lah etna te Pode 8 tgs a teting 32 WA ieta ins bes Spectr br thiere decane re NPR BEN a ah ey, oleitote = ¢ ts Siete ig Tet =o ee . a ¢ “" 3 yy ? ne . ue esate satel aS Re tofeitrte $< "e's Ieane eel” = 2 > > ee BEE ae D : ee Pha beeen * madera st pen ahd * mee z, Laem wees ee ~b © stengin’ re a Pe Sepa? et hee rie wee OF oe Be be wkiee a Pe hy Puente, - + PY pape ems i er Oe. ~ oe ee PN em eNom S fk Po I ae i i age PAD fe 0 PS OT ee eS Ee te ge SSRN Bete cto PacPantrern ® ig Woven FP tm ern ona ate m “peewee a viet Pa honed ae each aaa ohm ceed nee ee PA-4 ge PPO ne ee ® : ie edbend ere capes ie See ere a ae be EE Oe Fang Poe = Ne na at ae Nh PIR Tp NT aL eB ath hg chm anee tt Aer re EE eee PT EP AERA Rn RMT OF LOLA LR LLIN I oe hn Ott tbe Poe RE IT ie eat VC he WY FS eT edt ng PM Mtn? mS. a E - ey all — S in Rod = irs eb TS op EA OP PES ON 2S aA We eel e rer a eee ——. : Gere ee Xa SSR eee Te eee eer urate 4h APACE STE eH SREY t AOE Nera pig hese OS pa pts aw : Sig es E = a Pam eee ee Teer peureseten bear Ad a cea Ty ech ee net Pa 1a: } eee ree oot é = eee — Sr Saas eer re aT ern buted 4 ¢ nimi by righ abe PROCEEDINGS OF THE California Academy of Sciences FOURTH SERIES Vol. XXXVII SAN FRANCISCO PUBLISHED BY THE ACADEMY 1969-71 COMMITTEE ON PUBLICATION Dr. GEorcE E. LinpsAy, Chairman Dr. Epwarp L. KersseL, Editor Dr. Leo G. HERTLEIN No. No. No. No. No. 10. sea ia CONTENTS OF VOLUME XXXVII WILLIAMS, STANLEY C. Birth activities of some North American scorpions (19 figures). Published October 22, GO me sees 2 oe NES ee Nee ANDERSON, STEVEN C., and ALAN E. Leviton. Amphibians and reptiles collected by the Street Expedition to Afghan- istan, 1965 (8 figures; 1 table). Published October 22, 1969 AppicoTT, WARREN O. Late Pliocene mollusks from San Francisco Peninsula, California, and their paleogeographic significance (3 figures; 4 plates; 4 tables). Published Decemoer 1 Ol 1969 eee: Pee ee io Harris, M. P. The biology of storm petrels in the Galapagos Islands (19 figures; 15 tables). Published December 10, PR pO hae Ee ee eee TT wt ete “aed Hanna, G Dattas. Fossil diatoms from the Pribilof Islands, Bering Sea, Alaska (105 figures; 2 maps). Pub- Pestieds WMiareiei6. LOW Ope see et ee te td ed Lreecu, Hucu B. Copelatus glyphicus (Say) and Suphi- sellus bicolor (Say), water beetles new to California and presumably introduced (Coleoptera: Dytiscidae and No- teridae) (8 figures). Published March 6, 1970 LEvITON, ALAN E., and STEVEN C. ANDERSON. Review of the snakes of the genus Lytorhynchus (25 figures). Pub- MS HedeViacehyG. el Qn: ae ewe fs Ei, WILLIAMS, STANLEY C. Scorpion fauna of Baja California, Mexico: eleven new species of Vejovis (Scorpionida: Vejovidae) (31 figures; 11 tables). Published March 6, 1970 MINTON, SHERMAN A., STEVEN C. ANDERSON, and JEROMIE A. ANDERSON. Remarks on some geckos from southwest Asia, with descriptions of three new forms and a key to the genus Tropiocolotes (13 figures; 1 table). Published May eRPnO Og Rete: museee come el Str Fh eA Le ae Batt, Ropert W., and Davin L. JAMESON. Biosystem- atics of the canyon tree frog Hyla cadaverina Cope (= Hyla californiae Gorman) (5 figures; 5 tables). Pub- Lele iNiay Piserioomun cert Vier statin hsbucs tt | Orr, RoBErT T., JACQUELINE SCHONEWALD, and Kari W. KeENnyoN. The California sea lion: skull growth and a com- Ce = LIBRARY 57-93 95-166 167-234 237-248 249-274 275-332 333-362 363-380 No. No. No. No. Index to Volume XX XVII tad: , 14. SUSE AQ): 5 2s Zap 7seoXe parison of two populations (11 figures; 3 tables). Published June 15,1970]. eee WitiiAMs, STANLEY C. New scorpions belonging to the Eusthenura group of Vejovis from Baja California, Mexico (Scorpionida: Vejovidae) (10 figures; 7 tables). Published June:5,. V970 t-.22. le eee LouKASHKIN, ANATOLE S. On the diet and feeding behavior of the northern anchovy, Engraulis mordax (Girard) (8 fig- ures: 9 tables). Published October 13, 1970 _________.____ Licut, Witi1Am J. Polydora alloporis, new species, a com- mensal spionid (Annelida, Polychaeta) from a hydrocoral off central California (7 figures). Published October 13, MOO ig teow ewe ht ef aes cio Oe Fottett, W. I. Benthic fishes cast ashore by giant waves near Point Joe, Monterey County, California (2 figures; 1 table). Published October 13, 1970 _-. = ea Orr, W. N., JupI EHLEN, and J. B. Zaitzerr. A late Tertiary diatom flora from Oregon (30 figures). Published July 23) VOW: p05 eee ho en 6 ee EscCHMEYER, WILLIAM N. Two new Atlantic scorpionfishes (2 figures). Published July 23, 1971 2 = =a CuTLER, Bruce. Darwinneon crypticus, a new genus and species of jumping spider from the Galapagos Islands (Araneae: Salticidae) (3 figures). Published July 23, WONG 2 co ee ee ee ee Se ESCHMEYER, WILLIAM N., and GERALD R. ALLEN. Three new species of scorpionfishes (Family Scorpaenidae) from Easter Island (4 figures; 3 tables). Published July 23, 1971 PutHz, VoLKER. East asiatic and oriental species of Stenus represented in the collection of the California Academy of Sciences (Coleoptera: Staphylinidae). Published November DB MOW bac ceew weet ft 2 Sw BRATCHER, Twita, and R. D. Burcu. The Terebridae (Gastropoda) of Clarion, Socorro, Cocos, and Galapagos islands... Published November 23, 1971 .... eee SmMiTH, ALLYN G. The Carboniferous genus Glyptochiton de Koninck, 1883 (Mollusca: Polyplacophora). Published INovember-23097 1.2.8) a a a RotH, Barry, and EUGENE V. Coan. Marginellidae (Mol- lusca: Neogastropoda) from the Galapagos Islands and Cocos Island.“Published, November 23, 197] aes 381-394 395-418 419-458 459-472 473-488 489-500 501-508 509-513 515-527 529-535 537-566 567-574 575-584 585-600 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XXXVII, No. 1, pp. 1-24; 19 figs. October 22, 1969 BIRTH ACTIVITIES OF SOME NORTH AMERICAN SCORPIONS |) By Stanley C. Williams Assistant Professor of Biology Department of Ecology and Systematic Biology, San Francisco State College San Francisco, California 94132 Apstract: The birth and post-birth behaviors of 14 species of North American scorpions belonging to four families are discussed. Birth occurred at all hours of the 24-hour cycle, but was restricted to a limited season of the year. The birth season was longer for some species than others. Birth of an entire litter took up to 7% hours and consisted of several steps insuring that the young would escape from their birth membrane and ascend to the mother’s back without touching the ground. Post-birth associations of the first- and second-instar nymphs with the mother is described. INTRODUCTION Mating and other reproductive activities of terrestrial arthropods are often highly specialized and frequently reflect adaptation to a specific mode of life in occupied regions of the environment. The behavior of scorpions in regard to reproductive processes is no less specialized and distinctive. However, scorpions have long eluded investigators in regard to these processes because of their long and secretive life cycles, the difficulty of rearing the complete life cycle under laboratory conditions, and because of the uniqueness of the group. In recent years, however, considerable interest has been shown in the re- productive processes of scorpions. Courtship and mating studies have been re- ported by Maccary (1810), Fabre (1923), Serfat and Vachon (1950), Baerg (1954, 1961), Southcott (1955), Alexander (1955, 1957, 1959, 1962), Anger- mann (1955), Bucherl (1958), Shulov (1958), Shulov and Amitai (1959), Matthiesen (1960), Savory (1964), McAlister (1965), and Stahnke (1966). [1] 2 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Studies on the embryology of scorpions have been reported by Mathew (1959). Observations on parturition were reported by Shulov and others (1960). Post- birth growth and development have been discussed by Smith (1927), Schultze (1927), McAlister (1960), and Baerg (1961). Much interest has centered around scorpion meiosis and various aspects of gametogenesis in recent years: Lal Sareen (1961, 1962), Venkatanarasimhaiah and Rajasekarasetty (1964), Srivastava and Agrawal (1961), Sharma and others (1959, 1962), and Bedi (1962, 1963). The purpose of this paper is to describe behaviors and processes observed in the birth of some North American scorpions. The data presented here are the results of intermittent studies in western North America over the last 5 years. Observations were made on as wide a representation of the scorpion fauna as was reasonably possible. Reproductive data on 14 species belonging to seven genera and four families is here presented to permit the study to be as comparative as possible. The data on some species such as Vejovis spinigerus (Wood), Vejovts vorhiesi Stahnke, Uroctonus mordax Thorell, and Centruroides sculpturatus Ewing are much more extensive and based on numerous observations, while the data on other species are sometimes incomplete and may be based on single observations. Most of the research reported in this study was carried out in the field or in the laboratory facilities provided by the Department of Zoology of Arizona State University. This study was partially supported by a National Science Founda- tion Summer Fellowship. Special thanks to Mont A. Cazier of Arizona State University for suggestions and encouragement, to W. L. Minckley and Loren Honetschlager of Arizona State University for donation of some specimens yield- ing valuable information to the study. Thanks also to Charlene F. Williams for clerical assistance and to Gregory Fernald for field assistance. RESULTS Time of parturition. Parturition was observed in a number of North Ameri- can species belonging to the following taxonomic categories: Family BUTHIDAE: Centruroides gertschi Stahnke Centruroides sculpturatus Ewing Family CHACTIDAE Broteas alleni (Wood) Family DIPLOCENTRIDAE Diplocentrus whitei (Gervais) Family VEJOVIDAE Anuroctonus phaeodactylus (Wood) Hadrurus arizonensis Ewing VoL. XXXVIT] WILLIAMS: NORTH AMERICAN SCORPIONS 3 Uroctonus mordax Thorell Vejovis confusus Stahnke Vejovis mesaensis (Stahnke) Vejovis minimus Kraepelin Vejovis spinigerus (Wood) Vejovis vorhiesi Stahnke Female scorpions were observed delivering young in the field and laboratory at various times throughout the day and night. No pattern of preferred time of parturition was evident. Since these observations were made on a wide variety of species, is seems likely that most North American species deliver young during no specific portion of the 24-hour cycle. The length of time spent in parturition varied greatly among individuals of the same species and among members of different species. In general, the length of time spent in parturition seemed to depend upon the number of young delivered, size of the young, and occurrence of complications interfering with the process. The shortest recorded parturition was that of a H. arizonensis which delivered 10 young in one hour. The longest recorded case was that of a V. spinigerus which delivered 69 young in 7% hours. Scorpion birth occurred in the laboratory from early June through late November. Field observations indicated a similar but shorter birth season. This observed time interval of scorpion birth appears synchronized with the warmer seasons in temperate North America. Some species, such as C. sculpturatus, gave birth during a broader interval of time (June through September). Other species, such as V. vorhiesi and H. arizonensis, appear to deliver one litter during the year, this event occurring during a very restricted period of the season. As an example, during mid-July, 50 mature females of V. vorhiesi were collected in Portal, Arizona. Two weeks later they began delivering young (all were born in the same stage of development), the entire colony giving birth within a two- week period. No other birth activity was observed during the remainder of the season. A restricted parturition season was deduced for H. arizonensis by the observation that second instars were only observed in the field during a period of about 4 weeks during the summer season. THE BIRTH PROCESS. The birth of scorpions was strikingly similar in all the taxa observed. For this reason the birth process of scorpions can perhaps be best understood by examination of the following typical birth observation. In this case, a 1.27-gram Vejovis spinigerus female gave birth to 69 young in 7% hours. Birth began at 11:30 AM (MST) on 27 July, 1967. Prior to delivery, the mother raised her body as high as possible above the substrate by stilting on her last two pairs of walking legs. The first two pairs of walking legs had no contact with the substrate, and were held beneath the genital operculum with distal segments parallel to the under surface of the body. In this stilting posture the anterior end of the body was held distinctly higher than the 4 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47H Serr. Ficure 1. Stilting posture assumed by scorpions prior to giving birth. This posture is maintained until the young have ascended to the back and is characterized by metasoma on substrate and anterior part of body raised high. Photograph of Vejovis spinigerus, 27 July 1965. posterior of the body, resulting in a unique posture (fig. 1), which was main- tained throughout birth. In approximately one hour, the genital operculum opened and the babies began passing through the genital aperature, head first, and in a somewhat anterior direction (fig. 2). Each of the newly emerged young was tightly encased in a thin transparent birth membrane (fig. 3). They emerged continuously, one by one, and dropped into a “birth basket” formed by the distal segments of the first two pairs of walking legs which were held medially, parallel to the substrate and inferior to the genital opening (fig. 2). > Ficure 2. Scorpion parturition. A scorpion is now passing through the genital aperature and is being caught in the “birth basket.” The newly born escape from their birth membrane and climb the mother’s leg to ascend the back. During this process the female catches the young with her forelegs, the new born never touching the ground. The shed birth membranes are held in the “birth basket” and form a landing platform for other new born. Photograph of Vejovis spinigerus, 27 July 1965. Ficure 3. Newly emerged scorpions still encased in their birth membranes. Second scorpion from the left is about half free of its membrane. Wrapped in these moist membranes the scorpions pass quickly through the genital aperture of the mother with a minimum of birth complication. Photograph of Vejovis vorhiesi, 17 August 1965. VoL. XXXVIT] WILLIAMS: NORTH AMERICAN SCORPIONS 5 6 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 4. First instar orientation of Vejovis spinigerus. The entire litter has been born and all members ascended to the mother’s back without complication. Notice the distinctive pattern of orientation of the young in relation to each other and the mother. Photograph taken 2 August 1965, five days after birth. The young, never touching the substrate, became active within 5 to 15 minutes after birth. At this time the anterior end of the membrane split open and the newly born scorpion struggled free. After a pause of a minute or two, each baby then climbed up one of the mother’s walking legs to ascend the back (fig. 2), where they at first assumed a random orientation. However, as more and more new born reached this destination, they gradually assumed a definite non-random spatial orientation in regard to the mother and each other (figs. 4, 5). At this point, each baby faced anteriorly with prosoma down and metasoma curled so that the aculeus pointed toward the mother’s back. Newly ascended young crawled over the already oriented young until an available space was found. At this time they positioned themselves on the mother’s back in accord with the orientation of the other young (fig. 6). > Ficure 5. Newly ascended Vejovis spinigerus in the process of orienting themselves on the mother’s back. Once orientation is assumed, the young do not move until after the first molt. Photograph taken 27 July 1965. Ficure 6. First instar orientation of Vejovis vorhiesi. Note the distinctive vejovid orientation of the young. Photograph taken 7 August 1965. VoL. XXXVII] WILLIAMS: NORTH AMERICAN SCORPIONS 7 8 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 7. First instar orientation of Centruroides sculpturatus. The first instar nymphs are randomly positioned on the mother’s back. Note the lack of positional orientation characteristic of the species of Vejovis. Note also the circular curling of the metasoma in contrast to the “U-shaped” metasoma of the vejovids. Photograph taken 6 August 1965. This orientation was in general characteristic of the Vejovidae, in contrast to that of the Buthidae which showed no first instar orientation. In this family, the young of the first instar were roughly parallel to the mother’s back and piled up randomly in layers sometimes three to four individuals deep (fig. 7). No conclusions could be made regarding first instar orientation in H. arizonensis or A. phaeodactylus as only one birth was observed for each species, and during these the first instar young never ascended. This lack of ascent was probably due to unfavorable laboratory conditions at time of birth. Four mechanisms aided the young in mounting the mother’s back without touching the ground. First was “birth basket” formation. Second was a sticky substance on the outside of the birth membrane which caused the newly emerged young to stick together in the “birth basket” until they were able to shed their membrane and thus gain some degree of control over their movement (fig. 8). Third, the shed birth membranes remained in the “birth basket,” thus forming a sticky platform which received the newly emerged babies and minimized accidental fall to the ground. Fourth, the mother remained perfectly still during the entire delivery process and afterward until long after the last baby had ascended. VoL. XXXVII] WILLIAMS: NORTH AMERICAN SCORPIONS 9 Ficure 8. Newly born young clinging together by their sticky surfaces before emergence from their birth membranes. This mass of newly emerged scorpions was removed from the “birth basket” of the mother during birth. Photograph of Vejovis vorhiesi, 7 August 1965. The two greatest obstacles to the survival of the newly born were being unable to escape from the birth membrane and accidental fall to the ground. PosT BIRTH ASSOCIATION OF YOUNG WITH MOTHER. Within 45 minutes following parturition the first instar nymphs found their way to the mother’s back, where they secured a firm hold and remained during the first few weeks of post-birth life. The first stadium of all species lasted for one to two weeks, at the end of which time all members of a litter molt simultaneously on the mother’s back within a period of 24 hours. The first instar nymphs never left the back, nor was movement readily accomplished. Even upon accidental fall to the substrate they seldom regained their position. With the onset of the second stadium, distinct changes in the appearance of the litter occurred. Each of the young became distinctly more elongate (not accompanied by increase in body weight), thus causing the mother’s back to appear much more crowded and disorganized. Even the vejovids lost their dis- tinct first instar positional orientation during this stadium (figs. 9, 10, 11). 10 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Serr. Ficure 9. Second instar orientation of Vejovis spinigerus. Compare this with the photograph of the same litter while in the first instar stage (fig. 4). The young have aban- doned their distinctive first instar orientation and have assumed a random positioning. The cast first-instar exuviae are still on the mother’s back and visible along the edges of the litter. Photograph taken 17 August 1965, 21 days after birth. At first, the second instar nymphs showed little or no movement. Gradually more and more movement occurred, until in about two weeks they began drop- ping off the mother’s back one by one. At this time the fallen young departed from the mother and assumed an independent existence. The second instar nymphs of burrowers such as A. phaeodactylus, V. spinigerus, and H. arizonensis at first sought shelter under rocks and other surface objects. Soon they began their burrowing activities by the excavation of simple cells under these shelters (fig. 12). The second instar young of all > Ficure 10. Second instar orientation of Vejovis vorhiesi. The young have abandoned their distinctive first-instar orientation and have assumed random positioning. Compare with figure 6. Note that figures 6 and 10 are not photographs of the same litter. First instar exuviae are visible on the periphery of litter. Photograph taken 17 August 1965. Ficure 11. Second instar orientation of Centruroides sculpturatus. The young are randomly positioned on the mother’s back. First-instar exuviae are visible on the periphery of litter. Photograph taken 7 August 1965. VoL. XXXVIT] WILLIAMS: NORTH AMERICAN SCORPIONS 11 12 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. ed, La sare eat ty. pee y : Re 2 i FicureE 12. Second-instar nymph of Vejovis spinigerus excavating its first burrow. Photograph taken Fall, 1965. observed species showed aggressive predatory tendencies and fed readily on small insects (Drosophila in the laboratory). That early instar offspring are carried on their mother’s back would tend to insure that they are always located among environmental conditions favorable for survival, growth, and development. In no case was a mother seen to devour the young on her back. On some occasions, however, females did eat unfortunate young which had fallen from the back. It was also clearly established that the first instar nymphs do not feed, much less show cannibalism toward each other or the mother. There is a superstition that young scorpions derive nourishment by eating the mother, but this was clearly not the case in the species observed in this study as the mother’s integument was intact and undisturbed after the babies had departed (fig. 13). During the first month or so of life, which was spent on the mother’s back, the babies never ate or drank. The time spent in the first instar form was interpreted as an extension of the developmental process which had previously been going on inside the ovarian tube. The time spent on the mother’s back in the second instar state was interpreted not only as necessary for carrying out general phases of development, but for furnishing protection and transportation for the babies until their cuticle had sufficiently hardened to make self-locomo- tion and independent life possible. VoL. XXXVII] WILLIAMS: NORTH AMERICAN SCORPIONS 13 Ficure 13. Mother Vejovis spinigerus and her last second instar offspring. This is the same mother shown in figures 1, 4, and 9. The back, vacated by the departing second-instar nymphs, is smooth, shiny, and with no signs of cannibalism by the young. Photograph taken in late August, 1965. DESCRIPTION OF FIRST AND SECOND INSTARS. Early instars belonging to the following taxonomic categories were studied: Family BUTHIDAE Centruroides gertschi Stahnke Centruroides exilicauda (Wood) Centruroides sculpturatus Ewing Centruroides vittatus (Say) Family CHACTIDAE Broteas alleni (Wood) Family DIPLOCENTRIDAE Diplocentrus whitei (Gervais ) Family VEJOVIDAE Anuroctonus phaeodactylus (Wood) Hadrurus arizonensis Ewing Uroctonus mordax Thorell Vejovis confusus Stahnke Vejovis mesaensis (Stahnke) Vejovis minimus Kraepelin 14 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 14. First instar of Anuroctonus phaeodactylus. Older instars develop a very dark cuticle pigmentation which is absent in the first instar. The body is very plump and filled with reserve nutrient. Young of this genus although very large in size are born not as fully developed as those of Vejovis and Centruroides. Photograph taken in September, 1965. Vejovis spinigerus (Wood) Vejovis vorhiesi Stahnke The first instar of each species observed demonstrated a unique color and morphology which differentiated it from all other instars. The body of this instar was soft and whitish throughout the duration of this stadium since the cuticle, although firm, never attained a significant degree of hardening or pigmentation. First instars of even the darkest species, such as the black diplocentrids, were whitish. Some of the vejovids did, however, have a slight trace of dusky pigmentation during this stage, while some of the centrurids were light orangish- white in general coloration. First-instar weight and length measurements varied only slightly within any litter. These measurements varied slightly more when different litters of the same species were compared. The first instar nymphs of different species showed the greatest differences when average measurements were compared. The first instar telson of all species was unique in that the aculeus was blunt and not fully developed. Likewise, the distal ends of the walking legs were blunt and lacked the pretarsal claws characteristic of all older instars (ios. 14> Alls; Vo). VoL. XX XVII] WILLIAMS: NORTH AMERICAN SCORPIONS 15 Per ae ee a tae i e % * Ficure 15. First-instar nymph of Vejovis spinigerus. Photograph taken 6 August 1965 on millimeter grid paper. “wait aati Ficure 16. First-instar nymph of Centruroides sculpturatus. Photograph taken 6 August 1965, on millimeter grid paper. 16 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Immediately following the first molt, the second instar young were still soft and pale, but gradually, over a 2-week period, the cuticle hardened and took on the characteristic color and color pattern of the species. The pretarsi were now distinct and possessed the characteristic claws while the aculeus was also distinct, in shape and proportion (figs. 17, 18). The first instars of all species looked remarkably similar. They all showed incomplete development of the external morphology and lacked the ability to move, feed, defend themselves, or burrow. These instars, of different species, generally differed only vaguely in color hue, size, and weight. The nymphs were born very plump, the extra body volume was occupied with large amounts of stored nutrients needed for support of normal metabolic activities and growth until feeding could begin. During the first and second in- stars, prior to the initiation of feeding, the weight gradually and slowly decreased until the onset of feeding when weight again began to increase. After the onset of feeding, the weight fluctuated noticeably because of the periodic, predatory, engorgement type of food procurement. DISCUSSION AND CONCLUSION Most temperate North American species appeared to mate before the onset of winter; the females then overwintered. With the coming of spring, the mated females became active again and their bodies gradually became swollen in size with the developing young (fig. 19). During the summer the mesosoma of the females became greatly distended and a number of the developing embryos be- came visible through the cuticle. The young, prior to birth, appeared as small, whitish, oval bodies with distinct black eye spots within the mother’s mesosoma. Field and laboratory observations indicated that probably only one litter is born annually in each of the species of Vejovis, Anuroctonus, and Hadrurus studied. Members of these species gave birth during late summer and early fall (August, September, and October). Specimens reared in the laboratory sometimes gave birth earlier or later in the season than apparently occurs in the field. Individuals of some species, such as Centruroides sculpturatus, possibly gave birth to more than one litter during the summer and fall seasons. This species was observed in the field with young on the back from late May through late October. Females with litters on the back were also frequently observed to engage in the courtship dance with a nearby male. A specimen of the more tropical species Centruroides gracilis gave birth to two litters during the year. > Ficure 17. Second-instar nymph of Vejovis spinigerus soon after departing from mother. Photograph taken Fall, 1965. Ficure 18. Second-instar nymph of Centruroides sculpturatus at time of departure from mother. Photograph taken 2 August 1965 on millimeter grid. 17 NORTH AMERICAN SCORPIONS WILLIAMS: VoL. XXXVIT] i were! 7 a a ha eames Ve i re * (ate OAS a presen et! gat MEL GE ND ~ gree =p seal ‘i Sr eee 18 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 19. Female of Anuroctonus phaeodactylus with mesosoma swollen by internally developing young. The necessary increase in body volume is provided for by expansion of the pleural membranes and backward telescoping of the mesosomal tergites. This was interesting because the mother had no opportunity to mate before delivering the second litter. Embryonic development in temperate species probably varies in duration depending upon the environmental conditions present. This development occurs at a rate that insures that the babies will be born during a favorable season. Gestation was estimated by Baerg (1954) to be from 4’2 to 7% months in the Jamaican Centruroides insulanus and about 8 months in the Arkansas Centru- roides vittatus. These estimates were in accord with the minimal gestation data gathered during this study. Gestation probably varies among the species depend- ing on size and number of young born and depending upon the general climatic conditions endured by the population. It seems reasonable that a high-altitude species such as V. vorhiesi would have different reproductive and developmental cycles than low altitude species such as V. spinigerus. Vejovis vorhiesi is active for a much shorter time during the year because of the increased period of annual inactivity caused by the longer cold season of the higher mountains. Vejovis vorhiesi, then, must go through its entire reproductive and developmental cycle in a much shorter time than is the case with species living in habitats with a longer activity season. VoL. XXXVII] WILLIAMS: NORTH AMERICAN SCORPIONS 19 In each species, the birth process appeared to commence when the young reached a certain level of development. This level of development, however, was not necessarily the same for all species. Under natural conditions, scorpions appeared to deliver their young in pro- tected situations such as under rocks, under bark of trees, in burrows. In the last 5 years some 75,000 scorpions have been observed in western North America and none were seen delivering young on the ground surface or in places lacking good shelter. Observation of birth in burrowing species, such as the members of the Vejovidae, were rare in natural field situations. Only on rare occasions was a female to be seen giving birth in a simple cell constructed under a rock or. log. These species presumably gave birth in the seclusion of the burrows which they normally occupy. In the laboratory, members of the Vejovidae (most notably Anuroctonus and Hadrurus) exhibited conspicuous burrowing behavior for several days prior to birth. Individuals belonging to species of Centruroides were more frequently observed giving birth in the field because their preferred shelters (un- der logs and bark) were more readily accessible for observation. All newly born young were very weak and not sufficiently developed to live independently. The period of time spent on the mother’s back as first instar nymphs permitted extension of development until the young were sufficiently developed for independent life. Following the molt from first to second instar stage, the time spent on the mother’s back allowed the second instar cuticle to harden and develop pigmentation. When the second instar body was strong and hard enough for adequate locomotion, the young left the back. Such a premature birth, followed by instinctive association with the mother, is not common in invertebrates. In a way this situation is analogous to the premature birth of marsupials followed by instinctive migration to the mother’s pouch. Scorpions, however, have no analogous pouch, nor do the young require food while as- sociated with the mother. Schultze (1927) reported that the young of the Philippine scorpion Palamnaeus longimanus left the mother in the third instars and that food was procurred for the litter by the mother. This seeming exception may not really be an exception. Probably the mother and young were kept in confined quarters which prevented the young from completely leaving the mother’s association. These young then may have fed on the remains of prey left after the mother’s feeding, since food of suitable nature could have been lacking due to the caged conditions. The birth process and post-birth association of the young was surprisingly uniform among the species of the four families observed. The families, however, ' diverge in respect to post-birth orientation of the first instars. The Buthidae demonstrated a random positioning of first instar nymphs on the mother’s back, while in the Vejovidae the first instar individuals demonstrated a highly orga- nized and characteristic orientation in relation to each other and to the mother. The vejovid orientation allowed more young to be efficiently carried per unit 20 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. space on the mother’s back as well as allowing a firmer positioning of each mem- ber of the litter. This orientation perhaps minimized accidental fall to the ground and thus insured greater probability of reaching the second instar stage. Orienta- tion of first instar young of the Diplocentridae and Chactidae could not be ade- quately studied because of lack of sufficient data. An interesting difference between the observed vejovids and buthids oc- curred in relation to the behavior of mothers with young on their backs. In the field, Centruroides females were frequently seen on the ground surface while carrying young on their backs. They demonstrated little or no reduction in loco- motory activity, frequently accepted food, and even participated in the court- ship dance. In contrast, it was a rare occurrence to find a female vejovid on the ground surface at night while carrying a litter. In the laboratory, members of this family showed significant reduction of locomotor movement and feeding and no courtship was observed while litters were being carried. It was, therefore, concluded that the vejovids usually remained in their burrows or other shelters while carrying young on their backs. The parturition and post-parturition activities of the scorpions reported by Shulov and others (1960) in the old world were strikingly similar to those observed here in North America. Comparison of birth activities of individuals in different families (Vejovidae, Diplocentridae, Scorpionidae, Buthidae, and Chactidae) indicated that this aspect of scorpion biology must have been a well established and successful plan early in the evolutionary history of the order. Since the now existing families are believed to have begun their divergence in the Carboniferous (Birula, 1917), it seems reasonable that the basic pattern of gestation, parturition, and post-parturition parental association must have been well established at this time. It is interesting to note, however, that some interesting differences do occur between the Old World species reported by Shulov and the North American species observed. First is that the Old World species (Nebo hierochonticus, Orthochirus innesi, Leiurus quinquestriatus, and Compsobuthus acutecarinatus ) were reported to deposit their young directly on the ground. These young ap- parently were born in a somewhat precocious condition and were able to leave the ground and ascend the mother’s back to continue their initial development. The mothers (of at least O. innesi) aided the ascent by lowering their bodies close to the ground after parturition, and in holding the first pairs of walking legs on the ground under the genital aperture. The New World scorpions (Uroctonus mordax and species of Vejovis and Centruroides) all delivered their young in what appears to be a much earlier state of development. These were wrapped in a sticky birth membrane and were caught by the first and second pairs of walking legs before they touched the ground. These young were then held by the mother until they escaped from their membranes and began their ascent to the mother’s back, never touching the ground. VoL. XXXVII] WILLIAMS: NORTH AMERICAN SCORPIONS 21 The more precocious young of the Old World species took longer to pass through the genital aperture than the New World species. Each of the young of N. hierochonticus and O. innesi required a parturition time approximating 10 and 4 minutes respectively while parturition times in species of Vejovis and Centruroides were seldom over one minute. Species such as N. hierochonticus (Diplocentridae) and Scorpio maurus (Scorpionidae) apparently retained their young within the mother’s body until a more advanced stage of development (based on data reported by Shulov and others, 1960). In these two species, each of the young had apparently already escaped from their “birth membrane” before parturition. They were capable of considerable locomotor action at time of birth and did not molt for some 10 days after parturition. Other species had apparently not attained this level of development at parturition time. The young of O. innesi, L. quinquestriatus, and C. acutecarin- atus (all Buthidae) were born still contained in the birth membrane. The young showed no movement immediately after parturition, but escaped from their membrane within several minutes of birth. Within 8 to 10 minutes of birth, individuals of these species went through their first molt (before ascending to the mother’s back). All the species of Vejovis (Vejovidae) and Centruroides (Buthidae) observed and apparently Euscorpius italicus (Chactidae) (based on data re- ported by Angermann, 1957) delivered young in what seems to be an even earlier stage of development. This group of scorpions delivered young still encased in the “birth membrane.” These young, after escape from this membrane, then ascended the mother’s back and underwent their first molt several days to over a week later. The available observations on scorpion birth indicate three types of young are born: those in an early state of development, those in a precocious state of development, and those in an intermediate group. The level of development at birth seems to be characteristic of the species and perhaps similar within related genera and families. Observations of more species will probably show that no three definite developmental groups occur, but that a continuous spectrum of development levels occur at time of birth within the order. For this reason, the terms viviparous, ovoviviparous, and oviparous may not be appropriate here. REFERENCES CITED ALEXANDER, ANNE J. 1955. Mating in scorpions. Nature, vol. 178, pp. 867-878. 1957. The courtship and mating of the scorpion, Opisthophthalmus latimanus. Proceed- ings of the Zoological Society of London, vol. 128, no. 4, pp. 529-544. 1959. Courtship and mating in the buthid scorpions. Proceedings of the Zoological Society of London, vol. 133, pp. 145-169. 1962. Courtship and mating in a scorpion. African Wildlife, vol. 16, no. 4, pp. 313-319. bo Bo CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER. ANGERMANN, HARTMUT 1955. Indirekte spermatophorenuebertragung bie Euscorpius italicus (Hbst) (Scorpiones, Chactidae). Naturwissenschaften, vol. 42, no. 10, p. 303. BaeErc, W. J. 1954. Regarding the biology of the common Jamaican scorpion. Annals of the Entomological Society of America, vol. 47, no. 2, pp. 272-276. 1961. Scorpions: Biology and effect of their venom. Agricultural Experiment Station, Division of Agriculture, University of Arkansas, Fayetteville, Bulletin 649, pp. 1-34. Bepi, USHA 1962. Studies on the male germ cells of scorpions Palamnaeus bengalensis and Palamnaeus fulvipes, with particular reference to the morphology and cyto- chemistry of the cytoplasmic inclusions. Research Bulletin of the Panjab University of Science, vol. 13, nos. 3/4, pp. 213-225. 1963. ‘“Chromatoid body” in the spermatogenesis of scorpions. Experientia, vol. 19, no. 2, pp. 90-91. Biruta, A. A. BYALYNITSKII 1917. Fauna of Russia and Adjacent Countries; Arachnoidea-Scorpions. Israel Program for Scientific Translations. 154 pp. BUCHERL, WOLFGANG 1958. Escorpioes e escorpionismo no Brasil. V. Observacoes sobre o aparelho reprodutor masculino e o acasalamento de Tityus trivittatus e Tityus bahiensis. Memorias do Instituto Butantan, vol. 27, pp. 121-155. Fasre, J. H. 1923. The Life of the Scorpion. Translation published by Dodd, Mead and Co., New York. LAL SAREEN, MApDAN 1961 (1962). Morphological and cytochemical studies on the female germ cells of the scorpions Buthus hendersoni Pocock and Buthus macmahoni Pocock. Research Bulletin of the Panjab University of Science, vol. 12, nos. 3/4, pp. 221-236. 1962. Morphological and histochemical studies on the female germ cells of scorpions Palamnaeus fulvipes Koch and Palamnaeus bengalensis Koch. Research Bul- letin of the Panjab University of Science, vol. 13, nos. 1/2, pp. 71-83. Maccary, A. 1810. Memoire sur le scorpion qui se trouve sur le Montagne de Cette, etc. Paris: Gabbon. (Quoted from Alexander, 1956). MatTHEw, A. P. 1959. Some aspects of the embryology of scorpions. Journal of the Zoological Society of India, vol. 11, no. 1, pp. 85-88. MATTHIESEN, F. A. 1960. Sobre o acasalamento de Tityus bahiensis (Perty, 1834) (Buthidae, Scorpiones). Revista de Agricultura, vol. 35, no. 4, pp. 341-346. MCALISTER, WAYNE H. 1960. Early growth rates in offspring from two broods of Vejovis spinigerus Wood. Texas Journal of Science, vol. 12, nos. 3/4, pp. 158-162. 1965. The mating behavior of Centruroides vittatus Say (Arachnida: Scorpionida). Texas Journal of Science, vol. 17, no. 3, pp. 307-316. Savory, T. 1964. Arachnida. Academic Press, New York. 291 pp. VoL. XXXVIT] WILLIAMS: NORTH AMERICAN SCORPIONS 23 SCHULTZE, W. : 1927. Biology of the large Philippine scorpion. Philippine Journal of Science, vol. 32, no. 3, pp. 375-388. SerFAT, A., and M. VAcHOoN 1950. Quelques remarques sur la biologie d’un scorpion de |’Afghanistan: Buthotus alticola (Pocock). Bulletin du Museum National d’histoire naturelle. Paris, vol. 22, pp. 215-218. SHARMA, G. P., RAM ParsHAD, and RAj:NDER HANDA 1962. Meiosis in two species of Palamnaeus (Scorpiones-Scorpionidae). Research Bulletin of the Panjab University of Science, vol. 13, nos. 1/2, pp. 85-89. SHARMA, G. P., Ram ParsuHap, and M. G. JoneEJA 1959. Chromosome mechanism in the males of three species of scorpions (Scorpiones- Buthidae). Research Bulletin of the Panjab University of Science, vol. 10, no. 2, pp. 197-207. SHULOV, A. 1958. Observations on the mating habits of two scorpions, Leiurus quinquestriatus H. and E. and Buthotus judaicus. Proceedings of the Tenth International Congress of Entomology, Montreal, 1956 (1958), pp. 877-880. SHuLov, A., and P. Amirar 1959. On the mating habits of two species of scorpions. Bulletin of the Research Council of Israel. Section B: Zoology, vol. 8, no. 1, pp. 41-42. SHuLov, A., R. Rosin, and P. Amirtat 1960. Parturition in scorpions. Bulletin of the Research Council of Israel. Section B: Zoology, vol. 9B, no. 1, pp. 65-69. SMITH, FRANK R. 1927. Observations on scorpions. Science, vol. 65, no. 1673, p. 64. SoutucorTt, R. V. 1955. Some observations on the biology, including mating and other behavior of the Australian scorpion Urodacus abruptus Pocock. Transactions of the Royal Society of Australia, vol. 78, pp. 145-154. Srivastava, M. D. L., and UMa AGRAWAL 1961. Absence of chiasmata and formation of a complex chromosomal body in the spermatogenesis of the scorpion Palamnaeus longimanus. Caryologia, vol. 14, no. 1, pp. 63-77. STAHNKE, HERBERT L. 1966. Some aspects of scorpion behavior. Bulletin of the Southern California Academy of Sciences, vol. 65, no. 2, pp. 65-80. VENKATANARASIMHAIAH, C. B., and M. R. RAJASEKARASETTY 1964. Contributions to the cytology of Indian scorpions. Chromosomal behavior in the male meiosis of Palamnaeus gravimanus. Caryologia, vol. 17, no. 1, pp. 195— 201. vy PROCEEDINGS | OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XX XVII, No. 2, pp. 25-56; 8 figs.; 1 table. October 22, 1969 AMPHIBIANS AND REPTILES COLLECTED BY THE STREET EXPEDITION TO AFGHANISTAN, 1965 By Steven C. Anderson and Alan E. Leviton California Academy of Sciences, San Francisco 94118 INTRODUCTION In June 1965, Mr. and Mrs. William S. Street began their expedition to Afghanistan. This was the second in their series of expeditions to survey the mammalian fauna of Southwest Asia and to secure specimens for the Field Museum of Natural History. An account of their first expedition to Iran in 1962-63 has been given by Lay (1967). Mr. Jerry Hassinger (1968), who accompanied the Afghanistan expedition, published an itinerary, a gazetteer, a description of the collecting localities, and a list of the mammals collected. Although concerned primarily with the mammalian fauna, the expedition also made valuable collections of amphibians and reptiles in the areas visited; these were sent to the California Academy of Sciences for study. The stations in Afghanistan where herpetological material was obtained (the numbers do not necessarily correspond with station numbers for mammals presented by Has- singer) are shown on the accompanying map (fig. 1). COLLECTING SITES The following are the Street Expedition collecting sites for amphibians and reptiles in Afghanistan, given with location and elevation. 1. Jalalabad, 34°26’ N., 70°25’ E. 732 meters. Paghman, 34°36’ N., 68°56’ E. 2440 meters. 12 miles north of Jalalabad 2600 meters. 64 miles by road east of Faizabad (?Zebak, 36°32’ N., 71°21’ E.). 12 miles east of Eskhsham Wakhan (? = Ishkamish, 36°43’ N., 71°34’ E.). [25] ne wh 26 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. of Feet “ 72 e " 3 ° = - Ficure 1. Map of Afghanistan with collecting stations of expedition numbered (refer to list of collecting stations, p. sup. 1a). Herat, 34°20’ N., 62°10’ E. 1800 meters. Mazar-i-Sharif, 36°43’ N., 67°05’ E. 457 meters. Maimana, 35°54’ N., 64°43’ E. 884 meters. Kamdesh, 35°25’ N., 71°26’ E. Oo wD VoL. XXXVII] ANDERSON & LEVITON: AFGHAN HERPETOFAUNA 27 10. Kandahar, 31°36’ N., 65°47’ E. 1425 meters. 11. 30 miles west of Dilaram (48 kilometers west), 32°15’ N., 62°50’ E. 823 meters. 12. 16 kilometers (10 miles) south of Qala-i-Kang, 30°58’ N., 61°54’ E. The present collection of 247 specimens includes six species of amphibians and 38 species of reptiles. The collection includes two new agamid species, and an additional seven species are recorded definitely from Afghanistan for the first time. The second known specimen from Afghanistan of the rare snake Eryx elegans was also collected by the expedition. Other records further extend our knowledge of the distribution of the Afghan herpetofauna. . ACKNOWLEDGMENTS We wish to thank Dr. Robert Inger and Hymen Marx of the Field Museum of Natural History for permitting our study of this material. Dr. Ilya Darevsky of the Zoological Institute, Leningrad, kindly provided us with comparative material of Cyrtodactylus fedtschenkoi. Jerry Hassinger, Expedition Fellow of the Street Expedition of the Field Museum, provided a list of the collecting localities prior to publication of his paper. Above all, we call attention to and recognize the continuing splendid efforts and avid interest of Mr. and Mrs. William S. Street, who are enhancing our knowledge of the fauna of Southwest Asia. Specimens in the collections of the Field Museum of Natural History, Chicago, are designated FMNH; California Academy of Sciences materials are coded CAS; specimens formerly in the collections of the Division of Systematic Biology of Stanford University and now on deposit at the California Academy of Sciences are cited as CAS-SU to indicate that the accompanying number is registered in the now closed Stanford catalogs. SYSTEMATIC NOTES Class AMPHIBIA Order SALIENTIA Family BUFONIDAE Bufo andersonii Boulenger. Pa Bufo andersonii BOULENGER, 1883, Ann. Mag. Nat. Hist., ser. 5, vol. 12, p. 161 (type locality: India: Rajputana: Ajmere). MATERIAL EXAMINED (21). CAS nos. 115902-115903 and FMNH no. 161037, 30°58’ N., 61°54’ E., 16 kilometers south of Qala-i-Kang [518 meters elevation], 17-19 November. FMNH nos. 161038—161042, 161044-161046, 161048-161050 and CAS nos. 115904—115907, Paghman, 34°36’ N., 58°56’ E. FMNH no. 161140 and CAS no. 115908, Jalalabad, 34°26’ N., 70°25’ E. sz meters elevation]. FMNH no. 161266, Kandahar, 31°36’ N., 65°47’ E. [1425 meters elevation ]. 28 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. REMARKS. This species was recently recorded from Afghanistan for the first time (Clark, Clark, Anderson, and Leviton, 1969). The larger of the two females from Jalalabad measures 73 mm. from snout to vent. Neither is in breeding condition. In the many specimens of Bufo viridis examined from various areas in South- west Asia (Afghanistan, northern, southwestern, southeastern Iran and Israel), the ridge on the inner side of the metatarsal is usually distinct, often a fold, and always smooth, lacking tubercles. In the specimens here identified as B. ander- soni, as well as a specimen from Nepal (CAS-SU no. 15315), this ridge consists of a row of distinct tubercles. The fourth toe is webbed to the base of the second phalanx (second from the proximal end of the toe), i.e. one-half or more the length of the toe (in the Nepal specimen slightly less than half, but more than one-third). In all of these specimens the digits are prominently tipped with dark pigmentation, whereas in B. viridis the digits usually lack dark tips. The parotoid glands extend back beyond the insertion of the forelimbs. The size and shape of the parotoids are variable in B. viridis, in which they are often much longer than wide, but may be as wide as long. Minton (1966), says that in West Pakistan the greatest width of the parotoids is approximately equal to the length in B. viridis and uses this as one of the characters for separating the two forms. In adults of B. andersonii from eastern Afghanistan and eastern Nepal, the dor- sum is irregularly mottled with gray, lacking the distinct spots usually found in females of B. viridis. The three specimens from the Seistan basin, the 15 from Paghman, and the one from Kandahar all are small immature animals, the largest being 34 mm. from snout to vent. They are assigned to B. andersonii with some reservation. They agree with those from eastern Afghanistan in the tuberculate nature of the metatarsal ridge; the extent of webbing on the toes; the size and shape of the parotoids; the width of the interorbital space which is equal to or greater than the greatest width of the eyelid; and the shape of the snout, the profile of which slopes down and back, the lower margin posterior to the forward extent of the rostral (in B. viridis the interorbital width is usually distinctly less than the width of the eyelid, and the profile of the snout slopes down and forward, or may approach the vertical). In these young specimens the height of the tympanum is one-half or less the diameter of the eye, in contrast to other specimens of B. andersonii examined and recorded in the literature, in which it is about two-thirds the diameter of the eye. In this respect they agree with B. oblongus Nikolsky in which the tympanum is said to be small. The distance between the fore and hind limbs is not dis- tinctly greater than the greatest width of the head, however, and the tarso- metatarsal articulation does reach the tympanum when the leg is laid forward. Bufo oblongus from “Ssaman-Schahi” in eastern Iran has been synonymized Vor. XXXVII] ANDERSON & LEVITON: AFGHAN HERPETOFAUNA 29 with B. andersoni by Nieden (1923) without comment. Bufo persicus Nikolsky from Seistan, Iran, is said to have small subquadrangular parotoid glands. Worthy of note is the fact that five specimens of B. viridis from Kandahar have small short parotoids, the width about equal to the length, whereas FMNH no. 161266, from the same locality, has such characters as much longer parotoids in addition to the more extensive webbing and a tuberculate metatarsal ridge. Qala-i-Kang is the northwesternmost locality recorded for this species, unless B. oblongus is in fact a synonym of B. andersonii as Nieden states. The status of the various nominal forms related to Bufo viridis is far from clear. Probably all of these, B. andersonii, B. oblongus, B. olivaceus, B. pentoni, B. persicus, and B. surdus represent post-Pleistocene isolation of populations of once more continuously distributed B. viridis. Bufo viridis Laurenti. Bufo viridis LAURENTI, 1768, Synop. Rept., p. 27 (type locality: Austria: Vienna). MATERIAL EXAMINED (17). FMNH nos. 161065—161068 and 161071, Paghman, 34°36’ N., 68°56’ E. [2440 meters elevation]. FMNH no. 161091 and CAS no. 115909, 19 kilometers east of Ishkamish, 36°42’ N., 71°46’ E., collected by S. Atakah. FMNH no. 161110, Mazar-i-Sharif, 36°43’ N., 67°05’ E. [457 meters elevation], 1-3 September. FMNH no. 161171, Paghman, 11-12 July. CAS no. 115910, Maimana, 35°54’ N., 64°43’ E. [884 meters elevation], 7 September. CAS nos. 115911—115912, Paghman, 16 July. FMNH nos. 161265, 161269, 161271 and CAS nos. 115913-115914, Kandahar, 31°36’ N., 65°47’ E. [1425 meters elevation ]. REMARKS. The largest female (78 mm. from snout to vent) is from Maimana and contains ripe ova. Other specimens do not appear to be in breeding condition. The largest male (68 mm.) is from Kandahar. Stomach contents consist mainly of beetles and ants. Family RANIDAE Rana cyanophlyctis Schneider. Rana cyanophlyctis SCHNEIDER, 1799, Hist. Amph., vol. 1, p. 137 (type locality: eastern India). MATERIAL EXAMINED (1). FMNH no. 161082, 48 kilometers west of Dilaram, 32°15’ N., 62°50’ E. [823 meters elevation], 15 November. Remarks. This widely distributed species was recently recorded for the first time in Afghanistan (Clark, Clark, Anderson, and Leviton, 1969) from Jalalabad and Khost in the eastern part of the country. The present record appears to be on the periphery of the known range. It is known from south- eastern Iran near the coast. Its present distribution in the low arid regions south and east of the Iranian Plateau is probably discontinuous. 30 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Rana ridibunda Pallas. Rana ridibunda Patuas, 1771, Reise Rus. Reich., vol. 1, p. 458 (type locality: USSR: Gurev, north coast of Caspian Sea). MATERIAL EXAMINED (7). FMNH no. 161055, Paghman, 34°36’ N., 68°56’ E. [2440 meters elevation]. FMNH nos. 161114-161115, CAS no. 115915, Herat area, 34°20’ N., 62°10’ E. [1800 meters elevation], 29 August. FMNH nos. 161124-161125 and CAS no. 115916, 102.4 kilometers by road east of Faizabad, northern Afghanistan (Zebak, 36°32’ N., 71°21’ E.), 18 August. Remarks. None of the mature females (all from Zebak) are in breeding condition. The largest measures 89 mm. from snout to vent. Rana sternosignata Murray. Rana sternosignata Murray, 1885, Ann. Mag. Nat. Hist., ser. 5, vol. 16, p. 120 (type locality: West Pakistan: Quetta). MATERIAL EXAMINED (5). CAS nos. 115917—115918, Paghman, 34°36’ N., 68°56’ E. [2440 meters elevation]. FMNH nos. 161221, 161224, Paghman, 16 July. FMNH no. 161270, Kandahar, 31°36’ N., 65°47’ E. [1425 meters elevation]. RemMarks. FMNH no. 161224 is a recently transformed young with the tail partially unresorbed. It measures 31 mm. The large female (97 mm.) from Kandahar contains ripe ova. The largest male measures 82 mm. from snout to vent. Class REPTILIA Order CHELONIA Family TESTUDINIDAE Testudo horsfieldii Gray. Testudo horsfieldii Gray, 1884, Cat. Tort. Crocod. Amphisb., British Mus., p. 7 (type locality: Afghanistan). MATERIAL EXAMINED (1). FMNH no. 161207, Paghman, 34°36’ N., 68°56’ E. [2440 meters elevation], 18 July. REMARKS. In this specimen, a female, the carapace measures 180 mm. in length, by 152 in width. Order SQUAMATA Suborder SAURIA Family AGAMIDAE Agama agilis Olivier. Agama agilis OLIVIER, 1807, Voy. Emp. Othoman, vol. 4, p. 394, and atlas, pl. 29, fig. 2 (type locality: Iraq: neighborhood of Baghdad). MATERIAL EXAMINED (15). FMNH nos. 161191-161192 and CAS no. 115921, Paghman vicinity, 34°36’ N., 68°56’ E. [2440 meters elevation], 12—22 VoL. XXXVII] ANDERSON & LEVITON: AFGHAN HERPETOFAUNA 31 FIGURE 2. Agama agrorensis (Stoliczka) from Jalalabad (FMNH no. 161161). July. FMNH nos. 161197—161199, 161201 and CAS nos. 115922-115923, Maimana, 35°54’ N., 64°43’ E. [884 meters elevation], 7 September. FMNH no. 161133 and CAS no. 115920, 102.4 kilometers by road east of Faizabad, northern Afghanistan (Zebak, 36°32’ N., 71°21’ E.), 18 August. FMNH nos. 161117-161119 and CAS no. 115919, Herat area, 34°20’ N., 62°10’ E. [1800 meters elevation |, 29 August. Remarks. None of the females contain large eggs in the ovaries. Agama agrorensis (Stoliczka). (Figure 2.) Stellio agrorensis STOLICZKA, 1872, Proc. Asiatic Soc. Bengal, July 1872, p. 128 (type locality: India: Sussel Pass, at the entrance to the Agror Valley, 6000 feet, Hazara District, north- western Punjab). Agama agrorensis, BOULENGER, 1885 Cat. Liz. British Mus., vol. 1, p. 363. MATERIAL EXAMINED (1). FMNH no. 161161, Jalalabad, 34°26’ N., 70°25’ E. [732 meters elevation]. ReMaARKS. This species, which occurs in Punjab, Kashmir, and Chitral, is here recorded within the borders of Afghanistan for the first time. Its occurrence in the valley of the Kabul River is not unexpected in view of the other recent additions to the fauna of Afghanistan based on collections from this same area (Clark, Clark, Anderson, and Leviton, 1969). The present specimen, a female with eggs in the oviducts, measures 90 mm. from snout to vent. The tail is regenerated. CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. 37 PEETELEEDEDE ETE VUET EDT ERD TED PPT EERE EEE EEE aa COCUeUee eee Cee EEE EE OEE OE EEE eee Eee Ficure 6. Agama badakhshana Anderson and Leviton, new species. A. Holotype from Mazar-i-Sharif (FMNH no. 161108). B. Paratype from 64 miles east of Faizabad (CAS no. 115924). Vor. XXXVII] ANDERSON & LEVITON: AFGHAN HERPETOFAUNA 33 CTT Ficure 7. Agama badakhshana Anderson and Leviton, new species. Ventral view of paratype (CAS no. 115924). Agama badakhshana Anderson and Leviton, new species. (Figures 6 A-B, 7.) Hototypr. FMNH no. 161108, female, Mazar-i-Sharif, northern Afghani- stan, 36°34’ N., 67°05’ E., [457 meters elevation], collected 1-3 September 1965 by Street Expedition to Afghanistan. Paratypes. CAS no. 115924, male, 64 miles by road east of Faizabad (Zebak, 36°32’ N., 71°21’ E., 2653 meters elevation), northern Afghanistan, collected 18 August 1965 by Street Expedition to Afghanistan. CAS no. 115925, female, and FMNH no. 161175, male, Paghman, Afghanistan, 34°36’ N., 68°56’ E., [2440 meters elevation], collected 11-12 July 1965 by Street Expedition to Afghanistan. Diacnosis. Head and body depressed; tail longer than head and body; head scales smooth; caudal scales forming distinct annuli; 19-25 scales around tail at level of approximately fifth whorl; tympanum large, superficial; mid- dorsal enlarged scales smooth or faintly keeled, distinctly larger than ventrals; patch of enlarged mucronate scales on flanks, distinctly larger than ventral and dorsolateral scales, about equal to largest mid-dorsal scales; scales on dorsum of thigh very large and strongly keeled; mature males with large patch of callose abdominal scales. DESCRIPTION OF HOLOTYPE. Head, body, and base of tail strongly de- pressed. Nostril in large nasal below canthus rostralis; upper head scales 34 : CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. smooth, slightly convex, largest on snout; 11/12 supralabials, 13 infralabials; group of enlarged spinose scales behind tympanum, similar group of somewhat smaller spinose scales on sides of neck, and small cluster on dorsolateral aspects of neck; all other scales of neck very small. Median dorsal scales of back enlarged, imbricate, somewhat heterogeneous in size, distinctly larger than dorsolateral and ventral scales, six to eight across middle of back, grading into dorsolaterals; mid-dorsal scales smooth, outer rows of enlarged scales weakly keeled; small scales of flanks grading into large patch of keeled, mucronate flank scales much larger than dorsolateral and ventral scales, about equal to largest mid-dorsals; small dorsals rounded or weakly keeled, subimbricate to juxtaposed; gular and ventral scales smooth, imbricate, gulars smaller than ventrals; no gular sac; distinct gular fold; skin of neck and sides of body loose, forming distinct dorsolateral fold from neck to groin and two transverse folds across neck in preserved specimen. No callose preanal or abdominal scales in holotype, which is a young female (both present in male paratypes). Hind limb reaches to eye. Tail depressed at base, covered with strongly keeled, mucronate scales much larger than any scales on head, limbs, or body; about 22 scales around tail at level of fifth whorl; scales arranged in regular annuli, three whorls to a tail segment, this segmental arrangement most distinct about a head- length posterior to vent; posteriormost whorl of each segment contains largest scales. Color (in alcohol, initial fixation in formalin) olive gray above, enlarged median dorsal scales uniform gray, bordered on either side by longitudinal row of irregularly shaped dark spots; dark-edged white ocelli arranged in oblique rows on back, much more distinct on anterior part of back; flanks mottled with dark gray; tail with small dark spots arranged to give impression of narrow crossbars; venter grayish-white, chin and throat marbled with gray; limbs marbled with gray; fingers and toes with light gray bars. The ovaries contain eggs, the largest about 2 mm. in diameter; oviducts are enlarged. Measurements of holotype (in mm.): Snout-vent length, 69; tail length, 105; head length (tip of snout to angle of jaw), 18; hind limb, 46. ParatyPes. Unfortunately, the skulls of the three paratypes have been crushed; all features of squamation are intact, however. CAS no. 115924, a male, has about eight rows of callose preanal scales, a large abdominal patch, and a small gular patch of callose scales. Dorsolateral and flank scales are much more strongly keeled and mucronate, the dorsal scales of limbs and tail having particularly prominent keels and mucrones. The ocelli on the dorsum are less distinct than in the holotype and the dorsal surfaces of head, body, and limbs have scattered small dark flecks, the venter is gray, the throat reticulated with darker gray. The male from Paghman has somewhat less strongly keeled scales than does Vor. XXXVII] ANDERSON & LEVITON: AFGHAN HERPETOFAUNA 35 CAS no. 115924, and the callose scales are less prominent, none appearing on the gular region; it also has a few scattered enlarged scales on the dorsum in addition to the enlarged mid-dorsal scales, resembling in this respect Agama caucasica. The female from Paghman has flatter, more distinctly imbricate dorsolateral scales than does the holotype. There are eggs in the ovaries. While not aS prominent as in the male, callose preanal and abdominal scales are present. The two Paghman specimens have the more prominently enlarged flank scales. Measurements of paratypes (in mm.): Museum number Snout-ventlength Taillength Headlength Hind limb CAS 115924 82 regenerated 22 62 CAS 115925 80 136 22 60 FMNH 161175 81 ea 7A 62 Remarks. This taxon is most closely related to Agama himalayana, from which it differs in having a large patch of strongly enlarged scales on the flank, a large abdominal patch of callose scales (in this regard like A. himalayana sacra of Tibet), fewer scales around the thickest part of the tail, and larger scales on the upper surface of the thigh. It is readily distinguished from Agama agrorensis in which the enlarged mid- dorsal scales are prominently keeled. Its range is to the west of A. himalayanum. We have no information as to its habitat, although its morphology suggests it is a rock-dwelling form as are other species in this group. Its elevational range appears to be considerable, since Mazar-i-Sharif lies at 457 meters, while Paghman and the area east of Faizabad are over 2400 meters. Hassinger (1968) lists the following biotopes for the localities where this species was taken: Mazar-i-Sharif—‘‘Clay and loess, Slope and plateau, Rock, Structure, and Watercourse, in dry, montane, and steppe habitat”; Zebak—‘‘Slope and plateau, Structure, Watercourse, and Rock, in dry, montane habitat”; Paghman—‘Slope and plateau, Structure, Watercourse, and Rock, in dry, montane habitat.” We recognize this form at the species level pending more detailed information regarding its distribution and variation. Agama caucasica (Eichwald). Stellio caucasicus EICHWALD, 1831, Zool. spec. Ross. Polon., vol. 3, p. 187 (type locality: USSR: Tiflis and Baku, Transcaucasia). Agama caucasica, BOULENGER, 1885, Cat. Liz. British Mus., vol. 1, p. 367. MATERIAL EXAMINED (20). FMNH nos. 161058-161062, Paghman, 34°36’ N., 68°56’ E. [2440 meters elevation]. FMNH nos. 161174, 161176 and CAS no. 115926, Paghman, 11-12 July. FMNH nos. 161209-161212, 161214— 161215, 161219 and CAS nos. 115927-115931, Paghman, 16 July. REMARKS. The smallest juvenile, collected in mid-July, has a snout-vent 36 , CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. RUUEECEUCESUV VOUT TAA TOUSTVAT STUUR SUESEREGERUEUOTTARE RAUCH SCEUUSUOUSEODSUESSUUDUAUECUIOSEUES MEELIS Ficure 3. Agama erythrogastra (Nikolsky) from vicinity of Paghman (FMNH no. 161187). length of 54 mm., tail 81. The largest male measures 157, the largest female 117 mm. from snout to vent. Stomachs examined contain blossoms and leaves as well as arthropod remains, mostly beetles and ants. The largest ovarian eggs are about 2 mm. in diameter. Agama erythrogastra (Nikolsky). (Figure 3.) Stellio erythrogastra Nikotsky, 1896, Ann. Mus. Zool. Acad. Imp. Sci. St. Pétersbourg, vol. 1, pp. 370-371 (type locality: eastern Iran: Kalender Abad and Ferimun). Agama erythrogastra, NIkotsKy, 1915, Faune Russie, vol. 1, pp. 119-121. MATERIAL EXAMINED (6). FMNH nos. 161187, 161189-161190 and CAS no. 115932, Paghman vicinity, 34°36’ N., 68°56’ E. [2440 meters elevation], 12-22 July. FMNH no. 161195 and CAS no. 115933, Maimana, 35°54’ N., 64°43’ E. [884 meters elevation], 7 September. Remarks. This constitutes the first record of this species from Afghani- stan. Its previously known distribution included northeastern Iran and south- eastern Turkmen at elevations of 3000-5000 feet. Its occurrence at Maimana is thus not unexpected, but its discovery at Paghman is particularly noteworthy. Terentjev and Chernov (1949) state that this species lives in clayey and sandy- loamy soils, found mainly in areas with colonies of Rhombomys, utilizing their burrows for retreat in the Badkhyz region of Turkmen. They were found on Vot. XXXVII] ANDERSON & LEVITON: AFGHAN HERPETOFAUNA 37 Ficure 4. Agama himalayanum (Steindachner) from 19.2 km. east of Eskhsham Wakhan (FMNH no. 161083). man-made rock piles in Iran (Clark, Clark, and Anderson, 1966), and apparently avoid vertical slopes and rock outcrops, in contrast to other species in this group of Agama. While there are no ecological data accompanying the herpetological materials acquired by the Street Expedition, it is noteworthy that Hassinger (1968) lists Rhombomys opimus as collected at Maimana, where one of the prevalent biotopes is clay and loess. Terentjev and Chernov (1949) say that A. erythrogastra feeds on insects, mainly beetles and caterpillars. The stomachs examined here contain grass- hoppers, ants, and beetles. No plant material is present. The largest male ’ measures 147 mm. from snout to vent, tail 186. The single mature female has a snout-vent length of 111 mm., tail 133, and contains small ovarian eggs. Agama himalayana himalayana (Steindachner). (Figure 4.) Stellio himalayana STEINDACHNER, 1869, Reptilia. Jn: Reise dsterr. Fregatte Novara. . .Zool. Theil., vol. 1, p. 22 (type locality: Kashmir: Ladakh). 38 ; CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. i a TELL LEE PUL Ficure 5. Agama lehmanni (Nikolsky) from Mazar-i-Sharif (FMNH no. 161109). Agama himalayana, BOULENGER, 1885, Cat. Liz. British Mus., vol. 1, p. 362. Agama himalayana himalayana, SmirH, 1935, Fauna British India, Reptilia, vol. 2 (Sauria), 1D ZS). MATERIAL EXAMINED (7). FMNH nos. 161083, 161085-161088 and CAS nos. 115934-115935, 19.2 kilometers east of Ishkamish, 36°43’ N., 71°34’ E., collected by S. Atakah. Remarks. This appears to be the first definite published record for Afghanistan. Smith (1935) gives the range as Kashmir and the adjacent country in eastern Turkestan and western Tibet. It is found in Chitral and Ladakh. Terentjev and Chernov (1949) state that it is found in the Himalayas, Trans- Himalayas, southern Tibet, Hindu Kush, the ridge system of the Pamiro-Alai west up to the Pamir inclusive and southern part of Tien Shan; not known north of the Fergana Valley. Agama lehmanni (Nikolsky). (Figure 5.) Stellio lehmanni NrxotskKy, 1896, Ann. Mus. Zool. Acad. Imp. Sci., St. Pétersbourg, p. xiv (type locality: USSR: Fergana and Bucharia). Agama lehmanni, BEpRIAGA, 1907, Rept. Przewalsky Exped., vol. 3, p. 126, pl. 3, fig. 2. VoL. XXXVII] ANDERSON & LEVITON: AFGHAN HERPETOFAUNA 39 MATERIAL EXAMINED (1). FMNH no. 161109, Mazar-i-Sharif, 34°43’ N., 67°05’ E. [457 meters elevation], 1-3 September. REMARKS. ‘Terentjev and Chernov (1949) give the distribution of this species as the mountains of the southeastern part of central Asia north to the Fergana Valley, west to the Nura Tau and Kugitang ridges, east to the Darvaz Ridge, south to northern Afghanistan inclusive. We find no previously published locality for Afghanistan, however. This species is said to be a mountain lizard, found at elevations up to 3400 meters, but also frequenting rocks of steep slopes of canyons, and on more or less gentle slopes covered with disintegrating rock, occasionally near mountain rivers and creeks and in loessy hills near mountains (Terentjev and Chernov, 1949). Prevalent biotopes in the Mazar-i-Sharif area include “Clay and loess, Slope and plateau, Rock, Structure, and Watercourse, in dry montane, and steppe habitat” (Hassinger, 1968). The present specimen, a male, measures 100 mm. from snout to vent, tail 166. It was compared with a specimen (CAS no. 94489) from the Gissarski Range of northwest Tajikistan. Both specimens differ from the descriptions given by Nikolsky (1915) and Terentjev and Chernov (1949) in that the abdominal and gular scales are faintly but distinctly keeled under magnification, as are the scales of the snout and forehead. In other respects they are in good agreement with the published descriptions. The stomach of the Afghan specimen contains grasshopper remains. Agama nupta nupta de Filippi. Agama nupta DE Fiviprt, 1843, Gior. Istit. Lombardo Sci., Lett., Arti, Biblioth. Italiana, vol. 6, p. 407 (type locality: Iran: Persepolis). Agama nupta nupta, MInTON, 1966, Bull. American Mus. Nat. Hist., vol. 134, pp. 91-92. MATERIAL EXAMINED (10). FMNH no. 161077, 48 kilometers west of Dilaram, 32°15’ N., 62°50’ E. [823 meters elevation]. FMNH nos. 161257- 161258, 161260, 161262-161263 and CAS nos. 115936-115938, Kandahar, 31°36’ N., 65°47’ E. [1425 meters elevation]. REMARKS. The largest female has a snout-vent length of 131 mm., tail 189. The largest male measures 146 mm. from snout to vent. The smallest juvenile is 74 mm. from snout to vent. Females from Kandahar and west of Dilaram have eggs up to 3 mm. in diameter in the ovaries. Agama nuristanica Anderson and Leviton, new species. (Figure 8.) HototypeE. FMNH no. 161136, adult male, Kamdesh, eastern Afghanistan, [1342 meters elevation], collected 13-17 October, 1965 by Street Expedition to Afghanistan. PARATYPE. CAS no. 115939, female, same data as holotype. 40 | CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 8. Agama nuristanica Anderson and Leviton, new species. Paratype from Kamdesh (CAS no. 115939). DiAcnosis. Head and body depressed; tail longer than head and body; segmentation into whorls composed of four regular annuli barely distinct; about 45 scales around thickest part of tail; tympanum large, superficial; largest scales of back not arranged in regular rows and somewhat heterogeneous in size, a few about twice as large as largest ventral scales, strongly keeled, dorsolateral surfaces covered by minute granules (much smaller than found either in 4. agrorensis or A. tuberculata; flanks with scattered enlarged scales, not grouped in patches, small scales granular; limbs above covered with strongly hetero- geneous scales, numerous greatly enlarged scales among much smaller ones, the smaller almost granular; large scales neither grouped in patches nor imbricate, but strongly spinose. DESCRIPTION OF HOLOTYPE. Head, body, and base of tail strongly depressed. Nostril in large scale below canthus rostralis; upper head scales unequal, rugose to bluntly keeled, largest on snout; 13 supralabials, 11/12 infralabials; three clusters of enlarged spinose scales behind tympanum, large cluster below, and cluster in front of tympanum, cluster above tympanum and two small clusters in line with this on sides of neck; all other scales of neck minute but spinose, mid-dorsal line of slightly enlarged scales on neck; median dorsal scales of back enlarged, largest about twice size of largest ventrals, strongly keeled, imbricate, heterogeneous, in 10-12 longitudinal rows, and strongly set off from adjacent minute scales of back and flanks; enlarged mid-dorsal scales begin at level of insertion of forelimbs becoming gradually larger posteriorly; scattered enlarged, conical, mucronate scales on flanks, somewhat linearly arranged, both transversely and longitudinally, many times larger than minute, granular, mucronate scales of Vot. XXXVII] ANDERSON & LEVITON: AFGHAN HERPETOFAUNA 41 back and flanks, largest tubercles being about equal to largest mid-dorsals; upper surfaces of limbs covered with flat, subimbricate, weakly keeled hetero- geneous small scales with numerous strongly keeled, tubercular large scales; gular and ventral scales smooth, imbricate, gulars smaller than ventrals; no gular sac; distinct gular and collar folds; skin of neck and sides of body loose, forming several transverse folds across neck and dorsolateral fold from neck to groin in preserved specimen. A preanal patch of callose scales involving about 7 transverse scale rows, and an abdominal patch involving about 17 transverse rows and 17 longitudinal rows at its widest point. Hind limb reaches eye. Tail depressed at base, covered with keeled mucronate scales about twice as large as largest mid-dorsals, not as large as largest tubercles on limbs; about 45 scales around thickest part of tail; scales arranged in regular annuli, but no distinctly differentiated tail segments as in other large agamas. Color (in alcohol, initial fixation in formalin) olive brown above, head and limbs dark gray, anterior and posterior corners of eyelids cream, three short dark bars on supracilliary region, middle one descending across eye; enlarged mid-dorsal scales light gray interspersed with dark flecks; enlarged tubercles on flanks very light gray, contrasting sharply with olive brown ground color; lower surfaces mottled gray and dirty white, throat medium gray with several cream spots; posterior two-thirds of tail very dark brown, nearly black above; fingers and toes dark cross-barred. Measurements of holotype (in mm.): Snout-vent length, 131; tail length, 245+ (tip missing); head length (tip of snout to angle of jaw), 37; hind limb, 110. PARATYPE. In the single paratype, a female, all of the scales are less strongly keeled, less mucronate; it lacks callose ventral scales, but agrees with the holo- type in all other characters mentioned. Measurements of paratype (in mm.): Snout-vent length, 93; tail length, 183; head length, 26; hind limb, 74. There are small eggs in the ovaries. Remarks. This species appears to be most closely related to Agama tuberculata, and Smith’s (1935, p. 215) record of the latter from Kabul may refer to this species. We have compared our specimens with nine specimens of A. tuberculata from Nepal and find several striking differences. The Nepal specimens are darker, with numerous light spots over the dorsum, whereas the _ Afghan specimens have only the scattered conical, enlarged scales of the flanks contrasting with the olive ground. The lateral scales are smaller in the Afghan specimens, the scales on the top of the head more rugose, and the nasal is sepa- rated from the rostral by one or two scales. In the Nepal specimens the nasal is in contact with the rostral. The enlarged mid-dorsal scales of the Afghan specimens are far more sharply set off from the small dorsals than in A. tuder- culata, and the caudal segments are not distinct, while in the Nepal specimens 42 ; CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. the caudal segments are well differentiated, each with four (plus one incomplete) annuli. The upper and lower portions of the arms and legs of the present species are much more strongly heterogeneous in squamation than are those of the Nepal animals. In A. nuristanica the gular scales are granular, while in 4. tuberculata they are pointed and imbricate; the caudal scales of the former are far more truncate than are those of the latter. The following information about the type locality, Kamdesh, is quoted from Hassinger (1968): ‘Prevalent biotopes: Conifer, Evergreen oak, Watercourse, and Structure in wet, montane habitat. ‘““Kamdesh is situated on a mountainside about 500 m. above the Bash Gal River, a tributary of the Kunar. Our camp was about 8 km. east of Kamdesh near the village of Kamu. Collecting was confined to elevations between 1300 and 2150 m. “The mean temperature for 48 records, one every two hours for Oct. 13, 14, 15, and 16, was 18.0° C. with a mean max. of 31.8° C. and a mean min. of 10.5° C. There were scattered showers on the 15th and 17th. “This locality is in that part of Afghanistan which receives precipitation sufficient to sustain forest from the left flank of the Indian monsoons. The terrain is mountainous with steep valleys and sheer cliffs. The climate and vegetation is unique for Afghanistan; nowhere else, except on other eastern mountains under the influence of the monsoons, can Afghan mountainside vegetation of comparable density be found.” Family ANGUIDAE Ophisaurus apodus (Pallas). Lacerta apoda Patas, 1775, Nov. Comment. Acad. Sci. Petropol., vol. 19, p. 435, pls. 9-10 (type locality: USSR: Naryn Steppe, on north coast of Caspian Sea). Ophisaurus apodus, MERTENS AND MUtter, 1928, Abh. Senckenberg Naturf. Ges., vol. 41, p. 26. MATERIAL EXAMINED (2). FMNH nos. 161121—161122, 102.4 kilometers, by road east of Faizabad, northern Afghanistan (Zebak, 36°32’ N., 71°21’ E., 2653 meters elevation), 18 August. Remarks. Both specimens are males, the longer 236 mm. from snout to vent, tail 462. Family GEKKONIDAE Agamura persica (Auguste Duméril). Gymnodactylus persicus DuMERIL, 1856, Arch. Mus. Hist. Nat. Paris, vol. 8, p. 481 (type locality: Persia). Agamura persica, BLANFoRD, 1876, Eastern Persia, vol. 2, Zool. Geol., pp. 358-359. MATERIAL EXAMINED (2). FMNH nos. 161053-161054, Paghman, 34°36’ N., 68°56’ E. [2440 meters elevation]. Vot. XXXVII] ANDERSON & LEVITON: AFGHAN HERPETOFAUNA 43 Remarks. The male measures 52 mm. from snout to vent, the female 54. In both specimens the tails have been broken off just posterior to the basal constriction. These specimens agree in squamation with specimens examined from Esfahan Province in Iran. They are less distinctly marked, however. While the Iranian specimens have five distinct dark bars on the dorsum including one on the nape, the Afghan animals have an indistinct dark bar on the nape, one behind the shoulders, and one anterior to the sacrum. The overall coloration of these specimens, including the ground color and the darker bars on the limbs, is much lighter than in the Iranian geckos. The male has two distinct preanal pores, while one of three Iranian males has two distinct pores. Minton’s (1966) material from West Pakistan lacked pores. Minton (1966) places Gymnodactylus agamuroides Nikolsky in the genus Agamura along with A. persica (Duméril) and A. femoralis Smith. We have compared these three species and are reluctant to accept this generic allocation. Agamura persica differs from all other cyrtodactyloid geckos in Southwest Asia in the unique arrangement of the mental, which is truncate or rounded posteriorly, never pointed, the anterior infralabials which are longer than broad, the first infralabials nearly as long as the mental in some specimens, and the lack of postmental shields. The circumference of the tail is sharply reduced posterior to a basal swelling, is subcylindrical, and blunt at the end, never longer than the snout-vent length. In the 11 individuals examined from Iran and Afghanistan the rostral is completely divided, although Blanford (1876) stated that it was only partially divided in specimens he described as A. cruralis Blanford. Both “femoralis” and ‘“agamuroides” have large postmentals in contact behind the sharply pointed mental, anterior infralabials broader than long, tail tapering, not so sharply reduced near the base, and longer than the head and body. The head is longer in these two species, the head width going four and one half times in the snout-vent length, as opposed to three and three quarters to four times in A. persica. The dorsal tubercles are weakly subtrihedral rather than hemispherical to subconical in A. persica. The scales of the head are polygonal, all edges in contact with adjacent scales, while in A. persica they are subcircular, tiny granules between the scales where they are not in contact. We regard “agamuroides” as a member of the genus Cyrtodactylus as presently defined. It is closely related to C. gastropholis (Werner), and these two species have darkly pigmented peritonea and lower viscera, a feature they share with A. persica, but not with “femoralis,’ in which the peritoneum and lining of the viscera are without dark pigment. The abdominal scales of both A. persica and “femoralis” differ from those of other cyrtodactyloids in that while they have a more or less free posterior margin, they do not overlap adjacent scales. At best they can be described as subimbricate, while those of Cyrtodactylus agamuroides, C. gastropholis, and 44 ; CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. other Southwest Asian species of Cyrtodactylus are strongly imbricate. In both A. persica and “‘femoralis” the dorsal tubercles are broader than long and not arranged in regular rows, while in C. agamuroides the tubercles are longer than broad and form regular longitudinal rows. From A. persica and all Southwest Asian Cyrtodactylus, “femoralis” differs in the distinctive development of the nostril region. The nostril sits atop a “turret-like” projection, the entire lower margin of which is formed by the extremely high first supralabial. Three strongly swollen nasals form the rest of the protuberance, and the rostral is excluded from contact with the nostril. In all Southwest Asian forms of Cyrtodactylus examined, and in Agamura persica, the rostral forms the anterior border of the nostril. In most Southwest Asian species of Cyrtodactylus (C. heterocercus is an exception) the tail is covered below, at least in its distal two thirds, with enlarged plates, two per segment, which rarely are longitudinally divided. This condition also maintains in Agamura persica, although the plates are narrower than in most forms of Cyrtodactylus and are more frequently partially or completely divided. In “femoralis,” on the other hand, these plates are completely divided longitudinally, and there are three, rather than two, transverse series per caudal segment. In view of these features, and pending a much needed revision of Southwest Asian gekkonid genera, we cannot assign the nominal species Agamura femoralis Smith to either the genus Agamura or the genus Cyrtodactylus. We are of the opinion that it is as divergent from Cyrtodactylus as is A. persica, and equally different from Agamura, but do not agree with Smith (1935) that it is intermediate between the two genera. Bunopus tuberculatus Blanford. Bunopus tuberculatus BLANFoRD, 1874, Ann. Mag. Nat. Hist., ser. 4, vol. 13, p. 454 (type locality: Iran: Bahu Kalat; Pishin; Isfandak; near Bampur; Rigan, Narmashir; Tumb Island. West Pakistan: Baluchistan: Mand; Saman; Dasht). MATERIAL EXAMINED (1). FMNH no. 161248, Kandahar, 31°36’ N., 65°47’ E. [1425 meters elevation |. Remarks. This small male measures 38 mm. from snout to vent, tail 40. It has three preanal pores and six dark bars on the dorsum. Cyrtodactylus caspius (Eichwald). Gymnodactylus caspius EICHWALD, 1831, Zool. spec., vol. 3, p. 181 (type locality: USSR: Baku, on the Caspian Sea). Cyrtodactylus caspius, UNDERWOOD, 1954, Proc. Zool. Soc., vol. 124, p. 475. MATERIAL EXAMINED (19). FMNH no. 161063, Paghman, 34°36’ N., 68°56’ E. [2440 meters elevation]. FMNH nos. 161092-161096, 161099- 161100, 161102—161103, 161106—161107 and CAS nos. 115940-115944, Mazar- i-Sharif, 36°43’ N., 67°05’ E. [457 meters elevation], 1-3 September. FMNH Vot. XXXVII] ANDERSON & LEVITON: AFGHAN HERPETOFAUNA 45 no. 161130 and CAS no. 115945, 64 miles by road east of Faizabad (Zebak, 36°32’ N., 71°21’ E. [2653 meters elevation], 18 August). Remarks. The largest male has a snout-vent length of 69 mm. The largest female measures 66 mm. from snout to vent, tail 89. The smallest juvenile, collected in early September, has a snout-vent length of 33 mm., tail 43. The females have small (one mm.) eggs in the ovaries. Cyrtodactylus fedtschenkoi (Strauch). Gymnodactylus fedtschenkoi StRAucH, 1887, Mem. Acad. Imp. Sci., St. Pétersbourg, ser. 7, vol. 35, pp. 46-47 (type locality: USSR: Samarkand; Bokhara; Gissar). Cyrtodactylus fedtschenkoi, UNDERWOOD, 1954, Proc. Zool. Soc., vol. 124, p. 475. MATERIAL EXAMINED (3). FMNH no. 161076, 48 kilometers west of Dilaram, 32°15’ N., 62°50’ E. [823 meters elevation], 15 November. FMNH nos. 161255—161256, Kandahar, 31°36’ N., 65°47’ E. [1425 meters elevation]. Remarks. All three specimens are males, the largest 65 mm. from snout to vent, tail 90. The specimen from west of Dilaram was taken in a cave. Cyrtodactylus scaber (Heyden). Stenodactylus scaber HEYDEN, 1827, in Riippell, Atlas N. African Rept., p. 15, pl. 4, fig. 2 (type locality: Arabia). Cyrtodactylus scaber, UNDERWOOD, 1954, Proc. Zool. Soc., vol. 124, p. 475. MATERIAL EXAMINED (5). FMNH nos. 161226, 161249, 161251 and CAS nos. 115946-115947, Kandahar, 31°36’ N., 65°47’ E. [1425 meters elevation]. REMARKS. Two of the specimens from Kandahar (CAS no. 115947 and FMNH no. 161251) are unusually dark, the dorsum brown with indistinct darker markings, and small, irregular white marks on many of the dorsal tubercles. The gular region and sides of belly are dusted with brown. In one juvenile from Kandahar there are seven indistinct dark crossbars on the dorsum, while in another of similar size the pattern is broken up into more or less quadrangular dark spots. The largest male is 45 mm. from snout to vent, the largest female is 49, and the smallest juvenile 28 mm. Females have eggs up to 2 mm. in diameter in the ovaries. Cyrtodactylus watsoni (Murray). Gymnodactylus watsoni Murray, 1892, Zool. Beloochistan and southern Afghanistan, pp. 68-69 (type locality: West Pakistan: Quetta). Cyrtodactylus watsoni, Minton, 1966, Bull. American Mus. Nat. Hist., vol. 134, p. 79. MATERIAL EXAMINED (4). FMNH nos. 161165—161166, 161168 and CAS no. 115948, Jalalabad, 34°26’ N., 70°25’ E. Remarks. This is the first record of this species from Afghanistan. Minton found it in West Pakistan on the Iranian Plateau from extreme northern Las Bela to Quetta and northeastward to Swat and the northern Punjab. 46 ; CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Serr. This species differs from C. scaber, to which it is closely allied, chiefly in the smaller ventral scales; our specimens have 34-37 across the middle of the belly, while C. scaber has 18-24. The males have six and seven preanal pores. Females have well developed eggs in the ovaries. Eublepharis macularius (Blyth). Cyrtodactylus macularius BLYTH, 1854, Proc. Asiatic Soc. Bengal, vol. 23, p. 747 (type locality: India: Salt Range, Punjab). Eublepharis macularius, JOHN ANDERSON, 1871, Proc. Zool. Soc., 1871, p. 163. MaTERIAL EXAMINED (1). FMNH no. 161142, Jalalabad, 34°26’ N., 70°25’ E. [732 meters elevation]. REMARKS. The testes measure only 7 mm. in length in this mature male (about 127 mm. from snout to vent). Hemidactylus flaviviridis Riippell. Hemidactylus flaviviridis RUPPELL, 1835, Neue Wirb. Faun. Abyssinia, p. 18, pl. 6, fig. 2 (type locality: Eritrea: Nassaua Island). MATERIAL EXAMINED (19). FMNH no. 161056, Paghman, 34°36’ N., 68°56’ E. [2440 meters elevation]. FMNH nos. 161143, 161145-161147, 161149, 161151-161152, 161154, 161156, 161159-161160 and CAS nos. 115949— 115955, Jalalabad, 34°26’ N., 70°25’ E. [732 meters elevation]. Remarks. This appears to be the first record of this wide-ranging species in Afghanistan. Much of its distribution, from the shores of the Red Sea and around the shores of the Arabian Peninsula and Iran, is due to human agency. The movement of caravans may also account for its presence in Afghanistan. These specimens lack dorsal tubercles, while the tails have at most two longitudinal rows, a dorsolateral and a ventrolateral row on each side. Almost all individuals are a uniform gray in preservative, wavy dark crossbars discernible in a few of the smaller specimens. The largest male measures 72 mm. from snout to vent, the largest female 69. There are no small juveniles, the smallest specimen measuring 45 mm. in snout-vent length. Females have eggs up to 3 mm. in diameter in the ovaries. One stomach examined contained a fly. Teratoscincus bedriagai Nikolsky. Teratoscincus bedriagai NiKoLsKy, 1899, Ann. Mus. Zool. Acad. Imp. Sci. St. Pétersbourg, vol. 4, pp. 146-147 (type locality: eastern Iran: Zirkuch and Seistan). MATERIAL EXAMINED (6). CAS no. 161033 and FMNH nos. 161032, 161034, 30°58’ N., 61°54’ E. 16 kilometers south of Qala-i-Kang [518 meters elevation], 17-19 November. FMNH nos. 161252, 161254 and CAS no. 115957, Kandahar, 31°36’ N., 65°47’ E. [1425 meters elevation]. REMARKS. The single male, FMNH no. 171032, has a snout-vent length of 52 mm., the tail 26. The largest female, FMNH no. 161252, measures 62 mm. from snout to vent, the tail 35. There are 46—51 scales around the middle of the Vot. XXXVII] ANDERSON & LEVITON: AFGHAN HERPETOFAUNA 47 body in these specimens. All have four dark transverse bars on the body, two on the tail; the crossbars of the back are crescentic, with paravertebral extensions reaching back to contact the succeeding crossbars. There are two vertical dark bars on the upper and lower lips and sides of chin anterior to the eye, and a vertical white bar on rostral and mental. The dark pattern contrasts more strongly with the ground color in juveniles than in adults. Adult females have developing ovarian eggs. One specimen has a solpugid in the stomach, while another has well macerated arachnid remains. Teratoscincus scincus (Schlegel). Stenodactylus scincus SCHLEGEL, 1858, Handl. Dierk., vol. 2, p. 16 (type locality: USSR: Turkestan: Ili River). Teratoscincus scincus, BOULENGER, 1885, Cat. Liz. British Mus., vol. 1, pp. 12-13, pl. 2, fig. 3. MATERIAL EXAMINED (9). FMNH nos. 161028-161030 and CAS nos. 115958115959, 16 kilometers south of Qala-i-Kang, 31°05’ N., 61°52’ E. [518 meters elevation], 17-19 November. FMNH nos. 161079—161081 and CAS no. 115960, 32°15’ N., 62°50’ E., 48 kilometers west of Dilaram [823 meters elevation ]. Remarks. As in the preceding species, the nostril is surrounded by the rostral and four nasal shields; in this species the nostril is actually separated from contact with the rostral and all but one nasal, not by a rim of the lowermost nasal, but by a very thin crescentic scale which is nearly circumnasal. In this series of specimens there are 29-34 scales around the middle of the body, six to eight longitudinal rows of large cycloid scales on the occiput, from about 43 to 50 granules across the head between the posterior angles of the orbits. Juvenile specimens have four distinct transverse dark bars on the body, three or four on the tail; in adults these crossbars fade out, and six narrow longitudinal stripes run the length of the body on dorsum and flanks. In a young female, CAS no. 115959, with a snout-vent length of 63 mm., tail 39, the bars are beginning to fade, the lineate pattern beginning to appear, most distinct in the areas of the crossbars. In a female of 91 mm. snout-vent length the lineate pattern is well developed, the crossbars still faintly visible. The females, collected in mid-November, have developing ovarian eggs. One specimen has numerous termites in the stomach. Both the largest male and largest female measure 91 mm. from snout to vent. The smallest juvenile has a snout-vent length of 51 mm., tail 35. Any decision regarding subspecific status of specimens from various areas of Southwest Asia awaits further analysis. According to Terentjev and Chernov (1949, p. 128), T. s. scincus differs from T. s. keyzerlingii Strauch from Iran in having 28-32 scales around the body as opposed to 31-36 in the latter, 30-48 scales between the eyes in “scincus,”’ 42-50 in “keyzerlingii.” According to Nikolsky (1915, p. 55), T. zarudnyi Nikolsky 1907 (the type of which comes 48 ; CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. from the same general locality as “keyzerlingi”’) has 50 granules between the eyes and six longitudinal rows of cycloid scales on the occiput, while “scincus” has 35 and nine respectively. He places “keyzerlingii” in the synonymy of ““scincus.” The type of T. zarudnyi no longer exists (Dr. I. Darevsky, personal communication ). Family LACERTIDAE Acanthodactylus cantoris Giinther. Acanthodactylus cantoris GUNTHER, 1864, Rept. British India, p. 73 (type locality: India: Ramnagar, Agra; holotype in British Museum). MATERIAL EXAMINED (1). FMNH no. 161164, Jalalabad, 34°26’ N., 70°25’ E. [732 meters elevation]. Remarks. The single specimen is a badly damaged juvenile. The status of populations in Afghanistan is under investigation. Meristic data indicate that lizards from southern Afghanistan differ significantly from both A. c. cantoris and A. c. blanfordi (Clark, Clark, Anderson, and Leviton, 1969). Eremias guttulata watsonana Stoliczka. Eremias (Mesalina) watsonana StoviczKA, 1872, Proc. Asiatic Soc. Bengal, 1872, pp. 86-87 (type locality: West Pakistan: Sind: between Karachi and Sakhar). Eremias guttulata watsonana, SmMirH, 1935, Fauna British India, Rept. Amphib., vol. 2, pp. 389-390. MATERIAL EXAMINED (23). FMNH no. 161170 and CAS no. 115961, Jalalabad, 34°26’ N., 70°25’ E. [732 meters elevation]. FMNH nos. 161227, 161229, 161231, 161235—-161238, 161240-161245, 161247 and CAS nos. 115961- 115968, Kandahar, 31°36’ N., 65°47’ E. [1425 meters elevation]. Eremias nigrocellata Nikolsky. Eremias nigrocellata NiKOLSKy, 1896, Ann. Mus. Zool. Acad. Imp. Sci., St. Pétersbourg, vol. 1, p. 371 (type locality: eastern Iran: Feizabad-Mondechi and Seistan). MATERIAL EXAMINED (3). FMNH nos. 161127—161129, 102.4 kilometers by road east of Faizabad, northern Afghanistan (Zebak, 36°32’ N., 71°21’ E. [2653 meters elevation], 18 August. Remarks. This is the first record of this species in Afghanistan. Its occur- rence is not surprising, however, since it is known from southwestern Tajikistan and southern Uzbekistan near the Afghan border in the U.S.S.R. as well as from northeastern Iran. These specimens agree well with two (CAS nos. 94485-94486) from Kurgan- Tepe, Tajikistan. All have two series of subdigital lamellae beneath the fourth toe, the two series of femoral pores widely separated, the second of the large supraoculars longer than the first, the parietal at least as broad as long, temporal shield small but distinct; no enlarged preanal plate. The row of frontal granules Vor. XXXVII] ANDERSON & LEVITON: AFGHAN HERPETOFAUNA 49 is complete in two specimens, incomplete in one; in one specimen the subocular reaches the mouth on one side. All are gray above with small black spots and dark-margined white ocelli longitudinally arranged. There are remains of various small arthropods in the stomachs, including ants, locusts, and a solpugid. The two females contain small ovarian eggs. Counts and measurements are as follows: snout-vent length 50-55 mm.; tail 59-71; transverse rows of ventral plates 31-33, 16-17 in the longest series; 48-50 dorsals; 28-34 gulars; 11—13 plates in collar; 12-16 femoral pores. Eremias velox persica Blanford. Eremias persica BLANFORD, 1874, Ann. Mag. Nat. Hist., ser. 4, vol. 14, p. 31 (type locality: Tran: near Isfahan). Eremias velox var. persica, BOULENGER, 1921, Monogr. Lacertidae, vol. 2, pp. 312-314. MATERIAL EXAMINED (1). FMNH no. 161064, Paghman, 34°36’ N., 68°56’ E. [2440 meters elevation]. REMARKS. This specimen has a snout-vent length of 75 mm., tail 122; there is a large egg in each oviduct. Family ScINCcIDAE Mabuya dissimilis (Hallowell). Euprepes dissimilis HALLOWELL, 1860, Trans. American Philos. Soc., vol. 11, p. 78 (type locality: Bengal). Mabuya dissimilis, Boulenger, 1887, Cat. Liz. British Mus., vol. 3, p. 175. MATERIAL EXAMINED (1). FMNH no. 161162, Jalalabad, 34°26’ N., 70°25’ E. [732 meters elevation]. Remarks. This is the second specimen of this skink from Afghanistan. It was recently reported for the first time within Afghan limits from a locality near Jalalabad in the Kabul River valley (Clark, Clark, Anderson, and Leviton, 1969). This female measures 72 mm. from snout to vent, tail about 114 mm. Family VARANIDAE Varanus bengalensis bengalensis (Daudin). Tupinambis bengalensis Daupin, 1802, Hist. Nat. Rept., vol. 3, p. 67 (type locality: Bengal). _ Varanus bengalensis, DUMERIL AND BrBRON, 1836, Erp. Gen., vol. 3, p. 480. Varanus (Indovaranus) bengalensis bengalensis, MERTENS, 1942, Abh. Senckenberg Naturf. Ges., vol. 462, p. 13. MATERIAL EXAMINED (1). FMNH no. 161208, 19.2 kilometers north of Jalalabad [2600 meters elevation], 3 August. Remarks. This adult male measures 390 mm. from snout to vent; the tail is 663 mm. long. [Proc. 4TH Serr. 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The type, a female, had no locality data other than “Afghanistan.” The species is also known from northeastern Iran and Turkmen, U.S.S.R., in the Kopet-Dag. Terentjev and Chernov (1949, p. 229) give the range of counts as: 36-41 scale rows, 162-184 ventrals, 24-25 subcaudals. The type has 36 scale rows, 184 ventrals, and 24 subcaudals (Boulenger, 1893, p. 128). Counts for our specimen, a male, are given in table 1. In the specimen at hand the second supralabial is highest. The spurs are well developed. Nikolsky (1916, p. 327) described the Kopet-Dag specimens as Eryx jaculus czarewskii, saying that they differed from E. elegans in having 38—40 scale rows, 149-169 ventrals, and 30-34 subcaudals, and in having the second, rather than third supralabial highest (as shown in Boulenger’s illustration of the type, 1893, pl. 5, fig. 1). Comparison of these data with our specimen clearly indicates Terentjev and Chernov (1949, p. 229) were correct in placing “‘czarewskii” in the synonymy of “‘elegans.” Eryx tataricus (Lichtenstein). Boa tatarica LICHTENSTEIN, 1823, in: Eversmann, Reise von Orenburg nach Buchara, p. 146 (type locality: U.S.S.R.: Aral Sea). Eryx tataricus, TERENTJEV AND CHERNOV, 1949, Opred. presmyk. zemnov., Moscow, p. 230, fig. 105. MATERIAL EXAMINED (4). FMNH no. 161183 and CAS no. 115969, Pagh- man vicinity, 34°36’ N., 68°56’ E. [2440 meters elevation], 12-22 July. FMNH no. 161123, 102.4 kilometers east of Faizabad, northern Afghanistan (Zebak, 36°32’ N., 71°21’ E. [2653 meters elevation], 18 August. FMNH no. 161205, Maimana, 35°54’ N., 64°43’ E. [884 meters elevation], 7 September. REMARKS. Counts and measurements are given in table 1. These specimens agree in counts with the subspecies E. t. speciosus Tzarewsky (scale rows 47-59; _ventrals 187-210; caudals 34-43 in males, 23-37 in females; Terentjev and Chernov, 1949, p. 231). Terentjev and Chernov (1949, p. 230) state that E. t. speciosus usually has two scales posterior to the internasals, while &. ¢. tataricus generally has not less than three; in the present specimens, one has 2, another has 3, and two specimens have 4 scales posterior to the internasals. Three specimens have a row of confluent dark spots on the ventrals, while one lacks such spots. 52 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Family COoLUBRIDAE Coluber ravergieri Ménétries. Coluber ravergieri MENETRIES, 1832, Cat. rais. obj. zool. Caucase. St. Pétersbourg, p. 69 (type locality: U.S.S.R.: Baku). MATERIAL EXAMINED (1). FMNH no. 161177, Paghman, 34°36’ N., 68°56’ E. [2440 meters elevation], 11-12 July. Remarks. This is a strikingly patterned individual, having 81 nearly black crossbars on a white (in alcohol, initial preservation in formalin) ground; there are about 44 spots on the tail, fusing into a dorsal stripe posteriorly; there are two rows of lateral dark spots, those on tail confluent to form a stripe; the venter is dusky, flecked with darker gray. See table 1 for counts and measure- ments. Coluber rhodorhachis (Jan). Zamenis rhodorhachis Jan, 1865, in: de Filippi, Note viag. Persia, Milan, p. 356 (type locality: Persia). Coluber rhodorhachis, PARKER, 1931, Ann. Mag. Nat. Hist., ser. 10, vol. 8, p. 516. MATERIAL EXAMINED (5). FMNH no. 161075, one unnumbered specimen and CAS no. 115970, Kandahar, 31°36’ N., 65°47’ E. [1425 meters elevation], 4 November. FMNH no. 161185 and CAS no. 115971, Paghman vicinity, 34°36’ N., 68°56’ E. [2440 meters elevation], 12—22 July. REMARKS. These specimens are referred to C. rhodorhachis with the same reservations expressed previously (Leviton, 1959, p. 455-456; Clark, Clark, Anderson, and Leviton, 1969). The two smallest specimens are gray with small dark spots, arranged in alternating rows (checkerboard fashion, but the spots not in contact with one another), these spots fading out on the posterior third of tail. In a larger female (FMNH no. 161075) these spots are confluent to form narrow crossbars, one scale row wide, with a row of vertical dark bars down each flank, alternating in position with those of the back; again, this pattern fades out on the posterior third of body and the tail. CAS no. 115971 is more or less uniform tan, with no distinct pattern, while FMNH no. 161185 (in which the head is missing) is olive-gray above, the posterior margin of each dorsal scale of anterior third of body black, with a pink vertebral stripe on the anterior half of body. In one specimen a postsubocular separates the sixth supralabial from the eye on the left side, while on the right this scale is fused with the sixth labial. One specimen has a mouse in the stomach, another a small Agama agilis. Counts and measurements are given in table 1. Natrix tessellata tessellata (Laurenti). Coronella tessellata LAURENTI, 1768, Syn. Rept., p. 87 (type locality: “in Japidia, vulgo Cars”). Natrix tessellata, BONAPARTE, 1834, Iconog. Faun. Ital., vol. 2, p. 11, pl. Natrix tessellata tessellata, SocHUREK, 1956, Burgenl. Heimatbl., Eisenstadt, vol. 18, p. 89. Vor. XXXVII] ANDERSON & LEVITON: AFGHAN HERPETOFAUNA 53 MATERIAL EXAMINED (4). FMNH no. 161112, Mazar-i-Sharif, 36°43’ N., 67°05’ E. [457 meters elevation], 1-3 September. FMNH no. 161180 and CAS no. 115972, Paghman, 34°36’ N., 68°56’ E. [2440 meters elevation], 11-12 July. FMNH no. 161204, Maimana, 35°54’ N., 64°43’ E. [884 meters eleva- tion], 7 September. RemMarRKs. Counts and measurements are given in table 1. Psammophis lineolatus (Brandt). Coluber (Taphrometopon) lineolatus BranpT, 1838, Bull. Sci. Acad. Imp. Sci., St. Péters- bourg, vol. 3, p. 243 (type locality: U.S.S.R.: Transcaspia). Psammophis lineolatus, SmirH, 1943, Fauna British India, Rept. Amphib., vol. 3, p. 367. MATERIAL EXAMINED (1). FMNH no. 161184, vicinity of Paghman, 34°36’ N., 68°56’ E. [2440 meters elevation], 12-22 July. REMARKS. Counts and measurements are given in table 1. Ptyas mucosus (Linnaeus). Coluber mucosus LINNAEUS, 1758, Syst. Nat., ed. 10, vol. 1, p. 226 (type locality: India). Ptyas mucosus, GUNTHER, 1864, Rept. British India, p. 249. MATERIAL EXAMINED (6). FMNH no. 161113, Herat area, 34°20’ N., 62°10’ E. [1800 meters elevation], 29 August. FMNH no. 161135, Kamdesh, 35°25’ N., 71°26’ E. [1342 meters elevation], 13-17 October. CAS no. 115973, Kanda- har, 31°36’ N., 65°47’ E. [1425 meters elevation], 4 November. CAS no. 115974, Maimana, 35°54’ N., 64°43’ E. [884 meters elevation], 7 September. FMNH nos. 161272—161273, Kandahar. REMARKS. Counts and measurements are given in table 1. Two specimens, FMNH nos. 161113 and 161273, are unusual in having the anal undivided. Sphalerosophis diadema schirazianus (Jan). Periops parallelus var. schirasiana JAN, 1863, Elenco Sist. delgi Ofidi, p. 60 (nomen nudum). Periops parallellus var. schiraziana JAN, 1865, in: de Filippe, Note Viag. Persia, Milan, p. 356 (original description; type locality Persia [Schiraz, according to Jan, 1863, op. cit. supra]). Sphalerosophis diadema schirazianus, MERTENS, 1956, Jahresh. Ver. Vaterl. Naturk. Wirtem- berg, vol. 111, p. 96. MATERIAL EXAMINED (2). FMNH no. 161057, Paghman, 34°36’ N., 68°56’ E. [2440 meters elevation]. FMNH no. 161206, Maimana, 35°54’ N., 64°43’ E. [884 meters elevation], 7 September. REMARKS. The stomach of FMNH no. 161057 contains the remains of two mice. Counts and measurements are given in table 1. Family ELAPIDAE Naja oxiana (Eichwald). Tomyris oxiana E1cHwatp, 1831, Zool. Spec., vol. 3, p. 171 (type locality: U_SSaRe: Transcaspia). Naja oxiana, BouLENGER, 1889, Trans. Zool. Soc., ser. 2, vol. 5, p. 103, pl. 11, fig. 2. 54 / CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. MATERIAL EXAMINED (1). FMNH no. 161138, Jalalabad, 34°26’ N., 70°25’ E. [732 meters elevation]. REMARKS. Examination of the present specimen and a juvenile from Khorassan, eastern Iran, reveals the presence of a single small tooth on the maxillary posterior to the two fangs. At the time of Bogert’s (1943) study of dentition in cobras, he had no specimens of this form, and his statement that N. oxiana had two teeth on the posterior part of the maxillary was based on the literature. He subsequently examined three specimens from the Tashkent Zoo (personal communication) and discovered that Eichwald’s specimen was aberrant or inaccurately examined. Thus JV. oxiana differs in this regard from the African members of the genus, which have two solid teeth, and from Naja naja from northern India, which have none. It resembles populations from Bengal east- ward in this character, but differs from these as well as from peninsular Indian specimens in the consistently uniform dorsal coloration of adults, and in the higher ventral and subcaudal counts (186-213 and 62-75 in JN. oxiana). The Jalalabad specimen has a small shield (the cuneate) at the lip margin between the fourth and fifth infralabials on the left side. There are 27 scale rows on the neck. Other counts and measurements are given in table 1. The question as to whether there is intergradation or overlap between NV. oxiana and N. naja naja in northern West Pakistan must await the examination of adequate samples from this area. Clearly the affinities of VV. oxiana are to the east rather than to the African cobras, however. Family VIPERIDAE Echis carinatus (Schneider). Pseudoboa carinata SCHNEIER, 1801, Hist. Amph., vol. 2, p. 285 (based on a figure by Russell, 1796, Indian Serp., vol. 1, pl. 2; type locality: India: Arni, near Madras). Echis carinata, WAGLER, 1830, Syst. Amph., p. 177. MATERIAL EXAMINED (2). FMNH nos. 161072-161073, Kandahar, 31°36’ N., 65°47’ E. [1425 meters elevation], 4 November. Remarks. Arthropod remains constitute the recognizable stomach contents in these two juveniles. Counts and measurements are given in table 1. Vipera lebetina obtusa Dwigubsky. Vipera obtusa Dwicupsky, 1832, Essay nat. hist. Russian Emp., Moscow, p. 30 (type locality: U.S.S.R.: Jelisawetpol, Transcaucasia). Vipera lebetina obtusa, TERENTJEV AND CHERNOV, 1940, Kratkii opred. presm. zemnov., Leningrad, ed. 2, p. 163. MATERIAL EXAMINED (1). FMNH no. 161139, Jalalabad, 34°26’ N., 70°25’ E. [732 meters elevation ]. REMARKS. Counts and measurements in table 1. Vor. XXXVII] ANDERSON & LEVITON: AFGHAN HERPETOFAUNA 55 SUMMARY The collection of amphibians and reptiles made by the Street Expedition to Afghanistan 1965 consists of 247 specimens, including six species of amphibians and 38 species of reptiles. Definite Afghan localities are recorded for the first time for the following species: Agama agrorensis, Agama erythrogastra, Agama himalayana, Agama lehmanni, Agama nuristanica (new species herein described) , Agama badakhshana (new species herein described), Cyrtodactylus watsoni, Hemidactylus flaviviridis, Eremias nigrocellata, Eryx elegans (type was recorded as coming from Afghanistan, but no precise locality was given). An analysis of the zoogeographic significance of this and other collections from Afghanistan is contemplated for the future. LITERATURE CITED BLANFORD, WILLIAM T. 1876. Eastern Persia, an account of the journeys of the Persian Boundary Commission, 1870-1872. Vol. 2. The zoology and geology. London, viii + 516 pp., pls. 14-28. Bocert, CHARLES M. 1943. Dentitional phenomena in cobras and other elapids with notes on adaptive modifications of fangs. Bulletin of the American Museum of Natural History, vol. 81, pp. 285-360, pls. 48-51. BOULENGER, GEORGE ALBERT 1893. Catalogue of the snakes in the British Museum. London, vol. 1, xiii + 448 pp., 28 pls. Crark, RicHaArD J., ErtcA D. CLARK, AND STEVEN C. ANDERSON 1966. Report on two small collections of reptiles from Iran. Occasional Papers of the California Academy of Sciences, no. 55, 9 pp., 1 fig. CiarkK, RicHARD J., ErtcA D. CLARK, STEVEN C. ANDERSON, AND ALAN E. Leviton. 1969. Report on a collection of amphibians and reptiles from Afghanistan, Proceedings of the California Academy of Sciences, 4th ser., vol. 36, pp. 279-316, 5 figs. HASSINGER, JERRY D. 1968. Introduction to the mammal survey of the 1965 Street Expedition to Afghanistan. Fieldiana: Zoology, vol. 55, pp. 1-81. Lay, Douctas M. 1967. A study of the mammals of Iran resulting from the Street Expedition of 1962-653. Fieldiana: Zoology, vol. 54, 282 pp. Leviton, ALAN E. 1959. Report on a collection of reptiles from Afghanistan. Proceedings of the California Academy of Sciences, 4th ser., vol. 29, pp. 445-463. Minton, SHERMAN A. 1966. A contribution to the herpetology of West Pakistan. Bulletin of the American Museum of Natural History, vol. 134, pp. 29-184, pls. 9-36. NIEDEN, F. 1923. Amphibia, Anura I. Subordo Aglossa und Phaneroglossa, Sectio 1 Arcifera. Das Tierreich, 46 Lieferung, xxxii + 584 pp. Nrikorsky, A. M. 1915. Faune de la Russie. Reptiles. Vol. 1, Chelonia et Sauria. Petrograd, 532 pp., 9 pls. 1916. Faune de la Russie. Reptiles. Vol. 2, Ophidia. Petrograd, 349 pp., 8 pls. 56 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. SmirH, Matcorm A. 1935. The fauna of British India. xiii + 440 pp., 1 pl. TERENTJEV, PAUL V., AND S. A. CHERNOV 1949. Opredelitel Presmykaiushchikhsia: Zemnovodnykh. 3rd ed., Moscow, 340 pp. Reptilia and Amphibia. Vol. 2, Sauria. London, PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XX XVII, No. 3, pp. 57-93; 3 figs.; 4 plates; 4 tables December 10, 1969 LATE PLIOCENE MOLLUSKS FROM SAN FRANCISCO PENINSULA, CALIFORNIA, AND THEIR PALEOGEOGRAPHIC SIGNIFICANCE = Warren O. Addicott U.S. Geological Survey, Menlo Park, California 94025 Asstract: Late Pliocene marine mollusks occur in exposures of the Merced(?) Formation and Santa Clara Formation in the foothills of the Santa Cruz Mountains near Stanford University, Santa Clara County, California. The fauna of 39 molluscan taxa lived in a sandy, level-bottom, depositional environment located high in the inner sublittoral zone of a protected inner-coast environment. Nearly all of the dominant elements are represented by at least closely related taxa in modern bays or estuaries of the central California coast. There is, however, a small but abundant element of southern mollusks that suggests water temperatures somewhat warmer than those of modern central California bays. This assemblage marks the northern- most occurrence of several Pliocene index species previously unreported from this area that are common in central and southern California. The late Pliocene fossil localities of northern Santa Clara County lie directly across the San Andreas fault from lower Pliocene marine strata of comparable ecology mapped as the Purisima Formation. The distribution and facies of the Pliocene Purisima Formation in the northern Santa Cruz Mountains on the west side of the San Andreas fault suggest that late Pliocene deposition was restricted to a marine embayment that did not reach as far east as the early Pliocene one, which is truncated by the fault. Mer- ced(?) strata of northern Santa Clara County may have been connected to the open ocean through an area on the southwest side of the fault that is now located some 20 to 25 miles to the northwest near Mussel Rock. The amount of right-lateral slip implied by this correlation is compatible with estimates of rates of movements along the fault during earlier parts of the Tertiary. 1 Publication approved by the Director, U. S. Geological Survey. [57] CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. OL oo INTRODUCTION Fossiliferous marine sandstone of late Pliocene age crops out in the limbs of a northwest-trending syncline in the northeastern foothills of the Santa Cruz Mountains near Stanford University. Fossil mollusks occur abundantly near the base of a thin marine sandstone unit mapped as the Merced(?) Formation by Dibblee (1966a) and at scattered localities in overlying conglomerate, sandstone, and siltstone that he mapped as the Santa Clara Formation. The latter may be marine tongues in a predominantly nonmarine sequence, as suggested by Page and Tabor (1967, p. 5). The molluscan fauna of this area, although long recognized as of Pliocene age (Arnold, 1906), has never been described, nor has it been considered in accounts of the Pliocene history of the Santa Cruz Moun- tains. It is a moderately large fauna, numbering almost as many molluscan taxa as the late Pliocene fauna of the lower part of the type Merced Formation (Glen, 1959) of the northwestern San Francisco Peninsula. It is significant zoogeographically because it marks the northernmost occurrence of warm- temperate elements in the nearshore late Pliocene molluscan faunas of California. Its occurrence on the east side of the San Andreas fault, separated by many miles from outcrops of the relatively extensive upper Pliocene strata of the northern San Francisco Peninsula to the northwest, but opposite a band of presumably older Pliocene strata (Cummings and others, 1962) on the southwest side of the San Andreas fault, has an important bearing on Pliocene paleogeography and history of movement along the fault. Principal collections upon which this report is based were assembled in 1962 and 1963 by several of the United States Geological Survey geologists, including me, from building excavations and pipeline trenches in the vicinity of Arastra- dero Road near the southern boundary of the Stanford University lands. I am indebted to J. G. Vedder, M. D. Crittenden, E. E. Brabb, E. H. Pampeyan, and T. W. Dibblee for stratigraphic information and assistance in making collections from these localities. Collections made from other localities by Stanford Univer- sity students over a period of many years were kindly made available for study by Dr. A. Myra Keen of Stanford University. Identifications of material from these are incorporated into the report. Included are collections made by Ralph Arnold and others prior to 1900 that were referred to, but not listed, in Arnold (1906, 1908) and Branner and others (1909). L. G. Hertlein, who collected much of the Stanford material as a student, lent a muricid from a California Academy of Sciences collection for inclusion in this report. R. A. Loney fur- nished material collected from a locality about one-third mile north of Stanford University locality C308. Collections from the Pliocene Purisima Formation in Portola Valley west of the San Andreas fault were made by J. C. Cummings, E. H. Pampeyan, and me. The manuscript has been read by A. M. Keen, Druid Wilson, E. H. Pampeyan, V. A. Zullo, and J. C. Cummings. Their Vor. XXXVII] ADDICOTT: LATE PLIOCENE MOLLUSKS FROM CALIFORNIA 59 comments and suggestions are deeply appreciated. R. E. Petit and W. K. Emerson kindly provided assistance in classification of cancellariids considered in this report. Eugene Coan has discussed Pliocene tellinid identifications with me. Fossil photography is by Kenji Sakamoto, of the U.S. Geological Survey. EARLIER STUDIES Initial notice of the occurrence of late Cenozoic marine strata in the southern part of the San Francisco Peninsula east of the San Andreas fault was through Conrad’s (1856) description of Schizopyga |Nassarius| californianus. This small gastropod was collected from a locality “‘12 miles back from Santa Clara” associated with a coal bed (Newberry, 1856, p. 67), presumably in the Santa Clara Formation. Nearly 40 years later, Ashley (1895a, p. 322) reported fossiliferous strata from the foothills south of Stanford University which he referred to his “Merced Series.” Judging by his faunal lists of mollusks associated with basalt in the “foothills” (p. 337), however, it seems likely that he was dealing with Miocene strata. Ashley’s Pliocene faunal assemblages from near Searsville and Coal Mine Canyon are from the Purisima Formation in Portola Valley to the southwest and across the San Andreas fault from the Merced(?) and Santa Clara Formation localities of this report. The 25 species reported from the so-called Merced Series near Stanford (Ashley, 1895b) are in collections from these areas and not from the Merced(?) Formation of the present report. Arnold (1906, p. 29) included strata near Felt Lake (fig. 1) in his Pliocene and early Pleistocene Merced Formation and listed three species from this area: Pecten latiauratus, Margarites pupilla, and Littorina planaxis. The first two taxa have not since been recognized from this area and are not in the stratigraphic collections at Stanford University; the latter probably is L. petricola of this report. Subsequent to this listing, Arnold (1908) figured three additional taxa from the locality near Felt Lake (SU locality C321): Thais ostrina, T. trancosana, and Littorina petricola. These figures were re- printed in Branner and others (1909) and two additional fossiliferous localities in the vicinity of Felt Lake were mentioned, although the strata were not discriminated from the underlying so-called Purisima [Monterey] Formation. An inarticulate brachiopod, Discinisca cumingi Broderip, from a locality along Arastradero Road south of Felt Lake (fig. 1), was figured by Hertlein and Grant (1944). Recently two species of Nassarius from localities near Felt Lake were figured by Addicott (1965a, 1965b). STRATIGRAPHIC OCCURRENCE The principal fossil localities are at the base of a late Pliocene marine- nonmarine sequence of conglomerate, sandstone, and siltstone most recently mapped as the Merced(?) Formation and Santa Clara Formation by Dibblee 60 ‘ CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. 37°25' 122°10 EXPLANATION Eo Santa Clara and Merced(?) Formations Dibblee, 1966a i. f 4 Anticline Syncline Crees Fold axes Trancos * MI7I5 *c308 USGS Stanford Fossil localities »)) Fossiliferous Pliocene exposure (not collected) Ficure 1. Map of part of northern Santa Clara County, showing distribution of the Merced(?) and Santa Clara Formations (after Dibblee, 1966a) and late Pliocene fossil localities [see fig. 2 for location]. (1966a). Nearly all of the faunal constituents occur in a basal 2- to 10-foot thick bed that crops out in the limbs of a northwest-trending syncline along Arastradero Road between Page Mill Road and Los Trancos Creek (fig. 1). This bed consists principally of broken pelecypod shells. Shell fragments and a few whole specimens indicative of Pliocene age have been collected or observed in fine-grained sandstone near the axis of this syncline (fig. 1) in strati- graphically higher beds mapped as the Santa Clara Formation by Dibblee (1966a). These occurrences seem best explained as marine tongues (Page and Tabor, 1967) within the dominantly fluviatile nonmarine Santa Clara Formation. The best fossil localities (USGS localities M1715, M1720, M1870) are from man- made cuts and excavations in otherwise poorly exposed fine-grained sandstone along Arastradero Road. Material can still be secured from a cut near the most productive locality (M1715) although the pipeline trench from which nearly all of the figured specimens were collected has long since been filled. The Vot.XXXVII] ADDICOTT: LATE PLIOCENE MOLLUSKS FROM CALIFORNIA 61 assemblage of 33 molluscan taxa from this locality is far richer than that of any of the type Merced localities of the northwestern San Francisco Peninsula, the largest of which has yielded only 14 species (Glen, 1959, p. 157). Both Earl Pampeyan and I tried unsuccessfully to relocate old Stanford University localities near Felt Lake; only a few scattered mollusk fragments were found as float in the ditch bank near Felt Lake. The fossiliferous stratum is about 10 feet thick near USGS locality M1715 (fig. 1). It rests unconformably upon diatomaceous siltstone mapped as the Monterey Shale by Dibblee (1966a). Foraminifers indicative of a late Miocene Delmontian age were collected from near the top of this unit in the vicinity of locality M1715 according to Cummings and others (1962). However, a diatom assemblage from tuffaceous diatomite about 200 feet stratigraphically below locality M1715 studied by K. E. Lohman suggests correlation with the upper part of the Luisian Stage (late middle Miocene) or the lower part of the Mohnian Stage (early late Miocene) (E. H. Pampeyan, written communication, June 1968). About 500 feet stratigraphically below the unconformity, a poorly pre- served molluscan assemblage of middle or possibly late Miocene age was collected by the writer from a cut northeast of the intersection of Junipero Serra Freeway and Page Mill Road (USGS locality M2547). At locality M1715 the contact between the Merced(?) Formation and the Monterey Shale is marked by a well-developed zone of pelecypod borings. Some of the holes contain articulated valves of the pholad Zirfaea (pl. 2, fig. 9) in life position. Basal fossiliferous strata of the Merced(?) Formation appear to grade upward through lithologically similar unfossiliferous sand and sandy gravel into strata composed of debris from the Franciscan Formation that have long been mapped as the Santa Clara Formation (Branner and others, 1909; Davis and Jennings, 1954; Dibblee, 1966a). The Merced (?) and Santa Clara Formations are folded into a gentle synclinal structure that trends northwest- ward through Felt Lake (Dibblee, 1966a). Occurrences of marine Pliocene mol- lusks stratigraphically well above the base of the Santa Clara Formation as mapped by Dibblee include, 1) SU locality C308 (Arnold, 1908, loc. 8) and USGS locality M3824 near the head of Purisima Creek several hundred feet above the contact with the Monterey Shale, and 2) localities along Arastradero Road about 1000 feet west of the intersection with Arastradero Creek (fig. 1). Nassarius californianus and fragments of several kinds of pelecypods occur at all of these localities. Taking into account the isolated occurrences of late Pliocene mollusks near the axis of the syncline, and therefore well above the base of the Santa Clara Formation, it seems apparent that the post-Miocene marine-nonmarine section includes rocks of approximately the same age and that if an unconformity occurs between the Merced(?) Formation and _ the overlying Santa Clara Formation it cannot be assigned appreciable time signifi- cance. 62 ’ CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Taste 1. Late Pliocene mollusks from northern Santa Clara County. USGS Stanford U. localities localities we oO fo) ‘oO s+ = = oO | ees = GASTROPODS Ss SoS (See (Pagrablenam, gsanoasa, (Coxe) > k Littorina petricola Dall (pl. 1, figs. 2-5) —.______ x Bitttum casmaliense Bartsch (pl. 1, figs. 9, 14) x Crepidula cf. C. princeps Conrad (pl. 2, fig. 10) _____ x x Cryptonatica aleutica Dall (pl. 1, fig. 12; pl. 4, fig. 2) _cf. K xX Neverita (Glossaulax) cf. N. (G.) recluziana (Deshayes) (Gol, i, ates, ill, 19) eee Dees ere ee x Ocenebra interfossa Carpenter (pl. 1, figs. 20, 21)* Thais (Nucella) cf. T. (N.) lima (Gmelin) (Fo Uc Ha Fee = (0) LL) eee a= Wir SOR een Pe ods PTS ys xX Thais (Nucella) trancosana Arnold Gol tea Eesti (eects aye ov ed (0) ty seca Seine Pa eee Ee x xe x Thais (Nucella) emarginata forma ostrina Gould __ xX Neptunea cf. N. tabulata (Baird) (pl. 1, fig. 17) — xX Mitrella cf. M. gouldi (Carpenter) (pial Satie FOF e ples. Gihicac4))) ee a eee Xo spr sp. X Nassarius| (Demondia) californianus (Conrad) ; (Calg 3 aT Ge 7p) ates Be a ee A ae X Kk GX XS SxXGexeeex Nassarius (Caesia) grammatus (Dall) (perio s all Sieg eeAem i pe 58) eee ee ee ee >, ee ae, < cit «cies Olivella biplicata (Sowerby) (pl. 3, fig. 9) ~~ XL. Xe x Cancellaria arnoldi Dall (pl. 1, figs. 15, 16, 22) x x >. Cancellaria, new species (pl. 1, figs. 23, 24) xX Ophiodermella graciosana (Arnold) (pl. 3, figs. 2, 3) _. X x x Megasurcula remondi (Gabb) (pl. 3, fig. 10) x OGDSEO TU 7A MSTC CLES are eee ee x PELECYPODS Anadara (Anadara) trilineata Conrad Diba oligss ines) hc seer es) ATU eae xX: ‘Spt xX Po Speman Modiolus species —_ pa Besned Bis..1ct ah tel see Wee x x NTU Sie SC CICS a aed oe, cae i ee Byres Xx Eepiopecten atauratus \(Contad) see ee Clinocardium cf. C. meekianum (Gabb) (po ees Si oes) ern ere En ee ee x Spisula albaria coosensis Howe (ol, Sy es, il, GO; Be fol, 4, ties, BO) — xX “shes se cle SPisulascia Smercedensts Packard. sees ee OX iS jregepomoys. (Comer) sa x Tellina (Peronidea) cf. T. (P.) lutea Wood TECHIE SPECIES) (OM Ane TGs 63) ees es x Macoma nasuta forma kelseyi Dall (pl. 4, fig. 6) —_ x Vor. XXXVII] ADDICOTT: LATE PLIOCENE MOLLUSKS FROM CALIFORNIA 63 TABLE 1. Continued. USGS Stanford U. localities localities a = & S& 1 = = s 2 CS) 0 GASTROPODS PE arciigulne tie | eae eS: = Macoma new species? aff. M. nasuta (Conrad) (pimecrmetiomeet tes pls 4 fio: 12) >.< sp. sp Macoma secta (Conrad) (pl. 3, fig. 4) x GI CH aCE SE STCONCUS GGOUIG 225s i X sp. sp. x ROL EXILE SDCCICS. MARE oo eee Se eS Dx Humilaria perlaminosa (Conrad) Arnold (1907) ? (gl & hm 1) aS ee x Protothaca staleyz (Gabb) (pl. 2, figs. 5, 6, 8) — er OX sp. sp. Cryptomya californica (Conrad) (plmeepatioseey 4. pl. 4 fie. 7) 2 2G 13,8 x TC OPESPeGleESm(plh 2, tips 9)\ See x 1 Listed by Arnold (1906). 2Jn collection from CAS locality 28617. 3 In collection from SU locality 2675. PALEONTOLOGY The molluscan fauna consists of 39 taxa (table 1), the majority of which are represented by material sufficiently well preserved to permit specific identifi- cation and illustration. The purpose of this section is to furnish supplementary documentation of specific identifications. To conserve space, conventional systematic treatment of each taxon is modified into systematically organized discussion in which only original descriptions of fossil species and other pertinent reference are indicated. This treatment seems appropriate here because nearly all of the species have been previously named and described and major taxo- nomic revisions are not set forth. Taxonomic notes are arranged systematically by families, and taxa are considered in the same order as on the faunal list (table 1). This abbreviated form of systematics was first employed in Pacific Coast Tertiary reports by Woodring and others (1940). Included in the molluscan faunal list are three species reported by Arnold (1906, 1908) that do not occur in recent collections: Pupillaria pupilla, Thais ostrina, and Leptopecten latiauratus. Invertebrates other than mollusks are ‘very rare. Scattered barnacle plates occur at USGS locality M1715 and an inarticulate brachiopod Discinisca cummingi, was reported from nearby exposures by Hertlein and Grant (1944). GASTROPODS TrocuwaE. Pupillaria pupilla (Gould) identified by Arnold (1906) from an unspecified locality near Felt Lake does not occur in later collections from 64 ’ CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. this area. His material is missing from the Pliocene stratigraphic collections at Stanford University. LirToRINIDAE. Two of the figured specimens of Littorina petricola Dall (1909) (pl. 1, figs. 4, 5) compare favorably with the holotype, a thick-shelled spirally sculptured specimen from the Pliocene Coos Conglomerate at Coos Bay, Oregon. A third specimen (pl. 1, figs. 2, 3) has closely spaced axial growth lamellae but no spiral ribbing. As such it resembles an abraded specimen from the Pliocene Cebada Fine-Grained Member of the Careaga Sandstone of the Santa Maria basin, California, figured by Woodring and Bramlette (1950, pl. 19, fig. 12) as Littorina cf. L. petricola. In other characteristics, such as the relatively small aperture and flat subsutural segment of the body whorl, this smooth specimen closely resembles spirally sculptured individuals from the same locality. It is therefore included with this species as “L. petricola smooth form.” PAGE a! Fricures 1, 6-8, and 10. Thais (Nucella) transcosana Arnold, USGS locality M1715. Figures 1 and 6, low-spired, spirally ribbed form; figure 1, height 16 mm., width 13.2 mm., USNM 650942; figure 6, height 13.6 mm., width 12 mm., USNM 650943. Figures 7 and 8, high-spired form; figure 7, height 29.7 mm., width 19.1 mm., USNM locality 650944; figure 8, height 25.3 mm., width 22.4 mm., USNM 650945. Figure 10, intermediate form, height 28.3 mm., width 19.8 mm., USNM 650946. Ficures 2-5. Littorina petricola Dall, USGS locality M1715. Figures 2 and 3, smooth form, height 16.3 mm., width 14.1 mm., USNM 650947; figure 4, height 8.3 mm., width 6.4 mm., USNM 650948; figure 5, height 10.4 mm., width 10.1 mm., USNM 950949. Ficures 9, 14. Bittium casmaliense Bartsch, USGS locality M1715. Figure 9, height 15.1 mm., width 5.8 mm., USNM 650950; figure 14, height 9.9 mm., width 4.4 mm. USNM 650951. Ficures 11, 13. Neverita (Glossaulax) cf. N. (G.) recluziana (Deshayes), USGS locality M1715. Figure 11, height 12.1 mm., width 13.4 mm., USNM 650952; figure 13, height 10.7 mm., width 10.5 mm., USNM 650953. Ficure 12. Cryptonatica aleutica Dall, USGS locality M1715. Height 11.2 mm., USNM 650954. Ficures 15, 16, and 22. Cancellaria arnoldi Dall, USGS locality M1715. Figure 15, height 21.5 mm., width 12.8 mm., USNM 650955; figure 16, height 31.5 mm., width 16 mm., USNM 650956; figure 22, height 23.5 mm., width 13 mm., USNM 650957. Ficure 17. Neptunea cf. N. tabulata (Baird), USGS locality M1715. Diameter of fragment 9.8 mm., USNM 650958. Ficure 18. Nassarius (Caesia) grammatus (Dall), USGS locality M1715. Height 29 mm., width 18.7 mm., USNM 648570. Ficure 19. Mitrella cf. M. gouldi (Carpenter), USGS locality M1715. Height 13.1 mm., width 6.2 mm., USNM 650959. Ficures 20, 21. Ocenebra interfossa Carpenter, CAS locality 28617. Height 13 mm., width 7.6 mm., CAS 13108. Ficures 23, 24. Cancellaria, new species, USGS locality M1715. Height 36.5 mm., width 19.5 mm., USNM 650960. PLATE 1 (ADDICOTT) PROC. CALIF. ACAD. SCI. 4TH SER., VOL. XX XVII, NO. 3 Vor. XXXVII] ADDICOTT: LATE PLIOCENE MOLLUSKS FROM CALIFORNIA 65 The condition of the shell suggests that the absence of spiral sculpture is due to morphologic variation rather than wear. This species was originally identified as Littorina planaxis by Arnold (1906) but was subsequently identified as L. petricola Arnold (1908, pl. 37, fig. 7). Littorina petricola differs from the Pliocene and Pleistocene species L. martana Arnold (1909, pp. 86-87, pl. 29, figs. 1, 2) from the Coalinga-Kettleman Hills area of the San Joaquin Valley by its thicker, much stouter shell and flatter whorl profile. CERITHIDAE. Specimens of Bittium casmaliense Bartsch (1911, p. 411, pl. 55, fig. 3) (pl. 1, figs. 9, 14) from USGS locality M1715 are characterized by four strong spiral cords crossed by much weaker, slightly arcuate axial ribs that form nodes at the intersection with the spiral cords. The two anterior spirals are of equal strength; the posterior spirals are weaker and more closely spaced. Superimposed on the primary sculpture is a network of microscopic spiral striae. All specimens are smaller than the holotype from the Pliocene Cebada Fine-Grained Member of the Careaga Sandstone of the Santa Maria basin, California. This is a northward range extension of this Pliocene index species which had not previously been reported from north of the Salinas Valley area (Durham and Addicott, 1965). CALYPTRAEIDAE. Apical fragments of a large, very thick-shelled Crepidula from localities M1715 and M1926 (pl. 2, fig. 10) probably represent the extinct Miocene and Pleistocene species C. princeps Conrad (1856). Miocene specimens of this species occur in the Monterey Shale about 500 feet stratigraphically below the base of the Merced(?) Formation of Dibblee (1966a) (USGS loc. M2547). It is also reported from many localities in the Merced Formation (Arnold, 1906; Martin, 1916; Dickerson, 1922; Glen, 1959). NATICIDAE. A small naticid with nontabulate whorls (pl. 1, fig. 12; pl. 4, fig. 2) is identified as the Pliocene to Holocene species Cryptonatica aleutica (Dall). It and two doubtfully identified fragments from locality M1715 also have the fairly heavy shell and thick umbilical plug bordered posteriorly by a narrow crescent-shaped depression characteristic of this species. Pliocene records of C. russa (Gould) by Glen (1959) and Faustman (1964) presumably are of the C. aleutica of this report. Use of C. russa for this taxon seems un- satisfactory in the light of discussions by Keen (in Burch, 1946, no. 56, p. 27) ~and Woodring (in Woodring and Bramlette, 1950, p. 72), indicating that what Dall (1919) presumed to be the type specimen of C. russa appears to be C. clausa; a rather thin-shelled northern species. Incomplete, abraded naticoid gastropods from USGS locality M1715 (pl. 1, figs. 11, 13) are similar to a form of Neverita recluziana (Deshayes) figured by Arnold (1907, pl. 54, figs. 14a, 14b) from the Pliocene Careaga Sandstone of the Santa Maria basin. The distinctive feature of this naticid is its restricted, 66 4 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. strongly grooved bilobed callus. It also has an open umbilicus within which are faint spiral grooves. A similar Neverita occurs in the lower Pleistocene part of the Saugus Formation in the western Ventura basin. A large collection from USGS locality M1753 contains a weakly shouldered low-spired form and a high-spired form that have deeply impressed callus grooves which terminate posteriorly in a deep pit near the margin of the inner lip. Available specimens from the Merced(?) Formation are too poorly preserved, however, to determine relation- ship to the Pleistocene specimens. MuriciaE. A muricid from CAS locality 28617 identified as Ocenebra interfossa Carpenter (pl. 1, figs. 20, 21) is similar to rugose late Pleistocene specimens from Point Ano Nuevo, southwestern San Mateo County, California (USGS locality M1690) but appears to have fewer, somewhat coarser axial ribs than the holotype (Palmer, 1958, pl. 23, fig. 1). This species has not previously been reported from the Merced Formation or coeval strata of late Pliocene age in the San Francisco Bay area, although it is doubtfully recorded from the Purisima Formation at Pillar Point, San Mateo County (Glen, 1959). THAISIDAE. A variable population of Thais from locality M1715 is identi- fied as T. trancosana Arnold (1908, p. 388, pl. 36, fig. 3). Only a few of the nearly 50 specimens available for study are closely comparable with Arnold’s holotype. These forms (pl. 1, figs. 7, 8) are characterized by a short laterally constricted aperture, extremely thick shell walls, and a moderately well-developed spire. One specimen with a narrow slot-like aperture (pl. 1, fig. 8) has uniform spiral ribbing comparable to the holotype. The typical form is contrasted with a group of small, low-spired, spirally sculptured specimens (pl. 1, figs. 1, 6) that cannot be consistently separated from specimens that seem to have morpho- logic characters linking them with 7. trancosana. On this form the penultimate > PLATE 2 (All specimens are from USGS locality M1715) Ficures 1, 3. Anadara trilineata (Conrad). Figure 1, length 57 mm., height 48.5 mm., USNM 650961; figure 3, length 53 mm., height 44 mm. USNM 650962. Ficures 2, 4. Cryptomya californica (Conrad). Figure 2, length 33.5 mm., height 24 mm., USNM 650963; figure 4, length 30.5 mm., height 22 mm., USNM 650964. Ficures 5, 6, and 8. Protothaca staleyi (Gabb). Figure 5, immature specimen, length 23.5 mm., height 19.2 mm., USNM 650965; figures 6, 8, height (incomplete) 49 mm., USNM 650966. Ficure 7. Tellina cf. T. lutea Wood. Length (incomplete) 63 mm., height (incomplete) 34 mm., USNM 650967. Ficure 9. Zirfaea? species. Length 26 mm., height 15 mm., USNM 650968, a rubber cast. Ficure 10. Crepidula cf. C. princeps Conrad. Diameter (incomplete) 36.5 mm., USNM 650969. Ficure 11. Macoma, new species? aff. M. nasuta (Conrad). Length 86 mm., height 63.5 mm., USNM 650970. PROC. CALIF. ACAD. SCI. 4TH SER., VOL. XXXVII, NO.3 (ADDICOTT) PLATE 2 Vot. XXXVII] ADDICOTT: LATE PLIOCENE MOLLUSKS FROM CALIFORNIA 67 whorl is angulated by a spiral ridge and the relatively large body whorl is faintly shouldered below the suture. The large intermediate group of specimens are smooth, thick-shelled, low-spired and usually have a thin coating of whitish shell material within the aperture (pl. 1, fig. 10; pl. 3, fig. 5). This coating is also found on typical forms of T. trancosana but not on the spirally ribbed form. Denticles and corresponding spiral lines are found within the aperture of all three forms. The typical high-spired form of Thais trancosana is similar to T. etche- goinensis Arnold (1909, p. 89, pl. 18, fig. 2) from the Etchegoin Formation of the San Joaquin Valley. It also resembles some of the smooth Holocene ‘‘morphs”’ (local populations) of T. /amellosa from the Puget Sound area figured by Kincaid (1957). On these taxa, however, the base of the body whorl is constricted into a longer, better defined anterior canal. Hertlein and Allison (1959) discuss other features by which these species can be differentiated. The small, spirally ribbed form of Thais trancosana is similar to a smooth, low-spired form of T. emarginata (Deshayes), 7. emarginata forma ostrina Gould, but its spirally ribbed, denticulate aperture and nonrecurved columellar lip serve to distinguish it from Gould’s taxon. Arnold (1908, pl. 36, fig. 7) figured a specimen of 7. ostrina from SU locality C321 near Felt Lake but additional specimens of this taxon were not found during the present study. Thais trancosana has been reported from the Pliocene Cantil Costero Formation of Santillan and Barrera (1930) in northwestern Baja California, Mexico, and is known from several localities in the Merced Formation of the San Francisco Bay area (Hertlein and Allison, 1959). A small, incomplete thaisid from USGS locality M1715 (pl. 4, figs. 10, 11) identified as Thais cf. T. lima (Gmelin) is distinguished from other species in the late Pliocene assemblages by its slender shell and unique sculpture of fine spiral ribs. Moreover, the aperture is smooth within and there is no evidence of spiral sculpture within the outer lip. The shell of Thais cf. T. lima is much thinner than on specimens of 7. trancosana of comparable size. NEPTUNEIDAE. A fragment of a small neptuneid consisting of two whorls with a cream-colored outer shell layer (pl. 1, fig. 7) has the spiral sculpture and strongly tabulate whorls of Neptunea tabulata (Baird), a widespread Pliocene to Holocene species. A large stout form of this species, NV. tabulata forma colmaensis Martin (1914, pp. 188-189, pl. 20, fig. 1), is reported from upper Pliocene strata in the northernmost part of San Mateo County (Martin, 1914; Glen, 1959). CoLUMBELLIDAE. Two specimens from USGS locality M1715 are doubt- fully identified as Mitrella cf. M. gouldi (Carpenter). The larger of these (pl. 1, fig. 19; pl. 4, fig. 4) has convex whorls and a large evenly rounded body whorl. These characteristics suggest identification as M. gouldi rather than M. gausapata (Gould), at least in the sense that this taxon has been recognized by 68 4 CALIFORNIA ACADEMY OF SCIENCES [| Proc. 4TH SER. Grant and Gale (1931, p. 695) and Abbott (1954, p. 222, pl. 20, fig. m). The doubtful identification is necessitated by the fact that the unfigured type specimen has been lost and the attendant uncertainties as to its correct identifi- cation. This taxon appears to be the same as Astyris richthofeni (Gabb) of Arnold (1908, pl. 36, fig. 8) and M. gouldi (Carpenter) of Touring (in Cum- mings and others, 1962, photo 16, no. 7), both from the Pliocene Purisima Formation of coastal San Mateo County. Judging by Gabb’s figure (1866, pl. 2, fig. 16) of the holotype of M. richthofeni, which has since been lost, this Pliocene species has a relatively high spire with flattened whorls. As such it seems more closely allied to M. gausapata than to M. gouldi, as suggested by others (Grant and Gale, 1931; Woodring and Bramlette, 1950). NASSARIIDAE. Two species of Nassarius, N. californianus (Conrad, 1856) and NV. grammatus (Dall, 1917), occur in the late Pliocene assemblages. There has been considerable confusion over the identification of these distinctive Pliocene species (Addicott, 1965a). The smaller species, N. californianus (pl. 3, fig. 7), the most abundant gastropod in these assemblages, was originally described from marine Pliocene strata in northern Santa Clara County pre- sumably interbedded with the Santa Clara Formation. It is readily distinguished from the larger, NV. grammatus (pl. 1, fig. 18; pl. 4, fig. 5), by its coarser sculpture, slender profile, and narrow umbilical callus. Nassarius grammatus is perhaps better, though incorrectly, known as NV. moranianus (Martin), a late Pliocene and early Pleistocene species characterized by a medially angulated > PLATE 3 (All specimens are from USGS locality M1715) Ficures 1, 6, and 8. Spisula albaria coosensis Howe. Figure 1, length 74 mm., height 58 mm., USNM 650971; figure 6, length 55.5 mm., height 45 mm., USNM 650972; figure 8, length 57.5 mm., height 44 mm., USNM 650973. Ficures 2, 3. Ophiodermella graciosana (Arnold). Height 17.8 mm., width 8 mm., USNM 650974. Ficure 4. Macoma secta (Conrad). Length 38.5 mm., height 26.5 mm., USNM 650975. Ficure 5. Thais (Nucella) trancosana Arnold. Intermediate form. Height 21.3 mm., width 16.4 mm., USNM 650976. Ficure 7. Nassarius (Demondia) californianus (Conrad). Height 15 mm., width 8.3 mm., USNM 648550. Ficure 9. Olivella biplicata (Sowerby). Height 19.5 mm., width 9.9 mm., USNM 650977. Ficure 10. Megasurcula remondi (Gabb). Height 34.5 mm., width 16.4 mm., USNM 650978. Ficure 11. Clinocardium cf. C. meekianum (Gabb). Length (incomplete) 46 mm., height (almost complete) 47.5 mm., USNM 650979. Ficure 12. Humilaria perlaminosa (Conrad) of Arnold (1907)? Length of fragment 66 mm., USNM 650980. PROC CALIF: ACAD: SCI. 4TH SER., VOL. XXCXVIT, NO. 3 (ADDICOTD) PLATE 3 VoL. XXXVII] ADDICOTT: LATE PLIOCENE MOLLUSKS FROM CALIFORNIA 69 body whorl and sculptural features that are more closely allied to the Quater- nary species JV. fossatus than to V. grammatus. CANCELLARIIDAE. Specimens of Cancellaria arnoldi Dall (1909, pp. 29-30, pl. 14, fig. 7) are characterized by strongly shouldered whorls and a well- developed siphonal fasciole. Twenty-five specimens of this slender cancellariid in the collection from USGS locality M1715 exhibit considerable variation in sculptural detail. Axial sculpture is dominant on some specimens (pl. 1, fig. 22) and weak on others (pl. 1, fig. 16). Secondary spiral sculpture is developed on some of the specimens. The thickness of the parietal callus and development of the siphonal fasciole are also variable. Collection from locality M1715 is a north- ward range extension of this species; it was previously reported only from localities in the southern part of California: Santa Maria basin (Woodring and Bramlette, 1950), doubtfully from the eastern Ventura basin (Winterer and Durham, 1962), and the San Diego area (Dall, 1909). It has recently been collected from the Purisima Formation near Capitola, Santa Cruz County, California (USGS locality M2462), where it occurs in a shallow water assemblage similar to Merced assemblages from the San Francisco Bay area. A finely sculptured cancellariid from USGS locality M1715 appears to be undescribed. It resembles Cancellaria lipara Woodring (in Woodring and Bramlette, 1950, p. 76, pl. 16, figs. 13, 14) but has a moderately high spire, a well-defined anterior canal, and a concave subsutural slope. Cancellaria new species seems to be conspecific with a small abraded specimen from the Pliocene San Diego Formation (USNM 56207) which Woodring (im Woodring and Bramlette, 1950, p. 76) compared with the holotype of C. /ipara. There is no suggestion of sculptural intergradation between this relatively smooth species and the more than 30 specimens of C. arnoldi in the collection from USGS locality M1715. OtivipAE. Specimens of Olivella biplicata (Sowerby) (pl. 3, fig. 9) characterized by a low spire, heavy parietal callus, and a grooved spiral fold at the base of the columella occur- abundantly at locality M1715. Some of the minute specimens resemble O. pedroana (Conrad) but can be distinguished by their bilobed basal columellar fold and lower spire. TURRIDAE. Specimens of Ophiodermella graciosana (Arnold, 1907, pp. 430-431, pl. 54, fig. 18) compare very closely with the abraded holotype from the upper Pliocene Careaga Sandstone of the Santa Maria basin. Drillia mercedensis Martin (1914, pp. 194-195, pl. 22, figs. 2a-2c) is a synonym of this species. Available records of D. graciosana (Glen, 1959; Woodring and Bramlette, 1950; Martin, 1916; Nomland, 1917; Waterfall, 1929; Soper and Grant, 1932) suggest that this Pliocene and early Pleistocene species ranges. no lower than formations generally designated as late Pliocene in a twofold pro- vincial division of the epoch. However, a satisfactory age assignment cannot be made for some occurrences of this species in bulk collections from the Purisima 70 . CALIFORNIA ACADEMY OF SCIENCES [PRoc. 4TH SER. Formation of the Santa Cruz Mountains (Martin, 1916) for which strati- graphic data are lacking. Megasurcula remondi (Gabb, 1866, p. 3, pl. 1, fig. 5), a low-spired and rather stout species, is represented by two well-preserved specimens from USGS locality M1715. The larger of these (pl. 3, fig. 10) is of the same size as the lectotype. It compares very closely with Stewart’s figure (1927, pl. 31, fig. 5) and has the general proportions of Gabb’s original figure, a poor line drawing. This species was described from the Merced Formation of Sonoma and Marin County northwest of San Francisco but was subsequently identified as M. carpenteriana fernandoana (Arnold) by Dickerson (1922). Although this short, weakly angulated turrid has been treated as a form of M. carpenteriana (Gabb), it seems sufficiently distinct to recognize it as a separate species. This species seems to be limited to strata of Pliocene age, although there is a single specimen reported (Grant and Gale, 1931, p. 496) from undifferentiated Pliocene and Pleistocene strata in the Ventura basin near Fillmore. PYRAMIDELLIDAE. The two strongly abraded, incomplete specimens of Odostomia from USGS locality M1715 are specifically indeterminate and too poorly preserved to be figured. PELECYPODS ArcipaE. About 25 large, disarticulated valves of Anadara trilineata (Conrad, 1856) were collected from USGS locality M1715. Although the exterior and perimeter of these specimens are considerably abraded (pl. 2, figs. 1, 3), there are small specimens in the collection that shows the beaded orna- mentation of ribs characteristic of this species. Most specimens have 26 ribs, although there are as few as 24 on some specimens and as many as 28 on others. MytTiLtwarE. Two incomplete, specifically indeterminate specimens of Modio- PLATE 4 Ficure 1. Protothaca staleyt (Gabb), USGS locality M1720. Length 18 mm., height 17 mm., USNM 650981. Ficure 2. Cryptonatica aleutica Dall, USGS locality M1720. Height 15.3 mm., width 8.5 mm., USNM 650982. Ficurr 3. Tellina species, USGS locality M1715. Height 21 mm., USNM 650983. Ficure 4. Mitrella cf. M. gouldi (Carpenter), USGS locality M1715. Height 13.1 mm., width 6.2 mm., USNM 650959. Ficure 5. WNassarius (Caesia) grammatus (Dall), USGS locality M1715. Height 27.6 mm., width 18.2 mm., USNM 648592. Ficure 6. Macoma nasuta forma kelseyi Dall, USGS locality M1715. Length 78 mm., height 50.5 mm., USNM 650984. Ficure 7. Cryptomya californica (Conrad), USGS locality M1715. Length 33.5 mm., width 24 mm., USNM 650963. Ficures 8, 9. Spisula labaria coosensis Howe, USGS locality M1715. Figure 8, Hinge 1, USNM 650972; Figure 9, hinge 1, USNM 650985. Ficures 10, 11. Thais (Nucella) cf. T. (N.) lima Gmelin, USGS locality M1715. Height 11.8 mm., width 8.8 mm., USNM 650986. Ficure 12. Macoma, n. sp.? aff. M. nasuta (Conrad), USGS locality M1715. Length 86 mm., height 63.5 mm., USNM 650970. PROC. CALIF. ACAD. SCI. 4TH SER., VOL. XXXVII, NO.3 (ADDICOTT) PLATE 4 <=a¥* -~ VoL. XXXVII] ADDICOTT: LATE PLIOCENE MOLLUSKS FROM CALIFORNIA 71 lus were collected from USGS locality M1715. The largest (110 mm.) is severely deformed. The fragment of a beak of a Mytilus with a broadly acute apical angle is in a Stanford University collection from a ditch connecting Felt Lake with Los Trancos Creek (SU loc. C321). PecTINIDAE. Arnold (1906) reported Leptopecten latiauratus (Conrad) from exposures near Felt Lake. This record has not been substantiated by subsequent collecting, nor has the material upon which the identification was based been found in Stanford University collections. CarpimwDaAE. A badly exfoliated cardiid (pl. 3, fig. 11) is identified as Clinocardium cf. C. meekianum (Gabb, 1866). Although poorly preserved, the trigonal shape, strongly inclined beaks, and the character of ribbing indicates a close degree of similarity to this Pliocene index species. Moreover, the size and density of radial ribs are comparable to doubtfully identified specimens from the Kettleman Hills Pliocene of the San Joaquin Valley (Woodring and others, 1940) and specimens in U.S. Geological Survey collections from the Empire Formation near Cape Blanco, Oregon. MacrripaE. Several specimens of Spisula albaria coosensis Howe (1922) were collected from USGS locality M1715. The absence of a distinct posterior ridge and the relatively weak development of the left anterior cardinal tooth (pl. 4, fig. 8) are features noted by Howe (1922, p. 100) that distinguish this taxon from Spisula albaria (Conrad), a Miocene species from the Astoria Formation of northwestern Oregon. There is considerable variation in outline of these specimens—from a form with a strongly produced anterior extremity (pl. 3, figs. 1, 8) to one which is almost equilateral (pl. 3, figs 6, 8). Similar variation in profile of middle Miocene specimens from the Astoria Formation of Oregon was described by Moore (1963, p. 83). The nonequilateral form compares very closely in external morphology with middle Miocene specimens in U.S. Geological Survey collections from the Astoria Formation of the Newport embayment, Oregon. The equilateral form is similar to a taxon from the Astoria Formation of southwestern Oregon named S. albaria goodspeedi by Etheringtcn (1931, pp. 86-87, pl. 9, fig. 3). One incomplete left valve of a Spisula in the collection from locality USGS M1715 is much larger than the others and has a subquadrate outline. Although not suitable for illustration, it is doubtfully identified as Spisula cf. S. merce- - densis Packard (1916) because of its unique profile and similar hinge. Although poorly preserved, it can be differentiated from the similar S. catilliformis Conrad because the anterior lateral tooth is separated from the anterior cardinal. Tresus pajaroanus (Conrad, 1857) is identified on the basis of a crushed right valve from USGS locality M1715 that has the broad siphonal gape characteristic of the genus and a very long posterior dorsal slope. The specimen is unsuitable for figuring. 72 ( CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. SOLENIDAE. Several broken, thin-shelled specimens of Solen collected from locality USGS M1715 are doubtfully identified as Solen cf. S. sicarius Gould. Two of the fragments seem to have a gently curved dorsal margin, the principal shell character by which this species is distinguished from the similar Recent species S. rosaceus Carpenter. TELLINIDAE. An incomplete left valve of a large, thick-shelled tellinid is identified as Tellina cf. T. lutea Wood. The concentric sculpture and broadly obtuse umbonal angle resemble the living circumboreal species, which ranges from northern Japan (Habe, 1964, p. 202) to Cook Inlet, Alaska. The Pliocene specimen (pl. 2, fig. 7) has a somewhat broader umbonal angle and a heavier shell and hinge plate than specimens of 7. dutea in U.S. Geological Survey Holo- cene collections from Alaska. Similar forms occur in the Pliocene of the Santa Maria basin (Woodring and Bramlette, 1950, p. 87) and the Salinas Valley (Durham and Addicott, 1965, p. Al4). The genus Tedlina is further represented by a fragment of a moderately small, thick-shelled specimen identified as Tellina species (pl. 4, fig. 3). Al- though the posterior part of the valve is missing, there is an impression in the sandstone matrix in which the shell is imbedded that indicates a short, steep posterior dorsal slope unlike that of Tellina cf. T. lutea with which it occurs. Indeed, the posterior location of the beak and outline inferred from the sand- stone cast are more similar to 7. bodegensis (Hinds), although the shell seems thicker for its size and is much more produced ventrally than that species. The smooth surface of the shell is sculptured by evenly spaced concentric bands that are slightly tilted ventrally, giving the surface a finely imbricate appear- ance. The left valve of a very large specimen of Macoma (pl. 2, fig. 11; pl. 4, fig. 12) allied to M. nasuta (Conrad) may represent an undescribed species, but there is insufficient material to confidently separate it from M. nasuta. The pallial sinus extends forward to the anterior muscle scar as in M. nasuta, but the ligamental groove is much longer and the posterior extremity is broadly truncated rather than acute. Moreover, the beak is situated posterior to the midline of the valve and the valves are strongly flexed. On the only well- preserved specimen, the flexure begins midway between the umbo and the posterior extremity in dorsal view, whereas it begins in the anterior quarter of the shell along the ventral margin. In M. nasuta flexing of the ventral margin usually takes place directly below the beaks near the middle or slightly anterior to the middle of the shell. The degree of flexing may be a variable charac- teristic because a broken right valve from USGS locality M1715 that seems to belong with Macoma new species? because of its relatively long ligamental groove and posteriorly situated beak appears to be very weakly flexed. There is an unfigured, poorly preserved right valve (about 70 mm. long) that has a strong posterior flexure and broadly truncate posterior extremity VoL. XXXVII] ADDICOTT: LATE PLIOCENE MOLLUSKS FROM CALIFORNIA 73 similar to the larger left valve described above. Yet there are too few specimens to provide an adequate measure of possible variation. Accordingly, one cannot rule out the possibility that it is an aberrant Macoma nasuta (Conrad), as it occurs with the large thick-shelled variety M. nasuta forma kelsevi (Dall). A few incomplete specimens identified as Macoma nasuta forma kelseyi Dall are differentiated from the previously described taxon by their acutely pointed posterior extremities. These also are large, thick-shelled individuals (pl. 4, fig. 6) similar to late Pliocene specimens from the Careaga Sandstone of the Santa Maria basin figured by Woodring and Bramlette (1950, p. 87, pl. 20, figs. 2, 8) but they have an apparently stronger posterior flexure of the valves. Macoma secta (Conrad), represented by a thin-shelled immature right valve (pl. 3, fig. 4), is differentiated from the other macomas with which it occurs by its very short ligamental area and characteristic arcuate dorsal margin above the posterior ridge. VENERIDAE. The genus Katherinella is represented by a poorly preserved specimen from an old Stanford University locality (SU loc. C308) on Purisima Creek about 1% miles southeast of USGS locality M1715. An incomplete, thick-shelled valve of venerid pelecypod (pl. 3, fig. 12) identified as Humilaria perlaminosa (Conrad)? has been compared with a holo- type (USNM 165252) figured by Arnold (1907, pl. 58, figs. 1a, 1b) from the Santa Barbara Formation of southern California. Both specimens have rugose primary concentric sculpture which, where well preserved, tends to conceal a secondary network of fine radial ribs. However, specific identity is uncertain because about a third of the valve, including the hinge, is missing. If identifica- tion as H. perlaminosa (Conrad) were undoubted, this would be the northern- most recorded occurrence of this species in addition to its first record in strata of Pliocene age. Previously reported occurrences are from rocks of early Pleistocene age in southern California (Arnold, 1907; Waterfall, 1929; Woodring and others, 1946).” Protothaca staleyi (Gabb, 1866) (pl. 2, figs. 5, 6, 8; pl. 4, fig. 1) is repre- sented by several thick-shelled valves and many fragments. The subquadrate outline of the valves and incised line defining these specimens are features which Howe (1922, p. 98, pl. 10, fig. 1) used to separate a form from the Pliocene Empire Formation at Coos Bay, Oregon (P. staleyi forma hannibali Howe), from the typical form. Yet the rather smooth exterior with only faint radial sculpture is characteristic of the neotype from the Merced Formation of Sonoma County (Stewart, 1930, p. 233, pl. 15, fig. 4). The hinge of this species differs from that of the Pliocene to Holocene P. staminea. The lower part of the anterior cardinal tooth of the left valve of P. staleyi is strongly reflected anteri- 2 There is an unreported occurrence in an unnamed Pliocene formation at Newport Bay, Orange County, California (J. G. Vedder, oral communication, January 1968). 74 4 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. orly, whereas in P. staminea it is straight, much shorter, and oriented at 90 degrees to the long axis of the valve. Myiaer. Several disarticulated valves of Cryptomya californica Conrad occur in the late Pliocene collections from northern Santa Clara County (pl. 2, figs. 2, 4; pl. 4, fig. 7). The outline of these specimens is variable; some are elongate, others are suborbicular. Some compare closely with C. ovalis Conrad (Arnold, 1908, pl. 36, fig. 9), one of several names proposed for late Tertiary cryptomyas which seem to be best regarded as forms of the Holocene C. cali- fornica (Grant and Gale, 1931, p. 417; Woodring and Bramlette, 1950, p. 91). PHOLADIDAE. Two incomplete external molds of a small pholadid pelecypod found in extensively bored blocks of Monterey Shale collected from locality M1715 are doubtfully identified as Zirfaea species. A rubber cast of the better of these is figured (pl. 2, fig. 9). AGE AND POSITION IN THE PROVINCIAL CHRONOLOGY More than a third of the taxa in the Merced(?) assemblages from northern Santa Clara County are extinct.* Thus in the context of classical Lyellian age classification (Keen, 1939; Bird, 1967), probably the only way by which the highly endemic Neogene molluscan faunas of the Pacific Coast can be directly compared with those of European type sections, a Pliocene age is clearly indicated. Distinctive extinct taxa include Littorina petricola, Thais trancosana, Cancellaria arnoldi, C. lipara, Anadara trilineata, Clinocardium ci. C. meekianum, Protothaca staleyi, Tresus pajaroanus, and Spisula albaria coosensis. None of these species have been reported from strata usually classified as Pleistocene in the Pacific coast provincial sequence (Weaver and others, 1944; Woodring, 1952; Valentine, 1961). Some workers now place the Pliocene-Pleistocene boundary within the upper part of the traditional marine Pliocene sequence of the Pacific Coast (Moore, 1949; Durham iz Durham and MacNeil, 1967; Repenning, 1967) on the basis of indirect correlation of mammalian assemblages and associated radiometric age determinations with the European nonmarine mammal chro- nology. Lowering of this boundary vitiates the long-standing criteria upon which the marine shallow water Pliocene of the Pacific Coast is discriminated ‘from the Pleistocene (Woodring, 1952). Accordingly, use of Pliocene in this report is in a provincial sense and is dictated by utilitarian considerations. Although it is in keeping with long-established standards that honor extinction percentages compatible with the Lyellian principle, it is not necessarily com- parable to European usage. Clearly, provincial stage-age classification of the Pacific Coast marine Pliocene would obviate the need for utilization of European epoch terminology, but unfortunately, adequately defined provincial time- stratigraphic terminology based upon larger invertebrates has not been proposed. Assignment to the late Pliocene, in the sense of a twofold division of the ® A comparable percentage of extinct species was determined for the Merced Formation by Smith (1919, p. 145). Vot. XXXVII] ADDICOTT: LATE PLIOCENE MOLLUSKS FROM CALIFORNIA 75 Pliocene molluscan sequence in California (Vedder, 1960; Durham and Addicott, 1965), is based upon faunal correlation with well known Pliocene sequences that are generally used as reference sections for the shallow water Pliocene of Calli- fornia. The principal one is the type section of the Merced Formation along Seven Mile Beach near the northwestern end of the San Francisco Peninsula. This section spans the provincial Pliocene-Pleistocene boundary as long recognized on the Pacific Coast (Arnold, 1906; Weaver and others, 1944; Woodring, 1952; Valentine, 1961). About two-thirds of the taxa either occur in the lower Merced fauna or are represented by closely related taxa, many of which are doubtfully identified forms that might be conspecific. The general aspect of the present faunal assemblages also corresponds closely to the ecologically comparable upper part of the classic Pliocene section of the Kreyenhagen Hills-Kettleman Hills area (Arnold, 1909; Nomland, 1917; Woodring and others, 1940) of the San Joaquin Valley about 125 miles to the southeast. More of the species occur in the diverse molluscan fauna of the upper part of the Pliocene sequence of the Santa Maria Basin (Woodring and Bramlette, 1950) located somewhat farther to the southeast than in other California sections. However, this seems to be in part a manifestation of the greater diversity of the late Pliocene molluscan fauna of the Santa Maria basin. An early Pliocene age seemingly can be ruled out by the fact that none of the early Pliocene index species characteristic of the Pancho Rico Formation of the Salinas Valley area or the Jacalitos Formation of the San Joaquin Valley occur in this fauna. Moreover, species such as Ophiodermella graciosana, Humilaria perlaminosa, Cancellaria arnoldi, C. lipara, and Leptopecten latiauratus are not known to occur in strata generally classified as lower Pliocene according to a twofold division of the Pliocene Series. In this regard, if one accepts limits of about 3 million and 10 or 11 million years for the Pliocene Series in California, there seems to be an unrealistically short interval of time to permit a three- fold division based upon evolutionary phenomena. The Miocene Series in Cali- fornia, for example, has been divided by recent workers (Corey, 1954; Repenning and Vedder, 1961; Addicott, 1965c) into only three, or possibly four, time- stratigraphic units based upon molluscan data, although its duration is more than twice that of the Pliocene. Assignment of this fauna to the late Pliocene is not a new concept. It was initially considered to be late Pliocene by Arnold (1906) by inclusion of expo- sures near Felt Lake in his upper Pliocene “zone” or “horizon’”—the Merced Formation. However, the only recorded taxa from this locality (Arnold, 1906; p. 25) were living species. 4 Despite the use of the term Formation, which is now considered to be a lithostratigraphic term, Arnold’s utilization of zone and horizon, together with type sections and references of fossiliferous sections from widely separated areas in California to his Tertiary Formations, very nearly fulfills our modern time- stratigraphic concept of stages. He included only the lower part of the type Merced Formation in the Pliocene. 76 , CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. PALEOECOLOGY By and large the molluscan assemblages represent a very shallow, water- nearshore environment not unlike modern bays and estuaries of the central Cali- fornia coast. Many of the species, for example, are still living in the intertidal zone or uppermost part of the inner sublittoral zone (Hedgpeth, 1957) of nearby San Francisco Bay: Pupillaria pupilla, Ocenebra interfossa, Thais ostrina, Olivella biplicata, Leptopecten latiauratus, Solen sicarius, and Cryptomya californica (Packard, 1918).° These and other taxa that are represented by closely related species in San Francisco Bay or nearby Elkhorn Slough (latitude 36.8° N.) comprise more than half of the specifically identified taxa in the late Pliocene assemblages (table 2). There are, however, relatively deep-water elements, as well as both northern and southern faunal elements, that suggest environmental conditions at least somewhat different from these modern, shallow- water bays. Pertinent molluscan bathymetric data on living species, modern analogs, or supraspecific taxa (chiefly from Packard, 1918; Oldroyd, 1924; Burch, 1944— 1946; Fitch, 1953; Abbott, 1954; Quayle, 1960; Coan im Keen, 1963; and Parker, 1964) suggest that the late Pliocene assemblages lived in the littoral (intertidal) zone or the highest reaches of the inner sublittoral zone. Although nearly all of the living species and comparable generic taxa range downward into the middle or lower reaches of the inner sublittoral zone (20—50 fathoms), the most abundant species—Thais trancosana, Nassarius californianus, N. grammatus, Olivella biplicata, Cancellaria arnoldi, Anadara trilineata, Proto- thaca staleyi, Macoma nasuta group, and Cryptomya californica—are certainly indicative of a very shallow-water, level-bottom community. Excepting the two extinct warm-water taxa that have no closely related survivors along the California coast, Anadara (Anadara) trilineata and Cancellaria arnoldi, each of these is either still living or is represented by an analogous species in San Francisco Bay or Elkhorn Slough. There are, however, three gastropods of deeper water aspect in the assem- blage: Cryptonatica cf. C. aleutica, Neptunea cf. N. tabulata, and Megasurcula remondi. These species are still living in the latitude of central California at depths of 20 fathoms or greater. The seemingly anomalous association of these with the shallow-water, nearshore assemblage which includes the intertidal gastropod Littorina is difficult to interpret. Perhaps these species are best regarded as strays that somehow ventured into unusually shallow water. That this is not their preferred environment is suggested by the fact that each is represented by only one or two individuals. The apparently equal representation of gastropods and pelecypods suggested by the faunal list (table 1) does not accurately reflect the numerical dominance 5 Packard’s study preceded the effects of urbanization of the San Francisco Bay area on the molluscan fauna, and the concomitant establishment of many introduced mollusks. Vor. XXXVI] ADDICOTT: LATE PLIOCENE MOLLUSKS FROM CALIFORNIA 77 TaBLe 2. Some late Pliocene mollusks that are still living, or are represented by closely related species, in central California bays or estuaries. GASTROPODS Thais trancosana Nassarius californianus PELECYPODS Occurrence (Packard, 1918; Fitch, 1953) Related form (T. lamellosa) living in San Francisco Bay Related form (N. mendicus) living in San Francisco Bay Nassarius grammatus Related form (JN. fossatus) living in San Francisco Bay Clinocardium cf. C. meekianum Related form (C. nuttallii) living in San Francisco Bay Protothaca staleyi Related form (P. laciniata) living in Elkhorn Slough Similar form (S. catilliformis) living in San Francisco Bay Spisula cf. S. mercedensis Tresus pajaroanus Similar form (T. nuttalli) living in San Francisco Bay Macoma aff. M. nasuta Similar form (M. nasuta) living in San Francisco Bay Macoma nasuta kelseyi Related form (M. nasuta) living in San Francisco Bay Macoma secta Living in Elkhorn Slough of pelecypods in the faunal assemblages. Factors of preservation and the mode of accumulation may account for a gastropod : pelecypod ratio that is not repre- sentative of comparable shallow-water, sand-flat biotopes of modern bays along the California coast. Broken pelecypod shells are the dominant faunal element, yet whole or nearly complete shells are rarely encountered. Entire pelecypod valves could be collected only from large blocks that were excavated by power equipment at construction sites. Excellent gastropod specimens, on the other hand, were relatively simple to collect because of their small size and simple, cylindrical shape. And in this assemblage, abundance figures taken from col- lections would be highly biased toward gastropods; this is simply a measure of their ability to withstand destruction prior to burial because of their stream- lined form. Thus gastropods probably represent a much more thoroughly collected faunal element than pelecypods, because of the limited availability of large blocks and the tendency for the disarticulated, essentially two- dimensional pelecypod shells to be broken by wave or current action prior to final burial. The large percentage of broken shells and the usual disarticulated condition of the valves of burrowing pelecypods suggests that the shell beds represent a 78 t CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. death assemblage that has undergone considerable modification by wave or current action. Yet the numerically dominant mollusks in the assemblage are allied to modern taxa that characteristically inhabit a sandy, level-bottom substrate in the intertidal zone or high inner-sublittoral depths of bays and estuaries. Only minor mixing of elements from another biotope is suggested by the occurrence of rare specimens of three taxa charactersitic of the middle or lower parts of the inner sublittoral zone. ZOOGEOGRAPHY Included in the late Pliocene assemblages from northern Santa Clara County are a few molluscan taxa whose descendants or modern analogs do not occur together in the molluscan fauna now living off the central and northern California coast. These extralimital mollusks consist of both northern and southern elements (table 3) whose modern endpoints of latitudinal range are separated by at least several hundred miles along the Pacific Coast. It is difficult to interpret the joint occurrences of molluscan taxa that do not have overlapping zoogeographic distributions in the modern, shallow-water provinces of the northeastern Pacific Ocean. One possibility is that some invertebrates may have become adapted to different environmental conditions through competitive pressure or physiologic change. Were both the northern and southern elements represented by large populations in the assemblages, this would seem a good likelihood. In this case, however, only the southern forms of Cancellaria and Anadara® are represented by large, apparently well-established reproductive populations. In addition, the genus Neverita is not definitely known to occur north of the Californian molluscan province (northern boundary near 34.5° N.) according to Burch (1946) and Smith and Gordon (1948). By way of contrast, the northern taxa Tellina cf. T. lutea and Macoma nasuta forma kelseyi are represented by only one or two specimens. Accordingly, it can be argued that these northern taxa represented a brief southerly range extension during relatively cool parts of the marine climatic cycle but were unable to establish reproductive populations because of unfavorable environ- mental conditions, presumably excessively warm water temperatures. Cancellaria and Anadara occur, then, in a temperate, or possibly warm- temperate, protected inner-coast assemblage in which additional warm-water taxa are not represented. Possibly this assemblage reflects discontinous late 6 This is about the northernmost occurrence of Amadara s.s. in the late Pliocene of the Pacific Coast. A more northerly occurrence in the Ohlson Ranch Formation of Higgins (1960) in Sonoma County (Peck, 1960) may be of early Pliocene age judging by the associated fauna. In the early Pliocene it occurred as far north as southwestern Oregon, where it is found in the Empire Formation. By early Pleistocene time the genus had disappeared entirely from California. A species referable to the subgenus Cunearca occurs in the late Pleistocene of southern California. Two modern records of A. (Larkinia) multi- costata from southern California (Burch, 1945) have not been verified by subsequent collecting and are regarded doubtful. Further doubt is cast upon these records by the fact that this species is not known from the late Pleistocene of southern California (J. G. Vedder, oral communication, 1968). VoL. XXXVII] ADDICOTT: LATE PLIOCENE MOLLUSKS FROM CALIFORNIA 79 Tasie 3, Extralimital late Pliocene taxa from northern Santa Clara County. Species Distribution of modern analog Taxa with southern affinities: 1. Neverita ci. N. recluziana Living from Point Conception, Calif. to Tres (Deshayes) Marias Islands, Mexico (Burch, 1946). Old records from central and northern California are doubt- ful (Burch, 1946; Smith and Gordon, 1948) 2. Cancellaria arnoldi Dall Presumably referable to Cancellaria, s. s. species of which range from Cedros Island, Baja Cali- 3. Cancellaria, new species fornia, southward (Keen, 1958). Cancellaria emydis Dall and Ochsner, a Quaternary species from the Galapagos Islands, seems to be most closely related to C. arnoldi. There are no closely related species in modern Californian or Oregonian molluscan provinces. 4. Megasurcula remondi (Gabb) Living from Monterey Bay to Todos Santos Bay, Baja California (Burch, 1946) 5. Anadara (Anadara) trilineata Subgenus ranges from Cedros Island, Baja Cali- (Conrad) fornia southward (Keen, 1958) Taxa with northern affinities: 1. Tellina cf. T. lutea Wood Living from northern Japan to Cook Inlet, Alaska (Habe, 1964; Burch, 1944) 2. Macoma nasuta kelseyi Dall Living from Puget Sound to Coos Bay, Oregon (Burch, 1944). Pliocene zoogeographic distributions in which isolated populations of warm-water taxa existed far north of their normal outer-coast distribution by inhabiting the relatively warm water of protected bays. The co-occurrence of northward and southward ranging mollusks is not unique in the California Neogene. Similar joint occurrence of late Pliocene mollusks with mutually exclusive modern geographic ranges is reported by Soper and Grant (1932) from the Los Angeles basin. The subject is also treated by Woodring and others (1946), Valentine (1955; 1961) and Emerson (1956) as it concerns late Pleistocene mollusks from southern California and northwestern Baja California. The sum of the zoogeographic evidence seems to point, therefore, to a shallow- water marine climate that was probably somewhat warmer than occurs in bays along the California coast today. This is at variance with Smith’s (1919) con- clusion that the coeval lower type Merced fauna indicated water temperatures that were much cooler than exist today in the vicinity of San Francisco. Durham (1950), noting the occurrence of the small warm-temperate element in the type Merced assemblage, suggested minimum water temperatures as warm as OF slightly warmer than at this latitude today. It is notable that the warm-water cancellariids have not been reported from the type Merced. The genus Veptunea, 80 ( CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. however, is relatively abundant at localities in this formation (USGS loc. M1661; UCMP loc. 1736), suggesting at least local deeper and cooler water conditions than occur in the northern Santa Clara County Pliocene to the southeast. LATE PLIOCENE PALEOGEOGRAPHY Correlation of the late Pliocene fauna of the northern Santa Clara County area has significant bearing on Pliocene paleogeography of the San Francisco Peninsula and on the history of movement along the Santa Cruz Mountains seg- ment of the San Andreas fault. Principal exposures of late Pliocene strata on and near the San Francisco Peninsula are, 1) Bolinas Bay (Martin, 1916), 2) San Francisco and northwestern San Mateo Counties (Lawson, 1893; 1895; Ashley, 1895; Arnold, 1906; Martin, 1916; Glen, 1959), 3) upper part of the Purisima Formation along the San Mateo County coast (Ashley, 1895; Haehl and Arnold, 1904; Martin, 1916; Cummings and others, 1962), and 4) northern Santa Clara County (Arnold, 1906; 1908; Branner and others, 1909; this report). Appar- ently correlative strata occur along the north shore of Monterey Bay and in the Sargent-Pajaro River area to the southeast (Martin, 1916) and in Sonoma County to the northwest (Osmont, 1905; Dickerson, 1922). As indicated on the paleogeographic map (fig. 2), these exposures define relatively narrow, northwest-trending embayments. Of particular concern is the relationship of the northern Santa Clara County Pliocene to the coeval lower Merced Formation of the northern San Francisco Peninsula. The continuous band of Merced Formation extends from San Francisco southward to near Burlingame, from which point Lawson (1914) theorized that it continued southeastward beneath San Francisco Bay. However, a comprehensive recent study of the subsurface stratigraphy and bedrock configuration of the South [San Francisco] Bay area (Finlayson and others, 1967) indicates that flat-lying Quaternary alluvium unconformably overlies a “basement” of Miocene and older rocks beneath this part of the Bay and surrounding alluvial plains. Moreover, earlier accounts of the Neogene history of San Francisco Bay (Louderback, 1951; Howard, 1951) concluded that the Pliocene Merced embayment was a narrow coastal feature that did not occupy the present southern part of San Francisco Bay. If the northern Santa Clara County Merced(?) exposures were not directly connected to the Merced Formation to the northwest, they must have formed the nearshore part of a westward-oriented embayment that was subsequently truncated by the San Andreas fault. The Merced(?) and Santa Clara Formations of the Palo Alto area are located less than 3 miles northeast of a 1600-foot section of Pliocene Purisima Formation (Cummings, 1960) situated less than one mile southwest of the San Andreas fault. These exposures have been cor- related with the lower Pliocene Tahana Member of the Purisima Formation by Cummings and others (1962). The fauna consists of 37 mollusks (table 4) including taxa such as Colus recurvus, Clementia, Dosinia, and Spisula moss- Vor. XXXVII] ADDICOTT: LATE PLIOCENE MOLLUSKS FROM CALIFORNIA 81 \ 3S eee . \ ‘Type Merced | “Formation T ©1 sonoma co."MERCED" SOURCES OF DATA W > |. Dickerson, 1922 rams ") N 2.Martin, 1916 = 3. Lawson, 1893, 1895 2 4. Ashley, 1895 So 5. Arnold, 1906 “ ~ R MUSSEL ROCK \ > 9 ae > LINA PEs S = ) 6.Glen, 1959 oS \ AAPA 7. Haehl and Arnold, = 1904 D TYPE MERCED Q 8.Cummings and others, Ss a) 1962 B 9. Arnold, 1908 => co) 10. This report Wye = ‘© PURISIM|A FM 4 A Sc SANTAXCRUZ AREA Yy INDEX MAP 37°30- HALFMOON BAY 10 MILES EXPLANATION Morine strata of late Pliocene age Marine strata of early Pliocene age Postulated limits of late Pliocene marine deposition ed Ficure 2. Distribution of marine Pliocene rocks and late Pliocene paleogeography in the northern Santa Cruz Mountains (geology modified from Jennings and Burnett, 1961). 122015! beachensis (Ashley, 1895; Cummings, 1960; Cummings and others, 1962) that are suggestive of an early Pliocene age. None of these taxa occur in the nearby exposures of Pliocene strata northeast of the fault.‘ The inferred non- contemporaneity between these assemblages on opposite sides of the San Andreas 7 The warm temperate to tropical genus Dosinia is suggestive of an early Pliocene or older age at this latitude. Its northernmost late Pliocene and Pleistocene occurrences are in the warmer water faunas of southern California. [Proc. 4TH Serr. CALIFORNIA ACADEMY OF SCIENCES , 82 xptmosdmoy ‘(es6gt ‘Aatysy) 2eqqey SNSOqgqi3s snmopixng | MK Ge fa, jee ee, a lo a a (tajuedieDy) ounydnoipqns vyauisayoy OG er ee ae ie en ee te oe aa (AeIn) psosapuog piuisog ‘dsg DR PR ees ee en Peluo) vI2UAs0f{yv2 DKmWoId KAD >: Ge Linnie eee ae ie ne er «d@PPIUOD DUNIOI DIaIDSSDAD,, [(Bg6st ‘Aalysy) qqey wnunryaam wnrpsn | CS eats ee ai oor ro ae ~~ (qqey)) mnuniyaaum wnipaprouy) Om ke cane a rca a a a a saeds = p2jyuamary¢ [(8S68t ‘Aglysy) peruo0g Djuoposom DIAF | ‘ds¢ ds¢ yp axe x De Ge RE ie i ae aly Cane ham ee a (PRIU0D) DynaUIL] DADpDUP Xn A UMD cr Sar eames pay (SPUIF{) S7suatjsDI (DpLIDIUNAT) DIP sdodA0dlag [(e8s6st ‘Agysy) petuog na2us0fyv2 DSSDN ¢ *(O096T ‘SsutuUIND) (UnIePL) snuniuds0M SNUADSSD AJ | xX XK: > rae RS ae a a ae ae (12d) snqowmwuns3s snispssp Ky 5% x ix Xn PMR 0 bo cee eiepamtaaic a Sg eer (peiuoy) snupimsofyns snispssp ny [(es6st ‘Aalysy) pinoy wszmay vuynunT] ‘ds EN ee aie hs ARES a eg ae (p[noy)) zs2ma7 sao2uz0g EI EE a sateds nya4qzpy [(es6st “AaTysy) JJ1opuappyAl szpuns3 ynpidasy] x D2) ie. > pau ena ce we Vg ae See peo) sd¢aaumud vjnpidasa [(esost ‘Aatysy) qqey vasnoas vnaunjdayy] NGG ac aoa aan Ey es gree i ag eee (qqey) snasngas smog ‘ds Ss RS Sara ia pie ie rier a ea diapoig supypmoum vansdxjog [(8061) ploury sesuanjozsog (snsnfowoong) snsny] ‘ds XG Me mee MRR eh ce peng ar ee os (plou1y) sesuanjozsog snapyyunoyp) Ti et ean aa a. A a. ee = a 5 5 S S S 5 3 . z SdOdOULSV‘) o on = o a sa EF = ‘< 5 SoTPTTBIOT SOSN a a eee ee = ee ‘SJ9YIDAG UL UMOYS A4D (OGL) SSurmungy pup /(806I) pjoucy <(voogr) Kajysp Kq pasn sampu quasaffiq “DIUAO {YD “KaYVA 010]40g WoOdf (Z96T ‘s4ayzO PUD SSUIMUND) UOIDULAOY DUISLNgG ay, {0 saquayy DUDUDT 941 WOAf SYSN}OPY ‘pb ATAVI, 83 Vov. XXXVIT] ADDICOTT: LATE PLIOCENE MOLLUSKS FROM CALIFORNIA ~ ‘ds ‘ds ae) ‘ds ‘ds a) 19 i a A ~ mM Mm OM a nm a Pa ate ee ee os oe bn gre oe a saieds nzpj0 x 72 Daeg, Nach eae ae aa Al Aa a ara an Vr aa (41910}S) Sszumsofaniap4y] DIpjoO x ti Mine e ee) pad cma van] es ae qqer) 24a¢009 pipj0 x _ [Cesest PEND peiuod) szsuaorvlng (snuaA Ou) vAéjoDp] Sue arg cee ae Ye eee ee me) ae (peiu0d)) snunosnlnd snsaa TZ, [(es681 eeolusy) PPAUOD YPMIINU SNAADY 10214 IC | > egy € Se eS Cet ame ae 2 eg gre (peIuo)) yoyINnU snsaaT ea he ELL RD) SEO SOUT. THUSRES mien maar aRCoae san sens enme nn (DENUIOS)) NOC. DO IDEN ITISZS [(8S6st ‘AeTysy) ¢peruoyg nynjnunpd vpapunqs | PS ae TP Soc oo eee (PInoy) vzn27nf vnsids _-LCes68T ‘AIYSY) PeAUOD vI2UALO{YYI DYapunys | Peg ae sae = aes eT —_—————— (priu0D)) suudofajnns ninsigs Set a ee CE ae ee p[noy sn21p91s Uajog I (CAGE NOLS jo Cluejoowtsy)) fyi PC! ia) jMOApe! Ea ay ee Te Fa (peluoD)) wnipsuap wnipspropnasg [ (0961 ‘ssurmuIng)) (t9}U0dI1eD) vynjNSUNIQ syaUc)og YF | = ais Sie aes rs a ee (dajuedie)) pynjnsunrg srjauisozy [(ese6st ‘Aa[ysy) toyusdaeg vu2t4auaz sadn f | SRS ae eas a eee ne ee (dajuadie)) Dulrtsauaz DIDYI0JOAG [ (es6st ‘Aalysy) peru0g vauzunys sadn] '(O96T ‘SssuIWIUIND) (peIUOD) DaUu2IUDIS DIDYI010Ag | seen cence cna nnn na nnn een nn nnn enna (qqe)) 2hajpjs DIDYI0I0Ag [ (eS68T ‘Aa[ysy) P[Nory vsosauas SisaUtKIK]4 | SS eg Ce Ee a EL ae pa ee (peiu0g) nidnaqn agdouvg Sa aa aaa gag eae Jajuodiey) vsojzf pAopung ge ga ane ae Ge Oe peiuo0g vj7pj9Uung vAéopung ah eee Sa ee eee (peruodD) snqIa4 “Py “JO snjorpo pW Sa.) a oe a Che es (peIU0D) DINSDU DULOID LY [(esost ‘Aapysy) PeLuoD yD}4NU DULINT | Seah y as aera was OSS RRR IS EP a rages Ua (peluoy) wjpIjInNU DMISWMINT [ (0961 ‘ssuruUIND)) (1ojuadIey) Dunydnrpqns 84 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. MILES 300 250 Cor Le ve O Va Yi 4 200 (of 7 a” / ¢ / 150 / i / / i 100 / if O / Lateral slip postulated 7s) by others / Vi®) 50 © e Yi This report O| 10 20 30 40 50 M.Y. QUA- PLIOCENE MIOCENE OLIGOCENE EOCENE TERNARY Ficure 3. Comparison of indicated lateral offset of marine late Pliocene strata on the San Francisco Peninsula with Tertiary offsets postulated by other workers (modified from Grantz and Dickinson, 1968). fault is supported by the fact that there is a much smaller number of species in common, 14 out of 33, than might normally be expected, since both represent comparable very shallow water inner sublittoral environments and are so close together. The upper Pliocene members of the Purisima Formation of Cummings and others (1962) of the western slope of the Santa Cruz Mountains seem to define an eastward oriented depositional basin of very shallow water aspect (upper Vor. XXXVII] ADDICOTT: LATE PLIOCENE MOLLUSKS FROM CALIFORNIA 85 inner sublittoral zone) that did not reach nearly as far inland as the much thicker, somewhat deeper water lower Pliocene member. The nearest exposures of Pliocene strata along the San Andreas fault occur along the coast between the fault and Mussel Rock (Glen, 1959), about 20 to 25 miles to the northwest. Although Higgins (1961) believes that these exposures have been displaced by landsliding across the San Andreas fault to their present position on the south- west side, it is noteworthy that they contain species such as Anadara trilineata (USGS loc. M1618) and Tresus pajaroanus that have not been reported from localities in the lower part of the type Merced Formation (Glen, 1959) on the northeast side of the fault. Most of the mollusks from exposures of Merced Formation on the southwest side of the San Andreas fault near Mussel Rock also are recorded from the Merced(?) Formation of northern Santa Clara County. Perhaps the most significant of these is A. trilineata which, interest- ingly, is not recorded from strata in the type Merced Formation northeast of the San Andreas fault (Glen, 1959). The indicated post-depositional right-lateral slip on the order of 20 to 25 miles is compatible with postulated rates of lateral slip along the San Andreas fault during earlier parts of the Neogene (Hill and Dibblee, 1953; Dibblee, 1966b; Galehouse, 1967; Addicott, 1968). It is also in close agreement with Cummings’ (1968) estimate of about 18 miles of dis- placement of distinctive nonmarine facies in the Pliocene and Pleistocene Santa Clara Formation in this area. The relationship of the apparently offset late Pliocene marine strata on the San Francisco Peninsula to earlier postulates of Tertiary slip along the San Andreas fault is shown on a recent graph (fig. 3) compiled by Grantz and Dickinson (1968). LOCALITIES United States Geological Survey Cenozoic localities (USGS) (Menlo Park Register ) M1618. In large cut on east side of new location of Highway 101 where it crosses a ridge trending northwestward through the center of the NE 7 sec. 23, T. 3 S., R. 6 W., San Francisco South 7%’ quadrangle, California. Altitude 525 feet Merced Formation. Collectors: M. Bonilla and W. O. Addicott, 1962. M1661. In large excavated area 950 feet S. 61° W. of intersection of Junipero Serra Boulevard and Arroyo Drive. Map coordinates: 1, 434,850 feet E., 423,600 feet N., California coordinate system, zone 3. San Fran- cisco South quadrangle, California. Merced Formation. Collectors: M. Bonilla and G. O. Gates, 1962; W. O. Addicott, 1963. M1715. Cut and pipeline trench on north side of Arastradero Road, 1200 feet west of intersection with Page Mill Road, Palo Alto 7'2’ quadrangle, California. Merced(?) Formation, late Pliocene. Collectors: E. H. Pampeyan, J. G. Vedder, and W. O. Addicott, 1963. 86 M1720. M1870. M1926. M2664. M2665 M3253. M3254. Mi3295: M3256. M3824. Stanford C-308. C-321 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Cut on south side of highest building site for American Institute for Research buildings on southwest side of Arastradero Road, 2200 feet due east of intersection with Alpine Road, Palo Alto 7%’ quadrangle, California. Collectors: M. D. Crittenden and W. O. Addicott, 1962. Cut on northeast side of Arastradero Road, 2450 feet N. 78° E. of intersection of Alpine and Arastradero Roads, Palo Alto 7'2’ quad- rangle. Merced(?) Formation. Collector: W. O. Addicott, 1963. About 2700 feet southeast of intersection of Arastradero and Alpine roads, 400 feet north of lat. 37°22’33” N., 6000 feet west of long. 122°10’ W., altitude 650 feet, Palo Alto 7%’ quadrangle, California. Collector: E. H. Pampeyan, 1963. Roadcut 450 feet south 2860 feet east of northwest corner of Mindego Hill 774’ quadrangle, California. Purisima Formation. Collectors: E. H. Pampeyan and W. O. Addicott, 1966. In Neils Gulch, 3800 feet east of southwest corner of Mindego Hill 72’ quadrangle, California. Purisima Formation. Collectors: E. H. Pampeyan and W. O. Addicott, 1966. In Hamms Gulch on unsectioned land, 1650 feet north, 4,000 feet east of SE. cor. sec. 36, T. 6 S., R. 4 W., Mindego Hill 7’2’quadrangle, California. Purisima Formation. Collector: J.C. Cummings, 1967. In Damiani Creek on unsectioned land, 2125 feet south, 6200 feet east of NE. cor. sec. 1, T. 7 S., R. 4 W., Mindego Hill 772’ quadrangle, California. Purisima Formation. Collector: J. C. Cummings, 1967. South side of Rapley Trail Road on unsectioned land, 600 feet north, 8400 feet east of SW. cor. sec. 6, T. 7 S., R. 3 W., Mindego Hill 77%’ quadrangle. Purisima Formation. Collector: J. C. Cummings, 1967. In Jones Gulch on unsectioned land 250 feet north, 3850 feet east of SE. cor. sec. 36, T. 6 S., R. 4 W., Mindego Hill 7727 quadranates Purisima Formation. Collector: J. C. Cummings, 1967. North side of E.-trending gully, 4600 feet north, 3700 feet west of SE. cor. proj. sec. 36, T. 6 S., R. 3 W., Mindego Hill 774’ quadrangle. Collector: Mrs. R. A. Loney, 1968. University (SU) localities About 1% miles southeast of the forks of Page Mill Road (southern intersection of Page Mill and Arastradero roads). Also listed as 2 miles up the east fork of Madera [Matadero] Creek between the ‘“‘e” and “p” of “Concepcion,” Santa Cruz 30’ quadrangle, California (= Arnold, 1908, loc. 8). Santa Clara(?) Formation. Collectors: R. Arnold and T. J. Hoover. East bank of Los Trancos Creek, 2 miles above junction of San Fran- cisquito Creek immediately southwest of Felt Lake, Santa Cruz 30’ VoL. XXXVII] ADDICOTT: LATE PLIOCENE MOLLUSKS FROM CALIFORNIA _ 87 quadrangle, California. Merced(?) Formation. (= Arnold, 1908, loc. 21). Collectors: R. Arnold and T. J. Hoover. 153. In cut to Felt Lake [ditch bank near mouth of Felt Lake, Palo Alto 74’ quadrangle]. Collectors: E. K. Jordan and L. G. Hertlein. 2675. Incut on north side of road between Los Trancos and Matadero Creeks about % mile east of Los Trancos Creek and '2 mile south of Felt Lake, San Mateo County, California. Merced(?) Formation. Collectors: R. Compton and R. G. Thomas, 1947. Probably same as locality 154 “along road back of Wonder.” Collectors: E. K. Jordan and L. G. Hertlein. REFERENCES AzgzorT, R. T. 1954. American seashells. Van Nostrand Company, Inc. New York, 541 p., 40 pls. Appicott, W. O. 1965a. On the identification of Schizopyga californiana Conrad, a California Pliocene gastropod. Proceedings of the California Academy of Sciences, 4th series, vol. 33, no. 2, pp. 47-58, 11 figs. 1965b. Some western American gastropods of the genus Nassarius. United States Geological Survey Professional Paper 503-B, 24 pp., 3 pls. 1965c. Miocene macrofossils of the southeastern San Joaquin Valley, California. United States Geological Survey Professional Paper 525-C, pp. C101-C109, 4 figs. 1968. Mid-Tertiary zoogeographic and paleogeographic discontinuities across the San Andreas fault, California, in Proceedings of conference on geologic problems of San Andreas Fault System, edited by W. R. Dickinson and Arthur Grantz. Stanford University Publications, Geological Sciences, vol. 11, pp. 144-165. ARNOLD, RALPH 1906. The Tertiary and Quaternary pectens of California. United States Geological Survey Professional Paper 47, 264 pp., 53 pls. 1907. New and characteristic species of fossil mollusks from the oil-bearing Tertiary formations of Santa Barbara County, California. Smithsonian Miscellaneous Collections, vol. 50, pt. 4, no. 1781, pp. 419-447, pls. 50-58. 1908. Descriptions of new Cretaceous and Tertiary fossils from the Santa Cruz Moun- tains, California. Proceedings of the United States National Museum, vol. 34, no. 1617, pp. 345-390, pls. 31-37. 1909. Paleontology of the Coalinga district, Fresno and Kings Counties, California. United States Geological Survey Bulletin 396, 173 pp., 30 pls. [1910]. AsHtey, G. H. 1895a. The Neocene stratigraphy of the Santa Cruz Mountains, [part] 1, Stratigraphy. California Academy of Sciences, second series, vol. 5, pp. 273-376 [1896]. 1895b. Studies in the Neocene of California. Journal of Geology, vol. 3, no. 4, pp. 434-454. BartscH, PAuL 1911. The Recent and fossil mollusks of the genus Bittiwm from the west coast of America. Proceedings of the United States National Museum, vol. 40, no. 1826, pp. 383-414, pls. 51-58. Brann_er, J. C., J. F. Newsom, ANd RALPH ARNOLD 1909. Description of the Santa Cruz quadrangle, California. United States Geological Survey Geologic Atlas, Folio 163, 11 pp. 3 maps. 88 ; CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Birp, S. O. 1967. The binomial distribution applied to the percentage method of stratigraphic correlation. Geological Society of America Bulletin, vol. 78, no. 12, pp. 1507-1514. Burcu, J. Q., editor 1944— 1946. Distributional list of the west American marine mollusks from San Diego, Cali- fornia to the Polar Sea. Conchology Club of Southern California Minutes, nos. 33-63. Conrap, T. A. 1856. Description of three new genera; twenty-three new species middle Tertiary fossils from California, and one from Texas. Proceedings of the Philadelphia Academy of Natural Sciences, vol. 8, pp. 312-316; reprinted im Dall, W. H., 1909, United States Geological Survey Professional Paper 59, appendix 8, pp. 173-175. 1857. Descriptions of the fossil shells [Williamson’s survey in California and Oregon]. United States Pacific Railroad Exploration Reports (United States 33d Congress, 2d session, Senate Executive Document 78 and House Executive Document 91), vol. 5, pt. 2, appendix, article 2, pp. 317-329, pls. 2-9. Corey, W. H. 1954. Tertiary basins of southern California [part] 8 im chapter 3 of Jahns, R. H., editor, Geology of southern California. California Division of Mines Bulletin 170, pp. 73-83. Cummincs, J. C. 1960. Geology of the Langley Hill-Waterman Gap area, Santa Cruz Mountains, Cali- fornia. Stanford, California, Stanford University, Doctor of Philosophy thesis, ZSSNps 1968. Santa Clara Formation and possible post-Pliocene slip on the San Andreas fault in central California, in Proceedings of conference on geologic problems of San Andreas Fault System, edited by W. R. Dickinson and Arthur Grantz. Stanford University Publications, Geological Sciences, vol. 11, pp. 191-207. Cummincs, J. C., R. M. Tourinc, anp E. E. Brass 1962. Geology of the northern Santa Cruz Mountains, California, 77 Bowen, O. E., Jr., editor, Geologic guide to the gas and oil fields of northern California. California Division of Mines Bulletin 181, pp. 179-220. Dat, W. H. 1909. Contributions to the Tertiary paleontology of the Pacific Coast, I. The Miocene of Astoria and Coos Bay, Oregon. United States Geological Survey Profes- sional Paper 59, 278 pp., 23 pls., 14 figs. 1917. Summary of the mollusks of the family Alectrionidae of the west coast of America. Proceedings of the United States National Museum, vol. 51, no. 2166, pp. 575-579. 1919. Descriptions of new species of Mollusca from the North Pacific Ocean in the collection of the United States National Museum. Proceedings of the United States National Museum, vol. 56, no. 2295, pp. 293-371. Davis, F. F., anp C. W. JENNINGS 1954. Mines and mineral resources of Santa Clara County, California. California Journal of Mines and Geology, vol. 50, no. 2, pp. 320-430. Vot.XXXVIT] ADDICOTT: LATE PLIOCENE MOLLUSKS FROM CALIFORNIA 89 DipsLeE, T. W., Jr. 1966a. Geology of the Palo Alto quadrangle, Santa Clara and San Mateo Counties, California. California Division of Mines Map Sheet 8. 1966b. Evidence for cumulative offset on the San Andreas fault in central and northern California, zn Bailey, E. H., editor, Geology of northern California. California Division of Mines Bulletin 190, pp. 375-384. Dickerson, R. E. 1922. Tertiary and Quarternary history of the Petaluma, Point Reyes, and Santa Rosa quadrangles [California]. Proceedings of the California Academy of Sciences, fourth series, vol. 11, no. 19, pp. 527-601, pls. 17-41. DuruaMm, D. L., anp W. O. Appicotr 1965. Pancho Rico Formation, Salinas Valley, California, in Geological Survey research 1965. United States Geological Survey Professional Paper 524-A, pp. Al-A22, 7 figs., 5 pls., table. DurHaAM, J. W. 1950. Cenozoic marine climates of the Pacific Coast. Geological Society of America Bulletin, vol. 61, no. 11, pp. 1243-1264. Duruam, J. W., AnD F. S. MacNein 1967. Cenozoic migrations of marine invertebrates through the Bering Strait region, in Hopkins, D. M., editor, The Bering Land Bridge. Stanford, California, Stanford University Press, pp. 326-349. Emerson, W. K. 1956. Upwelling and associated marine life along Pacific Baja California, Mexico. Journal of Paleontology, vol. 30, pp. 393-397, 1 fig. ETHERINGTON, T. J. 1931. Stratigraphy and fauna of the Astoria Miocene of southwest Washington. University of California Publications, Department of Geological Sciences Bulletin, vol. 20, no. 5, pp. 31-142, 14 pls., 1 map. FaustMan, W. R. 1964. Paleontology of the Wildcat Group at Scottia and Centerville Beach, California. University of California Publications, Department of Geological Sciences Bulletin, vol. 41, no. 2, pp. 97-160, 3 pls. Fintayson, D. J.,. W. R. Hansen, R. S. Forp, R. G. Scott, anp J. V. VANTINE 1967. Evaluation of ground water resources, South Bay [California]. California Department of Water Resources Bulletin 118-1, 153 p. Bier, J. E. 1953. Common marine bivalves of California. California Department of Fish and Game Fish Bulletin, no. 90, 102 pp., 63 figs. Gass, W. M. 1866. Tertiary invertebrate fossils. California Geological Survey, Paleontology, vol. 2, section 1, pt. 1, pp. 1-38, pls. 1-13. GALEHOUSE, J. S. 1967. Provenance and paleocurrents of the Paso Robles Formation, California. Geo- logical Society of America Bulletin, vol. 78, no. 8, pp. 951-978. GLEN, WILLIAM 1959. Pliocene and lower Pleistocene of the western part of the San Francisco Penin- sula. University of California Publications, Department of Geological Sciences Bulletin, vol. 36, no. 2, pp. 147-198, pls. 15-17. 90 ‘ CALIFORNIA ACADEMY OF SCIENCES [| Proc. 4TH SER. Grant, U. S., IV, anp H. R. GALE 1931. Catalogue of the marine Pliocene and Pleistocene Mollusca of California. San Diego Society of Natural History Memoirs, vol. 1, 1036 pp., 32 pls., 15 figs. GrRANTZ, ARTHUR, AND W. R. DicKINSON 1968. Indicated cumulative offsets along the San Andreas fault in the California Coast Ranges, in Proceedings of conference on geologic problems of San Andreas Fault System, edited by W. R. Dickinson and Arthur Grantz. Stanford University Publications, Geological Sciences, vol. 11, pp. 117-119. Hase, TADASHIGE 1964. Shells of the western Pacific in color, vol. II. Osaka, Japan, Hoikusha, 233 pp. 66 pls. Haeut, H. L., anp RatpH ARNOLD 1904. The Miocene diabase of the Santa Cruz Mountains in San Mateo County, Cali- fornia. Proceedings of the American Philosophical Society, vol. 43, pp. 16-53. HepcPetTH, J. W. 1957. Classification of marine environments, in Treatise on marine ecology and paleo- ecology. Geological Society of America Memoir 67, vol. 1, pp. 17-27. Herren, L. G., ann E. C. ALiison 1959. Pliocene marine deposits in northwestern Baja California, Mexico, with the description of a new species of Acanthina (Gastropoda). Southern California Academy of Sciences Bulletin, vol. 58, pt. 1, pp. 17-26, 8 pls. HErTLEIN, L. G., anp U. S. Grant, IV 1944. The Cenozoic Brachiopoda of western North America. University of California at Los Angeles, Publications in Mathematical and Physical Sciences, vol. 3, 236 pp., 21 pls., 34 figs. Hicews, C. G. 1960. Ohlson Ranch Formation, Pliocene, northwestern Sonoma County, California. University of California Publications, Department of Geological Sciences Bulletin, vol. 36, no. 3, pp. 199-232, pls. 18-20, 6 figs., 1 map. 1961. San Andreas fault north of San Francisco, California. Geological Society of America Bulletin, vol. 72, no. 1, pp. 51-68. Hint, M. L., anp T. W. Disster, Jr. 1953. San Andreas, Garlock, and Big Pine faults, California—a study of the character, history, and tectonic significance of their displacements. Geological Society of America Bulletin, vol. 64, no. 4, pp. 443-458. Howarp, A. D. 1951. Development of the landscape of the San Francisco Bay counties. California Division of Mines Bulletin 154, pp. 95-106. Howe, H. V. 1922. Faunal and stratigraphic relationships of the Empire Formation, Coos Bay, Oregon. University of California Publications, Department of Geological Sciences Bulletin, vol. 14, no. 3, pp. 85-114, pls. 7-12. JENNINGS, C. W., AND J. L. BuRNETT 1961. Geologic map of California, Olaf P. Jenkins edition, San Francisco Sheet. Cali- fornia Division of Mines, scale 1 : 250,000. Keen, A. Myra 1939. The percentage method of stratigraphic dating. Proceedings of the Sixth Pacific Science Congress, vol. 2, pp. 659-663. 1958. Sea shells of tropical west America; marine mollusks from Lower California to Columbia. Stanford, California, Stanford University Press, 624 pp., 10 pls. VoLt. XXXVII] ADDICOTT: LATE PLIOCENE MOLLUSKS FROM CALIFORNIA 91 1963. Marine molluscan genera of western North America. Stanford, California, Stan- ford University Press, 126 pp., illustrations. Kincar, TREVOR 1957. Local races and clines in the marine gastropod Thais lamellosa Gmelin; a popu- lation study. Seattle, Washington, Calliostoma Company, 75 pp., 65 pls. Lawson, A. C. 1893. The post-Pliocene diastrophism of the coast of southern California. University of California, Department of Geology Bulletin, vol. 1, no. 4, pp. 115-160, pls. 8-9. 1895. Sketch of the geology of the San Francisco Peninsula [California]. United States Geological Survey Annual Report 15, pp. 399-476. 1914. Description of the San Francisco district (Tamalpais, San Francisco, Concord, San Mateo, and Hayward quadrangles). United States Geological Survey Geologic Atlas, Folio 193, 25 pp., 15 maps, 10 pls. LoupErBACK, G. D. 1951. Geologic history of San Francisco Bay. California Division of Mines Bulletin 154, pp. 75-94. Martin, BRUCE 1914. Descriptions of new species of fossil Mollusca from the later marine Neocene of California. University of California Publications, Department of Geology Bulletin, vol. 8, no. 7, pp. 181-202, pls. 19-22. 1916. The Pliocene of middle and northern California. University of California Publications, Department of Geology Bulletin, vol. 9, no. 15, pp. 215-259, tables 1-3. Moors, E. J. 1963. Miocene mollusks from the Astoria Formation in Oregon: United States Geo- logical Survey Professional Paper 419, 109 p., 33 pls. Moorr, R. C. (chairman, Stratigraphic Committee) 1949. The Pliocene-Pleistocene boundary. American Association of Petroleum Geolo- gists Bulletin, vol. 33, no. 7, pp. 1276-1280. Neweerry, J. S. 1856. Report upon the geology of the route [Williamson’s survey in California and Oregon]. United States Pacific Railroad Exploration Reports (United States 33d Congress, 2d session, Senate Executive Document 78 and House Execu- tive Document 91), vol. 6, pt. 2, pp. 9-68. NomIanp, J. O. 1917. The Etchegoin Pliocene of middle California. University of California Publica- tions, Department of Geology Bulletin, vol. 10, no. 14, pp. 191-254, pls. 6-12, 2 figs. Otproyp, Ina S. 1924. Marine shells of Puget Sound and vicinity. University of Washington, Puget Sound Biology Station Publications, vol. 4, 272 pp., 49 pls. Osmont, V. C. 1905. A geological section of the Coast Ranges north of the Bay of San Francisco. University of California Publications, Department of Geology Bulletin, vol. 4, no. 3, pp. 39-87, pls. 6—7. Packarp, E. L. 1916. Mesozoic and Cenozoic Mactrinae of the Pacific coast of North America. Uni- versity of California Publications, Department of Geology Bulletin, vol. 9, no. 16, pp. 261-360, pls. 12-35. 92 ‘ CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. 1918. Molluscan fauna from San Francisco Bay. University of California Publications in Zoology, vol. 14, no. 2, pp. 199-452, pls. 14-60. Pace, B. M., anp L. L. Tasor 1967. Chaotic structures and décollement in Cenozoic rocks near Stanford University, California. Geological Society of America Bulletin, vol. 78, no. 1, pp. 1-12. PALMER, K. V. 1958. Type specimens of marine mollusca described by P. P. Carpenter from the west coast (San Diego to British Columbia). Geological Society of America Memoir 76, pp. 1-376, pls. 1-35. Parker, R. H. 1964. Zoogeography and ecology of macro-invertebrates, particularly mollusks, in the Gulf of California and the continental slope off Mexico. Videnskabelige Meddelelser fra Dansk naturhistorisk foreningen, Copenhagen, Bd. 126, 178 p., 15 pls. Pre Jj. Ee yr 1960. Paleontology and correlation of the Ohlson Ranch Formation. University of California Publications, Department of Geological Sciences Bulletin, vol. 36, no. 4, pp. 233-242, pl. 21. Quavte, D. B. 1960. The intertidal bivalves of British Columbia. Victoria, British Columbia Provin- cial Museum Handbook 17, 104 pp., illus. REPENNING, C. A. 1967. Palearctic-Nearctic mammalian dispersal in the late Cenozoic, in Hopkins, D. M., editor, The Bering Land Bridge. Stanford, California, Stanford University Press, pp. 288-311. SANTILLAN, M., AND T. BARRERA 1930. Las Posibilidades petroliferas en la costa occidental de la Baja California, entre los Paralelos 30° y 32° de latitud norte. Instituto Geologico de Mexico Anales, vol. 5, pp. 1-37, 6 pls., 12 figs. SmitH, A. G., anpD MacKenzie Gorpon, Jr. 1948. The marine mollusks and brachiopods of Monterey Bay, California, and vicinity. Proceedings of the California Academy of Sciences, fourth series, vol. 26, no. 8, pp. 147-245, pls. 3, 4, 4 text figs. SmitTH, J. P. 1919. Climatic relations of the Tertiary and Quaternary faunas of the California region. Proceedings of the California Academy of Sciences, fourth series, vol. 9, no. 4, pp. 123-173, pl. 9. Soper, E. K., ann U. S. Grant, IV 1932. Geology and paleontology of a portion of Los Angeles, California. Geological Society of America Bulletin, vol. 43, no. 12, pp. 1041-1068, 7 figs. STEWarT, R. B. 1927. Gabb’s California fossil type gastropods. Proceedings of the Philadelphia Academy of Natural Sciences, vol. 78, pp. 287-447, pls. 20-32. 1930. Gabb’s California Cretaceous and Tertiary type lamellibranchs. Philadelphia Academy of Natural Sciences, Special Publication no. 3, 314 p., 17 pls., 5 text figs. VALENTINE, J. W. 1955. Upwelling and thermally anomalous Pacific Coast (California-Mexico) Pleis- tocene molluscan faunas. American Journal of Science, vol. 253, no. 8, pp. 462— 474, 3 figs. Vor. XXXVII] ADDICOTT: LATE PLIOCENE MOLLUSKS FROM CALIFORNIA 93 1961. Paleoecologic molluscan geography of the Californian Pleistocene. University of California Publications, Department of Geological Sciences Bulletin, vol. 34, no. 7, pp. 309-442. VEDDER, J. G. 1960. Previously unreported Pliocene Mollusca from the southeastern Los Angeles basin [California], article 151 im Short papers in the geological sciences. United States Geological Survey Professional Paper 400-B, pp. B326-B328. WatTeERFALL, L. N. 1929. A contribution to the paleontology of the Fernando Group, Ventura County, California. University of California Publications, Department of Geological Sciences Bulletin, vol. 18, no. 3, pp. 71-92, pls. 5-6. Weaver, C. E., chairman 1944. Correlation of the marine Cenozoic formations of western North America. Geo- logical Society of America Bulletin, vol. 55, no. 5, pp. 569-598, 1 pl. WinTERER, E. L., ann D. L. DurHAM 1962. Geology of southeastern Ventura Basin, Los Angeles County, California. United States Geological Survey Professional Paper 334-H, pp. 275-366, pls. 44-49. Wooprinc, W. P. 1952. Pliocene-Pleistocene boundary in California Coast Ranges. American Journal of Science, vol. 250, no. 6, pp. 401-410. Wooprinc, W. P., anp M. N. BRAMLETTE 1950. Geology and paleontology of the Santa Maria district, California. United States Geological Survey Professional Paper 222, 185 pp., 23 pls., 9 figs. [1951]. Wooprinc, W. P., M. N. BRAMLETTE, AND W. S. W. Kew 1946. Geology and paleontology of the Palos Verdes Hills, California. United States Geological Survey Professional Paper 207, 145 pp., 37 pls., 16 figs. Wooprinc, W. P., RatpH Stewart, AND R. W. RICHARDS 1940. Geology of the Kettleman Hills oil field, California. United States Geological Survey Professional Paper 195, 170 pp., 57 pls., 15 figs. [1941]. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XXXVII, No. 4, pp. 95-166; 19 figs.; 15 tables. December 10, 1969 THE BIOLOGY OF STORM PETRELS IN THE GALAPAGOS ISLANDS* By M. P. Harris Edward Grey Institute of Field Ornithology, Oxford University, Oxford, England Asstract : Between November, 1965 and July, 1967, studies were made on the biology of Oceanodroma castro and O. tethys in the Galapagos Islands. Oceanodroma castro is a widespread species in subtropical parts of the Pacific and Atlantic oceans. Although there is considerable size variation between breeding popula- tions, the species is best treated binomially. Four specimens collected in the Gulf of Guinea may however belong to a distinct race. In the Galapagos Islands O. castro has two breeding seasons a year but there ap- peared to be two entirely separate populations, each breeding annually, one in the hot season, one in the cold. Even nonbreeders and failed breeders remained faithful to their annual cycle in the four seasons studied. A detailed study of the breeding biology showed marked differences in the seasons but little correlation with breeding in the hot or cold seasons. There was a decline in nesting success with date of laying within each season. The cause of this was obscure but it did not appear to be food shortage or predation of the adults by Asio galapagoensis. Wing molt occurred away from the breeding grounds and took most of the time between breeding seasons. This need not indicate that birds were prevented from breeding more rapidly as other factors might also be important. The factors preventing nesting throughout the year are obscure, but presumably there was some advantage to a pair in breeding when the majority of individuals did so, which more than compensated for losses due to competition for food and nest sites. Predation is probably important in the synchronization of breeding. Oceanodroma tethys is a diurnal species when visiting land. The colony studied on Tower Island was occupied by approximately 200,000 breeding pairs. Large numbers of birds frequented the colony throughout the year but all but a few eggs were laid from May to July. The vast numbers of birds flighting at the colony out of the breed- ing season were apparently nonbreeders as adults would have been away replacing their wing feathers. No apparent reason for the flighting was found. In another colony * Contribution of the Charles Darwin Foundation for the Galapagos. [95] 96 , CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. at Isla Pitt, there were no birds present during the nonbreeding season. The nesting colonies were extremely crowded and the low nesting success (less than 23 percent) was mainly due to intense intraspecific competition for nest sites. The third species of storm petrel was Oceanites gracilis. This is ecologically distinct from the Oceanodroma species as it is an inshore feeder. There was probably little competition between O. castro and O. tethys as the former feeds on fish and cephalo- pods by day, the latter on smaller fish, cephalopods, and crustacea caught by night. Their feeding areas may also be different. INTRODUCTION Three species of storm petrels are seen in the waters around the Galapagos Islands, the Madeiran storm petrel (Oceanodroma castro), the Galapagos storm petrel (Oceanodroma tethys), and Elliot’s storm petrel (Oceanites gracilis). The last species has not been found breeding but the results of gonad examination (Loomis, 1918), and the fact that it appears to be resident and an endemic race, show that it must breed in the archipelago. Ecologically however it is quite distinct. My wife and I were resident at the Charles Darwin Research Station in the Galapagos from November, 1965, to July, 1967, and for the majority of the time we were engaged on studies of the two Oceanodroma species. Observations were also made on the other species of sea birds present in the archipelago but these will be published separately except as far as they concern the storm petrels. There has always been confusion regarding the names of the various islands in the archipelago (Slevin, 1959) and I have compromised and used the names now in commonest usage. However for the sake of completion I give below the alternate names for those which have both Spanish and English names but are not direct translations. The alternate names are given in brackets and the official Ecuadorian title in italics. The islands are Santa Cruz |Indefatigable|; /sabela [Albemarle]; San Cristébal [Chatham]; Espanola [Hood]; Genovesa | Tower]. OCEANODROMA CASTRO Oceanodroma castro is a widespread species in the subtropical parts of both the Atlantic (summary in Bannerman, 1941) and Pacific oceans. It is known to breed in Hidejima and possibly Sanganjima off the eastern coast of Japan (Austin and Kuroda, 1953), Kauai in the Hawaiian Archipelago (Richardson, 1957), several islands in the Galapagos Archipelago (Loomis, 1918; Lévéque, 1964), the Azores (Hartert and Ogilvie-Grant, 1905), the Salvages (Lockley, 1952), Madeira (Bannerman, 1914), Cape Verde Islands (Murphy, 1924), Ascension (Allan, 1962), and St. Helena (Haydock, 1954). It may also nest on Sao Tomé in the Gulf of Guinea (Amadon, 1953) but there is no reason to suppose that it nests on Cocos Island, Pacific Ocean (Murphy, 1936). The only detailed breed- ing study is that of Allan (1962) on Boatswainbird Island off Ascension. Vot.XXXVII] HARRIS: BIOLOGY OF GALAPAGOS STORM PETRELS 97 SUBSPECIATION The classification of storm petrels is extremely difficult and has been the subject of much splitting and discussion. Several races of O. castro have been described, all on insufficient specimens, but their validity was successfully chal- lenged by Austin (1952). During the present study I examined far more specimens from Galapagos (the race “bangsi’” of Nichols, 1914) than anyone previous and I also measured the skins of O. castro in the American Museum of Natural History, the Peabody Museum (Yale), Museum of Comparative Zoology (Harvard), California Acad- emy of Sciences (San Francisco), the Los Angeles County Museum, the Smith- sonian Institution (Washington), and the British Museum (London). It seemed therefore desirable to attempt to reassess the amount of variation shown by this species. All measurements were made by myself (thus reducing error caused by differing techniques). Austin (1952) had a larger sample of birds from Japan but it has been thought advisable to use my own measurements; they are, how- ever, similar to those of Austin. The state of specimens and the time available for the work meant that totals of different measurements varied and details are given in table 1. Wing and bill lengths were measured on all but the few molting and damaged specimens. The depth of the bill at gonys, at nostrils, and at its minimum depth just in front of the nostrils and the length of the gonys were also measured but no marked differences were noted. Although any such measurement was likely to be inaccurate, an attempt was made to define the rump patch. The rump patch was measured in the mid-dorsal line from the base of the white, even if hidden by covering feathers, to the furthest limit of the white on the most posterior black-tipped feather. As there were no demonstratable sex- ual differences in any measurements the results have been lumped. Included among the measurements are four birds from, or very near to, Sao Tomé in the Gulf of Guinea. The full details (measurements in mm.) of these specimens collected by Correia and now in ihe American Museum of Natural History are: Male collected 10 December 1928. Wing 155, bill 17.0, rump patch 24. Gonads small. This is in extremely fresh juvenile plumage. Male collected 10 December 1928. Wing 160, bill 17.3, rump patch 25. Gonads swelling. Male collected 14 December 1928. Wing 160, bill 15.5, rump patch 25. In fresh plumage, gonads large. Female collected 16 November 1928. Wing 171, bill 16.8, rump patch 26. Plumage fresh. Dr. Dean Amadon has very kindly examined the details of other birds col- lected by Correia on S40 Tomé and it seems as though these petrels may well have been collected from the shore as other land birds were from the same locali- [ Proc. 4TH SER. CALIFORNIA ACADEMY OF SCIENCES 98 ee 080 €7 es iy It Sy OL g's 87ST Ol 97-SZ Ajnf{—ounf nemepy ee ese 9 ot’ Ly eT ee gest ial S¢-b2 Ayn f—oun [ uede f 9°7 Sys err $9" SST TZ ee s'est 101 ¢¢-7e | Ajn{—ady ‘‘qaq-AoN sosedryer) Ui Ose Gt 89° 9'9T b ss ¢19T t Of-€2 é vouinr) JO jn ee Ope Ze UG: v'ST ce og SOST ce L7-972 220-PO uOoIsuaISY ST 9°9¢ LT ols O'+rT a Ur O'SsT LT b7-€7 22d-VO eua[eH 3S 9°7 iets eb ¢s* est Is O'r O'esT zs 7-1 yoreyp—ue f ‘SE epraA adey rit ere = oy oF" LT f) (eas OSST i O¢-8T “AON—}das $9102 6S" vbr ST 9°¢ 7 Lb ST EMG Ajn f—9un [ SASBATES OT ce OL St" 9'PT 7Z er OTST 7Z Coal (avaA |B) BILAPP IN {ars asDidgnp “ON LOS asDdaap “ON ‘as aspdaap “ON Ne Surpaadg aAnypsaguay yoqog quny yq3ua] 117 yy5uay Sut A Dag Seen eee ee ee 838888] oo ‘(F96T) “MSS Q ‘saauaig fo Kmapvry ay] fo spyyy ayy most aap ‘soSpdypy s0f asoyy mol yavdgn ‘sadnosagumay 4ajDM aavfang “uupnos *) “4q qQ paydqns asamr YI1ym Sasvaywg ayy mosf asoy} sof ydaaxa fjaskKum KQ appus asamr Syuamasnspam Yp “suorDindod snowMma wmwodf OAYSI BUOIPOURIO fo (“MUm) SJUamadnsDayY “T ATAVL Vor. XXXVII] HARRIS: BIOLOGY OF GALAPAGOS STORM PETRELS 99 Ficure 1. A comparison of the bill of a specimen of Oceanodroma castro from the Gulf of Guinea (on the left) with the slightly shorter bill of a bird from the Azores (right). The bill of the Gulf of Guinea bird measured 16.8 mm. The nostrils of the left-hand bird may have collapsed with shrinkage due to drying. ties. There are however no definite data as to whether ‘“‘chummed” at sea or caught on land. Correia was a very experienced collector and it is more than likely that he would have mentioned the fact if they had definitely been breeding. Three of these birds are extremely large in wing and bill length (fig. 1), the female having a wing much longer than any other examined, and all have by far the smallest area of white in the rump of all examined (fig. 2). The differences between all four birds and those from the Cape Verde Islands are significant at the 5 percent level. It therefore seems likely that a subspecies of O. castro may be found in the Gulf of Guinea (possibly at Isla das Cabras where the white-tailed tropic-bird (Phaethon lepturus) probably breeds (Snow, 1950) but it would be premature to describe a new race on only four specimens, before a breeding colony has been found. Although this species shows considerable variation between the isolated breeding populations, especially in wing length, the differences are however not significant except for the Gulf of Guinea birds. I agree with Austin (1952) and Bourne (1955) that the existence of geographic variation should be recognized but that the species is best treated binomially. The populations are probably dis- tinct with little mixing. There is a suggestion that the tropical populations are 100 ’ CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Ficure 2. Photograph to show the relatively small rump patch in two specimens of Oceanodroma castro from the Gulf of Guinea (the two right-hand birds) contrasted with two birds from the Azores (on the left). larger than those of more temperate regions (especially if, as suggested by Bourne (1955), the Azores have warmer surface waters than Madeira). An interesting point concerning differing populations is the occurrence of white feathers in the normally dark parts of the plumage, usually about the head. Baptista (1966) found white feathers in four birds from the Galapagos Islands out of 30 Pacific skins he examined, but I noted only three among 143 Atlantic skins. Admittedly he may have been looking more closely than I did, but there seems to be a regional difference. Allan (1962) did not mention any such birds. In the Galapagos Islands I noted only conspicuous white markings as I was un- willing to disturb my birds more than necessary. Among 1015 birds handled, there were six with very obvious white patches on the head and neck. Two of these birds had white on the head which increased from one season to the next whereas another obtained its white area in a molt between successive breedings Vor. XXXVIT] HARRIS: BIOLOGY OF GALAPAGOS STORM PETRELS 101 (TOWER ) GENOVESA 1] BARTHOLOMEW ie DAPHNE eo GUY FAWKES e Wi CALDWELL os Ficure 3. Definite and possible breeding colonies of Oceanodroma castro in the Galapagos Islands. The present study was undertaken on Plaza. which suggests that the white may be associated with age. These markings are erratic and form no pattern except that they occur mainly on the head. Similar ' markings were noted on a single O. tethys. HABITAT AND METHOD OF STUDY The first Galapagos breeding record of O. castro was made by Beck on Cowley Island in 1906 (Loomis, 1918) and Lévéque (1964) also found it nesting on Plaza and Daphne (for locations see fig. 3). Between November, 1965, and July, 1967, I visited all the main islands and most of the climbable isolated rocks with 102 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. the exception of Culpepper, Crossman, and some of the Baimbridge Rocks, look- ing for nesting storm petrels. Oceanodroma castro was found nesting on North and South Plaza, Daphne major, Guy Fawkes, Cowley Islet, Onslow, Isla Pitt, and Tower. Dead birds or remains in pellets of the short-eared owl (Asio (flam- meus) galapagoensis) were also collected on Hood, Caldwell, Bartholomew, and Bainbridge Rocks; the species may well nest on all of these excepting Caldwell which, like the majority of the smaller isolated islands and rocks, is made of sandstone-like lava so smooth as to be virtually holeless. All the colonies were reasonably small and well spaced out, as appears typical of the species. General observations were made at several of the colonies but the main part of the study was undertaken on South Plaza. The Plazas are two small islets of basaltic lava about 400 yards off the eastern tip of Santa Cruz. Both are inhabited by sea birds but, as the larger South Plaza has more birds and lacks the extremely dense thorn scrub of North Plaza, all previous observers have restricted their activities to South Plaza. My main studies, likewise, were carried out on South Plaza, but regular checks were also made on the northern island and so, for the sake of brevity, the term Plaza where used will signify South Plaza. South Plaza is an elongated island about a half mile long by an average of 150 yards wide with the long axis running approximately east-west. The south side, which is exposed to the prevailing southeast winds and where all but a handful of the sea birds nest, has cliffs mainly 30-60 feet high but decreasing to 10 feet at the ends of the island (fig. 4). At the bases of the cliffs there are considerable quantities of boulders due mainly to cliff falls, which still continue. From the cliff tops the island slopes gently down to the sea on the very sheltered northern shore. The western third of the island has a covering, dense in places, of thorn scrub. Cacti (Opuntia species) are common in this area and also occur singly or in small groups scattered over the rest of the top of the island, which otherwise has only low ground vegetation. Five species of sea birds nest on Plaza, the swallow-tailed gull (Creagrus furcatus) (on the cliffs, among the boulders, and rarely inland), Audubon’s shearwater (Puffinus lherminieri) and Oceanodroma castro (both of them in holes in cliffs and boulders), red-billed tropic-bird (Phaethon aethereus) (in cliff holes), and a few brown noddies (Anous stolidus) (on exposed cliffs). Masked and blue-footed bobbies (Sula dactylatra and S. nebouxii) and frigates (mainly Fregata magnificens) are always to be seen on or around the island but none nested. Other nesting species were the finches (Geospiza fortis, G. fuligi- nosa, G. scandens), the yellow warbler (Dendroica petechia), and a single pair of yellow-crowned night-herons (Nyctanassa violacea). An egret (Casmerodius egretta), one or more grey herons (Ardea herodias), and one or more individuals of Asio galapagoensis were regularly present, and the owls may well have bred on Vot.XXXVII] HARRIS: BIOLOGY OF GALAPAGOS STORM PETRELS 103 Ficure 4. Part of the south-facing cliffs on Plaza Island during the hot (and calm) sea- son. Creagrus furcatus nests on the flatter area (one bird can just be seen incubating in the guano covered area), while Oceanodroma castro, Puffinus Iherminieri, and Phaethon aethereus nest in the cliff holes and among the boulders. The guano is mainly due to Sula nebouxii and S. dactylatra which however do not breed on Plaza. 104 ' CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. North Plaza. A surprising absentee was the mockingbird (Nesomimus parvulus), which would be a potential nest predator (Harris, 1968). There were three other large vertebrates which were known to be responsible for losses of sea-bird nests. The largest and most numerous was the sea lion (Zalophus californianus), whose population was in excess of a thousand, mainly on the north side but with haul-outs of unemployed males near the east and west tips. Plaza supports the densest colony of land iguanas (Conolophus subcristatus) remaining in Galapagos and also a smaller colony of marine iguanas (Amblyrhyn- chus cristatus). The former species is restricted to the flatter area of the island, the latter usually to the sea cliffs, but both species are not infrequently found in the nesting holes of sea birds. Nest SITES The nest sites of O. castro varied greatly in the Galapagos but the preferred site appeared to be a hole in a cliff or under a boulder, deep enough to hide the bird from sight but not so deep as to exclude all light. Direct access to the sea is extremely important and very few birds nest far from the cliff edge or the sea. Of 171 nest sites examined on Plaza, 55 were in holes in cliffs, 27 in holes at top of cliff, 38 in boulders at cliff base, 38 among boulders or large stones on cliff top, 6 in the back of small caves, 5 in holes excavated by the birds in soil, mud, or seal guano, and 2 inland under boulders. Of the cliff nest sites, half were under substantial overhangs and only 10 allowed a clear view of an incubating bird. These figures are probably biased towards the shallower sites, as these are more easily found, but it is unlikely that many birds nest in very deep holes. On Onslow there were very few holes indeed and birds nested very much more in the open than on Plaza. Onslow was not visited during a breeding season, but out of season there were many abandoned eggs and also the remains of adults eaten by owls (Asio galapagoensis). In a colony such as this, the incubating birds must be very liable to predation. The total population of any small nocturnal species is extremely difficult to determine, but on Plaza it was thought that about a third of the available nest sites had been found by the end of the study, which would put the number of oc- cupied holes in the cold season at about 600 and in the hot season approximately 300. However the data on owl predation and the estimate of the annual adult mortality taken together, suggest that my estimates were too low. Regular monthly visits were made to Plaza. At least 10 days a month were spent on the island (with only one exception), usually divided into a stay of a week or more, with several day visits timed, as far as possible, so that never more than 10 days (or a week at peak breeding times) elapsed without the nests being checked. In all a total of 208 days was spent on Plaza. All holes with birds found were given a description and a number which was painted on the rock alongside. The number of holes used in the study increased Vor. XXXVII] HARRIS: BIOLOGY OF GALAPAGOS STORM PETRELS 105 throughout the period, from 74 at the end of the first breeding season to just over 180 at the end of the study. Some birds were also found in burrows of Puffinus lherminieri, which was the subject of a separate study (Harris, 1969). Birds were marked with numbered bands and examined for wing molt and state ef develop- ment of the brood patch. When time allowed, they were also weighed and measured. Birds were sexed by cloaca inspection after the laying of an egg, and adults of some pairs marked with spots of paint on head, wings, and tail, so that incubation spells could be followed without handling the birds. Luckily, and un- like some other storm petrels, such as the British storm petrel (Hydrobates pelagicus), this species does not desert readily, so that it was possible to handle and even weigh some birds daily during incubation. Eggs were measured and young were weighed daily to obtain growth rates and twice daily to get weights of feeds and feeding frequencies. The majority of the nests were examined every day, but in some awkward or dangerous areas they were checked only two or three times a month. The terrain was such that night work was hazardous and restricted to a little mist-netting and a few searches for holes or specific birds. This part of the study was very similar to that on Ascension by Allan (1962). However, unlike those of Ascension, the adult and young petrels here were subject to much predation by owls, and young were also taken by herons (Ardea herodias) and the red crab (Grapsus grapsus). Some eggs and chicks were also destroyed by the two species of iguana. GENERAL BIOLOGY Virtually nothing is known about the biology or distribution of this species at sea except that it is seldom seen on or from ships, so that our knowledge of its biology is restricted to the breeding grounds. However, it does appear to feed well away from land, presumably mostly by day. In accordance with the strict conservation measures now enforced in the Galapagos, no attempt was made to collect large numbers of birds for the exam- ination of stomach contents, but during the study 15 stomachs, mainly from birds injured or killed by owls, were collected. Of these 14 had fish-eye lenses (up to 19) or otoliths, and 4 had cephalopod remains. One cephalopod beak was identi- fied as belonging to a myopsid (probably sepiolid) of estimated weight 3—4 grams. Two adults netted in April regurgitated fish of lengths 50 and 37 mm., and the droppings of a young bird on Plaza in August contained a single fish otolith and a smashed cephalopod beak. This evidence indicates that the main food is small fish (probably mainly of the size of the regurgitated sample to judge from otolith size) and squids, all caught on or near the surface. The inside of the mouth of O. castro has large numbers of relatively big back- wardly pointing spines (fig. 5), and is well adapted for catching and holding slip- pery prey. The stomach is large, approximately 55 mm. long by 38 mm. (width when cut open and flattened) and capable of large distention. The intestine 106 ! CALIFORNIA ACADEMY OF SCIENCES | Proc. 4TH SER. “« ae ewe @ a AVIS & ea ¢ —_ a*® @ aq ememeg=~@ a ry a Palate Ficure 5. ‘Tongue and palate of Oceanodroma castro to show backwardly pointing spines. measured about 250 mm. The roughly dissected flight muscles of a bird of 33 grams, weighed 6 grams or 18 percent of the body weight. As the method of feeding is presumably similar to that of other species of Oceanodroma, that is, catching prey at or just below the surface of the sea, it is hardly surprising that birds show injuries to the feet, presumably due to preda- tory fish. A sample of 100 birds examined closely included 7 with injured feet. The injuries were holed webs (2), only a single toe left on a foot (2), distal half of a foot missing (2), and leg missing completely from outside the body (1). Injuries seen in other birds were a badly torn web (1), a leg missing outside body (1), and a bird unable to straighten a leg (1). Allan (1962) found similar in- Vor. XXXVII] HARRIS: BIOLOGY OF GALAPAGOS STORM PETRELS 107 juries in 3% percent of his birds. These injuries did not seem to affect the re- productive success of the birds concerned. THE BREEDING CYCLE The majority of sea birds have fixed annual breeding cycles, usually assumed to be timed to coincide with the maximum availability of food for egg formation, feeding, and/or survival of young. However, in a small but varied group of equatorial species, breeding occurs at intervals of less than a year. In some spe- cies, such as Anous tenuirostris and the sooty tern (Sterna fuscata) (Ashmole, 1962, 1963) on Ascension, it seems clear that the time between the starts of suc- cessive layings is that needed to complete the breeding processes and undergo a molt, a period of just under 10 months in S. fuscata. In a few other parts of its range, S. fuscata has breeding seasons at 6-month intervals, and on Christmas Island (in the Pacific) successful breeders nest annually, but those which lose their egg or young chick may return for the next cycle 6 months later (Ashmole, 1965). Oceanodroma castro is a widespread species, but since it breeds on isolated islands and is nocturnal when visiting land, there is a dearth of precise data on its breeding cycle, except on Ascension, where there is an annual cycle with all but a few eggs laid from October to December (Allan, 1962). The Blossom col- lections taken on Ascension in 1925 (now at the Peabody Museum) include some young birds from eggs which must have been laid August-November which per- haps fits with the contention of Allan that there is a fixed annual cycle. The few eggs from St. Helena were of October and November origin (Benson, 1950; Hay- dock, 1954; Stonehouse, 1963b). In the Cape Verde Islands, breeding is recorded in the first half of the year (Bourne, 1957) whereas in the Salvages it is probably a little later with no breed- ing found at the end of April (Ogilvie-Grant, 1896), and fresh and incubated eggs and very few small young in July (Lockley, 1952; Dr. C. Jouanin, personal communication). In the Madeiran group, eggs have been found in every month but May (Ban- nerman, 1914 and Jouanin, personal communication), and Schmitz (Bannerman, 1914), thought that there was no definite breeding season. Bannerman (1914) interpreted the same results as suggesting that there are two breeding seasons, June-September and October-December, involving different individuals and a few birds laying in between. Lockley (1952) spent the 4 days 10-13 July 1939 at the colony at Chao Deserta and found birds present but no nests. Roux and Jouanin (personal communication) visited Baixo in July, 1967 and found the birds with fresh eggs, incubated eggs, and rarely with very young chicks. It is therefore uncertain whether there are two seasons a year, one season with some out-of-season nesting (as Ascension), or a less than annual cycle. Further north 108 , CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. 40 1966-67 Season Mid 1967 Season 1965-66 Season Mid 1966 Season Number of eggs /aid N D J F M A M J J A S O MONTHS Ficure 6. The distribution of egg-laying dates of Oceanodroma castro in the study bur- rows on Plaza November, 1965, to the start of July, 1967. in the Azores no breeding was found 1 June, but birds were coming ashore to breed in September (Hartert and Ogilvie-Grant, 1905). In Japan the birds arrive at the colonies in late May, breed through the sum- mer, and leave in October (Austin and Kuroda, 1953). The scant Hawaiian data of several barely flying young found in October in the late nineteenth century, suggest a similar cycle (Richardson, 1957). For many years it was thought from the single breeding record quoted by Loomis (1918) that egg laying in Galapagos took place May-July, but a single TaBLE 2. The numbers of birds ringed for Oceanodroma castro on Plaza in one season and of retraps from one season to another. The figures in brackets after the number of birds handled is the number of possible nonbreeders in the sample. Birds handled in season 1965-66 Mid-1966 1966-67 Mid-1967 1967-68 Mid-1968 1968-69 Mid-1969 118 305 213 410 15 0) 17 18 [30] [113] [66] [221] Retraps from season 1965-66 0 64 0) 5 8 0) mid-1966 0) 150 0 0 10 1966-1967 6) 5 6 0) mid-1967 0 0 6 Vot.XXXVII] HARRIS: BIOLOGY OF GALAPAGOS STORM PETRELS 109 chick found by Brosset (1963) indicated an egg laid around December. Snow and Snow (1966) have since shown that on Plaza there were two breeding sea- sons a year, with eggs laid March-June and December-January. They thought that the same birds were probably involved in successive cycles, and also that the December-January season was subsidiary, with less birds breeding and a lower nesting success. Details of all eggs found on Plaza between November 1965 and July 1967 are shown in figure 6, and it should be stressed that the same burrows were often used by different birds in successive cycles, a few in all four seasons for which records are available (see later). There were four sharply demarcated cycles, two in the hot seasons (November to February) and two in the cold (May to July). Note that the two peaks do not come quite at 6-month intervals, but closer to 5 and 7 respectively, the significance of which is not known. All birds ringed as breeders and nonbreeders in one season and retrapped in another are shown in table 2, which indicates that each individuals bird had an annual cycle and that there were no recorded interchanges of birds between the cold and hot seasons. Unfortunately, only 12 adults and a single chick of this species had been ringed previously on Plaza (in the cold season of 1960 by Lévéque) and only one of these was recorded. This, however bred again in the cold seasons of both 1966 and 1967. The species has such a long breeding cycle, involving at least a 3-week pre- egg period, 6 weeks’ incubation, up to 3 months feeding the young, and with the need to fit in a complete molt as well, that it is clear that successful breeders could not nest in successive seasons during the same year. But the faithfulness of failed breeders and nonbreeders to the same cycle was unexpected and con- trasts markedly with the results for S. fuscata where it breeds twice a year on Christmas Island (Ashmole, 1965). This complete separation of birds nesting in the hot and cold seasons, if as- sociated with young birds returning to breed at the same season of the year as they were raised, could potentially give rise to separate forms of the same species. However, no morphological differences were found between adults at different seasons, though there was a slight and constant difference in egg breadth (see later). CLIMATIC FACTORS Although the Galapagos Islands straddle the equator, the climate is only subtropical owing to the modifying influence of the Humboldt Current.* An ac- count of the rather complicated currents is given by Abbott (1966). Figure 7 shows the average monthly surface water temperature taken by the * The 1965 temperatures were in a year when the Humboldt Current lessened allowing ‘El Nino’ or warm current to affect Galapagos. The phenomenon was responsible for a very large mortality of sea birds from the guano islands of Peru. 110 q CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. 60 J F M A M J J A S OC Neat MONTH Ficure 7. Surface water temperatures taken at Academy Bay, Santa Cruz, Galapagos, by the Charles Darwin Research Station. Charles Darwin Research Station at Academy Bay, Santa Cruz—about 18 miles from Plaza. Despite a regular fluctuation in water temperatures I was unable to detect any large-scale seasonal variation in surface plankton, although the samples show that it appears to be very variable both in time and place. As a general rule, the seas and winds are calmest during the warm months, and strongest during the cold season, but there is little correlation of breeding success with time of year. ACTIVITY OF NONBREEDERS In all sea bird populations there are large numbers of nonbreeding birds which may or may not frequent the breeding colonies. In some instances, as in the Manx shearwater (Puffinus puffinus) and H. pelagicus, the total number of nonbreeders visiting the colonies throughout the breeding season may be a substantial pro- portion of all the birds there (Harris, 1966a, and personal observations). Unfortunately netting at night was all but impossible, so observations of nonbreeders were almost entirely restricted to those roosting in burrows by day. However night observations and estimates of the number of birds calling in flight agree very closely with the three seasons’ results given below. Detailed observations were made in all four seasons but only three seasons are treated here as many fewer burrows were available in the 1965-66 season. In the other Vot.XXXVII] HARRIS: BIOLOGY OF GALAPAGOS STORM PETRELS 111 70 > 60 Q Mutts = 350 [-] Incubating birds 8 FZ Young = 40 is) S g 30 = 8 ®& 20 10 oO 1966 iog7) ONaH S NON-BREEDERS lu 10 S G5 8 H = 3 © ps) Q w 8 OTHER BREEDERS RAY eh ADSM: J) 0) A Sl OUWN UD UR) ME. A OM 1966 1967 MONTH Ficure 8. The percentages of the study burrows of Oceanodroma castro occupied Jan- uary, 1966, to July, 1967, by a) incubating birds (unstippled area) and unattended young (stippled), b) nonbreeders, and c) breeders before laying or after losing egg or young. seasons, 43 percent, 43 percent, and 64 percent of all burrows had nonbreeders roosting in them at some time, and, as the burrows were checked on only about 1 day in 3, the true proportion must have been much higher. The visits by nonbreeders were not scattered throughout the year but were closely correlated with the reproductive cycles of the breeders, and to a lesser 112 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. extent with the phases of the moon. A full moon inhibited most nonbreeders from visiting the colonies. The monthly incidence of burrows occupied by nonbreeders is shown in figure 8b as a percentage of all the burrows examined in any period. The actual number of nest checks varied greatly between different periods but averaged about 400, with all but 8 involving more than 150. The proportion of burrows occupied by incubating birds, unattended young (fig. 8a), and birds known to have bred later in the same season or which had lost egg or young (fig. 8c) are also shown. Although nonbreeders will occupy a hole with an unattended egg, or rarely, with a chick, they do not usually use burrows occupied by adults or large young. An interesting point brought out by these figures is that the nonbreeders were most numerous at the same time of year as pre- and failed breeders, with a peak before most burrows were occupied by incubating birds and young. Some non- breeders indeed return with the first returning breeders. This is different to the pattern found in H. pelagicus (Davis, 1957, personal observations), and in the shearwaters P. puffinus (Harris, 1966a) and P. tenuirostris (Serventy, 1967), where the majority of nonbreeders arrive back long after the adults, are most numerous during the incubation time, and decrease only after the peak of hatch- ing is passed. At least in the shearwaters, older prebreeders possibly sexually mature but still not breeding, return with the adults and do not leave until the first young fledge. The three species mentioned above are long-distance migrants, however, so the results may not be directly comparable; although we have no definite information for O. castro, it may well have no regular migration. Both food and predation might be important in the evolution of the timing of the visits of immature petrels to the colonies. Given that the food supply is reasonably constant (see later), any competition for food would be greatest when the adults were feeding rapidly growing young. If there was competition for food it might be thought that the possibly less efficient immature birds would suffer, so reducing their chances of survival as compared with any not present at the colonies. However, an adult feeding young must collect at least one-and-a-half times as much food as a nonbreeder, and also use much energy finding this extra food and flying to and from the colony. Individual nonbreeders visit the colony only infrequently and can spend most of their time on the feeding grounds. As it appears unlikely that a nonbreeder is only half as efficient at feeding as a breeder, the first result of any competition for food would be food shortage for the nestlings. The adults themselves look after their own needs before those of their young (Harris, 1966b). Predation by owls is important in this colony (see later) and perhaps the nonbreeders, by returning with the displaying adults, increase their own chances of survival as the predators would take proportionately many fewer birds than if the return was spread over a longer time. Of these nonbreeders, some presumably Vot.XXXVII] HARRIS: BIOLOGY OF GALAPAGOS STORM PETRELS 113 TABLE 3. Weights in grams of Oceanodroma castro found in burrows on Plaza Island. Aver- Standard Aver- Standard No. age Range deviation No. age Range deviation 1965-66 season 1966-67 season October 5 42.1 39.0—-44.5 2.2 November 39 39.2 33.0-45.5 Bil December 6 42.5 35,0—53.0 5.6 37 38.7 31.5—50.0 3.8 January 16 47.3 41.0-55.0 4.8 20 40.2 32.5-56.5 4.2 February 2 46.0 38.0—54.0 4 39.5 28.5—41.5 4.7 April 1 39.5 Mid-1966 season Mid-1967 season April 33 39.3 31.0—49.0 Sint 3 40.8 38.5—43 .0 May 34 41.3 34.5—54.5 4.7 52 39.7 32.0—48.0 S35) June 102 45.3 33.5—-54.0 Sal 27 41.5 34.0—50.5 4.5 July 4 44.4 39.0-46.5 il 1 39.0 August 3 42.7 40.5—44.5 form pairs with birds that have lost their mates and others find and occupy empty burrows; the remainder, probably the majority, leave the colony so do not en- danger their survival. In any long-lived species, such as this must be, it pays an individual in terms of overall reproductive output, to postpone breeding for a year and leave the colony, rather than reduce its chances of survival. Nonbreeders were, with two exceptions, faithful to a hole from one season to another even if the hole was simultaneously occupied by a breeding pair. WEIGHTS OF ADULTS Many adults were weighed to .5 gram with spring balances but, as these soon became inaccurate, the observed weights were corrected by a callibration graph made for each visit to Plaza. Full details of the weights are given in table 3. The 12 birds weighed by Allan (1962), which averaged 43.5 grams, standard devia- tion 5.0, agree with my series. The differences between the four breeding seasons are not associated with birds breeding at different times of year and might indi- cate some difference in the availability of food (but see later). In all four seasons, the average weight increased as birds started incubating and had food stored in the stomach, reached peaks during January and June, and then decreased when the birds had small young. There were no significant differences in the weights of adults before laying, failed breeders, and intruders. VISITS TO BURROWS General observations showed that as soon as birds returned for the start of a season, some might remain in the burrows by day. In extreme instances, adults [Proc. 4TH SEr. CALIFORNIA ACADEMY OF SCIENCES 114 “BulAey 0} 101d A[ayeIpawUI SMOLIN UT PUNO] spllq OT Ipnj[out jou saop sIyy, y «80 08 L9 Lt OTZ vst aval $6 6 ST ss 87 Spiiq pexssun suIpnypout % 0 Ct 8 v9 OTT 68 IS oe aC on iS 9° Jay}e50} Y}0q skep % of iS oT iS 6¢ 87 14 0) S @) (0) 0 Ajuo ajeway skep % «VL Or se vs or £9 cr ST Gao 0) LT TT A[uo ajeu skep %, SOE ele OLE 967 To¢ 6LT LEZ 89T Sst £OT est oLT skep afqissod jo ‘oN et! Oo Sir 02-91 $2-1z —0€-92 SE-I€ — Ob-9E St-Ib S96 =~ S$-1S 09-95 surkn] asofag Skog “‘Suitp] asofaqg stop ZL puv [9 Uuaanrjzaq (sp4iq Y10q yym ¢) skop aarf uo quasaad Spdiq OsjD adam aday, “Susp asofag spoisad Kvp-¢ Buranp Sjadqag WAOys OAYSLD LWOIPOURID Kq smoOddng Busau ur quads skvq “pf ATAV, VoL. XXXVII] HARRIS: BIOLOGY OF GALAPAGOS STORM PETRELS 115 stayed in burrows still occupied by the previous season’s young. This was how- ever rare as there was little overlap between successive seasons. There were no detectable differences in the occupation of burrows before lay- ing in the four seasons, and all the results are treated together in table 4. The longest recorded interval between birds first roosting in the burrow by day and laying was 72 days, but in several burrows no birds were found until the egg was laid. After these early visits, the frequency of birds in the burrows increased rap- idly to a maximum 3-4 weeks before laying and then decreased to a minimum 2 weeks before laying. This decrease was most marked in the females, which were presumably far away at the time feeding hard in order to form the large egg. There is however no well-marked “honeymoon period” as in P. tenuirostris (Marshall and Serventy, 1956) when both birds are away from the colony. The longest recorded stay in a burrow was 5 days by a female which was joined by the male for 3 days. It was common to find both birds together in the burrow by day. The male continued to roost in the burrow occasionally, perhaps so as to keep possession, right up to the time of laying, even though it might have been expected that he would have been away building up his reserves in preparation for the first incubation spell. In the 5 days immediately prior to laying, birds of known sex were found in the burrow in 41 instances—only one of these was a female, which returned 2 days before laying. Of the males, 16 were found the day before the egg was laid (and it might be assumed that they were waiting for the first incubation spell), eight were 2 days prior, six were 3 days, three were 4 days and seven were 5 days prior to laying. Normally the female leaves after laying the egg and the male incubates. In some instances, however, probably when the male has not returned, she may either incubate for 1 or 2 days or leave the egg. It may well be that the arrival of the male at laying time is a matter of chance and perhaps at this time he is feeding near the colony by day and visiting the nest site most nights. The data of Pinder (1966, fig. 4) for the Cape pigeon (Daption capensis) lend support for this view. BEHAVIOR On Plaza the species flies only at night and in only a single instance, about 30 minutes before dusk, was a bird seen near the island by day. The first birds _arrived ashore about 1850 hours, that is, 50 minutes after sunset, but birds were never heard calling before 1900 hours. The peak activity was normally 2200 to 2400 hours on nights without moon but just before dawn on nights with a full moon. The effect of the moon was very marked, especially affecting the numbers of calling birds. The calls have been well described by Lockley (1952) and Allan (1962) and were uttered both in flight and from the burrows. Birds usually flew close to the cliffs and rarely over the flat top of the island 116 é CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. and there appeared to be two types of display. One was directed toward the holes, usually to a hole with a churring occupant. Presumably unattached birds are attracted by these calls emitted by a bird with a hole but not a mate. In sev- eral species of storm petrels, adults can be attracted by a tape recording of these churrs. (Huntington, personal communication and personal observation). The second display was aerial and usually between two birds which chased each other in reasonably constant circuits. This was very noticeable when nets were used, as some pairs would circle time and again, just missing the net, until one, or usually both, were caught. In one display, two birds repeatedly went through a 2-foot gap between the net and the cliff face until one was caught; the other then did its usual circuit before joining its mate in the net. A single mating was observed by day in a very open nest site. The prior dis- play was not observed, but the whole subsequent procedure was silent. Throughout the mounting, which lasted at least 3 minutes, the male gently pecked the female’s head, moving from side to side across the head with special emphasis at the base of the upper mandible. The egg was laid 33 days later. DEVELOPMENT OF THE Broop PATCH Little is known of the molt of the brood patch in sea birds and the only ob- servations on petrels appear to be those of Allan (1962), who thought that no bird molted its brood patch later than 20 days before laying, that some started at least 40 days before laying, that the actual molt might be completed in no more than 4 days, and that vascularization was complete 5 days before laying. In the present study there were 249 observations on birds prior to laying, 267 on nonbreeders, 44 on birds which had lost the egg, 10 on those with young, and 3 on those which had lost young. With a few exceptions, incubating birds were not examined. There was no apparent difference between the sexes so the results have been lumped. Similarly there was no difference between the various seasons. The state of the brood patch was classified and scored as follows: Score 0. No sign of brood patch. Score 1. Brood patch half defeathered. Score 2. Brood patch defeathered but unvascularized. Score 3 Score 4 . Blood vessels just visible. . Fully vascularized, with blood vessels “knotted” and obvious. No special note was made of oedema, but it was apparently only present in incubating birds. Apart from a netted sample which is discussed separately, all birds were taken from burrows and their breeding state was known. Any dubiously breeding birds have been omitted. The laying dates of most breeders were known to within 2 or 3 days, and if not so exactly, then the mean date was used between a check without and with egg provided that the interval was less than 8 days. Although it is realized that the scores 0-4 are probably not equivalent in Vor. XXXVII] HARRIS: BIOLOGY OF GALAPAGOS STORM PETRELS 117 TaBLeE 5. State of brood patch in early and late nesting representatives of Oceanodroma castro in relation to days before laying. Stute of brood patch 0 = no trace, 1 = half de- feathered, 2 = defeathered but unvascularized, 3 = blood vessels just visible, 4 = fully vascularized. State of brood patch and numbers of birds Early breeders Late breeders Overall 0) 1 2 3 4 Average 0 1 2 3 4 Average average Days before laying 46+ 1 2 1 3 2 3 9 1 1.6 1S) 41-45 z 1 5th 1 1 2.0 1.2 36-40 4 3 9 9 0} 18) 31-35 4 1 8 2 V5 6 4 4 1 1.6 1.6 26-30 11 4 8 1 Z eZ, 4 13 2 2.0 ies) 21-25 5 4 9 4 ez 7, 1 7 4 1 2.4 2.0 16-20 2 1 5 1 Ds, 5 13 4 Z 2 Med 11-15 1 3 3 3 2.8 2 2 2.5 ASH 6-10 1 1 2 8 3.4 3 1 He) 3.2 1-5 4 10 So 1 3 5 3.4 3.6 time or metabolic effort to the birds indicated, an average score for each 5-day period prior to or after laying was calculated and used below. BREEDING BIRDS At least some birds lose the feathers from the brood patch immediately after, or perhaps even before, coming to the colony at the start of a season. Of 10 adults caught more than 50 days before laying, only two were without a brood patch, and one even showed some enlargement of the blood vessels. However, these birds may have been slightly anomalous, as many other individuals come back without any trace of a brood patch. The progression of the losing of the feathers and vascularization is shown in table 5. It is difficult to find how long the various stages take as individual birds were irregular in their roosting in the holes. But the process is certainly very variable, as some birds have fully vascularized brood patches more than 6 weeks before laying, but others not even when the egg is laid. The minimum observed time for vascularization of a defeathered brood patch was 7 days. I agree with ‘Allan (1962) in finding that all birds had lost all the feathers of the brood patch at least 20 days before laying, but this is hardly surprising, as most nonbreeders also reach this stage. My minimal observed time for defeatherization was 12 days. The most complete record for any bird was (1) 32 days prior to laying, no brood patch, (2) 23 days prior to laying, defeathered but nonvascular brood patch, (3) 18 days prior, partially vascularized, and (4) 2 days before laying, a fully vascularized brood patch. 118 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. TaBLe 6. State of brood patches in nonbreeding members of Oceanodroma castro. Details of brood patch numbers as table 5. Numbers of birds with brood patches 0 1 2 3 4 Average October 1 1.0 November 4 4 23 2 le December 1 40 12 DD) January 20 5 1 2.3 February 2 1 es March 1 0 April 8 1 4 eit May 11 5 66 10 3 1.9 June Bil 7 2 2.4 July 1 i 2 1 2.2 August if 2.0 There is evidence to suggest (table 5) that birds laying early in the season arrive back with brood patches less developed relative to the date of laying than do later nesters, and that this is associated with quicker development. Regrowth of the feathers of the brood patch usually started about a week after hatching, but the process was again variable and the numbers of birds ex- amined small. Ten birds netted on Tower 16 February 1966 consisted of five birds with brood patches unvascularized, four with feathers growing, and a single bird with no brood patch. At least six of these were probably feeding young, as they regurgitated large quantities of food. NONBREEDERS The majority of nonbreeders (184 out of 267 observations) had defeathered but unvascularized brood patches, while many fewer (39) showed signs of vas- cularization. A few birds (7) had fully vascularized brood patches identical with those of breeders. The brood scores (table 6) show that there is a cyclic fluctua- tion related to the 6-monthly breeding. Unfortunately, I know little of birds coming ashore by night, but a sample of 36 birds caught at night, 2 and 3 June 1967 consisted of two birds with no brood patch (one bird had bred a year pre- viously), 32 with it defeathered (one bird had bred a year previously), and two with it fully vascularized (one had an egg about to be laid). This gave an aver- age brood score of 2.0, or 1.9 if the breeder is omitted, which is significantly lower than that for 30 birds taken from burrows in June (2.4). This is probably explained by the birds spending time in aerial display at this stage of the breed- ing cycle being younger than those which were overstaying in the burrows. Vor. XXXVII] HARRIS: BIOLOGY OF GALAPAGOS STORM PETRELS 119 TaBLe 7. Egg measurements (in mm.) of eggs of Oceanodroma castro. Average Standard deviation Number Length Breadth Length Breadth Galapagos 1965-66 season 16 Sly 22.8 1.8 .80 mid-1966 season 72 31.5 23.5 il8! Shs 1966-67 season 45 Si lei 22.6 1.4 eal mid-1967 season 61 31.4 23.6 95 .66 Cape Verde Islands (Bannerman, 1959) 2 31.0 23.8 St. Helena (Haydock, 1954) 8 Bors 24.2 Madeira (Brit. Mus. and Jouanin) 9 32.8 24.9 iil 48 Ascension (Stonehouse, 1963a) 44 322 23.5 ih) 1.0 Salvage Islands (Jouanin) 5 Sasi 25.0 91 Hil THe EGG-STAGE The egg was always laid at night and then either incubated by the male (32 instances), by the female (23 instances), or left unincubated (6 instances). The male took the first long incubation stint and the female only waited for him to appear before departing; if he did not arrive soon she either incubated for 1 or at the most 2 days or departed at once. The male took over from the female the night after laying in at least five cases. The single egg was white, often with a ring of faint pink marks (as in other storm petrels) at the blunt end which soon disappear with incubation. The mea- surements of 194 eggs from Galapagos and some from other colonies are shown in table 7. The differences between the seasons and the colonies are not signifi- cant. The extreme measurements of the Galapagos eggs (in mm.) were 34.8 X 22.1, 28.0 X 22.3, 32.7 X 24.9, 28.3 X 21.0 (which was also the overall smallest egg) and 34.0 X 24.0 (overall largest). Twenty-eight newly laid eggs averaged 8.5 grams, range 6.0 to 11.1 (standard deviation .93). Five newly laid, and hard boiled, eggs had yolk: albumen weights of 2.3 : 4.3, 2.5 : 4.9, 2.4 : 4.9, 3.6 : 6.8, 2.8 : 3.8. The shell weights were from .3 to .7 grams but my balance was not very accurate in this range. The female usually weighed least immediately after laying, 20 individuals weighed the morning after laying averaged 39.8 grams, range 36.5 to 45.5 (stan- dard deviation 2.6). One bird caught after laying in 2 successive years weighed 36.5 and 38.0 grams. This postlaying weight is still slightly heavier than non- breeders (average 38.9 grams) and only just under that of adults in the prebreed- ing period, 40.3 grams), which suggests that the female obtained the food reserves for the egg during the prelaying period at sea. The female is probably lighter than nonbreeders if the weight of the enlarged ovary is excluded. The egg was approxi- mately a fifth of the female’s weight (20 eggs, range 17 to 24 percent, average 21 percent) as compared to 26 percent in Oceanites oceanicus (Roberts, 1940) 120 f CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. and Pelagodroma marina (Richdale, 1943-44), 25 percent in H. pelagicus (Davis, 1957), and 20 percent in Oceanodroma leucorhoa (Huntington in Lack, 1967). The measurements of eggs laid by individual females in two seasons showed less variation than those of the population as a whole. Birds were extremely faithful to their burrows, and this could sufficiently explain this tendency to have the same mate in successive years. In only 10 in- stances was a bird ever caught in a different hole from that where it had been ringed. Two of these refer to a pair which were found breeding in the 1966-67 season in a hole 12 feet away from that occupied in the 1965—66 season. It is impossible to say if they moved as a pair or met again by chance in a new hole. One other breeder was found a second season in a hole a few feet from where it had previously bred. Four nonbreeders in one season were retrapped in another season in other holes, all within 10 feet; two of them were breeding with unringed birds. A single bird was caught once in each of two seasons in shearwater holes 150 yards apart. The remaining two cases referred to two holes only 6 inches apart in a smooth vertical rock face and could easily have been due to birds entering the wrong hole. All other retraps between seasons (194 birds) and within a season (many thousands) were in holes where the birds had been ringed. Obviously birds could have moved into burrows inaccessible to me but it is unlikely that burrow desertions occur at all frequently if both birds of a pair remain alive. Richdale (1965) found similar results in Pelagodroma marina. The laying dates of pairs were not influenced by the success or otherwise of the breeding attempts in the immediately previous season even if there had been a late chick present when the first adults returned for the new season. This sug- gests that the gap between breeding cycles was sufficient for all the prelaying activities. Pairs which had bred before laid an average of 4 days before new pairs but this difference was not significant. The relative shortness of my stays on Plaza (maximum 13 days) did not al- low many individual incubation spells to be calculated directly and the results are biased towards the shorter spells. Observations were however made on color- marked birds and the results showed that the normal spell was 4 to 7 days with, rarely, a prolonged stay of at least 11 days, or as short as 2. Such long stays are however very unusual as most birds leave the egg long before this if not relieved by the mate. Another method of calculating the average incubation spell (table 8) is by observing the proportion of nests where changeovers have occurred over- night, with the proviso that the observations are spread over enough nights to avoid bias due to many pairs changing over together. The average spell appeared to be 6 days as compared with 2 days in Oceanites oceanicus (Roberts, 1940), 3 to 5 days in Pelagodroma marina (Richdale, 1943-44) and 3 days in H. pelagicus (Davis, 1957). Vot.XXXVII] HARRIS: BIOLOGY OF GALAPAGOS STORM PETRELS 121 TaBLE 8. Number of change overs from one day to the next in incubating members of Oceanodroma castro. Number Number with Average incubation Year Month examined different bird shell (days) 1966 January 62 11 5.6 1966 June 392 62 6.3 1967 January 103 Al 4.9 1967 May 25 6 4.2 1967 June 170 24 Heal 1967 July 43 8 5.4 Total 877 146 6.0 As reported by Allan (1962), a strange bird may rarely be found in a burrow with an egg when the normal adults are missing. In this study such birds were never actually incubating, the egg being cold, and were certainly nonbreeders prospecting for holes. Some birds were weighed daily during incubation and the observed weight losses are shown in figure 9, the average loss being 1.6 grams a day. Expressed another way, for a normal incubation spell a bird must have stored food reserves equivalent to a quarter of its normal weight. Three birds were weighed on a day before a changeover and then the first day of their next incubation spell, the gains were 9.5 grams (in 5 days), 5.5 grams (7 days) and 8.5 grams (8 days) or 23 percent, 20 percent, and 12 percent of the weight of the birds at the end of the spell. Two of these increases were less per day than the daily losses during incu- bation, and it was unfortunate that the incubation spells of these returning birds were not known, as they may have been shorter than average. The weight of a returning bird did not increase, presumably after a minimum level, with the length of time spent away from the nest, suggesting that birds came back as soon as they had sufficient reserves to undertake another incubating spell. The average incubation period for 62 eggs was 42 days, with extremes of 39 and 51 days, much longer than the 38 days obtained indirectly by Allan (1962) but in line with the 43 days (39-48) for Oceanites oceanicus (Roberts, 1940), 41-42 days for Oceanodroma leucorhoa (Huntingdon in Palmer, 1962), and 41 days for H. pelagicus (Davis, 1957). In all these species, the incubation period may be prolonged because of eggs being left unincubated for several days at a time. As yet there appears to be only a single published record in a procellariiform of a repeat laying in the same season after the loss of an egg. This was in P. puffinus (Harris, 1966b) but Britton (personal communication) has now found another instance of repeat laying in this species. Huntington (in preparation) documents a case of repeat laying in O. leucorhoa. On Plaza the situation was complicated owing to competition for nest sites, 122 ( CALIFORNIA ACADEMY OF SCIENCES [PRoc. 4TH SER. 10 9 8 a = a 9 6 ~J = 5 © ue 3 2 | 6) | 2 3 4 0) 6 DAYS INCUBATION Ficure 9. Daily weight losses in incubating individuals of Oceanodroma castro. and in several instances two pairs were known to have laid in the same burrow during the same season. There were however six instances where repeat laying was a possibility but in no case was it certain. In the first instance, an egg was accidently broken when the female was in- cubating the day after an egg appeared in a burrow. Unfortunately the cloaca of the bird was not examined in detail but, as she had had a fully vascularized brood patch 20 days before, it is probable that she had laid this egg. This same female, with a swollen and distended cloaca, was found on a fresh egg a month later. This was the latest egg to be laid that season. No other bird was seen in the burrow. The second case was a female found with a vascularized brood patch and a broken egg a month before laying another egg. The third was of a male incubating an egg which was broken after 3 days’ incubation, almost certainly by a representative of P. /herminieri, which was also Vor. XXXVII] HARRIS: BIOLOGY OF GALAPAGOS STORM PETRELS 123 using the nest hole; the female (its mate from a previous breeding) was on another egg 6 weeks later, the last egg to be laid that season. No other bird was seen in the hole. The other cases refer to males, with fully developed brood patches, found on eggs which were almost immediately lost, and later on another egg, 27, 31, and 36 days respectively after the losses. There was no evidence to suggest that one of the birds early in the season had deserted and the remaining bird had to delay breeding while finding a new mate (Davis, 1957). It is impossible, however, to be sure that a nonresident had not visited the burrow and laid an egg which was then taken over by the resident pair, but the birds having had vascular brood patches very early (which on average occurred only 2 to 3 weeks prior to laying) sug- gested that some were probably true second eggs. Allan (1962) had similar oc- currences in two burrows but was unable to prove repeat layings. Two eggs were laid in two other holes but these were probably a result of 2 pairs. The numbers of birds found in a burrow did not affect the nesting success; this agrees with Allan’s (1962) suggestion that once ownership was established it was well nigh absolute. Nothing in the data suggested that females laid at the same date each year or that the nesting success of the pair using the site 6 months previously affected the date of laying. THE CHICK-STAGE Hatching was a very variable process, taking from 3 to about 7 days from the first denting of the shell. Davis (1957) suggested there was a changeover during chipping, but with short incubation spells as in H. pelagicus this is in- evitable. The only observations I have on the behavior of the adults near hatching were on those marked during incubation. These showed that there was no change- over prior to hatching, but once the chick emerged, changeovers were more fre- quent as the adults gave food to the young and probably depleted their own food reserves in the process. Many young were brooded for only 2 to 3 days, as com- pared to 6 in H. pelagicus (Davis, 1957), 2 to 4 in Pelagodroma marina (Rich- dale, 1943-44), and 1 or 2 in Oceanites oceanicus in a very cold climate (Roberts, 1940) ; it was rare to find an adult with the chick after this time. GROWTH OF THE YOUNG Growth of the young of many procellariiformes tends to be erratic, presum- ably due to scarcity and fluctuations in the available food, and the long fledging period has doubtless been evolved to cope with a food supply of this kind (Lack, 1948). If, however, a reasonable sample of chick weights are lumped, a uniform and typical growth pattern is found. This shows a steady increase to a maximum 124 ' CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. much higher than the adult weight, and then a gradual fall-off towards fledging. I have suggested (Harris, 1966b) that this large accumulation of fat reserves serves two complementary functions in allowing the adults to leave the colonies earlier than if they had to supply less food regularly but for a longer time and the chicks to have a better chance of surviving if the adults are forced to leave them due to food shortage. The former point is probably most important to a migratory species; the latter to some tropical species as P. /herminieri in Gala- pagos, which suffer from frequent food shortages (Harris, 1969). Unlike Allan (1962), I did not find it possible to readily separate the young into normal and retarded groups (though there were some of the latter so aptly described by him as ‘Mohawks’); the average growth curves for the three seasons (fig. 10) include all young which survived to an age when they might have fledged, and also young lost earlier due to predation. As far as possible, evening weighings have been used, as they were less affected by larger-than-average feeds, but for some instances when this was not possible, some morning weighings are also included. Any errors from the last cause are slight because the average decrease between morning and evening weighings was just under 2 grams. The growth curves for my three seasons and that given by Allan (1962) differ con- siderably, in fact the mid-1966 growth curve is far below that of “retarded” young on Ascension. These differences are also apparent in the maximum weights attained by the individual chicks. Surprisingly, these marked differences in growth are not correlated with corresponding differences in feeding frequency, weight of feeds, or nesting success. It is impossible to say if the “quality” of the food presented to the young varied from season to season, something which has yet to be demonstrated in any sea bird. Wing lengths of chicks were also measured and showed a slightly different pattern (fig. 11) in that the growth lines in the 1965-66 and mid-1966 seasons were almost identical despite dissimilar weight curves, whereas the 1966—67 chicks appear slightly advanced. I have too few data on actual fledging weights for any comparison between seasons. The fledging periods obtained varied in their accuracy, the majority having a possible error of + 2 days. Eight for the 1965-66 seasons averaged 69 days (range 60-72) and 11 in 1966-67 averaged 71 days (65-72). These from the hot seasons (average 70, standard deviation 4.0) were longer than the 58 days quoted by Allan (1962), but the 25 from the 1966 cold season (average 78, range 66-107, standard deviation 9.5) were longer still. Snow and Snow (1966 > Figure 10. The average growth curves of young of Oceanodroma castro on Plaza in the 1965-66 season (ten young), mid-1966 season (37 young), and 1966-67 season (16 young). The growth curves for young on Ascension are plotted from Allan (1962). VoL. XXXVIT] WEIGHT (g) 80 70 60 50 40 30 20 70 60 40 30 HARRIS: BIOLOGY OF GALAPAGOS STORM PETRELS 8 ASCENSION Vi o—o Normal young f e--e Retarded young I-5 HW-15) 21-25 31-35 41-45 51-55 61-65 7I- poet aT oe” : 5 a ee o @ @*. ee @ < pene aA oa ~4_ A +f Bo ye a4 A Rs Of ee “is Ui PLAZA GC: O---0 1965-66 js &----A Mid 1966 R e—e 1966-67 I-51) 21-25) 31-35 41-45 51-55 61-65 71-75 AGE IN DAYS 125 126 q CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. 150 3? / o--0 1965-66 Ye 140 e--e Mid 1966 | 9 +—+ 1966 -67 ones nm ul fo) Oo WING LENGTH (mm) ° I-S>) IRIS) 21-25 31-35 «41-45 51-55 6165 71-75 AGE IN DAYS FicurE 11. Average wing lengths of young of Oceanodroma castro in relation to age in the three seasons studied. and personal communication) gave four fledging periods for a cold season on Plaza as 86, 84, 83, and 81 days (all + 7 days), agreeing with my observations. The difference between the hot and cold seasons is significant at the 5 percent level. There is no desertion period as in some shearwaters (Lockley, 1930; Ser- venty, 1958; Richdale, 1963) as most birds were fed up until 2 nights before fledging. The observed times between the last feed and leaving the burrow were 1 night (seven cases), 2 nights (two), and at least 5 nights (two). Some of these young may not have actually fledged when last seen as occasionally almost- Vor. XXXVIT] HARRIS: BIOLOGY OF GALAPAGOS STORM PETRELS 127 TABLE 9. Feeding frequencies and average nightly increases in weight of young of Ocean- odroma castro. Average increase Season Month Baek — ae nae be ey 1965-66 January 36 23 64 6.8 February 30 22 73 eS March 9 4 44 8.4 Average 65 Ud mid-1966 July 112 67 60 57 August 56 31 55 6.2 September 70 36 51 6.0 October 10 3 30 4.7 Average 55 5.8 1966-67 January 16 7 44 6.6 February 65 24 37 6.7 March 61 32 52 4.9 April 18 7 39 3.9 Average 44 5.6 mid-1967 July 32 23 72 Dal fledged juveniles were found in other burrows or wandering about the colonies. Some of these wanderers were obviously half-starved, but others were doubtless on their way to fledging. FEEDING FREQUENCIES AND FEED WEIGHTS All easily accessible chicks in sheltered locations where accurate weighings were easily made, were weighed night and morning to obtain figures for feeding frequencies and average feed sizes (see table 9). In this table only increases in weight are taken as indicating a feed. This obviously overlooks small feeds which might not even make up for nightly losses due to metabolic processes. It is im- possible, however, to be sure of these feeds, or even if a feed has taken place, as the weight losses varied to some extent with the size of the previous feed, that is if a bird had a very large feed it would then lose weight very quickly. If any- thing above a loss of 2 grams was taken as indicating a feed, then the percentage of nights when a young was fed was 69 percent. The average feeding frequencies and feed sizes (by slightly varying techniques) for Pelagodroma marina were 72 percent and 6.4 grams (Richdale, 1943-44), and for H. pelagicus 83 percent and 6.4 grams (Davis, 1957). Both these species have shorter fledging periods than O. castro, probably because the food supplies were richer or nearer the colonies in these more temperate regions, so that the feed size, or the feeding frequency, or both, must be higher. 128 . CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. 80 70 60 50 40 WEIGHT (g) 30 20 Fi Io DEAD 8) 10 20 30 40 50 60 70 AGE IN DAYS Ficure 12. Growth curve of Oceanodroma castro young in an artificial set of twins in the 1965-66 season. EXPERIMENTS WITH TWINS Several small-scale experiments were carried out on the ability of this species to raise two young instead of the normal one. In the 1965-66 season a single pair were given an extra chick and, although it was fed at first, the introduced young did not grow normally and died after 41 days. The other young had a normal growth curve (fig. 12) and fledged at about 72 days. In the mid-1966 season, seven sets of twins were established and the results are set out below: 1. One young did not develop and died at 35 days; the other developed normally Vot.XXXVII] HARRIS: BIOLOGY OF GALAPAGOS STORM PETRELS 129 70 60 1966 (1) 50 ~ 40 & ~ ~ < 2 30 a Oy Ate 1966(3) Bee. “wae Gone 20 VOY | eee § & a Gone Dead ) 10 20 30 40 50 60 70 AGE IN DAYS Ficure 13. Growth curves of individual young of Oceanodroma castro in two sets of artificial twins in the mid-1966 season. Circles and solid lines refer to set 1 (see text) and tri- angles and dotted lines to set 3. (fig. 13) and fledged at 69 days; 2. One did not develop and died after 52 days; the other did not grow normally, in that it never reached the normal peak weight, but fledged above average weight at 87 days; 3. Neither developed very well and both died, at about 44 and 54 days (fig. 13); 4. One was lost at 45 days when weighing more than its foster sibling; remaining bird fledged; 5. One died after 12 days; the other was lost at 90 days when near fledging; 6. Both were lost within 10 days; 7. One was lost within 6 days; the other developed normally but did not fledge. In the 1966—67 season four more attempts were made but with similar results: 1. One did not develop and died after 30 days; the other fledged at 74 days; 2. Neither developed and both died at 45 and 51 days; 3. One died after 26 days; the other developed normally but was lost, probably to an iguana, after 45 days; 4. One died after 38 days; the other fledged at about 76 days. 130 ( CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. 160 140 120 100 WINGLENGTH (mm) 80 10 20, 50 40 50 60. 70 39se AGE IN DAYS Ficure 14. Increase in wing lengths with age in four successful and one unsuccessful Oceanodroma castro young from five sets of artificial twins. Also shown (dotted) are the average curves for normal young in the three seasons. The 12 pairs with 24 young succeeded in rearing 6 young, the same suc- cess per pair as with normal pairs. Of the young which did not fledge, four were lost (perhaps to predators) when developing normally, and the remainder died of starvation. This starvation was not due to the parents rejecting the chicks, but to one of the young becoming dominant and taking the majority of the food. It is significant that in the pairs where this did not occur, both young died. In only two instances were both young fed on the same night, possibly when the two adults returned together. The wing-growth curves of five of the twins which developed far enough to Vor. XXXVIT] HARRIS: BIOLOGY OF GALAPAGOS STORM PETRELS 131 grow primaries are shown in figure 14, along with the average for the controls in the three seasons. Only one, which did not fledge, shows a different slope from the controls, and three are almost identical with normal young; the fifth, which is much retarded, belonged to the mid-1966 pair 2; its growth was very abnormal. This last case indicates how very plastic the chick phase is in petrels. As with most experiments of this kind (Rice and Kenyon, 1962; Nelson, 1964; Harris, 1966b), the results are inconclusive in that the survival of the young immediately after fledging is not known. However, in the present cases the fledging weights were not different from those of single fledglings so that presumably their chances of survival to breed were as good as those of any other young. As with the previous experiments on shearwaters cited above, these sug- gest that there is not enough food available for the adults to raise more than a single young. NESTING SUCCESS Full details of nesting success (omitting experimental manipulations) are given in table 10. In the full three seasons studied, 60 percent of the eggs hatched and 50 percent of the young fledged, a low overall production of .3 young per pair but similar to the one season’s results at Ascension (Allan 1962). Most lost eggs just disappeared from the burrows (63 cases) but others were known to have been broken or dented (16), deserted (23), ejected by other petrels (13) or P. lherminieri (3), addled (13), and buried (4). The desertions appeared not to be due to my interference as many nests where adults were never disturbed by me were lost in this way; in six cases the egg was never incubated. Similarly most young were just noted as missing (31), but others died of starva- tion (6), were eaten by Asio galapagoensis (3) or Grapsus grapsus (3) or Ardea herodias (1), died due to sea spray (5), ejected by other storm petrels (1) or P. lherminieri (1), or just wandered from the nest site (1). Egg losses were evenly spread over the whole incubation period but about 40 percent of chick losses occurred within 10 days of hatching owing to a variety of causes, among which predation by crabs and accidental crushing by iguanas were probably important. PossIBsLE Factors INFLUENCING BREEDING SUCCESS 1. SEASON OF LAYING. There was no advantage in birds laying in either the hot or the cold season as far as fledging success was concerned. 2. DATE OF LAYING WITHIN A SEASON. This was important (table 10) as the proportion of eggs which gave rise to fledged young was highest early in the course of each breeding season studied. The decline was most marked in the sur- vival of young from hatching to fledging. The losses of eggs due to possible predation of the adults or failure to hatch because of being addled, did not increase as each season progressed. The causes of chick losses were mainly unknown, but starvation (as mea- 132 sf CALIFORNIA ACADEMY OF SCIENCES [PRoc. 4TH SER. Tasre 10. Breeding success of Oceanodroma castro im relation to season and date of lay- ing. Note that in the 1965-66 season the November egg success is too high as many nests were not found until December. Of ten other eggs laid before mid-January, one hatched but the young did not fledge. The mid-1967 figures show the range of nesting success after known losses are removed. Percent Third-month Eggs Young Per- Young Young Per- overall Season Month period laid _ hatched cent hatched fledged cent SUCCESS 1965-66 November 2 1 i 2Goo)a A 1 UuheaGe (78) 3 8 | 8 6 | December 1 7 5 | 4 0) | 2 14 7 , 61 7 1 13 8 3 5 4 4 1 January 1 2 2 | 67 2 0) 0 0) 2 1 | ) mid-1966 April 2 1 Gp tl NtGreao | #56 34 3 2 2)» 2 1 | May 1 11 6 | 3 1 2 34 22 , 64 15 8 58 37 3 33 PD | 18 12 June 1 12 6 | 6 2 | 2 7 3 , 41 3 1 33 14 3 3 (0) | 0) | July 1 6 3 50 3 (0) 0) 0) 1966-67 November 2 2 ge | =30 66 33 3 6 al 3 2 December 1 17 15 | 1 7 | 2 15 8 68 6 5 64 43 3 8 4 | 4 2 January 1 9 5 | 5 2 2 1 (0) 50 (0) 33 16 3 2 1 | 1 0 February 1 1 (0) | 0) 2 1 ) | mid-1967 April 3 1 0) 0) May 1 7 4 | 2 20 14-18 490-71 3 32 11-20 | June 1 17 5-14 | 2 9 1-7 19-79 3 7 O05 | July 1 A O=2 es eenitteeetee Vou. XXXVII] HARRIS: BIOLOGY OF GALAPAGOS STORM PETRELS 133 sured by weights of young) was unimportant and, to judge from the growth rates of the young, food was apparently uniformly available throughout the year. Predation was observed in only seven nests, but it might have been the reason for some other losses; however there was no correlation between the extent of possible predation and the date of hatching within any season. The very latest young may be evicted by the new pair returning to take over the nest, and some very small young may be lost through intruders, but the main causes of the de- cline of fledging success and its date are not apparent. It is possible, but unlikely, that purely social factors are important in that the birds breed more efficiently when other birds are also breeding. Against this however, Allan (1962) found that some out of season nesting produced at least third-grown young. 3. Foop sHorTAGE. This is hardly likely to affect hatching success unless severe enough to drive the birds from the colonies. Eggs were often found without an incubating bird, and usually it appeared that the bird due to take an incuba- tion spell was late returning, which certainly suggests it was hungry. Some eggs hatched after being left at least 6 days within the normal incubation period. However, the chances of an egg hatching decreased if left unattended; 116 of 163 eggs which were never seen without a bird (but on the average only checked 1 day in 3) hatched, the corresponding figures for eggs left on only 1 day were 24 out of 46 and for eggs left between 2 and 8 days the figures were 15 out of 51. Eggs laid in the hot seasons were more frequently left (11 percent of possible days for all birds) than those in the cold seasons (5 percent). There are three possible reasons: (a), the birds might be more willing to leave eggs during the warm season as there would be less chilling; (b), a food shortage might exist at this time of year; (c), more isolated birds may be more prone to leave the colonies, for some reason or other, during a time when there is less breeding activity. The reduced chances of eggs hatching due to being left unattended are probably the result of interference by nonbreeders or low enthusiasm in those parents. Associated with leaving eggs unincubated is the resistance to chilling of petrel embryos (Matthews, 1954). Some observations, made on eggs which came to hand, of several Galapagos sea birds, showed that resistance to chilling (table 11) was most pronounced in the storm petrels. One embryo of O. castro remained alive for 23 days without incubation, and a chick inside a chipping O. tethys egg continued to call for 16 days. Surprisingly two embryos of P. lherminieri were dead when first examined 10 days after the egg had been last incubated. Other species showed survival for up to 6 days, even in species such as F. minor, where there is little chance of an egg surviving predation if left uncovered for even a few minutes. Food shortage is doubtless important in determining the rate of growth of the young, and the twinning experiments suggest that there is not a super- abundance of food. However, it was extremely rare to find starving young, and 134 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47H Ser. TasBLE 11. Some observations on the ability of sea bird eggs to withstand chilling. “Less than 3” indicates that the embryo was dead when the egg was first opened 3 days after in- cubation ceased. Days after last incubation State of development Species embryo still showing movement of embryo Oceanodroma castro 6+- half developed 0-16 half developed 23 half developed Oceanodroma tethys 9-10 half developed 16 chipping egg Puffinus lherminieri less than 10 near hatching less than 10 third developed Phaethon aethereus less than 3 quarter developed less than 3 half developed less than 6 quarter developed less than 8 quarter developed 5-6 quarter developed Creagrus furcatus 3-5 three-quarters developed less than 3 quarter developed less than 3 half developed less than 4 third developed less than 5 quarter developed Fregata minor 5+ near hatching Bee three-quarters developed less than 3 three-quarters developed less than 3 three-quarters developed less than 3 three-quarters developed Sula sula less than 6 just started development less than 7 half developed those few could be due to one of the adults having died. Of the six young which died of starvation, one was very small, two were in holes where one of the adults did not reappear the next year so may have died during the first season, and two had both adults alive the next year. The parents of the other were not known. During my stay in the Galapagos, I saw evidence of severe food shortage in P. therminieri, Phaethon aethereus, and S. nebouxii, but never in O. castro. It ap- pears therefore that food shortage is not a common cause of chick losses. 4. PREDATION. Several large young were known to have been eaten by owls or herons and there were a few other cases in which this was suspected. Some few breeding adults may also have been killed by owls (see later). An indirect measure of the effect of predation can be obtained by comparing nesting success one year with the numbers of adults returning a full year later. The proportion of birds returning was much higher in those which had previously raised a chick (56 of 67 birds) than in those which had failed to hatch an egg (41 of 65) or raise a hatched chick (69 of 111). It is difficult here to determine Vor. XXXVII] HARRIS: BIOLOGY OF GALAPAGOS STORM PETRELS 135 cause and effect, but the faithfulness of adults to nest sites suggests that at least some observed differences were due to predation of adults at the colonies and not just to successful birds being more likely to retain the same nest sites. 5. CoMPETITION. Although some eggs (22 out of 72 known causes of failure) were lost due to competition for burrows, the numbers of birds caught in a bur- row during a season apparently had no effect on nesting success; of 183 eggs in burrows where no intruder was caught, 56 percent hatched and 25 percent of the young fledged; of 75 burrows where up to four intruders were found, 53 per- cent of eggs hatched and 21 percent of young fledged. As two eggs were found in a burrow very infrequently, it seems that once a pair has established itself, there is little chance of another pair laying in the hole the same season, although losses might still be caused by intruders and prospecting birds. 6. Nest sites. The nest sites were extremely varied but, apart from two very open and marginal sites where the adults may well have been killed by owls, it is difficult to see why the nest site should affect the nesting success. There was, however, a markedly high-nesting success in the most frequently used burrows. Of the 57 burrows where observations were carried out in all four seasons, 24 (42 percent) were used all four seasons and had an overall success rate of 40 percent as against 13 percent for all other layings. This difference was due almost entirely to the varying fledging success (65 percent to 23 percent) and not to any factor which might prevent the eggs from rolling out or being dented. There were different pairs involved in at least two of the three seasons, and also a few other changes due to mortality. There was no tendency for birds in these burrows to nest earlier than average. Indeed in all burrows there was no correla- tion between laying date and the success or fledging date from the immediately prior season. One is forced to the conclusion that these nest sites were in some way more attractive to the more efficient, perhaps older or more experienced, adults, and not to the actual physical potentialities of the burrows. 7. Socray Factors. Allan (1962) concluded that on Ascension in the year he studied that “no factor other than the behavior of the petrels during the breeding season was obviously a major cause of loss of eggs or young.”’ My more detailed study has explained many more of the losses, but there is still a substantial num- ber unaccounted for, especially of eggs and small young. Some losses were directly attributable to other birds, and I must agree with Allan in that disturbance due to intraspecific competition is an important source of loss. In O. tethys (later) 7 intraspecific competition was responsible for almost all the observed egg losses. 8. INTERSPECIFIC COMPETITION. There is some little conflict with P. lhermi- nieri but this is not severe enough to be important. On Tower, Nelson (1966) thought that interspecific competition with O. tetiys was an important source of egg loss of these two species. He was however mistaken as the species do not nest in the same area. 136 , CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. On Isla Pitt however interspecific competition is important as there the large colony of O. tethys apparently prevents O. castro from breeding during the cold season by occupying all the available nest sites. At this time O. castro does visit the island, but no nests were found even in sites used for breeding during the hot season. Mott Primary molt in sea-bird populations has, with a few important exceptions, been little studied, but it is known that it is extremely variable with regards to its timing in relation to the breeding season. Within the procellariiformes, some shearwaters, including P. tenuirostris (Marshall and Serventy, 1956) and P. gravis (Bourne im Palmer, 1962), delay their wing molt until they have migrated the long distance to the nonbreeding grounds. The same applies to some storm petrels including Oceanities oceanicus (Murphy, 1918) and Pelagodroma marina (Bourne in Palmer, 1962). Puffinus puffinus is an interesting species as the British race P. p. puffinus does not molt at the breeding grounds (personal observations) and presumably must molt in its winter quarters off South America, as the birds migrate immediately after the breeding season. The Mediterranean race P. p. mauretanicus, which has no extensive migration, molts immediately after it has finished breeding (Mayaud, 1931). Other species of the order may start the primary molt when feeding young, that is, H. pelagicus (personal observations) and the giant petrel (Macronectes giganteus) (Warham, 1962). The four species of oceanic terns breeding on Ascension and studied by Ash- mole (1962 and 1963), Dorward (1963), and Dorward and Ashmole (1963), show interesting variations in the primary molt. Sterna fuscata and Anous tenuirostris both usually finished their molt before returning to breed but some may then start the next cycle before the young are fully grown. The fairy tern (Gygis alba) was never found molting primaries when incubating or feeding young chicks and must therefore have completed the molt between breeding cycles. In this species, the wing molt is irregular with primaries in different parts of the wing being replaced at the same time. Anous stolidus molted during the breeding cycle as it also does in Galapagos (personal observations). MOoLtiInc As far as possible all nonincubating birds were examined for primary molt, and scattered observations were also made on body and tail feathers. In the following account the primaries are numbered in the standard way from the inside (shown on the left of the diagrammatic formulae) outwards, and the stages of molt are represented as 1 = empty socket or pin feather, 2 = vane up to one-third its final length, 3 = vane between one- and two-thirds grown, 4 = vane VoL. XXXVIT] HARRIS: BIOLOGY OF GALAPAGOS STORM PETRELS 137 two-thirds to full grown, N = new feather without any blood in calamus, 0 = old feather (Ashmole, 1962). The outermost, minute primary is ignored. The start of the primary molt of the breeding adults which is from the inner- most primary outwards is extremely variable, but appears never to occur before the young is well grown and some birds do not start until their young have fledged. One adult with an almost-fledged young had the two inner primaries on each wing a third grown on 5 April. On the same date, when only adults feed- ing young would be expected to be visiting the colonies, netting produced two molting adults — Zoe OO 8 0 0 0 O Mes OO 0 OO oO 0 0 and four with exceptionally worn plumage but with only body molt. Two males out of 10 birds netted at Tower on 16 February had one and two primaries respectively just starting regrowing in each wing. Both these birds were carrying food and so presumably were feeding young. Non- and failed breeders may undergo a body molt when at the colony but not a wing molt. The only possible exception was a bird caught on 13 February in a hole where it had not bred, which had its inner primary in pin. Almost all the primary molt occurs when the birds are away from the colonies, 6 to 7 months for successful breeders, but some newly arrived birds returning at the start of a breeding season have the outermost primary still growing. One bird had the fourth secondary half grown, the fifth old, and the remainder new, two growing rectrices and feather replacement of the upper parts. This bird laid exactly a month after this examination. It seems, therefore, that the primary molt of the population and certainly of some individuals, takes the whole time between breeding attempts and it might be, as suggested for S. fuscata (Ashmole, 1963), that the birds are prevented from breeding more frequently by the necessity of the whole population fitting in this molt, that is, the molt governs the intervals between breeding. However some individuals of Puffinus lherminieri, with more wing feathers to replace, are able to undergo a complete molt and breed again in 5 months after the end of a previous breeding attempt. Even allowing for the differences in feeding habits, this suggests that a storm petrel just might be able to complete its molt in a similar time. Possibly this elongated molt is adapted so as to spread any ineffi- ciency it brings about over the longest possible time; and there is some other fac- tor responsible for the timing of the breeding cycle. A third possibility is that this molt pattern was primarily evolved in an annual breeding, oceanic species (as O. castro on Ascension), as it is reasonable to suppose that a nonannual cycle has come from an annual, where the birds had to roam widely for food and could not hope regularly to find a rich feeding ground where rapid molt could be safely undertaken. In this case again it would be the molt preventing more frequent 138 ’ CALIFORNIA ACADEMY OF SCIENCES [PRoc. 4TH SER. breeding. In Galapagos, with two populations breeding 6 months out of phase, there is probably little advantage in the molt being shortened even if this were possible. Birds with body and/or tail molt were found throughout the breeding season. There was no orderly replacement of tail feathers and it was often difficult to classify individual rectrices as old or new. Unfortunately few specimens of O. castro have been collected away from the breeding colonies and only two from near the Galapagos show any primary molt. They are 11 April, near Galapagos N N N. N. NN N NGN@See 18 June, off Cocos Island N N-N.N NN. NOOR It is not known if O. castro breeds on Cocos Island (about 500 miles from Galapagos) but this bird could possibly be an immature from the Galapagos or Hawaiian populations. This sequence of molt is similar to that of birds on Ascension (Allan, 1962) and the few skins I have examined from other Atlantic colonies which show any primary replacements: 28 June, Cape Verde Islands N -N N N N N NSU 12 June, Madeira NN N N N N NON 15 June, Madeira N N N N N N WN 12 Sept., Madeira N 1 0 0 0 O (‘O23GR enero suggest that the molt pattern found in Galapagos is universal in this species. The mallophaga found on O. castro included Halipeurus pelagicus and Philoceanus species. An undescribed flea Parapsyllus species was found on both O. castro and O. tethys, this genus of flea is associated with sea birds in the Southern Hemi- sphere. PREDATION Oceanodroma castro in the Galapagos is heavily preyed on by Asio gala- pagoensis and counts of fresh prey remains on Plaza (table 12) showed that predation was greatest during the cold season. These peaks may be slightly re- tarded as some remains were 2 to 3 weeks old when found. Unfortunately remains often consisted of a pair of wings so that it was impossible to calculate the pro- portion of ringed to unringed birds. This was quite small, however as only a single ringed bird (a recently fledged juvenile) was found killed by an owl. Many owl pellets were collected on North and South Plaza during the study and consisted of remains of O. castro (547 pellets), P. lherminieri (51), Creagrus furcatus or Phaethon aethereus (16), Rattus rattus (14), finches (4), and insects (three entirely and they were present in five others). Among these a single ring was dis- covered, that from a breeding member O. castro, suggesting that birds regularly frequenting burrows (many ringed), were not so likely to be caught as nonbreed- Vot.XXXVII] HARRIS: BIOLOGY OF GALAPAGOS STORM PETRELS 139 TaBLe 12. Monthly incidence of kills of Asio galapagoensis found on Plaza. The figures in brackets are definite juveniles and are included in the main totals. At the start of the study the following older remains were found: O. castro (21), P. lherminieri (25), and during the study an additional 39 remains of O. castro and 74 of P. lherminieri which were too old when found to be classified as to month. One individual of Puffinus lherminieri weighs about the same as 41% of O. castro. Number of remains Oceanodroma Puffinus Year Month castra lherminieri Others 1965 December 36 15 1966 January 12 8 February 4 2 March 0 2 April 1 1 May 10 (0) June 23 0) July 81 2 August 25 6 Creagrus furcatus [2] September 1 {LS toy isi Lizard [2], P. pacificus [1] October 6 [6] 25 [6] Heteroscelus incanum [1] November 9 [6] 410) [Pile] December 5 ila 12 [6] Squatarola squatarola (1) 1967 January 15 19 [10] C. furcatus (1), Phaethon February 3 3 acthereus March 0 Slelal April 2 12] 0 May P| 1 June 21 0 July (1st week) 16 (6) ers (few ringed) which spent more time actually on the ground looking for holes. In P. puffinus it has been shown (Harris, 1966a) that these nonbreeders are much more likely to be caught on the surface than breeders, which are either in the burrow, and therefore safe from most predators, or actually going into or leaving the burrow. The owls did not breed, or roost at all frequently, on South Plaza during the study but were always present on North Plaza and were frequently seen at dusk and dawn flying between the islands. Indeed they may well have bred there in 1966, but it is extremely unlikely that they did so in 1967 when regular checks were made. There was no evidence that more than one pair of owls was involved in this predation. This pattern of kills found need not fit exactly with the predation, for, if the owls had unfledged young, they would take the intact prey back to the nest. When the young had fledged they could well follow the parents to South Plaza and the plucked remains of kills would then be found. It must be stressed that there is no 140 t CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. evidence for this speculation, as no birds were heard to give the typical food- begging calls. Although the numbers of eaten specimens of O. castro found fit very closely the number of eggs and young in the petrel colony, it does not follow the pattern of nonbreeders visiting the holes. The number of nonbreeders was highest just before and during the egg-laying period and then decreased before the majority of the young hatched. Indeed, by July and February there were very few non- breeders or failed breeders in the holes and probably very few visiting at night. It is therefore difficult to explain the peak of corpses in July, 1966, except that the owls may have been feeding young. Asio galapagoensis in Galapagos may breed in many months of the year (Lévé- que, 1964) so, although predation could obviously shorten the time that non- breeder petrels spend at the colonies, it could hardly affect the timing of the returning birds unless by coming back at the same time as the breeders, they “swamp” the predators and so reduce the chance of any individual petrel being killed. When there are few petrels in the colonies, the owls prey on other sea birds, especially P. lherminieri, but given the choice they appear to prefer to take storm petrels. The prey spectrum is wide and includes the introduced Rattus rattus from the mainland of Santa Cruz and migrant waders so that it is only on rare instances that the species on Plaza can be really short of food. During the study there were two periods when food may have been short for owls, February and March in both 1966 and 1967, when there were only a few petrels, mainly breeders feeding young frequenting the colonies, therefore unlikely to be caught on the surface; there were also very few shearwaters (Harris, 1969). In 1967 the absence of shearwaters was much longer than in 1966 and could have prevented the owls from breeding. In 1966 it might have delayed breeding so that the owls could lay only in April when the storm petrels and shearwaters returned. Then they would have missed the peak of storm petrels at the colonies and preyed on the nonbreeders which were not frequenting burrows but still prospecting for holes. From other studies on shearwaters (Serventy, 1967; Harris, 1966a) it seems that these might be younger than those which returned with the breeders early in the season. SURVIVAL OF ADULTS There has never been a satisfactory direct estimate of the annual mortality of a storm petrel species, and indeed the difficulties of obtaining one may be insur- mountable. Richdale (1963) has suggested on the basis, as he admits, of inade- quate data, an annual mortality of 45 percent for Pelagodroma marina which is obviously impossible for a bird having a single egg clutch, fairly low nesting suc- cess, and deferred maturity. For H. pelagicus, Davis (1957) had at least 60 out of Vor. XXXVII] HARRIS: BIOLOGY OF GALAPAGOS STORM PETRELS 141 74 breeding adults surviving overwinter, an 81 percent survival, which is again too low to allow the species to keep its population steady. Lack (1966) suggested a 7 percent annual mortality for this species but only used the successful breed- ers, which could be a biased sample as some adults may well have died at the colonies, and the period over which the measurement was made was less than 12 months. However to judge from other birds with a single egg clutch and de- ferred maturity, for example the royal albatross (Diomedea epomorpha) with a 3 percent annual mortality (Lack, 1954) and P. puffinus with 6 percent (Har- ris, 1966a) it should be in this region. In the 1966-67 hot season I retrapped 71 percent of breeding adults ringed in the 1965—66 season, in the mid-1967 season 65 percent of those from the mid-1966 season. These survival rates were too low to allow the population to remain stable. I know that I missed some adults in my study burrows and many more must have moved, perhaps only a few feet, into burrows where I could not find or reach them. The estimate of .3 young raised a pair means that 100 adults would produce 15 young to fledging every year, and even if all these survived to breed, the adult mortality could not exceed 15 percent, if the population was to remain constant. Unfortunately we have no data on the age of first breeding in this species. Allan (1962) using rather inadequate data calculated that four seasons are passed be- fore maturity is reached. At least one representative of O. leucorhoa (Gross, 1947) and one of H. pelagicus (a bird ringed by me and recovered by D. Scott) are known to have bred at 3 years, but the average age of first breeding is probably much older. Huntingdon (personal communication), working at the same colony as Gross, has found one individual of O. leucorhoa breeding at 4 years and four at 5 years. As it seems likely that birds do not breed until their third year or later and probably have a postfledging annual mortality higher than that of the adults, one would suspect that the annual mortality could not be higher than 5 to 7 percent. The causes of mortality of adults are varied. In the colonies I found five birds dead and jammed in holes (one hole had two dead birds wedged in the entrance), a female apparently egg-bound, two died after getting their wingtips caught in thorny bushes, and one was found badly pecked (? by a frigate). These were in addition to all those killed by owls. A Galapagos hawk (Buteo galapagoensis) was also reported (Dr. U. Eliasson) as killing an injured storm petrel (? species). Ritchie (1966) recorded an adult of Oceanites oceanicus as taken by a shark as it pattered along the water. DISCUSSION OF THE BREEDING SEASON It was shown earlier that there were two entirely separate populations of storm petrels nesting in the same nest holes approximately 6 months out of phase with each other, a situation which has not yet been described in any other bird. 142 d CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. It is also known that the seas around the Galapagos have a well-marked seasonal water temperature fluctuation, though my fortnightly surface-plankton sampling failed to show any associated differences. This latter point is however inconclu- sive because of the plankton being extremely patchy and the fact that the sam- pling may have been inadequate to pick up any changes. There is also the possibility that the birds may have been feeding in waters which bear little or no relationship to changes detected in the inshore waters of the Galapagos. Although there were a few marked differences between the various seasons studied, for instance in growth curves of the young, these may only show how variable are the conditions under which the species breeds. The only differences between the hot and cold seasons were egg diameters (not significant), fledging periods (70 days in the hot as compared to 78 days in the cold) and the fact that approximately half as many birds breed during the hot season as in the cold. The latter points might seem to be irreconcilable in that a longer fledging period suggests less available food, but it is at this time that the majority of birds are choosing to breed. However, this could be due to depletion of the food supply by the greater number of birds breeding, though this is less likely in forms like storm petrels which eat surface food. Nevertheless, any disadvantage to the young might be overcome by the advantage to the adults by reducing the proportion of birds taken by predation. Be this as it may, there was little difference between the hot and cold seasons studied. In O. castro on Ascension (Allan, 1962) and O. tethys on Tower (later), a few eggs were laid out of the normal breeding season. If out-of-season eggs were successful, this could quickly give rise to a situation in which birds were breeding throughout the year. Alternately, if there were some factors to separate them from the normal season, they might result in the situation found in O. castro. The latter situation assumes that it is advantageous for each individual to be syn- chronized in its breeding with others, and that it is incapable of breeding less than annually. Perhaps the timing of the molt is crucial, as it might be advanta- geous for this to be spread over the longest possible time. Even so, it is surprising that the earliest failed breeders should not return to relay until a year after their previous egg. Bourne (1957) has postulated that in Madeira, O. castro and Pterodroma mollis might each have twice colonized the island from different areas and the two populations now breed at different times of year. This might be true of P. mollis where the two populations are slightly morphologically different, but we have too little information on O. castro to make an evaluation. However, this is unlikely to have happened in Galapagos as the two nearest populations breed at similar times. The difficulty of explaining the two cycles is not so much how they may have come about, but how they remain separate. In every month there were some Vor.XXXVIT] HARRIS: BIOLOGY OF GALAPAGOS STORM PETRELS 143 birds producing eggs or feeding young, both of which activities require large quantities of food, which implies that food is available in every month of the year. It is conceivable that with the movement north and south of the Humboldt Current, the optimum feeding conditions could occur every 6 months or so, but this is contrary to the available evidence. If food is in fact uniformly available, why is not O. castro breeding throughout the year, like some other Galapagos species such as C. furcatus, P. lherminieri, Phaethon aethereus, S. sula, and S. nebouxii. , The detailed studies on tropic birds on Ascension by Stonehouse (1962) and the observations made in Galapagos by Snow (1965) and myself, show that in these continually breeding hole-nesting species, competition for holes resulted in heavy losses. For a storm petrel, which rarely lays a replacement egg, this would be a wasteful process, and the present situation allows a large number of birds to utilize an apparently limited number of the most suitable nest sites. But how this might be regulated by natural selection, and how it prevents any individual pair from attempting to nest away from the peak times, is not at all clear. Pre- sumably there is some advantage to a pair in nesting when the majority of in- dividuals do so, and that this is so great that it more than compensates for losses due to the additional competition for food and nest sites within each of the two seasons. Predation by Asio galapagoensis could well be one of the important factors in bringing about synchronization of breeding, as was predation on S. fuscata by cats and frigate birds on Ascension (Ashmole, 1963). The decline in breeding success within each season appeared not to be due to food shortage and is too marked to be explained by the death of the adults, so provides another puzzle. The sharp beginning of the breeding season is also difficult to explain, unless most individuals cannot breed much quicker and those which try cannot find mates, or are heavily preyed upon, or suffer heavy losses when the majority of the birds return. Whatever the selection forces involved, they are at present highly obscure. The present situation is probably the most economical for the species in this habitat, because a well-synchronized breeding season, brought about by display flighting and calling, might well help to reduce losses due to predation and conflicts between adjoining seasons, and so increase the chances of the individual producing young. OCEANODROMA TETHYS Unlike the previous species, O. tethys is normally restricted to the Humboldt Current, though a few birds have been recorded as far north as southern Cali- fornia (Murphy, 1936). The species has been divided into two subspecies ‘elsalli’ breeding in Peru and ‘tethys’ in Galapagos (Lowe, 1925). The Galapagos race is bigger in all di- 144 ‘ CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. ROCA REDONDA a ——— (TOWER) GENOVESA a ISEA PITT : Be 92° 97° SS go° Ficure 15. Colonies of Oceanodroma tethys in the Galapagos. Breeding has not been proved on Roca Redonda. mensions (100 adults measured had wings averaging 136.1 mm., standard devia- tion 3.4) than the other race (30 taken off South America averaged 124.6, stan- dard deviation 2.8). The few large individuals collected off the continent (wings 129, 129, 130, and 131 mm.) might in fact have belonged to O. t. tethys as it is likely that the two populations mix off Ecuador and Colombia. The species is very common at sea in Galapagos waters and nests on Tower Island and Isla Pitt, an islet off the eastern tip of San Cristébal (fig. 15). Sr. Miguel Castro told me that in March, 1966, he saw this species flying around Roca Redonda off the north of Isabela and it could well nest there too. Vor. XXXVII] HARRIS: BIOLOGY OF GALAPAGOS STORM PETRELS 145 3 : Se anak FicureE 16. Flighting in Oceanodroma tethys on Tower Island. Comparatively few of the birds nest in these cliffs but they are the focal point of flighting during the nonbreeding season (when this was taken). The extensive white rump patch and the lighter wing coverts can be seen on some of the birds (top left). Also in the photograph is a single individual of Phaethon aethereus and some nesting and roosting ones of Creagrus furcatus. The colony on the southeast coast of Tower has an extremely large popula- tion in two very different habitats, cliff and its adjoining lava field. The cliffs here are composed of lava flows a foot or so thick tiered like layers of a cake to a height of 50 feet (fig. 16). Some members of O. tethys nest in this area along with large numbers of Phaethon aethereus, Puffinus lherminieri, C. furcatus, and possibly the bulk of the island’s population of O. castro. Although the cliffs overhang and were frequently soaked in spray, the cracks are so deep that many nest sites are available to the birds. In spite of the fact that at some times of the year it appeared otherwise, the vast majority of petrels nested in a bare lava field stretching half-a-mile along the cliff top and a hundred yards inland (fig. 17). Inland the colony was bounded by a deep fissure, the other side of which was scrub composed of Cordia lutea, Croton scouleri, and Bursera graveolens, the nesting ground of large numbers of Sula sula, S. dactylatra, and Fregata minor. In the lava plain were a few stunted bushes and the cactus Brachycereus species. The lava was fissured and raised in bubbles which gave the birds access to a subterranean maze of passages from a few inches to a foot high; the petrels shared these with about 50 pairs of Phaethon aethereus, a few pairs of Puffinus lherminieri, and the Galapagos dove (Veso- 146 : CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 17. Mist-netting Oceanodroma tethys on Tower early in the morning in the non- breeding season before too many birds had arrived at the colony. In the foreground is the edge of the flat lava field under which these storm petrels nest. pelia galapagoensis). The finches G. magnirostris and G. conirostris, the mock- ingbird (Nesomimus parvulus), and the owl Asio galapagoensis regularly fed here, the last on sea birds. Nelson (1966) reported that the owls on Tower preyed on both O. tethys and O. castro, that there were two different calls heard from the burrows, and that there were two species of storm petrels flighting together over this colony. He therefore cast doubt on November observations by Castro, who thought that only O. tethys was involved, as it was based solely on sight observations. The owls do indeed prey on both species of Oceanodroma but catch representatives of O. castro when they come ashore at night. The two calls both refer to O. tethys (see later). Dr. D. Snow (in his notes at the Charles Darwin Research Station) during a visit to the colony with Nelson, saw only a single species flighting, O. tethys. There can be no doubt that only a single species is diurnal on Tower, and that this is O. tethys. Isla Pitt is an islet of crumbly lava about 50 yards across and 100 feet high, in part with bushes of Cryptocarpus, Malvastrum, and Periloba galapagensis on which nested Fregata minor and S. sula. Every Periloba bush had a pair of frigates nesting and their guano transformed all but the few topmost twigs into a solid mass, allowing the petrels to nest underneath. Numbers of seabirds also Vor. XXXVII] HARRIS: BIOLOGY OF GALAPAGOS STORM PETRELS 147 nested in the cliffs. During the warm season, O. castro bred on the island, but when O. tethys returned to breed in the cold season it appeared to oust O. castro entirely by sheer force of numbers. There was intense competition for nest sites as every available nook and cranny was occupied by several pairs of O. tethys. Foop I examined the stomachs of 13 adults and 3 young and the food regurgitated by approximately 50 netted birds. Fish (in thirteen stomachs and 27 regurgita- tions) was the commonest food followed by cephalopods (seven stomachs and 4 regurgitations), and crustacea (three stomachs and 6 regurgitations). Seven- teen fish ranged from 24 to 36 mm. long and averaged 28 mm. Almost all birds regurgitated large quantities of red oil which was probably stained by pigment from red planktonic crustacea. The species is normally a nocturnal feeder and birds caught early in the morning normally had undigested food. Many of the regurgitated fish had pro- portionately large eyes, suggesting that they may visit the surface only at night. Occasionally I have seen these birds feeding by day, when they dip or dart sideways to pick food from the surface of the sea. Once about 30 birds fed under the cliffs at Isla Pitt where a sea lion was killing a fish. Some birds then pattered on the surface of the water like species of Oceanites but this was atypical. This apparently is the only storm petrel which normally flies at its colonies by day and feeds at night. This is not due to the lack of predators so is pre- sumably adapted to allow the birds to exploit a rich nocturnal food resource. Several authors (for instance Murphy, 1936) have suggested that other storm petrels feed by day and night, but there is little direct evidence of this. There appear to be extremely few sea birds known to feed at night, Sterna fuscata (Bruyns and Voous, 1965), P. pacificus (Gould, 1967), Sula sula (Murphy, 1936), and possibly Fregata species (personal observation), but this mainly occurs on nights with a full moon. It is therefore surprising that Galapagos should have two species, O. tethys and C. furcatus, which feed at night, the latter species entirely so. There must be much food available to nocturnal feeders which is unavailable to diurnal species and more intensive observations will doubtless show up more species adapted to this niche. BREEDING CYCLE ON TOWER The evidence suggests that there is an annual breeding cycle on Tower, most of the eggs being laid in May and June, but birds visit the colony at other times and rarely lay eggs then. The colony was first visited 15-17 February 1966, when the numbers of birds present was so great as to be almost unbelievable; although it was impossible 148 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Taste 13. Numbers of Oceanodroma tethys storm petrels with brood patches in various stages of development in three samples caught on Tower. 16.2.66 17.1.67 8.3.67 State of brood patch Netted In burrows Netted In burrows Netted Not present 3 33 1 15 Half defeathered 7 1 10 Unvascularized 33 De, 57 2 28 Half vascularized 1 1 5 Vascularized 1* Regrowing feathers 10 1 26 Total birds 54 25 106 3 69 * Female with egg about to be laid. to get any worthwhile estimate of numbers, several observers agreed that there must have been several hundreds of thousands of birds flighting. The bulk of the birds were concentrated at the cliff edge, where they flew time and again near or actually into the cliffs. A much smaller number flew high over the lava and ex- tremely few dipped low as though prospecting for holes, but none actually landed in this area. Birds which did swoop down only circled once or twice before returning to the cliff edge or to the throng about 20 feet up. Many birds were present in holes in the cliff and in the base boulders, and any observer watching would probably think, as did Lévéque (1964) and myself, that the colony was centered on the cliffs. This impression was strengthened by finding two slightly incubated eggs, one in a small hole in the cliff, the other under a lava slab at the cliff edge. Despite a thorough search, no other evidence for breeding was ob- tained and of 54 birds caught (table 13), only one had any trace of blood vessels present in the brood patch. Three males and five females had undeveloped go- nads. A further visit on 27 February failed to produce any other evidence of breeding. At this time no birds were present in the burrows at night. On my return to the area 18-20 April, I confidently anticipated that breed- ing would have begun. However no eggs were found, but the pattern of flighting had altered and was now centered around the more inland flatter areas, where many birds were entering holes and even staying overnight. The numbers of birds present were reduced to perhaps a quarter of those in February, a decrease most marked at the cliff edge. It was not due to many of the birds having by then taken over holes, for whereas in February many hundreds of birds had been under the lava, a thorough search produced but one. By the last week in June there were many small young and eggs present, some of them newly laid. No birds were seen breeding in the cliffs, but some may have done so in the top few ledges, which were difficult of access. Whereas the number of birds present in June was similar to that in April, Vor. XXXVIT] HARRIS: BIOLOGY OF GALAPAGOS STORM PETRELS 149 by mid-July there were many fewer and these were going directly into and out of the holes, with little circling. It appeared as though at least some of the non- breeders and some failed breeders may have left the colony. By 17-18 August there were yet fewer inland but as many flighting at the edge. Unfortunately transport difficulties prevented any more visits during this breeding cycle but it is reasonable to suppose that it was coming to an end, as there was little display or fresh breeding activities. Birds are known to be pres- ent at the colony in small numbers in September (Loomis, 1918), October (Lévéque, 1964), and November (Brosset, 1963; M. Castro personal communi- cation). On 13-14 December 1966 large numbers of birds were again present but only near the cliff edge. A single female examined had an undeveloped ovary and unvascular brood-patch. No birds were seen in wing molt and there was no evidence of breeding. A similar situation was found 16-19 January 1967, except that a few birds were showing an interest in the inland lava. Of 106 birds netted and 25 taken from burrows, the majority had unvascularized brood patches (table 13). One female was caught with an egg about to be laid and another had recently laid; both these birds had only slightly vascularized brood patches. No birds were in wing molt but many of those without brood patches were regrowing body feathers. A fresh egg was found on 8 March, in a hole where an egg had been laid in February, 1966, suggesting that the same bird or birds may have been involved in both seasons. Twenty-six out of 69 birds netted were now refeathering brood patches, 11 were in body molt, and two had just started the replacement of the inner primaries. A short visit in July showed many young and some few eggs. One breeding adult had also bred a year previously, showing that at least some individuals have an annual breeding cycle. These data on out-of-season breeding are tantalizing, as they suggest that a very large number of birds come to the colonies out of the normal breeding season and go through many of the pre-breeding activities, but that only a minute proportion actually breed, and these in places not used in the normal season. It would seem that these birds present in the hot season were nonbreeders, as breeders from the cold season would be molting their primaries. To get into phase with the normal breeders, these hot-season birds would have to adjust or interrupt their molt for breeding (Ashmole, 1965). There is no evidence for an interrupted molt in O. fethys. Another possibility is that, as in O. castro, there are two populations breed- ing at different times of year, but for some reason hardly any of the population present during the hot season bred during January to April, 1966, or December, 1966 to March, 1967. It is at present impossible to give definite evidence to 150 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. prove or disprove this. Beck (in Loomis, 1918) found 200-300 petrels present and a single addled egg on 15 September, 1906 and collected a single bird with medium-sized gonads. Beck (1902) had previously seen several thousand birds but not eggs in the hot season of 1902, presumably March (Rothschild & Hartert, 1902). The notes of Lévéque (1964) for his visits in March and October, 1961, suggest that breeding was not taking place during those two hot seasons. Against these observations are those of Brosset (1963) which indicated that 11 birds caught in November, 1962, were physiologically ready for breeding. In view of the available evidence, I conclude that the species normally has an annual breeding cycle centered on the cold season with only extremely few breeding outside this time, but this needs checking by further observations. BREEDING CYCLE ON IsLA PITT This colony was discovered by Lévéque (1964) in June, 1961, when eggs but no young were found. Castro (personal communication) has revisited the colony in June and found a similar state of breeding. I visited the islet four times. On 19-20 December 1965 the colony was completely deserted and on 16 April 1966 laying had not begun, although several thousand birds were present. By 10-11 July the majority of eggs had hatched. The next visit was not until 26-28 May 1967 when most of the birds had recently laid. These observations indicate that there is an annual breeding cycle with the birds deserting the island during the nonbreeding season, and that the dates of breeding were very similar to those on Tower. THe DiurRNAL CYCLE The birds coming to land showed a very well-marked diurnal cycle in both colonies and at all visits. The first birds usually reached land just before sun- rise and the numbers increased to a peak in mid-morning before gradually de- creasing during the afternoon. The last birds flying about left at dusk. One bird was caught in a mist net at 1900 hours but this may have been spending the night in a burrow and been scared out by my activities. Nelson (1966) was mistaken in thinking that there were no marked daily or seasonal peaks in the numbers of birds visiting land. Considerable variations in the diurnal pattern did occur both with this month and with the varying weather conditions. On 27 February when, unlike most visits, the sky was exceptionally overcast, very few birds were ashore by 0745 hours. However the numbers soon increased and by 0930 hours the cloud of birds was visible a mile away. The morning peak is most marked prior to the breeding season and as the season advances birds spend more time at the colony so that the peak is less obvious. By the time the nonbreeders and failed breeders leave the peak is very small. Vor. XXXVIT] HARRIS: BIOLOGY OF GALAPAGOS STORM PETRELS 151 BEHAVIOR AT THE COLONY As so few observations have been made on the behavior of storm petrels, be- cause they are mainly nocturnal, it has seemed desirable to quote my few on this diurnal species in detail. The flight behavior was watched closest on Tower but the open nature of some of the nest sites on Isla Pitt allowed closer observa- tions of some burrow behavior. FLIGHT BEHAVIOR Nelson (1966) made three generalizations on the aerial display. (A), indi- viduals took part in the flighting for some time; (B), no two individuals were keeping together as in a courtship display; and (C), a few individuals descended repeatedly to the same piece of lava, pattered over the ground with raised wings, at the same time running their beaks over the lava. My observations agree with these and, like Brosset (1963), I was impressed with the relative lack of noise even though some birds did call “tchzz-te-tchzz” in flight and there were fre- quent aerial collisions. The general flight was slightly bouncing and tern-like and many birds spent much time doing nothing else but flying about. The flight at the cliff-edge was slightly different as birds hover, with raised wings and usually with tail spread, in the up-currents, or they prospect the topmost cliff holes, before either drop- ping away for another circuit or being blown upwards. The birds actually visiting the cliff have the rump patch very conspicuously displayed. When the birds were visiting the inland areas at the start of the breeding cycle, each bird was acting alone in its flighting. One bird when watched for 10 minutes flew in a circle of approximately 30 yards diameter and landed at six different holes before actually entering one and remaining there. In other in- stances birds were known to have repeatedly landed at the same hole before entering or flying away. Another common action was for birds to pause in flight as though looking for a hole in the lava. This was sometimes associated with birds calling from underground and at least once a bird called in flight and was an- swered by a bird from a burrow. Frequently birds landed in a completely hole- less area, which however had birds under the lava flow, and these too may have been attracted by calling. Birds on the ground either ran with wings raised high and tail spread, or with wings closed. A rare procedure was for birds to circle, possibly together, and land in turn at a hole. Once this was watched for 10 minutes before I investigated the hole and found a recently dead storm petrel which had jammed itself trying to come out of a very narrow crack. The bird was removed, some regurgitated oil being left behind, but the birds still continued to land, perhaps smelling the oil, or remembering the bird. A similar display with two birds ended by both trying to land together and rushing to the hole, pecking each other and leap-frogging 152 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. over each other. This is similar to an aerial chase in O. castro described by Allan (1962) but in my observation it was likely that this was a mere jostling for position and not a display. Some birds on landing appeared to peck at the lava or possibly were regurgi- tating oil. The reason or the function for this was not clear, perhaps it might be responsible for the slightly musty smell of the colony. BEHAVIOR AT THE NEST SITE The calls heard from underground were of two types. The commonest (here called the harsh call) was a slightly screeching churr split up by two quick squeaks, and resembled the calling of O. castro which was described by Allan (1962) as a “gutteral purring ‘urr-rrr-rrr’ interjected with a sharp ‘wicka’.” The other was a low purring (the soft call) with a quiet “Tchzee,” in the middle. This purr was made with the bill either open or closed, in the latter instance it was hardly audible 4 feet away. Rarely a chick-like cheep was heard. Any bird going underground, or under the bushes on Isla Pitt, elicitated much calling from other birds, especially if the intruder was forced to pass close to a bird with a nest site. However, disturbances of any sort would start this harsh calling. Each bird appeared to have a small territory which it would defend by calling and fighting if a strange bird came too close. One calling bird, without any egg, was sitting in a depression under a bush with another bird apparently trying to displace it. The “intruder” kept approach- ing to within 3 or 4 inches of the sitting bird which then attacked it and a fight developed with both birds calling, pecking, and holding each others wings as they rolled over and over together. After several such fights the intruder left and the remaining bird sat in its hollow and arranged a few pieces of twigs around it. Twice it went 18 inches away to nibble at small stones and twigs but never brought any of them back when it returned. In some pairs the egg was surrounded by small pieces of lava which must have been collected. An apparent courtship was watched for an hour at another nest. A bird (A), the assumed male. was threatening another (B), which however kept returning and trying to get underneath (A). After a few minutes (B) was accepted and began to pick up and move around stones and twigs but this lasted only a few minutes before it began to nibble gently at the head and especially the nape of (A). After 10 minutes (B) left and was followed by (A) to the edge of the bush (18 inches away). Bird (A) then returned to the scrape. A few minutes later (B), (or conceivably another individual), returned and was greeted by many threats and wing pulling. Again (B) was entirely submissive and soon they sat quietly side by side while (B) preened (A) and nibbled nest- material. Bird (B) then tried to incubate a small stone and immediately there was mutual head preening. Whenever (A) appeared to grow restive, (B) resumed Vor. XXXVII] HARRIS: BIOLOGY OF GALAPAGOS STORM PETRELS 153 nibbling at its nape, or its back if they happened to head to tail. Throughout these displays (A) usually stood whereas (B) moved on its tarsi, and the only calls heard were the soft call. A third bird arrived and was at first scared away by the harsh call but later, on its return, was forcibly evicted. Later in the day, and throughout the night, only a single bird was present. These observations suggest that there is much aggression between birds, perhaps due to intense competition tor nest sites, and that the female has to be submissive to be accepted. INCUBATION BEHAVIOR Notes were taken at a single nest under a Periloba bush. The incubating bird was extremely restless, constantly preened, pecked at various objects around the nest, moved the egg, and even walked up to a foot away for no apparent reason. When settling down on the egg, it either pushed it under with the bill or placed its foot on the egg so that it slid off one side or the other. If it was the wrong side the bird tried again. The bird threatened any intruder with the harsh call, which was often enough to drive it away; if not, the bird would leave the egg and advance with slightly raised and expanded wings and spread tail. No other call was heard from the incubating bird, but it does not seem that the harsh call is restricted to terri- torial defense as two birds visited the hole after hearing this note. The evidence above and some additional observations on the very densely crowded burrows on Tower, suggests that although this species nests in very large, and to our minds overcrowded colonies, there is some division into terri- tories. This is perhaps essential with such competition for nest sites. In the densest areas the size of the territories appears to be governed by the pecking range of the birds. In these very dense colonies adults are very loath to leave their nest sites for any reason. The very conspicuous white rump patch appears to be important in both aerial and terrestrial displays and it would be desirable to compare the behavior of this species with that of an all black storm petrel. As well as being used as a signal to other birds of the same species, it might conceivably serve as a deflec- tive mark to attract a predator’s attention to a nonvital part of the body. Associ- ated with this is the relative ease with which the white upper-tail coverts are pulled out. FEEDING OF THE YOUNG Observations were made on an adult brooding a very small young which had an additional young placed alongside it. The adult moved away 2 inches and sat for 40 minutes while both young called the usual ‘tweep-tweep-tweep—’ and pecked and nibbled at each other. The adults returned and fed the strange young for just over 6 minutes during which time the young gained 7.8 grams. 154 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. The feeding was by regurgitation at the back of the adult’s throat. The chick had its head sideways with the bill tip towards the adult’s throat. During the whole feed the young called continuously at a rate of about one call a second. This method of feeding was different from that described for O. castro (Allan, 1962) where the young was fed by six successive but separate rations of food, perhaps due to the larger prey taken by O. castro. The above case of an adult feeding a strange young may be unusual, in that in four sets of artificial twins, adults ejected the strange young, suggesting that they can recognize their own young. This is perhaps to be expected with dense crowding, especially as the chick is unusually mobile for a petrel, but is contrary to my results for P. puffinus, P. lherminieri, and O. castro. SIZE OF THE TOWER COLONY The immense numbers of birds flighting, which must have included an un- known proportion of nonbreeders, made a direct estimate of the population impossible. The only feasible, and probably the most accurate, method was to determine the density of nests in the colony and then calculate the total nest- ing area. The sample area measured 450 square yards (distances paced out on the flat lava) and was probably in the densest part of the colony. The 34 holes examined had a subterranean area of 37 square feet and held 105 eggs and young so that at least this many pairs were breeding. It was thought that approximately half the surface would cover open area suitable for the petrels which meant a density of about 13 pairs per square yard. The density of nests varied greatly throughout the colony, so that estimates were made in many places of the numbers of birds landing, and to a lesser degree circling, in comparison with the sample area. The areas were then paced out. Burrows were opened up in various parts to check that the scored densities were of the right order of magnitude. The colony was composed of: 3,000 square yards at the density of the examined sample (or 13 pairs per square yard) 18,200 square yards at 1/2 this density 10,800 square yards at 1/3 this density 700 square yards at 1/4 this density 1,200 square yards at 1/5 this density On this reckoning the total population was about 200,000 pairs. The cliff area was ignored as its investigation was impractical. BREEDING BIoLoGy The scattered visits made a detailed study impossible but a few data were collected. Adults were weighed on several dates and the results are summarized in Vor. XXXVII] HARRIS: BIOLOGY OF GALAPAGOS STORM PETRELS 155 TABLE 14. Weights in grams of full grown individuals of Oceanodroma tethys in relation to a) date and b) state of brood patch. Date Colony Number Average Range Pes a) 17.1.67 Tower 59 23.4 20.5—30.0 1.9 8.3.67 Tower 53 21.8 19.0—27.0 1.6 18—20.4.66 Tower 10 SS) 22.1—26.8 1.6 26.6.-3.7.66 Tower 6 28.5 26.0-32.5 Siu 27.5.67 Isla Pitt 7 2325 20.5-29.0 2.6 11.7.66 Isla Pitt 19 26.0 22.0-35.0 2.8 b) Birds with eggs 18 25.8 20.5-35.0 3.6 No brood patch 21 23.4 20.0-27.0 2.0 Unvascularized brood patch § 22.6 19.0-30.0 DD Regrowing feathers of brood patch 23 DRG: 19.0-27.0 2.0 With body molt 2 22.0 19.0-27.0 eA table 14. There were no significant changes in weights with date or breeding condition, except that incubating birds, presumably with stored food, tended to be heavier than nonbreeders. One hundred and ninety eggs measured (average length 27.8 mm., standard deviation 1.01; breadth 20.6 mm., standard deviation .60) showed no variation between years or colony and were similar to six taken from the Pescadores, Peru (average 27.3 X 19.7) (Murphy, 1936). Twenty-three newly laid eggs averaged 5.2 grams (range 3.6—6.3, standard deviation .72). The weight of yolk and albumen in seven freshly laid eggs ejected by birds competing for nest sites was approxi- mately equal. Two females, weights 23.0 and 26.0 grams, were caught on fresh eggs weighing 5.0 and 6.0 grams respectively. Both colonies were extremely crowded. At least two or three pairs were com- peting for virtually every hole, as shown by the numbers of eggs laid. The greatest density of pairs on Tower was found under a lava bubble of an approxi- mate area 8 square feet (fig. 18). In 1966 at least 21 eggs were laid, seven young hatched and no more than five could have fledged. In July, 1967 this same hole had 22 eggs and five young. This egg loss was typical of the whole colony. On Tower, nests were checked at 0800 hours, 1700 hours, and in some holes at 1200 hours. Some incubating birds were marked with paint and the average incubation spell was 5.3 days (changeovers in 14 out of 74 daily checks). There was no tendency for birds to change over at hatching but afterwards no adult brooded a chick for more than 2 days. Unlike O. castro young, these appeared to be blind at hatching and some did not seem to have full vision until about the tenth day. The weight increases of chicks are shown in table 15, and indicate that 156 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. 3 feet Ficure 18. Plan of the position of eggs and young in a crowded nesting area of Oceano- droma tethys under a lava bubble. No more than five young could have fledged from this hole in 1966. In July, 1967, this hole had 22 eggs and five young. feeds were most frequent during the day. Many of the overnight feeds were due to adults returning very early in the morning or staying overnight in the burrows. In 91 out of 181 days it was known that a chick was fed at some time in the 24 hours. The maximum recorded increase in the weight of a nestling was TABLE 15. Increases in weight of young of Oceanodroma tethys during the day. Tower weighings were made 26 June to 3 July, 1966, and 16 to 18 July, 1966; Isla Pitt weighings, 10 to 11 July, 1966. Time Number of Number ened Percent Average increase Colony (hours ) young in weight increased grams Tower 0800-1700 246 109 46 365 1700-0800 180 42 23 1.9 O800—1200 67 12 18 3.8 1200-1700 67 18 27 Sia 1700—O0800 67 8 12 l7/ Isla Pitt 0830-1630 17 7 41 D3 1630-0800 17 2 12 3.3 6 35 2:5 0800-1300 17 Vor. XXXVII] HARRIS: BIOLOGY OF GALAPAGOS STORM PETRELS 157 50 40 30 Weight(g) Wing length (mm) 10 20 30 40 50 60 70 AGE IN DAYS Ficure 19. Weight and wing length curves for a young of Oceanodroma tethys raised by a pair of Oceanodroma castro. 12.5 grams, perhaps due to it being fed by both its parents. Additional weigh- ings made on Isla Pitt showed a similar pattern with 13 out of 17 young fed at least once in the 30 hours covered by weighings. Four young of O. tethys were removed to Plaza and fostered under O. castro parents. One died soon after introduction but not of starvation as its stomach held food, one disappeared at 72 days but it was unlikely to have fledged, two were successfully reared. These two fledged at 66 and 86 days respectively (both + 2 days). One of these weighed 32 grams and had a wing length of 137 mm. The growth curve of one young is shown in figure 19. In the 34 holes on Tower for which I have adequate data, 193 eggs were laid, 63 young hatched and no more than 45 young could have fledged (23 per- cent of eggs laid). This success rate is not typical, however, as more eggs must have been laid and lost between my visits and the number of young that possibly fledged was a maximum (based on those alive on 17 August and whose remains were not found in December) and many more could have died unrecorded. Cause oF Ecc, Cuick, AND ADULT LOSSES Almost all the nesting losses appeared to be due to the intense intraspecific competition for nest sites. Nelson (1966) was wrong in thinking that the reason for the large numbers of ejected eggs was interspecific competition with O. castro for that species did not rest in the same area. Great care had to be taken when replacing stones on the roofs of burrows as eggs in some burrows were 158 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. eaten by Nesomimus parvulus predators after my inspections when these birds entered by seemingly impossibly small holes. Normally they probably only get eggs ejected by the birds. On Isla Pitt, where there are no mockingbirds, ejected eggs remain until rotten. As mentioned previously eggs of this species can normally undergo prolonged periods without incubation and still remain viable but this is of no advantage at present as any eggs left are immediately ejected by other pairs competing for nest sites. Due to my disturbance several pairs lost their eggs in this way. No reasons for chick losses other than this interspecific competition were known in Tower, but at Isla Pitt Phaethon aethereus and Puffinus lherminiert killed some young of O. tet/ys in open sites. This is unusual however as in such places, O. tethys nested in the back of the holes and the other species near the entrance. In both places Asio galapagoensis killed large numbers of O. castro and O. tethys but far more of the former than would have been anticipated by their relative abundance, probably because the owls preferred to hunt at night. When hunting for O. tethys the owls usually waited near a crack and jumped feet first at the petrel just as it took off. Once an owl was seen resting itself on bent wings while stretching both feet into a crack to try to get a petrel which it could see but not reach. On Tower the owls also killed prey of Puffinus lherminiert, C. furcatus, Phaethon aethereus, Anous stolidus, Pterodroma phaeopygia, and Sterna fuscata, the latter two species were not otherwise seen here. Other observed causes of death were: jammed in holes (5), eaten by Fregata minor (4) and Ardea herodias (3). Winc Mott Oceanodroma t. tethys appeared to molt its wing feathers away from the breeding grounds and between breeding seasons, as only two individuals were ever seen in primary molt. Both were from a sample of 80 netted 8 March; they were regrowing the two innermost primaries on each wing. Therefore it follows that the birds flighting in the nonbreeding season cannot be individuals that had bred or were going to breed during the main season. Of 82 skins col- lected in Galapagos waters, four collected in April (1), May (2), June (1), were just completing the growth of the outermost primaries, presumably just prior to breeding. Other molting birds taken at sea were: 1 August at 13°28’ N. 105°52’W. Primaries inner NN NNNNNNWN 3 1 August 13°28’ N. 105°52'W. NNNNNNN 41 0 5 August 10°N. 109°W. NNNNWNN NIN@2OO 8 August 10°N. 109°W. NNNNNNN 420 8 August 10°N. 109°W. NNN N N NeoNSeeeo 14 August 8°45‘N. 106°50’W. NN NN N N goo Vot.XXXVIT] HARRIS: BIOLOGY OF GALAPAGOS STORM PETRELS 159 1 September near Cocos Island NNNNNNNNN 4 1 September near Cocos Island NeNG ING NoG) Ny NY 32000 1 September near Cocos Island NNNNNNNN 4 3 All of these were perhaps nonbreeders. The pattern of molt appears to be the same in O. t. Relsalli for of 37 birds collected by Beck in May and June off Peru, including some breeding birds, three were just completing the primary molt, those birds were two males with enlarged gonads, and a female which had apparently just laid. Birds collected away from the breeding grounds are mainly in wing molt but the data are diffi- cult to interpret as they refer to a few series from scattered positions north of the equator. Many of these birds may have been nonbreeders, as the majority of them could not have finished their molts in time for the breeding season. Un- fortunately I have seen no skins collected from Peru in the nonbreeding season. Koepecke (1964) says that the species has a migration but it is not clear if this occurs in both adults and young. DiIscUSSION The two most impressive aspects of the biology of this species are the dense crowding of nests and the flighting behavior and these are the two most difficult aspects to explain. On Isla Pitt it was impossible for the colony to expand without emigration to the main island of San Cristébal, a distance of perhaps a half-mile, as all the good nest sites and many obviously sub-optimal were occupied. This overcrowd- ing on small islands because birds will not leave to start another colony, even a few hundred yards away, is typical of the majority of colonial sea birds. How- ever the Tower colony could expand at either end into apparently identical nesting habitat to that already occupied. Here at least, it is difficult to see what the birds would lose either from protection against predators or from any social factors which might be important by expanding the colony. Any slight losses would be more than offset by the increase in breeding success which might follow from lessening of intraspecific competition. The advantage to an individual bird of the prolonged display flighting out- side the breeding season is again difficult to understand. It might possibly bring all the birds into breeding condition together but other species synchronize their breeding with far less wastage of time and effort and anyway the advantage of synchronized breeding to O. tethys remains to be proved. This synchrony of breeding brings about large losses of eggs, but we do not know whether there might be only a short time each year when conditions are suitable for rearing young. Against any advantage must be placed the energy used in flight, the risk of damage from aerial collisions, and the risk of predation. Another possible explanation which much be considered is that of Wynne- 160 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Edwards’ (1962) theory of “epideictic” displays. Nelson (1966) noted several factors which he thought did not fit the flighting to this theory, namely no sharply demarcated daily or seasonal peaks of flighting when the population might be able to gauge its own numbers, and egg losses due to interspecific competition. Even though these conditions are, in fact satisfied, the breeders could hardly get an estimate of the total population as they are not present at the peak of flighting, but away molting. The flighting is therefore unlikely to be an epideictic display but its significance remains obscure. OCEANITES GRACILIS The commonest storm petrel seen by most people in Galapagos is Oceanites gracilis. As with Oceanodroma tethys, this is a species endemic to the Humboldt Current with two separate races, the larger of which “galapagoensis” is re- stricted to the Galapagos (Lowe, 1921). It is remarkable that despite the searchings of the older scientific expeditions (especially of Beck), Lévéque (1964), and myself, the breeding grounds have yet to be discovered. Similarly no nest of the South American form has been found. Presumably the species is nocturnal in its visits to land and the colonies situated in unusual locations, perhaps cliffs, as in Galapagos almost all the smaller islands have been investigated. From the examination of gonads it appears that breeding occurs during the cold season (Loomis, 1918). This would fit with the species being restricted to the Humboldt Current. I have seen many birds in wing molt from August on- wards, presumably after the breeding season. I examined a single female in June, 1967, which had an undeveloped ovary and no molt. The bird weighed 17 grams and its stomach contained nine very small fish eye lenses. Loomis (1918) recorded very small fish being taken from a bird shot in Galapagos waters. Flight and feeding habits appear to be similar to the closely related Oceanites oceanicus. Food is picked from the surface of the water as birds pattern into the wind — the familiar “walking on the water” of many long-legged storm petrels. Presumably the normal food is small plank- tonic fish and crustacea, but I have also seen this species eating scraps from the activities of fishermen, sea lions, sharks, and killer whales (Orcinus orca). ECOLOGICAL SEPARATION Although it is unfortunate that we know nothing of the breeding of Oceanites gracilis, which is a member of a group which for the most part breed in the sub- antarctic zone (Kuroda, 1954), it is ecologically quite distinct from the species of Oceanodroma. At least in Galapagos waters, Oceanites gracilis is an inshore species, I usually saw it feeding within a mile of the coast and extremely rarely more than 20 miles from land. VoL. XXXVII] HARRIS: BIOLOGY OF GALAPAGOS STORM PETRELS 161 Both the Galapagos species of Oceanodroma feed well out to sea, but there seems to be a food difference as O. tethys eats smaller fish than does O. castro and also takes some crustaceans. The basic separation may however, be in the time of feeding; O. castro is nocturnal when visiting the colonies so must feed mainly by day, while O. tethys must feed mainly at night. The feeding ranges may also be different. O. tethys, which is endemic to the Humboldt Current, is common among the islands and between the islands and the Ecuadorian coast. On the other hand, though we have no direct observations, it seems possible that O. castro might prefer the bluer, more oceanic waters to the west of Galapagos, as this would fit in with its general distribution in warm waters. This evidence strongly suggests that the three species of storm petrels resident in Galapagos are ecologically isolated and do not compete with each other for food. ACKNOWLEDGMENTS This study was carried out while I was receiving a research grant from the Scientific Research Council. My wife, who helped with the whole of this work, and I are extremely grateful for their support and also to the Royal Society whose research table at the Charles Darwin Research Station was made available to us. The Charles Darwin Foundation offered every help and Mr. Roger Perry, the director, and Mr. Tjitte de Vries put the resources of the research station at our disposal. Senor Miguel Castro, Mr. Rolf Sievers, and Mr. Bernhard Schreyer gave much help in the field. We should also like to thank our friends in Gala- pagos for their many kindnesses to us. The curators of the museums mentioned earlier courteously allowed me ac- cess to their collections on which parts of this paper are based. Dr. Dean Amadon of the American Museum of Natural History loaned two skins of O. castro collected on Sao Tomé; the other skins in figures 1 and 2 were from the British Museum. Dr. Christian Jouanin kindly measured skins in the Muséum National d’Histoire Naturelle (Paris) and sent me data on breeding seasons on the Salvages and Baixo. Dr. N. K. Johnson examined skins in the Museum of Verte- brate Zoology, University of California, Berkeley. The mallophaga were identified by Dr. Theresa Clay in the fleas by Dr. F.G.A. Smit. Dr. M. R. Clarke identi- fied the squid remains and Dr. David Snow supplied some notes on his study on Plaza. Dr. David Lack gave encouragement throughout and clarified the manu- script with his constructive criticism. Dr. C. M. Perrins, and Dr. W.R.P. Bourne read parts of the paper and offered advice. LITERATURE CITED Axpsott, Donatp P. 1966. Factors influencing the zoogeographic affinities of Galapagos marine fauna. The 162 CALIFORNIA ACADEMY OF SCIENCES [PRoc. 4TH SER. Galapagos, pp. 108-122, 8 figs. (editor Bowman), University of California Press. ACADEMY OF SCIENCES, U.S.S.R. 1964. Physical-geographic atlas of the world. 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W., AND B. K. Snow 1966. The breeding season of the Madeiran storm petrel in the Galapagos. Ibis, vol. 108, no. 2, pp. 283-284. STONEHOUSE, BERNARD 1962. The tropic birds (genus Phaethon) of Ascension Island. Ibis, vol. 103b, no. 2, pp. 124-161. 1963a. Egg dimensions of some Ascension Island sea-birds. Ibis, vol. 103b, no. 3, pp. 474-479, 1963b. The laying dates of some St. Helenan sea-birds, 1958-59. Ibis, vol. 103b, no. 3, pp. 480-482. WaArRHAM, JOHN 1962. The biology of the giant petrel Macronectes giganteus. Auk, vol. 79, no. 2, pp. 139-160. WyYNNE-Epwarps, V. C. 1962. Animal dispersion in relation to social behaviour. Oliver and Boyd, London, 653 pp. OO Vs , o @ | ora, theories nt ee ee _ a - a Vepsp arpa) Trea ; > pO) aa ie pol sibel re Ease e eS er ; 4 eC lane Rag AivSl Cesc eet SN QO, ie ; g y a ‘* nee 4 her : oe ent 2 “V2 % ae a oe to zune am 7 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XXXVII, No. 5, pp. 167-234; 105 figs.; 2 maps March 6, 1970 FOSSIL DIATOMS FROM THE PRIBILOF ISLANDS, BERING SEA, ALASKA By G Dallas Hanna California Academy of Sciences San Francisco, California 94118 The Pribilof group of islands is located near the center of Bering Sea, Alaska. The Aleutian Islands are about 200 miles to the south; the Alaska mainland is about an equal distance to the east; and St. Matthew Island lies about the same distance to the north. St. Paul Island is the largest of the group, about 12 miles long, and St. George is a little smaller. Otter Island is about a mile long, while Sea Lion Rock and Walrus Island are very small. All of these islands have been known since they were discovered to be the summer breeding ground of the Alaska fur seal. The climate is sub-Arctic; that is, there are no trees or woody shrubs of any kind, and there is no permafrost. Vegetation consists of a thick blanket of mosses, lichens, and grasses with a large flora of flowering plants, some of which are endemic. The plants effectively conceal some flat areas which may overlie surface sedimentary rocks. The fauna and flora have been more thoroughly explored than any other equal area in Alaska. Except for a few favorable sea-cliff exposures, all of the visible rocks are volcanic. No terraces have been identified and no surface evidence of glacial action is apparent except for the recent report of a small ice mass which was once present on the high part of St. George Island (Hopkins and Einarsson, 1966, pp. 343-344). There are numerous craters and a great succession of lava flows. The highest point is on St. George Island and is given as 994 feet by Barth (1956, p. 102). The highest point on St. Paul is Rush Hill, 662 feet. There are a few lava caves on St. Paul, but no fumaroles or hot springs have been found on any of the islands. The surrounding sea is relatively shallow, 30 to 50 fathoms, but the depth increases rapidly to the southwest. The Bering [167] mieten Ler 4 " ATES: Marine Bin Yiral rot 4 re r | Laboratory f 1 & Fe i MAR 1 61970 WJODS HOLE, MASS. SOMES" G-Star emegene 2s > a ee 168 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. 39602 Map 1. Index map of Alaska showing location of Pribilof Islands in Bering Sea. Sea ice sheet reaches the Islands once every 3 or 4 years and is a minor factor in erosion. The region is one of violent storms, one effect of which is extensive sea cliff erosion. Thus most of the shore lines are near vertical walls of more or less stratified layers of lava. Some of the former craters have been dissected so that only a landward portion remains. On St. Paul the headlands have been eroded away to such an extent that there are some extensive sand beaches between. It was my good fortune to be stationed on the Pribilof Islands from 1913 to 1920. During this period I visited and collected extensively at the well-known locality, Black Bluff on St. Paul Island. This is a more than half dissected cinder cone at East Landing. The volcanic material evidently was extruded through some sediments because among the cinders there is an occasional block of hard gray mudstone containing abundant mollusks. These were the basis for all published reports on the paleontology of this group of islands prior to my visit. The presence of these fossils led me to make as extensive a search as pos- sible of other sea cliffs for exposures of fossil-bearing sediments. In this I was successful to some extent as shown in the list of localities following. It was found upon careful examination of some of the Black Bluff sedimen- tary blocks that large numbers of diatoms were present and that they could be obtained for study by the usual treatment with acids. This led to an examination of the newly discovered exposures and a short list of species from the one found VoL. XXXVIT] HANNA: PRIBILOF FOSSIL DIATOMS 169 at Tolstoi Point, St. Paul. This list has been published (Hanna, 1919, pp. 223, 224). A restudy of that material and that from all of the other diatom-bearing sediments thus far found on the islands is the basis for the present paper. ACKNOWLEDGMENTS Much inspiration for the final preparation of this report was derived from consultations with F. Stearns MacNeil, David Hopkins, and the late Don K. Miller of the U.S. Geological Survey. In early work on the islands I was helped by my colleagues, the late George Haley and E. C. Johnston. And in 1960 St. Paul Island localities were visited again through the help of Howard Baltzo, Roy Hurd, and Kenneth K. Bechtel. My original interest in fossil diatoms was initiated by the study of material from these northern deposits through the encouragement of Dr. Albert Mann of Washington, D.C. I am under deep obligation to Dr. A. L. Brigger for much assistance in the later stages of the work and especially in the making of individual mounts of specimens for museum cataloguing and storage. He also very kindly made the photographs, figures 49, 102-105. Dr. Joseph F. Burke identified the Awlaco- discus, one of the species being checked by Dr. Paul S. Conger. This assistance is greatly appreciated. History oF GEOLOGICAL WoRK A general history of the Pribilof Islands is long overdue because of their being the breeding ground of Alaska fur seals, an economic asset of international significance. They are almost as well known for the vast rookeries of sea birds. Hundreds of books and papers of scientific and popular appeal have been written about the Islands, but there are scarcely a dozen which deal with the geology. Fewer yet are concerned with fossils. A late work is by Tom F. W. Barth in 1956, U.S. Geological Survey Bulletin, no. 1028-F. This contains descriptions of the various types of volcanic and igneous rocks, locations of fossil deposits, a short list of Mollusca, and an excellent bibliography. Although the volcanic nature of the islands had been mentioned by many travellers and naturalists, the first geologist to treat the subject was J. Stanley Brown in 1892. Dall followed in 1899 with a list of Mollusca found in the cliffs of Black Bluff, and again in 1919 with an account of those found by me at Tolstoi Point on St. Paul Island and at the Point having the same name on St. George Island. Also in 1919 I published the short note referred to above. In 1930 H. S. Washington gave an extensive account of the rocks of the Pribilofs. A very important work concerned with the geology of the Pribilofs is that by Hopkins, MacNeil, Merklin, and Petrov (1965, pp. 1107-1114). This deals primarily with the ‘Quaternary correlations across Bering Strait,” but some of the important information was obtained on St. Paul and St. George islands. The paper has an excellent bibliography. 170 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER. This was followed by the results of additional field work included in the note on glaciation at St. George Island by Hopkins and Einarsson (1966). AGE Some of the most common species of diatoms in these deposits are extinct in so far as available records indicate. Much work has been done on the living flora of Bering Sea and adjacent waters by Mann (1907), and several Soviet workers. The assemblages of these Pribilof deposits are scarcely comparable to those of living material. On the other hand they are similar in many ways to the species recorded by Jousé, Zabelina, and others from Sakhalin Island and Kamtschatka Pliocene. The very extensive and useful work on this area by V. S. Sheshukova-Poretskaia was received too late for profitable use in connection with the Pribilof study. It seems inescapable that the sediments which contain this assemblage of species can be no younger than Pliocene and it seems likely that they are well down in that period. Thus the diatoms confirm the age determination made by Dall from a study of the Mollusca. They do not bear out the belief of Barth and others that the age of the islands is Pleistocene. METHOD OF STUDY An attempt has been made herein to select individual diatoms representing all of the species found in the various samples studied. This seems to be the best method of preserving illustrated materials because each specimen is readily located inside a small black circle on a microscope slide. Thus it becomes a museum specimen which can be cataloged and referred to by later workers at will. The samples from Tolstoi Point, St. Paul Island were searched more thoroughly than any of the others for individual mounts. In addition to these, several strewn slides were made from each sample cleaned. These were scanned with mechanical stage and served in compiling the lists of species from each of the samples. As for the drill cuttings from Navy water well no. 2, long experience in the study of such samples from wells drilled for oil has indicated that the depths recorded for such cuttings cannot be relied upon exactly. There is bound to be some mixing due to slumping or other causes. Also, surface waters are often used in various ways in the drilling operations so that there is probability of mixing of freshwater living diatoms with the fossil forms. In the case of the present study, the few freshwater forms found have not been listed. In spite of these difficulties pertaining to the use of cuttings from a well drilled with cable tools, it is believed that the samples serve a useful purpose. The most obvious, of course, is that they show sediments extending to 400 feet below sea level at St. Paul Island village, a condition which could hardly have been expected from surface geology. VoL. XXXVIT] HANNA: PRIBILOF FOSSIL DIATOMS 171 Ardiguen Rookery Reef Pt + Lukanin Pt ~~ she” Sy z ee Kitovi Rookery | | Sze (fed Black Bluffs on Lion Rock Map 2. Southwest end of St. Paul Island showing locations of the three most important localities for fossil diatoms. COLLECTING LOCALITIES The Pribilof Island collecting localities recorded in the register of the Cali- fornia Academy of Sciences are as follows: 715 ALT i233 £322 36829 Tolstoi Point, St. George Island, Alaska. Edward C. Johnston, collector, 1920. Tolstoi Point, St. Paul Island, Alaska. G Dallas Hanna, collector, 1920. Black Bluff, St. Paul Island, Alaska. G Dallas Hanna, collector, 1920. Navy water well no. 2. Samples of cuttings down to 400 feet below sea level. A. Christofferson, collector, 1921. Tolstoi Point, St. Paul Island, Alaska. G Dallas Hanna, Kenneth Bechtel, and Howard Baltzo, collectors, 1960. 172 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. 36830 Zapadni Point, St. Paul Island, Alaska. G Dallas Hanna, Kenneth Bechtel, and Howard Baltzo, collectors, 1960. 36832 Black Bluff, St. Paul Island, Alaska. G Dallas Hanna, Kenneth Bechtel, and Howard Baltzo, collectors, 1960. 37724 Tolstoi Point, St. Paul Island, Alaska. G Dallas Hanna, collector, 1916. 37725 Garden Cove, St. George Island, Alaska. G Dallas Hanna, collector, 1917. 37726 Navy hand-dug well, St. Paul Island, Alaska. A. Christoffersen, collector, 1927. 38597 Einahnuhto Bluffs, St. Paul Island, Alaska. Diatoms from a boulder in gravel which contains fossil shells. David Hopkins, collector, 1962. The U.S. Geological Survey has extensive collections of fossil mollusks from both St. Paul and St. George islands. These have been collected by numerous individuals and include all such material which I was able to find up to and including 1919. WELLS The U.S. Navy built and maintained radio stations on both St. Paul and St. George prior to World War I. From time to time they were enlarged and finally the one on St. Paul reached a critical stage because of a shortage of water. Before that, all of the water for domestic purposes was obtained from a shallow well about half a mile from the village. This location had been well selected and engineered. It was on flat land only a few feet above sea level, but adjacent to a lava flow. It was sunk just about to sea level. The water was excellent but the supply was extremely limited. In 1918 a pump was installed at Ice House Lake under the direction of Agent and Caretaker, H. C. Fassett. A wood pipe line was laid to the village and to a tank house on top of the village hill. For the first time in the history of the island the inhabitants had an adequate supply of water without hauling or carrying it. This lake, however, was not sufficient to supply the enlarged radio facilities. Therefore, in 1920, the Navy brought to the island a water well drilling outfit of the usual cable tool variety. The location for the first well was close to the present radio station; that is, between it and the entrance to Salt Lagoon. Scanty records indicate that total depth reached was about 160 feet when the bit was stuck. No samples have been preserved. In 1921 a second well was drilled a little farther out toward the old village well (letter, Roy Hurd, April, 1962). Somewhat better equipment was used than on the first well. Mr. A. Christoffersen saved a suite of bailer samples for me from the various depths indicated below. The samples were collected fresh, excess water was removed, and the solids were sealed in tin cans with solder. In each case the depth was stamped in the lid of the can with steel figures. Mr. Vout. XXXVIT] HANNA: PRIBILOF FOSSIL DIATOMS 173 Christoffersen was then the manager of the by-products plant located close to the well site. Descriptions of the various samples are given below. In many of them there was well-preserved diatom material even down to the deepest one, 400 feet. Some of these samples have been cleaned by the usual technique and the species have been listed. The Navy drilled a third well between the first and second (map 2) at a later date, but no detailed information as to depth and section has been found. None of these three wells produced fresh water, at least not in requisite quantity for the purpose intended. Therefore, in 1927, the same organization dug a shallow well just beyond (north) the old village well. Evidentally it turned out to be useful because a windmill was installed and a pipe line was laid to the radio station. Remnants of the mill were still present in 1960. Mr. Christoffersen collected and sealed a sample for me from this well. It contains a very large assemblage of species of diatoms. These are obviously brackish water forms and belong to a much later period than those from the other wells; it has been decided not to include them in the present paper. The flora from this dug well deserves to be made the basis of a separate study. In 1960 a completely adequate supply of excellent water for all purposes was obtained from wells on the northeast flank of Telegraph Hill. The following are records of ditch samples recovered from water well no. 2, drilled near St. Paul Island village in 1921 by U. S. Navy, A. Christoffersen, collector. The figures indicate depth in feet. 60 Gray silty sand with many marine diatoms. 80 Black volcanic sand. 90 Black volcanic sand and scorria pebbles. A minor amount of gray silt contains marine diatoms. 130 Gray and brown sand with fragments of very hard fine grained gray sandstone. 144 Fragments of black and brown volcanic material. 164 Fine dark sand. Looks like beach sand. 177. Fine well sorted sand, not as dark as 164. 196 Fine gray well sorted sand. 198 Abundant fragments of Spisula alaskana and Balanus species in a fine, well sorted brown sand. Fragments of hard igneous pebbles up to 2 inches across, waterworn. 200 Fine gray silty sand. 206 Fine gray silty sand. 206 Fine gray sand with lumps of light gray silt and a few black waterworn pebbles. Fragments of Spisula species. 300 Fine gray silty sand. Abundant diatoms. 325 Fine gray silt with minor amount of fine sand. 174 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. 345 Fine gray silt with some fine sand. Abundant diatoms. 400 Same as for the 345-foot depth. Stained brown from rusty container. One pebble 2 inches across. Abundant diatoms. The following are common diatoms found in various samples from Navy well no. 2, drilled on St. Paul Island, Alaska. Locality 1322 (CAS). Depth 60 feet Coscinodiscus apiculatus Ehrenberg Coscinodiscus lineatus Ehrenberg Coscinodiscus undulosus Mann Cosmiodiscus insignis Jousé Rhaphoneis amphiceros Ehrenberg Thalassionema nitzschoides Grunow Depth 80 feet Arachnoidiscus ehrenbergi Bailey Coscinodiscus marginatus Ehrenberg Coscinodiscus pustulatus Mann Coscinodiscus radiatus Ehrenberg Cosmiodiscus insignis Jousé Depth 90 feet Actinoptychus senarius (Ehrenberg) Arachnoidiscus ehrenbergii Bailey Coscinodiscus apiculatus Ehrenberg Cymatotheca weissflogii (Grunow) Hendey Coscinodiscus lineatus Ehrenberg Coscinodiscus marginatus Ehrenberg Coscinodiscus pustulatus Mann Cosmiodiscus insignis Jousé Thalassiosira punctata Jousé Diploneis ornata Schmidt Stephanopyxis appendiculata Ehrenberg Trachyneis aspera (Ehrenberg) Xanthiopyxis ovalis Lohman Depth 105 feet Actinoptychus senarius (Ehrenberg) Chaetoceros species Cocconeis, 2 species Coscinodiscus apiculatus Ehrenberg Cymatotheca weissflogii (Grunow) Hendey Coscinodiscus marginatus Ehrenberg VoL. XXXVIT] HANNA: PRIBILOF FOSSIL DIATOMS Coscinodiscus pustulatus Mann Coscinodiscus radiatus Ehrenberg Thalassiosira punctata Jousé Coscinodiscus undulosus Mann Cosmiodiscus insignis Jousé Dictyocha fibula Ehrenberg (Silicoflagellata) Dictyocha speculum (Ehrenberg) (Silicoflagellata) Ebriopsis antiqua (Schulz) (Silicoflagellata) Melosira clavigera Grunow Melosira sulcata Ehrenberg Navicula lata Brebisson Navicula, 3 species Rhaphoneis amphiceros (Ehrenberg) Rhizosolenia species Xanthiopyxis ovalis Lohman Depth) 213; feet Actinoptychus senarius (Ehrenberg) Arachnoidiscus ehrenbergii Bailey Chaetoceros species Cocconeis species Coscinodiscus apiculatus Ehrenberg Cymatotheca weissflogii (Grunow) Hendey Coscinodiscus marginatus Ehrenberg Cosmiodiscus insignis Jousé Thalassiosira punctata Jousé Dictyocha fibula Ehrenberg (Silicoflagellata) Dictyocha speculum (Ehrenberg) (Silicoflagellata) Ebriopsis antiqua (Schulz) (Silicoflagellata) Melosira clavigera Grunow Melosira sulcata Ehrenberg Rhaphoneis amphiceros Ehrenberg Thalassionema nitzschoides Grunow Xanthiopyxis species Depth 400 feet Actinoptychus senarius (Ehrenberg) Cymatotheca weissflogii (Grunow) Hendey Coscinodiscus marginatus Ehrenberg Cosmiodiscus insignis Jousé Coscinodiscus pustulosus Mann Coscinodiscus radiatus Ehrenberg Thalassiosira punctata Jousé 175 176 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47H SER. Dictyocha fibula Ehrenberg (Silicoflagellata) Dictyocha speculum Ehrenberg (Silicoflagellata) Ebriopsis antiqua (Schulz) (Silicoflagellata ) Melosira clavigera Grunow Melosira sulcata Ehrenberg Stephanopyxis appendiculata Ehrenberg Thalassionema nitzschoides Grunow Xanthiopyxis ovalis Lohman EINAHNUHTO BLuFFs, ST. PAUL ISLAND Locality 38479 (CAS) A series of marine clastic sediments is exposed under volcanic rocks along the northeast shore of St. Paul Island. This exposure was studied in some detail by Dr. David Hopkins and associates of the U.S. Geological Survey in 1962 and among other things he collected a fine grained well cemented calcareous silt- stone boulder. This contained a few mollusks and through his permission an examination of it was made for siliceous fossils. The material broke down readily in the usual treatment of such rocks with acids and was found to contain an abundance of diatoms. In three nights spent searching slides, the following species were picked out and identified: Actinoptychus senarius (Ehrenberg) Cymatotheca weissflogii (Grunow) Hendey Coscinodiscus marginatus Ehrenberg Cosmiodiscus insignis Jousé Coscinodiscus pustulatus Mann Coscinodiscus radiatus Ehrenberg Thalosiosira punctata Jousé Coscinodiscus undulosus Mann Melosira clavigera Grunow Melosira sulcata Ehrenberg Stephanopyxis appendiculata Ehrenberg The first four listed species are by far the most abundant. They are also abundant at Black Bluff, Tolstoi Point, and in the material from Navy Well no. 2 samples at various depths. All of the other species are likewise found in the same deposits which strongly indicate that all are of approximately the same age. In addition to the diatoms found in the Einahnuhto Bluffs sample, the silicoflagellates Ebriopsis antiqua and Dictyocha were present. ZAPADNI PoINT, ST. PAUL ISLAND Locality 36830 (CAS) I first saw the sediments at this locality in 1916 but did not find any fossils. In 1960 I visited it again in company with K. K. Bechtel, Howard Baltzo, and VoL. XXXVIT] HANNA: PRIBILOF FOSSIL DIATOMS 177 Margaret M. Hanna. In the gray silty ash there is a small proportion of fossil diatoms. The species are the same as those found in other Pribilof Island localities. A description of the outcrop is as follows: A fault, upthrown along the west side of Antone Lake, has produced a steep talus slope of heavy blocks of volcanic rock. This material is also exposed on the sea cliff just to the west of the lake and is followed by 20 feet of gray fissile shale with a 1-inch layer of pebbles near the bottom. This shale is underlain by a heavy conglomerate with boulders of volcanic material up to 6 inches in diameter. Two smali pebbles of greenstone were found in the conglomerate. Below this the only visible remaining material was heavy black and red scorria with blocks of basalt. The entire section dips to the east 65 degrees. The list of species found at Zapadni Point, St. Paul Island, locality 36830 (CAS), consists of the following: Actinoptychus senarius (Ehrenberg) Cymatotheca weissflogii (Grunow) Hendey Coscinodiscus marginatus Ehrenberg Coscinodiscus imsignis Jousé Coscinodiscus pustulatus Mann Coscinodiscus radiatus Ehrenberg Thalossiosira punctata Jousé Coscinodiscus undulosus Mann Dictyocha fibula Ehrenberg (Silicoflagellata) Ebriopsis antiqua (Schulz) (Silicoflagellata) Melosira clavigera Grunow Melosira sulcata Ehrenberg Stephanopyxis appendiculata Ehrenberg Thalassionema nitzschoides Grunow EXPOSURE AT ToLtsTo1 Point, ST. PAUL ISLAND The exposure of sediments is perhaps best seen from the base of Lagoon Reef. It starts along the shore about 4 mile from there and lies below 150 feet or more of basaltic cliffs. It extends down below sea level an unknown distance. The outcrop extends northward about ’2 mile and the maximum exposed thick- ness is estimated to be 70 feet. The strata are nearly horizontal for the most part, although there is a slight dip both north and south at the ends of the out- crop. Thus the exposure is an anticline but it is faulted near the center. To the south of the fault the sediments are brown sandstones, friable to firmly cemented, containing abundant fossils and interspersed layers of volcanic pebbles. Most of the fossils are bivalve shells belonging to Astarte. One layer of gray sandy shale is very fossiliferous and may contain diatoms. North of the fault and extending to the end of the exposure there is very fine gray shale also about 70 178 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. feet thick above sea level. This contains abundant fossil diatoms. Both the sands south of the fault and the shale north of it are overlain by nearly horizon- tally bedded basalt. Therefore, it is obvious that the fault is pre-volcanic. However, a crevice in the vertical cliff at that point indicates some horizontal movement on the fault after the basalt was deposited. In my earlier work I found the fine gray diatom-bearing shale only at extreme low tide. This area is now obscured by large blocks of talus basalt which have fallen from the cliff above. Since most of the species shown on the following plates are from this locality, it seems unnecessary to list them separately. It is obvious from comparing the various lists that there is no difference in the diatom flora of any age significance among the several localities. Brack Buiurr, ST. PAuL ISLAND Locality 1233 (CAS) The following species were taken at this locality: Actinoptychus senarius (Ehrenberg Coscinodiscus apiculatus Ehrenberg Cymatotheca weissflogi (Grunow) Hendey Coscinodiscus marginatus Ehrenberg Cosmiodiscus insignis Jousé Coscinodiscus pustulatus Mann Coscinodiscus radiatus Ehrenberg Coscinodiscus undulosus Mann Dictyocha fibula Ehrenberg (Silicoflagellata) Distephanus speculum (Ehrenberg) (Silicoflagellata) Melosira clavigera Grunow St. GEORGE ISLAND Locality 37725 (CAS) Seven species were taken here: Thalassiosira punctata Jousé Coscinodiscus marginatus Ehrenberg Cosmiodiscus insignis Jousé Coscinodiscus pustulatus Mann (The above four species were very abundant) Coscinodiscus radiatus Ehrenberg Rhaphoneis amphiceros Ehrenberg Stephanopyxis apendiculata Ehrenberg VoL. XXXVII] HANNA: PRIBILOF FOSSIL DIATOMS 179 SPECIES FOUND ON PRIBILOF ISLANDS Actinoptychus splendens (Shadbolt). (Figures 25, 26, 28, 40, 44.) Actinophaenia splendens (Shadbolt) in BricHTwet1, Quart. Journ. Micr. Sci., vol. 8, 1860, p. 94, pl. 6, fig. 18. Actinoptychus splendens (Shadbolt), Ratrs in Pritchard, Hist. Infus. ed. 4, 1861, p. 840. ScumipT, Atlas Diat., pl. 153, 1890, figs. 3, 16-17. Wortr, Diat. N. Amer., 1894, pl. 92, figs. 9-12. Actinoptychus solist HANNA and Grant, Proc. Calif. Acad. Sci., 4th ser., vol. 15, no. 2, 1926, p. 123, pl. 12, figs. 1-3. Maria Madre Island, Mexico. The specimens found at Tolstoi Point and in Navy well no. 2 both on St. Paul Island, Alaska, do not differ in any constant character known to me from material customarily assigned to A. splendens from Tertiary deposits in California and elsewhere. It would seem better to consider the species to be one of high variability rather than to try to fit each specimen to one of a multitude of named and unnamed forms. Figure 28 approaches the variation which at one time was given the genus name Debya. Actinoptychus senarius (Ehrenberg). (Figures 38, 39, 45, 76.) Actinocyclus senaritus EHRENBERG, Infusionsthierchen, 1838, p. 172, pl. 21, fig. 6. Actinoptychus senarius (EHRENBERG), Abh. Akad. Wiss. Berlin., 1841 [1843], p. 400, pl. 1, fig. 21. Hernprey, Discovery Reports, vol. 16, 1937, p. 271. Actinocyclus undulatus BAtLey, Amer. Journ. Sci. Arts, 1842, pl. 2, fig. 11, Richmond, Virginia. Actinoptychus undulatus (Bailey), HANNA and Grant, Proc. Calif. Acad. Sci., 4th ser., vol. 15, 1926, p. 124, pl. 12, fig. 4. Wore, Diat. N. Amer., 1894, pl. 92, figs. 4-6. If any species of marine diatom may be considered to be “universally distrib- uted,” it is this representative of Actinoptychus. Probably those forms which lived prior to the Tertiary can be distinguished but those of the Miocene and Pliocene do not seem to differ in any way from those living at the present time. There are many variations which have received names and many more which have not. The species is very common in the Pribilof Island deposits. Arachnoidiscus ehrenbergii Bailey. (Figure 41.) Arachnoidiscus ehrenbergii Bailey, Wortr, Diat. N. Amer., 1894, pl. 91, fig. 2. Brown, Arachnoidiscus, 1933, p. 55, pl. 4, fig. 5. The species is reasonably common in the heavier fractions of almost every Pribilof sample. Arachnoidiscus ornatus was not found in any of the material. Brown’s extensive study of this genus resulted in his recognition of several species and varieties of somewhat questionable utility after mild criticism of earlier authors for useless multiplication of names. However, in the study of west American fossil forms I find great difficulty, after many years of observation, 180 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. in being convinced that there is any escape from allowing wide limits of variation under each of a few species names. I have not found any constant differences between Miocene and living material from western North America. The type locality of the species appears to be Puget Sound, but it is very common along the west American coast. I have collected it from Bering Sea near St. Matthew Island as far south as Maria Madre Island, Mexico. It is also found in most of the California fossil marine diatom deposits which are Miocene or later but the specimens found in the upper Eocene were called Arachnoidiscus indicus, rightly or wrongly. (Hanna, 1927, p. 109.) Arachnoidiscus indicus Ehrenberg. (Figures 77, 80.) Arachnoidiscus indicus EHRENBERG, Ber. Akad. Wiss., Berlin, p. 47, pl. 5, fig. 7 [after Mills], [1854, p. 165, after Mann]. EnRenBerc, Mikrogeologie, 1854, pp. 163-165, 171, pl. 36, C, fig. 34, Noncoury Island, Nicobar Islands, Fossil. Scumuipt, Atlas, Diat., pl. 68, fig. 6, 1886. Brown, Arachnoidiscus, 1933, p. 66, pl. 5, figs. 8, 9. Nicobar Islands, Fossil. The diatoms here identified with the above name were present but not common in the deposit at Tolstoi Point, St. Paul Island. If this identification be correct it is apparently the only record of the species having been found at other than the type locality according to N. E. Brown who made an extended study of the genus. In general, diatomists were inclined to call small specimens A. indicus, which were not so boldly marked as A. ehrenbergii, thus recording the species from a great many widely spread fossil localities. Ehrenberg’s original figure is a very good drawing and our specimens, it would seem, agree sufficiently to be regarded as the same species. However, some of the drawings which others have called A. indicus, but Brown rejected, also seem to be sufficiently close. The original mention of the name Arachnoidiscus indicus and perhaps a description, may have been during 1854 in the Berichte of the Berlin Academy, a copy of which I have not seen. Cited references do not agree. Asteromphalus darwinii Ehrenberg. (Figure 90.) Asteromphalus darwiniti EHRENBERG, Ber. Akad. Wiss. Berlin, 1844, pp. 198, 200, fig. 1. Pelagic “64° N. Lat., 160° W. Long.” Scumupr, Atlas Diat., pl. 38, 1876, fig. 16. “Mon- terey,” Calif. Enrensperc, Mikrog. 1854, pl. 35A, group 21, fig. 4. WoLte, Diat. N. Amer. 1894, pl. 93, figs. 8, 9. “Monterey.” [Upper Miocene. ] Asterolampra darwinii (Ehrenberg), Grevit_E, Trans. Micr. Soc. London, vol. 8, n. s., 1860, p. 116, pl. 4, figs. 12, 13, “Monterey Stone.’ Scuuttze and Karn, Bull. Torrey Bot. Club, vol. 23, 1896, p. 498. “Santa Monica” Calif. [Malaga Cove, Los Angeles Co.] Ehrenberg described and illustrated five species in 1844 which he included in his new genus Asteromphalus: A. darwinii, A. hookerii, A. rossii, A. buchii, and A. humboldti. All of these appear to be one species and it is therefore logical to use the first name. VoL. XXXVIT] HANNA: PRIBILOF FOSSIL DIATOMS 181 Very few specimens of this pelagic genus were found in the Pribilof material, probably in part because of their fragility. The one photographed seems to be close to A. darwinii as illustrated by Greville. It is characterized especially by the few divisions and the structure of the division lines in the central area. This species was selected by Boyer, (1927, p. 72.) as the genotype of Asteromphalus. It, and many other species of the group, have been shifted back and forth by diatomists between Asterolampra and Asteromphalus. The chief difference used to separate the two genera is the presence of one slim radial bar in Asteromphalus. Aulacodiscus laxus (Mann). (Figures 102, 103, 104.) Tripodiscus laxus Mann, Cont. U. S. Nat. Herb., vol. 10, pt. 5, 1907, p. 280, pl. 54, fig. 3. “Station 4029 (U.S.S. Albatross), Bering Sea, 913 fms.” Long search of the heavier fractions of a large sample of material from Tolstoi Point, St. Paul Island by Dr. A. L. Brigger, resulted in the finding of a very few specimens of this rare species. It was submitted by Dr. Brigger to Dr. Joseph F. Burke, who is engaged in a thorough study of Aulacodiscus and the temporary identification was due to him. In order to be certain, however, Dr. Burke submitted a specimen to Dr. Paul Conger of the Smithsonian Institu- tion for comparison with Dr. Mann’s type and the identification was confirmed by him. Aulacodiscus tripartitus Tempére and Brun. (Figures 46, 49, 105.) Aulacodiscus tripartitus TEMPERE and Brun, in Brun and Tempere, Diatomées Fossiles de Japan, Mém. Soc. Phys. D’Hist. Nat. Genéve, vol. 30, no. 9, 1889, p. 21, pl. 4, fig. 3. Scumipt, Atlas, Diat., pl. 169, 1892, figs. 8, 9. “Japan.” No locality was cited in the original descriptions of this species. The authors compared it to A. kilkellyanus Greville from Barbados, A. septus Schmidt from Simbirsk and A. schmidti Witt. Aulacodiscus kilkellyanus appears to be the closest. All of the specimens from Tolstoi Point, St. Paul Island, Alaska, which have been found to date have been studied by Dr. Joseph F. Burke of Staten Island Museum in connection with his review of the genus Awlacodiscus, and it is his opinion that they belong to the species ¢ripartitus. It is a very rare diatom in this Alaska deposit and unfortunately the first one found (fig. 46) was corroded. This gave a false impression of specific characters and it was only after protracted search that additional specimens were found by Dr. A. L. Brigger which showed the true markings (figs. 49, 105). 182 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Biddulphia aurita (Lyngbye), Brébisson and Godey. (Figure 78.) Diatoma aurita LyNcByE, Hydrophytologie Danicae, . . . etc., 1819, p. 182, pl. 62, fig. D. Biddulphia aurita (Lyngbye), BREBISSON AND GopEy, Considerations sur les Diatomées ... etc. 1838, p. 12. ScumipT, Atlas, Diat., pl. 122, 1888, fig. 6. Wore, Diat. N. America, 1894, pl. 96, figs. 8-10. It is probable that not all diatoms which have been given the above name actually belong to the same species. However, the nomenclature is in a state which makes it seem best to adhere to this early name until someone finds time to unravel the tangle. Biddulphia baltzoi Hanna, new species. (Figure 82.) Frustule in end view more slender than in most Biddulphia, divided into three parts, the center one with two strong spines; end processes long and heavy, projecting slightly away from center; surface with heavy, discrete dots, finer in the central section. Length .040 mm. Holotype no. 3639 (CAS) Dept. Geol. Type Coll., from locality no. 36829 (CAS), Tolstoi Point, St. Paul Island, Alaska. The slender shape of this diatom and the coarseness of the surface markings set it aside from any of the other species I have encountered. There is a temp- tation to place it in Hemialus which is a genus of even more slender species and found in large part lower in the Tertiary. For the present it seems best to leave it in Biddulphia because that genus contains so many diverse diatoms. The species is named for Mr. Howard Baltzo of the U. S. Fish and Wildlife Service, who made it possible to secure the 1960 collections at Tolstoi Point and elsewhere on St. Paul Island. Biddulphia roperiana Greville. (Figure 79.) Biddulphia roperiana GREVILLE, Quart. Journ. Micr. Sci., vol. 7, 1859, p. 163, pl. 8, figs. 11-13; Monterey, Calif., living on seaweeds, Calif. guano. Boyer, Proc. Acad. Nat. Sci. Phila- delphia, 1900, p. 700; living on Pacific Coast; Calif. Miocene. The Pribilof Island specimens agree fairly well with the illustrations which have been published. A similar form from the Miocene of New Jersey, (B. cookiana Kain and Schultze, Bull. Torrey Bot. Club, vol. 6, no. 8, 1889, p. 73, pl. 89, fig. 4) seems to be very close. Chaetoceros didymus Ehrenberg. (Figures 62, 97, 98.) Chaetoceros didymus EHRENBERG, Ber. Akad. Wiss. Berlin, 1845 [1846], p. 75. Mikrog. 1854, pl. 35A, group 17, fig. 5; group 18, fig. 4. Hustepr in Schmidt, Atlas, Diat., pl. 323, 1920, fig. 7; pl. 326, 1920, figs. 2, 6, 7. Hustept, Rabenhorst’s Krypt. Flora, Kieselalgen, vol. 7, pt. 1, 1930, p. 688, fig. 390. Vout. XXXVII) HANNA: PRIBILOF FOSSIL DIATOMS 183 The determination of fossil Chaetoceros is difficult because of the dismem- bering of the chains in which they grow. Chaetoceros didymus has been recorded from many wide-spread localities, especially in northern regions and this seems to be the most reliable identification for the Pribilof Island material at present. The drawing, figure 65, has been doubtfully referred to Chaetoceros. Several of these objects were found in the light fractions from Tolstoi Point. Cocconeis antiqua Tempére and Brun. (Figure 48.) Cocconeis antiqua TEMPERE AND BRUN in Brun and Tempére, Diat. Foss. Japan, 1889, p. 32, pl. 8, fig. 5. Sendai and Jedo, Japan. Kanava, Sci. Repts. Tohoku Univ., ser. 2, Geoll., vol. 30, 1959, p. 107, pl. 10, figs. 1, 2 [Reprint of Tempére and Brun] Miocene, Japan. Scum, Atlas, Diat., pl. 191, 1894, figs. 49-52. Cocconeis japonica PANTOcSEK, Diat. Foss. Ungarns, pt. 3, pl. 42, 1893, fig. 582. Our specimens from Tolstoi Point, St. Paul Island, are very close to the figures given by Schmidt. Cocconeis formosa Brun. (Figure 89.) Cocconeis formosa Brun, Diat. Esp. Nouv. Mar. Fossiles ou Pélagique. Mém. Soc. Phys. Hist. Nat. Genéve, vol. 31, pt. 2, 1891, no. 1, p. 16, pl. 18, fig. 6. SrNpa1, Japan, fossil. Indian Ocean. Scumipt, Atlas, Diat., 1894, pl. 193, figs. 42-47. Hokkaido and Sendai, Japan. Pantocsek, Beit. Kennt. Foss. Bac. Ungarns, 1893, pl. 32, fig. 457 [See next reference for locality distribution]. Kanaya, Sci. Repts. Tohoku Univ. ser. 2, Geol., vol. 30, 1959, p. 109, pl. 10, figs. 4, 5. [Reprint of Brun] Miocene, Japan. This beautiful diatom does not seem to be common anywhere. Cocconeis maxima (Grunow). (Figure 31.) Mastogloia maxima Grunow, Verh. Zool.-Bot. Ges., Wien, vol. 13, 1863, p. 136, pl. 4, fig. 1. “Lower valve.” (Mills.) Cocconeis lorenziana GRUNOW, Schmidt, Atlas, Diat., pl. 191, 1894, figs. 28-34. Cocconeis maxima (Grunow), PERAGALLO, H. & M., Diat. Mar. France, pl. 3, 1897, figs. 1-4. Very few specimens referable to Cocconeis were found in any of the deposits studied. The one illustrated here came from cuttings in Navy Well no. 2, at 90-foot depth. It belongs to a group of species of this genus which is very widely dispersed and has many variations. It is marine in habitat and probably was not a contaminating factor in the sample from the well. Schmidt (1894, pl. 191, fig. 54) illustrated a specimen from Hokkaido, Japan, which agrees almost exactly with our specimen. He said that Cleve would refer it to Cocconeis scutellum but he would not. Upon comparison with Ehrenberg’s original figure (1838, pl. 14, fig. 8), it appears that Schmidt was probably correct. 184 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Cocconeis pribilofensis Hanna, new species. (Figure 34.) Valve ovate with a single row of heavy beads around the margin; through the center, where the raphe would be, there are two straight rows of beads the same size as those at the margin; these rows are interrupted near the center by the omission of various numbers of beads. Hotorype no. 3574, (CAS) Dept. Geol. Type Coll. from locality no. 36829 (CAS), Tolstoi Point, St. Paul Island, Alaska. Length, .0302 mm. A reasonably thorough search of the literature has failed to disclose a described species to which this may be referred. Coscinodiscus fimbriatus Ehrenberg. (Figure 81.) Coscinodiscus fimbriatus EHRENBERG. Ber. Akad. Wiss. Berlin, 1844, p. 78. EHRENBERG, Mikrog. 1854, pl. 22, fig. 2. Grunow, Diat. Franz Joseph Land, Denk. Math.-Natur. Cl. Kais. Akad. Wiss. Wien, vol. 48, 1884, p. 74 [22], and var. californica, op. cit. Rattray, Rev. Coscinodiscus, 1890, p. 105 [553]. Husrept, Kieselalgen, vol. 7, pt. 1, 1928, p. 424, fig. 227. It is believed that the peculiar border markings of the disk should be used to distinguish this species from others of similar size. The presence or absence of a coarse rosette of aerolae in the center is not believed to be of sufficient constancy to be relied upon. Hustedt’s figure of 1928 seems to bear this distinction out. The species has been repeatedly reported from upper Miocene strata of Cali- fornia. Coscinodiscus kiitzingii Schmidt. (Figure 7.) Coscinodiscus kiitzingii ScuHMipT, Atlas, Diat., 1878, pl. 57, figs. 17, 18. Hustepr, Kieselalgen, vol, 7, pt. 1, 1928, p. 398, fig. 209. The diatom is fragile and it is difficult to find perfect specimens, although fragments are present on most strewn slides. The beading is small but scattered over the disk. There are a few heavier, irregular dots. The zone of fine border markings, together with the division of the disk into sectors, seems to set the species apart, although Hustedt remarked that it stands between C. excentricus and C. rothii. Coscinodiscus marginatus Ehrenberg. (Figures 1, 2.) Coscinodiscus marginatus EHRENBERG, Abh. Akad. Wiss. Berlin, 1841 [1843], p. 142. Scumipr, Atlas, Diat., pl. 62, 1878, figs. 1-5, 9, 11, 12. Mann, Cont. U. S. Nat. Herb., vol. 10, pt. 5, 1907, p. 253, pl. 49, fig. 2. This large and heavy diatom is one of the most common species in the Pribilof Island deposits. The coarse markings are arranged in no geometric VoL. XXXVIT] HANNA: PRIBILOF FOSSIL DIATOMS 185 order. The margin is wide and transversely marked. Secondary markings are usually present on the coarse aerolae as shown in figure 1. Coscinodiscus oculus-iridis Ehrenberg. (Figure 18.) Coscinodiscus oculus irvidis EHRENBERG, Abh. Akad. Wiss. Berlin, 1839, p. 147. EHRENBERG, Mikrog. 1854, pl. 18, fig. 42; pl. 19, fig. 2; pl. 21, fig. 3. Scmipr, Atlas, Diat., pl. 63, 1878, figs. 6, 7, 9. Hustept, Kieselalgen, vol. 7, pt. 1, 1928, p. 454, fig. 252. This large diatom is widely distributed in present seas and has a long geo- logical record. It is common in the heavy fractions of cleaned material from St. Paul Island. Coscinodiscus pustulatus Mann. (Figures 12, 19-24.) Coscinodiscus pustulatus MANN, Cont. U. S. Nat. Herb., vol. 10, pt. 5, 1907, p. 257, pl. 48, fig. 3. “Bering Sea.” Hanna, Amer. Journ. Sci., vol. 48, 1919, p. 224. Coscinodiscus sp. Hanna, 1951, Bull. Calif. State Division of Mines 154, p. 283, figs. 2, 3, St. Paul Island, Alaska, Pliocene. This is by far the most easily recognized diatom in all of the Pribilof deposits studied in connection with this report. The species was originally described from a depth of 1866 fathoms in Bering Sea and it is entirely possible that it came from an exposed subsurface outcrop of strata of Pliocene age. Nothing like it has been seen in any of the collections of living marine diatoms I have made in the Arctic and subarctic. A selection of many variations has been illustrated. The two valves differ. One approaches a hemisphere; the other is roundly conical like a Korean hat. Variation in size of beads is very great. It is rare that a geometrically perfect arrangement is found. Another similar species is C. nano-lineatus Mann (1925, p. 68, pl. 14, fig. 4) from the Philippine Islands. It is likewise convex, contrary to C. lineatus which is flat, but does not flare out laterally as does C. pustulatus. Coscinodiscus radiatus Ehrenberg. (Figures 4, 8, 17.) Coscinodiscus radiatus EHRENBERG, Ber. Akad. Wiss. Berlin, 1839, p. 148, pl. 3, fig. 1 a-c. [From Mills.]. Enrenserc, Mikrog., 1854, pl. 19, fig. 1. Wore, Diat. N. Amer., 1894, pl. 81, fig. 7. Hanna anp Grant, Proc. Calif. Acad. Sci., 4th ser., Wl, 5, wo), 7, Iewey. p.142, pl. 15, fig. 12. Hustept, Kieselalgen, vol. 7, pt. 1, 1928, p. 420, fig. 225. The specimens illustrated are from the sediments at Tolstoi Point, St. Paul Island, locality no. 36829 (CAS). They are representative of many which were recovered in the separations of coarser species in other localities. Almost always the markings are arranged in geometrical order with rarely some slight imper- 186 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. fection. The center is slightly depressed. As species in Coscinodiscus go, this one may be considered as lacking in great variability. At least, that opinion has been formed from the study of material from this limited area. However, it will be found that many names have been given to variations of the species else- where. (see Mills, Index, 1933-1935). Coscinodiscus rothi (Ehrenberg). (Figure 36.) Heterostephania rothii EHRENBERG, Mikrog., 1854, pl. 35A, group 13B, figs. 4, 5. Coscino- discus symmetricus GREVILLE, Schmidt, Atlas, Diat., pl. 57, 1878, figs. 25-27. Coscino- discus rothii (Ehrenberg), Grunow im Schneider, Naturwiss. Beitr. Kennt. Krauk- auslander, 1878, p. 125. Hustept, Kieselalgen, vol. 7, pt. 1, 1928, p. 400, fig. 211. In the deposit at Tolstoi Point, St. Paul Island, there are many small and highly variable diatoms which belong to this general group. Because of the inadequacy of many early figures, such as those on plate 57 of Schmidt’s Atlas, species are difficult to separate, more so perhaps than any other part of this difficult genus. Coscinodiscus undulosus Mann. (Figure 3.) Coscinodiseus undulosus Mann, Cont. U. S. Nat. Herb., vol. 10, pt. 5, 1907, p. 259, pl. 49 ee il This species was found in the first sample studied in 1916 from Tolstoi Point, St. Paul Island, and was identified by Dr. Mann among the slides I made then. He had found it in only one dredging of the U.S.S. Albatross, Station 3526 in Bering Sea. In later work it was found frequently in material from the Navy Well, Black Bluff, and Zapadni Point on St. Paul Island. Cosmiodiscus insignis Jousé. Gigures Os Os i 30532.) Cosmiodiscus insignis JousE, Diatomeae Marinae Mioceni et Plioceni ex Oriente Extreme. Notula Systematicae e Sectione Cryptoganiea Instituti Botanici Nomine V. L. Komarovii Academiae Scientiarum USSR, 1961, p. 67. Valve strongly convex with radial rows of beads; these are heaviest toward the center but the center is usually a large hyaline area without markings or, if present, very sparingly distributed. The margin is heavy and hyaline; just inside it there is a series of slightly elongate spines or low ridges. This species and Coscinodiscus pustulatus Mann are extremely abundant in all of the fossil bearing shales found on the Pribilofs, and, it would appear, they can be used as characteristic fossils for this northern horizon. The highest elevation in which the formation was found is about 50 feet above sea level, but the fact that it extends downward 400 feet shows that it is a formation of con- VoL. XXXVII] HANNA: PRIBILOF FOSSIL DIATOMS 187 siderable magnitude. The species was described by Mrs. Jousé from lower Pliocene strata of Sachalin Island and apparently has been found living in the littoral zone of the Kamtschatka Peninsula. She compared the species to Coscinodiscus (Cestodiscus) intersectus Brun (1891, p. 22, pl. 20, fig. 5) from the Miocene (?) of Sendai, Japan. This seems to be a flatter diatom and with a much smaller hyaline central area, although C. insignis at the Pribilof localities is a highly variable species. It is interesting to note that Mrs. Donahue has recently found this species in the Pleistocene of the Pacific Sector of the southern ocean. (Donahue, 1967, p55, pl. 1, figs. e, f.). Cymatotheca weissflogii (Grunow), Hendey. (Figures 15, 16.) E{[uoda] (genus novum?) weissflogii GRUNow in Van Heurck, Syn. Diat. Belgique, pl. 126, 1883, fig. 13. West Africa, Bengal, China, Brazil. Euodia weissflogii (GRUNOW) in Leuduger-Fortmorel, 1898. Diat. Mar. Cote Afrique, p. 18, pies ties 13> Euodia ratabouli Brun in Leuduger-Fortmorel, 1898. Diatomées Marines de la Cote Occi- dentale d’Afrique, p. 24, pl. 5, fig. 3. Coscinodiscus asymmetricus MeEIstErR, Kieselalgen aus Asien, 1932, p. 19, pl. 4, figs. 32, 33. Belewan. Hemidiscus weissflogi (Grunow) Hustept, 1953. Marine littoral diatoms of Beaufort, North Carolina, Duke Univ., p. 11, pl. 1, figs. 6, 7. Found on piles in the harbor. Cymathotheca weissflogii (Grunow) HENpEy, Journ. Roy. Micr. Soc., vol. 77, 1957 [1958], p. 41, pl. 5, fig. 9. Vorct, Journ. Roy. Micr. Soc., vol. 78, nos. 3-4, 1958 [1960], p. 93, joll, i, weal, We The literature indicates that this diatom is widely distributed in the southern hemisphere and especially along the west coast of Africa and the southeast coast of Asia. It is surprising to find it fairly common in the Pliocene deposits of the Pribilof Islands. The species was not widely known among diatomists until details of its structure were studied by Hendey and published in 1958. Voigt added much additional distributional information in 1960 and described a variety and an additional species, C. minima. The Pribilof specimens do not vary as widely in shape and sculpture as has been described for the recent collections, yet their characters are not very constant. There is an allied species (undescribed) in Upper Miocene diatomite at Monterey, California. This one is circular in shape with the two halves in different planes as in C. weissflogii, but the sculpture is essentially as in many Coscinodiscus. Another similar species is “Coscinodiscus temperei” Brun, (1889, p. 33, pl. 8, fig. 2) from “Sendai, Japan.” This was illustrated again by Schmidt, (1891, pl. 163, fig. 9). Jousé recorded it from eastern USSR fossil deposits in 1959, (p. 47, pl. 2, fig. 7). 188 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Dicladia capreola Ehrenberg. (Figure 63.) Dicladia capreola EHRENBERG, Mikrog. 1854, pl. 35A, group 18, figs. 5, 8. WoLtz, Diat. N. Amer. 1894, pl. 64, figs. 5-7. Dicladia pylea HANNA AND Grant, Proc. Calif. Acad. Sci., 4th ser., vol. 15, no. 2, 1926, p. 142, pl. 16, figs. 4, 5. Maria Madre Island, Mexico, Miocene. Objects such as this are common in the deposit at Tolstoi Point, St. Paul Island, although in order to recover them special care needs to be taken in separating the light fraction during preparation. Diploneis bombus Ehrenberg. (Figure 14.) Diploneis bombus Eurenserc, Mikrog. 1854, pl. 19, fig. 31. Scumupt, Atlas, Diat., pl. 13, 1875, figs. 4-6; pl. 69, figs. 28, 29. Navicula bombus densistriata ScHMmuipT, Atlas, Diat., pl. 13, 1875, figs. 11, 12. “California.” Navicula densistriata (Schmidt), HANNA AND Grant, Proc. Calif. Acad. Sci., 4th ser., vol. 15, no. 2, 1926, p. 150, pl. 17, figs. 8-10. “Maria Madre Island, Mexico.” Very few diatoms of this form were found in the Pribilof Island deposits. The one illustrated came from 90 feet depth in Navy Well no. 2, St. Paul Island. Diploneis ornata Schmidt. (Figure 13.) Navicula ornata Scumipt, Atlas, Diat., pl. 69, 1881, fig. 5. “Monterey.” Wotte, Diat. N. Amer., 1894, pl. 15, fig. 20. This particular form of Diploneis has a long geological range; that is, from middle Miocene to the present. It is rare in the Pliocene deposits of the Pribilof Islands, the example illustrated having come from locality no. 36829, Tolstoi Point, St. Paul Island. Material from this locality was searched more extensively than any of the others. Dossetia temperei Azpeitia. (Figures 59, 60.) Dossetia temperei AzpritiA, Asoc. Espanola Progreso de las Ciencias; Congreso de Zaragoza, vol. 4, sect. 4a, 1911, p. 203, pl. 9, figs. 3, 7. “Serrata de Lorca,” Spain. Xanthiopyxis lacera Forti, Cont. Diat. XIII. Atti d. Reale Ist. Veneto di Sci. Lett. Art., vol. 72, pt. 2, 1913, p. 21 [1555], pl. 2, [12], figs. 14-18. Miocene, Italy. KoroTKEviEz, Species novae Diatomacearum e Neogeno Peninsulae Kamczatka, Akad. Nauk SSSR, 1964, p. 108, pl. 3, figs. 3, 4. Diatoms of this general shape are common in the finer fractions from most marine deposits of west American Miocene and Pliocene age. There appear to be more than one species. Those found in some Pliocene deposits of California lack VoL. XXXVIT] HANNA: PRIBILOF FOSSIL DIATOMS 189 the marginal fringe. Azpeitia considered these diatoms to be sufficiently distinct to warrant generic separation, a decision which seems to have been well taken. Hemidiscus cuneiformis Wallich. (Figure 35.) Hemidiscus cuneiformis Wauuicu, Trans. Micr. Sci. London, ns., vol. 8, 1860, p. 42, pl. 2, figs. 3, 4. Husrept, Kieselalgen, vol. 7, pt. 1, 1930, p. 904, fig. 542. Henpey, British Coastal Algae, pt. 5, Bacill. 1964, p. 94, pl. 22, fig. 9. Hemidiscus simplicissimus HANNA AND GRANT, Proc. Calif. Acad. Sci., 4th ser., vol. 15, no. 2, 1926, p. 147, pl. 16, fig. 13. This widely distributed plankton species is confined to tropical and temperate waters at present but it was common in Bering Sea at the time of deposition of the sediments on St. Paul Island. The latter are rather finely marked as com- pared to living forms, but this difference is so slight that it probably does not have taxonomic significance. Hemidiscus rotundus Janisch. (Figure 86.) Hemidiscus rotundus JANISCH, Diat. gesam. Reise der “Gazelle,” 1874-1876 [1888], pl. 1, fig. 6. Hustept in Schmidt, Atlas, Diat., pl. 438, 1940, fig. 1. The report by Janisch was never actually published, although a few copies of the plates (except plates 10 and 14) were distributed to selected diatomists. The California Academy of Sciences’ copy appears to be the one described in detail by Junk (1926-1936, pp. 153, 217-218). It is accompanied by a type- written list for each plate, showing the identification of each figure by Janisch on one side and at times by Rattray or Norman on the other. In the case of H. rotundus, Rattray’s identification was “Coscin: curvatus var. recta Rattr.” Hemidiscus bicurvans Barker and Meakin (1949, p. 302, pl. 38, fig. 7) from the Eocene of “‘Conset, Barbados” may be ancestral to H. rotundus. It seems to lack the marginal ocellus according to the figure. Hercotheca inermis Mann. (Figure 84.) Hercotheca inermis Mann, U. S. Nat. Mus. Bull. 100, vol. 6, pt. 1, 1925, p. 82, pl. 18, figs. 1, 2. Philippine Islands, living. The St. Paul Island specimen seems clearly to be allied with another species of this genus, viz., H. mammalaris Ehrenberg. The presence in the case of the Alaska specimen of two frustules attached in chain fashion is a strong argument against such objects being endocysts of some very unlike diatoms. The name H. brevispina Grunow has not been formally published and cannot be identified except by referring to a slide; it is strictly a nomen nudum. Mann offered good evidence against such objects being endocysts. 190 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Lithodesmium undulatum Ehrenberg. (Figure 91.) Lithodesmium undulatum EHRENBERG, Abh. Akad. Wiss. Berlin, 1840, p. 75, pl. 4, fig. 13. Scumipt, Atlas, Diat., pl. 151, 1890, fig. 41; pl. 152, 1890, figs. 1-3. The variation is so great in members of this pelagic genus that it seems best to refer our Pribilof Island species to the oldest name. A later genus name, Ditylum Bailey, seems to be a synonym. Melosira clavigera Grunow. (Figures 52, 54.) Melosira clavigera Grunow in Schmidt, Atlas, Diat., pl. 74, 1876, figs. 13-15. “Monterey,” [Calif.; Miocene.]. Grunow in Van Heurck, Syn. Diat. Belgique, 1882, pl. 91, figs. 1, 2. “Monterey and San Francisco,” California. This very common diatom is found in many west American fossil deposits. The radial bars vary in fineness and length somewhat but usually not enough to cause the identification to be questioned. It resembles Melosira sol. Melosira sulcata (Ehrenberg). (Figures 50, 51, 53.) Gallionella sulcata EHRENBERG, Ber. Akad. Wiss. Berlin, 1837, p. 61. EHRENBERG, Infusion- thierken, 1838, p. 170, pl. 21, fig. 5. “Oran,” North Africa. EHRENBERG, Abh. Akad. Wiss. Berlin, 1840, pl. 3, fig. 5 [Reference after Kiitzing, 1844]. This species of Melosira has characteristic heavy wedge-shaped radial bars; they are highly variable, but seem always to be present. Individuals were common in all of the Pribilof deposits which were studied in detail. Figure 50, is from Einahnuhto Bluffs, and figure 51 is from Tolstoi Point, both on St. Paul Island, Alaska. These Pribilof Island specimens resemble most Cyclotella antiqua, W. Smith, (1853, p. 28, pl. 5, fig. 49) as illustrated by Grunow in Van Heurck, (1882, pl. 92, fig. 1). That species, however, is very small and seems always to be found in fresh water or freshwater deposits. Navicula semen Ehrenberg. (Figure 96.) Navicula semen ERENBERG, Abh. Akad. Wiss. Berlin, 1843, pl. 1, group 2, fig. 17. Hustepr im Schmidt, Atlas, Diat., pl. 299, 1913, figs. 18-20. This freshwater diatom is widely distributed in the northern hemisphere. Its presence in the marine deposit at Tolstoi Point, St. Paul Island, Alaska, is probably due to drift from rivers. The above identification of the species was Vor. XXXVIT] HANNA: PRIBILOF FOSSIL DIATOMS 191 suggested by Dr. Ruth Patrick from an examination of the photograph. Cleve (1894, p. 139) stated that Ehrenberg’s original figure was not recognizable. Apparently the species is found living only on rare occasions although it is reported to be common in Pleistocene deposits. Pinnularia lata (Brébisson). (Figure 33.) Frustulia lata BreBisson, Mem. Soc. Sci. Nat. Falaise, 1838, p. 18. Navicula lata (Brébisson), W. Smirn, Syn. Brit. Diat., vol. 1, 1853, p. 55, pl. 18, fig. 167. Mann, Cont. U.S. Nat. Herb., vol. 10, pt. 5, 1907, p. 346. Published records indicate that this is an exceedingly widely distributed species. Many varities and forms have been named and nearly all refer to living material. Since it is strictly a freshwater species, I hesitate to include it in this report. It was found only in cuttings (ditch samples) from a depth of 90 feet in Navy Well no. 2 (locality no. 1322, CAS). It could very well have been present in surface waters used in bailing the well and it is included solely to illustrate the caution which should always be taken in the study of “ditch” or “bailer” samples. Pinnularia ruttneri Hustedt. (Figure 101.) Pinnularia ruttnerti Hustept in Schmidt, Atlas, Diat., pl. 390, 1934, figs. 6-8. The presence of scattered freshwater diatoms in marine deposits of late Tertiary and Quarternary ages is not unusual and can be attributed to material carried out to sea from rivers. This species is one which was originally found living in lakes of Sumatra and the identification was suggested by Dr. Ruth Patrick from an examination of the photograph. A similar species, widely dis- tributed in the northern hemisphere is P. streptoraphe Cleve (see Hustedt, 1930, p. 337, fig. 620) which differs in some details. Pseudopyxilla dubia (Grunow) Forti. (Figures 66, 68.) Pyxilla dubia Grunow in Van Heurck, Syn. Diat. Belgique, pl. 83, 1882, figs. 7, 8; pl. 83 bis, fig. 12 “Monterey” upper Miocene. Wotte, Diat. N. Amer., 1894, pl. 65, fig. 24. Pseudopyxilla dubia (Grunow), Fort1, Nuova Notarisia, vol. 20, 1909, p. 12, pl. 1, figs. 1-3. Several simple forms such as those illustrated are common in the light frac- tions of Pribilof Island samples. Whether they should all be called the same or if two or more species should be recognized remains to be determined when a careful study can be made from several localities. Forti, 1909, described P. tempereana in the paper cited usually as “Pilla.” The illustrations do not differ greatly from P. dubia. 192 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Pyxilla americana Grunow. (Figure 61.) Pyxilla americana GRuNoW in Van Heurck, Syn. Diat. Belgique, 1882, pl. 83 bis, figs. 1-3. Wo tte, Diat. N. Amer., 1894, pl. 65, figs. 8, 17, 18. Objects similar to this one are common in light fractions of many preparations of fossil diatoms. Their exact nature is not known in most cases and have some- times been considered to be “‘resting spore cases” of such diatoms as Rhizosolenia. They were found in most of the deposits on St. Paul Island, Alaska. Rhabdonema crozieri (Ehrenberg), Grunow. (Figure 94.) Striatella crozieri EHRENBERG, Ber. Akad. Wiss. Berlin, 1854, pp. 524, 529. EHRENBERG, Mikrog., 1854, pl. 38A, group 23, figs. 14-16. Rhabdonema crozierii (Ehrenberg), GRuNow, Verh. Zool. Bot. Ges. Wien, vol. 12, 1862, p. 423. FricKE in Scumupt, Atlas, Diat., pl. 220, 1889, figs. 4-9; pl. 221, fig. 1. This species was first found at ‘“‘Assistance Bay,’’ Cornwallis Island, Canadian Arctic, N. Lat. 73°50’. Our specimen is slightly wider in proportion to length than Ehrenberg’s original figures indicate, but this is not believed to be significant in this group of diatoms. Mills, (1934, p. 1386) suggested that R. crozierit was the same as R. arcuatum Kiitzing and that Ralfs’ interpretation was R. adri- aticum Kitzing. Upon comparison of original illustrations neither of these interpretations seem very sound. Rhabdonema japonicum Tempére and Brun. (Figures 47, 92, 93, 95.) Rhabdonema japonicum TEMPERE and Brun, Mém. Soc. Phys. d’Hist. Nat. Geneve, vol. 30, no. 9, 1889, p. 53, pl. 1, fig. 6. Fricke im Schmidt, Atlas, Diat., pl. 230, 1899, figs. 18-21. Fricke’s excellent drawings on plates 217-221 of the Atlas show several of the named varieties of this species. They span the series from St. Paul Island, Alaska, and probably should not be considered as separate taxa. The cross bars of the species bear a single row of prominent dots and the valves are not usually curved in edge view. These characters serve to separate members of the genus from the Entopyla-Gyphria complex. Rhaphoneis amphiceros Ehrenberg. (Figures 29, 55, 56.) Rhaphoneis amphiceros EHRENBERG, Ber. Akad. Wiss. Berlin, 1844, p. 87. EHRENBERG, Mikrog., 1854, pl. 33, group 14, fig. 22; group 15, fig. 20. Wore, Diat. N. Amer., 1894, pl. 37, fig. 20. HANNA AND GRANT, Proc. Calif. Acad. Sci., 4th ser., vol. 15, no. 2, 1926, p. 165, pl. 20, fig. 8. Hanna, Proc. Calif. Acad. Sci., 4th ser., vol. 20, no. 6, 1932, p. 211, pl. 15, figs. 3-5. Most of the specimens of Rkaphoneis amphiceros found in the Pribilof Island deposits were the short sparsely marked forms such as those illustrated by figures VoL. XXXVIT] HANNA: PRIBILOF FOSSIL DIATOMS 193 55 and 56. Many published figures show much finer markings for this species, but all of the genus are so highly variable that it is believed a measure of stability will be effected by assuming rather wide limits to variation. The diatom illus- trated by figure 29 is confusing. The print looks as if two exposures had been made with the negative offset, but this is not the case. It is believed that this is a nearly typical example of R. amphiceros but in a much corroded condition. A somewhat similar situation was found in Aulacodiscus tripartitus, figure, 46. Rhaphoneis lancettula Grunow. (Figure 57.) Rhaphoneis lancettula GRUNOW in PANTOCSEK, Beit. Kennt. Foss. Bac. Ungarns, pt. 1, 1886, p. 35, pl. 27, fig. 271 “Richmond, Petersburgh, Naparima.” (Grunow.) Rhaphoneis lancettula var. jutlandica GruNowW, Pantocsek, Beit. Kennt. Foss. Bac. Ungarns, pt. 1, 1886, p. 35, pl. 30, fig. 321. “Mors Jutlandiae.” Raphoneis asiatica BRuN, Mém. Soc. Phys. d’Hist. Nat. Geneve, vol. 30, no. 9, 1889, p. 51, pl. 1, fig. 8. “Calcaire de Yédo [Japan] et dépot d’Onianino (Russie).” Several excellent specimens of this greatly elongated representative of Rhaphoneis were found in the Pribilof Island deposits. According to the draw- ings, R. asiatica differs from the earlier-named one only by having fewer dots on the surface of the valve. The species dates back to the Eocene if Pantocsek’s record may be accepted. Rhaphoneis nitida (Gregory). (Figure 27.) Cocconeis nitida Grecory, Trans. Roy. Soc. Edinburgh, vol. 21, pt. 4, 1857, p. 20, pl. 1, fig. 26. Hustept, Kieselalgen, vol. 7, pt. 2, lief. 2, p. 177, fig. 683a. It would seem that the very coarse beads, each separated by a considerable space would make this a very easily identifiable species. This has not been the case. The name selected above is as satisfactory as any I have been able to get, but a closer match than this would be preferable. Rhaphoneis rhombus Ehrenberg. (Figure 83.) Rhaphoneis rhombus EHRENBERG, Ber. Akad. Wiss. Berlin, 1844, p. 87. EHRENBERG, Mikrog., 1854, pl. 33, group 13, fig. 19 (not pl. 18, figs. 84, 85). Hanna, Proc. Calif. Acad. Sci., Ath ser., vol. 20, no. 6, 1932, p. 212. Louman, U. S. Geol. Surv. Prof. Ppr. 189-C, 1938, 10) We This species was not common in any of the Pribilof Island samples studied. The last two papers cited above indicate some of the problems involved in working with this group of Rkaphoneis. Stephanopyxis appendiculata Ehrenberg. (Figures 58, 99, 100.) Stephanopyxis appendiculata Enrenserc, Mikrog., 1854, pl. 18, fig. 13b. Scumipt, Atlas, Diat., pl. 130, 1888, fig. 34. Wortr, Diat. N. Amer., 1894, pl. 62, figs. 12-15. 194 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. The spines are often missing on the crown of specimens found in the Pribilof Island deposits. They have probably been broken either in cleaning or in the compaction of the shale after deposition. Otherwise the species seems to be typical of many localities and with an extensive geological range. Many varietal names have been given to diatoms of this species and S. turris, but most of them are based upon what seem to be very slight differences. Unless such variations have some constant biological, geographical, or geological significance, it seems best to use the earliest name for the group. This is not to imply that variation per se should not be studied, because it is fundamental to all studies of biology and paleontology, especially when classification is involved. Thalassionema nitzschoides (Grunow). (Figure 37.) Synedra nitzschoides GRUNOW, Verh. Zool.-Bot. Ges. Wien, vol. 12, 1862, p. 403, pl. 8, fig. 18. This small diatom was found to be very abundant in all of the Pribilof Island deposits when the finer materials were closely examined. The only mark- ings on the valve consist of a single row of beads around the border. Length varies considerably but in this region general shape is quite consistent. There are no cross bars or pseudoraphe, this being the main reason for recognizing the diatom as distinct from members of the genus Synedra in which it was originally placed. When the genus Thalassionema was first proposed (Grunow in Van Heurck, 1881, pl. 43, figs. 7-10) the name was printed in very small type under the heading ‘“Thalassiothrix? ? nitzschoides.” No other species was included but several “varieties” have been. Some confusion exists in the literature as to the application of the names Thalassionema, Thalassiosira, and Thalassiothrix. Thalassiosira is a monotypic genus formed by Cleve (1873, p. 6) for the species nordenskioldit (p. 7, pl. 1, fig. 1), from “Davis Strait,’ a circular diatom. Thalassiosira punctata Jousé. (Figures 5, 6.) Thalassiosira punctata Joust. The main phases in the development of the flora of marine diatoms in the far eastern seas of the USSR at the end of the Tertiary, and during the Quaternary period. Akad. Nauk. SSSR. Bot. Inst., vol. 44, no. 1, 1959, pp. 44-55, pl. 4, fig. 17. Joust, Diatomeae Marinae Mioceni et Plioceni ex Oriente extremo. Notulae Systematicae e Sectione Cryptogramica, Instituti Botanici Nomine V. L. Kromarovii Academiae Scientiarum USSR, vol. 14, 1961, p. 64, pl. 1, figs. 7, 8; pl. 3, fig. 3.) This very distinct diatom was found frequently in the earliest samples from St. Paul Island, which were studied and it is common in most of the material which has been prepared. At first it might be suspected that this is a freak or a superposition of one diatom on another, but this is not the case. When picking out specimens from a dry strewn slide, the species is very conspicuous owing to its deep brown prismatic VoL. XXXVITI] HANNA: PRIBILOF FOSSIL DIATOMS 195 color. Mounted specimens were brought to Washington in 1917, and Dr. Albert Mann, who studied them at some length, was unable to assign them definitely to any known species. Triceratium condecorum Brightwell. (Figures 42, 88.) Triceratium condecorum BRIGHTWELL, Quart. Journ. Micr. Sci., vol. 1, 1853, p. 250, pl. 4, fig. 12. Scumut, Atlas, Diat., pl. 76, 1882, fig. 27. Hanwa, Proc. Calif. Acad. Sci., 4th Sele pVOlNZO moon 19320 p. 221e pl. 17, tgs: 1 3) It is believed that the Pribilof Island diatoms illustrated by figures 42 and 88 belong to this widely distributed species. It goes back to middle Miocene. There is considerable variation in the coarseness of the surface markings but the absence of spines or processes in the angles is a constant character. Triceratium montereyi Brightwell. (Figure 43.) Triceratium montereyi BRIGHTWELL, Quart. Journ. Micr. Sci., vol. 1, 1853, p. 251, pl. 4, fig. 8. Scumipt, Atlas, Diat., pl. 150, 1890, fig. 21, [See pl. 153 for name]; pl. 159, fig. 7; pl. 165, fig. 4. The specimens from the Pribilof Islands are very close to those found in the vicinity of Monterey, California, both living and fossil. Triceratium validum Grunow. (Figure 85.) Triceratium validum GRruNow in Schmidt, Atlas, Diat., pl. 94, 1886, fig. 5. “Santa Monica” [= Malaga Cove, Los Angeles County, California]. The diatom from Tolstoi Point, St. Paul Island differs somewhat from available figures of T. validum; this specimen has much more crowded surface markings. The original figure in the Atlas shows very scattered beads. In spite of this difference it seems best not to separate the Alaska form until more specimens can be made available for study. The name “T. tripolaris” was given to a similar form from Japan, Miocene by Tempére and Brun (1889, p. 66, pl. 6, fig. 7). It was also used by Schmidt, (1891, pl. 166, figs. 1, 2). This has crowded surface markings and a central zone where they are very scattered. Xanthiopyxis globosa Ehrenberg. (Figure 74.) Xanthiopyxis globosa EHRENBERG, Ber. Akad. Wiss. Berlin, 1844, p. 273. Forti, Cont. Diat. XIII, Atti R. Inst. Veneto, Sci. Lett. Art., 1913, vol. 72, pt. 2, p. 1556 (22) oll, 4 (2). figs. 39-49. Hanna, Proc. Calif. Acad. Sci., 4th ser., vol. 20, no. 6, 1932, p. 224; pl. 18, fig. 3. Sharktooth Hill, California, Middle Miocene. Apparently no named illustration of this common species appeared from 1844 until 1913 when Forti gave good photographs of Miocene specimens from 196 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Italy, unless the diatom which Azpeitia (1911, p. 207, pl. 11, fig. 3.) listed as Hercotheca? caput-medusae Azpeitia can be the same. The spines appear on his photograph to be confined to the marginal area whereas they cover the disk in X. globosa. Omphalotheca californica Hanna, (1927, p. 117, pl. 20, figs. 6, 7; 1930, p. 192) is a much more symmetrical form. It came from upper Eocene. There is considerable confusion among generic names which have been given to diatoms of this general form. Xanthiopyxis dates from 1844, (Ehrenberg, p. 264) where the name appears as a subgenus of Pyxidicula. Four species were included on page 273, under the names, “alata,” “globosa,” “oblonga,” “constricta.” Bailey, (1845, p. 327, pl. 4, fig. 14) illustrated one of these, ‘oblonga,’ and it should therefore be the genotype. It was 10 years before Omphalotheca appeared (Ehrenberg, 1854, p. 132, pl. 35A, IX, fig. 4) for Omphalotheca hispida. The species, O. ampliata and O. laevis were included in the genus, but have not been illustrated. The spines of the specimen illustrated here are long and sharply pointed. The species suggests Liradiscus minutus which Greville (1865, p. 47, fig. 6) recorded from Barbados, but in that one the spines are quite short. Xanthiopyxis lohmani Hanna, new species. (Figure 69.) Valve a long symmetrically oval shape with a single row of low spines forming a slightly broader oval on the disk. Length .0306 mm. Holotype no. 3609 (CAS), Dept. of Geol. Type Coll., from locality no. 36829 (CAS), Tolstoi Point, St. Paul Island, Alaska. While this species was found rarely in the finer fractions from locality no. 36829 (CAS), it, or a closely similar species, has been noted in great abundance in deposits of Pliocene age in California, notably those exposed on Harris Grade, between Santa Maria and Lompoc, Santa Barbara County. The central ring of spines seems to be a constant character, although they are not always as regularly spaced as in the Alaska specimens. Xanthiopyxis ovalis Lohman. (Figures 64, 70.) Xanthiopyxis ovalis Louman, U. S. Geol. Surv. Prof. Paper 189C, 1938, py 91> plZOshieaor oll, BAS vines, 112. This distinctive and not highly variable diatom was common in the lighter fractions from Tolstoi Point, St. Paul Island. In this case the density of the spines on the disk varies somewhat but, as Lohman pointed out, they do not extend outward over the border and the disks are symmetrically oval. Lohman also indicated the impossibility of assigning species of Xanthiopyxis to species of Chaetoceros as spore cases during studies of deposits of fossil material. The Vor. XXXVIT] HANNA: PRIBILOF FOSSIL DIATOMS 197 assumption that such objects are spores is not warranted at this time and even if it should be found to be true with some living individuals, this should have no effect whatsoever upon the use of the fossils in stratigraphy and the use of the names assigned to them. So far as is known X. ovalis is confined to beds of Pliocene age. Xanthiopyxis umbonata Greville. (Figure 87.) Xanthiopyxis umbonata GREVILLE, Trans. Micro. Soc. London, n-s., vol. 14, 1866, p. 2, pl. 1, fig. 5. Monterey, California. Woxte, Diat. N. Amer., 1894, pl. 74, fig. 16. The species is rare in the collections made at the Pribilof Islands. It was originally described from upper Miocene diatomite from Monterey, California, and a study of many specimens from there failed to disclose any which have the disk so completely covered with spines as the one here illustrated. The Monterey specimens have a hyaline zone of variable width inside the border before spines appear. Nevertheless, it seems best to refer the Alaska material to that species. Xanthiopyxis cingulatus Ehrenberg (Hanna and Grant, 1926, p. 169, pl. 21, fig. 9) from the upper Miocene of Maria Madre Island, Mexico, has spines scat- tered sparsely over the entire disk but in that species a series also projects outwardly from the margin. SILICOFLAGELLATA AND EBRIATA The members of these groups have siliceous skeletons and therefore become concentrated along with diatoms, in the preparation of fossil materials. They are small in size and rare in present seas. Many of the genera are extinct. The oldest known forms are late Cretaceous in age. Maximum development seems to have been reached in the Miocene. The two groups differ in that the Silicoflagellata are said to have hollow skeletal struc- tures whereas these are solid in the Ebriata. Currently the first is placed in the plant kingdom and the latter in the animal kingdom, a distinction of somewhat questionable value so far as applied paleontology is concerned. Members of both groups have definite stratigraphic value. They are placed in the kingdom Protista along with Diatomaceae, Foraminifera, and other groups in the Treatise on Invertebrate Paleontology currently being published under the editorship of R. C. Moore. Three species of Silicoflagata have been recognized in the Pribilof Island deposits and there is one Ebriata, Ebriopsis antiqua (Schulz). SILICOFLAGELLATA Since members of this group are normally present in most collections of cleaned marine, fossil diatoms, it is convenient to select representatives of the 198 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. various species during the preparation of specimen slides. Some of the forms have very significant stratigraphic value. Most of the genera are extinct. In the Pribilof Island collections only one species, Ebriopsis antiqua Schulz, belonging to the Ebriata was found. It apparently is confined to the Miocene and Pliocene. Dictyocha fibula Ehrenberg. (Figure 75.) Dictyocha fibula EHRENBERG, Mikrog., 1854, pl. 18, figs. 54, 55; pl. 20, fig. 45. GEMEINHARDT, Krypt.-Fl., vol. 10, pt. 2, Silicoflagellatae, 1930, p. 47, fig. 39. The species occurs with Distephanus and Ebriopsis in about equal abundance in the finer fractions of preparations from nearly all of the Pribilof Island diatom- bearing deposits. Distephanus speculum (Ehrenberg). (Figures 71, 72.) Dictyocha speculum EHRENBERG, Mikrog., 1854, pl. 18, figs. 5-7. Distephanus speculum (Ehrenberg), GEMEINHARDT, Rabenhorst’s Kryptogamen-Flora von Deutschland, Osterreich und der Schweiz, vol. 10, pt. 2, Silicoflagellatae, 1930, p. 61, figs. 53 a-h. The three genera of silicoflagellates are fairly common in the lighter separa- tions of material from most of the Pribilof Island localities. This species is somewhat more complicated than the preceding and some modern students have considered the two to belong to the same genus. Mesocena corona Hanna, new species. (Figure 73.) This is a heavy circular ring with 12 strong, sharp spines. These point alternately slight upward and downward from the central horizontal plane of the ring. Diameter .036 mm. HoLotype no. 3611 (CAS), Dept. Geol. Type Coll. from locality no. 36829 (CAS), Tolstoi Point, St. Paul Island, Alaska. Pliocene. This is much heavier than any other species of Mesocena I have seen. Ebriopsis antiqua (Schulz), Hovasse. (Figure 67.) Ebria antiqua Scuuttz, Bot. Archiv, vol. 21, pt. 2, 1928, p. 273, fig. 69. Miocene, Japan, California, Maryland. Ebriopsis antiqua (Schulz), Hovasse, Bull. Soc. Zool. France, vol. 57, 1932, p. 120, fig. 1. “Santa Monica.” [California, Miocene. ] Usually all three genera of the common silicoflagellates are found, a few on each strewn slide, and all are present in about equal numbers. VoL. XXXVIT] HANNA: PRIBILOF FOSSIL DIATOMS 199 REFERENCES AzpeITIA Moros, D. FLoRENTINO 1911. La Diatomologia Espanola en los comienzos del Siglo XX Asociacién Espafiola para el Progreso de las Ciencias, Congreso de Zaragoza, vol. 4, Sec. 3a, Ciencias Naturales Segunda parte, Madrid, 320 pages, 12 plates. Barry, J. W. 1845. Notice of some new localities of Infusoria, fossil and Recent. The American Journal of Science and Arts, vol. 48, no. 2, pp. 321-343, pl. 4. Barker, J. W., and S. H. Meakin 1949. New and rare diatoms. The Journal of the Quekett Microscopical Club, ser. 4, vol. 2, no. 6, pp. 301, pls. 37 and 38. BartH, T. F. W. 1956. Geology and petrology of the Pribilof Islands, Alaska. Bulletin U. S. Geological Survey, no. 1028F, pp. iv, 101-160, 2 maps, figs. 17-36. Boyer, Cuartes S. 1927. Synopsis of North American Diatomaceae, Part I.—Coscinodiscatae, Rhizo- selenatae, Biddulphiatae, Fragilariata. Proceedings of the Academy of Natural Sciences of Philadelphia, vol. 78, Supplement, 228 pp. Brun, JACQUE et TEMPERE, J. 1889. Diatomées Fossiles du Japon, Espéces Marines u Nouvelles des Calcaires Argileux de Sendai & de Yedo. Mémoires de la Société de Physique et D’Histoire Naturelle de Genéve, vol. 30, no. 9, 73 pp., 9 pls. BruN, JACQUES 1891. Diatomées especes nouvelles marines, fossiles ou pélagiques. Mémoires de la Société de Physique et D’Histoire Naturelle de Genéve, vol. 31, pt. 2, no. 1, 47 pp., 12 pls. CLEVE, P. T. 1873. On diatoms from the Arctic Sea. Bihang till Kongl. Svenska Vetenskaps- Akademiens Handlingar, vol. 1, no. 13, 28 pp., 4 pls. 1894. Synopsis of the Naviculoid Diatoms, pt. 1, Kongl. Svenska Vetenskaps- Akademiens Handlingar, vol. 20, no. 2, 194 pp., 5 pls. Dat1, W. H. 1896. Report on coal and lignite of Alaska. U. S. Geological Survey, 17th Annual Report, pt. 2, pp. 763-908. 1899. The mollusk fauna of the Pribilof Islands, in the Fur Seals and Fur Seal Islands of the North Pacific Ocean, pt. 3, pp. 539-546. [The fossil species found at Black Bluff, St. Paul Island are listed and the age is given as Pliocene.] 1919. On some Tertiary fossils from the Pribilof Islands. Journal Washington Academy of Sciences, vol. 9, pp. 1-3. [A list of the mollusks found at Tolstoi Point, St. Paul Island, and Tolstoi Point, St. George Island is given.] Dawson, G. M. 1894. Geological notes on some of the coasts and islands of Bering Sea and vicinity. Bulletin Geological Society of America, vol. 5, pp. 117-146; Pribilof Islands, pp. 130-133. DONAHUE, JESSE G. 1967. Diatoms as indicators of Pleistocene climatic fluctuations in the Pacific sector of the southern ocean. Progress in Oceanography, vol. 4, (The Quaternary History of the Ocean Basins), pp. 133-140, 1 pl. 5 text figs. 200 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47H SER. Etior, H. W. 1875. A report on the condition of affairs in the Territory of Alaska. U. S. Treasury Department, p. 70. [Also published as House Miscellaneous Document no. 83, 44th Congress, 1st Session. ] 1886. Our Arctic Province, p. 229. GREWINGK, CONSTANTINE 1850. Beitrag zur Kenntniss der orgraphischen, etc., der Nordweste Kuste Amerikas. Verhandlungen der Russisch Kaiserlichen Mineralogischen Gesellschaft zu St. Petersburg. [Pribilof geology, p. 190. Wossnessensky’s observations]. EHRENBERG, G. C. 1838. Die Infusionsthierchen als volkommene Organismen. Leipzig 548 pp., 64 pls. 1841. Verbreitung und einflus der mikroskopischen lebens in Siid-und Nord-Amerika. [1843.] Abhandlungen der K6niglichen-Akademie der wissenschaften zu Berlin. Erster Theil, pp. 291-446, pls. 1-4. [Often referred to as “Amer.” or “America” ]. 1844. Beitrage zur Kenntniss des Kleinsten Lebens im agadischen Meere am Euphrat und [1845.] auf den Bermuda-Inseln. Monatsberichten der Berliner Akademie der Wissen- schaften, p. 264. 1854. Mikrogeologie. Das Erden und Felsen Schaffende Wirken des Unsichtbar Kleinen Selbststandigen Lebens auf der Erde. Leipzig, verlag von L. Voss in two parts. Text pp. I-X XVIII, 1-374; Atlas, pp. 1-31, pls. I-XXXX. GREVILLE, R. K. 1865. Descriptions of new and rare diatoms, series 16. Transactions of the Micro- scopical Society of London, new series, vol. 13, pp. 43-57, pls. 5, 6. Hanna, G DALLAs 1919. Geological notes on the Pribilof Islands, Alaska, with an account of the fossil diatoms. American Journal of Science, vol. 48, pp. 216-224. 1927. The lowest known Tertiary diatoms in California. Journal of Paleontology, vol. 1, no. 2, pp. 103-127, pls. 17-21. 1930. The growth of Omphalotheca. Journal of Paleontology, vol. 4, no. 2, p. 192. Hanna, G Darras, and W. M. Grant 1926. Miocene marine diatoms from Maria Madre Island, Mexico. Proceedings of the California Academy of Sciences, 4th ser., vol. 15, no. 2, pp. 115-192, pls. 12-21. HeEnbeEy, N. I. 1937. The plankton diatoms of the southern seas. Discovery Reports, vol. 16, pp. 151-364, pls. 6-13. Hopkins, DAavip M., F. MacNem, R. MeErK1iIn, and O. PETROV 1965. Quaternary correlations across Bering Strait. Recent Soviet and American studies cast new light on the history of the Bering Sea land bridge. Science, N.S., vol. 147, no. 3662, pp. 1107-1114, map. Hopkins, Davip, and THorRLEIFUR EINARSSON 1966. Pleistocene glaciation on St. George, Pribilof Islands. Science, n.s., vol. 152, no. 3720, pp. 343-344, 2 text figs. Hustept, FRIEDRICH 1930. Die Stsswasser-Flora Mitteleuropas, Heft 10 : Bacillariophyta (Diatomeae), 466 pages, 875 text figures. Wows, Ay 12: 1959. The main phases in the development of the flora of marine diatoms in the far eastern seas of the USSR at the end of the Tertiary and during the Quaternary period. Academy of Sciences, USSR. Botanical Journal, vol. 44, no. 1, pp. 44-55, pls. 2-4. VoL. XXXVI] HANNA: PRIBILOF FOSSIL DIATOMS 201 1961. Diatomeae Marinae Mioceni et Pliocenei ex Oriente Extremo. Akademiia Nauk SSSR. Lenigrad. Botanichesii Institut. Otdel Sporovykh Rastinii Botanicheskie Materialy, vol. 14, pp. 59-70, 3 pls. Junk, WILHELM 1900-1913. Rara historica-naturalia et mathematica, Berlin; vol. 1, 121 pp., vol. 2, 1926-1936, pp. 123-240. KANnAvYA, Taro, and ITaru Koizumi 1966. Interpretation of Diatom Thanatocoenoses from the North Pacific applied to a study of core V 20-130 (Studies of a Deep-sea Core V 20-130, Part IV.) Science Reports, Tohoku University, Sendai, ser. 2, (Geol.), vol. 37, no. 2, pp. 89-130, 6 text figs. : Mann, ALBERT 1925. Marine diatoms of the Philippine Islands. Bulletin of the U. S. National Museum NOOO VOl On pial, 182 pp: 39epls: Muts, F. W. 1933-1935. An index to the genera and species of the Diatomaceae and their synonyms, 21 parts, 1726 pages. Wheldon & Wesley, Ltd. London. Muir, JoHN 1883. Arctic Cruise of the Revenue Cutter Corwin in 1881, p. 140. [Notes and obser- vations on supposed general glaciation of the Pribilofs.] ScumiptT, A., and others 1874-1959. Atlas der Diatomaceenkunde, 480 plates. ScHott, Davin W., Epwin G. Burrincton, AnD Dav M. Hopkins 1966. Exposure of basement rock on the continental slope of the Bering Sea. Science, n.s., vol. 153, August 26, pp. 992-994, 3 text figures. SHESHUKOVA-PoRETSKAIA, V. S. 1967. Neogene marine diatoms of Sakhalin and Kamchatka. 432 pp., 50 pls. Leningrad University Press. SmitH, W. 1853. A synopsis of the British Diatomaceae; with remarks on their structure, functions and distribution; and instructions for collecting and preserving specimens, vol. 1, pp. IL-X XXIII, 1-92, pls. 1-31. London. STANLEY-Brown, J. 1892. Geology of the Pribilof Islands. Bulletin, Geological Society of America, vol. 3, pp. 496-500. VAN HeurckK, HENRI 1880-1883. Synopsis des Diatomées de Belgique, Anvers, Atlas, 141 plates, 2 pages introduction, 6 pages index. WasHINcTON, H. S. 1930. Rocks on the Pribilof Islands. American Journal of Science, vol. 20, pp. 321-338. ZABELINA, M. 1934. Trudy Neftianogo Geologo-Razvedochnogo Institute Ser. A, no. 48, pp. 1-19, 9 test figs. [Diatoms from the Tertiary deposits of the eastern coast of Kamchatka. Transactions of the Petroleum Prospecting Institute Ser. A, No. 48, pp. 1-19, 9 text figs.] [State Scientific-Technical Mining-Geological- Petroleum Publishing House, Leningrad, Moscow, Novosibirsk, 1934.] 202 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 1. Coscinodiscus marginatus Ehrenberg. Hypotype no. 3578 (Calif. Acad. Sci., Dept. Geol. Type Coll.), from locality no. 36829 (CAS), Tolstoi Point, St. Paul Island, Alaska. Diameter .0870 mm. Ficure 2. Coscinodiscus marginatus Ehrenberg. Hypotype no. 3579 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 1. Diameter .200 mm. Ficure 3. Coscinodiscus undulosus Mann. Hypotype no. 3580 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 1. Diameter .0860 mm. VoL. XXXVII] HANNA: PRIBILOF FOSSIL DIATOMS 203 204 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 4. Coscinodiscus radiatus Ehrenberg. Hypotype no. 3581 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality no. 36829 (CAS), Tolstoi Point, St. Paul Island, Alaska. Diameter .1490 mm. Ficure 5. Thalassiosira punctata Jousé. Hypotype no. 3582 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality no. 1233 (CAS), Black Bluff, St. Paul Island, Alaska. Diameter .0456 mm. Ficure 6. Thalassiosira punctata Jousé. Hypotype no. 3583 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 1. Diameter .0570 mm. Ficure 7. Coscinodiscus kiitzingii Schmidt. Hypotype no. 3584 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality no. 717 (CAS), Tolstoi Point, St. Paul Island, Alaska. Diameter .0506 mm. 205 HANNA: PRIBILOF FOSSIL DIATOMS VoL. XXXVII] 206 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 8. Coscinodiscus radiatus Ehrenberg. Hypotype no. 3593 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality no. 1322 (CAS), St. Paul Island, Alaska, Navy Well no. 2, depth 90 feet. Diameter .1156 mm. Ficure 9. Cosmiodiscus insignis Jousé. Hypotype no. 3594 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality no. 1322 (CAS), St. Paul Island, Alaska, Navy Well no. 2, depth 400 feet. Diameter .0640 mm. Ficure 10. Cosmiodiscus insignis Jousé. Hypotype no. 3595 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 9. Diameter .0660 mm. Ficure 11. Cosmiodiscus insignis Jousé. Hypotype no. 3595A (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 9. Diameter .0726 mm. Ficure 12. Coscinodiscus pustulatus Mann. Hypotype no. 3596 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality no. 36829 (CAS), Tolstoi Point, St. Paul Island, Alaska. Diameter .0416 mm. Ficure 13. Diploneis ornatus Schmidt. Hypotype no. 3597 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 12. Diameter .0510 mm. Ficure 14. Diploneis bombus (Ehrenberg). Hypotype no. 3598 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 8. Length .060 mm. 207 HANNA: PRIBILOF FOSSIL DIATOMS VoL. XXXVII] PV rereree 74 OOF che rete yy) dawei. ly Oe SSE Fe-<: =conhane >)??? ‘ nAAT) . SS} yan} 208 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 15. Cymatotheca weissflogii (Grunow), Hendey. Hypotype no. 3586 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality no. 36829 (CAS), Tolstoi Point, St. Paul Island, Alaska. Length .0860 mm., width .0768 mm. Ficure 16. Cymatotheca weissflogii (Grunow), Hendey. Hypotype no. 3587 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality no. 1233 (CAS), Black Bluff, St. Paul Island, Alaska. Length .040 mm., width .0344 mm. Ficure 17. Coscinodiscus radiatus Ehrenberg. Hypotype no. 3588 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality no. 1322 (CAS), St. Paul Island, Alaska, Navy Well no. 2, depth 90 feet. Diameter .1332 mm. Ficure 18. Coscinodiscus oculus-iridus Ehrenberg. Hypotype no. 3589 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 15. Diameter .240 mm. Vor. XXXVIT] HANNA: PRIBILOF FOSSIL DIATOMS 209 a. aos ie? gn * “% * lw og e Brea e, ~ . 2 . eae! a “ ar ae eo” rr) 210 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 19. Coscinodiscus pustulatus Mann. Hypotype no. 3599 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality no. 717 (CAS), Tolstoi Point, St. Paul Island. Alaska. Diameter .0526 mm. Ficure 20. Coscinodiscus pustulatus Mann. Hypotype no. 3600 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 21. Diameter .0356 mm. Ficures 21, 22. Coscinodiscus pustulatus Mann. Hypotypes nos. 3601, 3602 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality no. 36829 (CAS), Tolstoi Point, St. Paul Island, Alaska. Diameter .0940 and .0656 mm. Ficure 23. Coscinodiscus pustulatus Mann. Hypotype no. 3603 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figures 21, 22. Diameter .060 mm. Ficure 24. Coscinodiscus pustulatus Mann. Hypotype no. 3603A (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figures 21, 22. 211 HANNA: PRIBILOF FOSSIL DIATOMS VoL. XXXVII] i © ‘oe a 88eg 212 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 25. Actinoptychus splendens (Shadbolt). Hypotype no. 3549 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality no. 36829 (CAS), Tolstoi Point, St. Paul Island, Alaska. Diameter .0864 mm. Ficure 26. Actinoptychus splendens (Shadbolt). Hypotype no. 3550 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 25. Diameter .0892 mm. Ficure 27. Rhaphoneis nitida Gregory. Hypotype no. 3577 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 25. Length .0360 mm. FicurE 28. Actinoptychus splendens (Shadbolt). [= Debya.] Hypotype no. 3551 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 25. Diameter .0824 mm. Ficure 29. Rhaphoneis amphiceros Ehrenberg. Hypotype no. 3621 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 25. Length .1024 mm. VoL. XXXVIT] HANNA: PRIBILOF FOSSIL DIATOMS 213 nd ra) u w~ am (>> CTA ye: - a ra aD ~ 47 = cs = t IOS) Fear “a eae Se Be es ys > eri ~~ a —_— NTN Wy i I on” “im U a 5 Fal 214 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. FicurE 30. Cosmiodiscus insignis Jousé. Hypotype no. 3585 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality no. 717 (CAS), Tolstoi Point, St. Paul Island, Alaska. Diameter .0476 mm. Ficure 31. Cocconeis maxima (Grunow). Hypotype no. 3590 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality no. 1322(CAS), St. Paul Island, Alaska. Navy Well no. 2, depth 90 feet. Length .0360 mm. Ficure 32. Cosmiodiscus insignis Jousé. Same specimen as figure 30, but with a lower focus. Ficure 33. Pinnularia lata (Brébisson). Hypotype no. 3591 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 31. Length .0728 mm. Ficure 34. Cocconeis pribilofensis Hanna, new species. Hypotype no. 3574 (CAS), from locality no. 36829 (CAS), Tolstoi Point, St. Paul Island, Alaska. Length .0302 mm. VoL. XXXVIT] HANNA: PRIBILOF FOSSIL DIATOMS PAIS) 216 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 35. Hemidiscus cuneiformis Wallich. Hypotype no. 3556, (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality no. 36829 (CAS), Tolstoi Point, St. Paul Island, Alaska. Greater diameter .0900 mm.; lesser diameter .0472 mm. Ficure 36. Coscinodiscus rothii (Ehrenberg). Hypotype no. 3629 (Calif. Acad. Sci. Dept. Geol. Type Coll.) from same locality as figure 35. Diameter .0744 mm. Ficure 37. Thalassionema nitzschoides (Grunow). Hypotype no. 3572 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 35. Length .0620 mm. Ficure 38. Actinoptychus senarius Ehrenberg. Hypotype no. 3571 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 35. Diameter .1150. FicurE 39. Actinoptychus senarius Ehrenberg. Hypotype no. 3552 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 35. Diameter .1140 mm. VoL. XXXVI] HANNA: PRIBILOF FOSSIL DIATOMS 217 ry 0 Re G 3. 218 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 40. Actinoptychus splendens (Shadbolt). Hypotype no. 3547 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality no. 36829 (CAS), Tolstoi Point, St. Paul Island, Alaska. Diameter .0734 mm. Ficure 41. Arachnoidiscus ehrenbergit Bailey. Hypotype no. 3616 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality no. 1322 (CAS), Navy Well no. 2, St. Paul Island, Alaska. Diameter .1828 mm. Ficure 42. Triceratium condecorum Brightwell. Hypotype no. 3618 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 40. Length of base to apex .0852 mm. Ficure 43. Triceratium montereyi Brightwell. Hypotype no. 3619 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 40. Length of one side .1154 mm. Ficure 44. Actinoptychus splendens (Shadbolt). Hypotype no. 3548 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 40. Diameter .1068 mm. Ficure 45. Actinoptychus senarius Ehrenberg. Hypotype no. 3553 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 40. Diameter .0682 mm. VoL. XXXVI] HANNA: PRIBILOF FOSSIL DIATOMS 219 bo bo oO CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 46. Aulacodiscus tripartitus Tempére and Brun. Hypotype no. 3575 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality no. 36829 (CAS), Tolstoi Point, St. Paul Island, Alaska. Diameter .060 mm. Ficure 47. Rhabdonema japonicum Tempére and Brun. Hypotype no. 3576 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 46. Length .0760 mm. Ficure 48. Cocconeis antiqua Tempére and Brun. Hypotype no. 3573 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 46. Length .0950 mm. Ficure 49. Aulacodiscus tripartitus Tempere and Brun. Hypotype no. 3637 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 46. Diameter .0732 mm. VoL. XXXVII] HANNA: PRIBILOF FOSSIL DIATOMS 221 bo bo bo CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 50. Melosira sulcata (Ehrenberg). Hypotype no. 3604 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality no. 38579 (CAS), Einahnuhto Bluffs, St. Paul Island, Alaska. Diameter .0350 mm. Ficure 51. Melosira sulcata (Ehrenberg). Hypotype no. 3605 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality no. 36829 (CAS), Tolstoi Point, St. Paul Island, Alaska. Diameter .0426 mm. Ficure 52. Melosira clavigera Grunow. Hypotype no. 3555 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 51. Diameter .0888 mm. Ficure 53. Melosira sulcata (Ehrenberg). Hypotype no. 3606 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 51. Diameter .0256 mm. Ficure 54. Melosira clavigera Grunow. Hypotype no .3604 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 51. Diameter .0270 mm. Ficure 55. Rhaphoneis amphiceros (Ehrenberg). Hypotype no. 3607 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 51. Length .0320 mm. Ficure 56. Rhaphoneis amphiceros (Ehrenberg). Hypotype no. 3607A (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 51. Length .0306 mm. Ficure 57. Rhaphoneis lancettula Grunow in Pantocsek. Hypotype no. 3617 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 51. Length .1420 mm. FicuRE 58. Stephanopyxis appendiculata Ehrenberg. Hypotype no. 3610 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality no. 1322 (CAS), Navy Well no. 2, St. Paul Island, Alaska. Depth, 90 feet. Diameter .0284 mm. VoL. XXXVII] HANNA: PRIBILOF FOSSIL DIATOMS 223 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. bo LS) _ Ficure 59. Dossetia temperei Azpeitia. Hypotype no. 3625 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality no. 36829 (CAS), Tolstoi Point, St. Paul Is!and, Alaska. Length of base .0568 mm. FicurE 60. Dossetia temperei Azpeitia. Hypotype no. 3625A (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 59. Greatest diameter .0560 mm. Ficure 61. Pyxilla americana Grunow. Hypotype no. 3626 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 59. Length .0948 mm. Ficure 62. Chaetoceros didymus Ehrenberg. Hypotype no. 3620 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 59. Length .0572 mm. Ficure 63. Dicladia capreola Ehrenberg. Hypotype no. 3622 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 59. Length of base .0332 mm. Ficure 64. Xanthiopyxis ovalis Lohman. Hypotype no. 3608A (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 59. Length .0280 mm. Ficure 65. Chaetoceras ? (sp.). Hypotype no. 3627 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 59. Length .0320 mm. Ficure 66. Pseudopyxilla dubia (Grunow) Forti. Hypotype no. 3623 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 59. Length .1092 mm. Ficure 67. Ebriopsis antiqua (Schulz), Hovasse. Hypotype no. 3612 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 59. Diameter .0270 mm. Ficure 68. Pseudopyxilla dubia (Grunow), Forti. Hypotype no. 3624 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 59. Diameter .0568 mm. Ficure 69. NXanthiopyxis lohmani Hanna, new species. Holotype no. 3608 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 59. Length .0448 mm. Ficure 70. Xanthiopyxis ovalis Lohman. Hypotype no. 3609 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 59. Length .0306 mm. Ficure 71. Distephanus speculum Ehrenberg. Hypotype no. 3613 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 59. Diameter, exclusive of spines, .0228 mm. Ficure 72. Distephanus speculum Ehrenberg. Hypotype no. 3614 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 59. Diameter .0244 mm. Ficure 73. Mesocena corona Hanna, new species. Holotype no. 3611 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 59. Diameter .0340 mm., exclusive of spines. Ficure 74. Xanthiopyxis globosa Ehrenberg. Hyptoype no. 3592 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 59. Diameter of base .0256 mm, Ficure 75. Dictyocha fibula Ehrenberg. Hypotype no. 3615 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from same locality as figure 59. Horizontal length .0280 mm., exclusive of spines. VoL. XXXVII] HANNA: PRIBILOF FOSSIL DIATOMS 225 i) in) fon) CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 76. Actinoptychus senarius Ehrenberg. Hypotype no. 3631 (Calif. Acad. Sci. Dept. Geol. Type Coll.), from locality no. 36829 (CAS), Tolstoi Point, St. Paul Island, Alaska. Diameter .0928 mm. Ficure 77. Arachnoidiscus indicus Ehrenberg. Hypotype no. 3635 (CAS), from same locality as figure 76. Diameter .0644 mm. Ficure 78. Biddulphia aurita (Lyngbye) Brébisson and Godey. Hypotype no. 3656B (CAS), from same locality as figure 76. Diameter .0540 mm. Ficure 79. Biddulphia roperiana Greville. Hypotype no. 3638 (CAS), from same locality as figure 76. Length .0748 mm. Ficure 80. Avrachnoidiscus indicus Ehrenberg. Hypotype no. 3634 (CAS), from same locality as figure 76. Diameter .0692 mm. Ficure 81. Coscinodiscus fimbriatus Ehrenberg. Hypotype no. 3641 (CAS), from same locality as figure 76. Diameter .1088 mm. Ficure 82. Biddulphia baltzoi Hanna, new species. Hypotype no. 3639 (CAS), from same locality as figure 76. Length .040 mm. Ficure 83. Rhaphoneis rhombus Ehrenberg. Hypotype no. 3651 (CAS), from same locality as figure 76. Length .0564 mm. Vor. XXXVIT] HANNA: PRIBILOF FOSSIL DIATOMS 227 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. bo bo ee) Ficure 84. Hercotheca inermis Mann. Hypotype no. 3648 (CAS), from locality no. 36829 (CAS), Tolstoi Point, St. Paul Island, Alaska. Length .0372 mm. Ficure 85. Triceratium validum Grunow. Hypotype no. 3655 (CAS), from same locality as figure 84. Height .1040 mm. Ficure 86. Hemidiscus rotundus Janisch. Hypotype no. 3646 (CAS), from same locality as figure 84. Length .0944 mm., width .0816 mm. Ficure 87. Xanthiopyxis umbonatus Greville. Hypotype no. 3656 (CAS), from same locality as figure 84. Diameter .1104 mm. Ficure 88. Triceratium condecorum Brightwell. Hypotype no. 3654 (CAS), from same locality as figure 84. Height .0420 mm. FicurE 89. Cocconeis formosa Brun. Hypotype no. 3640 (CAS), from same locality as figure 84. Length .0376 mm. Ficure 90. Asterolampra darwinii Greville. Hypotype no. 3636 (CAS), from same locality as figure 84. Diameter .0560 mm. Ficure 91. Lithodesmium undulatum Ehrenberg. Hypotype no. 3653 (CAS), from same locality as figure 84. Height .030 mm. VoL. XXXVIT] HANNA: PRIBILOF FOSSIL DIATOMS 229 ART * a a Ps RSENS OD CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. bdo is) oO Ficure 92. Rhabdonema japonicum Tempére and Brun. Hypotype no. 3643 (CAS), from locality no. 36928 (CAS), Tolstoi Point, St. Paul Island, Alaska. Length .1460 mm. Ficure 93. Rhabdonema japonicum Tempeére and Brun. Hypotype no. 3644 (CAS), from same locality as figure 92. Length .1892 mm. Ficure 94. Rhabdonema crozieri (Ehrenberg). Hypotype no. 3645 (CAS), from same locality as figure 92. Length .0812 mm. Ficure 95. Rhabdonema japonicum Tempére and Brun. Hypotype no. 3642 (CAS), from same locality as figure 92. Length .1280 mm. Ficure 96. Navicula semen Ehrenberg. Hypotype no. 3649 (CAS), from same locality as figure 92. Length .0832 mm. FicurEe 97. Chaetoceros didymus Ehrenberg. Hypotype no. 3632 (CAS), from same locality as figure 92. Length .1440 mm. Ficure 98. Chaetoceros didymus Ehrenberg. Hypotype no. 3630 (CAS), from locality no. 1322 (CAS), Navy Well no. 2, St. Paul Island, Alaska. Length .0656 mm. FicurEe 99. Stephanopyxis appendiculata Ehrenberg. Hypotype no. 3652 (CAS), from same locality as figure 92. Diameter .0508 mm. Ficure 100. Stephanopyxis appendiculata Ehrenberg. Hypotype no. 3652A (CAS), from same locality as figure 92. Diameter .0548 mm. Ficure 101. Pinnularia ruttneri Hustedt in Schmidt. Hypotype no. 3650 (CAS), from same locality as figure 92. Length .120 mm. HANNA: PRIBILOF FOSSIL DIATOMS Vor. XXXVIT] SUT , UCIT: QUT INV 101 bo Ww bo CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 102. Aulacodiscus laxus (Mann). Hypotype no. 3950 (CAS), from locality no. 36829 (CAS), Tolstoi Point, St. Paul Island, Alaska. Diameter .084 mm. Ficure 103. Aulacodiscus tripartitus Tempére and Brun. Hypotype no. 3951 (CAS), from same locality as figure 102. Diameter .080 mm. Ficure 104. Aulacodiscus laxus (Mann). Hypotype no. 3952 (CAS), from same locality as figure 102. Diameter .093 mm. Ficure 105. Aulacodiscus tripartitus Tempere and Brun. Hypotype no. 3953 (CAS), from same locality as figure 102. Diameter .074 mm. Figures 102-105 are from photographs made by Dr. A. L. Brigger, Research Associate, California Academy of Sciences. | Vor. XXXVII] 233 HANNA: PRIBILOF FOSSIL DIATOMS OE ee, Ax aA f ta | oN Bone ~ M77, }y {i WD ? it se! am 7 ’ a - ’ @ 7 % ; es ae uy. ; 7 - PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XXXVII, No. 6, pp. 237-248; 8 figs. March 6, 1970 COPELATUS GLYPHICUS (SAY) AND SUPHISELLUS BICOLOR (SAY), WATER BEETLES NEW TO CALIFORNIA AND PRESUMABLY INTRODUCED (COLEOPTERA: DYTISCIDAE AND NOTERIDAE) By Hugh B. Leech California Academy of Sciences, San Francisco 94118 INTRODUCTION In August, 1963, a living specimen of the small dytiscid Copelatus glyphicus (Say) was found in a clear, cold mountain stream at an altitude of 6600 feet in Tuolumne County, California. In October three more were found in a murky pool near sea level in Marin County. These two localities are over a thousand miles west of any published records for the species, and provide very unlike habitats. In April, 1968, one male was taken in a tributary to Deer Creek, Tulare County, at 4000 feet altitude, and a female in Deer Creek itself some 20 air- line miles away, at 550 feet altitude. In this last spot I collected also a series of the little noterid Suphisellus bicolor (Say), type species of the genus, here at least 1200 miles west of its known distribution. One explanation might be that the two species (and perhaps others not yet found) have been introduced, possibly in aquaria materials. This may be plausible for S. bicolor taken in series from one stream, but hardly accounts for C. glyphicus, which has been found sparingly in each of four places over a northwest-southeast distance of some 250 miles. [237] paca enn ae | Marine Biological Laboratory | MAR 1 61970 | f 4 | WOODS HOLE, MASS. == Se SB as oe -_ ae | CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER. iS) Ww co Family DyTISCIDAE The species of Copelatus of the United States and Canada are small brownish dytiscid beetles, often flattened, which have regular, clearly impressed _ longi- tudinal discal striae (not series of punctures) on the elytra, in both sexes (fig. 1). In 1956 I recorded C. chevrolati renovatus Guignot from Imperial County, California, and listed C. impressicollis Sharp as a possible California species. Since then F. N. Young has published a revisional study of the Nearctic species in which he tentatively placed C. impressicollis as a synonym of C. distinctus Aubé, remarking of the latter “I believe that it will eventually prove to form an Artenkreis of isolated species.” It seems not to have been taken in the United States from much west of a line drawn from Flagstaff to Nogales in south central Arizona, and should be dropped from the California list. On the other hand an unlikely second species, C. glyphicus (Say), has been taken here four times in recent years, a thousand miles west of its nearest recorded locality. Key To THE CALIFORNIA SPECIES OF Copelatus 1. Each elytron with 10 discal striae, and a submarginal stria in posterior half (fig. 1). Prosternal process slightly evenly inflated, not keeled. Inner margin of front tibia of male narrowed and weakly notched in basal third (fig. 3), that of female slightly sinuate in same region. Smaller, flattened species, 4.0 to 4.75 mm. ONG, see ee ee C. glyphicus (Say), 1823. Each elytron with 8 discal striae, and a submarginal in posterior half. Prosternal process strongly but not sharply keeled. Inner margin of front tibia of male straight (fig. 5), as in female. Larger, more convex species, 5.25-6.50 mm. On gdp eee eee oe Se Ee ee See ee ee C. chevrolati renovatus Guignot, 1952. Copelatus glyphicus (Say). Colymbetes glyphicus Say, 1823, Trans. Amer. Philos. Soc., new ser., vol. 2, no. 1, p. 99. Say in LeConte, 1859 [Complete writings of Thomas Say], vol. 2, p. 512. Copelatus glyphicus Say, SHARP, 1882, Sci. Trans. Royal Dublin Soc., (2) vol. 2, p. 589. SCHAEFFER, 1908, Jour. New York Ent. Soc., vol. 16, no. 1, p. 17. BLATCHLEY, 1910, Coleoptera . . . Indiana, p. 223. Younc, 1954, Univ. Florida Studies, Biol. Sci. Ser., vol. 5, no. 1, p. 106. Younc, 1963, Quarterly Jour. Florida Acad. Sci. vol. 26, no. 1, p. 60 (key), 62 (text). This common eastern species has not been reported from further west than eastern New Mexico (Sublette and Sublette, 1967, p. 369). Knowing its gen- eral distribution, I simply couldn’t believe my eyes when I took my first example in northern California. A female was collected in a little stream a hundred yards from its confluence with Niagara Creek, at an altitude of 6600 feet on the western slope of the Sierra Nevada. This was at the Niagara Creek Forest Campground adjacent to Highway 108 in Tuolumne County, on 11 August 1963. On 21 September my wife and I returned to the same spot, but Vor. XXXVII] LEECH: WATER BEETLES NEW TO CALIFORNIA 239 FicurEs 1-2. Aquatic beetles new to California. Ficure 1. Copelatus glyphicus (Say) from Deer Creek at the Porterville - Fountain Springs road, Tulare County, California. < 16.5. Ficure 2. Suphisellus bicolor (Say), same data. X 26.5. Photographs by David G. Kissinger. failed to get any more. We did vow never again to go into the mountains on the first day of deer shooting season, for our collecting site had become the campground for a swarm of hunters. With their teen-age progeny many were busy “sighting in” their guns, and it was no place for entomologists to be stooped over at the edge of a stream. On 6 October 1963, I found two males and a female of C. glyphicus in a pool filled with dead grass in a ditch just to the south side of the old railroad grade which went under Highway 101 at Forbes Overhead, San Rafael, Marin County, only 8 feet above sea level. This marshy area, tributary to Gallinas Creek, has since been “improved” with rock fill. The third and fourth localities are some 250 airline miles southeast of San Francisco, at the eastern edge of the San Joaquin Valley. One female was collected on 9 April 1968, in Deer Creek at the Porterville-Fountain Springs 240 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. road, a few miles south of Porterville, Tulare County, altitude 550 feet; a male was taken some 20 airline miles away in a tributary to Capinero Creek (itself an affluent of Deer Creek), 2 miles east of California Hot Springs, at an altitude of 4000 feet, on 10 April. It is surprising to find C. glyphicus a thousand miles west of its cited range, but the known distribution and habitats in California are even more puzzling. Niagara Creek is a rapid, small, clear stream in a boulder and gravel bed in the forested sierra; the San Rafael locality was a pool full of dead organic matter near sea level; Deer Creek at the Fountain Springs road is turbid with silt, shallow, and spread over a broad sandy bed in open meadow- land, with a good current along the bank where the beetle was found; its seasonal affluent at 4000 feet is a tiny rapid stream over a sand and silt bed in a gully. If the species was originally introduced, it has spread remarkably and is surely here to stay. Perhaps it has been actively spreading westward recently, and is still so scarce in western New Mexico, Arizona, and adjacent areas as to have escaped collectors. It can hardly be so widespread in Califor- nia and yet be unreported by other collectors, unless it has been here for only a short time. Another little species, smaller than C. glyphicus but with similarly modified front tibiae in the male, has been taken in Brownsville, Texas, and on the west coast of Mexico in Nayarit and Sinaloa. This is C. debilis Sharp, in which each elytron has five striae. Copelatus chevrolati renovatus Guignot. Copelatus chevrolati var. australis SCHAEFFER, 1908, Jour. New York Ent. Soc., vol. 16, no. 1, p. 17 (not C. australis (Clark), 1863, p. 14). LrEcH, 1948, Proc. Calif. Acad. Sci., 4th ser., vol. 24, no. 11, p. 406. Copelatus chevrolati ssp. schaefferi GuicNotT, 1952, Rev. franc. Ent., vol. 19, no. 1, p. 23 (not C. schaeffert Young, 1942, p. 92.) Copelatus chevrolati ssp. renovatus GuIGNoT, 1952, Rev. frang. Ent., vol. 19, no. 3, p. 170. Copelatus chevrolati renovatus Guignot, Lrerecu, 1956, in Usinger, Aquatic insects of Calif... . , p. 321. Younc, 1963, Quarterly Jour. Florida Acad. Sci., vol. 26, no. 1, p. 74. Young gives the distribution of the typical subspecies as the Atlantic and Gulf coastal plain area and the Bahamas. He mentions having seen specimens of C. chevrolati renovatus from “Arizona, Arkansas, California, Kansas, Loui- siana, New Mexico, Oklahoma, Texas, Sonora, Baja California, Sinaloa, Ta- maulipas, and Nuevo Leon.” As C. chevrolatii, C. chevrolati, or var. australis, C. chevrolati renovatus has been recorded from California but never with actual localities, by Crotch, 1873, p. 413; Sharp, 1882, p. 584; Woodworth, 1913, p. 184; Sherman (im Leng), 1920, p. 80; and Leech, 1948, p. 406. Vout. XXXVII] LEECH: WATER BEETLES NEW TO CALIFORNIA 241 There are 64 California specimens at hand, all but one from the extreme south in the drainage or flood plain of the Colorado River. The following lo- calities are given from south to north. IMPERIAL County: Calexcio, 24. VIII. 1937 (light trap); El Centro, 4.X11.1927 (F. E. Blaisdell), and some dated only 1937; Holtville, 15.V1I.1934 (E. C. Van Dyke), 15.VII.1934 (M. Cazier), VI.1936 (E. S. Ross), 7.V.1940 (R. P. Allen), 23.VII.1946 (E. C. Van Dyke), 5.1V.1949 (P. D. Hurd. Col- lected at light); Meloland, 25.VII.1949 (Ray F. Smith); Imperial, 3.I1V.1924; Imperial Valley, 28.V.1925 and .V.1926; Brawley, 23.X.1936 (A. T. McClay), 18.111.1939 and 9.V.1940 (R. P. Allen); Calipatria 1.VI.1959 (at argon light). RIVERSIDE CouNTy: Coachella, 24.V.1928 (E. C. Van Dyke); Coachella Val- ley, 25.V.1928 (E. C. Van Dyke); Indio, 24.IV.1952 (O. Bryant), IV.1952 (N. Lewis). TurarE County: Deer Creek at Porterville—Fountain Springs Road, 9.I1V.1968 (H. B. Leech). Also one example labeled only “S. Cal.” The Deer Creek locality is more than 200 miles northwest of the others, north of the Tehachapi Mountains, and in the “landlocked” Tulare drainage basin. But in the spring of 1969, as occasionally in other years of excessive rain and snow, Tulare Lake flooded and drained northward. Some people went from there to San Francisco by boat, via the San Joaquin River in mid-June (Zane, 1969, p. 5). Copelatus distinctus is of the same size as C. chevrolati renovatus but darker, flatter, more parallel-sided, and each elytron has 10 discal striae. The front tibiae of the male are straight on the inner margin. Copelatus distinctus is known as far northwest as central southern Arizona, as mentioned at the be- ginning of this paper. Family NOTERIDAE Suphisellus bicolor (Say). Noterus bicolor Say, 1831", Descr. n. spp. . . insects. . ., Louisiana. . ., p. 5. Say, 1834, Trans. Amer. Philosophical Soc., vol. 4, p. 446. Say in Le Conte (editor), 1859, [Complete writings of Thomas Say], vol. 2, p. 561. ScuppER, 1899, Psyche, vol. 8, no. 273, p. 307. Suphisellus bicolor (Say), LEEcH, 1948, Proc. Calif. Acad. Sci., 4th ser., vol. 24, no. 11, p. 403. Younc, 1954, Water beetles of Fla., p. 131. In 1948 I designated Noterus bicolor Say, 1831, p. 5, as the type of Sup- hisellus Crotch, 1873, p. 397, so a present-day identification of the species is of some interest. Zimmermann had overlooked Crotch’s obscure proposal 1 This March, 1831, paper by Say, published at New Harmony, Indiana, is not included in LeConte’s 2-volume edition of Say’s writings; see terminal bibliography for further comment. There is a copy of Say’s 1831 paper in the library of the U. S. Department of Agriculture, Washington, D.C.; I have a photographic copy of the title page and pages 5 to 7 before me. The description in Say’s 1834 paper is usually cited as the original; it is word for word the same, but differs in 9 instances of punctuation, and it has the added line, ‘For this species I am indebted to Mr. Barabino,’’ which was of course covered by the title of the 1831 paper. CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. bo a bo ELY TRON 55>=--EDGE OF MESOSTERNAL GROOVE PSS GUN Mib= METASTERNAL AREA SS FeAT; METACOXAL PROCESS~* SUPHISELLUS AGABINUS Ficures 3-8. Structures of Copelatus, Suphisellus, and Agabinus species. FIGURE 3. Front tibia of male of Copelatus glyphicus, spines and setae omitted, with arrow pointing to notch. Ficure 4. Lateral view of apparent 5th abdominal sternite of male of Suphisellus gtbbulus (Aubé) (stippled), with arrow indicating the position of a transverse impression. Ficure 5. Front tibia of male of Copelatus chevrolati renovatus. FicuRE 6. Lateral view of apparent 5th abdominal sternite of male of Suphisellus bicolor, showing the non- impressed median line (though the sides may have impressions). FicureE 7. Ventral view of Suphisellus bicolor to show the conjoined median metasternal area and hind coxal plates; spines and setae omitted. Ficure 8. Part of undersurface of Agabinus glabrellus to show the plate-like hind coxal processes; spines and setae omitted. VoL. XXXVII] LEECH: WATER BEETLES NEW TO CALIFORNIA 243 of the generic name, and in 1921, p. 187 coined the same name for what is in fact the same generic concept. Though he was at the time writing about and specifically named only South American species, he stated that his new generic name applied to all American (in the broad sense) species which he had for- merly listed in Canthydrus in Junk’s Coleopterorum Catalogus (1920), (except for C. buqueti (Laporte) and the two species C. octoguttatus and C. uniformis he was about to describe, all from South America and all three true species of the otherwise Old-World genus Canthydrus). Since he did not actually list S. bicolor by name when proposing his generic name Suphisellus, his name and Crotch’s are objectively different taxa, and Swphisellus Zimmermann, 1921, is a junior homonym, as well as a junior synonym, of SupAisellus Crotch, 1873. Guignot (1946, p. 116) designated S. varicollis Zimmermann, 1921, as the type species of Suphisellus Zimmermann. Say described S. bicolor as from “Louisiana,” collected by Joseph Barabino, a resident of New Orleans who was Say’s agent there for shipments received (Weiss and Ziegler, 1931, p. 176; according to the same authors, p. 110, Say also had collected in Louisiana; see their map on p. 189, taken from Barber, 1928, p. 16. Barber, p. 19, presumed a landing at New Orleans for Say). It seems reasonable to designate New Orleans as the restricted type locality for S. bicolor, and I hereby do so. Of the various references to this species in our literature (usually as Suphis or Canthydrus bicolor), probably most do not apply to the true S. bicolor at all. Their cited distributions alone will disqualify those of Crotch, 1873, p. 397; Sharp, 1882, p. 271; Blatchley, 1910, p. 208. The taxonomy of the Nearctic species of Swphisellus is difficult and in need of further study. Young (1954, p. 131) did not recognize S. bicolor amongst more than 2700 specimens of the genus from Florida. He remarked that S. gibbulus (Aubé) is closely allied and possibly only a geographic variant or race, but also said that his concept of S. gibbulus might include several forms and perhaps species. I have not seen S. bicolor from further east than Mobile, Alabama; it is interesting that Young gives Mobile County, Alabama, for his furthest west example of S. gibbulus, and Léding (1945, p. 27) lists both for the county. It appears that adequate collections from southern Alabama should clarify the status of S. gibbulus Aubé, though in a recent letter F. N. Young mentions having taken both fairly typical S. bicolor and the unicolorous S. gibbulus-like form in southern Indiana and in Illinois. Following are the locality records for the 34 examples of S. bicolor at hand. Lourstana: New Orleans, 4 collected by Bock’, but without date, and 3 taken 2The collector, “Bock,” was presumably Dr. George W. Bock of St. Louis, Missouri. In a short obituary notice E. P. Meiners wrote, ‘“He was at one time a very enthusiastic beetle collector and carried on a rather extensive correspondence with many of the older collectors.” See Entomological News, vol. 52, no. 4, p. 119, issue for April, 1941. 244 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47H SER. in May but without a collector’s name, all from the A. Fenyes collection; 2, Port Sulphur, 4 April 1944, D. E. Beck coll. no. 301 (H. P. Chandler collec- tion). ALABAMA: 2, Carrie’s Lake, Mobile, 17 September, 1945, Mike Wright collector (H. B. Leech collection). ARKANSAS: 1, Hope, 28 June 1932, Louise Knobel collector (H. B. Leech collection). Trxas: 8, Fish Lake, Brazos County, 9 December 1964, and several dates in March and April, 1965, F. S. Conte collector. CALIFORNIA: 5, Deer Creek at Porterville-Fountain Springs Road a few miles south of Porterville, Tulare County, 9 April 1968, H. B. Leech collector; 9, same data, 12 April. Through the kindness of Mr. Conte I have also seen an additional large series from his Fish Lake locality. All specimens are conspecific with those from New Orleans, and represent the only Nearctic species known to me which agrees with Say’s description of S. bicolor. The yellow to reddish yellow head and pronotum, the rich black elytra with contrasting brownish yellow apices, faintly aeneous surface reflec- tions and relatively coarse punctation, define it well (fig. 2). The eyes may be all black, as described by Say, or partially or completely white, depending on methods of preservation. The pronotum is usually more reddish than the head, and may be tinged with piceous discally at the front and back margins; the elytra may be narrowly reddish at and behind the humeral angles; the under-surface varies in color, the raised plate-like metasternal keel and ad- joining laminate inner processes of the hind coxae being darkest. Only one specimen, that from Hope, Arkansas, has the elytra appreciably reddish near the suture. These color characters readily separate S. bicolor from even the darkest examples of S. gibbulus I have seen. In addition there is a structural difference commented on by Young: in S. gibbulus the last abdominal sternite is shallowly but distinctly transversely impressed at or just before the middle (fig. 4); in S. bicolor the sternite may or may not be impressed on each side of the median line, but the median area is on a plane from front to back, not impressed (fig. 6). Suphisellus bicolor and species of the genus Hydrocanthus may be easily separated from all other aquatic beetles known from California. In these two genera the prosternal process and the median area of the meso- and metasternum form a continuous raised flat plate, triangular in shape (fig. 7). In Suphisellus the apex of the prosternal process is at least twice as wide as its width between the front coxae, but not broader than long; in Hydrocanthus°* it is at least two and one-half to three times its width between the coxae, and broader than long. 3 As mentioned in Aquatic Insects of California, p. 326, Zimmermann in 1928 cited ‘‘Californien”’ for one of his specimens of Hydrocanthus similator Zimmermann, but no other California examples of the genus are known and its occurrence here is in doubt. The species resemble those of Suphisellus but are larger, over 4.0 mm. as against usually less than 3.0 mm. for the latter. In addition there are com- plete lateral marginal lines on the pronotum in Hydrocanthus, while they occur only in the basal half in Suphisellus. VoL. XXXVIT] LEECH: WATER BEETLES NEW TO CALIFORNIA 245 The only genus in the western United States having a structure with any re- semblance to this is the dytiscid Agabinus; it contains two black species, 6 to 8 mm. in length but here only the metacoxal processes are differentiated and flattened (fig. 8). ADDENDUM After the above was set in type, Mr. Joe Schuh of Klamath Falls, Oregon, happened to mention that he had a specimen of Suphisellus from Porterville, California. It agrees exactly with the preceding description and figure of S. bicolor and is labelled ‘Porterville, Calif., Tulare County, April 16, 1961. Ed Ball, coll.” So the species has been resident in that area for some years, but how it reached there remains unexplained. ACKNOWLEDGMENTS F. N. Young was so kind as to verify the identification of two California specimens of Copelatus glyphicus; F. S. Conte allowed me to cite a Texas lo- cality where he took Suphisellus bicolor in numbers; W. I. Follett was con- sulted on a point of synonymy; J. A. Powell and N. Bell loaned specimens from the California Insect Survey collection, Berkeley; and B. A. Waldron, District Ranger of the Forest Service, gave local information on the California Hot Springs area; their help is much appreciated. LITERATURE CITED Barber, H. S. 1928. Thomas Say’s unrecorded journey in Mexico. Entomological News, vol. 39, no. 1, pp. 15—20, 1 map. BEQUAERT, J. 1951. A bibliographic note on Say’s two tracts of March, 1831, and January, 1832. Psyche, vol. 57, no. 4, p. 146. (This is the issue of the journal for Decem- ber, 1950, but according to the notice on the inside front cover of vol. 58, no. 1, the December issue was actually published on April 23, 1951.) BLATCHLEY, W. S. 1910. An illustrated descriptive catalogue of the Coleoptera or beetles (exclusive of the Rhynchophora) known to occur in Indiana, with bibliography and description of new species. Indiana Department of Geology and Natural Resources, Bulletin 1. Nature Publishing Co., Indianapolis. 1386 pp., 595 figs. in text, 1 map. (The last numbered figure, on p. 1364, is 590; this is an error for 595, as there is a fig. 590 on p. 1359. Actually, there are no figs. 257 to 265 inclusive, but there are nine ‘“a’’ supplementary figures, so the total is 595. Figures 374 and 375 are out of sequence, but present.) Crark, H. 1863. Catalogue of the Dytiscidae and Gyrinidae of Australasia, with descriptions of new species. Part IJ. Journal of Entomology, vol. 2, pp. 14-23. (Not seen by H.B‘L.) CrotcyH, G. R. 1873. Revision of the Dytiscidae of the United States. Transactions of the American Entomological Society, vol. 4, no. 3-4, pp. 383-424. 246 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SEr. GuIcnor, F. 1946. Génotypes des Dytiscoidea et des Gyrinoidea. Revue francaise d’Entomologie, vol. 13, pp. 112-118. 1952a. Description de dytiscides inédits de la Collection Régimbart. Revue frangaise d’Entomologie, vol. 19, no. 1, pp. 17-31, 8 figs. in text. 1952b. Description de dytiscides inédits de la Collection Régimbart et de quelques autres espéces et variétés nouvelles. Revue francaise d’Entomologie, vol. 19, no. 3, pp. 166-171, 1 fig. in text. LeContTeE, J. L. 1859. [As editor of] American entomology: A description of the insects of North America, by Thomas Say, with illustrations drawn and colored after nature. Edited by John L. LeConte, M.D., with a memoir of the author, by George Ord. Bailliere Bros., New York, Vol. 1: xxxiv +412 pp., 54 col. pls., 1 plain pl. Vol. 2: vi+ 814 pp. (Reprinted in 1869 by Estes and Lauriat, Boston; in 1885 by S. E. Cassino, Boston; in 1891 by Foote, Philadelphia. The re- print editions lack the plain plate of Cicindelidae.) Leecu, H. B. 1948. Coleoptera: Haliplidae, Dytiscidae, Gyrinidae, Hydrophilidae, Limnebiidae. No. 11 zm Contributions toward a knowledge of the insect fauna of Lower California. Proceedings of the California Academy of Sciences, 4th series, vol. 24, no. 11, pp. 375-484, incl. pls. 20, 21. 1956. In H. B. Leech and H. P. Chandler, Aquatic Coleoptera, Chapter 13 im R. L. Usinger, ed., Aquatic insects of California with keys to North American genera and California species. University of California Press, Berkeley and Los Angeles. (Chapter 13, pp. 293-371, 61 figs. in text.) Lenc, C. W. 1920. Catalogue of the Coleoptera of America, north of Mexico. John D. Sherman, Jr., Mount Vernon, N.Y. viii+ 470 pp., 1 folding chart. (Includes Cata- logue of North American Coleoptera described as fossils. By H. F. Wickham, pp. 347-365. According to Mr. Leng’s introduction to the main catalogue, “The text for the water beetles was prepared by Mr. John D. Sherman, Jr.”) Lopinc, H. P- 1945. Catalogue of the beetles of Alabama. Geological Survey of Alabama, Mono- graph 11, pp. 1-172. SAveue 1823. Descriptions of insects of the families Carabici and Hydrocanthari of Latreille, inhabiting North America. Transactions of the American Philosophical So- ciety, New Series, vol. 2, pp. 1-109. 1831. Descriptions of new species of North American insects, found in Louisiana by Joseph Barabino. 19 pp. “March 1831. Indiana, printed at The School Press, New-Harmony.” (NOT “New species of North American insects found by Joseph Barabino chiefly in Louisiana,” a 16-page paper by Say, published at New Harmony in January, 1832. This latter is the one included in the LeConte edition of Say’s complete writings, vol. 1, pp. 300-309. The former is omitted entirely. See Scudder, 1899; Bequaert, 1951.) 1834. Descriptions of new North American insects and observations on some already described. Transactions of the American Philosophical Society, New Series, vol. 4, pp. 409-470. (According to LeConte in a footnote on p. 521 of vol. 2 of his edition of Say’s works, this paper was issued as part of an 8 vo. pamphlet published in New Harmony, Indiana, and dated 1829-1833.) Vor. XXXVII] LEECH: WATER BEETLES NEW TO CALIFORNIA 247 1859. In J. L. LeConte, editor, American entomology, a description of the insects of North America, by Thomas Say, with illustrations drawn and colored after nature, vol. 2, pp. vi+ 814. (See entry under LeConte, 1859.) SCHAEFFER, C. 1908. On North American and some Cuban Copelatus. Journal of the New York Entomological Society, vol. 16, no. 1, pp. 16-18. ScHuH, J. 1969. (Letter of December 23, to H. B. Leech.) Scupp_er, S. H. 1899. An unknown tract on American insects by Thomas Say. Psyche, vol. 8, no. 273, pp. 306-308. (Pages 307-308 include signed parts by P. R. Uhler and C. R. Osten Sacken.) SHARP, D. 1882. On aquatic carnivorous Coleoptera or Dytiscidae. Science Transactions of the Royal Dublin Society, series 2, vol. 2, pp. 179-1003, pls. 7-18. SHERMAN, J. D., Jr. 1920. (See comment in entry under C. W. Leng, supra.) SUBLETTE, J. E., and M. S. SuBLETTE 1967. The limnology of playa lakes on the Llano Estacado, New Mexico and Texas. Southwestern Naturalist, vol. 12, no. 4, pp. 369-406, 15 figs. in text. WeIss, H. B., and G. M. ZIEcLER 1931. Thomas Say, early American naturalist. Charles C Thomas, Springfield, Illinois, and Baltimore, Maryland. Pp. xvi + 260, 8 unnumbered figs. in text, frontis- piece and 18 unnumbered pls. WoopwortTH, C. W. 1913. Guide to California insects. The Law Press, Berkeley. Pp. vi + 360, 361 figs. in text. Younc, F.N. 1942. New species of Copelatus from the West Indies (Coleoptera: Dytiscidae). Pro- ceedings of the New England Zoological Club, vol. 20, pp. 79-94, pl. XIII. 1954. The water beetles of Florida. University of Florida Studies, Biological Series, vol. 5, no. 1, pp. ix + 238, 31 figs. in text. 1963. The Nearctic species of Copelatus Erichson (Coleoptera: Dytiscidae). Quar- terly Journal of the Florida Academy of Sciences, vol. 26, no. 1, pp. 56-77, 11 figs. in text. 1968. (Letter of May 23, to H. B. Leech.) ZANE, M. 1969. From Bakersfield by boat, the put-put odyssey. San Francisco Chronicle, issue for June 19, 1969. 105th year, whole no. 169, p. 5. Z:MMERMANN, A. 1921. Beitrage zur Kenntnis der siidamerikanischen Schwimmkdaferfauna nebst 41 Neubeschreibungen. Archiy fiir Naturgeschichte, vol. 87, no. 3, pp. 181-206. 1928. Neuer Beitrag zur Kenntnis der Schwimmkafer. Wiener Entomologische Zeitung, Bd. 44, Heft 3 & 4, pp. 165-187. gemiee 99 } a ei a sah. 2 1 este tear My ; , 7 4“ OQ srr) 7 we kd aN vi at es! ae | ¥ i> 4 7 = a 18 mae s 4a v \ ai pe 4 . Th. jm 26 > ea call — 7 ‘ t vi aA 4 4s oP : i; P| a A log ea a 1 © ths ‘ Veer aX — p iS Fs , Pa ane | 4 } : as haa if " PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XXXVII, No. 7, pp. 249-274; 25 figs. March 6, 1970 REVIEW OF THE SNAKES OF THE GENUS LYTORHYNCHUS By Alan E. Leviton and Steven C. Anderson California Academy of Sciences, San Francisco, California 94118 The nonvenomous snakes of the genus Lytorhynchus are distributed from the Atlantic coast of North Africa eastward to the west Indian frontier. Included at one time or another in this group are species currently referred to Phyllorhynchus, a small distinctive genus of deserticolous North American snakes with which the Southwest Asian forms really have no affinities. The lytorhynchids, though not especially diversified, seem best adapted to arid habitats. Reported from rocky as well as sandy deserts of the Palearctic region, members of the genus have been taken at or at least near coastal areas at sea level in Saudi Arabia and North Africa, and at altitudes in excess of 8000 feet (according to Minton, 1966, p. 131, but see our remarks on distribution under L. ridgewayi). With the exception of two nominal species, L. diadema and L. ridgewayi, the seven remaining nominal species, and three additional nominal subspecies presently assigned to the genus, were known until recently from less than two dozen specimens. Through the kindness of museum curators of several institutions we have been privileged to examine the types of most of the nominal taxa. Further, due to the efforts of Mr. John Gasperetti, Field Associate of the Department of Herpetology of the Academy, and Mr. Jeromie Anderson, we have seen substantially more material of the eastern populations of this genus than has anyone heretofore. We are indebted to several individuals: to Mr. John Gasperetti, first and foremost for depositing at the Academy his excellent Southwest Asian collections, including two separate series of L. gaddi; Dr. Sherman A. Minton, Jr., for providing additional data on the specimens of Lytorhynchus he collected while [249] aa } Marine Rid| eral Labora.ory LieprRaAar |} MAR 1 61970 | WOODS HOLE, MASS, jae ae ren oo 250 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. in Pakistan and which were loaned to us by Dr. Charles M. Bogert and Dr. Richard G. Zweifel of the American Museum of Natural History, New York (AMNH); Dr. Robert F. Inger and Mr. Hymen Marx for the loan of specimens in the collections of the Field Museum of Natural History (FMNH); for similar courtesies by Dr. Donald W. Tinkel of the Museum of Zoology, University of Michigan (UMMZ), Dr. Robert C. Stebbins, Museum of Vertebrate Zoology, University of California at Berkeley (MVZ), Dr. James A. Peters, United States National Museum, Washington, D.C. (USNM), Mr. J. C. Daniel, Bombay Natural History Society, Bombay, India (BNHS), Dr. Ilja S. Darevsky, Zoo- logical Institute, Academy of Sciences, Leningrad, USSR (ZIAS), and Dr. Yehudah L. Werner, Hebrew University, Jerusalem, Israel (HUJ). David Leviton assisted in making counts and measurements of specimens examined. All drawings were executed by Miss Betsey Hutchings. Genus Lytorhynchus Peters Lytorhynchus Peters, 1862, Monatsb. Akad. wiss. Berl., p. 273 (type species Heterodon diadema Dumeéril, Bibron and Duméril, by monotypy). Chatachlein Jan, 1863, Arch. Zool. Anat. Phys., vol. 2, p. 228 (type species Heterodon diadema Dumeéril, Bibron and Dumeéril, by monotypy). Acantiophis GUNTHER, 1875, Proc. Zool. Soc. London, p. 232 (type species Acantiophis para- doxus Ginther, by monotypy). Catachlaena BLANFORD, 1881, Proc. Zool. Soc. London, p. 678 (emendation of Chatachlein Jan, therefore takes same type species). Lythorhynchus DouMERGUE, 1901, Essai Fauna Herp. Oranie, p. 267 (emendation of Lyto- rhynchus Peters, therefore takes same type species). DerFINiITION. Maxillary teeth 6—9, last two longer than others and separated from them by an interval; palatine teeth 3-5; pterygoid teeth 0-3. Head slightly distinct from neck, with cuneiform, projecting snout; premaxillary either nar- rowed and pointed, or truncate with concave depression at tip, lateral horns (maxillary processes) arise about midway between ends, project posteriorly and form acute angle with longitudinal axis; supraoccipital does not border foramen magnum; eye moderate or large, with vertically elliptic pupil; rostral large, projecting, angularly bent in profile, concave inferiorly; nostril oblique slit between two large nasals. Body elongate, cylindrical; scales smooth or feebly keeled, without apical pits, in 19: 19: 17 or 15 rows; ventrals obtusely angulate laterally; tail moderate or short, subcaudals paired. Hemipenes symmetrical, spinose for half their length with shallow calyces in the distal half. Remarks. Described in 1862 by Karl Peters, the type species is Heterodon diadema, by monotypy. Various authors have sought to include in Lytorhynchus the North American species of leaf-nosed snakes, currently referred to Phyllo- rhynchus by North American herpetologists, the most recent being that of Andrushko and Mikkay (1964). We cannot subscribe to this view. The two nominal genera differ in many ways. For example, the premaxillary of Phyllo- Vor. XXXVII] LEVITON & ANDERSON: REVIEW OF LYTORHYNCHUS 251 FicurE 1. Ventral view of premaxilla of Phyllorhynchus decurtatus perkinsi (CAS 79687). rhynchus (fig. 1) is a large stout structure with the maxillary processes rising from its anteriormost end and projecting at almost right angles from the body of the premaxillary bone. The structure in Lytorhynchus (figs. 2 and 3), besides being narrower, has the maxillary processes rising from a point about at the middle of the core and extending posteriorly at an acute angle. We might also cite the differences in the supraoccipital which in Phyllorhynchus borders the Ficure 2. Ventral view of premaxilla of Lytorhynchus gaddi (CAS 97609). CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. bo UL LSS) ote “A a :_— Ficure 3. Ventral view of premaxilla of Lytorhynchus ridgewayi (CAS 101409). foramen magnum and prevents the exoccipitals from meeting in the midline (fig. 4), but in Lytorhynchus is excluded from the border of the foramen magnum by the exoccipitals (fig. 5). The pterygoid in Phyllorhynchus (fig. 6B) is shorter and stouter and the minute teeth are so positioned that the dentigerous Ficure 4. Dorsal view of occipital region of skull of Phyllorhynchus decurtatus perkinsi (CAS 79687). VoL. XXXVII] LEVITON & ANDERSON: REVIEW OF LYTORHYNCHUS 253 Ficure 5. Dorsal view of occipital region of skull of Lytorhynchus gaddi (CAS 97609). portion about equals the non-dentigerous portion; in Lytorhynchus (fig. 6A) the minute pterygoid teeth occupy only about one-fifth of the pterygoid and are positioned near its anterior end. Underwood (1967) has suggested, by reason of his placement of PAyllorhyn- chus in his family group ‘“Colubridae” and Lytorhynchus in the group “Nat- ricidae,” that the hemipenes in the former are asymmetrical, while in the latter he suspected they were symmetrical, this requiring confirmation. At hand we have one male L. paradoxus with a fully everted symmetrical hemipenis. How- ever, we find in another character used by Underwood that the anterior Vidian foramen is not closer to the lateral border of the basisphenoid in Lytorhynchus than in Phyllorhynchus, his statement to the contrary notwithstanding, in our material (3 skulls of Phyllorhynchus decurtatus and 4 of Lytorhynchus, L. gaddi, L. ridgewayi, L. maynardi, and L. paradoxus). Thus, while we agree completely that Phyllorhynchus and Lytorhynchus are entirely distinct genera, we cannot confirm the basis for his allocation of these genera to different family groups. The nominal genus Lytorhynchus is restricted to the Old World, specifically North Africa and Southwestern Asia. Lytorhynchus diadema, L. kennedyi, and L. gaddi are members of one section of the genus; we are not at all certain how 254 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Figure 6. Ventral view of palatopterygoid arch of: A. Lytorhynchus gaddi (CAS 97609) ; B. Phyllorhynchus decurtatus perkinsi (CAS 79687). Vout. XXXVII] LEVITON & ANDERSON: REVIEW OF LYTORHYNCHUS 255 to treat L. ridgewayi, L. paradoxus, and L. maynardi, but it seems likely that the latter two are more closely allied to one another than is either to L. ridgewayi. Lytorhynchus diadema is the most widely distributed member of the genus, being found in North Africa from the Atlantic coast to the Red Sea and thence in the Sinai and northward to the west of the “Jordan Rift” along the Mediterranean coast at least to Jerusalem. Kramer and Schnurrenberger (1963, p. 505) give counts which suggest the possibility that geographical races might be defined, but on the basis of data and material available to us we cannot recognize taxonomically distinct subunits. Lytorhynchus gaddi appears to be widely distributed in the Arabian Peninsula (fig. 7) south and east of the “Jordan Rift,” north into Iraq and around the head of the Persian Gulf into extreme southwestern Iran. The last member of the ‘“‘diadema” group, L. ken- nedyi, is known with certainty only in Syria from between Homs and Palmyra, and if we are correct in presuming Werner’s specimen of L. diadema is a “‘ken- nedyi,” then it is known from west of Damascus too, both localities for L. kennedyi being in the lowland desert north and east of the “rift.” The distributions of the eastern and western species complexes of Lyto- rhynchus overlap in the Province of Fars in southwestern Iran. Lytorhynchus ridgewayi has managed to extend its distribution as far west as Ahram, near Bushire, in Iran, and north into Transcaspia. The easternmost species, Lyto- rhynchus paradoxus, has been taken at a number of localities, mostly along the trend of the Indus River, though not necessarily in its immediate vicinity. Lytorhynchus maynardi appears isolated in the interior basins of northern Baluchistan and southern Afghanistan; so few specimens have been taken, how- ever, that the extent of its distribution is still uncertain. Kry TO THE SPECIES OF THE GENUS Lytorhynchus la. Rostral shield broadly truncate, as broad at its base as at its anteriormost projection. 2a. Pattern of black crossbars very sharply defined, bars with intense black pigment, Pe=Vablotchessonsbodya-s On tall. =. Ee L. kennedyi 2b. Pattern of dark brown to black ovoid bars distinct or not, but if distinct not intense black, each scale in dark bar usually with light center; 30-55 blotches on body, 9-13 on tail. 3a. Ventrals 155-177; total of ventrals and subcaudals 197-215 _______ L. diadema 3b. Ventrals 173-195; total of ventrals and subcaudals 212-240 _____________. L. gaddi 1b. Rostral shield narrowed or pointed, broader at its base than at its anteriormost projection. 4a. Anal shield usually single, occasionally divided; prefrontals usually united ; ventrals 165-188; ground color grayish brown with series of darker brown, not Pivelombinbcies OL GrOsSbars: 2 L. ridgewayi 4b. Anal shield usually divided; prefrontals paired; ventrals variable. 5a. Ventrals 169-185; ground color grayish brown with series of darker brown, not black, blotches, not crossbars ———___—_ L. paradoxus Sb. Ventrals 187-202; ground color yellowish green, with distinct well defined SERIESMO TET AGRO Walla Ger GhOSS asm L. maynardi [Proc. 4TH Serr. CALIFORNIA ACADEMY OF SCIENCES so | in T [ikea ee VISV LSAMHLNOS t\ADmMoaBpis “JT x snxoppipd ‘] v ipapuAnw ‘7 1\Apauuay “1 Oo Ippob 7 e@ pDWAPPIP “J x 1 of NVINVUUNFZLIGIGA | ese | ey ee erat NS IE 00 i= ares a Vor. XXXVIT] LEVITON & ANDERSON: REVIEW OF LYTORHYNCHUS 257 “DIADEMA” group DEFINITION. Rostral truncate, as broad at its base as at its anteriormost projection; anterior portion of premaxillary broad, slightly bilobate, choanal portion broad and deeply notched (fig. 1). Included species: Lytorhynchus diadema (Duméril, Bibron and Duméril) ; Lytorhynchus gaddi Nikolsky; Lytorhynchus kennedyi Schmidt. Lytorhynchus diadema (Duméril, Bibron, and Duméril). (Figures 8-10.) Heterodon diadema DuMERIL, BiBRoN, and DuMERIL, 1854, Erp. Gén., vol. 7, p. 779 (type loc: Algeria; type in Paris Museum [MHNP 7560]). Gervais, 1857, Mém. Ac. Montpellier, vol. 3, p. 511, pl. 5, fig. 1. Kiunzincer, 1878, Zeitschr. Ges. Erdk. Berl., vol. 13, p. 95 (not seen). Simotes diadema, GUNTHER, 1858, Cat. Snakes British Mus., p. 26 (Algiers; N. Africa). STRAUCH, 1862, Erp. Algeria, p. 53 (not seen). Bortrcrr, 1879, Ber. Senckenberg Mus., p. 61 (not seen). Chatachlein diadema, JAN, 1863, Arch. Zool. Anat. Phys., vol. 2, fasc. 2, p. 228 (18 of reprint) (Algerie) ; 1865, Icon. Gen., livr. 10, pl. 6, fig. 2. Lytorhynchus diadema, PETERS, 1862, Monatsber. Akad. wiss. Berlin, p. 272, pl. 1, fig. 1 (new combination). BOULENGER, 1891, Trans. Zool. Soc. London, vol. 13, pp. 145-146 (Tunisia {Mraier; Ferriana] description; distribution). Konic, 1892, Verh. (S. B.) nat. Verh. Bonn, p. 22 (not seen). Matscutr, 1893, Sitzb. Ges. Naturf. Berlin, p. 31 (not seen). BouLeENGER, 1893, Cat. Snakes British Mus., vol. 1, p. 415 (part: Algeria; Egypt [Gizeh, Abou-Roash (= Abu Rawash)]). Otivirer, 1894, Mém. Soc. Zool. France, vol. 7, p. 119 (not seen). FRANCAV-GLIA, 1896, Bull. Soc. Rom. Stud. Zool., vol. 5, p. 36 (not seen). ANDERSON, 1896, Herp. Arabia, pp. 87, 107 (part: Algerian Sahara to Egypt [W. bank Suez Canal between Suez and Ismailia, Abu Roash (= Rawash), Gizeh], Senaar District). BoetTTcer, 1898, Kat. Rept. Senckenberg Mus., II. Schlangen, p. 46 (Jaffa; Siid-Tunis). ANDERSON, 1898, Zool. Egypt., Rept. and Batr., p. 271, pl. 37, fig. 3 (part: Egypt [be- tween Suez and Ismailia, Abu Roash (= Rawash), Gizeh]; Ain Sefran, Algeria; Jaffa). FLow_Er, 1933, Proc. Zool. Soc. London, 1933, p. 815 (Egypt [Bir Hooper, Giza Pyramids, Eiu el Shams, Ezbet el Zeitun, Abu Zabal, Kantara, Ismailia, Mahadet, Mohammadia] ; Palestine [Gaza, Rashleh]). Vir1rers, 1950, Bull. Inst. Francaise Afr. Noire, Initiations Africains II, p. 79 (part: “Afrique du Nord . . . Mauritanie”). ScumiptT AnD Marx, 1956, Fieldiana: Zool., vol. 39, p. 30 (Sinai). SocHuREK, 1956, Aquar. u. Terr., Leipzig, vol. 3 (W. Algeria; not seen). Vitters, 1956, Bull. Inst. Francaise Afrique Noire, vol. 18A (Mauritanie). Sarnt-Girons, 1956, Arch. Sci. Inst. Cherifien, Rabat, no. 8 (Mo- rocco). ScHmiptT anD Marx, 1957, Bull. Zool. Soc. Egypt, vol. 1955-56, no. 13, p. 25 (Egypt [2 mi. N Bir Kansisrob]). ScHmMipt AND INGER, 1957, Living Reptiles of the World, figure on p. 223. KRAMER AND SCHNURRENBERGER, 1958, Aquar. Terr. Zeits., vol. 11, p. 57 (Libya [El Hameimat; sudlich von El Adem]) ; 1959, Mitt. naturf. Ges. Bern, N.S., vol. 17 (Libya). Domercur, 1959, Arch. Inst. Pasteur Tunis, vol. 36 (N. Africa). ViriiErS, 1963, Inst. Afrique Noire, Initiations Africaines II, ed. 2, p. 111, text figs. 136-138 (part: “Afrique du Nord .. . Mauritanie”). Marx, 1968, U.S. Naval Med. Res. Unit No. 3, Cairo, Egypt, Spec. Publ., p. 34 (Egypt [Sinai: St. Catherine’s Monastery Ficure 7. Distribution of species of Lytorhynchus in southwestern Asia. 258 CALIFORNIA ACADEMY OF SCIENCES (Proc. 47H Serr. Ficure 8. Dorsal view of head of Lytorhynchus diadema (USNM 134681). area, Raba. Southeastern Desert: Gebel Elba, 3.2 km. N Bir Kansisrob. Giza: Abu Rawash area, Manshiyet, Radwan, El Qatta. Faiyum: Kom O Shim, Fanus, Wadi Muwellih Bir Dakaar area. Matruh: Burg el Arab]). Lytorhynchus diadema diadema, Haas, 1952, Copeia, no. 1, p. 22 (Palestine). PASTEUR AND Bons, 1960, Trav. Inst. Sci. Cherifien Rabat, zool. ser., no. 21, p. 84 (“Sahara et Haubs- Plateaux”). SCHNURRENBERGER, 1962, Vischr. naturf. Ges. Zurich, vol. 107 (Libya; not seen). KRAMER AND SCHNURRENBERGER, 1963, Rev. Suisse Zool., vol. 70, p. 504 (Libya). Lithorhynchus diadema, DOUMERGUE, 1901, Essai Fauna Herp. Oranie, p. 267. Lithorhynchus diadema hirouxii DOUMERGUE, 1901, Essai Fauna Herp. Oranie, p. 269, pl. 20, fig. 5a (type loc: d’Ain Sefra or Mecheria, Algeria; type unknown, probably in Paris Museum). Lytorhynchus diadema hoggarense ANGEL, 1944, Bull. Mus. Hist. Nat. Paris, ser. 2, vol. 16, p. 419 (type loc: Hoggar, Sahara; type in Paris Museum). MATERIAL EXAMINED (57). EGYPT: lower Nile River Valley, Cairo to Alexandria (USNM 128205). Cairo (USNM 56152, 134706). Abu Rawash (FMNH 63121-63122; USNM 124726-124727, 131120-131127, 134675134679, 134681—134683, 136274). 4 mi. NW of Abu Rawash (FMNH 63123-63125). 4 mi. W of Abu Rawash (USNM 131119). 6-7 mi. W of Abu Rawash (USNM 131115—131118, 134680). El Mansuriya (USNM 134844-134847). Manshiyet Radwan (FMNH 75338). Gizeh (MCZ 21056). El Qatta, Imbaba (FMNH 129885). Sakkara (FMNH 63120). Shamama Halt [railroad station about midway between El] Almein and El Haman] (MCZ 46843). Km. 121 along road from Cairo to Alexandria, about 21 mi. NW of Wadi Natrun (MCZ 46842). Kom-O-Shim (FMNH 67246-67247). Fanus, Tamiya (FMNH 82668). Burg el Arab, Mariut (FMNH 67245). St. Catherine Monastery, El Roba, Sinai Vot.XXXVII] LEVITON & ANDERSON: REVIEW OF LYTORHYNCHUS 259 Ficure 9. Lateral view of head of Lytorhynchus diadema (USNM 134681). (FMNH 72111). 2 mi. N Bir Kansisrob, Sudan Government Administrative Area (FMNH 73538). ISRAEL: Holon (FMNH 74403). Rehovot (MCZ 52262). LIBYA: Tripoli (USNM 56153, UMMZ 67206). MOROCCO: Aglat Cedr. (MCZ 29917). NIGER: Air: 39 mi. N Tanout [= Tarzut] (MCZ 67901). [PALESTINE: coastal plain (FMNH 48502-48503).| DISTRIBUTION. Reported from Morocco on the northwest coast of North Africa to Egypt, thence north along the coast to Israel. From the southern Sahara it is reported for the first time in this paper from Niger. Confirmed localities include: ALGERIA: Ain Sefran. EGYPT: Gizeh, Abu Rawash, west bank of Suez Canal between Suez and Ismailia, Senaar District, Giza Pyramids, Ein el Shams, Ezbet el Zeitun, Abu Zabal, Kantara, Ismailia, Mahadet, Moham- madia, between Cairo and Alexandria, E] Mansuriya, Manshiyet Radwan, Sak- kara, E] Qatta, Shamama Halt, Kom-O-Shim, Fanus, Burg El] Arab, El Raba, Bir Kansisrob. ISRAEL: Jaffa, Holon, Rehovot. LIBYA: Tripoli, El] Hameimat, south of E] Adem. MAURITANIA: without exact localities. MOROCCO: Aglat Cedr. NIGER: N of Tanzut. Diacnosis. Rostral shield broadly truncate, as broad at its base as its width at its anteriormost projection; pattern of brown to black ovoid blotches or cross- bars usually distinct but never intense black, each scale in dark bars usually with lighter centers; ventrals 152-177 (164.0 = 0.58 SE [N = 65]);' subcaudals 35-49 (42.1 + 0.38 SE [N = 58]);7 ventrals plus subcaudals 197-215 (206.6 + 1Ventrals: o [30] 155-167 (161.9 + 0.55 SE); 2 [27] 161-177 (167.2 = 0.77 SE). 2Subcaudals: of [27] 37-49 (43.6 + 0.52 SE); 9 [24] 35-44 (40.7 + 0.45 SE). 260 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 10. Variation in dorsal color pattern among a series of specimens of Lytorhyn- chus diadema from Abu Rawash, Egypt. 0.51 SE [N = 57]);* body blotches 30-52 (36.6 + 0.5 SE). Standard length (of the largest male and female examined, in mm.); ¢ 429, 2 391; tail length (of the largest male and female examined, in mm.); 4 80, 2 67. Remarks. Most of our specimens came from the vicinity of Abu Rawash, Egypt. We believe we have seen enough material from other localities and have culled enough data from the literature to gain a fairly comprehensive idea of range of variation within this species. We have not seen sufficient samples to comment on the possibility of there being geographically and taxonomically distinct populations within the species, distinguished perhaps by scale counts or small but constant color pattern differences. We must note, though, that the range of pattern variation (size of dark blotches relative to light interspaces) is quite large, even within a single sample from Abu Rawash (fig. 10). Data from Kramer and Schnurrenberger (1963) suggest the Libyan population and perhaps those from western North Africa may be taxonomically distinct at the subspecies level from the Egyptian-Sinai population. We can add nothing to the information already given by Anderson (1898) and Flower (1933) about the habitats or behavior of these interesting animals. 8 Ventrals plus subcaudals: ¢& [27] 197-215 (206.1 + 0.99 SE); 9 [23] 203-215 (207.8 + 0.72 SE). Vot.XXXVII] LEVITON & ANDERSON: REVIEW OF LYTORHYNCHUS 261 Lytorhynchus gaddi Nikolsky. (Figures 11-15.) Catachlaena diadema, BLANForD, 1881, Proc. Zool. Soc. London, 1881, p. 678, figs. 2a—b (Iran [25 mi. S Bushire]; counts and measurements of material examined). Lytorhynchus diadema, BOULENGER, 1887, Ann. Mag. Nat. Hist. ser. 5, vol. 20, p. 408 (Mus- cat); 1893, Cat. Snakes British Mus. vol. 1, p. 415 (in part; Arabia [Muscat], Iran [Bushi]; composite description, counts of material examined). ANDERSON, 1896 Herp. Arabia, pp. 82 and 87 (Arabia [Maskat = Muscat], Aden) ; 1898, Zool. Egypt, Rept. Batr., vol. 1, p. 272 (in part; Iran [Bushi], Arabia [Maskat = Mus- cat]; counts of material examined). BouLeNncer, 1920, Jour. Bombay Nat. Hist. Soc., vol. 27, p. 26 (Iraq [Shaiba, Faleya], Iran [Awaz]). WERNER, 1929, Zool. Jahrb., vol. 57, p. 62 (listed from Iran and Arabia) ; 1936, Festschr. Dr. Embrik Strand, vol. 2, p. 201 (listed from Iran). Scumupt, 1939, Field Mus. Nat. Hist. Zool., vol. 24, p. 74, fig. 6 (Iraq [Baghdad]; measurements and counts; suggests possible affinity with L. gaddi). KuHaAtar, 1960, Publ. Iraq Nat. Hist. Mus., p. 78 (brief composite description based on literature). ANDERSON, 1963, Proc. Calif. Acad. Sci., 4th ser., vol. 31, p. 478 (listed in Iran). Lytorhynchus gaddi N:koisxky, 1907, Ann. Mus. Zool. St. Pétersbourg, vol. 10, p. 294 (type locality: Dizful [= Dezful], Khuzistan, Iran; syntypes [2] in Zoological Institute, Academy of Sciences, Leningrad). ANDERSON, 1963, Proc. Calif. Acad. Sci., 4th ser., vol. 31, p. 478 (listed from Iran). Lytorhynchus diadema mesopotamicus Haas, 1952, Copeia, 1952, p. 22 (type locality: Addaye, 40 km. W of Mosul, Iraq; type in Department of Zoology, Hebrew University, Jerusalem, Israel). KHALAF, 1960, Publ. Iraq. Nat. Hist. Mus., p. 78 (brief description). Lytorhynchus diadema arabicus Haas, 1952, Copeia, 1952, p. 22; Haas, 1957, Proc. Calif. Acad. Sci., 4th ser., vol. 29, p. 80 (type locality: Abqaiq, Arabia; type in California Academy of Sciences; Arabia [Abqaiq, Dhahran, Moreiwa Post]). MATERIAL EXAMINED (23). IRAN: Awaz (BNHS 790); Dezful (ZIAS 10288[1], one of the two syntypes of L. gaddi). IRAQ: Addaye, 40 km. W of Mosul (HUJ 3551, holotype of L. d. mesopotamicus); Baghdad (FMNH 20859); Shaiba (BNHS 789). KUWAIT: (BNHS 791). ARABIAN PENIN- SULA: Abu Dhabi region (CAS 97809-97810, 98092); Dhahran (CAS 84487 [holotype of L. d. arabicus |, 84504, 84526, 84557, 84579, 84507, FMNH 73995 [all paratypes of L. d. arabicus|; Jidda (BNHS 792 [693, 700, 704]; Al Qurayn [26°02’ N, 43°23’ E] (CAS 97580); Riyad al Khabra [26°03’ N, 43°35’ E] (CAS 97584); Buleariyah [26°07’N, 43°38’ E] (CAS 97575); Riyad al Khabra [26°04’ N, 43°36’ E] (CAS 97588). DistrRisuTION. ARABIAN PENINSULA: Abu Dhabi region; Dhahran; Al Qurayn; Jidda; Riyad al Khabra; Buleariyah; Muscat. IRAN: Awaz; Dezful; Bushi. IRAQ: Addaye; Baghdad; Faleya; Shaiba. KUWAIT. 4This name and a very brief indication first appear in Haas’ discussion of the relationships of Eid. mesopotamicus, the author’s conclusions being based on the examination of a series of specimens from northern Arabia in the collections of the California Academy of Sciences and referred to in a later paper (Haas, 1957). Haas made no mention of his specimens at this time nor of any locality other than “‘Arabia’’; nevertheless, the use of the name appears to satisfy the criteria of availability, as detailed in Articles 10-17, International Code of Zoological Nomenclature. If the name is recognized as being available then the entire series of specimens Haas had before him must be regarded as syntypes. Therefore, to insure stability in the application of this name we hereby select CAS 84478 as lectotype inasmuch as this is also the designated holotype of L. d. arabicus Haas, 1957. 262 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser. Ficure 11. Dorsal view of head of Lytorhynchus gaddi (ZIAS 10288, syntype). Diacnosis. Rostral shield broadly truncate, as broad at its base as its width at its anteriormost projection; pattern of brown to black ovoid blotches or cross- bars usually distinct but never intense black, each scale in dark bars usually with lighter centers; ventrals 173-195 (184.7 + 1.22 SE [N = 26]);° sub- 5 Ventrals: of [11] 173-192 (180.8 + 1.83 SE); 9 [15] 180-195 (187.6 + 1.16 SE). Ficure 12. Lateral view of head of Lytorhynchus gaddi (ZIAS 10288, syntype). Vor. XXXVIT] LEVITON & ANDERSON: REVIEW OF LYTORHYNCHUS 263 Ficure 13. Dorsal color pattern of Lytorhynchus gaddi (ZIAS 10288, syntype). caudals 33-47 (42.12 + 0.72 SE |N = 14]);°® ventrals plus subcaudals 212-240 (2276-5 = 136 SE ||N = 26]);* body blotches 33-55 (40.7 = 1.30 SE [N = 21]). Standard length of largest male and female, in mm.): ¢ 428, @ 436; talmensth (in mm.); 3¢ 78, 2 71. Remarks. We have been fortunate in being able to examine the type specimens of each of the nominal forms we refer to this taxon. Each has been photographed, and these are included here (figs. 13-15). Having examined the types of the three nominal forms with care, and having seen a more extensive series of specimens than has heretofore been available, we do not find evidence to justify recognizing any as being distinct from L. gaddi Nikolsky. That L. gaddi is related to L. diadema is obvious. The shape of the rostral, the relation of head shields to one another, and color pattern clearly attest to their closeness. They differ most strikingly in ventral and subcaudal counts, and since we have no reason to suspect the two overlap in these characteristics, or indeed in distribution, we believe it reasonable to recognize the two taxa as distinct species. Lytorhynchus kennedyi, also belonging to this group, differs from L. gaddi in color pattern (fig. 15) in addition to those character differences already mentioned for L. diadema. Worthy of note is the fact that L. gaddi possesses a greater number of sub- caudal plates than either L. diadema or L. kennedyi, and in this character agrees with members of the “ridgewayi” group. However, rostral shape and color pat- tern leave little doubt that L. gaddi must be grouped with L. diadema. 6 Subcaudals: ¢ [11] 39-47 (43.7 £ 0.85 SE); @ [15] 36-46 (40.9 + 1.02 SE). 7 Total of ventrals plus subcaudals: of [11] 212-239 (22.5 + 2.25 SE); @ [15] 220-240 (228.5 + 1.45 SE). 264 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. wi eet “ase Ficure 14. Holotype of Lytorhynchus diadema mesopotamicus (HUJ 3551) [= L. gaddi). Ficure 15. Holotype of Lytorhynchus diadema arabicus (CAS 84487) [= L. gaddi]. VoLt.XXXVII] LEVITON & ANDERSON: REVIEW OF LYTORHYNCHUS 265 Ficure 16. Holotype of Lytorhynchus kennedyi (FMNH 19586). This species has a particularly interesting distribution. Based on available records we must conclude that it is the common lytorhynchid of the Arabian Peninsula east and south of the Jordan rift, and that it has extended its range around the head of the Persian Gulf into southwestern Iran and south and south central Iraq. We have very little information about its habitat, but based on data sent us by John Gasperetti we presume it is a dune dweller, or at least is found in and on loose sand. Lytorhynchus kennedyi Schmidt. (Figure 16.) Lytorhynchus diadema, ANDERSON, 1896, Herp. Arabia, p. 87 (listed from Syria). BOETTGER, 1898, Katalog Rept. Samml. Mus. Senckenberg, II, Schlangen, p. 46 (in part; listed from Syria). WeErNER, 1929, Zool. Anz., vol. 81, p. 242 (Syria [“Syrische Wiiste, 160 km. west- lich Damaskus auf der Route der ehemaligen Wiistenpost bei Saba-Biar’’]; counts on specimen examined). Corxit1, 1932, Snakes and snake bite in Iraq, London, pl. 10. Lytorhynchus kennedyi Scumipt, 1939, Fieldiana, Zool., vol. 24, p. 75, fig. 6 (type locality: between Homs and Palmyra, Syria; type in Field Natural History Museum, Chicago). Lytorhynchus diadema kennedyi, Haas, 1952, Copeia, 1952, p. 22 (listed). MATERIAL EXAMINED (1). FMNH 19586 [holotype] from between Homs and Palmyra, Syria. DistRIBUTION. SYRIA: 160 km. W of Damascus; between Homs and Palmyra. Dracnosis. Rostral shield broadly truncate, as broad at its base as its width at its anteriormost projection; pattern of black crossbars sharply defined, cross- bars narrower than lighter interspaces and intense black; ventrals 163-175 |N = 2]; subcaudals 39-44 [N = 2]; ventrals plus subcaudals 202-219 [N = 2]; body blotches 23-27 [N = 2]; standard length (in mm.): ¢ 327; tail length: S$ 56. 266 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Remarks. We have seen the type specimen only. Werner’s record is referred to this species on the strength of its geographic position and lower number of body blotches. Lytorhynchus kennedyi is very close to L. diadema, apparently differing only in color pattern, though based on our scanty material, this is rather pronounced. We do suspect future collecting will turn up individuals intermediate between L. kennedyi and L. diadema, and at that time its taxonomic status will have to be reviewed. “RIDGEWAYI” group DEFINITION. Rostral narrowed, or pointed, not truncate, broader at its base than its width at its anteriormost projection; anterior portion of premaxillary narrow, projecting; posterior portion also narrow and projecting, not notched (igs) INCLUDED SPECIES. Lytorhynchus maynardi Alcock and Finn; Lytorhynchus paradoxus (Gunther); Lytorhynchus ridgewayi Boulenger. Lytorhynchus maynardi Alcock and Finn. (Figures 17-19.) Lytorhynchus maynardi ALcocK AND FINN, 1896, Jour. Asiatic Soc. Bengal, vol. 65, p. 562, pl. 14 (type locality: near Robat I, Afghanistan-Baluchistan border [see map plate 11]; syntypes [4] in Indian Museum, Calcutta, and British Museum [Natural History], London). ANNANDALE, 1904, Jour. Asiatic Soc. Bengal, vol. 73, p. 208. WALL, 1923, Jour. Bombay Nat. Hist. Soc., vol. 29, p. 619 (Baluchistan). WERNER, 1929, Zool. Jahrb., vol. 57, pp. 62 and 63 (N. Baluchistan; synonymy, distribution, key characters). SMITH, 1943, Fauna British India, vol. 3, Serp., p. 192, fig. 59 (synonymy, description, hemi- penes). Minton, 1966, Bull. American Mus. Nat. Hist., vol. 134, p. 129, pl. 28, fig. 2. (Pakistan [vicinity of Nushki]; description, natural history). MATERIAL EXAMINED (16). PAKISTAN: Nushki (CAS 101359-101360, 101405—101406, 101475). 1.5 mi. W of Nushki (AMNH 88458-88460). 2 mi. W of Nushki (UMMZ 123435). 9-10 mi. NW of Nushki (AMNH 88461- 88463). 1.5 mi. N of Ahmad Wal (AMNH 96241-96242). AFGHANISTAN: 10 km. NE of Darweshan (CAS 120493). 56 km. S and 10 km. E of Darweshan (CAS 120494). DistripuTION. AFGHANISTAN-PAKISTAN FRONTIER: Robat I [alt. 4055 ft.]|. PAKISTAN: vicinity of Nushki; vicinity of Ahmad Wal. AF- GHANISTAN: Registan desert region. Dracnosts. Rostral shield narrow and pointed, not truncate, broader at its base than above; prefrontals divided; upper labial shields do not border eye; ground color yellowish with distinct, well defined series of black crossbars; ventrals 187-202 (195.9 = 0.99 SE [N = 16]) [¢ + 2];8 subcaudals 53-62 (58.7 + 0.65 SE [N = 15]) [¢ + @];°® ventrals plus subcaudals 246-264 8 Ventrals: o& [4] 186-198 (192.3 + 2.1 SE): 9? [12] 9 Subcaudals: & [4] 60-62 (61.0), 9 [11] 53-62 Vor.XXXVII] LEVITON & ANDERSON: REVIEW OF LYTORHYNCHUS 267 Ficure 17. Dorsal color pattern of Lytorhynchus maynardi (CAS 101360). (253.8 = 1.07 SE [N = 16]) [¢ + 2]; black crossbars on body 38-51 (43.9 + 0.89 SE [N = 16]); standard length (in mm.) for largest male and female examined: ¢ 298, 2 334; tail length (in mm.): ¢ 71, 2 70. Remarks. The highest ventral counts combined with its distinctive color pattern makes this species the most striking member of the genus. It is related to the “vidgewayi” group on the basis of its snout and general habitus as well Ficure 18. Dorsal view of head of Lytorhynchus maynardi (CAS 101406). 268 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 19. Lateral view of head of Lytorhynchus maynardi (CAS 101406). as its distribution. Apparently restricted to the Helmand Basin, the major interior drainage of southern Afghanistan and northern Baluchistan, the species lies within the general range of L. ridgewayi (see fig. 7). Its snout structure is much closer to that of L. paradoxus, another member of the “rvidgewayi” group found further east. Interestingly, L. paradoxus possesses the basic color pattern and ventral counts of L. ridgewayi but the head structure of L. maynardi. Dr. Sherman Minton collected in West Pakistan during 1958-1962. In a letter dated 12 October 1966, he made the following comments: “Concerning Lytorhynchus in Pakistan, maynardi and paradoxus seem to be ecological equivalents, both being restricted to fine, wind-blown sand dune habitat. All my maynardi have come from the region around Nushki where it is a common snake. Without referring to notes or map, I guess elevations here to be about 3000-5000 feet. I have paradoxus from several localities in the western Thar (including the desert between Mianwali and Muzzaffargarh which I regard as a western outlier of the Thar) at elevations below 500 feet.” Lytorhynchus paradoxus (Giinther). (Figures 20-22.) Acontiophis paradoxa GUNTHER, 1875, Proc. Zool. Soc. London, 1875, p. 232, fig. 5 (type locality: northern India; type in British Museum [Natural History], London). Murray, 1884, Ann. Mag. Nat. Hist., ser. 5, vol. 14, p. 110 (Sind [Thool Talooka, at Zungipur] ; color pattern, counts) ; 1886, Rept. Sind, p. 84 (Sind [Thool Talooka, at Zungipoor]; description). Lytorhynchus paradoxus, BOULENGER, 1890, Fauna British India, Rept. & Batr., p. 323, fig. 98 (Sind; description, figure of head) ; 1893, Cat. Sn. British Mus., vol. 1, p. 416 (Sind [Zangipur]; description, counts of material examined). Watt, 1923, Jour. Bombay Nat. Hist. Soc., vol. 29, p. 619 (Sind; Punjab [Multan]). Myers, 1947, Herpetologica, vol. 3. p. 167 (listed). Minton, 1962, American Mus. Novitat., no. 2081, p. 16, fig. 52; 1966, Vor. XXXVII] LEVITON & ANDERSON: REVIEW OF LYTORHYNCHUS 269 4 ‘ Ficure 20. Dorsal color pattern of Lytorhynchus paradoxus (CAS 101412). Bull. American Mus. Nat. Hist., vol. 134, p. 130, pl. 29, fig. 1 (Pakistan [Burra; Umar- kot; 11 mi. S. Fatehpur; Baran Nai; near Jamrao Head]; description, habits). Lytorhynchus monticornis WERNER, 1906, Sitzb. Akad. Wiss. Wien, vol. 135, p. 243 (type locality: Sind; type in Vienna). MATERIAL EXAMINED (7). PAKISTAN: Jungshai [= Jungshahi] (CAS 101361, 101411—-101413). 11 mi. S of Fatehpur, Muzaffargarh District (AMNH 88446). Baran Nai, near Bholari, Dadu District (AMNH 89295). Sandhills near Jamrao Head, Sanghar District (AMNH 87461). DistriBUTION. WEST PAKISTAN: Zangipur, Jungshahi, vicinity of Fateh- pur, Baran Nai (SW of Kotri), vicinity of Jamrao Head, Burra, Umarkot. A FicureE 21. Dorsal view of head of Lytorhynchus paradoxus (CAS 101361). 270 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47H Ser. Ficure 22. Lateral view of head of Lytorhynchus paradoxus (CAS 101361). specimen in the collection of the Bombay Natural History Society (BNHS 5188, not examined by us) is from PAKISTAN: Thar and Pakar District. Diacnosts. Rostral shield narrowed and pointed, not truncate, broader at its base than above; prefrontals divided; fifth upper labial usually borders eye; light brown with series of medium brown blotches on dorsum; ventrals 169-185 (176.5 = 1.99 SE [N = 6]) [¢ + 2]; subcaudals 47-51 (48.7 [N= 3))if¢ + 2]; total of ventrals plus subcaudals 220-227 (224.0 [N = 3]) |¢ = 2]; dorsal body blotches 44-51 (48.3 + 1.15 SE [N = 6]) [¢ + 2]; standazd length (of largest male and female examined, in mm.): ¢ 298, 2 325; tail length: ¢ 58, 2 49+ (tip missing; this is the only female seen). REMARKS. We have already recorded Dr. Minton’s observations on L. paradoxus (personal correspondence) in our discussion of L. maynardi. Regret- fully, little is known of this small, secretive snake, few specimens having been taken; those available for study do not shed any further light on its habits or behavior. Indeed, we have seen but seven specimens and have data for three others, the type reported on by Giinther in 1875 and two examined by Murray (1884) who stated that the collector, Mr. F. Gleadow, dug the specimens up from depressions in an area of wind-blown sand forming small hills 20 to 30 feet high. Lytorhynchus ridgewayi Boulenger.!° (Figures 23-25.) Lytorhynchus ridgewayi BOULENGER, 1887, Ann. Mag. Nat. Hist., ser. 5, vol. 20, p. 413 (type loc: Chinkilok, Afghanistan; type in British Museum). Borttcer, 1888, Zool. Jahrb., Syst. Abt., vol. 3, p. 924 (Transcaspia). BoULENGER, 1889, Trans. Linnaean Soc. London, vol. 5, p. 102, pl. 11, fig. 1; 1883, Cat. Sn. British Mus., vol. 1, p. 415. ALcocK AND Finn, 10Tn the accompanying synonymy a number of Russian references are included to which we have not had access. We are unable to confirm these references. They were abstracted from various sources, especially the Zoological Record. VoL. XXXVII] LEVITON & ANDERSON: REVIEW OF LYTORHYNCHUS Ayal Ficure 23. Dorsal color pattern of Lytorhynchus ridgewayi (CAS 84899). 1896, Jour. Asiatic Soc. Bengal, vol. 65, p. 526 (Afghanistan [Saindak, 3000 ft.]). Nrxotsky, 1899, Herp. Turan, p. 59 (not seen). MicuHamovsku, 1904, Ezhegodnik Zool. Mus. Akad. Nauk, vol. 9, p. 43 (not seen). NrkorsKy, 1905, Herp. Rossica, p. 244 (Persia [Terra Zirkuch: Atkaul]; Transcaspia); 1907, Opredel. presm. zemnovod. Rossiskoi imperii Kharkov, p. 114 (not seen). WALL, 1911, Jour. Bombay Nat. Hist. Soc., vol. 20, p. 1037. Nrxkoisky, 1916, Fauna Russie, p. 111, figs. 22-23 (Persia oriental [Terra Zirkuch: Atkaul; Dunin, Transcaspia]). SHKarr, 1916, Izv. Kavkazsk. otdela RGO, vol. 24, p. 181 (not seen). TzArEWwskt, 1917, Ann. Mus. Zool. Leningrad, vol. 22, p. 88 (not seen). Bosrinsku, 1923, Opredel. zmei Turkestan. kraya. Izd-vo SAGU, Tashkent, p. 10 (not seen). WALL, 1923, Jour. Bombay Nat. Hist. Soc., vol. 29, p. 619 (Transcaspia; Afghan- istan; Baluchistan [Man, Gusht, Kacha, Sib, Kanki, Quetta]). WERNER, 1929, Zool. Jahrb., vol. 57, p. 62. CHeErnov, 1934, Presm. Turkmenii, Trudy SOPS, Seriya Turkmen- skaya, vol. 6, p. 274 (not seen). WERNER, 1936, Festschrift Strand, vol. 2, p. 201. TERENT- JEV AND CHERNOV, 1936, Kratkii opredel. zemnovod. presm. kaiushchikhsa SSSR, p. 59 (not seen). Lapreyv, 1937, Dikie zhivot. Kopet-Daga . . ., Ashkabad-Baku, p. 36 (not seen). ANDRUSHKO, et alii, 1939, Voprosi ekol. biochem., vol. 4, pp. 211, 234 (not seen). SHIBANOV, 1939, Presm. v. kn. Zhizn. zhirot . . ., p. 739 (not seen). TERENTJEV AND CHERNOV, 1940, Kratkii opredel. zemovod. presm. kaiushchikhsa SSSR, p. 142 (not seen). SmitH, 1943, Fauna British India, Rept. and Amph., vol. 3, p. 190 (Baluchistan [Man, Gusht, Kacha, Sib, Kanki, Quetta], Afghanistan and southern Turkestan to Transcaspia). CHERNOV, 1948, Presm. Zhivotnii SSSR, vol. 2, p. 154 (not seen). TERENTJEV AND CHER- Nov, 1949, Diag. Rept. Amphib., pp. 246-247, 320, map 31 (distribution compiled). WETTSTEIN, 1951, Sitzb. Oster. Akad. Wiss. Wien, vol. 160, p. 444 (Iran [Khanu, 100 km. siidl. Sabzawaran, Prov. Kerman]). Rustamov, 1954, Ptichi pustini Kara-Kum, Izd-vo AN TSSR, Ashkabad, p. 324 (not seen). KarTASHEV, 1955, Uch. zap. MGU, Biol. CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. bo ~r iS) Ficure 24. Dorsal view of head of Lytorhynchus ridgewayi (CAS AMNH 126355). Izd-vo MGU, vol. 171, p. 196 (not seen). Lrviron, 1959, Proc. Calif. Acad. Sci., 4th ser., vol. 29, pp. 456-457 (Afghanistan [Chah-i-Angir]). Bocpanov, 1962, Herp. Turkmen, p. 188, figs. 35-36. ANDERSON, 1963, Proc. Calif. Acad. Sci., 4th ser., vol. 31, p. 478. ANDRUSHKO AND MIKKAy, 1964, Vestnik Leningradskogo Universiteta, no. 9, biol. ser., pp. 5-19 (ecology). Minton, 1966, Bull. American Mus. Nat. Hist., vol. 134, p. 131, pl. 28, fig. 1 (Pakistan [vicinity of Kolpur; near Quetta]; description, natural history). Litorhynchus ridgewayi, NrKotsKy, 1899, Ann. Mus. Zool. Acad. Imp. Sci. St. Pétersbourg, vol. 4, pp. 403-404 (Iran [Atkaul, in Terra Zirkuch]). Lytorhynchus ridgewayi var. rosent ELPATJEVSKI AND SABANEJEV, 1906, Zool. Jahrb., vol. 24, p. 257, pl. 19, figs. 6-7 (type loc: Nachduin, Transcaspia; type whereabouts unknown). Lytorhynchus gabrielis WERNER, 1938, Zool. Anz., vol. 121, p. 268, figs. 2-3 (type loc: Unter Ziarat, Baluchistan; type in Vienna). MATERIAL EXAMINED (12). AFGHANISTAN: Chah-i-Angir (CAS 84639) ; 35 km. S Darweshan (CAS 120495); without data (AMNH 74577). IRAN: Ahram, Fars Prov. (FMNH 141602). PAKISTAN: 5 mi. NW Kolpur (AMNH 86897); near Kolpur (AMNH 96223-96224); 3 mi. NE Kolpur (UMMZ 126355); Nushki (CAS 101407-101409, 101476); Quetta (USNM 52140). RANGE. (See fig. 7.) Reported from southern and eastern Iran, lowland and intermediate elevations of western and southern Afghanistan and adjacent areas of Pakistan, and the Transcaspian region of the Soviet Union. The following specimens (not examined by us) are in the collections of the Bombay Natural History Society. IRAN: Pusht, 42 mi. N of Dizak (BNHS S184); Kanki, 20 mi. SW of Sib (BNHS S185); Sib (BNHS $186). PAKISTAN: Baluchistan: Kacha (BNHS S183); Mand (BNHS S187). All these localities are near the Iran-Pakistan border. Vor. XXXVII] LEVITON & ANDERSON: REVIEW OF LYTORHYNCHUS 273 Ficure 25. Lateral view of head of Lytorhynchus ridgewayi (AMNH 126355). Dracnosis. Rostral narrowly truncate anteriorly; prefrontal single or divided; upper labials usually separated from eye by suboculars; prefrontal single or divided; light buff or grayish above with series of brown, black-edged squarish or transverse spots on dorsum; anchor-shaped marking on head from prefrontal(s) to neck; ventrals 165-188,'' subcaudals 41—54,!2 dorsal body blotches 36-47;" standard length (of largest male and female examined, in Imm) c O15, 2 347; tail length: ¢ 65, 2 65. Remarks. Dr. Minton (personal correspondence) has provided us with the following information on the specimen of L. ridgewayi he collected in Pakistan: “All my ridgewayi come from the valley between Quetta and the top of Bolan Pass near Kolpur. Elevations are 5000-6000 feet. If the type locality of gabrielis is correct, it must range to about 8000 feet, however. The only ones I have personally collected were on the highway. Surrounding soils varied from almost pure clay (perhaps an old Jake bed) to clay liberally admixed with sand and gravel. Definitely not a dune snake in my experience, however. In my experience, the three forms are allopatric, but I have no reason not to consider all as full species.” Minton (1966, p. 131) apparently assumes Werner’s type locality for L. gabrielis to refer to Ziarat near Quetta, West Pakistan. There are a number of localities of this name in Iranian Baluchistan as well, and Werner gives no indication of the elevation or precise locality for this site. An elevation of 8000 feet seems to us unlikely, particularly since Werner also lists such species as Agama nupta and Pseudocerastes persicus from this locality. Most of Werner’s identifiable localities are in eastern Iran. 4 Ventrals: ¢ [2] 167.0 (165-169), 9 [7] 179.0 + 1.5 SE (174-188). 12 Subcaudals: <& [2] 47.5 (47-48), @ [5] 48.4 (45-53). 18 Body blotches: [8] 42.1 + 1.3 SE (36-47). CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser. bo ~r - SUMMARY The Asian snakes of the genus Lytorhynchus are reviewed. The genus is clearly distinct from the North American leaf-nosed, Phyllorhynchus, with which it has been confused in the past. Six nominal species of Lytorhynchus are recognized. These are referred to two species groups, the ‘“‘diadema” group, including L. diadema, L. gaddi, and L. kennedyt, and the “ridgeway” group of L. maynardi, L. paradoxus, and L. ridgewayi. The latter group is distributed throughout southwestern Asia, from Pakistan and northwestern India to Iran, the former from Iraq and southwestern Iran, through the Arabian Peninsula, to Egypt and across North Africa to Morocco. Synonymies appropriate to each of the nominal species recognized are presented and available names allocated. Little is known about the habits of these snakes beyond the obvious that they are found in arid and semiarid environments, on either sandy or rocky soil (although they have been taken on the Batna Plateau, a grassy plain at 3350 feet in Algeria), are nocturnal, probably lay eggs, and probably feed mostly on lizards. LITERATURE CITED ANDERSON, JOHN 1898. Reptilia and Batrachia. In: Zoology of Egypt, vol. 1. London (Bernard Quaritch), lviii + 371 pp., 50 pls. ANDRUSHKO, A. M., anp N. E. MIKKay 1964. [Distribution and way of life of the Afghanistan lytorhynchid (Lytorhynchus ridgewayi Boulenger, 1887) with an ecological geographic survey of the genus Lytorhynchus Peters, 1862.) Vestnik Leningradskogo Universiteta, no. 9, Biol. Ser., Issue 2, pp. 5-19. FLOWER, STANLEY 1933. Notes on the reptiles and amphibians of Egypt, with a list of species known to occur in that kingdom. Proc. Zool. Soc. London, 1933, pp. 735-851, map. GUNTHER, ALBERT 1875. Second report on collections of Indian reptiles obtained by the British Museum. Proc. Zool. Soc. London, 1875, pp. 224-234, 4 pls. Haas, GEORG 1957. Some amphibians and reptiles from Arabia. Proc. California Acad. Sci., 4th ser., vol. 29, pp. 47-86. KraM_Er, E., AnD H. SCHNURRENBERGER 1963. Systematik, Verbreitung und Okologie der Libyschen Schlangen. Rev. Suisse Zool., vol. 70, pp. 453-568, pls. 1-4. Minton, SHERMAN A., Jr. 1966. A contribution to the herpetology of West Pakistan. Bull. American Mus. Nat. Hist., vol. 134, pp. 27-184, pls. 9-36. Murray, JAMEs A. 1884. Additions to the reptilian fauna of Sind. Ann. Mag. Nat. Hist., ser. 5, vol. 14, pp. 106-111. UnpDERWOop, GARTH 1967. A contribution to the classification of snakes. British Museum (Natural History), Publ. no. 653, x + 179 pp. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XX XVII, No. 8, pp. 275-332; 31 figs.; 11 tables March 6, 1970 SCORPION FAUNA OF BAJA CALIFORNIA, MEXICO: ELEVEN NEW SPECIES OF VEJOVIS (SCORPIONIDA: VEJOVIDAE) ny Stanley C. Williams Research Associate in Entomology California Academy of Sciences San Francisco, California 94118 ABSTRACT: Eleven new species of scorpions of the genus Vejovis were collected on the Baja California Peninsula and are here described. None of these species are known to occur outside the Baja California Peninsula but several have close relatives in California, Arizona and mainland Mexico. INTRODUCTION The purpose of this paper is to describe and name 11 new species of scorpions belonging to the genus Vejovis, all from the Baja California Penin- sula of western Mexico. The descriptions of these species are based on several thousand specimens collected during the summer of 1968. A number of other undescribed species of Vejovis were also collected on this trip but will not be discussed until further study can be made. In June of 1968, the collecting trip departed for the Baja California Penin- sula for two months of intensive field study. Two separate parties made up the cooperative expedition. One party was composed of Mont A. Cazier of Arizona State University assisted by three graduate students, Joe Bigelow, Jerry Davidson, and Norman Leppla. The other party was composed of my- self and two graduate students, Michael Bentzien and William Fox. The Cazier party was primarily concerned with study of the insect fauna, especially flies of the Family Apioceridae, while my party was primarily concerned with the [275] ee — = i Se ee =’ BAA. ape : a -eeee ; : Marine Did;OsICa! Laporator h : \ - MAH | 619/70 | 276 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. study of arachnids, especially scorpions. The two parties worked together very closely and cooperated to insure greatest success for all interests. During the months of June and July the entire length of the Baja California Peninsula was traversed overland and collections were made in as many different habitat types as was possible in the limited time. Special efforts were made to visit type localities to secure series of topotypes. Most of the scorpion specimens were collected by means of the ultraviolet detection method (Williams, 1968a; Hadley and Williams, 1968). Rock rolling and turning other surface cover were surprisingly unproductive except for a few species. This was perhaps due to the hot dry conditions existing during the summer. The scorpions were killed and fixed each morning, using the process and fixative described by Williams (1968b). Measurements given in this paper are the standard ones used in scorpion taxonomy, with a few useful modifications as described by Williams (1968c). ACKNOWLEDGMENTS Sincere thanks and appreciation to Mont A. Cazier for his help and sug- gestions in the field. Thanks also to Michael Bentzien and Joe Bigelow for their many efforts and long hours of work throughout the entire expedition, and to William Fox, Jerry Davidson, and Norman Leppla for their assistance during the earlier part of the summer. Thanks are due to Charlene F. Williams for clerical assistance and to Vincent F. Lee for his technical assistance in the laboratory. Thanks and appreciation are also due Richard and Mary Lou Adcock, of the La Paz Skin Diving Service, for providing transportation to Isla Partida and Isla Espiritu Santo on their boat Marisla, and for their in- valuable aid in pointing out habitats of special interest. This study was supported by the Systematics Section of the National Science Foundation through research grant number GB-7679. THE BajA CALIFORNIA PENINSULA Baja California, Mexico, is an 800-mile long peninsula surrounded by the Pacific Ocean on the west and separated from the Mexican mainland by the Gulf of California. The whole peninsula of Baja California is a primitive re- gion. There are few roads, except in border areas and in the extreme south. The peninsula is divided into two political units, the state of Baja California in the north and the Federal Territory referred to as “Baja California Sur” in the south. Baja California varies from tropical habitats in the south to harsh Colorado desert in the northeast, and chaparral forest in the northwest. Many other diverse habitats include isolated canyons, high mountains (covered with coniferous forest), volcanic ranges, sedimentary basins, foggy deserts, and shifting sand dunes. VoL. XXXVIT] WILLIAMS: NEW SPECIES OF VEJOVIS 277 NEw SPECIES Vejovis grandis Williams, new species. (Figures 1, 2.) Diacnosis. One of the largest species of Vejovis known both in body length and body mass. Denticles on the inferior border of the cheliceral finger reveal that this species is a member of the subgenus Paruroctonus. Readily identified by the numerous stout reddish bristles on the inferior intercarinal spaces of the metasoma and more or less dark markings on the carapace and mesosomal dorsum. Pectine teeth of females varies from 22 to 26, those of males 30 to 35. Very closely related to Vejovis vachoni (Stahnke) and Vejovis mesaensis (Stahnke). Differs from V. mesaensis by the presence of dark dorsal markings, presence of red pedipalp fingers, and slightly more swollen pedipalp palms. Differs from V. vachoni in the following ways: telson more elongate and less swollen, vesicle with hairs longer and less bristle like in males; pedipalp chela without the deep, conspicuous, double dentate proximal scallop. DESCRIPTION OF HOLOTYPE (male). Coloration: Base color of carapace, mesosoma, metasoma, pedipalps, and walking legs uniform bright yellow; cara- pace with underlying, intricate, dark color pattern, dark pigment most con- centrated lateral to ocular tubercule; interocular triangle without dark mark- ings; mesosomal tergites also with intricate dark markings, posterior tergites with dark markings more faded and not as extensive; pedipalp fingers reddish brown; eyes black; cheliceral fingers reddish brown; aculeus dark reddish brown; pectines white; tips of pretarsal claws light brown. Carapace: Anterior margin essentially straight, but with a very subtle sug- gestion of a median emargination, this margin with three pairs of erect reddish bristles. Lateral eyes three per group, anterior two eyes about equal in size, posterior eye smallest of three. Median diad distinctly greater than 4 carapace width at that point; one pair of short red bristles on ocular tubercule above and between eyes. Carapace surface densely covered by large granules. Mesosoma: Tergites not densely granular, granules present but mostly re- stricted to posterior % of each tergite; tergite 7 with two pairs of irregularly dentate lateral keels and a single broad, low median keel, this set with a few isolated granules. Sterna agranular except on last sternite; last sternite with one pair of dentate keels, median intercarinal space densely granular, lateral areas with a few distinct granules. Stigma short slits. All sternites with an abundance of conspicuous reddish bristles. Metasoma: All dorsal and dorsolateral keels serrate to deeply crenulate except that dorsal keels of segment V absent, dorsolateral keels I to IV each begin anteriorly as a distinctly broadened wing; lateral keels dentate on pos- terior % of segment I, crenulate on posterior % of II and % of III, absent on 278 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 47H Serr. IV, crenulate on anterior half of V; inferior lateral keels irregularly serrate to deeply crenulate; inferior median keels irregularly crenulate on segments I to III, irregularly crenulate to serrate on IV, irregularly serrate on V. Inferior intercarinal space of V basically smooth but with several distinct scattered granules. Space between inferior lateral keels on segments I to IV densely covered with stout reddish bristles, these very conspicuous. Telson: Ventral aspect of vesicle smooth and agranular; this surface with about 10 pairs of conspicuous stout reddish hairs. Very subtle, broad subaculear tubercule. Vesicle slightly wider than widest dimension of metasomal segment V. Pectines: Very elongate and curved; 31/33 pectinal teeth; triangular fulcra; ventral surface conspicuously hirsute, hairs reddish and most abundant on fulcra, middle lamellae, and anterior margin. Genital operculum: Completely divided longitudinally; large genital pa- pillae visible externally; surface of operculum conspicuously hirsute with about 10 pair of stout reddish bristles. Chelicerae: Inferior border of movable finger with three small unpigmented denticles; terminal tooth of superior border subequal to terminal tooth of in- ferior border, but terminal tooth reduced in size. Inferior border of fixed finger with two small unpigmented denticles. Pedipalps: Hand long and swollen, all keels distinct and granular; inter- carinal spaces smooth and agranular; movable finger approximates carapace in length but fixed finger distinctly shorter than carapace; grasping edges of both fingers distinctly scalloped; teeth do not quite extend to proximal bases of each finger; teeth do not touch when chela closed over much of proximal region but no large open spaces occur. Standard measurements and photographs: Table 1 and figures 1 and 2. DESCRIPTION OF ALLOTYPE (female). Essentially the same as holotype in color and morphology with the following exceptions: Distinctly greater in total body length and most other body dimensions; dark color pattern on carapace and mesosoma similar in pattern, but much lighter; aculeus proportionally longer; pectines slightly shorter, with about % fewer teeth (22/22 instead of 31/33), pectinal teeth each much shorter; no genital papillae; genital oper- culum not completely divided longitudinally; teeth on inferior border of moy- able cheliceral finger larger and more distinct; inferior border of fixed cheliceral finger with three small denticles; inferior keels of metasoma slightly less promi- nent and not quite as distinctly crenulate or serrate; chela not distinctly scal- loped. Standard measurements: Table 1. VARIATION WITHIN Paratypes. Study of 273 paratopotypes (122 males, 151 females) indicated little variation from the structure of the holotype and VoL. XXXVIT] WILLIAMS: NEW SPECIES OF VEJOVIS 279 Ty FicurE 1. Vejovis grandis, new species. Dorsal view of holotype. allotype. Males varied in total length from 28 to 77 millimeters while females varied from 21 to 77 millimeters. Pectinal tooth count varied from 22 to 26 (predominantly 24) in females and from 30 to 35 (predominantly 32) in males. Base cuticle color similar in most specimens, but a large degree of variability existed in the dark underlying markings. Some individuals had very dark Ficure 2. Vejovis grandis, new species. Ventral view of holotype. 280 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Taste 1. Measurements (in millimeters) of Vejovis grandis, new species, holotype and alloty pe. Holotype Allotype (male) (female) Total length 64.9 76.0 Carapace, length Us 9.5 width (at median eyes) 6.2 tell Mesosoma, length 16.3 20.1 Metasoma, length 33.5 37.1 segment I (length/width) 4.5/3.2 5.0/3.9 segment II (length/width) 5.8/2.8 6.3/3.3 segment III (length/width) 6.1/2.7 6.7/3.2 segment IV (length/width) 7.6/2.4 8.3/2.9 segment V (length/width) 9.5/2.4 10.8/2.9 Telson, length 7.8 9.3 Vesicle (length/width) 4.9/2.5 Silom depth 22 Ae) Aculeus, length 2.9 3.6 Pedipalp Humerus (length/width) UPD 8.3/2.8 Brachium (length/width) 6.8/2.7 7.8/3.4 Chela (length/width) 13.0/3.4 16.2/4.2 depth 4.5 5.4 movable finger, length 7.4 9.3 fixed finger, length 5.6 U8 Pectines teeth (left/right) 33/31 22/22 markings on the carapace and mesosoma dorsum while other individuals had faint markings, most individuals being intermediate. Smaller individuals with dark markings more distinct, larger individuals showing tendency toward fading of dark markings. Interocular triangle always lacking dark markings. On in- dividuals with dark pigment obsolete on the carapace a dusky ocular crescent remains. TYPE DATA AND ETYMOLOGY. The holotype, allotype, and 273 paratopo- types were collected at Oakies Landing, 27 miles south of Puertecitos, Baja California Norte, Mexico, 12 June 1968 by S. C. Williams, M. A. Cazier, and party. The holotype and allotype are permanently deposited in the California Academy of Sciences. This species is named “grandis” because of its very long and heavy body. MarterIAL. In addition to the holotype, allotype, and 273 paratopotypes, 1622 additional paratypes were studied. These were from the following 12 locations in Baja California Norte, Mexico: Puertecitos and for 2 miles south, Vou. XXXVIT] WILLIAMS: NEW SPECIES OF VEJOVIS 281 10 June 1968 (S. C. Williams, M. A. Cazier, and party), 14 males, 14 females; Puertecitos, elevation 50 feet, 11 June 1968 (S. C. Williams, M. A. Cazier, and party), 5 males, 1 female; Puertecitos and for 4% mile north, 11 June 1968 (S. C. Williams, M. A. Cazier, and party), 6 males, 10 females; 8 miles north of Gonzaga Bay, 13 June 1968 (S. C. Williams), 28 males, 45 females; 6 miles north of Gonzaga Bay, 13 June 1968 (S. C. Williams), 71 males, 75 females; 10 miles north of Gonzaga Bay, 13 June 1968 (S. C. Williams), 55 males, 83 females; 1 mile north of Gonzaga Bay, elevation 100 feet, 14 June 1968 (S. C. Williams), 48 males, 49 females; 5 miles north of Gonzaga Bay, elevation 150 feet, 14 June 1968 (S. C. Williams), 27 males, 26 females; 6 miles north of Gonzaga Bay, elevation 100 feet, 14 June 1968 (S. C. Williams), 35 males, 49 females; 7 miles north of Gonzaga Bay, 14 June 1968 (S. C. Williams), 81 males, 90 females; Calamajue Arroya, elevation 1000 feet, 15 June 1968 (S. C. Williams, M. A. Cazier, and party), 64 males, 40 females; Calamajue Arroya, elevation 1000 feet, 16 June 1968 (S. C. Williams, M. A. Cazier, and party), 65 males, 59 females; Bahia de Los Angeles, elevation 25 feet, 17 June 1968 (S. C. Williams, M. A. Cazier, and party), 39 males, 30 females; Bahia de Los Angeles, elevation 25 feet, 18 June 1968 (S. C. Williams, M. A. Cazier, and party), 16 males, 15 females; Bahia de Los Angeles, elevation 25 feet, 19 June 1968 (S. C. Williams, M. A. Cazier, and party), 219 males, 263 females. CoMMENT. ‘This species was found along the eastern side of the peninsula between Puertecitos and Bahia de Los Angeles where it appeared to prefer rocky volcanic habitats where considerable soil had accumulated (fig. 3). This is apparently a burrowing species, and was not found under rocks during the day. All specimens were collected by ultraviolet detection. This species was the numerically predominant species in most of the samples taken in its habi- tats. The samples contained a great variety of different age groups with the exception that the youngest instars were not frequently encountered. This species forms local color races throughout its distribution, being dark on dark soils and light on light soils. On light soils, the only dark pigmentation was inconspicuous, faint dusky markings underlying the translucent cuticle. The adults of even the lightest races retained the reddish pedipalp fingers, however. Its nocturnal activity patterns were very similar to those of Vejovis mesaensis and Vejovis vachoni. Vejovis harbisoni Williams, new species. (Figure 4, 5.) Dracnosis. Large species of Vejovis in the “wupatkiensis” group. Adults with body color dark brown, sometimes becoming darker at the distal end of the metasoma; juveniles with more dirty yellow coloration, this with underlying dusky pigmentation. Metasoma with inferior lateral keels crenulate; inferior median keels smooth to obsolescent on segments I to III, better developed and 282 CALIFORNIA ACADEMY OF SCIENCES [Proc. 471 SER. Ficure 3. Type locality and habitat of Vejovis grandis, on plateau above Oakies Landing, Baja California Norte, Mexico, 13 June 1968. smooth to crenulate on segments IV and V. Telson with swollen vesicle and with subaculear tubercule. Chelicerae with denticles completely lacking on in- ferior border of movable finger. Pedipalps with very elongate fingers, fixed and movable finger each longer than carapace, each finger terminates in long conspicuous tooth, these overlap when chela closed. Pectines with 21 to 23 teeth in females and 25 to 27 teeth in males. Related to Vejovis wupatkiensis Stahnke from which it can be distinguished by larger adult body size, darker adult coloration, and greater number of pec- tinal teeth (V. wupatkiensis has male counts of 17 and female counts of 15 and 16). DESCRIPTION OF HOLOTYPE (male). Coloration: base color of carapace, mesosoma, metasoma, and pedipalps uniform dark yellowish brown; last meta- somal segment and telson darker; walking legs slightly lighter; pectines white; eyes black; aculeus black; some dusky markings on legs. Carapace: Anterior margin faintly crenulate; with median emargination; set with three pairs of erect brown bristles. Lateral eyes three per group, an- terior eye largest in each group. Median eyes on low ocular tubercule; one pair of reddish bristles on tubercule behind diad, no similar pair on tubercules anterior to diad; diad less than % carapace width at that point. Carapace slightly lustrous; area of interocular triangle not conspicuously granular, re- mainder of carapace densely granular. VoL. XXXVITI] WILLIAMS: NEW SPECIES OF VEJOVIS 283 Mesosoma: Tergites with abundant granules; tergite 7 with two pairs of serrate to dentate lateral keels; one median keel set with irregular low rounded granules. Sternites smooth, lustrous and agranular; segment 7 with one pair of keels, these smooth to slightly crenulate; stigma long oval. Metasoma: All dorsal and dorsolateral keels serrate to deeply crenulate except that dorsal keels of segment V absent; lateral keels serrate on posterior % of posterior I, anterior % of II, posterior % of III, absent on IV, crenulate on anterior % of V; all inferior lateral keels well developed and deeply crenulate; inferior median keels smooth to obsolete on segments I to III, smooth to crenu- late, but approaching obsolescence on IV, smooth to crenulate on V. Inferior intercarinal space of V agranular. Telson: Vesicle broad and bulbous; ventral surface densely covered with low rounded granules obsolescent in appearance; broad subaculear tubercule; one pair of long brown hairs lateral to subaculear tubercule, a number of much shorter, less conspicuous hairs also present; dorsolateral aspect of vesicle granu- lar. Aculeus short and well curved. Pectines: Long and thick; 25/26 pectinal teeth; fulcra triangular; each middle lamella with one long reddish hair. Genital operculum: Completely divided longitudinally; distinct genital papillae visible externally. Chelicerae: Inferior border of movable finger completely lacking denticles; terminal tooth of superior border and terminal tooth of inferior border sub- equal with superior tooth smallest. Long brown bristle on superior base of fixed finger, this almost as long as fixed finger. Pedipalps: Hand elongate with only slight swelling of palm; keels broad, rounded and agranular on the dorsolateral aspect of hand, keels granular me- dially; intercarinal spaces agranular; fingers elongate; fixed finger and mov- able finger each distinctly longer than carapace. Grasping edges of fingers not scalloped; very elongate, conspicuous terminal tooth on each finger; teeth ex- tend to proximal bases of fingers; teeth do not meet proximally when chela closed, but leave narrow elongate space. Standard measurements and photographs: Table 2 and figures 4 and 5. DESCRIPTION OF ALLOTYPE (female). Coloration and patterns essentially the same as holotype. Structure essentially the same as holotype except: total body length longer; most body dimensions slightly greater; pectines with slightly fewer teeth (23/23 instead of 26/25); without genital papillae; genital oper- culum not completely divided longitudinally; vesicle hairs much longer, heavier and more conspicuous; pectinal teeth very short, proximal tooth in each comb longer than adjacent teeth. Standard measurements: ‘able 2. VARIATION WITHIN PARATYPES. Study of the 10 paratopotypes (6 males, 4 females) indicated little significant variation from the description of the holo- 284 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Ficure 4. Vejovis harbisoni, new species. Dorsal view of holotype. type and allotype. Males varied in total length from 35 to 51 millimeters, while females varied from 26 to 50 millimeters. Pectine tooth counts varied from 22 to 23 (predominantly 23) in females and from 25 to 27 (predominantly 25 and 26) in males. Most specimens were mature, but about one-fifth were juveniles. Ficure 5. Vejovis harbisoni, new species. Ventral view of holotype. VoL. XXXVIT] WILLIAMS: NEW SPECIES OF VEJOVIS 285 TABLE 2. Measurements (in millimeters) of Vejovis harbisoni, new species, holotype and alloty pe. Holotype Allotype x (male) (female) Total length 43.0 50.5 Carapace, length 5.4 6.0 width (at median eyes) 4.0 4.6 Mesosoma, length 12.4 17.3 Metasoma, length 20.0 Di Pe segment I (length/width) 30/3 BIS segment II (length/width) 3.5/3.1 3.6/3.0 segment III (length/width) 3.5/3.0 Bal Pell segment IV (length/width) 4.5/2.8 4.7/3.1 segment V (length/width) 5.5/2.9 6.0/3.1 Telson, length Sif) 6.0 Vesicle (length/width) B53 Dail 3.7/2.8 depth 2.0 2.2 Aculeus, length ile 23 Pedipalp Humerus (length/width) 6.2/1.3 6.6/1.5 Brachium (length/width) 6.5/1.5 6.9/1.7 Chela (length/width) 10.2/2.0 10.8/2.2 depth 2.3 Beil movable finger, length 6.8 6.3 fixed finger, length 5.9 7.2 Pectines teeth (left/right) 26/25 23/23 Mature males and mature females attained about the same size in total body length but males tended to have slightly more elongate metasomas where as females had carapace and mesosoma both longer and wider. Adult females had vesicles differing from adult males in that they were less swollen, not as con- spicuously tubercular, more hirsute in appearance, and with a slightly more prominent subaculear tubercule. Juveniles appeared morphologically similar to adults with the following exceptions: body color light dirty yellow not dark brown; cuticle with underlying dusky pigment over much of body; smaller in body size; telson lacking sexual dimorphism, not greatly swollen, relatively hirsute in both sexes. Juvenile and adult males with genital papillae visible externally. Specimens from outside the type locality were essentially the same in color and morphology with the following exceptions: one female specimen from near Bahia San Luis Gonzaga had a pectine tooth count of 21 (all other paratype females had 22 to 23 pectinal teeth); specimens from Bahia de Los Angeles had inferior median keel of metasomal segment V more smooth, 286 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. TYPE DATA AND ETYMOLOGY. The holotype, allotype, and 10 paratopotypes (6 males, 4 females) were collected in Calamajue Arroyo, Baja California Norte, Mexico, 15 and 16 June 1968 by S. C. Williams and M. A. Cazier. The holo- type and allotype are permanently deposited in the California Academy of Sciences. This species is named ‘‘harbisoni” after Charles F. Harbison, Curator of Entomology, San Diego Museum of Natural History. Mr. Harbison has spent many years making biological explorations of Baja California, collected many of the first scorpion specimens known from Baja California, and over the years has encouraged many biology students to study the arthropod fauna of this region. MatTERIAL. In addition to the holotype, allotype, and 10 paratopotypes, 29 additional paratypes were studied from three other localities in Baja California. These records are as follows: Mexico, Baja California Norte: 6 miles north of Bahia San Luis Gonzaga, 13 June 1968 (S. C. Williams, M. A. Cazier, and party), 1 male, 1 female; 7 miles north of Bahia San Luis Gonzaga, 14 June 1968 (S. C. Williams, M. A. Cazier, and party), 2 females; Bahia de Los Angeles, elevation 25 feet, 18 June 1968 (S. C. Williams, M. A. Cazier, and party), 2 females; Bahia de Los Angeles, elevation 25 feet, 19 June 1968 (S. C. Williams, M. A. Cazier, and party), 1 male, 4 females. CoMMENT. This species lives in the rocky volcanic habitats occurring in the north eastern regions of the Baja California Peninsula (fig. 6). Here it is found in rock slides, and along rocky slopes where it occurs in the spaces created by accumulated fragmented rock. In the evenings, individuals of this species may come to the surface rocks. During the day, however, this species appar- ently resides deep enough in the rock deposits that collection by rock rolling is unproductive. Even in the most favorable habitats known, this species has never been found in abundance. Males and females were found in approxi- mately equivalent numbers and juveniles were encountered about as frequently as adults. All specimens were collected using the ultraviolet detection method. Vejovis minutis Williams, new species. (Figures 7, 8.) Diacnosis. One of the smallest species of Vejovis known; uniform brown- ish in color, lacking stripes or other conspicuous markings; chelicerae lacking denticles on inferior margin of movable finger; pedipalps with very elongate fingers, each terminating in a long tooth; pedipalp palms not greatly swollen; pectinal teeth 11 to 13 in females, 12 to 15 in males; inferior lateral keels of metasoma crenulate; inferior median keels of metasoma obsolete on segments I to III. Vejovis minutis belongs to the “wupatkiensis” group of Vejovis. It is the smallest known species in this group and is related to Vejovis gertschi Williams. Vor. XXXVIT] WILLIAMS: NEW SPECIES OF VEJOVIS 287 Ficure 6. Type locality and habitat of Vejovis harbisoni, in Calamajue Arroyo, Baja California Norte, Mexico, 15 June 1968. It is easily distinguished from V. gertschi by its smaller pectines, reduced pec- tine tooth count, and less distinctly developed inferior metasomal keels. DESCRIPTION OF HOLOTYPE (male). Coloration: Base color of entire body uniform light yellow; walking legs and pectines slightly lighter; carapace and Ficure 7. Vejovis minutis, new species. Dorsal view of holotype. 288 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 8. Vejovis minutis, new species. Ventral view of holotype. dorsum of mesosoma with underlying dusky markings; eyes black; aculeus red- dish brown; some dusky markings on regions of pedipalps and legs. Carapace: Anterior margin with broad median emargination and three pairs of fine brown hairs. Three lateral eyes in each group; anterior two eyes largest in each group; median diad less than % carapace width at that point; a pair of long brown hairs on ocular tubercule posterior to diad. Carapace surface smooth to finely granulate. Mesosoma: Tergites smooth to finely granulate; last tergite with two pairs of granulate lateral keels and a smooth median keel. Sternites smooth and agranulate; sternite 7 with one pair of keels, these smooth to finely granulate. Stigma small and oval. Metasoma: Dorsal and dorsolateral keels crenulate on all segments except that dorsals absent on segment V; lateral keels crenulate on posterior % of seg- ments I to II, posterior ’ of III, absent on IV, obsolete to slightly crenulate on V; inferior lateral keels crenulate; inferior median keels obsolete on segments I to III, reduced and smooth to slightly crenulate on IV, crenulate on V. In- ferior intercarinal space of segment V agranulate. Telson: Basically smooth and agranulate; ventral surface of vesicle with about 13 to 15 pairs of long brown hairs, many approaching aculeus in length; very subtle subaculear tubercule. Pectines: Small with relatively long teeth; 13 pectinal teeth; inferior sur- face with little hirsuteness. Genital operculum: Completely divided longitudinally; genital papillae visible externally but not conspicuous. Chelicerae: Inferior border of movable finger completely lacking denticles. VoL. XXXVIT] WILLIAMS: NEW SPECIES OF VEJOVIS 289 Taste 3. Measurements (in millimeters) of Vejovis minutis, new species, holotype and allotype. Holotype Allotype (male) (female) Total length 14.7 17.8 Carapace, length 2.3 Dail width (at median eyes) 17) 2.0 Mesosoma, length 3:3 5)! Metasoma, length al (hos segment I (length/width) 1.0/1.4 1.0/1.5 segment II (length/width) 1.1/1.4 WA segment III (length/width) eZ ales 1.2/1.4 segment IV (length/width) 1.6/1.3 1.7/1.4 segment V (length/width) BY NS 2.4/1.4 Telson, length 2.0 2.3 Vesicle (length/width) 1.3/0.9 1.4/1.1 depth 0.7 0.7 Aculeus, length 0.7 0.9 Pedipalp Humerus (length/width) 1.9/0.7 2.2/0.7 Brachium (length/width) 2.0/0.7 2.4/0.9 Chela (length/width) 32/07) 3.8/0.9 depth 0.7 0.9 movable finger, length 2.0 25 fixed finger, length e7 2.0 Pectines teeth (left/right) 13/13 12/12 Pedipalps: Chela distinctly elongate; palms only slightly swollen; keels on palm smooth to obsolete except that those on medial aspect show some granu- lation; fingers elongate but movable and fixed finger each shorter than carapace; grasping edges of chela not scalloped, each finger terminates distally in a con- spicuously long, overhanging, terminal tooth. Standard measurements and photographs: Table 3 and figures 7 and 8. DESCRIPTION OF ALLOTYPE (female). Coloration and structure essentially the same as holotype except: overall length greater; most body dimensions slightly greater; pectines with slightly fewer teeth (12 instead of 13); without genital papillae; genital operculum not completely divided longitudinally; vesicle with some low rounded granules. Standard measurements: ‘Table 3. VARIATION WITHIN PARATYPES. Study of 36 paratopotypes (21 males, 15 females) indicated little variation from the color and structure of the holotype and allotype. Males varied in total length from 13 to 17 millimeters, while 290 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH Serr. females varied from 15 to 20 millimeters. Pectine tooth counts varied from 11 to 13 (predominantly 12) and from 12 to 15 (predominantly 13) in males. Little sexual dimorphism occurs. Males and females have overlapping pectinal tooth counts, and are very similar in size, color, and morphology. Males may be distinguished by the following characters: genital papillae present (some- times hard to see); genital operculum completely divided longitudinally; slightly slimmer mesosoma; pectinal teeth longer in length; hairs on vesicle not as stout. TYPE DATA AND ETYMOLOGY. The holotype, allotype, and 36 paratopotypes were collected 5 miles southwest of La Paz, Baja California Sur, Mexico, 3 August 1968 by S. C. Williams, M. A. Cazier, and party. The holotype and allotype are permanently deposited in the California Academy of Sciences. This species is named “‘minutis” because of its very tiny size. MatTeRIAL. In addition to the holotype, allotype, and 36 paratopotypes, 19 additional paratypes were studied. These paratypes came from the following three localities in Baja California Sur, Mexico: 2.9 miles northwest of San Antonio, elevation 1400 feet, 24 July 1968 (S. C. Williams and M. Bentzien), 2 males, 1 female; Las Cruces, 29 July 1968 (S. C. Williams, M. A. Cazier, and party), 9 females; Las Cruces and for 5 miles southwest, 30 July 1968 (S. C. Williams, M. A. Cazier, and party), 7 females. CoMMENT. This species appears to have a rather specialized habitat re- quirement judging by the few samples which included it. It was most abundant in the type locality near La Paz where it was found in relatively dense vegeta- tion growing on low, flat, poorly drained terrain (fig. 9). The soils were fine textured, moderately well packed and with a moderate covering of litter under the denser vegetation. This species was, even in this habitat, one of the least abundant species in the samples which were dominated by scorpions of the genus Bioculus. Young instars were completely absent in the samples, and all specimens were collected by means of ultraviolet detection. Vejovis vittatus Williams, new species. (Figures 10, 11.) Diacnosis. Medium to small species of Vejovis with yellow base color and two pairs of dorsal mesosomal stripes; inferior median and inferior lateral keels of metasoma with underlying irregular dark pigmentation giving the appearance of two pairs of dotted lines; blotches of dusky pigment over much of the body gives this species a speckled appearance. Pedipalp palms not very swollen; fingers shorter than carapace; frontal margin of carapace with four or five pairs of bristles and one median hair; inferior border of movable cheliceral finger completely lacking denticles; pectinal teeth 18 to 22 in males, 15 to 18 in females. Very closely related to Vejovis punctatus punctatus Karsch from which it differs in the following ways: mature individuals with smaller body size; frontal Voit. XX XVII] WILLIAMS: NEW SPECIES OF VEJOVIS 291 Ficure 9. Type locality and habitat of Vejovrs minutis, 5 miles southwest of La Paz, Baja California Sur, Mexico, 31 July 1968. 292 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 47H SER. margin of carapace with four or five pairs of bristles and one median bristle (compared with three pairs and no median bristle); inferior lateral keels of metasoma crenulate (these smooth on punctatus); lacks the proximal scallop on the pedipalp chela of mature males. DESCRIPTION OF HOLOTYPE (male). Coloration: Base color of carapace, mesosoma, metasoma, pedipalps, and walking legs uniform yellow; telson slightly darker; pectines whitish. Carapace with underlying, conspicuous black color markings; dorsum of mesosoma with black blotches on tergites forming two pairs of distinct longitudinal stripes—these approaching obsolescence; dorsal keels of metasoma with obsolescent underlying dusky pigment; inferior median and inferior lateral keels of metasoma with distinct but obsolescent underlying dark pigmentation; inferior surface of metasomal segment V with a number of irregular but distinct dusky blotches; ventral surface of vesicle with a number of faint dusky blotches, pedipalps and walking legs with a number of dusky areas. Ocular tubercule black; eyes black; chela fingers amber; cheliceral teeth reddish amber; aculeus black. Carapace: Anterior margin slightly crenulate, with subtle median emargi- nation, this margin set with nine reddish bristles (four pairs and one median). Lateral eyes three per group, anterior eye largest. One pair of bristles on ocular tubercule anterior to diad, and a similar pair on ocular tubercule posterior to diad; diad slightly less than 4 carapace width at that point. Carapace surface densely granulate. Mesosoma: Tergites densely granulate, segment 7 with two pairs of dentate lateral keels and one crenulate median keel. Sternites smooth, agranulate and lustrous; one pair of crenulate keels on sternite 7; stigma long oval. Metasoma: All dorsal and dorsolateral keels serrate except that dorsal keels of segment V absent, dorsal keels of segments I to III each terminate posteriorly as an enlarged spine; dorsolateral keels on segments II and III each terminate posteriorly as an enlarged spine; lateral keels serrate on posterior % of segment I, posterior “ of II, posterior 4 of III, absent on IV, irregularly crenulate on anterior “s of V; inferior lateral keels irregularly serrate to crenulate on seg- ments I to IV, serrate on segment V; inferior median keels completely absent on segments I to III, obsolete on IV, appearing as several isolated irregular granules, serrate on V. Entire inferior intercarinal space of V granulate. Telson: Ventral surface of vesicle smooth and lustrous, lacking granulation; with about 12 to 15 pairs of fine, long, brownish hairs; distinct subaculear tubercule; dorsal surface of vesicle with broad anterior pair of keels, each ter- minating laterally as a distinct spine. Pectines: Pectines long and thick; 22 pectinal teeth; pectinal teeth very long; triangular fulcra; abundant reddish hairs, these most numerous on fulcra, middle lamellae and anterior margin. VoL. XX XVII] WILLIAMS: NEW SPECIES OF VEJOVIS 293 Genital operculum: Completely divided longitudinally; genital papillae dis- tinctly visible externally. Chelicerae: Inferior border of movable finger completely lacking denticles; terminal tooth of inferior margin and terminal tooth of superior margin of mov- able finger subequal with superior tooth greatly reduced in size. I.arge reddish bristle on superior base of fixed finger, this approximating fixed finger in length. Pedipalps: Palms moderately swollen; keels obsolete; surface of palm smooth, agranulate, and lustrous. Fixed finger and movable finger each dis- tinctly shorter than carapace length. Grasping edges of chela not scalloped; teeth extend to proximal ends of fingers; long, narrow, proximal open space between fingers when chela closed. Standard measurements and photographs: Table 4 and figures 10 and 11. DESCRIPTION OF ALLOTYPE (female). Coloration and color patterns essen- tially the same as holotype. Structure essentially the same as holotype except: slightly longer total length and with most body dimensions slightly larger; pec- tine teeth fewer (17 instead of 22); with more bristles on anterior margin of carapace (five pairs of laterals and one median); much more pronounced sub- aculear tubercule: hairs on ventral surface of telson much coarser, longer and more conspicuous. Standard measurements: Table 4. VARIATION WITHIN PARATYPES. Study of 304 paratopotypes (222 males, 82 females) indicated little variation from the description of holotype and allo- type. Males varied in total length from 26 to 36 millimeters while females varied from 27 to 41 millimeters. Young instars were completely lacking in the samples. Mature females tended to reach slightly larger size in overall length and most body proportions. Pectine tooth counts varied in the males from 19 to 22 (predominantly 21), while female pectine tooth counts varied from 16 to 18 (predominantly 17). Little sexual dimorphism was apparent with the following exceptions: males with larger pectines, these with longer and more numerous teeth; females with more pointed subaculear tubercule and with area around subaculear tubercule with more irregular surface texture; males with genital papillae visible externally and with genital operculum completely di- vided longitudinally; ventral surface of female vesicle appearing more hirsute because of more stout development of hairs. One female, apparently a second or third instar, was collected 75 miles northwest of La Paz. This specimen had the cuticle somewhat lighter than the adults, but had the same patterns of stripes on the dorsum and on the inferior surface of the metasoma. Throughout the range of distribution, the paratypes were, in general, re- markably similar. In several locations, light color races occurred, notably on Isla Espiritu Santo, Isla Partida, and Las Cruces. These light phases were characterized by lighter yellow base cuticle and dorsal stripes being less distinct. The populations on Isla Espiritu Santo and Isla Partida also differed from 294 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Ficure 10. Vejovis vittatus, new species. Dorsal view of holotype. those of the adjacent peninsula in having inferior lateral keels of metasoma slightly more reduced and less deeply crenulate. In the southern part of the range the pectinal tooth count tended to be slightly less than in the northern part of the range. Several males were found to have pectinal tooth counts as low as 18 and several females were found to have counts as low as 15 (these low counts were not common, however). Ficure 11. Vejovis vittatus, new species. Ventral view of holotype. VoL. XXXVIT] WILLIAMS: NEW SPECIES OF VEJOVIS 295 TaBLeE 4. Measurements (in millimeters) of Vejovis vittatus, new species, holotype and allotype. Holotype Allotype (male) (female) Total length 32.8 35.3 Carapace, length 4.4 4.8 width (at median eyes) gay? 3.6 Mesosoma, length 8.2 10.3 Metasoma, length 15.3 15.5 segment I (length/width) 2.0/2.6 2.0/3.0 segment II (length/width) 2.3/2.6 2.4/2.9 segment III (length/width) 225)/2.0 2.6/2.8 segment IV (length/width) 3.5/2.6 SLO/2ES segment V (length/width) 5.0/2.6 5.0/2.7 Telson, length 4.9 4.7 Vesicle (length/width) 32/220 3.0/2.1 depth 1.5 155 Aculeus, length if le? Pedipalp Humerus (length/width) Syl Soy Al Brachium (length/width) 3.6/1.4 3.8/1.5 Chela (length/width) 5.6/1.7 Sei ples depth 1.9 il7/ movable finger, length 3.3 S25 fixed finger, length 2.5 2,5 Pectines teeth (left/right) 22/22 17/17 TYPE DATA AND ETYMOLOGY. The holotype, allotype, and 304 paratopotypes were collected 5 miles southwest of San Miguel de Comondu, Baja California Sur, 2 and 3 July 1968 by S. C. Williams, M. A. Cazier, and party. The holo- type and allotype are permanently deposited in the California Academy of Sciences. This species is name “‘vittatus’” because of the conspicuous striped appear- ance of the mesosomal dorsum. MatTERIAL. In addition to the holotype, allotype, and 304 paratopotypes, 1082 (636 males, 446 females) additional paratypes were studied from some 36 other locations. These specimens were collected in the following locations in Baja California Sur, Mexico: 75 miles northwest of La Paz, elevation 200 feet, 4 July 1968 (S. C. Williams, M. A. Cazier, and party), 50 males, 18 fe- males; 5 to 6 miles southwest of La Paz, elevation 25 feet, 5 July 1968 (S. C. Williams, M. A. Cazier, and party), 27 males, 11 females; 2 miles east of La Paz, elevation 50 feet, 5 July 1968 (S. C. Williams, M. A. Cazier, and party), 296 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. 4 males; 5 to 10 miles west of La Paz, elevation 25 feet, 6 July 1968 (S. C. Williams, M. A. Cazier, and party), 11 males, 17 females; 5 to 6 miles south- west of La Paz, elevation sea level, 6 July 1968 (S. C. Williams, M. A. Cazier, and party), 35 males, 15 females; 1 mile south of La Paz, elevation 100 feet, 6 July 1968 (S. C. Williams, M. A. Cazier, and party), 7 males, 11 females; Isla Espiritu Santo, elevation sea level, 7 July 1968 (S. C. Williams, M. Bent- zien, and W. Fox), 4 females; Isla Partida, 9 July 1968 (S. C. Williams, M. Bentzien, and W. Fox), 19 males, 6 females; Isla Partida, 10 July 1968 (S. C. Williams, M. Bentzien, and W. Fox), 1 male, 1 female; 1 mile east of Los Aripes, 8 July 1968 (M. A. Cazier, J. Bigelow, and N. Leppla), 24 males, 8 females; 14 miles northeast of La Paz, 10 July 1968 (M. A. Cazier, J. Bigelow, and N. Leppla), 2 males; 14 miles northeast of La Paz, 14 July 1968 (S. C. Williams and party), 3 males, 2 females; 1.5 mile northeast of Punta Palmilla, 16 July 1968 (S. C. Williams, M. A. Cazier, and party), 89 males, 56 females; 1.5 mile northeast of Punta Palmilla, 17 July 1968 (S. C. Williams, M. A. Cazier, and party), 10 males, 14 females; 2.2 miles southwest of Punta Palmilla, elevation 30 feet, 17 July 1968 (S. C. Williams, M. A. Cazier, and party), 3 females; 3.5 miles southwest of Punta Palmilla, elevation 30 feet, 17 July 1968 (S. C. Williams, M. A. Cazier, and party), 10 males, 10 females; 3.9 miles southwest of Punta Palmilla, elevation 30 feet, 17 July 1968 (S. C. Williams, M. A. Cazier, and party), 47 males, 42 females; 3 miles east of Cabo San Lucas, elevation 30 feet, 18 July 1968 (S. C. Williams, M. A. Cazier, and party), 2 males; Cabo San Lucas, elevation 30 feet, 19 July 1968 (S. C. Williams, M. A. Cazier, and party), 9 males, 4 females; Cabo San Lucas, elevation 30 feet, 20 July 1968 (S. C. Williams, M. A. Cazier, and party), 7 males, 3 females; 2 miles north of Cabo San Lucas, elevation 30 feet, 20 July 1968 (S. C. Williams, M. A. Cazier, and party), 6 males; 6 miles north of Cabo San Lucas, elevation SO feet, 21 July 1968 (S. C. Williams, M. A. Cazier, and party), 10 males, 4 females; 5 miles north of Cabo San Lucas, elevation 50 feet, 21 July 1968 (S. C. Williams, M. A. Cazier, and party), 1 male, 2 females; 4 miles north of Cabo San Lucas, elevation 50 feet, 21 July 1968 (S. C. Williams, M. A. Cazier, and party), 7 males, 1 female; 3 miles east of Cabo San Lucas, elevation 50 feet, 22 July 1968 (S. C. Williams, M. A. Cazier, and party), 4 males, 2 females; 16.5 miles northwest of Cabo San Lucas, 23 July 1968 (S. C. Williams, W. Fox, and M. Bentzien), 1 male; 2 miles northwest of Los Pozos, 23 July 1968 (S. C. Williams, M. Bentzien, and W. Fox), 1 male, 3 females: 4 miles north of Tinaja, 23 July 1968 (S. C. Williams, M. Bentzien, and W. Fox), 3 males, 1 female; 3.5 miles south of El Pescadero, elevation 20 to 25 feet, 23 July 1968 (S. C. Williams, M. A. Cazier, and party), 80 males, 33 females; 5.9 miles north of Todos Santos, elevation 500 feet, 24 July 1968 (S. C. Williams, W. Fox, and M. Bentzien), 7 males, 4 females; 14.8 miles north of Todos Santos, elevation 500 feet, 24 July 1968 (S. C. Williams, W. Fox, and M. Bentzien), 1 male, 1 VoL. XXXVIT] WILLIAMS: NEW SPECIES OF VEJOVIS 297 female; 2.9 miles northwest of San Antonio, elevation 1400 feet, 24 July 1968 (S. C. Williams and M. Bentzien), 25 males, 13 females; 6.8 miles southeast of San Antonio, elevation 1400 feet, 24 July 1968 (S. C. Williams and M. Bentzien), 17 males, 24 females; 0.5 mile east of San Bartolo, elevation 1200 feet, 24 July 1968 (S. C. Williams and M. Bentzien), 4 males, 1 female; 2.5 miles east of San Bartolo, elevation 1200 feet, 24 July 1968 (S. C. Williams and M. Bentzien), 1 male, 4 females; 9.6 miles west of Los Aripes, elevation 400 feet, 25 July 1968 (S. C. Williams, M. Bentzien, and J. Bigelow), 1 female; 11.9 miles west of Los Aripes, elevation 800 feet, 25 July 1968 (S. C. Williams, M. Bentzien, and J. Bigelow), 11 males, 10 females; 21.4 miles west of Los Aripes, elevation 900 feet, 25 July 1968 (S. C. Williams, M. Bentzien, and J. Bigelow), 5 males, 10 females; 31 miles west of Los Aripes, elevation 800 feet, 25 July 1968 (S. C. Williams, M. Bentzien, and J. Bigelow), 5 males, 4 females; 10.5 miles west of El Crucero, 26 July 1968 (S. C. Williams, M. A. Cazier, and party), 1 male, 2 females; 10.3 miles southeast of Santa Rita, 27 July 1968 (S. C. Williams, M. Bentzien, and J. Bigelow), 10 males, 9 females; 35.3 miles northwest of Los Aripes, 27 July 1968 (S. C. Williams, M. Bentzien, and J. Bigelow), 1 male, 1 Bentzien, and J. Bigelow), 1 male, 1 female; 15.0 miles northwest of Los Aripes, female; 49.3 miles southeast Santa Rita, 27 July 1968 (S. C. Williams, M. 27 July 1968 (S. C. Williams, M. Bentzien, and J. Bigelow), 5 males, 2 females; Las Cruces, 29 July 1968 (S. C. Williams, M. A. Cazier, and party), 39 males, 64 females; Las Cruces and for 5 miles southwest, 30 July 1968 (S. C. Williams, M. A. Cazier, and party), 27 males, 19 females. CoMMENT. This species was found throughout the southern part of the peninsula from Comondu to Cabo San Lucas. It was one of the most frequently encountered of all the Vejovis species and was commonly a dominant in the samples. It tended to prefer rocky habitats where considerable fine textured soil also occurred (fig. 12) but was also found in areas away from rocky out- crops. The lack of young instars in the samples was conspicuous. Only one young instar, probably a second or third instar, was collected in all of the samples. The sex ratios significantly favoring males in most of the samples were also conspicuous. Vejovis pumilis Williams, new species. (Figures 13, 14.) Dracnosis. Very minute in size ranking among the smallest species of Vejovis known; total length less than 30 millimeters. Base color of body uni- form pale yellow; mature males with distinctive dark orange telson; both sexes with reddish pedipalp fingers; no other contrasting markings. Pedipalps with short fingers and swollen hands; pectines with 7 to 8 teeth in females, 12 to 16 teeth in males; no denticles on inferior border of movable cheliceral finger. DESCRIPTION OF HOLOTYPE (male). Coloration: carapace, mesosoma, meta- 298 CALIFORNIA ACADEMY OF SCIENCES - [Proc. 47H SEr. el Ficure 12. Type locality and habitat of Vejovis vittatus and Vejovis bruneus, 5 miles southwest of San Miguel de Comondu, Baja California Sur, Mexico, 3 July 1968. soma, pedipalps, and walking legs uniformly yellowish white. Only contrasting markings: telson orange; pedipalp fingers pinkish; pectines almost white; cara- pace and dorsum of mesosoma with faint underlying dusky pigmentation; eyes black, some dusky markings on legs. Carapace: Anterior margin convex; no median emargination; seven reddish bristles. Lateral eyes small and three per group; median diad % carapace width at that point. Carapace surface irregular and finely granulate. Mesosoma: Tergites densely covered with fine granules; two pairs of lateral keels on tergite 7, these not well developed, but set with irregular dentate granules; tergite 7 with obsolescent median keel. Sternites agranulate and lus- trous; stigma inconspicuous and long oval; one pair of poorly developed keels on sternum 7, these smooth to crenulate. Metasoma: Dorsal and dorsolateral keels of segments I to IV set with irregular short dentate granules, segment V lacking dorsal keels and with dorso- laterals almost smooth, no enlarged terminal spines; lateral keels serrate on posterior % of segment I, *% of segment II, %4 of segment III, absent on IV, these keels rounded and approaching obsolescence on anterior 4% of V; all in- ferior lateral keels crenulate to serrate; inferior median keels smooth to ob- solete on I, smooth on II, smooth to crenulate on III and IV, irregularly serrate Vor. XXXVII] WILLIAMS: NEW SPECIES OF VEJOVIS 299 Ficure 13. Vejovis pumilis, new species. Dorsal view of holotype. on V. Inferior intercarinal space of V granulate; inferior median keels of seg- ments I to IV set with two to four reddish bristles. Telson: Ventral surface of vesicle smooth, lustrous and agranulate; this surface with about 12 pairs of short reddish hairs; no subaculear tubercule; aculeus very short, only about % vesicle length; vesicle not bulbous but elon- gate. Pectines: Long and thick; 13 pectinal teeth; fulcra triangular. Genital operculum: Completely divided longitudinally; distinct genital pa- pillae visible externally. Chelicerae: Inferior border of movable finger completely lacking denticles; terminal tooth of inferior and superior borders not subequal, superior terminal Ficure 14. Vejovis pumilis, new species. Ventral view of holotype. 300 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. tooth greatly reduced. Long brownish hair extends from superior base of fixed finger to beyond distal end of finger. Pedipalps: Palms swollen; fingers short compared with palm, fixed finger about % the length of chela; keels absent, hand smooth, agranulate and lus- trous; movable finger and fixed finger each distinctly shorter than carapace. Grasping edge of fingers not scalloped except for one large scallop over the proximal % of fingers, fingers do not meet in this scallop but leave large open space when chela closed; teeth do not extend to proximal ends of fingers. Standard measurements and photographs: Table 5 and figures 13 and 14. DESCRIPTION OF ALLOTYPE (female). Coloration and patterns essentially same as holotype with the following exceptions: lacks bright orange telson, telson similar to metasoma in color; mesosoma slightly darker. Structure es- sentially the same as holotype except: pectine teeth about % fewer (7/8 instead of 13/13); pedipalp chela not so conspicuously swollen; no large proximal space when chela closed; dorsum relatively smooth and lustrous; keelation of meta- soma distinctly reduced, inferior median keels of segments I to IV obsolete, inferior lateral keels of segments I to IV obsolete to slightly crenulate, dorsal and dorsolateral keels more rounded and reduced; vesicle bulbous and not elongate, ventral surface appearing very hirsute with numerous coarse reddish hairs; aculeus proportionally much longer; genital operculum not completely divided longitudinally; no genital papillae. Standard measurements: Table 5. VARIATION WITHIN PARATYPES. Study of 106 paratopotypes (97 males, 9 females) indicated little variation from the description of holotype and allo- type. Males varied in total length from 21 to 27 millimeters (predominantly 25 millimeters), while females varied from 20 to 24 millimeters (predominantly 23 millimeters). Most specimens of both sexes appeared to be mature. Mature females appeared to approximate the mature males in most body proportions. Females showed a definite tendency toward the obsolescence of inferior meta- somal keels as compared with the males. Sexual dimorphism was distinct in that the males had a conspicuous orange telson whereas the females had telsons colored similar to the rest of the cuticle. Other sexual dimorphism occurred as follows: males with distinctly more swollen palms, enlarged pectines, greater pectine tooth counts, genital papillae, completely divided genital operculum, and better developed inferior metasomal keels. The younger instars were con- spicuously few but were essentially similar to the adults with the following exceptions: younger males with pedipalp palms not quite so swollen, and with telson not as deep an orange as adult males. Pectine teeth varied from 12 to 16 (predominantly 13 and 14) in males, and from 7 to 8 in females. The male pectine tooth count of 15 and 16 was a rare occurrence and was represented by a single male with a 15/16 count. VoL. XXXVIT] WILLIAMS: NEW SPECIES OF VEJOVIS 301 TaBLe 5. Measurements (in millimeters) of Vejovis pumilis, new species, holotype and alloty pe. Holotype Allotype (male) (female) Total length 26.5 24.4 Carapace, length il 3.4 width (at median eyes) DD) 2.5 Mesosoma, length led 7.3 Metasoma, length 12.4 10.7 segment I (length/width) elon eS yale segment II (length/width) 2.0/1.7 1.8/1.7 segment III (length/width) BD Nell 1.9/1.6 segment IV (length/width) 2.8/1.6 DW S segment V (length/width) elles BANAL Telson, length 3.8 3.0 Vesicle (length/width) Solel 1 O//i 3 depth 1.0 1.0 Aculeus, length 0.6 ill Pedipalp Humerus (length/width) 2.2/0.8 1.9/0.8 Brachium (length/width) Zroyleal DP i\ il Chela (length/width) 4.0/1.4 3.4/1.0 depth 1.4 1.0 movable finger, length 2.0 1.8 fixed finger, length 1.3 EZ Pectines teeth (left/right) 133// 1183 7/8 TYPE DATA AND ETYMOLOGY. The holotype, allotype, and 106 paratopotypes were collected 26.8 miles west of El Crucero, Baja California Sur, Mexico, 26 July 1968 by S. C. Williams, M. A. Cazier, J. Bigelow, and M. Bentzien. The holotype and allotype are permanently deposited in the California Academy of Sciences. This species is named “‘pumilis” because of its characteristic and distinctive dwarf appearance. MATERIAL. In addition to the holotype, allotype, and 106 paratopotypes, 135 additional paratypes were studied from six other localities in Baja Cali- fornia. These records are as follows: Mexico, Baja California Sur: 28.9 miles west of El Crucero, elevation 25 feet, 26 July 1968 (S. C. Williams, M. A. Cazier, and party), 13 males; 21.4 miles west of El Crucero, elevation 75 feet, 26 July 1968 (S. C. Williams, M. A. Cazier, and party), 104 males, 10 females; 35.5 miles northwest of Los Aripes, elevation 600 feet, 27 July 1968 (S. C. Williams, M. Bentzien, and J. Bigelow), 1 male; 21.4 miles west of Los Aripes, 302 CALIFORNIA ACADEMY OF SCIENCES - [Proc. 47H SER. Ficure 15. Habitat of Vejovis pumilis and Vejovis diazi near type locality. Photograph was taken on Magdalena Plain, 26 miles west of El Crucero, Baja California Sur, Mexico, 27 July 1968. elevation 900 feet, 25 July 1968 (S. C. Williams, M. Bentzien, and J. Bigelow), 1 male, 1 female; 11.9 miles west of Los Aripes, elevation 800 feet, 25 July 1968 (S. C. Williams, M. Bentzien, and J. Bigelow), 4 males; 8 miles northwest of San Raymundo, elevation 500 feet, 30 June 1968 (S. C. Williams, M. A. Cazier, and party), 1 female. CoMMENT. This species appears to have a very limited geographic dis- tribution occurring from near San Raymundo to a few miles north of La Paz. Its populations reached the greatest density on the Magdalena Plain (fig. 15) whereas populations located outside of this area appeared to occur in significantly reduced densities. This is apparently a burrowing species preferring habitats with sparse plant cover and fine textured sedimentary soils. Males were en- countered significantly more often than females, females being conspicuously rare in the samples taken. Juveniles were also conspicuously lacking in the samples taken on the ground surface at night. All specimens were collected on the soil surface at night using the ultraviolet detection method. Vejovis schwenkmeyeri Williams, new species. (Figures 16, 17.) Diacnosis. Medium sized species of Vejovis. Body base color pale yellow; completely lacking stripes and other distinctive markings except for underlying dusky markings on carapace—this most prominent in juveniles and almost ob- VoL. XXXVI] WILLIAMS: NEW SPECIES OF VEJOVIS 303 solete in adults. Pedipalp hands with keels smooth to obsolete; males with open proximal scallop when chela closed; movable finger approximates carapace in length. Metasoma with inferior median keels smooth to obsolete on segments I to III and smooth on segment IV; inferior lateral keels smooth to subtly crenu- late on segments I to III, and crenulate on segment IV. Pectines with 14 to 17 (predominantly 15) teeth on females, and 17 to 19 teeth on males. Inferior border of movable cheliceral finger completely lacking denticles. Appears related to Vejovis confusus Stahnke, from which it differs in the following characters: V. confusus lacks the large proximal open scallop on pedipalp chela of adult males; teeth of pedipalp chela extend to proximal border of both fingers in V. confusus; V. confusus tends to have slightly smaller body size. DESCRIPTION OF HOLOTYPE (male). Coloration: carapace, mesosoma, meta- soma, and pedipalps of uniform pale yellow; walking legs similar but lighter than pedipalps; pectines white. Only contrasting color markings are: eyes black, teeth of chelicerae and fingers of pedipalps reddish, aculeus dark reddish brown. Cuticle otherwise unpigmented. Carapace: Anterior margin essentially straight but with subtle median emargination, set with six erect bristles. Lateral eyes three per group, anterior eye in each group largest. Median eye diad slightly more than '% carapace width at that point. Carapace surface densely covered with large conspicuous granules. Mesosoma: All dorsal plates densely covered by large granules; segment 7 with two pairs of lateral keels with large dentate granules. Sternites smooth and lustrous; one pair of crenulate keels on last sternite. Stigma small and iong oval. Metasoma: All dorsal and dorsolateral keels complete and with large gran- ules giving serrate appearance, except that the dorsals absent on segment V. Lateral keels present and serrate on posterior % of segment I, and posterior % of segments II and III, absent on IV, present and crenulate on anterior half of V. Inferior lateral keels present on all segments; smooth to subtly crenulate on I, II and III; crenulate on IV; serrate on V. Inferior median keels smooth to obsolete on I to III: smooth on IV; serrate on V. Inferior median keels set with 3, 3, 3, 4 pairs of bristles on segments I to IV respectively. Inferior inter- carinal space of V densely covered with conspicuous granules. Telson: Ventral side lacking conspicuous long hairs, but with numerous, short, inconspicuous, whitish hairs apparent under higher magnification. Ven- tral side of vesicle densely covered with conspicuous granules; subaculear tuber- cule present. Pectines: Long and thick; triangular fulcra; 18/19 pectinal teeth. Inferior surface with short, red hair, especially dense on fulcra, middle lamallae and anterior margin. 304 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47H SER. FicurE 16. Vejovis schwenkmeyeri, new species. Dorsal view of holotype. Genital operculum: Completely divided longitudinally; large distinct genital papillae visible externally. Chelicerae: Inferior border of movable finger completely lacking denticles; terminal tooth of superior border of movable finger much shorter than terminal tooth of inferior border. Pedipalps: Palm of hand only slightly swollen, all keels smooth to obsolete. Fixed finger distinctly shorter than carapace, movable finger same length as carapace. Internal margin of fingers with large proximal scallop in which op- posing teeth do not meet when fingers closed; teeth do not extend to proximal end of either finger. Standard measurements and photographs: Table 6 and figures 16 and 17. DESCRIPTION OF ALLOTYPE (female). Morphologically the same as holo- type with the following exceptions: body heavier, carapace and mesosoma broader; chela slightly longer but not as wide; pectine teeth fewer (15/15 in- stead of 18/19); pectines smaller in overall size; no genital papillae; chela with no large proximal scallop when fingers closed; genital operculum not com- pletely divided longitudinally; ventral side of vesicle with numerous long red- dish conspicuous hairs; frontal margin of carapace without distinct median emargination; inferior metasomal keels not as heavily developed. Standard measurerments: Table 6. VARIATION WITHIN PARATYPES. Study of 153 paratopotypes (75 males, 78 females) indicated little variation from the description of holotype and allotype. VoL. XXXVIT] WILLIAMS: NEW SPECIES OF VEJOVIS 305 FicurE 17. Vejovis schwenkmeyeri, new species. Ventral view of holotype. Males varied in total length from 25 to 54 millimeters while females varied from 27 to 56 millimeters. Most specimens appeared to be mature, but young instars of both sexes were present. Little sexual dimorphism occurred with the follow- ing exceptions: females with slightly wider mesosoma and heavier body in adults; male pectines tended to have longer and more numerous teeth; genital papillae present in males; genital operculum of males completely divided longi- tudinally; vesicle of female conspicuously more hirsute than that of male; adult males with proximal open scallop on pedipalp chela. Young instars looked similar to adults except that the cuticle was lighter yellow and had dusky under- lying marking on the dorsum. Dusky markings of juveniles was generally very distinct but variable among individuals. Pectine teeth varied from 14 to 17 in females (predominantly 15) and 17 to 19 in males (predominantly 18 and 19). TYPE DATA AND ETYMOLOGY. The holotype, allotype, and 153 paratopotypes were collected at Bahia de Los Angeles, Baja California Norte, 19 June 1968 by S. C. Williams, M. A. Cazier, and party. The holotype and allotype are permanently deposited in the California Academy of Sciences. This species is named “schwenkmeyeri” in honor of Richard C. Schwenk- meyer who has encouraged many students to pursue studies in field biology and who has also made it possible for many students to travel to various inaccessible regions of the Baja California Peninsula. MatTerIAL. In addition to the holotype, allotype, and 153 paratopotypes, an additional 490 specimens were studied. These came from the following lo- cations in Baja California Norte, Mexico: Puertecitos and for 2 miles south, 10 June 1968 (S. C. Williams, M. A. Cazier, and party), 5 males, 2 females; 306 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. TaBLe 6. Measurements (in millimeters) of Vejovis schwenkmeyeri, new species, holotype and allotype. Holotype Allotype (male) (female) Total length yer 54.3 Carapace, length 5.8 6.8 width (at median eyes) 4.2 5.0 Mesosoma, length 13.8 14.9 Metasoma, length 25.9 24.9 segment I (length/width) 3.6/3.5 3.4/3.8 segment II (length/width) 4.3/3.4 4.0/3.8 segment III (length/width) 4.5/3.3 4.2/3.6 segment IV (length/width) 5.8/3.2 5.5/3.5 segment V (length/width) Wath Sof! 7.8/3.3 Telson, length 7.2 Hell Vesicle (length/width) 5.0/2.9 5.4/3.3 depth 24 2.6 Aculeus, length 2.2 Pe) Pedipalp Humerus (length/width) 525/165 Sa(l//les Brachium (length/width) 5.9/1.9 6.4/2.1 Chela (length/width) 9.1/2.3 9.4/2.0 depth 2.6 Dl movable finger, length 5.8 6.3 fixed finger, length 4.5 5.0 Pectines teeth (left/right) 18/19 15 y/al5 Puertecitos, 10 June 1968 (M. A. Cazier, J. Davidson, and N. Leppla), 2 males, 1 female; Puertecitos, elevation 50 feet, 11 June 1968 (S. C. Williams, M. A. Cazier, and party), 2 males, 2 females; Oakies Landing, 12 June 1968 (S. C. Williams, M. A. Cazier, and party), 45 males, 1 female; 8 miles north of Bahia de San Luis Gonzaga, 13 June 1968 (S. C. Williams, M. A. Cazier, and party), 17 males, 19 females; 6 miles north of Bahia de San Luis Gonzaga, 13 June 1968 (S. C. Williams, M. A. Cazier, and party), 17 males, 19 females; 10 miles north of Bahia de San Luis Gonzaga, 13 June 1968 (S. C. Williams, M. A. Cazier, and party), 10 males, 5 females; 1 mile north of Bahia de San Luis Gonzaga, 14 June 1968 (S. C. Williams, M. A. Cazier, and party), 13 males, 5 females; 6 miles north of Bahia de San Luis Gonzaga, 14 June 1968 (S. C. Williams, M. A. Cazier, and party), 30 males, 26 females: 7 miles north of Bahia de San Luis Gonzaga, 14 June 1968 (S. C. Williams, M. A. Cazier, and party), 20 males, 15 females; Calamajue Arroyo, elevation 1000 feet, 15 June 1968 (S. C. Williams, M. A. Cazier, and party), 11 males, 3 females; Calamajue VoL. XXXVITI] WILLIAMS: NEW SPECIES OF VEJOVIS 307 Arroyo, elevation 1000 feet, 16 June 1968 (S. C. Williams, M. A. Cazier, and party), 7 males, 2 females; Mission San Borjas, elevation 1500 feet, 20 June 1968 (S. C. Williams, M. A. Cazier, and party), 25 males, 12 females; 2 miles northwest of Miller’s Landing, elevation sea level, 21 June 1968 (S. C. Williams, M. A. Cazier, and party), 8 males, 3 females; 3 miles north of Manuela, eleva- tion 500 feet, 22 June 1968 (S. C. Williams, M. A. Cazier, and party), 13 males, 6 females. The following specimens came from Baja California Sur, Mexico: 3 miles southeast of Rancho Tablon, 23 June 1968 (S. C. Williams, M. A. Cazier, and party), 21 males, 11 females; San Ignacio, elevation 500 feet, 24 June 1968 (S. C. Williams, M. A. Cazier, and party), 3 males, 2 females; San Ignacio, elevation 500 feet, 25 June 1968 (S. C. Williams, M. A. Cazier, and party), 1 male, 2 females; San Ignacio, elevation 500 feet, 26 June 1968 (S. C. Williams, M. A. Cazier, and party), 9 males, 2 females; San Angel, 27 June 1968 (S. C. Williams, M. A. Cazier, and party), 7 males, 5 females; San Angel, 28 June 1968 (S. C. Williams, M. A. Cazier, and party), 10 males, 15 females; 35 miles south of San Angel, elevation 50 feet, 29 June 1968 (S. C. Williams, M. A. Cazier, and party), 1 male; 8 miles northwest of San Raymundo, elevation 500 feet, 30 June 1968 (S. C. Williams, M. A. Cazier, and party), 54 males, 24 females; 4 miles west of La Purisima, elevation 375 feet, 1 July 1968 (S. C. Williams, M. A. Cazier, and party), 4 males, 5 females. CoMMENT. This is a relatively widely distributed species, occurring from Puertecitos in Baja California Norte to La Purisima in Baja California Sur. It occurs on the Gulf coast, the coast of the Pacific, and in inland habitats where rocky and sandy habitats occur together (fig. 18). It was a numerically abun- dant species in a number of the habitats sampled, but was rare in a number of habitats as well. Juveniles were conspicuously rare in most of the samples. Specimens collected in the northern part of the range (Puertecitos to Bahia de San Luis Gonzaga) were distinct from typical specimens in that the telson of the adult male had a relatively hirsute appearance and both sexes had dusky markings on the dorsum more distinct. In a few other localities the dusky dorsal markings were more variable also, but this always faint and indistinct. All specimens were collected by means of ultraviolet detection. Vejovis diazi Williams, new species. (Figures 19, 20.) Diacnosts. Medium-sized species of Vejovis. Base body color uniform pale yellow; carapace and mesosoma dorsum may or may not have evident underlying dusky markings; mature individuals of both sexes with contrasting reddish telson and pedipalp fingers. Pedipalp palms moderately swollen and with keels obsolete; movable finger distinctly shorter than carapace. Inferior lateral keels of metasoma basically smooth to crenulate on segments I to IV; inferior median keels obsolete on segments I to IV. Inferior border of movable 308 CALIFORNIA ACADEMY OF SCIENCES [| Proc. 4TH SER. FicureE 18. Type locality and habitat of Vejovis schwenkmeyeri, Bahia de Los Angeles, Baja California Norte, Mexico, 19 June 1968. cheliceral finger completely smooth and lacking denticles. Pectinal teeth vary from 14 to 16 in females and from 17 to 20 in males. Related to Vejovis eusthenura (Wood) from which it differs in the follow- ing characteristics: adults with more reddish pedipalp fingers and more reddish telson; hand of adults more swollen; three pairs of bristles on frontal margin of carapace (five pairs in V. eusthenura); vesicle with fewer basal granules; base color of cuticle slightly more reddish; closed pedipalp chela with distinctly larger proximal open space. DESCRIPTION OF HOLOTYPE (male). Coloration: Carapace, mesosoma, meta- soma, and pedipalps of uniform pale yellow; walking legs similar but lighter than pedipalps; pectines almost white. Carapace with faint, dusky, underlying markings. Only contrasting color markings are: eyes black, teeth of chelicerae reddish orange, fingers of pedipalps light amber, entire vesicle reddish orange, aculeus black, tips of pretarsal claws reddish orange. Cuticle almost trans- parent. Regions of walking legs with faint dusky markings. Carapace: Anterior margin straight, with six erect red bristles. Lateral eyes three per group, most anterior eye in each group largest. One large, erect bristle lateral to posterior margin of each median eye; diad slightly less than “4 cara- pace width at that point. Carapace covered with large granules and slightly lustrous. VoL. XXXVITI) WILLIAMS: NEW SPECIES OF VEJOVIS 309 Ficure 19. Vejovis diazi, new species. Dorsal view of holotype. Mesosoma: All dorsal plates densely covered by large granules; segment 7 with two pairs of lateral keels with large dentate granules; lateral margin of segment 7 abruptly flattened horizontally and with dentate granules. Sternites smooth and lustrous; one pair of keels on last sternite smooth to crenulate. Stigma long oval. Metasoma: All dorsal and dorsolateral keels serrate. Lateral keels present and serrate on posterior % of segment I, posterior % of II, posterior 4 of III, absent on IV, present and serrate to crenulate on anterior '2 of V. Inferior lateral keels present and complete on all segments; segments I to III smooth to crenu- late; IV crenulate to serrate; V serrate. Inferior median keels obsolete on I to IV, serrate on segment V. Area of inferior median keels with three pairs of reddish bristles on segments I to IV. Inferior median keel of segment V with regularly spaced serrate granules. Intercarinal spaces with abundant, moder- ately sized granules. Telson: Ventral side densely covered by long reddish hairs that approach aculeus in length. Vesicle smooth and lustrous and lacking subaculear tubercule. Pectines: Long and thick; large subcircular to triangular fulcra; 19/19 pectinal teeth. Inferior surface with short, red hair, especially dense on fulcra, middle lamellae and anterior margin. Genital operculum: Completely divided longitudinally; large genital pa- pillae visible externally. Chelicerae: Inferior border of movable finger completely lacking denticles; terminal tooth of superior border of movable finger subequal to terminal tooth of inferior border. 310 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 20. Vejovis diazi, new species. Ventral view of holotype. Pedipalps: Palm moderately swollen, all keels obsolete. Fixed finger and movable finger each shorter than carapace. Internal margin of fingers faintly scalloped; teeth do not extend to proximal end of fingers. Proximal teeth do not meet in last scallop when chela closed. Standard measurements and photographs: Table 7 and figures 19 and 20. DESCRIPTION OF ALLOTYPE (female). Morphologically the same as holo- type with the following exceptions: slightly smaller in total length; slightly smaller in carapace width; larger in mesosomal length and smaller in meta- somal length; chela slightly shorter and narrower; pectine with about 25 per- cent fewer teeth (14/13 instead of 19/19); dusky markings of carapace faintly detectable; carapace and tergites not as granular; carapace and tergites more lustrous; pectines smaller, no genital papillae, genital operculum not completely divided longitudinally; vesicle not as dark reddish orange; lacking basal scallop of pedipalp fingers. Standard measurements: Table 7. VARIATION WITHIN PARATYPES. Study of 19 paratopotypes (10 males, 9 females) indicated little significant variation from the descriptions of holotype and allotype. Males varied in total body length from 29 to 48 millimeters while females varied from 23 to 43 millimeters. The sample contained both adult and subadult instars but younger instars were totally lacking. Little sexual dimorphism occurred except for structure of pectines, presence of genital pa- pillae in males, complete division of genital operculum in males, and large proxi- mal open space in mature male pedipalp chela. Subadults were similar to adults VoL. XXXVIT] WILLIAMS: NEW SPECIES OF VEJOVIS 311 TABLE 7. Measurements (in millimeters) of Vejovis diazi, new Species, holotype and alloty pe. Holotype Allotype (male) (female) Total length 42.5 41.6 Carapace, length 52 5.0 width (at median eyes) 3.8 3.6 Mesosoma, length 10.5 13.8 Metasoma, length 20.6 eS segment I (length/width) 10/32 2.4/3.0 segment II (length/width) SV? BAIL DS segment III (length/width) 3.4/3.2 2.9/2.8 segment IV (length/width) 4.6/3.2 3.7/2.8 segment V (length/width) 6.3/3.2 5.6/2.8 Telson, length 6.2 By Vesicle (length/width) 42/23 Defy) Af) depth 1.9 1.8 Aculeus, length 2.0 lee Pedipalp Humerus (length/width) 4.2/1.4 3.8/1.4 Brachium (length/width) 4.8/1.8 4.3/1.6 Chela (length/width) TBVOM 6.6/1.7 depth 2.2 1.9 movable finger, length 4.5 4.0 fixed finger, length Bs) BP Pectines teeth (left/right) 19/19 14/13 except that young males lacked open space on pedipalp chela. Most subadults of both sexes showed some very faint dusky pigmentation lateral to ocular tubercule—this was not apparent in adults. Adults of both sexes had pedipalp fingers and vesicle reddish brown, subadults by contrast did not. Pectine teeth varied from 14 to 16 (predominantly 15) in females and from 17 to 20 (nre- dominantly 19 and 20) in males. TYPE DATA AND ETYMOLOGY. The holotype, allotype, and 19 paratopotypes were collected 21.4 miles west of El Crucero, elevation 75 feet, 26 July 1968, by S. C. Williams, M. A. Cazier, and party. The holotype and allotype are permanently deposited in the California Academy of Sciences. This species is named “diaz” in honor of A. Diaz Najera of the Instituto de Salubridad y Enfermedades Tropicales of Mexico, because of his significant contributions to the knowledge of the North American scorpion fauna. MatTERIAL. In addition to the holotype, allotype, and 19 paratopotypes, 312 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. 1394 (568 males, 826 females) other specimens were studied. These specimens were from the following localities in Baja California Sur, Mexico: San Ignacio, elevation 500 feet, 25 June 1968 (S. C. Williams, M. A. Cazier, and party), 3 males, 5 females; San Ignacio, elevation 500 feet, 26 June 1968 (S. C. Williams, M. A. Cazier, and party), 5 females; 35 miles south of San Angel, elevation 50 feet, 29 June 1968 (S. C. Williams, M. A. Cazier, and party), 5 males, 4 females; 8 miles northwest of San Raymundo, elevation 500 feet, 30 June 1968 (S. C. Williams, M. A. Cazier, and party), 12 males, 22 females; 8 miles northwest of San Raymundo, elevation 500 feet, 30 June 1968 (S. C. Williams, M. A. Cazier, and party), 76 males, 28 females; 4 miles west of La Purisima, elevation 1375 feet, 1 July 1968 (S. C. Williams, M. A. Cazier, and party), 29 males, 23 females; 5 to 10 miles southwest of San Miguel de Comondu, 3 July 1968 (S. C. Williams, M. A. Cazier, and party), 16 males, 22 females; 75 miles northwest of La Paz, elevation 200 feet, 4 July 1968 (S. C. Williams, M. A. Cazier, and party), 20 males, 16 females; 5 to 6 miles southwest of La Paz, elevation 25 feet, 5 July 1968 (S. C. Williams, M. A. Cazier, and party), 5 males, 6 females; La Paz, elevation 25 feet, 5 July 1968 (S. C. Williams, M. A. Cazier, and party), 2 males, 8 females; 5 to 10 miles west of La Paz, elevation 25 feet, 6 July 1968 (S. C. Williams, M. A. Cazier, and party), 10 males, 12 females; 5 to 6 miles southwest of La Paz, elevation 25 feet, 6 July 1968 (S. C. Williams, M. A. Cazier, and party), 7 males, 9 females; 1 mile south of La Paz, elevation 100 feet, 6 July 1968 (S. C. Williams, M. A. Cazier, and party), 28 males, 39 females; Isla Espiritu Santo, 7 July 1968 (S. C. Williams, M. Bentzien, and W. Fox), 23 males, 56 females; Isla Partida, 9 July 1968 (S. C. Williams, M. Bentzien, and W. Fox), 15 males, 31 females; 1 mile east of Los Aripes, 8 July 1968 (M. A. Cazier, J. Bigelow, and N. Leppla), 7 males, 12 females; 14 miles northeast of La Paz, 9 July 1968 (M. A. Cazier, J. Bigelow, and N. Leppla), 23 males, 39 females; 14 miles northeast of La Paz, 10 July 1968 (M. A. Cazier, J. Bigelow, and N. Leppla), 11 males, 20 females; % mile north of La Paz airport, 12 July 1968 (M. A. Cazier and J. Bigelow), 11 males, 27 females; ’ mile north of La Paz airport, 13 July 1968 (S. C. Williams, M. A. Cazier, and party), 42 males, 72 females; 4 mile north of La Paz airport, 13 July 1968 (S. C. Williams, M. A. Cazier, and party), 1 male, 1 female; 14 miles northeast of La Paz, 14 July 1968 (S. C. Williams, J. Bigelow, W. Fox, and J. Davidson), 3 males, 3 females; 7 miles west of El Triunfo, elevation 1800 feet, 24 July 1968 (S. C. Williams and M. Bentzien), 2 males, 2 females; 9.6 miles west of Los Aripes, elevation 400 feet, 25 July 1968 (S. C. Williams, M. Bentzien, and J. Bigelow), 50 males, 90 females; 11.9 miles west of Los Aripes, elevation 800 feet, 25 July 1968 (S. C. Williams, M. Bentzien, and J. Bigelow), 5 males, 11 females; 21.4 miles west of Los Aripes, elevation 900 feet, 25 July 1968 (S. C. Williams, M. Bentzien, and J. Bigelow), 7 males, 11 females; 31.0 miles west of Los Aripes, elevation 800 feet, 25 July 1968 (S. C. Williams, M. Bentzien, and J. Bigelow), 3 males, VoL. XXXVII] WILLIAMS: NEW SPECIES OF VEJOVIS 313 4 females; La Paz, 25 July 1968 (M. A. Cazier), 2 females; 28.9 miles west of E] Crucero, 26 July 1968 (S. C. Williams, M. A. Cazier, and party), 1 male, 2 females; 26.8 miles west of El Crucero, 26 July 1968 (S. C. Williams, M. A. Cazier, and party), 7 males, 10 females; 10.5 miles west of El Crucero, 26 July 1968 (S. C. Williams, M. A. Cazier, and party), 1 male, 1 female: Santa Rita, 27 July 1968 (S. C. Williams, M. Bentzien, and J. Bigelow), 3 males, 2 females; 10.3 miles southeast Santa Rita, 27 July 1968 (S. C. Williams, M. Bentzien, and J. Bigelow), 11 males, 17 females; 27.3 miles southeast of Santa Rita, 27 July 1968 (S. C. Williams, M. Bentzien, and J. Bigelow), 2 males, 5 females; 35.3 miles northwest of Los Aripes, 27 July 1968 (S. C. Williams, M. Bentzien, and J. Bigelow), 9 males, 13 females; 35.3 miles northwest of Los Aripes, 27 July 1968 (S. C. Williams, M. Bentzien, and J. Bigelow), 1 male; Las Cruces, 29 July 1968 (S. C. Williams, M. A. Cazier, and party), 38 males, 72 females; Las Cruces and southwest for 5 miles, 30 July 1968 (S. C. Williams, M. A. Cazier, and party), 79 males, 121 females; 5 miles southwest of La Paz, elevation 25 feet, 31 July 1968 (S. C. Williams and M. Bentzien), 1 male, 2 females. CoMMENT. In the type locality on the Magdalena Plain, this species lived in relatively flat, rock-free terrain (fig. 15). The soils here were fine textured, moderately packed sediment. The vegetation was of a xeric type with many cacti and ocotillos. The ocotillos were often densely covered by hanging lichen. In other parts of the geographic range, this species also was found in predominantly rocky, volcanic habitats in regions where loosely packed soils had accumulated. Three morphological characteristics tended to vary over the geographic range: telson hirsuteness, body size, and emphasis of underlying dark pigment on the carapace and mesosomal dorsum. In some habitats, this pigment was greatly emphasized, especially in younger instars, while in other habitats this pigment formation was obsolescent and barely detectable. Females were significantly more abundant than males in most of the samples. All specimens were collected by ultraviolet detection. Vejovis hoffmanni Williams, new species. (Figures 21, 22.) Diacnosis. Medium sized species of Vejovis with mature females attaining slightly larger body sizes than mature males. Body base color pale golden yellow; with underlying faint dusky markings on carapace near ocular tubercule. Inferior lateral keels of metasoma smooth to crenulate (basically smooth) on segments I to IV; inferior median keels absent or obsolete on segments I to IV. Pectines with 15 to 18 teeth in females, 18 to 21 teeth in males. Carapace slightly longer than movable pedipalp finger; palm of chela not greatly swollen, lacks keels and agranulate. Inferior border of movable cheliceral finger com- pletely lacking denticles. Related to Vejovis schwenkmeyeri from which it differs in the following 314 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47H SER. characteristics: metasoma with inferior median keels absent or obsolete, inferior lateral keels more smooth; pedipalp fingers of adults not reddish; smaller body size; slightly stouter metasomal segments; adult males and females do not differ in vesicle hirsuteness. DESCRIPTION OF HOLOTYPE (male). Coloration: Carapace, mesosoma, metasoma, pedipalps, and walking legs uniform, pale golden yellow; underlying faint dusky markings on carapace later and posterior to ocular tubercule; pec- tines whitish; eyes black; aculeus reddish brown; dusky marking in places on legs. Carapace: Anterior margin straight, no emargination, set with three pairs of erect brown bristles. Lateral eyes three per group, anterior two largest in each group; median diad % carapace width at that point; one pair of long brownish hairs on ocular tubercule posterior to diad. Carapace densely covered with granules. Mesosoma: Terga densely granular; segment 7 with two pairs of lateral keels and one median keel, these irregularly dentate. Sterna smooth, lustrous and agranulate; sternite 7 with one pair of finely crenulate keels; stigma long oval. Metasoma: All dorsal and dorsolateral keels serrate except that dorsal keels of segment V absent, dorsal keels of segments I to II each terminate posteriorly in a slightly enlarged spine; lateral keels irregularly serrate on posterior % segment I, posterior % of II, posterior % of III, absent on IV, crenulate on anterior ’ of V; inferior lateral keels smooth to crenulate on segments I to IV, serrate on V; inferior median keels obsolete on segments I to IV, serrate on V. Inferior intercarinal space of segment V densely granular. Telson: Vesicle appears very hirsute; ventral surface of vesicle with about 20 pairs of long conspicuous reddish hairs; ventral surface of vesicle basically smooth and lustrous, but with numerous low, rounded, obsolete granules; sub- aculear tubercule subtle. Pectines: Long and thick; 19 long pectinal teeth; triangular to subcircular middle lamellae. Gential operculum: Completely divided longitudinally; genital papillae visible externally. Chelicerae: Inferior border of movable finger completely lacking denticles; terminal tooth of inferior border and terminal tooth of superior border of mov- able finger subequal, superior tooth being shorter. Pedipalps: Palms only slightly swollen; keels absent, surface of palm smooth, agranulate and lustrous. Fixed finger and movable finger each shorter than carapace; fingers elongate, fixed finger slightly longer than palm. Grasp- ing edge of fingers with only one slight scallop, this along proximal % of fixed finger; opposing fingers touch only at distal end when chela closed: teeth do not extend to proximal ends of fingers. Standard measurements and photographs: Table 8 and figures 21 and 22. VoL. XXXVIT] WILLIAMS: NEW SPECIES OF VEJOVIS 315 Ficure 21. Vejovis hoffmanni, new species. Dorsal view of holotype. DESCRIPTION OF ALLOTYPE (female). Essentially the same as holotype in color and structure with the following exceptions: slightly larger in most body dimensions; dusky markings on carapace reduced; slightly fewer pectinal teeth (16/17 instead of 19/19); genital papillae absent; genital operculum not com- pletely divided longitudinally; scallop on fixed finger of chela not well devel- oped; pectines smaller and with considerably shorter teeth; carapace and tergites not as granular and with more lustrous appearance. Standard measurements: Table 8. VARIATION WITHIN PARATYPES. Study of 137 paratopotypes (74 males, 63 females) indicated little variation from the description of holotype and allotype. Males varied in length from 13 to 35 millimeters while females varied from 18 Ficure 22. Vejovis hoffmanni, new species. Ventral view of holotype. 316 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SEr. TABLE 8. Measurements (in millimeters) of Vejovis hoffmanni, new species, holotype and alloty pe. Holotype Allotype (male) (female) Total length 3325 36.5 Carapace, length 4.0 4.8 width (at median eyes) 3.0 3.6 Mesosoma, length 8.8 10.2 Metasoma, length 16.0 16.1 segment I (length/width) 2.2/2.4 OY Ot segment II (length/width) Desf Des) 2.5/2.6 segment III (length/width) Dif 2s Delf PS) segment IV (length/width) 3.6/2.3 Se 5/255 segment V (length/width) 5.0/2.4 5.2/2.6 Telson, length 4.7 5.4 Vesicle (length/width) Ses 3.4/2.2 depth 1.4 1.8 Aculeus, length 1.6 2.0 Pedipalp Humerus (length/width) 3 VAleil SHoyAll Brachium (length/width) Sil) led, 4.0/1.6 Chela (length/width) 5.6/1.3 6.3/1.5 depth 1.4 1.6 movable finger, length Bi 4.1 fixed finger, length 2.9 Soil Pectines teeth (left/right) 19/19 16/17 to 40 millimeters. The specimens were variable in size and ranged from about third instars to sexually mature adults. Mature females reached longer body lengths and had a slightly heavier mesosoma. A definite tendency toward ob- solescence of the inferior median keels of the metasoma was apparent. The main sexual dimorphisms were: pectines smaller and with fewer teeth in the female; males with genital papillae; males with genital operculum completely divided longitudinally; adult males with open scallop on pedipalp chela. Both sexes were similar in coloration and other body proportions. Young instars were similiar to adults in color and structure, but smaller in size. Pectine teeth varied from 15 to 18 (predominantly 16 and 17) in females and from 18 to 21 (pre- dominantly 19 and 20) in males. TYPE DATA AND ETYMOLOGY. The holotype, allotype, and 137 paratopotypes were collected 3 miles north of Manuela, elevation 500 feet, Baja California Norte, Mexico, 22 June 1968 by S. C. Williams, M. A. Cazier, and party. The VoL. XXXVITI] WILLIAMS: NEW SPECIES OF VEJOVIS 317 holotype and allotype are permanently deposited in the California Academy of Sciences. This species is named “‘hoffmanni” in honor of the late Carlos C. Hoffmann who devoted many years to studying the scorpion fauna of mainland Mexico. His systematic monograph on the scorpions of Mexico has been an outstanding and very useful contribution to the knowledge of North American scorpions. MATERIAL. In addition to the holotype, allotype, and 137 partopotypes, 304 (161 males, 143 females) paratypes were studied from four other localities. These specimens came from the following places in Baja California Norte, Mexico: 2 miles northwest of Miller’s Landing, elevation 25 feet, 21 June 1968 (S. C. Williams, M. A. Cazier, and party), 54 males, 35 females. The remainder of these specimens came from the following localities in Baja California Sur, Mexico: 3 miles southeast of Rancho Tablon, 23 June 1968 (S. C. Williams, M. A. Cazier, and party), 28 males, 26 females; San Ignacio, elevation 500 feet, 24 June 1968 (S. C. Williams, M. A. Cazier, and party), 6 males; San Ignacio, elevation 500 feet, 26 June 1968 (S. C. Williams, M. A. Cazier, and party), 17 males, 13 females; San Angel, elevation 500 feet, 27 June 1968 (S. C. Wil- liams, M. A. Cazier, and party), 14 males, 12 females, San Angel, elevation 500 feet, 28 June 1968 (S. C. Williams, M. A. Cazier, and party), 42 males, 57 females. CoMMENT. This species was found predominantly in moderately packed sedimentary soils along the western side of the Baja peninsula from San Ignacio to near La Purisima. It often occurred in habitats completely lacking surface rocks or other surface cover, and appears to be a burrowing species (fig. 23). It has the tendency to form local color races in accord with the predominant substrate. This was most conspicuous around the San Angel sand dunes where the cuticle had a lighter base color and the dusky carapace pigment was very faint. At San Ignacio this species had a cuticle with a darker base color and more prominent dusky pigmentation on the dorsum. All specimens were collected by means of ultraviolet detection. Vejovis bruneus Williams, new species. (Figures 24, 25.) Dracnosis. Medium-sized species of Vejovis with mature females attain- ing larger body proportion than mature males. Body base color uniform light brown; pedipalps with contrasting reddish brown fingers; pedipalp hands with greatly swollen palm; palm with well developed and granulate keels; movable finger distinctly shorter than carapace; internal border of fingers not scalloped and no distinct open space when chela closed. Inferior lateral and inferior median keels of metasoma distinctly developed and very finely crenulate to finely serrate. Vesicle of both sexes with a number of fine, inconspicuous hairs. Chelicerae with inferior border of movable finger completely lacking denticles. 318 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Ficure 23. Habitat of Vejovis hoffmanni, on plateau overlooking San Juan Arroyo, 8 miles northwest of San Raymundo, Baja California Sur, Mexico, 30 June 1968. Related to Vejovis punctipalpus (Wood) and Vejovis insularis Williams. Differs from V. punctipalpus in that neither sex with scalloped pedipalp fin- gers or open space when chela closed; no distinct sexual dimorphism of pedipalp hand; V. bruneus has telson with emphasized aculeus but this not as distinctly so as with V. punctipalpus; cuticle and pedipalp fingers more brownish. Vejovis bruneus differs from Vejovis insularis in the following ways: vesicle of male and female not densely covered with long robust hairs; mature female without scal- loped pedipalp fingers; telson without vesicle so greatly reduced and aculeus so greatly elongate. DESCRIPTION OF HOLOTYPE (male). Coloration: Carapace, mesosoma, metasoma, and pedipalps of uniform light brown; walking legs similar but lighter than pedipalps; pectines white. Only contrasting color markings are: eyes black, fingers of pedipalps dark reddish brown, aculeus dark brown; various cuticular keels darker than surrounding cuticle. Carapace: Anterior border with distinct, broad median emargination produc- ing rounded frontal lobes, this border set with six short erect bristles. Lateral eyes three per group, most anterior eye in each group largest. Median eye diad % carapace width at that point. Carapace surface densely covered with large granules. Mesosoma: All dorsal plates densely covered by large granules; segment 7 Vor. XXXVIT] WILLIAMS: NEW SPECIES OF VEJOVIS 319 Ficure 24. Vejovis bruneus, new species. Dorsal view of holotype. with two pairs of lateral keels and one medial keel, these keels set with large dentate granules; each inner lateral keel terminates posteriorly with enlarged tooth. Sternites relatively smooth and lustrous; one pair of crenulate keels on sternite 7. Stigma long oval. Metasoma: All dorsal and dorsolateral keels complete and with large granules giving serrate appearance, except that the dorsals absent on segment V; each dorsal keel terminates posteriorly in a slightly enlarged tooth. Lateral keels present and serrate on posterior % of segment I, posterior % of II and III, absent on IV, present and serrate on anterior 7 of V. Inferior lateral and in- ferior median keels distinct and finely serrate to crenulate on all segments. In- ferior intercarinal space of segment V without noticeable granulation. Telson: Ventral surface of vesicle smooth and lustrous, lacking granulation; this surface with a number of fine inconspicuous hairs; no subaculear tubercule. Pectines: Long and thick; triangular fulcra; 18 pectinal teeth. Inferior surface with short, red hair, especially dense on fulcra, middle lamellae and anterior margin. Genital operculum: Completely divided longitudinally; large distinct genital papillae visible externally. Chelicerae: Inferior border of movable finger completely lacking denticles; terminal tooth of superior border of movable finger and terminal tooth of inferior border subequal. Pedipalps: Palm of hand greatly swollen; all keels well developed and granular. Fixed and movable finger each distinctly shorter than carapace. In- CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Os bo (=: Ficure 25. Vejovis bruneus, new species. Ventral view of holotype. ternal margin of fingers not scalloped; no open gap exists when chela closed; fingers with teeth extending to proximal border. Standard measurements and photographs: Table 9 and figures 24 and 25. DESCRIPTION OF ALLOTYPE (female). Morphologically the same as holo- type with the following exceptions: strikingly larger in all body dimensions; pectines with slightly fewer teeth (16 instead of 18); no genital papillae; genital operculum not completely divided longitudinally; ventral surface of vesicle ap- pears more hirsute; aculeus proportionally longer. Standard measurements: Table 9. VARIATION WITHIN PARATYPES. Study of 173 paratopotypes (124 males, 49 females) indicated little significant variation from the description of holo- type and allotype. Males varied in total length from 22 to 41 millimeters while females varied from 19 to 50 millimeters. Most specimens of both sexes ap- peared to be mature. Mature females appeared to attain larger body proportions than did mature males. Pectines varied from 15 to 17 (predominantly 16) in females and from 17 to 19 (predominantly 18) in males. Little sexual di- morphism occurred except for body size, smaller female pectines, females with fewer pectinal teeth, presence of genital papillae in males, and completely divided male genital operculum. Both sexes were very similar in color, struc- ture and proportions. Young instars differed from adults in that they were more whitish in color, lacked reddish brown pigments on pedipalp fingers and had pedipalp palms not swollen. VoL. XXXVIT] WILLIAMS: NEW SPECIES OF VEJOVIS 321 TABLE 9. Measurements (in millimeters) of Vejovis bruneus, new species, holotype and alloty pe. Holotype Allotype (male) (female) Total length 38.3 il Carapace, length bee 6.7 width (at median eyes) 3.6 4.6 Mesosoma, length 9.9 11520 Metasoma, length 18.4 2355) segment I (length/width) 2.6/2.7 322/322 segment II (length/width) 2.9/2.5 3.8/3.1 segment III (length/width) 3.0/2.4 4.0/2.9 segment IV (length/width) 4.2/2.2 Seen segment V (length/width) Sia I BYOS Telson, length 4.8 6.0 Vesicle (length/width) 3.0/1.6 3.3/2.0 depth es 1.6 Aculeus, length 1.8 Bot Pedipalp Humerus (length/width) 4.0/1.5 4.2/1.9 Brachium (length/width) 4.1/1.8 5.4/2.3 Chela (length/width) 7.4/2.8 10.3/3.4 depth 3.0 3.6 movable finger, length 4.1 5.8 fixed finger, length 3.0 4.4 Pectines teeth (left/right) 18/18 16/16 TYPE DATA AND ETYMOLOGY. The holotype, allotype, and 173 paratopo- types were collected 5 miles southwest of San Miguel de Comondu, elevation 1000 feet, 2 July 1968 by S. C. Williams, M. A. Cazier, and party. The holo- type and allotype are permanently deposited in the California Academy of Sciences. This species is named ‘“‘bruneus” because of its brownish coloration. MatTerRIAL. In addition to the holotype, allotype, and 173 paratopotypes, 173 additional paratypes were studied. These were from the following locations in Baja California Sur, Mexico: 8 miles northwest of San Raymundo, elevation 500 feet, 30 June 1968 (S. C. Williams, M. A. Cazier, and party), 23 males, 3 females; 5 to 10 miles southwest of San Miguel de Comondu, elevation 1000 feet, 3 July 1968 (S. C. Williams, M. A. Cazier, and party), 117 males, 30 females. ComMMENT. This species was found only in the samples taken around Comondu and near San Raymundo. The habitats were predominantly rocky volcanics where some fine textured soil had accumulated (fig. 12). This species 322 CALIFORNIA ACADEMY OF SCIENCES [| Proc. 4TH SER. appears to be a burrower. Young instars and adult females were conspicuously rare in the samples. All specimens were collected by means of ultraviolet de- tection. Vejovis insularis Williams, new species. (Figures 26, 27.) Dracnosis. Medium sized species of Vejovis with mature females attain- ing larger body size than mature males. Base color of body bright yellow, with pedipalp fingers dark reddish brown. Pedipalp hands with short fingers and swollen palms; movable finger distinctly shorter than carapace; keels well de- veloped and granular on palm; neither sex with scalloped fingers or open space when chela closed. Chelicerae with inferior border of movable finger lacking denticles; inferior median and inferior lateral keels of metasoma well developed and finely serrate; telson with conspicuously reduced vesicle and conspicuously elongate aculeus; vesicle of both sexes densely covered with long conspicuous reddish hairs; pectines with 18 to 19 teeth in males and 16 to 17 teeth in females. Closely related to Vejovis punctipalpus (Wood) from which it differs in the following characteristics: hirsuteness of telson; absence of scalloped pedipalp fingers on both sexes; absence of proximal space when chela closed. DESCRIPTION OF HOLOTYPE (male). Coloration: Carapace, mesosoma, metasoma and pedipalps uniform bright yellow; walking legs similar to pedi- palps but lighter; pectines white. Only contrasting color markings: eyes black, cheliceral teeth light brownish, pedipalp fingers light reddish brown, aculeus dark reddish brown, tips of pretarsal claws light brown. Cuticle otherwise un- pigmented. Carapace: Anterior margin with six red bristles and with subtle median emargination; lateral eyes three per group, most anterior eye in each group largest; median eye diad slightly greater than % carapace width at that point; carapace surface densely covered with large granules. Mesosoma: All tergites densely covered by large granules; tergite 7 with two pairs of coarse granular lateral keels and a short granular median keel; inner lateral keels each terminate posteriorly in a slightly enlarged spine. Ster- nites agranular; one pair of keels on sternite 7, this crenulate to serrate; stigma short slits. Metasoma: All dorsal and dorsolateral keels serrate except that dorsal keels of segment V absent, these keels on segments I to IV each terminate posteriorly as an enlarged spine; lateral keels serrate on posterior % of I, % of II and III, absent on IV, serrate to crenulate on anterior “% of V:; inferior median and in- ferior lateral keels all well developed and finely serrate. Inferior median keels each set with three or four pairs of reddish bristles on segments I to IV; inferior intercarinal space of segment V mostly smooth and not conspicuously granular. All dorsal intercarinal spaces granular. VoL. XXXVII] WILLIAMS: NEW SPECIES OF VEJOVIS 323 FicurE 26. Vejovis insularis, new species. Dorsal view of holotype. Telson: Ventral aspect of vesicle smooth and lustrous, lacking granulation; densely covered with long, conspicuous reddish hairs, each approximating aculeus in length; no subaculear tubercule; aculeus long and curved, about 7” length of vesicle. Pectines: Long and broad; 18/17 pectinal teeth; with abundant reddish hairs, these most abundant on fulcra, middle lamellae, and anterior margin; fulcra triangular. Ficure 27. Vejovis insularis, new species. Ventral view of holotype. 324 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. TaBLe 10. Measurements (in millimeters) of Vejovis insularis, new species, holotype and alloty pe. Holotype Allotype (male) (female) Total length 38.0 48.1 Carapace, length 5.0 6.3 width (at median eyes) 35) 4.4 Mesosoma, length 10.0 14.2 Metasoma, length 18.2 21.7 segment I (length/width) 25/25 2.8/3.0 segment II (length/width) 3.0/2.4 3.6/2.8 segment III (length/width) SP 8 3.8/2.8 segment IV (length/width) 4.0/2.2 4.8/2.6 segment V (length/width) 5.5/2.0 6.7/2.4 Telson, length 4.8 5.9 Vesicle (length/width) 2.8/1.6 3.3/1.8 depth te? 1.4 Aculeus, length 2.0 2.6 Pedipalp Humerus (length/width) 3.8/1.3 Stl filed Brachium (length/width) 4.0/1.7 DYDD Chela (length/width) 7.4/2.5 9.5/3.0 depth 2.7 3.4 movable finger, length 4.1 5.5 fixed finger, length 2.8 4.2 Pectines teeth (left/right) 18/17 17/17 Genital operculum: Completely divided longitudinally; distinct genital papillae visible externally. Chelicerae: Inferior border of movable finger completely lacking denticles. Pedipalps: Palms swollen; keels well developed and with low rounded granules. Fixed finger and movable finger each distinctly shorter than cara- pace; grasping edges of fingers not scalloped, no gaping space between fingers when chela closed. Standard measurements and photographs: Table 10 and figures 26 and 27. DESCRIPTION OF ALLOTYPE (female). Coloration and patterns essentially the same as holotype except allotype with more brownish base color and darker pedipalp fingers. Structure essentially the same as holotype except: body size much larger in all dimensions; lacks genital papillae; genital operculum not completely divided longitudinally. Standard measurements: Table 10. VARIATION WITHIN PARATYPES. Study of 3 paratopotypes (2 males, 1 Voi. XXXVII] WILLIAMS: NEW SPECIES OF VEJOVIS 325 female) indicated little variation from the description of holotype and allotype. Males were all about the same body size and varied in total length from 29 to 33 millimeters, while females also were about the same body size and varied from 44 to 48 millimeters. Pectine tooth counts varied from 16 to 17 (pre- dominantly 16) in females and from 18 to 19 (predominantly 19) in males. The males were distinctly shorter in body length than the holotype and had hairs on vesicle somewhat shorter. TYPE DATA AND ETYMOLOGY. The holotype, allotype, and 3 paratopotypes were collected on Isla Partida, in the large central valley, Baja California Sur, Mexico, 10 July 1968 by S. C. Williams, M. Bentzien, and W. Fox. Isla Partida is the small separated island adjacent to the north end of Isla Espiritu Santo (several other islands are sometimes also referred to as ‘“‘Partida’’). The holo- type and allotype are permanently deposited in the California Academy of Sciences. This species is named “‘insularis” because it is known only from Isla Espiritu Santo and Isla Partida. It may occur on other Gulf islands but it appears not to occur on the peninsula of Baja California. MatTeriAL. In addition to the holotype, allotype, and 3 paratopotypes, 4 additional paratypes were studied. These came from the following localities in Baja California Sur, Mexico: Isla Espiritu Santo, north shore facing Isla Partida, 8 July 1968 (S. C. Williams, M. Bentzien, and W. Fox), 1 male, 1 female; Isla Partida, large central valley, 9 July 1968 (S. C. Williams, M. Bentzien, and W. Fox), 2 females. CoMMENT. This species is known only from Isla Partida and Isla Espiritu Santo in the southern end of the Gulf of California. The preferred habitat ap- peared to be places where rocky volcanic hillsides met open fine textured soils at their base (fig. 28). This species was not abundant on these islands judging by the samples and was not widely distributed among the available habitats. The specimens on Isla Espiritu Santo were very similar to those of Isla Partida. The two islands have apparently only recently become separated by a narrow channel of sea. The scorpion populations on the two islands are probably bio- logically isolated at this time, but this isolation has not been long enough for the population to diverge to a distinguishable degree. All specimens were col- lected by ultraviolet detection. Vejovis gravicaudus Williams, new species. (Figures 29, 30.) Diacnosis. Medium to large species of Vejovis with mature females at- taining larger and heavier bodies than mature males. Carapace and mesosomal dorsum brownish yellow with underlying dusky markings; two pairs of dark stripes underlying inferior keels of metasoma. Pedipalp with palm slightly swollen and with keels obsolete; palm agranular; movable finger shorter than 326 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 28. Type locality and habitat of Vejovis insularis on Isla Partida, Baja Cali- fornia Sur, Mexico, 9 July 1968. carapace. Metasoma with inferior lateral keels greatly reduced and smooth; inferior median keels obsolete. Pectines with 19 to 23 teeth in females, 24 to 27 teeth in males. Inferior border of movable cheliceral finger lacking denticles; mature males with vesicle appearing bald while mature females have very hirsute vesicle. Related to Vejovis spinigerus (Wood) but differs in the following character- istics: vesicle of mature female distinctly more hirsute than in V. spinigerus; mature male with vesicle distinctly less hirsute than in V. spinmigerus; very definite sexual dimorphism exists in mature instars in regard to telson hirsuteness (this is not the case in V. spinigerus); greater proximal space exists between fingers when chela closed; dorsal stripes on mesosoma not distinct—even in young instars. DESCRIPTION OF HOLOTYPE (male). Coloration: Carapace and mesosoma brownish yellow with underlying dusky markings; metasoma pale yellow with keels contrastingly reddish orange; pedipalps and walking legs pale yellow; pec- tines yellowish white. Contrasting color markings are: eyes black, teeth of chelicerae reddish brown, fingers of pedipalp light amber, vesicle reddish brown, aculeus dark red to black, position of inferior keels of metasoma outlined with dark pigment, tips of pretarsal claws light orange. Carapace: Anterior margin straight with 7 erect bristles. Lateral eyes three VoL. XXXVIT] WILLIAMS: NEW SPECIES OF VEJOVIS 327 Ficure 29. Vejovis gravicaudus, new species. Dorsal view of holotype. per group, anterior two in each group largest. One large, erect bristle lateral to posterior margin of each median eye; diad slightly more than % carapace width at that point. Carapace surface covered with large granules. Mesosoma: All dorsal plates covered with large granules, segment 7 with two pairs of lateral keels with large dentate granules; lateral margin of segment 7 abruptly flattened horizontally and with serrate granules. Sternites relatively smooth and lustrous; one pair of obsolete keels on last sternite. Stigma long oval. Metasoma: Dorsal and dorsolateral keels strongly developed and irregularly serrate and end in enlarged terminal tooth (except for dorsolateral keel of seg- ment IV which ends in broad wing) on segments IV, dorsal keels of V com- posed of broadly arranged granules. Lateral keels present and crenulate on posterior % of segment I, smooth to crenulate on posterior % of II, smooth on posterior % of III, absent on IV, and obsolete on V. Inferior lateral and inferior median keels obsolete on all segments. Position of inferior median keels marked with three pairs of short reddish hairs on each segment. Inferior lateral and median keels of segment V serrate. Telson: Ventral side with numerous, short, brownish hairs. Vesicle basically smooth but with numerous punctate depressions, and small subtle subaculear tubercule. Pectines: Long and thick; large subcircular to triangular fulcra; 25 pectinal teeth. Inferior surface with short, reddish hairs especially dense on fulcra, middle lamellae, and anterior margin. CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH Serr. wW bo oo Ficure 30. Vejovis gravicaudus, new species. Ventral view of holotype. Genital operculum: Completely divided longitudinally; large genital papillae visible externally. Chelicerae: Inferior border of movable finger completely lacking denticles; terminal tooth of superior border of movable finger subequal to terminal tooth of inferior border. Pedipalps: Palm swollen, keels obsolete, entire palm smooth and lustrous. Fixed finger and movable finger each distinctly shorter than carapace. Internal margin of fingers not distinctly scalloped; teeth do not extend to proximal end of fingers. Proximal teeth do not meet in last scallop when chela closed. Standard measurements and photographs: Table 11 and figures 29 and 30. DESCRIPTION OF ALLOTYPE. Morphologically the same as holotype with the following exceptions: larger in all body dimensions; pectine with about 20 percent fewer teeth (21 instead of 25); carapace and tergites not as granular; carapace slightly lustrous; pectines smaller, no genital papillae, genital oper- culum not completely divided longitudinally; ventral side of vesicle densely covered with about 40 long reddish hairs. Standard measurements: Table 11. VARIATION WITHIN PARATYPES. Study of 27 paratopotypes (13 males, 14 females) indicated little variation from the description of holotype and allotype. Males varied in total length from 26 to 56 millimeters, while females varied from 43 to 64 millimeters. Most individuals of both sexes appeared to be ma- ture, young instars were conspicuously absent. Mature females had a slightly wider and heavier mesosoma than mature males. Males and females similar in appearance with the following exceptions: female pectines smaller and with VoL. XXXVIT] WILLIAMS: NEW SPECIES OF VEJOVIS 329 TABLE 11. Measurements (in millimeters) of Vejovis gravicaudus, new species, holotype and allotype. Holotype Allotype (male) (female) Total length 59.7 65.1 Carapace, length Uc? 8.2 width (at median eyes) Sy? 6.3 Mesosoma, length 17.4 19.6 Metasoma, length 26.3 27.7 segment I (length/width) 3.7/4.6 Sell! Dail segment II (length/width) 4.2/4.5 4.3/5.0 segment III (length/width) 4.4/4.5 4.5/5.0 segment IV (length/width) 5.8/4.6 6.1/5.3 segment V (length/width) 8.2/4.6 9.1/5.6 Telson, length 8.8 9.6 Vesicle (length/width) 5.8/3.7 6.6/4.6 depth 2.8 BS Aculeus, length 3.0 3.0 Pedipalp Humerus (length/width) 5.8/1.8 O5)//Pal Brachium (length/width) 6.4/2.3 UP Dll Chela (length/width) 9.7/3.0 11.4/2.7 depth 3a 2.8 movable finger, length 6.5 7.0 fixed finger, length 4.8 5.4 Pectines teeth (left/right) 25/25 yD fewer teeth; males with genital papillae and with completely divided genital operculum; female vesicle hirsute, male vesicle bald in appearance. Pectine teeth varied in number from 19 to 23 (predominantly 20 and 21) in females and from 24 to 27 in males. There was a tendency for the dark stripes of the inferior metasomal keels to approach obsolescence on segments I and II. Smaller individuals had these stripes slightly darker than larger individuals. Largest individuals of each sex (presumably mature instars) had the telson with slight rose base color, smaller individuals had telson colored same as metasoma base color. There was some degree of individual variation in the expression of the underlying dusky pigmentation of the dorsum—some individuals with this ob- solescent, others with it more prominent. TYPE DATA AND ETYMOLOGY. The holotype, allotype, and 27 paratopotypes were collected 21.4 miles west of Los Aripes, elevation about 900 feet, 25 July 1968 by S. C. Williams, J. Bigelow, and M. Bentzien. The holotype and allo- type are permanently deposited in the California Academy of Sciences. 330 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 31. Type locality and habitat of Vejovis gravicaudus, 21.4 miles west of Los Aripes, Baja California Sur, Mexico, 26 July 1968. This species is named “gravicaudus” because of its relatively robust meta- soma. MateriAL. In addition to the holotype, allotype, and 27 paratopotypes, 170 additional specimens were studied. These came from the following locations in Baja California Sur, Mexico: 4 miles west of La Purisima on bank of Rio de La Purisima, elevation 375 feet, 1 July 1968 (S. C. Williams, M. A. Cazier, and party), 3 males, 3 females; 5 miles southwest of San Miguel de Comondu, elevation 1000 feet, 2 July 1968 (S. C. Williams, M. A. Cazier, and party), 37 males, 27 females; 5 to 10 miles southwest of San Miguel de Comondu, eleva- tion 1000 feet, 3 July 1968 (S. C. Williams, M. A. Cazier, and party), 24 males, 24 females; 75 miles northwest of La Paz, elevation 200 feet, 4 July 1968 (S.C. Williams, M. A. Cazier, and party), 7 males, 8 females; 31 miles west of Los Aripes, elevation 800 feet, 25 July 1968 (S. C. Williams, M. Bentzien, and J. Bigelow), 6 males, 17 females; 35 miles northwest of Los Aripes, 27 July 1968 (S. C. Williams, M. Bentzien, and J. Bigelow), 2 males, 2 females; 15 miles northwest of Los Aripes, 27 July 1968 (S. C. Williams, M. Bentzien, and J. Bigelow), 5 males, 5 females. CoMMENT. This species was collected in the southern half of the peninsula in rocky, volcanic habitats (fig. 31). It was found only in inland areas, and was conspicuously absent in similar habitats near the coast. This species was most abundant in samples taken in the bottom of canyons and washes where VoL. XXXVIT] WILLIAMS: NEW SPECIES OF VEJOVIS 331 outcrops of volcanic rock and moderately packed soils were adjacent to each other. All specimens were collected by means of ultraviolet detection. DISCUSSION AND CONCLUSIONS The 11 new species described here double the known species of Vejovis from Baja California, Mexico. It is very likely that most of these new species will prove to be endemic to this peninsula. Several of the new species have close relatives in California, Arizona, or mainland Mexico but several are so different that no close relatives have been found yet. It is interesting that some species, most notably Vejovis pumilis, were found to inhabit only soils in a region of very recent geologic origin. This would indicate that at least some species of scorpions may be genetically plastic and ecologically quite adaptable. The species belonging to the genus Vejovis appear to be uncommonly diverse in Baja California and also very abundant. LITERATURE CITED Haptey, N. F., anp S. C. WmxLIAMsS 1968. Surface activities of some North American scorpions in relation to feeding. Ecology, vol. 49, no. 4, pp. 726-734. WILLIAMS, S. C. 1968a. Methods of sampling scorpion populations. Proceedings of the California Academy of Sciences, 4th ser., vol. 36, no. 8, pp. 221-230. 1968b. Scorpion preservation for taxonomic and morphological studies. Wasmann Journal of Biology, vol. 26, no. 1, pp. 133-136. 1968c. Scorpions from Northern Mexico: Five new species of Vejovis from Coahuila, Mexico. Occasional Papers of the California Academy of Sciences, no. 68, pp. 1-24. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XXXVII, No. 9, pp. 333-362; 13 figs.; 1 table. May 15, 1970 REMARKS ON SOME GECKOS FROM SOUTHWEST ASIA, WITH DESCRIPTIONS OF THREE NEW FORMS AND A KEY TO THE GENUS TROPIOCOLOTES By Sherman A. Minton Department of Microbiology, University of Indiana Medical Center and Steven C. Anderson California Academy of Sciences, San Francisco and Jeromie A. Anderson P.O. Box 121, Quetta, West Pakistan INTRODUCTION Systematic treatment of the geckos inhabiting the vast arid and semiarid region between northwestern Africa and western India involves numerous prob- lems at both the genus and species level. During the past decade, collections made by the authors and others have resulted in the rediscovery of two geckos previously known only from the type descriptions and have indicated the need to describe three new taxa. Blanford’s genus Bunopus was reinstated by Strauch (1887) because of the tuberculate and denticulate nature of the subdigital lamellae of B. tuberculatus Blanford and B. blanfordi Strauch. Leviton and S. C. Anderson (1963) again resurrected this generic name for species with dorsal tubercles usually included in 1333] 334 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Alsophylax. Owing to early confusion in the literature, the type species of Alsophylax, Lacerta pipiens Pallas, had long been considered a tuberculate species by most workers. Actually, as Bedriaga (1912) showed, it is a nontuberculate form. Species with enlarged dorsal tubercles and tuberculate, pectinate subdigital lamellae are: Bunopus tuberculatus, B. blanfordi, B. crassicauda, and a fourth recently described species, B. abudhabt. Bunopus spatulurus John Anderson has been tentatively reassigned to a monotypic genus (Leviton and S. C. Anderson, 1967). Tuberculate species with smooth nontuberculate nonpectinate lamellae are Alsophylax loricatus, A. microtus, A. prezwalskii, A. spinicauda, and A. tibet- anus. Alsophylax pipiens and A. laevis are nontuberculate species with smooth, nonpectinate, nontuberculate subdigital lamellae. Further comments on the status of these two nominal genera are made by Mertens (1965) and Leviton and S. C. Anderson (1967). Final disposition is deferred until the Southwest Asian gecko- nid genera are subjected to general review. ACKNOWLEDGMENTS We wish to thank the following individuals for permission to examine ma- terial in their care: Dr. J. Eiselt, Naturhistorisches Museum, Wien; Dr. Charles M. Bogert and Dr. Richard G. Zweifel, American Museum of Natural History; Dr. Robert F. Inger and Mr. Hymen Marx, Field Museum of Natural History; Dr. Donald W. Tinkle, Museum of Zoology, University of Michigan; Dr. James A. Peters, U. S. National Museum. We are indebted to Dr. Alan E. Leviton, California Academy of Sciences, for comments on the manuscript and for taking the photographs used in figures 2 and 8-13. Miss Linette Sabre, Department of Herpetology, California Academy of Sciences, made the drawings in figures 4—7. We are particularly grateful to Mr. Robert Tuck, assistant in the Herpetology Section of the U. S. National Museum, for calling to our attention the series of Tropiocolotes helenae which he and John Neal collected in southwestern Iran. The following abbreviation symbols are used in the text: AMNH_ American Museum of Natural History CAS California Academy of Sciences FMNH Field Museum of Natural History NMW Naturhistorisches Museum, Wien RSM Royal Scottish Museum SAM Sherman A. Minton UMMZ University of Michigan Museum of Zoology USNM _ United States National Museum Bunopus crassicauda, described by Nikolsky in 1907 from the western part of the central plateau of Iran and known only from three syntypes, has recently been collected near Tehran. It is herewith redescribed. Vor. XXXVIT] MINTON ET AL.: ASIAN GECKOS 335 50 55 easean AS \ ate y Ie Hamadan es [IR aASsIN IRA gee Soleyman ach an oc id Zard i oe op Saas = i , @Mirjawa Soren rN --~ a ~ Dehak Dadu AA PERSIAN Y__ a eo Sehwan x re Pip Bela Diwa Wa GULF tS 7 e 20 er, a £ oO Karachi de ae KILOMETERS FiGuRE 1. Southwest Asian localities mentioned in text. Bunopus crassicauda Nikolsky. (Figure 2.) Bunopus crassicauda NiKousky, 1907, Ann. Mus. Zool. Acad. Imp. Sci. St. Pétersbourg, vol. 10 (1905), pp. 261-264, pl. 1, fig. 2 (type locality: Iran: Irak-Adschemi Province: Kum [Qom]; Maljat-Abad; Chara-Magommed-Abad. Syntypes: Zoological Institute Lenin- grad 10230, 10345, 10233). WERNER, 1936, Festschrift Strand, vol. 2, p. 200 (listed). S. C. ANDERSON, 1963, Proc. Calif. Acad. Sci., 4th Ser., vol. 31, p. 476 (listed). MATERIAL EXAMINED (2). SAM 930, AMNH 99663, Iran: Tehran Province, 15 miles southwest of Rey, collected 5 August 1965 by Sherman A. Minton, G. Possehl, W. Fairservis, and J. Fairservis. Dracnosis. A pair of postmentals in contact behind the pentagonal mental. Margins of subdigital lamellae pectinate. DEscrRIPTION. Rostral slightly broader than high, partially cleft dorsally; nostril bordered by rostral, first supralabial and three small scales; scales of snout and top of head subequal, juxtaposed; 6 to 8 supralabials, all anterior to posterior edge of eye; 8 or 9 infralabials, second touching anterior postmental; 336 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. mental shield-shaped, followed by a pair of large postmentals in contact with each other and a second smaller pair of postmentals widely separated; scales of mental and lateral gular region larger than those of median gular region. Vertical diameter of ear opening about 2/3 vertical diameter of eye. Dorsal scales small, juxtaposed, granular, separating about 13 irregular rows of enlarged tubercles. Ventral scales larger than dorsals, imbricate, smooth, not sharply differentiated from lateral scales, in 23—26 rows across mid-venter. Forelimb reaches to anterior margin of eye; hind limb reaches almost to axilla. Digits cylindrical, not angulate between last and next-to-last phalanx. Subdigital lamellae in single row with margins pectinate: 14-16 lamellae on underside of fourth toe. Basal portion of tail with segments each consisting of four whorls of imbricate scales, those of the most distal whorl being slightly larger than the others. This arrangement is not so evident in the distal half of the tail. No femoral or preanal pores or enlarged preanal or femoral scales; no enlarged platelike subcaudal scales. Color in life pale brownish gray with darker transverse bands on body and tail; five bands on body counting one in nuchal and one in femoral region; nine dark bands on tail about equal to interspaces. Both specimens apparently are adult females, snout-vent lengths 42 and 45.5 mm.; tail of larger specimen 44 mm. complete and unregenerated. Bunopus crassicauda has the same type of dorsal tubercles and subdigital 930 a a —_—, Ficure 2. Bunopus crassicauda (SAM 930). VoL. XXXVIT] MINTON ET AL.: ASIAN GECKOS 337 lamellae as B. tuberculatus, B. blanfordi and B. abundhabi, a recently discovered species from the Persian Gulf coast of Arabia (Leviton and S. C. Anderson, 1967). It differs from these forms in having two large postmentals in contact behind the apex of the mental. The two specimens of B. crassicauda were collected 5 August at the edge of a cultivated alluvial plain during mid-afternoon with the shade temperature 95° F. One animal was under a large clod of earth; the other was spotted moving under cover of small shrubs. It may, however, have been dislodged from some superficial hiding place. ; The genus Tropiocolotes (Peters, 1880) was originally established for the species “‘tripolitanus.” Subsequently the species ‘“‘steudneri” (Peters, 1869) and “natterert” (Steindachner, 1901) were added. Nikolsky (1907) established the monotypic genus Microgecko for the species “‘helenae.’’ Mertens (1956) synony- mized Microgecko with Tropiocolotes. S. C. Anderson (1963) reinstated Micro- gecko to include both “helenae” and “‘steudneri.” Minton (1962) referred a population in Sind and Las Bela, West Pakistan to Tropiocolotes helenae, and Minton and J. A. Anderson (1965) described 7. depressus from the Baluchistan mountains near Quetta. Guibé (1966b) recognized the two genera as distinct on the basis of his study of nine specimens from West Pakistan, which he identified as Microgecko helenae (herein described as a new taxon, Tropiocolotes persicus euphorbiacola) and over 50 specimens of other species assigned to Tropiocolotes. He questioned the identity of the specimen from southwestern Iran identified as M. helenae by S. C. Anderson (1961; 1963), and Tuck’s (in press) recent discussion of a series of M. helenae from southwestern Iran substantiates Guibé’s doubts. Guibé also examined Tropiocolotes depressus Minton and J. A. Anderson, and concluded that it failed to fit his generic definitions of either Tropiocolotes or Microgecko. He did not reassign it, however, pending further study. The forms Alsophylax persicus Nikolsky and Tropiocolotes scortecci Cherchi and Spano’ were not con- sidered. Guibé regarded Microgecko as a monotypic genus, and Tropiocolotes as embracing four species, 7. steudneri, T. tripolitanus, T. somalicus, and T. occiden- talis. Kluge (1967) recognized both Microgecko and Tropiocolotes in his discus- sion of familial relationships among the Gekkonidae. He stated that Microgecko has fused nasal bones and the hyoid cornu present, while in Tropiocolotes the nasal bones are paired and the hyoid cornu absent. One of us (S.C.A.), along with Dr. Alan E. Leviton, examined the hyoid apparatus of specimens of T. per- sicus euphorbiacola and T. tripolitanus algericus and found that the hyoid cornu (see Kluge, 1967, p. 37, fig. 7) is clearly present and well developed in both forms. 338 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Subsequently, Kluge examined our specimen of 7. ¢. algericus and confirmed our findings. The condition of the nasal bones is as stated by Kluge, who found it an interspecifically variable character in Cnemaspis, Cyrtodactylus, and Phyl- lodactylus. In light of present knowledge, we believe that the ends of taxonomy are better served if those forms variously referred to Microgecko and Tropiocol- otes are all united under one genus. Once the gekkonid genera of Southwest Asia have been defined adequately, it may prove desirable to include the forms here referred to Tropiocolotes under Alsophylax along with the small species from Afghanistan, northern Iran, and the USSR, perhaps recognizing subgeneric groupings. Certainly Tropiocolotes heter- opholis (herein described) appears to differ more from T. depressus, for ex- ample, than the latter differs from the gecko from Kabul, Afghanistan, referred to Alsophylax pipiens by Leviton and S. C. Anderson (1963). Such a restruc- turing of the nominal genera cannot be undertaken, however, until our knowl- edge of the northern forms is more complete, and the identity of Pallas’ Lacerta pipiens more definitely established. Tropiocolotes Peters, 1880. Tropiocolotes Peters, 1880, Mon. Berlin Akad., p. 306, pl.—, fig. 1 (type species: T. tri- politanus Peters, by monotypy). Stenodactylus BOULENGER (in part, not Fitzinger), 1885, Cat. Liz. British Mus., vol. 1, p. 16. Microgecko Nrkorsky, 1907, Ann. Mus. Zool. Acad. Imp. Sci. St. Pétersbourg, vol. 10 (1905), pp. 264-265 (type species: M. helenae Nikolsky, by monotypy). DEFINITION. A gekkonine genus (in the sense of Kluge, 1967) rarely exceed- ing 35 mm. in snout-vent length; digits slightly angularly bent (but not so strongly as are those of, for example, Cyrtodactylus, Agamura), not dilated, fringed, webbed, or ornamented, covered below with a single series of transverse lamellae, either smooth or keeled (but not tuberculate in the sense of Bunopus) ; pupil vertical; dorsal scales imbricate to subimbricate, never juxtaposed gran- ules; postanal sacs present; preanal and femoral pores usually absent (two pre- anal pores in male 7. steudneri according to John Anderson, 1898, and in T. de- pressus); postmentals usually well developed, shieldlike (except in T. depressus). The genus includes eight species distributed from Spanish Sahara across northern Africa and the Middle East to the Indian Desert. KEY TO THE SPECIES AND SUBSPECIES OF 7‘ROPIOCOLOTES la. Dorsals smooth or faintly keeled, ventrals smooth _____ = sssssSsS—SssSsS 2 1b. Dorsals and ventrals strongly keeled 2a. Scales along dorsal midline from axilla to groin 50 or less; four scales border nostril; Sulbdiritalslamellaesdistinetly. itrrcaninate, see eee en 3 Vor. XXXVIT] MINTON ET AL.: ASIAN GECKOS 339 2b. Sar 3b. 4a. 4b. 5a. 5b. 6a. 6b. hae 7b. 8a. 8b. 9a. 9b. 10a. 10b. lla. 11b. 12a. 12b. Scales along dorsal midline from axilla to groin 60 or more; four or five scales border nostril; subdigital lamellae smooth or very indistinctly tricarinate 4 Adpressed hind limb reaches elbow or nearly to axilla; range: Israel west through Egypt and Sudan to Algerian Sahara _. Tropiocolotes steudneri Peters Adpressed hind limb reaches beyond shoulder; range: eastern Sinai Peninsula, pos- SiDlyawies tell Owe by aye ts er ee T. nattereri Steindachner Internasals not differentiated from adjacent scales; four scales border nostril; post- mental shields absent, or cne small pair not in contact; males with two preanal pores; range: West Pakistan: Baluchistan mountains near Quetta sa T. depressus Minton and J. A. Anderson Internasals large, followed by a second pair of enlarged shields; five scales border nostril; postmental shields well developed; males without prenanal pores 5 A single pair of postmentals, not in contact; dark crossbars of body absent or in- distinct, sometimes two dorsolateral series of white spots; range: Iran: western footnllsvot Zagros Mountains ee T. helenae (Nikolsky) Two pairs of postmental shields; dark crossbars on tail distinct 6 Dark dorsal crossbars of body and tail broader than interspaces; range: Iran: west- ConmOOtnl SsoteZacross Mountains, = ee eee eee T. persicus bakhtiari Minton, S. C. Anderson, and J. A. Anderson Dark dorsal crossbars of body and tail narrower than interspaces _. 7 Dark crossbars less than half width of interspaces; 82 scales along dorsal midline from axilla to groin; range: Iran-West Pakistan border region ana T. persicus persicus (Nikolsky) Dark crossbars as wide or slightly narrower than interspaces; 62-76 scales along dorsal midline from axilla to groin; range: West Pakistan: coastal plain from Bela to Haleji near Tatta, upper Hab River Valley, and Nabisar in the Thar Desert east of the Indus __ 7. persicus euphorbiacola Minton, S. C. Anderson, and J. A. Anderson Dorsal scales strongly heterogeneous; range: Iraq: Salahedin (known only from the ie) T. heteropholis Minton, S. C. Anderson, and J. A. Anderson DorsalascalesmhomogeneOus) 9, ee eee 9 Postmentals not in contact; 50-52 midbody scale rows; range: Hadramaut, Arabia T. scortecct Cherchi and Spano’ ROS tmentalSeingco lita Cieseeer: kn es aes ee we BP. ae Bee eee 10 Wessaehanes Okmid bodyuscaley OWS i -<- 25 oe ee 11 Midbody scale rows 46-54 (mean 52); range: Mali T. tripolitanus subspecies! A pair of very large postmental shields in contact, followed by a second, smaller FORE ats eS ee eee eee eee ee CT es ae ee a eee 12 A single pair of large postmental shields in contact, no second pair distinguishable ELSES Gl PONE eirce See eee so a Dect hn ee ee Ee 13 Anterior postmental shields extending backwards to form a long suture with lower labial thus excluding posterior postmentals from contact with first lower labial; midbody scale rows 42-48; range: Egypt west through Libya to Tunisia _.___ T. tripolitanus tripolitanus Peters Anterior postmental shields usually not extending backwards as far as second lower labial so that posterior postmental shields are in contact with first lower labial; mid- body scale rows 35-41; range: Somalia —__ eae Ee ae T. tripolitanus somalicus Parker 1 An undescribed population from Mali appears most closely allied to T. t. tripolitanus, but having a greater number of scale rows at midbody (Ted Papenfuss, personal communication). 340 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. i3a. Postmental shields extending backwards to form a short suture with second lower labial; supraorbital scales imbricate, smaller than those on interorbital region; mid- body scale rows 44-46; range: Algerian Sahara ____ T. tripolitanus algericus Loveridge 13b. Postmental shields failing to reach the second labial; supraorbital scales imbricate, larger than those on interorbital region; midbody scale rows 40-41; range: Spanish S aa ais cera a ee eee T. tripolitanus occidentalis Parker In the above key we have followed Loveridge’s (1947) key to the subspecies of T. tripolitanus, as we have seen little north African material. In treating the forms allied to T. persicus as subspecies of this form we follow Loveridge’s treat- ment of the “‘tripolitanus” group for the sake of consistency. We do not imply that we have any knowledge of a zone of intergradation between any two of the currently recognized taxa. The distributional limits of these various forms re- main to be elaborated in all cases. The dwarf geckos of north Africa, Tropiocolotes tripolitanus and its subspe- cies, differ strikingly from the eastern forms in body lepidosis, both dorsal and ventral scales being strongly keeled. The scales are larger than in the eastern forms. Counted along the dorsal midline from the level of the axilla to a point on line with the insertion of the hind legs, they number 32 to 34 in the subspecies “algericus” and 33 to 37 in the subspecies “‘tripolitanus.” With regard to other characters studied, the subdigital lamellae are strongly tricarinate and number 14 to 15 on the fourth toe; the postrostral and internasal scales are not markedly larger than the surrounding scales; there are 6 or 7 upper labials; the anterior postmentals are large and in contact with each other; the posterior postmentals are half or more the size of the anterior pair in Lybian specimens, about one fourth as large as the anterior in Moroccan specimens. Moroccan specimens show 6 or 7 irregular transverse dark bands on the body; Lybian specimens are dark speckled. In both, the dark tail bands are wider than the interspaces sep- arating them and number 11 in the only specimen with an intact, unregenerated tail. A population related to the “tripolitanus” group has recently been dis- covered in Mali by Mr. Ted Papenfuss of the University of San Francisco. He is preparing a report on this form. Tropiocolotes scortecci Cherchi and Spano’, re- cently described from the Hadramaut of southern Arabia, appears to be closely allied to T. tripolitanus, differing from the latter in having the postmentals sep- arated by one or two scales, 50-52 scales around the middle of the body, and 15-17 lamellae under the fourth toe. > Ficure 3. A. Tropiocolotes steudneri from Wadi Raman, Negev, Israel (FMNH 74431); B. From Luxor, Egypt (SAM 809); C. From Hurghada, Red Sea Governate, Egypt (FMNH 78393); D. Tropiocolotes persicus persicus from Mirjawa, West Pakistan (SAM 937); E. Tropiocolotes depressus from Kolpur, West Pakistan, paratype (RSM 1964.58.1); F. Tropiocolotes persicus euphorbiacola from Haleji, West Pakistan (SAM 600). VoL. XXXVITI] MINTON ET AL.: ASIAN GECKOS 341 608 ms ea [Proc. 4TH SER. CALIFORNIA ACADEMY OF SCIENCES 342 Ficure 4. A. Tropiocolotes persicus bakhtiari, holotype (CAS 86408), scales of snout; B. Scales of chin. (Vertical lines represent 1 mm.) VoL. XXXVIII MINTON ET AL.: ASIAN GECKOS 343 SS RII Vy QR oe é Ficure 5. A. Tropiocolotes persicus euphorbiacola, holotype (CAS 93939), scales of snout; B. scales of chin. 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EM sooeds.19}Ul sajeos Juade[pe y0B}UOD UT 1dA3q uey} 1opIM ATV ystys WO1f PI}eIVUI IJCULIBITTY Aqyensn 4y}iOq Q spout g spol 1dduUpNnaqs ZI podojaAap Aj1o00d -layjip Aj1ood ‘T[ews 61-bI Os-8¢ ied a31e] 7 8-9 6-9 ZI $a]0]09014 04 J [IRL Apog s[eseusajuy 90} Yip Ul01s 0} s[ejuawysog sjeiqe[, sjeiqe, sSuawtoads a[aweyT elpixe -eljuy -eidng jo JaquinN sreqssoio Ye ‘QUI[PIW [BSIOp BuO[e sa[eog 344 ‘avuajey “| pup ‘snssaidap “y ‘snotsied “J, ‘iwaupnays sajopoooidory fo suoszpdmoy “ alav VoL. XXXVIT] MINTON ET AL.: ASIAN GECKOS 345 In Troptocolotes steudneri the dorsal and ventral body scales are smooth and imbricate, smaller than in T. tripolitanus, numbering 38 to 50 along the dorsal midline from axilla to groin. Subdigtal lamellae are tricarinate, numbering 14 to 19 on the fourth toe. The internasals are slightly larger than the adjacent snout scales. The posterior pair of postmentals are more than half the size of the ante- rior pair; both pairs are usually in contact with each other in the midline or the posterior pair may be just separated. Transition in size from postmentals to gular granules is gradual with 3 to 6 small scales behind the postmentals. The dorsal pattern may be one of crossbands or mottling; occasional individuals are almost unicolor. The distribution of 7. steudneri is from Egypt and northern Sudan through Sinai and most of Israel and presumably into parts of Jordan and Saudi Arabia. On the basis of material examined, there appears to be significant difference be- tween specimens from Egypt and those from Israel (see table 1 and fig. 3a-c). Being unfamiliar with the full range of variation in these areas, we prefer to call attention to the apparent differences and refer the problem to those with more material at their disposal and some field knowledge of the animals involved. Recently, Mr. Robert Tuck (in press) has recorded a series of specimens col- lected in southwestern Iran. These specimens agree in their pholidosis with Nikol- sky’s (1907) description of Microgecko helenae. Nikolsky’s type series of nine specimens has been destroyed (S. C. Anderson, 1961). Tropiocolotes helenae (Nikolsky). (Figures 6A-B, 8, 9.) Microgecko helenae Nikotsky, 1907, Ann. Mus. Zool. Acad. Imp. Sci. St. Pétersbourg, vol. 10 (1905), pp. 265-268, pl. 1, fig. 4, 4a (type locality: Iran: Khuzistan Province: Alchor- schir; Aguljaschker; Isfagan; Bidezar [Bid Zard]. Syntypes destroyed). WERNER, 1936, Festschrift Strand, vol. 2, p. 200 (listed). Tropiocolotes helenae, Tuck (in press). MATERIAL EXAMINED (20). USNM 153693-153703, 153705-153710, CAS 120795—-120796, Iran: Khuzistan Province: 35 km. east of Gach Saran. USNM 153731, Iran: Khuzistan Province: 16 km. south of Masjed Soleyman. Dracnosis. Dorsal scales homogeneous, smooth, subdigital lamellae smooth; a single pair of postmental shields, not in contact with one another; back with or without indistinct, undulating dark transverse bars, narrower than interspaces; back sometimes uniform, sometimes with two dorsolateral series of white spots; tail with narrow dark transverse bars narrower than interspaces and bordered posteriorly with white; regenerated portion of tail uniform black. REMARKS. See table 1 for further comparison with other taxa. Examination of a recently collected specimen of Tropiocolotes from the Iran- Pakistan border region indicates that it is identifiable with Alsophylax persicus (Nikolsky, 1903). We accordingly reassign this species to Tropiocolotes, the 346 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. REE TAN ry see Ficure 6. A. Tropiocolotes helenae (CAS 120795), scales of snout; B. Scales of chin. (Ver- tical lines represent 1 mm.) MINTON ET AL.: ASIAN GECKOS 347 VoL. XX XVII] ut; B. Scales s, holotype (AMNH 93003), scales of sno 7. A. Tropiocolotes depressu present 1 mm.) of chin. (Vertical lines re FIGURE 348 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER. LTTE ETE EEE Ficure 8. Tropiocolotes helenae (USNM 153693), showing original tail. name “‘persicus” apparently being the oldest one available for members of the genus hitherto referred to ‘“Aelenae” by Mertens (1956), S. C. Anderson (1961, 1963), Minton (1962, 1966), and Guibé (1966a, 1966b) (not Nikolsky, 1907). Tropiocolotes persicus persicus (Nikolsky), new combination. (Figure 3d.) Alsophylax persicus NrkoutskKy, 1903, Ann. Mus. Zool. Acad. Imp. Sci. St. Pétersbourg, vol. 8, p. 95 (type locality: Iran: Degak [Dehak], in the region of Dizak. Holotype: Zoologi- cal Institute Leningrad 10005. WerNeER, 1936, Festschrift Strand, vol. 2, p. 200 (listed). ? Tropiocolotes helenae, MERTENS, 1956, Jh. Ver. Naturk. Wirttemberg, vol. III, pp. 92—93 (Iran: 20 km. SW Pip. Synonymy of Microgecko with Tropiocolotes). Bunopus persicus, S. C. ANDERSON, 1963, Proc. Calif. Acad. Sci., 4th ser., vol. 31, p. 474 (listed; new combination, in error). ? Microgecko helenae, GuiBr, 1966, Bull. Mus. Nat. Hist. Natur., ser. 2, vol. 38, p. 98 (Iran: 100 km. N. Iranshar). MATERIAL EXAMINED (1). SAM 937, 2, West Pakistan: Chagai District: Mirjawa, near the Iranian border, collected 9 June 1965, by Jeromie A. Anderson. Dracnosis. Dorsal scales homogeneous, smooth; subdigital lamellae smooth; dark transverse dorsal bars of body narrow, less than half the width of inter- spaces, and with distinct light margins; tail bars narrower than interspaces. Vor. XXXVIT] MINTON ET AL.: ASIAN GECKOS 349 MEET Lill biiatiibe Ficure 9. Tropiocolotes helenae (USNM 153699), showing black regenerated portion of tail. DEscriPTION. Rostral broader than high, partially divided by midline dorsal furrow; nostril bordered by rostral, first supralabial and three small scales; inter- nasals large, broadly in contact with each other and with rostral; behind them another pair of enlarged shields, also in contact; 10 supralabials; 8/7 infralabials, second barely touching anterior postmental; mental shield-shaped with long pos- terior projection; anterior postmentals large, separated by single scale behind point of mental and followed by second much smaller pair of postmentals; re- 350 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER. mainder of mental and gular region covered with small scales of approximately equal size. Ear opening an oblique slit less than half vertical diameter of eye. Dorsal scales small, subequal, smooth, weakly imbricate, 82 along dorsal mid- line from axilla to groin; ventral scales similar to dorsals in size and arrangement. Digits cylindrical, toes with slight angulation between last and next to last phalanx. Subdigital lamellae smooth, margins not pectinate; 15 lamellae on underside of fourth toe. No femoral or preanal pores. Scales of tail and limbs similar to those of body. Dorsal ground color in life lemon-yellow; five dark transverse bars on body, the first on the neck and the fifth just anterior to the hind limbs, each band much less than half width of interspace between it and next band and bordered pos- teriorly by distinct whitish band at least half its width. Nine dark bands on tail, much narrower than interspaces, extreme tip of tail black. Dark stripe from tip of snout through eye and along side of head and neck to level of second dorsal band. Limbs without dark markings; venter white. Measurements of the specimen in millimeters are as follows: snout-vent 30.7; tail 32.6; head (to angle of jaw) 7.8; eye 1.9; snout 3.3; forelimb 11.3; hind limb 11323: Remarks. The fact that five, rather than four, scales border the nostril dis- tinguishes T. persicus and its subspecies along with T. helenae from all other members of the genus. The uppermost nasal shield (internasal) is large and in contact with its fellow on the midline of the snout. In the other species of the genus, a pair of small internasals lie posterior to the rostral and do not border the nostril. Smaller body scales, smooth subdigital lamellae, lack of contact between the second pair of postmentals, and abrupt transition in size from post- mentals to small granular gulars further distinguishes this form from 7. steud- nert. The narrow dark transverse bars with relatively wide light posterior bord- ers distinguish this subspecies from T. depressus, T. p. bakhtiari, and T. p. eubhorbiacola. The presence of a second pair of postmentals distinguishes it from T. helenae. This specimen represents a rediscovery of Nikolsky’s species known pre- viously only from the tvpe. Mertens’ (1956) specimen which he tentatively identi- fied as Nikolsky’s Microgecko helenae may belong to this taxon, as may Guibé’s from southeastern Iran. We have been unable to examine these specimens. Wett- stein’s (1960) record of Cyrtodactylus persicus from Kabul is based on the form referred to Alsophylax pipiens by Leviton and Anderson (1963). We have veri- fied this by examination of his specimen (NMW 15972). The specimen identi- fied as Alsophylax persicus Reed and Marx (1959) represents an undescribed form treated elsewhere in this paper. The known range of Tropiocolotes p. persi- cus occupies western Baluchistan in the region of the Iran-Pakistan boundary. VoL. XXXVIT] MINTON ET AL.: ASIAN GECKOS 351 A specimen from Khuzistan Province, southwestern Iran, was previously identified in error as Microgecko helenae Nikolsky (S. C. Anderson, 1961, 1963). Because of the many similarities of this specimen to T. persicus (Nikolsky), we propose to treat it as a subspecies of the latter. Tropiocolotes persicus bakhtiari* Minton, S. C. Anderson, and j. A. Ander- son, new subspecies. (Figures 4A-B, 10.) Tropiocolotes helenae (not Microgecko helenae Nikolsky, 1907), S. C. Anderson, 1961, Was- mann Jour. Biol., vol. 19, pp. 287-289 (Iran: Khuzistan Province: between Masjid-i- Suleiman [Masjed Soleyman] and Sar-i-Gach. Remarks on synonymy of Microgecko with Tropiocolotes). Microgecko helenae (not Nikolsky, 1907), S. C. ANpErson, 1963 Proc. Calif. Acad. Sci., 4th ser., vol. 31, pp. 440-441, 474, fig. 10 (notes on collection, photograph). Hototype. CAS 86408, adult male (?), between Masjid-i-Suleiman (Masjed Soleyman) and Sar-i-Gach, Khuzistan Province, Iran, collected 13 May 1958, by S. C. Anderson. Dracnosis. A subspecies of 7. persicus in which the dark transverse bars of body and tail are broader than the interspaces separating them; anterior pair of postmentals in contact. For additional points of comparison see table 1. DESCRIPTION OF HoLotyPeE. Habitus moderately depressed; head length (tip of snout to angle of jaw) contained 3 1/4 times in snout-vent length; length of snout greater than distance between posterior corner of eye and angle of jaw; diameter of eye slightly greater than distance betwen orbit and nostril; ear open- ing oval, about 1/5 diameter of eye; rostral pentagonal, 1 2/3 times broader than high, cleft in upper half of its height, bordered by first supralabials, nostrils, two postrostrals, and a small scale at its apex. Nostril surrounded by five scales, including rostral, first supralabial, enlarged postrostral, and two additional small nasal scales; enlarged postrostrals seperated by small scale in contact with apex of rostral, and followed by additional pair of enlarged scales, also separated in mid- line by small scale; this second pair of enlarged scales slightly larger than post- rostrals. Snout covered with small juxtaposed granules slightly larger than those covering top and sides of head and occiput, granules on side of snout somewhat larger than those on midline. Twenty-two scales across top of head between centers of eyes, 10 supralabials on right, 9 on left, posterior labials distinctly larger than succeeding small granules, but not as sharply set off as in T. helenae; 7 infralabials. Mental pentagonal, sharply pointed behind, about 1 1/3 times longer than broad, followed by pair of trapezoidal postmentals, larger than first infralabial, in contact, and forming short suture behind mental; second smaller 2 Named for the Bakhtiari tribe, which gives its name to the general region of Khuzistan in which this gecko was found. Ww On bo CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. pair (greatest length about 1/3 greatest length of first pair) lies between first postmentals and posterior portion of second infralabials; scales of chin and throat granular, juxtaposed to subimbricate, subequal to scales on sides of head, and distinctly larger than scales on occiput. Dorsum covered with equal, smooth, imbricate small scales; scales of venter 1 1/2 times larger than those of dorsum, smooth, flat, imbricate; scales of upper surfaces of limbs similar to those of back, scales of lower surfaces of hind limbs like those of venter; no femoral or preanal pores. Tail slender, tapering to point, covered above and below with smooth scales, larger than those of dorsum, arranged in regular transverse series, but not ar- ranged to give distinctly segmented appearance to tail, scales of lower surface larger than those of dorsal surface of tail. Two enlarged (but not tubercular nor pointed) scales on either side of swelling at base of tail just posterior to level of vent. Digits covered above with small, imbricate, smooth scales, below with single series of lamellae, weakly tricarinate (in sense of obtusely bent, not sharply keeled as in 7. tripolitanus and T. heteropholis; perhaps due to preservation, as suggested by Guibé, 1966b). Toe somewhat angularly bent, but not distinctly so as in Cyrtodactyvlus and Agamura. Color in life (S. C. Anderson, 1961): Cream-colored on dorsal surfaces with four chocolate-brown transverse bars in area between shoulders and pelvis, bars broader than interspaces, heaviest pigmentation on posterior margin of each bar; dark brown bar on nape, posterior corners of which extend back and down to meet next crossbar, anterior corners extending forward, meeting dark marking which lies just posterior to eye and over ear opening. Eleven dark crossbars on tail. Limbs, snout, and labials lightly dusted with brown. Ventral surfaces im- maculate creamy white. Remarks. Examination of Tuck’s series of Tropiocolotes helenae empha- sizes the differences between this specimen and Nikolsky’s original description of Microgecko helenae. CAS 86408 has ten and nine upper and seven lower labials, while 7. helenae has six or seven upper and five or six lower labials. The pos- terior labial shields are more readily distinguished from the following small shields in T. helenae than in T. p. bakhtiari. Nikolsky describes “helenae” as having a single pair of postmental shields, not in contact, the condition in Tuck’s series, while T. p. bakhtiari has two pairs, the anterior pair in contact. The tail of CAS 86408 is thin, while Nikolsky’s and Tuck’s specimens have thick tails. The dif- ference, however, is undoubtedly due to the emaciated condition of the specimen of T. p. bakhtiari, as specimens of T. p. euphorbiacola from West Pakistan (see below) starved before preservation show similarly thin tails. The caudal scales are more conical, the free margins projecting out from the tail to a greater extent in T. helenae than in T. p. bakhtiari, possibly also an artifact of the poor condi- Vor. XXXVIT] MINTON ET AL.: ASIAN GECKOS 353 # 96408 Ficure 10. Tropiocolotes persicus bakhtiari, holotype (CAS 86408). tion of the latter. The abdominal scales are proportionately larger than the dor- sals in T. helenae than in T. p. bakhtiari (50-55 from level of axilla to groin in the former species, about 62 in the latter). In 7. helenae the pair of scales pos- terior to the enlarged postrostrals on the midline of the snout are smaller than the postrostrals, while in 7. p. bakhtiari they are larger. The crossbars in T. p. bakh- tiart are much broader than the interspaces, while in Tuck’s series of T. helenae, the back lacks crossbars while those of the tail are narrower than the interspaces; Nikolsky’s illustration shows the dorsal crossbars to be narrower than the inter- spaces. Measurements (in mm.) of the present specimen are as follows: snout-vent 28.0; tail 28.5; head (to angle of jaw) 7.8; eye 2.2; snout 2.8; forelimb 11.1; hind limb 16.1. The position of the nostril and the nature of the scales of the snout are similar to the specimen of Microgecko persicus persicus. The scales of the chin differ only in that the anterior postmentals of CAS 86408 are in contact. There are 71 scales along the dorsal midline from axilla to groin. The fourth toe has 12 subdigital lamellae. No preanal or femoral pores. The dark transverse bars of the back are five in number, the first on the neck, the fifth just anterior to the hind limbs. They are 12 scales wide, the inter- spaces 7 or 8 scales wide. The posterior two scale rows of each bar are darkest. The population of Tropiocolotes inhabiting Sind and eastern Las Bela, al- though closely resembling T. p. bakhtiari, is separated from the known range of that subspecies by approximately a thousand miles with part of the intervening territory occupied by P. persicus. These zoogeographical considerations add weight to the observed morphological differences between the populations and, in our opinion, justify nomenclatural recognition of the Sind population. Because 354 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. the characteristic habitat of this small gecko is clumps of the giant Euphorbia caducifolia, we propose the name: Tropiocolotes persicus euphorbiacola Minton, S. C. Anderson, and J. A. Anderson, new subspecies. (Figures 3F, 5A—B, 11.) Tropiocolotes helenae (not Nikolsky, 1907), Minton, 1962, American Mus. Novitates, no. 2081, p. 11, fig. 39; 1966, Bull. Amer. Mus. Nat. Hist., vol. 134, p. 81, pl. 15, fig. 1. Hototype. CAS 93939, adult male, Lower Pab Hills, Hab Chowki, Las Bela District, West Pakistan, about 350 feet elevation, collected 27 December 1962, by Jeromie A. Anderson. PaRATYPES (26). CAS 93933-93938, 93940-93951, same data as holotype; CAS 99871, 99873-99878, Hab Chowki, Las Bela District, West Pakistan, col- lected 25 March 1965, by Jeromie A. Anderson; UMMZ 122007 (2 specimens), 7 miles northwest of Karachi, Karachi District, West Pakistan, collected 20 De- cember 1958, by Sherman A. Minton. Dracnosis. A subspecies of 7. persicus in which the dark dorsal bands on the body are as wide as, or slightly narrower than the interspaces between them but are always more than half the width of the interspaces. The tail, when regen- erated, is white (yellow in life), without dark transverse bars. For additional points of comparison see table 1. DESCRIPTION OF HOLoTyYPE. Scales of snout as in other two subspecies; supra- labials 8/9; infralabials 8; anterior postmentals in contact with each other; pos- terior postmentals less than half size of anterior and not in contact. Scales of body, limbs and tail similar to those of other two subspecies; 74 scales along dorsal midline from axilla to groin; 14 smooth lamellae on underside of fourth toe; no preanal or femoral pores. Five dark transverse bands on body, anterior border of each being slightly lighter than postanterior; bands in middle of back 8-12 scales wide and separated by interspaces of 10-12 scales; 8 dark bars on tail, narrower than interspaces, extreme tip of tail black. Dark stripe from tip of snout through eye and along side of head, neck, and body to level of third dorsal band. Measurements (in millimeters) are as follows: snout-vent 28.5; tail 32.5; head to angle of jaw 7.7; eye 2.0; snout 2.9; forelimb 8.8; hind limb 13.7. Remarks. In the paratype series, there are 7 to 9 supralabials, 6 to 8 infra- labials, 62 to 75 scales along the midline of the back from axilla to groin and 12 to 15 lamellae under the fourth toe. Examination of eight specimens from locali- ties in Dadu, Tatta and Thar Parkar District of Sind does not significantly ex- tend the range of variation (see table 1). The dorsal ground color in life is usually chrome yellow but varies from pale straw to amber. The anterior edges of the bands are irregular and lighter in color, often tending to blend into the ground VoL. XXXVIT] MINTON ET AL.: ASIAN GECKOS 355 \ ‘ FicureE 11. Tropiocolotes persicus euphorbiacola, holotype (CAS 93939) from Lower Pab Hills, Hab Chowki, Las Bela District, West Pakistan. color; the posterior edges are dark, straight and often edged by a narrow white line. Specimens preserved soon after capture have thick, rather depressed tails, while individuals kept in captivity and not feeding well soon utilize the fat stored in this appendage. The tails of such animals are thin and more cylindri- cal. This dwarf gecko is abundant in the coastal plain from the vicinity of Bela to Haleji near Tatta. Peripheral localities are the Lakhi Hills south of Sehwan, Diwana in the upper Hab River valley, and Nabisar in the Thar Desert east of the Indus. In the Lakhi Hills it has been collected at an elevation of about 700 feet. These lizards are most readily found in the dry, decayed stems and roots of Euphorbia caducifolia after the clump has died and collapsed, forming a low mound. They probably also frequent the living clumps but are difficult to ob- serve among the closely spaced thorny stems. When exposed the geckos try to escape into the plant debris by wriggling the body and tail, making little use of the limbs. They may also flee in agile leaps and bounds, this escape behavior being particularly characteristic when they are surprised in the open at night. Their locomotion in a terrarium is slow and deliberate with bobbing and twisting movements of the head and shoulders when peering about and sinuous movements of the body and tail. Captive individuals show arboreal tendencies, but this has not been observed in the field. Mature adults are usually found in pairs. A single egg, about 8 X 5.7 mm., is laid in decayed hollow roots, in crevices, or beneath rocks from April to August. The egg, soft when laid, hardens in a few hours. It is elongated at the ends and 356 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. glossy white with longitudinal striations that disappear in about four weeks as the egg enlarges slightly. Hatching requires five to eight weeks. The hatchlings are 13-15 mm. in snout-vent length and are marked similarly to the adults. In the terrarium, these lizards lap water from a watch glass and feed in- discriminately on arthropods of suitable size, for example, newly hatched spiders and small termites. One captive ate more than 20 small termites in an hour. The tail is very easily broken, and about a third of the lizards examined have regenerated tails. Tropiocolotes depressus Minton and J. A. Anderson. (Figures 3E, 7A-B.) Tropiocolotes depressus MINTON AND J. A. ANDERSON, 1965, Herpetologica, vol. 21, pp. 59-61, fig. 1 (type locality: West Pakistan: Quetta Division: Kach). Minton, 1966, Bull. America Mus. Nat. Hist., vol. 134, p. 82, pl. 15, fig. 2 (description, key, habitat, pho- tograph). REMARKS. Since collecting the two original specimens of this species, one of us (J. A. A.) collected three additional specimens on the eastern slope of Chiltan Mountain, 17 miles south of Quetta and 14 miles north of Mastung 30-31 Octo- ber 1967. These were collected at about 14.30 from between large granite slabs at about 8500 feet elevation on a 45° hillside, very arid, where the only plants are small shrubs and grasses. They were found while digging out the Afghan rock pika, Ochotona r. rufescens, from under rocks two to eight cubic meters in size. A male and female were collected together under one rock. These geckos move much more rapidly than do those of Tvropiocolotes persicus euphoribacola from Sind and Las Bela. When alarmed they assume a “serpentine” position for a few seconds before commencing a steady jerky advance, coming back to a serpentine position at each halt. They also climb rock faces. Daytime temperatures in this area reach about 15° C. at this time of year and get down to about —4° at night. This and similar areas have been investi- gated in the same manner in the summer without encountering these geckos. Numerous small potential predators are present in the immediate vicinity, including scorpions (Androctononus australis and Buthus occitanus); solpugids (Galeodes caspius) ; centipedes (Scolopendra species); young snakes (Coluber rhodorhachis, C. karelini, Vipera lebetina obtusa, and Echis carinatus); lizards such as Eumeces schneideri blythianus; Baluch shrews; and hedgehogs (Hemi- echinus megalotus ). The series of three specimens was sent to Professor Robert Mertens at Senck- enberg, Frankfurt-am-Main. In examining specimens previously identified as Alsophylax persicus, we found a highly distinctive small gecko that appears to be undescribed. We pro- pose for it the name: VoL. XXXVIT] MINTON ET AL.: ASIAN GECKOS 357 Tropiocolotes heteropholis Minton, S. C. Anderson, and J. A. Anderson, new species. (Figure 12.) Alsophylax persicus (not Nikolsky, 1903), REED AND Marx, 1959, Trans. Kansas Acad. Sci., vol. 62, p. 97 (Iraq: Erbil Liwa: Salahedin Nahiya: Salahedin: Pirman Hotel). HorotyPe. Field Museum of Natural History 74549, immature female, Iraq; Erbil Liwa: Salahedin Nahiya: Salahedin: Pirman Hotel, 1080 meters elevation, collected 1954-1955 by Charles A. Reed. Diacnosis. A species of Tvropiocolotes with strongly carinate subdigital lamellae and strongly keeled dorsal scales of highly variable size. DESCRIPTION OF HoLotyPe. Snout longer than distance between eye and ear opening; rostral nearly twice as broad as high, cleft above; nostril between ros- tral, first supralabial, and two nasal shields; supranasal shields separated by single internasal; scales of snout and top of head subequal, strongly keeled, sub- imbricate; scales of occipital and temporal region heterogeneous, strongly en- larged scales intermixed with smaller ones; 12/11 supralabials, posterior two or three strongly keeled; seven infralabials; mental pentagonal, pointed behind; pair of large, roughly pentagonal postmentals broadly in contact behind mental, followed by smaller but well developed pair; diameter of eye shorter than dis- tance between eye and nostril or eye and ear opening; ear opening vertical oval, diameter equal to about 1/3 diameter of eye. Scales of back imbricate, strongly keeled, heterogeneous, enlarged scales in about 12 more or less regular longitudinal rows, separated by much smaller scales of diverse sizes. Scales of venter smaller than enlarged dorsals, uniform, imbri- cate, strongly keeled, keels forming regular longitudinal lines; about 53 scales round middle of body and 38 along dorsal midline from axilla to groin. Forelimb reaches to center of eye, and 2/3 distance to groin; hind limb reaches to axilla. Forelimb covered above with more or less homogeneous small, keeled scales; hind limb covered above with heterogeneous keeled scales, strongly enlarged scales intermixed with smaller. Digits cylindrical, covered above with uniform keeled scales, below with single series of transversely broadened lamellae, each with three to five strong keels, ends of which form mucronations. Third toe longest, second and fourth about equal, 15 lamellae under fourth toe. Tail covered above with heterogeneous keeled scales, greatly enlarged scales arranged in regular longitudinal and transverse series, not in contact; covered below with smaller, unequal keeled scales, no enlarged subcaudal plates. No preanal or femoral pores. All scales of head, body, limbs, and tail strongly keeled, with exception of rostral, mental, postmentals and labials. Unfortunately, the specimen is rather strongly stained with rust. There ap- pear to be five narrow dark bars on the body and alternating light and dark bars 358 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 12. Tropiocolotes heteropholis, holotype (FMNH 74549) from Salahedin, Salahedin Nahiya, Erbil Liwa, Iraq. of about equal width on the tail. The melanophores are contracted and more numerous on the dorsal than on the ventral surface. Measurements (in mm.): snout-vent 20.4; tail 19; head (to angle of jaw) 6.7; eye 1.4; snout 2.3; forelimb 6.6; hind limb 8.8. Remarks. The affinities of 7. heteropholis appear to be with T. tripolitanus, although 7. hketeropholis differs from all other members of the genus in having strongly heterogeneous dorsal scales. According to Reed and Marx (1959), the specimen was collected under a bit of bark on the cement floor of the back porch of a hotel, and may have come in with a load of scrub-oak firewood. This specimen clearly does not belong to Alsophylax persicus Nikolsky as originally identified. Alsophylax persicus is a nontuberculate, barred gecko with uniform, smooth scales (see above). Reed and Marx (1959) state that this speci- men can be distinguished readily from Gymnodactylus heterocercus Blanford by the keeled ventral scales and the fact that the forelimb does not reach the tip of the snout. Only the latter is a valid distinction, as no mention of the nature of the ventral scales is made by Blanford (1876). While Blanford states that his specimens of Gymnodactylus heterocercus were gray throughout, with markings, his illustration (pl. 22, fig. 3) shows a gecko with six indistinct dusky transverse bars. Blanford’s description, with the exception of the length of the forelimb, is insufficient to clearly separate G. heterocercus from the lizard described here. Wettstein (1951) had two specimens from Hamadan, Iran, the type locality of G. heterocercus, and which he identified as such. These had dark violet-gray bars on the dorsum. No mention is made of the ventral scales or the nature of the sub- digital lamellae. We have not been able to examine Blanford’s types of Gymnodactvlus hetero- cercus, which are in the Turin Museum. We append herewith a redescription of VoL. XXXVIT] MINTON ET AL.: ASIAN GECKOS 359 Gymnodactylus heterocercus based upon specimen number 7286 in the Vienna Museum. Cyrtodactylus heterocercus (Blanford). (Figure 13.) Gymnodactylus caspius (in part; not Eichwald, 1831), Dr Firippr, 1865, Viagg. in Persia, p. 352. Gymnodactylus heterocercus BLANFORD, 1874, Ann. Mag. Nat. Hist., ser. 4, vol. 13, pp. 453- 454 (type locality: Iran: Hamadan; two syntypes: Turin Museum). 1876, Eastern Persia, pp. 345-347, pl. 22, fig. 3. Diacnosis. Dorsum with enlarged trihedral tubercles; no enlarged subcaudal plates, tail covered below with strongly keeled, imbricate scales in 5—7 longitud- inal rows; sides of tail with greatly enlarged, sharply keeled, mucronate scales. DESCRIPTION OF NMW 7286 (Iran: Hamadan, collected June 11, 1950 by H. Loffler). Habitus depressed; snout 1 1/3 times distance between eye and ear opening; rostral 1 1/2 times broader than high, cleft above, upper portion curved backward onto surface of snout; nostril between rostral, first upper labial, and three nasals; separated by small internasal; scales of snout and top of head some- what heterogeneous in size; juxtaposed tubercular, smooth to weakly keeled or conical; temporal and occipital region with scattered keeled or conical tubercles; 9/10 supralabials, 7 infralabials; mental pentagonal, pointed behind; a pair of large, unequally hexagonal postmentals broadly in contact behind mental, followed by two smaller, but well developed pairs, the second partially, and the third en- tirely separated from infraliabials by smaller scales; diameter of eye slightly less than distance between eye and nostril or eye and ear opening; ear opening sub- circular, about 1/4 diameter of eye. Scales of back imbricate, most smooth, a few weakly keeled; enlarged strongly keeled, tubercular, weakly trihedral scales in 14 regular longitudinal rows (outermost row smallest); enlarged scales about as broad as wide, much larger than interspaces. Scales of venter smaller than dorsal tubercles, uniform, imbricate, weakly keeled (keels most noticeable on outer rows); about 59 scales round middle of body, about 87 along dorsal midline from axilla to groin (not counting enlarged tubercles). Forelimb reaches nostril or tip of snout, 2/3 distance to groin; hind limb reaches axilla. Upper arm covered above with equal, keeled, imbricate scales, lower arm with scattered enlarged, keeled tubercles; hind limbs with numerous, greatly enlarged trihedral, mucronate, keeled tubercles. Both fore and hind limbs covered below with keeled homogeneous scales. Digits cylindrical, or even slightly depressed at base, terminal phalanges compressed; fingers and toes strongly angularly bent, claws (at least in this specimen) very short, scarcely projecting beyond surrounding ungual scales; subdigital lamellae smooth, in single trans- versely broadened series. Fourth toe longest, bearing 21 subdigital lamellae. 360 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 13. Cyrtodactylus heterocercus (NMW 7286). Tail covered above with heterogeneous keeled scales, enlarged tubercular scales arranged in regular verticils, those immediately on either side of midline being smaller than enlarged tubercles of back, outer three on either side being progressively enlarged, very sharply keeled, mucronate, largest being nearly as long as diameter of eye; tail covered below with small, imbricate, sharply keeled scales in 5—7 longitudinal rows, not arranged in distinct verticils, outermost row enlarged, tubercular though smaller than lateral tubercles. No preanal or femoral pores (specimen is apparently a female). About 8 indistinct dark angular crossbars on dorsum, apices pointing cau- dad, first bar on neck, 8th across sacral region; these tend to fuse and coalesce; 13 dark bars on tail. REMARKS. There is a pronounced cutaneous fold from axilla to groin; this may be an artifact of preservation, as the specimen is not particuarly well pre- served. Although we have seen neither Gymnodactylus danilewsky Strauch 1887, nor G. colchicus Nikolsky 1902, the affinities of Cyrtodactylus heterocercus would appear to be with these forms, judging from the descriptions (Nikolsky, 1915). Terentjev and Chernov (1949) considered both G. danilewsky and G. colchicus only subspecifically distinct from Cyrtodactylus kotschyi. Cyrtodactylus heter- ocercus is distinguished from the typical form of C. kotschyi by the lack of smooth enlarged subcaudal plates, and from G. danilewsky by lack of enlarged internasals and the nature of the very large lateral tubercular caudal scales VoL. XX XVII] MINTON ET AL.: ASIAN GECKOS 361 (compare with Nikolsky, 1915, text-fig. 19, and Terentjev and Chernov, 1949, text-fig. 57). In the descriptions of G. danilewsky and G. colchicus, no mention is made as to whether the subcaudal scales are smooth or carinate. The latter form should be examined to determine whether or not it may be identical to Cyrtodactylus heterocercus. Cyrtodactylus russowi (Strauch) has smooth sub- caudal scales. LITERATURE CITED ANDERSON, JOHN 1898. Zoology of Egypt, vol. 1 Reptila and Batrachia. London, Ixv + 371 pp., 50 pls. ANDERSON, STEVEN C. 1961. A note on the synonymy of Microgecko Nikolsky with Tropiocolotes Peters. Was- mann Journal of Biology, vol. 19, pp. 287-289. 1963. Amphibians and reptiles from Iran. Proceedings of the California Academy of Sciences, 4th ser., vol. 31, pp. 417-498. BEDRIAGA, J. VON 1912. Amphibia and Reptilia. In: Nauchenui Rezul’ tatui puteshestvii N. M. Przheval’ skagho po tsentral’noi Azii .. . vol. 3, St. Pétersbourg, vi + 769 pp., 10 pls. BLANFORD, WILLIAM T. 1876. Eastern Persia, an account of the journeys of the Persian Boundary Commission, 1870-1872. Vol. 2. The zoology and geology. London, viii + 516 pp., pls. 14-28. CHERCHI, MARIA ADELAIDE, AND SILVIO SPANO’ 1964. Un nuova specie di Tropiocolotes del Sud Arabia Spedizione Scortecci nell’Hadra- mut (1962). Bollettino dei Musei e degli Istituti Biologici dell’Universita di Genova, vol. 32 (1962-1963), pp. 29-34, 4 figs. GuIBE, JEAN 1966a. Reptiles et amphibiens récoltés par la Mission Franco-Iranienne. Bulletin du Mu- séum National d’Histoire Naturelle, ser. 2, vol. 38, pp. 97-98. 1966b. Contribution a l’étude des Microgecko Nikolsky et Tropiocolotes Peters (Lacertilia, Geckonidae). Bulletin du Muséum National d’Histoire Naturelle, ser. 2, vol. 38, pp. 337-346. KiucE, ARNOLD G, 1967. Higher taxonomic categories of gekkonid lizards and their evolution. Bulletin of the American Museum of Natural History, vol. 135, pp. 1-60, pls. 1-5. Leviton, ALAN E., AND STEVEN C. ANDERSON 1963. Third contribution to the herpetology of Afghanistan. Proceedings of the California Academy of Sciences, 4th ser., vol. 31, pp. 329-339. 1967. Survey of the reptiles of the Sheikhdom of Abu Dhabi, Arabian Peninsula. Part II. Systematic account of the collection of reptiles made in the Sheikhdom of Abu Dhabi. Proceedings of the California Academy of Science, 4th ser., vol. 35, pp. 157-192. LOovERIDGE, ARTHUR 1947. Revision of the African lizards of the family Gekkonidae. Bulletin of the Museum of Comparative Zoology, vol. 98, pp. 1-469, pls. 1-7. MERTENS, ROBERT 1924. Ein neuer Gecko aus Mesopotamia. Senckenbergiana, vol. 6, p. 84. 1956. Amphibien und Reptilien aus S.O.-Iran, 1954. Jahreshefte des Vereins fur vater- landische Naturkunde in Wiirttemberg, vol. 111, pp. 90-97. 362 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. 1965. Bemerkungen iiber einige Eidechsen aus Afghanistan. Senckenbergiana Biologica, vol. 46, pp. 1-4. MINTON, SHERMAN A., JR. 1962. An annotated key to the amphibians and reptiles of Sind and Las Bela, West Pak- istan. American Museum Novitates, no. 2081, pp. 1-60. 1966. A contribution to the herpetology of West Pakistan. Bulletin of the American Museum of Natural History, vol. 134, pp. 29-184, pls. 9-36. Minton, SHERMAN A., JR., AND JEROMIE A. ANDERSON 1965. A new dwarf gecko (Tropiocolotes) from Baluchistan. Herpetologica, vol. 21, pp. 59-61. Nrixkotsky, A. M. 1903. On three new species of reptiles collected by Mr. N. Zarudny in eastern Persia in 1901. Annuaire du Musée Zoologique de l’Academie Impériale des Science de St. Pétersbourg, vol. 8, pp. 95-98. 1907. Reptiles et amphibiens recueillis par M. N. A. Zarudny en Perse en 1903-1904. An- nuaire du Musée Zoologique de l’Academie Impériale des Sciences de St. Péters- bourg, vol. 10 (1905), pp. 260-301, pl. 1 [in Russian]. 1915. Faune de la Russie. Reptiles. Petrograd, vol. 1, 532 pp., 9 pls. [in Russian; see 1963 translation by L. and E. Kochva, Israel Program for Scientific Translations Ltd., Jerusalem, cat. no. 838, 352 pp.|. PASTEUR, GEORGES 1960. Redecouverte et validité probable du Gekkonidé Tro piocoltes nattereri Steindachner Comptes Rendus des Séances Mensuelles, Societé des Sciences Naturelles et Phy- siques du Maroc, no. 8, pp. 143-144. ReeEpD, CHARLES A., AND HyMEN MARX 1959. A herpetological collection from northwestern Iraq. Transactions of the Kansas Academy of Sciences, vol. 62, pp. 91-122. STRAUCH, ALEXANDER 1887. Bemerkungen iiber die Geckoniden-Sammlung in zoologischen Museum der Kaiser- lichen Akademie der Wissenschaften zu St. Pétersbourg. Memoirs de l’Academie Impeéria!e des Sciences de St. Pétersbourg, ser. 7, vol. 35, 72 pp., 1 pl. TERENTJEV, PAUL V., AND SeRGIUS A. CHERNOV 1949. Opredelitel Presmykaiushchikhsia i Zemnovodnykh. Moscow, 3rd ed., 340 pp. [in Russian; see 1965 translation by L. Kochva, Israel Program for Scientific Trans- lations Ltd., Jerusalem, cat. no. 1255, 315 pp.] WETTSTEIN, OTTO 1951. Ergibnisse der Osterreichischen Iran-Expedition 1949/1950, Amphibien und Repti- lien. Versuch einer tiergeographischen Gliederung Irans auf Grund des Repti- lien-Verbreitung. Sitzungsberichten der Osterreichische Akademie Wissenschaf- ten Wien. Mathematisch-naturwissenschaftliche, vol. 160, pp. 427-448. 1960. Drei seltene Eidechsen aus Siidwest-Asien. Zoologischer Anzeiger, vol. 165, pp. 190— 193. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XXXVII, No. 10, pp. 363-380; 5 figs.; 5 tables May 15, 1970 BIOSYSTEMATICS OF THE CANYON TREE FROG HYLA CADAVERINA COPE (= HYLA CALIFORNIAE GORMAN) By Robert W. Ball San Diego Mesa College and David L. Jameson University of Houston; Research Associate, Department of Herpetology, California Academy of Sciences INTRODUCTION This paper reviews the taxonomy of the canyon tree frog, Hyla cadaverina, and describes its geographical variation. Baird and Girard (1852, p. 174) described the Pacific tree frog, Hyla regilla, from collections made on the Sacramento River, in Oregon, and in Puget Sound by the U. S. Exploring Expedition under the command of Captain Charles Wilkes. Hallowell described Hyla scapularis (1852, p. 183) from Oregon collections and Hyla scapularis var. hypochondriaca (1854) from Dr. A. L. Heermann’s collections from Tejon Pass in southern California. Hallowell (1854) also described a second species from his analysis of Dr. Heermann’s Tejon Pass collection as being distinctive from Hyla scapularis and used the name Hyla nebulosa as the species designa- tion. Baird and Girard (1852, p. 301) referred to Hla scapularis as a syn- onym of Hyla regilla after examining specimens collected by Dr. John L. LeConte in San Francisco. These early collections disclosed the presence on the Pacific Coast of two distinct species, Hla regilla and Hyla nebulosa. Cope (1866, p. 84) substituted Hyla cadaverina for the preoccupied Hyla nebulosa (Spix, 1824). Yarrow (1882) and Cochran (1961) list the type [363] lh = —- ae | 364 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. specimen of Hyla cadaverina as USNM 3230. However, Jameson (1966) notes that USNM 3230 is from Fort Vancouver, Washington, and assigned this specimen to Hyla regilla. Cope (1889) states that two specimens of Hyla collected by Dr. Heermann from Tejon Pass, southern California, are located in the Museum of the Academy of Natural Sciences of Philadelphia. Test (1898) attempted to rectify the confusion in the general taxonomy of Hyla regilla, reviewed the taxonomic history of the species, and lumped Hyla nebulosa and Hyla cadaverina as synonyms of Hyla regilla. Test noted that the specimen of Hyla nebulosa which he examined was collected by Dr. A. L. Heermann and that, of the two originally collected by Dr. Heermann, only one (USNM 3230) still existed in the museum collection. Cope (1889) described a specimen of Hyla nebulosa (= Hyla cadaverina) that was collected from Fort Tejon, California, under the synonymy of Hyla regilla var. regilla and described the specimen of Hla nebulosa as one of the three color variations. The specimen was in poor condition and was said to be ash gray, with dorsal blotches wanting, and similar to Hyla squirella. Cope questioned its synonymy with Hyla regilla. The specimens described by Cope and Test are obviously not any of the cotypes originally described by Hallowell as Hyla nebulosa (Jameson, et al. 1966). Storer (1925) and Slevin (1928) also place Hyla nebulosa and Hyla cadaverina in synonymy with Hyla regilla. Baird (1854) described Hyla affinis from the northern Sonoran region and Cope (1866), noting the preoccupancy of Hyla affinis (Spix, 1824), substituted Hyla arenicolor as a synonym. Yarrow (1882) and Cope (1889) refer to two specimens of Hla arenicolor from the collections of H. W. Henshaw in 1875 from southern California. Richardson (1912) described Hyla arenicolor and listed specific locales of several collections in southern California in which this species was found. Wright and Wright (1949), Stebbens (1951), and others noted differences between Hyla arenicolor collections from east and west of the Mojave and Imperial deserts and Gorman (1960) separated the western populations of Hla arenicolor into a separate species based on morpho- logical and ethological comparisons with the eastern populations. The western populations were designated by Gorman as Hyla californiae. The type specimen was an adult female, number 31,773, in the University of California Museum of Vertebrate Zoology collected at Canyon de Llanos, 9 miles (14.5 km.) SSW. of “Alaska” (La Rumorosa), Partido del Norte, Baja California, Mexico. The historical survey of the literature and the examination of the type specimens reveal that the following names have been used to describe the “canyon tree frog” inhabiting the range from San Luis Obispo County, central California, to approximately latitude 29° north, Baja California, Mexico: Hyla nebulosa Hallowell, 1854; Hyla cadaverina Cope, 1866; Hyla regilla Cope, 1889; Hyla arenicolor Yarrow, 1882; Hyla californiae Gorman, 1960. We submit that the first valid name describing the canyon tree frog from VoL. XXXVII] BALL & JAMESON: CANYON TREE FROG SYSTEMATICS 365 the Pacific Coast is Hyla cadaverina Cope. Two specimens of Hyla both listed as number 3230 in the Museum of the Academy of Natural Sciences of Phil- adelphia, appear to be the two cotype specimens of Hyla nebulosa described by Hallowell. We have examined the qualitative and quantitative morphological variation of the available specimens and we have studied the nonmorphoiogical and ecological characteristics of the several populations in order to determine whether or not biologically distinguishable races exist. MATERIALS AND METHODS MEASUREMENTS. A total of nine individual structural measurements were taken from 225 males representing 26 distinct populations and 72 females representing 10 distinct populations. Vernier calipers were used to measure snout-vent length, head width, forearm length, and shank length. A Bausch and Lomb dissecting microscope with an ocular micrometer insert was used to measure nostril-lip height, finger pad width (third digit), length of the fourth toe, and webbing development (between the fourth and fifth toe). The measurements were recorded in millimeter units. The localities sampled (fig. 1), locality codes, and the number of individuals measured from each population sampled are given in table 1. An analysis of morphological measurements of Hyla cadaverina has shown a highly integrated system of positively correlated characters (Ball and Jame- son, 1966). The morphological characters selected for this study reflect total body shape and therefore form a discriminate complex for the differentiation of various localities. MATHEMATICAL ANALYSIS. Multivariate statistics, which treats the total variation of (n) characters within and between groups around a grand mean on (n) characteristic vectors, were used to discriminate morphologically between the 26 populations sampled. The biological advantages of the use of multi- variate statistical techniques are comprehensively discussed by Jolicoeur (1959), Cooley and Lohnes (1962), and Jameson ef al. (1966). The variation of characters within the group was calculated by the within group covariance matrix W. The dispersion of group means was determined by the between group covariance matrix B. The characteristic vectors (discriminant functions) are the principal axes of matrix B after the standardization by matrix W. The standardization of matrix B by matrix W corrects the population positions on the discriminant graphs by adjusting each measurement by the variances of the other measures allowing such variants as age and size to be minimized and the interpopulational differences to be emphasized. The length of the resultant vector from the grand mean expresses the maximum directional displacement of a specific character by specific populations. The data were programmed for and processed by the IBM 1620 computer. The 366 CALIFORNIA ACADEMY OF SCIENCES [PRroc. 4TH SER. Ficure 1, Localities of Hyla cadaverina populations sampled. Table 1 gives locality codes. characteristic vectors are coded as follows on the discriminant graphs: V1, snout-vent length; V2, head width; V3, head length; V4, nostril-lip height; V5, forearm length; V6, finger pad width; V7, shank length; V8, length of fourth toe; V9, degree of webbing between fourth and fifth toe. We appreciate the loan of specimens of Hyla cadaverina from the following institutions: University of California Museum of Vertebrate Zoology, Cali- fornia Academy of Sciences, Stanford University Natural History Collection, San Diego Museum of Natural History, and the San Diego State College Re- search Collection. We have examined specimens from and extend our thanks to the appropriate staff members at the Los Angeles County Museum of Natural History, University of Southern California, University of California at Los Angeles, American Museum of Natural History, Academy of Natural Sciences of Philadelphia, Museum of Comparative Zoology, Harvard University, Uni- VoL. XXXVII] BALL & JAMESON: CANYON TREE FROG SYSTEMATICS 367 TaBLE 1. Geographic localities sampled, locality codes, and the number of specimens measured from each population. Number of individuals Forearm length measured "Shank length Locality Code Females Males Males Pine Valley, San Diego County A 5 406 DeLuz, San Diego County B 7 399 Coyote Creek, San Diego County Cc 5 391 Forty-nine Palms Canyon, San Bernardino County D 10 A24 Agua Caliente Springs, San Diego County E 5 416 La Puerta, San Diego County F 6 15 406 El Capitan, San Diego County G 10 5 410 Tubbs Springs, San Diego County H 4 9 404 Sentenac Canyon, San Diego County I 8 A12 Ojos Negros, Baja California Norte J 11 414 Mesquite Springs, Valle Trinidad, Baja California Norte K 10 11 A418 Borrego Palm Canyon, San Diego County L 6 11 416 Little Morongo Canyon, San Bernardino County M 8 409 La Crescenta, Los Angeles County N 9 383 Mojave River (West Fork), San Bernardino County O 7 402 Sespe, Ventura County P 6 390 Tujunga River, Los Angeles County Q 11 394 Canyon de Llanos, Baja California Norte R 6 423 Fillmore, Ventura County S 11 389 San Jose, Baja California Norte wD 8 A21 El Cajon Canyon, Baja California Norte U 9 445 Aqua Dulce, Baja California Sur V 8 10 425 Ensenada, Baja California Norte W 11 420 Soboba Springs, Riverside County xX 6 A36 Bahia de los Angeles, Baja California Sur WY 6 11 436 Indian Cove, San Bernardino County Z, 10 428 San Dimas Canyon, Los Angeles County $ 6 Boulder Creek, San Diego County & 8 Whitewater Canyon, Riverside County ! 8 Totals 72 225 versity of Michigan Museum of Zoology, Chicago Natural History Museum, and the United States National Museum. E. G. Bauer, D. Bacon, and M. Anderson assisted with the computation at the San Diego State College Com- ‘puter Center. R. C. Jameson performed the illustrations assisted by the com- puter. Financial assistance for parts of this study has come from the San Diego State College Foundation, San Diego Natural History Museum, California Academy of Sciences, National Science Foundation (G-15558), and the National Institutes of Health (RG-8489). 368 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. TaBLte 2. Character grand means, standard deviations from the grand mean, and population means of the 26 male populations sampled. Characters V1 V2 V3 V4 V5 V6 V7 V8 V9 Grand Means 31.80 11.98 9.62 2.46 6.89 1.44 16.55 6.54 3.63 Pooled Standard Deviation from the Grand Mean SoZ 0.62 0.30 0.03 0.24 0.29 1.14 0.23 0.15 Population Means Population A 30.84 11.52 8.76 2.36 6.68 1.42 16.44 6.16 3.32 B 32.61 12.23 9.30 2.59 6.86 1.67 17.19 6.86 3.70 C 33.96 12.84 9.82 2.60 7.02 1.70 17.94 6.82 3.86 D 33.86 13.07 10.05 2.67 7.18 1.52 16.92 6.81 3.87 E 33.04 12.86 9.78 2.54 7.16 1.78 tiene 6.90 3.64 F 32.29 12.10 9.76 2.54 6.92 1.65 17.05 6.73 3.65 G 34.18 12.54 10.10 2.50 7.28 1.64 17.76 6.86 3.84 H 30.31 10.91 9.02 Dron 6.41 1.46 15.88 6.16 3.50 I 30.71 11.44 9.08 2.31 6.61 1.43 16.01 6.23 3.65 i] 31.24 11.78 9.44 Prov 6.93 1.50 16.72 6.47 3.61 K 33.41 11.93 9.79 2.66 7.16 1.66 7/1) 6.86 3.90 L 34.42 12.83 10.57 2.67 7.45 1.63 17.91 6.95 3.99 M 30.50 11.00 8.95 DS) 6.44 1.23 67/8! 5.80 3.50 N 29.38 11.19 9.02 2.29 6.10 1.32 15.52 6.10 3.66 O 27.94 10.95 8.79 2.29 6.04 1.21 15.03 5.90 3.69 P 29.92 11.18 9.20 Deo 6.32 1.40 16.20 6.52 Sai/s3 Q 28.85 11.08 8.90 22s 5.88 1.39 14.92 5.86 SBy// R 34.12 12.85 10.52 2.58 7.48 1.52 17.67 7.28 S/H) S 28.83 11.04 9.06 2.16 6.15 12 15.80 6.37 3.59 als 28.85 10.89 9.06 2.24 6.29 1525 14.91 5.76 3.43 U 31.58 12.04 9.94 2.41 ele E32 16.12 6.34 3.80 V 36.41 14.25 11.15 2.76 8.23 13H 19.38 7.48 4.10 W 31.95 11.89 9.85 2.36 6.99 1.39 16.63 6.71 3.63 xX 28.67 11.10 9.08 2.25 6.92 1.30 15.88 6.25 S50 We S175 2 et2 10.15 2.47 6.83 1.10 15.66 6.15 2.28 Z B5),7/1 1355) 10.38 2.89 7.66 1.57 17.91 7.12 3.91 DESCRIPTION For gross morphological and ethological descriptions of Hyla cadaverina, we refer the reader to Hallowell (1854), Wright and Wright (1949), Stebbins (1951), Gorman (1960), and Ball and Jameson (1966). The tadpole of Hyla cadaverina is described by Gaudin (1964, 1965). We deliberately limit our description to new material and to that necessary to document our systematic conclusions. VoL. XXXVII] BALL & JAMESON: CANYON TREE FROG SYSTEMATICS 369 TaBLeE 3. Character grand means, standard deviations from the grand mean, and population means of the 10 female populations sampled. Characters V1 V2 V3 V4 V5 V6 V7 V8 V9 Grand Means 38.25 14.03 11.16 2.74 8.35 1.76 19.70 7.72 3.41 Pooled Standard Deviation from the Grand Mean 4.71 0.97 0.46 0.04 0.32 0.29 1.60 0.18 0.14 Population Means Population G 37.91 13.52 10.75 2.70 8.70 1.75 19.73 7.65 3.44 H 39.60 13.85 11.15 2.70 7.95 2.00 20.27 8.20 3.40 V 42.43 15.90 12.01 3.06 9.49 17S 21.70 8.10 3.95 F 36.90 14.50 11.16 2.62 8.47 1.68 19.90 7.65 3.42 L 39.83 14.96 11.73 DiS 8.78 1.90 20.66 7.80 3.40 & 34.55 13.02 10.26 2.50 7.73 1.64 18.45 6.96 2.89 K 36.73 13.05 10.29 2.65 7.80 1.67 18.44 7.55 3.23 $ 38.23 14.10 11.15 2.85 8.18 1.82 20.05 8.10 3.82 ny: 39.78 14.45 12.58 2.93 8.82 1.68 19.66 7.68 3.21 ! 36.58 13.02 10.52 2.68 7.60 Le 7/l 18.16 7.50 3.30 The color patterns of Hyla cadaverina are highly variable. The mountain forms generally are dark in color and have large melanophore patches which blend into the granite rock patterns along the streams. The coastal and desert forms usually display diffuse melanophore patterns and are generally lighter in color than the montane forms. The diffusion of coloration on the dorsal surface of many of the coastal and desert forms appears to correlate with the prevalent native stone within their habitats. The character grand means, standard deviations from the grand means, and populations means of the 26 populations of males and 10 populations of females sampled in this study are given in tables 2 and 3. An examination of the grand means of the male and female populations sampled reveals the sexual dimorphic characters of Hyla cadaverina. The snout-vent measurement is the best overall measure of size (Ball and Jameson 1966), and the female is 6 to 7 mm. mean larger than the male in snout-vent length. The females are larger in all of the measurements taken except for the degree of webbing. DISCRIMINATORY ANALYSIS. Male populations. The sum of the variance components of the first nine discriminant axes was 1572.98 (X*° with 225 degrees of freedom), which is highly significant (table 4). The first two discriminant axes contain 59.6 percent (36.9 + 22.7) of the total variation within and between the populations. The multivariate dispersion plots on the 370 CALIFORNIA ACADEMY OF SCIENCES | Proc. 47H SER. K5= 22.7 2s e xT Y e -10 U e e @ WwW p s e i I 23 - 9 y : V 8 A 7 ° : 6 =) 2 D ye Gale Kis 2i6a e e F e : B © 10,x 12Z 14 s e e Ficure 2. Discriminant analysis of 225 male canyon tree frogs. The mean of each sample is shown on the first (Ki = 36.9%) and second (Kz = 22.7%) discriminant axes. The original coordinates are represented by the direction of each vector emanating from the grand mean. The length of the vector is representative of one standard deviation of the measurement. The measurements are labeled as indicated in the text. Localities are shown in figure 1. K,—Ky discriminant axes (fig. 2) demonstrates a general north-south (left to right) geographical distribution. The samples from the geographically isolated populations at Agua Dulce (V) and Bahia de los Angeles (Y) appear as isolates, occurring on the right portion of the plot. Both of these populations (V and Y) are among geographical isolates occupying the southern extremes of the dis- tribution of Hyla cadaverina. The populations on the right half of the dispersion plot are separated into a distinct group by generally having larger head dimensions (V2 and V3), longer forearms (V5), and increased length of the fourth toe (V8). Larger finger pad widths (V6), shank lengths (V7), and increased webbing (V9) char- acterize the populations on the left half of the dispersion plot. Samples from populations at E] Capitan (G), Agua Caliente (E), and Borrego Palm Canyon (L) display a larger relative snout-vent length (V1) and nostril-lip height (V4). The plots of the third and fourth discriminant axes (fig. 3) contain 18.1 percent of the total variation and demonstrate some of the characters by which closely grouped populations on the K,—K» plot differ. A shorter shank length Vor. XXXVIT] BALL & JAMESON: CANYON TREE FROG SYSTEMATICS 371 Z e K ,-8.6 Ficure 3. Discriminant analysis plot of third (Ks; = 9.5%) and fourth (Ki = 8.6%) discriminant axes of males. See figure 2 for labels. (V7) and increased webbing (V9), a longer snout-vent length, (V1) and a higher nostril-lip height (V4) differentiate the population at Bahia de Los Angeles (Y) from that at Agua Dulce (V). Ojos Negros (J) differs from populations Agua Caliente (E) and Coyote Creek (C) by demonstrating a longer snout-vent length (V1), head length (V3) and a small finger pad width (V6). DISCRIMINANT ANALYsIS. Female populations. An examination of the K,—Kzs discriminant axes of the female populations sampled (fig. 4) displays a population distribution similar to the male K,—Kz discriminant plot. The total variance of the female plot is 438.54, significant at the 1 percent level (table 5). The plots of the first (variance 178.99, 40.8 percent) and the second axes (variance 90.13, 20.5 percent) again demonstrate that the southern isolates Agua Dulce (V) and Bahia de los Angeles (Y) are isolated on the plot. The sample from Tubbs Springs (H) appears along with Whitewater Canyon (!) Mesquite Springs (K) and San Dimas Canyon ($). These populations can ios) 72 CALIFORNIA ACADEMY OF SCIENCES [PRroc. 4TH SER. Ka220'5 Ficure 4. Discriminant analysis of plot of first (Ki = 40.8%) and second (Kz = 20.5%) discriminant axes of 72 female Canyon tree frogs. See figure 2 for labels. be distinguished by having longer fourth toe lengths (V8) and larger finger pad widths (V6). Longer shank lengths (V7) and shorter nostril-lip height (V4) discriminate frogs from El Capitan (G) and Boulder Creek (&) from the other populations. Animals from Bahia de los Angeles (Y) Aqua Dulce (V) and La Puerta (F) are characterized by having larger facial dimensions (V2 and V3), increased webbing (V9), and longer forearms (V5). Taste 4. The percent of significant information and the characteristic roots (variance component) of the nine morphological measurements taken from 26 male populations. Discriminant Variance Percent Degrees Axis Component of Total of Freedom Probability 1 581.26 36.9 33 < 0.01 2 357.19 DoT 31 << 0:01 3 150.28 9.5 29 < 0.01 4 136.90 8.6 27 < 0.01 5 110.71 all 25 < 0.01 6 79.41 540) 23 <(O}{0)u! 7 76.67 4.8 21 < 0.01 8 SSeval 3.3 19 < 0.01 9 26.85 1.6 7) > 0.05 Vor. XXXVII] BALL & JAMESON: CANYON TREE FROG SYSTEMATICS 373 Ky>8.7 2 at: 3 4 : K3:18.0 e 5 : -2 : Kee 3 ] Ficure 5. Discriminant analysis of plot of third (K; = 18.0%) and fourth (Ki = 8.7%) discriminant axes of females. See figure 2 for labels. The plots of the third and fourth discriminant axes (fig. 5) contain 26 percent of the total variation. A longer head length (V3) along with reduced webbing (V9) differentiates frogs from Bahia de los Angeles (Y) from those from Agua Dulce (V). Populations at Tubbs Springs (H) and San Dimas Canyon ($) differ from those at Mesquite Springs (K) and Whitewater Canyon (!) by having larger head widths (V2). TABLE 5. The percent of significant information and the characteristic roots (variance component) of the nine morphological measurements taken from 10 female populations. Discriminant Variance Percent Degrees Axis Component of Total of Freedom Probability 1 178.99 40.8 17 < 0.01 2 90.13 20.5 15 < 0.01 3 79.31 18.0 13 < 0.01 4 38.51 8.7 11 < 0.01 5 32.21 7.3 4 < 0.01 6 14.62 3) 7 < 0.01 7 4.77 1.0 5 > 0.05 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. WwW “I TS DISCUSSION The population dispersion within the first two discriminant axes reveals the degree of adaptive convergence and divergence occurring within Hyla cadaverina. The secular changes in the thermal environment and available moisture since the last pluvial maximum, 10,000 to 11,000 years ago, can assist in the interpretation of populational morphological variations and the apparent geographical isolation of the populations discriminated in the analysis. During the periods of glacial maxima the temperatures within the existing range of Hyla cadaverina were lowered and there was an increase in humidity. Savage (1960) indicates that amphibian forms with high rainfall requirements probably invaded the moist coastal regions during the several pluvial maxima of the late Pleistocene. The existing east-west and southern ranges of Hyla cadaverina may be understood as remnants of the geographical dispersion during the last pluvial maximum. As the temperatures and aridity increased during the postglacial periods the distribution of Hyla cadaverina was restricted generally north and south within the coastal mountains. Many authors have stressed the effects of postglacial cycles on the bio- geographical abundance and distribution of southwestern forms. The period of “thermal maximum” between 4000 and 8000 years ago is thought to be responsible for the isolation of southwestern desert and montane populations (Martin, 1963) and probably contributed to the development of fragmented populations of Hyla cadaverina to the east (desert) and in the high desert of the northern Baja California peninsula. Amphibians such as Bufo microscaphus, Scaphiopus hammondiu, and Hyla regilla deserticola share similar geographic ranges with Hyla cadaverina and may have undergone similar evolutionary patterns. As the temperature and aridity increased postglacially, amphibian populations became isolated in the south and eastern limits of their distribution to desert springs and cismontane within stream habitats. Eastern isolates of Hyla cadaverina (Forty-nine Palms Canyon and Indian Cove) and southern isolates sampled (Agua Dulce and Bahia de los Angeles) are limited to spring- fed oases. The southern populations have probably been isolated from the intact species population the longest since the last altithermal period and this is reflected by the discriminant separation of these populations (fig. 2). Martin (1963), using pollen records, indicates that post pluvial droughts were probably limited to the winter months and that the interglacial period was relatively wet from summer rains. The coefficients derived using populations means of forearm length/shank length suggests appendage adaptations accompanying aquatic habitat differences and isolation in desert springs areas (see table 1). The populations with relatively shorter shank lengths and larger forearm lengths (larger coefficients in table 1) are characteristically found in isolated springs within the southern Vor. XXXVII] BALL & JAMESON: CANYON TREE FROG SYSTEMATICS 375 or eastern distributional limits or along the steep canyons of the eastern Sierra slopes. The populations displaying proportionally significantly larger appendage characters, i.e., shank length, finger pad width, and toe webbing, are generally from aquatic environments requiring the ability to adapt to swift currents. The populations from more xeric habitats are proportionally larger in their snout-vent lengths, suggesting an adaptation for reduced water loss capacity. Jameson (1966), using experimental evidence from a series of desiccation experiments with Hla regilla, indicated that organismal size was not sufficiently explained by latitudinal or elevational distribution. Genetic characters adapting Hyla cadaverina both morphologically and physiologically appear to have been playing a role in its evolution. Montane and coastal forms are usually smaller than their more xeric counterparts. The distribution of Hyla cadaverina suggests that it has adapted to a variety of weathers. The mountain forms are subjected to extreme temperature ranges and have been vocally recorded at air tempratures of 7°C. Several high desert populations have been observed calling in late June and early July with air temperatures around 30°C. High elevational forms must hibernate in winter months to avoid freezing temperatures and desert forms must aestivate in late summer and fall or face desiccation. The primary differentiation in western canyon tree frogs since the last pluvial period appears to have been within the xeric populations. There is little doubt that many of the populations of the desert regions are genetically isolated from those in the mountains. Obviously animals in one mountain range are spatially isolated from those in other ranges. While we can generally recognize that a given collection is from the desert or from a particular mountain range, the translation of the cues to a key or to a workable description has been intractable. Additionally, we have not discerned significant differences in the life history of the animals from the several habitats. We conclude that the isolated populations are continuing to evolve in parallel paths as a result of the common genetic background and the restriction to a relatively narrow ecological niche. We refrain from describing subspecies of Hyla cadavenia for we feel that to do so at this stage of knowledge would merely obscure biological truths. (Jameson, et al., 1966, p. 552.) The evolutionary relationship of Hyla cadaverina to the other southwestern species of tree frogs has not been firmly established. Gorman (1960) suggests Hyla arenicolor as the probable sibling or parent species of Hyla cadaverina. The present distribution of Hyla arenicolor is from Texas to middle Arizona and including northern Mexico. Contact between Hyla cadaverina and Hyla arenicolor could have been established during the last pluvial period. The adult morphology of both Hyla cadaverina and Hyla arenicolor is very similar to Hyla regilla. Gaudin (1965) states that larval characters, especially larval 376 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. mouth parts, of Hyla regilla and Hyla cadaverina more closely resemble each other than either does Hla arenicolor. The calls suggest that Hyla cadaverina and H. regilla are closely related. Discriminant analysis of the relationships of North American Hyla reveals a closer morphological relationship between Hyla arenicolor and Hyla cadaverina that either have to Hyla regilla. Hybridiza- tion experiments using H. regilla, H. cadaverina, and H. arenicolor fail at very early stages. Conclusive assumptions of the evolutionary relations of south- western species of Hyla await further study. SYNONYMY Hyla cadaverina, (Cope). Hyla nebulosa, Hallowell, 1854. Proceedings of the Academy of Natural Sciences of Philadelphia, vol. 7, p. 96; type description; type locality, Tejon Pass (California). Hyla cadaverina, Cope, 1866. Journal of the Academy of Natural Sciences of Philadelphia, second series, vol. 6, p. 84. Substitute name for preoccupied H. nebulosa Spix, 1824. Hyla regilla, Baird and Girard. Cope, 1889. Bulletin U. S. National Museum, vol. 34, p. 359. Hyla arenicolor, Cope. Yarrow, 1882. Bulletin U. S. National Museum, vol. 24, pp. 1-249. Hyla californiae, Gorman. 1960. Herpetologica, vol. 16, pp. 214-222. Holotype. Number 3230. Museum of the Academy of Natural Sciences of Philadelphia. Collected by Dr. Heermann. TYPE SPECIMENS The following type specimens were examined: United States National Museum: 3230 (Hyla nebulosa by error), 3235 (Hyla scapularis hypochordriaca), 9182 (Hyla regilla, Puget Sound), 15409 (Hyla regilla, Sacramento River). Academy of Natural Sciences of Philadelphia: 1978 (Hyla scapularis), 3230 (Hyla nebu- losa). Museum of Vertebrate Zoology of the University of California: 31773 (Hyla californiae). TEJON PASS SAMPLE The description of a male-specimen sample from Tejon Pass, Los Angeles County, California (Los Angeles County Museum No. 26340-26348) follows: A moderate to small population typical of the northern montane and coastal forms attaining an average snout-vent size of approximately 30.4 mm.; dorsal skin moderately pustulate; volmerine teeth distinct; moderate pectoral granula- tion becoming fine anteriorly and posteriorly. A small vestige of webbing exists between the first three fingers; finger pads large; first finger of males possessing nuptial callosities; toes at least three quarters webbed; tibiotarsal VoL. XXXVII] BALL & JAMESON: CANYON TREE FROG SYSTEMATICS Si articulation to mid or anterior portion of the eye. The specimens color in alcohol a dark grayish brown with small diffuse melanophore patches closely dorsolaterally associated; abdomen a dark cream color; the vocal sac of the male not heavily pigmented with melanin; back of the thighs usually con- taining three melanin patches. The average morphological measures in mm. are as follows: snout-vent length 30.4, head width 11.3, head length 9.0, nostril to lip 2.4, forearm length 6.8, finger pad width (third digit) 1.4, shank length 15.6, length of fourth toe 6.3, and the webbing between the fourth and fifth toe 3:0. SPECIMENS ANALYZED VENTURA County. California Academy of Sciences: 50309, 50311—-15, 50396, 50399, 50409-10, 50418, 50440, 50446, 50453, 50455, 50467-68. Los ANGELES County. California Academy of Sciences: 39887-88, 39891, 39906, 39913, 39915, 39923, 39944, 39948, 3995455, 40009, 40012, 40016, 40019-20, 40023-26. Los Angeles County Museum: 26340-48. San Diego Natural History Museum: 19251—52, 19255-57, 19259. RIVERSIDE County. California Academy of Sciences: 43563-65, 43567, 43569-70. San Diego State College: 725-26, 734, 1033-34, 1116, 1143, 1164. SAN BERNARDINO CouNTy. University of California, Museum of Vertebrate Zoology: 14061, 14064, 28245-46, 28250, 28252-55, 41052-59, 52455-60, 5246263, 52470, 52480-81, 58434-36, 58438-39, 58440, 58442. SAN Dreco County. San Diego Natural History Museum: 342, 956, O5e=s0ne6l, 963-66, 971, 973, 1112, 1114, 7517-19, 7522-25, 11521-24. 11527-31, 11536—43, 11592, 11987-89, 12008, 12499, 13256-59, 13331 -32, 13334-38, 14256-58, 14260-65, 14267, 14272, 14274-75, 14497-98, 17134, 21302-03, 21306, 21308-09, 21311-14, 21316-17, 25177-80, 31359-66, 37250- 52, 37254-56, 37258-60, 37263-64. BajA CALIFORNIA DEL Norte. San Diego Natural History Museum: 1041, 1044, 1049-54, 16955-62, 16964-65, 16967-74, 16976, 16978-79, 23463-64, 23466-67, 23469, 23472, 23474-77, 23479. University of California, Museum of Vertebrate Zoology: 9572, 9574-80, 9582, 31766—68, 31772, 31774, 31777. California Academy of Sciences: 13424, 13428-30, 13432-35, 13437-38, 13442, 57460—64, 57468, 57518-19. San Diego State College: B10—-13, 682, 687-88, 694, 697, 719-20, 724, 727-29, 732-33, 735, 1038-40, 1042-43, 1046-48. LITERATURE CITED BALL, R. W., AND D. L. JAMESON 1966. Premating isolating mechanisms in sympatric and allopatric Hyla regilla and Hyla californiae. Evolution, vol. 20, pp. 533-551. Barrp, S. F. 1854. Descriptions of new genera and species of North American frogs. Proceedings of the Academy of Natural Sciences of Philadelphia, vol. 7, pp. 59-62. 378 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Bairp, S. F., anp C. GIRARD 1852. Description of new species of reptiles collected by the U. S. Exploring Expedition under the command of Captain Charles Wilks, U.S.N. Proceedings of the Academy of Natural Sciences of Philadelphia, vol. 6, pp. 174 and 301. Cocuran, D. M. 1961. Type specimens of reptiles and amphibians in the U. S. National Museum. United States National Museum, Bulletin 220, cv + 289 pp. CooLtry, W. W., anp P. R. LOHNES 1962. Multivariate procedures for the behavioral sciences. John Wiley & Sons, Inc., INBYS) x2 Lipp: Corr, E. D. 1866. On the structure and distribution of the genera of the Arciferous anura. Journal of the Academy of Natural Sciences of Philadelphia, 2nd. ser., vol. 6, pp. 67-112. 1866. The reptilia and batrachia of the Sonoran Province of the Nearctic Region. Proceedings of the Academy of Natural Sciences of Philadelphia, vol. 18, pp. 301. 1889. The Batrachia of North America. United States National Museum, Bulletin 34, pp. 1-525. (P. 359.) Gaupin, ANTONY. J. 1964. The tadpole of Hyla californiae Gorman. Texas Journal of Science, vol. 1, pp. 80-84. 1965. Larval development of the tree frogs Hyla regilla and Hyla californiae. Herpe- tologica, vol. 21, pp. 117-130. GORMAN, JOE 1960. Treetoad studies, Hyla californiae, new species. Herpetologica, vol. 16, pp. 214— ihe HALLOWELL, E. 1852. Descriptions of new species of reptiles inhabiting North America. Proceedings of the Academy of Natural Sciences of Philadelphia, vol. 6, pp. 177-183. 1854. Descriptions of new reptiles from California. Proceedings of the Academy of Natural Sciences of Philadelphia, vol. 7, pp. 91-97. JAMEsON, D. L. 1966. Rate of weight loss of tree frogs at various temperatures and humidities. Ecology, vol. 47, pp. 605-613. Jameson, D. L., J. P. Mackey, anp R. C. RicHMonD 1966. The systematics of the Pacific tree frog, Hyla regilla. Proceedings of the California Academy of Sciences, 4th Ser., vol. 19, pp. 551-620. JOLICcOEUR, P. 1959. Multivariate geographic variation in the wolf, Canis lupus L. Evolution, vol. 13, pp. 283-299. Martin, P. S. 1963. The last 10,000 years: a fossil record of the American southwest. University of Arizona Press, Tucson. Ricuarpson, C. H., Jr. 1912. The distribution of Hyla arenicolor with notes on its habits and variation. American Naturalist, vol. 46, pp. 605-611. SAVAGE, J. M. 1960. Evolution of a peninsular herpetofauna. Systematic Zoology, vol. 9, pp. 184-212. Vor. XXXVII] BALL & JAMESON: CANYON TREE FROG SYSTEMATICS 379 SLEVIN, J. R. 1928. The amphibians of western North America. Occasional Papers of the California Academy of Sciences, no. 16, pp. 1-152. (Pp. 110-118). Spx, J. B. DE 1824. Animalia nova, spec. noy. Testundinorum et Ranarum quas in Itinere per Brasiliam Coll. et descr. Munich 4to. STEBBINS, J. R. 1951. Amphibians of Western North America. University of California Press, Berkeley, California. Storer, T. H. 1925. A synopsis of the amphibia of California. University of California Publications in Zoology, no. 27, pp. 1-343. Test, FREDRICK C. 1898. A contribution to the knowledge of the variations of the tree frog Hyla regilla. Proceedings of United States National Museum, vol. 21, no. 1156, pp. 477-492. Wricut, A. H., ano A. A. WRIGHT 1949. Handbook of frogs and toads of the United States and Canada. Comstock, Ithaca, New York. Yarrow, H. C. 1882. Checklist of North American reptilia and batrachia with catalogue of specimens in the United States National Museum. United States National Museum, Bulletin vol. 24, pp. 1-249. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XXXVII, No. 11, pp. 381-394; 11 figs.; 3 tables June 15, 1970 THE CALIFORNIA SEA LION: SKULL GROWTH AND A COMPARISON OF TWO POPULATIONS By Robert T. Orr and Jacqueline Schonewald California Academy of Sciences, San Francisco, California 94118 and Karl W. Kenyon Bureau of Sport Fisheries and Wildlife, U.S. Fish and Wildlife Service, Seattle, Washington 98115 ApstrAct: Teeth and cranial characters denoting the age of 121 male California sea lion skulls (76 from the Pacific coasts of California and Baja California and 45 from the Gulf of California) were studied. Ages ranged from 1 to about 15 years. Great individual variations in cranial characters were found, but no significant differences between the two populations were revealed. The relation between age and suture closure was fairly constant until about age 15 when all 9 sutures studied are completely closed. In length and width, skull growth continues until about age 10 years. Sagittal crest growth commences in the 5th year and ceases in about the 10th year. INTRODUCTION The California sea lion (Zalophus californianus californianus) in the eastern North Pacific breeds on suitable offshore islands from San Miguel, the northern- most of the California Channel Island group, south along the west coast of Baja California to the San Benitos Islands, and on certain islands in the Gulf of Cali- fornia and along the mainland of Mexico, south possibly to Mazatlan. Between 1964 and 1968, in the months of April, June, August, and Novem- ber, Orr made six trips to islands in the Gulf to secure data on the sea lion pop- [381] 382 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Taste 1. Localities, from north to south, in the Gulf of California at which Zalophus californianus was observed by the authors between 1960 and 1968. Locality Date Number Composition Isla Coloradito 24 Nov. 1960 500-600 Both sexes, all ages Bahia San Luis Gonzaga 10 Nov. 1965 75 Both sexes, all ages Isla Granite 20 Apr. 1966 1,000-+ Both sexes, all ages Isla Angel dela Guarda 21 Apr. 1966 1,000++ Unknown, observed from a distance Bahia de los Angeles 13 Nov. 1965 75 Both sexes, all ages Isla Partida (Norte) 28 Apr. 1966 485 30% ad. 6 6, 30% subad. 6 6, 40 % 2 2 and imm. Isla San Esteban 27 Apr. 1966 151 125 ad) ¢ 4, 26 2 2 omimmeosc Isla San Pedro Martir 25 Apr. 1966 2,700+ Both sexes, all ages " 29 Apr. 1968 300+ Unknown Isla Santa Catalina 24 June 1964 several No breeding activity Isla Las Animas 27 June 1964 8 5 ad. 24, 3 292 ormmmecece no breeding activity " 12 Aug. 1965 30 ~=Both sexes, all ages " 7 Oct. 1964 500 Unknown? Los Islotes 30 June 1964 187. 12 ad. 64, incl. 3 dominant bulls, 2° subads =@\d GanliSaee Roe imams Breeding ? " 10 Aug. 1965 190+ Several ad. ¢ @ incl. 4 dominant bulls, 150 2 2 with large nipples, and imm. " 4 Oct. 1964 200-300 Unknown? 5 Isla Cerralvo 25 Apr. 1966 75+- 6-8 Ad) 6°65 larzesnumbersomegucn some imm. ¢ 4 Cabo San Lucas 8 Feb. 1965 il Nake 1Cf. Lindsay, 1962. 2 Observed by George Tsegeletos. ulation of that area. From 3 to 15 days per visit were spent in the field. Kenyon observed sea lions and collected skulls in the Gulf in November 1960 and Feb- ruary 1965. California sea lions were observed from Isla Coloradito in the north to Cape San Lucas in the south (table 1). Mixed populations of males, females, and im- matures were noted at essentially all times. This suggested a difference in be- havior between the Gulf population, which may be rather sedentary, and the Pacific coastal population, in which the majority of the males move north of the areas occupied by the females and young after the breeding season (Orr and Poulter, 1965; Peterson and Bartholomew, 1967). Fully adult males move farthest north, regularly at least to Sea Lion Caves near Florence, Oregon (Ken- yon and Scheffer, 1962). On only one occasion, at San Esteban Island on 27 April 1966, was there some indication of at least local sexual segregation. On that date a group composed of 125 males and 26 other individuals, which might have been females or immature males, was observed hauled out together. Vol. XXXVIT] ORR, SCHONEWALD, & KENYON: SEA LION 383 Ficure 1. A section of canine tooth of a Zalophus californianus male showing annual growth lines. University of California photograph by Alan Donaldson. Courtesy of Richard S. Peterson. 384 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. TaBLe 2. Age composition of 35 California sea lion skulls based on growth layers of an upper canine. Age Specimens Age Specimens Years Number Years Number 1 1 9 3 2 y) 10 4 3 2 11 5 4 1 12 2 5 1 13 3 6 1 14 1 7 3 15 4 8 2 The authors made a special effort to collect skulls, primarily those of adult (1) to determine if there were any cranial differ- ences between the Gulf of California population and that occurring on the Pa- cific coastal side of Baja California and the California coast, and (2) to study skull growth and suture closure in relation to age. males. Our objectives were: TABLE 3. Comparison of cranial measurements of adult male skulls (suture ages 25-36) of Zalophus californianus from the Gulf of California (G) and Pacific coasts of Baja Cali- fornia and California (P). Measurement Population N Range in mm. Mean Condylobasal length P 44 262 -300 284.38 + 1.42 G 28 264 -302 286.03 + 1.95 Gnathion to posterior |p 44 92.5-113 104.42 + 0.70 end of nasals G 28 92.5-115 103.82 + 1.15 Gnathion to posterior border P 45 199 -226 215.40 + 1.10 of postglenoid process G 28 201 -229 218.00 + 1.52 Gnathion to posterior border P 44 93 -108 Oily s= O57 of preorbital process G 29 88 -108 99.28 + 0.93 Breadth at preorbital 12 38 75.5— 95.5 81.82 + 0.78 processes G 24 67.0— 90.0 S2A 20g Interorbital constriction JP 45 36.0— 59.5 46.57 + 0.56 G $jil 39.0— 52.5 45.59 = 0.57 Breadth at supraorbital iP 43 56.5— 87.0 68.99 + 1.08 processes G 27 50.5— 82.5 65.68 + 1.48 Zygomatic breadth P 44 140 -171 ISS) SE 105 G 26 136 -171 157.15 + 1.46 Mastoid breadth 1p 44 127 -164 143.20 + 0.36 G 30 123 -153 142.33 + 0.43 Sagittal crest* 12 45 11 — 34 25.48 + 0.81 G 32 13 — 36.5 26.87 + 1.00 CV 3.30 3.60 4.50 5.83 3.42 3.68 3.42 5.06 5.86 6.10 8.15 7.01 10.29 11.72 4.41 4.74 5.30 5.26 235 PRES * This measurement is included here to show that it is equally variable in both populations. Vol. XXXVIT] ORR, SCHONEWALD, & KENYON: SEA LION 385 Pacific Coast a Gulf of California 8] Galapagos Ids. 7) ——— +} Ficure 2. Statistical analysis of the condylobasal length in adult male skulls (suture index 25-36) of Zalophus c. californianus from the Pacific coast and Gulf of California. Horizontal lines represent observed range, open rectangles the standard deviation, and the solid black is twice the standard error of the mean. For further comparison the range and mean of a small sample of Zalophus c. wollebaeki is shown. The number of specimens used is given in parentheses. MATERIALS AND METHODS SPECIMENS EXAMINED. A total of 121 skulls of males (76 from the Pacific coast of California and Baja California and 45 from the Gulf of California, Mexico) were studied:* California: Marin County: Point Reyes Peninsula, 1 (CAS); San Francisco County: San Francisco, 2 (RB); San Mateo County: Pacifica, 1 (RB); Afio Nuevo Island and Ano Nuevo Point, 42 (RB, 31; CAS, 11); Santa Cruz County: Waddell Beach, 7 (RB); Santa Barbara County: Santa Cruz Island, 15 (CAS, 14; CAS-SU, 1); Mexico: Baja California: Isla San Martin, 2 (LACM); vicinity of Bahia de Sebastian Viscaino, 4 (LACM, 2; SDNHM, 2); Bahia Tortugas, 2 (SDNHM); Puertecitos, 1 (LACM); Isla Coloradito, 17 (KWK); Isla Granite, 3 (SDNHM); Isla Angel de la Guarda, 5 (CAS, 1, SDNHM, 4); Bahia de los Angeles, 2 (CAS); Isla Carmen, 1 (CAS); Isla Santa Cruz, 1 (SDNHM); Isla San Diego, 1 (SDNHM); Isla San José, 1 (CAS); Isla Espiritu Santo, 1 (CAS); Isla Cerralvo, 1 (CAS); Rancho Chenque, 1 (SDNHM); Sonora: Isla San Jorge, 3 (LACM); Isla San Esteban, 2 (CAS); Isla San Pedro Martir, 5 (CAS, 4; SDNHM, 1). Most of the skulls were from animals that had recently died. A few were weathered skulls which were found on beaches. * Abbreviations used: CAS—California Academy of Sciences collection; RB—Raymond Bandar private collection: CAS-SU—Stanford University collection, now incorporated in CAS collection; LACM—Los An- geles County Museum collection; SDNHM—San Diego Natural History Museum collection; KWK—Karl W. Kenyon collection, now incorporated in CAS collection, 386 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. UII! CI es a ae 80 90 100 110 Pacific Coast (44) Gulf of California ———— Galapagos Ids. (6) —+— Ficure 3. Statistical analysis of the skull from gnathion to posterior end of nasals of adult males from Pacific coast and Gulf of California populations of Zalophus c. califor- nianus. The range and mean of a small sample of Zalophus c. wollebaeki are shown for further comparison. = a De Sa ian Cie Taare) (amet Comes | 130 140 150 160 170 mm Pacific Coast (44) Gulf of California a jase Olen Ids. (6 8 Ficure 4. Statistical analysis of zygomatic breadth of skull of adult males from Pacific coast and Gulf of California populations of Zalophus c. californianus. The range and mean of a small sample of Zalophus c. wollebaeki are shown for further comparison. Vol. XXXVIT] ORR, SCHONEWALD, & KENYON: SEA LION 387 200 250 300 mm Ficure 5. Condylobasal length of 31 skulls of Zalophus c. californianus males, selected to show a wide range in age, and plotted against approximate known age determined by number of annual tooth rings. AGING TECHNIQUE. Scheffer (1950) demonstrated that annual growth layers could be counted in the teeth of fur seals (Callorhinus ursinus). The teeth of sea lions exhibit similar growth layers (fig. 1). For the present study we selected 35 skulls, representing a broad spectrum of ages. A single upper canine was re- moved from each, and sectioned and polished to reveal the growth layers. Ages, based upon tooth ring counts, are accurate only to within a year or two. The age groups represented are shown in table 2. SUTURE INDEX. All specimens were suture indexed according to the method 388 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. 10 120 140 160 mm Ficure 6. Zygomatic breadth of 33 skulls of Zalophus c. californianus males, selected to show a wide range in age, and plotted against approximate known age determined by number of annual growth rings. used by Sivertsen (1954), which is a modification of that first suggested by Doutt (1942). Nine sutures were used. These are as follows: I, occipito- parietal; II, squamoso-parietal; III, interparietal; IV, interfrontal; V, coronal; VI, basioccipito-basisphenoid; VII, maxillary; VIII, basisphenoid-presphenoid; IX, premaxillary-maxillary. A value of 1 to 4 is given on the basis of degree of closure: 1, open; 2, less than half closed; 3, more than half closed; and 4, com- pletely closed. An immature skull, therefore, could not have an index of less than 9 and a mature skull of more than 36, Vol. XXXVII] ORR, SCHONEWALD, & KENYON: SEA LION 389 yrs. 90 110 130 150 alt Ficure 7. Mastoid breadth of 33 skulls of Zalophus c. californianus males, selected to show a wide range in age, and plotted against approximate known age determined by number of annual growth rings. MEASUREMENTS. Ten cranial measurements were taken, as shown in table 3. These were submitted to statistical analysis. Studies of individual variation and variations between populations were made. DISCUSSION AND CONCLUSIONS GEOGRAPHICAL VARIATION. Analysis of the 10 cranial measurements showed great individual variability, but there were no significant statistical differences in these characters between the California sea lion population of the Gulf and [ Proc. 4TH SER. ACADEMY OF SCIENCES CALIFORNIA 390 dy} UO 4Sa1d JUSUTOId 9Y} BION “puRIS[ OAINN OUYW UO 3NO paNeyY SUOT] vas RIUIOJI[ED sew y[Npe jo Ajasiey posodwios dnois y ‘g aunog — hie i * . . ail P er AjoggpeceelS Keg feet. Gli i (feo ‘10 “L ‘UW Aq 1961 API SZ paydeisojoyg ‘[[NyYs oy} UO 4satd [e}JISeS ay} JO JUaUIdO[aAaP 9Y} YIM poyefatiod st yoIyM ‘pray Pores Vol. XX XVII] ORR, SCHONEWALD, & KENYON: SEA LION 391 yrs. 6) 5 10 15 20 25 mm Ficure 9. Height of sagittal crest on skull of 34 specimens of Zalophus c. californianus males, selected to show a wide range in age, and plotted against approximate known age de- termined by number of annual growth rings. that of the Pacific coast. Three measurements (condylobasal length, gnathion to posterior end of nasals, and zygomatic breadth) which were among those with the lowest coefficient of variability are shown in figures 2, 3, and 4. By way of comparison, the range and mean for these same measurements in six adult skulls of Zalophus c. wollebacki from the Galapagos Islands are included. SKULL GROWTH AND SUTURE CLOSURE. The indicated ages of 35 animals were plotted against four measurements: condylobasal length, zygomatic breadth, mastoid breadth, and height of sagittal crest. These measurements indicate 392 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. suture number 10 15 20 25 30 35 suture index Ficure 10. Suture index from beginning to end of closure of sutures I to IX. growth until about the 10th year of life, following which there is essentially no marked change (figs. 5, 6, and 7). Similar findings for the Steller sea lion (Eumetopias jubata) male were reported by Fiscus (1961). Although it is dif- ficult to obtain an accurate measurement of the height of the sagittal crest, it was used because it is one of the best indicators of age in male California sea lions (fig. 8). No crest is apparent on 2-, 3-, and 4-year-old individuals. A small crest appears in the 5th year and from then on the crest increases in height up to the 10th year of life, after which little change is evident (fig. 9). The time at which suture closure begins as well as the length of time involved depend upon the suture. Sutures I, II, V, and VI are the first to show signs of closing. Sutures VII, VIII, and IX are last to begin closure. The greatest length of time required for closure is shown by sutures IV and V, the shortest time by sutures III, VI, and VII (fig. 10). By plotting the known tooth age in 35 skulls of male Zalophus californianus against the suture index for each specimen, it is evident that there is a fairly constant correlation between age and suture index; the latter increases grad- ually until the 15th year of life, at which time all nine sutures are completely closed (fig. 11). ACKNOWLEDGMENTS We are grateful for the loan of skulls from: Raymond M. Bandar, his per- sonal collection; the San Diego Natural History Museum; and the Los Angeles County Museum. Clifford H. Fiscus of the Marine Mammal Biological Labo- Vol. XXXVII]_ ORR, SCHONEWALD, & KENYON: SEA LION 393 15 10 14 18 22 26 30 34 suture index Ficure 11. Suture index of 35 skulls of Zalophus c. californianus males, selected to show a wide range in age, and plotted against approximate known age determined by number of annual growth rings. ratory, Bureau of Commercial Fisheries, Seattle, Washington, sectioned teeth and assisted in obtaining readings of age. For making possible field studies related to this work, we wish to thank the following individuals or organizations: Bechtel Corporation, Belvedere Scientific Fund, Victor J. Bergeron, California Academy of Sciences, Mr. and Mrs. Bing Crosby, Mr. and Mrs. Lawrence C. Kuebler, Dr. Thomas C. Poulter, San Diego Natural History Museum, Stanford Research Institute, and Mr. and Mrs. George Wetmore. 394 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. LITERATURE CITED Doutt, KENNETH J. 1942. A review of the genus Phoca. Annals of the Carnegie Museum, vol. 29, pp. 61-125. Fiscus, Ciirrorp H. 1961. Growth in the Steller sea lion. Journal of Mammalogy, vol. 42, pp. 218-223. Kenyon, Kart W., AND VICTOR B. SCHEFFER 1962. Wildlife surveys along the northwest coast of Washington. Murrelet, vol. 42, pp. 29-37. Linpsay, GEORGE E. 1962. The Belvedere Expedition to the Gulf of California. Transactions of the San Diego Society of Natural History, vol. 13, pp. 1-44. Orr, Ropert T., AND THOMAS C. POULTER 1965. The pinniped population of Ano Nuevo Island, California. Proceedings of the California Academy of Sciences, ser. 4, vol. 32, pp. 377-404. Peterson, RicHarD S., AND GEORGE A. BARTHOLOMEW 1967. The natural history and behavior of the California sea lion. American Society of Mammalogists special publication no. 1, xi + 79 pp. SCHEFFER, VICTOR B. 1950. Growth layers on the teeth of Pinnipedia as an indication of age. Science, vol. 112, pp. 309-311. SIVERTSEN, ERLING 1954. A survey of the eared seals (family Otariidae) with remarks on the antarctic seals collected by M/K “Norvegia” in 1928-1929. Det Norske Videnskaps- Akademi i Oslo. Scientific results of the Norwegian Antarctic Expeditions 1927-1928 et sqq., instituted and financed by Consul Lars Christensen, no 36. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XXXVIT, No. 12, pp. 395-418; 10 figs.; 7 tables June 15, 1970 NEW SCORPIONS BELONGING TO THE EUSTHENURA GROUP OF VEJOVIS FROM BAJA CALIFORNIA, MEXICO (SCORPIONIDA: VEJOVIDAE). By Stanley C. Williams Department of Ecology and Systematic Biology San Francisco State College, San Francisco, California, and Research Associate, California Academy of Sciences. ABSTRACT: Seven new species and subspecies of scorpions in the genus Vejovis are described and discussed. These new taxa belong to the eusthenura group of the genus. One of the new species, Vejovis viscainensis Williams, appears to represent a distinct systematic branch of this group. INTRODUCTION Recent biological exploration of the Baja California peninsula has resulted in the finding of much important information about the scorpion fauna of western North America. The species Vejovis eusthenura (Wood), which has caused much taxonomic confusion for many years, has now been collected in large numbers and defined in terms of morphology and zoogeographic distribution. In addition, three other new species closely related to Vejovis eusthenura have been described from Baja California: Vejovis schwenkmeyeri Williams, Vejovis hoffmanni Williams, and Vejovis diazi Williams. Since this more recent work, a whole group of related Vejovis scorpions has been discovered in Baja California, southern California, and Arizona. Seven of these are described and named in this paper. These findings have special importance in that the evolution and systematic relationships of those scorpions belonging to the genus we now call Vejovis are beginning to be understood. [395] Marine Biological |aharatany ] Dor all JUN 231970 396 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Scorpions of the “eusthenura” group of Vejovis are found in western North America with greatest species diversity in the Baja California peninsula and the desert country of southern California and Arizona. Most are uniform pale yellow in color and found in more or less sandy habitats. Most of the species are burrowers, some of which appear to be found on the ground surface only during a very restricted time of the year, presumably the mating season. Several of the species appear to have several sexually mature instars. Generally, members of this group may be recognized by the following characteristics: metasomal seg- ment V distinctly longer than either carapace or movable finger; pedipalp fingers elongate, movable or fixed finger distinctly longer than underhand; palm narrow, with keels lowly developed to obsolete; dorsal and dorsolateral keels of metasoma well developed and coarsely crenulate to serrate; inferior lateral keels of metasoma present, these smooth, crenulate or intermediate; carapace and tergites densely granular; chelicerae with inferior border of movable finger lacking denticles; carapace anterior margin with subtle median emargination to es- sentially straight; lateral eyes small, three per group; pectines well developed, teeth numbering about 13 to 22; females’ pectines smaller and with fewer teeth than those of male; adult males with distinct genital papillae, and completely divided genital operculum; females lacking genital papillae, genital operculum not completely divided longitudinally. The species of the ‘‘ewsthenura” group are very similar in basic morphology and coloration. The following characteristics, however, appear to be sensitive indicators of speciation and are, therefore, considered good diagnostic character- istics: elongation of the pedipalp fingers; reduction of palm width; size and structure of proximal scallop of pedipalp fingers when chela closed; open space remaining between pedipalp fingers when chela closed; hirsuteness of vesicle, especially of mature males; surface texture of vesicle; structure of inferior median and inferior lateral keels of metasoma; ratio of length to width of metasomal segment I; setation of anterior margin of carapace. The measurements used in this paper are essentially the standard ones in current scorpion systematics with minor modification (Williams, 1968a). Most of the specimens studied were collected by the ultraviolet detection method (Williams, 1968b), on two expeditions through the Baja California peninsula during 1968 and 1969. During these two expeditions, some 50,000 scorpions were collected including several thousand belonging to the “eusthenura”’ group. These specimens were killed, preserved, and processed by the methods recently recom- mended by Williams (1968c). ACKNOWLEDGMENTS Appreciation is greatly acknowledged to the following individuals and institu- tions for the loan of specimens: W. J. Gertsch, American Museum of Natural Vol. XXXVII] WILLIAMS: NEW VEJOVIS TAXA FROM BAJA CALIFORNIA 397 History; Paul H. Arnaud, California Academy of Sciences; M. A. Cazier, Arizona State University. The following people significantly aided the study by assistance during the field work: M. A. Cazier, J. Bigelow, J. D. Davidson, M. M. Bentzien, W. K. Fox, V. F. Lee, and H. L. Heringhi. Thanks are also acknowledged to V. F. Lee for technical assistance and to C. F. Williams for clerical assistance. This research was partially sponsored by the National Sci- ence Foundation through research grant GB-7679 and by San Francisco State College by a Faculty Research Leave during the spring of 1969. The Mexican government, Department of Wildlife, is gratefully acknowledged for permission to collect and study scorpions in Mexico. Much appreciation is acknowledged to Mr. and Mrs. Thomas P. Hearne for providing transportation on their boat the Muy Pronto, to otherwise inaccessible regions of Baja California. NEw SPECIES Vejovis waeringi Williams, new species. (Figures 1, 2.) Dracnosis. Relatively large, uniform pale yellow species of the “eusthenura”’ group of Vejovis. Closely related to Vejovis schwenkmeyeri Williams and Vejovis coloradensis Williams by the essentially bald and conspicuously tubercular vesicle of the male. Differs from V. schwenkmeyeri in the following ways: lack of conspicuous pedipalp scallop; the presence of more elongate pedipalp fingers; metasoma with inferior median keels smooth to crenulate on segments I to ITI, crenulate on IV: metasoma with inferior lateral keels crenulate (not essentially smooth with a few crenulations). More closely related to V. coloradensis from which it differs in the following characteristics: females with vesicle conspicu- ously hirsute (not approaching inconspicuous hirsuteness to baldness); females with dorsal keels of metasomal segments IV and V conspicuously hirsute; inferior median keels of metasoma more smooth than crenulate (not definitely crenulate) ; movable finger of pedipalp approximates carapace length (not distinctly longer than carapace). DESCRIPTION OF HOLOTYPE (male). Coloration: Entire body uniform pale yellow, pectines whitish; no distinctive contrasting markings. Carapace: Anterior margin essentially straight, with very subtle median emargination; anterior margin set with about three or four pairs inconspicuous reddish bristles; entire surface densely granular. Mesosoma: Tergites densely granular; 7th tergite with two pairs well de- veloped serrate keels. Sternite 7 with one pair crenulate lateral keels. Stigma long oval. Metasoma: Inferior lateral keels crenulate on segments I to IV. Inferior median keels smooth on segment I, smooth with several crenulations on IJ, 398 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Ficure 1. Vejovis waeringi Williams, new species. Dorsal view of holotype male. smooth to crenulate on III, irregularly crenulate on IV. Segment I wider than long. Telson: Ventral surface with numerous short inconspicuous hairs, not ap- pearing hirsute. Ventral surface conspicuously tubercular. Vesicle more than twice aculeus length. Pedipalps: Chela long and slender. Keels low and inconspicuous with low, rounded granules. Fingers with long shallow proximal scallop; fingers touch only at distal ends when chela closed leaving long open space. Movable finger shorter than metasomal segment IV, movable finger equal to carapace in length, longer than brachium. Fixed finger longer than metasomal segment III. Standard measurements and photographs: Table 1 and figures 1 and 2. DESCRIPTION OF ALLOTYPE (female). Morphologically the same as holotype with the following exceptions: larger in total length; ventral surface of vesicle abundantly covered with long conspicuous hairs; pectines smaller, with fewer teeth; keels on pedipalp hand obsolescent; tarsomeres with long conspicuous hairs on posterior side; vesicle more swollen; dorsal keels of metasomal segments IV and V set abundantly with long hairs; inferior median and inferior lateral keels of metasoma set with several long conspicuous hairs. Standard measurements; able 1, Vol. XXXVIT] WILLIAMS: NEW VEJOVIS TAXA FROM BAJA CALIFORNIA 399 Ficure 2. Vejovis waeringi Williams, new species. Ventral view of holotype male. VARIATION WITHIN PARATYPES. Study of 233 paratopotypes (179 males, 54 females) indicated little variation from the descriptions of the holotype and allotype. Males varied in total length from 32 to 47 millimeters while females varied from 30 to 58 millimeters. Most specimens of both sexes appeared to be mature. Females tended to be larger than males in most body proportions. Pectine teeth varied from 12 to 14 in females and from 15 to 18 in males (based on 20 males, 20 females). The holotype and allotype appear typical of the paratopotype sample. TYPE DATA AND ETYMOLOGY. The holotype, allotype, and 233 paratopotypes were collected at Okies Landing, 27 miles south of Puertocitos, Baja California Norte, Mexico, 12 June 1968, by S. C. Williams, M. A. Cazier, and party. The holotype and allotype are permanently deposited in the California Academy of Sciences. This species is named “‘waering?” after Erik Kjellsvig-Waering, because of his contributions toward the understanding of the evolution and paleontology of scorpions, and because of his recent contributions toward defining the scorpion fauna of the West Indies. DISTRIBUTION. Known only from the upper gulf region of Baja California, Norte, Mexico, from the Rio Hardy Fishing Camp south to Okies Landing, and east of the Sierra San Pedro Martirs and Sierra Juarez. 400 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. TABLE 1. Measurements (in millimeters) of Vejovis waeringi Williams, new species, holotype and allotype. holotype allotype (male) (female) Total length 46.0 54.0 Carapace, length 5.6 6.7 width (at median eyes) 4.0 4.6 Metasoma, length 22.6 23 segment I (length/width) BAe 3.3/3.8 segment II (length/width) 3.7/3.4 3.7/3.7 segment III (length/width) 3.8/3.3 AL /3. segment IV (length/width) 52/34 Seite segment V (length/width) Gniy/ SEZ 6.8/3.8 Telson, length 6.2 7.3 Vesicle (length/width) AP 5.1/3.4 depth Pail 2.8 Aculeus, length 1.9 2.2 Pedipalp Humerus (length/width) 4.9/1.4 5.4/1.5 Brachium (length/width) 32 / Aoi Soo Chela (length/width) 8.1/1.4 9.2/2.0 depth 7 Deed movable finger, length 5.6 6.5 fixed finger, length 4.6 5.0 Pectines teeth (left/right) 15/16 13/13 Recorps. In addition to the holotype, allotype, and 233 paratopotypes, 53 other paratypes, all from Baja California Norte, Mexico, were available for study: San Felipe, elevation 25 feet, 8 June 1968 (S. C. Williams, M. A. Cazier, and party), 12 males, 4 females; San Felipe, elevation 25 feet, 9 June 1968 (S. C. Williams, M. A. Cazier, and party), 10 males, 3 females; Puertecitos, eleva- tion 25 feet, 10 June 1968 (S. C. Williams, M. A. Cazier, and party), 6 males, 2 females; Puertecitos, elevation 25 feet, 11 June 1968 (S. C. Williams, M. A. Cazier, and party), 9 males, 5 females; 4 miles west of Rio Hardy Fishing Camp, elevation 100 feet, 19 July 1969 (S. C. Williams, and V. F. Lee), 2 females. CoMMENT. The conspicuous sex ratios, significantly in favor of the males, indicate that this species must have a predominantly subsurface living habit, even at night. The conspicuous male surface activity was probably due to courtship surface migrations. Vol. XXXVII] WILLIAMS: NEW VEJOVIS TAXA FROM BAJA CALIFORNIA 401 Vejovis coloradensis Williams, new species. (Figures 3, 4.) Dracnosis. Large uniform pale yellow species of the eusthenura group of Vejovis. Distinctive characteristics include: movable finger distinctly longer than carapace; metasomal segment I wider than long; no pedipalp scallop; male vesicle appearing essentially bald and conspicuously tubercular, female only slightly more hirsute; related to Vejovis waeringi Williams and Vejovis confusus Stahnke. Differs from V. confusus in the following ways: chela more elongate and slender; males with palm of chela distinctly less than two times width of movable finger base, females with palm width two times movable finger base width (palm proportionally wider in V. confusus); males with vesicle more laterally swollen and more conspicuously tubercular; females with a few long conspicuous hairs on vesicle but not conspicuously hirsute; movable finger dis- tinctly longer than carapace. Differs from V. waeringi in the following ways: females with vesicle not conspicuously hirsute; females with dorsal keels of metasomal segments IV and V not conspicuously hirsute; inferior median keels of metasoma definitely crenulate (not more smooth than crenulate); movable finger of pedipalp definitely longer than carapace (not approximating carapace length). DESCRIPTION OF HOLOTYPE (male). Coloration: Body uniform pale yellow; pectines whitish; pedipalp finger slightly darker yellow. Carapace: Anterior margin with slight median emargination, this border with three pairs of short stout hairs; entire surface coarsely granular. Mesosoma: Tergites densely granular; 7th tergite with two pairs of serrate lateral keels; 7th sternite with one pair of serrate lateral keels; stigma small, long-oval. Metasoma: Inferior lateral keels crenulate to serrate; inferior median keels crenulate; segment I definitely wider than long. Telson: Ventral surface laterally swollen and conspicuously covered with large rounded tubercules, this surface with about three pairs of relatively long but inconspicuous hairs (vesicle definitely not hirsute in appearance); aculeus short, % vesicle in length. Pedipalps: Long and slender; palm width 1.5 times width of movable finger base; movable finger longer than carapace; palm with inconspicuous keels, these smooth to faintly crenulate. Standard measurements and photographs: Table 2 and figures 3 and 4. DESCRIPTION OF ALLOTYPE (female). The same as holotype in coloration and structure with the following exceptions: considerably longer in body length; pec- tines with fewer teeth; palms slightly more swollen; palm twice as wide as base of movable finger; vesicle venter with about seven or eight pairs of long hairs 402 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Ficure 3. Vejovis coloradensis Williams, new species. Dorsal view of holotype male. (vesicle still not hirsute in appearance); hairs on anterior border of carapace longer. Standard measurements: Table 2. TYPE DATA AND ETYMOLOGY. The holotype, allotype and one male paratopo- type were collected at the California Agricultural Inspection Station, 2 miles west of Andrade, Imperial County, California, 6 July 1969 by S. C. Williams and V. F. Lee. The holotype and allotype are permanently deposited in the California Academy of Sciences. This species is named ‘“‘coloradensis” because of its distribution in the Colo- rado Desert of North America. DisTRIBUTION. This species is found in the Imperial Valley of the Colorado Desert in southern California and in northern Baja California, Mexico. It was found in open sandy habitats but was not abundant in the samples taken. Recorps. In addition to the holotype, allotype, and one male paratopotype, 12 other paratypes were available for study: California (U. S. A.), Imperial County: Glamis, 14 October 1967 (M. A. Cazier, J. Bigelow, and S. Goroden- ski), 2 males; 19 miles west of Calexico, 6 July 1969 (S. C. Williams and V. F. Lee), 2 males; 10 miles east of Calexico, 6 July 1969 (S. C. Williams and V. F. Lee), 2 males. Mexico, Baja California Norte: 2 miles south of Algodones, 18 July 1969 (S. C. Williams and V. F. Lee), 2 males, 1 female; Algodones, 18 July 1969 (S. C. Williams and V. F. Lee), 3 males. Vol. XXXVIT] WILLIAMS: NEW VEJOVIS TAXA FROM BAJA CALIFORNIA 403 TaBLe 2. Measurements (in millimeters) of Vejovis coloradensis Williams, new species, holotype and allotype. holotype allotype (male) (female) Total length 46.0 52.0 Carapace, length SE 5.8 width (at median eyes) 3.7 4.3 Metasoma, length 20.2 PV, segment I (length/width) 2.8/3.2 2.8/3.4 segment II (length/width) 3.4/3.2 3.3/3.3 segment III (length/width) SMS 3.6/3.3 segment IV (length/width) 4.8/3.3 4.8/3.4 segment V (length/width) Se SVisies: 6.7/3.4 Telson, length 5.8 6.7 Vesicle (length/width) 4.2/2.6 4.6/3.0 depth 1.8 2.4 Aculeus, length 1.6 pel Pedipalp Humerus (length/width) 4.8/1.3 5.2/1.4 Brachium (length/width) Eyes 5.7/1.9 Chela (length/width) 8.3/1.4 9.2/1.9 depth 1.5 Beal movable finger, length 5.6 6.4 fixed finger, length 4.6 5:3 Pectines teeth (left/right) 18/18 14/14 Vejovis terradomus Williams, new species. (Figures 5, 6.) Diacnosis. Medium sized species of the eusthenura group of Vejovis. Base color yellow with light dusky markings on dorsum of carapace and mesosoma; pedipalp fingers of medium length; males with vesicle bald and tubercular; pedipalp fingers of adult male with proximal scallop. Closely related to Vejovis galbus Williams and Vejovis schwenkmeyeri Williams by the bald, conspicuously tubercular male vesicle and hirsute female vesicle. Differs from V. schwenk- meyeri by the following characteristics: distinctly smaller body size; completely crenulate inferior lateral keels of metasoma; pedipalp fingers proportionally slightly longer. Differs from V. galbus in the following characters: somewhat larger species; metasoma with segment I slightly longer than wide (not slightly wider than long); pedipalp fingers proportionally longer, movable finger slightly longer than carapace (not shorter); pedipalp fingers with distinct scallop (scallop 404 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Ficure 4. Vejovis coloradensis Williams, new species. Ventral view of holotype male. not obsolescent) ; males with movable pedipalp finger twice as long as underhand (not 1.5 times as long as underhand). DESCRIPTION OF HOLOTYPE (male). Coloration: Base color of body pale yellow; carapace and mesosomal dorsum with irregular dusky underlying mark- ings, tergites terminate with posterior black seam; pedipalp fingers light orange; pectines whitish. Carapace: Anterior margin straight, set with six erect hairs; entire surface densely granular. Mesosoma: Tergites densely granular; tergite of segment 7 with two pairs of well developed and serrate lateral keels; 7th sternite with one pair of crenulate lateral keels; stigma long oval. Metasoma: Inferior lateral keels distinctly crenulate on segments I to V; inferior median keels smooth on segments I and II, smooth to lightly crenulate on III and IV; segment I longer than wide; segment V longer than telson. Telson: Ventral surface of vesicle with a number of very short inconspicuous hairs, not hirsute in appearance; venter of vesicle densely covered with rounded tubercules; vesicle twice aculeus length. Pedipalps: Fingers of chela with proximal scallop; fingers touch only at distal ends when chela closed, leaving elongate open space; palm with keels obsolete; movable finger shorter than metasomal segment V but longer than carapace. Vol. XXXVII] WILLIAMS: NEW VEJOVIS TAXA FROM BAJA CALIFORNIA 405 TaBLE 3. Measurements (in millimeters) of Vejovis terradomus Williams, new species, holotype. holotype (male) Total length 44.0 Carapace, length 4.9 width (at median eyes) 3.5 Metasoma, length 21.3 segment I (length/width) 2.9/2.7 segment II (length/width) S\o5y) Doll segment III (length/width) 3.7/2.6 segment IV (length/width) ADS) segment V (length/width) 6.5/2.4 Telson, length 5.9 Vesicle (length/width) ALAA) A0) depth 1.6 Aculeus, length 1.8 Pedipalp Humerus (length/width) 4.4/1.3 Brachium (length/width) 4.6/1.6 Chela (length/width) 7.7/1.8 depth 1.9 movable finger, length 5.1 fixed finger, length 4.0 Pectines teeth (left/right) 19/19 Standard measurements and photographs: Table 3 and figures 5 and 6. VARIATION WITHIN PARATYPES. Study of nine paratopotypes (6 males, 3 females) indicated little significant variation from holotype in color and struc- ture. An allotype was not erected because doubt existed whether any of the females were mature. Males varied in total body length from 33 to 41 milli- meters, females varied from 23 to 35 millimeters. Pectine tooth counts varied from 14 to 15 in females, and from 18 to 19 in males. TYPE DATA AND ETYMOLOGY. The holotype and nine paratopotypes were collected 1 mile southwest of Rancho Canipolé, Baja California Sur, Mexico on 15 May 1969 by S. C. Williams. The holotype is permanently deposited in the California Academy of Sciences. This species is named ‘“‘terradomus” which means soil dweller. DIsTRIBUTION. This species is known only from the limited distribution between Rancho Canipolé and Loreto, Baja California Sur, Mexico. 406 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47H Serr. Ficure 5. Vejovis terradomus Williams, new species. Dorsal view of holotype male. Ficure 6. Vejovis terradomus Williams, new species. Ventral view of holotype male. Vol. XXXVIT] WILLIAMS: NEW VEJOVIS TAXA FROM BAJA CALIFORNIA 407 Recorps. Mexico, Baja California Sur: 5 miles south of Loreto, elevation 100 feet, 16 May 1969 (S. C. Williams), 7 males, 3 females; 1 mile south of Loreto, elevation 25 feet, 17 May 1969 (S. C. Williams), 1 male, 3 females. CoMMENT. The sizes and sex ratios of the samples indicate that this species probably spends most of its diurnal and nocturnal life inside more or less per- manent burrows. Surface activity may be limited to courtship and mating seasons and then would be expected to be predominantly that of sexually mature males. Vejovis galbus Williams, new species. (Figures 7, 8.) Diacnosis. Medium sized, pale yellow species of Vejovis belonging to the “eusthenura” group. Metasomal segments I to IV with inferior lateral keels more smooth than crenulate; inferior median keels smooth to obsolete; male vesicle bald and tubercular, female vesicle hirsute; pedipalp fingers relatively short for eusthenura group, movable finger distinctly shorter than carapace and only 1.5 times longer than underhand; pedipalp fingers with scallop obsolescent, male fingers appearing much like that of female. Somewhat related to Vejovis terradomus Williams from which it differs in the following characteristics: smaller body size; pedipalp fingers with obsolescent scallop; pedipalp fingers considerably shorter in proportion. DESCRIPTION OF HOLOTYPE (male). Coloration: Dorsum pale yellow with faint underlying dusky markings on carapace and mesosoma; pectines whitish; walking legs whitish yellow with faint dusky marking on femur and patella. Carapace: Anterior border essentially straight but with subtle median emargination; six inconspicuous hairs; carapace surface densely granular; lateral eyes three per group. Mesosoma: Tergites densely granular; last tergite with two pairs of serrate lateral keels; last sternite with one pair of lateral keels, these smooth to subtly crenulate; stigma small, long oval. Metasoma: Inferior lateral keels smooth to irregularly crenulate on segments I to IV; inferior median keels smooth to obsolete on segments I to IV; segment I slightly wider than long; segment III longer than fixed finger; segment IV slightly longer than movable finger. Telson: Vesicle with a few short inconspicuous hairs ventrally, appearing bald and densely tubercular; subtle subaculear tubercule; vesicle more than twice aculeus length; vesicle distinctly narrower than metasomal segments. Pedipalps: Chela long and slender; palm with keels obsolescent; fingers with obsolescent scallop; narrow but distinct space between proximal 7 of fingers when chela closed; movable finger distinctly shorter than either carapace or metasomal segment V. Standard measurements and photographs: Table 4 and figures 7 and 8. 408 CALIFORNIA ACADEMY OF SCIENCES | [ Proc. 4TH SER. Ficure 7. Vejovis galbus Williams, new species. Dorsal view of holotype male. Ficure 8. Vejovis galbus Williams, new species. Ventral view of holotype male. Vol. XXXVIT] WILLIAMS: NEW VEJOVIS TAXA FROM BAJA CALIFORNIA 409 TaBLe 4. Measurements (in millimeters) of Vejovis galbus Williams, new species, holotype and allotype. holotype allotype (male) (female) Total length 34.0 38.0 Carapace, length 4.3 4.9 width (at median eyes) 3.0 7 Metasoma, length 16.5 16.6 segment I (length/width) 2.3/2.4 2.3/2.6 segment II (length/width) Dt Des 2.7/2.6 segment III (length/width) DISD Ah 2.8/2.5 segment IV (length/width) 3.6/2.3 316/225 segment V (length/width) 5.1/2.4 Seay 2S Telson, length 4.6 By Vesicle (length/width) 3/9 35/23 depth iL5) 1.8 Aculeus, length 14 wa Pedipalp Humerus (length/width) SAY 3.7/1.2 Brachium (length/width) Bell fos) 4.2/1.6 Chela (length/width) 5.7/1.4 6.3/1.8 depth 1.6 Beil movable finger, length 3.4 3.8 fixed finger, length DS 2.7 Pectines teeth (left/right) 17/18 iA DESCRIPTION OF ALLOTYPE (female). Essentially the same as holotype in color and morphology with the following exceptions: body size larger; hairs on patella and tarsomeres of walking legs longer; inferior lateral keels of metasoma more crenulate; dorsal keels of metasomal segments IV and V with several long conspicuous hairs (hairs lacking in holotype); vesicle set with abundant long conspicuous hairs ventrally; metasoma proportionally broader, segment I much wider than long, segment II slightly longer than wide; metasomal segment V approximates telson in length; pectines smaller, with fewer teeth. Standard measurements: ‘Table 4. VARIATION WITHIN PARATYPES. Study of 18 paratopotypes (9 males, 9 fe- males) indicated little variation from the description of holotype and allotype. Males varied in total length from 18 to 35 millimeters, females varied from 19 to 37 millimeters. Pectine tooth counts varied from 12 to 15 in females and from 16 to 18 in males. Younger instars have dusky dorsal pigmentation more distinct. 410 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. TYPE DATA AND ETYMOLOGY. The holotype, allotype and 18 paratopotypes were collected 5 miles south of Loreto, Baja California Sur, Mexico, on 16 May 1969 by S. C. Williams. The holotype and allotype are permanently deposited in the California Academy of Sciences. This species is named “‘galbus” because of its more or less uniform yellow coloration. ReEcorps. Known only from holotype, allotype, and paratopotype location. Vejovis viscainensis Williams, new species. (Figures 9, 10.) Diacnosts. Medium to large pale yellow species of Vejovis generally related to scorpions of the “eusthenura” group but differing by the very fine crenulation of the dorsal, dorso-lateral, and inferior lateral keels of the metasoma. Diagnostic characteristics are: male and female with vesicle not conspicuously hirsute; vesicle distinctly elongate, this more so in male; aculeus elongate and well curved; movable finger of pedipalp longer than carapace; pedipalp with hand very elongate and narrow; metasoma with inferior lateral keels very finely crenulate along entire length; each dorsal keel of segment II with about 25 to 30 fine crenulations (about half this number in other members of the eusthenura group) ; fingers of pedipalp lacking scallop, fingers meet only at distal ends when chela closed. Is somewhat related to Vejovis schwenkmeyeri Williams and Vejovis diazi Williams but differs by having more elongate pedipalp fingers and a lack of scallop on fingers. DESCRIPTION OF HOLOTYPE (male). Coloration: Entire body pale yellow; carapace and mesosomal tergites with very faint dusky underlying pigmentation; some dusky pigmentation concentrated in interocular area; pectines whitish. Carapace: Essentially straight but with subtle median emargination; anterior border set with two pairs short inconspicuous bristles; entire surface densely granular. Mesosoma: ‘Tergites densely granular; tergite 7 with two pairs of finely serrate keels; sternite 7 with one pair crenulate keels; stigma long oval. Metasoma: Dorsal and dorso-lateral keels very finely serrate; serrations irregular; inferior lateral keels finely crenulate on segments I to IV; inferior median keels smooth on segments I and II, smooth to finely crenulate on III, finely crenulate on IV; inferior median and inferior lateral keels of segment V low and very finely crenulate; segment I slightly longer than broad. Telson: Long and slender, ventral surface with a number of short incon- spicuous hairs, vesicle generally not appearing hirsute; ventral surface finely tubercular with broad subaculear tubercule; vesicle about three times aculeus in length. Vol. XXXVIIT] WILLIAMS: NEW VEJOVIS TAXA FROM BAJA CALIFORNIA 411 Ficure 9. Vejovis viscainensis Williams, new species. Dorsal view of holotype male. Pedipalps: Chela long and slender, palm not swollen; fingers not scalloped; movable finger more than twice underhand length; small proximal space between fingers when chela closed; keels on palm smooth to obsolete; movable finger slightly longer than brachium. Standard measurements and photographs: Table 5 and figures 9 and 10. DESCRIPTION OF ALLOTYPE (female). Morphologically the same as holotype with the following exceptions: larger in total body size; pectines smaller and with fewer teeth; inferior median keels smooth to obsolete on segment I, smooth on II; ventral surface of vesicle with about five pairs of moderately conspicuous hairs, vesicle still does not appear distinctly hirsute; aculeus distinctly more elongate; vesicle more swollen; keels on palm obsolete. Standard measurements: Table 5. VARIATION WITHIN PARATYPES. Study of nine paratopotypes (five males, four females) indicated little significant variation from the description of holo- type. Males varied in total length from 28 to 42 millimeters, females varied from 27 to 39 millimeters. Because of some doubt about the maturity of the females, the allotype was selected from another location. Pectine tooth counts varied from 16 to 17 in males and from 14 to 15 in females. TYPE DATA AND ETYMOLOGY. The holotype and nine paratopotypes were collected 2 miles northwest of Miller’s Landing, Baja California Norte, Mexico, + fair bo CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Ficure 10. Vejovis viscainensis Williams, new species. Ventral view of holotype male. 21 June 1968, by S. C. Williams, M. A. Cazier, and party. The allotype was collected 14 miles south of Guerrero Negro, Baja California Sur, Mexico, 16 April 1969, by S. C. Williams. The holotype and allotype are permanently deposited in the California Academy of Sciences. This species is named ‘‘viscainensis” because it appears widely distributed through, and endemic to the Viscaino Desert of Baja California. DISTRIBUTION. Viscaino Desert of Baja California from 2 miles northwest of Miller’s Landing, Baja California Norte to San Angel, 13 miles west of San Ignacio, Baja California Sur, Mexico. Recorps. Mexico, Baja California Sur: San Angel, 13 miles west of San Ignacio, elevation 500 feet, 27 June 1968 (S. C. Williams, M. A. Cazier, and party), 4 males, 5 females; San Angel, 13 miles west of San Ignacio, elevation 500 feet, 28 June 1968 (S. C. Williams, M. A. Cazier, and party), 1 male, 1 female; 1 mile east of Las Bombas, elevation 100 feet, 16 April 1969 (S. C. Williams), 20 males, 10 females; 2 miles east of Las Bombas, elevation 100 feet, 16 April 1969 (S. C. Williams), 8 males, 13 females. CoMMENT. This species is apparently restricted to the sandy habitats of the Viscaino Desert to which it appears endemic. The low frequency of encounter in the samples taken on the ground surface at night combined with the significant predominance of males in some of the samples indicate this species to spend most of its life underground in burrows. This species is of systematic interest because it appears to represent a distinct branch from the eusthenura group of Vejovis. Vol. XXXVII] WILLIAMS: NEW VEJOVIS TAXA FROM BAJA CALIFORNIA 413 TABLE 5. Measurements (in millimeters) of Vejovis viscainensis Williams, new species, holotype and allotype. holotype Pine. (male) (female) Total length 44.0 49.0 Carapace, length 5.2 5.6 width (at median eyes) 3.6 3.8 Metasoma, length 21.6 Ml segment I (length/width) 3.0/2.8 3.0/2.9 segment II (length/width) 3.4/2.6 3.6/2.8 segment III (length/width) 3.7/2.6 3.8/2.7 segment IV (length/width) 4.8/2.5 4.9/2.6 segment V (length/width) 6.7/2.4 6.8/2.4 Telson, length 6.0 6.4 Vesicle (length/width) 4.3/1.8 4.1/2.2 depth 5 1.8 Aculeus, length led 23} Pedipalp Humerus (length/width) 4.6/1.2 5.0/1.5 Brachium (length/width) 5.0/1.6 5.6/1.9 Chela (length/width) 1A/1.2 8.2/1.4 depth 1.4 1.6 movable finger, length Bed 5.9 fixed finger, length 4.2 4.8 Pectines teeth (left/right) 16/15 14/14 Vejovis hoffmanni fuscus Williams, new subspecies. Diacnosis. Related to Vejovis hoffmanni hoffmanni Williams in body size and basic morphology, especially in pedipalp and telson structure, but readily distinguished by the following characteristics: carapace and dorsum conspicu- ously marked with dusky coloration; telson with more hairs on vesicle; inferior lateral keels of metasoma with tendency to be more crenulate than smooth (not more smooth than crenulate). DESCRIPTION OF HOLOTYPE (male). Coloration: Base color of cuticle light yellow; dorsum of carapace and mesosoma with dark dusky underlying markings, these somewhat marbled; dorsal intercarinal spaces of metasoma with faint dusky markings; vesicle light orange; walking legs with femur and patella with irregular dusky markings; pedipalps with light dusky markings; pectines whitish; inferior median keels of metasoma with faint dusky outline on segments III and IV. Carapace: Anterior margin essentially straight; entire surface densely granular. 414 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. TABLE 6. Measurements (in millimeters) of Vejovis hoffmanni fuscus Walliams, new subspecies, holotype and allotype. holotype allotype (male) (female) Total length 34.0 31.0 Carapace, length 4.1 4.0 width (at median eyes) 3.0 3.0 Metasoma, length 15.9 13.5 segment I (length/width) 2.1/2.4 1.8/2.0 segment IT (length/width) 2.6/2.3 2.1/2.0 segment III (length/width) De8/ 23 DPV AGS segment IV (length/width) BIW 2L 3.0/1.9 segment V (length/width) 4.9/2.3 4.4/1.9 Telson, length 45 4.2 Vesicle (length/width) 2.9/1.6 2.7/1.6 depth i) 1.2 Aculeus, length 1.6 125 Pedipalp Humerus (length/width) 3.4/1.0 Soily/il0) Brachium (length/width) 3.8/1.4 3.6/1.2 Chela (length/width) 6.0/1.5 5.4/1.2 depth 1.6 1.3 movable finger, length 3.6 35) fixed finger, length Doll 2.6 Pectines teeth (left/right) 20/18 16/17 Mesosoma: Tergites densely granulate; last tergite with two pairs of serrate lateral keels; last sternite with one pair of lightly crenulate keels; stigma small, long oval. Metasoma: Inferior lateral keels lightly and irregularly crenulate; inferior median keels obsolete; segment I wider than long; segment IV shorter than carapace; dorsal keels of segments IV and V with long conspicuous hairs. Telson: Ventral surface of vesicle with obsolescent tubercules also abun- dantly covered with long conspicuous hairs; several long hairs on dorsum of vesicle at base of aculeus; vesicle almost twice aculeus length; vesicle narrower than either caudal segments or pedipalp palm. Pedipalps: Fingers of chela with large proximal scallop; narrow open space between proximal *4 of fingers when chela closed; keels lacking on palm; movable finger slightly shorter than brachium and slightly longer than metasomal seg- ment IV. Standard measurements: Table 6. Vol. XXXVII] WILLIAMS: NEW VEJOVIS TAXA FROM BAJA CALIFORNIA 415 DESCRIPTION OF ALLOTYPE (female). Essentially the same as holotype in color and structure with the following exceptions: slightly smaller in body size; inferior median keels of metasoma smooth to slightly crenulate; no scallop in pedipalp fingers; pectines smaller and with fewer teeth. Standard measurements: Table 6. VARIATION WITHIN PARATYPES. Study of 12 paratopotypes (10 males, 2 females) indicated little variation from holotype and allotype. Males varied in total body length from 12 to 36 millimeters while females were 22 millimeters. Pectine tooth counts were 15 in females, and varied from 18 to 21 in males. Most of the specimens appeared to be subadults. Younger instars usually were more darkly pigmented than older ones, especially the terminal metasomal segment, and lacked the scalloped pedipalp fingers of the mature males. TYPE DATA AND ETYMOLOGY. The holotype, allotype, and 12 paratopotypes were collected 24 miles northeast of San José de Comondu, Baja California Sur, Mexico, 15 May 1969 by S. C. Williams. The holotype and allotype are per- manently deposited in the California Academy of Sciences. This subspecies is named “fuscus’” because of its distinctive dark dorsal pigmentation. DISTRIBUTION. Found only in the volcanic area of Comondu, Baja California Sur, Mexico. Recorps. Mexico, Baja California Sur: 21 miles south of San Miguel de Comondu, elevation 500 feet, 14 May 1969 (S. C. Williams), 1 male, 2 females; 21 miles south of San Miguel de Comondu, elevation 500 feet, 15 May 1969 (S. C. Williams), 2 males. CoMMENT. This subspecies appears to represent an invasion of V. hoffmanni, a predominantly sandy desert species, into the volcanic canyon bottoms and oases regions of the peninsula. Considerable isolation within the subspecies ap- pears to occur as evidenced by apparent local color race formation in each of the known localities. Vejovis diazi transmontanus Williams, new subspecies. DiaAcnosis. Essentially the same as Vejovis diazi diazi Williams in structure and size but readily recognized by lack of red coloration to the vesicle, vesicle similar to metasoma in color. DESCRIPTION OF HOLOTYPE (male). Coloration: Base color of body and appendages more or less uniform pale yellow; pedipalp fingers light orange- yellow; obsolescent dusky marking on carapace near median eyes; pectines whitish. Carapace: Anterior margin straight, set with three pairs of long reddish hairs; entire surface densely granular. Mesosoma: Tergites densely granular; last tergite with two pairs of serrate 416 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. TaBLe 7. Measurements (in millimeters) of Vejovis diazi transmontanus Williams, new subspecies, holotype. holotype (male) Total length 39.0 Carapace, length 4.9 width (at median eyes) By Metasoma, length 19.2 segment I (length/width) 2.6/2.8 segment II (length/width) 2.9/2.7 segment III (length/width) 3.4/2.7 segment IV (length/width) 4.3/2.6 segment V (length/width) 6.0/2.7 Telson, length Safl Vesicle (length/width) 3.9/2.1 depth 1.8 Aculeus, length 1.8 Pedipalp Humerus (length/width) 4.0/1.2 Brachium (length/width) 4.5/1.7 Chela (length/width) 7.0/2.1 depth 2.4 movable finger, length 4.2 fixed finger, length Sil Pectines teeth (left/right) 18/19 lateral keels; last sternite with one pair of smooth to slightly crenulate lateral keels; stigma small, long oval. Metasoma: Inferior lateral keels of segments I to IV smooth to lightly crenulate (predominantly smooth); inferior median keels obsolete on segments I to IV; segment I slightly wider than long; segment IT slightly longer than wide; segment IIT longer than fixed pedipalp finger. Telson: Vesicle smooth and not tubercular; ventral surface abundantly covered with long conspicuous hairs; vesicle twice aculeus length. Pedipalps: Fingers of chela with deep proximal scallop; fingers touch only at distal ends when chela closed; palm without distinct keels; palm smooth and lustrous; fixed finger shorter than metasomal segment III; movable finger shorter than carapace; movable finger longer than brachium. Standard measurements: Table 7. VARIATION WITHIN PARATYPES. Study of 10 paratopotypes (7 males, 3 females) indicated little significant variation from the holotype. Males varied Vol. XXXVII] WILLIAMS: NEW VEJOVIS TAXA FROM BAJA CALIFORNIA 417 in total length from 19 to 40 millimeters, females varied from 18 to 23 milli- meters. None of the females appeared to be mature. Pectine tooth count was 15 in females, and varied from 18 to 20 in males. TYPE DATA AND ETYMOLOGY. The holotype and 10 paratopotypes were collected on the coastal sand dunes at Punta San Telmo, Baja California Sur, Mexico, 26 May 1969 by S. C. Williams. The holotype is permanently deposited in the California Academy of Sciences. This subspecies is named “transmontanus” because of its only known geo- graphical location on the eastern side of the Sierra de la Giganta which possibly isolates this subspecies. RecorpDs. Known only from the type locality. LITERATURE CITED WILLIAMS, S. C. 1968a. Scorpions from Northern Mexico: Five New Species of Vejovis from Coahuila, Mexico. Occasional Papers of the California Academy of Sciences, no. 68, pp. 1-24. 1968b. Methods of Sampling Scorpion Populations. Proceedings of the California Academy of Sciences, 4th ser., vol 36, no. 8, pp. 221-230. 1968c. Scorpion Preservation for Taxonomic and Morphological Studies. Wasmann Journal of Biology, vol. 26, no. 1, pp. 133-136. 1970. Scorpion Fauna of Baja California, Mexico: Eleven new species of Vejovis (Scorpionida: Vejovidae). Proceedings of the California Academy of Sciences, 4th ser., vol. 37, no. 8, pp. 275-332. : PGS BM. wh al ratieete oF bbengtied 7. as Che = wie a ea li i oa ria ra | ar. eI pee tee eh! ine 3 et sy oe _" i : s vat ie | _ tie aa ata : 7 7 — - <= a Fs PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XXXVII, No. 13, pp. 419-458; 8 figs.; 9 tables. October 13, 1970 ON THE DIET AND FEEDING BEHAVIOR OF THE NORTHERN ANCHOVY, ENGRAULIS MORDAX (GIRARD)* By Anatole S. Loukashkin California Academy of Sciences San Francisco, California INTRODUCTION Plankton has long been known to constitute the main food of such com- mercially important pelagic fishes as herring and sardines. However, existing literature on this subject, in regard to fish preference either for zooplankton or phytoplankton, quite often presents contradictory conclusions by different au- thors. Sometimes this is true for one and the same species from the same area and period, and even when investigation is based on the same material collected, as was the case of the Lewis-Parr controversy. It was especially difficult to ascertain whether there was a definite preference for zooplankton over phyto- plankton, or vice versa, in the diet of the species investigated. For example, food habits of the Pacific sardine, Sardinops caerulea, have been investigated by several researchers, of whom Lewis (1929) was the first. On the basis of 207 stomachs, he concluded that this fish is primarily a phytoplankton feeder. Parr (1930), using Lewis’ own data, reversed his conclusion by recognizing the zoo- plankters found in the stomach contents as “the only real objects of special pursuit.” Presence of phytoplankton in the contents, he explained, was ‘merely due to incidental ingestion with other elements of the diet.” Hart and Wailes (1932) examined 285 sardine stomachs from the waters of British Columbia. They found 2 dominant types of food: Diatomaceae and Crustacea, thus indicating the sardine to be omnivorous in its food habits. Radovich (1952), on the basis of 42 sardine stomachs from Baja California and * Manuscript received October 29, 1969. [419] 420 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. -2 hy X, a Ficure 1. Diagrammatic representation of the alimentary tract of the northern anchovy. O.—Oesophagus, C—Cardiac stomach, P—Pyloric stomach, F—Fundus of stomach, I—In- testine, PC—Pyloric caeca, A—Anus. southern California, regarded this fish as a carnivorous animal living mostly on zooplankters (chiefly copepods). An anonymous author (1952), after examination of 273 stomachs, agreed in part with Radovich’s conclusion by saying that the chief food of sardine larvae is “very early stages of small copepods.” In stomachs of the adult sardines, zooplankton was found in 100 percent and phytoplankton in only 75 percent. Hand and Berner (1959) examined 571 sardines, reporting that crustaceans are the “major food” (copepods being the most important item) which, on the average, contributed 89 percent of the organic matter, while phytoplankton con- tributed only 11 percent. This proportion of food types in the stomach contents showed a high correlation with plankton samples taken at the same place and time. Regardless of demonstrated preference in favor of zooplankters in the pre- ceding 2 cases, the sardine exhibited a dual mode of feeding, consuming both zoo- and phytoplankton, and therefore should be considered omnivorous in its food habits. The same contradictory conclusions were made by the investigators who studied food habits of the Japanese sardine—the iwashi, Sardinops melanosticta. Kishinouye (1907) concluded that the juvenile fish appeared to be carnivorous in food habits, being dependent on zooplanktonic food, and that the adult fish becomes vegetarian, feeding on phytoplankton. Deriugin (1933) assumed that the iwashi eats both types of plankton without specific discrimination, according to which is present in the water mass of a given season. In other words, he identified the iwashi as an omnivorous animal. Gail (1934) at first was of the opinion that the phytoplankters were the only basic food of the sardine in the northwestern parts of the Sea of Japan. He admitted that sardine stomachs sometimes were found filled with copepods, but Vot. XXXVII] LOUKASHKIN: ANCHOVY DIET AND FEEDING BEHAVIOR 421 n” - o ~ =x c 0 a ° ° N Other Ficure 2. Percentage distribution of dominating food items in 667 anchovy stomachs. said that this “in no measure would diminish the role of phytoplankton as the basic food of iwashi.” Two years later (1936), apparently after examination of a greater number of stomachs, he came to regard the zooplankton (copepods) as the preferred food of the Japanese sardine. Koganovskaia (1934) recorded dominance of one or the other type of plankton in the sardine stomachs by season (zooplankton in May, June, and October, and phytoplankton in August and November) in direct relation to the presence of these food types in the associated plankton samples. Koganovskii (1935) concluded that the iwashi is not selective in its food habits. “It feeds on phyto- and zooplankton with mass predominance of either [in their stomachs] in direct dependence on their predominance in the surrounding water. In cases where fish descended to near-bottom layers in shal- lows, benthic crustaceans and even detritus were found in the fish stomachs.” Brodskii and Iankovskaia (1935) reported predominance of zooplankters (chiefly copepods and protozoans) in 110 stomachs examined. The number of distended stomachs filled with the crustaceans bore direct relation to the amount of zooplankton in the sea-masses and of copepods in the plankton hauls. With phy- 422 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. toplankton they observed a nonconformity between the quantities in the plankton samples and the stomachs. These authors assumed that the role of phytoplankton in the diet of the Japanese sardine “‘can be fairly large in general (although second to copepods), but is reduced to the role of the so-called ‘forced diet’ in the absence of concentrations of zooplankton.” Brodskii (1936) speaking of the food of the iwashi in Possiet Bay, mentioned that it included almost all the microplankton of the area investigated, but that stomachs ‘“‘filled to capacity were noted only in the presence of Copepoda.”’ Predominance of copepods over the other food items was at times exceptionally high, reaching 95—100 percent. Iankovskaia (1937) confirmed the outstanding role of zooplankters, chiefly copepods, in the sardine diet in conformity with the abundance of them in the sea; in almost every case, greater abundance of plant organisms in the sea coincided with low filling of the stomachs. The above investigations clearly indicated a dual mode of feeding in the Japanese sardine, with, however, preference for the animal food. The Indian sardines Sardinella gibbosa and S. longiceps are regarded as carnivorous animals feeding on zooplankton, chiefly on copepods (Madras Fish- eries Department, 1933, 1936). According to Davies (1957), the South African pilchard, Sardinops ocellata, consumes phytoplankton “‘to far greater extent than the zooplankton during the greater part of the year.” The year’s mean ratio was found to be 2:1. He examined 16,664 stomachs during 1953-56, and concluded the South African sardine to be a vegetarian which also feeds on zooplankton “at times when abundance of phytoplankton has become diminished.” In reference to zoo- plankton Davies stated that “the most important group in the zooplankton eaten by pilchards is the copepoda: copepods were eaten throughout the year but occurred in stomach contents in largest numbers in summer and in winter.” In the opinion of the present writer, if plant food is equal to 67 percent of the total, and animal food estimated at 33 percent, the South African sardine falls in the category of an omnivorous animal, regardless of a statistical “preference”’ for phytoplankton. Regarding the feeding habits of the (Atlantic) European sardine, Sardina pilchardus, we refer to the investigations conducted by Hickling (1945) who examined 2,400 stomachs of pilchard taken off the coast of Cornwall in 1935-38. He established that these fish feed seasonally on either of the two types of plankton. Among the most important and consistently recurring organisms, Hickling listed (in order of importance) copepods, euphausiids, mysids, amphi- pods, and many other larval and young crustaceans. Next in importance were found to be diatoms and peridinians. Once again we can see a dual mode in food habits, thus letting us classify Cornish pilchard as an omnivorous animal dis- playing preference toward animal food. Vor. XXXVIT] LOUKASHKIN: ANCHOVY DIET AND FEEDING BEHAVIOR 423 Ficure 3. Percentage frequency of food type incidence in 840 filled stomachs of the anchovy. With such a confusing picture of the food habits of the ecologically related pelagic species in mind, the present studies on feeding behavior of the Northern Anchovy, Engraulis mordax, were initiated in 1965 and concluded in 1968. It was intended to find out, as accurately as possible with the opportunities avail- able, the actual diet of our anchovy, its mode of feeding, and its ability to dis- criminate food types. Material presented on the following pages is based on the collections of stomachs, on visual observations in aquaria and in the ocean, and on review of the available, though scant, literature. MATERIAL Three basic collections of anchovy stomachs were made by the writer, and 3 supplementary ones were provided by other collectors. The first collection was made in Monterey Bay, central California, in May 1965, through assistance of Mr. Tom Arcolea, a live-bait fisherman in Monterey. One hundred and thirty- 424 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. nine live anchovies, ranging in size from 87 mm. to 215 mm. standard length, were randomly picked out of his night catches in lampara nets. The second basic collection was made during the survey cruise of M/V Alaska of the De- partment of Fish and Game on September 15—October 3, 1965 in northern Baja California, Mexico, between Acme Rock and the California-Mexico border. Electric light and blanket net, and midwater trawl were used as collecting gear. Four hundred and nine anchovies, ranging in size from 43 mm. to 149 mm. standard length, were obtained and their stomachs preserved for analysis. The third basic collection was made in 1966 during the October survey cruise of the same research vessel from Geronimo Island in northern Baja California to Point Conception in southern California, when 220 stomachs were obtained from an- chovies ranging from 50 mm. to 155 mm. standard length which were collected using chiefly the midwater trawl. In addition to these basic collections, 3 small supplementary ones were made by other persons. Dr. Reuben Lasker contributed 2 sets of stomach contents from northern Baja California, one consisting of 20 specimens from fish 121 mm. to 138 mm. standard length collected at Bahia Soledad in the month of June, 1965, and the other one numbering 40 specimens from Ensenada Fisheries, collected in October of the same year (no information on the size of the fish was provided). The third supplement came from southern California waters. It consisted of 98 frozen anchovies, 60 mm. to 134 mm. standard length, collected and preserved by Mr. Deon L. Hamilton while aboard M/V Alaska, surveying fish populations in January-February, 1968, mostly in Santa Monica Bay. METHODS Fishing gear used during this study consisted of midwater trawl, lampara net, blanket net, electric light and blanket net (at night)!, dip net, and hook-and-line (treble) for snagging fish. The trawl was used most generally. The other types of fishing gear mentioned were used occasionally. Immediately after the fish were caught, 10 to 25 specimens were randomly picked out. Their stomachs (cardiac, pyloric, and fundus) including esophagus were removed, (fig. 1), freed from fat, slightly rinsed in cold salt water, and preserved in 4 percent formalin. All stomachs of a given sample were kept in a jar bearing neces- sary information on the label inserted in, or attached to, the jar. No con- current plankton sampling was done. In the laboratory, the contents of each stomach were removed and placed in individual vials for macroscopic and micro- scopic examination. All items of the contents of individual stomachs were identi- fied in as great detail as practicable, their relative proportions determined by wet volume, and recorded under 3 categories: dominating, second in order, and insignificant. Then all individual records were grouped in a sample table to 1The blanket net and its use in sampling populations of commercially important pelagic fishes was described in detail by Radovich and Gibbs (1954). Vor. XXXVIT] LOUKASHKIN: ANCHOVY DIET AND FEEDING BEHAVIOR 425 show the frequency of occurrence within the sample, and finally all samples of a given collection were tabulated in 6 tables (nos. 2 through 7). Evaluation of preferential tendencies of the anchovy in its diet was based on those stomach contents in which predominance of a certain food type over the others was clearly evident. RESULTS Altogether, contents of 926 stomachs were examined and, in accordance with the amount of food found, were classified as empty, very poor, poor, filled to full capacity, % capacity, 2 capacity, and % capacity. The majority of the stomachs fell into categories “poor” (299) and “very poor” (218) and 86 stomachs were empty (table 1). The best filled stomachs were found in fishes caught during the day or in fishes caught at night attracted by an electric light under which they were feed- ing on plankton swarming within the illuminated zone. Less filled stomachs and most of the empty ones were found in fishes caught in the midwater trawl during night sampling. Baxter (1967) also recognized anchovies ‘‘chiefly” as daytime feeders. A dominance of one type of food over the others (by wet volume) was recorded in 667 stomachs (tables 1 and 8). In a spring collection from Monterey Bay (May 1965), dominance was recorded in 81 out of 139 stomachs. The zooplanktonic forms (64.2 percent) definitely exceeded the phyto- planktonic organisms (35.8 percent) (table 2). In the fall collection from north- ern Baja California (September 1965) of 409 stomachs, dominance was recorded in 283 in favor of zooplankters (98.5 percent), and foreign matter (1.41 per- cent). On no occasion were the phytoplankters found in significant volume (table 3). In the autumn collection from southern California made in October of 1966, dominance was recorded in 165 out of 220 stomachs. This dominance was exclusively in favor of zooplankters. The phytoplanktonic organisms on only one occasion occupied second place (table 4). In supplementary collections, dominance of certain food types was recorded as follows: in the June collection of 20 stomachs (1965) from northern Baja California dominance was recorded in 18, with precedence of the phytoplankters (83.33 percent) over the zoo- planktonic organisms (16.67 percent) as shown in table 5. In the fall collection of 40 stomachs from the same area, dominance was evident in 23 stomachs in favor of zooplankters alone (table 6). In the winter collection from southern California (January—February 1968) dominance was displayed in 97 out of 98 stomachs in favor of zooplanktonic forms alone; however, in 33 cases the phyto- plankton occupied second place (by wet volume) (table 7). In 667 cases of recorded dominance of zooplankters the top priority belongs to crustaceans (71.81 percent), the second place being taken by other zooplank- ters (11.54 percent), and the third place by indeterminate zooplanktonic remains and fleshy parts (9.46 percent). The fourth in subordinate order was phyto- [Proc. 4TH Ser. 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XXXVII] LOUKASHKIN: ANCHOVY DIET AND FEEDING BEHAVIOR 427 TABLE 2. Frequency of occurrence of components of anchovy stomach contents, Monterey Bay, central California, collected in May, 1965. Of 139 stomachs in 9 samples, 26 were empty, 73 very poor, 5 poor, 17 half-filled, 2 filled to *, capacity, and 16 to capacity (2 being “gorged”). Dominance of one type of food over the others was found in 81 stomachs. Frequency of occurrence Description Dominating Second Place Insignificant Total (Number and Percent) Copepods: large = = 1 1 small = = 8 microcalanids = = 1 Al eggs - = 15 15 fleshy parts - - 6 6 Mysids = = 1 1 Shrimps, small adults = = 2 2 Crustacean eggs (?) = = 11 11 fleshy parts (?) 19 (23.46%) 2 16 Bil remains (?) 11 (13.58%) = = 11 Thaliaceans: Salpa 5 @6xi7%)) 6 16 27 Polychaete worms = = 1 1 Chaetognath worms = = 1 1 Hemicordata: Enteropneusta larvae 4 ( 4.94%) 2 29 35 Fish larvae - - 1 1 Pteropods: Limacina sp. - ~ 1 1 Medusae _ - 1 1 Indeterminate fleshy material (?) 13 (16.05%) 14 3 30 Eggs (?) - - 1 1 Protozoans: Silicoflagellates - = 1 1 Algae (kelp-like) fragments 1122395) - 4 5 Diatoms 28 (34.57%) 25 16 69 Anchovy scales (foreign matter) - - 8 8 Total 81 (100.00% ) 49 144 274 Broken by Groups: Crustaceans 30 (37.04%) 2 61 93 Other zooplankters 9 (11.11%) 8 51 68 Indeterminate flesh and eggs 13 (16.05%) 14 4 il 52 (64.20%) 24 116 192 Phytoplankton 29 (35.80%) 25 20 74 Foreign matter = = 8 8 Total 81 (100.00% ) 49 144 274 428 CALIFORNIA ACADEMY OF SCIENCES TABLE 3. Frequency of occurrence of components of anchovy stomach contents from northern Baja California, Mexico, collected in September, 1965. samples collected, 35 were empty, 122 very poor, 127 poor, 26 filled to “4 capacity, 40 half- filled, 33 filled to *4 capacity, and 26 filled to capacity (2 being “gorged”). Dominance of one type of food over the others was found in 283 stomachs. [Proc. 4TH Ser. Frequency of occurrence Of 409 stomachs in 29 Dominating Second Description (number and percent) Place Insignificant Total Euphausiids: adult 40 14.14% 1 7 48 eggs _ _ - iW) 17 larvae - _ - 45 45 fleshy parts 8 2.82% 5 13 26 (mixed)—adult and fleshy parts 80 28.27% 4 5 89 128 45.23% 10 87 225 Copepods: large 18 6.36% 10 43 (Al small 1 0.35% 4 71 76 microcalanids 2 0.71% - 100 102 small and microcalanids 11 3.89% 1 6 18 eggs ~ - - 68 68 larvae - - - 18 18 fleshy parts 20 7.06% 4 29 53 adult all types and flesh 47 16.61% 1 - 59 remains 1 OSI - 8 9 100 35.33% Sil 343 474 Other crustaceans: Mysids: adult 1 0.36% - 7 8 larvae ~ - - 3 3 (mixed)—adult and fleshy parts Be (0) (NGG - = 2 Amphipods: adult 2 OGG 4 8 14 larvae in capsule - - - if 1 Isopods: adult = = = 5 5 Cumaceans: adult = = _ 2 2 Porcellanid crab larvae - - ~ 2 2 Hermit crab larvae - = - 1 1 Brachyuran zoea “= - = 32 32 Brachyuran megalopa = = = 16 16 Small shrimp, adults ~ - - 2 2 Small shrimp, larvae - - - 2 2 Ostracods - — - i 7 Balanus (cyprid stage) larvae - - - 8 8 Cladocerans (Podon) = = = 3 3 Unidentified crustacean larvae - - - 6 6 Crustacean fleshy parts (ee ke) - - 6 Ant: 4 105 120 Vot. XXXVII] LOUKASHKIN: ANCHOVY DIET AND FEEDING BEHAVIOR 429 TABLE 3 ( Continued) Frequency of occurrence Dominating Second Description (number and percent) Place Insignificant Total Pelagic worms: Sagitta = = = 10 10 Other Chaetognaths 1 0.36% - 14 15 Polychaetes, adult 10 3.53% 6 12 28 Polychaetes, eggs = = = ii 7 Polychaetes, larvae = = = 9 a) Polychaetes, egg pouches and eggs ~ - - 2 2 Hemichordata (Enteropneusta) larvae - = 1 62 63 Nemertean worms (pelagic stage) ~ ~ ~ 2 2 11 3.89% 7 118 136 Mollusca: Pteropods (Limacina sp.) = ~ ~ 18 18 Heteropods ~ - - 3 3 Gastropod larvae - = = Bivalve larvae - = ~ 12 12 Cephalopods: squid larvae - - - 20 20 Cephalopods: octopus larvae - ~ ~ 2 2 - - - 63 63 Vertebrates: Fish eggs ~ - - 19 19 Fish eggs with larva inside - - - 2 2 Fish larvae 4 143% ~ 7 11 Fish remains 1.06% 2 6 11 Anchovy eggs - . - 1 1 Anchovy eggs with larva inside ~ — - 1 1 7 249% 2 36 45 Other zooplanktonic forms: Cephalocordate larvae - - = 8 8 Protozoans: Tintinnids - - ~ 9 9 Protozoans: Foraminiferans - - - 3 3 Protozoans: Radiolarians ~ - - 4 4 . Protozoans: Silicoflagellates - ~ 1 1 Hydroid, chunks - - - 1 1 Appendicularians (Ozkopleura) HUG 6 57 68 Thaliaceans: (Salpa) 3 1.06% 4 84 91 Thaliaceans: (Salpa) with embryo - - - 7 7 Thaliaceans: Doliolids - = — 8 8 Bryozoan “cyphonautes” larvae D2 Onl 5 72 79 430 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. TABLE 3 (Continued) Frequency of occurrence Dominating Second Description (number and percent) Place Insignificant Total Jellyfish (medusae, ctenophores, etc.) - - - 19 19 Echinoderms: sea cucumbers - — 1 10 11 sea urchin parts - - - 1 1 Brachiopod larvae - - - 10 3.52% 16 287 Sil} Phytoplankton: Diatoms (“present” only) - = = 65 65 Dinoflagellates (“present” only) ~ - - 19 19 Algae (Phaeocystis) - - - 6 6 Algae tissues (kelp-like) - - - 45 45 - - - 135 135 Unidentified material: . Fleshy parts (?) 12 4.24% - 41 53 | 12 4.24% - 41 53 Foreign matter: Insect = = = 1 1 Fine sand grains 035% - 27 28 Fine mica flakes — - 1 24 25 Fine sand and mica, mixed 3 1.06% 12 8 WS) Minute pebbles ~ - - 2 2 Pollen grains - - - 1 1 Fish scales - - - 1 1 4 141% 13 64 81 Parasites: Nematodes ~ ~ - 18 18 Total 283 100.00% 83 1297 1663 Broken by groups: Crustaceans, all 239 84.45% 45 535 819 Other zooplanktonic forms 28 9.90% DS 504 57 Indeterminate fleshy matter 12 4.24% ~ 41 53 Total, zooplanktonic material 279 98.59% 70 1080 1429 Phytoplankton - - - 135 135 Foreign matter 4 141% 13 64 81 Grand Total 283 100.00% 83 1281 1645 Parasites - - = 18 18 Vor. XXXVII] LOUKASHKIN: ANCHOVY DIET AND FEEDING BEHAVIOR 431 plankton (6.59 percent), and the last place was taken by foreign matter (only 0.60 percent) (fig. 2). As to predation of the northern anchovy on small fish, analysis of stomach contents revealed 104 cases of fish eggs and larvae, mostly of its own kind. In 8 cases these items were found to be the major food, 3 times they occupied second place, and 93 times they were present in meager quantities. Baxter (1967) also observed this pattern. Among crustaceans found in dominating volumes in the stomach contents, the first place belongs to copepods and the second to euphausiids (all developmental stages from egg to adult form in both groups) .” The same trend is illustrated in table 9 which shows food-type incidence in 840 filled stomachs (recapitulation of tables 2 through 7) regardless of the quantity of food present in the stomach. In this case, out of 98 food type entries on the menu of our anchovy, only 11 represented phytoplankton and _ foreign matter. By contrast zooplankters constituted 86.54 percent, of which 50.78 percent were crustaceans. In the latter group, copepods were again most abun- dant (29.32 percent),* while the euphausiids were second in frequency (9.54 percent). Incidence of each of the rest of the 24 forms of crustaceans was very low. The phytoplankters made up only 10.99 percent of the total (fig. 3). OBSERVATION ON FEEDING BEHAVIOR From actual visual observations both in the field and in the tanks of Steinhart Aquarium, and from the examination of the stomach contents, it is clearly evident that the northern anchovy in all size categories is both a filter feeder and a particulate feeder,! depending on the size of the available food organisms (Miller, 1967, 1968). As an example of filter feeding, reference is made to a field ob- servation of the writer recorded on September 26, 1965, at San Jose Point in Baja California, which reads as follows: “at 9:00 a.m. a huge school of juvenile anchovies around anchored Alaska was actively feeding on some microplanktonic organisms very close to the surface. For two hours we watched this interesting behavior. Water was very clear and we could see all movements and feeding pattern of the school. It was a sort of a loose formation in which individual members were sparsely located, however all being orientated toward east. With mouths wide open and gill-covers stretched out, they strained water. They did this while rising toward the surface in a rhythmic wave-like pattern. When they headed down, they closed their mouths and tightly pressed gill-covers to the gills, as if they were swallowing organisms trapped by gill-rakers. With line and treble hook I snagged 12 young fish 67-91 mm. standard length.” 2 Baxter (1968) recorded euphausiids as the most common food object in the stomach contents of the anchovies collected at night. It can be added that this is especially true when electric light and blanket net are used as collecting gear. 3 A list of partially identified copepods is presented in appendix A. 4 Experimental studies, conducted at the California Current Resources Laboratory of the U.S. Bureau of Commercial Fisheries in La Jolla, revealed that predation is by filtering on organisms less than 1 mm. in length and by particulate biting on organisms a few mm. in length. The larger organisms are preferred. (Anonymous, 1967). CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. bo 46 .98% Ficure 4. Percentage frequency of occurrence of various types of organisms found in the stomachs of 273 sardines in relation to total number of occurrences. Based on table 1 of Cadet H. Hand and Leo Berner, Jr. (1959). Later, stomach contents of these fish were examined in the laboratory. Ten of them were filled up to *% capacity, and 2 were “poor” in contents. Their stomachs contained small copepods, microcalanids, copepod eggs and nauplii, and bryozoan ‘‘cyphonautes” larvae. Diatoms (Coscinodiscus species) were also present in 5 stomachs, but in very insignificant numbers. Selective or particulate feeding also was observed in the sea many times, usually when electric light at night was used as attractant and blanket net as catching gear. As an example, a record is given below dated October 19, 1966, for night anchorage of the Alaska in Government Cove (near Santa Barbara): “”. . About 11:00 p.m. Alaska dropped her anchor in Government Cove (southern Cal- ifornia). A 1500-watt lamp was suspended from the boom about 12 feet above the water surface. Soon after midnight swarms of planktonic organisms appeared within the illuminated Vor. XXXVII] LOUKASHKIN: ANCHOVY DIET AND FEEDING BEHAVIOR 433 zone. Large euphausiids, chaetognaths, large copepods, and many kinds of other smaller organisms were abundantly represented. About 2:00 a.m. a school of anchovies made its appearance, and after a while a wild chase started. The illuminated area seemed boiling. The fish feeding on the surface uninterruptedly splashed water with their tails. There was a “disorderly pattern” of feeding; it seemed that each member of the school acted individually, swimming back and forth, and up and down, and attacking its prey and snapping at it at irregular short-time periods. The fish were so wildly “preoccupied” with their chase that I had no difficulty in catching them with a scoop net. Twelve anchovies 135-145 mm. stan- dard length were so obtained. Two of the stomachs were * filled, while the rest were simply “gorged,” and when cutting the abdomens the stomachs “blew up,” pouring the con- tents on the table.” Examination of the contents later in the laboratory disclosed that euphausiids were the dominating food component in all 12 stomachs. They represented 75 percent to 95 percent of the wet volume of the contents in the individual stom- achs. Their numbers varied from 11 to 15 in the stomachs filled up to %4 capacity, and from 38 to 180 (including young and larvae) in the stomachs filled to extreme capacity (“gorged’’). Second place was occupied mostly by the euphausiid soft parts (8 times), 3 times by large copepods, and once by a polychaete worm broken to pieces. Many other zooplankters of smaller size, found in the stomach contents in meager numbers, might have been ingested by the anchovy while feed- ing on larger animals. No diatoms were found at all. Visual observations on feeding behavior of the schools of northern anchovy and Pacific sardine kept in the past for experimental purposes in the 1000-gallon tanks of the Steinhart Aquarium, revealed the very same dual tendency in feeding patterns of these 2 species. When live brine shrimps, Artemia salina, were added to the tank, the school (sardines or anchovies alike) would rush at the dense cloud of shrimps, disperse them at once, and then begin to snap up the individual animals. When just-hatched brine shrimps were added to the tank, the sardines or the anchovies would proceed with filter-feeding. Again, when these fishes were fed with finely chopped or ground horse heart, ground fish, or commercially manufactured dry fish pellets containing protein, they demonstrated ability to turn to particulate feeding. The examination of the contents of the entire set of stomach collections described above exposed a great diversity of food objects on the menu of the anchovy. It became evident that it feeds on many types of planktonic organisms, both of animal and plant origin. This examination of the components of the stomach contents confirmed once again that the anchovy is both a filter feeder and a particulate one. In addition to this, an observation was made during the present study which may indicate an aggressive character in the feeding behavior of the anchovy. In several stomachs of a single sample there were found chunks of larger poly- chaete worms, a fact that may suggest the possibility of a group attack by several anchovies on a larger animal which they tore to pieces, a single fish being unable to swallow the entire worm. 434 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Taste 4. Frequency of occurrence of components of the anchovy stomach contents from northern Baja California, Mexico, and southern California waters collected in October, 1966. Of 220 stomachs in 21 samples collected, 23 were empty, 21 very poor, 53 poor, 44 filled to 14 capacity, 37 half-filled, 24 filled to *4 capacity, and 18 to full capacity (8 being “gorged”). Dominance of one type of food over the others was found in 165 stomachs. Frequency of occurrence Dominating Second Description (number and percent) Place Insignificant Total Euphausiids: Adult 37 22.43% 9 23 69 eggs ae = 9 9 larvae — - ~ 17 17 fleshy parts 1 0.60% 24 17 42 (mixed)—adult and fleshy parts Dil WPS 1 ~ 22 59 35.76% 34 66 159 Copepods: large 3 1.83% 19 52 74 small 6 3.64% 4 98 108 microcalanids = ~ 2 85 87 eggs - _ - 72 72 larvae - - ~ 35 35 Herpacticoid adult - = — 1 if fleshy parts 29 17.57% 12 38 79 (mixed)—large adult and fleshy parts 4 242% - ~ 4 (mixed)—small adult and fleshy parts 9 5.45% 2 _ 11 51 30.91% 39 381 471 Other Crustaceans: Mysids: adult 1 0.60% 5 11 17 larvae - - - 4 4 Amphipods: adult Bo NBG: 7 44 53 larvae in capsules - - - 6 6 Phyllopods (Spiny lobster larvae) ~ ~ - 4 4 Tsopods: adult - - 1 11 12 Porcellanid crab larvae - — - 10 10 Hermit crab larvae — = = Sand crab larvae - - - 1 1 Brachyuran zoea - = = 45 45 Brachyuran megalopa - _ - 35 35 Coridean shrimp = = = 1 1 Small shrimp - ~ - 1 Sergestid shrimp larvae — - - 1 1 Ostracods 1 0.61% _ 12 13 Balanus (cyprid stage) larvae = = = 9 9 Cladocerans (Podon) = = = 10 10 Crustacean eggs (?) = = = 1 1 Crustacean larvae (?) = = = 24 24 Vor. XXXVII] LOUKASHKIN: ANCHOVY DIET AND FEEDING BEHAVIOR 435 TABLE 4 (Continued) Frequency of occurrence Dominating Second Description (number and percent ) Place Insignificant Total Crustacean fleshy parts (?) 6 3.64% 1 3 12 Crustacean remains 3 1.82% ~ 1 4 13 7.88% 14 242 269 Pelagic worms: Sagitta = = = 9 Other chaetognaths - ~ - 8 8 Polychaete adults 8 4.85% 3 25 36 eggs - - 1 2 3 larvae - ~ - 3 3 egg pouch with eggs 2 1.21% — ~ 2 fleshy chunks 1 0.60% 2 - 3 Hemichordata (Enteropneusta) larvae ~ ~ — 35 35 11 6.66% 6 82 99 Mollusca: Pteropods (Limacina sp.) - - - 40 40 Heteropods ih (OMG: - 9 10 Gastropod larvae ~ - - 11 11 Bivalve larvae - ~ ~ 5 5 Cephalopod larvae - = = 1 0.61% = 74 75 Vertebrates: Fish eggs ~ - 1 35 36 larvae and fish flesh - - ~ 7 7 post-larval stage 1 0.61% - - 1 Anchovy eggs ~ - - 1 1 with larva inside - - - 1 1 1 0.61% 1 44 46 Other zooplanktonic forms: Protozoans: Tintinnids - - - 9 9 Dictocystis - ~ ~ 1 1 Salpiginella ~ - - 2 2 Parafavella - - = 1 1 Ciliates - = = 3 3 Foraminiferans ~ - = 1 1 Radiolarians Be WANG - 20 22 Silicoflagellates - - - 2 Appendicularians: Oiko pleura 13 7.88% 7 54 74 Fritillaria - - = 1 1 436 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. TABLE 4 (Continued) Frequency of occurrence Dominating Second Description (number and percent) Place Insignificant Total Thaliaceans: Salpa 9 545% 10 85 104 with embryos 1 061% 1 5 7 with aggregates 2 PIG 1 - 3 Doliolids - - - 13 13 Bryozoan “cyphonautes” larvae - - - 30 30 Jellyfish (medusae, ctenophores, etc.) - - ~ 31 31 Echinoderm larvae (sea urchins) - - - 6 6 Echinoderm adult (sea cucumbers) - - - 2 iz 27 16.36% 19 266 312 Indeterminate zooplanktonic matter: Fleshy soft parts (?) 2 1.21% - 4 6 Eggs (?) - = - 14 14 Larvae (?) ~ - - 4 4 2 W200 = 22 24 Phytoplankton: Diatoms - - 1 M51 (PES) 76 Dinoflagellates - - - Pi (Gave) Dil Algae: Phaeocystis - = = 4 (pr.) 4 tissue (fragments of kelp-like algae) - ~ = 25 25 - - 1 130 132 Foreign matter: Fish scales - - ~ 1 1 Fine sand grains - - - 6 6 Fine mica flakes — - - 2 2 - - - 9 9 Parasites: Nematodes - - = 3 3 Total 165 100.00% 114 1320 1599 Broken by groups: Crustaceans, all 123 74.55% 87 689 899 Other zooplanktonic forms 40 24.24% 26 466 532 Indeterminate fleshy material Bo WANG 6 - 22 24 Total zooplanktonic material 165 100.00% 113 1177 1455 Phytoplankton - — 1 131 132 Foreign matter = = = 9 9 Grand total 165 100.00% 114 1317 1596 Parasites - - ~ 3 3 Abbreviation: pr. = present. Vor. XXXVII] LOUKASHKIN: ANCHOVY DIET AND FEEDING BEHAVIOR 437 TABLE 5. Frequency of occurrence of components of anchovy stomach contents from northern Baja California, Mexico, collected on June 12, 1965, by Dr. Reuben Lasker. Of 20 stomachs in collection, 1 was empty, 2 poor, 7 half-filled, 4 filled to *4 capacity, and 6 filled to full capacity. Dominance of one type of food over the others was found in 18 stomachs. Frequency of occurrence Second Description Dominating Place Insignificant Total (Number and Percent) Euphausiids: Adult = 1 1 2 fleshy parts 1 (5.56%) - = Fel! Copepods: large = = 1 1 small - = 1 1 microcalanids — = 1 i eggs - - 8 8 larvae = = 1 1 Crustacean soft parts (?) 2 (lil ie) _ 2 4 Hemichordata: Enteropneusta larvae = = 3 3 Protozoans: Tintinnids = 11 ili Indeterminate fleshy material ~ - 2 2 Diatoms 15 (83.33%) 2 2 (pr.) 19 Dinoflagellates - 1 1 (pr.) 2 Total 18 (100.00% ) 4 34 56 Broken by groups: Crustaceans 3 (16.67%) 1 15 19 Other zooplankters - - 14 14 Indeterminate fleshy material = ~ 2 Phytoplankton 15 (83.33%) 3 3 21 Total 18 (100.00% ) 4 34 56 Abbreviation: pr. = present. As to the phytoplankton in relation to 840 filled stomachs (926 minus 86 empty), diatoms were found in 295, taking dominating position in 43 cases, second place in 64, and being present in insignificant quantities in 188. Dino- flagellates once took second place and 67 times were recorded as “present.” Algae and algal tissues of kelp-like type were found once in dominating position, and 85 times were present in insignificant quantities. In certain collections, dia- toms consumed by the anchovy played a very important role in its diet. In the stomach collection from Monterey Bay, made in the month of May, 1965, dominance of diatoms over the other food types was equivalent to 34.57 percent of the total. Several stomachs were filled to capacity and because of uniform food components the stomachs were of green color. In some of these stomachs, diatoms were the only food item found, and quite often 99 percent of the diatoms in such stomachs belonged to a single form (Chaetoceros species, for instance). 438 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. In the Monterey collection, the anchovy ranged in size from 85 mm. to 215 mm. standard length, and diatoms were present in all size groups (85-100, 100— 150, 160-215). In certain samples they were present in every stomach in various quantities. In other stomachs of this Monterey collection phytoplankters and zooplankters were present more or less in equal amounts, but in a much greater number of cases, only a few specimens of diatoms were found. Dinoflagellates were recorded in meager quantities. In Dr. Lasker’s collection from Bahia Soledad made in the month of June, 1965, diatoms were found in 19 out of 20 stomachs—in 15 cases they were first in abundance, twice they took second place, and twice they were present in insignificant quantities. In some stomachs the contents were composed of diatoms exclusively, or diatoms with a small admixture of dinoflagellates. Among 11 generic forms of diatoms recorded from this collection, Chaetoceros, Coscinodis- cus, and Thalassiosira were most numerous. An explanation for the abundance of diatoms in the anchovy stomachs from Monterey, and from Bahia Soledad as well, may be found in the observations of Abbott and Albee (1967) on seasonal distribution of phytoplankton in Monterey Bay. They stated that “for the thirteen-year period, 1954-1966, maximum crops appeared most frequently in June, but in different years peaks fell in all months from March through July.” They maintain that Chaetoceros “is usually the predominant genus in the Bay in spring and early summer.” This explains why in our collection certain stomach contents were composed of diatoms of the genus Chaetoceros in amounts up to 99 percent of the total. Again, in the Bahia Soledad collection, Chaetoceros was dominant among the 11 forms of diatoms recorded for the said collection.° In studying behavioral responses of the northern anchovy to various intensi- ties of white light under laboratory conditions at the California Academy of Sciences, (Loukashkin and Grant, 1965), it was found that the fish always dis- played preference for the lower intensity regardless of any combination of the values of contrasting light intensity zones in the experimental tank. Therefore, similar behavior was expected of the anchovy in its natural habitat. Field ob- servations made by the present writer, and by observers on the staff of the California State Fisheries Laboratory of the Department of Fish and Game, often expressed in their Cruise Reports, confirmed this natural response of the anchovy. In bright sunshine schools of anchovies as a rule descend to deeper layers of water than they do in overcast weather or at times when upper layers of water are full of plankton, which would provide both a filter from the sunlight and a source of food. Presence of bottom-dwelling animals, such as certain foraminiferans, worms, © For a complete list of diatoms and dinoflagellates identified chiefly by Dr. G Dallas Hanna, see ap- pendix B. Vor. XXXVII] LOUKASHKIN: ANCHOVY DIET AND FEEDING BEHAVIOR 439 \ Ficure 5. Stomach contents of the northern anchovy no. 124 collected on September 27, 1965, 4-5 miles WNW. of Soledad Rock in Baja California, Mexico. Stomach of this 136 mm. fish (standard length) was filled with food to capacity, and its contents consisted exclusively of adult and juvenile euphausiids and their parts (84) with 3 large copepods. Only a part of the contents is shown in this picture, being greatly magnified. Picture courtesy of Mr. Allyn G. Smith, California Academy of Sciences, April, 1969. and amphipods, and an accumulation of fine sand grains and mica flakes (even tiny pebbles) in the anchovy stomachs demonstrated that this fish not in- frequently descends to near-bottom layers, especially in shallower waters, and behaves there apparently as an indiscriminate filter feeder. Referred to a category of foreign matter in figures and tables, sand grains, mica flakes, and these 2 materials mixed together were found in 85 stomachs in different samples, being present in dominating volumes on 4 occasions, taking second place 13 times, and recorded in insignificant quantities 68 times.° Within the group of foreign matter, pollen grains were included on the basis of a single occurrence in stomach no. 714 from anchovy collected near Point Dume on September 14, 1966. Presence of pollen in the stomachs of plankto- phagous species has been recorded for anchoveta and sardine: Bayliff (1963) 6 For instance, in the sample VIII, taken on September 21, 1965, at 6:00 p.m. in San Quintin Bay, 12 stomachs were filled from ‘“‘poor’’ to ‘14 capacity.’’ Presence of sand grains and mica flakes was noted in all of them: in dominant position in four, in second place in two, and in insignificant quantities in three. 440 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. TABLE 6. Frequency of occurrence of components of anchovy stomach contents from northern Baja California, Mexico, collected by Dr. Reuben Lasker on October 20 and 27, 1965. Of 40 stomachs in the collection, 2 were very poor, 18 poor, 19 half-filled, and 1 filled to capacity. Dominance of one type of food over the others was found in 23 stomachs. Frequency of occurrence Second Description Dominating Place Insignificant Total (Number and Percent) Euphausiids: eggs only ~ - 1 1 Copepods: large - - 3 3 small - - 4 4 microcalanids - ~ 3 3 eggs — = + 4 fleshy parts 20 (86.95%) ~ - 20 Amphipods - - 1 1 Thaliaceans: Salpa ~ - 2 2 Appendicularians: Oikopleura ~ - 2 2 Hemichordata: Enteropneusta larvae - - Z 2 Chaetognaths: Sagitta - - 1 1 Fish egg shell - ~ 1 1 Jellyfish larvae - ~ 1 1 Protozoans: Tintinnids - 1 3 4 Radiolarians - - 1 1 Silicoflagellates = - 1 Ht Indeterminate fleshy material 3 (13.05%) - 16 19 Diatoms - 3 iMOi((ove5)) 13) Dinoflagellates - - 2 2 Total 23 (100.00% ) 4 58 85 Broken by groups: Crustaceans 20 (86.95%) - 16 36 Other zooplankters - 1 14 15 Indeterminate fleshy material 3 (13.05%) ~ 16 19 Phytoplankton = 3 12 15 Total 23 (100.00% ) + 58 85 Abbreviation: pr. = present. found pollen grains in the diet of the anchoveta, Cetengraulis mysticetus, from the Gulf of Panama; Gail (1936) observed adult Japanese sardines in the USSR territorial waters of the Sea of Japan actively feeding on pollen grains of the Korean pine, Pinus koraiensis, (which he classified as “pseudoplankton”’). Dur- ing the blooming season he found sardine stomachs filled with masses of this pollen, rendering the stomachs bright yellow in color. It would be of interest to know if such a mass appearance of pollen on the surface of inshore waters in Vor. XXXVII] LOUKASHKIN: ANCHOVY DIET AND FEEDING BEHAVIOR 441 the California Current system has ever taken place, and how this supply has been utilized by the anchovy and other planktophagous fish. It is not unlikely that, if a greater number of stomachs were obtained, all the components of the plankton in the anchovy’s environment could be found in its intestinal tract, save for the extremely minute planktonic forms and those too large for it to ingest. The sampling problem inherent in food studies is usually inadequacy of the sampled material. Better definition of general food habits of any animal species would require the use of a greater number of stomachs. Hand and Berner (1959) listed 34 food objects found in the stomachs of 273 Pacific sardines. In our 6 collections numbering 926 stomachs the list of food items recorded reached 96, including all 34 found by them, or approximately threefold. These results are almost in direct proportion to the number of stom- achs examined. And yet, the present writer does not believe that the material collected presents a complete picture of the food habits of the anchovy. There- fore, he considers his account a preliminary one until more abundant data have been accumulated and reported. Nevertheless, despite a limited number of stom- achs in the collections taken sporadically and randomly within the area of Cal- ifornia Current System, it is clearly seen that the anchovy feeds indiscriminately on a variety of planktonic organisms available within the area occupied. The other planktophagous pelagic fish occurring in the vicinity, or mixed with the anchovy schools, would certainly feed on the same plankton available to the anchovy. This was well demonstrated in the catches of mixed lots of fish during the author’s collecting cruises aboard M/V Alaska in 1965 and 1966. On Sep- tember 21, 1965, at 10:15 p.m., using an electric light as attractant and the blanket net as fishing gear, in San Quintin Bay, Baja California, 2.8 miles off- shore in waters 17 fathoms deep, 3 species of pelagic fish were caught at a single operation. Among them were 3,000 jack mackerel, Trachurus symmetricus, 200 Pacific sardines, and 10 anchovies. All 10 stomachs of the latter were filled to capacity mostly by large copepods, with insignificant amounts of other zoo- plankters. Examination of stomach contents of 11 randomly picked sardines (3 stomachs were filled to extreme capacity—‘gorged,’ 2—% capacity, 2—'% capacity, and 4 were poor in contents) revealed identical food items in the same proportions (appendix C). In 12 stomachs of the jack mackerels 7 were filled to full capacity, 2 to % capacity, and 3 to % capacity. The dominating food item was large copepods, with euphausiids taking second place (appendix D). In sample LVIII taken on October 18, 1966, at 8:13 p.m. near Point Conception, southern California, 4 miles offshore, 7 anchovies and 2 hake, Merluccius productus, were caught in the midwater trawl. In 3 out of 7 anchovies selected (nos. 783, 785, and 788), euphausiids were in dominating position; the stomachs of the other three were filled with copepods, salps, and polychaete worms; and the seventh stomach was empty. The hake stomachs were filled with euphausiids 442 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. alone’ (appendix E). The above examples may serve as a basis for the general- ization that certain, if not all, pelagic fishes regarded as planktophagous in their food habits are indiscriminate omnivorous animals. As to the northern anchovy, the same examples present a clear picture of existence of interspecific food competition.® Just about at the outset of the present study, when no exact information on the food of the adult anchovy existed, Murphy (1966) published his exhaustive analysis of the population biology of the Pacific sardine in the California Current System. In discussion of the increase of anchovy population concurrent with simultaneous decrease of that of the Pacific sardine, he stated his personal im- pression that these 2 species “eat about the same things,” and therefore it was hypothetically reasonable to consider anchovy and sardine “fundamental com- petitors.”” This impression and postulation appear to be correct. Berner (1959), in his report on food studies of the larvae of the northern anchovy said that larvae longer than 7 mm. “‘actually ate the greatest variety of food.’’® The major food item in all larval groups consisted of crustaceans in various developmental stages, of which copepods were most important. Copepod eggs and nauplii were met with most frequently and most abundantly. The other items found in the stomachs were adult copepods, euphausiid eggs and nauplii, clam larvae, ciliates, dinoflagellates, tintinnids, coccolithophores, for- aminiferans, unidentified eggs, miscellaneous plant material, and miscellaneous unidentified material. Shumann (1963) in reference to findings by D. kK. Arthur!® states that food of the larvae of northern anchovy, Pacific sardine, and jack mackerel is ‘“‘sub- stantially the same.” From Berner and Schumann’s studies, and from our investigations as well, it appears that the northern anchovy in all age stages remains a planktophagous animal, and that it is located in a low trophic level, near the primary production link. For the Japanese anchovy, Engraulis japonica, according to Kubo (1961) and Hayasi (1967), copepods comprise the major food throughout the life span. The size of the food objects is linearly correlated with the size of the fish. Nakai et al. (1962) provided more specific details. They found out that ‘“‘postlarvae of less than 5 mm. total length, just after absorbing yolk, take mainly eggs and 7 Euphausiids are abundant and important organisms in the ocean plankton available to a variety of planktophagous pelagic fishes, ‘‘ranging from sardine and jack mackerel to tunas and salmon, and they are the chief food of baleen whales’? ((Anonymous, 1967). To this listing we may add our anchovy, hake, Pacific herring, Clupea harengus pallasi, (Mikulich, 1960), and Argentine anchovy (de Ciechomski, 1967). SIn addition to sardine, jack mackerel, and hake as anchovy competitors, we may include juvenile sable- fish, Anoplopoma fimbria, which, according to Conway ef al. (1968), feeds ‘‘primarily on small pelagic organisms, including copepods, amphipods, euphausiids, and the urochordate, Oikopleura sp.’? They charac- terize sablefish at all ages, including juveniles, as “highly opportunistic in feeding habits, ingesting whatever suitable prey species are most abundant or available in their immediate surroundings.” ® Berner examined 13,620 larvae of which only 211 had ingested food. 10“The particulate food and food resources of the larvae of three pelagic species, especially the Pacific sardine, Sardinops caerulea.’’ Doctoral thesis, University of California, Scripps Institution of Oceanography, 231 pp. (typewritten), 1956. Vor. XXXVII] LOUKASHKIN: ANCHOVY DIET AND FEEDING BEHAVIOR 443 Ficure 6. Photomicrograph of diatoms from the contents of the stomachs of the northern anchovy (Engraulis mordax) nos. 544-553. The slender spines are from various species of Chaetoceros. The dark circle on the right is Stephanopyxis turris. Monterey Bay, California, May 27, 1965. The diatoms were the chief item in the stomach contents (from 90-100 percent) rendering bright green color to the stomach. Photograph by Dr. G Dallas Hanna, California Academy of Sciences, June, 1969. nauplii of copepods. Only a few of them were found with protozoans, small mollusca, and diatoms in their digestive tracts. Postlarvae of 5-10 mm. total length still mainly eat nauplii of copepods, but copepodid larvae increase in number from before. Generally, sizes of postlarvae and food organisms are correlated with each other. Number of postlarvae with food in the digestive tract occupy less than 20 percent on the average. They take the food most actively in the day time.”'! The juveniles through adult stages (Nakai et al., 11 The same is true of the Black Sea anchovy, Engraulis encrasicholus (Duka, 1961). 444 CALIFORNIA ACADEMY OF SCIENCES [PRroc. 4TH SER. Taste 7. Frequency of occurrence of components of anchovy stomach contents from southern California waters collected by Mr. Deon L. Hamilton in January-February, 1968. Of 98 stomachs in 7 samples collected, 1 was empty, 94 poor, and 3 half-filled. Dominance of one type of food over the ethers was found in 97 stomachs. Frequency of occurrence Dominating Second Description (number and percent) Place Insignificant Total Euphausiids: adult - - - 1 1 eggs = = - 1 1 Copepods: large = - 2 18 20 small - - 4 34 38 microcalanids ~ = - 38 38 eggs - - - 17 17 larvae - - - 8 8 fleshy parts 34 35.05% - - 34 (mixed)—adult and fleshy parts 20 20.62% - - 20 54 55.67% 6 lil7 177 Other crustaceans: Amphipod larvae - - - 2 2 Cumacean adult - - ~ 1 1 Brachyuran megalopa = - — 1 1 Porcellanid crab larvae - - - 2 Zz Small shrimp larvae - - - 2 2 Sergestid shrimp larvae ~ - ~ 1 1 Balanus (cyprid stage) larvae - - - 3 3 Crustacean larvae (?) - - - 10 10 Crustacean fleshy parts (?) 10 10.31% - - 10 10 10.31% - 2 32 Pelagic worms: Sagitta = S a 1 1 Polychaete worm - - 1 2 3 Nemertean worm larvae (pelagic stage) ~ ~ ~ 1 Hemichordata: Enteropneusta larvae - - - 50 50 - ~ 1 54 55 Mollusca: Pteropods (Limacina sp.) - - - 1 1 Bivalve larvae - - - 2 2 Cephalopod larvae ~ - - 2 2 - - ~ 5 5 Vertebrates: Fish eggs - = - 11 11 Vor. XXXVIT] LOUKASHKIN: ANCHOVY DIET AND FEEDING BEHAVIOR 445 TABLE 7 (continued) Frequency of occurrence at Dominating Second Description (number and percent) Place Insignificant Total Other zooplanktonic forms: Protozoans—Tintinnids = = = 2 2 Radiolarians = = = 6 6 Appendicularians—Oiko pleura — = ~ 7 7 Thaliaceans—Salpa = = = 10 10 Jellyfish = = = 7 eel Bryozoan “cyphonautes” larvae = = = 2 2 ———— = 34 34 Unidentified zooplankton: Fleshy material (?) 33 34.02% - - 3g Eggs (?) - - = 8 8 33 34.02% a 8 41 Phytoplankton: Diatoms - - 33 (v.m.) 10 (m.) 10 pr. 53 Dinoflagellates - - _ 18 pr. 18 Algae tissues (kelp type) ~ - - 1 1 = — 33 39 72 Foreign matter: Fish scales - ~ - 2 2 Fine sand grains _ _ ~ <= = 3 3 Parasites: Nematodes - - ~ 1 1 Total 97 100.00% 40 294 431 Broken by groups: Crustaceans 64 65.98% 6 139 209 Other zooplanktonic forms - - 104 105 Unidentified zooplanktonic material 33 34.02% ~ 8 41 Phytoplankton - - 33 39 HZ Foreign matter - - - 3 3 Total 97 100.00% 40 293 430 Parasites = = = 1 1 Abbreviations: v.m. = very many; m. = many; pr. — present. 1955, and Kubo, 1961) are also dependent on copepods. Other food organisms are diatoms, various types of small crustaceans, larval mollusks, chaetognaths, and other small animals. The adult Japanese anchovy eats eggs and larvae of fishes including its own, as is the case with our anchovy. 446 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47H SER. TABLE 8. Frequency of occurrence of dominating food types in 667 anchovy stomachs. Group Description Number Percent Zooplankton—Crustaceans Copepods, all stages, flesh 225 33.73 Euphausiids, all stages, flesh 188 28.18 Other crustaceans all stages, flesh 9} s5 Indeterminate crustacean flesh 43 6.45 Indeterminate crustacean remains 14 2.10 479 71.81 Zooplankton—Vertebrates Fish larvae 4 0.60 Fish remains 4 0.60 8 1.20 Zooplankton—Other animals Protozoans: Radiolarians 2 0.30 Appendicularians 18 2.70 Salps 20 3.00 Worms: Hemichordata larvae 4 0.60 Chaetognaths 1 0.15 Polychaetes 21 Bell) Mollusca: Heteropods 1 0.14 Bryozoa: Cyphonautes larvae 2 0.30 69 10.34 Zooplankton— Indeterminate Fleshy material 63 9.46 Total for Zooplankton 619 92.81 Phytoplankton Diatoms 43 6.45 Algae (kelp-like) tissues 1 0.14 Total for Phytoplankton 44 6.59 Foreign matter Sand grains, mica flakes, etc. 4 0.60 Grand Total 667 100.00 1 Mysids, Amphipods and Ostracods. A detailed account on the food habits of larvae and juveniles of the Argentine anchovy, Engraulis anchoita, was recently published by de Ciechomski in 2 arti- cles (1967a and 1967b). She stated that the basic food of fish 35-80 mm. long consists of copepods in all stages of development: eggs, nauplii, and adult forms. Their diet includes also juvenile decapods and other crustaceans, eggs of various marine organisms, and sometimes fish and mollusk eggs. In the stomach contents of juveniles of 40 mm. length she found radiolarians and acantharians, but in scarce amounts. In juveniles 50 mm. long there appeared diatoms and dinoflagel- lates, always with zooplankters present. However, she referred to other investi- Vor. XXXVII] LOUKASHKIN: ANCHOVY DIET AND FEEDING BEHAVIOR 447 FicurE 7. Brown pelicans and sea-gulls feeding on a large school of anchovies (Engraulis mordax) at sunset in San Quintin Bay, Baja California. September 21, 1965. gators whose findings were different from her own (Fuster de Plaza, 1964, and Angelescu and Fuster de Plaza, 1962). According to de Ciechomski, they found the stomach contents of the juveniles to be composed primarily of phytoplankton. In fish 50 mm. to 100 mm. they found a “predominance of diatoms,” but as soon as juveniles approach the adult stage their diet becomes ‘preferentially zooplanktonic.”’ According to the said investigators, in the stomachs of the fish 150-190 mm. long, copepods (especially calanids), pelagic amphipods, sergestid shrimps, and sometimes juveniles of anchovy and other fish compose the bulk of the contents. But contrary to the statement of the above mentioned authors, de Ciechomski in her own investigations disclosed a quite different situation. She found out that ‘‘the larvae and juveniles of the Argentine anchovy are zoo- planktophagous from their earliest stages. The contribution of the phytoplank- ton to their diet is quite small.” The diet of the adult fish is made up almost exclusively of zooplankters. The most important food items for larvae and juveniles 22-90 mm. long are “various groups of Copepoda, especially the Calanoida”; and least in importance are Cyclopoida.'* Second in order of 12 Among calanids the most frequently encountered forms were Paracalanus parvus and Centropages spp., and among cyclopoids—Oithona minuta and Corycaeus spp. Herpacticoid copepods were represented only by a single species—Eutherpina acutiformis. The calanids and herpacticoid copepods frequently and abundantly found in the stomach contents of the larval and juvenile Argentine anchovy were the most abundant forms in the plankton samples taken concurrently. 448 CALIFORNIA ACADEMY OF SCIENCES [PRroc. 4TH SER. Taste 9. Food type incidence in 840 filled stomachs of northern anchovies collected in 1965-1968 in northern Baja California, southern, and central California. Number of Incidence food type entries in a Percent Group Sub-group sub-group Number to total Zooplankton Crustaceans Copepods! 10 1198 29.32 Zooplankton Crustaceans Euphausiids! 5 390 9.54 Zooplankton Crustaceans Others! 24 487 11.92 39 2075 50.78 Zooplankton Other zooplankters Pelagic worms 9 333 8.15 Zooplankton Other zooplankters Thaliaceans 4 ZZ 6.66 Zooplankton Other zooplankters Appendicularians 2 152 3oZ Zooplankton Other zooplankters Molluscs 6 144 Ses! Zooplankton Other zooplankters Bryozoan “cyphonautes” 1 111 Bete Zooplankton Other zooplankters Fish eggs and larvae" 7 104 Doss) Zooplankton Other zooplankters Other miscellaneous 14 175 4.28 Zooplankton Other zooplankters Indeterminate soft parts and remains 3 170 4.16 46 1461 35.76 Total 85 3536 86.54 Phytoplankton Diatoms Diatoms 1 295 fe2e Phytoplankton Other phytoplankters - 3 154 Shi Total 4 449 10.99 Foreign matter - - 7 101 2.47 Grand total 96 4086 100.00 1 All stages and remains. “Included anchovy eggs and larvae. importance as food items are caladocerans (Podon polyphemoides and Evadne nordmannt).'* Larvae of decapods and bivalves were found in quantities of “slight significance in respect to total food supply.” The same was true of the eggs of various marine organisms. Most of the fish eggs in the stomach contents were eggs of anchovy and of Prionotus species.'4 Among the phytoplankters found in the stomach contents, species of the diatoms Coscinodiscus and Triceratium, and of the dinoflagellate Exuviaella were the plant organisms most frequently encountered. De Ciechomski observes that “although these organisms are found in many individuals and are occasionally 18 The great quantities of cladocerans in the intestinal contents, especially during some winter and summer months, coincided with the abundance of these microcrustaceans in the plankton. 4 Presence of fish eggs in the stomachs, especially during the spring and summer, coincided with the appearance of these eggs in the plankton. Vor. XXXVII] LOUKASHKIN: ANCHOVY DIET AND FEEDING BEHAVIOR 449 Ficure 8. Not the stars in the sky! This is a mass concentration of juvenile northern anchovy (Engraulis mordax) feeding on the microplankton consisting mainly of microcalanids, their eggs and nauplii, with insignificant admixture of bryozoan “cyphonautes” larvae, and occasional diatoms (Coscinodiscus species), as uncovered by examination of the fish stomach contents. With gill covers stretched wide they filtered water when raising to the surface in vertical wave pattern. The sun reflection from the gill covers and fish sides is seen in the picture. Reaching close to the surface, the fish shut their mouths, and submerge for about 7-10 feet, and then repeat the sequence. San Jose Point, Baja California, Mexico, close to the shore line; early morning hour, September 26, 1965. Specimens in the collection numbered 39-50. Size range of the fish in this school was within 67-91 mm. (standard length). very abundant, their significance in the total weight is low. During the summer a lesser contribution of the phytoplankton organisms is observed in the anchovy diet.” Among other things, de Ciechomski mentioned that in juveniles 41-90 mm. long she found amounts of detritus and very fine sand grains with great quantity of Triceratium, a diatom abundant in water layers close to the bottom. In the spring the skeletons of benthic foraminiferans were found in the food of the in- dividuals.!° In closing, de Ciechomski identified the Argentine anchovy as a 15 This, as in case with our northern anchovy, exposes behavioral tendency to descend to near-bottom horizon in bright sunshine. 450 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. zooplanktophagous animal in comparison with the phytoplanktophagous Peruvian anchoveta, Engraulis ringens, and concludes that “‘all of the information obtained suggests that the larvae and juveniles of the | Argentine] anchovy do not select much of their food, and that they feed upon the food which is present in greater abundance.'!® As a consequence, their diet may be quite dependent upon the patterns of plankton dispersal. Another very important factor is the accessibility of the food as determined by the size of the prey and dimensions of the mouth of the fish.” ACKNOWLEDGMENTS This work has been carried on with funds provided by the Marine Research Committee as a part of the California Cooperative Oceanic Fisheries Investiga- tions. The author expresses his appreciation to Dr. Robert C. Miller, Senior Scientist of the California Academy of Sciences, for his keen interest in the studies and for encouragement and valuable suggestions during the progress of the research; to Dr. G Dallas Hanna, Dr. Martin W. Johnson, and Messrs. Allyn G. Smith, Dustin D. Chivers, and William J. Light for their willing counsel and identification of the more difficult plankton organisms; to Dr. Reuben Lasker of the U.S. Fish and Wildlife Service, and Mr. Dion L. Hamilton of the California Department of Fish and Game, for their assistance in obtaining materials for the present study. CONCLUSIONS On the basis of the materials analyzed and visual observations incorporated in the present report, and referring to the literature cited, the following sum- marized conclusions can be reached: (1) The northern anchovy, Engraulis mordax Girard, is a planktophagous species. (2) Itis an omnivorous animal living either on phytoplanktonic or zooplanktonic organisms, or on both at the same time. (3) Zooplankters seem to be preferred in the anchovy diet. Among zooplankters, crustaceans in all stages, from egg to adult form, appear to be the most dominant food objects found in the stomach contents. (5) Of crustaceans, the copepods and euphausiids are most frequently and abundantly found in the stomachs, and they appear to be the most im- portant food items in the diet of the northern anchovy, as well as in the diet of other anchovies whose food habits were already reported by other investigators. (6) Though phytoplanktonic organisms were also found in the stomachs (some- times they constituted a dominating item, or they even contributed up to 16 All 30 food objects listed by de Ciechomski for the larval and juvenile Argentine anchovy are also present on the menu of the northern anchovy. Vor. XXXVII] LOUKASHKIN: ANCHOVY DIET AND FEEDING BEHAVIOR 451 100 percent of the contents), their role in the anchovy diet seems on the whole negligible. (7) In mode of feeding, the northern anchovy is primarily a filter feeder, but may also be a particulate or selective feeder, depending on the size of the available food. (8) In relation to feeding habits, the northern anchovy is a diurnal animal, feeding mostly during the day. (9) From the larval stages through adult life, the northern anchovy shows an apparent preference for the second link in the food chain. This is in con- trast to the food of the Peruvian anchoveta, Engraulis ringens, which is reported to be chiefly phytoplankton. LITERATURE CITED ApBott, Donatp P., and RICHARD ALBEE 1967. Summary of thermal conditions and phytoplankton volumes measured in Monte- rey Bay, California, 1961-1966. California Cooperative Oceanic Fisheries In- vestigations Reports, vol. XI, pp. 155-166, |Sacramento, California]. ANGELESCU, V., and M. L. FUSTER DE PLAzA 1962. El papal de la anchoita en la bioeconomia general del mar Argentino (Sector bonaerense, resultados preliminares). 1st Reunion, Comision Consultiva Re- gional de Pesca Atlantico Sud-Occidental, Rio de Janeiro, [Brazil], Tema 6/6, pp. 1-13 [After Janina Dz. de Ciechomski, 1968—a, as listed below]. ANONYMOUS 1952. Food of the sardine. California Cooperative Sardine Research program, Progress Report, 1 January 1951 to 30 June 1952, pp. 23-24, and 1 figure on p. 25, [Sacramento, California |. ANONYMOUS 1967. Behavior studies on anchovies. California Cooperative Oceanic Fisheries In- vestigations Reports, vol. XI, pp. 19-20, [Sacramento, California]. BAXTER, JOHN L. 1967. Summary of biological information on the northern anchovy—Engraulis mordax Girard. California Cooperative Oceanic Fisheries Investigations Reports, vol. XI, pp. 110-116, 3 figures and 6 tables in text. 1968. California Department of Fish and Game Pelagic Fish Investigations (agency report). California Cooperative Oceanic Fisheries Investigations Reports, vol. XII, pp. 10-13. Bay irr, WILLIAM H. 1963. The food and feeding habits of the anchoveta, Cetengraulis mysticetus, in the Gulf of Panama. Inter-American Tropical Tuna Commission Bulletin, vol. 7, no. 6, pp. 399-419; 14 tables and 1 figure on pages 420-432, La Jolla. BERNER, LEO, JR. 1959. The food of larvae of the northern anchovy Engraulis mordax. Inter-American Tropical Tuna Commission Bulletin, vol. 4, no. 1, pp. 3-15, 5 text figures and 4 tables. Bropsku, K. A. 1936. Kratkii predvaritel’nyi otchet o planktonnyh issledovaniiah po _ pitaniiu dal’nevostochnyh sardin vy 1935 g. (Preliminary brief account of plankton in- CALIFORNIA ACADEMY OF SCIENCES [| Proc. 4TH SER. as mn bo vestigations on feeding behavior of the Far Eastern sardines in 1935). Vestnik Dal’nevostochnogo Filiala Akademii Nauk SSSR, no. 18, pp. 155-160, Vladivostok. Bropsku, K. A., and A. J. IANOVSKAIA 1935. O pitanii dal’nevostochnoi sardiny (On feeding of the Far Eastern sardine). Vestnik Dal’nevostochnogo Filiala Akademii Nauk SSSR, no. 13, pp. 103-114, Vladivostok. Conway, J. B., D. A. Farris, and R. F. Forp 1968. Studies on the sablefish and the Pacific hake. California Cooperative Oceanic Fisheries Investigations Reports, vol. XII, pp. 14-15, [Sacramento, California]. Davirs, D. H. 1957. The South African pilchard (Sardinops ocellata)—Preliminary report on feeding off the West Coast, 1953-56. Division of Fisheries Investigational Report No. 30—A reprint from the “Commerce and Industry,’ November, 1957, pp. 1-40 including 17 tables, Pretoria, [South Africa ]. DE CIECHOMSKI, JANINA Dz. 1967a. Present state of the investigations on the Argentine anchovy Engraulis anchoita (Hubbs, Marine). California Cooperative Oceanic Fisheries Investigations Re- ports, vol. XI, pp. 58-71, 5 text figures and 2 tables. |Sacramento, California]. 1967b. Investigations of food and feeding habits of larvae and juveniles of the Argentine anchovy Engraulis anchoita. California Cooperative Oceanic Fisheries Investiga- tions Reports, vol. XI, pp. 72-81, 8 text figures and 3 tables. DERIUGIN, PROF. 1933. Tihookeanskie ekspeditzii GGI vy 1922 g. (Pacific expedition of the State Hydro- graphic Institute in 1922). Issledovaniia morei SSSR, Issue 17, Moscow [After A. J. Iankovskaia, 1937, as listed below]. IDG, Wis AN 1961. Food of the anchovy larvae in the Black Sea (In Russian). Trudy Sevastopol’skoi Biologischeskoi Stantsii, vol. 14, pp. 244-258. [After Schumann, 1965 listed below]. FUSTER DE Praza, M. L. 1964. Algunos datos sobre la biologia de la anchoita del sector bonaerense (resultados preliminares). The Food and Agriculture Organization 2a Reunion. Comisidn Asesora Regional de Pesca para el Atlantico Sud-Occidental, 2, Documento téchnico 12, pp. 1-11, [After De Ciechomski, 1967a, as listed above]. Gar, G. I. 1934. Fitoplankton—pistcha ivasi (Phytoplankton—food of the iwashi). Rybnoe Hoziaistvo Dal’nego Vostoka, no. 1-2, pp. 52-54, Moscow—Habarovsk. 1936. Raspredelenie fitoplanktona y poverhnostnyh sloiah vod severozapadnoi chasti Iaponskogo moria (Distribution of the phytoplankton in the surface layers of waters of the north-western part of the Sea of Japan). Vestnik Dal’nevostoch- nogo Filiala Akademii Nauk SSSR, no. 18, pp. 107-108, Vladivostok. Hanp, Capvet H., and Leo Berner, JR. 1959. Food of the Pacific sardine (Sardinops caerulea). Fishery Bulletin No. 164 from Fishery Bulletin of the Fish and Wildlife Service, vol. 60, pp. 175-180, Wash- ington, D.C. Hart, Joun Lawson, and Grorce Hersert WAILES 1932. The food of the pilchard, Sardinops caerulea (Girard), off the coast of British Columbia. Contribution to Canadian Biology and Fisheries being studied from Vor. XXXVII] LOUKASHKIN: ANCHOVY DIET AND FEEDING BEHAVIOR 453 the Biological Stations of Canada, New Series, vol. VII, nos. 15 to 23, Series A, General, nos. 12 to 20. Biological Board of Canada, Toronto, [Hart’s con- tribution numbered 19 (Series A, General, no. 16, pp. 1-10) J. HayaAst, SIGETO 1967. A note on the biology and fishery of the Japanese anchovy Engraulis japonica (Houttuyn). California Cooperative Oceanic Fisheries Investigations Reports, vol. XI, pp. 44-57, 3 text figures and 3 tables. [Sacramento, California]. HICKLING, C. F. 1945. The seasonal cycle in the Cornish pilchard, Sardina pilchardus Walbaum. Journal of the Marine Biological Association of the United Kingdom, vol. XXVI, no. 2 (issued August, 1945), pp. 115-138, text figures 1-8, Cambridge, at the Uni- versity Press. JTaANKovskala, A. J. 1937. Zooplankton i pitanie ivasi v severo-zapadnoi chasti Iaponskogo moria (Zoo- plankton and feeding of the iwashi in the Northwestern part of the Sea of Japan). Vestnik Dal’nevostochnogo Filiala Akademii Nauk SSSR, no. 27, pp. 63-83, illustrations, Vladivostok. KISHINOUYE, KAMAKICHI 1907. Notes on the natural history of the sardine (Clupea melanosticta Schlegel). Journal of the Imperial Fisheries Bureau, vol. 14, no. 3, pp. 71-105, Tokyo. [ After Brodskii and Iankovskaia, 1935, as listed above ]. KocanovsKala, S. M. 1934. Promysel ivasi v raione o. Putiatina v 1933 godu (Iwashi fishery in the region of Putiatin island in 1933). Rybnoe Hoziaistvo Dal’nego Vostoka no. 12, pp. 34-44, Moscow-Habarovsk. KocAnovskil, A. G. 1935. Materialy k poznaiiu Dal’nevostochnoi sardiny-ivasi (Materials toward the knowledge of the Far Eastern sardine-iwashi). Rybnoe Hoziaistvo Dal’nego Vostoka, no. 13, pp. 35-38, Habarovsk. Kuso, I. 1961. Suisan Sigen Kauron (Summary of fishery biology, particular to living resources in Japan), 396 pp. Tokyo, Koseisya-Koseikaku. [After Hayasi, 1967, as listed above]. Lewis, Ratpu C. 1929. The food habits of the California sardine in relation to the seasonal distribution of microplankton. Bulletin of the Scripps Institution of Oceanography, La Jolla, California, Technical Series, vol. 2, no. 3, pp. 155-180, 2 text figures, University of California Press, Berkeley, California. LOUKASHKIN, ANATOLE S., and NoRMAN GRANT 1965. Behavior and natural responses of the Northern anchovy, Engraulis mordax Girard, under the influence of light of different wave lengths and intensities and total darkness. Proceedings of the California Academy of Sciences, Fourth Series, vol. XX XI, no. 24, pp. 631-692, 11 figs., San Francisco. Mapras FISHERIES DEPARTMENT 1933. Administrative Report for the year 1931-32. Madras [After Brodskii, 1936, as listed above]. 1936. Administrative Report for the year 1934-35. Madras [After Iankovskaia, 1937, as listed above]. 454 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. MIKULIcH, L. V. 1960. O raspredelenii planktona v severnoi chasti Ohotskogo moria letom 1955 goda (On distribution of the plankton in northern part of the Sea of Okhotsk in the summer of 1955). Izvestiia Tihookeanskogo Nauchno-issledovatelskogo Instituta Rybnogo Hoziaistva 1 Okeanografii, vol. XLVI, pp. 41-64, illus- trations, Vladivostok. Mirter, R. C. 1967. California Academy of Sciences (Agency activities). California Cooperative Oceanic Fisheries Investigations Reports, vol. XI, p. 9. [Sacramento, Cal- ifornia]. MiIcter, R. C. 1968. California Academy of Sciences (Agency reports). California Cooperative Oceanic Fisheries Investigations Reports, vol. XII, p. 10. [Sacramento, California]. Murpuy, Gartu I. 1966. Population biology of the Pacific sardine (Sardinops caerulea). Proceedings of the California Academy of Sciences, Fourth Series, vol. XXXIV, no. 1, pp. 1-84; 17 figs.; 20 tables. San Francisco. Nakat, Z., K. Honjo, T. Kipacut, H. Suzuxt, T. Yokota, T. Tsujita, E. Ozasa, Y. SHOyJOMA, and S. NISHIMURA 1962. Iwasirui kokisigyo no syokuzi to kanyuryo tono kankei (Relationships between food organisms and size of recruitment of iwasi). Suisan Sigen ni kasuru kyodo kenkyu suisin kaigi hokokunsyo, Syowa 36 nendo, pp. 102-121. Norin Suisan Gizyuto Kaigi, Tokyo [After Hayasi, 1967, as listed above]. Naxkal, Z., S. Usami, S. Hattori, K. Honjo, and S. HAyas1 1955. Progress report of the Cooperative iwashi resources investigations, 1949—1951, 116 pp., Tokai Regional Fisheries Research Laboratory, Tokyo [After Hayasi, 1967, as listed above]. Parr, A. E. 1930. Is the presence of phytoplankton in the stomach content of the California sardine caused by special pursuit or merely due to incidental ingestion? Ecology, vol. XI, no. 2 (Notes and comments), pp. 465-468, Brooklyn, N.Y. Rapovicnu, JOHN 1952. Food of the Pacific sardine, Sardinops caerulea, from Central Baja California and Southern California. California Fish and Game, vol. 38, no. 4, pp. 575-585, Sacramento, [California ]. Rapovicu, JoHN, and Eart D. Gripes 1954. The use of a blanket net in sampling fish populations. California Fish and Game, vol. 40, no. 4, pp. 353-365, 8 figs., Sacramento. SCHUMANN, GEORGE O. 1965. Some aspects of behavior of clupeoid larvae. California Cooperative Oceanic Fisheries Investigations Reports, vol. X, pp. 71-77. [Sacramento]. Vor. XXXVII] LOUKASHKIN: ANCHOVY DIET AND FEEDING BEHAVIOR 455 APPENDIX A. A list of partially identified copepods from the stomachs of the Northern anchovy—Engraulis mordax (Girard). Identified by Professor Martin W. Johnson 1. Acartia spp. 11. Metridia sp. 2. Calanus helgolandica 12. Microcalanus spp. 3. Calanus spp. 13. Microsetella sp. 4. Candacia sp. 14. Oithona sp. 5. Centropages sp. 15. Onacea sp. 6. Corycaeus sp. 16. Paracalanus spp. 7. Cyclopoid copepods 17. Pseudocalanus spp. 8. Eutherpina sp. 18. Rhyncalanus sp. 9. Herpacticoid copepods 19. Temora sp. 10. Labidocera trispinosa 20. Calanoid copepods APPENDIX B. A list of diatoms and dinoflagellates found in the stomach contents of northern anchovy (Engraulis mordax) collected in Baja California and southern and central California waters in 1965-1968. Identifications were made chiefly by Dr. G Dallas Hanna, California Academy of Sciences. DIATOMS 1. Bacteriastrum sp. 14. Distephanus speculum 2. Biddulphia longicruris 15. Eucampia sp. 3. Chaetoceros decipiens 16. Lithodesmium sp. 4. Chaetoceros sp. 17. Navicula distans 5. Coscinodiscus asteromphalus 18. Nitzschia sp. 6. Coscinodiscus excentricus 19. Pleurosigma sp. 7. Coscinodiscus oculis-iridis 20. Rhizosolenia hebatica 8. Coscinodiscus radiatus 21. Rhizosolenia sp. 9. Coscinodiscus stellaris 22. Skeletonema costatum 10. Coscinodiscus wailesii 23. Skeletonema sp. 11. Coscinodiscus sp. 24. Stephanopyxis turris 12. Dicladia capreolus 25. Thalassiosira sp. 13. Diploneis smithii 26. Thalassiothrix sp. DINOFLAGELLATES 1. Ceratium furca 4. Ceratium tripos ae Ceratium longipes 3. Ceratium macroceros 6. Ceratium sp. Dinophysis sp. 7. Peridinium spp. 456 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser. APPENDIX C. SAMPLE LXIV. On September 21, 1965, at 10:15 p.m. in San Quintin Bay, Baja California, Mexico, 2.8 miles offshore, 200 Pacific sardines and 10 anchovies were captured using electric light as attractant and a blanket net as collecting gear. Depth of water was 17 fathoms. Stomachs of 11 sardines 203-235 mm. standard length and 10 stomachs of anchovy 105-117 mm. standard length were preserved. Rec- ords of examination of the contents of stomachs are given below. Northern anchovy Engraulis mordax Nos. 395-404 Pacific sardine Sardinops caerulea Nos. 405-414 and 414a All stomachs were filled to capacity In all stomachs large copepods constituted dominating food item (up to 98%). Among these copepods other zooplankters were scat- tered in insignificant numbers, such as small copepods, decapod oikopleura, etc. larvae, amphipods, No diatoms were found. Three stomachs were filled to full capacity (“gorged”) ; two—%4 capacity; two—half- filled; and four—poor. In all stomachs, even with poor filling, large copepods were dominating food item (up to 98-99%). Among these copepods there were a few other zooplankters present either by single individuals or in small numbers. To this category belonged small copepods, brachyuran zoea and megalopa, isopods, amphipods, euphausiid larvae, copepod eggs, copepod fleshy parts, chunks of polychaete worms, salps, hemicordate larvae, oikopleura. Small copepods and copepod eggs were more numerous than the rest of the other zoo- plankters. In one stomach a fragment of algae (kelp-like type) was found. No diatoms were present. Foreign matter, such as many fine mica flakes and a few fine sand grains, was found in stomach No. 413. Similarity in food items is evident. Vou. XXXVII] APPENDIX D. LOUKASHKIN: ANCHOVY DIET AND FEEDING BEHAVIOR 487 Frequency of occurrence of various food items in the stomach contents of 12 jack mackerel, Trachurus symmetricus (Ayres) 87-102 mm. standard length from the catch of 3,000 fish, using electric light and blanket net, on September 21, 1965, at 10:15 p.m. in San Quintin Bay, Baja California. Seven stomachs were filled to full capacity, 2 to 4 capacity, and 3 stomachs to “% capacity. Position Description Dominating Second Place Insignificant Total MOO MEMOdsSmmlALees) 7 4 1 12 Paauausnds, adit 3 1 1 5 Vity Sic] pues Pt 1 = 4 5 Indeterminate fleshy material — 1 3 1 5 @rustacean’ fleshy parts 2 ~ = 5 5 OikGpletnamepe ee 8 = 1 3 4 brachiyunam ZOCa = = 10 10 emicordata larvae = = 10 10 Jellyfish larvae and parts —.. - = 6 6 (ChaACiOR iN 2 ee ~ = 6 6 “URED Ut) Gath) Sei - - 6 6 Fine sand grains (foreign matter) — - = 6 6 AVI ERO Ga) An Se - = 4 4 Algae (kelp-like fragments) __ ee - _ 4 4 Foraminiferans (bottom living) — _ = 4 4 Minute pebbles (foreign matter) ~~ - = 4 4 IS@oOGIS Le - — 3 3 SANDS eens - ~ 3 3 Brachyvopod Warvae - - 3 3 Fine mica flakes (foreign matter) — - = 3 3 [EES CP eee - - 3 3 ILAGae (CiP)) 9 al ee - ~ 2 2 IMEC Sdepeeeen re e - ~ 2 2 Small copepods) in 52 ne re ee = = 2 2 Nemertean worm larvae (pelagic) = — - 2 2 cero WOCSm(CEZMAGING Sp) = - - 2 2 Cephalocordatay larvae 222 - - 2 2 SACHEM (WOEMS)) 22 8 - - 2 2 Porcellanid crab larvae eee ee sees! ~ - 1 1 BOLO Smee CLS: LAL VAG oe - - 1 1 BOlGhaectem WwOLMS js 6 - - 1 1 Rolyehacte- worn’ eggs 8 - = 1 1 KGastropodilakvae. 2.2 oe ~ - 1 1 A\naiol MOG LAN ~ ~ 1 1 Shiai mesma |e Lae ae ~ - 1 1 Mamntiseshmmap larvae Se 2 = — 1 1 (Onin Ce ANS wee eeeeen ee ~ - 1 1 BUpPhAUsidacalyptOpis) see - = 1 1 1 Includes adult copepods, their fleshy parts and remains (carapaces). CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. = U1 o APPENDIX D. (Continued) Position Second Description Dominating Place Insignificant Total Roraminiterans i (oe!a crc) en a - ~ 1 1 Bish: eges 32 Sa te ee es ee - - 1 1 Pisin Gees wali lemye, woe — — - 1 1 Lait O11 Sa em ie Eee ee ee es er = = 1 1 ishwscalesm(fonergre aia iter) enn - - 1 1 Rough sand grains (foreign matter) —— - ~ 1 i TOTAL 12 6 121 142 APPENDIX E. SAMPLE LVIII. October 18, 1966, 8:13 p.m.; 14 mi. E. xX N. Point Concep- tion, southern California, 4 m. offshore; 17 anchovies 107-131 mm. standard length and 2 hakes 129-132 mm. standard length caught by midwater trawl net. Stomach contents of 7 anchovies and 2 hakes examined. Dominating food items eo (Engraulis jordan) Hake (Merluccius productus) No. 783 Euphaustids No. 790 Euphausiids No. 784 Copepods No. 791 Euphausiids No. 785 Euphausiids No. 786 Salps No. 787. Polychaete worms No. 788 Euphausiids No. 789 Empty Euphausiids 50% Euphausiids 100% Other 50% TOTAL 100% TOTAL 100% PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XXXVII, No. 14, pp. 459-472; 7 figs. October 13, 1970 POLYDORA ALLOPORIS, NEW SPECIES, A COMMENSAL SPIONID (ANNELIDA, POLYCHAETA) FROM A HYDROCORAL OFF CENTRAL CALIFORNIA By William J. Light Department of Biological Sciences Simon Fraser University Burnaby 2, British Columbia Apstract: A new species of polychaetous annelid, Polydora alloporis (Spionidae), is described from central California. It is always abundantly found in burrows bored into the coenosteum of the hydrocoral Allopora californica Verrill, 1866, and is one of the largest of the polydorids, reaching a length of 75 mm. or more. It is related to Polydora hornelli Willey, 1905, from India, and to P. cavitensis Pillai, 1965, and P. pacifica Takahashi, 1937, both from the western Pacific. INTRODUCTION Biologists have long known that the subtidal hydrocoral Allopora californica Verrill, 1866, from central California is always found to harbor a burrowing spionid, whose paired burrow-openings are scattered throughout the branches of both the orange-red and the purple color phases of the hydocoral. Its identity, however, has until now remained obscure. Preserved material of Allopora calif- . ornica collected by Mr. Dennis Sullivan, then of the California Academy of Sci- ences, in August, 1962, off San Jose Creek Beach (Monastery Beach), Carmel, Monterey County, California, in 35 to 40 feet of water, was deposited in the Academy’s collections in the Department of Invertebrate Zoology (CASIZ). This material was examined and the worms it contained were diagnosed simply as Polydora species, inasmuch as only a few isolated fragments could be with- [459 | Marine Biologi ne Biological Laboratory OCT 301970 WOODS HOLE, MASS, 460 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. drawn from the preserved hydrocoral. This, however, was sufficient for a generic determination. On 19 September 1968, Mr. Don Wobber of San Francisco State College secured live material with scuba equipment from the same general vicinity near Carmel, California, from a depth of 80 feet. We were able to obtain entire living worms from this sample of Adlopora by carefully crushing it with a pair of vice-grip pliers. It was readily apparent that these polychaetes represented an undescribed species closely allied to Polydora cavitensis Pillai, 1965, from the Philippine Islands and to P. hornelli Willey, 1905, from India. Further investigation also revealed affinities with Polydora pacifica Takahashi, 1937, from Palau Island, Caroline Islands, in the Philippine Sea. The last mentioned is a species commonly found burrowing in a species of Montipora, a branched coral growing in the reefs of that region, as well as in the shells of the pearl oyster, Pinctada margaritifera (Linné, 1758). On 26 October 1968, Mr. Wobber and I returned to the Monterey area and obtained additional living material off 17-Mile Drive near Carmel, California from 125 feet of water in an area of submerged seastacks known locally to scuba divers as ‘“‘the Pinnacles.” The type series contains material from all three of the above collections, and includes the holotype and 12 paratypes. Family Sp1ionmae Grube, 1850 Genus Polydora Bosc, 1802 Polydora alloporis Light, new species. DiAcnosis. Length to 75 mm. or more. Modified major spines of fifth setiger simple and falcate, with hollowed out distal ends; companion setae situated immediately ventrad of the major spine series, normally lanceolate, but when worn resembling narrow clubs with many small spiny projections. Fifth setiger heavily folded and creased, dorsally. A prominent band of cilia running transversely across the dorsum of each segment from the seventh setiger to the pygidium; similar cilia on the medial surfaces of the branchiae. Prostomium anteriorly entire, formed of two membranes, a superior one forming a dorsal sheath over an inferior process which projects anteriorly beyond the forward margin of the superior membrane; prostomial caruncle narrow, extending poste- riorly to the middle of the fourth or the anterior border of the fifth setiger. Median and posterior segments ventrally marked with distinctive greenish-or reddish-brown butterfly or fleur-de-lis pattern, the “wings” of which may form open rings, two or more parallel lines, or even more or less solid blotches. Pygidium a widely flaring dusky silver, fleshy disc with a deep dorsal cleft and many small papillae on the posterior surface. Found exclusively and in large numbers in deep burrows in the coenosteum of Allopora californica at subtidal depths. Vor. XXXVIT] LIGHT: A COMMENSAL SPIONID FROM A HYDROCORAL 461 TYPE MATERIAL. The holotype and six paratypes have been placed in the California Academy of Sciences, Department of Invertebrate Zoology (CASIZ) Type Series as numbers 381 (the holotype) through 387, inclusive. Additional material, including the modified spines of the fifth setiger, neuropodial crotchets, palpi, and longitudinal and cross sections of various segments and parapodia have been mounted separately as CASIZ microscope slide numbers 353 through 364, inclusive. Paratypes have also been deposited in the British Museum (Natural History), American Museum of Natural History, the Zoological Institute of the Academy of Sciences (Leningrad), United States National Museum, and. the Allan Hancock Foundation at the University of Southern California. DistRIBUTION. Polydora alloporis has been recovered only from samples of Allopora californica taken at various subtidal depths off the Carmel area, Monte- rey County, California. Hasitat. This polychaete lives as a commensal in deep burrows in the coenosteum of Allopora californica where it may form large colonies. The paired openings of the burrows are distinctive and the worms may be seen in living material with the palpi projecting from one of the apertures, and the pygidium just visible below the lip of the other (fig. 1). There is a distinctive fauna as- sociated with this hydrocoral, including the gastropod Pedicularia californica Newcomb, 1864, which matches the color of its host, and a variety of small reddish caprellids and other amphipods. A distinctive balanomorph barnacle, Balanus (Armatobalanus ) nefrens Zullo, 1963, was also recovered from the same samples of Allopora as were the worms. This barnacle, which was identified by Dr. Victor A. Zullo, California Academy of Sciences, is only known to exist in the coenosteum of Allopora californica and Errinopora pourtalesi Dall, 1884, from Monterey Bay and the Channel Islands, California (Zullo, 1963, p. 593). It has its closest affinities with animals from the Australasian region (Zullo, 1966, p. 142). In addition, a number of nonhost specific organisms have been taken from these samples, including the following polychaetes: Halosydna johnsoni (Dar- boux, 1899) and H. brevisetosa Kinberg, 1855 (Polynoidae), and Anaitides species (Phyllodocidae). EXTENDED DESCRIPTION. Polydora alloporis is extremely long for the genus and specimens commonly attain lengths of 65 to 70 mm. The longest specimen, the holotype with 188 segments, measured 75 mm. in length by 2 mm. wide. The greatest number of segments (236), was noted in an individual measuring 67 mm. in overall length. The most striking features of the living animals are the prominent dorsal and ventral blood vessels through which pulsates bright-red blood. The dorsal vessel bifurcates anteriorly and is continued in the palpi, where it forms convoluted loops proximally. The ground color is fleshy but there is often a faint greenish tinge to the body in living specimens. Distinctive 462 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 1. Polydora alloporis, new species. Living animal protruding from paired holes in coenosteum of Allopora californica. Photograph by Allyn G. Smith. patterns of pigmentation are exhibited on the venter and the palpi. These are discussed below. The pygidium is a striking dusky silver in complete contrast to the rest of the body. The animal is somewhat dorsoventrally compressed in the anterior third of its length, but the somites become gradually attenuated and laterally swollen posteriorly, where they acquire a barrel-like configuration. The sixth to about the twelfth setigers appear to be affixed to the modified fifth segment at a lower level than are the preceding somites in many individuals, so that the animals appear to be ventrally distended over this region (fig. 5). Prostomium. The prostomium projects well beyond the anterior border of the lateral cephalic lobes of the peristomium. It is bluntly rounded and entire at its anterior margin where it forms a small roll of tissue passing from one side of the prostomium to the other, imparting a ‘“‘Shognosed”’ appearance to the snout (fig. 5). Immediately behind this lies the anterior border of a superior membrane which forms a sheath over the prostomium that extends laterally onto the lateral cephalic lobes on either side of the prostomium for about one-fourth of the width VoL. XXXVII] LIGHT: A COMMENSAL SPIONID FROM A HYDROCORAL 463 of these lobes (figs. 2, 5). Posteriorly, this sheath passes back to the point of insertion of the palpi. This membrane is medially continuous with the prostomial caruncle and appears to represent a forward extension of that structure. The prostomial portion underlying this superior sheath is widest distally and tapers evenly to the level of the first setiger, or to the midpoint of insertion of the palpi. At this point are situated one pair of small, but prominent eyespots, which are overlain by the superior prostomial sheath. At the level of the eyespots the caruncle forms two successive dorsally project- ing humps or bosses and then passes posteriorly as a rather narrow band of raised tissue to the middle of the fourth or the anterior border of the fifth setiger (to the anterior margin of the third in one specimen). The caruncle is dorsally raised into a strip of corrugated glandular tissue extending from the posterior border of the palpi to the posterior tip of the caruncular extension (figs. 2, 5). From a dorsal perspective the two anterior dorsal bosses appear to project slightly for- ward, but there is no trace whatsoever of an occipital cirrus. Peristomium. The peristomium is laterally projected on either side of the prostomium into two widely flaring cephalic lobes which are not fused with the first setiger, contrary to the condition occurring in some polydorids. Anteriorly, it sweeps downward to enclose a converging U-shaped buccal opening, which is lined on each side by two fleshy strips of tissue (figs. 2, 5). These two fleshy borders are confluent with a folded and lobulated eversible proboscidial mass which lies inside the oral cavity. This proboscis is well extended in one of the paratypes. Posteroventrally, the peristomium extends to a point below the level of the second parapodia and generally, though not always, completely supresses the ventral development of the first setiger. The palpi are thick, long, and highly lobulated along the longitudinal ciliated grooves. They are densely studded over their entire surface with minute, bell- shaped, cellular protuberances, each of which bears a single whip-like flagellum. The median longitudinal groove is heavily ciliated and is bordered on each side by thick pads of mucus-secreting glandular tissue which are raised into heavy mounds and ridges. This glandular region is densely colored with a sooty-black pigment which forms a more or less continuous border on either side of the groove in most specimens, but which in some animals forms discrete blotches or rectangular bars along the groove (figs. 1, 2). The proximal sixth of the palpi lacks both pigment and glands and the grooves at this level are nonciliated. Modified fifth setiger. The modified fifth setiger is twice the length of the segments immediately adjacent to it, and two and one-half times as wide as it is long. The dorsum of this segment is heavily ridged and folded. From the region of attachment of the modified spines and setae, a large fold passes dorsally and somewhat posteriorly over the dorsum, extends along the posterior margin of 464 CALIFORNIA ACADEMY OF SCIENCES 0.1 [Proc. 4TH SER. Vor. XXXVII] LIGHT: A COMMENSAL SPIONID FROM A HYDROCORAL 465 the somite, and is continuous with its companion from the opposite side (figs. 2, 5). A slightly smaller fold extends transversely across the segment along its anterior border, and the two folds enclose a central spindle-shaped depression (fig. 2). The modified spine series forms a dorsally concave anteroposteriorly sweep- ing arc and the spines themselves project far out from the body wall. There are usually six or seven of these spines (rarely eight) which are visible externally, but six or seven additional embryonic spines are generally found embedded in the tissues. i These spines are quite long and laterally expanded along the mid-portion of their lengths; they are distally falcate and quite pointed at their tips when un- worn. Each spine bears a deep, hollowed-out, subdistal concavity, which is bordered laterally and ventrally by a continuous slightly raised flange (fig. 4). This condition is very similar to that found in both Polydora cavitensis and P. hornelli. The embedded proximal tips of these spines are heavily striated with transverse ridges which become obscure distally. The core is considerably ex- panded in the middle part of the spine and fills the greater part of the cavity. It extends beyond the distal concavity almost to the terminal point. Each spine alternates with a clear transparent companion seta. These setae project laterally from the body wall in an even row lying immediately adjacent, but inferior to the major spine series. These setae are generally quite worn and closely resemble those figured by Takahashi (1937, p. 160) for Polydora pacifica in forming narrow, club-shaped, distal tips (the “lancet-like forms” of Takahashi) which are covered by many minute spiny projections (fig. 4). These setae taper proximally into a long, thin, whip-like, medial stalk. It is apparent, however, that these club-shaped companion setae are but worn versions of the lanceolate setae which remain unmodified in some animals. One or two such unworn setae can also be found farther posteriorly among the younger spines of most individuals. A vertical tuft of from seven to nine lanceolate setae emerges from near the base of the anteriormost modified major spine, and a much smaller fascicle of four or five similar setae projects from an isolated position ventral to the major spine series. Unmodified segments and parapodia. Both the dorsal and ventral lamellae are well developed on the first four setigers, but the entire parapodium of the first setiger has moved somewhat dorsally, so that the neuropodium lies at an < Ficure 2. Polydora alloporis, new species. Dorsal view of animal with left palp removed. Ficure 3. Polydora alloporis, new species. Neuropodial hooded crotchet. Ficure 4. Polydora alloporis, new species. Distal tip of major spine of fifth setiger, showing worn (left) and unworn (right) companion setae. 466 CALIFORNIA ACADEMY OF SCIENCES [Proc, 4TH SER. intermediate position between the notopodia and neuropodia of the succeeding somites. This differs from the situation in most members of this genus, in which the first neuropodium lies approximately at the same level as the following notopodia. There are no notosetae on the first setiger and only a small fascicle emerges from the neuropodium. This somite is generally ventrally supressed by the posterior prolongation of the peristomium. The postsetal lamellae of both rami are well developed from setigers two to four, but the notopodial postsetal lobes are distinctly larger than those in the neuropodia. The notosetae from setigers two to about fourteen emerge in two, closely appressed, vertical rows as a prominent light-colored, U-shaped fascicle. The superior portion of the anterior row consists of long capillaries which merge ventrally with unilimbate setae of an intermediate length. The latter are abruptly bent in a posterior direction shortly after emerging from the body wall and the wings are distinctly striated. At the base of the U-shaped fascicle appear very small spearshaped setae, which again blend into unilimbate setae of the inter- mediate type in the posterior vertical row of the fascicle. These setae resemble those found in the more generalized genus, Pseudopolydora, Czerniavsky, 1881, although the arrangement is somewhat different in the latter. From about the fourteenth somite, the smaller bladed setae drop out and the long capillaries and unilimbate setae become reduced in number. The two rows of the fascicle are reduced to one, and in the posterior segments the capillaries alone remain. The neuropodial setae of the first six setigers (including the small ventral tuft on the fifth) consist of bladed lanceolate forms. Simple hooded neuropodial crotchets appear on setiger seven where they are usually seven or eight in number, but occasionally as few as five or as many as ten. Within a very few segments they increase to 10 or 11 per row, this number persisting over from 6 to 10 somites. Thereafter, they gradually decrease by one crotchet per row (this may fluctuate by one or two hooks in a given series), each number being maintained over a long region, up to the segments immediately preceeding the pygidium, which usually contain from two to five each. These crotchets resemble those of Polydora cavitensis in having a bidentate tip in which the beak forms an angle of slightly less than 90° back against the main shaft (fig. 3). As in P. cavitensis, the point where the beak emerges from the shaft is smoothly rounded, and the apical tooth is more closely appressed against the beak than is usual among members of this genus considered in the strict sense. Below the level of the transparent hood, the shaft forms a curved neck and waist, followed by a slight swelling which then tapers evenly to the pointed proximal tip. The neck is much shorter than in the hooks of P. cavitensis, so that the waist lies at the distal fourth of the shaft, rather than at the distal third, as in Pillai’s species. The crotchets are extremely small for so large an Vor. XXXVII] LIGHT: A COMMENSAL SPIONID FROM A HYDROCORAL 467 animal, measuring less than 0.5 mm. in length, and the neurosetal lamellae are poorly developed. The branchiae are well developed from the outset, beginning on the seventh setiger and continuing to the pygidium. The inner or medial surfaces are heavily ciliated and this ciliary band is continued over the dorsum of each segment on a raised transverse ridge, where it merges with the cilia of the opposite gill. A well developed postsetal lobe arises at the posterior edge of each gill. Dorsolateral nephridial openings are evident just behind the septa dividing the somites in a position anteroventral to the branchiae. These openings are surrounded by a highly glandular epithelial ridge which runs dorsally parallel to the intersegmental septum, and which terminates in the vicinity of the dorsal transverse ciliary band. The glandular regions surrounding the nephridial openings are often pigmented and appear as reddish spots along the sides of the animal. They begin on the eighth segment and continue over as many as 40 somites in some individuals. The dorsum in preserved material exhibits a series of longitudinal, dark, reddish-brown, rectangular blotches over many of the segments, but this appears to be due to coagulated material in the massive dorsal blood vessel. A con- spicuous and typical pattern commonly appears on the ventral surfaces of the median and posterior segments, however. This consists of either a sooty-green or reddish-brown butterfly or fleur-de-lis configuration made up of closely paral- lel lines of pigment (fig. 6). The lateral “wings” of this pattern may form more or less open rings, consist of two or more longitudinal lines, or even appear as a solid blotch. This pigmentation is absent on some animals. Pygidium. The pygidium is a widely flaring, thick, fleshy disc, with a deep dorsal notch (figs. 6, 7). The fleshy pad is flattened on the posterior surface and is densely studded with many minute papillae. There is no posterior cup- like invagination so characteristic of the polydorids, and the anus opens somewhat dorsally. Polydora cavitensis also has a flattened anal disc which, however, is triangular in shape and which bears projecting dorsolateral lobes. Discussion. Polydora alloporis is closely related to P. cavitensis and P. hornelli by virtue of the specialized spines on the fifth setiger, which in all three species have hollow distal concavities. Polydora hornelli, however, has a distinct neck proximal to the concavity which is not found in the other two forms. The arrangement of the setae of the fifth setiger is similar in all of these species. The heavy dorsal folding of this segment seen in Polydora alloporis is not mentioned for either of the Asian forms. However, it is indicated in the illustrations of P. cavitensis and appears to be in an incipient stage in the figures of P. hornelli. An examination of paratypic material of P. cavitensis reveals the 1Pillai (1965, p. 152) mentions a similar pigmentation on the dorsum of P. cavitensis, but an examina- tion of two paratypes on loan from the British Museum (Natural History) indicates that this is also due to the dorsal aorta and its contents. 468 CALIFORNIA ACADEMY OF SCIENCES | [Proc. 47TH SER. Ficure 5. Polydora alloporis, new species. Left lateral view of anterior end. Ficure 6. Polydora alloporis, new species. Lateral view of pygidium, showing papillae. Ficure 7. Polydora alloporis, new species. Posterior view of pygidium, showing papillae. Scale indicates 2 mm. dorsal folding to be present, although poorly developed. The neuropodial crotchets are very similar to those of P. cavitensis; to a lesser extent they also resemble those of P. hornelli. The prostomium is distinctly bifurcated anteriorly in P. cavitensis and slightly notched in P. hornelli, whereas it is distally entire and exhibits a distinctive superior caruncular sheath in Polydora alloporis. The caruncle extends to the end of the second or third setiger in P. hornelli and to the end of the third in P. cavitensis. In Polydora alloporis it is almost always found as far back as the middle of the fourth or anterior border of the fifth setiger. Polydora cavitensis possesses an occipital cirrus and two pairs of eyes, as compared with no such cirrus and but one pair of eyes in P. alloporis. There is no evidence of an occipital cirrus or eyes in P. hornelli, but all three species VoL. XXXVII] LIGHT: A COMMENSAL SPIONID FROM A HYDROCORAL 469 show one or more raised bosses in this region. The anal segments of both P. alloporis and P. cavitensis are described above; that of P. hornelli remains un- known. Polydora alloporis also shows affinities with P. pacifica from Palau Island. In size, it approaches the latter species and reaches 75 mm. or more ii length, as against 85 mm. for the Palau animals. Both Polydora hornelli (31 mm. for an incomplete specimen of 92 segments) and P. cavitensis (15 mm.) are consider- ably smaller. The prostomium of P. pacifica is anteriorly notched and the caruncle reaches the anterior margin of the third setigerous segment. No eyes are indicated and the fifth segment does not appear to exhibit the dorsal folding of the other three forms. The modified major spines of the Palau specimens are described as having a lateral subdistal spur, as in Polydora ciliata (Johnston, 1838), and the companion setae appear much like the worn companion setae of P. alloporis, being termed “‘lancet-like’” by Takahashi. As we have seen, these setae, at least in P. alloporis, are but the worn lanceolate setae of the same basic design generally found in polydorids. The hooded crotchets of Polydora pacifica are described as possessing a distally directed spine on the main shaft at the lower point of attachment of the transparent hood. In these crotchets, the portion of the hood which shields the beak is less pronounced, and that covering the apical fang more flared than in any of the other three species in this complex. It is of interest, however, that one slide containing the crotchets of P. alloporis which were distorted by improper mounting of the coverslip, showed a very similar configuration to those described for P. pacifica. The pygidium of P. pacifica more nearly resembles that of P. alloporis than does that of P. cavitensis, except that the dorsal lobes in P. pacifica do not ap- proach one another so closely and the dorsal cleft is much wider, leaving the anus directly exposed above. Both Polydora pacifica and P. alloporis are found burrowing in branched corals and hydrocorals, Montipora species and Allopora californica, respectively, but the former is also commonly known to burrow into the inner nacreous layer of the pearl oyster, Pinctada margaritifera (Linné, 1758). Polydora hornelli is known only from crevices in oyster shells from fresh water at Manikpatna, Chilka Lake near Calcutta, India, and the Gulf of Manaar, Ceylon. Polydora cavitensis was collected from the Government Oyster Farm in Binakayan, Cavite, - Manila Bay, Luzon, Philippine Islands, likewise from among oysters. We have, then, two marine and two estuarine populations. According to Fisher (1938, p. 496), Allopora californica occurs at tempera- tures between 46° F. and 52° F. (7.7° C. to 11.1° C.) from the Farallon Islands south to Lower California at depths of up to 25 fathoms. It would not be sur- prising to find a similar distribution for Polydora alloporis. In this connection, 470 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. however, it is noteworthy that a sample of Allopora, near A. californica from an unknown depth off Anacapa Island near Santa Barbara, California, in the dry collections of the Department of Invertebrate Zoology, California Academy of Sciences, shows no evidence of the characteristic burrow-openings made by these animals. Allopora venusta Verrill, 1868, another hydrocoral which ranges from Wash- ington to Monterey Bay and which is also represented in the dry collections of the Academy, does exhibit the paired holes of Polydora, as do several other hydrocorals from Vancouver Island and Alaska. These holes are much smaller than those typically found in the coenosteum of Allopora californica, and since none of the polychaetes are available for study, it is uncertain whether these other hydrocorals harbor Polydora alloporis, another previously described polydorid, or an as yet new and undescribed species. An examination of the plates in a monographic work on northern Pacific hydrocorals (Fisher, 1938) reveals paired burrow openings in the coenosteum of the following species: Allopora porphyra (Fisher, 1931, pl. 59, fig. 2), A. stejnegeri Fisher, 1938 (pl. 56, fig. 2), and A. boreopacifica (Broch, 1932 [pl. HO, Hie.) 3) i ACKNOWLEDGMENTS I wish to express my gratitude to Messrs. Dennis Sullivan and Don Wobber for their collections of the Allopora from which the polychaetes were extracted. I am also indebted to Miss Gay Little of the Department of Zoology, University of California at Berkeley, for verifying the identity of Allopora californica, and to Dr. Victor A. Zullo and Mr. Allyn G. Smith, both of the California Academy of Sciences, for identifying the barnacle, Balanus (Armatobalanus) nefrens, and the gastropod, Pedicularia californica, respectively. Mr. Smith also took the remarkable photograph of the living speci- men of Polydora alloporis (fig. 1). I am especially grateful to Dr. J. David George, Curator of the Porifera and Polychaeta Section of the British Museum (Natural History) for his generous loan of two paratypes of Polydora cavitensis. NOTE. After the manuscript had gone to press, additional spionids were recovered from specimens of Allopora venusta from Monterey County, California (kindly provided by Gay Little), as well as from samples of an encrusting hydrocoral, Allopora species near A. petrograpta Fisher, 1938, from the lower littoral of the west coast of Vancouver Island, British Columbia. These worms are, in both cases, referred to Polydora alloporis. LITERATURE CITED FIsHEeR, WALTER K. 1938. Hydrocorals of the North Pacific Ocean. Proceedings of the United States National Museum, vol. 84, no. 3024, pp. 493-554, pls. 34-76. VoL. XXXVII] LIGHT: A COMMENSAL SPIONID FROM A HYDROCORAL 471 Pirrat, T. GOTTFRIED 1965. Annelida polychaeta from the Philippines and Indonesia. Ceylon Journal of Science (Biological Sciences), vol. 5, no. 2, pp. 110-177, 24 figs. SOUTHERN, ROLAND 1921. Fauna of the Chilka Lake. Polychaeta of the Chilka Lake and also of fresh and brackish waters in other parts of India. Memoirs of the Indian Museum, vol. V, pp. 563-659, pls. 19-31, 18 figs. TAKAHASHI, KEIzo 1937. Notes on the polychaetous annelid, Polydora pacifica, n. sp. Palao Tropical Biological Station Studies, Tokyo, no. 2, pp. 155-167, 9 figs., 5 tables. ZULLO, VicTor A. 1963. A review of the subgenus Armatobalanus Hoek (Cirripedia: Thoracica), with the description of a new species from the California coast. Annals and Magazine of Natural History, series 13, vol. 6, pp. 587-594, 1 fig., 1 key. 1966. Zoogeographic affinities of the Balanomorpha (Cirripedia: Thoracica) of the eastern Pacific. In: The Galapagos; Proceedings of the Symposia of the Galapagos International Scientific Project. Robert I. Bowman, ed., University of California Press, Berkeley, pp. 139-144, 2 tables, 1 map. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XXXVII, No. 15, pp. 473-488; 2 figs.; 1 table. October 13, 1970 BENTHIC FISHES CAST ASHORE BY GIANT WAVES NEAR POINT JOE, MONTEREY COUNTY, CALIFORNIA By W. IL. Follett California Academy of Sciences, San Francisco, California 94118 INTRODUCTION In the vicinity of Point Joe, Monterey County, California, on the morning of February 9, 1960, giant waves driven by winds of gale force cast ashore a number of fishes that normally inhabit one or more zones of the benthic environment. The powerful winds, which for three days battered the California coast from Humboldt County to San Luis Obispo County, generated waves up to 40 feet in height,’ causing extensive damage along the shoreline of the Monterey Peninsula. More than 50 boulders were cast onto or across the Seventeen Mile Drive near Point Joe (these were shown in photographs published on the front page of the Monterey Peninsula Herald, February 9, 1960). An automobile being driven along the Seventeen Mile Drive near Point Joe was picked up by a wave and moved about 150 feet. The Seventeen Mile Drive was partly torn out north of Cypress Point. A huge wave entered a third-story window of a waterfront hotel at Cannery Row, Monterey. A 300-foot pier and its boathouse were destroyed at Stillwater Cove. Suspecting that fishes might have been cast ashore by these enormous waves, Miss E. Eugenia Patten and Miss Priscilla J. Ferguson, of Carmel, California, immediately began to search the region south of Point Joe, and continued their search of that area from February 9 to February 24, 1960. By their assiduous 1No tsunami is known to have occurred in the eastern Pacific during February 1960. See Talley and Cloud (1960, p. 36). [473] 474 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 1. Offshore view of area where benthic fishes were cast ashore, showing logs that giant waves deposited over the channel of the intermittent stream. Photograph by E. Eugenia Patten, February 1960. efforts, they collected 36 fishes cast ashore by this storm, all but one of which rep- resent species that have been recorded from the outer sublittoral zone (depths from about 50 to 200 meters; see Zonation, p. 482). The relatively flat unforested area where these fishes were cast ashore is approximately 0.9 mile (by road) southward from Point Joe. It extends for some 90 yards in a northerly and southerly direction between the shoreline and the Seventeen Mile Drive, and varies in width from 20 to 25 yards. This area slopes gently toward the ocean, terminating at the top of an irregular bluff about 10 feet above normal high water. The narrow channel of a short intermittent stream which enters the Pacific Ocean at 36°35’47” N., 121°57’37” W. (see Monterey Sheet 1657 III SW, Army Map Service Series V895), crosses the area from east to west. (From the stream, the area extends some 75 yards northward and 15 yards southward.) An offshore view of this area (fig. 1) shows several logs that the giant waves deposited over the channel of the intermittent stream; an inshore view (fig. 2) shows a number of boulders that the waves cast ashore and lodged in two branches of the stream. The locality where these fishes were cast ashore is about four miles (by road) from Stillwater Cove, where thousands of blue lanternfish, Tarletonbeania crenu- laris (Jordan and Gilbert), were found dead on October 17, 1952, along some 200 yards of beach, at the high-tide level. The mass stranding of that bathypela- gic species may have been caused by attacking predators, since it occurred during VoL. XXXVIT] FOLLETT: BENTHIC FISHES CAST ASHORE 475 Ficure 2. Inshore view of area where benthic fishes were cast ashore, showing boulders that giant waves lodged in two branches of the intermittent stream. Photograph by E. Eugenia Patten, February 1960. clear weather, with only a slight wind and less than one-foot surf action (Aughtry, 1953, p. 190). FISHES REPRESENTED The collection comprises 36 fishes, representing seven species, five genera, and five families: Family EmprotocipAE—Surfperches Micrometrus aurora (Jordan and Gilbert )—reef perch Family ScoRPAENIDAE—Scorpionfishes Sebastes flavidus (Ayres)—yellowtail rockfish Sebastes rosaceus Girard—rosy rockfish Sebastes helvomaculatus Ayres—rosethorn rockfish Family GopripAE—Gobies Coryphopterus nicholsu (Bean)—blackeye goby Family BroTuLIDAE—Brotulas Brosmophycis marginata (Ayres)—red brotula Family OpHipiipAE—Cusk-eels Chilara taylori (Girard)—spotted cusk-eel 476 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. DISCUSSION The meristic data were taken from radiographs, and are expressed as by Hubbs and Lagler (1958). The abbreviation “CAS” precedes a catalog number of the Department of Ichthyology of the California Academy of Sciences. Family EmMprorocipAE—Surfperches Micrometrus aurora (Jordan and Gilbert). The reef perch (see Tarp, 1952, fig. 31), was originally described from 15 examples “taken in the bay of Monterey, and purchased by us in the San Fran- cisco market” (Jordan and Gilbert, 1880, p. 299, as Abeona aurora). Micrometrus aurora ranges from Tomales Bay, Marin County, California (Tarp, 1952, p. 86), to the first point south of the mouth of Rio San Telmo, Baja California (CAS 27142). The specimens from the vicinity of Rio San Telmo represent an extension of the recorded range from the southernmost locality noted by Tarp (1952, p. 86: 1’ miles south of the mouth of Rio San Isidro). This species has been recorded from Point Pinos (Jordan and Gilbert, 1881, p. 51), which is about 2'’2 miles generally northward from Point Joe. According to Hubbs (1921a, p. 183, as Amphigonopterus aurora), this species is restricted to the reefs, and commonly lives and breeds in the pools and channels of medium tidal height, particularly those that are largely open, free of eelgrass and algae, and floored with sand. Its habitat differs widely from that usual for the other species of this family, most of which live in the surf along sandy beaches, or in sheltered bays. The reef perch attains a standard length of 141 mm. (Hubbs, 1921a, p. 201). Material. One specimen (small adult), standard length 74 mm., CAS 26760. The characters and meristic data are in accord with those noted by Tarp (1952, p. 84). Family ScoRPAENIDAE—Scorpionfishes I concur with Matsubara (1943, p. 178) in regarding Sebastodes Gill (1861, p. 165) as a subgenus of Sebastes Cuvier (1829, p. 166). The vernacular “rockcod” for the species of this genus was criticized more than a century ago by Ayres (1854-1855, p. 94): “They are taken in rocky localities along the coast and in the Bay of San Francisco, and the title Rock Fish applies to them very well. One more inappropriate, on the contrary, than that of Rock Cod, could scarcely have been selected, inasmuch as they are very widely removed from the family in which the Codfishes are classed.” Neverthe- less “rockcod”’ still persists among sportsmen and commerical fishermen. Be- cause of the extent of this popular usage, “rockcod” may eventually be recognized as the “official” vernacular despite its impropriety—as was the vernacular “‘ling- cod” for Ophiodon elongatus Girard, also widely removed from the cods. VoL. XXXVI] FOLLETT: BENTHIC FISHES CAST ASHORE 477 Sebastes flavidus (Ayres). The yellowtail rockfish (see Phillips, 1957, fig. 17) was originally described from the San Francisco market (Ayres, 1863, p. 209, fig. 64). Sebastes flavidus ranges from the vicinity of Forrester Island, Alaska (West- rheim, 1966, p. 1469), to San Diego, California (Phillips, 1964, p. 28). The yellowtail rockfish occurs over reefs, especially in depths greater than 15 m. (SO feet), and also over deep sand and mud bottoms to depths of 183 m. (100 fathoms), but it is usually found at depths between 24 m. and 46 m. (be- tween 13 and 25 fathoms), according to Odemar, Wild, and Wilson (1968, p. 126). The young do not normally inhabit the tide-pools (Hubbs and Schultz, 1933, pp. 20, 21). I collected a young specimen (standard length 62 mm., CAS 25897) from the Monterey municipal wharf in 4.9 m. Specimens have been re- corded from Cordell Bank, Marin County, California, in 51 m. to 72 m. (Follett, 1952, p. 415; “28 to 40 fathoms’’); from Baynes Sound, British Columbia, in 55 m. to 72 m. (Clemens and Wilby, 1961, p. 254; “30 to 40 fathoms”); from the southern edge of Astoria Canyon, Oregon, in 110 m. to 133 m. (Pereyra, Pearcy, and Carvey, 1969, p. 2211; “60-73 fath’’). It has been taken in depths of 187 m. to 194 m. (Westrheim, 1966, p. 1469) and 366 m. or more (Alverson, Pruter, and Ronholt, 1964, fig. 43; ‘200-299 fathoms’’). The yellowtail rockfish attains a length of 66 cm. (Phillips, 1957, p. 56; Clemens and Wilby, 1961, p. 253; Miller and Gotshall, 1965, p. 117; ‘26 inches”’), as determined from Canadian specimens (see Phillips, 1968, Appendix A-5). The maximum length recorded from California is about 58 cm. (Phillips, 1964, pp. 19, 23 25 inches’). Material. One specimen (young), standard length 73 mm., CAS 26761. The characters and meristic data are in accord with those noted by Phillips (1957, p. a7. 1964) p. 28). Sebastes rosaceus Girard. The rosy rockfish (see Phillips, 1957, fig. 47), was originally described from San Diego, California (Girard, 1854, p. 146). Sebastes rosaceus ranges from Cordell Bank, Marin County, California (Follett, 1952, p. 417), to Turtle Bay, Baja California (Phillips, 1957, p. 116). Records from farther north than Cordell Bank appear to have been based on misidentified specimens of S. helvomaculatus (Lo-chai Chen, personal correspon- dence, January 20, 1970). This species has been recorded from reefs in and near Monterey Bay, Cali- fornia, in less than 15 m. to more than 46 m. (Miller, Odemar, and Gotshall, 1967, pp. 95, 96, 102, 103; “less than 50 feet to more than 150 feet”); from Cordell Bank, Marin County, California, in 51 m. to 62 m. (Follett, 1952, p. 417; “28 to 34 fathoms’’); from off San Nicolas Island, California, in 58 m. to 60 m. 478 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. (Gilbert, 1915, p. 336; “32-33 fathoms”’). It has been taken at depths as great as 130 m. to 140 m. (Lo-chai Chen, personal correspondence, December 1, 1969). The rosy rockfish attains a length of 36 cm. (Miller and Gotshall, 1965, p. 116; “14.2 inches’’). Material. Four specimens (young), standard length 53 mm. to 67 mm., CAS 26762; identification confirmed by Lo-chai Chen. Four other specimens, stan- dard length 37 mm. to 60 mm., may represent Sebastes rosaceus, but they have not been identified with certainty. Sebastes helvomaculatus Ayres. The rosethorn rockfish (see Phillips, 1957, fig. 48) was long ago described by Ayres (1859, p. 26, fig. 8), from a specimen obtained in the San Francisco mar- ket. However, this species was for nearly a century thereafter relegated to the synonymy of Sebastes rosaceus Girard—as by Gill (1864, p. 147), Jordan and Gilbert (1883, p. 667), Eigenmann and Beeson (1894, p. 395), Jordan and Evermann (1898a, p. 1808), and Jordan, Evermann, and Clark (1930, p. 367). The distinctive coloration of this species was noted by Carl L. Hubbs (personal communication). Later, Phillips (1957, pp. 30, 31, 118, 119) noted (in addition to the coloration) distinctive morphometric characters, which, however, were considered inadequate by Heyamoto and Hitz (1962, p. 848) and by Westrheim (19655. 232). Sebastes helvomaculatus ranges from the Gulf of Alaska (Westrheim, 1965, p. 232) to Guadalupe Island, Baja California (Phillips, 1957, p. 118). This species has been recorded from depths of 133 m. (Westrheim, 1965, p. 232; “73 fathoms”) to 457 m. (Heyamoto and Hitz, 1962, p. 848; “250 fathoms’’). The rosethorn rockfish attains a length of 33 cm. (Phillips, 1957, p. 118; “S.mnches’”)). Material. One specimen (large young), standard length 89 mm., CAS 27139; identified by Lo-chai Chen. One or more of the four small specimens noted under Sebastes rosaceus as not identified with certainty may represent Sebastes helvo- maculatus. Family GopiraAr—Gobies Coryphopterus nicholsii (Bean). The blackeye goby (see Barnhart, 1936, fig. 245, as Rhinogobiops nicholsi; see also Ebert and Turner, 1962, fig. 1), also called “bluespot goby” and “crested goby,” was originally described from a specimen taken at Departure Bay, British Columbia, in 37 m. (Bean, 1882, p. 469, as Gobius nicholsii; “20 fathoms”). Coryphopterus nicholsti ranges from Skidegate Channel, Queen Charlotte Islands (Clemens and Wilby, 1961, p. 359), to San Martin Island, Baja Califor- nia (Limbaugh, 1955, p. 120). Vor. XXXVIT] FOLLETT: BENTHIC FISHES CAST ASHORE 479 This species is generally known as an inshore, shallow-water, benthic inhab- itant (Berry and Perkins, 1966, p. 676). According to Gilbert (1915, p. 359), this species “has also been taken in shallow water, but not between tides, at Monterey.” Two specimens were collected within the littoral (intertidal) zone, by Rolf L. Bolin, on August 18, 1947, at a rocky point 0.5 mile south of Malpaso Creek, Monterey County. Using small pieces of prawn for bait, I collected with rod and line two specimens (Stanford Natural History Museum catalog nos. 35015 and 35016) on March 16, 1940, in San Francisco Bay, at a depth of 6 m., from the Van Ness Avenue pier, San Francisco. It has been recorded from off San Elijo Lagoon, San Diego County, in 12 m. and 18 m. (Turner, Ebert, and Given, 1965, p. 109; “40 feet and 60 feet”), and from off Hermosa Beach and Santa Monica in 18 m. (Ebert and Turner, 1962, p. 250; ‘60 feet”). Two speci- mens, standard length 36 mm. and 37 mm. (CAS 27140), were collected south- west of South Coronado Island, Baja California, in 18 m. to 25 m. It has been recorded from near Pacific Grove, California, in 18 m. or 27 m. (Snyder, 1913, p. 459; “10 or 15 fathoms”’); from British Columbia at six localities—in depths of 37 m. or less (Clemens and Wilby, 1961, p. 359; “20 fathoms or less”): from Anacapa Passage, California (Jordan and Evermann, 1898b, p. 2218; ‘Albatross station 2944”), in 55 m. (see Tanner, 1892, p. 493; ‘30 fathoms”; see also U.S. Coast and Geodetic Survey Chart no. 5202, 1937); and from the vicinity of San Diego in 91 m. (Starks and Mann, 1911, p. 16, as Rhinogobius nicholsii; “50 fathoms’). Also, it has been reported as sometimes taken in more than 640 m. (Barnhart, 1936, p. 81; “2100 feet”). Its pelagic prejuvenile stages have been taken at depths of approximately 1,863 m. and 2,234 m. (Berry and Perkins, 1966, pp. 626, 627, 676). The blackeye goby attains a length of about 15 cm. (Ebert and Turner, 1962, p2250;) Ganches’’)). Material. Seven specimens (young to adult), standard length 33 mm. to 65 mm., CAS 26763. (The stomach of a 54-mm. specimen contained a gastropod, Turbonilla species, 2.8 mm. in length, identified by Allyn G. Smith.) In all seven specimens, the sixth dorsal spine is remote from the fifth, and the attenuate, unsegmented first element of the second dorsal fin, as well as that of the anal fin, appears to represent a spine. So interpreted, the dorsal formula is VI-I,13 in four specimens and VI-I,12 in three; the anal formula is I,12 in three specimens and IJ,11 in four. There are 17 segmented caudal rays (counting _ both unbranched and branched rays) in six specimens. There are 2 unbranched segmented caudal rays above and 1 below in the three specimens in which these rays are discernible. The vertebral formula is 10 + 16 = 26 in all seven specimens. The interpretation of the first element of the second dorsal fin and of the anal fin as a spine is in accord with that of Hubbs (1943, p. 134), Ginsburg (1953, p. 18), and Bohlke and Robins (1960, p. 104). The dorsal, anal, and 480 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. vertebral counts are in accord with those of Clothier (1951, p. 73). The caudal- ray counts are in accord with those of Ginsburg (1945, pp. 136, 137). Family BrotuLIpDAE—Brotulas Pending further study of the ophidioids, I retain the family Brotulidae Swainson (1838, p. 317, as Brotulinae*) as distinct from the family Ophidiidae Bonaparte (1831, p. 38, as Ophididae*). On the basis of the otoliths, these two nominal families should not be merged according to John E. Fitch (personal communication, May 7, 1969). Brosmophycis marginata (Ayres). The red brotula (see Best, 1957, fig. 1), was originally described from a 305-mm. (12-inch) specimen collected more than a century ago near the entrance to San Francisco Bay (Ayres, 1854, p. 202, as Brosmius marginatus). Brosmophycis marginata ranges from Petersburg, Alaska (Schultz and De- Lacy, 1936, p. 142), at least to Ensenada, Baja California (Fitch, 1968, p. 24). This species was recorded from Puget Sound by Gilbert and Thompson (1905, p. 985); from five other localities in Washington, by Schultz and DeLacy (1936, p. 142); and from British Columbia at four localities, including English Bay at 55 m. (‘30 fathoms’), by Clemens and Wilby (1961, p. 394). Ten specimens, taken at seven localities along the coast of California, from off the mouth of the Klamath River to Ship Rock, Santa Catalina Island, in depths of 37 m. to 91 m., were recorded by Best (1957, p. 97; “120 to 300 feet”). It is said to occur mostly in depths of 18 m. to 122 m. (Fitch, 1968, p. 24; “60 to 400 feet”’). A specimen deposited at the Cabrillo Marine Museum, San Pedro, California, was washed up in a severe storm on December 3, 1939, at Cabrillo Beach. Larvae have been collected north of Santa Catalina Island, California, and off E] Des- canso, Baja California (Ahlstrom, 1959, p. 110). Selle (1924, p. 143) recorded a specimen from “90 miles south of Los Angeles Harbor.” This record, published under the title “San Pedro Rarities,” may constitute the basis for the statement by Jordan, Evermann, and Clark (1930, p. 479) that this species had been “recently taken at San Pedro, Calif.” This “San Pedro” record was apparently accepted by Barnhart (1936, p. 92), Schultz (1936, p. 197), and Schultz and DeLacy (1936, p. 142). The red brotula has been said to reach a length of nearly 46 cm. (Jordan and Gilbert, 1881, p. 65; “18 inches’’). Material. Eleven specimens (large young to small adult), standard length 101 mm. to 250 mm., CAS 26764: 2 The priority of the name of a taxon in the family-group is not affected by elevation in rank within the group (International Code of Zoological Nomenclature, 1964, Article 23c). ® A family-group name of which the suffix is incorrect is available with its original date and authorship, but in properly emended form (International Code of Zoological Nomenclature, 1964, Article 11 (e) (ii). VoL. XXXVIT] Standard length (mm.) 101 103 107 112 119 130 133 149 165 216 250 FOLLETT: BENTHIC FISHES CAST ASHORE 481 Dorsal Anal rays rays Caudal rays Vertebrae 101 73 16 (8 + 8) 16 + 48 = 64 99 76 16 (8 + 8) 17+47= 64 101 75 16 (8+ 8) 16 + 47 = 63 101 74 16 (8 + 8) 16 + 48 = 64 101 75 16 (8 + 8) 16+ 48 = 64 104 74 16 (8+ 8) 16 + 49 = 65 105 81 16 (8 + 8) 16+ 48 = 64 = 76 16 (8+ 8) 16+ 49 = 65 104 15 16 (8 + 8) 16 + 48 = 64 — 73 16 (8 + 8) 16 + 48 = 64 110 77 17 (9 + 8) 16 + 49=65 The caudal-ray counts of 16 (8 + 8) in ten specimens and 17 (9 + 8) in one specimen suggest that Brosmophycis marginata normally has twice the number of caudal rays noted for two other brotulids known from California: 8 (4+ 4) 8 (444) 9(5+4) 8 (4+4) 5 (sey in a specimen of Cataetyx rubrirostris Gilbert, standard length 60 mm. (CAS 27145), picked up at the drift-line on Moss Beach, Monterey County, California, on April 5, 1959, by Miss E. Eugenia Patten; in a specimen of Cataetyx rubrirostris collected off northern Oregon (Grinols and Greenfield, 1966, p. 212); in a specimen of Cataetyx rubrirostris collected off northern Oregon (Grinols and Greenfield, 1966, p. 212); in a specimen of Parabassogigas grandis (Gunther), standard length 127 cm. (CAS 25724), collected some 40 miles west-south- west of Farallon Lighthouse, California, on September 22, 1952, by Warren Beadle and R. G. Hamilton; in a specimen of Parabassogigas grandis, total length 135 cm. (cata- log no. 39255 in the Zoological Institute, University of Tokyo), collected February 28, 1938 (?), off Katsu-ura, Wakayama Pre- fecture (eastern coast of Kii Peninsula), south of Nagoya, Japan. Ray counts of D. 92, A. 70 taken by Jordan and Evermann (1898b. p. 2502) from a specimen of Brosmophycis marginata collected off San Francisco were corrected to D. 100, A. 76 by Gilbert and Thompson (1905, p. 985). Vertebral counts of 16 + 47 = 63, 16 + 48 = 64, 17 + 47 = 64, and 17 + 48 = 65 were noted by Best (1957, p. 98). Family OpHip1ipAE—Cusk-eels Chilara taylori (Girard). The spotted cusk-eel (see Herald, 1953, figs. 1-3, as Otophidium taylort) was originally described from six small specimens, the largest 3% inches in 482 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER. length, taken in “the sands of Monterey Beach,” California (Girard, 1858, p. 138, as Ophidion taylori). This species has long been referred to the genus Otophidium Gill in Jordan (1887, p. 914), but its high vertebral counts indicate that it should be referred instead to the genus Chilara Jordan and Evermann (1896, p. 482). The type- species of Otophidium (Genypterus omostigma Jordan and Gilbert, 1882, p. 301; by original designation) has a relatively low vertebral count (14 + 43 = 57, as determined from a radiograph of the holotype; see Bohlke and Robins, 1959, p. 42). In contrast, the type-species of Chilara (Ophidion taylori, by original des- ignation) has a high vertebral count (18 + 69 = 87, etc.; see vertebral counts, below). Therefore, on the basis of the vertebral count, the genus Chilara appears worthy of recognition. The binomen Chilara taylori is well known from the usage of Jordan and Everman (1896, p. 482; 1898b, p. 2489), Starks and Morris (1907, p. 240), and Snyder (1913, p. 460). Chilara taylori ranges from northern Oregon to San Cristobal Bay, Baja California (Fitch and Lavenberg, 1968, p. 75). Recorded depths of capture are 1.2 m. (Herald, 1953, p. 381; “4 feet”); 13 m. to 126 m. (Gilbert, 1895, p. 472; “‘7 to 69 fathoms”) ; 18 m. or 22 m. (Snyder, 1913, p. 460; “10 or 12 fathoms”); 18 m. to 244 m. or more (Fitch and Laven- berg, 1968, p. 75; “60 to 800 feet or more’’) ; 55 m. (Hubbs, 1916, p. 166). A specimen of this species has been taken from the stomach of Sebastes auri- culatus Girard (Hubbs, 1921b, p. 28), and another specimen, about 25 cm. in length, from the stomach of a 115-cm. specimen of Alepisaurus ferox Lowe (Roedel and McCully, 1939, p. 35, as Alepisaurus aesculapius ). The spotted cusk-eel has been said to reach a length of about 36 cm. (Jordan and Gilbert, 1881, p. 65, as Ophidium taylori; “14 inches’’). Material. Seven specimens (large young to small adult), standard length 74 mm. to 190 mm., CAS 26765. Six specimens have each 4 + 5 caudal rays, the articulation of these rays with the hypurals somewhat resembling that in Ophidion barbatum Linnaeus (see Whitehouse, 1910, pl. 50, fig. 30, as Ophidium barbatum). Four specimens have each a vertebral formula of 18 + 69 = 87; one, 18 + 70 = 88; one, 18 + 71 = 89; and one, 19 + 71 = 90. (Of the specimens noted by Clothier, 1951, p. 79, pl. 23, two had each a total of 87 vertebrae; eight, 88; and one, 89.) ZONATION I follow the terminology of Hedgpeth (1957, fig. 1) for the classification of the benthic environment below the littoral (intertidal) zone: Inner sublittoral, between low water and a depth of about 50 meters; Outer sublittoral, depths from about 50 to 200 meters; Bathyal, depths below 200 meters (to about 4000 meters). VoL. XXXVI] FOLLETT: BENTHIC FISHES CAST ASHORE 483 Of the seven species represented among the fishes cast ashore near Point Joe, Sebastes helvomaculatus is noteworthy as an inhabitant of the bathyal zone, al- though it has been recorded as well from the outer sublittoral zone: Sebastes rosaceus and Brosmophycis marginata have been recorded from both the outer and the inner sublittoral zone; Coryphopterus nicholsii and Chilara taylori are known to range from the outer sublittoral into the littoral (intertidal) zone; Sebastes flavidus descends to the lower limits of the outer sublittoral, but a young example, of a size comparable to that of the specimen in this collection, has been taken within the inner sublittoral zone; only Micrometrus aurora appears re- stricted to the littoral (intertidal) zone. ACKNOWLEDGMENTS I wish to express my deep appreciation to Miss E. Eugenia Patten, member of the California Academy of Sciences, and to Miss Priscilla J. Ferguson, for this remarkable collection; to Dr. Tokiharu Abe, of Tokaiku Suisan Kenkyujo, Tokyo, for radiographs of the Katsu-ura specimen of Parabassogigas grandis; to Dr. Lo-chai Chen, of San Diego State College (formerly of the Scripps Institu- tion of Oceanography ), for determination of the specimens referable to Sebastes rosaceus and S. helvomaculatus; to Mr. William K. Cloud, of the U.S. Coast and Geodetic Survey, for records of tsunamis during 1960; to Mrs. Lillian J. Demp- ster, of the California Academy of Sciences, for assistance with the manuscript; to Mr. Maurice C. Giles, of the California Academy of Sciences, for enlargements of the photographs; to Dr. Carl L. Hubbs, of the Scripps Institution of Ocean- ography, for assistance and advice and for the Cabrillo Beach record of Brosmo- phycis marginata; to Dr. C. Richard Robins, of the University of Miami, for advice on Chilara; and to Mr. Allyn G. Smith, of the California Academy of Sciences, for determination of the gastropod found in the stomach of a speci- men of Coryphopterus nicholsit. LITERATURE CITED AHLSTROM, ELBertT H. 1959. Vertical distribution of pelagic fish eggs and larvae off California and Baja Cali- fornia. United States Fish and Wildlife Service Fishery Bulletin, vol. 60, no. 161, pp. i-iv, 107-146, figs. 1-13. ALVERSON, D. L., A. T. PRUTER, AND L. L. RONHOLT 1964. A study of demersal fishes and fisheries of the northeastern Pacific Ocean. H. R. MacMillan Lectures in Fisheries. Institute of Fisheries, The University of Brit- ish Columbia, Vancouver, 190 pp., 72 figs. AuGHTRY, Ropert H. 1953. A note on mass mortality of the myctophid fish Tarletonbeania crenularis. Copeia, 1953, no. 3, pp. 190-192, fig. 1. AYRES, WILLIAM O. 1854. [Descriptions of Brosmius marginatus and Syngnathus griseo-lineatus.| The Pacific, vol. 3, no. 51, Oct. 27, p. 202 [misprinted “102” ]. 484 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. 1854-1855. New species of Californian fishes. Proceedings of the Boston Society of Nat- ural History, vol. 5, pp. 94-103. 1859. [New fishes of the Pacific coast.]| Proceedings of the California Academy of Natural Sciences, vol. 2, pp. 25-32. 1863. [Descriptions of Sebastodes flavidus and Sebastodes ovalis.| Proceedings of the California Academy of Natural Sciences, vol. 2, pp. 209-211, figs. 64-66. BARNHART, PERCY SPENCER 1936. Marine fishes of southern California. University of California Press, Berkeley, vi + 209 pp., 290 figs. BEAN, TARLETON H. 1882. Notes on a collection of fishes made by Captain Henry E. Nichols, U.S.N., in British Columbia and southern Alaska, with descriptions of new species and a new genus (Delolepis). Proceedings of the United States National Museum, vol. 4, pp. 463-474. 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Fisheries Research Board of Canada, Bulletin 68 (second edition), 443 pp., colored frontispiece, 6 colored pls., 281 figs. CLOTHTER, CHARLES R. [1951]* A key to some southern California fishes based on vertebral characters. State of California Division of Fish and Game, Fish Bulletin 79, 1950, 83 pp., 23 pls., 22 figs. CuviER, GEORGES 1829. Le regne animal distribué d’aprés son organisation, pour servir de base a Vhistoire naturelle des animaux et d’introduction a lanatomie comparée. Vol. 2 (new edition). Paris, xv + 406 pp. EBERT, EARL E., AND CHARLES H. TURNER 1962. The nesting behavior, eggs and larvae of the bluespot goby. California Fish and Game, vol. 48, no. 4, pp. 249-252, figs. 1-4. EIGENMANN, Cart H., anp CHARLES H. BEESON 1894. A revision of the fishes of the subfamily Sebastinae of the Pacific coast of Amer- ica. Proceedings of the United States National Museum, vol. 17, pp. 375-407. ‘ Date from Follett (1952, p. 424). Square brackets indicate extrinsic evidence of the date, in accordance with Recommendation 22A(3) of the International Code of Zoological Nomenclature (1964). Vor. XXXVIT] FOLLETT: BENTHIC FISHES CAST ASHORE 485 Fitcu, Joun E. 1968. Otoliths and other fish remains from the Timms Point silt (early Pleistocene) at San Pedro, California. Los Angeles County Museum, Contributions in Science, no. 146, 29 pp., 4 figs. Fitcu, JOHN E., AND RoBert J. LAVENBERG 1968. Deep-water teleostean fishes of California. University of California Press, Berke- ley and Los Angeles, 155 pp., 74 figs. FoueTtT, W. I. 1952. Annotated list of fishes obtained by the California Academy of Sciences during six cruises of the U.S.S. Mulberry conducted by the United States Navy off central California in 1949 and 1950. Proceedings of the California Academy of Sciences, 4th ser., vol. 27, no. 16, pp. 399-432, pls. 21-26. GILBERT, CHARLES H. 1895. The ichthyological collections of the steamer Albatross during the years 1890 and 1891. Report of the United States Commissioner of Fish and Fisheries for the Year Ending June 30, 1893, part 19, pp. 393-476, pls. 20-35. 1915. Fishes collected by the United States Fisheries steamer “Albatross” in southern California in 1904. Proceedings of the United States National Museum, vol. 48, pp. 305-380, pls. 14-22. GILBERT, CHARLES H., AND JosEPH C. THomMPpsoNn 1905. Notes on the fishes of Puget Sound. Proceedings of the United States National Museum, vol. 28, pp. 973-987, figs. 1-3. GILL, THEODORE 1861. Notes on some genera of fishes of the western coast of North America. Proceed- ings of the Academy of Natural Sciences of Philadelphia, vol. 13, pp. 164-168. 1864. Critical remarks on the genera Sebastes and Sebastodes of Ayres. Proceedings of the Academy of Natural Sciences of Philadelphia, 1864, pp. 145-147. GInsBurG, ISAAC 1945. Contribution to a methodology in the caudal fin ray count of fishes and its use in classification. Copeia, 1945, no. 3, pp. 133-142. 1953. Ten new American gobioid fishes in the United States National Museum, including additions to a revision of Gobionellus. Journal of the Washington Academy of Sciences, vol. 43, no. 1, pp. 18-26. GIRARD, CHARLES 1854. Observations upon a collection of fishes made on the Pacific coast of the United States, by Lieut. W. P. Trowbridge, U.S.A., for the Museum of the Smithsonian Institution. Proceedings of the Academy of Natural Sciences of Philadelphia, vol. 7, pp. 142-156. 1858. Fishes. Jn General report upon the zoology of the several Pacific railroad routes. Reports of explorations and surveys...for a railroad from the Mississippi River to the Pacific Ocean. U.S. Senate Executive Document no. 78, 33rd Congress, 2nd Session, vol. 10, part 4, pp. i-xiv, 1-400, pls. 7, 8, 13, 14, 17, 18, 22, 26, 29, 30, 34, 37, 40, 41, 48, 53, 59, 61, 64, 65, 71. Grinots, RicHArRD B., AND DAvip W. GREENFIELD 1966. New records of Cataetyx rubrirostris Gilbert from the northeastern Pacific Ocean. California Fish and Game, vol. 52, no. 3, pp. 211-213, fig. 1. HEpDGPETH, JOEL W. 1957. Classification of marine environments. Geological Society of America. Memoir 67, vol. 1, pp. 17-27, figs. 1-5. 486 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER. HERALD, Ear S. 1953. Spotted cusk-eel, the strange fish that stands on its tail. California Fish and Game, vol. 39, no. 3, pp. 381-384, figs. 1-3. Heyamorto, Hrromu, AnD CuHartes R. Hitz 1962. Northern range extensions of three species of rockfish (Sebastodes rubrivinctus, S. aurora, and S. helvomaculatus). Copeia, 1962, no. 4, pp. 847-848, fig. 1. Husss, Caru L. 1916. Notes on the marine fishes of California. University of California Publications in Zoology, vol. 16, no. 13, pp. 153-169, pls. 18-20. 1921a. The ecology and life-history of Amphigonopterus aurora and of other viviparous perches of California. Biological Bulletin, vol. 40, no. 4, pp. 181-209, figs. 1-5. 1921b. Notes on Sebastodes. Copeia, no. 93, pp. 27-28. 1943. [Review of] Fishes of the Phoenix and Samoan islands collected in 1939 during the expedition of the U.S.S. “Bushnell,” by Leonard P. Schultz. Copeia, 1943, no. 2, pp. 133-134. Husss, Cart L., anp Kart F. LAGLER 1958. Fishes of the Great Lakes region. Bulletin of the Cranbrook Institute of Science, vol. 26 (revised edition), xiii + 213 pp., 44 colored pls., 251 figs. Hugss, Cart L., AND LEONARD P. SCHULTZ 1933. Descriptions of two new American species referable to the rockfish genus Sebas- todes, with notes on related species. University of Washington Publications in Biology, vol. 2, no. 2, pp. 15-44, pls. 1-2. JorDAN, DAvip STARR 1887. A catalogue of the fishes known to inhabit the waters of North America, north of the Tropic of Cancer, with notes on the species discovered in 1883 and 1884. Report of the United States Commissioner of Fish and Fisheries for 1885, part 13, pp. 789-973. Jorpan, DAvip Starr, AND BARTON WARREN EVERMANN 1896. A check-list of the fishes and fish-like vertebrates of North and Middle America. Report of the United States Commissioner of Fish and Fisheries for the Year Ending June 30, 1895, part 21, pp. 207-584. 1898a. The fishes of North and Middle America: a descriptive catalogue of the species of fish-like vertebrates found in the waters of North America, north of the Isthmus of Panama. Part 2. Bulletin of the United States National Museum, no. 47, pp. i-xxx, 1241-2183. 1898b. The fishes of North and Middle America: a descriptive catalogue of the species of fish-like vertebrates found in the waters of North America, north of the Isthmus of Panama. Part 3. Bulletin of the United States National Museum, no. 47, pp. i-xxiv, 2183a-3136. JorDAN, Davip STARR, BARTON WARREN EVERMANN, AND Howard WALTON CLARK 1930. Check list of the fishes and fishlike vertebrates of North and Middle America north of the northern boundary of Venezuela and Colombia. Report of the United States Commissioner of Fisheries for the Fiscal Year 1928, with Appen- dixes, part 2,iv + 670 pp. Jorpan, Davin S., AND CHARLES H. GILBERT 1880. Description of a new embiotocoid (Abeona aurora), from Monterey, California, with notes on a related species. Proceedings of the United States National Museum, vol. 3, pp. 299-301. 1881. Notes on the fishes of the Pacific coast of the United States. Proceedings of the United States National Museum, vol. 4, pp. 29-70. Vor. XXXVIT] FOLLETT: BENTHIC FISHES CAST ASHORE 487 1882. Notes on fishes observed about Pensacola, Florida, and Galveston, Texas, with description of new species. Proceedings of the United States National Museum, vol. 5, pp. 241-307. 1883.° Synopsis of the fishes of North America. Bulletin of the United States National Museum, no. 16, lvi + 1018 pp. LIMBAUGH, CONRAD 1955. Fish life in the kelp beds and the effects of kelp harvesting. University of Califor- nia, La Jolla, Institute of Marine Resources, IMR Reference 55-9, iii + 158 pp., 21 figs. MAtTsupara, KivoMAtsu 1943. Studies on the scorpaenoid fishes of Japan. Anatomy, phylogeny and taxonomy (II). The Transactions of the Sigenkagaku Kenkyusyo, no. 2, pp. 171-486, pls. 1-4, figs. 67-156. Miter, Dante J., AND DANIEL GOTSHALL 1965. Ocean sportfish catch and effort from Oregon to Point Arguello, California, July 1, 1957—June 30, 1961. California Department of Fish and Game, Fish Bulletin 130, 135 pp., 16 figs. Miter, Danter J., Metvyn W. OpEMAR, AND DANIEL W. GoTsHALL 1967. Life history and catch analysis of the blue rockfish (Sebastodes mystinus) off central California, 1961-1965. California Department of Fish and Game, Marine Resources Operations, MRO Reference no. 67-14, iii + 130 pp., 46 figs. OpEMAR, MEtvyn W., Paut W. WILbD, AND KENNETH C. WILSON 1968. A survey of the marine environment from Fort Ross, Sonoma County, to Point Lobos, Monterey County. California Department of Fish and Game, Marine Resources Operations, MRO Reference no. 68-12, v + 238 pp., 28 figs. PEREYRA, WALTER T., WILLIAM G. PEARCY, AND Forrest E. Carvey, JR. 1969. Sebastodes flavidus, a shelf rockfish feeding on mesopelagic fauna, with consider- ation of the ecological implications. Journal of the Fisheries Research Board of Canada, vol. 26, no. 8, pp. 2211-2215, figs. 1-2. PuHitiires, JuLtus B. 1957. A review of the rockfishes of California (family Scorpaenidae). State of Califor- nia Department of Fish and Game, Fish Bulletin 104, 158 pp., 66 figs. 1964. Life history studies on ten species of rockfish (genus Sebastodes). California Department of Fish and Game, Fish Bulletin 126, 70 pp., 44 figs. 1968. Review of rockfish program. California Department of Fish and Game, Marine Resources Operations, MRO Reference no. 68-1, 27 pp., 8 figs., 8 appendices. ROEDEL, PHL M., AnD Howarp McCULLyY 1939. Occurrence of the wolf-fish, Alepisaurus aesculapius (Bean) in southern Califor- nia. California Fish and Game, vol. 25, no. 1, pp. 35-37. ScHuttz, Leonarp P. 1936. Keys to the fishes of Washington, Oregon and closely adjoining regions. Univer- sity of Washington Publications in Biology, vol. 2, no. 4, pp. 103-228, figs. 1-50. ScHuLtTz, LEonarD P., AND ALLAN C. DELAcy 1936. Fishes of the American Northwest. A catalogue of the fishes of Washington and Oregon, with distributional records and a bibliography. Mid-Pacific Magazine, vol. 49, no. 2, pp. 127-142. S[ELLE], W[iBur] A. 1924. San Pedro rarities. California Fish and Game, vol. 10, no. 3, pp. 143-144. 5 The date “1883” on the original cover was confirmed by Jordan (1887, p. 789), who noted the date of publication as ‘‘about April 1, 1883.” CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. : = tr = ad SxvpnR; Bieabotocksens or 1913. 4Notes | on Rauzania makua Jenkins and other species of fishes of rare occurrence es »on the California coast. Proceedings of the United States National Museum, eric = st Nole44, no. 1961, pp. 455-460, pl. 63. Starks, Epwin CHAPIN, AND WILLIAM M. MANN ——— = 19 New and rare fishes from southern California. pe of California Publica- _ tions in Zoology, vol. 8, no. 2, pp. 9-19, figs. 1-2 STARKS, EDWIN Carin, AND Eart Leonarp Morris 1907. The marine fishes of southern California. University of California Publications, Zoology, vol. 3, no. 11, pp. 159-251, pl. 21. SwAINnson, WILLIAM 1838. The natural history of fishes, amphibians, & reptiles, or monocardian animals, Vol. 1. London, vi + 368 pp., 100 figs. TaLLey, H. Carrorr, Jr., AND WiLLIAM K. CLoup 1962. United States earthquakes, 1960. U.S. Department of Commerce, Coast and Geo- detic Survey, Washington, iv + 90 pp., 8 figs. Tanner, Z. L. 1892. Report upon the investigations of the U.S. Fish Commission steamer Albatross for the year ending June 30, 1889. Report of the United States Commissioner of Fish and Fisheries for 1888, part 16, pp. 395-512, pls. 50-52. Tarp, Frep HarALp 1952. A revision of the family Embiotocidae (the surfperches). State of California Department of Fish and Game, Fish Bulletin 88, 99 pp., 32 figs. TURNER, CHARLES H., EArt E. Epert, AND ROBERT R. GIVEN 1965. Survey of the marine environment offshore of San Elijo Lagoon, San Diego County. California Fish and Game, vol. 51, no. 2, pp. 81-112, figs. 1-16. WESTRHEIM, S. J. 1965. Northern range extensions for four species of rockfish (Sebastodes goodei, S. helvomaculatus, S. rubrivinctus, and S. zacentrus) in the North Pacific Ocean. Journal of the Fisheries Research Board of Canada, vol. 22, no. 1, pp. 231-235, figs. 1-2 1966. Northern range extensions for three species of rockfish (Sebastodes flavidus, S. paucispinis, and S. pinniger) in the North Pacific Ocean. Journal of the Fish- eries Research Board of Canada, vol. 23, no. 9, pp. 1469-1471. WHITEHOUSE, RIcHARD H. 1910. The caudal fin of the Teleostomi. Proceedings of the Zoological Society of London, 1910, pp. 590-627, pls. 47-50, fig. 57. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XXXVII, No. 16, pp. 489-500; 30 figs. uly 23, 1971 A LATE TERTIARY DIATOM FLORA FROM OREGON By W. N. Orr University of Oregon, Eugene, Oregon 97403 Judi Ehlen Army Map Service, Brookmont, Maryland and J. B. Zaitzeff Manned Spacecraft Center, Houston, Texas ApstRACT: Siliceous microfossils including diatoms, radiolaria and _ silicoflagellates were extracted from an exposure of diatomite in southwest Oregon. The diatom flora here is indicative of a Late Miocene (Delmontian) age and represents a coastal temperate environment. INTRODUCTION A small exposure of marine diatomite in the vicinity of Bandon, Oregon, was sampled and processed for microfossils. In addition to a diverse radiolarian fauna and a few silicoflagellates, some 15 species of diatoms were successfully extracted. Although a number of diatom floras have been described from nonmarine sediments in Oregon, marine diatomites are rare in this state and elsewhere in the Pacific Northwest, and little has been published on siliceous marine microfossils from this area. By comparison, exposures of marine diatomites in California are well known, and much has been published with regard to siliceous plant and animal microfossils in that state. Recent papers by Wornardt (1967a, b) are a particularly useful review of the biostratigraphy, taxonomy, and distribution of late Tertiary diatoms in California. [489] Marine Biological Laboratory LIBRARY AUG3 1971 | WOODS HOLE, MASS. 490 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. DIATOM LOCALITY Ficure 1. Locality map in southeast Oregon. LOCALITY AND LITHOLOGY Approximately 30 stratigraphic feet of diatomite is exposed just above the high-tide mark on the beach adjacent to Bandon State Park on Beach Loop Road three miles south of Bandon, Oregon (fig. 1). Steps down to the beach have been carved in the soft diatomite at the locality. The base of the unit which dips gently to the north is not exposed, and the top is unconformably overlain by sand dunes. The rock is cream colored on weathered exposures and buff where it is fresh. Considerable limonite staining occurs along fractures running through TABLE 1. Percentage occurrence of genera from three localities: University of Oregon Museum of Natural History, Localities 2569, 2570, 2571. Genera Percent Coscinodiscus spp. Stephanopyxis sp. Lithodesmium sp. Arachnoidiscus sp. Stictodiscus sp. Actinocyclus spp. Actinoptychus spp. l TASTES TRIS VoL. XXXVII] ORR, EHLEN, AND ZAITZEFF: OREGON DIATOMS 491 ACTINOPTYCHUS SPLENDENS var. INCISA STEPHANOPYXIS APPENDICULATA ARACHNOIDISCUS DECORUS ACTINOPTYCHUS BISMARKII LITHODESMIUM CORNIGERUM var. ? STICTODISCUS BURYANUS ACTINOCYCLUS CUBITUS RECENT PLEISTOCENE @ @| coscinopiscus ASTEROMPHALUS @ @ @ @ @| CoOSscINODIScUS ANTiQUUS eee @ PLIOCENE @e@e0e @ eee @ LATE MIOCENE @eeeeeed @ @ @ @ @ OO @ | ACTINOPTYCHUS SPLENDENS vor. SOLIS! @@@ @ @ @ @/| COSCINODISCUS PACIFICUS MIDDLE MIOCENE LOWER MIOCENE OLIGOCENE EOCENE PALEOCENE @e@e@e0e00608086086 @ @ @ @/| COSCINODISCUS MARGINATUS ©0686 © OS @ O @ @/ CoSsciNnoviscus oBscuRUS @e e086 @ OO OO @ @ @|ACTINOPTYCHUS SENARIUS LATE CRETACEOUS Ficure 2. Published age distribution of diatom species from Bandon. the fine-grained massive diatomite. Using estimates of acid-treated residues and thin sections, the rock was found to exceed 75 percent organically precipi- tated silica. Of this siliceous fraction, more than 95 percent is in the form of diatom frustules and the remaining five percent is skeletal material from radio- laria and silicoflagellates. The remaining 25 percent nonsiliceous portion of the rock is very fine terriginous material which occurs in thin layers and imparts the only visible stratification to the rock. Three samples were collected from evenly spaced stratigraphic intervals and processed separately. These samples were later found to be very homogeneous with regard to the diatom flora. 492 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. @.ee8% Feeeses ty of Oregon Museum of Natural History. Hypotype number 27466. Diameter 0.115 mm. Univ. of Oregon Mus. Loc. 2569. Ficure 4. Coscinodiscus marginatus Ehrenberg. University of Oregon Museum of Natural Ficure 3. Coscinodiscus marginatus Ehrenberg. Univers VoL. XXXVII] ORR, EHLEN, AND ZAITZEFF: OREGON DIATOMS 493 FLORAL COMPOSITION AND ENVIRONMENT Species of Coscinodiscus (table 1) clearly dominate the Bandon flora which is characterized by the abundance of C. marginatus, C. asteromphalus, and C. pacificus. Of the seven species in this flora that range into the Recent, C. marginatus Ehrenberg and C. asteromphalus Ehrenberg are ubiquitous in modern oceanic plankton at various latitudes (Hendey 1937, 1964; Lohman 1941). Two other species, C. obscurus Schmidt and Actinoptychus senarius Ehrenberg, are most common in temperate seas, but are also widely distributed (Hendey, 1957; Cupp, 1943). Almost all of the modern species in the present flora are either neritic pelagics or benthonics. The excellent preservation of the specimens at Bandon and the lack of evidence of diagenetic alteration indicate that the fossil assemblage fairly represents the original flora. Because many of the forms are geographically wide ranging, precision is difficult, but the abun- dance of benthonics and temperate species and the absence of dominantly tropical or polar species imply a shallow coastal environment not markedly different from the adjacent ocean. GEOLOGIC AGE Seven of the species in this flora are stratigraphically long-ranging forms which appear in sediments from the Cretaceous or Tertiary to the Recent and are thus of limited biostratigraphic use (fig. 2). The remaining species of diatoms in the Bandon flora are unreported from rocks older or younger than the late Tertiary. Four of the Bandon species, Stephanopyxis appendiculata Ehrenberg, Arachnoidiscus decorus Brown, Actinocyclus cubitus Hanna and Grant, and Actinoptychus bismarkii Schmidt, are reported by Wornardt (1967) as strati- graphically restricted to the late Miocene in California. The presence of these four species in the flora by themselves would clearly imply a late Miocene age. How- ever, also present in the flora are fragments and small valves of a diatom resem- < History. Hypotype number 27467. Diameter 0.135 mm. Univ. of Oregon Mus. Loc. 2571. Ficure 5. Coscinodiscus asteromphalus Ehrenberg. University of Oregon Museum of Natural History. Hypotype number 27468. Diameter 0.184 mm. Univ. of Oregon Mus. Loc. 2569. Ficure 6. Coscinodiscus asteromphalus Ehrenberg. University of Oregon Museum of Natural History. Hypotype number 27469. Diameter 9.205 mm. Univ. of Oregon Mus. Loc. 2569. Ficure 7. Coscinodiscus obscurus Schmidt. University of Oregon Museum of Natural History. Hypotype number 27470. Diameter 0.112 mm. Univ. of Oregon Mus. Loc. 2570. Ficure 8. Coscinodiscus pacificus Rattray. University of Oregon Museum of Natural History. Hypotype number 27471. Diameter 0.145 mm. Univ. of Oregon Mus. Loc. 2569. Ficure 9. Coscinodiscus pacificus Rattray. University of Oregon Museum of Natural History. Hypotype number 27472. Diameter 0.128 mm. Univ. of Oregon Mus. Loc. 2571. 494 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 10. Coscinodiscus antiquus Grunow. University of Oregon Museum of Natural History. Hypotype number 27473. Univ. of Oregon Mus. Loc. 2569. Diameter 0.052 mm. Ficure 11. Coscinodiscus antiquus Grunow. University of Oregon Museum of Natural History. Hypotype number 27474. Univ. of Oregon Mus. Loc. 2571. Diameter 0.065 mm. Ficure 12. Stephanopyxis appendiculata Ehrenberg. University of Oregon Museum of VoL. XXXVIT] ORR, EHLEN, AND ZAITZEFF: OREGON DIATOMS 495 bling Lithodesmium cornigerum Brun. This latter species is regarded by authors (Wornardt, 1967; Hanna, 1930; Lohman, 1938) as a guide fossil to the Pliocene. Contamination of the Bandon flora by the four late Miocene species seems un- likely and the identification of L. cornigerium Brun is discussed under system- atics. Because all of the species are regarded as representative of the original flora and not contaminants, the evidence suggests a late Miocene (Delmontian) age for the Bandon flora. The co-occurence of the four late Miocene species and the Pliocene L. cornigertum Brun in the Bandon flora may extend the range of this latter species into the upper Miocene. The silicoflagellate flora extracted from these sediments (Orr and Zaitzeff, 1970) corroborates this diatom correlation. SYSTEMATICS Because of the limited number of most species in the Bandon flora, this section on systematics is not intended to be a detailed examination of the species. The reader is directed to Wornardt (1967) for a more comprehensive taxonomic treatment of the species identified here. < Natural History. Hypotype number 27475. Univ. of Oregon Mus. Loc. 2569. Diameter 0.105 mm. Ficure 13. Lithodesmium cornigerum Brun. var.? University of Oregon Museum of Natural History. Hypotype number 27478. Univ. of Oregon Mus. Loc. 2570. Diameter 0.079 mm. Ficure 14. Stephanopyxis appendiculata Ehrenberg. University of Oregon Museum of Natural History. Hypotype number 27476. Univ. of Oregon Mus. Loc. 2569. Diameter 0.112 mm. Ficure 15. Stephanopyxis appendiculata Ehrenberg. University of Oregon Museum of Natural History. Hypotype number 27477. Univ. of Oregon Mus. Loc. 2570. Diameter 0.135 mm. Ficure 16. Lithodesmium cornigerum Brun. var.? University of Oregon Museum of Natural History. Hypotype number 27479. Univ. of Oregon Mus. Loc. 2571. Diameter 0.111 mm. Ficure 17. Lithodesmium cornigerum Brun. var.? University of Oregon Museum of Natural History. Hypotype number 27480. Univ. of Oregon Mus. Loc. 2569. Diameter 0.085 mm. Ficure 18. Stictodiscus buryanus Grunow. University of Oregon Museum of Natural History. Hypotype number 27482. Univ. of Oregon Mus. Loc. 2569. Diameter 0.144 mm. Ficure 19. Arachnoidiscus decorus Brown. University of Oregon Museum of Natural History. Hypotype number 27481. Univ. of Oregon Mus. Loc. 2569. Diameter 0.165 mm. Ficure 20. Stictodiscus buryanus Grunow. University of Oregon Museum of Natural History. Hypotype number 27482. Univ. of Oregon Mus. Loc. 2569. (Same specimen as fig. 18 at higher magnification.) 496 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. FiGuRE 21. Actinocyclus species. University of Oregon Museum of Natural History. Hypotype number 27483. Univ. of Oregon Mus. Loc. 2569. Diameter 0.085 mm. FicuRE 22. Actinoptychus splendens var. incisa (Grunow). University of Oregon Museum of Natural History. Hypotype number 27485. Univ. of Oregon Mus. Loc. 2570. Diameter 0.095 mm. VoL. XXXVIT] ORR, EHLEN, AND ZAITZEFF: OREGON DIATOMS 497 PLANT KINGDOM Division CHRYSOPHYCOPHYTA Class BACILLARIOPHYCEAE Fritsch, 1935 Order BaciLiariates Schiitt, 1896 Family CoscinopiscacEaE Kiitzing, 1844 Genus Stephanopyxis Ehrenberg, 1845 Stephanopyxis appendiculata Ehrenberg. (Figures 12, 14, 15.) Stephanopyxis appendiculata EHRENBERG, 1854, pl. 18, fig. 4; WoRNARDT, 1967, p. 17, figs. 1213: Genus Coscinodiscus Ehrenberg, 1838 Coscinodiscus antiquus (Grunow) Rattray. (Figures 10, 11.) Coscinodiscus antiquus (Grunow) Rattray, 1889, p. 461; WorNarpt, 1967, p. 20, fig. 23. Coscinodiscus asteromphalus Ehrenberg. (Figures 5, 6.) Coscinodiscus asteromphalus EHRENBERG, 1854, pl. 18, fig. 45; WorNARDT, 1967, p. 20, figs. 14-18. Ficure 23. Actinocyclus cubitus Hanna and Grant. University of Oregon Museum of Natural History. Hypotype number 27486. Univ. of Oregon Mus. Loc. 2569. Diameter 0.075 mm. Ficure 24. Actinoptychus senarius Ehrenberg. University of Oregon Museum of Natural History. Hypotype number 27487. Univ. of Oregon Mus. Loc. 2569. Diameter 0.075 mm. Ficure 25. Actinocyclus species. University of Oregon Museum of Natural History. Hypotype number 27484. Univ. of Oregon Mus. Loc. 2570. Diameter 0.065 mm. FicurE 26. Actinoptychus bismarkii Schmidt. University of Oregon Museum of Natural History. Hypotype number 27488. Univ. of Oregon Mus. Loc. 2571. Diameter 0.075 mm. FicuRE 27. Actinoptychus splendens var. solisit Hanna and Grant. University of Oregon Museum of Natural History. Hypotype number 27489. Univ. of Oregon Mus. Loc. 2569. Diameter 0.090 mm. FicurE 28. Actinoptychus splendens var. solisi Hanna and Grant. University of Oregon Museum of Natural History. Hypotype number 27490. Univ. of Oregon Mus. Loc. 2569. ' Diameter 0.115 mm. Ficure 29. Actinoptychus splendens var. solisi Hanna and Grant. University of Oregon Museum of Natural History. Hypotype number 27491. Univ. of Oregon Mus. Loc. 2570. Diameter 0.155 mm. Ficure 30. Actinoptychus splendens var. solisi Hanna and Grant. University of Oregon Museum of Natural History. Hypotype number 27492. Univ. of Oregon Mus. Loc. 2570. Diameter 0.125 mm. 498 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Coscinodiscus marginatus Ehrenberg. (Figures 3, 4.) Coscinodiscus marginatus EHRENBERG, 1854, pl. 18, fig. 44; WORNARDT, 1967, p. 26, figs. 27, 28. Coscinodiscus obscurus Schmidt. (Figure 7.) Coscinodiscus obscurus SCHMIDT, 1878, pl. 61, fig. 16; WORNARDT, 1967, p. 27, fig. 32. Coscinodiscus pacificus Rattray. (Figures 8, 9.) Coscinodiscus pacificus RATTRAY, 1890, p. 563; WorNaArpt, 1967, p. 30, figs. 36, 37. Genus Actinocyclus Ehrenberg, 1838 Actinocyclus cubitus Hanna and Grant. (Figure 23.) Actinocyclus cubitus HANNA AND GRANT, 1926, p. 118, pl. 11, fig. 3. Wornarpt, 1967, p. 33, figs. 47, 48. Actinocyclus species. (Figures 21, 25.) Specimens of this species of Actinocyclus in the Bandon flora closely resemble a diatom figured and identified as A. ehrenbergii Ralfs by Lohman (1938) plate 22, figure 1. Lohman (1941) places this same reference in synonymy with A. octonarius Ehrenberg as does Wornardt (1967). Specimens figured and identified as A. octonarius in these latter two references do not, however, re- semble the Bandon specimens. Family ACTINODISCACEAE Schutt, 1896 Genus Stictodiscus Greville, 1861 Stictodiscus buryanus Grunow. (Figures 18, 20.) Stictodiscus buryanus Grunow, Hustept, in Schmidt, 1940, pl. 441, fig. 9, pl. 442, fig. 1; Wornarpt, 1967, p. 38, fig. 52. Genus Arachnoidiscus Bailey, from Ehrenberg, 1849 Arachnoidiscus decorus Brown. (Figure 19.) Arachnoidiscus decorus Brown, WoRNARDT, 1967, p. 40, fig. 53. Genus Actinoptychus Ehrenberg, 1943 Actinoptychus bismarkii Schmidt. (Figure 26.) Actinoptychus bismarkii ScHMipT, 1886, pl. 91, fig. 4; WorNarpT, 1967, p. 42, fig. 65. Actinoptychus senarius Ehrenberg. (Figure 24.) Actinoptychus senarius EHRENBERG, 1838, p. 172, pl. 21, fig. 6; WorNaARDT, 1967, p. 44, fig. 67. VoL. XXXVIT] ORR, EHLEN, AND ZAITZEFF: OREGON DIATOMS 499 Actinoptychus splendens var. incisa (Grunow). (Figure 22.) Actinoptychus incisa GRUNOW, in Schmidt, 1890, pl. 154, figs. 2, 3. Actinoptychus splendens var. incisa (GRUNOW) ; WorNarDT, 1967, p. 48, figs. 68-71, 73. Actinoptychus splendens var. solisi (Hanna and Grant). (Figures 27, 28, 29, 30.) Actinoptychus solisi HANNA AND GRANT, 1926, p. 123, pl. 12, figs. 1-3. Actinoptychus splendens var. solist (Hanna and Grant), WorRNARDT; WoRNARDT, 1967, p. 48, figs. 74-77. Family BippuLPHIACEAE Kiitzing, 1844 Genus Lithodesmium Ehrenberg, 1840 Lithodesmium cornigerum Brun var.? (Figures 13, 16, 17.) Lithodesmium cornigerum Brun, Hanna, 1930, p. 189, pl. 14, figs. 9, 10; Wornarpt, 1967, p. 67, fig. 131. Specimens identified as L. cornigerum var.? from the Bandon flora differ from Pliocene forms reported from California (Hanna, 1930; Wornardt, 1967) in that the valves of most of those in the present flora frequently have more club- shaped processes (figs. 13, 17). Typical “propeller” shaped forms mentioned by Wornardt (1967) are nonetheless present (fig. 16). The transition between club and propeller shape appears to be complete within the Bandon flora and the smaller specimens tend to be club shaped. The processes of typical “propeller” shaped forms in the present flora (fig. 16) are shorter than those on Pliocene specimens illustrated by Wornardt (1967) and Hanna (1930). BIBLIOGRAPHY Cupp, EASTER 1943. Marine plankton diatoms of the west coast of North America. Bulletin of Scripps Institute of Oceanography of the University of California, La Jolla, California, vol. 5, no. 1, pp. 1-235. EHRENBERG, C. G. 1838. Die Infusionsthierchen als vollkommene Organismen, Ein Blick in das Tiefere Organische Leben der Natur: Leipzig, verlag von Leopold Voss, Text und Atlas. Text, pp. I-XVIII, 1-547; Atlas von Vier und Sechzig Kupfertafeln, tables I-LXIV. 1854. Mikrogeologie, Das Erden und Felsen Schaffende Wirken des Unsichtbar Kleinen Selbststandigen Lebens auf der Erde. Leipzig, verlag von L. Voss, in two parts. Text, pp. I-X XVIII, 1-374; Atlas, pp. 1-31, tables I-XXXX. FritscH, F. E. 1935. The structure and reproduction of the algae, vol. I, class IV. Bacillariophyceae, pp. 564-651. New York, The Macmillan Co.; Cambridge, England, The Uni- versity Press. Hanna, G Datras 1930. Observations on Lithodesmium cornigerum Brun. Journal of Paleontology, vol. 4, no. 2, pp. 189-191, pl. 14, figs. 9-10, June. 500 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Hanna, G Darras, AND WitL1AM M. GRANT 1926. Miocene marine diatoms from Maria Madre Island, Mexico, in 1924. In Expe- dition to the Revillagigedo Islands, Mexico, in 1925. Proceedings of the Cali- fornia Academy of Sciences, 4th ser., vol. 15, no. 2, pp. 115-193, pls. 11-21, April 16. HENDEY, INGRAM N. 1937. The plankton diatoms of the southern seas. Discovery Reports, vol. XVI, pp. 151-364, pls. VI-XIII, Cambridge. 1957. Marine diatoms from some west African ports. Journal of the Royal Micro- scopical Society, vol. 77, pp. 28-84, pls. I-VI. 1964. An introductory account of the smaller algae of British coastal waters. Part V, Bacillariophyceae (Diatoms). Fishery investigations, ser. IV, London. pp. I- XXII, 1-317, pls. 1-45, text-figs. 1-9. KUTZING, FRIDERICUS TRAUGOTT 1844. Die kieselechaligen Bacillarien oder Diatomeen. Nordhausen, Kohne W., intro- duction pp. 1-4, pp. 1-152, tables 1-30. LOHMAN, KENNETH E. 1938. Pliocene diatoms from the Kettleman Hills, California. United States Geological Survey, Professional Paper 189-C, pp. 81-102, pls. 20-23. 1941. Diatomaceae, Part 3. Jn Bradley and others, Geology and biology of North At- lantic deep-sea cores between Newfoundland and Ireland. United States Geo- logical Survey, Professional Paper 196-B, pp. 55-87, pls. 12-17, pp. XVII-XX. Orr, W.N., AND J. B. ZAIrzerr. 1970. Miocene Silicoflagellates from Southeast Oregon. Northwest Science, vol. XLIV, no. I. RATTRAY, JOHN 1889. A revision of the genus Coscinodiscus Ehrenberg and some allied genera. Pro- ceedings of the Royal Society of Edinburgh, vol. XVI, pp. i-viii, 449-692, pls. I-III. Edinburgh, Neill and Company. 1890. A revision of the genus Actinocyclus Ehrenberg. Journal, Quekett Microscopical Club, ser. 2, vol. 4, pp. 137-212. pl. II. ScumipT, ADOLF 1874-1959. Atlas der Diatomaceen-Kunde (continued by Martin Schmidt, Friedrich Fricke, Heinrich Heiden, Otto Muller, Friedrich Hustedt). Heft 1-120 (1874— 1959), tables 1-480. Leipzig. Scuuwrt, F. 1896. Bacillariales (Diatomeae), 7 Engler, A., und Prantl, K., Die natiirlichen Pflan- zenfamilien nebst ihren Gattungen und wichtigeren Arten, insbesondere den Nutzpflanzen, unter Mitwirkung zahlreicher hervorragender Fachgelehrten, Teil I. Abteilung 1b, pp. 1-153. Leipzig. WorNARDT, WALTER 1967a. Miocene and Pliocene marine diatoms from California. California Academy of Sciences, Occasional Papers, no. 63, pp. 1-108, fig. 217. 1967b. Siliceous microfossils from the Bickmore Canyon diatomite and the Pancho Rico formation. Jn Guidebook, Pacific Section, A.A.P.G.-S.E.P.M. 1967 Gabilan Range, pp. 48-53, pl. 1-2. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XXXVII, No. 17, pp. 501-508; 2 figs. July 23, 1971 TWO NEW ATLANTIC SCORPIONFISHES' Ly William N. Eschmeyer California Academy of Sciences, San Francisco, California 94118 ABSTRACT: Two new species of scorpionfishes are described from the central Atlantic Ocean. Scorpaenodes insularis is known only from Saint Helena and Scorpaena ascensionis from Ascension Island. Descriptions and figures are given. INTRODUCTION The scorpionfishes of the Atlantic Ocean were the subject of recent studies by the author (Eschmeyer, 1965, 1969b). Examination of additional specimens in many museums has resulted in the discovery of two new species from isolated central South Atlantic islands. One, a species of the genus Scorpaena from Ascension Island (7°57’ S, 14°22’ W), was misidentified as Scorpaena scrofina in the Discovery reports (Norman, 1935). The second is a species of Scorpa- enodes from the island of Saint Helena (15°57’ S, 5°42’ W). The most recent general study of the fishes of Saint Helena and Ascension was by Cadenat and Marchal (1963); they treated a total of 124 species, 96 from Saint Helena and 78 from Ascension, of which 50 occurred at both islands. Methods follow those used earlier (Eschmeyer, 1969b). The specimens of both species were found in the collection of the British Museum of Natural History, and thanks are due the BMNH staff, especially Alwyne Wheeler and Peter J. P. Whitehead, for courtesies extended during a visit there and for loaning the specimens for further study. Katherine Smith, Pearl Sonoda, Lillian Dempster, and Terry Arambula, California Academy of Sciences, assisted in the study. Maurice Giles did the photography for the illustrations. The drawings were made by Katherine Smith. 1 Funds for travel to museums were provided by National Science Foundation Grant GB-15811. [501] [Proc. 4TH SER. CALIFORNIA ACADEMY OF SCIENCES 502 —— TPs + Ek eye" ‘adAjeied ay} UO AT]SOW paseq SUIMeBIp ‘pur[s] UOISUdISY “T'S “WU GL ¢—-9' PE ‘seworsUaIsD DUaDg 41095 Se ‘| TaN VoL. XXXVIT] ESCHMEYER: NEW SCORPIONFISHES 503 Scorpaena ascensionis Eschmeyer, new species. (Figure 1.) Scorpaena scrofina, Norman, 1935, p. 55 (not of Valenciennes in Cuvier and Valenciennes, 1833; 2 specimens from Ascension Island). Remarks. The name “‘scrofina” has been used for one of the species of Scorpaena occurring at Saint Helena and Ascension. Eschmeyer (1969b, p. 59) included these records under the species S. plumieri. Norman (1935, p. 56, table) evidently was confused over the type locality of S. scrofina, since he shows S. scrofina to be restricted to Saint Helena and Ascension. The type locality of Scorpaena scrofina Valenciennes in Cuvier and Valenciennes, 1833, is Brazil, and the original description is clearly of a specimen referable to S. plumieri, a western Atlantic and eastern Pacific species which also occurs at Saint Helena and Ascension. The specimen in the Museum National d’Histoire Naturelle (MNHN 6702), which according to Blanc and Hureau (1968, p. 17) is the type of S. scrofina, is not the type. Now in bottle MNHN 6702, with the locality “Brazil,” is a specimen of Scorpaena laevis, an eastern Atlantic species which does not agree in diagnostic color features with information in the original de- scription of S. scrofina. It is likely that a switch of specimens occurred. In any event, S. scrofina is a synonym of S. plumieri, which does occur at Saint Helena and Ascension. The only other species of scorpionfishes known from either Saint Helena or Ascension are Scorpaena mellissii Giinther (1868, p. 225, pl. 19) and Pontinus nigropunctatus (Giinther, 1860, p. 145). Synanceja horrida has also been reported from Saint Helena (Gunther, 1860, p. 145) and is represented by a dried specimen in the British Museum collection; I assume that the locality information for this Indo-Pacific species is probably in error. The new species is compared with the other species from Saint Helena and Ascension in the “Comparisons” section. Hototyee. BMNH 1935.5.2.33 (37.5 mm. S.L.) Ascension Island, Dis- covery Station 1, 16 November 1925, medium rectangular net, depth 16-27 meters. ParatyPe. BMNH 1935.5.2.34 (33.6 mm. S.L.) same data as holotype. DescripTION. A small species with the following distinguishing features: very shallow occipital pit, cycloid scales, about 60 vertical scale rows, chest and pectoral-fin base scaled, and two preorbital spines over maxillary. Dorsal fin with 12 spines and 9 soft rays (last double). Anal fin with 3 spines and 5 soft rays (last double). Pectoral fin with 20 rays, rays 3 through 4 or 5 branched dis- tally; pectoral fin reaching to over the first or second anal spine. Gill rakers including rudiments 14-15, 5 on upper arch, 9-10 on lower arch; all gill rakers fairly short, the upper 4 and most lower ones as rudiments. Preorbital bone with 2 spines over maxillary; anterior spine directed forward, posterior spine to rear. Suborbital ridge without spinous points. Supplemental preopercular spine pres- CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. imen). in spec In missing Scorpaenodes insularis, holotype, 54.7 mm. S.L., Saint Helena (much of caudal fi FIGURE 2. VoLt. XXXVIT] ESCHMEYER: NEW SCORPIONFISHES 505 ent, slightly dorsal to first preopercular spine. First preopercular spine short; second through fourth preopercular spines present, about one-half length of first; fifth preopercular spine very small and covered by a dermal flap. Other spines present typical for the genus Scorpaena (Eschmeyer, 1969b, fig. 1, and p. 54) and include the following: nasal, preocular (low), supraocular, postocular, tym- panic, parietal, nuchal, pterotic, sphenotic, upper and lower posttemporal, supra- cleithral, and cleithral. Most spines low, short, and frequently blunt. Vertebrae 24. Scales on sides cycloid or slightly emarginate; vertical scale rows about 60 (about 58-62 in available material, some scales rubbed off); lateral-line scales 26 + 2; chest, pectoral-fin base, postorbital area of head, opercle, and cheek with cycloid scales. Tentacles and skin flaps on head and body short; small supra- ocular tentacle less than about one-tenth orbit diameter; short flaps on some lateral-line scales and body scales; a few small flaps on eye, on preopercular spines, and on posterior preorbital spine. Measurements in millimeters are as follows (paratype in parentheses; holo- type with upper jaw slightly distended thereby affecting some measurements) : total length 37.5 (33.6); head length 17.9 (15.7); body depth 11.8 (12.0); orbit diameter 5.9 (5.0); snout length 4.0 (4.0); interorbital width 1.8 (1.6); jaw length 8.2 (8.0); predorsal fin length 15.5 (13.6); length third dorsal spine 6.6 (5.5); length first anal spine 3.3 (2.7), second 7.0 (5.9) and third 5.5 (5.4); length pectoral fin 11.1 (10.3); length pelvic fin 9.7 (8.6); length caudal fin OwACLO.1). Body shape and coloration as in figure 1. Axil of pectoral fin with no dis- tinctive color pattern, mostly pallid, a few brown spots. The holotype is a gravid female. It is likely that the species does not exceed about 50 mm. in total length. DISTRIBUTION. The species is known only from Ascension Island. CoMPARISONS. Scorpaenodes insularis, Pontinus nigropunctatus, Scorpaena mellissu, S. plumieri, and S. ascensionis constitute the scorpionfishes known from Saint Helena and Ascension. The first three are known only from Saint Helena. Scorpaena plumieri occurs at both islands, and S. ascensionis is known only from Ascension. Information on P. nigropunctatus, S. plumieri, and S. mel- lissii may be found in Eschmeyer (1969b). Scorpaenodes insularis can be sep- arated on the basis of 13 dorsal spines and no palatine teeth. Scorpaena plumieri is most easily characterized by the presence of large white spots on a black _ background in the axil of the pectoral fin. Pontinus nigropunctatus is a large species characterized by no branched pectoral rays. Scorpaena mellissii is easily distinguished on the basis of coloration, especially in the lack of bars on the body and the presence of scattered dark brown spots on the fins (see Gunther, 1868, pl. 19). Scorpaena ascensionis may be separated on the basis of coloration (fig. 1) and, unlike the other species above, it has no spinous points on the 506 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. suborbital ridge. Scorpaena ascensionis does not seem to be especially closely related to any other species of the genus Scorpaena. Name. The specific name “‘ascensionis” has been used several times in other families of fishes. It is unlikely that the present species of Scorpaena will occur elsewhere, with the possible exception of Saint Helena; so this geographical name is a meaningful one, and is used for the present species. Scorpaenodes insularis Eschmeyer, new species. (Figure 2.) Hototyree. BMNH 1969.3.10.9 (54.7 mm. S.L.) South Atlantic Ocean, Saint Helena Island, James Bay, wreck of Papanu, collected June 1968, presented by A. Loveridge. DeEscripTION. A species of Scorpaenodes characterized by 3 spines on the suborbital ridge, another spine below the main suborbital ridge, pectoral rays 18— 19, total gill rakers 17-18, normally (?) 10 soft dorsal rays, interorbital spines present, and no dark spot at end of spinous dorsal fin. Dorsal fin with 13 spines and 10 soft rays (last double). Anal fin with 3 spines and 5 soft rays (last double). Pectoral fin with 18 (left) or 19 (right) rays, rays 3 through 8 or 9 branched in available specimen. Gill rakers including rudiments 17-18, 6 on upper arch, 9 on ceratobranchial and 2-3 flat rudiments on hypobranchial. Preorbital bone with 2 lobes over maxillary, posterior one as a broad spine. Sub- orbital ridge with 3 spinous points, first on preorbital, second below posterior part of eye, and third at end of ridge. Supplemental preopercle spine as a lump on base of first preopercular spine; first through third preopercular spines present, fourth and fifth absent. Interorbital spines present; coronal (post- frontal) spines present on right side; upper posttemporal spines absent; nasal spines present; spine on extrascapular bone present. Other spines present typical for the genus Scorpaenodes (Eschmeyer, 1969a, p. 4; 1969b, pp. 85-86, fig. 1) and include the following: preocular, supraocular, postocular, tympanic, parietal, nuchal, sphenotic, lower posttemporal, opercular (2), supracleithral, and clei- thral. Vertebrae 24. Scales on sides slightly ctenoid; vertical scale rows about 47; lateral-line scales 23 + 2; chest, pectoral-fin base, lower portions of fins, and most of head scaled. Tentacles and skin flaps inconspicuous; short, simple tentacles associated with most head spines. Measurements in millimeters are as follows: total length 64.5, caudal dam- aged distally; head length 22.8; body depth 17.3; orbit diameter 6.7; snout length 5.3; interorbital width 2.5; jaw length 10.8; predorsal fin length 20.6; length fourth dorsal spine 7.0; length first anal spine 4.5, second 11.8, and third 8.5; length pectoral fin 16.0; length pelvic fin 15.7. Body shape and coloration as in figure 2. Body with brown patches and spots on a pale background. Head mostly dusky. Dorsal and pectoral fins faintly spotted with brown. VoL. XXXVI] ESCHMEVER: NEW SCORPIONFISHES 507 DISTRIBUTION. The species is known only from the island of Saint Helena. Comparisons. Scorpaenodes insularis is compared with other species from Saint Helena and Ascension in the account of Scorpaena ascensionis. Compari- son with eastern and western Atlantic species may be made by consulting Esch- meyer (1969b: pp. 86-91). Scorpaenodes insularis most resembles S. africanus in coloration, but S. insularis has 3 spines on the suborbital ridge while S. afri- canus has 2 (usually a stable feature). Also, S. insularis has a spine present under the main suborbital ridge while S. africanus does not. The soft dorsal ray count of 10% for S. insularis may be one more ray than normal; most species of Scorpaenodes normally have 9%, and some 8'%. Name. The specific name is based on the Latin adjective “insularis,” of an island, alluding to the occurrence of this species at the island of Saint Helena. LITERATURE CITED Branc, M., ano J. C. HuREAU 1968. Catalogue critique des types de piossons du Muséum National d’Histoire Naturelle (poissons a joues cuirassées). Publications diverses du Muséum d’Histoire Naturelle, no. 23, 71 pp. CADENAT, JEAN, AND E, MARCHAL 1963. Résultats des campagnes océanographiques de la Reine-Pokou aux iles Sainte- Hélene et Ascension. Poissons. Bulletin de l’Institut Francais d’Afrique Noire, vol. 25, ser. A, no. 4, pp. 1235-1315, 42 figs. Crark, R.S. 1913. Scottish National Antarctic Expedition—“Scotia” collection of fishes from St. Helena. Proceedings of the Royal Physical Society of Edinburgh, vol. 19, pp. 47-53. CUNNINGHAM, JOSEPH THOMAS 1910. On the marine fishes and invertebrates of St. Helena. Proceedings of the Zoo- logical Society of London, 1910, pp. 86-131, pls. 1-4, figs. 1-4. Cuvier, GEORGES, AND ACHILLE VALENCIENNES 1833. Histoire naturelle des poissons. F. G. Levrault, Paris, vol. 9, 512 pp. EsSCHMEYER, WILLIAM N. 1965. Western Atlantic scorpionfishes of the genus Scorpaena, including four new species. Bulletin of Marine Science, vol. 15, no. 1, pp. 84-164, 12 figs. 1969a. A new scorpionfish of the genus Scorpaenodes and S. muciparus (Alcock) from the Indian Ocean, with comments on the limits of the genus. Occasional Papers of the California Academy of Sciences, no. 76, 11 pp., 1 fig. 1969b. A systematic review of the scorpionfishes of the Atlantic Ocean (Pisces: Scor- paenidae). Occasional Papers of the California Academy of Sciences, no. 79, 130 pp., 13 figs. GUNTHER, ALBERT 1860. Catalogue of the acanthopterygian fishes in the collection of the British Museum, vol. 2, xxi + 548 pp. 1868. Report on a collection of fishes made at St. Helena by J. C. Melliss, Esq. Pro- ceedings of the Zoological Society of London, 1868, no. 15, pp. 225-228, pls. 18-19. 508 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. MELLIss, JOHN CHARLES 1875. Class V—Pisces. (Jn) Sancta Helena: a physical, historical, and topographical description of the island, including its geology, fauna, flora, and meteorology. L. Reeve and Co., London, Pisces on pp. 100-113, 3 pls. NorMan, JOHN R. 1935. Coast fishes. Part I. The South Atlantic. Discovery Reports, vol. 12, pp. 1-58. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XXXVITI, No. 18, pp. 509-513; 3 figs. July 23; 1971 DARWINNEON CRYPTICUS, A NEW GENUS AND SPECIES OF JUMPING SPIDER FROM THE GALAPAGOS ISLANDS (ARANEAE: SALTICIDAE)! By Bruce Cutler? Department of Entomology, Fisheries and Wildlife University of Minnesota St. Paul, Minnesota 55101 ABSTRACT: Darwinneon crypticus, new genus and species, is described from three female and three immature specimens. All were taken on the south side of Santa Cruz Island in the Galapagos Islands. They are very small jumping spiders related to Neon and Neonella, particularly to the species Neon nigriceps Bryant. Recent expeditions to the Galapagos Islands have uncovered a new genus and species of salticid spider from Santa Cruz Island (Indefatigable). The spider is described below, and its taxonomic placement is discussed. Darwinneon Cutler, new genus Very small spiders closely related to Neon and Neonella. Carapace longer than wide, truncated anteriorly, rounded posteriorly, hardly rounded at sides, height less than two-thirds width. As viewed laterally, anterior and posterior steeply sloping from just behind row III eyes. Diameter of anterior median 1 Paper no. 7085, Scientific Journal Series, Minnesota Agricultural Experiment Station, St. Paul, Minnesota 55101. 2I wish to thank Mr. Vincent Roth of the Southwestern Research Station of the American Museum of Natural History, Portal, Arizona, for bringing these spiders to my attention; Mr. Terry Dillon of the Department of French and Italian, University of Minnesota, for translating the labels on the Tervuren specimens; and my wife, Lucy E. Cutler, for assistance in the preparation of this paper. [509] Marine Biological Laboratory LIBRARY AUG3 1971 WOODS HOLE, MASS. 510 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser. pn es Z Z d 3 Ficures 1-3. Darwinneon crypticus Cutler, new genus and species. 1. Lateral view of holotype. 2. External view of epigynum of paratype in California Academy of Sciences. 3. Internal view of epigynum of paratype in California Academy of Sciences. eyes very large for the size of the spider, less than one-fourth the length of the prosoma. Eyerow I recurved. Row III eyes face laterally and smaller than anterior lateral eyes. Eyefield wider than long, as wide behind as in front. Median groove of carapace nearly obsolete. Clypeus very narrow. Abdomen suboval, dorsum without scutum, integument slightly iridescent and _ hairy. Retromargin of chelicera with a small single tooth. Legs of moderate length, formula 4312, first pair slightly heavier. Trochanteral limuli rounded. Leg spination complex, but with two pair of prominent spines on the metatarsi of the first three legs. Spination reduced on posterior legs, but prominent spines still present. Epigynum consisting of two anterior atria without an obvious septum separating them. The seminal receptacles appear to be two semicircular tubes mostly posterior to the atria. Vor. XXXVII] CUTLER: A NEW GALAPAGOS JUMPING SPIDER 511 Darwinneon may be separated from related genera as follows: from Chalcoscirtus by the presence of a retromarginal cheliceral tooth, possession of posterior leg spines, smaller size, and different epigynal structure; from Neon by usual smaller size, different body shape viewed laterally, extreme size of anterior median eyes, possession of strong posterior leg spines, lack of a median epigynal septum, and relative larger size of retromarginal cheliceral tooth; from Neonella by lack of usual Neonella color pattern on abdomen, different structure of internal ducts and receptacles of epigynum, possession of posterior leg spines, and more rounded lateral profile of carapace; from Semiopyla by presence of retromarginal cheliceral tooth, and possession of two anterior epigynal atria instead of one. Darwinneon crypticus Cutler, new species. Dracnosis. The characters of the species are the same as those of the genus. All types are mature females. DESCRIPTION OF HOLOTYPE. Measurements are in millimeters. Total length 1.72, prosoma 0.83 long, 0.65 wide. Height of carapace at eyerow III 0.40. Clypeus 0.02 high. Diameter of anterior median eyes 0.20, of anterior lateral eyes 0.15, of row II eyes 0.02, of row III eyes 0.11. Distance of eyerow II from eyerow I 0.11, of eyerow II from eyerow III 0.13. Eyefield length 0.39. Sternum length 0.32, width 0.24. The major leg spines occur as follows: metatarsi I-III have two pair of ventral spines, metatarsus IV has three spines at the distal end; tibia I and II have two pair of ventral spines, tibia III and IV have two spines, one pair midway along the length on the prolateral and retrolateral sides; there are no femoral spines. Leg lengths are: I II Ill IV Femur 0.39 Orsi 0.42 0.48 Patella 0.20 0.21 0.18 Oil Tibia 0.24 0.22 0.27 0.36 Metatarsus 0.18 0.18 0.21 0.34 Tarsus 0.21 0.18 0.21 0.20 Total 122 1.10 1.29 1.59 Opisthosoma 0.88 long, 0.60 wide. Color of carapace brown, ocular quadrangle brownish black, lateral margin with thin black stripe and pale brown stripe just above, middorsally a white stripe from the posterior margin of carapace to just forward of row III eyes, scattered white hairs in ocular quadrangle. Chelicerae as viewed from front, brown proximally with gray tips and gray along the medial edge. Endites and labium pale brown with gray tips. Sternum brown. The pedipalps are slightly swollen and strikingly colored; tarsus, femur, and trochanter black; tibia and patella milk white. All legs similarly 512 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser. colored; tarsi light grayish brown; metatarsi light grayish brown on distal half, dark grayish brown on proximal half; tibia dark grayish brown on distal three-fourths, proximal fourth light grayish brown; patella as in tibia but proportions half and half; femora all dark grayish brown with distal ventral yellow spot; all dorsal midlines of all specimens are pale yellow. Dorsum of opisthosoma gray with slight iridescence, cardiac stripe of thick white hairs on anterior half, scattered small white spots forming faint chevron pattern posteriorly. Many long hairs on dorsum and along sides. Venter grayish brown, no iridescence or pattern, few hairs. Holotype in the collection of the California Academy of Sciences. DESCRIPTION OF OTHER SPECIMENS. Paratype in the California Academy of Sciences, total length 1.67; paratype in Royal Museum of Central Africa, Tervuren, Belgium, total length 1.75. The immature specimens in the California Academy of Sciences are 1.70 and 1.62 in total length, and are probably penultimates, the Tervuren specimen is smaller, 1.45 in total length. All immatures look alike. They are very pale brown, have a flatter lateral carapace profile compared to the adults, and have the legs and palpi annulate brown and white. DIsTRIBUTION. All specimens are from the south central side of Santa Cruz Island in the Galapagos Archipelago. The ecological amplitude of the species appears great as one specimen was taken from Scalesia, a genus of tree- size composites, while two others were taken from humid detritus at the bottom of a crevasse. Hototyrr. Galapagos Archipelago, Santa Cruz, Darwin Research Station, 12 February 1964, Cavagnaro and Schuster. Paratype in the California Academy of Sciences collection: Galapagos Archipelago, Isla Santa Cruz, Bella Vista Trail, 11 February 1964, on Scalesia, R. O. Schuster. Paratype in Tervuren: Galdpagos: Ile de Santa Cruz, A 800 m de l’embarcadére de la St. Darwin; Au fond d’une crevasse, profunde de 10 m;-dans eboulis de pierres avec terre et détritus humides -XII-1964 J. et N. Leleup. The two immatures in the California Academy of Sciences were taken with the holotype. The immature in the Tervuren Museum was taken with the paratype. DERIVATION OF NAME. Darwinneon is a composite name honoring Charles Darwin with Neon, the name of a closely related genus. The specific name refers to the hidden habits of this species. Discussion. The only species to which D. crypticus shows a close rela- tionship is known as Neon nigriceps Bryant. This species is found in Cuba and the Bahama Islands. The major differences are the larger size of WN. nigriceps, the different lateral profile of the carapace, and the structure of the epigynum. There are certain structural similarities which cannot be ignored: both species have a similar retromarginal cheliceral tooth; both species have similar leg spination; and the internal genitalia of D. crypticus could be Vor. XXXVII] CUTLER: A NEW GALAPAGOS JUMPING SPIDER 513 derived from N. nigriceps by shortening of the seminal receptacles; of course the converse may also be true. However, NV. nigriceps appears to have a faint septum separating the atria, and has a distinct posterior epigynal notch. Neon nigriceps, as Gertsch and Ivie (1955) have indicated, does not belong in the genus Neon. It shows certain affinities to Neonella and to Darwinneon. The question of the placement of Neon nigriceps will have to await further study, and a new genus may have to be erected for it. LITERATURE CITED GertscH, W. J., AND W. IviIEe 1955. The spider genus Neon in North America. American Museum Novitates, no. 1743, pp. 1-17. oe PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XXXVIT, No. 19, pp. 515-527; 4 figs.; 3 tables. July, 23; 1971 THREE NEW SPECIES OF SCORPIONFISHES (FAMILY SCORPAENIDAE) FROM EASTER ISLAND LS William N. Eschmeyer California Academy of Sciences, San Francisco, California and Gerald R. Allen University of Hawaii and Bernice P. Bishop Museum, Honolulu, Hawaii ABSTRACT: Three new species of the fish family Scorpaenidae, Scorpaena orgila, S. pascuensis, and Scorpaenodes englerti, are described from Easter Island in the south- eastern Pacific Ocean. These species are known only from Easter Island, and are the only scorpaenids occurring there. INTRODUCTION Easter Island (Rapa Nui or Isla de Pascua) was discovered by the Dutch admiral Jacob Roggeveen on Easter Sunday in 1722. Since that time the world has marveled at the mysterious stone giants which rise above the island’s barren slopes. Although this isolated outpost of Polynesia has been the focus of much archeological study, only a token amount of natural history research has taken place. The Canadian Medical Expedition to Easter Island in 1964-65 made the first comprehensive biological collections. Prior to the Canadian collections, only 40 species of fishes had been recorded from Easter Island (Kendall and Radcliffe, 1912; Regan, 1913; Fuentes, 1914; Rendahl, 1921; Wilhelm and Hulot, 1957; de Buen, 1963); no scorpaenids had been recorded from the island. The Canadians collected about 70 species, many of them new. An earlier col- lection was made at Easter Island by Ramsey Parks and the crew of the ketch Chiriqui in 1958. John E. Randall and the junior author, using SCUBA gear [515] WOODS HOLE, MASS. 516 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. to depths of 40 meters, collected fishes at Easter Island for one month in 1969, adding additional new species. Easter Island is of volcanic origin and is characterized by an extremely rugged and rocky coastline. In many places the cliffs rise almost vertically from the edge of the sea. There is only one beach of any consequence, that of Anakena, which is a mere 200 meters in length. Occasional surge pools are encountered, with several in the vicinity of Hanga Roa, the only village on the island. The typical underwater habitat is that of rocky boulders usually covered with brown algae. Coral growth is sparse, but two species, representing the genera Porites and Pocillopora, are commonly observed in some localities. Strangely enough there is often abundant coral growth (Porites) in the shallow surge pools, sometimes covering more than 25 percent of the bottom. The abundance of rocky substrate provides an ideal habitat for scorpionfishes. The fish fauna at Easter Island is very restricted and shows a relationship to both the eastern Pacific and Indo-west-Pacific faunas. As one might expect, the high degree of isolation results in a correspondingly high rate of endemism. Preliminary investigations reveal that the rate for fishes may be as high as 40 percent. The three species of scorpionfishes occurring at Easter Island are endemics. ACKNOWLEDGMENTS Ian E. Efford, University of British Columbia, provided the scorpionfishes which he and Jack A. Mathias collected during the Canadian Medical Expedi- tion. John E. Randall, Bernice P. Bishop Museum, made available specimens collected by him and the junior author in 1969. Specimens collected by the ketch Chiriqui in 1958 were sent to us by Robert J. Lavenberg, Los Angeles County Museum of Natural History. Part of the junior author’s expenses for a trip to San Francisco to work on Easter Island scorpionfishes was paid by the California Academy of Sciences. The junior author acknowledges the financial assistance of the National Geographic Society for his collecting trip to Easter Island. The senior author received financial support from National Science Foundation grant GB-15811, which permitted him to examine scorpionfishes, particularly types, in most major museums. Special thanks are due Dr. Randall for permission to reproduce his photographs, for providing color slides of speci- mens, and for his comments on the manuscript. We thank Lillian Dempster for her help with the manuscript. The drawings of head spination were made by Katherine Smith. Pearl Sonoda and Terry Arambula aided in curatorial ways. METHODS Measurements, counts, and terminology of head spines are as used by Esch- meyer (1969). Abbreviations are as follows: BC—University of British Colum- bia; BPBM—Bernice P. Bishop Museum; CAS—California Academy of Vor. XXXVII] ESCHMEYVER & ALLEN: EASTER ISLAND SCORPIONFISHES 517 TABLE 1. Counts and measurements for type specimens of Scorpaena orgila from Easter Island. (Measurements are in millimeters ; numbers in parentheses are percentages of standard length.) CAS BPBM CAS CAS USNM BPBM BPBM 24809 6772 24810 24810 205209 6771 6776 Standard length 248 78.7 109 60.0 212 67.4 48.3 Dorsal fin rays ieeo ior gto 861049 6p =eG igo aero Anal fin rays 3+5 345 3+5 345 3+5 3+5 3+5 Pectoral fin rays 174+17) 17417) «17417 17417) 17417) « 17417) 17417 Head length 111 (45) 33.4 (42) 46.4 (43) 25.1 (42) 100(47) 29.0(43) 21.0°(43) Body depth 82/6183) 925.2) (32) 341 G1)! 19339(32)) 74:2) GS) 1Okn (29) 172k) Orbit diameter 20508). 9-727. (10) 10:1 (09). 96.5.(11) 20.7 (10), Fale) 5.310) Snout length S82 (IS) OSGI) WEED) Wel) ABO CD) esi (lz) Se (2) Interorbital width 13.7(06) 3.8(05) 5.6(05) 2.8(05) 12.1 (06) 3.0(04) 2.7 (06) Jaw length 345 (22) ICH (AW) 228 (@iD) IS (aD) 237 (23) 1843 (20) eK (BB) Predorsal-fin length 95.6 (39) 30.3 (38) 39.5 (36) 21.8 (36) 86.1 (41) 24.7 (37) 19.4 (40) Pectoral fin length 68.8 (28) 24.1 (31) 32.7 (30) 16.7 (28) 61.7 (29) 20.0(30) 14.5 (30) Pelvic fin length 62.7 (25) 21.8 (28) 30.2 (28) 16.5 (28) 56.1 (26) 19.2 (28) 14.1 (29) Caudal fin length 59.5 (24) 23.4 (30) 32.5 (30) 17.4 (29) 57.6(27) 19.5 (29) 14.2 (29) BPBM CAS BPBM LACM BC BC BC 6777 24811 6774 31198-1 65-438 65-439 65-457 Standard length 202 76.5 Wi, 214 113 194 145 Dorsal fin rays WesOm eO 19-6 12 ozo” uveson = Miz=s0 Anal fin rays ee 315 315 gees cS 3.15 3-45 Pectoral fin rays 7-07 0 174-170 17-17) 17417) 174-17) 16+ 17-17 Head length 92.1 (43) 32.4 (42) 56.5 (44) 101 (47) 48.2 (43) 93.2 (48) 61.7 (43) Body depth 71.0 (33) 23.1 (30) 43.0 (34) 73.4 (34) 37.6 (33) 64.7 (33) 50.0 (34) Orbit diameter 18.9 (09) 7.8(10) 11.6 (09) 21.9(10) 10.0(09) 19.9 (10) 12.5 (09) Snout length 255013). 9.2112). 17.1-(13) 28:4°(13), 14.2 (13): 626.7 (14) 19:.01(13) Interorbital width 11.0(05) 3.4(04) 6.7 (05) 12.2 (06) 5.0(04) 10.5 (05) 6.7 (05) Jaw length 46.8 (22) 16.1(21) 27.2 (21) 48.8 (23) 23.0(20) 46.7 (24) 29.9 (21) Predorsal-fin length 77.2 (36) 28.6 (37) 48.6 (38) 86.7 (40) 40.6 (36) 80.2 (41) 54.1 (37) Pectoral fin length 60.0(28) 23.0(30) 36.6 (29) 60.8 (28) 34.5 (31) 59.1 (30) 41.4 (29) Pelvic fin length 52.8 (25) 21.8 (28) 36.1(28) 54.8 (26) 29.3 (26) 51.6 (27) 41.0 (28) Caudal fin length 53.4(25) 23.5 (31) 37.8 (30) 56.7 (27) 29.7 (26) 50.8 (26) 39.5 (27) Sciences; LACM—Los Angeles County Museum of Natural History; USNM— United States National Museum. Scorpaena orgila Eschmeyer and Allen, new species. (Figures 1a, 2; table 1.) MATERIAL EXAMINED. Holotype: SPECIES ACCOUNTS CAS 24809, formerly BPBM 6770, a specimen 248 mm. in standard length, collected offshore of Ahu Akapu in 70 [Proc. 4TH SER. CALIFORNIA ACADEMY OF SCIENCES 518 ‘yaajsua Sapouang40rg “ “TS “Wu ce ‘ynpe ‘7849 Wadd Apsout paseq ‘ny340 Duapds0Ig “VY ‘saystjuordioos purys “TS wu 19 WNpe ‘8449 NAda UO AT}sour paseq q uo Asou paseq ‘szsuanasp¢ vuang4o9g “A “TS “uu zoz ‘yWnpe ‘4419 Wadd UO I Joyseq Mou ut sautds pray jo uoneyuasaidar dT}eULURISeIPIMag “~T AXNOTY VoL. XXXVIT] ESCHMEYER & ALLEN: EASTER ISLAND SCORPIONFISHES 519 feet, by J. E. Randall on 3 February 1969. Paratypes: BPBM 6772 (1 speci- men, 79 mm. in standard length), between Hanga Roa and Hanga Piko in 1-3 feet, J. E. Randall, G. R. Allen, B. A. Baker and E. Edwards, 25-26 January 1969. CAS 24810, formerly BPBM 6773 (2, 60-109), offshore from south end of Hanga Roa in 40 feet, J. E. Randall and G. R. Allen, 10 February 1969. USNM 205209, formerly BPBM 6769 (1, 212), off Motu Tautara in 60 feet, J. E. Randall, 7 February 1969. BPBM 6771 (1, 67), Hanga Piko in 1-3 feet, tidepools, J. E. Randall and G. R. Allen, 29 January 1969. BPBM 6776 (1, 48), offshore of Ahu Akapu in 80 feet, J. E. Randall and G. R. Allen, 5 February 1969. BPBM 6777 (1, 202), Motu Iti in 40 feet, J. E. Randall, 4 February 1969. BPBM 6774 (1, 127), boat channel at Hanga Piko in 6 feet, J. E. Randall, 18 January 1969. CAS 24811, formerly BPBM 6775 (1, 76), wreck about 20 meters offshore between Hanga Roa and Hanga Piko in 10-15 feet, J. E. Randall and G. R. Allen, 27 January 1969. LACM 31198-1, formerly BC 65-421 (1, 214), Hanga Piko in 180 feet, lobster trap, local fisherman, 3 January 1965. BC 65-438 (1, 113), Hanga Piko in 23 feet, I. E. Efford and J. A. Mathias, 14 Janu- ary 1965. BC 65-439 (1, 194), caught by islanders, no other data, [1965]. BC 65-457 (1, 145), Hanga Roa, subtidal, I. E. Efford and J. A. Mathias, no date [1965]. DeEscripTION. A large species, with slightly ctenoid scales on the sides, very shallow occipital pit, and chest and pectoral-fin base scaled. Dorsal fin with 12 spines and 9 soft rays (last double). Anal fin with 3 spines and 5 soft rays (last double). Pectoral fin short, not reaching level of anal fin; pectoral rays usually 17 (table 1), rays 2 through 6 or 7 branched in larger specimens, branching probably begins at about 45 mm. standard length. Gill rakers 17-19 (including rudiments), 5 on upper arch, 12—14 on lower arch. Spines as in figure la. Pre- orbital bone usually with 3 or 4 spinous points over the maxillary; first directed forward, with 1 or 2 tiny spines at its base, followed by large spine (sometimes as 2 close-set spines) which points down; small specimens with only anterior and posterior spines (not unusual for juveniles). Suborbital ridge usually with 5 spinous points, 2 on lateral face of preorbital bone and 3 on the suborbital bones; first spine on preorbital bone small and pointing up, second long and directed posteriorly and slightly below level of first spine on suborbital. Other spines present include nasal, pre-, supra-, and postocular, nuchal, parietal, tympanic, supplemental and 5 preopercular, upper and lower posttemporal, opercular, pterotic, sphenotic, and cleithral. Scales on sides slightly ctenoid; scales on belly and chest mostly without ctenii; head mostly naked, a few buried scales below suborbital ridge, behind eye, and on opercular bone. Vertical scale rows about 50-55. Vertebrae 24 (5 specimens) or 25 (1 specimen). Swimbladder absent. Palatine teeth present. Head and body with numerous filaments and tentacles; supraocular tentacle frilly, its length about equal to orbit diameter, but sometimes reduced or absent. Orbit smaller than snout, orbit diameter into snout length CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Un iw) =) FicurRE 2. Scorpaena orgila. A. Holotype, CAS 24809, adult, 248 mm. S.L. B. Paratype, BPBM 6772, juvenile, 79 mm. S.L. 1.1-1.6 times (lower values usually in smaller specimens); orbit diameter into head length 3.9—5.4 (lower values usually in smaller specimens). Color pattern in alcohol about as in figure 2 (taken from fresh specimens). Largest specimens tend to have less intense dark coloration. In life coloration may depend on depth of capture; juvenile specimens (shallower stations) everywhere mottled with gray, white, black, and brown, with dark pigmentation concentrated as in figure 2b. Individuals from deeper water (20 meters or more) mostly red with numerous small brown spots on head and body; areas appearing pallid in figure 2 white or translucent in life. CoMPARISONS. Scorpaena orgila seems to be more closely related to eastern VoL. XXXVII] ESCHMEYVER & ALLEN: EASTER ISLAND SCORPIONFISHES 521 TABLE 2. Counts and measurements for some type specimens of Scorpaena pascuensis. (Measurements are in millimeters ; numbers in parentheses are percentages of standard length.) CAS CAS CAS USNM USNM BPBM BC 24812 24813 24813 205210 205210 6784 65-451 Standard length 44.6 53.6 35.6 50.9 39.2 54.7 SN Geaieonerayss 1249 1249 1249 1249 1249 12-9 12-49 Anal fin ray* 3+5 3+5 345 3-+5 3+5 345 345 Pectoral fin ray* 16+16 16+16 16+16 15416 16+16 17+16 16416 Head length 18.9 (42) 23.2 (43) 15.4 (43) 21.5 (42) 16.4 (42) 22.2 (40) 15.0 (41) Body depth 16.3 (37) 20.5 (38) 12.7 (36) 19.2 (38) 13.6 (35) 19.6 (36) 13.1-(35) Orbit diameter De i2) OO (I1) -4.6413)_ 5:9.(12)" 4:7°(12) “62 1D) 4512) Snout length 5.3) (IZ) e312) Zonas) SGA) SECO) Sasaudy 33g) Interorbital width 2.2(05) 2.5 (05) 1.5(04) 2.3(05) 1.8(05) 2.5 (05) 1.9 (05) Jaw length DAE (A) WEA) 7/sor(0)) AES) 7 (BO) 2 WOE (al) 75 (BD) Predorsal-fin length 15.4 (34) 19.0(35) 12.7 (36) 19.5 (38) 12.7 (32) 18.2 (33) 12.3 (34) Pectoral fin length 15.1 (34) 17.9 (33) 12.6(35) 16.2 (32) 13.7 (35) 18.3 (33) 12.4 (34) Pelvic fin length 11.4 (26) 13.8(26) 9.9(28) 14.0(27) 10.4 (27) 14.5 (26) 10.3 (28) Caudal fin length 13)95) (GO) USI GL) OO (Gib) AGO) wy (Gad) Wee (oO) iho (O)) *Counts of 23 additional specimens: Dorsal rays 12 +9 (22 specimens) or 12+ 8 (1); anal rays 3+ 5 (23); pectoral rays 16-+ 16 (19), 15+ 16 (2), 16+ 15 (1), and one with 12 on left and 16 on right. and southern Pacific species than to those occurring in the central Pacific to east Africa. As now recognized, the genera Parascorpaena and Sebastapistes are included in the synonymy of Scorpaena, but the limits of the genus are poorly understood. Parascorpaena includes Indo-Pacific species which have cycloid, or at least feebly crenulate, scales on the body and have the posterior preorbital spine hooked forward; S. orgila does not possess these characters. Sebastapistes, also an Indo-Pacific group, contains species which have mostly ctenoid scales, partially scaled head, and the posterior preorbital spine pointing to the rear; the preorbital spination is different in S. orgila (fig. la). The genus Scorpaena con- tains eastern Atlantic species and a few Indo-Pacific species which have a naked chest. The chest in S. orgila is mostly scaled. Western Atlantic and most eastern Pacific species of Scorpaena have cycloid scales on the body, but they do not appear to be particularly closely related to the Indo-Pacific subgroup Para- scorpaena. Scorpaena orgila seems to be most closely related to S. cooki Gunther, 1873, from Norfolk Island and the Kermadec Islands, S. uncinata de Buen, 1961, from San Ambrosio and S. thomsoni Giinther, 1880, from Juan Fernandez Island, at least in squamation, body shape, and preorbital spination. Scorpaena cooku has more pectoral rays (probably 18 normally) and smaller scales (about 70 verti- cal scale rows) than does S. orgila. Scorpaena orgila is distinguished from S. uncinata by the presence in S. uncinata of a black patch on the distal third of the dorsal fin at the posterior dorsal spines. Scorpaena orgila also differs from S. thomsoni in coloration; the body is brownish with white spots in S. thomsont, while S. orgila lacks the white spots and tends to have dark brown spots on a pale background. In any event, these four species, three from the offshore 522 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 3. Scorpaena pascuensis. A. Holotype, CAS 24812, adult, 45 mm. SL. B. Paratype, CAS 24813, adult, 53 mm. S.L. islands of western South America and one from islands north of New Zealand, appear to be more closely related to each other than to other species. Scorpaena pascuensis, the other species of Scorpaena at Easter Island, is easily separated from S. orgila by having cycloid scales on the body, and it does not appear to be closely related on other features. EcoLocy. Scorpaena orgila was collected in tidepools and in depths to 60 meters. The large adults were taken at the deeper stations. This species is the largest and most colorful of the Easter Island scorpaenids, and is also the most Vor. XXXVII] ESCHMEYVER & ALLEN: EASTER ISLAND SCORPIONFISHES 523 aggressive towards the diver. Large specimens of approximately 250 mm. in standard length were frequently observed resting under ledges or seemingly guarding the entrances to caverns. On one occasion a specimen lunged out towards John Randall, actually striking the spear which he held in his hand. The natives refer to this species as the “‘nohu,” a name often associated with members of the genera Scorpaenopsis and Synanceia throughout the rest of Polynesia. Etymotocy. The specific name is based on the Greek adjective orgilos, prone to anger. Scorpaena pascuensis Eschmeyer and Allen, new species. (Figures 1b, 3, table 2.) MATERIAL EXAMINED. Holotype: CAS 24812, formerly BPBM 6783, 45 mm. in standard length, wreck about 20 meters offshore between Hanga Roa and Hanga Piko, depth 10-15 feet, J. E. Randall and G. R. Allen, 27 January 1969. Paratypes: BPBM 6782 (3 specimens, 26-32 mm. in standard length); CAS 24813 (3, 22-53), CAS 24814 (1, 40, cleared and stained), and USNM 205210 (2, 39-51), all formerly BPBM 6782; off Tahai, small rock ledge adjacent to sand patch in 35 feet, J. E. Randall and G. R. Allen, 7 February 1969. BPBM 6784 (5, 23-56), west shore, between Hanga Piko and Hanga Roa, depth 1-3 feet, tidepools, J. E. Randall, G. R. Allen, B. A. Baker and E. Edwards, 25-26 January 1969. CAS 24815, formerly BPBM 6785 (1, 26), Hanga Piko in 1-3 feet, tidepools, J. E. Randall and G. R. Allen, 29 January 1969. CAS 24816, formerly BPBM 6786 (3, 21-25), tidepools between Hanga Piko and Hanga Roa in 1-3 feet, J. E. Randall and G. R. Allen, 6 February 1969. BC 65-451 (4, 20- 36), Hanga Roa in 1-5 meters, I. E. Efford and J. A. Mathias, 2 February 1965. BC 65-417 (1, 21), Hanga Piko in 1 meter, I. E. Efford and J. A. Mathias, 31 December 1964. LACM 6560-14 (6, 33-45), east side of Anakena Cove, 100 yards NE. of sand beach, ketch Chiriqui, 0-15 feet, 1 October 1958. DescripTIoN. A small species, with cycloid scales, no well defined occipital pit, and chest and pectoral-fin base scaled. Dorsal fin usually with 12 spines and 9 soft rays (last double). Anal fin with 3 spines and 5 soft rays (last double). Pectoral fin reaching to or slightly beyond level of first anal spine; pectoral rays 15-16, usually 16, rays 2 or 3 through 5 or 6 branched in specimens greater than 39 mm. standard length, no branched rays in smaller specimens. Gill rakers 14— 16 (including rudiments), 4 on upper arch, 10-12 on lower arch. Location of head spines as in figure 1b. Preorbital bone with 2 spines over maxillary, first directed forward, second directed down and to rear. Suborbital ridge with 2 spines, first under eye, second at end of suborbital ridge. Other spines present include nasal, pre-, supra-, and postocular, nuchal, parietal, tympanic, supplemen- tal and 5 preopercular, upper and lower posttemporal, opercular, pterotic, sphenotic, and cleithral. Scales on body cycloid; vertical scale rows about 45; lateral line scales 23-24 plus 1 on caudal fin; most of head scaled. Vertebrae 524 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. TaBLe 3. Counts and measurements for some type specimens of Scorpaenodes englerti from Easter Island.(Measurements are in millimeters; numbers in parentheses are per- centages of standard length.) CAS BPBM BPBM CAS CAS CAS USNM 24806 6778 6778 24807 24807 24807 205211 Standard length 67.9 66.7 53.3 50.0 43.0 34.8 72.4 Dorsal fin rays* 13-+ 10 13+10 13+10 13+10 13-410 13+9 13+ 10 Anal fin rays* 345 345 345 3+5 345 3+5 3+5 Pectoral fin rays*¥ 19+19 19+19 19+19 20420 19419 19419 19419 Head length 29.2 (43) 28.4 (43) 23.7 (43) 21.8 (44) 18.8 (44) 15.2 (44) 31.7 (44) Body depth 24.3 (36) 24.0(36) 16.6(31) 16.4 (33) 14.9 (35) 11.9 (34) 21.3 (29) Orbit diameter 8:8 (13) 8:73) 7.314) (6:5°(13)) 52713) Ae ore len) Snout length 6.7(10) 65(10) 54(10) 4.9(10) 44(10) 3.6(10) 7.5 (10) Interorbital width 3.1(05) 3.1(05) 2.2 (04) 2.0(04) 1.9(04) 1.5 (04) 4.3 (06) Jaw length 14.021) 15.0:@22)) 11:5°(22)) 1101(22)) 957523) Vie G2elOrOnee.) Predorsal-fin length 27.3 (40) 27.5 (41) 21.2 (40) 20.7 (41) 16.7 (39) 13.8(40) 28.3 (39) Pectoral fin length 24.2 (36) 25.4 (38) 19.0(36) 18.7 (37) 15.6 (36) 13.3 (38) 27.1 (37) Pelvic fin length 17.1(25) 19.2 (29) 15.1 (28) 14.9(30) 12.0(28) 9.7 (28) 19.3 (27) Caudal fin length 18:3 (27) 20.1°G0) 15:8 (0)! 15.0)” 12.4929) SOS) eae aso) *Counts of 5 additional specimens: Dorsal rays 13-+10 (3 specimens), 13-+9 (1) and 14+9 (1); anal rays 3 +5 (5); pectoral rays 19 + 19 (4) and 18 + 18 (1). 24 (6 specimens). Swimbladder absent. Palatine teeth present. Dermal flaps and tentacles inconspicuous except for supraocular tentacles; length supraocular tentacles about equal to orbit diameter, sometimes reduced. Orbit diameter slightly greater than snout length in smaller specimens or about equal to snout length in larger specimens, orbit diameter into snout length .7—1.0; orbit diameter into head length 3.3-3.9 (smaller specimens tend to have lower ratio). Color pattern in alcohol about as in figure 3 (taken from fresh specimen) ; most con- spicuous feature a black spot on the opercle just posterior to first 3 preopercular spines (this feature also in Scorpaenodes englerti); body mostly light brown to gray on a pallid background, with a few small dark spots on fins usually present, but many specimens with fins mostly clear; a black spot on the dorsal fin between about spines 7-10, more restricted or absent in some specimens. Small specimens tend to have large pale areas between soft dorsal and anal fin and on caudal peduncle. In life, body mostly brown to greenish brown, mottled with light and dark areas; head brownish with small black dots, the dots extending onto the body; supraocular tentacles and other skin flaps on head and body pallid; spot behind preopercular spines brown to black; rays of soft dorsal fin, caudal fin, and anal fin alternately banded with brown and white; membranous portions of fins mostly unpigmented, except where body coloration extends onto bases of fins; base of pectoral fin strongly mottled with brown; belly and chest whitish. COMPARISONS. Scorpaena pascuensis may be separated from the other species of Scorpaena from Easter Island quite easily on the basis of coloration, but it also differs in spination (fig. 1) and other features as discussed in the text, e.g., VoL. XXXVII] ESCHMEYVER & ALLEN: EASTER ISLAND SCORPIONFISHES 525 Ficure 4. Scorpaenodes englerti, holotype, CAS 24806, adult, 68 mm. S.L. ctenoid scales on the body in S. orgila and cycloid scales in S. pascuensis. Scor- paena pascuensis does not seem to have a particularly close relative in the Indo- Pacific, and the species does not fall into either the subgroup Parascorpaena or Sebastapistes. No eastern Pacific species appears closely related either. EcoLtocy. Scorpaena pascuensis appears to be confined primarily to the rocky pools of the intertidal zone, where both adults and juveniles were collected. Several specimens were taken or observed in depths to about 15 meters, but none was collected or observed at the deeper depths sampled by Randall and Allen. Erymo.tocy. This specific name is based on the Spanish name for Easter Island, Isla de Pascua. Scorpaenodes englerti Eschmeyer and Allen, new species. (Figures 1c, 4; table 3.) MATERIAL EXAMINED. Holotype: CAS 24806, formerly BPBM 6780, 68 mm. in standard length, west shore of Easter Island, between Hanga Roa and Hanga Piko, in tidepools, J. E. Randall and G. R. Allen, 25 January 1969. Para- types: BPBM 6778 (2, 54-67), offshore from Ahu Akapu in 70 feet, J. E. Ran- dall and B. A. Baker, 3 February 1969. CAS 24807 (3, 34-50), CAS 24808 (1, 70, cleared and stained), and USNM 205211 (2, 52-71), all formerly BPBM 6781, off Motu Tautara in 125 feet, over sand bottom near ledge, J. E. Randall and B. A. Baker, 12 February 1969. BPBM 6779 (1, 60), Mataveri o Tai, depth 20 feet, boulder and sand bottom, J. E. Randall and G. R. Allen, 2 February 1969. BC 65-455 (1, 47), Hanga Roa, subtidal, I. E. Efford and J. A. Mathias, 5 February 1965. LACM 6560-44 (1, 74), east side of Anakena Cove, 100 yards NE. of sand beach, 0-15 feet, ketch Chiriqui, 1 October 1958. 526 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Description. Measurements and counts in table 3; location of head spines in figure 1c; body shape and coloration in figure 4. Dorsal fin with 13 spines and normally 9 soft rays (last ray double). Anal fin with 3 spines and 5 soft rays (last double). Pectoral fin with 18-19 rays, some rays branched. Gill rakers 18-20 (including rudiments), usually 6 on upper arch and 12-14 on lower arch. Vertebrae 24 (7 specimens). Preorbital bone with 2 broad lumps over maxillary. Suborbital ridge usually with 2 spinous points, first under eye, second at end of suborbital ridge; no secondary row of spines below the suborbital ridge. Preopercular bone with small supplemental spine and first three preopercular spines, fourth and fifth preopercular spines virtually absent. Upper posttemporal spine absent. Interorbital spines usually present; two spines on frontal bones near midline usually present. Other spines present include nasal, pre-, supra-, and postocular, anterior and posterior parietal, lower posttemporal, opercular, tympanic, pterotic, sphenotic, cleithral, and a small spine between tympanic and pterotic spines. Scales weakly ctenoid; verti- cal scale rows about 45—55; pectoral fin base, chest, and head scaled. Skin ap- pendages inconspicuous; usually small simple tentacles associated with most head spines. Color pattern in alcohol as in figure 4 (taken from fresh specimen) ; body and head with brown pigment on a pallid background, pigment mostly in large patches with paler areas between; a dark spot on opercle just behind upper preopercular spines usually well marked (as in Scorpaena pascuensis); brown patches on body concentrated in five or six saddle-shaped areas along the back, one across nape, three under spinous dorsal fin, one or two under soft dorsal fin, and one at base of caudal fin; dorsal fin with dusky or brown pigment as exten- sion from brown saddle-shaped areas; pectoral fin usually with brown pigment at base; other fins mostly clear, sometimes dusky distally. In life body light brown to reddish brown, mottled with tan areas; opercular spot brown; dorsal, anal, and pelvic fins with distal fourth red or reddish brown; pectoral fin mostly pale with distal third of fin rays reddish; caudal fin rays banded with red, with pale areas between. CoMPARISONS. Scorpaenodes englerti belongs with the group of species of the genus Scorpaenodes which lack an additional spine or spines below the main sub- orbital ridge, and with those species which tend to lack the interorbital spines. The species is closely related to S. xyris from the eastern Pacific, to S. littoralis from Japan, and to S. littoralis-like specimens recently collected by John E. Ran- dall at Hawaii and by the Australian Museum staff on the Barrier Reef off Aus- tralia. These species agree in body shape and spination, and they are quite simi- lar in coloration, especially in having the dark spot on the opercle behind the upper preopercular spines. Scorpaenodes englerti appears to average lower in pectoral ray counts. A thorough study of the S. littoralis-like species is needed, however. Vor. XXXVII] ESCHMEYER & ALLEN: EASTER ISLAND SCORPIONFISHES 527 EcoLocy. Scorpaenodes englerti exhibits a wide range of vertical distribu- tion, specimens being collected at all sampled depths (surge pools to 40 meters). Etymo.ocy. This species is named in honor of Father Sebastian Englert, who lived on Easter Island for over 30 years and was an avid student of the archaeology and natural history of the island. He died in New Orleans, Louisiana, while on a tour of the United States to raise funds for the restoration of archaeo- logical sites on Easter Island. He was buried in the courtyard of his small church on January 18, 1969, the same day that Randall and Allen arrived on the island. LITERATURE CITED DE BUEN, F. 1961. Peces chilenos. Familias Alepocephalidae, Muraenidae, Sciaenidae, Scorpaenidae, Liparidae y Bothidae. Montemar [continuation of Revista de Biologia Ma- rina], no. 1, pp. 1-52, figs. 1-11. 1963. Los peces de la Isla de Pascua, catalogo descriptivo e ilustrado. Boletin de la Sociedad de Biologia de Concepcidn (Chile), vols. 35-36 (1960-61), pp. 3-80, 33 figs. EscHMEYER, W. N. 1969. A systematic review of the scorpionfishes of the Atlantic Ocean (Pisces: Scor- paenidae). Occasional Papers of the California Academy of Sciences, no. 79, 130 pp., 13 figs. FUENTES, F. 1914. Contribucién al estudio de la fauna de la Isla de Pascua. Boletin del Museo Nacional de Chile, 1914, 37 pp., 11 pls., 1 map. GunTue_r, A. C. L. G. 1873. Andrew Garrett’s Fische der Siidsee. Vol. 1. Journal des Museum Godeffroy, vol. 2, Heften 3, 5, 7, and 9, 128 pp., 83 pls. 1880. Report on the shore fishes procured during the voyage of H.M.S. Challenger in the years 1873-1876. In Report on the scientific results of the voyage of H.M.S. Challenger during the years 1873-76..... Zoology, vol. 1, part 6, 82 pp., 32 pls. KENDALL, W. C., AND L. RADCLIFFE 1912. Reports on the scientific results of the expedition to the eastern tropical Pacific, in charge of Alexander Agassiz, by the U. S. Fish Commission steamer “Alba- tross,” from October, 1904, to March, 1905, Lieut. Commander L. M. Garrett, U. S. N., commanding. XXV. The shore fishes. Memoirs of the Museum of Comparative Zoology, Harvard, vol. 35, no. 3, pp. 75-171, 8 pls. Recan, C. T. 1913. A collection of fishes made by Professor Francisco Fuentes at Easter Island. Proceedings of the Zoological Society of London, 1913, pp. 368-374, pls. 55-60. RENDAHL, H. 1921. The fishes of Easter Island. Jn The Natural History of Juan Fernandez and Easter Island, edited by Carl Skottsberg. Vol. 3, Zoology, part 1, pp. 59-68. Uppsala. WILHELM, O. E., anp A. L. HuLor 1957. Pesca y peces de la Isla de Pascua. Boletin de la Sociedad Biologia de Concep- cidn (Chile), vol. 32, pp. 139-152, 4 figs. . | = 7 ® Ox * 1 he Ua Meigs preaeTsny ) @ @) Of eine ati LA es ee pid - i 7e—< G vg WC — mie ten | p40 baer aie ae - a] ois Pad - Pi ) PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XXXVII, No. 20, pp. 529-535; 1 figure. November 23, 1971 EAST ASIATIC AND ORIENTAL SPECIES OF STENUS REPRESENTED IN THE COLLECTION OF THE CALIFORNIA ACADEMY OF SCIENCES (COLEOPTERA: STAPHYLINIDAE) By Volker Puthz 6407 Schlitz, Limnologische Flusstation, Germany Through the kindness of Mr. Hugh B. Leech I borrowed a small number of west hemisphere species of Stenus. Amongst the material which is quoted here, there was a new species from Taiwan. This species is described below. Although our knowledge of the Taiwan Stenus fauna is very poor, a key to all species known from that island is given. Stenus (sensu stricto) alienus Sharp. Stenus alienus SHARP, 1874, Trans. Ent. Soc. London (1874), p. 81. MATERIAL. One male: Tokyo, 17 May 1931, Gressitt; one female: Man- churia: Mukden, 14 August 1923, Van Dyke. This species is widely distributed in the southeast palearctic; it is also known from Taiwan: Takakiyama, Sauter (Zoological Museum, Berlin). Stenus (sensu stricto) comma LeConte. Stenus comma LEConrE, 1863, Smithson. Miscell. Coll., no. 167, p. 50. MATERIAL. Two females: Nanking, 14 September 1923, Van Dyke; one female: Manchuria: Kirin, 1 September 1923, Van Dyke; one female: Man- churia: Halling NE, 30 August 1923, Van Dyke. This species occurs over all the northern hemisphere. [529] Laboratory LIBRARY DEC1 1971 Woeds Hicde. Mass. 530 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 1. Stenus (sensu stricto) imsulanus Puthz, new species (paratype), ventral aspect of aedeagus. Stenus (sensu stricto) insulanus Puthz, new species. (Figure 1.) This new species belongs to the species complex which has the abdomen feebly margined and in which abdominal keels are lacking. Black, feebly shining, coarsely, partly rugosely, and very closely, punctured, feebly pubescent. Antennae dark brown. Palpi reddish yellow, 3rd joint infus- cated. Legs dark brown. Length 4.7—5.1 mm. Description. Head: distinctly narrower than elytra (1,000 mw : 1,100 p), front moderately broad (average distance between eyes: 602 mu), deeply con- cave with two longitudinal furrows, median part narrower than each of the side-pieces, feebly elevated, deeply under the level of the inner margin of eyes. Punctation moderately coarse and very dense, diameter of puncture larger than section of 3rd antennal segment, interspaces at most as large as half of a punc- VoL. XXXVIT] PUTHZ: ORIENTAL SPECIES OF STENUS 531 ture. Antennae: damaged in the types, presumably constructed as in S. sau- terianus. Prothorax: distinctly longer than broad (884 : 756 &), broadest some- what behind middle, sides to anterior margin nearly straight, to posterior margin distinctly concave. A feebly horizontal constriction behind anterior margin and a short longitudinal impression on posterior part discally. Punctation coarse and rugose, somewhat coarser than on head. Elytra: distinctly broader than head (1,100 w : 1,000 4), about as broad as long, humeri prominent, sides roundly enlarged, restricted in posterior third, posterior margin moderately deeply emarginate. Sutural impression deep, humeral impression distinct. Punctation coarse and very dense, coarser than on pronotum, rugosely in lateral direction in outer two-thirds. Abdomen: feebly narrowed towards apex, sides very feebly margined, para- tergites almost smaller than one puncture, basal restrictions of first tergites deep without any cusp. Punctation moderately fine and very dense, somewhat finer than on the front of the head. Legs robust, hind tarsi about as long as two- thirds of the tibiae, Ist segment longer than 2nd and 3rd together, longer than the last. The dorsal surface of the whole insect lacks ground sculpture. Male: metasternum deeply coriaceous. Tibiae without apical spines. Fourth to 6th sternites very feebly emarginate at posterior margin; 7th sternite with a shallow but distinct emargination at posterior margin, in front of it shallowly impressed and between the punctures deeply coriaceous; 8th sternite with a moderately deep notch in posterior tenth; 9th sternite with strong apicolateral teeth, between them deeply emarginate and serrate; 10th tergite at smooth posterior margin, broadly rounded, slightly concave in the middle. Aedeagus (fig. 1): broad, the median lobe with a broad lancet-like anterior portion set with short setae ventrally. In the inner lobe there are broad longi- tudinal expulsation bands and a membranous internal sac with short spines or clasps having a stronger sclerotized portion apically. The parameres are shorter than the median lobe, enlarged anteriorly, and with many setae. Female: eighth sternite rounded apically; valvifera with a strong lateral tooth; 10th tergite as in male; no sclerotized spermatheca. Type MATERIAL. Holotype, male, and male and female paratypes (female partly damaged): Taiwan, Keelung (= Chi-lung), June 1954, T. C. Maa. Holotype in the California Academy of Sciences, paratypes in the California Academy of Sciences and in the collection of the author. At present I do not know the sister-species (sensu Hennig) of Stenus insulanus. Its closest relatives known are S. duplex Benick (China) and S. sauterianus Bernhauer (Taiwan). For identification see the key below. Key TO THE SPECIES OF STENUS FROM TAIWAN 1 (24) Abdomen margined 2 (11) 4th tarsal segment simple 532 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Elytra with a reddish yellow spot. Length 4.2-5.0 mm. __.____. S. alienus Sharp Elytra unicolorous Abdomen without carinae in basal restrictions of tergites Abdomen broadly margined, elytra near suture irregular and smooth. Median lobe nearly straight-sided to acute apex. Length 5.0-5.5 mm, — Stl I a a Te 2 ee ee ee S. sauterianus Bernhauer 7 (6) Abdomen very finely margined, elytra near suture coarsely and very densely punctured. Median lobe as in fig. 1. Length 4.7-5.1 mm. —____ a Bo a ee S. insulanus Puthz, new species Num & W 8 (5) Abdomen with keels in basal restrictions of tergites 9 (10) Abdominal carinae 3-cuspid; male sexual characters as in fig. 4 of Benick, 1041s Wength 5055:8 mine 2 ae eee eee S. formosanus L. Benick (synonyms S. formosae Wendeler, S. forterugosus Bernhauer, S. submarginatus Bernhauer) 10 (9) Abdominal carinae 4-cuspid. Aedeagus (compare Benick 1915, fig. 5). Length 20-3 SMM = ee S. melanarius verecundus Sharp 11 (2) 4th tarsal segment bilobed 12 (15) Elytra immaculate 13. (14) Larger species, 6.0 mm. long. Median lobe broadly truncate at apex, apical POLLOnNOngen shan basaluonc eee ee S. rugosipennis Cameron 14 (13) Smaller species, 3.74.2 mm. long. Median lobe rounded at apex, parameres enlarged strumpetslikestoy apex: eee S. rugipennis Sharp (Stenus sharpianus Cameron, 1930 (Ent. Monthly Mag., vol. 66; A 205), new synonymy.) I borrowed the types of S. rugipennis and S. sharpianus from the British Museum (Natural History); they are conspecific! 15 (12) Elytra with yellow or reddish spots 16 (19) Head nearly as broad or broader than elytra with very large eyes 17 (18) Elytral spot larger, as large as 4-5 punctures together. Aedeagus as in fig. 12 of Puthz, 1968a. Length 6.0-7.2 mm. —— S. gestroi taiwanensis Puthz (in the diagnosis I regarded this as a subspecies of S. submaculatus Bernhauer ; since then I have found that S. submaculatus and S. tatwanensis both are sub- species of the polytypic S. gestroi Fauvel) 18 (17) Elytral spot smaller, as large as 2 punctures together. Median lobe extending distinctly beyond the parameres. Length 5.2-5.7 mm. _ S. stigmatipennis L. Benick 19 (16) Head distinctly narrower than elytra, eyes smaller 20 (21) Larger, abdomen very finely bordered laterally. Aedeagus as in fig. 19 of eyed, MOE, Welt SOO) tere, S. miwai Bernhauer 21 (20) Smaller, not exceeding 5.3 mm. 22 (23) Abdominal punctation fine and very dense. Aedeagus as in fig. 62 of Puthz (969) Helbencthy425=5 Otmmr) 2) a ee ee eee S. virgula Fauvel (synonym S. kwantungensis Cameron) 23 (22) Abdominal punctation moderately coarse and moderately close. Aedeagus nar- rower, median lobe truncate apically. Length 4.5-5.3 mm. __ S. arisanus Cameron 24 (1) Abdomen emarginate. Prothorax and abdominal segments 3 to 6 reddish yellow. Aedeagus as in fig. 3 of Scheerpeltz, 1957. Length 5.0-5.6 mm. ___ ee eee ee one ee aera ioly Sh Ree ee S. flavidulus paederinus Champion (one male in British Museum from Taiwan). Stenus (Hypostenus) mercator Sharp. Stenus mercator SHARP, 1889, Ann. Mag. Nat. Hist., 6th ser., vol. 3, no. 16, p. 333. VoL. XXXVIT] PUTHZ: ORIENTAL SPECIES OF STENUS 533 MATERIAL. One male: Nanking, 14 March 1923, Van Dyke. Known from China to Japan. Stenus (Hypostenus) cicindeloides (Schaller). Staphylinus cicindeloides SCHALLER, 1783, Abh. Hallisch. Naturforsch. Ges., vol. 1, p. 324. MATERIAL. Six females: Nanking, 23 March and 21 April 1923, Van Dyke; two females: Suisapa, 1,000 m., Lichuan District, West Ho-pei, 25 July 1948, Gressitt and Djou. This species occurs over the whole Palearctic region, south to Vietnam. . Stenus (Hypostenus) tropicus Bernhauer. Stenus tropicus BERNHAUER, 1915, Philippine Jour. Sci., ser. D, vol. 10, p. 119. MATERIAL. One male: Manila, May 1906, J. C. Thompson. Well known from the Philippines. Stenus (Hypostenus) nothus L. Benick. Stenus nothus L. Benicx, 1929, Dtsch. Ent. Zeitschr., vol. 4, p. 264. MATERIAL. Three males, two females: San Jose, Mindoro, April, October 1945, E. S. Ross. Known from Mindoro, also distributed over Borneo and Celebes (British Museum). Stenus (Hypostenus) ambiguus L. Benick. Stenus ambiguus L. BENICK, 1929, Dtsch. Ent. Zeitschr., Heft 4, p. 266. MATERIAL. Two males, two females: Luzon: Mabatobato, Pili Camarines Sur, 16 May 1931, E. E. Schneider. The aedeagus of this species strongly resembles that of S. nothus Benick, but the median lobe extends somewhat beyond the parameres. Stenus (Hypostenus) spinosus L. Benick. Stenus spinosus L. BENIcK, 1921, Ent. Mitt., vol. 10, no. 6, p. 193. MATERIAL. One male: Mt. Makiling, Laguna, 800 ft., 5 May 1932, F. C. Hadden. This species was described from Mt. Makiling. Stenus (Hypostenus) subtropicus Cameron. Stenus subtropicus CAMERON, 1949, Proc. U.S. Nat. Mus., vol. 99, no. 3247, p. 464. MATERIAL. One female: Hai-nan: Ta Hiau, 16 June 1935, Gressitt. This species has the aedeagus of S. basicornis Kraatz and might represent an eastern subspecies of it. A male has also been collected from Pho Vi, Tonkin (in collection of Benick). 534 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Stenus (Parastenus) bicolon javanicus Bernhauer. Stenus (Hemistenus) javanicus BERNHAUER, 1915, Tijdschr. Ent., vol. 58, p. 216. Stenus (Parastenus) javanicus L. BENIcK, 1938, Stettiner Ent. Zeitung, vol. 99, no. 1, pp. 10, 11 (fig.). Stenus (Mesostenus) bernhaueri CAMERON, 1925 (nec S. bernhaueri Poppius, 1907), Treubia, vol. 6, no. 2, p. 177. Stenus (Hemistenus) maximiliani SCHEERPELTZ, 1933, Coleopt. Cat., vol. 129, p. 1192. Stenus (Parastenus) bicolon javanicus PuTHz, 1968b, Notulae Ent., vol. 48, p. 100. MarTeERIAL. One female: Bali: Bretam Lake, 3,600 ft., 1 October 1956, J. Sedlacek. Until now only known from Java. LITERATURE CITED Benick, L. 1915. Ueber Stenus morio Grav. und melanarius Steph., nebst Beschreibung einer neuen deutschen Art (Col.). Entomologische Mitteilungen, vol. 4, pp. 226-234. 1921. Nomenklatorisches tiber Steninen (Col., Staph.). Entomologische Mitteilungen, vol. 10, pp. 191-194. 1929. Die Stenus—Arten der Philippinen. (Col., Staphyl.). Deutsche Entomologische Zeitschrift, Jahrgang 1929, Heft 4, pp. 241-277, figs. 41-82. 1938. F. C. Dreschers Steninen—Sammlungen von Java und Sumatra (Col., Staph.). Stettiner Entomologische Zeitung, Jahrgang 99, Heft 1, pp. 1-49, 12 figs. in text. 1941. Weitere ostchinesische Steninen (Col., Staph.). Stettiner Entomologische Zeitung, Jahrgang 102, pp. 274-285. BERNHAUER, M. 1915a. Zur Staphylinidenfauna der Philippinen. VI. Beitrag zur Kenntnis der Indo- Malayischen Fauna. Philippine Journal of Science, Section D, vol. 10, pp. 117- 129. 1915b. Neue Staphyliniden aus Java und Sumatra (7. Beitrag zur indo-malayischen Staphylinidenfauna). Tijdschrift voor Entomologie, vol. 58, pp. 213-243. CAMERON, M. 1925. New Staphylinidae from the Dutch East Indies. Treubia, vol. 6, livraison 2, pp. 174-198. 1930. New species of Staphylinidae from Japan. Entomologist’s Monthly Magazine, vol. 66, pp. 181-185, 205-208. 1949. New species and records of staphylinid beetles from Formosa, Japan, and South China. Proceedings of the United States National Museum, vol. 99, no. 3247, pp. 455-477. HEnnIc, W. 1965. Phylogenetic Systematics. Annual Review of Entomology, vol. 10, pp. 97-116. LeConrTeE, J. L. 1863. New species of North American Coleoptera. Part I. Smithsonian Miscellaneous Collections, vol. 6, no. 167, pp. 1-168. (Pages 1-86 published March 1863; pp. 87-168 in April 1866.) PuTHz, V. 1968a.On some East Palearctic Steni, particularly from Japan (Coleoptera, Staphylin- idae). 52. Contribution to the knowledge of Steninae. Entomological Review of Japan, vol. 20, nos. 1-2, pp. 41-51. VoL. XXXVIT] PUTHZ: ORIENTAL SPECIES OF STENUS 535 1968b. Neue Steninen aus der Sowjetunion nebst synonymischen Bemerkungen (Coleop- tera, Staphylinidae). 53. Beitrag zur Kenntnis der Steninen. Notulae Ento- mologicae, vol. 48, pp. 93-102. 1969. Revision der Fauvelschen Stenus-Arten, exklusive madagassische Arten. 55. Bei- trag zur Kenntnis der Steninen. Bulletin de ]’Institut royal des Sciences naturelles de Belgique, vol. 45, no. 9, pp. 1-47. SCHALLER, J. G. 1783. Neue Insekten. Abhandlungen der Naturforschungs Gesellschaft, vol. 1, pp. 217- 332. (Dessau & Leipzig.) SCHEERPELTZ, O. 1933. Staphylinidae VII: Supplementum 1. Coleopterorum Catalogus (W. Junk, ed. S. Schenkling), Pars 129, pp. 991-1500. 1957. Staphylinidae (Col.) von Sumba und Flores (4. Beitrag zur Kenntnis der Staphy- liniden der orientalischen Region). Verhandlungen der Naturforschenden Gesell- schaft in Basel, vol. 68, pp. 217-357. SHarp, D. 1874. The Staphylinidae of Japan. Transactions of the Entomological Society of London for the year 1874, pp. 1-103. 1889. The Staphylinidae of Japan (continued from p. 267). Annals and Magazine of Natural History, ser. 6, vol. 3, no. 16, pp. 319-334. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XXXVII, No. 21, pp. 537-566; 33 figs. November 23, 1971 THE TEREBRIDAE (GASTROPODA) OF CLARION, SOCORRO, COCOS, AND GALAPAGOS ISLANDS By Twila Bratcher and R. D. Burch Invertebrate Zoology, Los Angeles County Museum of Natural History The Ecuador-owned Galapagos archipelago touches the equator 600 miles west of Ecuador. In spite of lying in the equatorial zone, the Humboldt Current from the Antarctic, flowing north along South America’s coast and swinging west through the Galapagos, brings water cold enough to support penguins and seals. The Islands, which are scattered over an area of 3,000 square miles, now have Spanish names. In this study the Spanish names will be used with their English or secondary names in parentheses. Since Charles Darwin’s day, the Galapagos Islands have fascinated naturalists as being the ‘cradle of new species” for both vertebrate and invertebrate fauna, so it is not surprising that some of the Terebra species should prove to be new. Some of the species from this area are so variable that it would be difficult to assign specimens at the ends of the variability range to the same species if there were not intergrades between the two extremes available for study. This study originally was intended to include only the Galapagos Islands, but when additional material was made available, we included Cocos Island, Costa Rica, and Socorro and Clarion islands, Mexico. It is almost to our embarrassment that we discovered four new species among the 16 species we examined from the off-shore islands (Bratcher and Burch, 1970, pp. 1-6, figs. 1-9). Each of these new species was checked against the original description, original figure (if any), and wherever possible, the holotype or photograph of the holotype of each of the more than 180 Recent and fossil species of Terebra described from the eastern Pacific area and related species from the [537] LIBRARY DEC 1. 1074 Weeds Hole, Maes. 538 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Indo-Pacific and western Atlantic areas. Many of the names proposed for the eastern Pacific Recent and fossil species of Terebra are synonyms, but only those concerned with the areas included in this paper will be assessed as to validity here. This list is not expected to be all-inclusive, but it contains each of the species we have encountered in the above-mentioned material. ACKNOWLEDGMENTS We wish to express our thanks to Dr. Leo G. Hertlein for his encouragement, for the loan of material from the California Academy of Sciences, for making literature available, and for making holotypes accessible for study and photog- raphy. In the same institution, Maurice Giles prepared the photographs (with the exception of figure 20) used as text figures; Mrs. Margaret Hanna kindly retouched the original photographs shown in figures 2, 4, 7, 9, and 29; and Barry Roth aided in the arrangement of the illustrations. Acknowledgments also are due to Dr. James McLean for the loan of Hancock Expedition material from the Los Angeles County Museum of Natural History; Drs. Harald Rehder and J. Rosewater of the National Museum for the loan of types and material; Dr. George Radwin of the San Diego Museum of Natural History for loan of material; and William Old, Jr. and Dr. William K. Emerson of the American Museum of Natural History for the loan of material. We also appreciate the loan of speci- mens for this study by the Ben Purdys of San Diego, California, the Carl Shys of Westminster, California, and Jacqueline DeRoy of Academy Bay, Santa Cruz (Indefatigable) Island, Galapagos Islands. We wish to thank Dr. Norman Tebble for making available holotypes in the British Museum (Natural History) for study and photography; Dr. Stillman Berry, the John Q. Burches, and Dr. R. W. Barker for the use of their fine libraries; the Hancock Library and the California Academy of Sciences library personnel for their help and cooperation; Allyn Smith for his encouragement; and Ford Bratcher for understanding the time needed for research in this study. SYSTEMATICS TOXOGLOSSA Troschel, 1848 Family TEREBRIDAE H. and A. Adams, 1853 Genus Terebra Bruguiére, 1789 Terebra stohleri Bratcher and Burch. (Figures 1, 2.) Terebra stohleri BRATCHER and Burcu, Los Angeles County Mus. Nat. Hist., Contrib. in Sci., no. 188, p. 5, figs. 7, 8, May 4, 1970. Type. Holotype, Los Angeles County Museum of Natural History—Allan Hancock Foundation no. 1180. Vor. XXXVII] BRATCHER AND BURCH: GALAPAGOS TEREBRIDAE 539 TYPE LocaLity. “Braithwaite Bay, Socorro Island, Mexico, 18° 42’ 20” N., 110° 56’ 15” W., sand and red mud bottom.” 18 to 37 meters (10 to 20 fathoms). DisTRIBUTION. Cabo Pulmo, Baja California, Mexico, to Socorro Island, Mexico, 2 to 23 fathoms. DEscriPTION. Shell medium-small, sturdy; color and appearance of ivory with faintly darker blotches; color of nucleus slightly darker than the following whorls; whorls convex; sutures deeply channeled, constricted; subsutural band inconspicuous; whorls slightly shouldered anterior to suture; sculpture of sharp, slightly curved axial ribs about equal to the interspaces crossed by evenly spaced spiral grooves; body whorl of medium length; posterior to the periphery sculp- ture remains the same; anterior to periphery axial ribs continue, and _ spiral grooves become more numerous and close set; aperture elongate; outer lip thin with sculpture pattern showing through; columella straight and simple; siphonal fasciole striate; anterior canal short, recurved. Discussion. No species known to us can easily be confused with this beauti- fully sculptured shiny terebra. Terebra purdyae Bratcher and Burch. (Figures 3, 4.) Terebra purdyae BRATCHER and Burcu, Los Angeles County Mus. Nat. Hist., Contrib. in Sci., no. 188, p. 5, figs. 5, 6, May 4, 1970. Type. Holotype, Los Angeles County Museum of Natural History—Allan Hancock Foundation no. 1182. Type Locatity. “North of Santa Maria (Charles) Island, Galapagos Islands, Ecuador, 0° 59’ S., 90° 25’ W., 70-80 fms., sand and rock bottom.” DIsTRIBUTION. Panama to Peru, 16 to 146 meters (8 to 80 fathoms). DeEscriPTION. Shell small and slender; shiny pale cream color with incon- spicuous fulvous blotches; nucleus of four purple-beige convex whorls; remainder of whorls almost flat; suture fairly well marked; barely evident subsutural band of beading slightly more prominent than the bands of beading on remainder of whorl; sculpture finely cancellate (remarkably consistent from second _ post- nuclear whorl) crossed by cords giving a beaded effect; axial ribs about equal to interspaces; body whorl of medium length; sculpture continuing anterior to periphery but less well defined; aperture elongate and slender, interior being the same color as exterior; columella straight with one microscopic plication; faint siphonal fasciole with posterior keel; anterior canal short, recurved. Discussion. Although the sculpture of Terebra purdyae is one of the most finely cancellate known to us, it does have some general resemblance to several other species. Terebra panamensis Dall (1908) is larger, with a coarser cancellate sculpture, fewer ribs, and with interspaces wider than the almost equal inter- spaces of T. purdyae. The fossil species T. gatunensis Toula (1908) and the 540 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 1. Terebra stohleri Bratcher and Burch. Holotype no. 1180, Los Angeles County Museum of Natural History, Type Collection. Braithwaite Bay, Socorro Island, Mexico. Length 21.4 mm., width 5.1 mm. FicurE 2. Nucleus, same shell as figure 1. Ficure 3. Terebra purdyae Bratcher and Burch. Holotype no. 1182, Los Angeles County Museum of Natural History, Type Collection. Santa Maria (Charles) Island, Galapagos Islands. Length 13.9 mm., width 3.3 mm. Ficure 4. Nucleus, same shell as figure 3. subspecies 7. g. kugleri Rutsch (1934) from the Neogene of Panama and Vene- zuela, are much larger species with less cancellate sculpture and wider subsutural bands. Terebra stohleri Bratcher and Burch (1970) has more convex whorls and spiral grooves rather than spiral cords. The sculpture of 7. purdyae most closely resembles that of 7. shyana Bratcher and Burch (1970) but T. purdyae is a smaller, more slender species. Average size about 13 mm. Terebra jacquelinae Bratcher and Burch. (Figures 5, 6, 7.) Terebra jacquelinae BRATCHER and Burcu, Los Angeles County Museum, Contrib. in Sci., no. 188, p. 2, figs. 3, 4, May 4, 1970. Type. Holotype, California Academy of Sciences, Department of Geology, Type Collection no. 13215. Type LocaLity. “Academy Bay, Santa Cruz (Indefatigable) Island, Gala- Vot. XXXVIT] BRATCHER AND BURCH: GALAPAGOS TEREBRIDAE 541 pagos Islands, Ecuador, 0° 46’ 16” S., 90° 19” 38” W., CAS loc. 39585, about 10 fathoms, on hard packed coralline sand bottom.” DISTRIBUTION. This species seems to be confined to the Galapagos Islands. Specimens have been taken from the vicinity of Santa Cruz (Indefatigable), San Salvador (James), and Baltra (South Seymour) islands in water from 4 to 37 meters (2 to 20 fathoms). DescriPTION. Shell of medium size; usually shiny cream color throughout; one and one-half shiny opaque nuclear whorls, the first somewhat bulbous; the early postnuclear whorls flat, the remaining whorls being very concave: sculpture on first postnuclear whorls consists of weak axial ribs ending in small nodes at suture with obsolete spiral grooves which cross the ribs; apical angle increases after about the sixth postnuclear whorl and ribs begin fading at center of whorl while nodes at rib endings become more prominent, those anterior to suture being slightly more prominent than the posterior ones; interspaces on later whorls contain minute microscopic growth striae, body whorl of medium length with obsolete axial ribs ending in nodes at periphery; anterior to row of nodes, weakly incised spiral grooves cross equally weak axial ones; aperture elongate, white within; columella white, straight with rounded plication which continues as keel setting off well developed siphonal fasciole; anterior canal short, recurved. Discussion. Although most specimens we have examined are of a light cream color, we have seen a number of individuals, most of which were collected at San Salvador (James) Island, which range in color from beige to light brown with cream colored nodes. While there is little variation among individuals of this spe- cies except that some are more slender than others, some small or immature specimens have the peripheral nodes forming a sharp keel differing in appearance from adults. There are several species, concave in profile, which somewhat resemble Terebra jacquelinae in general appearance. Terebra frigata Hinds differs from T. jacquelinae in that the former is smaller, less concave, and has continuous ribs from suture to suture. The ribs are less bulbous at their anterior endings, and there is more pronounced spiral sculpture and a more slender profile. In the latter the ribs fade out at the middle of the whorl; the spiral sculpture is weak or barely evident; and the nodes at the suture are more inflated. The Gulf of Mexico species T. concava Say (1827) is a much more slender form with small sharp nodes at the subsutural band and periphery and more numerous spiral grooves than T. jacquelinae. The East Indian species T. constricta Thiele (1925) is considerably smaller than 7. jacquelinae and has fine spiral striae, and the Persian Gulf species T. contracta E. A. Smith (1873) is also a smaller species with more numerous spiral striae and a laminated columella not evident in 7. jacquelinae. The average size of this species is about 30 mm. The largest specimen 542 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. ~ Ficure 5. Terebra jacquelinae Bratcher and Burch. Holotype no. 13215, California Academy of Sciences, Department of Geology, Type Collection. Academy Bay, Santa Cruz (Indefatigable) Island, Galapagos Islands. Length 33.2 mm., width 8.4 mm. Ficure 6. Middle whorls, same shell as figure 5. Ficure 7. Nucleus, same shell as figure 5. examined by us is 36.9 mm. in length, 10.8 mm. in diameter. It is in the collection of Mrs. Jacqueline DeRoy. Terebra hertleini Bratcher and Burch. (Figures 8, 9.) Terebra hertleini BrRatcHER and Burcu, Los Angeles County Museum, Contrib. in Sci., no. 188, p. 1, figs. 1, 2, May 4, 1970. Type. Holotype, California Academy of Sciences, Department of Geology, Type Collection no. 13222. TYPE Locatity. “Academy Bay, Santa Cruz (Indefatigable) Island, Gala- pagos Islands, Ecuador, 08° 46’ 16” S., 90° 19’ 38” W., CAS loc. 27536, 3.5 to 5.5 fms, dredged.” Vor. XXXVIT] BRATCHER AND BURCH: GALAPAGOS TEREBRIDAE 543 Ficure 8. Terebra hertleini Bratcher and Burch. Holotype no. 13222, California Academy of Sciences, Department of Geology, Type Collection. Santa Cruz (Indefatigable) Island, Galapagos Islands. Length 11.8 mm., width 5 mm. FicureE 9. Nucleus, same shell as figure 8. DISTRIBUTION. All the specimens of this species examined by us came from the Galapagos Islands in depths from 6 to 46 meters (3.5 to 25 fathoms). DEscRIPTION. Shell small, white, turreted; two glassy, convex, nuclear whorls, the first one constricted; postnuclear whorls flat; sculpture consists of spiral cords, about three per whorl, and obsolete axial ribs ending in large nodes anterior to suture; sculpture on body whorl of three spiral cords crossing obsolete ribs that end in faint nodes at periphery, these nodes being less prominent than those at the suture; anterior to periphery spiral cords cross minute axial stri- ations; aperture somewhat quadrate; outer lip thin, white within; columella straight with one weak plication; well developed siphonal fasciole with sharp keel; anterior canal of medium length, recurved. Discussion. The spiral cords on shells of this species are variable, being well 544 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. defined on some individuals and almost obsolete on others. The nodes at the periphery of the body whorl are inconspicuous in some specimens and more prominent in others. The apertures of all specimens examined exhibit a rather quadrate look, and some have a definite flair at the periphery. Although Terebra hertleini is one of the smaller species of terebrids, it presents some general superficial resemblance to larger species which also are profoundly turreted. Terebra tiarella Deshayes (1857) is a larger species, with more numer- ous and finer spiral striae and an elongate aperture that does not exhibit the peripheral swelling of T. hertleini. The distinct axial ribs of T. frigata Hinds (1844) along with its more slender and longer nucleus and spiral grooves dis- tinguish that species from 7. hertleini, which has indistinct, fading axial ribs, a broad dome-like nucleus, and spiral cords. Juvenile and subadult specimens of 7. hertleini begin to show a crenulated, turreted, subsutural band about the fourth postnuclear whorl. Terebra armillata Hinds (1844) and the fossil subspecies, T. armillata sheppardi Pilsbry and Olsson (1941) along with T. nelsoni Hanna and Israelsky (1925) are all much larger in size than T. hertleini. The last two species are fossil in the Neogene of Peru and Ecuador. Terebra jacquelinae Bratcher and Burch (1970) is much larger. The holotype of T. hertleini with ten whorls measures 11.8 mm. in length while the holotype of 7. jacquelinae with 13 whorls measures 33.2 mm. Terebra jacquelinae has a row of large nodes posterior to the suture which 7. Aertleini does not have. Terebra maculata maculata (Linnaeus). (Figures 12, 13.) Buccinum maculatum LINNAEUS, Syst. Nat., ed. 10, p. 741, 1758. Ref. to Buonanni, Recr. and Mus. Kircher, 3, fig. 317, 1684; Rumphius, Amboin. Rariteit., pl. 30 (in part), fig. A. (only; not figs. B, D), 1705; Gualtieri, Index. Test., pl. 56, fig. I (all; not fig. B), 1742; Argenville, Conchyl., pl. 14, fig. A, 1742. Dopcr, Bull. Amer. Mus. Nat. Hist., vol. 111, Art. 3, pt. 4, p. 218, 1956. Not Buccinum maculatum Linnaeus var. KAEMMERER, Cab. Rudolstadt, p. 152, no. 1la, 1786. (Referable to Terebra.) Not Buccinum maculatum Linnaeus vars. B and C, GMELIn, Syst. Nat., ed. 13, p. 3499, 1791. (Referable to Terebra.) Acus columna trajana HumMPuHREY, Mus. Calonnianum, p. 31, 1797. (Invalid for Nomencla- ture, International Commission of Zoological Nomenclature: Opinion 51). Not Terebra maculata PERRY, Conch., pl. 16, fig. 2, 1811. Terebra maculosa Lamarck, K1ener, Icon. Coq. Viv., vol. 8, pt. 5, Index to species of Terebra, 1839. Terebra maculata var. confluens DAUTZENBERG, Mem. Mus. Nat. Hist. Belg., Ser. 2, vol. 17, p. 31, 1935. Terebra maculosa (Linnaeus) HANNA and Hert etn, Proc. Calif. Acad. Sci., Fourth series, vol. 30, no. 3, p. 67, 1961. Terebra cf. T. maculata (Linnaeus), Emerson and Otp, Nautilus, vol. 77, no. 3, p. 91, 1964. Type. Holotype, probably in Linnaean Society collection in London (Dodge). VoL. XXXVII] BRATCHER AND BURCH: GALAPAGOS TEREBRIDAE 545 FicureE 10. Terebra maculata roosevelti Bartsch and Rehder. Hypotype no. 13228, Califor- nia Academy of Sciences, Department of Geology, Type Collection. Binner’s Cove, Socorro Island, Revillagigedo Islands, Mexico. Dorsal view. Length 23 mm., width 7 mm. Ficure 11. JTerebra maculata roosevelti Bartsch and Rehder. Same specimen as figure 10. Apertural view. Ficure 12. Terebra maculata maculata (Linnaeus). Oahu, Hawaii. Collected by C. S. Weaver. Bratcher and Burch Collection no. 522. Dorsal view. Length 19 mm., width 5.7 mm. Ficure 13. Terebra maculata maculata (Linnaeus). Same specimen as figure 12. Apertural view. Type Locatity. “In O. Africae, Asiae.” DistTRIBuTION. Cocos Island, Costa Rica, and Indo-Pacific. DEscrRIPTION. Shell large, sturdy; color varies from shiny dark cream through orange buff to almost melanistic coloration, all ornamented with irregu- lar, squarish, brown markings on subsutural band and brown dots, also irregular, on whorls anterior to subsutural band; nucleus of three inflated, shining opaque whorls; whorls flat to slightly convex, shouldered anterior to subsutural band which is set off by deeply channeled suture and clearly defined spiral groove; early sculpture of regular, straight, slightly oblique axial ribs on whorl and subsutural band; interspaces appear smooth; axial ribs fade out on later whorls until only axial growth striae are evident; spiral groove marking subsutural band also disappears though color pattern of band continues through body whorl; body whorl of medium length with rounded periphery; color of body whorl bro- 546 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. ken by light spiral bands which are crossed by light axial bands, giving body whorl a tessellated appearance in most specimens; aperture elongate, sturdy; columella short with one oblique fold, cream in color; anterior canal slightly recurved. Discussion. This is the largest species of Terebra with specimens measuring over 300 mm. in length. Melanistic coloration mentioned above is rare with most specimens being a dark cream color. We have examined seven specimens of 7. maculata maculata in this study. American Museum of Natural History no. 14613, three specimens, were collected dead and had suffered natural damage as by wave action or predator and showed subsequent unusual distortion with elongation of body whorl. One specimen showed damage on three occasions. We have not encountered this damage to specimens in other areas, but this may be because of collectors selecting only better specimens. Four specimens in the col- lection of the U. S. National Museum, no. 568100, did not show the above- mentioned damage. All specimens were collected at Cocos Island, Costa Rica. The difference between this species and the subspecies Terebra maculata roosevelti Bartsch and Rehder (1939) is that the subspecies retains definite axial ribs in adult specimens, while 7. m. maculata loses the ribs on later whorls. Terebra m. maculata also reaches a much greater size. Terebra maculata roosevelti Bartsch and Rehder. (Figures 10, 11.) Terebra (Subula) roosevelti BartscH and REHDER, Smithsonian Miscell. Coll., vol. 98, no. 10)(Publ, 3535), pa Le ple 1) tie. 6; June) 13591939: Type. Holotype no. 472534, United States National Museum. TYPE LocALity. “It was dredged on Socorro Island, Mexico, in 7—8 fathoms on sandy bottom, off the landing beach toward the rocky point forming the east side of the cove.” DIsTRIBUTION. We have examined specimens of this species from the fol- lowing collection stations in addition to the type lot: California Academy of Sciences, locality 34112, Binner’s Cove, Socorro Island, Revillagigedo Islands, Mexico, in 20 meters (11 fathoms). Allan Hancock Pacific Expedition collection station, 922-39, Braithwaite Bay, Socorro Island, Revillagigedo Islands, Mexico, in 18 to 37 meters (10 to 20 fathoms). This is currently considered to be a rare shell, and we have encountered no specimens except from Socorro Island, Mexico, the type locality of this species. DEscRIPTION. Shell medium large, sturdy; color shiny dark cream to orange- buff, ornamented with irregular squarish brown markings on subsutural band and small brown dots, also irregular, on whorls anterior to subsutural band; nucleus three inflated, shining opaque whorls; whorls flat to slightly convex, shouldered anterior to subsutural band which is set off by deeply channeled suture and Vor. XXXVII] BRATCHER AND BURCH: GALAPAGOS TEREBRIDAE 547 clearly defined spiral groove; early sculpture of regular, straight, slightly oblique axial ribs on whorl and subsutural band, ribs becoming less close-set and regular on later whorls, fading out more on subsutural band than on remainder of whorl; interspaces appear smooth; body with rounded periphery and axial ribs con- tinuing to sharp keel which sets off siphonal fasciole; aperture elongate, same color as body whorl with brown markings showing through; columella short with one oblique fold, yellowish cream in color; anterior canal slightly recurved. Discussion. Terebra maculata roosevelti is indistinguishable from specimens of T. m. maculata (Linnaeus, 1758) of the same size, including the nucleus under magnification. The difference between the species and the subspecies is that the former grows to a much greater size and loses the axial ribs in the later whorls while the subspecies retains the ribs as do some specimens of 7". strigata Sowerby (1825). We believe this to be a subspecies which has developed during geographic isolation. In even the largest specimens of 7. m. maculata, the shape of the aperture, the columella, and the very sharp keel setting off the siphonal fasciole remain the same as in 7. m. roosevelti. Average size about 50 mm. Terebra frigata Hinds. (Figures 16, 17.) Terebra frigata H1nps, Prov. Zool. Soc. London for 1843, p. 162, issued June, 1844. A new name for Terebra gracilis Gray [in part, Galapagos Islands record only]. [Not. T. gracilis Gray, Proc. Zool. Soc. London for 1834, p. 61, November 25. “Hab. ad Africae oras.” (= T. grayi E. A. Smith, 1877) ] [Not T. gracilis Lea, 1833.] Hunps, Thes. Conch., vol. 1, p. 163, pl. 44, fig. 71, January 15, 1845. Locality same as original. Reeve, Conch. Icon., vol. 12, Terebra. pl. 24, fig. 132, 1860. “Hab. Galapagos Islands (in coral sand) ; Cuming.” [Not a good figure of this species.] Tryon, Man. Conch., fol. 7, p. 26, pl. 7, fig. 26, 1885. [A copy of Reeve’s illustration. ] Terebra galapagina Datu and OcHsNER, Proc. Calif. Acad. Sci., ser. 4, vol. 17, no. 4, p. 100, pl. 2, fig. 2, June 22, 1928. [Fossil.] Holotype no. 2897, California Academy of Sciences, Department of Geology Type Collection. “114 miles northeast of Vilamil, Albemarle Island, Galapagos Group. Probably Pleistocene.” Type. Holotype British Museum (Natural History). Type Locatity. “Galapagos Islands; in six fathoms, coral sand.” DisTRIBUTION. Galapagos Islands, Ecuador. DescRIPTION. Shell small, slender; color shiny white; nucleus of three in- flated, shining, opaque whorls with a constriction between nucleus and first post- nuclear whorl; whorls slightly concave; sculpture of strong, straight, axial ribs extending over entire whorl swelling into elongate nodes anterior to suture which in some specimens gives the appearance of a convex subsutural band set off on the anterior by punctations between the nodes; interspaces slightly narrower than the axial ribs with about five rows of well defined spiral striae which faintly cross ribs. This is a variable species. In some specimens the sculpture is con- 548 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 14. Terebra plicata Gray. Hypotype no. 13230, California Academy of Sciences, Department of Geology, Type Collection. Locality 23154 (CAS), Espanola (Hood) Island, Galapagos Islands. One end of variability range. Apex lacking. Length 15.6 mm., width 5.1 mm. Ficure 15. Terebra plicata Gray. Tagus Cove, Isabela (Albemarle) Islands, Galapagos Islands. Bratcher and Burch Collection no. 728. Collected by Jacqueline DeRoy. Other end of variability range. Length 48 mm., width 12.3 mm. Ficure 16. Terebra frigata Hinds. Tagus Cove, Isabela (Albermarle) Island, Galapagos Islands. Los Angeles County Museum of Natural History, Hancock Collection no. 118i (324-35). Juvenile specimen, end of variability range. Length 8.6 mm., width 2.8 mm. Ficure 17. Terebra frigata Hinds. Hypotype no. 13229, California Academy of Sciences, Department of Geology, Type Collection. Post Office Bay, Santa Maria (Charles; Floreana) Island, Galapagos Islands. Typical form for adult of this species. Length 18.3 mm., width 4.4 mm. sistent throughout. In others, the early sculpture consists of more numerous small axial ribs crossed by spiral lines, which break into tiny nodes. Specimens with this sculpture, mostly immature, look like a different species of Terebra. Medium length body whorl; anterior to periphery of body whorl, the axial ribs are broken into small nodes by spiral grooves, usually three to five. Body whorl is rather squared off at base; aperture is elongate; outer lip thin with shadow of ribs showing through; columella white and shiny, straight with one plication; parietal wall with very thin callus; inconspicuous siphonal fasciole. Vot. XXXVII] BRATCHER AND BURCH: GALAPAGOS TEREBRIDAE 549 Discusston. This is the second-most-abundant species of Terebra in the Galapagos collections we have examined, having been taken at 19 collecting sta- tions of the Allan Hancock Pacific expeditions and 10 collecting stations of the California Academy of Sciences at depths from 4 to 82 meters (2 to 45 fathoms). Though it was collected at many stations, the lots contained fewer individuals than some of the large lots of Terebra plicata Gray (1834). Most of the speci- mens were live taken, and the most abundant form of this variable species matches well with the holotype of 7. galapagina Dall and Ochsner, described from beds of Pleistocene age in the Galapagos Islands. In addition to the above men- tioned collections, we have examined specimens of T. frigata in the DeRoy collection and the Bratcher and Burch collection. Average size about 20 mm. Largest specimen examined is 29.9 mm. in length and 6.9 mm. in diameter. Terebra plicata Gray. (Figures 14, 15, 18, 19, 20.) Terebra plicata Gray, Proc. Zool. Soc. London for 1834, p. 61, issued November 25, 1834. [No illustration, no locality.] Hinps. Thes. Conch., vol. 1, Terebra, p. 165, pl. 43, fig. 61, January 15, 1845. “Guayaquil; in seven fathoms, sandy mud: Cuming.” Reeve, Conch. Icon., vol. 12, Terebra, pl. 17, fig. 76, 1860. Same locality as preceding reference. Tryon, man. Conch., vol. 7, p. 24, pl. 7, fig. 20, 1885. Same locality as given by Hinds. Type. Holotype in British Museum (Natural History). Type Locatity. No locality originally cited. ‘‘Guayaquil, in seven fathoms, sandy mud: Cuming.” (Hinds, 1845). DISTRIBUTION. This is the most abundant species in the Galapagos material examined by us. It has been taken, sometimes in large lots, at 31 collecting stations of the Allan Hancock Pacific Expeditions and numerous California Academy of Sciences locations in depths from intertidally to 110 meters (60 fathoms). The type locality is Guayaquil, Ecuador, but at this writing we have examined no specimens of this species except those from the Galapagos Islands. It has been conspicuously absent from material we have examined from other offshore islands included in this study. DESCRIPTION. Shell medium sized, color from flesh to dark brick red, the red, like periostracum or a wash of varnish, is not removable with lye or purex; nucleus of three glassy, lavender, rather flat whorls. In most specimens ex- amined the first four postnuclear whorls also are lavender. Whorls flat with prominent subsutural band set apart by well defined suture and subsutural groove; sculpture variable and consisting of axial ribs, straight on the early whorls, flexuous on the later ones, with interspaces being wide in some specimens and narrow in others; interspaces crossed by spiral grooves, usually three in number, but occasionally only one, which sometimes cross ribs strongly, giving them a beaded appearance. In the majority of individuals examined, the spiral 550 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 18. Terebra plicata Gray. Hypotype no. 13231, California Academy of Sciences, Department of Geology, Type Collection. From Locality 27232 (CAS), Conway Bay, Santa Cruz (Indefatigable) Island, Galapagos Islands. Length 25.6 mm., width 7 mm. Typical form. Ficure 19. Terebra plicata Gray. View of middle whorls. Ficure 20. Terebra plicata Gray. Specimen showing variation in sculpture in comparison with specimens illustrated in figures 14 and 18. grooves cross the ribs weakly, if at all. Spiral grooves are clearly punctate in some specimens, microscopically in others. In the early whorls the subsutural band is nodulous at rib endings. On later whorls the nodes become elongate or become mere flexuous plications. In the latter case the band becomes flat and less prominent. Body whorl is of medium length with axial ribs fading somewhat at periphery. Spiral lines become more numerous at periphery and continue to the sharp keel posterior to the siphonal fasciole. Aperture elongate; outer lip thin with rib pattern showing through; columella ivory colored, straight with two plications; siphonal fasciole well developed; anterior canal broad, recurved. Discussion. Specimens with a dark beige to brick red varnish-like coating have been taken in quantities at various collecting stations from intertidal to deep water and on many types of bottom. Some of these lots have a few normal flesh-colored specimens among them. VoL. XXXVII] BRATCHER AND BURCH: GALAPAGOS TEREBRIDAE 551 FicurE 21. Terebra strigata Sowerby. Smooth form. Isabela (Albemarle) Island, Gala- pagos Islands. Los Angeles County Museum of Natural History, Hancock Collection no. 1185 (146-34). Length 87.3 mm., width 27.6 mm. Ficure 22. Terebra strigata Sowerby. Ribbed form. Baltra (South Seymour) Island, Galapagos Islands. Los Angeles County Museum of Natural History, Hancock Collection no. 1186 (790-38). Length 80.8 mm., width 23 mm. Ficure 23. Terebra strigata Sowerby. Without brown axial stripes. Dredged by shrimp fishermen of Guaymas, Mexico. No. 92105, Purdy Collection, San Diego, California. Length 79.2 mm., width 21.6 mm. Ficure 24. Terebra ornata Gray. Hypotype no. 13232, California Academy of Sciences, Department of Geology, Type Collection. Venedig, northern Santa Cruz (Indefatigable) Island, Galapagos Islands. Collected by Carmen Angermeyer. Length 67 mm., width 16.6 mm. Average size about 35 mm. Largest specimen examined 70.7 mm. in length, San Diego Museum of Natural History. Terebra strigata Sowerby. (Figures 21, 22, 23.) Acus zebra HumpureY, Mus. Colonnianum, p. 31, 1797. “India.” [Invalid for Nomenclature, International Commission on Zoological Nomenclature: Opinion 51.] 552 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Terebra strigata SowERBy, Catalogue of Shells in Collection of Earl of Tankerville, Ap., p. XXIII, no. 1984, 1825. [No fig.] Hzinps, Thes. Conch., vol. 1, pt. 5. Terebra, p. 151 (bis), pl. 41, fig. 10, 1845. “West coast of America between Panama and Realejo.” Buccinum elongatum Woon, Index Test., Suppl., p. 13, pl. 4, fig. 25, 1828. “Tndia.” [Locality erroneus according to Hinds, 1845.] Terebra flammea Lamarck, Lesson, IIlustr. Zool., pl. 48, 1832. “vit sur les cotes de listhme de Panama.” [Not “Hab in mare Antillarum? .. . habite le golfe des Antilles.” ] Not Terebra flammea Lamarck, Hist. Nat. Anim. s. Vert., vol. 7, p. 284, 1822. Terebra zebra KiEnER, Spéc. Gén. et Icon. Coq. Viv., vol. 8, Terebra, p. 5, no. 2, pl. 3, fig. 5, 1839. “Les cotes de l’isthme de Panama.” [Not “habite le golfe des Antilles.” ] Type. Location unknown to present authors. TYPE Locality. “Panama.” DisTRIBUTION. In addition to the locality records cited above, Reeve (1860) gives as locality, “Panama, Galapagos, and Philippine Islands; Cuming, “Mo- luccas etc.”; and Hidalgo (1904, p. 348) repeated the Philippine citation with doubt as to its correctness. Burch (1964) states, “We have not seen Terebra strigata from any area other than the western Americas, where it ranges from Mexico to Peru.” Grant and Gale (1931) record this species as a fossil in the Pleistocene of Ecuador and Mexico. DeEscRIPTION. Shell large and heavy; color dull cream ornamented, after about the seventh postnuclear whorl, with axial stripes of brown or black (rarely with stripes of paler cream); nucleus of two and a half beige, translucent, slightly convex whorls; whorls flat with flat subsutural band clearly defined by a spiral groove; early sculpture of very slightly curved, close-set axial ribs on both whorl and subsutural band which sometimes fade so that later whorls are smooth. Where sculpture continues through body whorl of mature specimens, ribs become small, low, and close-set, flexuous and less regular than in the early whorls; aperture elongate, cream colored with stripes of body whorl usually showing through; columella straight, simple, pale cream color: anterior canal straight, parietal wall with very thin callus in fully adult specimens. Discussion. Though most specimens have brown axial markings, we have examined four specimens which at first glance appeared to have no stripes at all, but on closer examination revealed a very faint axial stripe of slightly lighter color. One of these specimens, trawled by shrimp fishermen of Guaymas, Mexico, is in the Purdy collection, San Diego, California. Another was found tracking in sand at 6.5 to 8 meters (20 to 25 feet) at Venado Island near Guaymas, Sonora, Mexico, by Laura Shy and is in the Shy collection at Westminster, California. The others were collected at Manzanillo, Mexico, 8 to 24 meters (4.5 to 13 fathoms), by Lawrence Thomas of Morro Bay, California. This species has been placed by Grant and Gale (1931) and Keen (1958) in Terebra, s.s., a subgenus characterized by the last whorls being smooth except Vor. XXXVII] BRATCHER AND BURCH: GALAPAGOS TEREBRIDAE 553 for the subsutural band. While many specimens have smooth last whorls, the majority of specimens examined by us have axial sculpture which continues through the body whorl. Both smooth and ribbed forms have been collected from Guaymas, Sonora, Mexico, to the Galapagos Islands. Average size about 90 mm. Largest specimen examined by us 143.8 mm., Lawrence Thomas collection, Morro Bay, California. Terebra ornata Gray. (Figure 24.) Terebra ornata Gray, Proc. Zool. Soc. London for 1834, p. 62, November 25, 1834. HInps, Proc. Zool. Soc. London for 1843, p. 160, issued June, 1844. Hrinps, Thes. Conch., vol. 1, pt. 5, Terebra, p. 152 (bis), no. 6, pl. 42, fig. 34, 1844. “Galapagos Islands; in five to seven fathoms, coral sand: Cuming. Panama in seven fathoms, mud.” Hanna and HERTLEIN, Proc. Calif. Acad. Sci., Ser. 4, vol. 30, no. 3, p. 71, pl. 6, fig. 1961. Earlier records cited. Not Buccinum ornatum Martyn, Universal Conch., vol. 3, pl. 92, fig., 1786. [Referable to the genus Terebra.] [Invalid for nomenclature. International Commission on Zoological Nomenclature, Opinion 456.] Type. Holotype, in British Museum (Natural History). TypE Locality. No locality originally cited. ‘Galapagos Islands; five to seven fathoms, coral sand; Cuming. Panama; seven fathoms, mud; H.,” cited by Hinds (1844). DIsTRIBUTION. Baja California, Mexico to the Galapagos Islands, intertidal to 96 meters (45 fathoms). DEscrRIPTION. Shell large, moderately heavy, broad apical angle; color dull white to pale cream ornamented with spiral rows of rather square dark brown spots, three rows on body whorl plus one row on subsutural band; whorls shouldered anterior to deeply constricted groove which defines the subsutural band; subsutural band nodulated on early whorls, the nodes fading completely or to slight swellings on later whorls; early whorls with very faint axial ribs sometimes swelling into nodes at the posterior end; short body whorl; aperture elongate and the same color as body whorl with the brown spots showing through; columella short, often more orange in color, with two plications, the posterior one faint and the anterior one very sharp and placed at the extreme anterior end; siphonal fasciole elongate; anterior canal recurved. Discussion. We have examined specimens of Terebra ornata from Santa Cruz (Indefatigable) Island, California Academy of Sciences locality no. 38898 and from South Baltra (South Seymour) Island, Galapagos, 9 meters (5 fathoms), Allan Hancock Pacific Expeditions collecting station no. 173-34. Average size about 60 mm. Largest specimen examined 93 mm., Purdy col- lection, San Diego, California. 554 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Terebra robusta Hinds. (Figure 25.) Terebra robusta Hinps, Proc. Zool. Soc. London for 1843, p. 149, issued June, 1844. “Hab. West coast of America, between 8° 57’ and 21° 32’ north latitude; namely Panama, Gulf of Nicoya, Gulf of Papagayo, and San Blas; in from four to eighteen fathoms, sandy mud.” Huinps, Zool. Voy. Sulphur, Moll., pl. 2, p. 32, October, 1844. [Same localities cited in previous reference.] Hinps, Thes. Conch., vol. 1, Terebra, p. 153 (bis), pl. 42, fig. 35, 1845. [Same localities cited in previous reference.] DrsHayes, Proc. Zool. Soc. London for 1859, p. 307, 1859. “Panama; Golfe de Nicoya, Golfe de Papayo; San Blas.” Reeve, Conch. Icon., vol. 12, Terebra, sp. 10, pl. 3, fig. 10, 1860. [Same localities origi- nally cited.] Morcu, Malakazool. Blatter, Bd. 7, p. 105, 1860. “Realejo.” [Nicaragua.] Tryon, Man. Conch., vol. 7, p. 11, pl. 2, fig. 16 (Hinds figure), 1885. “W. Coast of Central America.’”’ GRANT and GALE, Mem. San Diego Soc. Nat. Hist., vol. 1, p. 465, 1931. “Pleistocene: Lower Quaternary at Magdalena Bay, Lower California, Mexico (Jordan). Recent: Gulf of California, Mexico, to Panama.’ HERTLEIN and STRONG, Bull. Amer. Mus. Nat. Hist., vol. 107, art. 2, p. 214, 1955. “Off Cape Pasado, Ecuador,” also localities in that area. “Range: Guaymas, in the Gulf of California to the Rio Esmeraldas, Ecuador. Galapagos Islands.” Keren, Seashells of Tropical West America, 1958, p. 489, no. 948. “Guaymas, Gulf of California to Ecuador and the Galapagos Islands.” HANNA and HeErTLEIN, Proc. Calif. Acad. Sci., ser. 4, vol. 30, no. 3, p. 72, pl. 6, figs. 3, 7, 8 and pl. 7, fig. 1, Aug. 1961. “Guaymas, Sonora, Mexico, in the Gulf of California, to the Rio Esmeraldas, Ecuador, and the Galapagos Islands, in 7 to 33 meters, (4 to 18 fathoms). Also Pliocene to Recent.’’ Otsson, Paleont. Research Inst., Ecuadorian Neogene Mollusks, 1964, p. 76, pl. 10, fig. 9. “Angostura formation; Telembi Cayapas.” Not Terebra robusta Hinds, Gass, Trans. Amer. Philos. Soc., vol. 15, p. 224, 1873, renamed Terebra gabbi Dall, 1895. Caribbean, Miocene. Maury, Bull. Amer. Paleo., vol. 5, no. BE), SECs to joo, BH (AUG), Wily. Terebra lingualis Htnps, Proc. Zool. Soc. London for 1843. p. 153, issued June, 1844. Terebra loroisii GUERIN-MENEVILLE, Mag. de Zool., 1854, p. 218, pl. 4, fig. 5, 1854. Not Terebra loroisii DESHAYES, Proc. Zool. Soc. London for 1859. “Hab... ?” 1859. Terebra insignis DESHAYES, Journ. Conchyl., vol. 6, p. 70, pl. 3, fig. 2. 1857. Terebra macrospira Li, Bull. Geol. Soc. China, vol. 9, no. 3, p. 273, pl. 8, fig. 66, 1930. “Brought up by marine dredge from depths varying from 10 ft. to 40 ft. in the mud at the mouth of the Rio Grande near La Boca about one mile from the mainland in Panama ” Bay. Type. Holotype, in British Museum (Natural History). TYPE LocALITY. “Hab. West coast of America, between 8° 57’ and 21° 32’ north latitude; namely at Panama, Gulf of Nicoya, Gulf of Papagayo, and San Blas; in from four to eighteen fathoms, sandy mud.” ‘Panama’ selected as type locality by Hertlein and Strong (1955). DISTRIBUTION. Outer coast of Baja California, Mexico to Ecuador and the Galapagos Islands, intertidal to 90 meters (50 fathoms). Recorded as a fossil in Neogene formations of Panama by Li (1930), from the Quaternary at Magda- lena Bay, Baja California, Mexico by Jordan and from the Neogene of Ecuador by Olsson (1964). DescripTIon. Shell large; color whitish to beige decorated with brown spots Vor. XXXVII] BRATCHER AND BURCH: GALAPAGOS TEREBRIDAE 555 which coalesce to form axial lines; whorls rather flat, subsutural band convex on early whorls, more flattened on later ones, set off by well marked suture and constricted subsutural groove; heavy early sculpture of many tiny axial riblets ending in small nodes anterior to subsutural band and larger ones on the band itself; later sculpture in mature specimens smooth except for subsutural band; body whorl with same coalesced brown spots posterior to periphery and one row of squarish spots anterior to periphery; aperture elongate; outer lip with color pattern showing through faintly; columella slightly curved, with one plication; thin callus on parietal wall; anterior canal very short; shiny siphonal fasciole. Discussion. This species varies greatly in apical angle, often within a single population, by some individuals being slender while others are obese. In some specimens the brown spots coalesce into stripes, while in others they do not. We have examined specimens of Terebra robusta taken in the Galapagos Islands by the California Academy of Sciences, the Allan Hancock Pacific Expeditions, and by the DeRoys who live in the Galapagos Islands. Average size about 60 mm. Largest specimen examined 140 mm. Terebra crenulata (Linnaeus). (Figure 26.) Buccinum crenulatum LINNAEUS, Syst. Nat., ed. 10, p. 741, 1758. Ref. to Gualtieri, Index Test., pl. 57, fig. L, 1742; Argenville, Conchyl., pl. 14, fig. Y, 1742. Dodge, Bull. Amer. Mus. Nat. Hist., vol. 111, Art. 3, p. 221, 1956. Buccinum luteola MARTYN, Universal Conchologist, vol. 3, pl. 92, figure at bottom. [Invalid for Nomenclature, International Commission on Zoological Nomenclature, Opinion 456.] CHENU, Universal Conchologist, in Bibl. Conchyl., premiére ser., tome 2, p. 25, pl. 33, fig. 1, 1845. “Chine.” Buccinum varicosum GMELIN, Linn. Syst. Nat., ed. 13, pars. VI, p. 3505, 1791. “Habitat .. .” Ref. to Seba, Mus., 3.t.56.f.17. Acus coronata HumpuHREY, Mus. Calonnianum, p. 31, 1797. [Invalid for Nomenclature, Inter- national Commission on Zoological Nomenclature, Opinion 51.] Terebra maculata PERRY, Conchology, pl. 16, fig. 2, 1811. [Not the locality “The Brazils and the West Indies.” ] Not Buccinum maculatum Linnaeus, 1758, p. 741. [A Terebra.] Terebra fimbriata DresHAYES, Journ. de Conchyl., vol. 6, p. 71, pl. 5, fig. 1, 1857. [Cotypes (2) in British Mus. (Nat. Hist.) “Patria inconnue.” Not found in Deshayes’ collection at the Ecole des Mines, Paris. ] Terebra interlineata DresHAyEs, Proc. Zool. Soc. London for 1859, p. 277, issued between July and October, 1859. “Hab. Les Iles Sandwich.” Not figured. Holotype in British Museum (Natural History). Terebra crenulata Linnaeus var. booleyi Mretvmt and Sykes, Proc. Malacol. Soc. London, vol. 3, p. 42, pl. 3, fig. 5, April, 1898. Andaman Islands. Type. Holotype probably in Linnaean Society collection in London (Dodge). Type Locatity. “Habitat in O. Africano.”’ DIsTRIBUTION. Clarion and Socorro islands, Revillagigedo Islands, Mexico; Red Sea, and Indo-Pacific. 556 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. DescripTIoN. Shell large, shining; flesh colored or light grey, often with darker blotches. After about the 12th postnuclear whorl two rows of orange- brown dots decorate each whorl and orange-brown axial lines ornament the inter- spaces between the nodes on the subsutural band; body whorl with a third row of dots anterior to the rounded periphery continuing into the aperture; whorls flat, with convex subsutural band; nucleus of two convex, shining whorls; early whorls white with sculpture of sharp axial ribs (narrower than interspaces) and nodose subsutural band set apart from remainder of whorl by suture and microscopically punctate subsutural groove; later sculpture with axial ribs continuing to periph- ery of body whorl in some individuals, fading out in others; body whorl smooth anterior to periphery except for axial growth striae; aperture quadrate; outer lip thin with orange-brown dots showing through; columella straight with one weak plication; parietal wall with thin wash of callus; well developed, striate siphonal fasciole with sharp keel; anterior canal short, broad recurved. Discussion. This species exhibits extreme variations in form and sculpture. The form described by Deshayes (1857) as Terebra fimbriata does not have the large and prominent nodes on the subsutural band and is generally more slender than the more common form with which it usually can be found in all areas where the species occurs. The form described as T. interlineata by Deshayes (1859) continues the axial sculpture of ribs to adult size. This latter form, which the Clarion Island specimens exhibit, is not uncommon in Hawaii where it is found with the more common form. We are unable to separate the Clarion Island material from this form of 7. crenulata collected in Hawaii. Two speci- mens from Clarion Island, dredged at 73 meters (40 fathoms), California Acad- emy of Sciences locality 49692, appear identical to the holotype of the form described by Deshayes as T. interlineata. One dead and badly damaged speci- men was dredged off Cape Rule, Socorro Island, by the Allan Hancock Pacific Expedition. These, along with another specimen in the Los Angeles County Museum of Natural History collection and one in the American Museum of Natural History collection, both from Clarion Island, are the only specimens of T. crenulata taken in the eastern Pacific which we have examined. The largest eastern Pacific specimen examined is 57.1 mm., though they are much larger in the Indo-Pacific area. Terebra armillata Hinds. (Figure 27.) Terebra armillata Hinps, Proc. Zool. Soc. London for 1843, p. 154, issued June, 1844. [No fig.] Hrnps, Thes. Conch., vol. 1, Terebra, p. 173, pl. 43, fig. 49, 1845. KrEn, Veliger, vol. 8, no. 4, p. 273, pl. 47, fig. 18, 1966. (Photograph of syntypes.) Not Terebra armillata Hinps of Menke, Zeitschr. f. Malakozool., Jahrg. 8, no. 3, p. 34, 1851; Not of Reeve, Conch. Icon., vol. 12, Terebra species 72 (in part), pl. 16, fig. 72a (only), 1860. —— Vor. XXXVII] BRATCHER AND BURCH: GALAPAGOS TEREBRIDAE oat on ~ Ficure 25. Terebra robusta Hinds. Punta Colorado, Guaymas, Sonora, Mexico. Bratcher and Burch Collection no. 729. Collected by Twila Bratcher. Length 90 mm., width 17.5 mm. Ficure 26. Terebra crenulata (Linnaeus). Hypotype no. 13233, California Academy of Sciences, Department of Geology, Type Collection. Clarion Island, Revillagigedo Islands, Mexico. Length 66.3 mm., width 12.5 mm. Ficure 27. Terebra armillata Hinds. Guaymas, Sonora, Mexico. Collected by G. Jacobs. Bratcher and Burch Collection no. 90. Length 40 mm., width 9.1 mm. Ficure 28. Terebra berryi Campbell. Hypotype no. 1184, Los Angeles County Museum of Natural History, Type Collection. Wafer Bay, Cocos Island, Costa Rica. Length 16.8 mm., width 4.2 mm. FicureE 29. Nucleus, same shell as figure 28. Terebra albicostata ADAMS and REEVE, Zool. Voy. Samarang, Moll., pt. 2, p. 30, pl. 10, fig. 21, 1850. “Hab. China Sea.” [Locality erroneous. | Terebra marginata DrESHAYES, Journ. Conchyl. vol. 6, p. 86, pl. 4, fig. 8, 1857. “Hab. L’embouchure de la Gambie.’ ’ [ Locality erroneous. ] Type. Syntypes, British Museum (Natural History). TyPeE Locality. “Hab. abundant in various localities on the west coast of America between Panama and the Bay of Magdalena in Lower California, in from five to thirteen fathoms: also at the Galapagos, in ten fathoms: chiefly in sandy situations.” 558 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. DIsTRIBUTION. Magdalena Bay, west coast of Baja California, Mexico, to northern Peru and the Galapagos, intertidal to 73 meters (40 fathoms). DESCRIPTION. Shell medium in size; color usually light brown with cream colored nodes on subsutural band but sometimes is light tan or yellowish; whorls slightly convex with convex subsutural band set off by well defined suture and constricted subsutural groove; sculpture quite consistent throughout of narrow, rather sharp, flexuous ribs with much wider interspaces; interspaces divided by spiral grooves (usually about five to eight) which faintly cross axial ribs; body whorl of medium length, often with a faint light stripe at periphery where axial ribs end. In most specimens, axial sculpture becomes obsolete anterior to periph- ery; aperture quadrate; columella curved with two plications, the posterior one continuing as a sharp keel to the well developed striate siphonal fasciole; anterior canal broad, short, curved. Discussion. In some individuals the subsutural band is decorated with large, round, pearl-like nodes; in others they are small and less conspicuous. In still others the nodes are more like elongate ribs. Average specimen about 40 mm. Largest specimen examined 67 mm. Terebra berryi Campbell. (Figures 28, 29.) Terebra (Strioterebrum) berryi CAMPBELL, Veliger, vol. 4, no. 1, p. 26, figs. 5, 6, July 1, 1961. Terebra berryi CAMPBELL, DUSHANE, and SpHON, Veliger, vol. 10, no. 3, p. 244, January 1, 1968. Puertecitos, Gulf of California. Type. Holotype, no. 12352, California Academy of Sciences, Department of Geology, Type Collection. TYPE LOCALITY. ‘“‘Puertecitos, Baja California.”’ [ Mexico. | DisTRIBUTION. East coast of Baja California, Mexico to Guatamala, inter- tidal to 36 meters (20 fathoms). DeEscriPTION. Shell medium, color marbled pale lavender-gray and brownish; whorls very slightly convex; convex subsutural band set off by well defined suture and impressed groove which in some specimens becomes a sharply cut groove; sculpture of sharp flexuous axial ribs which swell into nodes on the subsutural band; ribs much narrower than interspaces on early whorls, occasion- ally more closely placed in later whorls, spiral grooves, about four in addition to the subsutural groove, mark the interspaces and sometimes faintly cross axial ribs; sculpture continuing on body whorl, but becoming less well defined from periphery to anterior canal which is marked with fairly deep spiral grooves; aperture elongate; outer lip thin with marbled color showing through; columella curved with one plication; well developed siphonal fasciole with sharp posterior keel; anterior canal broad, short, curved; aperture moderately laminated in adult specimens. Vor. XXXVIT] BRATCHER AND BURCH: GALAPAGOS TEREBRIDAE 559 Discussion. Color varies among the specimens examined. Some are almost entirely lavender-grey, while some are quite brownish, though all have a more or less marbled appearance. The sculpture also varies in number of ribs and width of interspaces. On one specimen taken at Cocos Island, Costa Rica, the ribs are not sharp and clearly defined, and the spiral sculpture is more noticeable than the axial sculpture. Largest specimen examined 57.1 mm. Terebra litorea Dall and Ochsner. (Figure 30.) Terebra litorea Dati and Ocusner, Proc. Calif. Acad. Sci., ser. 4, vol. 17, no. 4, p. 101, pl. 2, fig. 3, June 22, 1928. Type. Holotype, California Academy of Sciences, Department of Geology, Type Collection no. 2904. TYPE LOCALITY. “One and one-fourth miles northeast of Vilamil, Albemarle Island, Galapagos Group. Probably Pleistocene.” DISTRIBUTION. Known only from the Galapagos Islands. DEscrIPpTION. Shell large; color buff in fossil state; whorls flat, slightly con- stricted at weak spiral groove which sets off convex subsutural band; early sculpture weak axial ribs with nodulous subsutural band; later sculpture of flexuous axial ribs which cross subsutural band and distinct suture, ribs con- tinuing over body whorl; aperture elongate; columella with two strong plications, the posterior of which is carried outside the aperture as a sharp keel posterior to the siphonal fasciole. Discussion. This species was described as fossil, probably of Pleistocene age, and has not been seen by us as a Recent species. Length of the holotype is 56 mm. Terebra albemarlensis Dall and Ochsner. (Figure 31.) Terebra albemarlensis DALL and OcHsNER, Proc. Calif. Acad. Sci., ser. 4, vol. 17, no. 4, p. 99, pl. 2, fig. 1, June 22, 1928. TyprE. Holotype, California Academy of Sciences, Department of Geology, Type Collection no. 2894. TYPE LOCALITY. “One and one-fourth miles northeast of Vilamil, Albemarle Island, Galapagos Islands. Probably Pleistocene.” DISTRIBUTION. Known only from the Galapagos Islands. DEscrRIPTION. Shell large, whorls flat, slightly shouldered anterior to suture, with flat subsutural band which is nodulose in early whorls, flat in later ones; early sculpture of axial ribs, which become slightly nodulose at anterior end of 560 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 30. Terebra litorea Dall and Ochsner. Holotype no. 2904, California Academy of Sciences, Department of Geology, Type Collection. Isabela (Albemarle) Island, Galapagos Islands. Pleistocene. Length 56 mm., height of last whorl 21 mm., width 13 mm. Ficure 31. Terebra albemarlensis Dall and Ochsner. Holotype no. 2894, California Academy of Sciences, Department of Geology, Type Collection. Isabela (Albemarle) Island, Galapagos Islands. Pleistocene. Length 85 mm., height of last whorl 23 mm., maximum diameter 15 mm. whorl, crossed by spiral grooves; later sculpture of flat ribs, on both subsutural band and remainder of whorl, crossed by weak spiral grooves; body whorl long, gently rounded at periphery; outer lip thin; aperture elongate; columella twisted, with two plications, the posterior of which forms a keel to siphonal fasciole. Discussion. This species was described as probably Pleistocene from Isabela (Albermarle) Islands, Galapagos. We have seen no specimens of this species except for the type material. Some of the larger specimens of Terebra plicata Gray at the end of the sculpture variability range with flatter subsutural band and more obsolete sculpture somewhat resemble this species, but we have seen none with profile and subsutural band as flat or body whorl as elongate. We Vor. XXXVII] BRATCHER AND BURCH: GALAPAGOS TEREBRIDAE 561 believe the fossil, T. albemarlensis, to be the progenitor of the Recent T. plicata Gray (1834). Hastula albula (Menke). (Figures 32, 33.) Terebra albula MeNKE, Moll. Novae Hollandiae, p. 30, 1843. [No figure. ] Terebra aciculina Lamarck, KIeNER, Spéc. et Gén. Icon. Coq. Viv., Fam. Purpuriféres, Terebra, p. 18 (in part), pl. 7, fig. 136b (only), 1838-1839. “Habite la mer des Indes.” REEVE, Conch. Icon., vol. 12, Terebra, species 121 (in part), pl. 23, fig. 121a (only). 1860. . Not Terebra aciculina LAMARCK, Hist. Nat. anim. s. Vert., vol. 7, p. 290, 1822; not Tryon, Man. Conch., vol. 7, p. 32, pl. 10, fig. 81, 1885. Terebra casta Hinps, Proc. Zool. Soc. London for 1843, p. 156, issued June, 1844. “Hab. Ilo- ilo, Island of Panay, Philippines, at low water.” Hinps, Thes. Conch., vol. 1, Terebra, p. 165, pl. 44, fig. 84, 1844. Holotype in British Museum (Natural History). Terebra eburnea DUNKER, Zeitsch. f. Malakozool., Jahrg. 10, no. 6, p. 96, 1853. “Patria ignota.” [No figure.] (Not of Hinps, Proc. Zool. Soc. London for 1843 (1844), p. 153.) Terebra incolor DESHAYES, Proc. Zool. Soc. London for 1859, p. 283, issued between July and October, 1859. ‘Hab. Iles Philippines.” Holotype in British Museum (Natural History). Terebra bipartita DESHAYES, Proc. Zool. Soc. London, for 1859, p. 284, issued between July and October, 1859. [No figure.] “Hab. Iles Sandwich.” Holotype in British Museum (Natural History). Not Terebra bipartita SowERBY, Quart. Journ. Geol. Soc. London, vol. 6, p. 47, 1849. San Domingo. Tertiary. Not Terebra bipartita Goutp, Proc. Boston Soc. Nat. Hist., vol. 7, p. 330, September 1860. Terebra philippiana DESHAYES, Proc. Zool. Soc. London for 1859, p. 289, issued between July and October, 1859. [No figure.] ‘Hab. Iles Marquises?” Holotype in British Museum (Natural History). Not Strioterebrum pedroanum philippianum Dati, U.S. Nat. Mus., Bull. 112, p. 67, February 24, 1920. “San Pedro, California, to head of Gulf of California.” A new name for Terebra simplex Carpenter, 1865, not 7. simplex Conrad, 1830. Holotype in British Museum (Natural History). Terebra (Hastula) casta, nov. var. natalensis E. A. Smiru, Proc. Malacol. Soc. London, vol. 5, no. 6, p. 360, October 31, 1903. [Not figured.] “Hab. Umkomaas, Natal.” Terebra medipacifica Pirspry, Proc. Acad. Nat. Sci. Philadelphia, vol. 72, p. 308, pl. 12, figs. 8, 9, 10; issued January 7, 1921. “Off Honolulu, 6 to 8 fathoms.” Terebra medipacifica melior Pirspry, Proc. Acad. Nat. Sci. Philadelphia, vol. 72, p. 308, pl. 12, fig. 11; issued January 7, 1921. “Kaneohe Bay, Oahu.” [Hawaii.] Hastula gnomon Keen, Trans. San Diego Soc. Nat. Hist., vol. 10, no. 2, p. 47, pl. 4, fig. 11, December 30, 1943. Loc. 2121 (LSJU), “California, Kern Co., Caliente Quadrangle, near center of southwest quarter of sec. 6, T. 29 S., R. 30 E., Mount Diablo B.L.M., in small gully close to terrace contact. Stratigraphic horizon: lowermost part of Round Mountain silt.’ Miocene. Holotype no. 7536, Stanford University Paleontology type collection. Not Terebra gnomon JreFrreys, “Paetel, Catal. Conchyl—Sammlung. Abt. 1, p. 251, 1888.” Ref. to Proc. Zool. Soc. London, 1878, p. 412. “Oc. atl.” [Jeffreys’ reference concerns Atretia gnomon Jeffreys, a brachiopod. ] CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Ficure 32. Hastula albula (Menke). Hypotype no. 1183, Los Angeles County Museum of Natural History. Socorro Island, Mexico. Hancock Collection no. 291.34. Length 12.3 mm., width 3.6 mm. Ficure 33. Hastula albula (Menke). Kailua Bay, Oahu, Hawaii. Collected by C. S. Weaver. Bratcher and Burch Collection no. 530. Length 12.5 mm., width 3.9 mm. TypeE Locatity. ‘Hab. in litore occidentali.”’ DISTRIBUTION. East coast of Africa through the Indian and Pacific oceans to Clarion and Socorro islands, Mexico. Of the 15 lots of Recent shells we examined, all were in the vicinity of Clarion and Socorro islands taken at depths ranging from 7 to 110 meters (4 to 60 fathoms). Recorded as fossil in Neogene formations of California (Keen, 1943). DescriPTION. Shell small to medium, sturdy; shining, with color ranging from very pale beige with a hint of darker stripe to dark chestnut brown with white stripe anterior to suture and at periphery; nucleus conical with four rather flat dark purple whorls with wide apical angle. Nucleus has the appearance of being set into postnuclear whorls at slight angle; first postnuclear whorl of all specimens examined are white; whorls flat; sculpture varies from specimens with straight narrow well defined axial ribs (averaging about 20 per whorl) which cover entire shell from postnuclear whorls through body whorl to specimens with axial ribs showing only as deep plications at suture and fading out on remainder of whorl; body whorl elongate with (in most specimens examined) a light stripe at periphery; aperture elongate; outer lip of medium thickness; columella straight with one microscopic fold; parietal wall with thin callus: anterior canal very short; shiny siphonal fasciole. Vot. XXXVII] BRATCHER AND BURCH: GALAPAGOS TEREBRIDAE 563 Discussion. Of all species of Hastula, H. albula is the most variable in color, sculpture, and size. This variability may show up in a single dredged lot. Thus it has been possible for authors to describe as many as three ‘‘new species” in a single paper (Deshayes, 1859) when all are forms of H. albula. Specimens of H. albula collected in west America are indistinguishable from specimens collected in Hawaii except that the colorless or albino form is not present in west American material examined. Average size about 18 mm. Largest west American specimen examined 27.9 mm. | LITERATURE CITED BarTscH, PAUL, AND HARALD ALFRED REHDER 1939. Mollusks collected on the Presidential Cruise of 1938. Smithsonian Miscellaneous Collections, vol. 98, no. 10, pp. 1-18, pls. 1-5, June 13. Burcu, Ropert DONALD 1964. Notes on the Terebridae of the Philippine Islands. The Veliger, vol. 6, no. 4, pp. 210-218, April 1. BRATCHER, TwItA, and Rospert DONALD BuRCH 1970. Four new terebrid gastropods from eastern Pacific islands. Los Angeles County Museum, Contributions in Science, no. 188, pp. 1-6, figs. 1-8, May 4. DALL, WILLIAM HEALY 1908. Report on the dredging operations off the west coast of Central America to the Galapagos, to the west coast of Mexico, and in the Gulf of California. XIV. The Mollusca and the Brachiopoda. Bulletin of the Museum of Comparative Zoology, Harvard College, vol. 43, no. 6, pp. 205-487, pls. 1-22, October. DALL, WILLIAM HEALY, and WASHINGTON HENRY OCHSNER 1928. Tertiary and Pleistocene Mollusca from the Galapagos Islands. Proceedings of the California Academy of Sciences, Fourth Series, vol. 17, no. 4, pp. 89-139, text figures 1-5, pls. 2-7, June 22. DesHAYES, GERARD PAuL 1857. Description d’espéces nouvelles de genre Terebra. Journal de Conchyliologie, vol. 6, pp. 65-102, pls. 3-5. 1859. A general review of the Genus Terebra, and a description of new species. Pro- ceedings of the Zoological Society of London, part 27, pp. 270-321. (Issued between July and October.) GRANT, Utysses S., IV, and Hoyt RopNnrey GALE 1931. Catalogue of the marine Pliocene and Pleistocene Mollusca of California and adjacent regions. Memoirs of the San Diego Society of Natural History, vol. 1, pp. 1-1036, pls. 1-32, text figures 1-15, November 3. Gray, JoHN EDWARD 1834. Enumeration of the species of the genus Terebra, with characters of many hitherto undescribed. Proceedings of the Zoological Society of London, Part 2, pp. 59- 62, November 25. Hanna, G Darras, AND MERLE C. ISRAELSKY 1925. Contributions to the Tertiary paleontology of Peru. Proceedings of the California Academy of Sciences, Fourth Series, vol. 14, no. 2, (Terebra), pp. 55-56. HERTLEIN, LEO Georce, and A. M. STRONG 1955. Marine mollusks collected during the “Askoy” Expedition to Panama, Colombia, 564 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. and Ecuador in 1941. Bulletin of the American Museum of Natural History, vol. 107: art. 2, pp. 159-318, pls. 1-3. Hmatco, JOAQUIN GONZALES 1904. Catalogo de los molluscos testaceos de las islas Filipinas, Jol6 y Marianas. Revista de la Real Academia de Ciencias, vol. 1, no. 5, (Terebra), pp. 345-349, September. Hinps, RICHARD BRINSLEY 1844. Descriptions of new shells, collected during the voyage of the Sulphur, and in Mr. Cuming’s late visit to the Philippines. Proceedings of the Zoological Society of London, pp. 149-168, 1843 (June, 1844). 1845. Monograph of the genus Terebra. [In] George Brettingham Sowerby. Theasaurus Conchyliorum, or Monographs of Genera of Shells, vol. 1, part 5, pp. 147-190, pls. 41-45. i JorDAN, Eric KNIGHT 1924. Quaternary and Recent molluscan faunas of the west coast of Lower California. Bulletin of the Southern California Academy of Sciences, vol. 23, p. 149. Keen, A. Myra 1943. New mollusks from the Round Mountain silt (Temblor) Miocene of California. Transactions of the San Diego Society of Natural History, vol. 10, no. 2, pp. 25-60, pls. 3-4, figs. 1-5. December 30. 1958. Sea shells of tropical West America; marine mollusks from Lower California to Colombia. Stanford University Press, Palto Alto, California, pp. 1-624, 946 figures, pls. 1-9. 1966. West American mollusk types in the British Museum (Natural History), part 2. Species described by R. B. Hinds. The Veliger, vol. 8, no. 4, pp. 265-275, pl. 47, April 1. Li, CHTH CHANG 1930. The Miocene and Recent Mollusca of Panama Bay. Bulletin of the Geological Society of China, vol. 9, no. 3, pp. 249-296, pls. 4-8, 1 map. LINNAEUS, CAROLUS 1758. Systema naturae per regina tria naturae, Edition 10, Reformata, Holmiae, 1, pp. 1-824. Pitspry, HENry AuGusSTUS, and AXEL ADOLF OLSSON 1941. A Pliocene fauna from western Ecuador. Proceedings of the Academy of Natural Sciences of Philadelphia, vol. 93, pp. 1-79, pls. 1-19, September 9. REEVE, LOVELL AUGUSTUS 1860. Monograph of the genus Terebra. Conchologia Iconica, vol. 12, species 1-155, pls. 1-27; descriptive letter press plus index. RutscuH, ROLF 1934. Die Gastropoden aus dem Neogen der Punta Gabilan in Nord-Venezuela. Zweiter Teil. Abhandlungen der Schweizerischen Palaeontologischen Gesellschaft, vol. 55, no. 1, pp. 89-169, pls. 8, 9, figs. 13-20 in text. (Terebra, pp. 106-109, 153.) Say, THOMAS 1822. Marine shells of the United States. Journal of the Academy of Natural Sciences of Philadelphia, First Series, vol. 2, (Terebra), p. 235. 1827. Descriptions of marine shells recently discovered on the coast of the United States. Journal of the Academy of Natural Sciences of Philadelphia, First Series, vol. 5, (Terebra), pp. 207-208. VoLt. XXXVII] BRATCHER AND BURCH: GALAPAGOS TEREBRIDAE 565 SmitH, EpcAr ALBERT 1903. List of South African marine Mollusca, with descriptions of new species. Pro- ceedings of the Malacological Society of London, vol. 5, no. 6, (Terebra), pp. 360-361, pl. 15. SOWERBY, GEORGE BRETTINGHAM (first of name) 1825. A catalogue of the shells contained in the collection of the late Earl of Tanker- ville, Appendix, (Terebra), pp. XXIII-XXIV. THIELE, JOHANNES 1925. Wissenschaftliche Ergebnisse der Deutchen Tiefsee-Expedition auf dem dampfer “Valdivia” 1898-1899, Jena, vol. 17, part 2, Gastropoda, (Terebra), pp. 343- 348, pls. 29-30. TOULA, FRANZ VON 1908. Ein Jungtertiare Fauna von Gatun am Panama-Kanal. Jahrbuch der Kaiserlich- Konigliche Geologische Reichsanstalt, Wien, Bd. 58, Heft 4, (Terebra), pp. 705-706, plates 25, 28. [1909.] PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XXXVII, No. 22, pp. 567-574; 15 figs. November 23, 1971 THE CARBONIFEROUS GENUS GLYPTOCHITON DE KONINCK, 1883 (MOLLUSCA : POLYPLACOPHORA) a Allyn G. Smith California Academy of Sciences The genus Glyptochiton was established by L. G. de Koninck in 1883 for some unusual fossils from the Carboniferous beds at Tournai, Belgium. These were described originally by him as Chiton? cordifer, which he designated as the type of his genus. Subsequently, four additional species of fossil chitons were de- scribed that can be assigned to Glyptochiton. These are: Chitonellus youngianus Kirkby in Young, 1865, from the Lower Carboniferous Main (or Hurlet) lime- stone at Craigenglen, Campsie, Ayrshire, and from Cunningham Baidland, near Dalry, both in western Scotland; Chitonellus subquadratus Kirkby and Young, 1867, from the same strata; Chitonellus quadratus Etheridge, 1882, and C. kirkbyanus Etheridge, 1882, also from the same strata. All of the earlier illustrations of the valves of Glyptochiton are stylized drawings. For this reason advantage is taken to provide photographs of a series of valves labeled “Glyptochiton cordifer de Kon./carbe-Kalenskalk/Tournai/ Belg.” in the collection of the United States National Museum (Natural His- tory), Division of Paleontology, no. 63404, which were loaned for study through the courtesy of Dr. G. Arthur Cooper. Because of the nature and the considerable age of the original label (headed “Comptoir Belge de Mineralogie & de Paleon- tologie/Ad. Piret-Tournai”’) it seems reasonable to assume that these valves are from the type locality of G. cordifer, perhaps identified by de Koninck himself. The six specimens in the set include a head and tail valve and four intermediate valves. So far as I know they are the only currently available representatives of the species in North America. [567] alory LIBRARY REC1 1971 Weeds Hole, Mass. [Proc. 4TH SEr. CALIFORNIA ACADEMY OF SCIENCES Vor. XXXVII] SMITH: CARBONIFEROUS GENUS GLYPTOCHITON 569 The tegmentum of the head valve (fig. 1) has no particularly marked sculp- ture but the dorsal side of the rather wide insertion plate has very faint, irregular, closely spaced grooves across it. This valve is about as wide as long; it is not much thickened on the ventral side (fig. 2). The tegmentum of intermediate valve A (fig. 3) consists of a raised, slightly overhanging, heart-shaped ridge forming an area 4.0 mm. long and 3.4 mm. wide, the point of the heart directed toward the anterior end of the valve; the area inside the ridge is much depressed with a prominent elongated tubercle situated at its center and extending anteriorly until it reaches the point of the heart- shaped ridge. On the dorsal side of the articulamentum of this valve there appears to be slight evidence of faint grooves similar to but weaker than those on the articulamentum of the head valve. However, across the porterior sinus (at the broad end of the heart-shaped ridge) there is a series of 8 closely spaced, low, narrow, short ridges bounded by narrower channels that extend under the small overhang of the tegmentum. Like the head valve, intermediate valve A is not appreciably thickened on the ventral side (fig. 4). The angle of divergence of the side slopes is 86° (fig. 9). Intermediate valves B, C, and D (figs. 5-8, 10-11, inclusive) have much the same sized heart-shaped tegmentum, although the preservation of these is such that there is only the faintest suggestion of the occurrence of the series of grooves, or ‘pectinations,’ across the posterior sinus described for intermediate valve A. The tail valve (fig. 12) has a semicircular posterior margin, with side margins that taper inward slightly toward what remains of a shallow anterior sinus. The tegmentum on this valve is well marked, with the edges slightly overhanging the articulamentum, forming a distinct eave, and has a prominent, laterally pinched- up mucro placed centrally and extending longitudinally almost to the anterior tegmentum margin. The shape of the tegmentum area is somewhat heart-shaped also, but with a semicircular posterior end opposite the point of the heart. The area occupied by the insertion plate is extensive toward the anterior end of the valve but becomes quite narrow about its middle and extends all the way around the semicircular posterior margin as a short, almost vertical ledge equal to the < Ficure 1. Glyptochiton cordifer (de Koninck, 1844). Head valve; dorsal view. U. S. National Museum no. 63404. Length, 6.1 mm. Ficure 2. Head valve; ventral view. FIGURE 3. Intermediate valve A; dorsal view. Length, 11.4 mm. Ficure 4. Same; ventral view. Ficure 5. Intermediate valve B; dorsal view. Length, 10.4 mm. Ficure 6. Same; ventral view. Ficure 7. Intermediate valve D; dorsal view. Length, 9.3 mm. Ficure 8. Same; ventral view. Ficure 9. Intermediate valve A; posterior end view showing valve thickness and an 86° angle of divergence of side-slopes. Height, 3.4 mm. The pectinations across the posterior sinus under the slight overhang of the heart-shaped tegmentum show faintly in this view. cn CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Vor. XXXVII] SMITH: CARBONIFEROUS GENUS GLYPTOCHITON 571 thickness of the valve’s articulamentum layer. This insertion-plate ledge is coarsely but closely pectinated, as shown in figs. 14 and 15. The measurements of the valves are as follows: Intermediate Measurement (mm.) Head Valve A Valve B Valve C Valve D Tail Length 6.1 11.4 10.4 7.9 9.3 10.0 Width 6.5 4.6 4.0 4.0 4.7 5.9 Height Shee) 3.4 S59! 2.8 3.5 - 34 Older paleontologists doubted that Glyptochiton cordifer was, in fact, a chiton. de Ryckholt (1845) thought it might be a crinoid plate and so also did Dall (1882) and Rochebrune (1883). According to Salter (1847), even de Koninck himself admitted it might be “an encrinital plate”, although he considered it definitely to be chitonoid when he established the genus Glyptochiton for it. There seems to be little doubt now that Glyptochiton is polyplacophoran. What is considered to be the tegmentum layer of the valves is similar in general configuration to that in the Recent genus Amicula in which the valves are buried deeply in a thick, muscular girdle with only small, heart-shaped areas exposed in the intermediate valves. The tegmentum of the valves in the Recent genus Choriplax from Australia also are somewhat similar, although the areas of the articulamentum of these valves are not at all similar to those in Glyptochiton and the overall valve outlines are not at all comparable. No species of chiton similar or referable to Glyptochiton has been reported in the Paleozoic other than from the Carboniferous of Belgium and western Scot- land. There are no comparable Mesozoic species known. This suggests that the evolutionary line represented by Glyptochiton may have been an offshoot from the main trend of chiton development, which came to an abrupt dead end in the Paleozoic for reasons that certainly are presently unknown. It seems likely, also, that the heart-shaped tegmentum areas in the intermediate valves of Amicula and Choriplax are of relatively recent origin. The only fossil record for the former genus is Amicula vestita altior Carpenter in Pilsbry, 1893, from the Pleistocene drift of Lower Canada; valves of Choriplax so far have not been reported to occur as fossils. The intermediate valves of Glyptochiton are unique in configuration com- pared with all known Recent and fossil Polyplacophora. No other chiton valves < Ficure 10. Glyptochiton cordifer. Intermediate valve C; dorsal view. Length, 7.9 mm. Ficure 11. Same; ventral view. Ficure 12. Tail valve; dorsal view. Length, 10.0 mm. Fic- URE 13. Same; ventral view. Ficure 14. Same; end view showing insertion-plate pectination. Width, 5.9 mm. Ficure 15. Same; from left side showing extent of the anterior portion of the articulamentum, pectination of the insertion plate, and position of the mucro. CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. z bo are known that are so highly elongate and, at the same time, have rather deep sinuses at both ends terminating laterally in quite pointed processes at the cor- ners. Even of greater significance is the occurrence, apparently for the first time in chiton evolutionary history, of grooved or pectinated insertion plates. Chitons with this type of insertion-plate sculpture so far have been considered by chiton specialists to represent the highest evolutionary development charac- terized by species placed in the family Chitonidae as now restricted. The oldest species with such insertion plates have been reported from the Cretaceous (Berry, 1939; Smith, Yochelson, and Sohl, 1968). In modern chitons with pectinated insertion plates, the pectinations occur usually on both head and tail valves and along the side margins of the intermediate valves; also, the insertion plates generally are cut by a varying number of slits, which divide the plate into “teeth.” In Glyptochiton, however, the tail valve is the only one seen with well developed pectinations in the insertion plate (figs. 14-15); in the intermediate valves such pectinations as there are occur in a small area at the base of the posterior sinus where they are not nearly so well developed. Glyptochiton valves have no slits or ‘“‘teeth” unless the sculptural character here interpreted as pectinations (which are more rounded and less sharply defined than in species in the Chitonidae) is in reality the forerunner of extremely closely spaced ‘‘teeth.”’ The following summarizes all of the known published references to the four species of Glyptochiton mentioned earlier in this report: Glyptochiton cordifer (de Koninck, 1844) Chiton? cordifer de Koninck,1844, p. 324, pl. 22, figs. 5a-b. de Ryckholt, 1845, p. 60, pl. 4, figs. 9-16. Salter, 1847, p. 49. Bronn, 1848, p. 291. Bigsby, 1878, p. 319. Dall, 1882, p. 283. Rochebrune, 1883, pp. 31-32. Chiton (Chitonellus) cordifer de Koninck. de Koninck, 1857, p. 196 (as “condifer’’). Baily, 1860, p. 95. Chitonellus cordifer (de Koninck). d’Orbigny, 1850, p. 127. Kirkby, 1862, p. 237 (foot- note). Etheridge, 1882, p. 97. Glyptochiton cordifer (de Koninck). de Koninck, 1883, p. 213; type species of Glyptochi- ton de Koninck, 1883, by original designation. P. H. Fischer, 1957, p. 12. Smith, 1960, p. 72, figs. 44, 5a-c (ex de Koninck, 1883). Glyptochiton youngianus (Kirkby in Young, 1865) Chitonellus youngianus Kirkby in Young, 1865, pp. 14-15, pl. 1, fig. 2. Kirkby and Young, 1867, pp. 341-342, pl. 16, figs. 2-4. Etheridge, 1882, pp. 101-102, pl. 2, figs. 23-24. Chiton youngianus (Kirkby in Young). Rochebrune, 1883, pp. 33-34 (as “yungianus”). Glyptochiton youngianus (Kirkby in Young). Smith, 1960, p. 72, figs. 44, 6a-b (ex Etheridge, 1882). Glyptochiton subquadratus (Kirkby and Young, 1867) Chitonellus subquadratus Kirkby and Young, 1867, p. 342, pl. 16, fig. 5. Etheridge, 1882, pp. 96-97, pl. 2, figs. 4-5. Glyptochiton quadratus (Etheridge, 1882) Chitonellus sp. indet. Etheridge, 1882, pp. 97-98, pl. 2, figs. 6-7. Chitonellus quadratus Etheridge, 1882, p. 98. Vor. XXXVII] ROTH & COAN: GALAPAGOS AND COCOS MARGINELLIDAE — 573 Glyptochiton kirkbyanus (Etheridge, 1882) Chitonellus kirkbyanus Etheridge, 1882, pp. 100-101, pl. 2, figs. 14-22. The preceding account of a peculiar group of Paleozoic chitons no doubt raises more evolutionary questions than can be answered with the small amount of factual information now at hand. Nevertheless, it is presented with the hope that more careful study of Glyptochiton valves preserved in European museum collec- tions will be encouraged, and that new material can be collected in order that the basis for answering at least some of these questions can be provided. REFERENCES CITED Barry, W. H. 1859. On the occurrence of detached plates of the shell of a new species of Chiton in the Carboniferous limestone at Lisbane, County of Limerick. Natural History Review and Quarterly Journal of Science, vol. 6, 1859, pp. 330-334, pl. 27, figs. 2a-c. [Also in: Journal of the Geological Society of Dublin, 1860, vol. 8, 1857-60, pp. 167-171, pl. 4, figs. 2a-c.] 1860. Observations on two new species of Chiton from the Upper Silurian ‘Wenlock Limestone’ of Dudley. Annals and Magazine of Natural History, ser. 3, vol. 6, no. 32, pp. 91-98, pl. 2, figs. la-d, 2a-c. London. [An English translation of de Koninck, 1857, q.v., with additional footnotes. ] Berry, CHARLES T. 1939. Some fossil Amphineura from the Alantic Coastal Plain of North America. Pro- ceedings of the Academy of Natural Sciences of Philadelphia, vol. 91, 1939, pp. 207-217, pls. 9-12. Bicssy, J. J. 1878. Thesaurus devonico-carboniferus. The flora and fauna of the Devonian and Carboniferous periods. Pp. 317, 319. London. Bronn, HEINRICH GEORG 1848. Index Paleontologicus oder Ubersicht der bis jetzt bekannten Fossilen Organis- men, Ist halfte, Chiton, p. 291. Stuttgart. Dati, WILLIAM HEALEY 1882. On the genera of chitons. Proceedings of the United States National Museum, vol. 4, 1881, pp. 279-291. ETHERIDGE, R., JR. 1882. A contribution to the study of the British Carboniferous Chitonidae. Proceedings of the Natural History Society of Glasgow, 1880-81, vol. 5, pt. 1, pp. 84-107, pls. 1-2. FiscHer, P. H. 1957. Revision des Amphineures de la Collection de l’Ecole des Mines de Paris. Journal de Conchiliologie, vol. 97, no. 1, pp. 1-24. Kirxsy, J. W. 1862. Onsome remains of Chiton from the Mountain limestone of Yorkshire. Quarterly Journal of the Geological Society, vol. 18, pp. 233-237, figs. 1-10. London. Kirxsy, J. W., and Joun Younc 1867. Notes on some remains of Chiton and Chitonellus from the Carboniferous strata of Yorkshire and the west of Scotland. Geological Magazine, vol. 4, no. 38, pp. 340-343, pl. 16. 574 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. OrBIGNY, ALCIDE DESSALINES D’ 1850. Prodrome de Paléontologique Stratigraphique Universelle des Animaux Mollusques et Rayonnés, vol. 1, 1850, p. 127. ROCHEBRUNE, A. T. DE 1883. Monographie des espéces fossiles appartenant a la Classe des Polyplaxiphores. Annales des sciences géologiques, vol. 14, art. 1, pp. 1-74, pls. 1-3. Paris. RYCKHOLT, BARON PHILIP DE 1845. Résumé geologique sur le genre Chiton Lin. Bulletin de |’Academie royale des Sciences et de Belle-Lettres de Bruxelles, vol. 12, no. 7, pt. 2, pp. 36-62, pls. 1—4. SALTER, JOHN W. 1847. Description of a fossil chiton from the Silurian rocks, with remarks on the fossil species of the genus. Quarterly Journal of the Geological Society (London), vol. 3, pp. 48-52, figs. 1-6. SmitTH, ALLYN G. 1960. Amphineura. Treatise on Invertebrate Paleontology (Raymond C. Moore, ed.), Part I, Mollusca 1, pp. 41-76, figs. 31-45. SmitH, ALLYN G., NorMAN F. Sout, and Eriis L. YOCHELSON 1968. New Upper Cretaceous Amphineura (Mollusca). United States Geological Sur- vey, Professional Paper 593-G, pp. 1-9, pl. 1. Younc, JOHN 1865. Provisional notice of a new Chiton, and a new species of Chitonellus, from the Carboniferous rocks of western Scotland; with descriptions by Mr. James W. Kirkby. Transactions of the Geological Society of Glasgow, vol. 2, pt. 1, pp. 13-15, pl. 1, figs. 1-2. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XXXVII, No. 23, pp. 575-584; 5 figs. November 23, 1971 MARGINELLIDAE (MOLLUSCA: NEOGASTROPODA) FROM THE GALAPAGOS ISLANDS AND COCOS ISLAND By Barry Roth Scientific Assistant, Department of Geology California Academy of Sciences and Eugene V. Coan Research Associate, Department of Geology California Academy of Sciences Eighteen species of the gastropod family Marginellidae have been reported from the eastern Pacific area (Coan and Roth, 1966; Roth and Coan, 1968). The present paper describes four more, recognized in the course of a bio- geographic study of the family; three of these receive names. Marginellid records for the Galapagos Islands (Hertlein and Strong, 1939, 1955) and Cocos Island, Costa Rica (Hertlein, 1963), are in part revised. For generous aid in locating and lending specimens, the authors are indebted to personnel of the Division of Mollusks, United States National Museum, and the Department of Malacology, Academy of Natural Sciences, Philadelphia. Dr. Leo G. Hertlein and Mr. Allyn G. Smith of the California Academy of Sci- ences contributed valuable opinions and advice. The shell illustrations were prepared by Miss Linette Sabre, Department of Herpetology, California Academy of Sciences. [575] LIBRARY MEC1 1971 Weods Hole, Mass. oat oO CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. . ( Ficure 1. Volvarina nyssa Roth and Coan, new species. Holotype no. 13713, California Academy of Sciences, Department of Geology, Type Collection. Isla Pinta, Galapagos Islands. Length 5.7 mm. DESCRIPTIONS OF SPECIES Volvarina nyssa Roth and Coan, new species. (Figures 1, 2.) DESCRIPTION OF HOLOTYPE. Shell small, ovate-cylindrical, narrower an- teriorly; highly polished, translucent white; body whorl with three spiral orange- brown bands, the first just anterior to suture, the second on broadest portion of body whorl, the third occupying posterior half of anterior third of shell (as seen in apertural view); spire low, rounded-conical, with portion of posterior color band visible, covered by a transparent glaze which leaves sutures visible but impalpable; outer lip shallowly sinuous medially, white, thickened along its outer margin; aperture with indistinct posterior notch, narrow, widening an- teriorly, color bands showing within; anterior margin evenly rounded, without indentation; columella slightly convex, with four equally spaced, sharp, oblique folds on anterior portion, most anterior fold at base of columella. Length 5.7 mm., width 3.0 mm. TyPE LOCALITY. Southwest side of Isla Pinta (Abingdon), Galapagos Is- lands, Ecuador (0° 31’ N; 90° 46’ W), intertidal area, collected by David Q. Cavagnaro, 25 May 1964. TYPE MATERIAL. Holotype no. 13713, California Academy of Sciences, De- partment of Geology, Type Collection. Four paratypes, in alcohol, no. 451, and one paratype, a radula slide, no. 452, California Academy of Sciences, De- partment of Invertebrate Zoology, Type Collection. One paratype, United States National Museum, Division of Mollusks. One paratype, Academy of Natural Sciences of Philadelphia, Department of Malacology. All paratypes are from the same locality as the holotype. VoLt. XXXVII] ROTH & COAN: GALAPAGOS AND COCOS MARGINELLIDAE _ 577 Ficure 2. Volvarina nyssa Roth and Coan, new species, rachidian radular plate. Para- type no. 452, California Academy of Sciences, Department of Invertebrate Zoology, Type Collection. Isla Pinta, Galapagos Islands. Longest dimension 42 uw. REFERRED MATERIAL. One specimen, California Academy of Sciences locality no. 38871, Darwin Bay, Isla Genovesa (Tower), Galapagos Islands (0° 19’ N; 89° 58’ W), in tide pools, collected by Allyn G. Smith, 30 January 1964. Discussion. In outline, intense coloring, and small size, Volvarina nyssa resembles Volvarina albolineata (d’Orbigny, 1842) of the Caribbean fauna. The central brown band of the latter species is narrower than that of V. nyssa (cf. Warmke and Abbott, 1961, plate 23, figure i). Tryon (1882) reported ‘Mar- ginella” albolineata as occurring both in the West Indies and in Lower California, but Tomlin (1916) correctly rejected the eastern Pacific record. All specimens we have seen from both coasts of Baja California belong to the species Volvarina taeniolata Moerch, 1860. Volvarina nyssa is readily distinguished from the pink Volvarina taeniolata rosa (Schwengel, 1938), the other Galapagos species of the genus, on the basis of color. Erosion and sunbleaching of dead specimens may reduce the normal pink color of the latter, revealing brown banding which is not, however, as clear and uniform in density as that of V. myssa. The specimens on which Hertlein and Strong (1955) based their Galapagos record of ““Marginella (Hyalina) cali- fornica Tomlin” |= Volvarina taeniolata taeniolata| all appear referable to V. taeniolata rosa. Specimens of Volvarina taeniolata taeniolata from southern California, which may have color bands disposed as in V. myssa, are larger as adults (length 7.5 to 9 mm.), more inflated and less cylindrical, and have a ground color of pale straw—seldom the clear white of V. myssa. Color bands of the California shells are frequently edged with darker brown, as in the form named Marginella cali- fornica var. parallela Dall, 1918. This seems to be without taxonomic signifi- cance. Banded species of Volvarina occur in the western Pacific, in both temperate and tropical waters, but none appears similar enough to V. myssa to require special comparison. The radula of this new species consists of straight comblike rachidian plates, on which large cusps alternate with groups of three and four smaller cusps (fig. 2). A total of 21 cusps are present on the figured specimen, taken from a shell about 5.5 mm. in length. Present collecting records of Volvarina nyssa are restricted to Isla Pinta and 578 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Isla Genovesa, outlying members of the Galapagos Archipelago north of the equator. Museum collections contain specimens of Volvarina taeniolata rosa from Islas Santa Cruz, Pinzén, Rabida, and San Cristobal, and the portion of Isla Isabela south of the equator. Abbott (1966, figure 2) has plotted the an- nual surface temperature regimes for the Galapagos area; temperatures north of the equator average higher than those south, particularly during the months June to December. If the apparent geographic separation of V. nyssa and V. taeniolata rosa is not merely the result of insufficient collecting, temperature may be a cause. The name “nyssa” is a Greek noun meaning a goal, or starting post. Granula achenea Roth and Coan, new species. (Figure 3.) Marginella minor C. B. Adams, BArTscH and REHpER, Smithson. Miscell. Coll., vol. 98, no. 10 (Pub. 3535), p. 18, 1939. Cocos Island, Costa Rica. HErTLEIN, Proc. Calif. Acad. Sci., ser. 4, vol. 32, no. 8, p. 240, 1963. Cocos Island. Not Marginella minor C. B. Apams, Catalogue of shells collected at Panama, p. 264, 1852. Panama City, Panama. DESCRIPTION OF HOLOTYPE. Shell minute, ovate, broader posteriorly; almost transparent, highly polished, colorless but with bluish-white glaze; spire low, convexly conical, unsculptured; outer lip nearly straight, on last whorl advanc- ing high on spire, thickened by callus evenly along its outer edge, with a few faint denticles internally; suture appressed, indistinct, rendered irregular by narrow, closely spaced, longitudinal, raised ridges which extend somewhat protractively across body whorl, becoming fainter and disappearing just behind anterior mar- gin of shell; “false suture” (internal trace of body whorl’s junction with previous whorl) visible under enamel of body whorl, just anterior to actual suture; aper- ture with posterior notch, narrow, widening slightly anteriorly; parietal wall evenly convex, with slight, transparent callousing posteriorly; columella with five evenly spaced folds (including the one at base of columella), and one very faint smaller fold posterior to them; folds thick, nearly square in outline; sharply defined opaque siphonal fasciole present on body whorl adjacent to columellar folds, sculptured with minute papillae; anterior margin of shell indented into small siphonal notch. Length 2.6 mm., width 1.6 mm. Type LocALity. Chatham Bay, northeast side of Isla del Coco (Cocos Is- land), Costa Rica (5° 35’ N, 87° 02’ W), in 80 feet (24.5 meters), collected by Waldo L. Schmitt, 24 April 1941. The bottom of Chatham Bay is said to consist of coral and sand. The shells in the type lot were dead when collected. TYPE MATERIAL. Holotype no. 568110, United States National Museum, Division of Mollusks. Thirty-six paratypes, no. 681406, United States National Museum, Division of Mollusks. Paratypes also have been distributed to the California Academy of Sciences; Museum of Comparative Zoology, Harvard Uni- versity; Los Angeles County Museum of Natural History; Academy of Natural Vor. XXXVII] ROTH & COAN: GALAPAGOS AND COCOS MARGINELLIDAE _ 579 Ficure 3. Granula achenea Roth and Coan, new species. Holotype no. 568110, United States National Museum, Division of Mollusks. Chatham Bay, Isla del Coco. Length 2.6 mm. Sciences, Philadelphia; Geology Department, Stanford University; and the American Museum of Natural History. The paratypes differ from the holotype mainly in development of callus around the aperture. Younger specimens show less callousing along the edge of the outer lip and on the parietal wall; their columellar folds are sharp instead if thick and somewhat squared as in the holo- type, and they do not have denticles inside the outer lip. REFERRED MATERIAL. One specimen, United States National Museum, no. 427596, Chatham Bay, Cocos Island, collected during Presidential Cruise, 1938, from bottom sample. Twenty-seven specimens, Academy of Natural Sciences of Philadelphia, no. 155005, Chatham Bay, Cocos Island, collected by H. A. Pilsbry, 1929; these show some variation in height of spire. Discussion. As indicated by McLean (1969), the genus Granula Jous- seaume, 1875, is a prior synonym of Kogomea Habe, 1951, the name which was applied to this group by Coan and Roth (1966). From all other eastern Pacific species of Granula, this species is immediately recognizable by the strong and numerous longitudinal ridges which corrugate its surface. The occasional thicken- ing of one or more growth lines has been observed as a variation in some individ- uals of other species, but not the regular development of a plicate surface over almost the entire shell. Surface ornamentation of any kind is rare among members of the subfamily Cystiscinae, being otherwise confined to a few species of the recent genera Pugnus and Marginellopsis and the Miocene genus Topaginella, none of which is known from the eastern Pacific region. In outline Granula achenea is not as conspicuously conical as Granula minor (C. B. Adams, 1852); its spire is more inflated than that of Granula polita (Carpenter, 1857). 580 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. ““Marginella minor C. B. Adams” was the only marginellid listed by Hertlein (1963) from Cocos Island, following Bartsch and Rehder (1939); Coan and Roth (1966) repeated this record. On re-examining Bartsch and Rehder’s specimen and other Cocos Island material in the Academy of Natural Sciences of Philadelphia and United States National Museum collections, we were unable to find any specimens which closely match the typical form of Granula minor from Panama. Granula achenea is the most numerous Cocos Island species in the collections. With it in some lots are specimens of a species of Granula re- sembling the lectotype of Gibberula coniformis Moerch, 1860 |= Granula polita|, from Costa Rica (Coan and Roth, 1966, plate 51, fig. 75), but with a lower spire and differences in outline. As presently construed, Granula polita is a species with considerable geographic variation; further study may permit the discrimination of localized subspecies, or even more extensive reclassification. The name proposed is a noun formed from the botanical term, achene, a dry seed, derived from the Greek achanes, not gaping. Granula insularum Roth and Coan, new species. (Figure 4.) Marginella (Cystiscus) minor C. B. Adams, HERTLEIN and StRoNG, Essays in Nat. Sci. in Honor of Capt. Allan Hancock, p. 127, 1955 (in part). Marginella (Cystiscus) polita Carpenter, HERTLEIN and Stronc, Essays in Nat. Sci. in Honor of Capt. Allan Hancock, p. 128, 1955 (in part). Marginella (Cystiscus) regularis Carpenter, HERTLEIN and StrRONG, Essays in Nat. Sci. in Honor of Capt. Allan Hancock, p. 128, 1955 (in whole or part). DESCRIPTION OF HOLOTYPE. Shell minute, elongate-ovate, broader poste- riorly; translucent, white, polished, without sculpture; spire very low, evenly dome-shaped; outer lip nearly straight, on last whorl extending high on body whorl, smooth within, thickened by callus along its outer edge, most strongly thickened on posterior portion; suture very fine and faint; “false suture” (internal trace of body whorl’s junction with previous whorl) visible through body whorl anterior to suture; aperture with oblique, curved posterior notch; posterior third of aperture moderately narrow, anterior portion wider; parietal wall gently and evenly convex, irregularly calloused on posterior half; columella with five folds (including the one at base of columella), anterior two subequal and strong, most posterior one very faint; minute granulation visible on parietal area adjacent to columellar folds; anterior margin of shell indented into a broad, somewhat oblique, siphonal notch. Length 2.5 mm., width 1.5 mm. Type Locaity. California Academy of Sciences locality no. 27221, Caleta Black (Black Bight), about one mile west of Tagus Cove at south end of Banks Bay, Isla Isabela (Albemarle), Galapagos Islands (0° 15’ S, 91° 23’ W), on black sand beach, collected by Leo G. Hertlein, 5 January 1932. The shells in the type lot were dead when collected. Vor. XXXVII] ROTH & COAN: GALAPAGOS AND COCOS MARGINELLIDAE 381 Ficure 4. Granula insularum Roth and Coan, new species. Holotype no. 13715, California Academy of Sciences, Department of Geology, Type Collection. Caleta Black, Isla Isabela, Galapagos Islands. Length 2.5 mm. TYPE MATERIAL. Holotype no. 13715, California Academy of Sciences, De- partment of Geology, Type Collection. Five paratypes, nos. 13716-13720, Cali- fornia Academy of Sciences, Department of Geology, Type Collection. Paratypes also have been distributed to the Museum of Comparative Zoology, Harvard University; United States National Museum; American Museum of Natural History; Academy of Natural Sciences, Philadelphia; Geology Department, Stan- ford University; San Diego Natural History Museum; and the Los Angeles County Museum of Natural History. All paratypes are from the same locality as the holotype. The length/width ratio is inconstant in this species, and some paratypes are shorter, in proportion to greatest diameter, than the holotype. REFERRED MATERIAL. Galapagos Islands: Fifteen specimens, California Academy of Sciences locality no. 27232, Conway Bay, Isla Santa Cruz (In- defatigable) ; four specimens, California Academy of Sciences locality no. 38910, beach at west end of south channel of Isla Santa Cruz, collected by V. A. Zullo, 14 February 1964; about thirty specimens, California Academy of Sciences locality no. 27249, yellow beds in cliffs on west side of Isla Baltra (South Sey- mour), collected by Leo G. Hertlein, 16-18 January 1932, late Pliocene. Four specimens, Academy of Natural Sciences, Philadelphia, no. 154787, Seymour Bay, Isla Santa Cruz; about seventy specimens, Academy of Natural Sciences, Philadelphia, nos. 153220, 153222, and 170349, Wreck Bay, Isla San Cristobal (Chatham). Discussion. The very low spire and elongate outline of Granula insularum are sufficient to distinguish it from other eastern Pacific species of the genus. Granula minor (C. B. Adams), which may be low-spired, is smaller (length 2.3 mm.) and much more conical. Granula polita (Carpenter, 1857), while elongate in some populations, especially in the southwestern portion of the Gulf CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. V1 co bdo Ficure 5. Persicula (?) species. Hypotype no. 13721, California Academy of Sciences, Department of Geology, Type Collection. Caleta Black, Isla Isabela, Galapagos Islands. Length 3.5 mm. of California, has not been seen with the very low, dome-shaped spire of G. insularum. The outer lip of G. polita is also characteristically more sinuous medially, and often faintly denticulate within. Members of the genus Cystiscus, which may resemble the new species in general outline, do not have the anterior margin indented into a siphonal notch. Granula insularum (including the type lot) constitutes part of the Gala- pagos records by Hertlein and Strong (1955) of “Marginella (Cystiscus) minor,’ “M. (C.) polita,’ and “M. (C.) regularis,’ and is the most numerous marginellid species present in that material. Specimens tentatively referable to Granula minor and Granula polita are also present. Their relationship to typical mainland specimens of those species is still under study. As shown by Coan and Roth (1966), Marginella regularis Carpenter, 1864, is a synonym of Granula subtrigona (Carpenter, 1864), a species not present in any Galapagos material we have examined. The name Marginella regularis has also been applied to Cystiscus politulus (Dall, 1919), another species which apparently does not occur in the Galapagos Islands. In late Pliocene material from Isla Baltra, examined through the kindness of Dr. Leo G. Hertlein, who assigned its age, Granula insularum is the most common marginellid species represented. The specimens show a similar range of variation to that observed in the type lot. The new species is apparently not present in the collection reported on by Hertlein and Strong (1939) from a late Pleistocene raised beach on Isla San Salvador (James). The specimens they recorded as “Marginella minor C. B. Adams” resemble Recent Granula polita. The name proposed for the new species is Latin, meaning ‘“‘of the islands” and is a noun in the genitive case. Vor. XXXVII] ROTH & COAN: GALAPAGOS AND COCOS MARGINELLIDAE _ 583 Persicula (?) species. (Figure 5.) Small white specimens possibly belonging to this genus have been seen from two Galapagos localities and Panama. Three were apparently mature, with thickened outer lip, at a length of 3.5 mm. or less. Although the shells are small for the genus, the flat spire immersed in a pad of enamel and the pattern of callousing on the parietal wall are typical of Persicula. REFERRED MATERIAL. One specimen, United States National Museum no. 509123, Panama, collected by James Zetek. One specimen, California Academy of Sciences locality no. 27221, Caleta Black, west of Tagus Cove, Isla Isabela, Galapagos Islands, collected by Leo G. Hertlein, 1932; one specimen, California Academy of Sciences locality no. 43261, north side of Bahia Academy, Isla Santa Cruz, on gorgonians, collected by André and Tui De Roy, 1964. A second, immature specimen from the last locality was broken in an unsuccessful attempt to remove the radula for study. Two specimens from the late Pliocene of Isla Baltra, California Academy of Sciences locality 27249, are also tentatively referred here. REFERENCES Axsportt, D. P. 1966. Factors influencing the zoogeographic affinities of Galapagos inshore marine fauna. Jn: Bowman, R. I., editor, The Galapagos. Proceedings of the symposia of the Galapagos International Scientific Project (Berkeley: Uni- versity of California Press), pp. i-xviii, 1-318 [Abbott, pp. 108-122, figs. 1-8 in text]. BartscH, P., AnD H. A. REHDER 1939. Mollusks collected on the Presidential Cruise of 1938. Smithsonian Miscellaneous Collections, vol. 98, no. 10 (Pub. 3535), pp. 1-18, pls. 1-5, June 13. Coan, E. V., AnD B. RotTH 1966. The west American Marginellidae. The Veliger, vol. 8, no. 4, pp. 276-299, pls. 48-51, figs. 1-5 in text, April 1. Hert LeEIN, L. G. 1963. Contribution to the biogeography of Cocos Island, including a bibliography. Proceedings of the California Academy of Sciences, ser. 4, vol. 32, no. 8, pp. 219-289, figs. 1-4 in text, May 20. HeErTLEIN, L. G., anp A. M. STRONG 1939. Marine Pleistocene mollusks from the Galapagos Islands. Proceedings of the California Academy of Sciences, ser. 4, vol. 23, no. 24, pp. 367-380, pl. 32, July 20. 1955. Marine mollusks collected at the Galapagos Islands during the voyage of the Velero III, 1931-1932. In: Essays in the natural sciences in honor of Captain Allan Hancock on the occasion of his birthday, July 26, 1955 (Los Angeles: University of Southern California Press), pp. 111-145, pl. A, July 26. McLean, J. H. 1969. Marine shells of southern California. Los Angeles County Museum Science Series 24, Zoology no. 11, pp. 1-104, figs. 1-54, October. 584 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Roru, B., ano E. V. Coan 1968. Further observations on the west American Marginellidae with the descriptions of two new species. The Veliger, vol. 11, no. 1, pp. 62-69, pl. 7, figs. 1-2 in text, 1 map, July 1. Tomuin, J. R. LEB. 1916. Note on the Marginella varia of Sowerby. Nautilus, vol. 29, no. 12, pp. 138-139, April 8. Tryon, G. W., Jr. 1882. Family Marginellidae. Jn: Manual of conchology; structural and systematic, vol. 5, Marginellidae, part 17, pp. 5-58, pls. 2-13, December 21. WarMkEgE, G. L., ano R. T. ABBOTT 1961. Caribbean seashells (Narberth, Pennsylvania: Livingston Publishing Company), pp. i-x, 1-346, pls. 1-44, figs. 1-34 in text, maps 1-19. INDEX TO VOLUME XXXVII FOURTH SERIES New names and principal reference in boldface type Abeona aurora, 476 Acanthodactylus cantoris, 48 cantoris blanfordi, 48 cantoris cantoris, 48 Acartia, 455 Acila (Truncacila) castrensis, 82 Acontiophis, 250 paradoxa, 250, 268 Actinocyclus, 490, 496, 497, 498 cubitus, 491, 493, 496, 497, 498 ehrenbergii, 498 octonarius, 498 senarius, 179 undulatus, 179 Actinophaenia splendens, 179 Actinoptychus, 490, 498 bismarkii, 491, 493, 496, 497, 498 incisa, 499 Senarius, 174, 175, 176, 177, 178, 179, 216, 218, 226, 491, 493, 496, 497, 498 solisi, 179, 499 splendens, 179, 212, 218 splendens incisa, 491, 496, 499 splendens solisi, 491, 496, 497, 499 undulatus, 179 Acus columna trajana, 544 coronata, 555 zebra, 551 Addicott, Warren O., Late Pliocene mollusks from San Francisco peninsula, Cali- fornia and their paleogeographic significance, 57-93. Agabinus, 242, 245 glabrellus, 242 Agama, 37 agilis, 30, 52 agrorensis, 31, 35, 40, 55 badakhshana, 32, 33, 55 caucasica, 35 erythrogastra, 36, 37, 55 himalayana, 35, 38, 55 himalayana himalayana, 37, 38 himalayana sacra, 35 himalayanum, 35, 37 lehmanni, 38, 55 nupta, 39, 273 nupta nupta, 39 nuristanica, 39, 40, 42, 45, 55 tuberculata, 40, 41, 42 Agamura, 43, 44, 338, 352 cruralis, 43 femoralis, 43, 44 persica, 42, 43, 44 Alepisaurus aesculapius, 482 ferox, 482 Allen, Gerald R., see Eschmeyer, William N. Allopora, 460, 461, 470 boreopacifica, 470 californica, 459, 460, 461, 462, 469, 470 petrograpta, 470 porphyra, 470 stejnegeri, 470 venusta, 470 Alsophylax, 334, 338 laevis, 334 loricatus, 334 microtus, 334 persicus, 337, 345, 348, 350, 356, 357, 358 pipiens, 334, 338, 350 prezwalskii, 334 spinicauda, 334 tibetanus, 334 Amblyrhynchus cristatus, 104 Amicula, 571 vestita altior, 571 Amphigonopterus aurora, 476 Anadara, 78 (Anadara) trilineata, 62, 76, 79 (Larkinia) multicostata, 78 trilineata, 66, 70, 74, 76, 82, 85 Anaitides, 461 Anderson, Jeromie A.,see Minton, Sherman A. Anderson, Steven C., and Alan E. Leviton, Amphibians and reptiles collected by [585] 586 CALIFORNIA ACADEMY OF SCIENCES the Street Expedition to Afghanistan, 1965, 25-56. Anderson, Steven C., see Leviton, Alan E. Anderson, Steven C., see Minton, Sherman A. Androctononus australis, 356 (Annelida, Polychaeta) from a hydrocoral off central California, Polydora al- loporis, new species, a commensal spionid, by William J. Light, 459-472. Anoplopoma fimbria, 442 Anous stolidus, 102, 136, 158 tenuirostris, 107, 136 Anuroctonus, 16, 19 phaeodactylus, 2, 8, 10, 13, 14, 18 Apolymetis biangulata, 83 Arachnoidiscus, 179, 180, 490, 498 decorus, 491, 493, 494, 495, 498 ehrenbergi, 174, 175, 179, 218 indicus, 180, 226 ornatus, 179 (Araneae: Salticidae), Darwinneon crypticus, a new genus and species of jumping spider from the Galapagos Islands, by Bruce Cutler, 509-513. Arca microdonta, 82 Ardea herodias, 102, 105, 131, 158 Artemia salina, 433 Asio (Flammeus) galapagoensis, 102 galapagoensis, 95, 102, 104, 139, 140, 143, 146, 158 Astarte, 177 Asterolampra, 181 darwinii, 180, 228 Asteromphalus, 180, 181 buchii, 180 darwinii, 180, 181 hookerii, 180 humboldtii, 180 rossi, 180 Astyris ricthofeni, 68 Atlantic scorpionfishes, Two new, by William N. Eschmeyer, 501-508 Atretia gnomon, 561 Aulacodiscus, 169, 181 kilkellyanus, 181 laxus, 181, 232 schmidti, 181 septus, 181 tripartitus, 181, 193, 220, 232 131, 188 [Proc. 4TH SER. Bacteriastrum, 455 Balanus, 173, 428, 434, 444, 457 (Armatobalanus) nefrens, 461, 470 Ball, Robert W., and David L. Jameson, Biosystematics of the canyon tree frog Hyla cadaverina Cope (= Hyla californiae Gorman), 363-380. Benthic fishes cast ashore by giant waves near Point Joe, Monterey County, California, by W. I. Follett, 473-488. Biddulphia, 182 aurita, 182, 226 baltzoi, 182, 226 cookiana, 182 longicruris, 455 roperiana, 182, 226 Bioculus, 290 Biology of storm petrels in the Galapagos Islands, The, by M. P. Harris, 95-165. Biosystematics of the canyon tree frog Hyla cadaverina Cope (= Hyla californiae Gorman), by Robert W. Ball and David L. Jameson, 363-380. Bittium casmaliense, 62, 64, 65 Boa tatarica, 51 Brachycereus, 145 Bratcher, Twila, and R. D. Burch, The Te- rebridae (Gastropoda) of Clarion, So- corro, Cocos, and Galdpagos islands, 537-566. Brosmius marginatus, 480 Brosmophycis marginata, 475, 480, 481, 483 Broteas alleni, 2, 13 Buccinum crenulatum, 555 elongatum, 552 luteola, 555 maculatum, 544, 555 maculatum var. B, 544 maculatum var. C, 544 ornatum, 553 varicosum, 555 Bufo andersonii, 27, 28, 29 microscaphus, 374 oblongus, 28, 29 olivaceus, 29 pentoni, 29 persicus, 29 surdus, 29 viridis, 28, 29 VoL. XXXVIT] Bunopus, 333, 338 abudhabi, 334, 337 blanfordi, 333, 334, 337 crassicauda, 334, 335, 336, 337 persicus, 348 spatulurus, 334 tuberculatus, 44, 333, 334, 337 Burch, R. D., see Bratcher, Twila Bursera graveolens, 145 Buteo galapagoensis, 141 Buthus occitanus, 356 Calanus, 455 helgolandica, 455 Calicantharus portolaensis, 82 California Academy of Sciences (Coleoptera: Staphylinidae), East asiatic and ori- ental species of Stenus represented in the collection of the, by Volker Puthz, 529-535. California, Benthic fishes cast ashore by giant waves near Point Joe, Monterey County, by W. I. Follett, 473-488. California sea lion: skull growth and a com- parison of two populations, The, by Robert T. Orr, Jacqueline Schonewald, and Karl W. Kenyon, 381-394. Callorhinus ursinus, 387 Calyptraea mamillaris, 82 Cancellaria, 62, 64, 69, 78, 79 arnoldi, 62, 64, 69, 74, 75, 76, 79 emydis, 79 lipara, 69, 74, 75 Candacia, 455 Canthydrus, 243 bicolor, 243 buqueti, 243 octoguttatus, 243 uniformis, 243 canyon tree frog Hyla cadaverina Cope (= Hyla californiae Gorman), Bio- systematics of the, by Robert W. Ball and David L. Jameson, 363-380. Carboniferous genus Glyptochiton de Koninck 1883 (Mollusca: Polyplacophora), The, by Allyn G. Smith, 567-574. Cardium meekianum, 82 Casmerodius egretta, 102 Catachlaena, 250 diadema, 261 Cataetyx rubrirostris, 481 INDEX 587 Centropages, 447, 455 Centruroides, 14, 19, 20, 21 exilicauda, 13 gertschi, 2, 13 gracilis, 16 insulanus, 18 sculpturatus, 2, 3, 8, 10, 11, 13, 15, 16, 17 vittatus, 13, 18 Ceratium, 455 furca, 455 longipes, 455 macroceros, 455 tripos, 455 Cetengraulis mysticetus, 440 Chaetoceros, 174, 175, 183, 196, 224, 437, 438, 443, 455 decipiens, 455 didymus, 182, 183, 224, 230 Chalcoscirtus, 511 Chatachlein, 250 diadema, 257 Chelonia, 30 Chilara, 482, 483 taylori, 475, 481, 482, 483 Chiton (Chitonellus) cordifer, 572 cordifer, 567, 572 youngianus, 572 Chitonellus, 572 cordifer, 572 kirkbyanus, 567, 573 quadratus, 567, 572 subquadratus, 567, 572 youngianus, 567, 572 Choriplax, 571 Clarion, Socorro, Cocos, and Galapagos islands, The Terebridae (Gastropoda) of, by Twila Bratcher and R. D. Burch, 537-566. Clementia, 80, 82 Clinocardium, 62, 68, 71, 74, 77 meekianum, 62, 68, 71, 74, 77, 82 nuttalli, 77 Clupea harengus pallasi, 442 Cnemaspis, 338 Coan, Eugene V., see Roth, Barry Cocconeis, 174, 175, 183 antiqua, 183, 220 formosa, 183, 228 japonica, 183 lorenziana, 183 588 CALIFORNIA ACADEMY OF SCIENCES maxima, 183, 214 nitida, 193 pribilofensis, 184, 214 scutellum, 183 Cocos, and Galapagos islands, The Terebridae (Gastropoda) of Clarion, Socorro, by Twila Bratcher and R. D. Burch, 537-566. Cocos Island, Marginellidae (Mollusca: Neo- gastropoda) from the Galapagos Is- lands and, by Barry Roth and Eugene V. Coan, 575-584. (Coleoptera: Dytiscidae and Noteridae), Co- pelatus glyphicus (Say) and Suphi- sellus bicolor (Say), water beetles new to California and presumably intro- duced, by Hugh B. Leech, 237-247. Coluber karelini, 356 mucosus, 53 ravergieri, 50, 52 rhodorhachis, 50, 52, 356 (Taphrometopon) lineolatus, 53 Colus recurvus, 80, 82 Colymbetes glyphicus, 238 Compsobuthus acutecarinatus, 20, 21 Compsomyax subdiaphana, 82, 83 Conolophus subcristatus, 104 Copelatus, 238, 242 chevrolati, 240 chevrolati australis, 240 chevrolati renovatus, 238, 240, 241, 242 chevrolati schaefferi, 240 debilis, 240 distinctus, 238, 241 glyphicus, 237, 238, 239, 240, 242, 245 impressicollis, 238 Copelatus glyphicus (Say) and Suphisellus bicolor (Say), water beetles new to California and presumably introduced (Coleoptera: Dytiscidae and No- teridae), by Hugh B. Leech, 237-247. Cordia lutea, 145 Coronella tessellata, 52 Corycaeus, 447, 455 Coryphopterus nicholsii, 475, 478, 483 Coscin curvatus recta, 189 Coscinodiscus, 184, 186, 187, 432, 438, 448, 449, 455, 490, 493, 497 antiquus, 491, 494, 497 apiculatus, 174, 175, 178 [Proc. 4TH SER. asteromphalus, 455, 491, 492, 493, 494, 497 asymmetricus, 187 (Cestodiscus) intersectus, 187 excentricus, 184, 455 fimbriatus, 184, 226 fimbriatus californica, 184 kitzingii, 184, 204 lineatus, 174, 185 marginatus, 174, 175, 176, 177, 178, 184, 202, 491, 492, 493, 498 nano-lineatus, 185 obscurus, 491, 492, 493, 494, 498 oculus-iridis, 185, 208, 455 pacificus, 491, 492, 493, 494, 498 pustulatus, 174, 175, 176, 177, 178, 185, 186, 206, 210 radiatus, 174, 175, 176, 177 lies oo: 204, 206, 208, 455 rothii, 184, 186, 216 stellaris, 455 symmetricus, 186 temperel, 187 undulosus, 174, 175, 176, 177, 178, 186, 202 wailesii, 455 Cosmiodiscus insignis, 174, 175, 176, 178, 186, 187, 206, 214 Crassatella collina, 82 Creagrus furcatus, 102, 103, 134, 138, 139, 143, 145. 04758 Crepidula, 65, 66 grandis, 82 princeps, 62, 65, 66, 82 Croton scouleri, 145 Cryptocarpus, 146 Cryptomya californica, 63, 66, 70, 74, 76, 82 ovalis, 74 Cryptonatica, 76 aleutica, 62, 64, 65, 70, 76 clausa, 65 russa, 65 Cunearca, 78 Cusoria elegans, 51 Cutler, Bruce, Darwinneon crypticus, a new genus and species of jumping spider from the Galapagos Islands (Araneae: Salticidae), 509-513. Cyclotella antiqua, 190 —_ VoL. XXXVIIT] Cymatotheca minima, 187 weissflogii, 174, 175, 176, 178, 187, 208 Cyrtodactylus, 44, 338, 352 agamuroides, 43, 44 caspius, 44 fedtschenkoi, 27, 45 femoralis, 44 gastropholis, 43 heterocercus, 44, 359, 360, 361 kotschyi, 360 macularius, 46 persicus, 350 russowi, 361 scaber, 45, 46 watsoni, 45, 55 Cystiscus, 582 politulus, 582 Daption capensis, 115 Darwinneon, 509, 511, 512, 513 erypticus, 509, 510, 511, 512 Darwinneon crypticus, a@ new genus and species of jumping spider from the Galapagos Island (Araneae: Saltic- idae), by Bruce Cutler, 509-513. Dendroica petechia, 102 diatom flora from Oregon, A late Tertiary, by W. N. Orr, Judi Ehlen, and J. B. Zaitzeff, 489-500. Diatoma aurita, 182 diatoms from the Pribilof Islands, Bering Sea, Alaska, Fossil, by G Dallas Hanna, 167-234. Dicladia capreola, 188, 224 capreolus, 455 pylea, 188 Dictocystis, 435 Dictyocha, 176 fibula, 175, 176, 177, 178, 198, 224 speculum, 175, 176, 198 diet and feeding behavior of the northern anchovy, Engraulis mordax (Girard), On the, by Anatole S. Loukashkin, 419-458. Dinophysis, 455 Diomedea epomorpha, 141 Diplocentris whitei, 2, 13 Diploneis, 188 bombus, 188, 206 INDEX 589 ornata, 174, 188, 206 smithii, 455 Discinisca cumingi, 59, 63 Distephanus, 198 speculum, 178, 198, 224, 455 Ditylum, 190 Dosinia, 80, 81 ponderosa, 82 Dossetia temperei, 188, 224 Drillia graciosana, 69 mercedensis, 69 Drosophila, 12 East asiatic and oriental species of Stenus represented in the collection of the Calfornia Academy of Sciences (Cole- optera: Staphylinidae), by Volker Puthz, 529-535. Easter Island, Three new species of scor- pionfishes (Family Scorpaenidae) from, by William N. Eschmeyer and Gerald B. Allen, 515-527. Ebria antiqua, 198 Ebriopsis, 198 antiqua, 175, 176, 177, 197, 198, 224 Echis carinatus, 50, 54, 356 Ehlen, Judi, see Orr, W. N. Engraulis anchoita, 446 encrasicholus, 443 japonica, 442 mordax, 419, 423, 443, 447, 449, 450, 455, 456 ringens, 450, 451 Engraulis mordax (Girard), On the diet and feeding behavior of the northern an- chovy, by Anatole S. Loukashkin, 419- 458. Enteropneusta, 427, 429, 435, 437, 440, 444 Entophyla, 192 Eremias guttulata watsonana, 48 (Mesalina) watsonana, 48 nigrocellata, 48, 55 persica, 49 velox persica, 49 Errinopora pourtalesi, 461 Eryx elegans, 27, 50, 51, 55 jaculus czarewskii, 51 590 CALIFORNIA ACADEMY OF SCIENCES tataricus, 50, 51 tataricus speciosus, 51 tataricus tataricus, 51 Eschmeyer, William N., Two new atlantic scorpionfishes, 501-508. Eschmeyer, William N., and Gerald R. Allen, Three new species of scorpion- fishes (Family Scorpaenidae) from Easter Island, 515-527. Eublepharis macularius, 46 Eucampia, 455 Eumeces schneideri blythianus, 356 Eumetopias jubata, 392 Euoda weissflogii, 187 Euodia ratabouli, 187 weissflogii, 187 Euphorbia caducifolia, 354, 355 Euprepes dissimilis, 49 Euscorpius italicus, 21 Eusthenura group of Vejovis from Baja California, Mexico (Scorpionida: Vejo- vidae), New scorpions belonging to the, by Stanley C. Williams, 395-418. Eutherpina, 455 acutiformis, 447 Evadne nordmanni, 448 Exuviaella, 448 feeding behavior of the northern anchovy, Engraulis mordax (Girard), On the diet and, by Anatole S. Loukashkin, 419-458. Florimetis biangulata, 83 Follett, W. I., Benthic fishes cast ashore by giant waves near Point Joe, Monterey County, California, 473-488. Fossil diatoms from the Pribilof Islands, Bering Sea, Alaska, by G_ Dallas Hanna, 167-234. Fregata, 147 magnificens, 102 minor, 133, 134, 145, 146, 158 Fritillaria, 435 Frustulia lata, 191 Fuscus (Buccinofuscus) portolaensis, 82 Galapagos Islands and Cocos Island, Mar- ginellidae (Mollusca: Neogastropoda) from the, by Barry Roth and Eugene V. Coan, 575-584. [Proc. 4TH SER. Galépagos Islands (Araneae: Salticidae), Darwinneon crypticus, a@ new genus and species of jumping spider from the, by Bruce Cutler, 509-513. Galapagos Islands, The biology of storm petrels in the, by M. P. Harris, 95-165. Galapagos Islands, The Terebridae (Gastrop- oda) of Clarion, Socorro, Cocos, and, by Twila Bratcher and R. D. Burch, 537-566. Galeodes caspius, 356 Gallionella sulcata, 190 (Gastropoda) of Clarion, Socorro, Cocos, and Galapagos islands, The Terebridae, by Twila Bratcher and R. D. Burch, 537-566. Geckos from southwest Asia, with descrip- tions of three new forms and a key to the genus Tropiocolotes, Remarks on some, by Sherman A. Minton, Steven C. Anderson, and Jeromie A. Anderson, 333-362. Genypterus omostigma, 482 Geospiza conirostris, 146 fortis, 102 fuliginosa, 102 magnirostris, 146 scandens, 102 Gibberula coniformis, 580 Glycymeris generosa, 83 Glyptochiton, 567, 571, 572, 573 cordifer, 567, 568, 569). S/05sS/leeo2 kirkbyanus, 573 quadratus, 572 subquadratus, 572 youngianus, 572 Glyptochiton de Koninck 1883 (Mollusca: Polyplacophora), The Carboniferous genus, by Allyn G. Smith, 567-574. Gobius nicholsii, 478 Granula, 579 achenea, 578, 579, 580 insularum, 580, 581, 582 minor, 579, 580, 581, 582 polita, 579, 580, 581, 582 subtrigona, 582 Grapsus grapsus, 105, 131 Gygis alba, 136 Gymnodactylus agamuroides, 43 VoL. XXXVII] caspius, 44, 359 colchicus, 360, 361 danilewsky, 360, 361 fedtschenkoi, 45 heterocercus, 358, 359 persicus, 42 watsoni, 45 Gyphria, 192 Hadrurus, 16, 19 arizonensis, 2, 3, 8, 10, 13 Halipeurus pelagicus, 138 Halosydna breviosetosa, 461 johnsoni, 461 Hanna, G Dallas, Fossil diatoms from the Pribilof Islands, Bering Sea, Alaska, 167-234. Harris, M. P., The biology of storm petrels in the Galapagos Islands, 95-165. Hastula, 563 albula, 561, 562, 563 gnomon, 561 Hemialus, 182 Hemidactylus flaviviridis, 46, 55 Hemidiscus bicurvans, 189 cuneiformis, 189, 216 rotundus, 189, 228 simplicissimus, 189 weissflogi, 187 Hemiechinus megalotus, 356 Hercotheca brevispina, 189 caput-medusae, 196 inermis, 189, 228 mammalaris, 189 Heterodon diadema, 250, 257 Heteroscelus incanum, 139 Heterostephania rothii, 186 Humilaria perlaminosa, 63, 68, 73, 75 Hydrobates pelagicus, 105, 110, 112, 120, iieeis3s. 127 136, 138;+ 140, 141 Hydrocanthus, 244 similator, 244 Hyla, 364, 365, 376 affinis, 364 arenicolor, 364, 375, 376 cadaverina, 363, 364, 365, 366, 368, 369, 370, 374, 375, 376 californiae, 363, 364, 376 INDEX 591 nebulosa, 363, 364, 365, 376 regilla, 363, 364, 375, 376 regilla deserticola, 374 regilla regilla, 364 scapularis, 363, 376 scapularis hypochondriaca, 363, 376 squirella, 364 Hyla cadaverina Cope (=Hyla californiae Gorman), Biosystematics of the can- yon tree frog, by Robert W. Ball and David L. Jameson, 363-380. Jameson, David L., see Ball, Robert W. jumping spider from the Galapagos Islands (Araneae: Salticidae), Darwinneon crypticus, a new genus and species of, by Bruce Cutler, 509-513. Katherinella, 63, 73 subdiaphana, 82 Kenyon, Karl W., see Orr, Robert T. Kogomea, 579 Labidocera trispinosa, 455 Lacerta apoda, 42 pipiens, 334, 338 Late Pliocene mollusks from San Francisco peninsula, California, and their paleo- graphic significance, by Warren O. Addicott, 57-93. Leech, Hugh B., Copelatus glyphicus (Say) and Suphisellus bicolor (Say), water beetles new to California and presum- ably introduced (Coleoptera: Dytis- cidae and Noteridae), 237-247. Leiurus quinquestriatus, 20, 21 Leptopecten latiauratus, 62, 63, 71, 75, 76 Leviton, Alan E., and Steven C. Anderson, Review of the snakes of the genus Lytorhynchus, 249-274. Leviton, Alan E., see Anderson, Steven C. Light, Wiliam J., Polydora alloporis, new species, a commensal spionid (Annelida, Polychaeta) from a hydrocoral off central California, 459-472. Limacina, 427, 429, 435, 444, 457 Liradiscus minutus, 196 Lithodesmium, 455, 490, 499 cornigerum, 491, 494, 495, 499 undulatum, 190, 228 592 CALIFORNIA Lithorhynchus diadema, 258 diadema hirouxii, 258 Litorhynchus ridgewayi, 272 Littorina, 64, 76 mariana, 65 petricola, 59, 62, 64, 65, 74 planaxis, 59, 65 Loukashkin, Anatole S., On the diet and feeding behavior of the northern an- chovy, Engraulis mordax (Girard), 419-458. Lucina nuttalli, 83 Lucinisca nuttalli, 83 Lunatia lewisii, 82 Lythorhynchus, 250 Lytorhynchus, 249, 250, 251, 252, 253, 255, 257, 268, 274 diadema, 249, 253, 255, 256, 257, 258, 259, 260, 261, 263, 265, 266, 274 diadema arabicus, 261, 264 diadema diadema, 258 diadema hoggarense, 258 diadema kennedyi, 265 diadema mesopotamicus, 261, 264 gabrielis, 272, 273 geeks, 220) AS, PSs, OY, AS, 2G, 257, 261, 262, 263, 264, 274 kennedyi, 253, 255, 256, 257, 263, 265, 266, 274 maynardi, 253, 268, 270, 274 monticornis, 269 paradoxus, 253, 255, 256, 266, 268, 269, 270, 274 ridgewayi, 249, 252, 253, 25 266, 267, 268, 270, 271, 2 ridgewayi roseni, 272 Lytorhynchus, Review of the snakes of the genus, by Alan E. Leviton and Steven C. Anderson, 249-274. 255, 256, 266, 267, Mabuya dissimilis, 4.9 Macoma, 63, 66, 70, 72, 77 gouldi, 68 NASUtayOS OO, OW Mey Sen 10h fi, 83 nasuta kelseyi, 62, 70, 73, 77, 78, 79 richthofeni, 68 secta, 63, 68, 73, 77 Macronectes giganteus, 136 Mactra pajaroensis, 83 ACADEMY OF SCIENCES [Proc. 4TH SER. Malvastrum, 146 Margarites pupilla, 59 Marginella albolineata, 577 californica parallela, 577 (Cystiscus) minor, 580, 582 (Cystiscus) polita, 580, 582 (Cystiscus) regularis, 580, 582 (Hyalina) californica, 577 minor, 578, 580, 582 regularis, 582 Marginellidae (Mollusca: Neogastropoda) from the Galapagos Islands and Cocos Island, by Barry Roth and Eugene V. Coan, 575-584. Marginellopsis, 579 ‘Mastogloia maxima, 183 Megasurcula carpenteriana, 70 carpenteriana fernandoana, 70 remondi, 62, 68, 70, 76, 79 Melosira, 190 clavigera, 175, 176, 177, 178, 190, 222 sol, 190 sulcata, 117551164 iia LOO e222 Merluccius productus, 441 Mesocena, 198 corona, 198, 224 Metridia, 455 Microcalanus, 455 Microgecko, 337, 338, 348, 351 helenae, 337, 338, 345, 348, 350, 351, 352 persicus persicus, 353 steudneri, 337 Micrometrus aurora, 475, 476, 483 Microsetella, 455 Minton, Sherman A., Steven C. Anderson, and Jeromie A. Anderson, Remarks on some geckos from southwest Asia, with descriptions of three new forms and a key to the genus Tropiocolotes, 333-362. Mitrella, 62, 64, 67, 70, 82 gausapata, 67, 68 gouldi, 62, 64, 67, 68, 70 Modiolus, 62, 70, 71, 83 rectus, 83 (Mollusca: Neogastropoda) from the Gald- pagos Islands and Cocos Island, Mar- ginellidae, by Barry Roth and Eugene V. Coan, 575-584. VoL. XXXVII] (Mollusca: Polyplacophora), The Carbonif- erous genus Glyptochiton de Koninck 1883, by Allyn G. Smith, 567-574. mollusks from San Francisco peninsula, Cali- fornia, and their paleographic signifi- cance, Late Pliocene, by Warren O. Addicott, 57-93. Monterey County, California, Benthic fishes cast ashore by giant waves near Point Joe, by W. I. Follett, 473-488. Montipora, 460, 469 Mytilus, 62, 71 Naja naja, 54 naja naja, 54 oxiana, 50, 53, 54 Nassa californica, 82 Nassarius, 59, 68 (Caesia) grammatus, 62, 64, 70 californianus, 61, 68, 76, 77, 82 (Demondia) californianus, 62, 68 fossatus, 69, 77 grammatus, 68, 69, 76, 77, 82 mendicus, 77 moranianus, 68, 82 Natrix tessellata, 50, 52 tessellata tessellata, 52 Navicula, 175 bombus densistriata, 188 densistriata, 188 distans, 455 lavareeli/5 019) ornata, 188 semen, 190, 230 Nebo hierochonticus, 20, 21 Neogastropoda) from the Galapagos Islands and Cocos Island, Marginellidae (Mol- lusca:, by Barry Roth and Eugene V. Coan, 575-584. Neon, 509, 511, 512 nigriceps, 509, 512, 513 Neonella, 509, 511, 513 Neptunea, 62, 64, 76, 79 recurva, 82 tabulata, 62, 64, 67, 76 tabulata colmaensis, 67 Nesomimus parvulus, 104, 146, 158 Nesopelia galapagoensis, 146 Neverita, 66, 78, 79 INDEX 593 (Glossaulax), 62, 64 (Glossaulax) recluziana, 62, 64 recluziana, 65, 79 New scorpions belonging to the Eusthenura group of Vejovis from Baia California, Mexico (Scorpionida: Vejovidae), by Stanley C. Williams, 395-418. Nitzschia, 455 northern anchovy, Engraulis mordax (Girard), On the diet and feeding behavior of the, by Anatole S. Loukashkin, 419- 458. Noterus bicolor, 241 Nyctanassa violacea, 102 Oceanites, 147 gracilis, 96, 160 gracilis galapagoensis, 160 Oceanicus, 119, 120, 121, 123, 136, 141, 160 Oceanodroma, 96, 106, 146, 160, 161 castro, 95, 96, 97, 98, 99, 100, 101, 102, 103, 104, 105, 106, 107, 108, 111, 112, 113, 114, 107 1S hT9, Wate, de2s 174 126, 127, 128, 129) 130; 1324133) 134, 136,137, 138) 139, 140; 142,9143. 9145) 1465) 149) 140, 152.9 154) 155-eudsve 158, 161 castro bangsi, 97 leucorhoa, 120, 121, 141 tethys, 95, 96, 101, 133, 134, 135, 136, 138, 142, 143, 144, 145, 146, 147, 148, 149, 155, 156, 157, 158, 159, 160, 161 tethys kelsalli, 143, 159 tethys tethys, 143, 144, 158 Ocenebra interfossa, 62, 64, 66, 76 Ochotona rufescens rufescens, 356 Odostomia, 62, 70 Oikopleura, 429, 435, 440, 442, 445 Oithona, 455 minuta, 447 Olivella biplicata, 62, 68, 69, 76 pedroana, 69 Omphalotheca, 196 ampliata, 196 californica, 196 hispida, 196 laevis, 196 Onacea, 455 594 CALIFORNIA ACADEMY OF SCIENCES On the diet and feeding behavior of the northern anchovy, Engraulis mordax (Girard), by Anatole S. Loukashkin, 419-458. Ophidion barbatum, 482 taylori, 482 Ophidium barbatum, 482 taylori, 482 Ophiodermella graciosana, 62, 68, 69, 75 Ophiodon elongatus, 476 Ophisaurus apodus, 42 Opuntia, 102 Orcinus orca, 160 Oregon, A late Tertiary diatom flora from, by W.N. Orr, Judi Ehlen, and J. B. Zaitzeff, 489-500. Orr, Robert T., Jacqueline Schonewald, and Karl W. Kenyon, The California sea lion: skull growth and a comparison of two populations, 381-394. Orr, W. N., Judi Ehlen, and J. B. Zaitzeff, A late Tertiary diatom flora from Oregon, 489-500. Orthochirus innesi, 20, 21 Otophidium, 482 taylori, 481 Palamnaeus longimanus, 19 Pandora filosa, 83 punctata, 83 Panope abrupta, 83 Parabassigigas grandis, 481, 483 Paracalanus, 455 parvus, 447 Parafavella, 435 Parapsyllus, 138 Parascorpaena, 521, 525 Pecten latiauratus, 59 Pedicularia californica, 461, 470 Pelagodroma marina, 120, 123, 127, 136, 140 Peridinium, 455 Periloba, 146, 153 galapagensis, 146 Periops parallellus schiraziana, 53 parallelus schirasiana, 53 Persicula, 582, 583 Phaeocystis, 430, 436 [Proc. 4TH SER. Phaethon aethereus, 102, 103, 134, 138, 139, 143, 145, 158 lepturus, 99 Philoceanus, 138 Phyllodactylus, 338 Phyllorhynchus, 249, 250, 251, 252, 253, 274 decurtatus, 253 decurtatus perkinsi, 251, 252, 254 Pinctada margaritifera, 460, 469 Pinnularia lata, 191, 214 ruttneri, 191, 230 streptoraphe, 191 Pinus koraiensis, 440 Pleurosigma, 455 Pocillopora, 516 Podon, 428, 434 polyphemoides, 448 Point Joe, Monterey County, Calfornia, Benthic fishes cast ashore by giant waves near, by W. I. Follett, 473-488. Polinices lewisii, 82 Polydora, 459, 460, 470 alloporis, 459, 460, 461, 462, 464, 465, 467, 468, 469, 470 cavitensis, 459, 460, 465, 466, 467, 468, 469, 470 ciliata, 469 hornelli, 459, 460, 465, 467, 468, 469 pacifica, 459, 460, 465, 469 Polydora alloporis, new species, a commensal spionid (Annelida, Polychaeta) from a hydrocoral off central California, by William J. Light, 459-472. Pontinus nigropunctatus, 503, 505 Porites, 516 Pribilof Islands, Bering Sea, Alaska, Fossil diatoms from the, by G Dallas Hanna, 167-234. Prionotus, 448 Protothaca laciniata, 77 staleyi, 63, 66, 70, 73, 74, 76, 77, 83 staleyi hannibali, 73 staminea, 73, 74, 83 tenerrima, 83 Psammophis lineolatus, 50, 53 Pseudoboa carinata, 54 Pseudocalanus, 455 Pseudocardium densatum, 83 VoL. XXXVIT] Pseudocerastes persicus, 273 Pseudopolydora, 466 Pseudopyxilla dubia, 191, 224 tempereana, 191 Pterodroma mollis, 142 phaeopygia, 158 Ptyas mucosus, 50, 53 Puffinus gravis, 136 Iherminieri, 102, 103, 105, 122, 124, 131, $38, 134> 335,°137,138, 139, 140; 143, 145, 154, 158 pacificus, 139, 147 puffinus, 110, 112, 121, 136, 139, 141, 154 puffinus mauretanicus, 136 puffinus puffinus, 136 tenuirostris, 112, 115, 136 Pugnus, 579 Pupillaria pupilla, 62, 63, 76 Puthz, Volker, East asiatic and oriental species of Stenus represented in the collection of the California Academy of Sciences (Coleoptera: Staphylin- idae), 529-535. Pyxidicula, 196 (Xanthiopyxis) alata, 196 (Xanthiopyxis) constricta, 196 (Xanthiopyxis) globosa, 196 (Xanthiopyxis) oblonga, 196 Pyxilla, 191 americana, 192, 224 dubia, 191 Rana cyanophlyctis, 29 ridibunda, 30 sternosignata, 30 Raphoneis asiatica, 193 Rattus rattus, 138, 140 Remarks on some geckos from southwest Asia, with descriptions of three new forms and a key to the genus Tropi- ocolotes, by Sherman A. Minton, Steven C. Anderson, and Jeromie A. Anderson, 333-362. Review of the snakes of the genus Lytor- hynchus, by Alan E. Leviton and Steven C. Anderson, 249-274. INDEX 595 Rhabdonema adriaticum, 192 arcuatum, 192 crozierli, 192, 230 japonicum, 192, 220, 230 Rhaphoneis, 193 amphiceros, 174, P12 222 lancettula, 193, 222 lancettula jutlandica, 193 nitida, 193, 212 rhombus, 193, 226 Rhinogobiops nicholsii, 478, 479 Rhizosolenia, 175, 192, 455 hebatica, 455 Rhombomys, 36 opimus, 37 Rhyncalanus, 455 Roth, Barry, and Eugene V. Coan, Marginel- lidae (Mollusca: Neogastropoda) from the Galapagos Islands and Cocos Island, 575-584. 175) 198. 1192 © fos Sagitta, 429, 435, 440, 444 Salpa, 427, 429, 436, 440, 445 Salpiginella, 435 Sardina pilchardus, 422 Sardinella gibbosa, 422 longiceps, 422 Sardinops caerulea, 419, 442, 456 melanosticta, 420 ocellata, 422 Saxidomus gibbosus, 82 Scalesia, 512 Scaphiopus hammondii, 374 Schizopyga (Nassarius) californianus, 59 Schizothaerus nuttalli, 83 Schonewald, Jacqueline, see Orr, Robert T. Scolopendra, 356 Scorpaena, 501, 503, 505, 506, 521, 522, 524 ascensionis, 501, 502, 503, 505, 506, 507 cookii, 521 laevis, 503 mellissii, 503, 505 orgila, 515, 517, 518, 520, 521, 522, 525 pascuensis, 515, 518, 521, 522, 523, 524, 525, 526 plumieri, 503, 505 scrofina, 501, 503 596 CALIFORNIA ACADEMY OF SCIENCES thomsoni, 521 uncinata, 521 Scorpaenidae) from Easter Island, Three new species of scorpionfishes (Family, by William N. Eschmeyer and Gerald R. Allen, 515-527. Scorpaenodes, 501, 506, 526 africanus, 507 englerti, 515, 518, 524, 525, 526, 527 insularis, 501, 504, 505, 506, 507 littoralis, 526 xyris, 526 Scorpaenopsis, 523 Scorpio maurus, 21 Scorpion fauna of Baja California, Mexico: eleven new species of Vejovis (Scorpi- onida: Vejovidae), by Stanley C. Williams, 275-332. scorpionfishes (Family Scorpaenidae) from Easter Island, Three new species of, by William N. Eschmeyer and Gerald R. Allen, 515-527. scorpionfishes, Two new Atlantic, by William N. Eschmeyer, 501-508. (Scorpionida: Vejovidae), New scorpions belonging to the Eusthenura group of Vejovis from Baja California, Mexico, by Stanley C. Williams, 395-418. scorpions belonging to the Eusthenura group of Vejovis from Baja California, Mexico (Scorpionida: Vejovidae), New, by Stanley C. Williams, 395-418. scorpions, Birth activities of some North American, by Stanley C. Williams, 1-24. Sebastapistes, 521, 525 Sebastes, 476 auriculatus, 482 flavidus, 475, 477, 483 helvomaculatus, 475, 477, 478, 483 rosaceus, 475, 477, 478, 483 Sebastodes, 476 Semiopyla, 511 Siliqua, 83 alta, 83 Simotes diadema, 257 Skeletonema, 455 costatum, 455 Skull growth and a comparison of two popu- lations, The California sea lion:, by [Proc. 4TH SER. Robert T. Orr, Jacqueline Schonewald, and Karl W. Kenyon, 381-394. Smith, Allyn G., The Carboniferous genus Glyptochiton de Koninck 1883 (Mol- lusca: Polyplacophora), 567-574. Socorro, Cocos, and Galapagos islands, The Terebridae (Gastropoda) of Clarion, by Twila Bratcher and R. D. Burch, 537-566. Solen, 63, 72 perlaminosa, 63 rosaceus, 72 sicarius, 63, 72, 74, 76, 83 Sphalerosophis diadema schirazianus, 50, 53 spionid (Annelida, Polychaeta) from a hy- drocoral off central California, Poly- dora alloporis, new species, a com- mensal, by William J. Light, 459-472. Sow, 2, 7, dil, as alaskana, 173 albaria, 71, 83 albaria coosensis, 62, 68, 70, 71, 74 albaria goodspeedi, 71 catilliformis, 71, 77, 83 falcata, 83 mercedensis, 62, 71, 77 mossbeachensis, 80, 81, 83 Squatarola squatarola, 139 Standella californica, 83 planulata, 83 Staphylinus cicindeloides, 533 Stellio agrorensis, 31 caucasicus, 35 erythrogastra, 36 himalayana, 37, 38 lehmanni, 38 Stenodactylus, 338 scaber, 45 scincus, 47 Stenus, 529 alienus, 529, 532 ambiguus, 533 arisanus, 532 basicornis, 533 bernhaueri, 534 comma, 529 duplex, 531 flavidulus paederinus, 532 formosae, 532 VoL. XXXVIT] formosanus, 532 forterugosus, 532 gestroi, 532 gestroi taiwanensis, 532 (Hemistenus) javanicus, 534 (Hemistenus) maximiliani, 534 (Hypostenus) ambiguus, 533 (Hypostenus) cicindeloides, 533 (Hypostenus) mercator, 532 (Hypostenus) nothus, 533 (Hypostenus) spinosus, 533 (Hypostenus) subtropicus, 533 (Hypostenus) tropicus, 533 insulanus, 530, 531, 532 kwantungensis, 532 melanarius verecundus, 532 mercator, 532 (Mesostenus) bernhaueri, 534 miwai, 532 nothus, 533 (Parastenus) bicolon javanicus, 534 (Parastenus) javanicus, 534 rugipennis, 532 rugosipennis, 532 sauterianus, 531, 532 sharpianus, 532 spinosus, 533 stigmatipennis, 532 submaculatus, 532 submarginatus, 532 subtropicus, 533 taiwanensis, 532 tropicus, 533 virgula, 532 Stenus represented in the collection of the California Academy of Sciences (Cole- optera: Staphylinidae), East asiatic and oriental species of, by Volker Puthz, 529-535. Stephanopyxis, 490, 497 appendiculata, 174, 176, 177, 178, 193, 222, 230, 491, 493, 494, 495, 497 turris, 194, 443, 455 Sterna fuscata, 107, 109, 136, 137, 143, 147, 158 Stictodiscus, 490, 498 buryanus, 491, 494, 495, 498 storm petrels in the Galapagos Islands, The biology of, by M. P. Harris, 95- 165. Street Expedition to Afghanistan, 1965, Am- INDEX 597 phibians and reptiles collected by the, by Steven C. Anderson and Alan E. Leviton, 25-56. Striatella crozieri, 192 Strioterebrum pedroanum philippianum, 561 Sula dactylatra, 102, 103, 145 nebouxil, 102, 103, 134, 143 sula, 134, 143, 145, 146, 147 Suphis, 243 bicolor, 243 Suphisellus, 241, 242, 243, 244, 245 bicolor, 237, 239, 241, 242, 243, 244, 245 gibbulus, 242, varicollis, 243 Suphisellus bicolor (Say), water beetles new to California and presumably intro- duced (Coleoptera: Dytiscidae and Noteridae), Copelatus glyphicus (Say) and, by Hugh B. Leech, 237-247. Synanceia, 523 Synanceja horrida, 503 Synedra, 194 nitzschoides, 194 243, 244 Tapes staminea, 83 tenerrima, 83 Tarletonbeania crenularis, 474 Tellina, 62, 66, 70, 72, 78, 79 bodegensis, 72 lutea, 66, 72, 78, 79 (Peronidea), 62 (Peronidea) lutea, 62 Temora, 455 Teratoscincus bedriagai, 46 scincus, 4:7 scincus keyzerlingii, 47, 48 scincus scincus, 47, 48 zarudnyl, 47, 48 Terebra, 537, 538, 544, 546, 547, 548, 549, 552, 553, 554, 556, 561 aciculina, 561 albemarlensis, 559, 560, 561 albicostata, 557 albula, 561 armillata, 544, 556, 557 armillata sheppardi, 544 berryi, 557, 558 On ie.) CALIFORNIA ACADEMY OF SCIENCES bipartita, 561 casta, 561 concava, 541 constricta, 541 contracta, 541 crenulata, 555, 556, 557 crenulata booleyi, 555 eburnea, 561 fimbriata, 555, 556 flammea, 552 frigata, 541, 544, 547, 548, 549 gabbi, 554 galapagina, 547, 549 gatunensis, 539 gatunensis kugleri, 540 gnomon, 561 gracilis, 547 grayl, 547 (Hastula) casta natalensis, 561 hertleini, 542, 543, 544 incolor, 561 insignis, 554 interlineata, 555, 556 jacquelinae, 540, 541, 542, 544 lingualis, 554 litorea, 559, 560 loroisii, 554 macrospira, 554 maculata, 544, 555 maculata confluens, maculata maculata, maculata roosevelti, maculosa, 544 marginata, 557 medipacifica, 561 medipacifica melior, 5 nelsoni, 544 ornata, 551, 544 544, 545, 546, 547 545, 546, 547 553 panamensis, 539 philippiana, 561 plicata, 548, 549, 550, 560, 561 purdyae, 539, 540 robusta, 554, 555, 557 540 simplex, 561 stohleri, 538, 540 strigata, 547, 551, 552 (Strioterebrum) berryi, 558 (Subula) roosevelti, 546 tiarella, 544 zebra, | shyana, [Proc. 4TH SER. Terebridae (Gastropoda) of Clarion, Socorro, Cocos, and Galapagos islands, The, by Twila Bratcher and R. D. Burch, 537-566. Tertiary diatom flora from Oregon, A late, by W. N. Orr, Judi Ehlen, and J. B. Zaitzeff, 489-500. Testudo horsfieldii, 30 Thais, 66, 67 (Demondia) californianus, 68 emarginata, 67 emarginata ostrina, 67 etchegoinensis, 67 lamellosa, 67, 77 lima, 67 (Nucella), 62, 70 (Nucella) emarginata ostrina, 62 (Nucella) lima, 62, 70 (Nucella) trancosana, 62, 64, 68 ostrina, 59, 63, 67, 76 trancosana, 59, 66, 67, 74, 76, 77 Thalassionema, 194 nitzschoides, 174, 175, 176, 177, 194, 216 Thalassiosira, 194, 438, 455 nordenskioldii, 194 punctata, 174, 175, 176, 177, 194, 204 Thalassiothrix, 194, 455 nitzschoides, 194 Three new species of scorpionfishes (Family Scorpaenidae) from Easter Island, by William N. Eschmeyer and Gerald R. Allen, 515-527. Tomyris oxiana, 53 Topaginella, 579 Trachurus symmetricus, 441, 457 Trachyneis aspera, 174 Tresus nuttalli, 77, 83 pajaroanus, 62, 71, 74, 77, 83, 85 Triceratium, 448, 449 condecorum, 195, 218, 228 montereyi, 195, 218 tripolaris, 195 validum, 195, 228 Tripodiscus laxus, 181 Tropiocolotes, 333, 337, 338, 345, 348, 351 353, 357 depressus, 337, 338, 339, 340, 344, 347, 350, 356 VoL. XXXVII] helenae, 334, 337, 339, 344, 345, 346, 348, 349, 350, 351, 352, 353, 354 heteropholis, 338, 339, 352, 357, 358 nattereri, 337, 339 occidentalis, 337 persicus, 340, 344, 348, 350, 351, 354 persicus bakhtiari, 339, 342, 344, 350, 351, 352, 353 persicus euphorbiacola, 337, 338, 339, 340, 343, 344, 350, 352, 354, 355, 356 persicus persicus, 339, 340, 344, 348, S50 o55 scortecci, 337, 339, 340 somalicus, 337 steudneri, 337, 338, 339, 340, 344, 345, 350 tripolitanus, 352, 358 tripolitanus algericus, 337, 338, 340 tripolitanus occidentalis, 340 tripolitanus somalicus, 339 tripolitanus tripolitanus, 339 Tropiocolotes, Remarks on some geckos from southwest Asia, with descriptions of three new forms and a key to the genus, by Sherman A. Minton, Steven C. Anderson, and Jeromie A. Ander- son, 333-362. Tupinambis bengalensis, 49 Turbonilla, 479 Two new Atlantic scorpionfishes, by William N. Eschmeyer, 501-508. 337, 338, 339, 340, 345, Uroctonus mordax, 2, 3, 13, 20 Varanus bengalensis, 49 bengalensis bengalensis, 49 (Indovaranus) bengalensis bengalensis, 49 Vejovis, 14, 16, 20, 21, 275, 277, 281, 286, 290,207. 302; 307, 313, 317, 322, 325, 331, 395, 396, 397, 401, 403, 407, 410 bruneus, 298, 317, 318, 319, 320, 321 coloradensis, 397, 401, 402, 403, 404 confusus, 3, 13, 303, 401 diazi, 302, 307, 309, 310, 311, 395, 410 diazi diazi, 415 diazi transmontanus, 4.15, 416 INDEX 599 eusthenura, 308, 395 galbus, 403, 407, 408, 409 gertschi, 286, 287 grandis, 277, 279, 280, 282 gravicaudus, 325, 327, 328, 329, 330 harbisoni, 281, 284, 285, 286, 287 hoffmanni, 313, 315, 316, 317, 318, 395, 415 hoffmanni fuseus, 413, 414 hoffmanni hoffmanni, 413 : insularis, 318, 322, 323, 324, 325, 326 mesaensis, 3, 13, 277, 281 minimus, 3, 13 minutis, 286, 287, 288, 289, 290, 291 (Paruroctonus), 277 pumilis, 297, 299, 301, 302, 331 punctatus punctatus, 290, 292 punctipalpus, 318, 322 schwenkmeyeri, 302, 304, 305, 308; 3135) 395; 397,403) 410 SPIMISELUS,) Jen 3504) D5) One al Onl emnlese 14s 15) doest val 8, 326 terradomus, 403, 405, 406, 407 vachoni, 277, 281 viseainensis, 395, 410, 411, 412, 413 vittatus, 290, 294, 295, 298 vorhiesi, 2, 3, 4, 5, 6, 7, 9, 10, 11, 14, 18 waeringi, 397, 398, 399, 400, 401 wupatkiensis, 281, 282, 286 Vejovis from Baja California, Mexico (Scor- pionida: Vejovidae), New scorpions belonging to the Eusthenura group of, by Stanley C. Williams, 395-418. Vejovis (Scorpionida: Vejovidae), Scorpion fauna of Baja California, Mexico: eleven new species of, by Stanley C. Williams, 275-332. 306, Venus, 83 Vipera lebetina obtusa, 50, 54, 356 obtusa, 54 Volvarina, 577 albolineata, 577 nyssa, 576, 57 taeniolata, 577 1-578 taeniolata rosa, 577, 578 taeniolata taeniolata, 577 water beetles new to California and pre- sumably introduced (Coleoptera: Dy- 600 CALIFORNIA ACADEMY OF SCIENCES tiscidae and Noteridae), Copelatus glyphicus (Say) and Suphisellus_bi- color (Say), by Hugh B. Leech, 237- 247. Williams, Stanley C., Birth activities of some North American scorpions, 1-24. Williams, Stanley C., New Scorpions belong- ing to the Eusthenura group of Vejovis from Baja California, Mexico (Scor- pionida: Vejovidae), 395-418. Williams, Stanley C., Scorpion fauna of Baja California, Mexico: eleven new species of Vejovis (Scorpionida: Vejo- vidae), 275-332. Xanthiopyxis, 175, 196 cingulatus, 197 globosa, 195, 196, 224 [Proc. 4TH SER. lacera, 188 lohmani, 196, 224 ovalis, 174, 175, 176, 196, 197, 224 umbonata, 197, 228 Yoldia, 83 cooperi, 83 thraciaeformis, 83 Zauizeff, J. B., see Orr, W. Ne Zalophus californianus, 104, 382, 383, 384, 392 californianus californianus, 381, 385, 386, 387; 388, 389, 301,1388 californianus wollebaeki, 385, 386, 391 Zamenis rhodorhachis, 52 Zirfaea, 61, 63, 66, 74 ERRATA for B. andersoni read B. andersoniti. for Agama himalayanum read Agama himalayana. for Discinisca cummingi read Discinisca cumingi. for Spisula labaria coosensis read Line 14 from top: for Arachnoidiscus ehrenbergi read Arachnoidiscus ehrenbergit. Line 3 from bottom: for Coscinodiscus pustulosus read Coscinodiscus pustulatus. Line 15 from bottom: for Thalosiosira punctata read Thalassiosira punctata. for Coscinodiscus insignis read Cosmiodiscus insignis. for Thalossiosira punctata read Thalassiosira punctata. Bottom line: for Stephanopyxis apendiculata read Stephanopyxis appendiculata. for Diploneis ornatus read Diploneis ornata. 50. Line 19 from top: for Acantiophis read Acontiophis and for Acantiophis para- Page 2. Line 14 from top: for ‘is’ read ‘are’. Page 2 Lines 8 and 9 from top: Page 35. Line 21 from top: for A. himalayanum read A. himalayana. Page Line 1 of figure caption: Page 63. Line 5 from bottom: Page 70. Line 5 from bottom of plate explanation: Spisula albaria coosensis. Page 102. Line 4 from top: for (flammeus) read (Flammeus). Page 161. Line 9 from bottom: for ‘in the fleas’ read ‘and the fleas’. Page 174 Page 175 Page 176 Page 177. Line 18 from top: Page 177. Line 21 from top: Page 178 Page 206. Line 4 from bottom: Page 224. Line 14 from top: for Chaetoceras read Chaetoceros. Page 25 doxus read Acontiophis paradoxa. Page 337. Top line: for B. abundhabi read B. abudhabi. Page 353. Line 3 from bottom: es) IBRARY |=> —] for P. persicus read T. p. periscus. iii mice Pr es nt Se , Ln” se LihctgN e e ~tmn ov Pe owt Se ee 2 A ee Print Daan fe FER aaaamee << FN EOE LN AA OOD AOL RG Se CP on LE LOOL APD PILI LDL ALATA et Per: ; ee eBanecega neta er eos . v4 - Naren teens PEP a Sie Gaia gS ure ~eanihs eee ae ete PL I 5 PPR a ee een one