Island in 1960 and 1961 ranged from 12.5 to 17.8 
gm. Six females averaged 14.5 gm. and seven males averaged 16.7 gm. 

Cactus Mouse. Peromyscus eremicus avius.— This subspecies was described by Osgood 
(1909) as endemic to Cerralvo Island, based on material collected there by Nelson and Gold- 
man in 1906. He recognized the animals as relatively large, with buffy underparts but without 
a pectoral spot. He presented average measurements of total length, tail, hind foot, and ear, 
of ten adult topotypes, without reference to their sex. He gave skull measurements only for 
the type. 

Averages which I calculated from the entire series taken by Nelson and Goldman (except 
the type) are considerably less than those given by Osgood, and I suspect that he may have 
averaged only the ten largest specimens. For example, Osgood (1909:247) gives 194 mm. for 
the average total length, with a range of 186 to 209 mm. The ten largest specimens available 
to me from that collection range from 187 to 200 mm., and average 193.5 mm. Fifteen adults, 
eight males and seven females, from Nelson and Goldman’s series are at hand, however, and 
range in total length from 176 to 200 mm.; the average is 189.4 mm. Males in that series 
average 187.5 mm. in total length, and females average 191.6 mm. 


1964} Banks: MAmMMALs OF CERRALVO ISLAND 403 


TABLE 2 

Measurements (in mm.) of Peromyscus eremicus avius from Cerralvo Island 

Males Females 
N Range Mean N Range Mean 
Wogalblencth e282 peas 21 176-208 190.2 18 182-216 195.5 
Miaiibletigt late ec 21 89-115 101.0 18 78-118 101.8 
Head and body length.......... 21 83-94 89.2 18 82-115 93.8 
Ieind toorlength=.<..ae.... 21 19-22 20.0 18 19.22 20.5 
Length of skull....-scces.- 20. -24.1-25.65 249-16 —Ss«(23.9-25.95 24.8 
Basilarslengthy. 22 2....2....--.. 20 19:4-21.2 20.4 16 19.6-21.4 20.4 
Width of braincase................ 20 11.5-12.5 12.0 17 11.3-12.4 12.0 
Interorbital constriction.......... 22 3.7-4.1 3.9 18 3.7-4.05 3.9 
Maxillary toothrow ..........-.-... 22 3.6-4.2 oi) 18 3.65-4.0 3.9 


A comparison of the sexes in both series available (that taken by Nelson and Goldman in 
1906 and that taken by me in 1960-1962) showed females averaging larger than males in total 
length and in head and body length. There were only slight and inconsistent differences in 
other features examined. Sex for sex, the specimens of the more recent series averaged larger 
than those of 1906 in the same characters, but I judge from tooth wear and from dates of 
collection that the specimens taken in 1960-1962 also averaged older. Table 2 summarizes my 
measurements of the Cerralvo Island population. 


This species was abundant in the arroyo on the east side of the island in 1960. In a single 
trap set near our food supply on October 28, ten mice were captured in approximately three 
hours. On the next evening, six more mice were taken in the same trap in about the same 
period of time. On the west side of the island, Peromyscus eremicus avius was taken most 
commonly in rocky areas and at the edges of the arroyos. 


Breeding seems to take place in the spring on Cerralvo Island. A female taken on May 
31, 1962, had two 19 mm. embryos, and three taken from May 30 to June 3 were lactating. 
A juvenile was taken on May 31. Males taken in late May and early June had testes 8 to 9 
mm. long. From October 25 to November 2, males had testes from 5 to 7 mm. in length. Four 
adult females obtained in the fall had no embryos, but a subadult, molting from the grey 
juvenile to the brown adult pelage, contained three 3-mm. embryos. This evidence indicates 
that although most breeding takes place in the spring, reproductive activity may continue 
sporadically at least into the autumn months. 


House Cat. Felis domesticus. — Feral house cats occur on many of the islands in the 
Gulf of California, where they are apparently abandoned by fishermen. The first mention of 
the species on Cerralvo Island was by Etheridge (1961). The abundance of tracks, scats, and 
skeletons indicates that the population is high, although few cats were seen. Tracks found in 
the moist sand suggest that the beaches are patrolled nightly. 


A large quantity of scats was gathered on the island, mostly on the southern end, ee 
Dr. Richar« 


was examined for remains which would indicate the feeding habits of the cats. 
Etheridge identified the remains of a bird, small mammals, ctenosaurs (Cfenosaura Demsog wid 
rattlesnakes (Crotalus sp.), whip snakes (Masticophus flagellum) and insects. Smaller | 


“ALM 
fc lizard the islan 
bones may have been from young ctenosaurs or from the smaller iguanid lizards on the island. 


404 San Drieco Society oF NaturaAL History {Vore13 


Goat. Capra hircus.— The presence of goats on Cerralvo Island probably dates back to 
the time of the Ruffo Ranch. I did not see any goats during my visits to the island, but their 
droppings and tracks were abundant, particularly on the northern two-thirds of the island. 
Fairly well-defined trails were seen on the higher slopes and ridges. One skeleton was found in 
a rocky arroyo. 

Sign of goats was most evident in October, 1960, on the east side of the island, where 
droppings were found from beach to ridge. Vegetation on the higher slopes, near the ridge, 
was obviously browsed, and low vegetation was grazed along the ridge at an elevation of ap- 
proximately 1500 feet. Even there, where evidence of feeding was most noticeable, there 
seemed to be no great damage. These observations suggest that the population of feral goats 
on Cerralvo Island is not large, and that it presents no serious threat to the flora at this time. 


LITERATURE CITED 


ALVAREZ, T. 


1958. Roedores colectados en el territorio de la Baja California. Acta Zool. Mexicana 
2 (8) :1-7. 


Banks, R. C. 
1962. A history of explorations for vertebrates on Cerralvo Island, Baja California. 
Proc. Calif. Acad. Sci., 4th ser., 30:117-125. 
1963a. New birds from Cerralvo Island, Baja California, Mexico. Occas. Pap. Calif. 
Acad. Sci. no. 37. 5 pp. 
1963b. Birds of Cerralvo Island, Baja California. Condor 65:300-312. 
1964. Range extensions for three bats in Baja California, Mexico. Jour. Mamm. 45:489. 


Banks, R. C., AND W. M. FarMER 
1962. Observations on reptiles of Cerralvo Island, Baja California, Mexico. Her- 
petologica 18:246-250. 


ETHERIDGE, R. 


1961. Additions to the herpetological fauna of Isla Cerralvo in the Gulf of California, 
Mexico. Herpetologica 17:57-60. 


Hatt, E. R., AND K. R. KELSON 
1959. The Mammals of North America. 2 vols. xxx + 1083 + 79 pp. Ronald Press 
Co., New York. 


NEtson, E. W. 
1921. Lower California and its natural resources. Mem. Nat. Acad. Sci. 16: 1-194. 


NeEtson, E. W., AND E. A. GOLDMAN 


1923. A new pocket mouse from Lower California. Proc. Biol. Soc. Wash. 36:159-160. 
1929. Six new pocket mice from Lower California and notes on the status of several 
described species. Proc. Biol. Soc. Wash. 42: 103-112. 


Oscoop, W. H. 


1907. Four new pocket mice. Proc. Biol. Soc. Wash. 20:19-21. 
1909. Revision of the mice of the American genus Peromyscus. North Amer. Fauna 
no. 28. 285 pp. 


TOWNSEND, C. H. 


1912. Mammals collected in Lower California, with descriptions of new species. Bull. 
Amer. Mus. Nat. Hist. 31:117-130. 


INDEX 


405 


INDEX TO VOLUME 13 


TRANSACTIONS OF THE SAN Disco Society of Natura. History 
PREPARED BY Mrs. Berry MackINTOsH AND Mrs. RosEMaRrIE FIs BIG 
New names are in boldface type. Page numbers for illustrations are in italic type 


abalone, 369-376 
Actidium sp., 273, 274 
Actitis macularia, 55 
Acanthina muricata, 340, 350 
Aequipecten abietis, 354 
tumbezensis, 355 
Agaphelus glaucus, 140 
Agathymus comstocki, 63, 70 
dawsoni, 64-70, 169, 170-172 
Agave goldmaniana, 62, 63, 169 
Agkistrodon, 185-268 
acutus, 189, 192, 195, 197, 202, 206, 214, 216, 
218, 220, 223-25, 227, 229, 231, 234-35. 
239, 240, 249, 260 
annamensis, 188, 239, 249 
bilineatus, 189, 192, 197, 201-4, 208, 212, 214, 
BIG. 218; 220! 223-5, 227. 229, 231, 234.5, 
239, 240, 249. 252, 260 
blomhoffi, 189, 190, 192, 214, 216, 218, 220, 
222-5, 227, 231, 235, 239, 240, 249 
contortrix, 189, 192, 200-4, 208, 209, 212, 214, 
BUG ie218;2220: 2225, 227, 22031, 235, 
239, 240, 249, 252, 255 
contortrix contortrix, 189 
contortrix laticinctis, 189 
contortrix mokeson, 189 
halysu801 19% 214 216, 218, 222-255.227, 
229, 231, 235, 239, 240, 249, 260 
himalayanus, 188, 239, 249 
hypnale, 189, 206, 214, 216, 218, 220, 222-5, 
227, 229, 231, 235, 239, 240, 249, 260 
millardi, 188, 239, 249 
mokeson, 218, 249 
monticola, 188, 239, 249 
nepa, 188, 239, 249 
piscivorus, 189, 192, 197, 198, 200-4, 206, 208, 
309, 212, 214. 216, 218, 220, 2235, 227, 
229, 231, 235, 239, 240, 249, 252, 260 
piscivorus leucostoma, 189, 192 
Piscivorus piscivorus, 189, 192 
thodostoma, 189, 214, 216, 218, 223-5, 229, 
231, 235, 239, 240, 249, 260 
steauchi, 189, 192, 206, 216, 218, 224-5, 229, 
235, 239, 249 
Aletes centiquadrus, 340, 350 
Algae 
calcareous, 339, 341, 344, 346, 348, 352 
Chondria nidifica, 285-300 
elk kelp, Pelagophycus, 301-308 
Amara jacobina, 275 
Ammospermophilus leucurus canfieldae, 103 
leucurus extimus, 103 
leucurus insularis, 103 
leucurus leucurus, 102 
leucurus peninsulae, 102 
Amphispiza belli belli, 182 
bilineata bangsi, 59, 60, 182 


bilineata belvederei, 60 
bilineata cana, 59 
bilineata carmenae, 60, 182 
bilineata deserticola, 182 
bilineata tortugae, 60 
Anadara multicostata, 340-42, 344, 346, 348 
cf. A. multicostata, 346, 348 
Anomalocardia subimbricata tumens, 340-41, 342, 
348 
Antigona isocardia, 343, 348 
Antilocapra americana americana, 151 
americana peninsularis, 151] 
Anthus spinoletta pacificus, 58 
Antilope americana, 151 
Antrozous minor, 98 
pallidus pacificus, 98 
Aphriza virgata, 54 
Apolymetis cognata clarki, 340-42, 344, 348 
Arctocephalus philippii townsendi, 149 
townsendi, 149 
Ardea herodias, 54 
Astraea unguis, 341, 344, 348, 350 
Atalapha teliotis, 97 
Atriplex barclayana, 31 
Auriparus flaviceps, 57 
flaviceps flaviceps, 58 
Aythya afhnis, 54 
Baja California 
Agathymus dawsoni, 64-72, 169-172 
Belvedere Expedition to the Gulf of California, 
1-44, 49-60, 313-332, 333-368 
Birds, 49-60, 177-184 
Cancellaria wigginsi, 362 
Carpelimus salinus, 278 ‘ 
Cerberilla pungoarena, 391 
Chromodoris banksi, 84 
Chromodoris norrisi, 81 
Conualevia alba, 383 
Conulaevia marcusi, 381 
Crotalus mitchelli angelensis, 75 
Eremarionta rowelli bechteli, 327 
fossils, 333-368 
Haliotis fulgens guadalupensis, 375 
mammals, 85-168, 177-184, 397-404 
mollusks, 81-84, 313-332, 333-368, 369-376, 3 


396 
Perognathus arenarius pa ralios, 113 
Perognathus arenarius sabulosus, 114 


Perognathus baileyi mesidios, 112 

Polycera alabe, 388 

Staphylinidae of marine mud flats, 269-284 
Balaena gibbosa, 140 

japonica, 141 

musculus, 141 

novae angliae, 141 


physalus, 


406 SAN Deco Society OF NATURAL History 


Balaenoptera acuto-rostrata, 141 
borealis, 141 
physalus, 140 
Balantiopteryx plicata pallida, 92 
Balanus sp., 339, 345, 352 
tintinnabulum californicus, 345, 346, 352 
trigonus, 342, 346, 352 
Banks, Richard C. 
Birds of the Belvedere Expedition to the Gulf of 
California, 49-60 
Birds and mammals of the voyage of the 
“Gringa’, 177-84 
The mammals of Cerralvo Island, Baja Califor- 
nia, 397-404 
Barbatia bramkampi, 343, 348 
reeveana, 339-42, 344, 348 
Bassariscus astutus insulicola, 144 
astutus octavus, 144 
astutus palmarius, 144 
astutus saxicola, 144 
saxicola, 144 
Basterotia hertleini, 341, 342, 348, 355, 358 
peninsularis, 344, 348, 357 
Bembidion ephippigerum, 273, 274 
Berosus metaliceps, 273, 274 
Berendtia digueti, 328 
Birds 
of Belvedere Expedition, 49-60 
of voyage of “Gringa”, 177-184 
Bledius densissimus, 280 
diagonalis, 274, 277, 280 
ferratus, 273-76, 280, 281, 282 
jacobinus, 275 
ornatus, 273, 274, 276, 277, 281, 282 
punctatissimus, 273, 274, 276, 277, 281, 282 
Bothriechis nigroviridis, 250 
Bothrops, 185-268 
alternata, 189, 193, 214, 218, 220, 223, 227, 
229, 231, 235-7, 241, 246-7, 249 
atrox, 189, 190, 193, 195, 196, 214, 216, 218, 
220) 223, 227) 2208 23k 235-7241, 246, 
247, 249 
bicolor 2411, 246. 250 
bilineatus, 189, 193, 206, 214, 216, 218, 220, 
223, 229, 231, 235-7, 241, 246, 247, 249 
brachystoma, 241, 246, 250 
castelnaudi, 241, 247 
cotiaray 189,°193,0214) 216) 2188220) 222.227, 
229, 231, 235-7, 241, 246, 249 
dunni, 189, 241, 246, 250 
ecdmant) 180) 193. 214 Bileioigia220\ 223. 
227, 229, 231, 236-7, 241, 246, 247, 249 
insularis, 241, 247, 249 
jararaca, 189, 193, 206, 214, 216, 218, 220), 
7733, Pf DIR MBN, PB, LENS, 23/2, PEN 
246, 247, 249 
jararacussu, 189, 193, 214, 216, 218, 220, 223, 
227, 229, 231, 235-7, 241, 246, 247, 249 
lanceolatus, 189, 193, 214,216, 218 220) 223, 
227, 229, 231, 235-7, 241, 246, 247, 249 
lateralis, 241, 246, 250 
nasuta, 189, 193, 201, 214, 218, 223, 227, 
235-7, 241, 246, 249 
neuwiedi, 189, 193, 201, 214, 216, 218, 220, 
223, 227-229, 23, 235-7.) 24 246-70 249 


nigroviridis, 241, 246, 250 
nummifer, 189, 193, 197, 199, 200, 203, 204, 
214, 218, 220, 223, 227-9, 231, 235-7, 241, 
246, 250 
picadoi, 241, 247, 249 
schlegeli, 189, 193, 197, 199, 200, 203-4, 206-8, 
214, 216, 218, 220, 223..227-98 231, 235-7, 
241, 246, 250 
Bovidae, 151-2 
Brattstrom, Bayard H. 
Evolution of the pit vipers, 185-268 
Broteas alleni, 34, 36, 37 
bryozoa, 339, 352 
Bubo virginianus, 56 
Buccinum lugubre, 361 
Bulimulus carmen, 322-24 
ceralboensis, 327 
chamberlini, 36, 318, 319 
dealbatus, 317, 325 
dentifer, 319 
exilis, 316 
guadalupensis, 316 
johnstoni, 326 
juarezi, 317 
lamellifer, 321, 322 
lardeus, 329 
montezuma, 317 
nitidulus, 329 
sanmarcosensis, 321, 322 
santacruzensis, 325, 326 
slevini, 317, 325, 326 
spirifer, 317, 318, 324 
sufflatus, 317 
veseyianus, 324 
vesicalis, 317 
ximenez, 322, 323, 324 
Bulla cf. B. punctulata, 340, 350 
Bursa californica, 40, 350 
Buteo jamaicensis, 54 
Cadlina evelinae, 380, 381, 382, 386 
Calamospiza melanocorys, 59 
calcarous algae, 339, 341, 344, 346, 348, 352 
Calliostoma bonita, 344, 350 
eximium, 344, 350 
Callorhinus ursinus cynocephalus, 148 
Calypte costae, 56 
Cameronium flavipenne, 176 
liebmanni, 176 
obockianum, 176 
sonorensis, 174, 175-6 
Campylorhynchus brunneicapillus, 58 
Cancellaria ellipsis, 362 
islacolonis, 362 
obesa coronadoensis, 340, 350 
oblonga, 362 
peruana, 362 
pulchra, 340, 350 
tessellata, 362 
wigginsi, 335, 344, 350, 362, 363 
Canis clepticus, 142 
latrans clepticus, 142 
latrans mearnsi, 142 
latrans peninsulae, 142 
mearnsi, 142 
peninsulae, 142 


INDEX 


Cantharus anomalus, 340, 350 
lugubris, 361 
pallidus, 344, 350 
cf. pallidus, 354 
Capra hircus, 404 
Cardita affinis californica, 341, 348 
megastropha, 340-41, 344, 348 
Cardium biangulatum, 340-44, 348 
consors, 340, 344, 348 
elense, 340-41, 342, 344, 348 
Carpelimus confinis, 273, 274, 277, 279, 280, 281 
282 
debilis, 273, 274, 277-79, 280, 281, 282 
providus, 278 
salinus, 274, 278, 279, 280 
Carpodacus mcgregori, 182 
mexicanus frontalis, 59 
mexicanus ruberrimus, 59 
Casmerodius albus, 54 
Castor canadensis repentinus, 124 
Cathartes aura, 54 
Catherpes mexicanus conspersus, 58 
Catoptrophorus semipalmatus, 55 
Cenchris mokeson, 249 
Centruroides exilicauda, 14, 23, 30, 36 
Centurus uropygialis brewsteri, 57 
uropygialis sulfuriventer, 57 
uropygialis tiburonensis, 57 
uropygialis uropygialis, 57 
Cepolis cepa, 324 
Cerberilla ambonensis, 393 
asamusiensis, 393 
pungoarena, 390, 391-93 
tanna, 393 
Cerithium gemmatum, 341, 344, 350 
maculosum, 340, 341, 344, 346, 348, 350 
sculptum, 340, 350 
uncinatum, 343, 350 
Cervidae, 150 
Chama buddiana, 348 
frondosa, 340, 348 
Chilomeniscus sp., 24 
Chione californiensis, 337, 340-342, 344, 348 
cf. californiensis, 348 
picta, 348, 357, 359 
undatella, 337, 348 
Ghlaeays abietis, 337, 342.43, 34547, 349, 354, 
358 
aff. abietis, 349 
cf. abietis, 343, 349 
circularis, 343, 349, 355 
revellei, 345-47, 349 
sp., 337, 343, 349 
tumbezensis, 339, 349, 355 
Chlorura chlorura, 59 
Choeronycteris mexicana, 93 
Chondria dasyphylla, 288, 290, 291 
nidifica, 286-300 
tenuissima, 288, 290 
Chromodoris banksi, 82, 84 
norrisi, 81, 82-84 
Cicindela gabbi, 275 
Citellus beecheyi nudipes, 103 
beecheyi rupinarum, 103 
tereticaudus apricus, 104 
tereticaudus vociferans, 104 


> 


407 
Clypeaster bowersi, 346, 347. 352. 364. 365 
sp., 343, 352, 364 
speciosus, 339, 352 


cf. speciosus, 346, 35 
testudinalis, 364 
testudinarius, 364 
Cnemidophorus tigris martyris, 1] 
Cochemiea poselgeri, 30 
Codakia distinguenda, 340, 341, 343, 344, 348 
349 
Coelocentrum digueti, 328 
insulare, 328 
oweni, 328-30 
vanduzeei, 328, 330 
Colaptes cafer, 181 
chrysoides, 181 
Coleoptera, see Staphylinidae 
Coluber lanceolatus, 249 
Collier, Clinton L., and Wesley M. Farmer 
Additions to the nudibranch fauna of the east 
Pacific and the Gulf of California, 377-96 
Comstock, John Adams 
The early stages of Stenaspilates apapinaria 
Dyar (Lepidopetra: Geometridae), 45-48 
The larva and pupa of Agathymus dawsoni 
(Lepidoptera: Megathymidae), 169-72 
Conualevia, 381-88 
alba, 383-85, 387, 388 
marcusi, 381-83, 385-386, 388 
Conualevinae, 381 
Conopeum commensale, 346, 352 
Conus brunneus, 344, 350 
diadema, 340, 350 
dispar, 343. 344, 350 
gladiator, 340, 343, 350 
nux, 344, 350 
patricius, 344, 350, 364 
perplexus, 344, 348, 350 
princeps, 340, 341, 344, 350 
purpurascens, 340, 344, 346, 350 
pyriformis, 364 
regularis, 340, 344, 350 
scalaris, 340, 350 
sp., 345, 350 
ximenes, 340, 350 
Cophias wagleri, 251 
Corbula bicarinata, 341, 349 
Corvus corax, 57 
Corynorhinus macrotis pallescens, 97 
Coulterella capitata, 27 
Crassispira cf. nymphia, 341, 350 
Creophilus maxillosus, 309 
Crepidula arenata, 340, 350 
onyx, 340, 351 
Cricetidae, 124-136 
Crotalus sp., 403 
adamanteus, 190, 194, 195, 197, 201, 203, 209, 
213, 215, 217, 219, 221, 226, 228, 230, 
232-234, 241, 242, 244, 245, 250, 252 
atrox, 12, 13, 35, 80, 190, 194, 195, 197, 199, 


732: 213. 215, Ziv 22), ees ee eo 
232-234, 241-244, 250 

basiliscus, 190, 194, 201, 203, 209, 212-215, 
217. 221, 226, 228, 230, 232-234, 242, 
245, 250 


catalinensis, 27, 80, 241, 250 


408 


cerastes, 190, 194, 196, 197, 198, 199, 212, 213, 
215, 217, 221, 226, 228, 230, 232-234, 242, 
244-245, 250 

cerastes cerastes, 190, 193, 228 

cerastes laterorepens, 190, 193, 215, 228 

confluentus mitchellii, 75-80 

durissus, 190, 194-196, 198, 201, 209, 213-215, 
217, 221-223, 226, 228, 230, 232:234, 242. 
243, 245, 250 

durissus durissus, 190 

durissus terrificus, 80, 190, 193, 241 

durissus tzabcan, 190, 193, 207, 209, 211, 212 

durissus unicolor, 190, 193, 219, 241 

enyo, 190; 194, 2095 213, 215, 217, 221, 226; 
228, 230, 232-234, 242, 245, 250 

enyo enyo, 29 

exsul, 80, 241, 250 

giganteus, 212, 241, 250 

horridus, 190, 194, 196, 199, 201-203, 212, 215, 
217, 218, 221, 226, 228, 230, 232-234, 
242, 250 

horridus atricaudatus, 190, 193, 219 

horridus horridus, 190, 193, 219 

intermedius 190, 194007215, 2308 234° 238 
250 

lepidus, 190, 194, 201, 203, 209, 213, 215, 217, 
221, 226, 228, 230, 232-234, 238, 241, 242, 
250 

lepidus klauberi, 190 

lepidus lepidus, 190 

lepidus muta, 214, 216 

miliarius, 250 

mitchelli, 6, 8, 75-80, 190, 194, 196, 199, 201, 
206, 209. 213, 215) 217.2211 228, 230, 
232-234, 242, 244-245, 250 

mitchelli angelensis, 74, 75-80 

mitchelli mitchelli, 29, 37, 75-80, 190 

mitchelli muertensis, 75-80 

mitchelli pyrrhus, 75-80, 190, 212 

mitchelli stephensi, 75-80, 190 

molossus, 194, 196, 199, 201, 206, 209, 213, 
215, 2170 221, 226,-228: 930, 232.234. 
242. 250 

molossus estebanensis, 80 

molossus molossus, 80, 190 

molossus nigrescens, 190 

mutus, 250 

polystictus, 190, 194, 195, 238, 239, 250 

potterensis, 250 

pricei, 190, 194, 201, 203, 209, 213, 215, 217, 
221, 226, 228, 230, 232-234, 238, 240, 
247 245250 

pusillus, 190, 194, 203, 213, 215, 217, 221, 
226, 228, 230, 232-234, 238, 239, 240, 
242. 250 

pyrrhus, 75-80 

ruber, 8, 21, 190, 194, 195, 201, 204, 206, 209, 
PAG), PNBY. PNG PRN PS, BAS, DIOL Dyn= 
234, 242, 244, 250 

ruber lucasensis, 24, 29, 244 

ruber ruber, 80, 186, 190, 191, 194, 195, 201, 
206, 209, 210, 211, 215, 219, 221, 226, 
228, 230, 232, 233, 234, 242-244 

scutulatus, 190, 194, 199, 209, 212, 213, 215, 
217; 219,221 226, 228, 230, 9232.234, 
242, 250 


scutulatus scutulatus, 80 


? 


San DzeGo Society oF NATURAL History 


stejnegeri, 190, 194, 195, 234, 238, 239, 250 

tioriss 190M 9 4 99209 21S ee2lo ela 2 lle 
226, 228, 230, 232-234, 242, 244, 245, 250 

tigris mitchellii, 75-80 

tortugensis, 18, 80, 190, 194, 201, 203, 204, 
2135 Z1Ds 275221, 2262228. 2305232-234. 
241, 242, 244, 250 

transversus, 250 

triseriatus, 190, 194, 201, 203, 204, 213, 215, 
DW ZS 2222 Oy 22 Bee 23 ON 252-254 
240, 242, 243, 249 

unicolor, 80, 241, 250 

viridis, 190, 192-196, 198, 199, 201, 203, 206, 
230, 232-234, 242-245, 250 

viridis caliginis, 80 

viridis concolor, 209 

viridis decolor, 190, 193, 194, 217, 221, 226, 
228 

viridis helleri, 80, 190, 192, 193, 194, 212-215, 
ZW iee22le 2260228245 

viridis lutosus, 190, 193, 194, 212, 213, 215, 
ING, 2PM, PES, DES) 

viridis oreganus, 190, 192-194, 213, 215, 217, 
218, 219, 221, 226, 228 

viridis viridis, 190, 193, 194, 209, 213, 215, 
PMH, LB PAS, PBs} 

willardi, 190, 194-196, 203, 209, 213, 215, 217, 
221, 226, 228, 230, 232-234, 241, 242, 250 


Crucibulum scutellatum, 341, 344, 351 
Ctena mexicana, 340, 341, 349 
Ctenosaura hemilopha, 403 


hemilopha conspicuosa, 14 


Cyathodonta undulata, 342, 349 
Cylindrella irregularis, 328 


turris, 328 


Cypraea albuginosa, 348, 351 


annettae, 341, 343, 344, 348, 351 
arabicula, 340, 351 


Cypraecassis coarctata, 340, 351 
Dasypterus ega xanthinus, 97 
Dawson, E. Yale, and Bilgin Tozin 


Structure and reproduction of the red alga 
Chondria nidifica Harvey, 285-300 


Dawson, E. Yale 


See Parker, Bruce C. 


Delphinus bairdii, 138 
Dendraster casseli, 345, 352, 363, 365 


granti, 365 

cf. granti, 345, 352, 363, 365 
vizcainoensis, 365 
vizcainoensis similaris, 365 


Dendrodoris atropos, 389, 390, 391 


nigra, 391 


Dendrocopos scalaris cactophilus, 57 


scalaris lucasanus, 57 
scalaris sinaloensis, 57 
scalaris soulei, 57 


Dentalium semipolitum, 344, 352 
Didelphis marsupialis virginiana, 91 


virginiana, 91 


Diodora alta, 341, 351 


inaequalis, 341, 348, 351 


Diomedea nigripes, 181 
Diplodonta sericata, 342, 349 


subquadrata, 340, 349 


INDEX 


Dipodomys agilis cabezonae, 119 
agilis latimaxillaris, 119 
agilis martirensis, 119 
agilis peninsularis, 120 
agilis plectilis, 120 
agilis simulans, 119 
antiquarius, 121 
deserti deserti, 124 
gravipes, 121 
insularis, 123 
margaritae, 123 
merriami annulus, 122 
merriami arenivagus, 121 
merriami brunensis, 123 
merriami Ilanoensis, 123 
merriami melanurus, 123 
merriami merriami, 121 
merriami platycephalus, 122 
merriami quintinensis, 122 
merriami semipallidus, 122 
merriami trinidadensis, 121 
paralius, 120 
peninsularis australis, 120 
peninsularis eremoecus, 120 
peninsularis pedionomus, 120 
peninsularis peninsularis, 120 
platycephalus, 122 
simulans peninsularis, 120 
Dipsosaurus catalinensis, 27 
Distichlis, 273, 274 
Divaricella eburnea, 341, 342, 344, 349 
Dosinia ponderosa, 344, 349 
Drupa lugubris, 361 
Echinanthus testudinarius, 364 
Echinocereus grandis, 14 
Echinometra vanbrunti, 348, 352 
Emballonuridae, 92 
Emerson, William K., and Leo George Hertlein 
Invertebrate megafossils of the Belvedere Expedi- 
tion to the Gulf of California, 333-368 
Emerson, William K., and Morris K. Jacobson 
Terrestrial mollusks of the Belvedere Expedition 
to the Gulf of California, 313-332 
Enaeta cumingii, 341, 351 
Encope california, 339, 341, 344, 352, 365-67 
grandis, 344, 352, 366, 367 
grandis inezana, 339, 340, 352 
micropora, 339, 367 
sp., 339, 342, 352 
Endomychura craveri, 9, 54, 55 
Enhydra lutris nereis, 147 
Enochrus hamiltonti pacificus, 274 
Eptesicus fuscus bernardinus, 96 
fuscus pallidus, 96 
fuscus peninsulae, 97, 399 
pallidus, 96 
Erolia minutilla, 55 
Eryngium nasturtiifolium, 23 
Eschrichtius gibbosus, 140 
glaucus, 140 
Eucidaris thouarsii, 337, 339, 344, 352 
Eutamias merriami meridionalis, 102 
merriami merriami, 102 
merriami obscurus, 102 
Falco peregrinus, 54 


409 


Farmer, Wesley M. 


Two new opisthobranch mollusks from Baja 
California, 81-84 
see also Collier, Clinton L. 
Fasciolaria princeps, 341, 351 


Felis aztecus browni, 147 
californica, 147 
concolor browni, 147 
concolor californicus, 147 
concolor improcera, 148 
domesticus, 403 
improcera, 148 

Ferocactus diguetii, 25, 

Ficus palmeri, 15, 31 

Fregata magnificens, 53 

Fusinus cinereus, 341, 351 
luteopictus, 361 

Fusus luteopictus, 361 

Geometridae 
Stenaspilates apapinaria, 45-48 

Geomyidae, 105 

Globicephala scammonii, 139 

Glycymeris gigantea, 340, 341, 344, 349 
multicostata, 340, 341, 344, 348, 349 

Gracilariopsis andersonii, 287 
sjoestedtii, 287 

Gulf of California 
see Baja California, Sonora 

Gyrineum spinosum, 360 
strongi, 341, 344, 348, 351, 360, 363 

Hadrotes crassus, 309-312 

Hadrurus sp., 14, 21 

Haematopus palliatus, 54 

Haliotis cracherodii californiensis, 375 
elsmerensis, 372 
fulgens, 340, 369-376 
fulgens fulgens, 373 
fulgens guadalupensis, 374, 375 
fulgens turveri, 373 
lasia, 372 
lomaensis, 372 
planilirata, 371, 373, 375 
splendens, 373 
varia, 372 
walallensis, 372 

Hannarabdotus, 325 

Harbison, Charles F. 

A second new species of megathymid from Baja 
California, Mexico (Lepidoptera: Megathy- 
midae), 61-72 

Heliacus robertsae, 341, 351 

Helix exilis, 316 


facta, 327 


Hertlein, Leo George 


see Emerson, William K. 


27, 35 


~ a ~—-~ > l 5 
Hesperomy s eremicus, yA 
gambelii, 127 
sonoriensis, 128 


Heteromyidae, 110 
Hipponix antiquatus, 341 
Huey, Laurence M. 
The mammals of Baja C lifornia, Mex 
168 
Hyla regilla 


Hylocharis xan 


410 SAN Drieco Society oF NATuRAL HIstory 


Hypsiglena, 27 
Icterus cucullatus trochiloides, 59 
spurius, 182 
Irus ellipticus, 357 
Isognomon janus, 340, 341, 349 
Jacobson, Morris K. 
see Emerson, William K. 
Janczewskia lappacea, 287 
Jeanneretia subtussulcata, 324 
Juncus acutus, 274 
Kerivoula pallida, 95 
Klauber, L. M. 
A new insular subspecies of the speckled rattle- 
snake, 73-80 
Knefastia funiculata, 341, 344, 351 
Koeberlinia spinosa, 10 
Kogia breviceps, 138 
Rachesis: mura, 1180) 191) 10472028 21845220) 
222-3, 227, 229, 231, 235, 250, 253 
Lagenorhynchus obliquidens, 139 
Lanius ludovicianus grinnelli, 58 
Larus californicus, 181 
delawarensis, 55, 181 
heermanni, 55, 181 
eccidentalis, 55, 181 
philadelphia, 55, 181 
Lasiurus borealis teliotis, 97 
cinereus cinereus, 97 
ega xanthinus, 97 
Latax lutris nereis, 147 
Latirus lugubris, 361 
Lemaireocereus thurberi, 30 
Lepidoptera 
see Geometridae, Megathymidae 
Leporidae, 99 
Leptotyphlops, 23 
Lepus alleni tiburonensis, 11 
arizonae confinis, 100 
bennettii, 100 
californicus bennettii, 100 
californicus deserticola, 101 
californicus magdalenae, 101 
californicus martirensis, 101 
californicus sheldoni, 101 
californicus xanti, 101 
cerrosensis, 99 
cinerascens, 99 
floridanus sanctidiegi, 100 
insularis, 37, 101 
martirensis, 101 
peninsularis, 100 
sylvaticus, 100 
texianus deserticola, 101 
Leucophoyx thula, 54 
Lichanura trivirgata, 11 
Lima tetrica, 340, 349 
Lindsay, George E. 
The Belvedere Expedition to the Gulf of 
California, 1-44 
Loomelania melania, 52 
Lophortyx californicus, 181 
gambelit pembertoni, 54 
Lucina lampra, 340, 342, 344, 349 
nuttalli, 340, 342, 344, 349 
Lynx baileyi, 148 
rufus baileyi, 148 


rufus californicus, 148 
rufus peninsularis, 148 
Lyrocarpa linearifolia, 14 
Machaerocereus gummosus, 10, 30 
Macrotus californicus, 93 
Mammals 
of Baja California, 85-168 
of Cerralvo Island, 397-404 
of voyage of “Gringa’”’, 177-184 
Perognathus arenarius paralios, 113 
Perognathus arenarius sabulosus, 114 
Perognathus baileyi mesidios, 112 
Mammillaria albicans, 30, 36 
Marsilea fournieri, 23 
Masticophus flagellum, 403 
Mazama hemionus peninsulae, 150 
Megaceryle alcyon, 56 
Megapitaria squalida, 340-44, 349 
Megaptera novaeangliae, 141 
Megathymidae 
Agathymus dawsoni, 64-70, 169-172 
Melanitta perspicillata, 54 
Meoma cf. M. grandis, 339, 352 
Mephitis estor, 146 
mephitis estor, 146 
mephitis holzneri, 147 
occidentalis holzneri, 147 
Mergus serrator, 54 
Mexico 
see Baja California, Sonora 
Micrarionta desertorum, 327 
rowelli bechteli, 327, 330 
rowelli mearnsi, 327, 328 
rowelli mexicana, 327, 328 
Microtus californicus aequivocatus, 136 
californicus grinnelli, 136 
californicus huperuthrus, 136 
californicus hyperythrus, 136 
californicus neglectus, 136 
californicus sanctidiegi, 136 
Miltha xantusi, 340, 349 
Mimus polyglottos, 58 
Mirounga angustirostris, 149 
Modulus cerodes, 341, 351 
Mollusks 
fossil, 333-368 
terrestrial, 313-332 
see nudibranchs 
Molossus mexicanus, 98 
Moore, Ian 
A new marine beetle from the Gulf of California 
(Coleoptera: Staphylinidae), 173-176 
The staphylinidae of the marine mud flats of 
southern California and northwestern Baja 
California (Coleoptera), 269-284 
The larva of Hadrotes crassus (Mannerheim) 
(Coleoptera: Staphylinidae) , 309-312 
Morula didyma, 361 
ferruginosa, 341, 351 
lugubris, 341, 351, 356, 361 
papillosa, 360 
Murex cf. M. elenensis, 341, 351 
gyrinus, 360 
Muricanthus princeps, 346, 351 
Muridae, 137 
Mustela frenata latirostra, 145 


Mus alexandrinus, 137 
domesticus, 137 
musculus domesticus, 137 
norvegicus, 137 
rattus, 137 

Myiarchus cinerascens cinerascens, 57 
cinerascens pertinax, 57 


Myotis californicus californicus, 95 
californicus pallidus, 95 
californicus stephensi, 95 
evotis evotis, 94 
longicrus interior, 95 
micronyx, 94 
orinomus, 95 
peninsularis, 94 
subulatus melanorhinus, 95 
thysanodes thysanodes, 94 
velifer peninsularis, 94 
vivesi, 95 
volans volans, 94 
volans interior, 95 
yumanensis lambi, 94 
yumanensis sociabilis, 93 
yumanensis yumanensis, 93 

Nassarius sp., 341, 351 
tiarula, 341, 342, 344, 351 

Natalus mexicanus, 93 
stramineus mexicanus, 93 

Natica unifasciata, 342, 351 

Neotoma abbreviata, 134 
albigula venusta, 131 
anthonyi, 135 
arenacea, 135 
bella felipensis, 132 
bryanti, 135, 184 
bunkeri, 135 
fuscipes macrotis, 135 
fuscipes martirensis, 135 
insularis, 132 
intermedia, 132 
intermedia gilva, 132 
intermedia notia, 134 
intermedia perpallida, 134 
intermedia pretiosa, 134 
intermedia ravida, 133 
intermedia vicina, 134 
lepida abbreviata, 134 
lepida arenacea, 135 
lepida aridicola, 133 
lepida egressa, 132 
lepida felipensis, 132 
lepida gilva, 132 
lepida insularis, 132 
lepida intermedia, 132 
lepida latirostra, 134, 184 
lepida marcosensis, 133 
lepida molagrandis, 133 
lepida notia, 134 
lepida nudicauda, 133 
lepida perpallida, 134 
lepida pretiosa, 134, 183 
lepida ravida, 133 
lepida vicina, 37, 134 
macrotis, 135 
martinensis, 135 


INDEX 


nudicauda, 133 

venusta, | | 
Nereocystis luetkeana, 307 
Nerita scabriocosta, 344, 35] 


Notiosorex crawfordi crawfordi. 92 


nudibranchs, 81-84, 377-396 

Nyctinomus femorosaccus, 98 

Ochthebius interruptus, 273, 274 
sp. near rectus, 273, 274 


Odocoileus cerrosensis, 150 
hemionus cerrosensis, 150 
hemionus fuliginatus, 150 
hemionus peninsulae, 30, 150 
hemionus sheldoni, 11 

Oliva incrassata, 344, 351 
polpasta, 342, 351 
porphyria, 341, 351 
spicata, 341, 343, 344, 348, 351 

Olivella dama, 341, 342, 344, 351 
Spica 351 

Ondatra zibethicus bernardi, 136 

Onychomys macrotis, 131 
pulcher, 130 
ramona, 131 
torridus macrotis, 131 
torridus pulcher, 130 
torridus ramona, 131 

Opuntia cholla, 30 

Orca rectipinna, 139 

Orcinus rectipinna, 139 

Oreoscoptes montanus, 58 

Oryzomys peninsulae, 124 


Ostrea angelica, 337, 341-343, 345, 346, 349, 


356 

cf. angelica, 353 

californica, 339, 353, 354 

cf. californica, 337, 349, 353 

chilensis, 354 

corteziensis, 354 

cumingiana, 353 

englekeyi, 354 

heermanni, 347, 354 

aff. heermanni, 346, 349 

iridescens, 353 

megodon, 340, 345, 349 

palmula, 340, 349 

sp., 337, 349 

vespertina, 353, 356 

virginica var. californica, 353 
Otraria californiana, 149 

gillespii, 149 
Ovis canadensis cremnobates, 151 

canadensis weemsi, 152 

cervina cremnobates, 151 
Pachycereus pringlei, 5, 12, 25, 27 
Pachycormus discolor, 5 
Pseudochama saavedrai, 344, 349 
Paleontology 

Invertebrate megafossils, 333-368 
Pandion haliaetus, 54, 181 
Parametaria dupontii, 341, 348, 351 
Parker, Bruce C., and E. Yale Dawson 


, 30, 44 


411 


353, 


Notes on variability and range in the elk kelp 


Pelagophycus, 301-308 


412 San Deco Society OF NATURAL History 


Passerculus sandwichensis, 59 
sandwichensis sanctorum, 182 
Pecten abietis, 355 
aspersus, 355 
cf. carrizoensis, 345, 349 
marquerensis, 342, 345, 349, 358 
sp., 346, 349 
cf. stearnsi, 343, 349 
subnodosus, 339-340, 343-346, 349 
tumbezensis, 355 


vogdesi, 339, 340, 342, 344, 347, 349 


Pelagophycus giganteus, 303, 304-307 
porra, 303-307 
Pelecanus occidentalis, 19 
occidentalis californicus, 52, 53 
Pelamis platurus, 24 
Peniocereus johnstonii, 27 
Periglypta multicostata, 344, 349 
Perityle robusta, 31 
Perodipus cabezonae, 119 
simulans peninsularis, 120 
streatori simulans, 119 
Perognathus anthonyi, 116 
arenarius, 399, 401-402 
arenarius albescens, 113 
arenarius albulus, 115, 401-402 
arenarius ambiguus, 113 
arenarius ammophilus, 115 
arenarius arenarius, 114, 400-402 
arenarius helleri, 113 
arenarius mexicalis, 113 
arenarius paralios, 113 
arenarius sabulosus, 114 
arenarius siccus, 115, 399-401 
arenarius sublucidus, 114, 400-401 
baileyi extimus, 112 
baileyi fornicatus, 112, 183 
baileyi hueyi, 111 
baileyi mesidios, 112 
baileyi rudinoris, 112 
bombycinus, 111 
bryanti, 118 
californicus femoralis, 116 
californicus mesopolius, 116 
evermanni, 117 
fallax fallax, 115 
fallax inopinus, 116 
fallax majusculus, 116 
fallax pallidus, 115 
fallax xerotrophicus, 116 
femoralis mesopolius, 116 
formosus cinerascens, 111 
formosus infolatus, 111 
formosus mesembrinus, 111 
helleri, 113 
knekus, 112 
longimembris aestivus, 110 
longimembris bombycinus, 111 
longimembris internationalis, 110 
longimembris venustus, 110 
margaritae, 119 
mesembrinus, 111 
penicillatus albulus, 115 
penicillatus ammophilus, 115 
penicillatus angustirostris, 113 
penicillatus arenarius, 400-401 


penicillatus siccus, 115, 399 
spinatus, 56, 183 
spinatus broccus, 118 
spinatus bryanti, 118 
spinatus evermanni, 117 
spinatus guardiae, 117 
spinatus lambi, 119 
spinatus latijugularis, 118 
spinatus magdalenae, 119 
spinatus marcosensis, 118 
spinatus margaritae, 119 
spinatus nelsoni, 118 
spinatus occultus, 118 
spinatus oribates, 117 
spinatus peninsulae, 119 
spinatus prietae, 117 
spinatus pullus, 118 
spinatus rufescens, 117 
spinatus seorsus, 118, 183 
spinatus spinatus, 117 
Peromyscus boylii rowleyi, 130 
californicus insignis, 125 
caniceps, 127 
cedrosensis, 126 
cineritius, 129 
crinitus pallidissimus, 125 
crinitus stephensi, 125 
dickeyi, 127 
dubius, 128 
eremicus avius, 126, 402-403 
eremicus carmeni, 126, 183 
eremicus cedrosensis, 126, 183 
eremicus cinereus, 126 
eremicus eremicus, 125 
eremicus eva, 127 
eremicus fraterculus, 126 
eremicus insulicola, 126 
eremicus pullus, 183 
eremicus polypolius, 126 
eremicus propinquus, 127 
eva, 127 
exiguus, 128 
gaurus, 130 
geronimensis, 129 
guardia, 14 
guardia guardia, 127 
guardia interparietalis, 127 
guardia mejiae, 127 
hemionotis, 130 
insignis, 125 
leucopus coolidgei, 129 
maniculatus assimilis, 128 
maniculatus cineritius, 129 
maniculatus coolidgei, 128 
maniculatus dorsalis, 129 
maniculatus dubius, 128 
maniculatus exiguus, 128, 183 
maniculatus gambelii, 127 
maniculatus geronimensis, 129, 183 
maniculatus hueyi, 129 
maniculatus magdalenae, 129 
maniculatus margaritae, 129 
maniculatus martinensis, 128 
maniculatus sonoriensis, 128 
cresterus, 128 
pseudocrinitus, 125 
sejugis, 36, 130 


slevini, 130 

stephensi, 125 

texanus medius, 128 

truei lagunae, 130 

truei martirensis, 130 
Petricola bulbosa, 359 

robusta, 340, 349, 359 

sinuosa, 359 

sp., 359 

venusta, 359 
Phacelia pauciflora, 8 
Phaethon aethereus, 52 
Phainopepla nitens, 58 
Phalacrocorax auritus, 53 

penicillatus, 53 
Phalaenoptilus nuttallii, 56 
Phoca richardii geronimensis, 149 

vitulina geronimensis, 149 
Phocoena sinus, 140 

vomerina, 140 
Phyllodactylus, 34, 36 
tuberculosus, 30 
Phyllostomidae, 93 
Physeter breviceps, 138 

catodon, 138 


INDEX 


Pinctada mazatlanica, 340, 341, 344, 346, 348, 349 


Pipistrellus hesperus australis, 37, 96, 183, 399 


hesperus hesperus, 96 
hesperus merriami, 96 
Piranga ludoviciana, 59 
Pitar newcombianus, 344, 349 
Pituophis catenifer affnis, 11 
Pizonyx vivesi, 16, 32, 95 
Placunanomia cumingii, 346, 349 
Plecotus townsendii pallescens, 97 
Plicatula sp., 339, 349 
Pocillopora robusta, 344, 346, 348, 352 
Podiceps caspicus, 51 
Polinices bifasciata, 341, 344, 351 
uber, 341, 344, 351 
cf. uber, 343, 351 
Polioptila caerulea obscura, 58 
melanura curtata, 58 
melanura lucida, 58 
melanura margaritae, 182 
Polycera alabe, 387, 388-389 
atra, 389 
hedgpethi, 389 
zosterops, 389 
Polymesoda sp., 44, 349 
Porites californica, 22, 341, 348, 352 
sp., 344, 347, 352 
Portulaca pilosa, 31 
Procyon lotor californicus, 145 
lotor grinnelli, 145 
lotor pallidus, 144 
lotor psora, 145 
pallidus, 144 
psora, 145 
Protothaca sp., 357 
Ptychoramphus aleutica aleutica, 181 
aleutica australe, 181 
Pufhnus puffinus opisthomelas, 181 
Pupa albilabris, 329 
beltiana, 329 
fallax, 329 


Pupoides albilabris, 329, 330 
catalinensis, 330 
marginatus, 329, 330 
Rabdotus baileyi, 325 
beldingi, 325 
ceralboensis, 325, 327, 330 
chamberlini, 318, 379. 320. 330 
dentifer, 319, 320, 330 
elatus, 325 
excelsus, 325 
hannai, 325 
inscendens, 325 
johnstoni, 325, 326, 330 
lamellifer, 319-324 
lamellifer lamellifer, 320, 327, 322, 330 


lamellifer ximenez, 320, 322, 323, 324. 330 


pallidior, 325 
sanmarcosensis, 32], 322 
santacruzensis, 325 
slevini, 325, 326, 330 
sufflatus, 317, 318, 330 
vegetus, 325 
veseyianus, 320, 324, 330 
Rattus norvegicus, 137 
rattus alexandrinus, 137 
rattus rattus, 137 
Reithrodon longicauda, 124 
megalotis, 124 


Reithrodontomys megalotis longicaudus, 124 


megalotis megalotis, 124 
megalotis peninsulae, 125 
peninsulae, 125 
Reptiles 
Crotalus mitchelli angelensis, 75 
evolution of pit vipers, 185-268 
Rhachianectes glaucus, 140 
Richmondena cardinalis ignea, 59 
cardinalis townsendi, 59 
Salicornia, 273, 274, 278 
Salpinctes obsoletus obsoletus, 58, 181 
obsoletus tenuirostris, 181 
Sator angustus, 35 
grandaevus, 33 
Sauromalus ater, 31 
hispidus, 7, 8 
varius, 14 
Scapanus anthonyi, 92 
latimanus anthonyi, 92 
latimanus occultus, 92 
Sciurus carolinensis carolinensis, 104 
fossor anthonyi, 104 
griseus anthonyi, 104 
hudsonius mearnsi, 105 
Scotophilus hesperus, 96 
Semele cf. bicolor, 341, 350 
flavescens, 341, 350 
cf. flavescens, 340, 350 
verrucosa, 340, 350, 356, 359 
Sibbaldus musculus, 141 
Sideroxylon leucophyllum, 14 
Sigmodon hispidus eremicus, 13! 
Siren cynocephala, 148 
Sistrurus, 185-268 


catenatus, 189, 
15. 217. 221. 226, 228, 


196. 200, 203 209 


413 


414 SAN Dteco Society oF NATURAL History 


catenatus catenatus, 189, 219 Tachys vittiger, 273, 274 
catenatus tergeminus, 189, 200 Tadarida brasiliensis mexicana, 98 
miliarius, 189, 196, 200, 203, 209, 213, 215, femorosacca, 98 
217, 221, 226, 228, 232-233, 241-243, 250 macrotis 99 
miliarius streckeri, 189 mexicana, 98 
ravus, 189, 196, 200, 203,.209) 213, 215, 217, melee, 6O 
218, 221, 226, 228, 230, 232-234, 239, Tagelus californianus, 344, 350 
240, 250 politus, 342, 350 
Sitomys americanus thurberi, 127 sp., 342, 350 
martirensis, 130 subteres, 344, 350 
rowleyi, 130 Talmadge, Robert R. 
Solecardia eburnea, 341, 344, 350 The races of Haliotis fulgens Philippi 
Solenastrea ecuadoriana, 347 (Mollusca: Gastropoda), 369-376 
fairbanksi, 347 Talpidae, 92 
sp., 347 Tamias asiaticus merriami, 102 
Sonora leucurus, 102 
Cameronium sonorensis, 175 leucurus peninsulae, 102 
Sorex californicus juncensis, 91 obscurus, 102 
crawfordi, 92 Tamiasciurus douglassii mearnsi, 105 
juncensis, 91 Tantilla sp., 23 
lagunae, 91 Tapes squamosa, 357 
oreinus, 91 Taxidea americana neglecta, 145 
ornatus lagunae, 91 berlandieri, 145 
ornatus ornatus, 91 taxus berlandieri, 145 
Southern California taxus infusca, 146 
Staphylinidae of marine mud flats, 269-284 taxus neglecta, 145 
Speotyto cunicularia, 56 Tegula mariana, 341, 351 
Spermophilus atricapillus, 103 Tellidora burneti, 342, 350 
beecheyi nudipes, 103 Tellina amianta, 342, 350 
beecheyi rupinarum, 103 cumingii, 340, 344, 350 
grammurus atricapillus, 103 meropsis, 341, 350 
tereticaudus apricus, 104 reeluca 3400344050 
tereticaudus tereticaudus, 104 Siowlees. 342, 344, 350 
Sphaeralcea hainesii, Lees viridotincta, 340, 341, 350 
Spilogale arizonae martirensis, 146 Tein Carne, Ss 25 
lucasana, 146 cf. specillata, 341, 344, 351 
ae ak variegata, 341, 344, 351 
pales Luersaia, Ue Thincbius fcizzelli, 273, 274, 2797280) 282 
putorius martirensis, 146 See: : ’ 
Spizella passerina, 60 Thinopinus pictus, 309 
Spondylus calcifer, 340, 350 Thomomys albatus, 105 
princeps, 339, 340, 343, 348, 350, aphrastus, 106 
cf. princeps, 345, 346, 350 bottae sspp., see umbrinus sspp. 


fulvus alticolus, 110 


Cameronium sonorensis, 175 fulvus anitae, 110 
Carpelimus salinus, 278 fulvus PSUS 107 
Hadrotes crassus, 309-312 fulvus nigricans, 105 
of marine mud flats, 269-284 magdalenae, 109 
Stenaspilates apapinaria, 45, 46-48 preike poeta 
Strombina maculosa, 344, 351 Sree Shae 105 
eee ye eT was 346, 351 umbrinus alticolus, 110 
acilior, : ' Ge : 11 
granulatus, 341, 343, 344, 348, 351 ce ace ase 


d umbrinus aphrastus, 106 
granulatus form cortezianus, 341, 351 umbrinus borjasensis, 108 


Staphylinidae 


Sula leucogaster, 53 umbrinus brazierhowelli, 107 
nebouxii, 12, 53 umbrinus cactophilus, 108 

Sylvilagus auduboni arizonae, 100 umbrinus catavinensis, 108 
auduboni confinis, 100 umbrinus cunicularius, 106 
auduboni sanctidiegi, 100 umbrinus homorus, 109 
bachmani cerrosensis, 99, 183 umbrinus imitabilis, 109 
bachmani cinerascens, 99 umbrinus incomptus, 109 
bachmani exiguus, 99 umbrinus jojobae, 106 
bachmani howelli, 99 umbrinus juarezensis, 106 
bachmani peninsularis, 100 umbrinus litoris, 109 
bachmani rosaphagus, 99 umbrinus lucidus, 106 


mansuetus, 100 umbrinus magdalenae, 109 


martirensis, 107 
nigricans, 105 
proximarinus, 106 
rhizophagus, 108 
ruricola, 108 
umbrinus russeolus, 109 
umbrinus sanctidiegi, 105 
umbrinus siccovallis, 107 
umbrinus xerophilus, 107 
Throscinus crotchi, 274 
Thryomanes bewickti cerroensis, 181 
Thalasseus elegans, 55 
Toxostoma cinereum, 58 
Toziin, Bilgin 
see Dawson, E. Yale 
Trianthema portulacastrum, 31 
Trimeresurus, 185-268 
acutimentalis, 251 


umbrinus 
umbrinus 
umbrinus 
umbrinus 
umbrinus 


INDEX 


albolabris, 189, 196, 202, 204, 214, 216, 218, 


220) 223, 227, 229, 231, 235, 244, 251 
anamallensis, 251 
borneensis, 251 
cantoris, 251 
chaseni, 251 
convictus, 251 
cornutus, 248, 251 
elegans, 251 
erythrurus, 251 
fasciatus, 251 


flaviviridis, 189, 195-196, 214, 216, 218, 220, 


223, 227, 229, 231, 235-237, 244, 247, 251 
flavomaculatus, 202, 244, 251 
gracilis, 251 
gramineus, 189, 236-237, 244, 247-248, 251 
halieus, 251 
jerdoni, 195, 244, 251 
kanburiensis, 251 
kaulbacki, 251 
labialis, 251 
macrolepis, 251 
malabaricus, 251 
mcgregori, 251 
monticola, 251 


mucrosquamatus, 189, 190, 196, 218, 223, 236- 


237, 244, 247-248, 251 
mutabilis, 251 
okinavensis, 251 
philippinensis, 248, 251 
popiorum, 251 


puniceus, 


189, 196, 202-203, 216, 218, 220, 


2232272, 229, 23 236-237, 244, 240-247, 


251 


purpureomaculatus, 189, 196, 202, 204, 214, 
ZUG AUS 2204 22355 227.9229) 235 23D- 


237, 244, 247-248, 251 
schultzi, 251 


stejnegeri, 189, 196, 202, 204, 218, 220, 223, 


227, 231, 235-237, 244, 247-248, 251 


415 
stigatus, 251 
sumatranus, 25] 
trigonocephalus, 251 
wagleri, 189, 190, 195-197, 200-204, 206, 214, 
Z16, 218; 220; 222, 223, 227. 220: 23 


235-237, 244, 246-248, 251, 255 
Trivia californica, 341, 351 
solandri, 341, 351 
Tropisternis salsamentus, 273, 274 
Tunga caecata, 183 
Turbo fluctuosus, 341, 
sp., 343 
squamiger, 341, 352 
Turritella gonostoma, 341, 352 
cf. imperialis, 354 
Sp), 342, 392 
Tursiops gillii, 139 
nuuanu, 139 
Tyto alba, 55 
Ulva, 277 
Urocyon cinereoargenteus californicus, 143 
cinereoargenteus peninsularis, 143 
cinereoargenteus scottii, 143 
virginianus scottii, 143 
Urosaurus, 34, 37 
microscutatus, 30 
magister, 143 
Uta, 36, 37 
palmeri, 11 
squamata, 27 
Vejovis sp., 30 
Vermivora celata lutescens, 58 
Vesperimus fraterculus, 126 
Vespertilio californicus, 95 
cinereus, 97 
evotis, 94 
fuscus peninsulae, 97 
melanorhinus, 95 
molossus, 99 
pallidus, 98 
volans, 94 
yumanensis, 93 
Vesperugo merriami, 96 
Vulpes arsipus, 142 
macrotis arsipus, 142 
macrotis devia, 143 
macrotis devius, 143 
macrotis tenuirostris, 142 
Wilcoxia diguetii, 10 
striata, 27 
Zalophus californianus californianus, 149 
Zenaida asiatica, 55 
Zenaidura macroura, 95 
Zephyranthes arenicola, 23 
Ziphius cavirostris, 138 
Zonotrichia leucophrys, 60 


344, 348, 352 


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