ey peena Ws Bras tear tose tue ia matbanaredahon — are . 4 - _ — _ EO — Ge GE Ae 9 Ome me ee Teeter ye Sy OTE PRIA PG en tag Sg HARVARD UNIVERSITY aa LIBRARY OF THE Museum of Comparative Zoology Wan a4. ee on fi ; faa ; ey ie Pum PAs Cine Na a feel hy Py Ski 4g ue _ n - \ a | VYuUI Lv tewe 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 — 10. Lie Pap 13. Ss 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: < +— oe 32° + ii —— See Bie A Angeles qi ee oO ° S . g 21+ ———— | Cc 26° ——+ + © —— 2 1s MONSERRATE 2 on. J. = ete re The BELVEDERE EXPEDITION 24°; to the = GULF OF CALIFORNIA Dash lines show route of the za San Agustin II eran re niga ns ee eae 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. L g aded 2as ‘qq va Jo ]re1ap 10,7 ‘aut] ysep ev Aq UMoYs UOIpadxa ay) Jo a1no4 ay; ‘spur]st J ocll aILAYW OYd27d NYS V7Ss/ ojuabivs Dp UuUNg- ocIT lr) PION] 7] Old ofIL VITVSON VINVS OL ¢ ( \ \ 1 \ -=<= - Se . ys a pecs V7S/ = \ Nv @sisz. eszosngisiys NYS Y7S/ + v7sl ~ valleyd sojabuy , SO] ap elueg obIT Li \sem v7s/ a2ng onby olog of IT CAG eS orbnfay ozLang : 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 GRANITE {Y Y fees Gy V7) 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. "a ~ oh ; v if ~ yt ig i 4 Y ‘ it : We Max’ Fic. 8. Reid Moran collecting plants at Puerto Refugio, Isla Angel de la Guarda. ee «i SS eee Mane 4 ‘yi ¥ TA Se SR a BA % UA 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 he ee aa 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. mY tet Meee 4 q : facie’ i oe "i Z a ¢ ih he Sut er hs rh SE ty og Tht A aia Saaieuee ae ct, 1 9 car ray a an ‘. P ; f- r ay} ha HAR | UNIVER 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 ‘puemsoyy ued “y Aq ydessoroyq *(¢6661 HHNSCS) 7961 TZ pudy ‘sony Yyieuuay “aq Aq parsayjoo ‘sisuajasup ipjayoqiu snjpjo4.,y ‘ayxeusayey pe]yx2edg purysy epsenyd x ep psuy ‘T ‘SIF A NEW INSULAR SUBSPECIES OF THE | | badd LCRS SPECKLED RATTLESNAKE =z } 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. 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The City and County of San Diego. The Pacific Press, Oakland and San Fran- cisco. March 1. [Antelope on Otay Mesa; last one killed 1883. } VAN GELDER, RICHARD G. 1959. A taxonomic revision of the spotted skunks (genus Spilogale). Bull. Amer. Mus. Nat. Hist. 117:229-392. June 15. YOuNG, STANLEY P., AND Hartiey H. T. JACKSON 1951. The Clever Coyote. The Stackpole Company, Harrisburg, Pa., and the Wildlife Management Institute, Washington, D.C. xv + 411 pp. 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, 138 165 in wv Hla "at oe rae wh. 941" phenth eh “i 4 + st We ree a> bad A sl = i ead Re ss ad Ate Ss 5 ha" : } 7 7 : i : 2) _ 7 F 7 7 A 7 a a _' : ae , | *, - ie er: i ¥ vA | aC is PUBLICATIONS OF THE SOCIETY The San Diego Society of Natural History provides publication outlets primarily for the staff members of the Museum and other members of the Society. 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A “Figure” is not part of a “Plate.” Taxonomic Papers. — Descriptions of new forms should give clear information on the type specimen, including its place of deposit and catalog number. Author's Responsibilities. — An author is responsible for the factual material presented in a paper, for the accuracy of illustrative material, and for the correctness of literature citations. Proofs. — Extensively edited or revised manuscripts, as well as galley and, when desired, page proofs, will be sent to the author for his review. Prompt action by the author will ac- celerate publication. The author must be charged, at cost, for extensive alterations after type has been set. Forms for Biological Abstracts will be sent with galley proofs. Reprints. — Fifty copies of the paper will be sent, gratis, to the author(s). Additional copies may be ordered, at cost, when galley proof is returned. 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, 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 + | 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 *ssad0id ay? JO aez 4aino ayi UO quasaid uoIssaidap JayiouR pue uoIssaidap e Aq pamoT[oy duasaad ssar0:d Ve ‘plosdaaid jo uonsod Jeseq ueyi Jadu] plosAsaidoig- “Bu0] ueY) JapiMm OT , "Jeaqued “]UaA JO “A ‘aepnsueny “14a ‘yepiozeden “den ‘ay8iems “ns farenbs “bs ‘aeynsueqes “3a1 ‘prosAsaid “Bid {ssad0ad_ “D03d ‘rol4aysod “ysod ‘auneyed ‘yed ‘ayppru “pr !(uawes0y uelayxsayy Joiaysod ayi 01 aduasayar ul) UeTTaxDayy “yay ‘saduo] “Buoy ‘Buoy “7 ‘yesayey] “3e] fayeSuoja ‘3uoja 3o ‘Ja ‘pro8Asaydoya ‘01a ‘]esiop “sop JO “p ‘A]iolzaque Surpuarxe uoissasdep “jue ‘dap ‘olajue “jue £(ABo]0aIso UO UOND—as 3xa} ay) UI passndsip Ayyensn aie suoneiAap) asesaae jo yeoidAi ‘,, ‘auou ‘Q Squasqe ‘— Squasaid ‘4 :suorpisasggr pur spoquik ¢ | suo] ~ I HEA || xE 7 “Suoyja ou L peolq Peyx40F uey) = a1 Wey 1) PINUL “7 6 | ropim a | | i | | qoys sa ‘dea, | “8uoja ing “suojo -- 11 1YINDAYS ° A | | a]duis “4 iv 4 | | < “Suoja "14) “Buoya a]duiis “Buoja — "a1 punoa uy PUuLO]SOpOYyA “py 5) | “suoja | Bb | 7 I €*Z € | qwoys 30Ys ou L “8u0ja ajduiis “Suoja — ‘den puno4 me) appuddy 'Y “Sua 5 | | eS | yous sa pee) 4105 “Suoja — ey me) skpPy ‘Y = qysiy]s 2 | | 2 Aqqnas joys ou 1 ajduis | ,."Buoja — 12] paaurod 1 X1LJLOJUOD * dA FF NSE <= 79 a0Ys “suoyja ; Ve I €27Z € Aqqnas “Buoja sa 10 ajduis “Suoja — mm paauiod 123 ifJoyuo]q 'y = | 29 yous uly : Q | ql z ——‘[enba 40 0 | yenba | aaoys “Buoya ou «L peoaq ajduns bs -~ a8) punos 1 | snsoansid “y a Got | | | Le I 0 € 27 Aqqnas woys 489A LL peoaq 4AOF ‘bs aS TER jpesntey i! | Snypautjig “py | 29 woys 1ysi]s | fb I | teed €7Z “Suoyja pains ou LL peozq ajduiis ‘bs —- 1 paauiod 1 snyNID “py ~ qsa1a0ys | pesny isaduo] adeys | adeys jequ0y adeys ssad01d adeys quasaid = adeys a8pa ssaoid | a sassan01q j sayono} jeqe1seg We] “Ue ssadoid | JO11aqUe ]ess0p sairadg > [eqwoayaig = || [equoazasog jequ014 : ]BsWeN e]pIxewiedg N “S1S9qYIP'] puke uopo.ysty3 py yo ABojoayso aaneaeduioy Ol ATaV.L 225 EvoLuTION OF Pir VIPERS BRATTSTROM 1964} gadep | — jyewus “]U9A — "901-"11) snjd c/ ysod peoiq duiny "123 ‘pal qouy ‘peal pynu “7 *do1d — “Buoja ]essop | MO] g2]duus sad | duiny “MO] "peal | + 01 IGINPLIS “Pp — “suoyja "UdA | a]dunis | ¢/| 3s0d | "a1 ~~ -— "Wa | PULOJSOPOYs “py | — “Suoja Jessop + | MO] a]duis €/T ‘pra peoaq ou MO] "aI | — a1 apoudxy py — “suoya “qUaA | MO] é2]duus | “‘Buoyja ou | duiny ‘mo] bE | = | peains sk]PY a | | | — “Suoja *]U9A alts 12] ‘MO] a]duis €/[ ‘piu | peoaq | ou MO] ‘eI | — ‘Deal XIAJLOJUOD “Py — “Buoja "JUaA le MO] 2]duis | %-sod | *Buoja ou | dunny ‘]]e3 "a1 | — paain> | iffoyuo)g 'y = “suoja [Jessop syle yal 2jduiis | €/[ ‘piu peoaq ou | dumny ‘]]P2) "3994 peoiq, + | ‘per sns0aisid ‘VY — “Buoja *UaA + Wa a]duns | ¢/T “pru peoiq ou dummy “ye3 We | — | Shep | SNJPIUING “py | } J | | t —= “Buoya jenba | + | 102 a]duiis €/1 ‘pra | “yy 28 “pq sa& yA05 “TTPI "s9ar | — ‘pal | INJNIP “py Moy]eq adeys “y01d | dwny | adeys adeys | = aasnd | ssad0ad zsid -q adeys adeys ssav0id = adeys | Il ATaVvL 233 EvoLUTION OF Pit VIPERS BRATTSTROM 1964] — | ‘Buoya “U3A _— | “per | — — + | paaurod | 7 + 2e] jp | ‘par | 06 | “nas SNAD4L -- | *Buoya "WUaA — | Mol | — — | + “pai ane “nas SMUD ITU —_ “Buoja "1UaA MO] | -- — a wunipaw ae Ie] a]ppma | ad ane us | Snjpuaypr “F = “Buoya “UaA + m | == —_— + moizeu + 2k] a|ppra 104 ainoe “as | snyjpisnd MO] | - “Suoja "uaa — | mop | — | — +. | winipaw | =f jenba | aypprw "Wad ainze ‘ns | Ipapypit _ 3 : = : it as - SN jPis95t uo qUaA MO] dey + + 1] | a]pprua a1 | 506 as IPILIS1AY | winipaut | | ‘Suoye “uaa = Mo] | = | —_ + | peoiq | = 2] | a]ppra | "yea peonpes “2s 19214 | | | MO] | | | P4205 — *Buoya "quaa — MO] | _- | — pte | peoiq | _ ie] | ayppra | “a4 ainoe *A12/*218 snpida] | | —= | *Buoja “]UaA | = MO] | + | + | + | paaurod + ueIpeul a] pp ‘Wal asnigo “nas okua _— | ‘Buoyja "uaa == 0] + | ak ate punoz | = 1] ‘380d 99 “pr “ad asniqo | -aano 59JSDL9I | | _— | ‘Buoja “1U3A = MO] + + = wunipaws | => Jenba ajpprmm "ad asnigo “ain 51481] + | "Buoja uaa = Pipe ht > | ak | at ak | punoa al 1] ‘sod 79 ‘piu pal | asmiqo 4 Aand Mjoqrqius | | | | s— “Buoyja on = | | + aL A. punoi xt Jenba a] pp ‘pal asmigo yas S1ptsta — | ‘Buoy "uaa | — | 12 ae ate aL | punos + ey] a]ppru *WDa1 asnaqo - | “ns/*AJ9 Snpojnjnrs hike) | | | —.06 + | *Buoja “uaa = "aI — +- at | Aypeaauaa | + wey] a|ppiw “Wea asmiqo “Aan 4aqn4 pawn | } _— ews "UaA == “pal — — | Ayyeaquaa "ye | ayppru "pel anode “1s sisuadnjso} I | ar Il + |) ARS : | | | pauina | | + yews uaa | — | mee) | — _ + | peoq + 1e] 7 ‘sod “pad asnigo “aan XO4D = jyeus quaa | | Gry | — — AL | wnipeus | ate jenba | 7%, ‘1sod “al aynoe 4 Aan snajuPuppp + yews quan | ae 1 — — + winipeur | Je 2] t/ ‘ysod “pal 306 “ain snssojou —= “Buoja qUaA + Wy = == + | winipad | on “2e] | & “ysod ‘We 06 *Aand Sn ptss0g = yews waa | AE 1 | — _ | athe | winipaur | alk ae] ZT, sod "W001 ane “ain SNIS1}I5DG Uf — yews quan ab 173 el| — — | | peozqs | ete ey a]pprw "Wal 06 “ain SNSSIAND “) | | . Mo]aq adeys 201g | dwiny adeys | “que quasaid quasaid | 201d 38] | uaamiag apim | = aaInd adeys ajaue adeys ssa20id qaq =| ssan0id | ‘dap uolssaidap | — ssa20id | adeys pare IY jeipaur | aeapend) | ‘202d "2e] waUIeIO¥ *¥D9]A) ysaduo7T | prouaydsiseg | AMAv) e]]IXe Ay : F - ploddsaidoisq ssad03g Uy plosdrard | [esowenbs (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 ‘passaidap equoay yo Jaques), I € 27 x£ 27 *Buoja yeao ou 1 ‘]2 L peoaq | 2]duns pyeeyw + "aI punoz uy) 149]8 D4 yews Japim “pal ou L peorq ajduis “Buoyja t4a8aulaqs I 0 Z Aqqnas yeao you "WDa1 x L Peo2q ajduns “Buoya a= “yal puno yyy snjpjnrpiuoasndind yews ‘12 "J? ajduis aaenbs =< “aI snanund I £7 £7 Aqqnas JG) PEE | P2725 “Buoja snqpupnbsossnu 0 «0 Ceca “Buoja paaan ou 12) "]2 aejnsuei pex205 “Buoja + "a1 punor 4rtya siptaratavy{ 0 0 Caza uly) "aI sad 12 "]2 jejnguel a]dunis aienbs = *Wa1 wy 4142 snauiups3 aT I 0 €~Z aaenbs ou "Wal LL peosq pays0y “J wey Ez: *Buoja punoa uryy 128a]425 “paains Japim I €~*7Z €27Z aienbs "Dal ou "a1 L peoq ajduus “Jy ueyi puno eae dafrumunu Japim I €97Z €727Z "yaI yews ou "1991 aejn3ue3 | ajduis “Buoja pynspu I €~7Z €~7Z “Buoyja "a1 ysiaenbs pex105 “Buoja — aienbs ‘yuiod 4prya PAp1J0) 29 “suoja ; I £27 £77 “Buoje EE ou ‘:DaI ‘den ajduns “Buoja =. esti? “qurod Pry? ipaiainau I £7 £7 le “Buoja sah "par | aeynBuel | — paxi0j “Buoja + “pe ‘2 “yurod pry 5njp]02 uP] 430y I €*Z €77 “duoyja “suoja ou "pal Jejndueiy | pax10j aaenbs + “Buje “yuiod 3prya nSSnIDAD apl I €727 €%7 “Suoja yews ou ‘a1 |, aeynBuein|) pax10j arzenbs — “Buoja yquiod uy) popap.apl I €*Z €27 “Buoyja yews ou “a1 aepnguel “Buoyja “duoja — “suoyje puno uly) 1upupos I €*7Z €77 “Buoja *Buoja sah "a1 ysizenbs Paxz0F aaenbs = "1aq "J9 punoa uly) snypautig I €727 €27Z “Bu0]a “Buoja 4 ou "al ysraenbs P2¥205 “Suoyja + “duoyja “yuiod yprya XOLJD I €27Z €797Z “Buoja “Buoye ou "yal ysiaenbs P2yxI0F “Buoy =. “pal "ya uiod uly} SNJDULINP “gf kK qjsaoys pasny qsasuo] adeys adeys ]e3u0ay adeys adeys — psezord adeys quasaid adeys sap ssa20id sainadg sassa20]q sayonoy | ayeipenQ) | yeiaieg qe] “ue ssad0id que Jessop [euosjarg [ewoayasog ]equ014 JPSeN e]pIxeulalg ‘Ol 793 ul se suoljerAaigqqe pue sjoquids “SNANSILIULA pue sdosqiog jo ABo]0a1S0 aarjereduioy cl YTaVL [e1paw 237 Jolaque J01J9jUe JOIaUe Jo1aque Joijaque Jola3ue Jol19jue Jolaque JOllejue 10119}Ue J0119jUe EvoLuTION oF Pit VIPERS 4O0j19jUe Jolaque 4olajue BRATTSTROM Jojiajue Jolajue Jolajue Joiajue Buey jo uaun] Jamo] jo uolsog + yews — aiesuoya — yews — aiesuoya —- aieBuoja — yews — aiesuoja mat yews 2 aiesuoya -- aiesuoyja -- aieBuoja — yews — aie8uoya — aiesuoja => * [ets — aie8uoja moyaq ssa0id UoweIO; UeT/IyIIJAT 1964} ]e23249A [eauoA ]eaU9A ]e23Ue8A ]eaUeA Jessop |® IJU9A yenba ]ea3u2a ye22ueA ]e23UeA ]e23UeA ]esz0p Jenba k ]ess0p ]e43ueA Arejuap qsaduo7] = 1 Rey — MO] duiny ITEtEs 7 mo] | padumy a ya | duiny yews = MO] AF “TBR paduiny — MO] peduiny a mop | paqqour Jo pedwiny + MO] ajduiis + a alts MO] ajduns ae MOT ajduns SII pel [etek [eee + ajduis + 1m ajduis + 0 ajduns + MO] ajduns a]duis - “pal ajduis + m1 duis dummy adeys aanjeaino ssad0id Aulae> plouaydsiseg eye 2 [Ppa %J/_‘ysod 2]Ppra ouou [PPI 2]PPrH [PPI yisue] enue 2]pprw PPI appr %_‘ysod [PPI [PPI e]PpIw 2]PpIw 2]PpIw eAIND jeipau plosAsaig proiq 29 2ey MOLeU peosq peo3q MoJIeU peolq peosq peosg peo3q wnipew “suoja lems «Pp uNp peosg uinipaur ‘P ‘ung “Ja ‘urya “1e] “Jue ‘dopgq (penunucd) 71 AIgV.L a -b *AInD “Joe1 “pel — “Buoya —_ *VDe1 —_ yeAo Aqqnas ‘pol + "W001 —_— peat n> — pe4ino Aqqnas —_— “pol + peAsins — “pel — “pol “ye ]esowenbs x Pe4and MO] duiny duiny duiny MO] ysoy ou ‘peduiny AF ou ‘padwny P244°F P247°F P20} pe 44°F P20} P2430] ouneled 149 [8p sua8aulays Snjpynrvuoaind ind snaniund snypupnbsosonu Stpistatavy{ Snauimups3 1adaqyos 4a fruiunu pynspu D4v1J02 ipaitinau Snjpj0a2uyy nssnopipspl popavavl 1upupos snyoautyiq XOLID SNJDULI]IV ‘d 238 San Disco Society oF NATURAL History { Vou. 13 CROTALUS ele - - a= =o5 -- SISTRURUS TERRESTRIAL BOTHROPS ARBORE AL BOTHROPS -- -_-" -- -- =- -- 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 Go Society oF NAtrurAt History E SAN D1 242 OTE 67-6 tsdnosd snunaysig pue snssunp ‘¢71 ‘67-¢ :sdno3B sniprasssy pue snssunp~ “uoWWOD UI siaj2eIeYD jo JUad Jad Pasn siaieaey> jo Jaquinu-uowwod ul siai2eIeY Jo Jaquinny, SMEAD ETE UE $28 Snjpuajp2> $717 7°64 $7Z snyisnd 7-61 67-17 - 7 7 —_ a tg = = 76L 0 f6L SS S'S9 Epsp ipa 7°61 67 £2 67-61 = F = = FIL F699 S19 EL snypisast4y IZ-sl 97-LI 97-91 9761 £99 $69 £8b $9 £99 691) 2781 STIL OTL ZHI OTOL AZT £99 802 ¢s$9 98S 179 £°6S $72 : | A@8lp7-Z1_ 676 ZAI OZBI LTO 671Z 1 ia 61S €86 986 LIS 069 $02 069 9°86 41 pebl 6ZATOZST 6ZOZ LZGI ZO OAT = a Vase ie ; OS SAY NE) ET EE TO Re OC $945D492 LUST best 67BI OHI TBI LZOI OZ IZ = 6ZOZ_—GZZZ : > ae z 4 < blr SCT SiCO 15981770169 B19 9 nn i CO CIC Mu 1c AEC 514314 yard LES PES OCB GC2EIS 62707 Ze8 I 6c6l) 676 Oe 62350 E pa zol fey |9So Sich 8h 88h Ih bbe Gre Rit ESI 1°79 9°86 1jaqoqim 67E OTT | AeST pe IL 6tHI ZHI ZL LTT ZOL TET = 6ZIT —6ZBI 6 Z“LI >, 5 8 ee : SyCS Ten iS9 Mn CLS Yn O1CCLO1C9 NI GCO MNCS CUS ICO MMENIELOZ, 78 = «1°79 SIPIAIA best 6@BI Gz 6ZZZ LTT TBI OTT OZHZ OZEZ ~—GZHZ __—OZ-BIL > ee . . y Gus a OS VANS EES ASS IG IR UD 069 8986 86669 SnpD]NINIS ‘ 2s bebl 67BI TAT 67ST AZO ZIT ZBI OT AI 6ZOZ_—GZOZ_—HZLI_—OGZ“EI : 2 Os eS tal 7S SASS WAS ES Way CHS ORS 7410S dogns bel OTT TTT TST ZHI OTOL ZHI TIT OTL OTBI GALI GZ“LE_— “SL 4 . Bil 8225 299. F0Z' (019) 999) ZIG Eos cise fos €65 Of Of £99 sisuadny1oy CSE} YS Sh STS 53 OY 45-3 GY 274 CY 5) SEDC 5) Gy 4) CY 45°) GY 4°) 147A CUNY 474 a gpesopua Ah a A ey ae es = eet 5 ee ee See ee ee dno 4q paseys 2 8 3 Sy scale ee me Cea BY se Oe meee so ak cae ae wes s2aDBIeY? JO Jaquiny] i BS = 5 SS = sh 5 9 ny ce x 5 = i a - 3 = § g “TT 21983 uo pase ‘sninizsig pue snppyory ut ,sdiysuonejes yo Areurums Jeoeump] bl ATAVL 243 Pir VIPERS BRATTSTROM: EVOLUTION OF 1964] 05 ‘Np Ga D & gnssi8® %, 2) , & 18 he ery ° Ss ey cy 3 a dNous 4° Ow SIGIUIA AN dnouo XOuLy yore e) o nih 2s = +» < ~ a < > Z w272/(5 & 5 ‘ay iS fod Cy (s) on ei ie N\® 7 s\ oh? =] < S ANS Se % a eles 2) SNYNYLSIS S Dp Sn, ¥ by 3? % ps go? a> Gp 3 c\ 7») Rte) Co “ty ys 7, AN ty) ‘ho 75 gy Sy % Hj % =) % 2% ¥ ro) 4% & < Oe %& 2% ay) <) SY, y ny 0 “iy 2 © ONY % NO Xy <7 4 C7 a\\\\ Sh % °o cS AG 4 > 2 4 39 ease /7) 7 J YOu, Sy Ry) yy. by Ny », % Sig % 14, oe 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 \ \ \ HALIEUS \ ERYTHRURUS SUMATRANUS | i! — ! / ' ' ScHULTZI \-~ ; ‘ \_-~” MGGREGORI y ALBOLABRIS FASCIATUS MAGCROLEPIS; ' ! 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. 13 ‘aeplyejoID) ay JO Aydeasoa800z pure uornjoaa yi jo Azeuruins IVeUTUTeABeIGG 6¢ ‘B14 { VoL. VOINSWY HLNOS SdOY¥H LOS WiIdlS3yy3L snoaovj1349 NW300 (SNW3V100IdOUL) L 1410 vd ahegs AVaIMadIA ope “YY y 2 fy Ro SP 4 ‘OIN-"09110 > 2 2 SAN Dteco Society oF NAturAL History Q1yOM a0 Sete er, 22 Pia 3 SNUNHLSIS -SNTVLONO we veee SISSHOV1 --<= ~— ~ ee NOGOULSINSY =e ame NOGOULSINSY Q1YOM M3N SdOYHLOB-SNYNSAY3INIY L 256 1964 | BRATISTROM: EVOLUTION OF Pit VIPERS 27, NC Se BA) Sea Ky mare 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. 258 SAN Disco Society of Natura History { Vousa3 cag eR © as ) \ 4 P) ne ae he ev. ’, es Ae ant } SS \ Va ——s SS oo ( ov >= WS ATROX GROUP SISTRURUS-TRISERIATUS ) Ee] VIRIDIS GROUP fy DURISSUS GROUP PALEOCENE —- EOCENE (Cs EOCENE (GS ce ne we ee no = __ oe Cae. Ca ti: VIRIDIS GROUP E::] viriois eRour [| norrious J DURISSUS BASILISCUS i) MOLOSSUS Eq OURISSUS -BASILISCUS ] HORRIDUS~ MOLOSSUS OLIGOCENE - MIOCENE MIOCENE ~ PLIOCENE ES Fig. 41. Hypothetical past distribution of Crotalus. Solid line, presumed boundary be- tween Tropical and Subtropical vs. Warm Temperate floras. 1964 } BRATTSTROM: EVOLUTION OF Pit VIPERS 259 Ey VIRIOIS GROUP J HORRIOUS {J ourissus C] BASILISCUS EC] MoLossus yeas HHH ATROX GROUP RECENT (> OLIGOCENE -MIOCENE Dee Ey ATROX ——— an RUBER q ADAMANTEUS 0 =| ATROX {||| rueer it GIGANTEUS NULLA | Eq ZA uli i = = SISTRURUS SHHHHHH! s i ; [XQ avamanteu th ) TRISERIATUS GROUP LN SISTRURUS ay TRISERIATUS GROUP MIOCENE — PLIOCENE NS RECENT Fig. 41 (Continued) 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. 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Sort ety’ T Dawhalt' We aOr * Ete el Orne es a 2oF : } te Nr egy LP eam i. 7 Sp liies Jeter 4 xhemaline ty ahvave ois Se Vow : = owe Be Odi ee wete 1s. lip kee : ipa rion 6 fares Lres Va - 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 7 ? i ol nL, 7 7 ee phi *h:¢ , \ ied eet a (beh oes s te sia re Five 7 Naa aay, bs wi a? ne oe ie a tyne e 5 Bai Pei! haus a oi aon Ny on) A 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. {VorsI3 San Disco Society oF NATURAL HIStory 272 TIJUANA WATS AN G 9 DIEGO o, <2 SOK <2 o O08, S50 ves SRO < Yes SRD | Pp EI —————————— Ot SO s SOIT HKD mmm 0000000000000 00 tactcse cee eenceaneegeege Sr rarorerereeenerteechehateietelcteieceeen Sse sesoctatatchctetatotstctceaconeneneracenene S ses eatenatenaten aacees i FERRO Ss revere, acetecatene Yy 800000 e , {) [x {> EE res pss ess 5 ones eee, ves we ‘es ee “ es x “ XS mM ZONE ZONE Il (QIisUANA ees 525 Celetetetetetels KR KKK? o, SSCS racteegtateetetctatctatenetes ed ZONE L/w — z a SS ORS R205 052525 OG SNS 555 25050525 BERKEL ID SESS sec oateontatonntseoneconatere ec eetatetstacatenee es res rene, OO S505 Vv FE] TRASH DUMP OCEAN AA Fig. 1. Above, diagram of salt marsh at La Salina; below, surface profile of section A-AA. 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 { Vor. 13 ee ‘ x af \ D000 oc, 0002 cia oe | 22 ee emis = QSEae ee e* en =: Sy. Wp 09 O, re) 0, A X) 0" (See o \ S Z XS $s NS SSS eam as ane YX’ 7; JPy Ma Fig. 2. Outline drawings of imagos. a, Bledius ferratus; b, B. ornatus; c, B. punctatissimus. Bar represents 1 mm. 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 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 en . sitet, inp hay te: A on ew rei | 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 ; y @ v . TA iv) - “! ? we ¥) it .? , a Ae é i er E eee be Mi ae : ; “f ) x i iv wi fo y/ “ ,. ( bad") : j wis X 4, - song a ; ao a “aA f . ‘ wy ay art : i y 7: y . - : a Reh iN 4 ti No 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 ¢ ' | ; re: ane a a ered) pet oe *E AT eth tre : tre Pes (an, ies Pte ita 4¢tere\ ya eee “Wie a nN a4 24 ber, eu Re ae a (apikaiadh nigh hi itil aoa cok Vaal Riek at ee ' es hs . : | y 7 a ve a he ‘) ~s = be ' me i y ‘ or ie ' cal ; ; “= oa. oe A x, hy ; ; : f : eat y} . ‘ee a 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) a 0 ODS: 00900 BOK Pay 00OKO»., O02 ‘© () We) oF BOE SRL oo ae OF, COPE ECSOGD Rb eee: ae Z OQADOD Sg) “6 6 Y, ene ©, v4 equa @Os, O setis b SS x s Sees SNES SSS SaS os wy { | J) ee: Ss: 0} Si Do! os : Koy CEG se ee 0e,.0c COOH Selah aes (rh NO se “5: tae Bj Vs 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 OQn0oc© o oy S& S ‘i ee \ 1 9990 FELON e o /, hye aa Ss. NX SY on a : 1 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 \ ne \ :) 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 : Po . ' =e aes 2) eT a sie f A ay - aa Sat ba Pha a eet aN a nee vant Mer It 7 : Rei? anh f 5 7 ng i ‘< ¢ ‘ j J Mes ms ie a us — LIBRARY 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 San Deco Society oF NATURAL HIsTory 320 G07 ¢9l 69°C GST Ctl Chl er 0°01 698 UPIpeA] 607 O' 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 A nary g 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 i : i? a ee Ph Dar Ls as) : 7 . ae ; open svt? ou aon oa ue Gee een oa ae _ — a ee ad : ere a a he bea pet 1 soit idiot 7 x > Ae - roar bhai ae am Loe : 5 3 4 ; ~e ’ AL han i i. a. y 7 , . om t wy ' - a ; ny . 2) i - d 8 “ff y ; 4¢@ = . Cr - - as ke a _ > Or “ra nae i aS ; > FOL ip ot af ya a io ue » & an 7 ‘aa «ao Nvireas thai ate a > we eo = a ahh a i” , a a 24nh al Leh LIBR 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 7 /? : ry) a oon. 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