z ’ ’ Ayer y serine Th Toe Se hw > i : \ y . BAT vate eh anes vies ies ’ s per eeternere ew ten} a iw) ss : ARES Crrrres is tins . yo toys Wee rane a rere a : at vis yee gin tala sev bee Ae ay ah ety veyeyns tes uae iy tee : a My ss oan ourpw erreass: © Tee Duras : NT A oe Mag Re ek SEN : : rere wes Css oA apere writ Sart Cor thr Patna re . . pret guest ae by yh ees Wess tag REDS ‘ - is SES WE he Nien SPA Pad er. . HEPAT Rd, epee Nt a i NEsss LACAN Gals tote oe 4, HA Cade Taner] fey tess Mats , : Ph 6 KLh We, > > 5 5 % ela la pd el Ha Moo gai ’ . Ow, Pre ne ere . i fanny etre Oi Be et Pare jak gt re jeren bas Weer me MP ages eh CA Mewes 9 hp ak wiry! pe an ee Pia ae ee teed 240-3 PKG tS aye i terebeurery eoats Jectiat ont od ¥, Peosvet a Bae CANE TIN 9: 9 hsyae A hyptet ta: Scant oy dak yo die ae “9 a ihegs a whe jee’ re iu at oe ress lat rh evar t ierorriceat yates Pieare dia rs iosest oe bt ek bat od peek orl BI e hohe. fe 49% abby if e ote esd Sete tata re eae et oH dace he studi, ap od a tglduraittate 1 : st Re gts Paci 4s ro Ry ret at f gue A: ae a ai Ly ed re cies lass je gales a yey P rat tf fet C acne ag deat vp Raita att Aid 4 sh tat ja, jatlan the ileeoy te pe oy dee aes 4b aie be vas ry Shee v3 pda dle connate eM gs vated Shite, 4, 452 ana t , meh po Beate Ss Ct) Pi a os ‘ . on tut By “ ie oon hye 4, ret ob iy ype af x3 ee De papacy He 1a [ ip wa 4 Was i} I ae i r Oh. 7 ne Faye ree | i - ie ie ‘og : > 'a~e 7 ates yeah) A i See Py a ‘A Pe, hak wd Fn ales us ' 4 _ : Hes: 7 B° " ie.’ ae. 7 ny ¥ 7 4 re Mie igh : i ee ; Bat 4 ales ale it e “ " iv i, { J Ut ; { mt { it ii \ Hoe ull sy he) i iy Wl Lon i Ay ht a z \ Lg Te Doaae iuiiarry | ‘ a i Ayn | ; Pg . a vn Cheha age PROCEEDINGS OF THE California Academy of Sciences FOURTH SERIES VOLUME XXXVIII — estschrift~ ee UL Se rague Myer Sn Honor gf Kofi Dirthday SAN FRANCISCO PUBLISHED BY THE ACADEMY 1970 | Mari ne Ay alc | FEB 1 2 1971 WOODS HOLE, MASS. a> Pe Hh gas eed aay ‘oh gee et ak Tah Heal a ea Se VN Washington Biologist’s Field Club; Retirement Photo 1970 Plummers Island, Md., 1933 Photo by A. K. Fisher With HIH Crown Prince Akihito in the Stanford Collection 1967 Palo Alto Times Photo GEORGE SPRAGUE MYERS PROCEEDINGS OF THE California Academy of Sciences FOURTH SERIES VOLUME XXXVIII Aestschrift— jor George Sprague Myers In Honor of Hoff Dirthday SAN FRANCISCO BLISHED BY THE ACADE 1970 COMMITTEE ON PUBLICATION GEORGE E. LinpsAy, Chairman Epwarp L. KeEsseEL, Editor Lro G. HERTLEIN NUMBER It 10. Ile 12: CONTENTS imtraduction.. By EARL S. HERALD << 2 ee On the natural history of George Sprague Myers. By PGND ee e VAL FORD 4-2) eo ee ee Annotated chronological bibliography of the publications of George Sprague Myers (to the end of 1969) _....... A new species of the doradid catfish genus Leptodoras, with comments on related forms. By JAMES E. BOHLKE — Systematics of the genus Hemitriakis (Selachii: Carcharhini- dae), and related genera. By L. J. V. ComMPAGNO __________- Notes on the natural history of snipe eels. By Gites W. ite caGle OWIEV TAG AL SIGARTRY se 522 Be The zoogeography of the herpetofauna of the Philippine Islands, a fringing archipelago. By WALTER C. Brown and PSN Tame Ose ANDY VANZA ees 00h oa 2 Ae ee Tropical shelf zoogeography. By JoHN C. Briccs A new species of glandulocaudine characid fish, Hysteronotus myersi, from Peru. By STANLEY H. WEITZMAN and JAMIE BRIBEIONERR SON) teeeete ORS OS 8 ea ee ee ee Rediscovery of the loricariid catfish, Acestridium discus Haseman, near Manaus, Brazil. By Ropert L. HAssur The amphibians and reptiles of Afghanistan, a checklist and key to the herpetofauna. By ALAN E. LEviToN and STEVEN (CAP NINE RSON a= seeks eter a ee oS eee Explosive spread of the oriental goby Acanthogobius flavi- manus in the San Francisco Bay-Delta region of California. By Martin R. BriTTAN, JOHN D. Hopxkirk, JERROLD D. GoOnNERS: and IVIIGHAEL IMIARTIN 2 = ee Some nerve patterns and their systematic significance in paracanthopterygian, salmoniform, gobioid, and apogonid fishes ay WARREN C. FREIBOFER 2.22 ee 19-52 53-62 63—98 99-104 105-130 131-138 139-156 157-162 163-206 207-214 215-264 13% 20. Zale Onda WS CONTENTS A new gymnotoid fish from the Rio Tocantins, Brazil. By MAARTEN JKORRINGA’ 22.3.6 eee On the trail of the golden frog: with Warszewicz and Gabb im Central America, By Av ME SAVACr EEE Teleost hybridization studies. By CLARK HuBBs — A revision of the fishes of the genus Notothenia from the New Zealand region, including Macquarie Island. By HucH ED Wt ce ee How many recent fishes are there? By Dante, M. CoHEN Description of a new subspecies of Rhabdophis auriculata in the Philippines, with comments on the zoogeography of Mindanao Island. By ALAN EE VarONs see A reinterpretation of the teleostean fish order Gobiesoci- formes: By, WALETAM Ae GOST INI: oisces seem oeeen see Scale-eating American characoid fishes with special reference to Probolodus heterostomus. By Tyson R. ROBERTS —— The dorsal and anal spine-locking apparatus of surgeon fishes (Acanthuridae)= By jars —————————E Amputation and replacement of marginal spines in ctenoid percoid: scales” By Llowarn Mic ———————— EE Size and distribution of proteins in elasmobranch plasma. By ROBERT J. Heckiy and BARS. HERALD 2a 265-272 273-288 289-298 299-340 341-346 347-362 363-382 383-390 391-410 411-414 415—420 421-438 INTRODUCTION The faculty appointment of George Sprague Myers as a Stanford associate professor in 1936 and full professor in 1938 marked the resurgence of what had informally been recognized as the Jordan school of ichthyology. In the thirty- four years since that time more than 104 graduate and special students as well as a large number of undergraduates, have come under Professor Myers’ guid- ance. Although the majority of these students were involved in the study of fishes, a very respectable number specialized in amphibians and reptiles. During this same period the Stanford Ichthyological Bulletin came into prominence as did a series of important herpetological reports published as Occasional Papers of the Natural History Museum of Stanford University. On behalf of the many students and colleagues who have carried out their studies at the Stanford Natural History Museum, the authors participating in this volume respectfully dedicate this Festschrift to George Sprague Myers in appreciation of his helpful leadership in the field of systematics of the lower vertebrates. EARL S. HERALD : August 1970 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Festschrift for George Sprague Myers Vol. XXXVIII, No. 1, pp. 1-18. December 31, 1970 ON THE NATURAL HISTORY OF GEORGE SPRAGUE MYERS!’ By Lionel A. Walford Sandy Hook Marine Laboratory Highlands, New Jersey 07732 At a very young age, George Sprague Myers manifested those qualities which were to remain his mark of distinction—an extraordinary sensitivity to the beauty of order in Nature, a boundless capacity to learn about what interested him, and a zest for collecting, arranging, and reasoning how things must fit to- gether. Given such an endowment, the place where he was born and spent his boyhood—Jersey City, New Jersey—and the epoch of his birth—early part of the twentieth century—were peculiarly right for guiding him towards and into his life work. For at that time many of the nineteenth century systematic zoologists were still flourishing (David Starr Jordan, for example) and there were plenty of roads from Jersey City leading to their doors and also to back country that was still unspoiled and beckoning. Jersey City in 1905, the year of Myers’ birth, was already well established as part of what was to grow into the great Atlantic megalopolis. Like its neigh- boring satellite communities, it did not share any part of New York’s splendor, 1T am deeply grateful to Mrs. Mary S. McKenzie, for providing information on the family background and early history of her nephew; and also to Dr. Alan E. Leviton through whose good offices I have been permitted to consult and use several autobiographical fragments which Myers wrote at various times and deposited along with his extensive file of biographical information on zoologists at the California Academy of Sciences. [1] 2 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. yet was close enough to attract the most unbeautiful features of industrial de- velopment. The house where Myers was born and spent much of his boyhood was a three-story brick structure at 283 Grove Street, directly opposite the front of the city hall. Even in 1905, this was part of a dwindling genteel neighborhood in the process of being eaten away by slums that were surrounding it. The biota of the Grove Street house was typical of relic residential sections in northeastern crowded cities. Near the woodshed in the back yard grew a large Rose-of-Sharon bush, a lilac which annually put out a few flowers, some South American spider plants (Cleome). There were a few Ailanthus trees in nearby back yards, and an ancient linden that grew out of a hole in the flagstone side- walk near the curb. Of insects that aroused some interest in young Myers, albeit a short-lived one, were bees of various sorts. ‘““What could you do with a bee?” he asked. The only birds were English sparrows. Mammals consisted of cats, dogs, rats, and bats which issued at dusk from a nearby church. One of the cats was a pet, the first of a series which Myers has had almost continuously ever since. Myers’ introduction to fish life was a minnow, probably Chrosomus neogaeus, living in a spring on a farm to which he was taken during a summer trip to Maine at the age of seven. Next he met some entrancing goldfish in a ‘pet shop where his mother and Aunt Marvy often took him during their visits to New York. He actually got to possess one or two goldfish in a bowl at various times. These experiences were no more remarkable than any other young city boy might have during his natural history phase. To young Myers, however, this was no phase, but rather a prologue to his great lifelong interest. This really began at age 12, when he first attended the Jersey City Aquarium Society’s annual ex- hibition in the public library. Enthralled by the colorful fresh-water fishes from all over the world, he promptly joined the Society and began to accumulate aquariums in which he kept not only exotic fishes but also native ones (Umbra, Fundulus, Enneacanthus, etc.). These he caught on trips made by train or trolley car to various places in northern New Jersey. At the same time he collected and kept amphibians and reptiles, these being as interesting to him as fishes. When he was about 15 years old, attending Lincoln High School in Jersey City, he sought advice of his biology teacher about a trip he was planning to the Pine Barrens of Lakehurst, New Jersey, to collect the beautiful rare tree frog Hyla andersonii. The teacher, rather out of his depth, suggested that Myers dis- cuss his problem with Dr. G. K. Noble at the American Museum of Natural His- tory in New York. By following that advice, Myers became introduced to the world of research zoologists. Noble, impressed, of course, introduced him to A. I. Ortenburger; and when the two had become well enough acquainted with this interesting young fellow, they took him on the last of Noble’s Lakehurst trips VoL. XXXVIII] WALFORD: HISTORY OF GEORGE S. MYERS 3 to study the life history of Hyla andersonii. Through Noble, Myers came to know Karl P. Schmidt, and, in the Museum’s fish department, John Treadwell Nichols, Arthur W. Henn, and Eugene W. Gudger. It was not long before he was spending so much time on his cold-blooded vertebrates that his school work slipped badly, for he took full advantage of the proximity of New York to meet most of the old guard zoologists at the Museum, the Zoo, and the Aquarium— Henry Fairfield Osborn, Bashford Dean, F. A. Lucas, Walter Granger, W. D. Matthew, Frank M. Chapman, Carl Akeley, Robert Cushman Murphy, Roy Chapman Andrews, C. H. Townsend, William Beebe, John Tee-Van, and Charles M. Breder. And when a young fellow from the University of Virginia was selected by Noble to go with Andrews to collect reptiles in China, Noble and Myers went out to Plainfield, New Jersey with him to teach him how to collect salamanders. His name was Clifford H. Pope. One day in 1924 while at the Museum, Myers was introduced to Dean Carl H. Eigenmann of Indiana University, the principal worker on the systematics of the fresh-water fishes of South America. Myers had then published a few short papers on fishes and had become especially interested in those of tropical Amer- ica. The result was an invitation by Eigenmann to come to Indiana as a student and have part of the cost defrayed by part-time work in caring for the Indiana fish collection. Myers had not done well in high school, and lacked several credits for grad- uation. Moreover, with the examples before him of several then well-known zool- ogists who had had no university preparation, he was uncertain even whether to go to college. However, he says that Noble gave him a thorough tongue-lashing about his refusal to get the necessary schooling, and this, coupled with Eigen- mann’s offer, decided him to go. In lieu of his missing high school credits, Dean Eigenmann arranged to have him granted credit for the research he had already_ accomplished. He has always felt a great deal of gratitude to Noble and Eigen- mann, without whose help his professional career might have been aborted. At Indiana, he neglected his academic studies to accomplish some field work and to complete a synopsis of the amphibians and reptiles of Indiana. At the same time he got a superb introduction to South American fishes, and also to curatorial methods for preserved research collections. Aside from Eigenmann, the man on the Indiana faculty who had the most influence on Myers (though Myers took no formal courses from him) was the entomologist, Professor Alfred C. Kinsey (later the student of human sexuality), whose forward-looking views of evolution and systematics were then finding expression in studies of cynipid wasps. From Kinsey, Myers began to gain a very broad view of systematics as a synthesis of the comparative aspects of all other biological disciplines, a view that finally found expression in a review paper in 1930, + CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. After Myers had been at Indiana one year, Eigenmann fell ill and was taken first to Florida and then to the San Diego region of California for his health. Since he obviously would not be able to return to the University, there seemed little reason for Myers to remain. However, he unpacked the large Ternetz South American collections as they arrived, and, with Eigenmann’s permission, described some of the novelties. Stanford University, founded in 1891, had become a center of research on fishes and their habits through the influence of the university’s first president, David Starr Jordan, and the first chairman of its zoology department, Charles H. Gilbert. In 1926, both men had been long retired, but Jordan was still active and he had noticed the papers published by Myers. When he heard of Eigen- mann’s illness, Jordan wrote to Myers asking whether he would like to transfer to Stanford with the same sort of part-time assistantship he had had at Indiana. Jumping at the opportunity, Myers arrived in California in September, 1926. It was shortly after his arrival at Stanford that I first met him there. As a biologist at the California State Fisheries Laboratory at Terminal Island, I had just begun a study of the California barracuda, and took advantage of an op- portunity to spend a few days at the Natural History Museum to search its library and to study the sphyraenid material in the fish collection. The Museum at that time was a center of quiet excitement such as I will never forget, for many of the biology faculty members focused their interests there, and the principal interest was the study of fishes. There is a vivid picture in my mind’s eye of a pleasant, fine-looking, young man, an undergraduate, working as an assistant in the Museum. “May I help you?” he asked. “Perhaps,” I replied, cataloging him as a library assistant. When I told him what I wanted, he poured a steady stream of information out of his head: “Of course, you must already have the essential papers about Sphyraena argentea and S. ensis. You may have missed Sphyraena idiastes. We have the original 1903 description by Heller and Snod- erass.” And so on through the whole family Sphyraenidae around the world. No, he was not working on sphyraenids himself. Never had. It seemed to me as I talked to this enthusiastic modest fellow that he knew everything about everything. He was already a learned person when most people are scratching about, trying to make up their minds what they want to learn about. The area surrounding the Stanford lands was then open country, and the University community (affectionately known as “The Farm” by faculty and student alike) moved at a relatively leisurely pace. Under Jordan’s influence it had been, and still was, a great center for studying systematic zoology, especially fishes. Jordan was still working on fossil fishes. John Otterbein Snyder was chairman of the zoology department, with Edwin C. Starks as morphologist, Harold Heath as invertebrate zoologist, G. C. Price as embryologist, and Isabel McCracken, R. W. Doane, and G. F. Ferris as entomologists. Aside from courses VoL. XXXVIII] WALFORD: HISTORY OF GEORGE S. MYERS 5 in other departments, Myers came to know Snyder, Starks, Heath, and Ferris best, as well as H. G. Schenk and S. W. Muller in the geology department, in which he eventually took his doctorate minor. The Dudley Herbarium, the Entomological Collections, and the Zoological Collections were then housed—temporarily, it was said—in the south end of the Stanford Museum where they remained as the ‘Natural History Museum,” and later the “Division of Systematic Biology,” for the entire period of Myers’ active association with the University. Professor LeRoy Abrams was in charge of the Herbarium, Ferris of the insects, and Snyder of the Zoological Collections. Although Snyder was his special mentor and friend, Myers visited Jordan, at Jordan’s warm invitation, at least once a week. Snyder, like Gilbert before him, had almost deserted systematics to work on the migrations of salmon and steelhead trout. In 1928, Willis H. Rich was appointed to the department, to teach ecology and fishery biology. A biweekly seminar in fishery biology in 1928-29, attended by Snyder, Rich, Frank Weymouth, Starks, and about a dozen serious students including Myers, most of whom were to become leaders in the study of fishes, was a whirlwind of lively discussion and argument among and between students and professors, such as I have rarely experienced since. Because the Stanford group then most interested in the theory of systematics was led by Schenk in geology, Myers gravitated to that quarter. In connection with one of Schenk’s seminars, Myers published a review of a recent botanical revision in Schenk’s Micropaleontology Bulletin, in which the ideas he had de- veloped after contact with Kinsey at Indiana and Schenk at Stanford were synthesized into a view of systematic biology that was unusually broad for its day. When it is recalled that Myers was then an undergraduate student, without much knowledge of what was then being done on theoretical systematics by several isolated men or groups in America and Europe, his statement is remark- able. Myers’ university work went slowly, not only because of his part-time em- ployment but also because he was able only slowly to force himself to neglect extracurricular work in ichthyology and herpetology enough to get good marks. One objection to granting him a bachelor’s degree was that he had not taken enough required courses in English. Snyder later confided to me (with much amusement), that he had demolished this hurdle simply by waving in the faces of the objecting English Department’s faculty members a handful of the publica- tions Myers had produced since he entered college, saying, “Look, see how much he has written!’’ He had published a good deal by the time he was granted his A.B. degree in June, 1930, seven years after he entered Indiana as a freshman. After that, things went more rapidly. He obtained an A.M. in 1931, and his PhD sine june; 1933: 6 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Of his Stanford student years, Myers has said that he learned more mor- phological zoology from Harold Heath and Tage Skogsberg at the University’s Hopkins Marine Station (where he spent the summer of 1929), than from all others. Nevertheless, Snyder, Starks, and Rich ranked high in his training years. For four years he saw and talked with Jordan almost weekly, gaining a great wealth of information about ichthyological workers and history. In 1928, Dr. Albert W. Herre, a former pupil of Jordan, was appointed to a non-faculty position as Curator of the Zoological Museum at Stanford with an arrangement by which he would be retired only upon the President’s pleasure. Myers says that he owes much broadening and maturing to Herre’s influence, not only while he was a student, but also after he returned to Stanford in 1936. In 1933, Myers was appointed Assistant Curator in charge of the Division of Fishes at the U.S. National Museum, with a first assignment to pack and ship many National Museum fishes that had been at Stanford in the hands of the late Charles H. Gilbert. He arrived in Washington in March, 1933. At the National Museum, Myers took charge of the most important ichthy- ological resarch collection in America but one which had suffered from nearly 40 years of impoverishment and neglect. Moreover, the great financial depression of the 1930’s had worsened the situation so much that for the first two years of Myers’ tenure, the Division lacked even the services of a typist. Besides Myers, the staff consisted only of one elderly but enthusiastic scientific aid, Earl D. Reid2, and a laborer who cared for the alcoholic collections. For the next three years, Myers and Reid spent most of their time in sorting, bottling, and register- ing an enormous backlog of specimens and putting the Division’s offices, files, and records into working condition. During most of this period they trained and supervised squads of up to a dozen temporary workers at a time, these having been supplied free to government bureaus by successive federal agencies set up to relieve unemployment. Although there was precious little time for research during these three years, Myers and Reid initiated a survey of the fresh-water fishes of Virginia. The only help from the impoverished Museum was for bottles and alcohol, but using Reid’s old automobile and paying all other expenses themselves, they made many collec- tions from the Dismal Swamp to the mountains of western Virginia. Myers says that in Washington he had the best and most cooperative superior administrators that a curator could have. His immediate superior was the late Leonhard Stejneger, Head Curator of Zoology and Curator of the Division of Reptiles and Amphibians, whose kindliness and enormous memory he has always remembered with pleasure. The other was Alexander Wetmore, then Assistant 2 Myers always expressed the greatest admiration for Reid, who was invalided out of the U.S. Marine Corps after being wounded in the eyes while in Nicaragua. Reid became a doorman in the Museum and worked his way to the Civil Service subprofessional grade of Aid by taking night courses in zoology at George Wash- ington University. ~I Vor. XXXVIII] WALFORD: HISTORY OF GEORGE S. MYERS Secretary of the Smithsonian and Director of the National Museum, whose administrative ability and thoughtfulness for his staff were boundless. In 1936, Myers was invited back to Stanford, and after considerable thought, accepted a position as Associate Professor of Biology and Head Curator of Zoological Collections, with the provisions that he be advanced to Professor by 1938, that half his time be spent on curatorial duties and half on teaching, and that Dr. Herre’s employment as Curator of Zoology not be terminated as the department head had planned. With Herre’s retention assured, Myers assumed his new position in September, 1936. George Myers has told me many times that his most important contributions to ichthyology and herpetology have been the help and guidance he has been able to give to the long line of graduate students who worked with him at Stanford. When he began teaching in 1936, no formal course dealing with more than the barest rudiments of taxonomic ichthyology appears to have been given anywhere. In those days, prospective taxonomists were supposed to pick up knowledge of their field without any formal guidance. The Stanford fish course initiated by Charles H. Gilbert and continued by John O. Snyder had consisted solely of identifying specimens with the aid of Jordan and Evermann’s “Fishes of North and Middle America.” Myers has told me that his own background in ichthy- ology and vertebrate evolution was very defective as a consequence, so that be- tween 1936 and 1938 he found it necessary to prepare himself by doing a great deal of reading and studying. The books of Goodrich and Romer and the papers of C. Tate Regan proved to be of the greatest help. In 1938, for the benefit of a small group of students, including W. A. Gosline and E. S. Herald, he attempted a general summary of fish classification and evolution, with emphasis on the literature and history and on major groups down to the family level. This first attempt developed into a more formal course, called ‘“‘Advanced Systematic Ichthyology,” which was usually given every other year, alternating with a shorter, somewhat less advanced course in herpetology. This course in ichthy- ology formed the genesis of other more or less similar courses, such as that given by W. A. Gosline first at the University of Michigan and later at the University of Hawaii. Myers also gave annually a course at first called ‘Vertebrate Paleontology” and, later, “Evolution of the Vertebrates,” and a short course on zoogeography. Myers’ most popular course, planned and given by himself and his botanical colleague, Professor Ira L. Wiggins, was a general survey of plant and animal ecology, including ecology of man. It was given for two or three years in the late 1940’s primarily for non-biologists. In this course, Myers was one of the first to emphasize the rapidly increasing danger to the human race caused by the unrestricted growth of human population. Unfortunately, other work forced Wiggins to withdraw from the course, and, as Myers felt himself incompetent to handle the botanical side, the course was regretfully dropped. 8 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Myers has maintained that graduate teaching is greatly helped by the presence of a “‘critical mass”’ of at least four enthusiastic students working under one professor on different dissertations in related fields. The presence of such a group was barely attained in his laboratory when World War II intervened. Students vanished, and he was sent to Brazil for two and one-half years. It was not reattained until the late 1940’s; but from that time until well into the 1960’s a “critical mass” was continuously present, rising at one point to as many as 12 or 13, usually two or three of them herpetologists and the rest ichthyologists. As he says: “Those were the most exciting and rewarding years that I have experienced. As a matter of principle, no graduate student was assigned a doctor- ate problem, or encouraged to choose one closely related to any of my own re- search. They were forced to select their own, my only requirements being that the problem be reasonably interesting and difficult but not impossibly time- consuming, and that it be concerned with areas within which I felt myself fairly competent to judge quality of performance. The atmosphere was never dull. Everybody helped and taught everybody else, the professor learning as much from the students as they did from him. Chores in the old Natural History Museum, such as registering and care of material, helping with the editing of the Stanford Ichthyological Bulletin, and the like, were often done almost as much by those who were not paid to do the work as those who were. My own contribu- tion was largely that of arbiter, critic, walking bibliographer, ruthless editor of often poorly expressed, first attempts to write up scientific results, father-con- fessor, cheerleader, and especially as the provider and keeper of a laboratory atmosphere conducive to hard work, cooperation, enthusiasm, and high attain- ment. There was little of the formality that often separates professor from stu- dent. Evening seminars or meetings often ended in a nearby Bierstube, and I was usually invited to student parties. Former students have often remarked on the uniqueness of human relations in the Museum and recall them with nostalgia —as do I. Yet, I must have commanded a modicum of respect for I have noted with some amusement that none of my former graduate students ever tried to address me by my first name (a common enough thing in the U.S.A.) until 10 or 15 years after obtaining their doctorates; and several have never been able to bring themselves to do so. But when the number of graduate students rose to eight, ten, or a higher number, I got comparatively little research of my own done, for I was available to all of them almost every day—an arrangement at which some other groups of graduate students, both at Stanford and elsewhere, marvelled. Yet near the height of my graduate student load, economic necessity forced me to write extensively for —and manage—a popular aquarium magazine (The Aquarium Journal). This very difficult regime went on for two years (1952-54) until my Stanford salary rose enough to make it possible to give up most such writing.” Vor. XXXVIII] WALFORD: HISTORY OF GEORGE S. MYERS 9 The Stanford fish collection was originally small, consisting largely of dupli- cates from the field work of Jordan and his pupils. Through the years it grew slowly through an unexpressed policy of growth in diversity without amassing long series of individual species, a policy which Myers enforced in more recent times. It gradually attained a diversity among American collections second only to the National Museum, although in numbers of specimens (between 750,000 and 1,000,000) smaller than several other large research collections. It has been especially useful for morphological work in systematics such as has been em- phasized in recent years. The herpetological collections were small when Myers took charge in 1936, consisting of fewer than 2,000 amphibians and 10,000 reptiles, mostly collected during early work in the days when John Van Denburgh was a student. Myers built these collections up judiciously until now they total about 60,000 speci- mens, half amphibians and half reptiles—numerous enough and diverse enough for many systematic purposes. For curatorial work and the management of the Zoological Collections pri- marily as a laboratory for graduate students, Myers was greatly aided by Margaret H. Storey. She had obtained her A.M. degree with Willis Rich while Myers was at the National Museum, had stayed on as a volunteer assistant later to be appointed Assistant Curator of Zoological Collections. She supervised the paper-work and curating, helped edit Stanford Ichthyological Bulletin and was a tower of strength and help to all those who worked in the Zoological Museum until her untimely death in 1960. Myers and Storey together worked out systematic methods, some of them new, for sorting, registering, bottling, labelling, arranging, installing, and finding bottled museum specimens. These methods, described chiefly in three of the Museum’s mimeographed circulars, made it possible for much of the work to be done by untrained student helpers, and to handle a large research-collection operation (up to a million specimens) with less than half the staff and funds usually available for such purposes. For all of Myers’ years on the faculty, the Zoological Collections had no more than four employees besides himself—A. W. Herre, until World War IJ, Margaret Storey and later Warren Freihofer, as aid or associate curator, one half-time student assistant, and, after World War II, one typist-secretary who also served entomology. This staff handled large and growing research collec- tions of fishes, amphibians, and reptiles as well as sizable collections of mammals, birds, and aquatic invertebrates. They were also responsible for the time-consuming processing of extensive loans to researchers elsewhere, running from about 500 to as many as 5,000 specimens annually. In one thing, Myers was adamant. Collections of animal groups in which he had no direct interest were also kept in good condition and order. Such curatorial conscience 10 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. is rare. Moreover, collections which had strayed away from the Museum to other parts of the University and elsewhere were retrieved and set in order. Types were rigidly labelled and segregated with blue (holotypes) or red labels. Species presumed to be extinct received green labels. The library on Systematic Ichthyology at the Museum was rich in the older literature and in reprints, all from Jordan’s personal library. It was kept up by exchanges for Stanford Ichthyological Bulletin and by judicious buying with the small funds available. Concurrently, Myers personally purchased many ichthy- ological and herpetological books that were not present in the Museum, and his library books and reprints admirably supplemented the Museum’s holdings. To- day, since Professor Carl L. Hubbs’ library has gone to Scripps Institution, Myers probably has the most extensive private fish library in the world. It is especially rich in reprints but lacks such expensive items as the great works of Bloch, Bleeker, and Agassiz. Myers started Stanford Ichthyological Bulletin in 1938. It was printed cheaply by offset (the text being typed by the Museum staff), since funds for this journal were always miniscule. They began at $133.00 annually and never rose over $750.00. All sorts of schemes to get outside funds were tried, usually with only moderate success. Of the eight volumes that eventually appeared, less than half were or could be paid for from regular funds. Myers and Miss Storey were the principal movers in two local groups. The old “Stanford Zoology Club,” which originated in the 1890’s and was supported by generations of Stanford students, was revived as the “Natural History Club” and survived until the 1950’s. A new, informal group, the “Fishverein,” com- posed of those at Stanford interested in fishes and the many local fishery biolo- gists working for the Federal Fish and Wildlife Service and the California Division of Fish and Game, was formed by Myers and met fairly regularly for many years. During his early preuniversity years (1920-24), Myers’ papers reflect the growing interests and ability of an untrained young man deeply interested in the habits and taxonomy of the lower vertebrates. He published his first articles on aquarium fishes at the age of 15, in 1920. These early attempts give an inkling of the extensive boyhood observations representing dozens of families of live fishes, and also amphibians and reptiles, either in captivity or in the field in New Jersey and North Carolina. As Myers says: “By the time I was 19, I knew in a general and sometimes specific way a great deal about fish behavior that has of late been ‘discovered’ and formally categorized by the fish behaviorists, in the same way that the field ornithologist becomes familiar with bird behavior.” By the end of 1923, Myers had published his first really scientific papers, one on a new poeciliid from Hispaniola with J. T. Nichols and others on the nomenclature of anabantids. By the end of 1924, he had published nine tax- Vout. XXXVIII] WALFORD: HISTORY OF GEORGE S. MYERS 11 onomic papers on fishes, and one herpetological paper. It was in 1924 that he made his first longer field trip, to Wilmington, North Carolina, where he made many observations and discovered what is now known as the common dusky shiner (Notropis cummingsi Myers) of the southeastern coastal plain, the de- scription of which he published in 1925. Although the beginning of university work in 1924 curtailed his output of papers, he continued publishing on a variety of ichthyological and herpetological subjects up to the time he finished his schooling at Stanford and went to the Smithsonian in 1933. To refer to only a few of the papers which he published during his student years at Indiana and Stanford (1924-33), there is a synopsis of Indiana amphibians and reptiles (1926), four papers on amphibians (1930— 31), descriptions of many South American fishes collected by Ternetz (1927), a revision of the genera of neotropical cyprinodontids (1927), three or four im- portant papers on Chinese fishes, and a prophetic paper on the phallostethids which foreshadowed some of the important features of Rosen’s radical reclassifi- cation of the atheriniform fishes in 1964. In addition, Myers found time in 1929 to write a sizable addendum to the final volume of Eigenmann’s ““The American Characidae.” Myers was faced with such exceptionally time-consuming curatorial duties at the Smithsonian that his research during those years (1933-36) suffered. However, he reviewed the genera of triacanthids in 1934, published on the cy- prinodonts of Hispaniola as well as the opistognathids (and owstoniids) in 1935, and revised the genera of Polynemidae in 1936. In that same year, in a report on fishes from Lake Tanganyika, he briefly pointed out for the first time some of the strange features of lake-fish evolution. After beginning his teaching and curatorial work at Stanford in 1936, Myers’ first paper was one that he had read before a meeting in New York in 1934 and which he based on observations made in the 1920’s. In this short paper, he arrived independently at the same conclusions as had C. M. Breder, Jr. in regard to the evolution of oral brooding in cichlid fishes. The most widely known and influential of Myers’ papers, prepared for the 1937 Smithsonian Report (1938) was his “Fresh-water Fishes and West Indian Zoogeography.” He had been highly dissatisfied with most writings on historical zoogeography, particularly the prevalence of the ideas of Matthew and others based largely on the tetrapod evidence, and especially with the use made of the evidence of fresh-water fishes. In this paper, dealing specifically with the West Indies but ranging over the fishes of all continents, Myers pointed out that what had previously been taken for true fresh-water fishes are really divisible into two physiologically different groups, one with considerable salt tolerance and the other (“primary fresh-water fishes”) much more strictly confined to fresh water. The primary fresh-water fishes are much less able to spread across continents 12 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. and sea gaps than are mammals and even amphibians, and thus their dispersal patterns provide a much more conservative and dependable guide to the past existence of these gaps than do those of tetrapods. Myers’ zoogeographical con- clusions, although stated only cautiously and tentatively, agreed with those of Matthew in regard to the absence of past continental connections of the West Indies, but disagreed with Matthew in the strong evidence provided by the primary fresh-water fishes for a past southern trans-Atlantic connection. It is notable that 20 years after publication this paper was acknowledged by P. J. Darlington in his great book “Zoogeography,” as the prime reference on which he built that part of his book dealing with fishes. Myers’ 1938 work combined with his later papers on salt tolerance of fresh-water fishes (1949) and East Indian zoogeography (1951), gave new direction to later studies on the his- torical zoogeography of continental vertebrates. Myers seemed more than half convinced of the truth of continental drift in 1938, and although he faltered in that conviction in his 1951 paper, he later reaffirmed it in 1966 and 1967, be- cause by then the weight of his evidence favored the primary fresh-water fishes as the most significant vertebrate indicators for establishing past continental connections. Early in 1938, Myers was able to accompany that year’s expedition of the Allan Hancock Foundation’s ship Velero I/I to the coasts of Mexico, Cocos Is- land, the Galapagos, Peru, Ecuador, and Panama. This resulted in collaborative papers with C. B. Wade on eels (1941), atherinids (1942), and other fishes (1946). In addition, a study on the zoogeography of the fishes of the Pacific Ocean appeared in 1941. Herpetological work had been impossible in the Division of Fishes in Wash- ington; but on Myers’ return to Stanford he began a number of smaller studies on amphibians and lizards which culminated in six herpetological papers in 1942. One of these described the now well-known black toad of Deep Springs Valley (Bufo exsul Myers), which has perhaps the smallest range of any living am- phibian. Following the entrance of the United States into World War IT, Myers was posted to the Museu Nacional in Rio de Janeiro, as part of a governmental plan to maintain good relations with Latin America in troubled times. He arrived in July, 1942, for a one-year period, which eventually lengthened to nearly 2% years. In Rio he helped with curatorial and library methods, with setting up civil service categories for the museum staff, with exhibits and with museum admin- istration. For the federal fish and game division and the Sao Paulo fish and game department, he helped by devising better methods of gathering fish-catch statistics. In addition, for a period of over a year, the Museum lent his services to the Rio office of the U.S. Coordinator of Inter-American Affairs. There was little time for research, and the wartime shortage of gasoline made travel by Vor. XXXVIII] WALFORD: HISTORY OF GEORGE S. MYERS 13 automobile next to impossible. Nevertheless, he managed to take many local trips, principally by tramway on weekends, to the wilder areas in the metro- politan region. These trips were mostly for frogs, in the company of Dr. Bertha Lutz and Joaquim Venancio or Antenor Carvalho. Eventually, there were longer trips with Carvalho or others by train and other conveyance, to the Rio Sao Francisco at Pirapora, to Santa Teresa in Espirito Santo, and southward along the coast to Rio Grande do Sul. Papers resulting from the Brazilian years were few, most of them appearing in 1944 and 1945. On Myers’ return in 1944, he hoped that the survey of Brazilian marine market fishes that he had helped to originate would result in taxonomic studies of these fishes at Stanford by Brazilian students; but the students did not ap- pear and the project languished after 1950. The sole results have been the amassing of an excellent representation of Brazilian shore fishes in the Museu Nacional, and a smaller duplicate set at Stanford. A trip to attend the Pacific Science Congress in New Zealand in 1949 re- sulted in two zoogeographical papers, one on East Indian fishes (1951 and 1954, published twice) and the other on East Indian amphibians (1954) both of which tended to firm up the concept of Wallace’s Line. Myers had become editor of an aquarium magazine for two years in the early 1950’s. Several of the articles published then have ichthyological interest, chief of them being “Annual Fishes” (1952), which brought together and greatly strengthened by original observa- tions what had consisted of scattered and mostly nonscientific reports of tropical cyprinodontid fishes which exhibit a diapause when no individuals are alive ex- cept as zygotes. At the 1958 International Congress of Zoology in London, Myers presented a paper on the endemic fishes of Lake Lanao having an important bearing on evolution. In this paper, published in 1960, he was able to show that this_ cyprinid fauna, now diverse enough to be alloted to several genera, almost cer- tainly evolved very rapidly from a single ancestral species, perhaps within 10,000 years. He also pointed out similarities in the evolution of other lake faunas, and was able to establish an evolutionary sequence: 1) an increasing number of very similar species belonging to a single genus, culminating 2) in a “species swarm;”’ then 3) the differentiation of a few species into new endemic genera, and finally 4) considerable reduction in the total number of species. Thus the number of species of the large genus gradually diminishes while the number of distinctive endemic genera increases. Myers also pointed out the strong possibility that on a grand scale the evolution of Amazonian fishes and of deep-sea fishes might parallel that of lake fish faunas and indeed, the original evolution of the animal phyla. In the 1960’s, Myers returned to zoogeographical studies of fresh-water fishes. His paper on the North American fauna (1963) was published only in an ab- 14 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER. stract which lacked the section on continental drift; but his 1966 paper on the derivation of the fresh-water fishes of Central America directly opposed Darling- ton’s idea that the ancestors of the South American fish groups originated in Holarctica, and suggested continental drift as an answer. In 1967 appeared his “Zoogeographical Evidence of the Age of the South Atlantic Ocean,” a brief exposition of his belief that the cypriniform fishes had originated in a South Atlantic continent which split in the Triassic or Jurassic to form the South Atlantic Ocean. In 1966 was published a collaborative work by Greenwood, Rosen, Weitzman, and Myers, ‘“‘Phyletic Studies of Teleostean Fishes, with a Pro- visional Classification of Living Forms,” which broke strongly with traditional classifications of the teleosts. Myers intended his series of apparently not directly related studies on fish zoogeography (1938, 1949, 1951, 1963, 1966, 1967), together with his two 1960 papers on lake fish evolution and the 1966 collaborative teleost study, to form an integrated whole indicating as nearly as can be done at present how and when the ostariophysan (and particularly the cypriniform) fishes evolved and dis- persed. In these papers the problem is attacked from several directions on the basis of the living world fauna and the few known fossils, ecological constitution of the fishes, their probable place and time of origin from the salmoniform fishes, their dispersal and evolutionary patterns as seen against the background of paleogeography, all within the strictures imposed by the greater known fossil evidence derived from tetrapods. Considered in this way, the nine papers con- cerned form an impressive contribution to knowledge of the fresh-water fishes of the world. One thing that Myers has said of his papers is that not many of them are as important, or represent as much thinking, as do a number of his reviews, mostly published since 1930 in Copeia. Many taxonomic and other conclusions first appeared in these reviews. Moreover, the column called by Myers “Phylax Telescopus,” which he maintained for a couple of years in Copeza during the 1960’s, contains some of the best biological criticism that has appeared any- where. Myers has said to me that if he is remembered for anything, he would like it to be for just a few things—his graduate pupils, his critical comments and reviews, his early espousal of the need for curtailment of human population growth, his pioneer urging of the conservation of non-food and game fishes, and his integrated series of papers on the evolution and dispersal of fresh-water fishes. Despite the number of publications listed in his bibliography (nearly 600), I doubt that he ever engaged in any research simply to increase the quantity of his publications. He has always avoided humdrum taxonomic questions unless they were of some special significance, for he is completely devoted to seeking and elucidating principles. Thanks in large measure to his scholarly creativeness, as well as to his subtle and boundless patient teaching, systematic ichthyology is Vot. XXXVIII] WALFORD: HISTORY OF GEORGE S. MYERS 15 alive and well today and the subject of vigorous teaching in many centers of learning where it is appreciated. It is a pity that Stanford has not appreciated the tradition it had inherited through Jordan or the treasure which he started in the Museum collection and libraries, and which Myers built up and organized. Instead, the university authorities have callously determined to give this treasure away and discontinue—discontinue—further teaching in this field! This is par- ticularly tragic at a time when the natural history of the earth and its resources is the most important thing we can know. CHRONOLOGY 1905 Born February 2, Jersey City, New Jersey, son of Harvey Derwood Myers and Lily Vale (Sprague) Myers. 1911-18 Public elementary schools, Jersey City. 1918-24 Public high schools, Jersey City. 1919-20 St. John’s Military School, Ossining, New York. 1922-24 Association with American Museum of Natural History, especially G. K. Noble and J. T. Nichols. 1924 Field work during May in vicinity of Wilmington, North Carolina. 1924-26 Indiana University, with Carl H. Eigenmann. Curatorial assistant, fish collection. 1926 Married Martha Ruth Frisinger, Decatur, Indiana, September 25. 1926 Entered Stanford University, October. Beginning of association with D. S. Jordan, C) He Gilbert, J; ©: Snyder, E: ©: Starks: 1926-31 Museum assistant, Stanford. 1929 Field work during April-June in western Texas and Arizona with Gregory M. Kranzthor. Rediscovery of Elaphe bairdii. 1930 Field work in Death Valley—Amargosa region—with Joseph H. Wales. Discovery of Cyprinodon diabolis. 1930 Bachelor of Arts, Stanford, June. 1931 Master of Arts, Stanford, June. 1931-32 Teaching assistant in comparative anatomy, Stanford. 1932-60 Associate editor, The Aquarium, Philadelphia, edited and published by William~ Thornton Innes, also scientific editor, 19 successive editions of Innes’ “Exotic Aquarium Fishes.” 1933 Appointed Assistant Curator, in charge, Division of Fishes, U.S. National Museum, Smithsonian Institution, Washington, D.C., January 1. 1933 Doctor of Philosophy, Stanford, June. 1934-36 Field work, freshwater fishes of Virginia, with E. D. Reid. 1935 Birth of first child, Thomas Sprague Myers, Washington, D.C., August 28. 1936 Awarded Silver Medal of the “Société National d’Acclimatation,”’ Paris, for work on acclimatization, habits, and taxonomy of exotic aquarium fishes. 1936 Resignation from Smithsonian. Appointed to faculty, Department of Biological Sciences, Stanford University, as Associate Professor and Curator of Zoological Collections, September. 1937 Birth of second child, John William Myers, Palo Alto, California, December 15. 1938 Member, Hancock Pacific Expedition, aboard M. V. Velero III, from January— March, visiting coasts of Mexico, Guatemala, Cocos Island, Galapagos Islands, Ecuador, Peru, Chinchas Island, Gorgona Island, Colombia, Panama. 1940-41 1942 1942-44 1944 1945-51 1946 1947 1949 1949-51 1950 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Co-leader, with Rolf L. Bolin, of Crocker-Stanford Deep-sea Expedition, aboard yacht Zaca, off California coast in September. Initiated Stanford Ichthyological Bulletin. Editor to end of volume 8 in 1967. Advanced to full Professor, Stanford, September. Member, Fishery Organizing Committee, 6th Pacific Science Congress, Berkeley, California. Intensive extracurricular work with William Allen White’s “Committee to Defend America by Aiding the Allies.” Elected Corresponding Member, Zoological Society of London. Posted to Rio de Janeiro (State Department funds) to aid Museu Nacional and Divisao de Caca e Pesca. Lecture course on ichthyology and fishery biology in Rio. Brief visits en route to Mexico City, Guatemala, Panama, Cali, Bogota, Mariquita, Lima, Arequipa, Santa Cruz (Bolivia), Corumba. Intermittent field work near Rio, and (with Antenor Carvalho and others) to Minas Gerais, Espirito Santo, Sao Paulo, Parana, Santa Catarina, Rio Grande do Sul, and Belém do Para. Return to Stanford, October. Vice-President and Council Member, California Academy of Sciences, San Francisco. Beginning of post-war upswing in graduate-student enrollment at Stanford. Bikini Scientific Resurvey, U.S. Navy, aboard U.S.S. Chilton. Field work on Bikini and Rongerik atolls. Plankton Survey, Bikini lagoon. Visits to Kwajalein and Honolulu, summer. Pacific Science Congress, Aukland and Christchurch, New Zealand. Some fish and reptile collecting on South Island and Aukland Harbor. Visits to Hawaii, Samoa, Noumea, Canton Island and Johnston Island en route. President, American Society of Ichthyologists and Herpetologists. Brief trip to Brazil during August and September, visiting Recife, Salvador (Bahia), Rio, Belém do Para, Manaus, and Puerto Rico. Special taxonomic work, U.S. Fish and Wildlife Service, Washington, D.C., summers. Managing editor, Aquartum Journal, San Francisco. European trip for Fish and Wildlife Service and FAO. Paris, with stop in London, December. Field work and fish collecting during February, upper Rio Caqueta basin, vicinity of Tres Esquinas, Colombia, with General Thomas D. White. Visit to Bogota. International Zoological Congress, London. Visits to Copenhagen and Hamburg, summer. Organizing Committee for First International Congress of Oceanography, held in United Nations headquarters, New York, summer of 1959. Elected honorary fellow, Zoological Society of India. Field work and fish collecting during February, upper Rio Guaviare basin, near Sierra Macarena, Colombia, with General T. D. White. Six-month study trip to Europe, visiting Hamburg, Copenhagen, Lund, Goteborg, Amsterdam, Leiden, Brussels, Frankfurt, Vienna, Lucerne, Paris, London. Field work and fish collecting during February in Nicaragua; Managua area, Lake Nicaragua, Rio San Juan, with General T. D. White. International Zoological Congress, Washington, D.C., August. International Conference on Tropical Oceanography, Miami. Arranger and con- vener, section of zoogeography, November. Marriage to Frances Edna Felin, Palo Alto, California. Vor. XXXVIII] WALFORD: HISTORY OF GEORGE S. MYERS 17 1967 Primer Foro Internacional sobre Planificacion y Desarrollo Pesquero, Caracas, Venezuela, August. Followed by brief travel in eastern Venezuela and lower Rio Orinoco with Agustin Fernandez-Yepez. Visits to Trinidad, Panama and Puerto Rico en route. 1969-70 Vice-President, Cactus and Succulent Society of California. 1970 Statutory retirement on August 31 from faculty, Stanford, August. 1970 Appointed Henry Bryant Bigelow Visiting Professor of Ichthyology, Harvard University. =e 1 a= - , - t ) pai 7 ws i he eweil - ; a ~ - : ee hs =e ee | ee ~wite @ . tram ee ° , -__=—e i ~, “Gren Re RS Peel as Hs ao f as ee . ae t e = '% Z - i ey - oe ol. ° % f vi wu. ; ay ; jee - _ rh RAlwh=-- © rm Gund Geax hare ‘ i ite ge a ay ie) oS = i 9 — - , is i a peaea) A 3 = 7 te 7 a) ie Si io 4 et (neal ae, oe |. 2 Pie pe alee = DY Fra ae 0 eee i err” i erin OUT er are LRaeite @ ier. Jara a = is sali reth al a hey dimen): | one : a) PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Festschrift for George Sprague Myers Vol. XXXVIITI, No. 2, pp. 19-52. December 31, 1970 ANNOTATED CHRONOLOGICAL BIBLIOGRAPHY OF THE PUBLICATIONS OF GEORGE SPRAGUE MYERS (to the end of 1969) This bibliography lists all known publications by George S. Myers, scientific and other, omitting only about two dozen ephemeral items such as newspaper articles. It has been compiled almost entirely from the mimeographed bibliogra- phies issued by Myers in 1950 and 1952 and his own card index of titles. Nearly all the bibliography and all the annotations are thus by Myers himself. The serial numbers of the papers have been used as annotations of mailed-out separata on Myers’ address cards of colleagues, and have not been changed even though a few previously missing entries have now been inserted in their chronological places. To avoid confusion, Myers almost always gave separate numbers even to subsequent reprintings or translations of original contributions and to successive installments of serially published papers. In all such cases cross references are given. The total of numbered entries is thus greater than the number of original papers, but the usefulness of this system is obvious in the annotations. The last numbered entry is 593, but with ten interpolations added, the total is 603. Aside from formal taxonomic papers, the largest classes of publications are articles on aquarium fishes and book reviews. Myers became interested in ichthyology through aquarium fishes and retained his interest in them perma- nently. By 1930 he was already becoming the recognized authority on the identi- [19] 20 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. fication and aquarium behavior of the smaller freshwater species from tropical America, Asia, and Africa. The number of Myers’ publications in this field be- came especially large during the 26 months (June 1952—August 1954) when he acted as editor of the Aquarium Journal. Although many of these papers were of a popular nature, many of them contain original observations on the taxonomy or behavior of the species concerned and could not be excluded even from a bibliography of scientific publications. In writing book reviews, Myers acted on a theory (reinforced, he says, by advice of the late Dr. Joseph Grinnell of the University of California) that care- fully done, critical reviews form a powerful instrument to weed out incompetence and raise the level of any scientific discipline. Myers obviously adhered to that theory, for his reviews are usually sharply critical and often embody original ideas or taxonomic views not expressed elsewhere. He has been known to say that he puts as much thought into reviews as into more formal contributions, and that some of his best writing has gone into them. A sampling shows this to be true; his reviews, editorials, and columns-of-comment give a broader view of his thought and critical abilities than do his formal papers. Many are worth reading at any time, especially by younger workers. There are also 49 papers dealing largely or wholly with zoological nomen- clature, ranging from his early attempts to modernize the names of aquarium fishes to such nomenclatural problems as the name of Culter (243), the family name of the characids (335), and some Neotropical frog names (560). Myers was asked by a former pupil if he could make a rapid analysis of his papers and came up with the following rough totals: (A) Papers of scientific interest on taxonomy, evolution, ecology, behavior, distribution, and nomencla- ture of fishes: 252. (B) Ditto of amphibians and reptiles: 51. (C) Historical zoogeography, chiefly of lower vertebrates: 8. (D) Formal book reviews: 104. (E) Editorials, comment, criticism: 11. (F) Taxonomic theory: 2. (G) Cura- torial, collecting, preserving: 5. (H) Popular articles on fishes, of little or no scientific interest: 95. (I) Botany: 2. (J) Station records: 1. (K) History of ichthyology: 2. (L) Biographies and obituaries: 14. (M) Translations and reprints of original Myers’ papers: 32. (N) Continuations of serial articles with- out any change of title: 6. (O) Nonfish aquarium articles: 6. (P) Verse, allegory, etc.: 3. (Q) Unimportant notes and corrections: 9. Myers says that several papers published in the aquarium literature are included under category A, but only when they contributed significantly to knowledge of the fishes con- cerned. For the benefit of herpetologists, all papers of all categories that are im- mediately concerned with amphibians and reptiles are preceded by an asterisk (*) in the bibliography. The form of the entries is as follows: first the serial number, followed by VoL. XX XVIII] MYERS: BIBLIOGRAPHY 21 the full title of the paper [untitled contributions have been given a title in brackets], the journal or other vehicle of publication, the volume number and issue number, and the pagination, followed by the precise date of publication so far as Myers knew it. Annotations by Myers are in brackets, usually at the end of the entry. The dates of papers, in cases where there is not even a month given, were known only as to year. Dates of publication of Copeza articles are the imprinted ones of the issues concerned. All dates given for the Stanford Ichthyological Bulletin papers, and for articles in the Aquarium Journal while Myers was editor (June 1952—August 1954) are the exact dates of mailing of the issues concerned. Where the imprinted year or month of a volume or article differs from the year placement or month placement in this bibliography (as with the proceedings of different Pacific Science Congresses) the date here given is the correct one for mailing of the volumes. Joint-authorship papers have the names of the authors given, in the original order, in parentheses immediately following the title. The only year in which no paper was published was 1968. Abbreviations of the names of journals frequently cited are as follows: AJ =The Aquarium Journal. Published monthly by the San Francisco Aquarium Society, 1928-1965. [Continued 1966-1967 as Ichthyologica, The Aquarium Journal, published by TFH Publications, Inc., Jersey City, N. J.] Edited by GS.M., 1952-1954. AL = Aquatic Life. Published by Joseph E. Bausman and edited by W. A. Poyser in Phila- delphia, 1915-1922; subsequently published and edited for many years by August M. Roth in Baltimore. Philadelphia issues usually published promptly; Baltimore issues often lagged badly. AMN = American Museum Novitates. Published by The American Museum of Natural History, New York City. Separate papers numbered consecutively; numbers separately paged. ANMG=The Annals and Magazine of Natural History. Published in London by Taylor and Francis, 1838-1966. Continued under the name Journal of Natural History. AP = Aquarium (Paris). Published in Paris, 1934—-at least to 1936. Many articles from TA translated into French with the original TA colored plates and other figures. BATK = Blatter fiir Aquarien- und Terrarienkunde. Published from 1890 until about 1937, first in Magdeburg and later in Stuttgart. Many important behavioral and taxonomic papers included. CO = Copeia. Journal of the American Society of Ichthyologists and Herpetologists. Pub- lished at various places in the U.S.A., 1913—present. To 1930, all issues numbered con- secutively, with no volume number. Later, the year is the volume number (4 issues per year). FC =The Fish Culturist. Published in Philadelphia by the Pennsylvania Fish Culturists’ Association, 1921—present. HAB = The Home Aquarium Bulletin. Published at first in Newark, N. J., by the Newark Aquarium Society, later by a group in East Orange, N. J., 1931-to at least March, 1936. Carl L. Hubbs and Myron Gordon were among the associated editors. LSJ = Lingnan Science Journal. (Continuation of Lingnaam Agricultural Review.) Pub. lished by Lingnan University, Canton, China. Edited by Wm. E. Hoffman and Robert Cunningham Miller. CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. bo bo PBSW = Proceedings of the Biological Society of Washington. Washington, D. C., 1880- present. SIB = Stanford Ichthyological Bulletin. Edited by G. S. M. and M. H. Storey. Published by Natural History Museum of Stanford University (later Division of Systematic Biology), Stanford, Calif., 1938-1967. TA=The Aquarium. Published by Innes Publishing Co., Philadelphia, beginning in 1932. Edited by Wm. Thornton Innes. G. S. M. was an associate editor for many years. TFH = The Tropical Fish Hobbyist. Published by TFH Publishing Co., Jersey City, N. J., beginning in 1952. WATK = Wochenschrift fiir Aquarien- und Terrarienkunde. Published weekly, 1904-1950, in Braunschweig. Much important behavioral and taxonomic information included. 1920 1. Phalloptychus januarius. AL, vol. 5, no. 7, p. 74. July. [Misidentification; species mentioned is spotted form of Phalloceros caudimaculatus.| 2. The red rivulus. AL, vol. 5, no. 7, pp. 79-80. July. [Xanthic form of Rivulus urophthalmus. | 3. Fundulus diaphanus. AL, vol. 5, no. 8, p. 91. August. [The figure, supplied by the editor, is of F. heteroclitus.| 4. Some fish suitable for home aquaria with suggestions concerning starting and main- taining an aquarium. 8vo, 16 pp., Hudson County Aquarium Society, Jersey City, N. J. September 10. [Circular distributed at 3rd annual exhibition of H.C.A\S.; pp. 2, 4, 6, and 8 are advertisements; author’s name omitted by printer. Copies in the libraries of Stanford University and the U. S. National Museum. ] 5. The Mexican swordtail. AL, vol. 5, no. 11, pp. 122-123. November. 1921 6. The labyrinth fishes. I. AL, vol. 6, no. 1, pp. 1-2. January. [Continued in nos. 13 and 19.] 7. Fundulus chrysotus. AL, vol. 6, no. 3, p. 18, September. 8. The common sunfish. AL, vol. 6, no. 4, pp. 19-20. October-December. 1922 9. The black banded sunfish. Aquarium News, Ridgewood, N. Y., vol. 1, no. 5, p. 2. January 15. 10. Chirodon arnoldi. AL, vol. 6, no. 5, p. 28. January—June. [The species mentioned is actually Astyanax mexicanus.| 11. Planting aquaria. FC, vol. 2, no. 4, pp. 157-158. September. 12. The aquarium and its denizens, being a brief exposition of the proper arrangement and maintenance of home aquaria, with a catalog of some of the fishes suitable for aquarium culture. 8vo, 22 pp. (cover is title-page); Hudson County Aquarium Society, Jersey City, N. J. September 8. [Copies in the libraries of Stanford Uni- versity, the British Museum, and the U. S. National Museum. See also no. 56.] 13. The labyrinth fishes. II. AL, vol. 6, no. 6, pp. 33-34. July—October. [Continuation of 6. See also 19.] 14. Interesting notes. FC, vol. 2, no. 6, pp. 172-173. November. [Notes on Fundulus gularis, Mesogonistius chaetodon, and Pterophyllum scalare; article includes six lines on p. 173, not well differentiated from other notes below. | 19. Ae 30. . XXXVIIT) MYERS: BIBLIOGRAPHY 23 Hudson County exhibition. AL, vol. 6, no. 7, p. 46. November. . A true fish story. Swastika, Jersey City, vol. 1, no. 7, pp. 4-5. December 10. [Con- cerns the climbing perch. See 18 for note.] . A recently described aquarium fish. CO, no. 113, p. 89. December 20. . The largest frog—the smallest frog. Swastika, Jersey City, vol. 1, no. 8, p. 5. December 25. [A brief note on the size of Rana goliath and newly hatched Eleutherodactylus. The Swastika was a student publication of the city high schools in Jersey City. Edited by Meyer Levin. ] 1923 The labyrinth fishes. III. AL, vol. 6, no. 9, pp. 63-64. January. [Continuation of 6 and 13. Never concluded. ] The characins. FC, vol. 2, no. 8, pp. 186-187. January. A note on the fighting fish. FC, vol. 2, no. 10, pp. 202-203. March. [Note on the identification of aquarium examples of Betta; in the third paragraph, line four, the word “small” was transferred by printer’s error from before “lake” in preceding line. | . Correct names. FC, vol. 2, no. 11, pp. 210-212. April. [An attempt to modernize the scientific names of aquarium fishes. See also 24.] . Aplocheilus chaperi. AL, vol. 7, no. 1, pp. 1-2, 12. May. Correct names. AL, vol. 7, no. 1, pp. 3-6. May. [A reprint of 22, with the addition of some name meanings. | . Notes on the nomenclature of certain anabantid fishes and a new generic name pro- posed. CO, no. 118, pp. 62-63. May 20. Ctenobrycon spilurus. FC, vol. 2, no. 13, p. 226. June. . On the subject of scavengers in the aquarium. FC, vol. 2, no. 12, pp. 228-229. June. A new limia from San Domingo. (John Treadwell Nichols and G. S. Myers.) AMN, no. 79, 2 pp. June 12. . Hyphessobrycon anisitsi, a new fish for the aquarist. FC, vol. 3, no. 3, pp. 250-251. November. [Identification erroneous; species mentioned is Hemigrammus caudovit- tatus.] Further notes on anabantids. CO, no. 124, pp. 111-113. November 20. 1924 . The labyrinth fishes. FC, vol. 3, no. 7, pp. 282-284. March. [Not part of the series: 6, 13, and 19.] . New genera of African poeciliid fishes. CO, no. 129, pp. 41-45. May 20. . A new rivulus from Rio de Janeiro. ANMG, ser. 9, vol. 13, pp. 588-590. June. [R. dorni; holotype in British Museum. This species is a synonym of R. brasiliensis Valenciennes, according to G. S. M.] . A new poeciliid fish from the Congo, with remarks on funduline genera. AMN, no. 116, 9 pp. June 6. . A new poeciliid fish of the genus Micropanchax from Ubangi. AMN, no. 112, 3 pp. June 24. . Amphibians and reptiles from Wilmington, N. C. CO, no. 131, pp. 59-62. June 30. . On the existence of the Japanese killifish, Fundulichthys virescens. ANMG, ser. 9, vol. 14, pp. 253-254. August. . Mutanda ichthyologica. Neoborus Boulenger and Barbus rubripinnis Nichols and 44. 45. 45a. 46. a (or) 49. CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER. Griscom. Revue Zoologie Africaine, Tervuren, Belgium, vol. 12, no. 3, p. 397. August 1. Lucania ommata in the aquarium. FC, vol. 4, no. 1, p. 314. September. A new poeciliid fish of the genus Rzvulus from British Guiana. AMN, no. 129, 2 pp. September 23. A new characin fish from Rio de Janeiro. FC, vol. 4, no. 3, pp. 330-331. November. [The legend for the figure was omitted by the printer; it is given correctly in 295. The two cotypes (syntypes) are now in the U. S. National Museum. See also 48.] On a small collection of fishes from Upper Burma. AMN, no. 150, 7 pp. November 13. The largest rivulus. CO, no. 135, p. 96. November 18. 1925 Concerning melanodimorphism in killifishes. CO, no. 137, pp. 105-107. January 15. [The specimens of Platypoecilus couchianus mentioned were misidentified in the Field Museum collection; they were Mollienisia sphenops.| Description of a new catfish from Abyssinia. CO, no. 139, pp. 12-13. February 15. [See 520 for corrections. | [Description of Rivulus rogoaguae.| (Nathan Everett Pearson and G. S. Myers.) In: Pearson, N. E., Fishes of the Rio Beni Basin (Indiana University Studies, “1924,” vol. 11, no. 64), p. 51. February. Results of some recent studies on the American killifishes. FC, vol. 4, no. 8, pp. 370-371. April. [Contains original diagnosis of Trigonectes strigabundus, n. gen. and n. sp. from the Rio Tocantins. Reprinted as 234.] Introduction of the European bitterling (Rhodeus) in New York and of the rudd (Scardinius) in New Jersey. CO, no. 140, pp. 20-21. April 14. Ein neuer Characinide von Rio de Janeiro. BATK, Jahrg. 36, no. 4, pp. 98-99, 1 colored plate. April 15. [Translation of 41, omitting the original figure, but ac- companied by a colored plate by Curt Bessiger and by an article on breeding and care in aquaria, by Wilhelm Schreitmiller. | Notropis cummingsi, a new minnow from Wilmington, North Carolina. AMN, no. 168, 4 pp. April 23. [Although named for a woman, Mrs. J. H. Cummings, the ending of cummingsi was intentional, the rationale being that Cummings was the name of her husband. ] Concerning mollienisias. AL, vol. 9, no. 1, pp. 3-4, 13-14. May. Astyanax fasciatus in the aquarium. AL, vol. 9, no. 3, p. 40. July. Description of a new cheirodontine characin from Rio de Janeiro. Annals of the Carnegie Museum, Pittsburgh, vol. 16, pp. 143-144, pl. 10. July 31. [Spintherobolus broccae, n. sp. = Phoxinopsis typicus Regan. | Fishes changing sex. AL, vol. 9, no. 4, pp. 56-57. August. The blue characin, Coelurichthys microlepis. Aquarium Bulletin, St. Louis, vol. 2, no. 3, pp. 3-4. September 1. | Reprinted as 95.] Labyrinth fishes for the aquarist. Aquarium Bulletin, St. Louis, vol. 2, no. 4, pp. 3, 6. October. The aquarium and its denizens, being a brief exposition of the proper arrangement and maintenance of home aquaria, with a catalog of some of the fishes suitable for aquarium culture. Second edition, revised and enlarged; 12mo, 45 pp., August M. Roth, Publisher, Baltimore. November. [A revised edition of 12.] Tridentopsis pearsoni, a new pygidiid catfish from Bolivia. CO, no. 148, pp. 83-86. November 25. VoLt. XX XVIII] MYERS: BIBLIOGRAPHY 58. *64. 66. 68. 69. bho OL Fishes and human disease. FC, vol. 5, no. 4, pp. 27-29. December. [An account of the part played by fishes in malaria and yellow fever control. The reference to Astroblepus (Arges) is a lapsus; the genus intended is Pygidium. | 1926 59. Notes on anabantids. III. CO, no. 150, pp. 97-100. January 25. On the correct names of the tetra from Buenos Aires, the haplochilus from Madras, and the mouthbreeder. FC, vol. 5, no. 8, p. 61. April. Two new genera of African characin fishes. Revue Zoologie Africaine, Tervuren, Belgium, vol. 13, nos. 3-4, pp. 174-175. April 1. Die Nomenklatur der Labyrinthfische. BATK, Jahrg. 37, no. 8, pp. 190-193. April 30. Descriptions of a new characin fish and a new pygidiid catfish from the Amazon basin. CO, no. 156, pp. 150-152. July 20. [Proof not seen by author; there are several bad typographical errors which are corrected in 71.] A synopsis for the identification of the amphibians and reptiles of Indiana. Proceed- ings of the Indiana Academy of Science, vol. 35 (1925), pp. 277-294. [Published in summer, 1926. ] . A cichlid fish that hangs its young on aquatic plants. AL, vol. 10, no. 4, pp. 60-61. August. Eine neue siidamerikanische Characinidenart der Gattung Pyrrhulina. BATK, Jahrg. 37, no. 18, pp. 441-442. September 30. [The type locality, left indefinite in this paper, was later found to be Rosario, Argentina; syntypes now in U. S. National Museum. | Alphabetical list of aquarium fishes, their breeding habits, care, etc. Ju: Innes, W. T., Goldfish varieties and tropical aquarium fishes, 9th ed., Philadelphia, pp. 264-283. October. [All of this chapter, save the introduction (pp. 264-265) and the conclud- ing remarks (pp. 283-284), is by G. S. Myers. This same list appeared in the later, cheaper edition of this book, called “The Complete Aquarium Book,” published by Halcyon House, N. Y., in 1936.] Eine neue Characinidengattung der Unterfamilie Cheiredontinae aus Rio de Janeiro, Brasilien. BATK, Jahrg. 37, no. 24, pp. 566-567. December 30. [Syntypes of Rachoviscus crassiceps now in U. S. National Museum. This fish was probably not from the Baixada Flumenense but from farther inland. Perhaps equals Oligobrycon? | 1927 . An analysis of the genera of neotropical killifishes allied to Révulus. ANMG, ser. 9, vol. 19, pp. 115-129. January. Note [correcting typographical errors in no. 63.] CO, no. 158, pp. 167-168. January. . Rana areolata at Bloomington, Indiana. (Herman P. Wright and G. S. Myers.) CO, no. 159, pp. 173-175. January 11. [First description of eggs. ] . On the identity of the killifish Fundulus meeki Evermann with Fundulus lima Vaillant. CO, no. 160, p. 178. January 12. . Puntis streeteri, a new cyprinoid fish from Borneo, and Cobitophis, a new genus of Bornean Cobitidae. AMN, no. 265, 4 pp. April 20. . The status of the darter Richiella brevispina (Coker). CO, no. 163, pp. 39-43. June. . The differential characters of Bufo americanus and Bufo fowlert. CO, no. 163, pp. 50-53. June. Descriptions of new South American fresh-water fishes collected by Dr. Carl Ternetz. Bulletin of the Museum of Comparative Zoology, Harvard, vol. 68, no. 3, pp. 107-135. 26 90. Or. 96. 97. *98. we). CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. July. [Types of Otothyris canaliferus now in U. S. National Museum, Paris, and London; paratypes of Bunocephalus salathei in Washington. See also 163 for corrections. | 8. A new genus of Brazilian characin fishes allied to Bivibranchia. (Carl H. Eigenmann and G. S. Myers.) Proc. National Academy of Sciences, Washington, vol. 13, no. 8, pp. 565-566. August. Notes on Indiana amphibians and reptiles. Proceedings of the Indiana Academy of Science, vol. 36 (1926), pp. 337-340. September. Rasboras. FC, vol. 7, no. 2, pp. 175-177. October. 1928 Carl H. Eigenmann—Ichthyologist. Natural History, New York, vol. 28, no. 1, pp. 98-101, portrait. The systematic position of the phallostethid fishes, with diagnosis of a new genus from Siam. AMN, no. 295, 12 pp. February 1. The species of Piabucina inhabiting Colombia. CO, no. 166, pp. 4-5. March 23. Two new genera of fishes. CO, no. 166, pp. 7-8. March 23. . Haplochilus cameronensis. AL, vol. 11, no. 12, p. 204. April. [A short note written in 1923 and published without author’s knowledge in 1928. See 89.] . The existence of cichlid fishes in Santo Domingo. CO, no. 167, pp. 33-36. June 28. . The urostyle in larval characin fishes. CO, no. 167, pp. 36-37, June 28. New fresh-water fishes from Peru, Venezuela, and Brazil. ANMG, ser. 10, vol. 2, pp. 83-90. July. [Two of the species are described by C. H. Eigenmann, and one jointly by Eigenmann and Myers. | “Haplochilus cameronensis.’ AL, vol. 12, no. 5, p. 94. September. [Correcting 85, above. | The characins. AL, vol. 12, no. 7, pp. 119-120, 122, 134-135. November. The characins. AL, vol. 12, no. 8, p. 152. December. [Continuation of 90.] 1929 The happy family tank. FC, vol. 8, no. 5, pp. 51-52. January. [On “community aquaria.”’ | The history of the veiltail fighting fish. AL, vol. 12, no. 10, pp. 195-199. February. [History of the original introduction of long tailed cultivated Betta splendens into the United States. ] Cranial differences in the African characin fishes of the genera Alestes and Brycinus, with notes on the arrangement of related genera. AMN, no. 342, 7 pp. March 2. . The blue characin, Mimagoniates microlepis. FC, vol. 8, no. 8, pp. 92-93. April. [A reprint of 54, above, with emendations. | Mutanda icthyologica. II. Heringia vs. Rhinosardinia (Clupeidae), Medipellona vs. Chirocentrodon (Clupeidae), and Entonanthias vs. Mirolabrichthys (Anthiidae). CO, no. 170, pp. 1-2. April 30. A note on the Formosan homalopterid fish, Crossostoma lacustre Steindachner. CO, no. 170, p. 2. April 30. Notes on the names of the spring peeper, the carpenter frog, and Anezdes aeneus. CO, no. 170, pp. 22-23. April 30. [See 121 for correction. ] On curimatid characin fishes having an incomplete lateral line, with a note on the peculiar sexual dimorphism of Curimatopsis macrolepis. ANMG, ser. 10, vol. 3, pp. 618-621. June. VOL. 100. 101. 102. * 103: *104. 105. *100. 107. 108. 109. 110. *118. 119. XXXVIIT] MYERS: BIBLIOGRAPHY Oe Notes on soles related to Achirus. CO, no. 171, pp. 36-38. June 28. The American Characidae. [Part 5] (Carl H. Eigenmann and G. S. Myers.) Memoirs of the Museum of Comparative Zoology, Harvard, vol. 43, part 5, pp. 429-558, 11 pls. September. [The supplement, pp. 516-550, is nearly all by G.S.M; the rest of the text is nearly all by Eigenmann. | Our aquarium fishes. I. The mouthbreeder. AJ, vol. 2, no. 8, p. 31. October 3. 1930 Amphibians and reptiles observed in the Palisades Interstate Park, New York and New Jersey. CO, no. 173, pp. 99-103. January 16. Notes on some amphibians in western North America. PBSW, vol. 43, pp. 55-64. March 12. Fishes from the upper Rio Meta basin, Colombia. PBSW, vol. 43, pp. 65-71. March 12. The status of the southern California toad, Bufo californicus (Camp). PBSW, vol. 43, pp. 73-77. March 12. On the occurrence and habits of ocean sunfish (Mola mola) in Monterey Bay, Cali- fornia. (G. S. Myers and Joseph Howe Wales.) CO, 1930, no. 1, pp. 11-12. April 30. The killifish of San Ignacio and the stickleback of San Ramon, Lower California. Proceedings of the California Academy of Sciences, ser. 4, vol. 19, no. 9, pp. 95-104. July 15. Ptychidio jordani, an unusual new cyprinoid fish from Formosa. CO, 1930, no. 4, pp. 110-113. December 31. |Type later found to represent a chance introduction in Formosa, originating from the pondfish-fry export industry centering at Wuchow. Genus and species are endemic to the Si Kiang (West River) system, near to and above Wuchow, Kwangsi, China. ] [Review of] Publications of the University of Oklahoma Biological Survey, vol. 1. By A. Richards, C. L. Hubbs, and A. I. Ortenburger. CO, no. 4, pp. 159-160. December 31. [Review, with critical comments, of] Hall, H. M., and Clements, F. E., The phylogenetic method in taxonomy. Micropaleontology Bulletin, Stanford Univer- sity, vol. 2, no. 3, pp. 55-58. December 31. 1931 Eigenmann, Carl H. Jn: Dictionary of American Biography, vol. 6, pp. 62-6 Charles Scribner’s Sons, New York. Killifishes in Hispaniola. FC, vol. 10, no. 6, pp. 103-104. February. ins) Ww . Ichthyological reminiscences of a trip east. AJ, vol. 4, no. 2, p. 9. February. Ichthyological reminiscences of a trip east (continued). AJ, vol. 4, no. 3, pp. 14-15. March 5. Ichthyological reminiscences of a trip east (concluded). AJ, vol. 4, no. 4, pp. 20-21. April 2. . The primary groups of oviparous cyprinodont fishes. Stanford University Publications, University Series, Biological Sciences, vol. 6, no. 3, pp. 241-254. [Copies first mailed April 7.] Ascaphus truet in Humboldt County, California, with a note on the habits of the tadpole. CO, 1931, no. 2, pp. 56-57. July 20. Fishes from southeastern China and Hainan. (Albert W. Herre and G. S. Myers.) LSJ, vol. 10, no. 2-3, pp. 233-254. August. [Key to Asiatic genera of Clupeidae by G.S.M. alone. See 126 for correction. ] 35). 136. 138. CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. On the fishes described by Koller from Hainan in 1926 and 1927. LSJ, vol. 10, nos. 2-3, pp. 255-262. August. The original descriptions of Bufo fowlert and Bufo americanus. CO, 1931, no. 3, pp. 94-96. October 30. Poeciliid fishes of the genus Mollienisia in Hispaniola, with notice of a new limia from the Samana Peninsula. AMN, no. 503, 2 pp. November 9. On the identity of Ophicephalus and Channa, two genera of labyrinth fishes. (G. S. Myers and Leo Shapovalov.) Peking Nat. Hist. Bull., Peiping, China, vol. 6 (1931- 32), pt. 2, pp. 33-37. November. [Vol. 6, part 2 has usually been considered to have been published in 1932. Copies of vol. 6, part 2 reached regular subscribers in California in December 1931, and publication in China must have been in November or earlier. | 1932 Gambusias in the aquarium. HAB, vol. 2, no. 1, pp. 2-5. March. [See also 150.] The osteoglossid fish Scleropages in the Malay Peninsula. CO, 1932, no. 1, p. 30. April 12. Nealosa Herre and Myers equals Konosirus Jordan and Snyder. CO, 1932, no. 1, p. 30. April 12. A new name for a Melanesian pseudochromid fish confused with Neszotes purpurascens de Vis. CO, 1932, no. 1, p. 30. April 30. A neglected description of a Mexican garter-snake, Thamnophis stejnegert McLain. CO, 1932, no. 1, p. 35. April 12. [Review of] Osborn, H. F. Cope: Master Naturalist, and Biographical memoir of Edward Drinker Cope. CO, 1932, no. 1, pp. 39-41. April 12. Danio analipunctatus identified as Brachydanio nigrofasciatus. TA, vol. 1, no. 2, p. 54. June. [Boulenger’s holographic description of Danio analipunctatus, which he sent to J. P. Arnold and which was published by Arnold in BATK, was sent to G.S.M. by Arnold, and is now in Library of U. S. National Museum. ] A note on the two Chinese paradise fishes. HAB, vol. 2, no. 5, p. 9. July. [Pre- liminary synopsis of 137.] Some new aquarium fishes from Panama. TA, vol. 1, no. 3, pp. 68-69, 82. July. Some notes on the characin, Astvanax mexicanus, in Texas. AL, vol. 16, no. 3, pp. 97-98. July. A new gonostomatid fish, Neophos nexilis, from the Philippines. CO, 1932, no. 2, pp. 61-62. July 1. A new whitefish, Prosopium snyderi, from Crescent Lake, Washington. CO, 1932, no. 2, pp. 62-64. July 1. Fundulus chrysotus, geschechte Abart. WATK, Jahrg. 29, no. 29, pp. 450-451. July 19. [Notes on black-spotted, melanic or melanodimorphic specimens of Fundulus, Gambusia, and Mollienisia. See also 44.] The two Chinese labyrinth fishes of the genus Macropodus. LSJ, vol. 11, no. 3, pp. 385-403, pls. 6-7. July 22. [A taxonomic revision of the genus. Proof not seen by author; many typographical and editorial errors, especially in explanation of plates. See also 131; see 576 for additional information and another synonym. | A rare deep-sea scombrid fish, Yenogramma carinatum Waite, on the coast of southern California. Trans. San Diego Society of Natural History, vol. 7, no. 11, pp. 111-117, VoL. 143. 144. 145. 146. #147, 148. 149. 150. TSE 1527 15S. 154. 155. 156. USE XXXVIIT] MYERS: BIBLIOGRAPHY 29 pl. 7. July 28. [First American record. First synonymization of several nominal species. Species later known as Lepidocybium flavobrunneum (A. Smith) 1849.] . Dangers in identifications. TA, vol. 1, no. 4, pp. 94-97, 110. August. . Pterophyllum, king of aquarium fishes. TA, vol. 1, no. 5, pp. 115-118, 140-141. September. [A systematic review of the 3 species. | . A new genus of funduline cyprinodont fishes from the Orinoco Basin, Venezuela. PBSW, vol. 45, pp. 159-162. September 27. Recent importations—Stethaprion innesi and Mylossoma aurem from the Amazon. TA, vol. 1, no. 6, pp. 149-150, 171. October. [This will probably have to stand as the original description of S. imnesi, which was formally described in 148. Re- printed as 262.]| A native fish, Notropis lutrensis, in the aquarium. AJ, vol. 5, no. 8, pp. 45-46. October 6. Notes on Colombian fresh-water fishes, with description of a new astroblepus. CO, 1932, no. 3, pp. 137-138. October 7. 1933 Pachypanchax, a new genus of cyprinodont fishes from the Seychelles Islands and Madagascar. AMN, no. 592, 1 p. January 23. A new genus of Chinese fresh-water serranid fishes. Hong Kong Naturalist, vol. 4, no. 1, p. 76. April. [Proposing the new genus Acroperca, which was antedated by Coreosiniperca Fang and Chong 1932. The difference in date was about three months. |] Two records of the leatherback turtle on the California coast. CO, 1933, no. 1, p. 44. April 3. Description of a new characid fish of the genus Stethaprion from the Lower Amazon. ANMG, ser. 10, vol. 11, pp. 604-605. May. [See 142.] Stevardia albipinnis 2? HAB, vol. 3, no. 4, p. 11. June. Gambusen im Aquarium. WATK, Jahrg. 30, no. 26, pp. 401-403. June 27. [Transla- tion of 124.] [Review of] Hora, S. L., Classification, bionomics and evolution of homalopterid fishes. CO, 1933, no. 2, p. 109. July 20. The classification of the African cyprinodont fishes with a discussion of the geo- graphical distribution of the Cyprinodontidae of the world. Stanford Univ. Bull., ser. 5, no. 158 (Abstracts of dissertations, vol. 8, 1932-33), pp. 10-12. July 31. [Brief abstract of doctorate thesis, which was more usefully abstracted as 157 and 214.] Anent Mr. Schoenfeld on scientific names. HAB, vol. 3, no. 6, pp. 6-11. August. New importations—“Jack Dempsey” unmasked. Aquarium, Philadelphia, vol. 2, no. 6, pp. 141-142. October. [Reidentifies fish previously known to aquarists as Cichla- soma nigrofasciatum—the “Jack Dempsey”—as C. biocellatum.| Note on the breeding habits of Corynopoma. FC, vol. 13, no. 3, p. 61. November. New importations—Leopard Corydoras. TA, vol. 2, no. 8, pp. 188-189. December. {Contains first diagnosis of Corydoras leopardus, which is compared to C. julii and C. trilineatus. The species is formally described in 178. Translated as 166.] The genera of Indo-Malayan and African cyprinodont fishes related to Panchax and Nothobranchius. CO, 1933, no. 4, pp. 180-185. December 27. [See also 180.] 169. Se 174. 17/5). 176. 177. CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. 1934 The identification of aquarium fishes related to Metynnis and Serrasalmus. FC, vol. 13, no. 5, pp. 120-122. January. [Generic identification of live specimens. ] New importations—The black-winged flying characin. (Carnegiella marthae Myers.) TA, vol. 2, no. 9, pp. 217-218. January. [First account of life colors and habits, first record from Amazon basin, and first photograph of living specimens. See also 186 and 289, and comments in 345.] [Radio interview on fish work in the National Museum.] AL, vol. 17, no. 9, pp. 239- 240. January. [Review of] Coates, W., Tropical fishes for a private aquarium. FC, vol. 13, no. 6, p. 154. February. Reports on the collections obtained by the first Johnson-Smithsonian Deep-sea Ex- pedition to the Puerto Rican Deep. Three new deep-water fishes from the West Indies. Smithsonian Miscellaneous Collections, vol. 91, no. 9, 12 pp., 1 pl. April 2. Corrections of the type localities of Metzia mesembrina, a Formosan cyprinid, and of Othonocheirodus eigenmanni, a Peruvian characin. CO, 1934, no. 1, p. 43. April 24. A new name for the Alaskan cottoid fish Ulca marmorata (Bean). CO, 1934, no. 1, p. 44. April 24. [Review of] Regan, C. T., and Trewavas, E., Deep-sea angler fishes (Ceratioidea). CO, 1934, no. 1, pp. 54-55. April 24. Le Corydoras Léopard. AP, no. 5, p. 77. May. [A re-edited translation of 156.] . Barbus partipentazona Fowler. TA, vol. 3, no. 4, p. 83. August. [See also 182.] . Our downtrodden helper, the snail. AL, vol. 18, no. 4, p. 74-76, 93. August. [Snails in aquaria. | Judging fish shows. TA, vol. 3, no. 5, pp. 103-106. September. Gnathocharax steindachneri, a new characin for the aquarium. HAB, vol. 4, no. 7, pp. 5, 29-30. September. [Records species from the Orinoco (Caicara) and from British Guiana (Rockstone) for the first time; also records Monocirrhus poly- acanthus from Rockstone, British Guiana. | Correct nomenclature. AL, vol. 18, no. 6, pp. 139-141. October. [Nomenclature of aquarium fishes. ] [Review of] Lederer, N., Tropical fish and their care. CO, 1934, no. 3, p. 143. October 31. Ueber den Namen des Zwergdrachenflossers, Corynopoma riisei Gill (=Stevardia albipinnis Gill). WATK, Jahrg. 31, no. 48, pp. 755-756. November 27. [Review of] Herre, A. W. C. T., Notes on fishes in the zoological museum of Stan- ford University, 1: The fishes of the Herre Philippine Expedition of 1931. CO, 1934, no. 4, pp. 196-197. December 31. 1935 Cichlasoma biocellatum. TA, vol. 3, no. 9, p. 196. January. The mouth-breeding fighting fish, Betta brederi. TA, vol. 3, no. 9, p. 210. January. [This must stand as the original description of B. brederi, which is formally des- cribed in 179. Translated as 181 and 191.] A new phallostethid fish from Palawan. PBSW, vol. 48, pp. 5-6. February. [Pro- poses new suborder Phallostethoidea. | Vor. XXXVIIT] MYERS: BIBLIOGRAPHY 31 178. Four new fresh-water fishes from Brazil, Venezuela and Paraguay. PBSW, vol. 48, pp. 7-13. February 6. [One new species, Corydoras leopardus, was first diagnosed in 156. ] 179. A new anabantid fish of the genus Betta from Johore. PBSW, vol. 48, pp. 25-26. February 6. [First diagnosed in 176.] 180. The genera of Indo-Malayan and African cyprinodont fishes allied to Panchax and Nothobranchius. Canadian Aquaria, London, Ontario, vol. 3, no. 3, pp. 42-45. March. [Unauthorized reprint of part of 157.] 181. Der maulbrutende Kampffisch Betta bredert. WATK, Jahrg. 32, no. 20, p. 307. May 14. [Translation of no. 176. See also 191.] 182. Barbus partipentazona, Fowler. AP, no. 19, p. 111. July. [Translation of 167.] 183. [Review of] Schreitmiiller, W., Leitfaden zur Pfilege und Zucht von einheimischen und fremlandischen Zierfischen, Seetieren, Schnecken und Wasserpflanzen nebst einem Anhang itiber Trocken- und Kunstfischfutterarten; 3rd edition. CO, 1935, no. Zap LOeeulyed's* 184. [Review of] Holly, M.; Meinken, H.; and Rachow, A., Die Aquarienfische in Wort und Bild. CO, 1935, no. 2, p. 107. July 16. 185. Fishes of the upper Potomac. Jn: Shosteck, Robert, The Potomac trail book (128 pp., 12mo; published by the Washington Post, Washington, D. C.), pp. 90-95. [Fish chapter edited by Shosteck and proof not seen by G.S.M. before publication. Para- graphs on the eel and the goldfish on pp. 94-95 were added by Shosteck. The term “upper Potomac” was insisted on by Shosteck; the paper concerns the fishes of the region above the estuary but below Great Falls. The book was published (placed on sale) on October 6. Copies in libraries of U.S. National Museum, Carl L. Hubbs, and G.S.M.] 186. Carnegiella marthae Myers. AP, no. 23, pp. 173-174. November. [Translation of 159. See also 289.] 187. An annotated list of the cyprinodont fishes of Hispaniola, with descriptions of two new species. Zoologica, New York, vol. 10, no. 3, pp. 301-316. November 29. 188. Reports on the collections obtained by the first Johnson-Smithsonian Deep-sea Ex- pedition to the Puerto-Rican Deep. A new genus of opisthognathid fishes. Smith- sonian Miscellaneous Collections, vol. 91, no. 23, 5 pp. December 24. [Contains notes on Owstoniidae, Macrurocyttus, etc.] 189. [Review of] Two new state ichthyological papers [Greene, C. W., the distribution of Wisconsin fishes, and O’Donnell, D. J., Annotated list of the fishes of Illinois]. CO, 1935, no. 4, pp. 196-197. December 31. [In this was attempted the only published history and evaluation of the unfortunate check-list of North American fishes by Jordan, Evermann, and Clark.] 1936 190. [The use of fishes in mosquito control.] Ju: Sweetman, H. L., the biological control of insects (Comstock Publishing Co., Ithaca, N. Y.), pp. 318-325. [This chapter, although not credited to G.S.M. as author, was written by him and is printed from the MS. he supplied to Sweetman, with the change of only a very few words. Most of the bibliography and references given in the MS., however, were omitted by Sweetman. | 191. Le Betta brederz. AP, no. 25, p. 15. January. [A translation of 176, with additional notes on habits and breeding. See also 181.] 198. 199. 200. NO (2) dN 206. 207. CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Note on Rhamphichthys cingulatus Brind. Aquarium News, Rochester, N. Y., vol. 3, no. 5, p. 68. January. 3. On the Indo-Australian fishes of the genus Scatophagus, with description of a new genus, Selenotoca. PBSW, vol. 49, pp. 83-85. July 3. 4. A new characid fish of the genus Hyphessobrycon from the Peruvian Amazon. PBSW, vol. 49, pp. 97-98. July 3. . A third record of the albulid fish Dixonina nemoptera Fowler, with notes on an albulid from the Eocene of Maryland. CO, 1936, no. 2, pp. 83-85. July 31. A note on the stephanoberycid fishes. CO, 1936, no. 2, p. 118. July 31. [Resuscitates Acanthochaenus luetkeni Gill = Stephanoberyx gilii.| [Review of] Herre, A. W. C. T., The fishes of the Crane Pacific Expedition. CO, 1936, no. 2, pp. 128-129. July 31. [Notes that Disparichthys is probably not a genus of eels and may be near the blennies, and that Alepideleotris equals Eleotrica.] A new genus of gymnotid eels from the Peruvian Amazon. PBSW, vol. 49, pp. 115- 116. August 22. [Oedemognathus exodon; also synonymizes Tateichthys duidae LaMonte with Steatogenys elegans (Steindachner). | A new polynemid fish collected in the Sadong River, Sarawak, by Dr. William T. Hornaday, with notes on the genera of Polynemidae. Journal of the Washington Academy of Science, vol. 26, no. 9, pp. 376-382. September 15. [Revision of genera of Polynemidae. | Report on the fishes collected by H. C. Raven in Lake Tanganyika in 1920. Proceed- ings of the United States National Museum, vol. 84, no. 2998, pp. 1-15, pl. 1. September 24. . Description of a new blennioid fish of the genus Acanthemblemaria from the Pacific coast of Panama. (G. S. Myers and Earl D. Reid.) Allan Hancock Pacific Expedi- tions, vol. 2, no. 2, pp. 7-10. December. [Note on the identity of the dead fish in the Tidal Basin at Washington, D. C.] PBSW, vol. 49, p. viii. [Oral communication; no title. It concerned thousands of adults of Opisthonema oglinum floating dead in the Tidal Basin after thawing of the heavy ice in spring, 1936. This section of the Proceedings was published at the end of the year. | 1937 . Notes on phallostethid fishes. Proceedings of the United States National Museum, vol. 84, no. 3007, pp. 134-143. January 6. . The deep-sea zeomorph fishes of the family Grammicolepidae. Proceedings of the United States National Museum, vol. 84, no. 3008, pp. 145-156, 3 pls. January 18. [A] possible method of evolution of oral brooding habits in the cichlid fishes. AJ, vol. 10, no. 4, pp. 4-6. April. [Printer omitted the “A” in the title. Reprinted in facsimile, except for title, as 218. See also 223.] A contribution to the ichthyology of the Malay Peninsula, Part II. Fresh-water fishes. (Albert W. C. T. Herre and George S. Myers.) Bull. Raffles Mus., Singa- pore, no. 13, pp. 53-75, pls. 5-7. August. [|The first part of this paper, on marine fishes, and the general introduction, are by Herre alone, although the title page would lead one to think otherwise. ] [Review of] Monographs on the fishes of the Iberian Peninsula and Madeira. [Buen, F. de, Catalogo de los peces Ibéricos; Lozano Rey, L., Los peces fluviales de Espana; Nobre, A., Fauna marinha de Portugal, I, Vertebrados; Noronha, A. C. de, VOL. 208. 209. 210. eve DMZ: 213. 214. Dale 216. Dili 218. XXXVIIT] MYERS: BIBLIOGRAPHY 33 and Sarmento, A. A., Os peixes dos mares do Madeira.] CO, 1937, no. 4, pp. 239-240. December 31. [Review of] Three new Asiatic check lists [Suvatti, C., Index to fishes of Siam; Roxas, H. A., and Martin, C., A check list of Philippine fishes; Mori, T., and Uchida, K., A revised catalogue of the fishes of Korea]. CO, 1937, no. 4, pp. 241-242. Decem- ber 31. 1938 [Review of] Hubbs, C. L., and Trautman, M. B., A revision of the lamprey genus Ichthyomyzon. CO, 1938, no. 1, p. 51. March 31. Foreword. SIB, vol. 1, no. 1, pp. 1-2. June 22. !Explains editorial policy and aims of the Bulletin.] Hydromantes platycephalus in Sonora Pass, California. CO, 1938, no. 2, p. 91, June 30. Notes on Ansorgia, Clarisilurus, Wallago, and Ceratoglanis, four genera of African and Indo-Malayan catfishes. CO, 1938, no. 2, p. 98. June 30. [See 322 for correc- tion. | Fresh-water fishes and West Indian zoogeography. Annual Report Smithsonian In- stitution for 1937, publication 3465, pp. 339-364, 3 pls. [volume appeared late in the summer of 1938.] Studies on the genera of cyprinodont fishes. XIV. Aplocheilichthys and its relatives in Africa. CO, 1938, no. 3, pp. 136-143. September 24. [Numbers 32, 34, 37, 46, 70, 84, 117, 141, 145, 157, 178, 187, and 200 are taken to be the first 13 papers in this series. ] [Review of] Wells, L. A., Tropical aquariums, plants and fishes. CO, 1938, no. 3, p. 152. September 24. Harvest of the sea. Stanford Illustrated Review, vol. 40, no. 2, pp. 20-21, 25-26. October. [Originally given as a radio address. | [Review of] Costen, H. E. T., Beneath the surface, the cycle of river life. CO, 1938, no. 4, p. 208. December 10. 1939 A possible method of evolution of oral brooding habits in cichlid fishes. SIB, vol. 1, no. 3, pp. 85-87. February 3. [A reprint, in photographic facsimile except for the title, of 205. See also 223.] . Notes on the labrid genus Lienardella. SIB, vol. 1, no. 3, pp. 87-88. February 3. . On the Brazilian characid fish Notropocharax difficilis Marini, Nichols and La Monte. SIEBy vol. Ie nol 3, pac. Hebruany: 3. [Review of] Norman, J. R., Discovery reports, coast fishes. CO, 1939, no. 1, p. 61. March 9. . A new owstoniid fish from deep water off the Philippines. PBSW, vol. 52, pp. 19-20. March 11. . Mouthbreeding in cichlid fishes. Aquarist and Pond-keeper, London, vol. 9, no. 3, pp. 90-91, 94. May. [Reprint of 205. See also 218.] . Hesperomyrus fryi, a new genus and species of echelid eels from California. (G. S. Myers and Margaret Hamilton Storey.) SIB, vol. 1, no. 4, pp. 156-159. May 24. . The possible identity of the Congo fish Teleogramma with the cichlid genus Leptolamprologus. SIB, vol. 1, no. 4, p. 160. May 24. 34 226. bo W bo 246. CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. A living coelacanth fish. CO, 1939, no. 2, p. 124. July 12. [A note on the discovery of Latimeria chalumnae in South Africa, with references to accounts in popular magazines. | [Review of] Watson, D. M. S., The acanthodian fishes. CO, 1939, no. 3, p. 178. September 9. [Review of] Tchernavin, V., Changes in the salmon skull. CO, 1939, no. 3, p. 178. September 9. [Obituary of Alipio de Miranda-Ribeiro, 1874-1939.] CO, 1939, no. 3, p. 184. Septem- ber 9. [See also 535.] [Review of] Moy-Thomas, J. A., Palaeozoic fishes. CO, 1939, no. 4, p. 239. Decem- ber 26. [Review of] Clements, F. E., and Shelford, V. E., Bio-ecology. (Lionel Albert Wal- ford and G. S. Myers.) CO, 1939, no. 4, p. 240. December 26. 1940 Suppression of Acaropsis and Chalcinus, two preoccupied names of South American fresh-water fishes. SIB, vol. 1, no. 5, p. 170. February 7. On the use of the generic name Barbus in ichthyology and ornithology. SIB, vol. 1, no. 5, p. 170. February 7. [Shows that “Barbus” of Cuvier is a plural vernacular term for a bird group, and does not preoccupy the fish name Barbus. |] Results of some recent studies on the American killifishes. SIB, vol. 1, no. 5, pp. 171- 172. February 7. [A photographic facsimile of 46.] Zoological nomenclature. Nature, London, vol. 145, no. 3668, pp. 264-265. February 17. [The word “many” in the MS. was misprinted “any” in fourth from last line of the last paragraph. | [Review of] Neave, S. A., Nomenclator zoologicus. CO, 1939, no. 1, p. 56. March 30. [Review of] Kuhne, E. R., A guide to the fishes of Tennessee and the mid-south. CO, 1939, no. 1, p. 58. March 30. [Remarks on resolution regarding dams and migratory fishes.] SIB, vol. 1, no. 6, p. 209. May 3. Cope as an ichthyologist. CO, 1940, no. 2, pp. 76-78, 1 pl. July 28. [In Edward Drinker Cope Centenary Number. ] A note on Monognathus. CO, 1940, no. 2, p. 141. July 28. [Selects a type species for Monognathus and proposes new genus Phasmatostoma for other species. | . The probable identity of Sphyraena chrysotaenia from the Red Sea and Arabia with S. aureoflamma from the Philippines, with notes on Naso vomer and N. lopez. CO, 1940, no. 2, p. 143. July 28. [Review of] Cutright, P. R., The great naturalists explore South America. CO, 1940, no. 2, p. 145. July 28. The nomenclatural status of the Asiatic fish genus Culter. CO, 1940, no. 3, pp. 199- 201. November 14. [See 516.] A note on the status of the generic name Corydoras. SIB, vol. 2, no. 1, pp. 11-12. December 23. [Inserted as part of paper by William Gosline on the Callichthyidae. | Suppression of some preoccupied generic names of fishes (Kessleria, Entomolepis, Pterodiscus, and Nesiotes), with a note on Pterophyllum. SIB, vol. 2, no. 1, pp. 33-36. December 23. [The cichlid generic name Pterophyllum is not preoccupied. | An American cyprinodont fish, Jordanella floridae, reported from Borneo, with notes on the possible widespread introduction of foreign aquarium fishes. CO, 1940, no. 4, Vou. XXXVIIT] MYERS: BIBLIOGRAPHY 35 247. 248. 249. 250. pp. 267-268. December 27. [First warning of the now worldwide introductions of tropical freshwater aquarium fishes in warmer regions. ] The neotropical anchovies of the genus Amplova. Proceedings of the California Academy of Sciences, 4th ser., vol. 23, no. 29, pp. 437-442. December 31. [This paper, written in 1926, follows Eigenmann’s system of giving total lengths (including caudal) of the specimens, but the length of the holotype of Amplova alleni, as given, is the standard length. The holotype was selected by Hildebrand from among what G.S.M. had intended to be syntypes of the species. | 1941 Suppression of Lissochilus in favor of Acrossocheilus for a genus of Asiatic cyprinid fishes, with notes on its classification. CO, 1941, no. 1, pp. 42-44. March 25. [Review of] Huxley, J., The new systematics. CO, 1941, no. 1, p. 61. March 25. The fish fauna of the Pacific Ocean, with especial reference to zoogeographical regions and distribution as they affect the international aspects of the fisheries. Proceedings, Sixth Pacific Science Congress, vol. 3, pp. 201-210. [Vol. 3 was issued in April 1941.] The work and program of the Natural History Museum of Stanford University in fisheries and general ichthyology. Proceedings, Sixth Pacific Science Congress, vol. 3, pp. 413-415. April. [See note under 250.] Four new genera and ten new species of eels from the Pacific coast of tropical America. (G. S. Myers and Charles Barkley Wade.) Allan Hancock Pacific Expeditions, vol. 9, no. 4, pp. 65-111, pls. 7-16. June 25. [ Review of] Schuchert, C., and Le Vene, C. M., O. C. Marsh, pioneer in paleontology. CO, 1941, no. 2, p. 121. July 8. [Review of] Sherborn, C. D., Where is the collection? An account of the various natural history collections which have come under the notice of the compiler. CO, USN ino), 25 jo. WHA, [irlby 3. [Review of] Fitch, H. S., A biogeographical study of the ordinoides Artenkreis of garter snakes (genus Thamnophis). CO, 1941, no. 2, pp. 122-123. July 8. [Review of] Phillips, W. J., The fishes of New Zealand. CO, 1941, no. 3, p. 187. September 30. [Review of] Berg, L. S., Classification of fishes, both recent and fossil. CO, 1941, no. 4, pp. 274-275. November 21. [Review of] Hubbs, C. L., and Lagler, K. F., Guide to the fishes of the Great Lakes and tributary waters. CO, 1941, no. 4, p. 275. November 21. [Review of] Parker, T. J., and Haswell, W. A., A text-book of zoology, 6th edition. CO, 1941, no. 4, pp. 275-276. November 21. [Comments on the decline of morphological zoology. | [Review of] Norris, H. W., The plagiostome hypophysis, general morphology and types of structure. CO, 1941, no. 4, p. 277. November 21. A new name for Taenionema, a genus of Amazonian siluroid fishes. SIB, vol. 2, no. 3, p. 88. November 27. 1942 . Stethaprion innesi and Mylossoma aureum. TA, vol. 10, no. 11, pp. 185-186. March. [A slightly altered reprint of no. 142.] [Review of] Longley, W. H., Systematic catalogue of the fishes of Tortugas, Florida, 36 bo oO On *274, Di Se 276. *283. CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. with observations on color, habits, and local distribution. CO, 1942, no. 1, pp. 57-58. March 24. [Gives a history of Dr. Longley’s submarine ichthylogical researches. | [Review of] Child, C. M., Patterns and problems of development. CO, 1942, no. 1, p. 58. March 24. . The Pacific American atherinid fishes of the genera Eurystole, Nectarges, Coleotropis, and Melanorhinus. (G. S. Myers and Charles Barkley Wade.) Allan Hancock Pacific Expeditions, vol. 9, no. 5, pp. 113-149, pls. 17-19. March 30. . A new frog from the Anamallai Hills, with notes on other frogs and some snakes from South India. PBSW, vol. 55, pp. 49-55. June 25. . A new frog of the genus Micrixalus from Travancore. PBSW, vol. 55, pp. 71-74. June 25. . Notes on Pacific Coast Tritwrus. CO, 1942, no. 2, pp. 77-82. July 10. [Review of] Beebe, W., Book of bays. CO, 1942, no. 2, p. 130. July 10. . [Description of Pimelodella peruana.| (Carl H. Eigenmann and G. S. Myers.) Jn: Eigenmann, C. H., and Allen, W. R., Fishes of western South America (University of Kentucky, Lexington), p. 101, pl. 3. Summer of 1942. . Studies on South-American fresh-water fishes. I. SIB, vol. 2, no. 4, pp. 89-114. August 24. . The “lungs” of Bothriolepis. SIB, vol. 2, no. 4, pp. 134-136. August 24. [Virtually predicts existence of such an agnathan genus as Jamoytius.| . The black toad of Deep Springs Valley, Inyo County, California. Occ. Pap. Mus. Zool. Univ. Michigan, no. 460, 13 pp., 3 pls. September 16. Neotropical lizards in the collection of the Natural History Museum of Stanford Uni- versity. (Charles E. Burt and G. S. Myers.) Stanford Univ. Publ., Univ. Ser., Biol. Sci., vol. 8, no. 2, pp. 273-324, portrait. October 6. Notes on some frogs from Peru and Ecuador. PBSW, vol. 55, pp. 151-155. October 17. [Review of] Marcgrave, J., Historia natural do Brasil. CO, 1942, no. 4, p. 269. December 28. [Portugese translation of Marcgrave. | 1943 The Myers Expedition. TA, vol. 11, no. 9, pp. 160-162. January. [Extracts from letter from G.S.M. to W. T. Innes on fishes of the Rio Japuhyba, near Angra dos Reis, State of Rio de Janeiro, Brazil. Figures are of aquarium fishes, supplied by editor. ] . The influence of Louis Agassiz on the ichthyology of Brazil. Revista Brasileira de Biologia, vol. 3, no. 1, pp. 127-133. March. [Review of] Eigenmann, C. H., and Allen, W. R., Fishes of western South America. CO, 1943, no. 1, pp. 60-61. March 31. [Includes comments on the Gregory and Conrad classification of Characidae. | [Review of] Santos, E., Anfibios e Répteis do Brasil. CO, 1943, no. 1, p. 60. March 31. . Notes on Rhyacotriton olympicus and Ascaphus truei in Humboldt County, California. CO, 1943, no. 2, pp. 125-126. June 30. George S. Myers reports. TA, vol. 12, no. 6, pp. 104-106. October. [Extracts from letter from G.S.M. to W. T. Innes on fishes in Minas Gerais, Brazil. ] Rediscovery of the Philippine discoglossid frog, Barbourula busuangensis. CO, 1943, no. 3, pp. 148-150. October 15. The lizard names Platyurus and Cosymbotus. CO, 1943, no. 3, p. 192. October 15. 283a. Sistematica geral de peixes e biologia da pesca. Rio de Janeiro. 84 pp. [This is a compiled set of notes taken by several students during a course of lectures given Vor. XX XVIII] MYERS: BIBLIOGRAPHY 37 *287. 288. 289. 290. 20 292. *300. by G.S.M. at the Museu Nacional in Rio de Janeiro, of which a large number of mimeographed copies were issued. The notes were not corrected by G.S.M. before mimeographing, and many errors are present. | 1944 . Field notes on fishes of the vicinity of Rio de Janeiro. TA, Philadelphia, vol. 12, no. 11, pp. 185-186. March. [Figure supplied by editor. ] . Field notes on fishes of the vicinity of Rio de Janeiro (concluded). TA, vol. 12, no. 12, pp. 204-206. April. [Figure supplied by editor. ] . A new species of carangid fish from the northeastern Pacific. (Lionel Albert Walford and G. S. Myers.) CO, 1944, no. 1, pp. 44-47. April 21. [Equals Trachurus symmetricus (Ayres), large adults. | California records of the western spade-foot toad. CO, 1944, no. 1, p. 58. April 21. Rhinobrycon negrensis, a new genus and species of characid fishes from the Rio Negro, Brazil. Proceedings of the California Academy of Sciences, ser. 4, vol. 23, no. 39, pp. 587-590. August 22. The black-winged flying characin (Carnegiella marthae Myers). TA, vol. 13, no. 7, pp. 105-106. November. [A reprint of 159. See also 186.] Two extraordinary new blind nematognath fishes from the Rio Negro, representing a new subfamily of the Pygidiidae, with a rearrangement of the genera of the family, and illustrations of some previously described genera and species from Venezuela and Brazil. Proceedings of the California Academy of Sciences, ser. 4, vol. 23, no. 40, pp. 591-602, pls. 52-56. November 7. Brazilian books of interest to ichthyologists and herpetologists. CO, 1944, no. 4, pp. 262-263. December 26. 1945 A new gurnard (Prionotus alipionis) from the coast of Brazil. (Gerard Warden Teague and G. S. Myers.) Boletim do Museu Nacional, Rio de Janeiro, n. s., zool., no. 31, 19 pp. January 24. [Teague wanted to describe this fish although G.S.M. was doubtful.] . A remarkable new genus of sexually dimorphic characid fishes from the Rio Paraguay Basin in Matto Grosso. (G. S. Myers and Paulo de Miranda-Ribeiro.) Boletim do Museu Nacional, Rio de Janeiro, n. s., zool., no. 32, 8 pp. January 25. . Possible introduction of Argentine toads into Florida. CO, 1945, no. 1, p. 44. March 31. . The habitat of Hyphessobrycon flammeus Myers. FC, vol. 24, no. 10, pp. 73-75. June. . A natural habitat of the house gecko (Hemidactylus mabouia) in Brazil. CO, 1945, no. 2, p. 120. June 30. . Notes on some new or little-known Brazilian amphibians, with an examination of the history of the Plata salamander, Ensatina platensis. (G. S. Myers and Antenor Leitao de Carvalho.) Boletim do Museu Nacional, Rio de Janeiro, n. s., zool., no. 35, 39 pp. August 25. [Original printed date of issue not correct. | . A third record of the Sonoran box turtle. CO, 1945, no. 3, p. 172. October 15. . A strange new leaf-nosed lizard of the genus Anolis from Amazonia. (G. S. Myers and Antenor Leitao de Carvalho.) Boletim do Museu Nacional, Rio de Janeiro, n. s., zool., no. 43, 22 pp. October 20. Nocturnal observations on sea-snakes in Bahia Honda, Panama. Herpetologica, vol. 3, no. 1, pp. 22-23. November 23. 38 *301. 307. *308. 309. 310. jilil 312. Sili3y 314, S1l5. 316. Sil 318. 319. CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. 1946 Lista provisoria dos anfibios do Distrito Federal, Brasil. Boletim do Museu Nacional, Rio de Janeiro, n. s., zool., no. 55, 36 pp. February 14. [Written in August 1944. Text in both Portugese and English. ] The introduction of the guppy (Lebistes) as an aquarium fish, and something on the origin of its name. TA, vol. 15, no. 3, pp. 46-48. March. On a recently proposed new family of deep-sea fishes (Barbourisiidae, Parr, 1945). CO, 1946, no. 1, pp. 41-42. April 30. [This and 290 both discuss absence of pelvic fins as a taxonomic character. | Occurrence of uranoscopid fishes of the western Pacific genus Gunathagnus in the American Atlantic fauna. CO, 1946, no. 1, p. 42. April 30. [Review of] Honig, P., and Verdoorn, F., editors, Science and scientists in the Netherlands Indies. CO, 1946, no. 1, p. 52. April 30. . New fishes of the families Dactyloscopidae, Microdesmidae, and Antennariidae from the west coast of Mexico and the Galapagos Islands with a brief account of the use of rotenone fish poisons in ichthyological collecting. (G. S. Myers and Charles Barkley Wade.) Allan Hancock Pacific Expeditions, vol. 9, no. 6, pp. 151-179 (pls. 20-23 included). December 16. 1947 The Amazon and its fishes. Part 1. The river. AJ, vol. 18, no. 3, pp. 4-9. March 3. Murray’s Reptiles of Sind, with a note on three forgotten descriptions of Indian sea- snakes, published therein. Herpetologica, vol. 3, no. 5, pp. 167-168. [The book mentioned is not Murray’s Vertebrate Zoology of Sind. |] The Amazon and its fishes. Part 2. The fishes. AJ, vol. 18, no. 4, pp. 13-20. April 1. [Review of] Smith, H. M., The fresh-water fishes of Siam, or Thailand. CO, 1947, no. 1, p. 69. April 20. [Review of] Hildebrand, S. F., A descriptive catalog of the shore fishes of Peru. CO, 1947, no. 1, p. 69. April 20. [Review of] Romer, A. S., Vertebrate paleontology. CO, 1947, no. 1, p. 70. April 20. [Review of] Woods, R. S., The naturalist’s lexicon. CO, 1947, no. 1, pp. 70-71. April 20. The Amazon and its fishes. Part 3. Amazonian aquarium fishes. AJ, vol. 18, no. 5, pp. 6-13, 32. May 1. The varieties of the Siamese fighting fish. AJ, vol. 18, no. 6, pp. 19-21. June 2. The Amazon and its fishes. Part 4. The fish in its environment. AJ, vol. 18, no. 7, pp. 8-19, 34. July. [Review of] Seale, A., Quest for the golden cloak. CO, 1947, no. 3, p. 213. September 12R Foreign introduction of North American fishes—Inadvisability of recommending North American fishes without careful appraisal of foreign fishes and ecology. Progressive Fish Culturist, Washington, D.C., vol. 9, no. 4, pp. 177-180. October. 1948 The ramirezi cichlid identified. (G. S. Myers and Robert Rees Harry, Jr.) TA, vol. 17, no. 4, p. 7. April. [This constitutes the original diagnosis of Apistogramma ramirezi; this species probably belongs to Geophagus. | VOL. 320. 324° S22) *323. Sie Soi) 336. 2Sfle XXXVIIT] MYERS: BIBLIOGRAPHY 39 [Review of] Hubbs, C. L., and Lagler, K. F., Fishes of the Great Lakes Region. CO; 1948, no. 2, p. 150. June 30. Apistogramma ramirezi, a cichlid fish from Venezuela. (G. S. Myers and R. R. Harry, Jr.) Proceedings of the California Zoological Club, vol. 1, no. 1, pp. 1-8. August. [See 319.] Notes on two generic names of Indo-Malayan silurid fishes, Wallago and Wallagonia. Proceedings of the California Zoological Club, vol. 1, no. 4, pp. 19-20. August. [Correction of 212.] The California plethodont salamander, Aneides flavipunctatus (Strauch), with descrip- tion of a new subspecies and notes on other western aneides. (G. S. Myers and Thomas Paul Maslin.) PBSW, vol. 61, pp. 127-135. September 3. . Proposed reprinting of Boulenger’s British Museum herpetological catalogues. Herpet- ologica, vol. 4, part 5, p. 180. September 13. [See also 325 and 352.] . Proposed reprinting of Boulenger’s British Museum herpetological catalogues. CO, 1948, no. 3, p. 229. [See also 324 and 352.] _ A list of common and scientific names of the better known fishes of the United States and Canada. (W. H. Chute, R. M. Bailey, W. A. Clemens, J. R. Dymond, S. F. Hildebrand, G. S. Myers and L. P. Schultz.) American Fisheries Society, Special Publication no. 1, 45 pp. [Fundulus in the West Indies.] In: Rivas, L. R., Cyprinodont fishes of the genus Fundulus in the West Indies (Proceedings of the United States National Museum, vol. 98, no. 3229, pp. 215-222), pp. 216-217. October 19. [Letter from G.S.M.] 1949 _ The Amazon and its fishes. Part 5. A monograph on the piranha [first part]. AJ, vol. 20, no. 2, pp. 52-61. February. [See 331.] Geographic variation in the ribbed frog, Ascaphus truei. (M. B. Mittleman and G. S. Myers.) PBSW, vol. 62, pp. 57-66. April 27. [Little save introduction is by G.S.M. | . A new frog of the genus Cornufer from the Solomon Islands, with notes on the endemic nature of the Fijian frog fauna. (Walter Creighton Brown and G. S. Myers.) AMN, no. 1418, 10 pp. May 9. _ The Amazon and its fishes. Part 5. A monograph on the piranha (concluded). AJ, vol. 20, no. 3, pp. 76-85. [Published May 31, although issue is for March; see 328.] Cichlid fishes in salt water. AJ, vol. 20, no. 6, pp. 147-149, 163. June. . Usage of anadromous, catadromous, and allied terms for migratory fishes. CO, 1949, no. 2, pp. 89-97. June 30. 4. Salt-tolerance of fresh-water fish groups in relation to zoogeographical problems. Bijdragen tot de Dierkunde, vol. 28, pp. 315-322. August. [J Festschrift for L. L. de Beaufort. | The family name of the characid fishes. CO, 1949, no. 3, pp. 195-204. September 15. Initial steps in the conservation of fresh-water fisheries in tropical South America, with remarks on fishery resources in general. Inter-American Conference on Con- servation of Renewable Natural Resources, held at Denver, Colorado, Sept. 7-20, 1948, pp. 501-506. October. [Proof not seen by author; references confused by editors. Published by the Department of State, Washington, D. C., October 1949.] A new frog of the genus Batrachylodes from the Solomon Islands. (Walter Creighton Brown and G. S. Myers.) Journal of the Washington Academy of Sciences, vol. 39, no. 11, pp. 379-380. November 15. 40 338. 354. 73508 356. SII 358. CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Status of the generic name Lioheterodon applied to Madagascan serpents. Herpetolog- ica, vol. 5, part 6, p. 146. December 15. [Text confused by printer; proof not seen by author; correctly reprinted as 341.] 1950 Systematic notes on some Amazonian clupeid fishes of the genus Jlisha. CO, 1950, no. 1, pp. 63-64. March 30. On the characid fishes called Hydrocynus and Hydrocyon by Cuvier. Proceedings of the California Zoological Club, vol. 1, no. 9, pp. 45-47. May 1. Status of the generic name Lioheterodon applied to Madagascan serpents. Herpetolog- ica, vol. 6, part 2, p.52. June 5. [No. 338, corrected. ] Manual of tropical herpetological collecting. 12 pp., mimeographed. Natural History Museum, Stanford University. June 15. Bibliography of the published papers of George Sprague Myers from 1920 to 1949 inclusive. 18 pp., mimeographed. June 15. Identity of the stromateid fish Centrolophus californicus with Icichthys lockingtoni. SIB, vol. 3, no. 4, p. 181. August 21. . Supplementary notes on the flying characid fishes, especially Carnegiella. SIB, vol. 3, no. 4, pp. 182-183. August 21. Studies on South American fresh-water fishes. II. The genera of anostomine characids. SIB, vol. 3, no. 4, pp. 184-198. August 21. A new lump-sucker of the genus Eumicrotremus from the northwestern Atlantic. (G. S. Myers and James Erwin Bohlke.) SIB, vol. 3, no. 4, pp. 199-202. August 21. Station records of the Crocker-Stanford Deep-sea Expedition, Coast of California, September 1938. (Rolf Ling Bolin and G.S. Myers.) SIB, vol. 3, no. 4, pp. 203-214. August 21. The “imitator catfish” which mimics a corydoras. (William Thornton Innes and G.S. Myers.) TA, vol. 19, no. 9, pp. 222-223. September 1. [See 366.] . The systematic status of Hyla septentrionalis, the large tree frog of the Florida Keys, the Bahamas and Cuba. CO, 1950, no. 3, pp. 203-214. September 5. [See also 447b.] | Review of] Bourret, R., Les batraciens de Indochine. CO, 1950, no. 3, pp. 243-244. September 5. Proposed reprinting of Boulenger’s herpetological catalogues. CO, 1950, no. 3, p. 244. September 5. [See also 324 and 325.] . A new genus of poeciliid fishes from Hispaniola, with notes on genera allied to Poecilia and Mollienisia. (Luis Rene Rivas and George S. Myers.) CO, 1950, no. 4, pp. 288-294. I plate. December 22. Flying of the halfbeak, euleptorhamphus. CO, 1950, no. 4, p. 320. December 22. [Review of] Liu, C. C., Amphibians of western China. CO, 1950, no. 4, p. 325. December 22. [Statement regarding A.S.I.H. Committee on Fish Classification.] CO, 1950, no. 4, p. 327. December 22. 1951 Notes on salamander voices. CO, 1951, no. 1, p. 76. March 21. [Review of] Hatch, M. H., editor, Studies honoring Trevor Kincaid. CO, 1951, no. 1, pp. 104-105. March 21. VOL. 360. 367. 368. 369. 370. Sle S12 = B/Se 374. $7153 376. SHU 378. 379. 380. XXXVIIT] MYERS: BIBLIOGRAPHY 41 [Review of] Tortonese, E., Gli animali superiori nella loro struttura e nella loro vita. CO, 1951, no. 1, p. 106. March 21. Study of fishes was the work David Starr Jordan loved best. Stanford Alumni Re- view, vol. 52, no. 7, pp. 13-15. March 27. . The Amazonian mottled knife-fish, Steatogenys elegans, and its strange vermiform organ. TA, vol. 20, no. 4, pp. 85-86. April 10. . The Amazonian checkerboard cichlid (Crenicara maculata). TA, vol. 20, no. 5, pp. 109-110. May 1. [See 367.] . Notas sobre la distribucion de los peces Sudamericanos del grupo Bivibranchia. Memorias Sociedad de Ciéncias Naturales La Salle, Caracas, tomo 10, no. 27 (for Sept—Dec. 1950), pp. 193-194. [Published in late Spring, 1951.] . The most widely heard amphibian voice. CO, 1951, no. 2, p. 179. June 8. [Voice of Hyla regilla in Hollywood sound-cinema. | . Asiatic giant salamander caught in the Sacramento River, and an exotic skink near San Francisco. CO, 1951, no. 2, pp. 179-180. June 8. . The “imitator catfish” which mimics a corydoras. (William Thornton Innes and G. S. Myers.) Jn: Innes, W. T. (editor), Aquarium Highlights, consisting of reprints of the most popular articles from the monthly magazine, the Aquarium, since 1932 (Innes Publishing Co., Philadelphia, 519 pp.), pp. 117-118. October. [Reprint of 349.] Amazon dwarf checkerboard cichlid. Jn: Innes, W. T. (editor), Aquarium Highlights (see 366 for full reference), pp. 120-121. October. | Reprint of 362.] Dangers in identifications. Jn: Innes, W. T. (editor), Aquarium Highlights (see 366 for full reference), pp. 366-371. October. [Reprint of 139.] Judging fish shows. In: Innes, W. T. (editor), Aquarium Highlights (see 366 for full reference), pp. 410-413. October. [Reprint of 169.] David Starr Jordan, ichthyologist, 1851-1931. SIB, vol. 4, no. 1, pp. 2-6. December 27. Fresh-water fishes and East Indian zoogeography. SIB, vol. 4, no. 1, pp. 11-21. De- cember 27. [See 483.] Some forgotten but available names for Indian fishes. SIB, vol. 4, no. 1, p. 26. De- cember 27. A new giant toad from Southwestern Colombia. (G. S. Myers and John W. Funk- houser.) Zoologica, New York, vol. 36, pt. 4, pp. 279-281, 1 pl. December 28. [Bufo blombergi, n. sp.] 1952 Tower of Babel. [Editorial.] CO, 1952, no. 1, pp. 57-58. June 2. [Obituary of] William G. Holbein. AJ, vol. 23, no. 6, pp. 118-119. June 12. Danio or Brachydanio, Barbus or Puntius? AJ, vol. 23, no. 6, pp. 121-122. June 12. [Vol. number erroneously given as 24 on the issue. | [Review of] Axelrod, H. R., Tropical fish as a hobby. CO, 1952, no. 2, pp. 120-121. June 26. [Review of] Ladiges, W., Der Fisch in der Landschaft. CO, 1952, no. 2, p. 121. June 26. [Review of] Steward, J. H., Handbook of South American Indians. Vol. 6. CO, 1952, no. 2, pp. 121-122. June 26. [Review of] Carter, G. S., Animal evolution, a study of recent views on its causes. CO, 1952, no. 2, p. 122. June 26. 398. 399. 400. 401. 402. 403. 404. 405. CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. [Review of] Kuenen, P. K., Marine geology. CO, 1952, no. 2, p. 122. June 26. [Review of] Suwatti, C., Fauna of Thailand. CO, 1952, no. 2, pp. 122-123. June 26. [Review of] Primer Congresso Nacional de Pesquerias Maritimas y Industrias Deri- vadas. CO, 1952, no. 2, p. 123. June 26. [Review of] Soljan, T., Fauna et Flora Adriatica, vol. 1, Pisces. CO, 1952, no. 2, p. 1235 june y2 OF [Review of] Stebbins, R. C., Amphibians of Western North America. CO, 1952, no. 2, pp. 123-124. June 26. [Obituary of] Chloe Leslie Starks. CO, 1952, no. 2, pp. 124-125. June 26. [America’s most able fish illustrator. | ._ A new dwarf toad from southeastern Brazil. (G. S. Myers and Antenor Leitao de Carvalho.) Zoologica, New York, vol. 37, no. 1, pp. 1-3. June 30. [Bufo pygmaeus, n. sp.] . Annual fishes. AJ, vol. 23, no. 7, pp. 125-141. July 11. [Obituary of] Louis L. Mowbray. AJ, vol. 23, no. 7, p. 141. July 11. On the problem of the status of names published by Scopoli in 1777 in his “Introduc- tio ad Historiam Naturalem.” Bulletin of Zoological Nomenclature, vol. 6, pt. 8, p. 255. July 23. [Reprinted in: Opinions and Declarations, vol. 9, pt. 23, p. 316.] Bibliography of the published papers of George Sprague Myers from 1920 to 1951, in- clusive. 20 pp., mimeographed. [Dated May 1951; issued during July 1952.] [Review of] Gohm, D., Tropical fish in the home. AJ, vol. 23, no. 8, pp. 152-153. August 7. [Review of] Carr, A., Handbook of turtles. AJ, vol. 23, no. 8, p. 153. August 7. . Hints to fish importers, no. 1. AJ, vol. 23, no. 8, pp. 156-157. August 7. [Obituary of] Johann Paul Arnold, 1868-1952. AJ, vol. 23, no. 9, pp. 169-170. Au- gust 20. Hints to fish importers, no. 2. AJ, vol. 23, no. 9, pp. 171-173. August 28. [Lake Tanganyika; helped initiate the reports of Tanganyikan aquarium fishes. | A note on the feathertail, an African characin (Phenacogrammus). AJ, vol. 23, no. 9, p. 173. August 28. [Review of] Hervey, G. F., and Hems, J., Freshwater tropical aquarium fishes. AJ, vol. 23, no. 9, pp. 174-177. August 23. The nature of systematic biology and of a species description. Systematic Zoology, vol. 1, pp. 106-111. September. Easiest of all to spawn and raise, the medaka. AJ, vol. 23, no. 10, pp. 189-194. Sep- tember 30. [Care, spawning, and raising Oryzias latipes. | Varieties of the three-spot gourami, Trichogaster trichopterus. AJ, vol. 23, no. 10, pp. 198-200. September 30. [Review of] Beck, P., Traité complét de la vie des animaux en aquarium. AJ, vol. 23, no. 10, p. 200. September 30. Amazonian tetras of the genus Thayeria. AJ, vol. 23, no. 10, pp. 206-207. September 30. [Species here called T. sanctae-mariae later described as T. boehlkez Weitzman. See also 429.] How the shooting apparatus of the archer fish was discovered. AJ, vol. 23, no. 10, pp. 210-214. September 30. Hints to fish importers, no. 3. AJ, vol. 23, no. 10, pp. 215-216. September 30. [Bur- mese fishes. ] A new Amazonian catfish for the aquarist. AJ, vol. 23, no. 11, pp. 224-225. October 31. [Corydoras elegans.] VOL. 406. 407. 408. 409. 410. 411. 412. 413. 414. 415. 416. 417. 418. 419. 420. 421. 422. 423. *424, 425. 426. 427. 428. 429. 430. 431. 432. 433. 434. XXXVI] MYERS: BIBLIOGRAPHY 43 Color schemes in your fishes. AJ, vol. 23, no. 11, pp. 228-230. October 31. Hints to fish importers, no. 4. AJ, vol. 23, no. 11, pp. 237-238. October 31. [His- paniola. ] [Review of] Ladiges, W., Der Fisch in der Landschaft. AJ, vol. 23, no. 11, p. 239. October 31. [Review of] Tropical Fish Hobbyist. AJ, vol. 23, no. 11, pp. 239-240. October 31. Corydoras elegans. TA, Philadelphia, vol. 21, no. 11, pp. 300-301. November. [Review of] L’Aquarium Exotique. AJ, vol. 23, no. 12, pp. 254-255. December 1. [Review of] Wendt, A., Die Aquarienpflanzen in Wort und Bild. AJ, vol. 23, no. 12, p. 255. December 1. [Review of] Holly, M., Meinken, H., and Rachow, A., Die Aquarienfische in Wort und Bild. AJ, vol. 23, no. 12, p. 255. December 1. [Review of] Lorenz, K., King Solomon’s ring. AJ, vol. 23, no. 12, p. 256. December 1. Hints to fish importers, no. 5. AJ, vol. 23, no. 12, p. 256. December 1. [Fundulus stellifer. | [Obituary of] Dr. Ernst Bade. AJ, vol. 23, no. 12, p. 264. December 1. [Review of] Innes, W. T., Exotic aquarium fishes, 15th edition. AJ, vol. 24, no. 1, p. 9. December 19. The miniature fish aquarium. AJ, vol. 24, no. 1, pp. 11-12. December 19. Hints to fish importers, no. 6. AJ, vol. 24, no. 1, p. 20. December 19. [Sierra Leone; Epiplatys annulatus.] Sharks and sawfishes in the Amazon. CO, 1952, no. 4, pp. 268-269. December 26. [Review of] Deraniyagala, P. E. P., A colored atlas of some vertebrates from Ceylon. CO, 1952, no. 4, p. 286. December 26. 1953 Spawning behavior of Polycentrus. AJ, vol. 24, no. 2, pp. 31-33. January 28. Hints to fish importers, no. 7. AJ, vol. 24, no. 2, pp. 33-34. January 28. [Chologaster.] [ Review of] Knight, M., Keeping reptiles and fishes. AJ, vol. 24, no. 2, p. 46. January 28. [Review of] Whitney, L. F., All about guppies. AJ, vol. 24, no. 3, p. 55. February. [Aquarium water-testers, filters, oil traps, and thermostatic heaters.] AJ, vol. 24, no. 3, pp. 60-61. February 23. Hints to fish importers, no. 8. Two beautiful little characins (Nematobrycon) trom western Colombia and their peculiar distribution. AJ, vol. 24, no. 3, pp. 64-65. Feb- ruary 23. [This initiated importation of the “Emperor tetra” as an aquarium fish. ] The living-fossil coelacanth fishes. AJ, vol. 24, no. 3, pp. 66-68. February 23. Vinkeltetrornas vetenskapliga nanm. Akvariet (Organ for Sveriges AkvariefOreningar), Argang 27, no. 3, pp. 42-44. March. [See also 402.] Queens of the water. AJ, vol. 24, no. 4, pp. 75-78. March 30. [An essay on water- lilies: Nuphar, Nelumbo, Nymphaea, Victoria, and Euryale.| Pets. [Verse.] AJ, vol. 24, p. 78. March 30. Hints to fish importers, no. 9. AJ, vol. 24, no. 5, p. 112. April 29. [Moenkhausia costae. | [Review of] Whitney, L. F., The complete book of home pet care. AJ, vol. 24, no. 5, p. 119. April 29. [Review of] Knowles, F. G. W., Freshwater and saltwater aquaria. AJ, vol. 24, no. 5, p. 119. April 29. CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. [Review of] Ladiges, W., Zierfisch Bilderbuch. AJ, vol. 24, no. 5, p. 119. April 29. [Review of] Kramer, K., and Weise, H., Aquarienkunde. AJ, vol. 24, no. 5, pp. 119- 120. April 29. The coelacanth fishes—living fossils. TA, vol. 22, no. 5, pp. 145-146. May. [Figure erroneously labeled. | . Aquarium difficulties with black mollienesias. AJ, vol. 24, no. 6, pp. 125-129. May 29. [This paper was later reprinted at least twice in the same journal, but references are not at hand. ] Hints to fish importers, no. 10. A strange glandulocaudine characin from the Rio das Velhas (Hysteronotus). AJ, vol. 24, no. 6, p. 137. May 29. [Review of] Leutscher, A., Vivarium life. AJ, vol. 24, no. 6, p. 141-142. May 29. [Review of] Evans, A., Aquariums. AJ, vol. 24, no. 6, p. 142. May 29. [Review of] Ichthys. AJ, vol. 24, no. 7, p. 157. June 29. [Review of] Tropical fish handbook-catalog. AJ, vol. 24, no. 7, pp. 157-158. June 29. [Review of] Taschenkalender fiir Aquarien und Terrarienfreunde, 1953. AJ, vol. 24, no. 7, p. 158. June 29. [Review of] Rounsefell, G. A., and Everhart, W. H., Fishery science; its methods and applications. AJ, vol. 24, no. 7, p. 158. June 29. [Review of] New Zealand Aquatic World. AJ, vol. 24, no. 7, p. 158. June 29. . Hints to fish importers, no. 11. Glass catfishes. AJ, vol. 24, no. 7, pp. 161-162. June 29. [Kryptopterus, Ailiichthys, Parailia, Physailia, Pseudepapterus. Evolutionary convergence in 3 families of catfishes.] 447a. [Quotation from letter regarding retention of original spellings of zoological names. ] Bulletin of Zoological Nomenclature, vol. 10, pt. 7, p. 216. July 14. *447b. [On the acceptance of certain names originally published in synonymy, especially 448. 449. 455, 456. 457. 458. 459. Eleutherodactylus and Hyla septentrionalis.| Bulletin of Zoological Nomenclature, vol. 10, pts. 10-11, p. 312. July 24. Piabucus in the aquarium. AJ, vol. 24, no. 8, pp. 172-174. July 28. [Identification of live examples of Pzabucus and similar genera. | Unbanded color variety of the Malayan “coolie” loach, Acanthophthalmus semicinctus. AJ, vol. 24, no. 8, p. 174. July 28. . The Florida pigmy topminnow, Leptolucania ommata, its history, and a record of the first California breeding. AJ, vol. 24, no. 8, pp. 184-187. July 28. . On aquarium magazines. |Editorial.] AJ, vol. 24, no. 8, pp. 188-189. July 29. . The Cuban green glass fish, Atherina evermanni. AJ, vol. 24, no. 9, pp. 195-197. Sep- tember. [For correction of name to Alepidomus evermanni, see same journal, vol. 24, no. 11, p. 170.] . Why show standards for most tropical fishes are unwise. AJ, vol. 24, no. 9, pp. 200— 202. September. . Publication dates of the Aquarium Journal. [In 1952-1953.] AJ, vol. 24, no. 9, p. 202. September. Hints to fish importers, no. 12. Rivulus ornatus. AJ, vol. 24, no. 9, p. 208. September. Neon tetra. AJ, vol. 24, no. 9, pp. 210-211. September. [Obituary of] Floyd S. Young. AJ, vol. 24, no. 9, pp. 211-212. September. What’s wrong with aquarists? Tropical Fish Tales, Springfield, Mass., vol. 2, no. 4, pp. 6-7. September. Zebra danio. AJ, vol. 24, no. 10, pp. 230-231. October. [Brachydanio rerio.] Vou. XXXVIIT] MYERS: BIBLIOGRAPHY 45 460. 461. 462. 463. 464. 465. 466. 467. 468. 469. 470. 471. 472. 473. 474. 475. 476. 477. 478. 479. 480. 481. #482. 483. 484. Classification of the danios. AJ, vol. 24, no. 10, pp. 235-238. October. [Critical taxo- nomic evaluation of four genera, Danio, Daniops, Allodanio, Brachydanio. See also 513a.] Where, oh where do the pictures come from? [Editorial.] AJ, vol. 24, no. 10, pp. 242-243. October. Hints to fish importers, no. 13. Rivulus sygonectes. AJ, vol. 24, no. 10, p. 244. October. Notes on selecting an aquarium. AJ, vol. 24, no. 11, pp. 254-256. November. [Review of] Innes, W. T., Exotic aquarium fishes, 16th Edition. AJ, vol. 24, no. 11, p. 258. November. Serpa tetra. AJ, vol. 24, no. 11, pp. 264-265. November. [Hyphessobrycon callistus.] Hints to fish importers, no. 14. Garmanella pulchra. AJ, vol. 24, no. 11, p. 266. No- vember. A note on the habits and classification of Corydoras hastatus. AJ, vol. 24, no. 11, pp. 268-270. November. [Subgenus Microcorydoras.]| The Christmas-tree fish. [Fantasy.] AJ, vol. 24, no. 12, p. 280. December. The secrets of the German fish breeders. [Editorial.] AJ, vol. 24, no. 12, pp. 287-288. December. Hints to fish importers, no. 15. Barbus candens. AJ, vol. 24, no. 12, p. 296. December. Habits of the spotted knife-fish (Steatogenys elegans) in the aquarium. AJ, vol. 24, no. 12, pp. 297-298. December. 1954 Hints to fish importers, no. 16. A fish that hasn’t been discovered yet. AJ, vol. 25, no. 1, p. 5. January. [Predicts the occurrence of a relative of Cubanichthys and Chrio- peopoides in Hispaniola. ] [Review of] Todd, R., The tropical fish book. AJ, vol. 25, no. 1, p. 7. January. Siamese fighting fish. AJ, vol. 25, no. 2, pp. 27-29. February. The fighting fish and its history. AJ, vol. 25, no. 2, pp. 30-33. February. [Betta splendens. | Hints to fish importers, no. 17. AJ, vol. 25, no. 2, p. 39. February. [Wild stock of Betta splendens. | [Review of] Emmens, C. W., Keeping and breeding aquarium fishes. AJ, vol. 25, no. 2, pp. 45-46. February. Fifty years of devotion to the aquarium hobby. TA, vol. 23, no. 2, pp. 35-39. Febru- ary. [Tribute to William Thornton Innes, on his 80th birthday.] Protective coloration in the leaf fish and Thayeria. AJ, vol. 25, no. 3, pp. 62-63. March. [Review of] Roberts, J. B., Jr., The pet shop manual. AJ, vol. 25, no. 3, pp. 77-78. March. [Review of] The Tropical Fish Magazine. AJ, vol. 25, no. 3, p. 78. March. Ability of amphibians to cross sea barriers, with especial reference to Pacific zoogeog- raphy. Proceedings 7th Pacific Science Congress [New Zealand, February 1949], vol. 4 (zoology), pp. 19-27. [Dated 1953; published March 1954.] Paleogeographical significance of fresh-water fish distribution in the Pacific. Proceed- ings 7th Pacific Science Congress [New Zealand, February 1949], vol. 4 (zoology), pp. 38-48. [When printing of the Proceedings had already been delayed for nearly three years, the present paper was printed in the United States as 371. Vol. 4 of the Proceedings finally was published (dated 1953) in March 1954.]| The protection of rare and vanishing fishes. Proceedings 7th Pacific Science Congress 46 485. 486. 487. 488. 489. 489a. 490. 50la. CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. [New Zealand, February 1949], vol. 4 (zoology), pp. 691-694. [Dated 1953; pub- lished March 1954. Aside from the Australian lungfish, this represents the first plea for the conservation of non-food, non-game fishes. ] Blue gularis |[Aphyosemion caeruleum.| AJ, vol. 25, no. 4, pp. 87-88. April. How to preserve fish specimens for study. AJ, vol. 25, no. 4, pp. 89-90. April. Another new corydoras from Brazil. (G. S. Myers and Stanley Howard Weitzman.) AJ, vol. 25, no. 4, pp. 93-94. April. [Corydoras cochui, n. sp.) Hints to fish importers, no. 18. AJ, vol. 25, no. 4, p. 102. April. [Poecilocharax bovalliz. | A new corydoras. (G.S. Myers and William Thornton Innes.) TA, vol. 23, no. 4, p. 105. April. [Corydoras cochui Myers and Weitzman. See 487 for original descrip- tion. | [Supplementary note.] Opinions and Declarations rendered by the International Com- mission on Zoological Nomenclature, vol. 4, pt. 15, p. 167. April 21. [Concerns Raphistoma versus Belone. | Hints to fish importers, no. 19. An unknown characin from the Cerro Duida, Vene- zuela. AJ, vol. 25, no. 5, pp. 111-112. May. [A lost and still undescribed relative of Poecilocharax.| . The Amazon longfin, Pterolebias longipinnis. (Fritz Mayer and G. S. Myers.) AJ, vol. 25, no. 5, pp. 113-115. May. [Review of] Fisher, E. L., Marine tropicals. AJ, vol. 25, no. 5, pp. 126-128. May. [Review of] La vita nell ’acquario. Manuale catalogo dell ’Acquario di Bologna. AJ, vol. 25, no. 5, p. 128. May. . The black-banded sunfish. AJ, vol. 25, no. 6, pp. 133-134. June. [Mesogonistius chaetodon.| The name of the Indian glassfish (Chanda lala). AJ, vol. 25, no. 6, pp. 149-150. June. The kissing gourami. AJ, vol. 25, no. 7, pp. 155-156. July. [Helostoma temmincki.. | . A new cyprinodont fish from the Peruvian Amazon. AJ, vol. 25, no. 8, pp. 175-177. July 28. [Pterolebias peruensis, n. sp.] [Review of] Nachstedt, J., and Tusche, H., Breeding aquarium fishes. AJ, vol. 25, no. 8, p. 183. July 28. . A beautiful new cyprinodont fish from the Amazon. TA, vol. 23, no. 8, pp. 236-237. August. [Pterolebias peruensis; see 497 for original description. | . The life and times of Polycentrus and the leaf-fish tribe. Tropical Fish Magazine, Springfield, Mass., vol. 4, no. 2, pp. 6-7. October. [Suggests possible relationship between Nandidae, Datnioides, and Lobotes.| . What fish is that? Tropical Fish Magazine, Springfield, Mass., vol. 4, no. 3, pp. 8-9. November. 1955 Notes on the freshwater fish fauna of middle Central America, with especial reference to pond culture of Tilapia. Fish Papers, FAO, Rome, no. 2. 1955. [Pagination and exact date lacking; warns of dangers in Tilapia introductions. | [Limericks] Zn: Martin, H. R. (editor), The little limerick book (Peter Pauper Press, Mt. Vernon, N. Y., 62 pp.). 1955. Notes on the classification and names of cyprinodont fishes. Tropical Fish Magazine, Springfield, Mass., vol. 4, no. 7, p. 7. March 1. [Includes diagnoses of Pantanodon podoxys Myers, n. g., n. sp., and Potamophylax pygmaeus Myers and Carvalho, n. gen.,n. sp. Proof not seen by author. There are a few editorial and printer’s errors. ] VoL. XXXVIITI MYERS: BIBLIOGRAPHY 47 503. 504. 505. Off *508. Oly on oe Sige, 514. 515. 516. Bly English names of aquarium cichlids. Tropical Fish Magazine, Springfield, Mass., vol. 4, no. 7, pp. 8-9. March 1. [On p. 9, two columns of print are transposed. | The wonderful world under the sea. Part 1. Tropical Fish Magazine, Springfield, Mass., vol. 4, no. 8, pp. 8-9. April. Gambusinos—a new term proposed for “live-bearing toothcarps.” TA, vol. 24, no. 5, pp. 149-152. May. . The wonderful world under the sea. Part 2. Tropical Fish Magazine, Springfield, Mass., vol. 4, no. 9, pp. 11, 14. May. 1956 Manual of tropical herpetological collecting. Ed. 2. Natural History Museum of Stan- ford University, Circular no. 4, 13 pp. [Mimeographed; very limited edition; second edition of 342.] Brief directions for preserving and shipping specimens of fishes, amphibians and rep- tiles. Natural History Museum of Stanford University, Circular no. 5, 3 pp. [Mime- ographed; very limited edition. | [Comments on Axelrod, H., and Schultz, L. P., Handbook of tropical fishes.] TA, vol. 25, no. 2, pp. 60-61. February. [Quotes from letter from G. S. Myers.] . Two new Brazilian fresh-water fishes. (G. S. Myers and Stanley Howard Weitzman.) SIB, vol. 7, no. 1, pp. 1-4. February 21. [Hyphessobrycon cardinalis and Hassar praelongus. H. cardinalis was also described by Schultz as Cheirodon axelrodi, in a publication bearing the printed date February 20.] . The name of the South American clupeid fish, Pristigaster. CO, 1956, no. 1, pp. 63-64. February 29. [Review of] Cochran, D. M., Frogs of southeastern Brazil. CO, 1956, no. 1, p. 69. February 29. Zoological results of the California Himalayan Expedition to Makalu, Eastern Nepal. I. Amphibians and Reptiles. (Alan E. Leviton, G. S. Myers, and Lawrence W. Swan.) Occasional Papers of the Natural History Museum of Stanford University, no. 1, 18 pp. March 9. Classification des danios. L’Aquarium et les Poissons, Paris, 6me Année, no. 4, pp. 5— 8. April. [Translation of 460.] Note on guppies in Mexico. TA, vol. 25, no. 4, p. 123. April Studies on the fishes of the family Characidae. No. 11. A new genus and species of hemiodontins from the Rio Orinoco in Venezuela. (James Erwin Bohlke and G. S. Myers.) Notulae Naturae, Philadelphia, no. 286, 6 pp. May 23. Request for a ruling as to the species to be accepted as the type species of the genera Culter and Nasus Basilewsky, 1855 (class Pisces). Bulletin of Zoological Nomencla- ture, vol. 12, pp. 136-138. July. [Francis Hemming, Secretary of the I.C.Z.N., re- worded 243, added certain requests to it, and published it under the name of G. S. Myers. Reprinted in: Opinions and Declarations, 1958, vol. 18, pt. 17, pp. 294-297.] The Xenurobryconini, a group of minute South American characid fishes with teeth outside the mouth. (G. S. Myers and James Erwin Bohlke.) SIB, vol. 7, no. 2, pp. 6-12. August 30. Copella, a new genus of pyrrhulinin characid fishes. SIB, vol. 7, no. 2, pp. 12-13. Au- gust 30. . Esomus rehi, an Indo-Malayan cyprinid fish. SIB, vol. 7, no. 2, pp. 13-14. August 30. [Pogonocharax rehi Regan, described as from “Argentina.” | 48 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. 520. A note on an Abyssinian catfish, Clarias depressus Myers. SIB, vol. 7, no. 2, p. 14. August 30. [See 45.] 521. Curatorial practices in zoological research collections. 2. System followed in filing specimens of Recent fishes in the Natural History Museum of Stanford University. (G. S. Myers and Margaret Hamilton Storey.) Natural History Museum of Stan- ford University, Circular no. 6, 44 pp. October. [Mimeographed; very small edi- tion. ] 1957 522. Exotic aquarium fishes—a work of general reference. By William Thornton Innes. 19th Edition, revised, enlarged, and edited by G. S. Myers. Innes Publishing Com- pany, Philadelphia; 541 pp., colored frontispiece, 90 colored figs., 366 black and white figs., 7 maps, 2 maps on linings. |All previous 18 editions of this book were also revised by G.S.M. before publication, but only in this edition was editorial re- sponsibility formally assumed. Previous editions are not listed in this bibliography. ] 1958 523. |Four world maps, showing extent of reef coral, current knowledge of species occur- rence of marine fishes, current knowledge of habits of marine food fishes, and the species composition of marine food-fish faunas.] Jn: Walford, L. A., Living re- sources of the sea—opportunities for research and expansion (New York, Ronald Press Co., xvi + 231 pp.), figs. 16-19, with discussions on pp. 246, 248, and 250. [Original maps and discussions by G.S.M. Other research for this book was also done by G.S.M.] 524. Trends in the evolution of teleostean fishes. SIB, vol. 7, no. 3, pp. 27-30. July 31. [Paper presented before Society for the Study of Evolution, August 1957.] 525. Nomenclator of certain terms used for higher categories of fishes. SIB, vol. 7, no. 3, pp. 31-40. July 31. 526. The priacanthid fish genus Pristigenys. SIB, vol. 7, no. 3, pp. 40-42. July 31. 1959 527. The endemic fish-fauna of Lake Lanao and the evolution of higher categories. Pro- ceedings XVth International Congress of Zoology (London, 1958), pp. 151-152. [Ab- stract of 537.] 528. A remarkable new genus of anostomin characid fishes from the upper Rio Xingu in central Brazil. (G. S. Myers and Antenor Leitao de Carvalho.) CO, 1959, no. 2, pp. 148-152. July 24. [Sartor respectus, n. gen., n. sp.| 529. A Caribbean chaetodont fish, Chaetodon eques Steindachner, now referred to Chaeto- don aya Jordan. CO, 1959, no. 2, p. 158. July 24. 530. [Review of] Simpson, G. G., and Roe, A., Behavior and evolution. CO, 1959, no. 3, pp. 270-271. October 9. 1960 531. Restriction of the croakers (Sciaenidae) and anchovies (Engraulidae) to continental waters. CO, 1960, no. 1, pp. 67-68. March 25. Voz. *532). XXXVIIT] MYERS: BIBLIOGRAPHY 49 Phylax telescopus. CO, 1960, no. 1, pp. 75-78. March 25. [A column of comment, criticism, and review dealing not only with ichthyology and herpetology but also broader subjects. | [Review of] Whitley, P. G., and Allen, J., The sea-horse and its relatives. CO, 1960, p. 78. March 25. [Review of] Poll, M., Les genres des poissons d’eau douce de |’Afrique. CO, 1960, no. 1, pp. 78-79. March 25. . Phylax telescopus, II. CO, 1960, no. 2, pp. 157-159. June 29. [Darwin; ideas versus data; the eel problem; Alipio de Miranda-Ribeiro, etc. In footnote, for Freund read Freud. | [Review of] McInerny, D., and Girard, G., All about tropical fish. CO, 1960, no. 2, p: 162. June 29. . The endemic fish fauna of Lake Lanao, and the evolution of higher taxonomic cate- gories. Evolution, vol. 14, no. 3, pp. 323-333. September. [Proof not seen by au- thor; p. 328, paragraph 2, for biotypes read biotopes; p: 330, delete entire sentence containing word “Triglopsis.”’ See also 527 for abstract; also 591 for reprintings.] . Fish evolution in Lake Nyasa. Evolution, vol. 14, no. 3, pp. 394-396. September. [Di- versity of freshwater fish faunas of the world, etc. ] Some reflections on phylogenetic and typological taxonomy. Systematic Zoology, vol. 9, no. 1, pp. 37-41. [March 1960; published September 1960. ] . Phylax telescopus, III. CO, 1960, no. 3, pp. 263-266. September 26. [On borrowing specimens; electric fishes; frog phylogeny, etc.] [Review of] Poll, M., Expedition océanographique Belge .. . l’Atlantic sud. Poissons. CO, 1960, no. 3, pp. 267-268. September 26. [Review of] Parr, A. E., Mostly about museums. CO, 1960, no. 3, p. 268. September 26. [Obituary of] Margaret Hamilton Storey (1900-1960). SIB, vol. 7, no. 4, p. 62a. Oc- tober 27. [See 557 for another obituary of M.HS.] . A new zeomorph fish of the family Oreosomatidae from the coast of California, with notes on the family. SIB, vol. 7, no. 4, pp. 89-98. October 27. . Two new fishes collected by General Thomas D. White in eastern Colombia. (G. S. Myers and Stanley Howard Weitzman.) SIB, vol. 7, no. 4, pp. 98-109. October 27. . The mormyrid genera Hippopotamyrus and Cyphomyrus. SIB, vol. 7, no. 4, pp. 123- 125. October 27. . The genera and ecological geography of the South American banjo catfishes, family Aspredinidae. SIB, vol. 7, no. 4, pp. 132-139. October 27. . A Brazilian pike-characid, Boulengerella lateristriga, rediscovered in the Rio Negro. (G. S. Myers and Stanley Howard Weitzman.) SIB, vol. 7, no. 4, pp. 201-205. Oc- tober 27. . The South American characid genera Exodon, Gnathoplax, and Roeboexodon, with notes on the ecology and taxonomy of characid fishes. SIB, vol. 7, no. 4, pp. 206— Zitien October 27. . Preface to any future classification of the cyprinid fishes of the genus Barbus. SIB, vol. 7, no. 4, pp. 212-215. October 27. . A forgotten account of a fresh-water belonid fish from northern India. SIB, vol. 7, no. 4, pp. 345-346. October 27. . The names of the South American catfish genera Conorhynchos and Diplomystes. SIB, vol. 7, no. 4, pp. 246-248. October 27. [See errata on p. 62b of same issue. Cono- rhynchos misspelled in three places on p. 247.] 50 BOO: *560. 561. *563. CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER. Phylax telescopus, IV. CO, 1960, no. 4, pp. 373-377. December 30. [Oceanography and the neglect of biological collecting; European centers; Denticeps, etc. ] 1961 . Phylax telescopus, V. CO, 1961, no. 1, pp. 117-120. March 17. [Conservationists neglect fishes; a new living perch from Europe; etc. ] The South American hylid frog names Sphaenorhynchus, Dryomelictes, and Spho- enohyla. (G.S. Myers and Alan Edward Leviton.) Herpetologica, vol. 17, pp. 61-62. April 15. . Phylax telescopus, VI. CO, 1961, no. 2, pp. 244-247. June 19. [Fishery biology and management; European museums, etc. Proof not seen by author; for Wandsee read Wansee. | [Obituary of] Margaret Hamilton Storey (1900-1960). CO, 1961, no. 2, pp. 261-263. June 19. [Not the same as 543.] Generic type species citation in taxonomic zoology. A guide for students. Natural History Museum of Stanford University, Circular no. 8, 7 pp. August. [Mimeo- graphed, very small edition. Also reproduced later by U.S. Bureau of Commercial Fisheries Ichthyological Laboratory, U.S. National Museum, Washington, D.C.] . The New Zealand lizard names Naultinus and Hoplodactylus. Herpetologica, vol. 17, no. 3, pp. 169-172. October 9. 1962 The American leptodactylid frog genera Eleutherodactylus, Hylodes (= Elosia), and Caudiverbera (= Calyptocephalus). CO, 1962, no. 1, pp. 195-202. April 11. [Suako 447b. | Statement regarding the argument of W. I. Follett and Daniel M. Cohen concerning the type species of the genus Bathylagus. Bulletin of Zoological Nomenclature, vol. 19, pp. 130-131. May 28. [Dealing principally with the type designations of Jordan and Evermann. | The Hong Kong newt described as a new species. (G. S. Myers and Alan Edward Leviton.) Occasional Papers, Division of Systematic Biology [formerly Natural His- tory Museum], Stanford University, no. 10, 4 pp. June 15. Generic classification of the high-altitude pelobatid toads of Asia (Scutiger, Aeluro- phryne, and Oreolalax). (G. S. Myers and Alan Edward Leviton.) CO, 1962, no. 2, pp. 287-291. July 20. 1963 Killifish identification. American Killifish Association, Killie Notes, Vol. 2, no. 2, pp. 7-10. March. Fresh-water fishes. Pacific Discovery, San Francisco, vol. 16, no. 4, pp. 36-39. July. [Freshwater fishes of the world, general facts, diversity, sizes, distribution, con- servation. | Comments on the proposed rejection of the type designations of Jordan and Evermann 1896-1900 and 1896. Bulletin of Zoological Nomenclature, vol. 20, part 4, p. 259. July 12. [See also 561.] . The fresh-water fauna of North America. Proceedings, XVI International Congress of Zoology, vol. 4, pp. 15-20. August. [Only abstracts were published in this volume. Vor. XXXVIIT] MYERS: BIBLIOGRAPHY 51 The paper as delivered at the Congress was much longer and argued in favor of continental drift. ] 568. Foreward. Jn: Jordan, D. S., The genera of fishes and a classification of fishes (re- print; Stanford University Press; xvi + 800 pp.), pp. vii—xvi. December 30. [Sets these two important works in perspective, gives considerable historical information on zoological nomenclature and fish classification, and warns against misuse of the Jordan papers. | 1964 *569. An electrophoretic survey of rattlesnake venoms. (Alan E. Leviton, G. S. Myers, and B. W. Grunbaum.) Jn: Leone, C. A. (editor), Taxonomic biochemistry and serology (New York; Ronald Press: x + 728 pp.), pp. 667-671. [On the basis of the venoms of 10 species, relations similar to those shown by morphology were found. ] 570. A brief sketch of the history of ichthyology in America to the year 1850. CO, 1964, no. 1, pp. 33-40. March 26. 571. Foreward. In: Albert W. Herre (1868-1962): a brief autobiography (Division of Systematic Biology, Stanford University, Circular no. 10, 20 pp.), pp. 1-2. March. 572. Bumblebee catfishes (Plotosus). TFH, vol. 13, no. 3, pp. 5-7, 75. November. [They are black-and-yellow, they buzz, they swarm, and they sting. ] 1965 573. Gambusia, the fish destroyer. TFH, vol. 13, no. 5, pp. 31-32, 53-54. January. [First indictment of Gambusia affinis, the mosquitofish, as a serious danger to other fishes, small and large, wherever introduced. See also 575.] 574. The body-wag, an innate behavioral characteristic of bony fishes. TFH, vol. 13, no. 9, pp. 21, 24-25. May. 575. Gambusia, the fish destroyer. Australian Zoologist, vol. 13, no. 2, p. 102. August. [Partial reprint of 573, with editorial note about Australian introductions. ] 576. Races of the Chinese paradise fish (Macropodus). TFH, vol. 14, no. 1, pp. 48-49. [Continuation of taxonomic revision begun in 137. Southernmost race in Indochina is M. opercularis concolor Ahl. Adds Polyacanthus yangye Dabry 1872 to synonymy of M. chinensis. | 1966 577. Foreword [to first issue]. Ichthyologica, the Aquarium Journal [continuation of AJ], vol. 37, no. 1, pp. 3-5. January. [G.S.M. resigned as editor after this issue appeared. | 578. How to become an ichthyologist. Part 1. TFH, vol. 14, no. 8, pp. 47, 50-51. April. [See 580, 581, 583. Advice for young prospective ichthyologists. ] 579. Phyletic studies of teleostean fishes, with a provisional classification of living forms. (P. Humphry Greenwood, Donn Eric Rosen, Stanley Howard Weitzman, and G. S. Myers.) Bulletin of the American Museum of Natural History, vol. 131, art. 4, pp. 339-446, pls. 21-23. April 18. 580. How to become an ichthyologist. Part 2. TFH, vol. 14, no. 9, pp. 47, 50-51. May. [See also 578, 581, 583.] 581. How to become an ichthyologist. Part 3. TFH, vol. 14, no. 10, pp. 28-30. June. [See also 578, 580, 583.] 52 587. 588. 589. 590. 593. 594. CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. . Two remarkable new trichomycterid catfishes from the Amazon basin in Brazil and Colombia. (G.S. Myers and Stanley Howard Weitzman.) Journal of Zoology [con- tinuation of the Proceedings of the Zoological Society of London], vol. 149, pp. 277-287. July. How to become an ichthyologist. Part 4. TFH, vol. 14, no. 2, pp. 29-31. August. [See also 578, 580, 581.] Megalomycteridae, a previously unrecognized family of deep-sea cetomimiform fishes based on two new genera from the North Atlantic. (G. S. Myers and Warren Curtis Freihofer.) SIB, vol. 8, no. 3, pp. 193-206. October 7. Derivation of the freshwater fish fauna of Central America. CO, 1966, no. 4, pp. 766-773. December 23. [Paper read before the American Society of Ichthyologists and Herpetologists in June 1964. | 1967 [Review of] Breder, C. M., and Rosen, D. E., Modes of reproduction in fishes. Natural History, N.Y., vol. 76, no. 2, pp. 66-67. February. [Suggests possibly primitive nature of nesting and parental care in bony fishes and notes its widespread oc- currence. | Note on the name of a Guatemalan cactus, Mammillaria voburnensis Scheer. Cactus and Succulent Journal of America, vol. 39, no. 4, p. 153. July. Zoogeographical evidence of the age of the South Atlantic Ocean. Studies in Tropical Oceanography, no. 5 (Miami, xx + 847 pp.), pp. 614-621. October 1. [Paper read at International Conference on Tropical Oceanography, Miami, two years previously. | Named main divisions of teleostean fishes. (P. Humphry Greenwood, G. S. Myers, Donn Eric Rosen, and Stanley Howard Weitzman.) PBSW, vol. 80, pp. 227-228. December 1. Note on the dentition of Creagrudite mavxillaris, a characid fish from the upper Orinoco-upper Rio Negro system. (G. S. Myers and Tyson Royal Roberts.) SIB, vol. 8, no. 4, pp. 248-249. December 5. 1969 . The endemic fish fauna of Lake Lanao, and the evolution of higher taxonomic cate- gories. In: Ehrlich, P. R., Holm, R. W., and Raven, P. H. (editors), Papers on evolution (Boston, Little Brown and Co., xii +564 pp.), pp. 247-261. [Reprint of 537, with corrections. | [Review of] Harden Jones, F. R., Fish migration. CO, 1969, no. 2, pp. 409-411. June 3: Peace! It’s wonderful! [Allegory.] Palo Alto Times, Thursday, November 27, 1969, p. 26. [Based on a true story. | The endemic fish fauna of Lake Lanao, and the evolution of higher taxonomic cate- gories. In: Laetsch, Watson M., the biological perspective (Boston; Little, Brown and Co., xii + 574 pp.), pp. 351-365. [There were two reprints of this paper during 1969, this being the second. See also 591. Both are reprints of 537.] aa eis Reon! 7: as : = - met 7 _ PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Festschrift for George Sprague Myers Vol. XXXVIII, No. 3, pp. 53-62; 3 figs.; 1 table. December 31, 1970 A NEW SPECIES OF THE DORADID CATFISH GENUS LEPTODORAS, WITH COMMENTS ON RELATED FORMS By James E. Bohlke Chaplin Chair of Ichthyology Academy of Natural Sciences of Philadelphia Asstract: Leptodoras myersi is described from a trawl haul made in Rio Amazonas near Iquitos, Peru. Leptodoras juruensis, previously known only from the holotype taken in Rio Jurua, Brasil, is recorded from the same haul. It was originally intended that this be a revision of the genus Leptodoras but at the last moment it became apparent that the species L. linnelli is a composite that will require further study. Also, in Eigenmann’s (1925) review of the Doradidae, the related genera Opsodoras, Hassar, and Leptodoras are perhaps the least well defined and thus require more attention. At present, I describe as new a well-marked species and comment on its relatives. The newly recorded specimens of Leptodoras myersi and L. juruensis were collected on the 1955 Catherwood Foundation Peruvian-Amazon Expedition by Charles C. G. Chaplin and Ruth Patrick of the Academy’s staff. They were taken with an otter trawl from the Amazonas (Maranon) between Isla Iquitos and Isla Lapuna. Only one downstream haul was made, because of the swiftness [53] 54 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. of the current and the many snags in the bottom, but this caught a fascinating group of mostly new and rare catfishes plus one specimen of Rhytiodus microlepis Kner. This suggests that more bottom sampling should be attempted in the large South American rivers. For the loan of important specimens, I thank P. H. Greenwood of the British Museum (Natural History) and W. I. Follett of the California Academy of Sciences. METHODS The standard length measurement was made with some difficulty but, by flexing the caudal fin and using reflected light, I believe fairly good accuracy was achieved. The head length includes the fleshy opercular flap. The eye and snout measurements involve the eyeball proper. The predorsal measurement, length of dorsal spine, and depth at dorsal-fin origin all have as one terminus the anterior groove of the small bony element at the base of the spine. The anterior end of the adipose dorsal-fin base is somewhat difficult to define, but measure- ments involving it are more easily duplicated in this species than in L. acipen- serinus for example. The greatest width of the peduncle is at the posterior end of the anal-fin base and includes the lateral spines that jut out from the body. The length of the pectoral spine is measured basally from the notch, where a needle-point fits in snugly, and not from the extreme base of the spine. Ventral- fin length is the greatest length of the fin, not of an individual ray. The greatest head width is of the bony portion and not of the rather indifferent fleshy portion posteriorly. The greatest scute depth is a vertical measurement of the area covered by scutes, rather than the diagonal measurement of an individual scute. The anal-ray counts are separated into anterior unbranched, small roman numerals, and posterior branched, arabic numerals; the last anal ray some- times is simple, sometimes consists of two rays united at their bases, but in either case is counted as one. The lateral scutes are all of those in the main lateral series, including the ones on the caudal-fin base but not the small ones in the humeral region, the tympanum. Leptodoras myersi Bohlke, new species. Dracnosis. This is an elongate, long-snouted species like L. acipenserinus, L. linnelli, and L. juruensis. Its dark color markings, particularly the broad nuchal band, are distinctive. Lateral scutes few, 36 or 37, each scute bearing few points. Total anal-fin rays few, 13 or 14 (except see discussion of L. linnelli below). Lacking extremely elongate first dorsal spine of L. juruensis. While anterior base of adipose dorsal fin is not sharply defined, it does not extend far forward as a fleshy ridge. Dorsal and pectoral spines with small hooks or spines along their anterior and posterior margins, these weakest on the dorsal spine and strongest on the posterior margins of the pectoral spines. Nuchal foramen VoL. XXXVIII] BOHLKE: NEW SPECIES OF LEPTODORAS On Jt Ficure 1. Leptodoras myersi: Holotype, 74.6 mm. standard length, ANSP 112318. present. Head covered by small, elongate, pale fleshy ridges, arranged in a pattern (see fig. 1). DescripTION. The body shape and dark color markings are shown on the photographs (fig. 1). Selected measurements and counts made on a series of 10 specimens appear in table 1. Dorsal rays I, 6. Total anal rays 13 or 14, nearly always 14. Pectoral rays I, 9 or I, 10, usually I, 10. Ventral rays i, 6/1, 6. Principal caudal rays i, 15, i, the ventral unbranched ray counted not extending back to the tip of the lobe as does the dorsal one. Number of lateral scutes 36 or 37 in equal numbers. Anterior dorsal serrae 7 to 12, posterior dorsal serrae 6 to 11, anterior pectoral serrae 19 to 23, and posterior pectoral serrae 12 to 15; the numbers of serrae apparently are not related to the length of the fish, at least within the limited size range examined. No teeth present. Nostrils both with raised margins, that of the rear nostril lowest posteroventrally. Anterior nostril nearer eye than tip of snout. Distance between the two nostrils on one side equal to that between posterior nostril and ‘ajduis Avi sey z ‘saseq JI9Y4} }@ pa}luN s}UIWa[a OM} JO SUISIsUOD API 4SPT _ [Proc. 47TH Ser. CALIFORNIA ACADEMY OF SCIENCES 9¢ 9¢ Le Le Le 9¢ Ms Le 9¢ 9¢ Saynos [P19] OV'T/OT'T OT‘I/O'I OLI/OL'L OL‘I/OlI 6'1/6% OfT/OT'I GI/Ol'I OT/OT'L OTI/O1'I Of T/T SABI UTJ-[P10}99q 04 204 Sta 204 04 20-4 eS tA 204 26° 34 SABI ULJ-[PUY +0 9'+0 t'+0 10 9°+0 s'+0 z'+0 z'+0 ¢'+0 9°40 yidap pueq aynos 4so}eoiry Sel 6°21 9°71 S21 Ne ST Ve eel 7 ¢T Z'eT UISLIO UlJ-[esIop ye Yy}doq 819 189 0°69 OL 9°69 £69 Z'OL 6°69 9°89 3°69 aouRysIp Ulf-[BURATg a St ZS a S'Sh SSb 19+ Ssh vSP eSb 19+ aouRsIp UlJ-[PIJUIAII T6I aA O'8T 7ST 781 O'ST PLT S81 ¢ST 161 YIPIM pesy 4so}eaI5) 9 LT esi T6r OST 6 LT Lt S81 61 S61 102 yi sue] Uly-[e1]UIA SIZ 12 6°72 912 Cae PZ 612 Ve? SIZ 7772 yysue] surds [e10}904 0'02 7 LT 0'61 0'6T Z'6r Z'6T S91 8'6T S61 O'8T yysua] ouids [esi0qd 9°€0 9°€0 9°€0 9°£0 9°¢0 6°£0 6'£0 s'€0 L°€0 6'£0 yideap gpunped jseoT LL0 180 9°L0 Z’80 S40 £80 6°40 9°40 vL0 180 Upra spunped 4sojee15) SEZ 9°72 [Gee Pe SZ vVrZ S22 9°72 Gee 9°72 UISIIO asodipe 0} uly [esIOd 8'0r 6'6¢ v6 €0r 60 L1v OTP SOF 6'Or SIP aueysIP [PSIOped z'+0 8°€0 6'£0 z'+0 0'+0 Z'+0 (a0) z'+0 z'+0 T'+0 [eyqrorajur Auog (ona) 6°20 8°20 3°20 6°20 3°20 T¢0 6°20 0'¢0 0'¢0 WSIoy IAG Z'+0 £70 £0 0'+0 Z'+0 T'+0 b'+0 Z'+0 9° +0 9°+0 YIpIM sAq 8°61 SST SSI © 6T ¢6T €°02 9°61 b'61 6°81 8°61 yysua] ynoug 6'be ree eee oe Ore ose Lye wre ore Z'S¢ yisue] prox LL 9°94 Srl +9 bl +69 8°89 $89 $89 eL9 1°09 (wut) YyySUa_ plrpueys 56 ‘sada wing aur pun (x) agkjojoy ay} {0 S]UNOD JAaras PUD YISUua] PADpUDIS {0 Juarsag UL SJUamadNSDayY :1Stahu se1opoydaT “T ATAVL, VoL. XXXVIIT] BOHLKE: NEW SPECIES OF LEPTODORAS 57 C Ficure 2. Leptodoras myersi: Paratype, 63.3 mm. standard length, ANSP 112320. A, outline of sixth lateral scute on left side of fish (central ridge not indicated), 3.1 mm. in longest dimension; B, dorsal spine with break-away tip, main spine 11.5 mm.; C, right pectoral spine, 14.2 mm. eye. Eyes distinctly horizontally elongate. Horizontal width of eye equal to least width of bony interorbit, or nearly so. The middorsal fontanel is long and narrow, extending from just behind a line drawn between the rear margins of the posterior nostrils to one drawn between the rear margins of the eyes. A middorsal groove extends in front of the fontanel on the snout and posteriorly from the fontanel to the dorsal-fin origin. Gill membranes connected to isthmus ventrally, the ventral ends of the two gill open- ings separated by a space of slightly more than the least interorbital width. Gill rakers low fleshy bumps. Two maxillary and 4 mental barbels present, all inter- connected to form a hood of considerable size when spread. Each maxillary barbel is divided, its outer portion with barblets along its outer margin; these barblets are in two series, the upper ones short and numerous, the lower ones long and fewer in number. The outer portion of the maxillary barbel usually fails to reach the ventral end of the gill opening but may just attain that level. The inner portion of the maxillary barbel forms the lateral margin of an ex- tensive fleshy lobe that connects the maxillary with the outer mental barbel; this lobe has short barblets around its margin. The mental barbels are studded with short barblets all around. The maxillary is ossified to the extent of about one-third the length of the outer division of the barbel. Branchiostegal rays 7. Upper end of gill opening just behind lower end of upper third of dark opercular spot shown on figure 1. On the dorsal spine, the anterior serrae are longer than the posterior; the anterior ones are crowded and overlapping, especially basally, are directed toward the spine tip, and are distributed along less than the basal half of the spine. The posterior serrae are short, widely spaced, their tips directed perpendicular to the main axis of the spine or slightly inclined either basally or distally; when their 58 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. tips are angled, the outer serrae are tilted basally and the lower serrae are tilted distally. On one individual the uppermost serration is directed distally. The posterior serrae extend along the distal one-half to three-quarters of the spine. There is a distinct ossified break-away tip on the spine when intact, this tip fitting into a median groove in the spine proper; the dorsal spine outlined in figure 2 shows this tip nearly completely disengaged. On the pectoral spines the anterior serrae are shorter than the posterior, are directed toward the spine tips, and the distalmost one counted forms the tip of each spine (not the case in several related species in which the spine tips are rounded or blunt and without projections). The posterior serrations are strong hooks, their tips directed toward the bases of the spines. There are pointed fleshy extensions beyond the ossified spine tips, indicating that there probably are no ossified break-away segments as on the dorsal spine of this species and the pectoral spines of L. jurwensis (the latter will be discussed and figured in a sub- sequent paper when the status of LZ. linnelli also will be treated). Pectoral spines, when depressed, extending back well beyond the ventral-fin bases. A single pectoral pore present on each side. Tips of ventral fins rounded, not extending back to the anal-fin origin. Anal and genital papillae placed between the ventral fins, at or slightly before mid-fin. Adipose dorsal fin well developed, short-based, its origin above points varying between the base of the sixth to the interspace between the seventh and eighth anal rays. Caudal fin distinctly forked, the lower lobe the longer. On the single stained individual there are 18 procurrent caudal rays above and 17 below the principal rays. In the humeral region there is a sharp spine on the posterior margin of the supraclavicle followed, across the center of the tympanum, by 3 narrow, elongate, mostly embedded ossifications. The first of the three is longest and has a rise in the middle that appears as a low hump externally; the second is completely embedded or with only a minute portion exposed; the third is shortest and bears a sharp projecting spine at its posterior end. The pointed coracoid processes ex- tend posteriorly beyond the base of the last pectoral ray, but not as far back as do the humeral processes; the last-mentioned are shallowly convex dorsally and terminate in a narrowly rounded tip. There is a horizontally elongate nuchal foramen on each side. The lateral series of scutes begins behind the tip of the humeral process and continues out onto the caudal fin basally. The scute outlined in figure 2 is much like those just before and after it (the outline does not indicate the median longitudinal ridge and the sharp outward angulation of the median spine itself) ; proceeding posteriorly the scutes become increasingly wider, less deep, less angular, and more overlapping. The terminal ossification in the series is a nar- row, elongate, tubular element that lacks a spine and usually is overlooked on UL No} VoL. XXXVIIT] BOHLKE: NEW SPECIES OF LEPTODORAS unstained specimens. This was not included in the counts of scutes recorded in this paper; apparently it also was not counted by Eigenmann (1925: 358), for I obtained the same count as he did on the same specimen of L. acipenserinus, omitting this element. The pattern of coloration in alcohol is shown in figure 1; however, the broad nuchal band is more definitely continuous across the dorsum than is suggested on the dorsal view, and the dark opercular margin frequently is more intense and continues farther ventrally. A faint dusky stripe is present on the upper half of the lower caudal-fin lobe, and sometimes there is an even fainter one on the lower half of the upper lobe. The dorsum is dusky between the dorsal fin and caudal-fin base. The top of the head is dusky before the nostrils and in a roughly circular middorsal patch immediately behind the eyes. The basal half or more of the pectoral fin exclusive of the spine (except sometimes the membrane en- casing the posterior serrae) usually is distinctly dark, sometimes only dusky and the extent of the dusky area variously more reduced than shown in the figure. RELATIONSHIPS. The elongate, long-snouted species L. acipenserinus, L. linnelli (both types if they represent more than one species) and L. juruensis are most closely related to L. myersi. Leptodoras juruensis is the most distinctive and most spectacular looking member of this group, with its extremely elongate anterior dorsal-fin element and distinctive black color markings (see fig. 3); it also has more lateral scutes than the others: 44 to 46 in L. juruensis, 42 in L. acipenserinus, 38 or 39 in the L. linnelli complex, and 36 or 37 in L. myersi. Leptodoras juruensis is distinctive in certain proportions, but these will be treated later. Leptodoras myersi has a lower anal-fin ray count than other species of Leptodoras excepting, evidently, the Guianan (typical) population of the L. linnelli complex. Eigenmann (1912: 192), in his original description of ZL. linnelli, gave an anal-ray count of 12 to 14; the specimen (ANSP 39734) from Rio Rupununi recorded by Fowler (1914: 264) has 12 rays. Leptodoras myersi has 13 or 14 rays, L. acipenserinus has 17 rays (16 recorded for the holotype by Giinther 1868: 230), L. juruensis has 16 or 17 rays, and the Peruvian specimens nearest L. linnelli have 15 to 17 rays. In the size range represented by the Peruvian material, the shapes of the lateral scutes are most similar in L. myersi and L. juruensis, those of L. acipen- serinus and nominal L. linnelli having more teeth above and below the median spine. While differing in numerous relative proportions (to be discussed further when the L. linnelli question is resolved), L. myersi has a distinctly smaller eye than L. linnelli (four or five percent of standard length versus seven or eight percent). Leptodoras linnelli has no color pattern except for a faint dusky stripe that extends out each caudal lobe, while L. myersi has the distinctive pattern described above. Leptodoras acipenserinus is described as devoid of a color 60 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser. Ficure 3. Leptodoras juruensis: 125.8 mm. standard length, ANSP 112321. pattern (Gunther 1868: 230) and IUM 15878 (Eigenmann 1925: 358) shows no trace of one. This color difference, coupled with the differences listed above between the elongate L. acipenserinus and L. myersi, plus numerous proportional differences that will be outlined in a subsequent paper, indicate how different are the two species. Leptodoras myersi has elongate, pale, raised ridges on the head, L. acipenserinus has pale low papillae, while L. linnelli has nothing of the sort. Name. For my professor and good friend, George S. Myers. MATERIAL EXAMINED. Holotype: ANSP 112318 (74.6 mm. standard length, photographed), Peru: vicinity of Iquitos, Rio Amazonas (Maranon) between Isla Iquitos and Isla Lapuna, near Isla Lapuna shore; to 12 ft. (3.66 meters) ; trawl; 9 October 1955; C.C.G. Chaplin, R. Patrick. Vor. XXXVIII] BOHLKE: NEW SPECIES OF LEPTODORAS 61 Paratypes: ANSP 112319 (9; 54.9-77.6), ANSP 112320 (1; 63.3, cleared and stained) and USNM 203816 (2; 68.5—74.8), taken with the holotype. Leptodoras juruensis Boulenger. Leptodoras juruensis BOULENGER 1898, p. 478 (Type locality: Rio Jurua, Brasil). E1GEn- MANN 1925, pp. 357, 358 (diagnosis based on type). Previously known only from the holotype from Rio Jurua, this species now is recorded from the same trawl haul that collected the type material of L. myerst. A fine Peruvian example is illustrated in figure 3. Peruvian specimens have been compared with the much larger holotype of the species and the results of this comparison will be forthcoming. MATERIAL EXAMINED. British Museum (Natural History) 1898—10—-11—25 (223 mm. standard length, holotype), Brasil: Rio Jurua; Goeldi. ANSP 112321 (1; 125.8, photographed), ANSP 112322 (6; 71.4-96.7) and USNM 203817 (1; 92.6), taken with the holotype of L. myersi. NOTE While this manuscript was in proof, a paper on Venezuelan doradids was received from Fernandez Yépez (1968, Boletin del Instituto Oceanographico, Universidad de Oriente, Cumana, Venezuela, vol. 7, no. 1, pp. 7-72). In it, he includes the species “leporhinus,” “Tinnelli,” and “notospilus” in the genus Opsodoras, whereas “Jinnelli” previously was in Leptodoras and “notospilus” was in Hassar. Subsequent correspondence with that author revealed that his rationale for making these and other nomenclatural changes is in a manu- script still in press. The new species, “myersz,’ is closest to “linnelli” and “acipenserinus,” which were placed by Eigenmann (1925: 357) in Leptodoras, so the name combination Leptodoras myersi is here published, with the realization that the species may later be transferred to a different genus. REFERENCES BouLenceEr, G. A. 1898. Descriptions of two new siluroid fishes from Brazil. Annals and Magazine of Natural History, ser. 7, vol. 2, pp. 477-478. EIGENMANN, C. H. 1912. The freshwater fishes of British Guiana, including a study of the ecological group- ing of species and the relation of the fauna of the plateau to that of the low- lands. Memoirs of the Carnegie Museum, vol. 5, xxii + 578 pp., 103 pls. 1925. A review of the Doradidae, a family of South American Nematognathi, or cat- fishes. Transactions of the American Philosophical Society, new series, vol. 22, no. 5, pp. 280-365, 27 pls. Fow ter, H. W. 1914. Fishes from the Rupununi River, British Guiana. Proceedings of the Academy of Natural Sciences of Philadelphia, vol. 66, pp. 229-284. GUNTHER, A. 1868. Descriptions of freshwater fishes from Surinam and Brazil. Proceedings of the Zoological Society of London, 1868, pp. 229-247, pls. 20-22. ——- be PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Festschrift for George Sprague Myers Vol. XXXVIII, No. 4, pp. 63-98; 8 figs. December 31, 1970 SYSTEMATICS OF THE GENUS HEMITRIAKIS (SELACHIT: CARCHARHINIDAE), AND RELATED GENERA By L. J. V. Compagno Division of Systematic Biology, Stanford University INTRODUCTION Herre (1923) described Hemitriakis leucoperiptera, a new genus and species of shark from the Philippine Islands. Hemitriakis was thought to differ from Triakis Miller and Henle in its dentition, snout, nasal valves, body, and caudal fin. However, Fowler (1941), Bigelow and Schroeder (1948), Garrick (1954), and Kato (1968) considered Hemitriakis a junior synonym of Triakis. Present data shows that Hemitriakis is a well defined genus with two species: H. leucoperiptera Herre, 1923; and H. japanica (Miller and Henle, 1841). This account is a review of the systematics of Hemitriakis and related genera in the family Carcharhinidae. ACKNOWLEDGMENTS Reeve M. Bailey (University of Michigan Museum of Zoology), Myvanwy M. Dick (Museum of Comparative Zoology, Harvard University), W. I. Fol- jett, William D. Eschmeyer, Lillian J. Dempster (Department of Ichthyology, California Academy of Sciences), and Stanley H. Weitzman (Division of Fishes, U. S. National Museum) loaned or gave specimens to me and provided working facilities at their institutions. C. G. Alexander (Department of Biology, San [63] 64 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. a OME Was or op) Ficure 1. A, dorsal view, and D, ventral view, of head of Hemitriakis japanica (SU-12677). B, dorsal view of head of Hypogaleus hyugaensis, adopted from Miyosi (1939). C, dorsal view, and F, ventral view, of head of Galeorhinus zyopterus (LJVC-0238 ; 847 mm. female.). E, ventral view of head of Hypogaleus zanzibariensis, adopted from Smith (1957b). Abbreviations: HHR, horizontal head rim; SR, subocular ridge. Vor. XXXVIII] COMPAGNO: SYSTEMATICS OF GENUS HEMITRIAKIS 65 Francisco State College), Robert P. Dempster (Steinhart Aquarium, California Academy of Sciences), and Louis Garibaldi (American Broadcasting Company Marine World, Redwood City, California) supplied many fresh and frozen car- charhinids for anatomical preparations. The late J. L. B. Smith (Department of Ichthyology, Rhodes University, Grahamstown, South Africa) sent specimens of Eridacnis sinuans and Scylliogaleus quecketti; Leslie W. Knapp (Smithsonian Oceanographic Sorting Center, Washington, D. C.), C. Richard Robins, and Phillip C. Heemstra (Institute of Marine Sciences, University of Miami) loaned other carcharhinids. In addition to providing numerous specimens and research facilities, Shelton P. Applegate (Division of Vertebrate Palaentology, Los An- geles County Museum of Natural History), Susumu Kato (Bureau of Commer- cial Fisheries Fishery-Oceanography Center, La Jolla, California), and Stewart Springer (Bureau of Commercial Fisheries Systematics Laboratory, U. S. Na- tional Museum) have discussed various aspects of carcharhinid taxonomy cov- ered in this paper with me. J. A. F. Garrick (Department of Zoology, Victoria University of Wellington, New Zealand) sent comments on several systematic problems concerning carcharhinid genera and species. George S. Myers critically reviewed the first draft of the manuscript, and Warren C. Freihofer (Division of Systematic Biology, Stanford University) offered useful suggestions. I am most grateful for the help offered by all of these people, without which this account could not have been written. STUDY MATERIAL Specimens mentioned in the text and figures are from the collections of the George Vanderbilt Foundation at the California Academy of Sciences (GVF); Division of Systematic Biology, Stanford University (SU); University of Michi- gan Museum of Zoology (UMMZ); U.S. National Museum (USNM); and of the writer (LJVC). Hemitriakis specimens examined are listed below, with number of specimens and total lengths in parentheses. Hemitriakis japanica: SU—12677, Nagasaki, Japan (1; 682 mm.); UMMZ-— 179060, Auraji (Osaki Market, Osaki), Japan (1; 650 mm.); UMMZ-179061, Ainoshima (Fukuoka Market, Fukuoka), Japan (1; 560 mm.); UMMZ-179062, Ezumi (Ezumi Market), Japan (1; 505 mm.); USNM-—191193, Taipeihsien, Taiwan (3; 651-685 mm.). Hemitriakis leucoperiptera: SU-27118, Dumaguete, Oriental Negros, Philip- pine Islands (2; 169-170 mm.). Hemitriakis species: SU-40097, Dumaguete, Oriental Negros, Philippine Islands (4; 161-180 mm.). Comparative material including most carcharhinid genera and species was examined. As the number of specimens in this sample is enormous, they are not listed here but will be given in a forthcoming revision of carcharhinid genera. 66 CALIFORNIA ACADEMY OF SCIENCES LOcimn. [Proc. 4TH SER. Vot. XXXVIII] COMPAGNO: SYSTEMATICS OF GENUS HEMITRIAKIS 67 Instead, the genera and species examined are listed. Carcharhinus species no- menclature is modified from Garrick (1967); that for Scoliodon, Rhizoprion- odon, and Loxodon is from V. Springer (1964). A prionodon isodon, Carcharhinus acronotus, C. albimarginatus, C. altimus, C. amblyrhynchus, C. borneensis, C. cauta, C. falciformis, C. galapagensis, C. leucas, C. limbatus, C. longimanus, C. maculipinnis, C. melanopterus, C. menisorrah, C. milberti, C. obscurus, C. pleurotaenia, C. porosus, C. remotus, C. sorrah, C. springeri, C. tyutjot, C. velox, Eridacnis barbouri, E. radcliffei, E. sinuans, Galeo- cerdo cuvier, Galeorhinus australis, G. chilensis, G. galeus, “G.” omanensis, G. zyopterus, Hemigaleus balfourt, H. macrostoma, H. microstoma, H. pectoralis, H. tengi, Hemipristis elongatus, Hypoprion hemiodon, H. macloti, H. signata, Isogomphodon oxyrhynchus, Lamiopsis temmincki, Leptocharias smithu, Loxodon macrorhinus, Mustelus antarcticus, M. asterias, M. californicus, M. canis, M. dorsalis, M. fasciatus, M. griseus, M. henlei, M. higmani, M. kanekonis, M. lenticulatus, M. lunulatus, M. manazo, M. mento, M. mustelus, M. norrisi, M. schmitti, Negaprion acutidens, N. brevirostris, N. forsteri, N. fronto, Prionace glauca, Proscyllium haberert, Rhizoprionodon acutus, R. lalandei, R. longurio, R. oligolinx, R. porosus, R. terraenovae, Scoliodon laticaudus, Scylliogaleus quecketti, Triaenodon obesus, Triakis acutipinna, “T.” fehlmanni, T. maculata, T. scyllia, T. semifasciata. TERMINOLOGY For descriptive purposes the morphological terminology of the head, eyes, dentition, vertebral column, and fins of carcharhinid sharks is discussed and elaborated here. Heap MorpuHo.ocy. The horizontal head rim (fig. 1) is the head margin in dorsal or ventral view. The subocular ridge is a ventrolateral expansion of the horizontal head rim beneath the eye. In Hemitriakis, Triakis, Mustelus, Fur- galeus, and other carcharhinid genera with well developed subocular ridges, the eyes appear medial to the horizontal head rim in dorsal view. A subocular ridge obscures the eyes in ventral view. NIcTITATING LOWER EYELID (fig. 2). Form and terminology of the carchar- hinoid ocular structures variously termed nictitating membranes, nictitating Ficure 2. Lateral views of carcharhinid eyes, showing nictitating lower eyelid types. A. Proscyllium habereri (UMMZ-179064; 535 mm. female), with rudimentary NLE. B. Mustelus canis (USNM-197676; 337 mm. female), with external NLE. C. Galeorhinus australis (USNM-176995; 385 mm. male), with transitional NLE. D. Leptocharias smithii (USNM- 202677; 570 mm. male), with internal NLE. Abbreviations: NLE, nictitating lower eyelid; SLE, secondary lower eyelid; SOP, subocular pouch; SP, spiracle; UE, upper eyelid. Dashed line is bottom of subocular pouch; dotted line in Leptocharias is edge of NLE inside palpebral aperture. 68 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. PUG RT BL TN? 3BG Ficure 3. Anteroposterior teeth of Hemitriakis japanica (SU-12677). A. Inner face of right lower tooth. B. Outer face of left lower tooth. Abbreviations: BL, basal ledge; CR, crown; FT, crown foot; PC, primary cusp; BG, basal groove; PLAS and PMAS, Vor. XXXVIII] COMPAGNO: SYSTEMATICS OF GENUS HEMITRIAKIS 69 folds, subocular folds, movable lower eyelids, and nictitans were reviewed by Gilbert (1963) and by Gilbert and Oren (1964). They used the term “nictitans”’ to cover all variations of the mobile eyelid of scyliorhinids and carcharhinids, but this term is not adopted here as the selachian structure is morphologically and developmentally unlike the true nictitans or nictitating membrane of tetrapods and in many cases is merely a little-modified movable lower eyelid. Instead, the term nictitating lower eyelid (NLE) is introduced to avoid some of the connota- tions of nictitans and to recognize the probable derivation of the structure from the original lower eyelid of precarcharhinoid sharks. The exterior fold formed by the groove below the NLE is termed the second- ary lower eyelid (SLE). The groove itself is the subocular pouch. Four nictitating lower eyelid types can be distinguished among carcharhinids if subdivisions are made in the morphological gradient seen in this structure. The rudimentary type is the least specialized. In it the NLE forms the ventral edge of the palpebral aperture and connects anteriorly and posteriorly with the upper eyelid. The SLE is a weak ridge below the NLE and does not connect with either the upper eyelid or the NLE. The upper edge of the SLE is not defined and the subocular pouch is a very shallow, external groove. The external type differs from the rudimentary in that the SLE is a strong flap with a well defined edge. The subocular pouch, although relatively shallow, is strongly differentiated. The internal type is the most advanced, with the NLE ends entirely internal to the palpebral aperture and not connected to the upper eyelid. The SLE replaces the NLE in contacting the anterior and posterior ends of the upper eyelid and forms the ventral edge of the palpebral aperture. The subocular pouch is entirely within the palpebral aperture and varies from moderately shallow (Leptocharias) to very deep (Carcharhinus). The transitional type covers intermediates between internal and external types. These often have the SLE attached by one of its ends (posterior or anterior) to the upper eyelid, while the NLE has its opposite end also attached to the upper eyelid. DENTITION (figs. 3-4). Tooth topography of selachians was discussed briefly by Applegate (1967). He divides the tooth into two external regions, the crown and the root. The crown is the enamel-covered region of the tooth distal to its attachment with the jaw. The proximal root lacks the enamel covering and has its component osteodentine exposed to the surface. The region of the crown proximal to the root is termed the foot. As used by Bigelow and Schroeder (1948), the term base includes both the root and the crown foot. The crown and root are both compressed in a plane with its horizontal sides < postlateral and premedial parts of attachment surface; PLC, postlateral cusplets; PLL, postlateral lobe of root; PME, premedial edge of crown; PML, premedial lobe of root; RT, root; TG, transverse groove; TN, transverse notch; TR, transverse ridges. [Proc. 4TH SER. CALIFORNIA ACADEMY OF SCIENCES 70 Vor. XXXVIII] COMPAGNO: SYSTEMATICS OF GENUS HEMITRIAKIS 71 parallel to the jaw axis and its vertical sides perpendicular to it. Teeth in upright functional positions at the edge of the jaw have outer and inner faces on their planes of compression. The orientations of these faces are reversed when the teeth are in replacement position but as a convention the functional orienta- tion is used here for any tooth. The root has its inner face partially formed into a flattened attachment sur- face that seats in the dental membrane against the jaw surface. The root has a vertical transverse groove that superficially divides the attachment surface into two lobes and may extend over the extreme rim of the root to form a transverse notch. The outer face of the root may have a strong basal groove extending horizontally across it that is overlapped by a strong basal ledge of the crown foot. A series of vertical transverse ridges may be present on the basal ledge and often extend distally on the outer face of the crown. In many species of Mustelus the crown inner face has a rounded protuberance or peg. The peg of one tooth extends into the basal groove of the next tooth in succession in the same row, an arrangement that may serve to interlock the teeth in the pavement dentitions of these forms. The distal part of the crown, as opposed to the foot, may have its margin in the plane of its compression formed into a sharp cutting edge, with or without serrations. Pointed projections from the crown edge are termed cusps or cusplets according to their size relative to each other. In carcharhinids a median primary cusp is commonly present and is usually larger than other projections of the crown edge (when such are present). The primary cusp may have its axis per- pendicular or oblique to the tooth base. Its proximal origin may occupy all or only part of the foot. When a primary cusp origin is restricted, the adjacent crown edges may be formed into other cusps or cusplets, sharp-edged blades, or rounded shoulders. The planes of compression in the teeth of carcharhinid sharks have their hori- zontal sides parallel to the jaw axis, but this axis changes from nearly perpendic- ular to the body axis at the symphysis to nearly parallel with the body axis at either end of the dental arcade. The horizontal sides of the planes of compression for tooth roots and crowns are therefore oriented in an anteromedial-to-postero- lateral direction relative to the anterior-to-posterior horizontal body axis along Ficure 4. Outer views of carcharhinid teeth. All teeth except C from left half of dental band. A, upper 8th tooth, and D, lower 10th tooth, of Galeorhinus zyopterus (LJVC-0114; 1670 mm. male). B, upper 10th tooth, and E, lower 9th tooth, of Hypogaleus zanzibariensis (1220 mm. male; modified from Smith, 1957b). C. Upper tooth of third row from end of dental band, Proscyllium haberert (GVF-Hong Kong-88; 523 mm. female). F. Same of Triakis semifasciata (LJVC-0137; 1097 mm. male). Abbreviations as in Figure 3, except for: PMC, premedial cusplets. 72 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Bt Le we be A Bx FicureE 5. Transverse views of carcharhinid vertebral calcification patterns, with calcified areas indicated in black. A. Eridacnis barbouri (“Silver Bay” 3514; 258 mm. female). B. Proscyllium habereri (UMMZ-179065; 565 mm. male). C. Mustelus henlei (LJVC-0020; 630 mm. female). D. Hemitriakis japanica (SU-12667). Abbreviations: DCL, diagonal calcified lamellae; I, intermedialia; NA, neural arch; NC, notochordal canal. most of the jaw, with the angle between the sides and the body axis decreasing from symphysis to rictus. It is possible with these orientations to distinguish anteromedial and posterolateral edges on the crowns and anteromedial and pos- terolateral lobes on the roots of most teeth. Exceptions occur at the symphysis, where teeth may have medial-to-lateral orientation, and at the ends of the dental arcade, where teeth can have anterior-to-posterior orientation. As a convention the anteromedial-to-posterolateral relations are used for all teeth. For brevity, structures having an anteromedial orientation on the tooth are termed premedial, whereas posterolateral structures are postlateral. Thus, carcharhinid teeth can have premedial and postlateral cusp edges, cusplets, serrations, blades, etc. The terms row and series were used almost interchangeably by Bigelow and Vor. XXXVIII] COMPAGNO: SYSTEMATICS OF GENUS HEMITRIAKIS 73 Schroeder (1948), but Applegate’s (1965) usage is followed here. A row is a single replicating file of teeth approximately transverse to the jaw axis that in- cludes both functional teeth and their replacements in various stages of develop- ment. The row represents an entire family of teeth derived from one germinal area on the jaw. The term “series” is used for a line of teeth along the jaws which is parallel to the jaw axis and includes teeth from all rows present. In some carcharhinids, especially those with pavement dentitions and very numerous teeth, the concept of series may be meaningless as all teeth are closely adpressed in quincunx formation and do not form distinct transverse lines. As indicated by Applegate (1965), there are two primary types of heterodonty, or differentiation between teeth in various positions on the jaws, that can be demonstrated in sharks. The first, here termed dignathic heterodonty, involves differences in morphology between teeth in opposition or approximate opposition in the upper and lower jaws. Dignathic heterodonty can apply to all opposing teeth in both jaws or to only some of them. The second type, monognathic heterodonty, involves differences between teeth in different positions on the same jaw series. Monognathic heterodonty is not restricted to situations in which ad- jacent teeth differ strongly in morphology, but also applies when a tooth in one position is different from that in another position on the same series but has a gradient of intermediate teeth between itself and the second tooth. The first condition can be called disjunct monognathic heterodonty; the second, gradient monognathic heterodonty. Applegate (1965) used a row-group terminology for implied disjunct monogna- thic heterodonty in the dentitions of Odontaspis taurus (Odontaspidae) and other sharks. The terms symphysials, alternates, and medials were used for dif- ferent tooth types in the region of the symphysis. Remaining teeth were grouped into anteriors, intermediates, laterals, and posteriors from premedial to postlateral along the dental band. Analogs of the intermediates in lamnoids do not exist in carcharhinids. However, some carcharhinid genera, especially those in the ad- vanced and intermediate groupings mentioned below, show strong disjunct monog- nathic heterodonty and have medials, alternates, symphysials, anteriors, laterals, and posteriors. Other genera (as Hemitriakis) have disjunct variation only be- tween the medials or alternates at the symphysis and the adjacent parasym- physial rows, which may be termed anteroposteriors. Two additional types of heterodonty can be defined here. Ontogenic hetero- donty is a gradient phenomenon in which tooth morphology at a functional series position in a single row or many rows changes with replacement of teeth during growth. Gynandric heterodonty, or dental sexual dimorphism, includes differ- ences in morphology of teeth in approximately similar series and row positions be- tween two individuals or groups of individuals of opposite sex and same species at about the same developmental stage. [Proc. 4TH SER. CALIFORNIA ACADEMY OF SCIENCES B. Caudal fin. Ficure 6. Carcharhinid fin terminology. A. Idealized first dorsal fin. Abbreviations: AM, anterior margin; AP, apex; BA, base; DM, dorsal margin; EL, HL, hypural lobe; IM, inner margin; IN, insertion; epural lobe; FRT, free rear tip; LOR, lower origin; LPVM, lower postventral margin; OR, origin; PM, posterior margin; Vor. XXXVIII] COMPAGNO: SYSTEMATICS OF GENUS HEMITRIAKIS ~I VERTEBRAE. Terminology for vertebral calcified parts follows Ridewood (1921). Springer and Garrick (1964) subdivided the vertebral complements of sharks into precaudal and caudal centra. They noted that an alternative grouping into monospondylic and diplospondylic centra was possible, but this was not utilized in their study. For present purposes the Springer and Garrick dichotomy of vertebral types is modified into a three-group system of monospondylic precaudal (MP), diplo- spondylic precaudal (DP), and diplospondylic caudal (DC) centra. This trichotomy is applicable to most carcharhinids, but breaks down in sharks such as Galeorhinus zyopterus where alternating long and short centra of a ‘‘stutter zone’ mark the transition from MP to DP centra. Springer and Garrick’s method of delimiting the caudal centra at the upper precaudal pit or upper caudal origin is followed here despite its shortcomings. In some instances it is useful to compare relative numbers of centra in dif- ferent vertebral groups of sharks with differing total vertebral counts. A system used here divides the MP, DP, and DC counts by the MP count to give DP/MP and DC/MP ratios that vary sufficiently between carcharhinid genera and species to be of systematic value (MP/MP = 1.00). An alternate system is to divide MP, DP, and DC counts by total count and multiply by 100 to obtain percent total count for each vertebral group. Fins. The terminology used here for carcharhinid fins is explained by fig. 6. The following terms apply to paired and unpaired fins other than the caudal: Origin; anterior margin; apex; posterior margin; free rear tip; inner margin; insertion; and base. The caudal fin terminology includes: Hypural lobe; epural lobe; terminal sector; subterminal notch; ventral lobe; dorsal margin; terminal margin; subterminal margin; upper postventral margin; lower postventral mar- gin; preventral margin; upper origin; and lower origin. Genus Hemitriakis Herre, 1923 Type spEcIES. Hemitriakis leucoperiptera Herre, 1923, by original designa- tion. Species. There are two named species: H. leucoperiptera, from the Philip- pine Islands (detailed distribution in Herre, 1953); and H. japanica (Miiller and Henle), from Japan, Taiwan, and Amoy (Chen, 1963). Hemitriakis japanica was placed in the genus Galeorhinus Blainville (or its junior synonyms, Galeus Cuvier, 1817, not Rafinesque, 1810, and Eugaleus Gill) < PRVM, preventral margin; STM, subterminal margin; STN, subterminal notch; TM, terminal margin; TS, terminal sector; UND, undulations in dorsal caudal margin; UOR, upper origin; UPVM, upper postventral margin; VL, ventral lobe. 76 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. by all previous writers (summarized in Fowler, 1941, and Chen, 1963). The species was named Galeus japanicus by Miller and Henle (1841). Specimens of Galeorhinus japanicus were compared with specimens of Gale- orhinus australis, G. chilensis, G. galeus (type species of Galeorhinus), G. zyopterus, and with specimens and Herre’s (1923) description of Hemitriakis leucoperiptera. This indicated that “japanicus” does not belong to Galeorhinus but is congeneric with Hemitriakis leucoperiptera. The two Hemitriakis species are close but H. leucoperiptera differs from H. japanica in having the first dorsal origin over inner pectoral margin (H. japanica with origin posterior to free rear tip of pectoral). The distance from pectoral free rear tips to pelvic origins about equal to first dorsal length from origin to free rear tip in H. leucoperiptera but much greater in H. japanica. Hemitriakis leu- coperiptera also has fewer vertebrae, with about 144-146 total count (2 speci- mens) and 34-35 MP centra (H. japanica with 156-161 total and 41-43 MP centra for 7 specimens). An undescribed Hemitriakis species may be represented by 4 specimens (SU— 40097) that differ from the sympatric H. leucoperiptera in various proportions, fin shapes, and in their strikingly barred and spotted coloration (H. leucopertp- tera and H. japanica have a nearly plain coloration). REDEFINITION AND DESCRIPTION OF THE GENUS Hemitriakis. Head flat- tened dorsoventrally, its length from snout tip to 5th gill opening about % of total length. Eyes high on sides of head, above horizontal head rim and level of nostrils by a space equal or greater than eye height (fig. 7A). Strong subocular ridge present, in dorsal view separating eyes from horizontal head rim by a wide space (fig. 1A). Eyes not visible in ventral view of head (fig. 1D). Eyes elon- gate, their apertures over twice as long as high, with a notch present posteriorly in adults and subadults. NLE external (fig. 7A), with its edge horizontal. Edge of SLE strongly differentiated. Subocular pouch shallow but well defined, with its interior surface covered with denticles. Spiracles present, slitlike or porelike, 1/5 to 1/7 of eye length. External gill openings short, the longest (3rd) less than eye length. Gill rakers absent from internal gill openings. Nostrils narrow, far apart, a nostril width about 2 times in internarial width. Anterior nasal flap a short rounded lobe, not a pointed barbel. Nostrils about half as far from mouth as from snout tip. Nasoral grooves absent. Mouth crescentic, broad, at least 2% times as wide as long. Large papillae absent from buccal cavity. Moderately long labial furrows present, upper about 1’ times as long as nostril width, the lower “% to 7% of upper. Upper labial fur- rows extending anteriorly to below first 4 of eye. Dignathic heterodonty weak, with upper anteroposterior teeth slightly larger and with higher crowns and more erect cusps than lowers; upper medials smaller Vot. XXXVIII] COMPAGNO: SYSTEMATICS OF GENUS HEMITRIAKIS ~I Ficure 7. Lateral view of carcharhinid heads. A. Hemitriakis japanica (SU-12677). B. Galeorhinus zyopterus (LJVC-0238). than lower ones. Disjunct monognathic heterodonty indicated by differentiation of 3 to 6 rows of medials in upper and lower jaws. Medial teeth differ from an- teroposteriors in their lesser size and erect primary cusps, flanked by 1 or 2 ~I 78 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. premedial and postlateral cusplets. The sharp-edged anteroposteriors are larger, compressed, bladelike cutting teeth with a strong oblique primary cusp and no premedial cusplets (fig. 3). Anteroposteriors show strong gradient monogna- thic heterodonty. In more premedial teeth in adults and subadults the primary cusp is large and has 2 to 4 postlateral cusplets flanking it. From premedial to postlateral the crowns of teeth become lower, the primary cusps become more oblique, and the postlateral cusplets become fewer and finally disappear. The most postlateral anteroposteriors have primary cusps reduced or absent and are very low and sharp-keeled. Ontogenic heterodonty present in more premedial teeth of anteroposteriors. These teeth gain more postlateral cusplets with growth, so that late embryos have no cusplets and adults have 2 to 4 cusplets. Gynandric heterodonty not apparent. Teeth moderately large, base width of longest lower anteroposteriors about 0.356-0.405 percent of total length in H. japanica. Tooth rows relatively few; Chen (1963) gives 23-29/27-33 (4 specimens) and Tang (1934) gives 35/33 (1 adult male) total tooth row counts for H. japanica. The 7 examples of H. japanica studied here have 33—38/29-33 rows. Herre (1923) gives 18/34 rows for the holotype (adult female) of H. leucoperiptera, but this may be erroneous as 33/30 rows were counted in one of the SU—27118 specimens (late embryo). One to 5 series functional along jaw edges. Teeth of adjacent rows in the alter- nate overlap pattern of Strasburg (1963). Serrations absent from crown edges. Crown premedial edge not indented and differentiated. Crown foot with a strong basal ledge overlapping a deep basal groove. Transverse ridges present on basal ledge only, not extending onto primary cusp. Roots low, deep, with strong transverse groove dividing attachment surface into 2 lobes and extending through extreme rim of root to form strong transverse notch. Teeth not notice- ably protruding when mouth is closed. Trunk not compressed, subcylindrical. Interdorsal ridge present. Lateral dermal keels absent from caudal peduncle. No precaudal pits. Head-trunk length from snout tip to cloaca equal to, or somewhat shorter than, tail length from cloaca to caudal tip. Denticles from sides of body below first dorsal fin small, with crowns much longer than wide at all sizes. A single strong medial cusp and bifurcated lon- gitudinal ridge with a weak lateral ridge on each side of crowns of adult denticles. In late embryos to subadults medial ridge not bifurcated and lateral ridges absent. A pair of weak lateral cusps often present on denticles, but these are not constant. Pectorals moderately large, pectoral area slightly greater than first dorsal area. Pectoral anterior margin about 1’ times as long as combined base and inner margin lengths. Apex of adpressed pectoral slightly posterior to its free rear tip when pectoral inner margin is held parallel to body axis. Origin of Vor. XXXVIII] COMPAGNO: SYSTEMATICS OF GENUS HEMITRIAKIS 79 pectoral below or slightly anterior to fourth gill opening. Pectoral skeleton projecting about “5 to 2 of pectoral anterior margin length into the fin. Distal pectoral radials slightly longer than proximal ones, with broad, truncate tips. Pelvics relatively small, their anterior margins less than % the length of pectoral anterior margins. Pelvic bases closer to Ist dorsal base than to 2nd dorsal base. Midpoint of 1st dorsal base almost equidistant between pelvic and pectoral bases or definitely closer to pectoral bases. First dorsal free rear tip anterior to pelvic origins. Second dorsal nearly as large as first one, with its height 70 to 80 percent of Ist dorsal height. Posterior margin of 2nd dorsal strongly concave. Anal much smaller than 2nd dorsal, its height ’ that of 2nd dorsal and its base only 7s to *4 of 2nd dorsal base. Anal posterior margin strongly concave in adults, shallowly concave in late embryos. Anal origin posterior to 2nd dorsal origin by about 43 of the 2nd dorsal base length. Anal insertion varying from under 2nd dorsal insertion to much less than 5 of 2nd dorsal base length posterior to it. Caudal with preventral and postventral margins expanded as a short ventral lobe in adults and subadults, but scarcely developed in late embryos. Preventral caudal margin over ’% of dorsal caudal margin in adults and subadults, slightly shorter in young. Postventral margin differentiated into upper and lower parts in subadults and adults, with upper postventral margin % to %5 of dorsal caudal margin. Subterminal caudal margin long, over ’% of terminal caudal margin length. Caudal short, dorsal margin about equal to head length and less than of total length. No lateral undulations in dorsal caudal margin. Terminal sector of caudal short; distance from subterminal notch to caudal tip only about 2’2 to 3 times in dorsal caudal margin. Vertebral axis of caudal slightly raised above body axis. Vertebrae moderately numerous, total count 144-161. Separation between MP and DP centra not sharp, gradual along two transitional centra. Vertebral calcification pattern of Applegate’s (1967) ‘“‘carcharhinoid” type. Chondrocranium very similar to that of Furgaleus ventralis as illustrated by Whitley (1948) and to that of Mustelus species described and illustrated by Gegenbaur (1872) and Holmgren (1941). Supraorbital crest of cranium strongly developed and entire. Intestinal valve of spiral type, with 6 to 8 turns in the spiral. Hemitriakis is livebearing and probably viviparous. Yolk-sac placentae are present on the SU—40097 (late embryo) specimens. FAMILIAL CLASSIFICATION OF HEMITRIAKIS The familial classification of the genus Hemitriakis is troublesome because one of its species, H. japanica, is conventionally placed in the family Carcharhin- 80 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. idae as delimited by Bigelow and Schroeder (1948). In contrast, H. leucoper- iptera is usually placed in the family Triakidae. Hemitriakis cannot simulta- neously reside in both families, but the problem goes beyond deciding in which family this genus belongs. This is because Hemitriakis is almost exactly inter- mediate between the Triakidae and Carcharhinidae as defined by modern writers. Hence the selection of a family for Hemitriakis is dependent on the validity of separating the Triakidae from the Carcharhinidae. According to Bigelow and Schroeder (1948) and to Garrick and Schultz (1963), triakids differ from carcharhinids only by NLE morphology and denti- tion. The triakids are supposed to have rudimentary, external, and transitional NLE types (except for Leptocharias and Triaenodon with an internal NLE), whereas carcharhinids have an internal NLE. The teeth of triakids are small, crushing molariform or bladelike multicuspidate types that are present in several functional series on the jaw sides. Carcharhinid teeth are small to large, blade- like, with not more than 1 or 2 series of teeth functional at the sides of the jaws. As noted by Garrick and Schultz, the separation of the two families is con- founded by the seemingly intermediate positions of 7Tviaenodon, Leptocharias, and Hemitriakis japanica. Triaenodon especially strains the classification by having “triakid” teeth and an internal NLE. However, Gohar and Mazhar (1964) claimed that T7riaenodon belonged in the Carcharhinidae because it has a scroll intestinal valve as in Carcharhinus and other advanced genera. An un- published study of the morphology of Tviaenodon obesus confirms Gohar and Mazhar’s results on the valvular intestine and also demonstrates that Tviaenodon is very different from other “triakids” in its cranial morphology, pectoral fin skeletal structure, head morphology, and many other characters. Of the various triakid and carcharhinid genera, Negaprion is evidently closest to Triaenodon. The teeth of Triaenodon superficially resemble those of other ‘‘triakids” only in having premedial and postlateral cusplets flanking a primary cusp, but are otherwise strikingly different in the advanced morphology of their crowns and roots. It is probable that the “‘triakid” characters of Triaenodon are convergent ones. Even without 7viaenodon to complicate the issue, the familial separation of Triakidae from Carcharhinidae, using the traditional characters, fails when other genera are considered. Thus, Hemitriakis had bladelike, sharp-edged anteroposterior teeth in 1 to 5 functional series that closely resemble those of Galeorhinus, but has medials with multiple premedial and postlateral cusplets closely resembling “‘triakid” teeth. Its NLE is external, as in many, but not all, supposed triakids. Furgaleus combines Galeorhinus-like upper anteroposterior teeth, Hemigaleus-like lower anteroposteriors with one erect primary cusp and no cusplets, and an external NLE. Furgaleus is conventionally placed in the Triakidae. Another triakid, Leptocharias, has an internal NLE and VoL. XXXVIII] COMPAGNO: SYSTEMATICS OF GENUS HEMITRIAKIS 81 anterolateral teeth with primary cusps, premedial cusplets, and _postlateral cusplets in the “triakid” pattern. In the genus Galeorhinus (a presumed carcharhinid), young specimens of G. australis, G. chilensis, G. galeus, and G. zyopterus have a transitional NLE, but half-grown to adult individuals have these structures internal. Finally, Triakis semifasciata and T. scyllia have an external NLE in young specimens but this changes to a transitional or fully internal one in adults and subadults. Adult and subadult Mustelus commonly have a transitional NLE, but large M. canis may have the internal type (Garman, 1913; Bigelow and Schroeder, 1948). The teeth of Triakis semifasciata are ar- ranged in only 3—4 functional series on the jaw edge. Also, in T. semifasciata the teeth show considerable ontogenic heterodonty, with loss of premedial and postlat- eral cusplets as the dentition is replaced until many to almost all of the teeth in adult specimens have only a strong, oblique primary cusp. 77iakis maculata also shows a similar type of ontogenic heterodonty (Kato, Springer, and Wagner, 1967). The orthodox distinction of Triakidae from Carcharhinidae is untenable at present because the supposedly diagnostic and traditional characters used to separate these families fail to do so. As the above examples show, there are enough transitional genera and species to make the retention of the two families Triakidae and Carcharhinidae an arbitrary choice based on tradition and con- venience. I prefer to submerge the Triakidae in the Carcharhinidae. This has the obvious disadvantage of creating a huge, unwieldy, and heterogeneous com- plex that combines advanced forms with scyliorhinoid genera. However, it may be possible eventually to divide the family Carcharhinidae as here constituted into a number of lesser families using new characters of comparative morphology that are now being investigated. Hence, the genus Hemitriakis is considered a member of the expanded family Carcharhinidae. COMPARISON WITH OTHER GENERA This section demonstrates the distinctness of Hemitriakis from other car- charhinid genera. A series of synoptic keys is presented in which allied groups of genera are compared and contrasted with Hemitriakis. A general key to car- charhinid genera is not offered here as revisional studies on the family are in- complete at present. To facilitate comparison of Hemitriakis with certain genera, it was necessary to include some species rearrangements within them in the following discussions. This primarily involved removal of some species from the heterogeneous genera Triakis and Galeorhinus and proposal of tentative new generic arrangements to accommodate them. ADVANCED AND INTERMEDIATE CARCHARHINIDS. A large proportion of car- 82 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. charhinid genera comprise the two groups here termed the advanced and inter- mediate carcharhinids. The advanced genera include A prionodon, Carcharhinus, Galeocerdo, Hypoprion, Isogomphodon, Lamiopsis, Loxodon, Negaprion, Prio- nace, Rhizoprionodon, Scoliodon, and Triaenodon. The intermediate genera are Hemigaleus (including Chaenogaleus, Negogaleus, and Paragaleus) and Hemi- pristis (including Dirrhizodon and Heterogaleus ). The advanced carcharhinids are so named because they depart furthest of all genera in the family from the morphology of generalized scyliorhinid genera widely thought to occupy the most primitive position among carcharhinoid sharks. The Sphyrnidae (hammerheads) is closely allied to the advanced car- charhinids but is not included for comparison with Hemitriakis because of its unique and obvious specializations. The intermediate genera are very similar to the advanced ones in many characters, but retain some generalized features in the morphology of the cranium, fins, dentition, and intestinal valve. The advanced and intermediate carcharhinids are grouped together for brevity to compare them with Hemitriakis. The following synopsis covers only a representative sample of numerous differences between these genera and Hemi- triakts. la. Eyes low on head, their ventral margins meeting or extending across the horizontal head rim. Subocular ridge weak or obsolete. NLE always internal, with slanted edge. Sub- ocular pouch very deep, with inner surface of SLE and bottom of pouch lacking denticles. Crowns of teeth without a strong basal ledge and groove (except in lower teeth of Hemz- galeus). Transverse ridges virtually absent from crown foot. Denticles of adults as wide or wider than long, with three or more subequal cusps and ridges. Precaudal pits present always at upper caudal origin and usually at lower origin also. Pectoral skeleton projecting at least 74 of pectoral anterior margin length into fin, with distal radials much longer than proximals. Distal radials with tapering, acute tips. Anal large relative to 2nd dorsal, its height 70 per cent or more of 2nd dorsal height. Lateral undulations present along dorsal caudal margin (except in young of some species and in Scoliodon where undulations are in- differently developed). Intestinal valve a scroll in advanced carcharhinids, but a spiral with only 2-6 turns in Hemigaleus and Hemipristis. Chondrocranium with isolated preorbital and postorbital processes only, without an intermediate supraorbital crest . . . Advanced and Intermediate Carcharhinids. 1b. Eyes high on head, their ventral margins widely separated from the horizontal head rim in ventral view. Subocular ridge very strong. NLE external, with edge horizontal. Subocular pouch shallow, with denticles covering its internal surface. Crowns of teeth with strong basal groove and ledge. Transverse ridges irregularly present on crown foot. Denticles of adults longer than wide, with a very strong medial ridge and cusp and flanking weak lateral ridges also; weak lateral cusps irregularly present. Precaudal pits absent. Pectoral fin skeleton projecting only 75 to % of pectoral anterior margin dis- tance into fin, with distal radials slightly longer than proximals. Distal radials with parallel articulating edges and truncate tips. Anal relatively small in relation to second dorsal, its height only % that of second dorsal height. Lateral undulations of dorsal caudal margin absent. Intestinal valve a spiral, with 6-8 turns. Chondrocranium with strong supraorbital crest between preorbital and postorbital processes = _.. Hemitriakis. VoL. XXXVIII] COMPAGNO: SYSTEMATICS OF GENUS HEMITRIAKIS 83 GALEORHINUS AND ALLIED GENERA. Recent workers have included the fol- lowing species in Galeorhinus: G. galeus (Linnaeus, 1758); G. japanicus (Muller and Henle, 1841); G. australis (Macleay, 1881); G. zyopterus (Jordan and Gilbert, 1883); G. chilensis (Perez Canto, 1886), including G. molinae (Philippi, 1887): G. omanensis (Norman, 1939); G. hyugaensis Miyosi, 1939; G. vita- minicus de Buen, 1950; and G. zanzibariensis Smith, 1957. Of these nine species, four are sufficiently different to require removal from Galeorhinus. ““Galeorhinus” japanicus has been already transferred to Hemitriakis. G. hyugaensis and the closely similar G. zanzibariensis are placed in the genus Hypogaleus and discussed below. “Galeorhinus” omanensis, as suggested by its describer (Norman, 1939), is not congeneric with Galeorhinus and will be discussed in a forthcoming paper by Mr. Stewart Springer and myself. It is included in the synopsis below to dis- tinguish it from Hemitriakis. The remaining 5 nominal species comprise the genus Galeorhinus as here delimited. Garman (1913), Fowler (1929, 1941), and Bigelow and Schroeder (1948) considered G. zyopterus a junior synonym of G. galeus, while Kato, Springer, and Wagner (1967) tentatively synonymized G. chilensis with G. zyvopterus. McCoy (1885) compared G. australis with G. galeus and listed several proportional differences between a few specimens of G. australis and one of G. galeus. However, comparison of a pair of equal sized specimens of G. galeus and G. australis suggests that most, if not all, of McCoy’s differences were allometric ones based on comparison of dissimilar sized specimens. G. vitam- inicus, as described by De Buen (1950), is hardly different from other Galeo- rhinus species. It may be that all 5 species are synonyms, as Smith (1957b) maintained, but the validity of this hypothesis cannot be tested at present because of insufficient material. Smith (1957b) proposed the subgenus Hypogaleus for its type, Galeorhinus (Hypogaleus) zanzibariensis Smith, 1957, and for Hemitriakis japanica. Accord- ing to Smith, Hypogaleus species have teeth without the transverse notch on their roots, but in Galeorhinus (including only G. galeus) the notch is present. In Galeorhinus the caudal terminal sector is large, about 4 caudal length, but much smaller and less than 2 caudal length in Hypogaleus. Hypogaleus has the second dorsal at least twice as great in area as anal, but Galeorhinus has these fins subequal in size. In Galeorhinus the pelvic fins of adults are inserted behind the middle of the total length; in Hypogaleus the pelvics of adults are inserted well in advance of the middle of the total length. Apparently Smith used only literature descriptions for Hemitriakis japanica. Although most of the fin characters for Hypogaleus fit Hemitriakis japanica, the dentitional character does not, as this species has a strongly developed transverse notch. Also, Smith’s tooth photographs of G. (Hypogaleus) zanzibariensis seem to indicate that this species has much higher roots and obsolete basal ledges and 84 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 8. Lateral views of carcharhinid sharks. A. Hemitriakis japanica (SU-12677). B. Hypogaleus zanzibariensis (modified from Smith, 1957b). C. Galeorhinus zyopterus (LJVC-0238). Vot. XXXVIII] COMPAGNO: SYSTEMATICS OF GENUS HEMITRIAKIS 85 grooves on its teeth (figs. 4B, 4E). Galeorhinus and Hemitriakis as presently delimited have low roots and strong basal ledges and grooves (figs. 3, 4A, 4D). Also, Smith did not examine Miyosi’s (1939) description of Galeorhinus hyugaensis. Comparison of the accounts of G. hyugaensis by Miyosi and by Chen (1963) with those of G. zanzibariensis by Smith (1957b) and by D’Aubrey (1964) indicates that these species are virtually identical in all important details of morphology (including dentition) and coloration. Indeed, it will be necessary to compare specimens of the two species to determine what differences, if any, exist between them. Galeorhinus zanzibariensis and G. hyugaensis are close to Galeorhinus proper and to Hemitriakis but are sufficiently different to merit generic status. Hence I propose to raise Hypogaleus Smith from subgenus to genus and include in it the two nominal species H. hyugaensis (Miyosi, 1939) and H. zanzibarien- sis (Smith, 1957). The Australian genus Furgaleus is included here because its two species, as described and illustrated by Whitley (1943a, 1943b, 1944, 1948), have upper anteroposterior teeth that strongly resemble those of Hemitriakis, Hypogaleus, and Galeorhinus. Furgaleus is closest to Hemitriakis but is easily distinguished. The following is a synopsis of Galeorhinus and allied genera (including Hemiutriakis ). la. Postlateral cusplets absent from anteroposterior teeth in upper jaw. Upper medial teeth without cusplets. First dorsal fin far forward, with origin over anterior half of pectoral base. Caudal fin without ventral lobe in adults _ “Galeorhinus” omanensis. 1b. Postlateral cusplets present on upper anteroposterior teeth. Upper medials with both premedial and postlateral cusplets. First dorsal origin posterior to pectoral base inser- Hone Candalawith moderate: toystrons: ventral) lobe im adults, = eee 2 2a(1b.). Nostrils larger and closer together, their widths about twice in internarial width. Nostrils equidistant between snout tip and mouth. Anterior nasal flap formed into a long, slender barbel. Dignathic heterodonty strong, with upper anteroposteriors having oblique primary cusps and postlateral cusplets while lowers have erect primary cusps and UMC ULS 19) US eee eer ac NEE Se ON Ee EN As Ee 5 eee oe eee, Oe Furgaleus. 2b(1b.). Nostrils smaller and farther apart, their width 214 times in internarial width or more. Nostrils much closer to mouth than to snout tip. Anterior nasal flap not produced into a barbel. Dignathic heterodonty weak; upper and lower anteroposteriors with oblique DAIRY CUS aravel joyoslewerrall CUO: $e 3a(2b.). Eyes high on sides of head, above level of nostrils by a space equal to or greater than eye height. Eyes over twice as long as high. NLE external in adults and sub- adults, with horizontal edge. Anterior nasal flap moderately large, expanded as a rounded lobe. Posteriormost anteroposterior teeth elongate, carinate. Interdorsal ridge present. Adult and subadult denticles with crowns much longer than wide and with lateral cusps and ridges weak. Caudal with short ventral lobe in adults and subadults (fig. 8A) ene nen Tb SE Eo Oe ee ee ee a Hemitriakis. 3b(2b.). Eyes lower on sides of head, Aion level of nostrils by a space less than eye height. Eyes twice as long as high or less. NLE internal in adults and subadults, with diagonal edge. Anterior nasal flap reduced, expanded as a minute, pointed lobe. Posteriormost 86 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. anteroposterior teeth not elongate, carinate. Interdorsal ridge absent. Denticles of adults and subadults with crowns nearly as long as wide and with strong lateral cusps and ridges. Caudal withvlong ventral lobesinwadulltssamne sulbyaic ul tse 4, 4a(3b.). Head very short, about 4% of total length in adults (fig. 8B). Subocluar ridge strong; in dorsal view eyes separated from horizontal head rim by a moderately wide space. Transverse notch absent from tooth roots (fig. 4B). First dorsal as large or larger than pectoral. Second dorsal about 73 as high as first dorsal and about twice as high as anal. Terminal sector of caudal about 2.6 in dorsal caudal margin. Upper postventral margin meni VY as lene as, Closeall @aclell imeyeainy Hypogaleus. 4b(3b.). Head longer, over ¥% of total length in adults (fig. 8C). Subocular ridge obsolete; in dorsal view ventral eye margins contact horizontal head rim. Transverse notch present on tooth roots (fig. 44). First dorsal much smaller than pectoral. Second dorsal less than half as high as first one and subequal to the anal in height. Terminal sector of caudal about 2.0 in dorsal caudal margin. Upper postventral margin only about %4 as long asvdorsalltcaudall mmarginis 22.3 2 ee ee eee eee eee Galeorhinus. LEPTOCHARIAS AND SCYLLIOGALEUS. Two aberrant, monotypic African genera, Leptocharias and Scylliogaleus, differ greatly from Hemitriakis. While Scylliogaleus is apparently closest to typical Mustelus species, the taxonomic po- sition of Leptocharias is quite isolated in the Carcharhinidae. Leptocharias, Hemitriakis, and Scylliogaleus are compared in the following synopsis. Additional data on Scylliogaleus is from Boulenger (1902) and Smith (CUISEKe) la. Eyes low on sides of head, above level of nostrils by less than eye height. Subocular ridge obsolete; ventral margin of eyes touching horizontal head rim in dorsal view. Eyes less than twice as long as high, with a slant-edged, internal NLE (fig. 2D). Spiracles minute, porelike, less than %4o of eye length. Anterior nasal flap expanded as a pointed barbel. Gynandric heterodontia strong, expressed by presence of about 4 tooth rows of hypertrophied ‘“anteriors” in both jaws on either side of weakly differentiated medials in adult males but not females. Teeth other than anteriors with slender, erect primary cusps and both premedial and postlateral cusplets, not bladelike or molariform. Vertebrae very numerous, 198-213 total centra (9 examples; data for 2 from Springer and Garrick, 1964). Spiral intestinal valve with about 16 turns. Supraorbital crest absent from cranium, with isolated preorbital and postorbital processes only __—----------------- Leptocharias. 1b. Eyes higher, above level of nostrils by an eye height or more. Subocular ridge strong ; eyes separated from horizontal head rim by a wide space. Eyes over twice as long as high, with horizontal-edged and external NLE (fig. 7A). Spiracles larger, 44 to 4% of eye length. Anterior nasal flap not formed into a barbel. Gynandric heterodonty not apparent. Teeth either molariform or bladelike, without premedial cusplets (except on medials of Hemi- triakis). Vertebrae fewer, 143-161 total centra (10 examples). Spiral valve with 6 to 8 (HUNTON, Srojouelondoyuernll Cree jOreeSerle Wil CHeNANNT — Dre 2a(1b.). Snout bluntly rounded, semicircular in shape. Anterior nasal flaps greatly enlarged as broad triangular lobes extending posteriorly to overlap mouth. Nostrils very large and separated by a distance much shorter than a nostril width. Deep nasoral grooves present. Teeth with crowns flattened and rounded to form a crushing pavement as in typical Mustelus species. Teeth not differentiated into medials and anteroposteriors. Tooth rows 60-72 in each jaw; 9-10 series functional in upper jaw, 16-17 in lower (Smith, 1957c). Vot. XXXVIII] COMPAGNO: SYSTEMATICS OF GENUS HEMITRIAKIS 87 Pelvics large, anterior margin lengths % or more of pectoral anterior margin lengths. Free rear tip of first dorsal over or posterior to pelvic origins _.... Scylliogaleus. 2b(1b.). Snout narrower, parabolic in shape. Anterior nasal flaps small truncate lobes, terminating far anterior to mouth. Nostrils smaller, further apart, their widths about 2% times in internarial width. Nasoral grooves absent. Teeth differentiated into medials and anteroposteriors, not forming a pavement. Anteroposteriors sharp-edged, bladelike teeth, with oblique primary cusps and postlateral cusplets; medials are not bladelike and have premedial cusplets also. Tooth rows fewer, 18(?)-39/27-34; only 1 to 5 series functional along jaw edges. Pelvics smaller, their anterior margins less than 4% length of anterior pectoral margins. Free rear tip of first dorsal anterior to pelvic origins Hemitriakis. TRIAKIS AND ASSOCIATED GENERA. Included here are those species placed by Bigelow and Schroeder (1948) and by Kato (1968) in the genera Triakis, Mus- telus, Eridacnis, and Calliscyllium. The systematics of Triakis and its relatives is unsatisfactory at present. This is in part due to the interpretations of Triakis by Garman (1913), Fowler (1929, 1941), Bigelow and Schroeder (1944, 1948), Garrick (1954), Kato (1968), and Springer (1968), which included several scyliorhinid-like species in this genus that are clearly not congeneric with typical Triakis species (as T. scyllia and T. semifasciata). Also, the separation of Triakis from Mustelus on differ- ences in tooth crown morphology seems untenable, as there are many dentitionally intermediate species between “typical” extremes of these genera. Bigelow and Schroeder (1940) and Kato (1968) have discussed the latter problem in detail, but left the two genera separate. Smith (1957a) proposed a solution to the Triakis heterogeneity problem. He removed Calliscyllium venustum Tanaka from Tvriakis and reinstated Calliscyl- lium Tanaka as a monotypic genus for it. Also, he proposed the genus NV eotriakis for his V. sinuans and for Triakis barbouri Bigelow and Schroeder. Finally, he transferred Triakis henlei (Gill) to Mustelus. Smith’s separation of Calliscyllium and Neotriakis from Triakis is undoubt- edly correct, as species included in these scyliorhiniform genera exhibit many dif- ferences from typical Triakis. Unfortunately, Smith retained two anomalous species, Triakis attenuata Garrick and Hemitriakis leucoperiptera Herre, in the genus Triakis. Triakis attenuata is closer to Calliscyllium and Neotriakis in the sense of Smith than to Triakis proper, and its presence in Triakis makes separation of that genus from Neotriakis especially difficult with the limited and ambiguous generic characters utilized by Smith to define these genera. Finally, placement of Triakis henlei in Mustelus further undermines the classical tooth crown differences purported to separate Triakis from Mustelus; however, T. hen- lei is closer to typical forms of Mustelus than to those typical of Triakis in many respects. Smith was evidently unaware of the Triakis-Mustelus continuity prob- lem, as he later (1957c) gave Mustelus familial separation from Tyriakis in his family Scylliogaleidae (along with Scylliogaleus). 88 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. A tentative reclassification of 77iakis and associated genera is presented here, subdividing these taxa into two groups: 1. Typical forms of Tviakis, intergrad- ing species, and typical Mustelus forms. 2. Scyliorhiniform triakoids, including as subgroups: A. Genus Proscyllium; B. Genus Eridacnis; C. Triakis fehlmanni; D. T. attenuata. Triakis, Mustelus, and intermediates (or Triakis-Mustelus) are closer to Hemitriakis than are other carcharhinids. Triakis-Mustelus includes Triakis scyl- lia, T. semifasciata, T. maculata, T. acutipinna, Mustelus henlei, and the various other Mustelus species. Typical species of Triakis, with strongly cuspidate teeth (7. scyllia, T. semi- fasciata), form one extreme of a dentitional continuum with molariform-toothed Mustelus species at the other extreme. The continuum is filled by a host of denti- tionally intermediate forms, as 7. maculata, T. acutipinna, Mustelus nigropuncta- tus, M. henlei, M. dorsalis, M. megalopterus, M. natalensis, and M. higmani, that exhibit various stages of cusp and cusplet reduction on tooth crowns. Also, ex- amination of small (150-450 mm. total length range) specimens of typical Mus- telus species, as M. canis, M. californicus, M. manazo, M. mustelus, M. lunulatus, and M. griseus, indicates that cusps are often well developed on the teeth of small individuals and that cusp obsoleteness in larger examples probably results from ontogenic heterodonty. Although condition of tooth cusps has been the only character regularly utilized in separating Triakis from Mustelus, extremes of the former genus differ from typical members of the latter by a number of additional characters. These include: 1. Absence of peg on inner face of crown and root. 2. Lesser number of tooth rows. 3. Lesser number of tooth series. 4. Absence of a tooth pavement. 5. Bluntly rounded, short snout (versus long, pointed or paraboloid snout in Mustelus). 6. Very short, arcuate mouth (versus longer, more angular mouth in many Mustelus species). 7. Reproduction ovoviviparous (viviparous in at least some Mustelus, including M. henlei). In addition, there are about a dozen cranial differences between Triakis semifasciata and 3 species of Mustelus (M. henlei, M. californicus, and M. lunulatus, which are virtually identical cranially). The brain, cranial nerves, and sense organs of 7. semzfasciata also differ in several respects from those of W/. henlei. The tooth peg is found in many Mustelus species (including M. henlez), but not in Triakis maculata, T. scyllia, or T. semifasciata. Its condition is not con- firmed for all species of Triakis-Mustelus and cannot be used to separate the two genera at present. Tooth row and series counts apparently vary along a continuum as in crown morphology, with an added complication that in at least some Mustelus (if not all forms) the tooth row and series counts increase with size increase. Tooth pavementization, snout shape, and mouth morphology evi- dently show a similar variation spectrum. Data on cranial, neural, and repro- Vou. XXXVIII] COMPAGNO: SYSTEMATICS OF GENUS HEMITRIAKIS 89 ductive characters is not available for many to most 7viakis-Mustelus species, making it impossible to judge their utility in separating the two genera. External morphology suggests that Triakis is not separable from Mustelus, but merging the two genera here would be premature with incomplete knowledge of promising anatomical characters. However, Triakis-Mustelus is treated as a single unit here for comparison with Hemitriakis. The scyliorhiniform triakoids include species formerly placed in the genera Proscyllium, Calliscyllium, Eridacnis, Neotriakis, and Triakis. They are divisable into four subgroups, two of which are provisionally ranked as genera. The first genus, Proscyllium, is a structural link between the Carcharhinidae and Scyliorhinidae but is placed in the former family because of its anteriorly positioned first dorsal fin. The systematic treatment of Proscyllium and its synonym, Calliscyllium, by various writers has been highly variable and extremely confusing. Hilgendorf (1904) proposed Proscyllium as a subgenus of Scyllium Cuvier (= Scylliorhinus Blainville), with a single new species, S. (Proscyllium) habereri, from Formosa. Later Tanaka (1912) described a new genus and species, Calliscyllium venustum, from Japan. Tanaka did not mention Hilgendorf’s very similar species in his account. Although Tanaka considered Calliscyllium a_ scyliorhinid, Garman (1913) placed it in his family Galeorhinidae (= Triakidae) and synonymized it with Triakis. Garman also placed Scyllium (Proscyllium) habereri in the Catulidae (=Scyliorhinidae) and raised the rank of Proscyllium to genus. Schmidt (1930) described and illustrated a Japanese specimen of Proscyllium habereri. His account is of special interest as he compared his specimen with measurements and photographs of the holotype of Hilgendorf’s species and found no significant differences between the two specimens. Schmidt’s account of Proscyllium habereri closely matches Tanaka’s description of Calliscyllium venustum, but for unknown reasons Schmidt did not refer to Tanaka’s account or to his own (1928) description of an Okinawan specimen of Calliscyvllium venustum. White (1937) recognized both Calliscyllium venustum and Proscyl- lium habereri as scyliorhinids in a broad sense but placed the former in her family Halaeluridae and the latter in her family Catulidae. Fowler (1929, 1941) followed Garman in placing Proscyllium in Scyliorhinidae and placed Triakis venusta (Tanaka) in the subfamily Triakiinae of the family Eulamiidae or Galeorhinidae (= Carcharhinidae). Bigelow and Schroeder (1948) placed Proscyllium habereri in the family Triakidae, but did not discuss its generic status in that family. These writers followed Garman’s synonymy of Calliscyllium with Triakis. Garrick (1954) discussed Triakis venusta, but not Proscyllium habereri. Smith (1957a) recognized Calliscyllium as distinct from Tvriakis, but also overlooked Proscyllium habereri. Lindberg and Legeza (1959) synonymized Proscvllium with Triakis, but considered 90 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Triakis haberert distinct from T. venusta. Chen (1963) placed both Proscyllium habereri and Triakis venusta in the family Triakidae, but separated Proscyllium from Triakis by supposed absence of the NLE in the former genus. Finally, Kato (1968) removed Calliscyllium from synonymy of Triakis on reproductive differences, but did not mention Proscyllium. Comparison of accounts of Proscyllium habererit and Calliscyllium venustum with each other and with specimens indicates that Lindberg and Legeza were correct in regarding these species as congeneric. However, ‘‘venustum” and “haberert” are not congeneric with typical species of Triakis and are placed here in the genus Proscyllium. Calliscyllium is therefore a junior synonym of Proscyl- lium. The two species P. venustum and P. habereri are possibly synonyms also, as the small differences between them listed by Lindberg and Legeza (1959) may be of only variational and allometric significance. The genus Eridacnis includes a few species of deepwater sharklets allied to Proscyllium but sufficiently different to merit generic status. Eridacnis was established by Smith (1913) for E. radcliffei, a new shark from the Philippine Islands. Eridacnis was supposed to differ from Tvriakis Muller and Henle by lacking labial furrows, but, as Kato (1968) pointed out, the holotype of E. radcliffei has vestigial labial furrows presumably overlooked by Smith. Bigelow and Schroeder (1944) described as Triakis barbouri a similar but specifically distinct shark from Cuba, but did not compare it with Evidacnis radcliffet. Misra (1950) described a third form, Proscyllium alcocki, from the Andaman Sea. Data from Misra’s account indicates that ‘“alcocki’’ does not belong in Proscyllium as here defined but falls in Eridacnis instead. The species “alcocki” closely resembles E. radcliffec and therefore it is quite possible that these two names are synonymous (Norman, 1939, reported E. radcliffe: from the Gulf of Aden, which is west of the type localities of both E. radcliffei and “alcocki”). Smith (1957a) described a new genus, Neotriakis, for his new South African species V. sinuans. Smith included Triakis barbouri in Neotriakis but overlooked Proscyllium alcocki and did not compare Neotriakis species with the closely similar Eridacnis radcliffei. Kato (1968) considered the characters used to separate Neotriakis and Eridacnis from Triakis to be untenable, and synonymized the three genera. However, Kato regarded the species ‘“‘radcliffei,” “barbourt,”’ and “sinuans” as closely related to each other within the genus Triakis. Kato’s synonymy was adopted unchanged by Springer (1968). The genus Eridacnis is revived here for the species E. racliffei, E. alcocki, E. barbouri, and E. sinuans, with Neotriakis considered as a junior synonym. Triakis fehlmanni, a small shark recently described by Springer (1968) from Somalia, forms a third group closely similar to Proscyllium and Eridacnis in many details. Its vertebral calcification pattern and relatively short broad caudal are as in Proscyllium, but its vertebral count, vertebral group ratios, short Vot. XXXVIII] COMPAGNO: SYSTEMATICS OF GENUS HEMITRIAKIS 91 body cavity, nostril spacing, pectoral fin position, first dorsal size, and anal base size fit Evidacnis. The blotched and spotted color pattern, extremely short pre- caudal tail (distance from cloaca to lower caudal origin about twice in distance from snout tip to cloaca), broad head, and stout body distinguish ‘“‘feklmanni”’ from both Proscyllium and Eridacnis. Mode of reproduction and clasper mor- phology are unknown for the species. Tviakis fehlmanni seems closer to Eridacnis than Proscyllium but may require subgeneric or generic separation from typical Eridacnis species. It does not belong to Triakis-Mustelus as here limited and cannot be confused with Hemitriakis. Generic placement of “‘feklmanni” is prob- lematical at present; therefore the species must be left as a tentative and possibly dubious appendage to Eridacnis. The New Zealand Triakis attenuata, as described by Garrick (1954), agrees with Proscyllium, Eridacnis, and T. fehlmanni in its NLE type, detailed tooth morphology, eye position, and large second dorsal, but differs from these forms in its elongate snout, narrower and more widely spaced nostrils, longer labial furrows, more numerous tooth rows, more anterior position of first dorsal fin, origin of second dorsal anterior to anal origin, exceptionally small anal fin with base only half length of second dorsal base, weak ventral caudal lobe, and larger size. Unfortunately nothing is known of its cranium, pectoral fin skeleton, verte- bral calcification pattern, vertebral counts, vertebral group ratios, clasper morphology, and buccal cavity. Triakis attenuata presumably forms a tentative fourth group of scyliorhiniform triakoids allied to, but distinct from, Proscyl- lium-Eridacnis-T. fehlmanni. The species is remote from Hemitriakis and is suf- ficiently different from Tviakis-Mustelus to be excluded from that group. Separate generic status may be required for 7. attenuata, but insufficient data on the species prohibits a decision on the matter for now. The following generic synopsis compares Hemitriakis to Triakis-Mustelus, Proscyllium, and Eridacnis (excluding T. fehlmanni). la. NLE rudimentary in adults (fig. 2A.). Labial furrows vestigial, confined to corners of mouth. All teeth with erect cusps and usually cusplets also (some species have teeth near symphysis lacking cusplets). Posterior teeth polycuspidate, comblike in shape (fig. 4C). Gradient monognathic heterodonty present, in which premedial cusplets increase in number from symphysis to rictus and displace primary cusp from central position on crown foot to a postlateral location. Cusps and cusplets do not become obsolete with age. First dorsal fin with midpoint of its base closer to pelvic origins than to pectoral insertions. Second dorsal origin over or posterior to anal origin. Pectoral fin skeleton as in scyliorhinids, with distal radials much shorter than proximal ones. Vertebral centra of thoracic region in adults with peripheral calcifications of the intermedialia only, not developed into strong lateral and vertical wedges between halves of calcified primary double cone (figs. 5A, 5B.). Diagonal calcified lamellae of double cone, when present, in form of rounded lobe opposite each basidorsal and basiventral. Large papillae present on dorsal and ventral surfaces of buccal cavity and pharynx posterior to teeth, forming dermal gill rakers along internal branchial ADEREUTCS eens awe eee, Soe ee tte ee ee ea A 2 1b. NLE external, transitional, or internal in adults. Labial furrows well developed, 92 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. extending far onto jaws. Teeth either cuspless or with cusps that range from erect to strongly oblique and often showing monognathic heterodonty in increasing obliqueness toward ends of dental band. Posterior teeth carinate, molariform, or weakly monocuspidate, not comblike (fig. 4F.). Increase of premedial cusplets and displacement of primary cusp postlaterally not apparent in species with cuspidate teeth, but instead cusps and cusplets tend to become less prominent postlaterally and may be completely absent on posteriormost teeth. Many species (not including Hemitriakis) tend to reduce or lose cusplets or even cusps with age. First dorsal fin with base midpoint equidistant between pectoral insertions and pelvic origins or closer to pectoral insertions. Second dorsal origin well anterior to anal origin. Pectoral fin skeleton with distal radials equal in length to, or longer than, proximal ones. Vertebral centra of thoracic region of adults and subadults with intermedialia extending as strong lateral and vertical wedgelike calcifications between halves of calcified primary double cone (figs. 5C, 5D.). Diagonal calcified lamellae well developed, extending as thin plates into the basidorsals and basiventrals. Papillae absent from buccal cavity, pharynx, and internal branchial apertures 2a(la.). Nostrils very close together, internarial width only % nostril width. Distance from hind edge of anterior nasal flaps to mouth only about \% of nostril width. Head length from snout tip to 5th gill opening shorter than body length from pectoral insertion to pelvic origin. First dorsal length from origin to free rear tip less than 4% length of interdorsal space. Anal base length only 1% of distance between anal insertion and lower caudal origin. Caudal short, less than 14 of total length. Greatest height of caudal about 14 of upper caudal margin. Vertebrae more numerous, total count 146-168 (6 examples). DP/MP ratios 1.58-1.82; DC/MP 1.08-1.28. Diagonal calcified lamellae of trunk centra present as four rounded lobes extending slightly into areas of basidorsals and basiventrals (fig. 5B.). Claspers of adult males with a row of recurved clasper hooks along external flap of hypopyle. Color pattern of scyliorhinid-like spots and _ stripes PLESCMEM REP FOGUCLIOMNTO;vAlfo ai OLS meanness Proscyllium. 2b(1b.). Nostrils farther apart, internarial width about equal to nostril width. Distance from hind edge of anterior nasal flaps to mouth about ™% of nostril width. Head length longer than body length from pectoral to pelvic. First dorsal length *%5 to 45 of interdorsal space. Anal base length subequal to distance between anal insertion and lower caudal origin. Caudal longer, over 14 of total length. Greatest height of caudal less than ¥% of upper caudal margin. Vertebrae less numerous, total count 113-135 (26 examples). DP/MP ratios 1.05-1.45; DC/MP 1.29-1.53. Diagonal calcified lamellae not developed in trunk centra (fig. 5A.). Claspers without hooks. Coloration plain or with a few obscure stripes confined to tail. Reproduction ovoviviparous as far as is known —_ Eridacnis. 3a(1b.). Nostrils narrow and farther apart; nostril width about 2% times in internarial width. Teeth larger, basal width of largest lower anteroposteriors about 0.356 to 0.405 percent of total length (H. japanica, 4 examples). Teeth differentiated into medials and anteroposteriors. The latter are strongly compressed, bladelike cutting teeth with an oblique primary cusp and a few small postlateral cusplets only. Fewer tooth rows present, ?18-38/24-34. Pelvic fins with anterior margins less than half as long as pectoral anterior TMA OUNS yee seek eee eee eed te ene Code oe 2 eee ee eee eee eee Pena) Soe rey Hemitriakis. 3b(1b.). Nostrils wider, closer together; nostril width 2 times or less in internarial. Teeth smaller, those of species with largest teeth (Triakis semifasciata) only 0.172 to 0.262 percent of total length (largest lower teeth, 11 examples) and considerably smaller in other species. Teeth not differentiated into distinct medials and anteroposteriors, but showing regular gradient monognathic heterodonty between rows in symphysial and parasymphysial regions. Teeth corresponding to anteroposteriors of Hemitriakis either cuspless or having Vor. XXXVIII] COMPAGNO: SYSTEMATICS OF GENUS HEMITRIAKIS 93 an erect or oblique median primary cusp and usually both premedial and _ postlateral cusplets when cusplets are present. Teeth more weakly compressed, not sharp-edged, modified for grasping or crushing. More tooth rows present, 44-80+/33-80+. Pelvic fins larger, with anterior margins over % as long as pectoral anterior margins —.- a ees as ie ED a a eT REN OP Triakis-M ustelus. SUMMARY The shark Galeus japanicus Miller and Henle, long considered a species of Galeorhinus Blainville (or one of its synonyms), is placed in the genus Hemitriakis Herre, which is removed from the synonymy of Triakis Miiller and Henle and redefined. Hemutriakis contains two described species, H. japanica (Muller and Henle) and H. leucoperiptera Herre. The familial position of Hemitriakis is discussed and the separation of the families Triakidae and Carcharhinidae is rejected on present evidence. Hemitriakis is placed in the expanded family Carcharhinidae. Other carcharhinid genera are compared with Hemitriakis in synoptic keys, and several tentative systematic rearrangements of species in certain genera are presented to facilitate comparison with Hemitriakis. The genus Galeorhinus is restricted to the nominal species G. galeus, G. australis, G. zyopterus, G. chilensis, and G. vitaminicus, while a former subgenus, Hypogaleus Smith, is accorded generic rank. Hypogaleus contains two nominal species, H. zanzi- bariensis (Smith) and H. hyugaensis (Miyosi). Consideration of Galeorhinus omanensis (Norman) is postponed for another paper. In addition to Hemitriakis, four tentative groups of scyliorhiniform triakoids are removed from 7viakis. The first is the genus Proscyllium Hilgendorf, of which Calliscyllium Tanaka is a junior synonym. Proscyllium has two nominal species, P. habereri (Hilgendorf) and P. venustum (Tanaka). The genus Eridacnis Smith forms the second group, with EF. radcliffei Smith, E. barbouri (Bigelow and Schroeder), E. sinmuans (Smith), and the dubious £. alcocki (Misra) as its constituent species. The last two groups contain Tviakis fehl- manni Springer and T. attenuata Garrick; these are not given genus-group names because of insufficient evidence on the generic relationships of their species. The problem of separating the restricted genus Tviakis from Mustelus Linck is discussed, but no solution is seen at present and the two genera are considered as one unit for comparison with Hemutriakis. A terminology for head morphology, nictitating lower eyelid structure, dentition, vertebral groups, and fin morphology is proposed for use with carcharhinids. LITERATURE CITED APPLEGATE, SHELTON P. 1965. Tooth terminology and variation in sharks with special reference to the sand shark, Carcharias taurus Rafinesque. Los Angeles County Museum of Natural History, Contributions in Science, no. 86, pp. 1-18, fig. 1-5. 94 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. 1967. A survey of shark hard parts. In Sharks, Skates, and Rays, edited by Perry W. Gilbert, Robert F. Mathewson, and David P. Rall. Johns Hopkins Press, Baltimore, Maryland. Part I, no. 2, pp. 37-67, figs. 1-7, pls. 14. BicrELow, Henry B., AND WILLIAM C. SCHROEDER 1940. Sharks of the genus Mustelus in the Western Atlantic. Proceedings of the Boston Society of Natural History, vol. 41, no. 8, pp. 417-438, pls. 14-19. 1944. New sharks from the Western North Atlantic. Proceedings of the New England Zoological Club, vol. 23, pp. 21-36, fig. 1, pls. 7-10. 1948. Sharks. Fishes of the Western North Atlantic. Memoir 1, Sears Foundation for Marine Research, Part 1, Vol. 1, pp. 59-576, figs. 6-106. BouLenGER, G. A. 1902. Description of a new South-African galeid selachian. Annals and Magazine of Natural History, ser. 7, vol. 10, pp. 51-52, pl. 4. CHEN, JoHNSON T. F. 1963. A review of the sharks of Taiwan. Department of Biology, College of Science, Tunghai University. Biological Bulletin 19, Ichthyological Series no. 1, pp. 1-102, figs. 1-28. D’ AUBREY, JEANNETTE D. 1964. Preliminary guide to the sharks found off the east coast of South Africa. South African Association for Marine Biological Research, Oceanographic Research Institute. Investigational Report no. 8, pp. 1-95, figs. 1-22, pls. 1-28. De BueEN, FERNANDO 1950. El tiburon vitaminico de la costa Uruguaya, Galeorhinus vitaminicus nov. sp., y algunas consideraciones generales sobre su Biologia. Servicio Oceanografico y de pesca, Montevideo, Uruguay. Publicaciones Cientificas, no. 4, Contri- buciones a la Ictiologia, pp. 152-162, figs. 1-2. Fow Ler, Henry W. 1929. A list of the sharks and rays of the Pacific Ocean. Proceedings of the Fourth Pacific Science Congress, Java, 1929, pp. 481-508. 1941. Contributions to the biology of the Philippine Archipelago and adjacent regions. The fishes of the groups Elasmobranchii, Holocephali, Isospondyli, and Ostariophysi obtained by the United States Bureau of Fisheries Steamer “Albatross” in 1907 to 1910, chiefly in the Philippine Islands and adjacent seas. Bulletin of the United States National Museum, no. 100, vol. 13, pp. i-x, 1-879, figs. 1-30. GARMAN, SAMUEL 1913. The Plagiostoma. Memoirs of the Museum of Comparative Zoology at Harvard College, vol. 36, pp. i-xiii, 1-515, pls. 1-77. Garrick, J. A. F. 1954. Studies on New Zealand Elasmobranchii. Part III. A new species of Triakis (Selachii) from New Zealand. Transactions of the Royal Society of New Zealand, vol. 82, part 3, pp. 695-702, figs. 1-2. 1967. A broad view of Carcharhinus species, their systematics and distribution. In Sharks, Skates and Rays, edited by Perry W. Gilbert, Robert F. Mathewson, and David P. Rall. Johns Hopkins Press, Baltimore, Maryland. Part I, no. 5, pp. 85-91. Garrick, J. A. F., anp LEONARD P. SCHULTZ 1963. A guide to the kinds of potentially dangerous sharks. In Sharks and Survival, edited by Perry W. Gilbert. D. C. Heath and Company, Boston. Pp. 3-60, figs. 1-33. Vor. XXXVIII] COMPAGNO: SYSTEMATICS OF GENUS HEMITRIAKIS 95 GEGENBAUR, CARL 1872. Untersuchungen zur Vergleichenden Anatomie der Wirbelthiere. Drittes Heft. Das Kopfskelet der Selachier, ein Beitrag zur Erkenntnis der Genese des Kopfskeletes der Wirbelthiere. Wilhelm Engelmann, Leipzig. Pp. i-x, 1-316, pls. 1-22. GILBERT, PERRY W. 1963. The visual apparatus of sharks. In Sharks and Survival, edited by Perry W. Gilbert. D. C. Heath and Company, Boston. Pp. 283-326, figs. 1-30. GILBERT, PERRY W., AND Mark E. OREN 1964. The selachian nictitans and subocular fold. Copeia, 1964, no. 3, pp. 534-535, fig. 1. Gowar, H. A. F., anp F. M. Mazar 1964. The Elasmobranchs of the North-Western Red Sea. Publications of the Marine Biological Station, Al-Ghardaqa (Red Sea), no. 13, pp. 1-144, figs. 1-81, pls. 1-16. HeErrE, ALBERT W. C. T. 1923. Notes on Philippine sharks. The Philippine Journal of Science, vol. 23, no. 1, pp. 68-73, pl. 1. 1953. Check list of Philippine fishes. United States Fish and Wildlife Service, Research Report no. 20, pp. 1-977. HILcGENpor®, F. 1904. Ein neuer Scyllivm-artiger Haifisch, Proscyllium habereri nov. subgen., n. spec. von Formosa. Sonder-Abdruk aus den Sitzungs-Berichten der Gesellschaft naturforschender, Jahrgang 1904, no. 2, pp. 39-41. HoLMcGrREN, NILs 1941. Studies on the head in fishes. Embryological, morphological, and phylogenetical researches. Part I]: Comparative anatomy of the adult selachian skull, with remarks on the dorsal fins in sharks. Acta Zoologica, vol. 22, part 1, pp. 1-100, figs. 1-74. Kato, SUSUMU 1968. Triakis acutipinna (Galeoidea, Triakidae), a new species of shark from Eucador. Copeia, 1968, no. 2, pp. 319-325, figs. 1-2. Kato, SUSUMU, STEWART SPRINGER, AND Mary H. WAGNER 1967. Field guide to Eastern Pacific and Hawaiian sharks. United States Fish and Wildlife Service, Bureau of Commercial Fisheries, Circular 271, pp. 1-47, figs. 1-75. LinpBeErG, G. U., anp M. I. LEGEZA 1959. Fishes of the Sea of Japan and the adjacent areas of the Sea of Okhotsk and the Yellow Sea. Part 1. Amphioxi, Petromyzones, Myxini, Elasmobranchii, Holocephali. Academy of Sciences of the Union of Soviet Socialist Republics. Keys to the Fauna of the USSR, no. 68, pp. 1-207, figs. 1-108. Translation by Israel Program for Scientific Translations, Jerusalem, 1967. McCoy, FREDERICK 1885. Prodromus of the zoology of Victoria. Melbourne, Australia. Decades 1-10, pls. 1-100, pages not numbered. Misra, K. S. 1950. On a new species of scyliorhinid fish from Andaman Sea, Bay of Bengal. Journal of the Zoological Society of India, vol. 2, pp. 87-90, pl. 1. MrvosI, Y. 1939. Description of three new species of elasmobranchiate fishes collected at Hyuga 96 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Nada, Japan. Bulletin of the Biogeographical Society of Tokyo, vol. 9, pp. 91-97, figs. 1-3. MULLER, JOHANNES, AND F. G. J. HENLE 1841. Systematische Beschreibung der Plagiostomen. Berlin. Pp. i-xxii, 1-200, pls. 1-60. Norman, J. R. 1939. Fishes. The John Murray Expedition, 1933-34, Scientific Reports. British Museum (Natural History). Vol. 7, no. 1, pp. 1-116, figs. 1-41. RIpDEWoop, W. G. 1921. On the calcification of the vertebral centra in sharks and rays. Philosophical Transactions of the Royal Society of London, Ser. B., Zoology, vol. 210, pp. 311-407, figs. 1-38. ScuHmpT, P. J. 1928. On a rare Japanese shark, Calliscyllium venustum Tanaka. Comptes Rendus de l’Académie des Sciences de ?URSS, 1928, pp. 65-67, figs. 1-3. 1929. On two rare Japanese sharks Proscyllium habereri Hilgendorf and A pristurus macrorhynchus Tanaka. Comptes Rendus de l’Académie des Sciences de VURSS, 1930, pp. 627-631, figs. 1-2. SmitH, Hucu M. 1913. Description of a new carcharioid shark from the Sulu Archipelago. Proceedings of the United States National Museum, vol. 45, pp. 599-600, figs. 1-3, pl. 47. Smira, J. L. B. 1957a. A new shark from South Africa. South African Journal of Science, vol. 53, no. 10, pp. 261-264, figs. 1-2. 1957b. A new shark from Zanzibar, with notes on Galeorhinus Blainville. Annals and Magazine of Natural History, ser. 12, vol. 10, pp. 585-592, figs. 1-2, pls. 18-19. 1957c. A preliminary survey of the scylliogaleid dogfishes of South Africa. South African Journal of Science, vol. 53, no. 14, pp. 353-359, figs. 1-2. SPRINGER, STEWART 1968. Triakis fehlmanni, a new shark from the coast of Somalia. Proceedings of the Biological Society of Washington, vol. 81, pp. 613-624, figs. 1-5. SPRINGER, VICTOR G. 1964. A revision of the carcharhinid shark genera Scoliodon, Loxodon, and Rhizoprion- odon. Proceedings of the United States National Museum, vol. 115, no. 3493, pp. 559-632, figs. 1-14, pls. 1-2. SPRINGER, Victor G., AND J. A. F. GARRICK 1964. A survey of vertebral numbers in sharks. Proceedings of the United States National Museum, vol. 116, no. 3496, pp. 73-96, pl. 1. STRASBURG, DonaLp W. 1963. The diet and dentition of Jststius braziliensis, with remarks on tooth replacement in other sharks. Copeia, 1963, no. 1, pp. 33-40, figs. 1-5. TANAKA, SHIGEHO 1912. Figures and descriptions of the Fishes of Japan, including Riukiu Islands, Bonin Islands, Formosa, Kurile Islands, Korea, and Southern Sakhalin. Tokyo. Vol. 10, pp. 165-186, pls. 41-50. TANG, D. S. 1934. The elasmobranchiate fishes of Amoy. The Natural Science Bulletin of the University of Amoy, vol. 1, no. 1, pp. 29-111, figs. 1-22. Wuitr, E. GRACE 1937. Interrelationships of the elasmobranchs with a key to the order Galea. Bulletin Vor. XXXVIII] COMPAGNO: SYSTEMATICS OF GENUS HEMITRIAKIS 97 of the American Museum of Natural History, vol. 74, no. 2, pp. 25-138, figs. 1-66, pls. 1-51. WHITLEY, GILBERT P. 1943a. Ichthyological notes and illustrations (Part 2). The Australian Zoologist, vol. 10, part 2, pp. 167-187, figs. 1-10. 1943b. A new Australian shark. Records of the South Australian Museum, vol. 7, no. 4, pp. 397-399. 1944. New sharks and fishes from Western Australia. The Australian Zoologist, vol. 10, part 3, pp. 252-273, figs. 1-6. 1948. New sharks and fishes from Western Australia. Part 4. The Australian Zoologist, vol. 11, part 3, pp. 259-276, figs. 1-7, pls. 24-25. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Festschrift for George Sprague Myers Vol. XXXVITI, No. 5. pp. 99-103; 1 fig. December 31, 1970 NOTES ON THE NATURAL HISTORY OF SNIPE EELS ey Giles W. Mead and Sylvia A. Earle Harvard University One cannot fail to be impressed by the adaptations for midwater life de- veloped by the mesopelagic eels, an assemblage doubtless derived from a benthic ancestor. In most the body has become far more attenuated than that of the most elongate of their benthic relatives, the terminal part of many being fila- mentous and apparently composed of little more than minute and poorly ossified vertebrae covered by thin skin and supporting fine and hair-like fin rays. This attenuation is accomplished by an increase in number of vertebrae rather than an increase in the length of each. An apparently intact specimen of Nemuichthys taken during the International Indian Ocean Expedition of 1964 had 670 verte- brae—certainly a record number among the vertebrates. Equally extreme are specializations in mouth parts. The teeth, for example, vary vastly in form, number and position. The more extreme genera such as Nemichthys, Labichthys, and Avocettina are of concern here. All have greatly prolonged jaws (fig. 1b) that bear numer- ous small teeth laterally as well as dorsally or ventrally (fig. 1d-f), and are usually tipped by flattened bony pads that bear teeth or rugosities on all sides. The jaw teeth are arranged in chevron-shaped consecutive series or, in others, in quincunx. The two halves of the lower jaw are loosely conjoined laterally for most of the length of the mandible. The principal part of the upper jaw, in [99] 100 CALIFORNIA ACADEMY OF SCIENCES | Proc. 4TH SER. NSS 1ocm YRS Vor. XXXVIIT] MEAD AND EARLE: SNIPE EELS 101 contrast to other fishes, is composed of the vomer; and the biting elements of other lower fishes, the maxillae, are reduced to lateral struts that support the base of the prolonged vomerine bar (Beebe and Crane, 1937). The positional relationship of these jaws has been particularly enigmatic, for they diverge for- ward from the gape so that their tips, and often as much as half of the total length of the jaws, cannot be brought into contact with each other when the mouth is as far closed as it can be. These fishes can but partially close their mouths, yet the distal ends of their jaws, that cannot possibly be brought into contact with each other, bear thousands of small chisel-shaped posteriorly in- clined teeth (fig. 1d-e) reminiscent of the shagreen of an elasmobranch. What can be the function of such a structure? That these jaws are used to funnel microplankton toward the mouth as the eel swims through the water seems unlikely. Lateral movement of the prey by but a millimeter or two would take it beyond the grasp of the predator. To feed in this way, structural adaptation similar to that in the herrings would be more in order. It has also been suggested that these prolonged jaws simply provide greater surface area, and might be considered adaptations for flotation. While we would consider the attenuated but fin-bearing shape of the body the result of selection toward greater surface area that serves the interests of flotation, we are reluctant to so consider the development of well ossified structures richly endowed with small but dense teeth. Such a beak would also seem to be the antithesis of a structure developed in aid of streamlining or locomotion. Nichols and Murphy (1944) repeated a report by Mowbray (1922) of a snapper cap- tured in Bermuda with a 265 mm. representative of Nemichthys scolopaceus attached by its slender jaws to the posterior margin of the snapper’s caudal fin. Mowbray concluded, ‘‘The specimen being taken in this way gives good reason to believe that grasping the tails of fishes is the function of the divergent man- dibles of these eels.” We can suggest an alternative function for these diverging and nonocclusable jaws, a suggestion emanating from observations made at mid-depths from the late D.S.R.V. Alvin and catches made concurrently by a supporting vessel, R.V. Gosnold, both of the Woods Hole Oceanographic Institution. These dives were made between October 2 and 6, 1967, in Slope Water of the western North Atlantic in an attempt to observe visually certain sound-scattering targets at Ficure 1. a. Snipe eel, Nemichthys scolopaceus, in typical vertical position as observed from the D.S.R.V. Alvin. b. Vertically oriented specimen of N. scolopaceus and sergestid shrimp, Sergestes (Sergestes) arcticus, drawn from specimens taken by R.V. Gosnold concurrent with D.S.R.V. Alvin observations. c. Distal portion of antenna of S. (S.) arcticus. d. Tooth from upper jaw of N. scolopaceus. e. Inner surface of upper jaw of N. scolopaceus. {. Inner surface of lower jaw of NV. scolopaceus. 102 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. mid-depth (see Backus e¢ al., 1968). Concurrent with these dives, R.V. Gosnold fished similar depths with a 10-ft. Isaacs Kidd Midwater Trawl. Observers aboard Alvin frequently saw snipe eels (Nemichthys) at depths below 300 m. and confirmed the observations of others that these eels are usually oriented vertically in the water, motionless or but slightly undulating, and usually with their divergent jaws directed upward. Among the other more spectacular animals seen at comparable depths were relatively large sergestid decapods. These too were often suspended vertically in the water, their bright orange-red stomachs and organs of Pesta prominent, the short pleopods beating furiously, and their long antennae extending upward and outward away from the body and then turning abruptly to follow a course parallel to the axis of the body to a point considerably below the tail. Neither eels nor sergestids appeared to be disturbed by the lights of the submarine. Both sergestids and snipe eels were caught by the nets of the Gosnold. The eels belong to Nemichthys scolopaceus Richardson, 1848, and the sergestids, kindly identified for us by Mr. Peter Foxton of the National Institute of Ocean- ography, Godalming, belong to Sergestes (Sergestes) arcticus Kroyer, 1855. Representatives of Nemichthys, as usual, were present in the catch with beaks entangled in everything present, living or not. Several were hanging by their beaks from the upper part of the netting as the trawl was raised above the water, the red stomachs of ingested sergestids visible through the semitransparent stomachs and body walls. This material was returned to Woods Hole and to the Museum of Comparative Zoology, Harvard University, for study and is deposited in the latter institution. The stomach contents of about 160 specimens of Nemichthys were examined. In addition to the Gosnold collection these included others variously collected in the western North Atlantic and those from the Indian Ocean and off central Chile that were caught during Cruises VI and XIII, respectively, of R.V. Anton Bruun. Most stomachs were empty. Those which were not, contained crustacean remains exclusively. An examination of the specimens of Sergestes which were available and published accounts of others (Burkenroad, 1934, 1937; Foxton, 1969; Hardy, 1956) revealed the complexity of the prolonged antennae with their multiple sensory hairs, structures admirably suited to aid in flotation. We believe that the function of the beak of the snipe eels can be added to the list of features in which these eels are unique among vertebrate animals, for we suggest here that these animals feed by entanglement. Given the vast extent and thread-like nature of some appendages of many mesopelagic crustaceans and the set and structure of snipe eel dentition, the evolution of structures adapted for the feeding of one on the other seems reasonable. The antennae of a sergestid if brushed across the bed of teeth of a snipe eel would almost certainly become entangled, and struggle by the prey would only worsen its plight, shorten the VoL. XX XVIII] MEAD AND EARLE: SNIPE EELS 103 distance between shrimp and fish; and ultimately bring the prey within that more posterior part of the jaws capable of crushing and swallowing movements. Such a feeding mechanism is consistent with present concepts of midwater ecology. Food in the deep ocean is scarce and energy precious. Hovering and darting, or luring types of predation tend to replace the roving activities more prevalent near the surface. Intake per unit of energy expended must be high if a predator is to survive. What finer an example of adaptation to these con- ditions can there be than that of these eels: hanging effortlessly with flotation facilitated through attenuation of the body, and with jaws covered by myriads of denticles exquisitely designed to entangle the appendages of passing crustacea, be they moving laterally or, with some, rising or descending as a part of their daily routine. ACKNOWLEDGMENTS This note owes its existence to an invitation to one of us to dive aboard Alvin and we thus record our grateful appreciation to the Woods Hole Ocean- ographic Institution and especially to Dr. Richard H. Backus of that institution for that opportunity. In addition to identifying the sergestids, Mr. Peter Foxton, National Institute of Oceanography, Godalming, reviewed the manuscript, as did Drs. Backus, R. L. Haedrich, and J. E. Craddock of the Woods Hole Ocean- ographic Institution. This paper is contribution number 2351 from the Woods Hole Oceanographic Institution. REFERENCES Backus, R. H., J. E. Crappock, R. L. Harpricu, D. L. SHores, J. M. Tear, A. S. WING, G. W. Meap, anp W. D. CLark 1968. Ceratoscopelus maderensis: peculiar sound-scattering layer identified with this myctophid fish. Science, no. 160, pp. 991-993. BEEBE, W., AND J. CRANE 1937. Deep-sea fishes of the Bermuda Oceanographic Expeditions. Family Nemichthy- idae. Zoologica (N.Y.), vol. 22, no. 4, pp. 349-383. BuRKENROAD, M. D. 1934. The Penaeidae of Louisiana with a discussion of their world relationships. Bulletin of the American Museum of Natural History, vol. 68, no. 2, pp. 61-143. 1937. The Templeton Crocker Expedition XII. Sergestidae (Crustacea, Decapoda) from the Lower California region, with descriptions of two new species and some remarks on the organs of Pesta. Zoologica (N.Y.), vol. 22, no. 4, pp. 315-329. Foxton, P. 1969. The morphology of the antennal flagellum of certain of the Penaeidae (Decapoda, Natantia). Crustaceana, vol. 16, no. 2, pp. 33-42, 1 pl. Harpy, A. C. 1956. The open sea. Boston. xv + 335 pages, 18 pls. Moweray, L. L. 1922. Habit note on snipe eel. Copeia no. 108, page 49. Nicuots, J. T., anp R. C. Murpuy 1944. A collection of fishes from the Panama Bight, Pacific Ocean. Bulletin of the American Museum of Natural History, vol. 83, no. 4, pp. 217-260. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Festschrift for George Sprague Myers Vol. XXXVIII, No. 6, pp. 105-130; 7 figs.; 11 tables | December 31, 1970 THE ZOOGEOGRAPHY OF THE HERPETOFAUNA OF THE PHILIPPINE ISLANDS, A FRINGING ARCHIPELAGO By Walter C. Brown and Angel C. Alcala Division of Systematic Biology, Stanford University and Menlo College, Menlo Park, California and Department of Biology, Silliman University, Philippines INTRODUCTION Inger, in his essay on the zoogeography of the Philippine amphibia (1954, pp. 448-510), presented the first major distributional paper for any part of the herpetofauna since Taylor’s essay (1928). The first part of Inger’s paper is concerned with the geological history of the Philippines, and the origins and degree of endemism exhibited by the amphibian fauna. Secondly, he discusses the pathways of entry into the Philippines in terms of the location of the nearest relatives and possible time of entry into the Philippines. He notes, for example, that the present distribution of the genus Platymantis (replaces Cornufer, Zweifel, 1967) suggests two speciation centers, one in the New Guinea-Solomons region and one in the northern Philippines; but by analogy, in comparison with some other amphibians, suggests a Papuan origin and subsequent dispersal into the Philippines. Inger therefore regards these two present centers as peripheral isolated concentrations of a once more widely distributed genus (1954, pp. 494, [105] 106 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47H SER. 497). He suggests that the bulk of the amphibians entered the Philippines by 2 major routes, the Palawan or Sulu-Mindanao routes. He discusses relative time of entry of different components of the amphibian fauna primarily in terms of extent of endemism and distance from areas occupied by presumed nearest relatives. Within the Philippines, Inger recognizes only 2 somewhat doubtful zoogeographic subdivisions. In his discussion of dispersal (pp. 475-484), he notes that both dispersal by way of earlier land connections and over-water dispersal must be considered, and he also notes, in general descriptive terms, possible routes within the Philippines. Leviton (1963) provides the most recent discussion of zoogeography of the terrestrial snake fauna of the Philippine archipelago. His discussion is primarily concerned with extraterritorial origins, time of entry and endemism, present distributions, and the taxonomic relationships of species within the Philippines. These are considered in terms of past changes in island configurations, and probable internal pathways. He states (p. 377), contrary to Inger’s views relative to the dispersal of the amphibians (Inger, 1954, p. 484), that the present distribution of the snakes can, for the most part, be explained on the basis of former land connections. He recognizes 5 faunal (serpentine) subregions within the archipelago at the present time. Both authors very ably discuss the present distribution of the faunal element with which they are concerned in terms of traditional concepts of extraterritorial origins, pathways of entry and internal dispersal as governed by probable geological changes, time of entry, and means of dispersal. Darlington (1957, pp. 476-541) discusses the Philippine ichthyological and herpetological faunas in the more general context of distributions on fringing archipelagoes. Immigrant patterns of distribution, where the species are distrib- uted along the migration route with dropouts occurring linearly as determined by distance and relative dispersal abilities, are, he believes, the primary patterns exhibited in fringing archipelagoes. This basic pattern is modified for older relict groups by concentration of species on distal or proximal islands within the archipelago (p. 533). MacArthur and Wilson (1963) propose the hypothesis that the number of species on an island represents a balance between number of species reaching the island and number of species becoming extinct per unit of time. They point out that a number of interacting variables will determine the point at which these 2 curves intersect. These include distance from source of immigrants, the species pool of immigrants, area of island concerned, or some other limiting factor such as unfavorable climate. They further propose that in time secondary radia- tion centers should increase with distance of islands from the major source of the fauna, when corrections are made for area or other limiting variables. They also note that the number of species decreases more rapidly for large than for small islands with increasing distance from source of colonization. VoL. XXXVIII] BROWN AND ALCALA: HERPETOFAUNA OF PHILIPPINES 107 ( = Mountain Peak ) 50 100 Miles °° O 50100 200 Kilometers Entryways and proposed 1o} =i = =< = ° Q oe = a oD of <— é fo! oa \o i=) _ | eo ie) Oo So | uw =) 2) = ne! vo Q. ~ vo » Ss o =) Be = oe = u ® BS a) & a = fe) te o 8 c OX = 3 = Q. [2] fey oa os = c f= oO S aa 4 8 2 c=) fe) ; 8, = oO © Lot a = : = = 8 3) » & x2 Ae n = = 2 2? oO = = oS a a ee Ea . rt 3} = ° Fg Boy ee ° 2 s gS v =) fey ae ye a) PROBLEMS AND METHODS Recent intensive exploration of the herpetofaunas of 5 Philippine islands, Palawan, Mindanao, Bohol, Negros, and Mindoro (fig. la), makes possible more critical examination of many of the zoogeographic hypotheses suggested by previous authors. Factors which we believe make this possible are: 1) intensive sampling techniques which provide more accurate estimates of species-diversity ; 2) choice of islands from proximal, distal, and intermediate regions of the archipelago (fig. la); 3) range in size from 5,000 to 95,000 square kilometers 108 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. TaBLe 1. Intensively explored areas on the five islands included in the recent survey. Altitude Island Area Mountain Region (meters) Island (sq. km.) Exploration Date Cuernos de Negros 1,903 Negros 12,700 March—May (southern part) 1958 (about 7 weeks) Mt. Canlaon 2,463 Negros " March—April (northern part) 1962 (about 4 weeks) Dapitan Peak 2,199 Mindanao 94,600 March—May (Zamboanga Peninsula) 1959 (about 6 weeks) Thumb Peak 1,286 Palawan 11,800 April—May (central part) 1961 (about 714 weeks) Mt. Halcon 2,580 Mindoro 9,750 April—May (northern part) 1963 (about 4 weeks) Sagungan Mountain 870 Bohol 4,100 April—May (southeastern part) 1962 (about 4 weeks) (table 1); 4) choice of islands which encompass sufficiently large areas of original and/or secondary lowland forest as to make negligible differences in diversity which might be due to major differences in the dominant type of plant community (see Brown and Alcala, 1964). The techniques stressed intensive sampling of arboreal, surface, and sub- terranean strata in the lowland forest whenever possible, as well as selected mountains. The expeditions to each of the 6 mountain areas on the 5 islands were carried out by crews of 8 to 10 men over 4 to 7’ week periods (table 1). TasLe 2. Number of species recorded for the islands included in this study. The number in parentheses is the number of species belonging to the group of 23 widely distributed species associated with man’s economy or beach communities. The number in brackets is the number of relict species. Palawan Mindanao Bohol Negros Mindoro Leyte Luzon Caecilians 1 1 Frogs 22(4)[1]- 34(4)[3] 2 4] 16(4)[4] 12(4)[2] 16(3)[3] 21(4)[6] Lizards 23M) SAGO) TG Nawal, 33a eae ae) 2 34(8)[ 7] Snakes 8((D Il@l| BOC) 2 1) 2 Ml QOCON2 1) TOC iay) TSA) So ))1 2 | Total 78 (24)[2] 125 (23)[11] 72(22)[9] 78 (24)[10] 58 (23)[5] 94 (21)[15] VoL. XXXVIIT] TABLE 3. BROWN AND ALCALA: and Luzon islands. HERPETOFAUNA OF PHILIPPINES 109 Amphibians known from Palawan, Mindanao, Bohol, Negros, Mindoro, Leyte, Palawan Mindanao Boikol Negros Mindoro Leyte Luzon 1. Ichthyophis monochrous x x 2. Barbourula busuangensis x 3. Ansonia mcgregori x 4. Ansonia mulleri x 5. Bufo biporcatus x 6. Leptobrachium hasselti x x x x 7. Megrophrys monticola x x x x 8. Platymantis cornutus x 9. Platymantis corrugatus x x x x x x 10. Platymantis dorsalis x x x x x x 11. Platymantis guentheri x x x x x 12. Platymantis hazelae x x 13. Platymantis ingeri x 14. Platymantis subterrestris x 15. Micrixalus mariae x 16. Ooeidozyga diminutiva x 17. Ooeidozyga laevis x x x x x x x 18. Rana cancrivora x x x x x x x 19. Rana erythraea x 20. Rana everetti x x x x 21. Rana leytensis x x x x 22. Rana limnocharis x x x x x 23. Rana magna x x x x x x x 24. Rana microdisca x< x 25. Rana nicobariensis x 26. Rana sanguinea < 27. Rana signata x x x x x x 28. Rana woodworthi xX 29. Staurois natator =< x x x 30. Philautus acutirostris x 31. Philautus bimaculatus S< SK 32. Philautus leitensis x x 33. Philautus longicrus x< 34. Philautus pictus x 35. Philautus schmackeri x 36. Philautus spinosus x x 37. Philautus williamsi x 38. Rhacophorus appendiculatus x x x 39. Rhacophorus everetti x 40. Rhacophorus emembranatus x 41. Rhacophorus leucomystax x x x x x x x 42. Rhacophorus lissobrachius x x 43. Rhacophorus pardalis x x x x 44. Rhacophorus surdus x x 45. Pelophryne albotaeniata SK 110 CALIFORNIA ACADEMY OF SCIENCES [| Proc. 4TH SER. TABLE 3. Continued. Palawan Mindanao Bohol Negros Mindoro Leyte Luzon 46. Pelophryne brevipes . Pelophryne lighti . Chaperina fusca x . Kalophrynus pleurostigma Kaloula baleata SK Kaloula conjuncta x Kaloula picta Xx Kaloula rigida Oreophryne annulata < aS a x aS ie) S Ne} x xX xX X x x fa 025 Se x x x x X X The belief that our sampling techniques for these selected mountain areas has provided a realistic estimate of the number of species present in the area is based upon experience in the Cuernos de Negros area in southern Negros Island. The initial survey expedition there in 1958, using the sampling methods noted above, resulted in the recording of 67 herpetofaunal species (Brown and Alcala, 1961, p. 631). Although extensive resampling of this area has occurred during the interim of several years, in connection with our population and other ecological studies, only 7 additional species (Hemiphyllodactylus typus, Lepidodactylus christiani, L. lugubris, Luperosaurus cumingi, Brachymeles tridactylus, Typhlops cumingt, and Boiga angulata) have been found in the southern Negros area. The 4 remaining species in the present list for Negros (tables 3, 4 and 5) are known only from the northern part of the island. Thus, even though Bohol, Palawan, and Mindoro are not yet widely explored, the presently available data on their herpetofaunal communities, based on past records and on our intensive sampling of selected mountain areas, are believed to be sufficient to realistically approximate their relative positions in terms of diversity. Utilizing primarily the data from the 6 recently intensively explored mountain areas on 5 islands (table 1), supplemented by available lists of species, largely based on data assembled by earlier explorers, for Leyte and Luzon islands as well as total distributional data for a few genera, we propose to examine: 1) the nature of the relict patterns for some of the older herpetofaunal elements; 2) the probability of secondary radiation zones; 3) the relative importance of probable internal migration routes, using Sorenson’s index of similiarity; 4) the probable importance of marine barriers in effecting the present distribution patterns; 5) the relation of island size and distance to diversity of species. RESULTS DIVERSITY OF SPECIES A total of 9,000+ herpetofaunal specimens, ranging from about 1,000 for Mindoro Island to 2,500 for Negros Island, were collected during our recent expeditions, Classification of the collected material when added to earlier rec- Vout. XXXVIII] BROWN AND ALCALA: HERPETOFAUNA OF PHILIPPINES 111 ords, reveals 197 species: 1 caecilian, 49 frogs, 82 lizards, and 66 snakes for these five islands (tables 2-5). Although the new records ranged from 61 for Bohol to 5 for Mindoro, only 14 were new species or species not recorded from the Philippines prior to our intensive sampling. Most of these 14 were from Palawan and Mindanao, the islands adjacent to Borneo. The number of species of snakes and frogs recognized from Luzon and Leyte islands are based primarily upon Inger’s review of the Philippine amphibia (1954) and Leviton’s review of the snakes (1963). The distribution of the species of the genus Calamaria, however, is from Inger and Marx (1965). The list of lizards for Luzon is based primarily on the earlier records of Taylor (1922a, 1922b, 1922c, 1923 and 1925) with a few recent additions by the present authors. The inclusion of Luzon adds 23 more species (5 amphibians, 10 lizards, and 8 snakes) to the 197, making a total of 220 species (tables 3-5). NATIVE FAUNA Twenty-three of the 220 species (4 frogs, 11 lizards, and 8 snakes) are re- garded as probably nonnative; that is, as possibly introduced or at least re- introduced by man. We do not presume that this list includes all species which have at any time been introduced by man, intentionally or otherwise. It probably, however, does include most of these species which are readily, perhaps often accidentally, transported from island to island. To be included in this category, the species must meet these criteria: 1) occur on at least 4 of the 5 islands included in our study; 2) exhibit no subspeciation except for the Palawan populations in some instances; 3) be associated with man’s habitations, and cultivated lands, or with other lowland beach communities (Brown and Alcala, 1964); 4) be widespread in Borneo and other adjacent areas. It is interesting to note (table 2) that the number of species of this nonnative group, whether we are concerned with frogs, lizards, or snakes, is essentially the same for each of the 7 islands irrespective of distance from entry-point or area of island. This we interpret as further evidence supporting their classification in this category. The 23 species include: Amphibians: Lizards: Snakes: Ooeidozyga laevis Cosymbotus platyurus Typhlops braminae Rana cancrivora Gehyra mutilata Python reticulatus Rana limnocharis Gekko gecko Ahaetulla prasina Rhacophorus leucomystax Hemidactylus frenatus Dendrelaphis pictus Hemiphyllodactylus typus Chrysopelea paradisi Draco volans Elaphe erythrura Dasia smaragdina Lycodon aulicus Emoia atrocostata Psammodynastes pulverulentus Lygosoma (Leiolopisma) quadrivittatum Mabuya multifasciata Mabuya multicarinata 112 CALIFORNIA ACADEMY OF SCIENCES [| Proc. 4TH SER. Taste 4. Lizards known from Palawan, Mindanao, Bohol, Negros, Mindoro, and Luzon islands. Palawan Mindanao Bohol Negros Mindoro Luzon 1. Cosymbotus platyurus x x x x x x 2. Cyrtodactylus agusanensis x 3. Cyrtodactylus annulatus x x 4. Cyrtodactylus philip pinicus x x x 5. Cyrtodactylus redimiculus x 6. Gehyra mutilata < x x Xx x x 7. Gekko athymus x 8. Gekko gecko x x x x x x 9. Gekko mindorensis x 10. Gekko monarchus x x x 11. Gekko palawanensis x< 12. Hemidactylus frenatus x x x x x x 13. Hemidactylus garnoti x x 14. Hemidactylus luzonensis x 15. Hemiphyllodactylus typus x x x x 16. Lepidodactylus aureolineatus x x 17. Lepidodactylus christiani x 18. Lepidodactylus herrei x 19. Lepidodactylus lugubris x 20. Lepidodactylus naujanensis x 21. Lepidodactylus planicaudus x< x 22. Luperosaurus cumingi x 23. Luperosaurus joloensis x 24. Perochirus ateles x 25. Pseudogekko compressicor pus << < x 26. Pseudogekko brevipes x< x 27. Ptychozoon intermedia x 28. Calotes cristatellus x x x 29. Calotes marmoratus x< x x 30. Draco bimaculatus x 31. Draco everetti x 32. Draco mindanensis SK 33. Draco ornatus x x 34. Draco quadrisi x 35. Draco rizali x x 36. Draco volans x x x x x x 37. Gonyocephalus interruptus x 38. Gonyocephalus semperi x SZ 39. Gonyocephalus sophiae x SK 40. Hydrosaurus pustulosus x x x x x 41. Varanus salvator x x x x x 42. Dibamus argenteus sé x< S< 43. Brachymeles bonitae x * 44. Brachymeles elerae x 45. Brachymeles gracilis x x x x x 46. Brachymeles pathfinderi x Vou. XXXVIII] BROWN AND ALCALA: HERPETOFAUNA OF PHILIPPINES | 113 TABLE 4. Continued. Palawan Mindanao Bohol Negros Mindoro Luzon 47. Brachymeles samarensis x 48. Brachymeles schadenbergi < x x 49. Brachymeles talinis xX x 50. Brachymeles tridactylus x 51. Brachymeles wrighti < 52. Brachymeles hilong x 53. Dasia griffini x 54. Dasia olivaceum x x 55. Dasia smaragdina x x x* x x x 56. Emoia atrocostata x x x x xK 57. Emoia caeruleocauda x 58. Emoia ruficauda x 59. Lygosoma (Leiolopisma) auriculatum x x 60. Lygosoma (Leiolopisma) pulchellum x x x x 61. Lygosoma (Leiolopisma) quadrivittatum x x < 62. Lygosoma (Leiolopisma) rabori x 63. Lygosoma (Leiolopisma) semperi < 64. Lygosoma (Leiolopisma) subvittatum x< 65. Lygosoma (Leiolopisma) vulcanium x 66. Lygosoma (Leiolopisma) zamboangensis x 67. Lygosoma (Lygosoma) chalcides x< 68. Lygosoma (Sphenomorphus) acutum x x 69. Lygosoma (Sphenomorphus) arborens XK 70. Lygosoma (Sphenomorphus) atrigularis x 71. Lygosoma (Sphenomorphus) coxi x< < 72. Lygosoma (Sphenomorphus) decipiens SK 73. Lygosoma (Sphenomorphus) diwati x 74. Lygosoma (Sphenomorphus) fasciatum x XK 75. Lygosoma (Sphenomorphus) jagori < x ~~ x x 76. Lygosoma (Sphenomorphus) luzonensis xX 77. Lygosoma (Sphenomorphus) mindanensis x x 78. Lygosoma (Sphenomorphus) palawanensis xX 79. Lygosoma (Sphenomorphus) steerei x x x x x 80. Lygosoma (Sphenomorphus) stejnegeri x 81. Lygosoma (Sphenomorphus) varigatum S< x 82. Lygosoma (Sphenomorphus) wrighti x 83. Lygosoma (Sphenomorphus) sp. SK 84. Mabuya bontocensis x 85. Mabuya multicarinata x x x x x x 86. Mabuya multifasciata x Ss x < x x 87. Otosaurus cumingi x x x x 88. Tropidophorus grayi x x 89. Tropidophorus leucospilos x 90. Tropidophorus misaminus x 91. Tropidophorus partelloi x 92. Tropidophorus sp. x 114 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. Taste 5. Snakes known from Palawan, Mindanao, Bohol, Negros, Mindoro, Leyte, and Luzon islands. Palawan Mindanao Bohol Negros Mindoro Leyte Luzon 1. Typhlops braminae x x x x x x 2. Typhlops cumingi x 3. Typhlops canlaonensis x 4. Typhlops dendrophis x 5. Typhlops jagori x 6. Typhlops longicauda x x 7. Typhlops luzonensis x x 8. Typhlops mindanensis x 9. Typhlops ruber x 10. Typhlops ruficauda x 11. Typhlops rugosa x 12. Xenopeltis unicolor x 13. Python reticulatus x x x x x x x 14. Ahaetulla prasina x x x x >< x 15. Aplopeltura boa x x 16. Calamaria bitorques x 17. Calamaria gervaisi x < x x 18. Calamaria lumbricoidea x x xX 19. Calamaria palawanensis ~< 20. Calamaria virgulata x x 21. Chrysopelea paradisi x x x x x x 22. Cyclocorus lineatus x x x x 23. Dendrelaphis caudolineatus *« x ~< x x x x 24. Dendrelaphis pictus x x x x x x 25. Dryophiops philip pina x x< xx 26. Dryocalamus tristrigatus x< 27. Dryocalamus subannulatus x 28. Elaphe erythrura x x x x x x x 29. Gonyosoma oxycephala x x x x< xX 30. Hologerrhum philippinum x 31. Hurria rynchops x x x xx x 32. Liopeltis philippinus x 33. Liopeltis tricolor x 34. Lycodon aulicus x x x x x x 35. Lycodon dumerili x 36. Lycodon mulleri x x 37. Lycodon subcinctus << 38. Lycodon tesselatus x 39. Myersophis alpetris x 40. Natrix auriculata x x x 41. Natrix chrysarga x 42. Natrix dendrophiops x x x x x 43. Natrix lineata x 44. Natrix spilogaster x 45. Oligodon ancorus x Vor. XXXVIII] BROWN AND ALCALA: HERPETOFAUNA OF PHILIPPINES 115 TABLE 5. Continued. Palawan Mindanao Bohol Negros Mindoro Leyte Luzon 46. Oligodon maculatus 47. Oligodon modestum 48. Oligodon vertebralis S< 49. Opisthotropis alcalai 50. Opisthotropis typica >< 51. Oxyrhabdium leporinum x x x 52. Oxyrhabdium modestum 53. Psammodynastes pulverulentus < 54. Pseudorabdion ater 55. Pseudorabdion mcnamarae 56. Pseudorabdion montanum 57. Pseudorabdion oxycephalum 58. Pseudorabdion taylori x 59. Sibynophis bivittatus x 60. Stegonotus miilleri x x 61. Zaocys carinatus x 62. Zaocys luzonensis x 63. Bioga angulata 64. Bioga cynodon 65. Boiga dendrophila 66. Boiga drapiezi 67. Boiga philippina 68. Calliophis calligaster x x 69. Maticora intestinalis < 70. Naja naja x 71. Ophiophagus hanna x 72. Trimeresurus flavomaculatus 73. Trimeresurus schultzei < 74. Trimeresurus wagleri x x x XS OX OK OX x X X x x x x x X X x xX X XS XK OK OK xX XK OK x X x x OK OK 2K OK KX IS, 0X ON IK x x x x x In other sections of this paper, certain of the indices are determined, both for the total fauna and for the presumed native fauna following the exclusion of these 23 species. ReLIctT PATTERNS AND SECONDARY RADIATION ZONES As noted by Darlington (1957, p. 505) endemism at the specific and sub- specific levels is high, but at the generic level very low for the Philippine herpeto- fauna. It is of interest to examine the distributional patterns of these endemic genera, as well as of a few other genera (the presumed earlier arrivals) which, though not limited to the Philippines, exhibit disrupted distributional patterns outside the Philippines, to determine their fit to typical relict or modified im- migrant patterns as postulated by Darlington (1957, p. 484 ff.). This selection of endemic genera and those with strongly disrupted distributional patterns does [Proc. 4TH SER. CALIFORNIA ACADEMY OF SCIENCES 116 ‘sourddiiyg 94} Ul SajamAyov4g snues pieziT 94} Jo satdeds Jo suizo}jed uoTNqIysIq ‘sourddiiyd 94} Ul SzqupmakyD]g SNUds B01} IY} JO Sotdeds Jo susaqyyed uoTyNqIy4sSIG € Bq $19j@WO|I4OOZ Ol OS O ie ets “a Taner ala G se SON OOl OS O ie ea Seas oc). o °o 9 2 o Xd ® udjId0Q Vv, O(N soJben ap € Seid an x "og aw NyAVIvd “Wd qunul “ht w(SOY93N a ) 9 + g 2 B o « AYN buosly sajawAyIog SIUI/O{ SA/9WKYIOIG SIIIDOID SAJABWAYIOIG JO6saqUaPOYIS Sa/AWAYIOIG YYblUM sajawsAyIosg Mapuly{od sajaWAyIosg 9013/9 SHOWAYIOIG sisuange? sajawAyIosg snj/Ajo0piy, SajawAyIoIg SISUGIOWOS Sa/AaWAYIOIG anjlu0g sajaWwKxYIo/g SIWJaA Sajal/AYIOIg ( 40aq UIOJUNOW =# ) OYOONIW 09|0H iW od0+sg84degnde z 54 siajawoliy OO2 OOlLOS O 0” 9° SAIN OOL OS O soJban ap Stee NwAWIvd Gy “Wd qUinuL oy. SOUOSN aise) o. _ &) a? . Dg WWNVd +g MY oh 8 Oo T\S \ Sey OHOGNIW 9 ) UOd|0H YW °, od o11I10d SINSIILA{GNS SIF{UDWA{O/g SISUB//1]OI SI{UOWAIO/ SN{NUIOI SI{UOWA{O/d (4abus SI{UOWALO/g 9O/AZOY SI{UOWA{D/q 1a4{uanb Ssif{uOWA{O/d SI/OSIOP SI{UOWA{O/qd SN{OBNLIOI SIFUOWAJO/g odose@dene (Dag UlOJUNOW = #) 0 0 ‘¢ GUN ‘? qANOIA Vor. XXXVIII] BROWN AND ALCALA: HERPETOFAUNA OF PHILIPPINES 117 not preclude the possibility that species in other, widely distributed genera may also be relicts of early immigrations, but provides objective criteria for selection. The endemic genera include Barbourula, an amphibian; Luperosaurus, Pseudogekko and Brachymeles, lizards; and Cyclocorus, Hologerrhum, and Oxyrhabdium, snakes. Platymantis in the amphibia and Perochirus in the squamata, though not endemic, exhibit overall disrupted patterns. Barbourula and Perochirus are limited to single species in the Philippines. Barbourula is known only from the Palawan group and Perochirus only from Mindanao and Leyte. This pattern of limitation to proximal islands permits 3 alternative explanations: 1) these genera have very low dispersal ability and did not go beyond these proximal islands; 2) they have reached but failed to establish themselves on more distal islands; 3) they still remain to be found on the other islands. Platymantis, Brachymeles, and Luperosaurus exhibit relict patterns (figs. 2, 3, and 4) of the type which may be interpreted as resulting from the partial extinction of an old widespread fauna (Darlington, 1957, p. 485). Each of these genera includes several species. The genus Platymantis (as noted by Inger, 1954, p. 496 exhibits a concentration of species on Luzon at the distal end of the archipelago, others with distribution limited to one or two scattered islands, and several species which are widespread throughout the Philippines, though none of these species are known from outside of the archipelago. As proposed by MacArthur and Wilson (1963, p. 386) and Inger (1957, p. 496), such a pattern may also be interpreted as due in part to secondary radiation from the distal island of Luzon as indicated by the present distribution of P. hazelae and P. polillensis, very closely related species, not necessarily as wholly due to chance extinctions of once widespread species. The genus Brachymeles, as reviewed by Brown and Rabor (1967), includes such widespread species as B. gracilis and B. schadenbergi, groups of species limited to the proximal or distal islands, and a third very interesting group (the tridactylus-vermis group) which strongly suggests an origin in and radiation from the center of the archipelago. Based on the presumed evolutionary relationships, indicated by the degree of reduction of the limbs and digits, the least specialized members of the group occur in the central islands and those with the greater reduction of limbs and digits in the northern and southern islands. These species include: Digits on Digits on Species Distribution fore limbs hind limbs Brachymeles tridactylus Negros 3 3 Brachymeles cebuensis Cebu 3 2 Brachymeles samarensis Leyte, Samar, and Luzon 2 2 Brachymeles bonitae Luzon and Mindoro 0-1 0-1 Brachymeles vermis Sulus 0 ) [Proc. 4TH SER. CALIFORNIA ACADEMY OF SCIENCES 118 ‘sourddyiyg ey} Ul wnipqnyAKxgQ pure ‘wnys1asoj0 FH ‘sns0I0]IKD eilaUas IYVUS 9Y} JO Satdads ay} JO susa}jed uoIyNqIIysIq ‘“¢ ANNI ‘sourddriyg ay} Ul oyYasopnasg pure SnAnvsSosagnT Bilduas pAeZIT IY} JO Suta}jed uoTyNGIUysIq ‘pf TINO g “Bly y Bly siajawo|!4 OO2 OO! 0S O > & . siajawolid OOZ OO! OS O ( oe SAl!W OO! OS (e) is 2 SiIN OO! OS 0 Ad upjidog soiban ap soiben ap souiend sousand NVAV 1¥d 3 wy | SOY9SN SOY9S3N of Ag Gp ; . ° YNVd AVNWd' 6 ., * 68 8 7 ae ° : OYOGNIW UO9|DH tW “SS XQ UO0D|DH JW "3, Sates! °o. 011110d snasoo Wh{sEpowW WhIpgoysAxow -/SSa/0WOI OyabOpNASdY ®@ wnulsode/ wnipqoys4xO Sadinaiq Oyyab0pNeasg @ wnujdaiyjiyd wnyssabojOH 7 obssb6owW sninoso1a0n7 Ww snjoeuy) sni0y90j9A9 O sisuaojol sninososaan7 @ ( yDaq UIDJUNOW = % ) J/Oujwnga sninososadny 7 snssiu/o sninososeon7 O 0 2 ( 408g UIDJUNO|W =# ) o i g ar r) Ws Vor. XXXVIII] BROWN AND ALCALA: HERPETOFAUNA OF PHILIPPINES 119 TasLe 6. Indices of similarity for amphibians. The index in parentheses is based on the native species after exclusion of common widespread forms associated with man’s economy or beach communities. The index in brackets is based on those species remaining after the relicit species are also excluded. Mindanao Leyte Bohol Negros Mindoro Tuan Palawan 0.429 0.368 0.372 0.263 0.412 0.326 (0.333 ) (0.258) (0.278 ) (0.067 ) (0.231) COs 7/8) [0.364 | [0.296 | [0.323 ] [0.080] [0.261] [0.214] Mindanao 0.600 0.691 0.520 0.476 0.509 (0.588 ) (0.667 ) (0.429) (0.368) (0.426) [0.486 | [0.634 | [0.343 | [0.303 | [0.368 ] Leyte 0.703 0.687 0.643 0.595 (0.645 ) (0.640) (0.571) (0.533) [0.583 | [0.556] [0.500 | [0.476] Bohol 0.541 0.485 0.429 (0.467) (0.385 ) (0.343) [0.364 | [0.300] [0.240] Negros 0.643 0.703 (0.500) (0.621) [0.429 | [0.526 ] Mindoro 0.606 (0.480) [0.471] Hologerrhum has a relict pattern exhibiting limitation to the distal island of Luzon (fig. 5). Pseudogekko, with two species, has a spotty distribution; Cyclocorus (1 species) and Oxyrhabdium (2 species) have less obvious relict patterns, being more widely distributed throughout the archipelago (figs. 4 and Se INTERNAL MIGRATION ROUTES When differences in composition as well as diversity are considered, it is possible to compare the relative effectiveness of probable dispersal routes indicated in fig. 1b. In comparing composition of the faunas, we have used the twice the number of species common to the two communities similarity index : a oa the sum of the species comprising each of the communities developed by Sorenson (1948) for comparing plant communities in northeast Greenland and in Denmark. A low index of similarity will be the result of either: (1) a large difference in diversity; or (2) when diversities are more or less equal, a small number of species in common. In addition to the 5 islands upon which this study is primarily based, Leyte Island, as well as Luzon, has been included in this section since the former lies 120 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. TaBLeE 7. Indices of similarity for lizards. (The index in parentheses is based on the species exclusive of the 23 widespread forms associated with man’s economy or beach communities, that in brackets is based on the species remaining after relicit species are also excluded ). Mindanao Bohol Negros Mindoro Luzon Palawan 0.400 0.481 0.500 0.440 0.316 (0.185 ) (0.125) (0.176) (0.000 ) (0.053 ) [0.213 ] [0.148 | [0.207 } [0.000] [0.067 | Mindanao 0.651 0.471 0.380 0.395 (0.548) (0.313) (0.207) (0.265) [0.539 ] [0.333 | [0.200 | [0.218] Bohol 0.562 0.552 0.491 (0.333 ) (0.278) (0.348) [0.294 | [0.267 | [0.286] Negros 0.600 0.478 (0.368) (0.333) [0.375 ] [0.323 ] Mindoro 0.590 (0.429) [0.424 ] closest to Mindanao on the most eastern dispersal route. The Leyte indices have been computed only for frogs and snakes, based on lists published by Inger (1954) and Leviton (1963). The lizards are not sufficiently well known to be included at this time, and the snakes are believed to be rather poorly known since several widely distributed species have not yet been reported from this island. This, however, would tend to introduce an error in the direction of a low rather than a high index. As is evidenced in table 2, Mindanao, at the proximal end of the eastern routes, exhibits a higher diversity for each of the major taxa considered, as well as for the herpetofauna as a whole, than do any of the other 4 islands included in our expeditions. This will depress the similarity index when comparisons are made with the other islands, whereas no such depressing effect will exist in the instance of Palawan at the proximal end of the western entryway, since the diversity of the Palawan herpetofauna is about the same as that of the more distal islands which range from 60 to 80 species in total herpetofauna. Thus, if indices between Mindanao and more distal islands are equal to or greater than the indices between Palawan and these same islands, less isolation of island faunas along eastern routes from Mindanao would be indicated. All indices, those for each of the taxa, (frogs, lizards, and snakes), as well as those for the total herpetofaunas (tables 6—9), are moderately high when Vor. XXXVIII] BROWN AND ALCALA: HERPETOFAUNA OF PHILIPPINES 121 TABLE 8. Indices of similarity for snakes. (The index in parentheses is based on species exclusive of the 23 widespread forms associated with man’s economy or beach communities, that in brackets is based on the species remaining after relicit species are also excluded). Mindanao Leyte Bohol Negros Mindoro Luzon Palawan 0.528 0.431 0.491 0.323 0.308 0.308 (0.373 ) (0.229) (0.278) (0.182 ) (0.171) (0.296) [0.385 ] [0.235 ] [0.286 ] [0.190 | [0.176] [0.314 ] Mindanao 0.596 0.644 0.500 0.414 0.538 (0.488 ) (0.524) (0.360) (0.293 ) (0.433 ) [0.474 | [0.513 | [0.348 ] [0.268 ] [0.436] Leyte 0.684 0.468 0.324 0.491 (0.609 ) (0.305 ) (0.182 ) (0.341) [0.571] [0.345 | [0.200] [0.378] Bohol 0.531 0.462 0.542 (0.315) (0.348) (0.333 ) [0.345 | [0.381 | [0.378 ] Negros 0.542 0.706 (0.516) (0.640) [0.429 | [0.622 ] Mindoro 0.621 (0.585) [0.541] Palawan and Mindanao are compared, even though the differences in diversity are relatively large. This is the result of the large number of species which these 2 entry-way islands have in common. These common species have either entered the 2 islands relatively recently from Borneo or have continued to reinvade from time to time. The fact that many of these species have not dispersed to more distal islands in the Philippines and yet are conspecific with Bornean populations tends, however, to support the first explanation of more recent entry. When indices between Mindanao and Mindoro, Leyte, Luzon, or Negros, based on the complete fauna (both native and nonnative species), are compared with the indices between Palawan and these same islands (tables 6—9), those with Mindanao are much higher, with one exception, even though the greater diversity of the Mindanao fauna, almost twice that of Palawan, tends to depress the similarity indices. The exception is the Palawan-Mindoro index for lizards. These differences are more pronounced and the exception ceases to exist when the 23 widespread, nonnative species (p. 111) are excluded. The differences are not quite as great, in most instances, when those classified as older relicts (p. 117) are also excluded; but these latter, small differences can be accounted for by the absence of the relict genera Platymantis and Brachymeles from Palawan. 122 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH SER. TasBLe 9. Indices of similarity for the total herpetofauna. (The index in parentheses is based on species exclusive of the 23 widespread forms associated with man’s economy or beach communities, that in brackets is based on the species remaining after relicit species are also excluded). Mindanao Bohol Negros Mindoro Luzon Palawan 0.453 0.453 0.372 0.382 0.372 (0.304 ) (0.245) (0.109 ) (0.165 ) (0.186 ) [0.331] [0.274] [0.122 ] [0.188 | [0.214 ] Mindanao 0.660 0.493 0.415 0.475 (0.584) (0.354) (0.286) (0.362) [0.567 ] hossrval [0.258] [0.331] Bohol 0.547 0.508 0.494 (0.377) (0.364) (0.368) [0.345 | [0.342 ] [0.337] Negros 0.588 0.616 (0.457) (0.511) [0.415] [0.481] Mindoro 0.605 (0.523) [0.505 ] This indicates that the primary dispersal routes for the native fauna were by way of Mindanao, and that the present herpetofauna of Palawan has not dis- persed widely into other areas in the Philippines. The high indices between Mindanao, Leyte, and Bohol are due to the very large number of species which they hold in common, even though diversity is greatly reduced. This suggests a very high rate of exchange between Mindanao and these 2 nearby islands on the eastern routes. The slightly higher indices between Mindanao and Bohol as compared to those between Mindanao and Leyte may be partly the result of lesser knowledge of the fauna of Leyte. How- ever, it may also be the result of some exchange across a direct Mindanao- Bohol route. The relatively high Mindoro-Negros index for each taxon, as well as for the total herpetofauna, may be in part due to their positions at the distal end of dispersal routes, and, consequently, the same species have tended to reach both. However, it also suggests an active Mindoro-Panay-Negros dispersal route. The richer fauna of Negros, on the other hand, also indicates that a part of the Negros fauna must have arrived by way of the shorter Mindanao-Leyte-Bohol, or Mindanao-Leyte-Cebu dispersal routes, or in some instances perhaps, a Mindanao-Bohol-Negros route. The progressively lower indices with Mindanao, as one progresses along the Vor. XXXVIII] BROWN AND ALCALA: HERPETOFAUNA OF PHILIPPINES | 123 eastern dispersal route to Mindoro, appears to be consistent with the decrease in diversity. The Mindoro-Mindanao index is higher for frogs than for either lizards or snakes. This would be expected in terms of a later arrival, which might be accounted for by their slower dispersal ability where marine barriers have presumably operated. EVIDENCE FOR OVER-WATER DISPERSAL In addition to the evidence of barriers to dispersal indicated by the reduction of numbers of species on distal islands as compared to proximal islands, when the size variable is minimized, as suggested in the section on diversity (fig. 2), the data may be further analyzed in terms of the probable differential effect of marine barriers. As noted at the close of the previous section, on biological grounds marine barriers would be expected to affect adversely dispersal of amphibians to a greater extent than that of reptiles. Therefore, on the assumption that the primary entryway into the Philippine archipelago was from Borneo by way of Mindanao, the amphibia might be expected to exhibit a more rapid reduction in number of species than do the reptiles when comparisons are made between Mindanao and the more distal islands such as Bohol, Negros, Mindoro, and Luzon if dispersal did take place wholly or in part across such barriers along these eastern routes. This should become even more evident if, as well as the nonnative, the presumably older endemic relict elements, those which exhibit the typical relict pattern of chance pockets of isolated species and/or secondary radiation centers on distal islands, were also excluded. To evaluate this, we propose a simple proportional-diversity index. This makes possible an objective comparison of changes in diversity for amphibia relative to changes in diversity for the reptiles. The index is calculated as the ratio of the number of species in the particular taxon (frogs, lizards, etc.) to the number of species in the total herpetofauna. This index for frogs is indeed lower for Luzon, Negros, and Mindoro, at the distal end, than for Mindanao or Bohol at the proximal end of the dispersal routes. The amphibian index for Bohol is noteworthy in that it is slightly higher than that for Mindanao. This may be, at least in part, a distortion due to a disproportionately poorly known snake fauna (see p. 119). Since indices for reptiles (both lizards and snakes) tend to increase as the index for amphibians decreases, it is interesting to note that the index for lizards exhibits a greater increase than that for snakes in all instances, except for Luzon Island, when the distal islands are compared with Mindanao at the entryway. For some reason, snakes appear to have been relatively more successful in their dispersal to Luzon, or have suffered fewer extinctions there. We interpret these changes in the proportional diversity indices as supporting the conclusion that much of the herpetofauna of the Philippines, with the ex- ception of that of Palawan, has been the result of waif dispersal across marine [Proc. 4TH SER. CALIFORNIA ACADEMY OF SCIENCES 4 2 fuozny (9) {so1saN (S$) {joyog (+) SezAe7T (¢) Sovuepurpy (44) Wau 000'001 (2) SAAVNS Ol Lets its ! i (,44 ) VauV 00000! 000'O! ooo! Toso a Line Cee es Ca aaa T [ea o} 4 SNVISIHDWY | | : 4 3 ol °) g 6 : = 2 e ean ao wen 7 aS bao 5 ee = allisnienie t | ett 001 S3193dS JO Y38WNN S3193dS 40 Y3A8WNN ‘oropuly, (2) [at eae 1 SUBMETed (T) :Spue[Sst Avy pue 1evou AO0F SdAIND sateds-va1y ‘QO AMNOLA (44) vay 000'00! 0000! oor InAs T Toalpeulaeal neal T ala Tall neal ane T i Le 6 et | al 4 L jl ¢ c= [ ae Ie 5 as {L~, —, 00in | @ 0 au Q t shee b | [ sayvzin 7 Sere ee eS eee 1 ear essuy i yer i =lroGl (Wy) vauV 000'00! 000'O! 0001, LUA 1s 1s T ee re if Tite (im one set | T \ [ | 4 8 S3193dS 40 Y38WNN VoL. XXXVIII] BROWN AND ALCALA: HERPETOFAUNA OF PHILIPPINES _ 125 Taste 10. Proportional diversity indices for caecilians, frogs, lizards, and snakes for selected proximal, intermediate, and distal islands for the total herpetofauna, the fauna with the presumed nonnative species excluded (in parentheses), and with both the nonnative and that element classified as relict included [in brackets |. Palawan Mindanao Bohol Negros Mindoro Luzon Number of Caecilian species 0.013 0.008 (Total herpetofaunal species) (0.018) (0.009 ) [0.019 | [0.011] Number of frog species 0.269 0.272 0.292 0.205 0.207 0.223 (Total herpetofaunal species) (0.309 ) (0.291) (0.352 ) (0.218) (0.216) (0.230) [0.302 ] [0.292 ] [0.333 [0.178] [0.188 ] [0.186] P; Number of lizard species 0.295 0.416 0.431 0.423 0.466 0.362 (Total herpetofaunal species) (0.218) (0.408 ) (0.392 ) (0.400) (0.459) (0.351) [0.208] [0391] [0.381] [0400] [0469] [0.322] Number of snake species 0.423 0.312 0.278 0.372 0.328 0.415 (Total herpetofaunal species) (@455)) (O80) (255) (0.382) (0.324) (0.419) [0.422 ] [0.315 | [0.286 | [0.422] [0.344 ] [0.492 ] barriers from Mindanao Island. This conclusion is most strongly evidenced when nonnative and isolated relict species are excluded leaving the so-called immigrant species (table 10). DIVERSITY AS RELATED TO AREA AND DISTANCE Area. When number of native species is plotted against area (fig. 6) for the compact group of islands included in the present study, the curves for amphibians and the total herpetofauna exhibit the expected pattern for overwater dispersal as postulated by MacArthur and Wilson (1963, 1967). Large islands do exhibit a greater diversity (number of species) than small islands and near islands a consistently greater diversity than more distant islands, in terms of the probable migration routes. The diversity of lizards on Palawan, a near small island, and the diversity of snakes of Mindanao are somewhat less than expected, however, based on the slope of the curves. The diversity of snakes on Luzon, the most distant large island, also appears slightly greater than might be expected. One explanation might be that the relatively short over-water distances obtaining in this compact archipelago have made possible a higher frequency of invasion of snakes along the eastern chain to Luzon or that there are more relict snakes on Luzon. Since lizards of the endemic genus Brachymeles are absent from Palawan, this probably accounts in part for the lower diversity of this faunal group on that island. The diversity for amphibians and lizards on Bohol Island is greater than expected and that of the snakes lower. It has been suggested that the latter may be due [Proc. 4TH SER. CALIFORNIA ACADEMY OF SCIENCES 126 ( 4X8} 895) X30NI JONVLSIO 000'0! 000'| (oleyt ( 4x9} 99S) X3Z0NI JONVLSIO 000'0! 000'! 00. r SNVIEIHGWV 4 S3193dS 3O Y3IB8WNN ool lo} S3103dS JO YAIBWNN fefe}| ‘o1opulyy (2) :uozny (9) ‘SOIBIN (S$) ‘foyog (+) ‘aJAeT (¢) ‘ovuepulyy (Z) ‘ueMeTeg (T) ‘SpUueIST [[eWIS pue aBIe] OJ SaAIND satads-xaput 9dueISIC, ( 4x8} 98S) X30NI JONVLSIO 000'0! 000'! O01 \TelUallan pal el oad Go a | a Ss L a : g : 7 o reas a r Passe 7 ¢ L Te ee | 0 m - a) @ C SQYVvZI1 a 8 Each hei tte Elna (4x8) 88S ) X30NI JONVLSIC 000'O! 0001! ool a ri === Sinaloa ee Ol s = 4 S ¥ Tews 1 i] : Se8 : = S55 001 = s LL | cr WNAVSOLSdHYSH TVLOL | Suen tesioo tha es | || i 9001 S3193dS 40 Y38WNN "L FINO Vor. XXXVIII] BROWN AND ALCALA: HERPETOFAUNA OF PHILIPPINES 127 TABLE 11. Calculated distance index for Philippine islands included in this study in rela- tion to Borneo. Weighted number Approximate over-water Weighted of effective distance by present distance Route marine barriers probable routes (in km.) index Borneo—Palawan (1) 150 150 Borneo—Sulus—Mindanao (1) 345 345 Borneo—Mindanao—Leyte (2) 395 790 Borneo—Mindanao—Leyte—Bohol (3) 420 1260 Borneo—M indanao—Leyte—Cebu —Negros (4-5=414) 505 2272.5 Borneo—Mindanao—Leyte—Samar —Luzon (4) 440 1760 Borneo—Mindanao—Leyte—Samar —Luzon—Mindoro (5) 455 DDS to a sampling bias. The high diversity of lizards and amphibians may possibly be the result of the very narrow water gap between Leyte and Bohol. Distance. Any attempt to measure over-water dispersal distances in this compact, nonlinear archipelago is difficult. If the effective distance is measured as the airline distance from Borneo to the various islands, Leyte is almost as distant as Luzon, and Bohol almost as distant as Mindoro (fig. 1b). If the effective distance is measured as the sum of the breadth of over-water distances between islands, by ways of eastern migration routes for all islands except Palawan, assuming that the islands themselves provide stepping stones of ecologically relatively uniform space, the effective over-water distance between Mindanao and Luzon, the northernmost island, is 75 kilometers as compared to 55 to 90 kilometers as the effective over-water distance between Mindanao and Bohol for example. In an attempt to minimize these sources of error we have derived a weighted index by multiplying the sum of the approximate over-water distances times a value for number of marine barriers (table 11). When this distance index is plotted against number of species (fig. 7), the shape of the curves are again, with slope opposite to that of the area curves, in keeping with that expected for amphibians and the herpetofauna as a whole, and the curves for large and small islands are nearly parallel. The diversities for snakes on Mindanao and Luzon and for lizards on Palawan also impose effects on the slopes of the curves which are comparable to the effects on the area curves. Bohol and Leyte also exhibit a very low diversity for snakes relative to the curve for other islands in the same general size-category. The general agreement between these curves and those based on area suggests that such a weighted distance index may be useful in the island faunas in similar compact archipelagos. 128 CALIFORNIA ACADEMY OF SCIENCES [| Proc. 4TH SER. SUMMARY AND CONCLUSIONS The existence of relict patterns and secondary radiation centers, the relative importance of possible internal migration routes from alternative entryways, the evidence for over-water dispersal of the ‘‘migrant” element, and the effects of island area and distance are considered for the herpetofauna of the Philippines, a compact, fringing archipelago. Evaluations are based primarily on diversities and relationships of the herpetofaunas of 7 of the islands, which have a total of 220 species (54 amphibians, 92 lizards, and 74 snakes). Only in the evaluation of relict distributions is the known herpetofauna of the total archipelago taken into consideration. The distributional patterns within the Philippines of the endemic, multi- species genera of lizards, Luperosaurus and Brachymeles, as well as the amphibian genus Platymantis, exhibit relict patterns of the type resulting from partial ex- tinction of an old fauna, which has existed as a number of isolated units. Brachymeles and Platymantis also give evidence of secondary radiation centers in distal islands. Patterns for the endemic genus of lizards Pseudogekko and snake genera Cyclocorus, Hologerrhum, and Oxyrhabdium are simpler relict patterns with only 1 or 2 species in each genus. These are rather widely distrib- uted, or, in some instances, limited to either the distal or proximal islands. These patterns are those postulated by Darlington (1957) as patterns which would develop within a chain of islands. Sorenson’s index of similarity is used to evaluate the relative effectiveness of dispersal routes. These indices, particularly when the nonnative fauna is excluded, indicate that the primary dispersal route or routes within the archi- pelago have been the eastern routes, by way of the Mindanao-Leyte—or possibly, in some instances the Mindanao-Bohol—pathways. The Palawan entryway has contributed very little to the herpetofauna of the rest of the Philippines. High indices of similarity between Negros and Mindoro suggest a relatively active migration route between these two islands. A proportional-diversity index, based on the presumed lower ability of am- phibians to disperse across marine barriers, is used to evaluate the probable effect of marine barriers. The evidence indicates that, with the exception of Palawan Island, much of the herpetofauna has apparently reached the inter- mediate and distal islands of the archipelago from Mindanao as a result of waif dispersal across marine barriers. When number of species for the total herpetofauna, and for the amphibians, lizards, and snakes independently, are plotted against area or against a weighted distance value the curves for amphibia and the total herpetofauna exhibit patterns consistent with those projected from MacArthur’s and Wilson’s thesis (1967) regarding faunal diversity along a chain of islands of varying size. The data indicate, however, that the diversity of the lizard fauna on Palawan island Vou. XXXVIII] BROWN AND ALCALA: HERPETOFAUNA OF PHILIPPINES 129 is lower than expected, and that the diversity of the snake fauna is probably somewhat lower for Mindanao and higher for Luzon island relative to the diversity exhibited by the fauna of other islands in the study. The effects of the narrow marine barriers on the present distribution of the amphibian fauna have produced a pattern of island diversities in general agreement with the MacArthur-Wilson hypothesis. The diversities of lizards and snakes for the sample group of islands included in this study exhibit several discrepancies. The relatively narrow over-water barriers between the islands of this compact archi- pelago, less effectual against reptiles than amphibians, and the several possible migration routes and secondary centers of radiation may be factors in the distribution patterns of these faunal elements. The reasons, however, for the low diversity of lizards of Palawan and for the effectiveness of the barrier be- tween Palawan and Mindoro, a marine channel only about 150 km. in breadth at the present time and broken by small islands, are not readily explained from data. ACKNOWLEDGMENTS This study was supported by National Science Foundation Grant GB-4156. Illustrations were prepared by Walter Zawojski, Stanford University. LITERATURE CITED Brown, W. C., and A. C. ALCALA 1961. Populations of amphibians and reptiles in the submontane and montane forests of Cuernos de Negros. Ecology, vol. 42, pp. 628-636. 1964. Relationship of the herpetofaunas of the non-dipterocarp communities to that of the dipterocarp forest of southern Negros Islands, Philippines. Senkenbergiana Biologica, Frankfurt am Main. vol. 45, pp. 519-611. Brown, W. C., and D. S. RABor 1967. Review of the genus Brachymeles (Scincidae), with descriptions of new species and subspecies. Proceedings of the California Academy of Sciences, vol. 34, no. 16, pp. 525-548. DariincrTon, P. J., JR. 1957. Zoogeography: the geographical distribution of animals. Wiley and Sons, New York, xi +675 pp. INGER, R. F. 1954. Systematics and zoogeography of Philippine amphibia. Fieldiana: Zoology, vol. 33, pp. 183-531. Incer, R. F., and H. Marx 1965. The systematics and evolution of the oriental colubrid snakes of the genus Calamaria. Fieldiana. vol. 49, pp. 1-304. Leviton, A. E. 1963. Remarks on the zoogeography of Philippine terrestrial snakes. Proceedings of the California Academy of Sciences. vol. 31, no. 15, pp. 369-416. MacArruor, R. H., and E. B. Wison 1963. An equilibrium theory of insular zoogeography. Evolution. vol. 17, pp. 373-387. 1967. The theory of island biogeography. Princeton University Press. xi +203. 130 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. SORENSON, T. 1948. A method of establishing groups of equal amplitude in plant sociology based on similarity of species content and its application to analyses of the vegetation on Danish commons. Danske Videnskabernes Selskab, Biologiske Skrifter, Copenhagen, vol. 5, no. 4, pp. 1-34, 6 tables, 1 figure. Taytor, E. H. 1922a. The lizards of the Philippine Islands. Philippine Bureau of Science. Manila. Publicaticn 17, 269 pp., 23 pis. 1922b. Additions to the herpetological fauna of the Philippine Islands I. Philippine Journal of Science. vol. 21, pp. 161-206, 7 pls. 1922c. Additions to the herpetological fauna of the Philippine Islands II. Philippine Journal of Science. vol. 21, pp. 257-303, 4 pls. 1923. Additions to the herpetological fauna of the Philippine Islands III. Philippine Journal of Science. vol. 22, pp. 515-557, 3 pls. 1925. Additions to the herpetological fauna of the Philippine Isalnds IV. Philippine Journal of Science. vol. 26, pp. 97-111. 1928. Amphibians, lizards and snakes of the Philippines. 7m: Dickerson, R., et al. 1928, pp. 214-241, pls. 27-32. ZWEIFEL, R. G. 1967. Identity of the frog Cornufer unicolor and application of the name Cornufer. Copeia, 1967, pp. 117-121. SPT CIEL As) i ——S =e - i j 7 — é lan ea PS - a3 ee . if 7 ] ide nr F Osi oi itd 4 jam sal i hifi 1 Vite Me >"l ae ; : - iéiu re, 20, pen —- i) Uw. OF, ee ihe mid i wel pees 9) .¢ >) OB eit “EY ie y ii PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Festschrift for George Sprague Myers Vol. XXXVIII, No. 7, pp. 131-138. December 31, 1970 TROPICAL SHELF ZOOGEOGRAPHY By John C. Briggs Depariment of Zoology, University of South Florida, Tampa, Florida INTRODUCTION The richest marine fauna is found in the shallow waters of the tropical oceans at depths generally less than 200 meters. Zoogeographically, four great regions may be identified, the Indo-West Pacific, the Eastern Pacific, the Western Altantic, and the Eastern Atlantic. Each region may, in turn, be subdivided into provinces, but these will not be discussed at this time. To the north and south, the tropics are bounded by the 20°C. isotherm for the coldest month in the year. Longitudinally, the tropical regions are separated from one another by barriers that are very effective since each region possesses, at the species level, a fauna that is highly endemic. By studying the operation of these longitudinal barriers, one can learn something about the interrelationship of the regions and can also obtain information leading to a better understand- ing of zoogeography and evolution. THE EAST PACIFIC BARRIER The Indo-West Pacific and Eastern Pacific regions are separated by the East Pacific Barrier, the vast stretch of deep-water that lies between Polynesia and America. In regard to the shore fishes, it was concluded that an eastward colonization movement was taking place across the Barrier and that successful reciprocal migrations were, at least, very rare and might be completely lacking [131] 132 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER. (Briggs, 1961, 1964, 1966). When the general relationship of the tropical shelf regions was first discussed (Briggs, 1967a), comparable information on the major groups of the shallow-water invertebrates was not available. Emerson (1967) published a revealing analysis of the distribution of those Indo-West Pacific species of mollusks that have succeeded in penetrating across the Barrier into the Eastern Pacific. He found that such trans-Pacific species were largely restricted, in the Eastern Pacific, to the oceanic islands, the greatest numbers being found at Clipperton (33 species) and at the Galapagos (25 species). It was noted that the gastropods, which greatly out- numbered the bivalves, belonged to groups that were known to have relatively long larval stages. Most important of all, Emerson pointed out that no mol- luscan species of apparent Eastern Pacific origin were known to occur in Polynesia. Data on the other invertebrate groups are not as complete, but it is significant that some of the littoral echinoderms (Ekman, 1946), holothurians (Deichmann, 1963), decapod crustaceans (Chace, 1962; Garth, 1965), and hermatypic corals (Emerson, 1967) found in the Eastern Pacific (especially around the offshore islands) are trans-Pacific species of apparent Indo-West Pacific origin. Therefore, it may now be said that for the tropical marine shore fauna in general, including both fishes and invertebrates, it seems likely that successful migration across the East Pacific Barrier takes place in one direction only—from west to east. THE NEW WORLD LAND BARRIER The New World Land Barrier, with the Isthmus of Panama forming its narrowest part, is virtually a complete block to the movement of tropical marine species between the Eastern Pacific and Western Atlantic. This state of affairs has existed since about the latest Pliocene or earliest Pleistocene (Simpson, 1965; Patterson and Pascual, 1968) so that, at the species level, the two faunas are well separated. The present Panama Canal has not notably altered this relationship since, for most of its length, it is a freshwater passage forming an effective barrier for all but a few euryhaline species. The New World Land Barrier is the most effective of the four zoogeo- graphic barriers that separate the tropical faunal regions. It has stood for approximately three million years, but it now appears that man is about to breach this barrier by excavating a sea-level canal somewhere in the vicinity of the Isthmus of Panama. If such a canal is constructed, it would present ample opportunities for marine animals to migrate in either direction. This could result in the Eastern Pacific being invaded by over 6000 species of fishes and invertebrates and the Western Atlantic being invaded by over 4000 species. Since the Western Atlantic species would apparently be competitively dominant, it has been predicted that a large scale extinction would take place in the VoL. XXXVIIT] BRIGGS: TROPICAL SHELF ZOOGEOGRAPHY 133 Eastern Pacific resulting in the irrevocable loss of a huge number (possibly thousands) of species ( Briggs, 1968, 1969). THE MID-ATLANTIC BARRIER The broad deep-water barrier that separates the Western Atlantic tropics from those of the West African coast functions in a very interesting manner. An impressive number of shore fishes have managed to traverse the Mid- Atlantic Barrier from west to east. It has been estimated (Briggs, 1967a) that about 118 shore fish species have trans-Atlantic distributions but that only about 24 of them came from the Indo-West Pacific via the Cape of Good Hope. The rest probably evolved in the Western Atlantic and successfully performed an eastward colonization journey across the ocean. Trans-Atlantic species comprise about 30 percent of the shore fish fauna of tropical West Africa. Works on some of the major groups of West African invertebrates also show that an appreciable number of the species are trans-Atlantic: Dekeyser (1961) found that about 25 percent of the ascidians showed this distribution; Burton (1956), 18 percent of the sponges; Monod (1956), 16 percent of the anomuran and brachyuran crabs; Knudsen (1956), 6 percent of the prosobranch mollusks; Ekman (1953), 16 percent of the starfishes, brittle stars, and sea urchins; and Marcus and Marcus (1966), 29 percent of the opisthobranch mollusks. Furthermore, Chesher (1966), who found 8 trans-Atlantic species of sea urchins in the Gulf of Guinea, stated that gene flow appeared to take place from west to east. It seems apparent that, in both the fishes and the invertebrates, the great majority of the trans-Atlantic species originated in the Western Atlantic and then migrated eastward. The westward colonization traffic appears to be re- stricted to certain dominant species that originated in the Indo-West Pacific and then gained access to the Atlantic by rounding the Cape of Good Hope. So far, there are no indications that species originating in the Eastern Atlantic, and belonging to genera typical of that area, have been successful in becoming established on the western side. THE OLD WORLD LAND BARRIER The Eastern Atlantic and the Indo-West Pacific regions are separated by the Old World Land Barrier. It has been estimated that the continental masses of Eurasia and Africa have been linked at least since the beginning of the Pleistocene (Gohar, 1954). The Suez Canal is a sea-level passage that has been open since 1869 but migration of marine animals has been inhibited for two reasons: first, the canal connects two areas that are separated by a temperature barrier, the Red Sea being tropical while the Mediterranean is warm-temperate; second, the Bitter Lakes, which form part of the Suez pas- sageway, have a high salinity (about 45 percent). Despite these difficulties, 134 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Serr. the limited migratory movements that have taken place through the Suez Canal do provide some significant information. The Mediterranean has been invaded by at least 24 species of Red Sea fishes (Ben-Tuvia, 1966), 16 species of decapod crustaceans (Holthuis and Gottlieb, 1958), and several species in other groups such as the tunicates (Pérés, 1958), mollusks (Engel and van Eeken, 1962), and stomatopod crus- taceans (Ingle, 1963). So, while there is ample evidence of intrusions into the eastern Mediterranean, there are no reliable data that indicate any success- ful reciprocal migration. Also, there are some indications that the invaders from the Red Sea (a part of the Indo-West Pacific Region) are replacing rather than coexisting with certain native species (George, 1966). The various circumtropical shore species have probably been able to pre- serve their genetic homogeneity by means of migration around the Cape of Good Hope (in addition to crossing the open ocean barriers in the Pacific and Atlantic). Talbot and Penrith (1962) remarked that surface temperatures of 21°C. are often present round the Cape outside a cold upwelling area. There are about 16 known species of circumtropical shore fishes (Briggs, 1960). Besides these, there are about 15 other species that apparently transgress the Old World Land Barrier at the Cape (Briggs, 1967a). Of the total of 31 fish species, eight are monotypic but all the rest represent genera that are best developed in the Indo-West Pacific. Apparently, only a few tropical invertebrate species have been able to migrate around the Cape of Good Hope. Monod (1956) in his monographic study of the West African decapods showed that 10 out of 176 shore species occurred in the Indo-West Pacific. Ekman (1953) noted that only 2 percent of the tropical Atlantic echinoderms (Asteroidea, Ophiuroidea, and Echinoidea) extended around the Cape. It appears, especially from the ichthyological evidence, that the colonization movement of tropical shore species around the Cape takes place entirely in a westerly direction, from the Indo-West Pacific into the Atlantic. RELATIONSHIPS OF THE SHELF REGIONS Evidence now available about the dispersal of the shallow-water marine invertebrates tends to substantiate the general nature of a remarkable distri- butional phenomenon that was discovered earlier for the shore fishes (Briggs 1961, 1964, 1967a). Successful (colonizing) migrations across the zoogeo- graphic boundaries that delimit the Indo-West Pacific can apparently take place in one direction only, outward into areas where the fauna is poorer and the competition is less. The realization that the East Pacific and Old World Land Barriers operate as one-way filters enables us to understand better how the Indo-West Pacific Region serves as the evolutionary and distributional center for the tropical shore animals of the world. We can see that competi- tively dominant species continue to migrate, as they probably have for mil- VoL. XXXVIIT] BRIGGS: TROPICAL SHELF ZOOGEOGRAPHY 135 lions of years, from the Indo-West Pacific eastward across the open ocean to America and westward around the Cape of Good Hope into the Atlantic; since 1869, some of them have also been able to pass northward through the Suez Canal into the Mediterranean. The Western Atlantic Region may be considered a secondary center of evolutionary radiation. Many species evolved in this area have proved ca- pable of migrating eastward to colonize the tropical Eastern Atlantic. How- ever, species originating in the Eastern Atlantic are apparently incapable of successfully invading the western side. Again, the advantage seems to lie with the area that possesses the richer fauna and the higher level of competi- tion. It can be seen that the completely eastward direction of successful mi- gratory movements across the East Pacific Barrier and the predominantly eastward movements across the Mid-Atlantic Barrier take place in a direction opposite to that of the main flow of the surface waters via the North and South Equatorial Currents. In contrast, the surface and subsurface counter- currents in the tropical Pacific and Atlantic are weakly developed but these smaller currents are obviously the principal means by which successful trans- port is achieved. Fell (1967) noted that certain groups of shore species apparently demon- strated speciation gradients in which the number of species gradually dimin- ished around the world in a westward direction. He interpreted this to mean that the direction of successful migrations had also been to the west and that such dispersals had been carried out by the North and South Equatorial cur- rents. Subsequently, it was pointed out that the existence of a gradient in numbers of species (or genera) across a major barrier did not necessarily indicate the direction of the original successful migration (Briggs, 1967b). The fact that colonizations do take place in a direction opposite to that of the major currents is a good indication that biological competition rather than passive transport is probably the most important factor controlling the successful dispersal of tropical marine shore animals. LITERATURE CITED Ben-Touvi, A. 1966. Red Sea fishes recently found in the Mediterranean. Copeia, no. 2, pp. 254-275, 2 figs. BRIGGS ANC. 1960. Fishes of worldwide (circumtropical) distribution. Copeia, no. 3, pp. 171-180. 1961. The East Pacific Barrier and the distribution of marine shore fishes. Evolution, vol. 15, no. 4, pp. 545-554, 3 figs. 1964. Additional transpacific shore fishes. Copeia, no. 4, pp. 706-708. 1966. Zoogeography and evolution. Evolution, vol. 20, no. 3, pp. 282-289. 1967a. Relationship of the tropical shelf regions. Studies in Tropical Oceanography, Miami, no. 5, pp. 569-578. 136 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. 1967b. Dispersal of tropical marine shore animals: coriolis parameters or competi- tion? Nature, vol. 216, no. 5113, page 350. 1968. Panama’s sea level canal. Science, vol. 169, no. 3853, pp. 511-513. 1969. The sea-level canal: potential biological catastrophe. BioScience, vol. 19, no. 1, pp. 44-47. Burton, M. 1956. The sponges of West Africa. Atlantide Report, no. 4, pp. 111-147, 4 figs. Cuace, F. 1962. The non-brachyuran decapod crustaceans of Clipperton Island. Proceedings of the United States National Museum, vol. 113, no. 3466, pp. 605-635, 7 figs. CuHESHER, R. H. 1966. Report on the Echinoidea collected by R/V Pillsbury in the Gulf of Guinea. Studies in Tropical Oceanography, Miami, no. 4, part 1, pp. 209-223. DEICHMANN, E. 1963. The holothurians of Clipperton Island in the Eastern Tropical Pacific. Breviora, no. 179, pp. 1-5. DEKEYSER, P. L. 1961. Liste provisoire des urocordés de la céte occidentale d’Afrique. Bulletin de la Institute francaise d’Afrique Noire, série A, vol. 23, no. 1, pp. 217-230. EKMAN, S. 1953. Zoogeography of the sea. Sidgwick and Jackson, London, xiv + 417 pages, 121 figs. Emerson, W. K. 1967. Indo-Pacific faunal elements in the tropical eastern Pacific, with special refer- ence to the mollusks. Venus, Japanese Journal of Malacology, vol. 25, nos. 3 and 4, pp. 85-93. ENGEL, H. and C. J. VAN EEKEN 1962. Red Sea Opisthobranchia from the coast of Israel and Sinai. Sea Fisheries Re- search Station, Haifa, Israel, Bulletin, no. 30, pp. 15-34, 7 figs. Idorit, JBl, 183, 1967. Resolution of coriolis parameters for former epochs. Nature, no. 214, pp. 1192- 1198. GarTH, JOHN S. 1965. The brachyuran decapod crustaceans of Clipperton Island. Proceedings of the California Academy of Sciences, vol. 33, no. 1, pp. 1-46, 26 figs. GeEorGE, C. J. 1966. A two year study of the fishes of the sandy littoral of St. George Bay, Leb- anon. Abstracts, 2nd International Oceanographic Congress, Moscow, page 130. GouHar, H. A. F. 1954. The place of the Red Sea between the Indian Ocean and the Mediterranean. Publications of the Hydrobiological Research Institute, University of Istanbul, Series B, vol. 2, no. 2-3, pp. 1-38, map. HoxtuHuts, L. B. and E. Gortiies 1958. An annotated list of the decapod Crustacea of the Mediterranean coast of Israel, with an appendix listing the decapoda of the eastern Mediterranean. Bulletin of the Sea Fisheries Research Station, Israel, no. 18, pp. 1-126, 15 figs. INGLE, R. W. 1963. Crustacea Stomatopoda from the Red Sea and the Gulf of Aden. Bulletin of the Sea Fisheries Research Station, Haifa, Israel, no. 33, pp. 1-69, 73 figs. Vor. XXXVIIT) BRIGGS: TROPICAL SHELF ZOOGEOGRAPHY 137 KNUDSEN, J ORGEN 1956. Marine prosobranchs of tropical West Africa (Stenoglossa). Atlantide Report, no. 4, pp. 8-110, 2 figs., 4 pls. Marcus, Ernst, and EvettIne Marcus 1966. Opisthobranchs from tropical West Africa. Studies in Tropical Oceanography, Miami, no. 4, part 1, pp. 152-208, 62 figs. Monon, Tu. 1956. Hippidea et brachyura ouest-africains. Memoires de la Instiute francaise d’Afrique Noire, no. 45, pp. 1-674, 884 figs., 1 map. PATTERSON, B. and R. PAscuaL 1968. The fossil mammal fauna of South America. The Quarterly Review of Biology, vol. 43, no. 4, pp. 409-451, 13 figs. PrrEs, J. M. 1958. Ascidies recoltées sur les cétes Mediterranéenar d’Israel. Bulletin of the Sea Fisheries Research Station, Haifa, Israel, no. 19, pp. 143-150. SIMPSON, G. G. 1965. The geography of evolution. Chilton Books, Philadelphia and New York. x + 249 pp., 45 figs. Tasot, F. H. and M. J. PenritH 1962. Tunnies and marlins of South Africa. Nature, vol. 193, no. 4815, pp. 558-559. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Festschrift for George Sprague Myers Vol. XXXVIII, No. 8, pp. 139-156; 6 figs.; 3 tables December 31, 1970 A NEW SPECIES OF GLANDULOCAUDINE CHARACID FISH, HYSTERONOTUS MYERSI, FROM PERU By Stanley H. Weitzman Smithsonian Institution, Washington, D.C. and Jamie E. Thomerson Southern Illinois University, Edwardsville, Illinois INTRODUCTION On August 23, 1964, one of the authors (Thomerson), Jerry Anderson, Albert J. Klee, Emanuel Ledecky-Janachek, Winfield Rayburn, and Dr. Rich- ard L. Stone made a collection of fishes taken from a small stream tributary to the Pachitea River (Amazon drainage) at the northeastern outskirts of Tournavista, Province of Huanuco, Peru. Some of these were kept alive for experimental purposes and some were preserved. Among the fishes taken were representatives of the new species described here. Hysteronotus is a small genus of glandulocaudine characids most recently reviewed by Bohlke (1958) who described a new species, Hysteronotus hes- perus, amplified our knowledge of the only other known species, Hysteronotus megalostomus Eigenmann (1911), and redefined the genus. The characters of the new species described here and an analysis of additional specimens of H. megalostomus require a reevaluation of Bohlke’s contribution. [139] 140 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Hysteronotus myersi Weitzman and Thomerson, new species. (Figures 1, 2, 3, 4, and 5.) MarTeRIAL. Holotype, a male USNM 203697, standard length 49.00 mm. (no. 13 in table 1) from a small stream directly tributary to Pachitea River (itself tributary to Ucayali River) at northeastern outskirts of Tournavista, Huanuco Province, Peru. Elevation approximately 200 meters. Paratypes, originally in two lots, one lot of 8 specimens (nos. 1—4, 7-9, and no. 14 in table 1) with same data as holotype. Second lot of 5 specimens (nos. 5, 6, and 10-12 in table 1) raised in aquaria by Thomerson and bred from specimens in lot 1 and the holotype. Disposition of these lots is as follows: specimens nos. 5, 6, 7, 11, and 14 to Academy of Natural Sciences, Philadelphia (ANSP no. 112326 for nos. 5, 6, and 11, and ANSP no. 112325 for nos. 7 and 14); nos. 1, 2, 3, 4, 8, and 9 to United States National Museum, (USNM no. 203698); nos. 10 and 12 to Tulane University Collections (TU no. 56456). DESCRIPTION. Proportions as thousandths of standard length appear in table 1. Body elongate, laterally compressed, especially in males; body depth just anterior to dorsal and anal fin 2.7-3.4 times in standard length. Predorsal body profile slightly convex with slight concavity at nape; concavity deepest at posterior termination of supraoccipital spine. Along base of dorsal fin, body surface slightly arched dorsally to accommodate inclinator and other muscles of fin. Posterior to dorsal fin, body profile nearly straight with gentle downward slope to adipose fin. Posterior to adipose fin, body profile a straight level line to procurrent caudal rays in males and a slightly downward slope to these rays in females (compare figs. 1-4). Ventral profile to anus usually gently rounded with steepest inclination ventral to jaws. Ventral profile protrudes ventrally its greatest distance at point ventral to midlength of adpressed pec- torals. At anal fin origin (anterior termination of fin base) body profile gently convex, more so in males, and slopes upward to beginning of caudal peduncle just posterior to posterior anal fin termination. At that point profile straight and level or sloping slightly downward to procurrent caudal fin rays. Caudal peduncle deeper in males, least depth in standard length 6.5—6.8 times in males and 7.5—8.5 times in females (compare figs. 1—4). Length of head 3.7—4.0 times in standard length, this proportion not chang- ing greatly in different sized specimens. Specimen 49.6 mm. (longest) and one 28.3 mm. in standard length both with head 3.9 times in standard length. Eye rather large, somewhat larger in small specimens, 2.8—3.3 times in head length. Snout short, equal to, or shorter than, eye in specimens at hand, 3.3-3.9 times in head length. Snout appears proportionally longer in small specimens (table 1). Least width bony interorbital 2.6—-3.0 times in head length, always longer than snout length. Maxillary long, relatively slender, sloping ventrally and posteriorly to form an angle of 60-80 degrees to longitudinal axis of specimens. Maxillary length 141 HYSTERONOTUS MYERSI WEITZMAN & THOMERSON: Vor. XX XVIII] £6 Vel Orl 9CI ver vel mvt toddn jo yysuey OC Ten (02 ene oa ora Glee OCL Hol Beil v1 coral 98° 160 660-S80 £60 7260 880 S80 680 980 £60 7260 +60 +60 460 666 680 $60 yqso1oyur AUog os eZ0 640-990 9140 $40 +20 890 T20 890 140 640 +40 +40 990 490 890 S40 jnous JO YySsUsT LS°$80 S60-9L0 440 920 640 640 €80 680 ¢80 7280 160 480 060 880 680 60 949 JO A9JOWPICT Os 8SZ SO7-Shz 9Sc 62 O92 8c 192 8h2 682 992. 982 882 882 852 19¢ 892 peoy jo yysuoT TL L61 pec-991 If2 re O02 991 ZLI 661 861 O02 v8t +8t O02 T6t 961 £02 [esiop Jo 3YsIOH 98°SEl VA-Vole TET “HEI OSI OCD “OT wOCIe OSI OST Oar2Is sce ee ce leurs clmesic emu izeD otAjed Jo Yyysuo'T 61°22 PVE GNGs CCGMMML Ge tGSG CLC meV.GG) RC Ch (QIGG sn S\C Canna Crum C Commnt:C CommO)1C mummy mmmmcnen e1oqyved Jo Yysus'T 00° 7rT RST, SL WSE SIE Gy Sih IE GIAL SAR sia SISIE SEITE hv Hel gpunped Jo yysueT SeTedl ST) Tile DSi [eee SITES Hz PCA Ci nO ChE Mie ©) NO\ClTos mG) Giles C/U hc) ek ©).C, eA apunped yo yydoq L0°S9r E8p-S7hp Sly Sh 8h OSh 88h O09 99h f8h Clb 9b L9b 8th f9h LO aseq uly [epnes 0} 9Sseq [es1op IOLOJUV CC eSV plLy-ver Obb Shr ly brr 9Shb O9F Sth L9b 68h LSb 69h 9h veh cr [esiop 0} 9Aq I7L8S HLO-90G) SCLGN CSSe 19) OS) 26S 98S LG) 065) OLS SEUSS OS, Lbs RO FD [eue 0} ynoUs 98 FOV I8h-rSp €9h @Sh Liv LShb Leh Sb 8h 6Ly Shr LSb lb OSb 68h 857 stajod 0} ynous VI Vhe SOE CVG OSG 89 Le T42 992 197 797 = 867 Cle Gi Ga OL CaO SIC CLE =. 89 [etoysed 0} yous CV ess ZO9-195 LOS ~cSS FES 995 F/G “98s 78S 1005 9ZS™ = VOs) 8209) OSS Ose e165 [esiop 0} ynoug VULZe TIS=Z00 OVS NOLS MGLG OLS GCOum OSS Occ COG ay Oem NOOl sumitcicemmmn Dc mumCOGuumcrte yidap yseyvain Or OWgir GOS Me Cee Cas Pes ES OO “Woe Woe Chie Wee fe (‘Wu) YSU] prepueys ? ? ? } fo) 3} 6 6 fo) ) 5 } } 65 X9S adAj}o[0H asuey st L 9 S v ¢ Z I uvoyN rel! a IT Or 6 8 ‘NAldId ‘AIULLOAG OIMUDNT “DISLADUANO J {0 sSjuysjno Udsa{spay]sOU 4D UlMaA]S DUS :qyp20T ‘Yy)3ua] pavpunjs fo sy ypuDsNoY] Ut ISADAVA SNYOUOII}SAH f{O SJuamainspayy “T ATAVL 142 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 1. Hysteronotus myersi, new species, holotype, USNM 203697, adult male, 49.0 mm. in standard length. Small stream (tributary to Pachitea River, tributary to Ucayali River) at northeastern outskirts of Tournavista, Huanuco Province, Peru. (measured from tip of snout to posteroventral end of maxillary) 1.9—2.2 times in head length. Teeth 7-10, tricuspid, in single row on maxillary. Four speci- mens with 7, three with 8, three with 9, and two with 10 teeth on one side. Teeth cover about 60—90 percent of free edge of maxillary. Premaxillary teeth in two series; outer row with 3 teeth except two specimens with 4 teeth on one side and 3 teeth on other side. Inner row with 4 tricuspid or quincuspid teeth in six specimens and 5 teeth in eight specimens. Usually 4 large, most often tricuspid, anterior teeth on each dentary (3 teeth on one side of one specimen). In large male specimens third tooth from anteromedian tooth larg- est and with secondary cusps reduced or absent. Sometimes other large den- tary teeth with reduced cusps. Large teeth followed by 9-13 abruptly smaller and usually tricuspid teeth. No teeth on vomer, palatines, or pterygoids. Fontanels almost absent, that part anterior to epiphyseal bar (often called frontal fontanel) not detectable, that part posterior to bar (often called pari- etal fontanel but almost always surrounded by frontal as well as parietal bones and supraoccipital) narrow, almost completely closed joint in all specimens. Gill rakers moderately short, pointed, longest less than 2 length of pupil, 6-8 in upper limb, 10-12 on lower limb. Two specimens with total of 16, four with 17, four with 18, three with 19, and one with 20 rakers on entire first arch of one side. Circumorbital bones well ossified, covering entire cheek area, so-called “great suborbital” (actually infraorbital 3) completely covers cheek, leaving no space between it and preopercle. Suprapreopercular process extends dorsally to level of dorsal fin of fourth infraorbital bone (postorbital of some authors). In large specimens posterior border of fourth infraorbital con- Vor. XXXVIII] WEITZMAN & THOMERSON: HYSTERONOTUS MYERSI 143 Ficure 2. Hysteronotus myersi, new species, paratype, USNM 203698, adult female 32.6 mm. in standard length. Same data as holotype. tacts suprapreopercular process. Small individuals with space between these bones. Fifth infraorbital not in contact with preopercle. Scales of moderate size, cycloid with concentric circuli and about 8—15 grooves or radii on the exposed posterior field. Lateral line complete, perfo- rating 39 scales in three specimens, 40 in four, 41 in three, 42 in four. Lateral line with slight ventral curve on side of body anterior to position of dorsal fin. Lateral line continues to caudal base along midline. Transverse scale rows between anterior bases of dorsal and anal fins 14-15, often 7 above and 7 below lateral line. Predorsal scale count 21—23; axillary scales present above pectoral and pelvic fins. Basal scale sheath at base of anal fin of about 27—29 scales, usually 2 obvious horizontal rows anteriorly with some accessory scales. One longitudinal scale row along posterior third of anal fin base, and 1% rows at midregion of fin. Between bases of pelvic fin and anus, scales of both sides of body meet at midline in elongate median acute angle. Scales overlap acute midline angle only anteriorly near base of pelvic fins. No sharp keel between pelvic bases and anus. Area from anterior and posterior medial bases of pelvic fins along midventral line to isthmus, covered by scales. Ventro- lateral bases of pectoral fin without greatly enlarged scales. Figure 3 diagrams scales around caudal gland at base of lower caudal fin lobe. Two lateral line scales illustrated just dorsal to posterior base of gland. Glandular tissue and fossa-like structure of gland entirely supported by modified scales, fibrous connective tissue, and skin. Dorsal fin with ii, 9 rays in ten specimens, ii, 10 in four specimens; origin usually vertically over anterior base of anal fin, sometimes somewhat posterior to anterior anal fin base, nearer margin of opercle than base of caudal fin. Distance from tip of snout to anterior base of dorsal fin 1.7—1.8 times in 144 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 3. Hysteronotus myersi, new species, holotype USNM 203697. ) standard length. Dorsal fin profile rounded, not “straight topped” as reported for Hysteronotus hesperus by Bohlke (1958). Length of longest fin ray (= height of dorsal in table 1) 4.1-6.0 times in standard length; large males with greatest dorsal fin height (4.1 and 4.2 vs. 4.9-6.0 for all other specimens) (see also table 1). Height of dorsal fin appears sexually dimorphic, but rela- tively short in females and small males. Anal fin with v, 34 rays in two specimens, v, 35 in eight specimens, and v, 36 rays in four specimens. First unbranched ray not visible externally. Origin at or slightly behind midpoint of standard length. Distance from tip of snout to anal fin origin 1.6-1.8 (1.7 in eleven of fourteen specimens) times in standard length. Ventral margin of anal fin nearly straight in females, con- vex in males (compare figs. | and 2). Males with small dorsally recurved hooks on fourth through eleventh or twelfth branched anal fin rays (see fig. 1). Pelvic fin rays i, 6 in all specimens, distal end always reaching anterior basal termination of anal fin. Length of pelvics sexually dimorphic, 5.8 times in standard length in largest males, 6.3 in smaller male and 7.4—8.1 in females. Two types of contact organs present, bony hooks and bony spinelets. Hooks of one large, thick, hooklike excresence per ray segment. Spinelets of small, slender spicules of bone, one or more per ray segment. Spinelets easily broken, hooks not easily broken. Hooks confined to anal fin. Retrorse bony spinelets on males very small, and primarily on the first and second branched ray, even in largest male; not nearly as well developed or common as on Hysteronotus hesperus. Two to 3 or 4 spinelets per bony segment of each fin ray. Caudal fin with 10/9 principal caudal rays (17 branched rays) in all specimens; fin deeply forked. Males with small antrorse spinelets on dorsal edge of caudal rays, especially of lower lobe. No caudal spur. Vertebral counts 38-39 including ural segment. Two specimens with 16 precaudals and 22 caudals, remainder (except for one abnormal specimen for which there is no count) with 16 precaudals and 23 caudals. CoLtor IN ALCOHOL. Humeral spot present, large, diffuse, and centered above fourth through sixth scales of lateral line. Single narrow, black, straight line extends from dorsal border of humeral spot to center of caudal peduncle Vor. XXXVIII] WEITZMAN & THOMERSON: HYSTERONOTUS MYERSI 145 FicuRE 4. Hysteronotus myersi, new species, paratype ANSP 112326, adult female 30.9 mm. in standard length. Bred from specimens collected at the type locality. where in males line arches dorsally to end at junction of center of upper caudal peduncle muscle mass with upper lobe of caudal fin (fig. 1). Line may be more diffuse than shown in fig. 1, or may be pale in some females as in fig. 2. Caudal blotch present, weak, sometimes absent as in fig. 1; weakly present in fig. 2. In male 36.5 mm. in standard length caudal blotch moderately well de- veloped at center of union of caudal fin with caudal peduncle. Anterior border of blotch diffuse but with some dark pigment extending onto central caudal rays. Never as much pigment as in Hysteronotus hesperus. Compare figs. 1 through 4 with fig. 2, plate 3 in Bohlke (1958) for H. hesperus and fig. 4, plate 58 in Eigenmann (1927) for H. megalostomus. Most of body of Ay- steronotus myersi pale brown, slightly darker dorsally and lighter ventrally. Top of head dark brown with a narrow band of dark pigment extending from head to dorsal fin base. COLOR IN LIFE. One of us (Thomerson) has kept two pair of H. myersi in aquaria for several months. Their color may be summarized as follows. Females silvery with no prominent markings. Males with humeral spot and dusky stripe or band extending length of body. Both sexes with a distinct greenish iridescence. When males excited, lateral band darkens and 2 distinct pinkish spots appear at upper and lower base of caudal fin. FURTHER AQUARIUM NOTES. Fertilization is internal. Eggs slightly oval, approximately 1 mm. in diameter, and translucent. Eggs distributed on aquar- ium glass, plants, and rocks. More eggs appear attached to underside of plant leaves than on top. Very few eggs deposited near bottom of tank, usually in upper 7% of tank (5, 15, and 20 gallon aquaria). Spawning probably occurred in early morning and eggs appear deposited individually. SPECIES NAME. This species is named in honor of George S. Myers in recog- nition of his long and continued interest in characid fishes, and his frequent and helpful council to students of this complicated but fascinating group. Type Locatity. Hysteronotus myersi is known only from the type locality, a small stream directly tributary to the Pachitea River (Amazon drainage) at the northeastern outskirts of Tournavista, Huanuco Province, Peru. Most 146 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. TasLte 2. Measurements of Hysteronotus hesperus in thousandths of standard length. All specimens from eastern Ecuador. See Bohlke (1958, p. 35) for localities; compare original numbers. 1 2 3 4 5 6 7 8 Range Mean Holotype Sex Q Q 3 Q $ g ) g Standard length (mm.) 61.0 62.3 63.4 68.2 75.0 76.0 76.5 81.8 Greatest depth 328 320 331 320 316 349 318 332 316-349 326.75 Snout to dorsal 640 644 640 650 657 663 612 621 612-663 640.88 Snout to pectoral Die 265 268 258 259 286 274 Dips 258-286 269.00 Snout to pelvic 492 475 470 469 4606 470 464 454 454-492 470.00 Snout to anal 623 623 613 640 597 627 4636 #606 597-640 620.63 Eye to dorsal 528 518 503 532 539 542 510 514 503-542 523.88 Anterior dorsal base to caudal peduncle 386 401 427 384 376 423 402 412 384-423 401.37 Depth of peduncle 107 106 121 113 127 126 118 127 106-127 118.13 Length of peduncle 134127 114 4 1B 126 13) 120) 4S IS Length of pectoral DPS) 2S 265. AGE OS Bil, AB — 258-279 269.14 Length of pelvic 139 135 142 137 136 150 140 142 135-150 140.13 Height of dorsal 134 156 164 180 189 191 191 183 134-191 173.50 Length of head 246 229 248 232 240 268 240 BSD 229-268 244.37 Diameter of eye 069 069 060 066 065 072 059 061 059-072 065.13 Length of snout 067, 075 O77 O70) 076) (083) 108t 0738) SOb7—08 sO 25 Bony interorbit 090 088 087 O88 084 099 089 089 084-099 089.25 Length of upperjaw 107 096 096 101 105 116 094 110 094-116 103.13 Original number P304 Pil001 Pi607 P308 P309 Pil002 P306 P307 of the specimens were taken from pools in an area of alternating shallow pools and riffles where the width of the stream varied from 1 to 5 meters and from a few centimeters to 0.5 meters deep. The bottom was gravel and sand, with a few snags and broken limbs but no macrophytic aquatic plants. The stream was in a shallow ravine and was shaded by a dense canopy of small trees, brush, and vines. Downstream were several small waterfalls leading to an area of deeper boulder filled pools. Elevation at the type locality is approxi- mately 200 meters. A popular account of this locality is given by Klee (1965a). Fishes were not abundant, either above or below the waterfalls. Most of the specimens of Hysteronotus myersi were taken from midwater in the shal- low pools. Representatives of Rivulus peruanus Regan and loricariid catfishes were taken from the same pools, but the most abundant macroorganism was a river shrimp, Macrobrachium brazilense (Heller). Klee (1965b) charac- terized the water at the type locality as “. . Clear, clean, cool, moving water containing little or no vegetation. It is very soft, well oxygenated, and contains little in the way of dissolved materials.”’ These observations and the collection of the specimens of Hysteronotus myersi were made during the Vor. XXXVIII] WEITZMAN & THOMERSON: HYSTERONOTUS MYERSI 147 last week of August and first week of September 1964, during the dry season. RELATIONSHIPS. Bohlke (1958) reviewed in detail our knowledge of Hy- steronotus. At that time Bohlke distinguished the two known species, H. hes- perus and H. megalostomus, by contrasting 14 characters. In most of these characters, H. myersi appears closest to H. megalostomus but differs from that species in many other respects. A new comparison is made of these 14 charac- ters plus additional characters based on new data for H. megalostomus, new counts and measurements of H. hesperus (so that all counts and measurements are consistent), and data from H. myersi. Tables 1, 2, and 3 present a com- parison of measurements as thousandths of standard length for the three species. Character 1, size of males: Standard length 63.4—81.8 mm. in Hysteronotus hesperus; 36.5-49.6 mm. in H. myersi; 29.0-41.8 mm. in H. megalostomus. All males at these various sizes appear fully adult. Both H. megalostomus and H. myersi appear to be relatively small species and the large adult males of H. myersi lived at least nine months in aquaria with little growth and are presumably large specimens of the species. Hysteronotus megalostomus may reach a larger size and perhaps these size differences between adult males of H. myersi and H. megalostomus do not reflect a real species difference. Char- acter 2, bony hooks on anal fin of male: H. hesperus with true hooks on last unbranched and first 8-9 branched rays. Bohlke (1958) reported hooks ex- tending back to third ray from posterior termination of fin; however, these are spinelets. Hysteronotus myersi with hooks confined to fourth through about twelfth branched rays, mostly on fifth through eleventh. Hysteronotus megalostomus with hooks on first through eleventh to twelfth branched rays. Character 3, pelvic fin rays: Rays i, 7 in H. hesperus; i, 6 in H. myersi and H. megalostomus. Character 4, humeral spot: Small, round, clearly defined in H. hesperus; diffuse and large in H. myersi; large, sharply defined, and vertically elongate in H. megalostomus. Character 5, outer and inner rows of premaxilliary teeth: Outer premaxillary teeth 4—6, usually 5 in H. hesperus; 3-4, usually 3 in H. myersi; and 3-5 in H. megalostomus. Inner premaxillary teeth 4—5, usually 4 in H. hesperus; 4-5, slightly more often 5, in H. myerst; and 5-6, usually 6 in H. megalostomus. Character 6, maxillary teeth: Teeth 6-9 and very strong, dorsal teeth sometimes quincuspid, ventral teeth tri- cuspid in H. hesperus; 7-10 strong, tricuspid teeth in H. myersi; 5—6 strong (especially dorsally in large specimens) tricuspid teeth in H. megalostomus. Character 7, caudal fin of males split to its base: Not split to base in male of H. hesperus and H. myersi but split to base in H. megalostomus. Character 8, pectoral rays: Normally i, 11 in H. hesperus; i, 9 in H. myersi; and i, 9 (12 specimens) or i, 10 (6 specimens) in H. megalostomus. Character 9, lower limb gill rakers: 12 or, usually, 13 in H. hesperus; 10-12, usually 11, in H. myersi; and 10-12, usually either 11 or 12, in H. megalostomus. Character 10, eye in head length: 3.4-4.3 times in H. hesperus; 2.8-3.3 in H. myersi; and 148 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. TaBLeE 3. Measurements of Hysteronotus megalostomus in thousandths of standard length. Specimens 1-15 are from 3 to 4 km. northwest of Logoa Santa, Minas Gerais, Brazil. Specimens 16-18 are from a tributary of Rio das Velhas near Lagoa Santa, Minas Gerais, Brazil. 1 2 3 4 5 6 7 8 9 10 Sex 2 $ fc) g 3 2 2 3} 2 a) Standard length (mm.) 28:4) 29:0) 2933) 2957, 29:8) 2919) 29.030 Seolesmrcie: Greatest depth 214: 314 324" 320° 329 278 291 348 294 344 Snout to dorsal 612 600 604 £616 584 586 612 604 600 596 Snout to pectoral 264, 268) 272) 262) 278) 251 2645256 eS OS Snout to pelvic 431 428 4 438 449 438 438 443 447 440 Snout to anal 545) 50 S50) S15 5457522) 55S SSS OSE Eye to dorsal Silo) Ars 485 482 474 458 478 482 489 468 Anterior dorsal base to caudal fin base 382 427 406 407 413 398 415 400 400 420 Depth of peduncle 104 128 123 118 WA) 100 106 125 105 124 Length of peduncle 139 155 154 135 138 137 134 148 137 146 Length of pectoral 243) ©2418) 9259) 7239. 92525) 22452400229 ee Ome 20 Length of pelvic 104 126 137 128 121 100 Tals} USI 102 131 Height of dorsal 182 186 188 192 185 177 174 194 179 185 Length of head 246 231 246 246 248 244 256 239 239 242 Diameter of eye 083 076 38085 084 084 080 O80 082 083 086 Length of snout 076 O79 082 O81 084 O77 080 082 083 £083 Bony interorbit 090 097 097 094 098 087 097 098 096 096 Length of upper jaw 115 107 123 118 128 107 110 121 118 111 Color of pelvics black part part black black TABLE 3. Continued. 11 12 13 14 15 16 17 18 Range Mean Sex ) cy 2 3 rc) ) 3) a) SLinokhnel eon Coron) Sil Sail Seo sway aul s7.3 B40) 350 Greatest depth 352 339 298 342 SO SD ON oSS 312 274-356 320.50 Snout to dorsal 614 592 582 598 630 597 606 600 582-630 601.83 Snout to pectoral 272 268 254 262 271 276 BO) Ao 251-278 265.94 Snout to pelvic 446 442 432 452 440 434 456 414 414-456 439.72 Snout to anal 544 534 534 534 524 545 535 503 503-565 535.61 Eye to dorsal 478 472 475 479 505 483 462 446 446-510 477.72 Anterior dorsal base to caudal fin base 422 436 422 424 411 440 400 374 374-440 410.94 Depth of peduncle 133 122 102 125 nw 124 124 all 100-133 117.33 Length of peduncle 149 157 137 137 146 149 144 140 134-157 143.44 Length of pectoral 250 — 229 232 254 251 248 223 223-259 241.41 Length of pelvic 1 ISI O96, 137 27 1277S 108) 0 96=13)/ OES O Height of dorsal 190 181 176 183 196 179 197 177 174-197 184.50 Length of head 246, 9242 °° 220' 250) 239 242)” 236)" 228 220=256eez aed Diameter of eye 089 O81 078 O86 084 O78 082 074 074-089 081.94 Length of snout 079 O75 O75 O79 O74 074 O74 069 069-084 078.11 Bony interorbit 101 097 O90 095 083 093 088 O80 080-101 093.16 Length of upper jaw 123 115 112 116 4100 118 118 114 100-128 115.22 Color of pelvics black black yellow red black Vor. XXXVIII] WEITZMAN & THOMERSON: HYSTERONOTUS MYERS!I 149 2.8-3.1 in H. megalostomus. Character 11, length of upper jaw in head length: 2.3-2.6 in H. hesperus; 1.9-2.2 in H. myersi; and 1.8—2.3 in H. megalostomus. Character 12, length of pelvics: 6.2—7.4 in males, 7.2—7.6 in females of H. hes- perus; 5.8-6.3 in males, 7.3-8.1 in females of H. myersi; and 7.3-9.2 in males, 7.8—10.4 in females of H. megalostomus. Character 13, anterior dentary teeth: Quincuspid in H. hesperus, tricuspid in H. myersi and H. megalostomus. Character 14, fine bony spinelets of male pelvic fins: Numerous and on both sides of ray segments, usually several per segment in H. hesperus; not numer- ous, 1 or 2 per segment (sometimes up to 4 in H. myersi) and on one side of ray only in both A. myersi and H. megalostomus. Other characters useful in comparing these species are as follows: Charac- ter 15, numbers of vertebrae: 40-42 vertebrae in H. hesperus with 17 pre- caudals in all specimens, 23 caudals in one specimen, 24 caudals in two speci- mens, and 25 caudals in five specimens; 38-39 vertebrae in H. myersi, with 16 precaudals in all specimens, 22 caudals in two specimens and 23 caudals in eleven specimens; 40-42 vertebrae in H. megalostomus with 15 precaudal vertebrae in almost all specimens and 25 caudal vertebrae in eight specimens, 26 in seven specimens, and 27 in two specimens. One specimen of H. megalos- tomus with 14 precaudal and 27 caudal vertebrae. Character 16, scales around caudal fin: H. hesperus with 14 (15 in one specimen) longitudinal rows of scales around caudal peduncle, H. myersi and H. megalostomus with 18. Character 17, predorsal scales: This count difficult and inaccurate but Z. hesperus with 23-25 scales, H. myersi with 20-23, and H. megalostomus with 21-25. Character 18, tip of snout to dorsal fin origin in thousandths of standard length (see tables 1-3): Range of H. hesperus (612-663), mostly beyond ranges of other two species, (561-602) for H. myersi and (584-614) for H. megalostomus. Character 19, snout to anal distance in thousandths of standard length: Ranges of H. mvyersi (566-614) and H. megalostomus (507-565) partly contiguous, that of H. hesperus (597-640) begins at upper limit of range of H. myersi, not approaching that of H. megalostomus. Character 20, eye to dorsal distance in thousandths of standard length: Range of H. hesperus (503-542) nearly falls outside that of other two species (434-474 for H. myersi and 446-510 for H. megalostomus). Character 21, distance between dorsal origin and base of caudal fin in thousandths of standard length: Ranges of H. hesperus (384-423) and H. megalostomus (374-440) broadly overlap; that of H. myersi (428-483) stands apart from that of H. hesperus and overlaps upper range of H. megalostomus. Character 22, length of caudal peduncle in thousandths of standard length: Ranges of H. myersi (131-151), and 4. megalostomus (134-157) broadly overlap, while that of H. hesperus (114-134) barely overlaps their lower limit. Character 23, caudal gland: This gland is different in H. myersi and H. megalostomus (compare figs. 5 and 6). The 150 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. gland of H. hesperus is very similar to that of H. myersi (compare fig. 5 with fig. 6 in Bodhlke 1958. Also see discussion below under Status of the Genus Hysteronotus). Character 24, caudal fin split to its base: The caudal fin is normally split to its base in Pseudocorynopoma doriae and Hysteronotus megalostomus but it is not split in H. hesperus or H. myerst. The determination of the closest relative of H. myersi is difficult. As can be seen in the above characters, for example spinelets on the pelvic and caudal fins, number of ventral fin rays, number of outer row premaxillary teeth, number of cusps on maxillary teeth and large dentary teeth, number of pectoral rays, size of eye in relation to head length, length of the upper jaw, number of longitudinal rows of scales around caudal peduncle, proportional distance between snout tip and dorsal fin origin, proportional length of the caudal peduncle, and small scales at pectoral base, H. myersi more closely approaches H. megalostomus than it does H. hesperus. In a very few presumably im- portant characters, for example caudal fin not split to its base and caudal gland structure, H. myersi more closely approaches H. hkesperus than it does H. megalostomus. In a few characters, for example length of pelvics in males, and relative distances between the dorsal origin and caudal fin base, H. hesperus and H. megalostomus are more similar to each other than either is to H. myersi. In some characters, for example in number of precaudal vertebrae, structure of glandular tissue within caudal gland, size and shape of the humeral spot, no bony hooks on first through third branched anal fin rays, relatively long pelvics in males, ventrally convex anal fin margin, and extremely rounded, convex male dorsal fin profile, H. myersi is unique and unlike either H. hes- perus or H. megalostomus. With our present unclear knowledge of the phyletic and genetic stability of the caudal fin organ, or gland, it is difficult to weigh the significance of this structure in showing a close relationship between H. hesperus and H. myersi in contrast to the many characters that indicate H. myersi is closer to H. megalostomus. The caudal glands of glandulocaudine characids are in need of detailed comparative study, both in their histology and gross struc- ture. The best review of this subject to date is by Nelson (1964). See Géry (1964, fig. 4) for figures of Glandulocauda, and Nelson (1964, figs. 3-5) for figures of Pseudocorynopoma, Argyropleura, Gephyrocharax, Landonia, Cory- nopoma, and Glandulocauda. Eigenmann and Myers (1927, plates 84, 86, and 88) illustrated Corynopoma, Landonia, Pseudocorynopoma, and Gephyro- charax. Unfortunately, at present we do not know enough about either caudal glands or other characteristics of glandulocaudine characids to utilize these glands as valid, generic differences. The formation of the glandular tissue and scales in Hysteronotus megalostomus on the one hand, and H. hesperus and H. myersi on the other, is very different (compare figs. 5 and 6). The VoL. XXXVIII] WEITZMAN & THOMERSON: HYSTERONOTUS MYERSI 151 Ficure 5. Lateral view of caudal gland of holotype of Hysteronotus myerst. gland of the latter two species is surrounded in part by several modified scales and the glandular tissue lies over the lateral surface of two modified scales which curve dorsally over the glandular tissue forming a deep longitudinally oriented fossa. This fossa is open on its lateral and ventral surface. The gland of H. megalostomus is very different and has a very modified scale oriented ventrally around glandular material. The gland in H. megalostomus most closely resembles that of Pseudocorynopoma, see Eigenmann and Myers (1927, pl. 84, figs. 4-5). In gross dissection of H. megalostomus no obvious modified glandular tissue is present, but thickened skin lies over the dorsal surface of the ventral, furrowed scale, this skin being also attached medially to the fin rays. The same structure is found in Pseudocoryvnopoma doriae Perugia. The pouch of the gland in H. megalostomus extends anteriorly four to five scale 152 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. e e ‘ . 200 Sepeeeacee © a = B55 0.0.0 0 , 25 So . , »& rn a | > oH 1 , al ph a o> ett BPs tury. & a > J == = i @ wis i» co pie ao © Mist dePAal ss, 400 aes. | ae Wide Ph. fides (a Oe - Nasal: 8 lames i sal tvs |Ponewiy 4 _ - re Wud 4 Selle rime, mate — Sieh bes, Scag, Tint a vt a i 7 a ; i > ; 7 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Festschrift for George Sprague Myers Vol. XXXVIII, No. 9, pp. 157-162; 3 figs. December 31, 1970 REDISCOVERY OF THE LORICARITD CATFISH, ACESTRIDIUM DISCUS HASEMAN, NEAR MANAUS, BRAZIL By Robert L. Hassur Division of Systematic Biology, Stanford University, California 94305 More than fifty years ago John D. Haseman (1911, p. 319, pls. 50 and 51) described and figured a very small elongate loricariid catfish which he col- lected near Manaus, Brazil, as a new genus and species, Acestridium discus. He distinguished the genus from Farlowella principally on the basis of the expanded, disclike end of the snout and the presence of many series of delicate, spiny ridges (with intervening depressions) on all of the scutes. Haseman had 3 examples, the largest (holotype) was 72 mm. in total length. The types are now in the Field Museum of Natural History, Chicago. With the excep- tion of a restatement by Miranda Ribeiro (1912) of Haseman’s original de- scription and the inclusion by Gosline (1945) of the species in his catalog of Central and South American catfishes, the species, so far as is known, has not been reported again. During a recent visit, Dr. Jacques Gery presented the Stanford Collec- tion with 2 examples of A. discus (fig. 1 a,b) collected on 23 October 1965, by E. Fittkau and himself in a small tributary of the Igarape Castanha, which meets the right (southwestern) bank of the Rio Negro at a point two hours by boat upstream from Manaus. The specimens (SU 64202) measure 49 and 51 mm. in total length. A search of the Stanford Collection revealed 9 other specimens of the same species collected by Dr. Carl Ternetz in 1924 ESiZal 158 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. ¢ ae a Ficure 1. Acestridium discus, SU 64202, 49 and 51 mm total length; a, ventral; b, dorsal; c, lateral. Scale 1 cm. from the Igarape do Mai Joana, also near Manaus. These 9 specimens (SU 64095) range in total length from 48 to 65 mm. The total known range of the species then includes 3 tributaries of the Rio Negro near Manaus. The specimens fit Haseman’s description and illustrations with but one exception. The length of the snout, as usually measured from the anterior of the orbit to its tip, is about 2.25 rather than 4 times in the distance from its tip to the anus. Apparently Haseman measured the snout length differently, as this same measurement made on his drawings is about 2.25. Perhaps he, like Regan (1904, p. 303), used the distance between the tip of the snout and the anterior border of the naked area containing the mouth. This snout measurement, as determined from Haseman’s drawings, does yield about 4 times in the distance from snout tip to anus. Several characteristics in Haseman’s description require discussion. The large retrose hooks on the dorsal and ventral surfaces of the expanded tip of the snout (fig. 2) are set in 4 rows of 3-4 hooks. One row lies on either side near the margin of the disc, the other parallel to it and situated a short distance from the margin. The first pelvic spine of many of the Loricariidae, including Farlowella, is covered with many small hooks. In Acestridium, the structure is comblike (fig. 3) with enlarged toothlike hooks in about 3 series confined to the median surface of the spine, the rest of the spine being naked. With the fins erected, the hooks point toward each other, and, with the at- Vor. XXXVIIT] HASSUR: ACESTRIDIUM DISCUS 159 Ficure 2. Pelvic fin. 38. tached fin membranes and supporting rays curving inward toward the midline, the whole structure appears like a small basket. This structure was common to all specimens of both sexes examined. Although sex was not easily deter- mined because of the small size of the fish and, in some specimens, a lack of any recognizable feature of the gonads, the structure does not appear to be a secondary sexual character. Regan (1904, p. 198) describes the sexual characteristics in males of this family as being generally confined to enlarged bristles on the sides and top of the head or on the pectoral fins. But he makes no mention of the pelvic fins. The body (fig. 1b) is finely striped with brown above, pale below (fig. la) with the black spots on the lower lateral sides (fig. 1c) forming a dark border in the abdominal region and continuing completely across on the snout and the caudal peduncle. A broad, brown, lateral stripe (fig. Ic) extends posteriorly on either side of the head from the snout through the eye and tapers to a point above the posterior edge of the pelvic base. Each of the rather bulging eyes bears a smooth-margined operculum on the dorsal part of the iris. Acestridium discus differs from all known species of Farlowella not only in the sharply expanded snout-tip with its series of hooks and the definite spiny 160 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 3. Expanded disclike snout with rows of retrose hocks. 60. VoL. XXXVIIT) HASSUR: ACESTRIDIUM DISCUS 161 ridges of the scutes, which are fewer in number (25-27) than in any known species of Farlowella (33-34), but also in its generally smaller size; the rounded caudal fin; the absence of filamentous extensions on the main, ter- minal upper and lower caudal rays; the rounded (rather than acutely pointed) dorsal, anal, and pelvic fins; the straight vertical margins of the lateral scutes of the long caudal peduncle; five rather than six plates between the dorsal plate and the supraoccipital; and in having the ventral surface of the abdomen covered with two rather than three series of plates. LITERATURE CITED GosLINE, WILLIAM A. 1945. Catalogo dos Nematognath os de Agua-Doce da America do Sul E. Central. Boletim do Museu Nacional do Rio de Janeiro, N. S., Zoologica, no. 33, 138 pp. HasEMAN, JOHN D. 1911. Descriptions of some new species of fishes and miscellaneous notes on others obtained during the expedition of the Carnegie Museium to Central South America. Annals of the Carnegie Museum, vol. 7, no. 304, pp. 315-328. Mrranpa RIBEIRO, ALEPIO DE. 1912. Loricariidae, Callichthyidae, Doradidae e Tricomycteridae. Comissao de Linhas Telegraphicas Estrategicas de Matto-Grosso ao Amazonas. Annexo No. 5, Historia Natural, Zoologica, 31 pp. 1 pl. Rio de Janeiro. Recan, C. TATE. 1903. A monograph of the fishes of the family Loricariidae. Transactions of the Zoological Society of London, vol. 17, pp. 191-351. ee 94 7. } ——— » > 1 a . . e e ne, s=" r a 2 uy ~~ - oo a —4 yaa! as m ; a! ; ss athe lie Seid) aa his a 14 @-& b' Ty. ne 2 » mh nee ia nes oy 4240 icp c= it _ 1% haiig’g hed isle as 7 ‘iy tas be) we reins ie io" a ae nd : ine Bea mit (ee cy.cloigd sini r cate ee ee 17 Back with small, irregular, dark spots and dots; dorsolateral dark stripe extending from behind eye to just beyond forelimb or about 1% distance between fore- and hind limbs; tail with dark blotches above Crossobamon eversmanni Back with dark crossbars or longitudinal stripes — 16 Back with dark crossbars; hind limb reaching beyond axilla; ventral scales smooth == RGSS AE 8 ee Ne Doe TEE Sa aa ee Crossobamon lumsdeni Back with longitudinal stripes (or dots arranged in regular longitudinal rows) ; hind limb reaching axilla; ventral scales keeled; dorsolateral stripe extending from behind eye to, or almost to, insertion of hind limb; light dorsal stripe extending length of posterior 74 of tail, bordered by serrated brown dorsolateral FS oY ara en ne ie ey Nas res, He a BAe eer ens ne SOS Ba AN I onc Crossobamon maynardi Cycloid scales on back extending on to hinder part of head _____ Teratoscincus scincus Cycloid scales on back not extending beyond shoulders — 18 Cycloid scales on back feebly imbricate; about 100 scales round middle of |00) 0 hi eaeek ere ee ee Deere a ee eee Ch) SN ee Oe Teratoscincus microlepis Cycloid scales on back strongly imbricate; about 50 scales round middle of BSG op Sate See SS See oP nc eet ze eB Teratoscincus bedriagat Dorsal scales uniform, small, juxtaposed; granular, without enlarged tubercles present (Afghanistan specimens only) = Alsophylax pipiens Enlarged dorsal tubercles present among granular scales 20 Chingshields(postmentals) meals mt eee een 21 Chingishieldss presents ee oie Abe ee Be oe ee 22 Tail tapering gradually, covered below by small scales only; subdigital lamellae with several small tubercles, or denticulate on distal margin __ Bunopus tuberculatus Tail cylindrical, very slender, of almost uniform diameter from base to tip, with median series of enlarged subcaudal plates; subdigital lamellae smooth Ai Te te ee uta eek A OE oo ele ee Be Agamura persica Rostral excluded from border of nostri] — eee Agamura femoralis Rostralgiorming anterior bong |r Ol 1 OS ExT pees eee rene een nee 23 Tubercles usually present among granules of lower surface of thigh, in short row of 1-6, some often in contact with posterior row of large imbricate scales; males with continuous series of preanal and femoral pores —. 24 No subfemoral tubercles; males with preanal pores only _.... 26 37-40 abdominal scales across middle of belly (19-25 in distance across belly equal to length of snout) ; series of pores separated on midline by 1-2 scales not con- taining pores; 35-46 pores in males (total of both sides) — Cyrtodactylus species 23-34 abdominal scales across middle of belly (10-16 in distance across belly equal to length of snout) ; no separation between right and left series of pores; 24-41 POLEsHIne males; ((lotaliote boty Isic es) see meee eee eee een 25 24-29 strongly keeled, nonmucronate trihedral or subtrihedral tubercles in para- Vor. XXXVIII] LEVITON & ANDERSON: AFGHANISTAN HERPETOFAUNA 167 26. We fe 28. 2), 30. 31. 32. SOE 34, 30 36. vertebral row from occiput to level of vent; males with 28-41 (32-40 in Afghan specimens examined) preanal and femoral pores (total of both sides) ee ee eet Dn E coves Ris psn wl oe SN ee Cyrtodactylus fedtschenkot 20-23 strongly keeled, distinctly mucronate trihedral tubercles in paravertebral row from occiput to level of vent; males with 23-31 (24-29 in Afghan specimens examined) preanal and femoral pores (total of both sides) Cyrtodactylus caspius Subcaudal scales 1 head-width behind vent small, not enlarged and platelike (in distal part of tail, more than 2 head-widths posterior to vent, median series becomes enlarged, but much narrower than width of tail); tubercles on dorsal surface of tail arranged around middle of each caudal segment, not in terminal Sale MEO Veen eee Aeneas Bere eee eS Pi AR Poe ee SS 2 eee Cyrtodactylus russowii* Subcaudal scales 1 head-width behind vent enlarged, platelike, in single median series covering nearly full width of tail; tubercles on dorsal surface of tail form- in eatenimninal erin caotseachs cantdallese pt erty seen ee 27 25-47 abdominal scales across middle of belly (more than 12 scales across belly AMMECIStancerequal toplengthrotesnOut) ye Cyrtodactylus watsoni 15-23 abdominal scales across middle of belly (7-10 scales across belly in distance equalgtovlengthtotesnout) === eee Cyrtodactylus scaber No large paired shields on top of head, which is covered by granules or smali scales OnmatUberclesy se ee he. 0 ee 29 Enlarged paired plates on top of head (some granules may be present, but enlarged Shields pPTEC ONIN Ate) eee 54 Venter covered by imbricate scales, not granules; tongue broad and short, smooth or covered with villose papillae, not deeply forked; dorsum covered by imbricate scales or combination of imbricate scales and granules (agamids) — ~~ 30 Venter covered by small granules or juxtaposed quadrangular scales; tongue deeply divided, long and slender, smooth, retractile into sheath at base; dorsum covered with numerous small granules or juxtaposed scales (varanids) — 53 Wellimarkedudorsallicresteat least on neck ee Calotes versicolor ING IC OGSalR CLES eae ere ee ee See ese Se Se ee 31 Femoral pores present; tail strongly depressed through most of length, dorsal surface of tail with transverse rows of very large spinous tubercles rounded AED ASCY pee ene es See ees) ES kei, od PE ee ek pot ee, He ote SE 32 Femoral pores absent; tail depressed only at base, without transverse rows of very large spinous tubercles rounded at base, although rings of large spiny scales may be present, forming more or less distinct caudal segments —---_-= 33 Back without greatly enlarged pointed tubercles; caudal spines small, 20-24 in CLOSS=seniestat, basevObstalll eee ce sate eee ee eee Uromastyx hardwickii Back with transverse series of large pointed tubercles; caudal spines large, 8-10 in GEOSS=serleswat bases Ol» tall 22 = ee ee eee ee Uromastyx asmussi {Myicoriniien Gqaoecok TH. Cale Opoerine: WC Se 34 Tympanum concealed or absent, no visible external ear opening —— 45 Caudal scales in oblique rows, not forming rings; tympanum deeply sunk 35 Caudal scales forming more or less distinct rings; tympanum large, superficial __ 36 Dorsal scales subequal in size and disposed in regular rows Agama acgilis Dorsal scales unequal, large dorsal scales twice as large as smallest, irregularly anian ge Cee ler Se AE a eee ee Agama ruderata megalonyx Brilarecdsd orsalescalesssmoothyox taint hiya k ee] ccleaner en 37 inmiarcedadorsalescalesestrongly, keeled) ae ee Eee 39 168 Sie 38. 39. 40. 41. 42. 43. 44. 45. 46. 47. CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Caudal segments with 2 whorls of scales 2 head-widths posterior to vent bee unl Ones Tee Ole ee See x on Rade tins Reet eee Agama caucasica Caudal segments with 3 whorls of scales 2 head-widths posterior to vent, or seg- mentation*of tallindistinct) 228) 8 2 en ee eee 38 19-24 scales around tail at level of 5th complete whorl; scales on flanks distinctly larger than ventral and dorsolateral scales _._..........---.--- Agama badakhshana 25-35 scales around tail at level of 5th complete whorl; scales on flanks small, not larger than ventrals, grading into granular dorsolateral scales _. Agama himalayana Caudal scales small, 30 or more in whorl 5 rows posterior to vent — 40 Caudal scales large, usually less than 30 in whorl 5 rows posterior to vent 41 Forelimb and tibial portion of hind limb covered above with scattered, greatly enlarged spinous scales surrounded by much smaller or granular scales, large scales neither grouped in patches nor imbricate; largest scales on dorsum twice as large as largest ventral scales, vertebral and paravertebral groups of scales Hetero pencousix 82-4) eee Ie a ee ae ie ee ee Agama nuristanica Forelimb and tibial portion of hind limb covered with regularly arranged, enlarged imbricate and strongly keeled scales; largest scales on dorsum not larger than ventrals, vertebral and paravertebral groups of scales homogeneous ea oh aan re eh ERE ancora dt Pd SR ne a ee Agama tuberculata Dikinks ikvolsines ~milkicaecl Genes Agama nupta Flanks with numerous enlarged scales intermixed with smaller scales — 42 Scales of chest and throat strongly keeled and mucronate —— Agama erythrogastra Scales of chest and throat smooth or weakly keeled, not mucronate — 43 Each caudal segment 1 head-length posterior to vent, with 2 whorls of scales BS 2 he Se UREN she OER SAO oS 5 Mach wd Sed Ot Ct AE eee ee Agama caucasica Each caudal segment 1 head- tenet posterior to vent with 3 whorls of scales 44 Scales on snout and forehead smooth or faintly keeled; large mid-dorsal scales in 4 longitudinal rows, intermixed with smaller scales; greatly enlarged dorsolateral and flank scales in small, separate clusters forming longitudinal rows Pc. Heat WEN EE Fees yO See TUPI OAs Oa A Piet ee Agama lehmanni Scales on snout and forehead SaGREly keeled; large mid-dorsal scales in regular longitudinal rows not intermixed with smaller scales, though vertebral row of small scales may separate enlarged scale rows into 2 paravertebral groups; no greatly enlarged dorsolateral scales; enlarged flank scales forming large patch ongmid=flaink: pees se deren Mom lea deck ds | Oe. fete Landen A ae eS Agama agrorensis Large cutaneous fold at corner of mouth Phrynocephalus mystaceus ING) JE WAEKY CUI OVEONOIS THONG eae Woyeovere Ci Toovoyot ela 46 Dorsal scales heterogeneous, small scales intermixed with strongly enlarged scales __ 47 Dorsal scales subequal, homogeneous (but in P. reticulatus clusters or single scales may appear to be of different size than surrounding scales because they are swollen and tubercular and with upraised posterior margins; if striking differ- ence1s) observed: se@e47))) ees ee ee ee ee ee eee 48 Enlarged dorsal scales flat, not tubercular, posterior border not sharply upturned; sides of back of head and neck with long, flexible, spinous or fringelike scales; both sides of 4th toe with long, well developed fringes; tail without dark cross- bars, tip black on ventral surface in adults, in very small juveniles not black but with single black spot on ventral surface of tail Phrynocephalus luteoguttatus Some enlarged dorsal scales nail-like, large portion of scale raised free of back; sides of back of head and neck without long spinous or fringelike scales; 1 side VOL. 48. 49. Sil 52. 53. 54. 58. 59. 60. 61. XXXVITI] LEVITON & ANDERSON: AFGHANISTAN HERPETOFAUNA 169 of 4th toe with short fringe; tail with dark crossbars always present at least on Wentralesuntaces tipo te lacke== se nee ee ee Phrynocephalus scutellatus Flexible, fringelike scales prominent in temporal region 49 INominingelikesscalessins temporal ei omer eee ee ee 50 No crossbars on tail, tip of tail black; large, prominent black spots on back and top of head, group of 4 especially conspicuous in scapular region wa te tn Sa he ee I aan lee hia ee Phrynocephalus euptilopus Tail with distinct black crossbars on eal surface, tip black; no conspicuous larees blacks spots: onl dorsum Phrynocephalus interscapularis Nasal shields separated by 1-3 series of scales eae a 51 Nasal shields in contact or partly separated __ Scattered scales or clusters of scales on dorsum with upraised posterior margins, often swollen, tubercular; scales of upper surfaces of limbs and midline of back pLOMINeNtlys Kee) ed ye Phrynocephalus reticulatus boettgeri No upraised, swollen scales on dorsum; scales of upper surfaces of limbs and back smooth to indistinctly keeled in young, scales of limbs distinctly keeled in SEVCGAWU NES lh ce Sa See le On Sr ert Phrynocephalus maculatus Distinct, dark-margined, light dorsolateral stripe from posterior angle of eye along body onto tail; single, very elongate suborbital scale 2-3 times as long as ad- PEON NUS SSCL UES oa ee a a ie Se Ba eee ey Phrynocephalus clarkorum No light stripe along side of body; 3 suborbital scales of about equal size I GEN UC a Fe eA IE I hy nhs SE So AR Mt Phrynocephalus ornatus Tail round in cross-section, or slightly compressed posteriorly, without double- toothed crest above; abdominal scales in 110-125 transverse series from collar LOLdatORenOlng eee. Os 2 ee Ses | ee Se ee ewe Varanus griseus Tail compressed, with low, double-toothed crest above; abdominal scales in 90-110 iransversesseries) trom) collar fold! to) froin) 2 ee Varanus bengalensis Abdominal scales similar to dorsals; no femoral or preanal pores (skinks) — 55 Abdominal scales subquadrangular or quadrangular, in 8-18 longitudinal rows across venter, very distinct from dorsal granules; femoral pores present (except rin. JOA DoUS CVOTFOSEAOS)) (exeeiatels)). = 65 Body elongate; limbs present but greatly reduced, 3-4 fingers, 3 toes 56 Bodyeno teserpentine 4 —5i tN Pers. 5 il OCS) eee ene 57 VESUSYEY ENS) S) specs eA ee De sr ee Ophiomorus tridactylus SING CrGR AS ee ee ee eee Be We ee Ophiomorus brevipes Lower eyelid scaly; palatine bones separated on midline of palate 58 Lower eyelid with transparent disc or lids not movable; palatine bones meeting (jal Taal Ths Koni OF AS a Se ee ee ee 60 21-23 scales around body; postnasal present; single broad vertebral scale row, MUCH pLOAdes thanaG) ACen ty nO WS) ene Eumeces taeniolatus 26-30 scales around body; no postnasal; 2 median rows of dorsal scales broader (Haveivey. Ctl aVoyeC Svea) ata 8) foie eee ae ee Eel ee ee eee 59 eAZY SOUS MPOStIMemCall exe ss ee ee Eumeces blythianus* DRAZY COU Sm OSE ER GAS ge } oe EP ae \ te. i hg Pcs ee Weal. calinhnes (pi atianiiog ae (ee ree PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Festschrift for George Sprague Myers Vol. XXXVIII, No. 13, pp. 265-272; 2 figs.; 3 tables. December 31, 1970 A NEW GYMNOTOID FISH FROM THE RIO TOCANTINS, BRAZIL By Maarten Korringa Divison of Systematic Biology, Stanford University INTRODUCTION Amongst the gymnotoids in the fish collection of the Cailfornia Academy of Sciences in San Francisco, I found three unusual specimens which were collected in 1924 at Porto Nacional, Rio Tocantins, Brazil, by Dr. Carl Ternetz. Accord- ing to the keys in Ellis (1913) and Schultz (1949), these specimens fall into the genus Sternopygus, but the unusual appearance of the new fish suggested that it might be different. Accordingly, the three specimens were radiographed, and, after morphometric data were taken, one was cleared and stained for osteological study. Although these fish superficially resemble Sternopygus, their affinities do not appear to lie with this genus. I believe that the species described below re- quires generic recognition. Osteological nomenclature follows Weitzman (1962). ACKNOWLEDGMENTS [ am grateful to Dr. W. I. Follett and Mrs. Robert Dempster of the Califor- nia Academy of Sciences in San Francisco for their kind assistance in obtaining study materials from the fish collection there. I am also indebted to Professor George S. Myers, Mr. Leonard Compagno, and Dr. Warren C. Freihofer for their encouragement and criticism of the work at various stages. [265] [Proc. 4TH SER. CALIFORNIA ACADEMY OF SCIENCES 266 “WU Sep YSU] [LIOL “EbLrZ SVO ‘edAjopoy ‘xn7q snwmanjoyr4y SSS SSS ‘| Fang S Vor. XXXVIII] KORRINGA: GYMNOTOID FISH FROM BRAZIL 267 Nill \| omm Ficure 2. Archolaemus blax, holotype. View of jaws, indicating shape of tooth patches and showing mandibular teeth outside mouth. 2a. Upper jaw; 2b. lower jaw. Archolaemus Korringa, new genus Type species. Archolaemus blax Korringa, new species. Elongate compressed gymnotoid fishes, lacking caudal fin and dorsal thong. Frontal and parietal fontanels large. Tail extending beyond end of anal fin. Body completely covered with scales, with several rows of large scales near and below lateral line; back and lower parts of sides covered with very small scales. Head naked. Orbital margin free. Mesopterygoid with approximately 10 small villi- form teeth. Premaxillaries and dentaries each carrying numerous villiform teeth in a single patch on each bone (see fig. 2); a number of dentary teeth are out- side the mouth and project forward in larger specimens. Premaxillaries small, about one and one-half times as long as wide, about one-fourth as long as the maxillaries. Maxillaries more or less straight, virtually parallel to the body length of the fish. Branchiostegal rays 5, 4 on ceratohyal, 1 on epihyal. Posterior process of the lower pharyngeal bones with a patch of approximately 12 teeth. The upper pharyngeals each with a patch of about 8 teeth. In both upper and lower pharyngeals these teeth are small and villiform. Mesocoracoid absent. Certain lateral line canals of the head are much larger in diameter than others, and larger even than the canal of the body. Especially prominent are the nasal- supraorbital canal (anterior to the eyes), the infraorbital canal (excepting the posterior 2 bones, which are of small diameter), and the preopercular-mandibular canal. In conjunction with this increase in diameter, there is a progressive re- duction of the superficial walls of the canals, so that they resemble troughs roofed over by arches. 268 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER. TaBLe 1. Measurements in millimeters. Figures in parentheses are measurements ex- pressed as percentage of the distance from tip of snout to end of anal fin base. Holotype Paratype Paratype CAS 24743 CAS 24744 CAS 24745 Total length 435 (127) 313 (124) 235 (137) Snout tip to end of anal fin 344 (100) 253 (100) 172 (100) Snout tip to origin of anal fin 46 (13.4) 45 (17.8) 26.5 (15.4) Longest anal fin ray 16.5 (4.8) 14.5 (5.7) 8.5 (4.9) Snout tip to vent 23 (6.7) 18 (Ca ciL)) 17 (9.9) Snout tip to occiput S15) (9.7) 30 (11.9) 20 (11.6) Snout tip to pectoral fin origin 47 (ls}37/)) 45 (17.8) 25.5 (14.8) Length of pectoral fin base 8 (2.3) 5 (2.6) 3.5 (2.0) Snout tip to tip of pectoral fin U5 (21.9) 71 (28.1) 42 (24.4) Snout tip to anterior margin of eye 19 (5.5) 18 (Hail) 11 (6.4) Snout tip to rictus! 6 (1.7) — — 3.5 (2.0) Snout tip to posterior edge of opercle 43.5 (12.7) 40 (15.8) 23 (13.4) Snout tip to anterior nostrilt 4.3 (1.3) — — DH (1.6) Posterior nostril to eye 13.5 (3.9) BLES) (4.5) 6 (3.5) Distance between orbital margins 4.5 (ale3)) 3 (LZ) 2.6 (1.5) Depth at eye 22.5 (6.5) 19 (7.5) 12.5 (7.3) Depth at occiput 29 (8.5) 25 (9.9) 15).8 (9.0) Maximum depth of body” 40.5 (11.8) 40 (15.8) 19.5 (11.3) Width of mouth 6 (1.7) 5.5 (2.2 Dil (1.6) Width of head at eye 13.5 (3.9) 11 (4.3) 6 (3.5) Width of head at occiput 15}. (4.5) 14 (5.5) 9 (5.2) Anal fin rays 218 202 202 Pectoral fin rays, right side 19 20 20 Pectoral fin rays, left side 19 20 19 Lateral line scales® 137 143 146 Abdominal vertebrae 15 14 14 Caudal vertebrae 59 51 75 1 Snout of this specimen too severely damaged to take accurate measurements. ° Maximum depth of body is approximately at tip of pectoral fins. % Counted between dorsal margin of gill opening and end of anal fin base. Abdominal vertebrae 14 or 15; caudal vertebrae 51 to 75 in the three speci- mens; actual range doubtless greater. Two pyloric caeca present. Vent and genital papilla lie directly below the eye. Snout moderately long, conical; dis- tance from snout tip to eye approximately equal to distance from eye to posterior margin of opercle. Archolaemus blax Korringa, new species. (Figures 1, 2.) STUDY MATERIAL. Three specimens. Holotype: CAS 24743, male; 435 mm. in total length; Porto Nacional, Rio Tocantins, Estado de Goias, Brazil; collected by Carl Ternetz, February 8, 1924, Paratypes: CAS 24744, female full of eggs, Vor. XXXVIII] KORRINGA: GYMNOTOID FISH FROM BRAZIL 269 313 mm.; and CAS 24745, 235 mm. (cleared and stained with alizarin, in glyc- erine), both collected with the holotype. DeEscriIPTION. See table 1 for counts and measurements. Dorsal profile of head straight; ventral profile slightly convex to slightly concave. Tail somewhat flattened and ribbonlike (this may be an artifact, as all the tails possess, instead of vertebrae, a slender flexible rod, indicating that regeneration has taken place). One paratype (CAS 24744) is a ripe female; the eggs are slightly ovoid, about 1.5 mm. in length. The first gill arch of the right side possesses 7 to 10 nubbins representing rakers (9 in holotype, 10 and 7 in paratypes). In Sternopygus each nubbin has imbedded in it a number of very small spines or teeth. I have not been able to determine if this is true for Archolaemus, as the spines or teeth are not present in the cleared and stained specimens. Color in alcohol an even tan; area around orbital margin and tip of snout pale. There is no evidence of the dark brown banding found in many gymnotoids. ORIGIN OF NAME. Greek, archos, anus; laimos, throat, from the location of the vent under the eye; Latin, b/ax, doltish, in reference to the fish’s general appearance. DISCUSSION Regan (1911) places all short-snouted gymnotoids with frontal and parietal fontanels, but lacking caudal fins and dorsal thongs, in the subfamily Sternopy- ginae of his family Sternarchidae (properly Apteronotidae). I assigned Archo- laemus to this subfamily on the basis of 1) absence of mesocoracoid, 2) presence of large fontanels, 3) absence of caudal fin and dorsal thongs, 4) presence of mesopterygoid teeth. Sternopygus and Archolaemus are the only members of the subfamily with a free orbital margin. However, Archolaemus is distinct from Sternopygus in several ways. Sternopygus has a pectoral girdle with a short suture between the coracoid and the scapula, the scapular foramen being open anteriorly. In Archolaemus, the suture between the two bones is considerably longer, and a distinct scapular foramen is present. The former type, as Regan points out, is found in Steatogenys, whereas the latter is characteristic of Eigen- mannia. The pectoral girdle of Hypopomus resembles that of Ezgenmannia and Archolaemus in possessing a long suture, but lacks a conspicuous scapular fora- men. Archolaemus is further distinguished in having the anus and genital pa- pilla directly below the eye; in all specimens of Sternopygus examined, these were well posterior to the eye. The eye of Archolaemus is considerably larger (about 4.5 to 5.5 times in the distance between the eye and the posterior margin of the opercle) than that of Sternopygus (6.5 to 11). Other distinguishing features are: premaxillaries small, about one-fourth the length of the maxil- laries; as opposed to slightly over one-half in S. macrurus; few anal rays, generally less than 220, whereas Sternopygus has 234 to 320 (counts between 260 and 280 are typical). The maxillaries are fairly long and almost horizontal, 270 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. whereas Sternopygus has oblique maxillaries. The body cavity of Archolaemus is short (6.3 to 8.3 times in length to end of anal fin base), and few (14 to 15) abdominal centra are present; Sternopygus has a longer body cavity (about 3.5 to 5 times in length to end of anal fin base) and more abdominal centra (gener- ally 20 to 25). Archolaemus has a much longer snout than Sternopygus, the distance from snout tip to eye being about 1.2 times the distance from eye to occiput, as opposed to 0.7 for the latter genus. Archolaemus blax is distinguished from all species of Eigenmannia by the presence of a free orbital margin. Except for E. virescens, no species of Eigen- mannia that I have examined has the vent as far forward as the eye. Archo- laemus has a significantly longer snout than does EHigenmannia; snout tip to center of eye is contained 13 to 16 times in the length to the end of anal fin base in the former, 20 to 28 times in members of the latter genus. Avcholaemus has a long maxillary disposed almost parallel to the body length, whereas the maxil- laries of EKigenmannia are much shorter and vary from 45° to perpendicular to the body length. Archolaemus appears to have a proportionately longer gape than Higenmannia. No specimen of Eigenmannia that I have examined has teeth outside the mouth. Rhabdolichops (Eigenmann and Allen, 1942) is distinguished from Archo- laemus principally by its squamation, gill rakers, and general body shape. The back of Rhabdolichops is naked to a point about two-thirds of the distance to the end of the anal; anteriorly, the entire region above the lateral line is naked; posteriorly, only the top of the back. The body of Archolaemus is entirely covered with scales. Rhabdolichops has long, well developed gill rakers, a short snout, and a concave upper head profile. I do not yet have osteological material of this genus and therefore am not certain of its affinities. The edentulous Steropyginae (Hypopomus, Steatogenys, and Parupygus) appear to be highly distinct from the foregoing genera. Most notably, Archo- laemus, Sternopygus, Eigenmannia, and Rhabdolichops have similar lateral line canals while in Steatogenys, Hypopomus, and Parupygus (Hoedeman, 1962) all the canals are tubelike and of small diameter. The circumorbital tubes lack platelike stays. Many of the canals seem to be isolated from the bones with which they are normally associated in other groups of fishes; e.g., there is no apparent bony connection between the dentary and the chain of tubules lying ventral to it. Furthermore, these 3 genera lack teeth on the premaxillaries, den- taries, and mesopterygoids and not one has a long snout. Archolaemus further differs from Steatogenys in the absence of ‘‘mental filaments’ and lacks the banded color pattern found in this genus and in some species of Hypopomus. COMPARATIVE MATERIAL EXAMINED Sternopygus macrurus (Bloch and Schneider): SU 21997, 2 specimens, Bo- tanic Garden, British Guiana. Eigenmannia virescens (Valenciennes): SU Vot. XXXVIII] KORRINGA: GYMNOTOID FISH FROM BRAZIL 2A 54508, 2 specimens, Lagoa Grande, Brazil. Eigenmannia macrops (Boulenger) : SU 54473, 2 specimens, Sao Gabriel Rapids, Rio Negro, Brazil. Eigenmannia conirostris Eigenmann and Allen: SU 54461, 1 specimen, Lagoa Grande, Lower Amazon, Brazil, in alcohol. Rhabdolichops longicaudatus Eigenmann and Allen: SU 54377, 1 specimen, Santerém, Amazon, Brazil; SU 64076, 1 specimen, Cu- cuhy, Rio Negro, Brazil; both in alcohol. Hypopomus brevirostris (Steindach- ner): SU 24769, 1 specimen, Lago Gatin, Three Rivers Plantation, Panama. Steatogenys elegans (Steindachner): SU 22445, 2 specimens, Belém do Para, Brazil. Parupygus savanensis Hoedeman: Zoological Museum of Amsterdam 106,074, 1 specimen, Botopasi, Surinam River, Surinam. Except as noted, all are alizarin preparations in glycerine. SUMMARY Archolaemus blax Korringa, a new genus and species of gymnotoid fish is described. It is one of the toothed Sternopyginae, a group consisting of Sterno- pygus, Eigenmannia, and Rhabdolichops. These are united by the possession of hypertrophied lateral line canal bones in the head. Archolaemus shares many characters with Sternopygus and Eigenmannia, though its affinites lie more with the latter. LITERATURE CITED EIGENMANN, C. H., AND W. R. ALLEN 1942. Fishes of Western South America. Lexington, Kentucky, xv + 494 pp., 22 pls., 1 map. Extis, M. M. 1913. The gymnotoid eels of tropical America. Memoirs of the Carnegie Museum, vol. 6, no. 3, pp. 109-204, pl. 19-23. HoEDEMAN, J. J. 1962. Notes on the ichthyology of Surinam and other Guianas, 9. New records of gym- notid fishes. Bulletin of Aquatic Biology, vol. 3, no. 26, pp. 53-60. REGAN, C. T. 1911. The classification of the teleostean fishes of the order Ostariophysi, 1. Cypri- noidea, Annals and Magazine of Natural History, ser. 8, vol. 8, no. 43, pp. 13=31u pla: Scuumnz ic. 2 1949. A further contribution to the ichthyology of Venezuela. Proceedings of the United States National Museum, vol. 99, pp. 1-211, pl. 1-3. WEITZMAN, S. H. 1962. The osteology of Brycon meeki, a generalized characid fish, with an osteological definition of the family. Stanford Ichthyological Bulletin, vol. 8, no. 1, pp. 1-77. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Festschrift for George Sprague Myers Vol. XXXVIII, No. 14, pp. 273-288; 3 figs. December 31, 1970 ON THE TRAIL OF THE GOLDEN FROG: WITH WARSZEWICZ AND GABB IN CENTRAL AMERICA Ly Jay M. Savage Department of Biological Sciences, Allan Hancock Foundation, University of Southern California Those who have viewed at first hand the steep, dark-green, forest-covered slopes of the Cordillera de Talamanca-Chiriqui of Costa Rica and Panama, with their ever changing aspect of sun and cloud, moon and mist, bright blue sky and bright green mantle, driving rain and boiling fog, come away with a feeling of overpowering awe and mystery at the variety of nature and the magic of the human soul. It is not surprising that the primitive peoples in this region also regarded the mountains and their forests with mystical reverence, so near and yet towering abruptly upwards to 4,000 meters from their lowland valley habitations. Among the Bribri, Cabécar, Boruca, Changina, and Chiriqui, when the chicha has been drunk, the night grows late and dark, and the fires die down to burning embers, the wisest old man of the tribe tells his engrossed listeners of a beautiful miraculous golden frog that dwells in the forests of these mystical mountains. According to the legends, this frog is ever so shy and retiring and can only be found after arduous trials and patient search in the dark woods on fog shrouded slopes and frigid peaks. However, the reward for the finder of this marvelous creature is sublime. Anyone who spies the glittering brilliance of the frog is at first astounded by its beauty and overwhelmed with the excitement and joy of discovery; almost simultaneously he may experience great fear. The story contin- [273] bo af 4 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. ~ Cplicd Ficure 1. Upper, Josef Warszewicz, original by Artura Grottgera, now the property of the Department of Plant Geography, Jagellonian University, Krakow; lower, William M. Gabb, original, the property of the Academy of Natural Sciences of Philadelphia. Vor. XXXVIIT) SAVAGE: WARSZEWICZ AND GABB bo ~I Ut ues that any man who finds the legendary frog finds happiness, and as long as he holds the frog happiness will follow him everywhere. The story tellers record many men who have scaled the highest peaks and searched the darkest forests for even a glimpse of the golden frog, but only a few ever see it. Fewer still cap- ture the cherished creature and hold him for a few moments, and a very few are able to carry him with them for a longer period of time. One story tells of the man who found the frog, captured it, but then let it go because he did not recog- nize happiness when he had it; another released the frog because he found happi- ness too painful. Like the Indians of Talamanca and Chiriqui, each human being is also on a mission searching for the golden frog. Field biologists in particular seem always to be searching for mystical truth and beauty in nature, and frequently at some unperceived level, for that happiness promised by the Indian seers. The present paper is appropriately about two 19th Century scientists who joined this search in the very regions where the golden frog abounds, and we may assume that for a time, at least, they captured that joy guaranteed to beholders of the frog. INTRODUCTION Both Josef Warszewicz and William M. Gabb (fig. 1) were pioneer collectors of herpetological materials from lower Central America. Since they were among the first to sample the region, many of the animals they collected became types of previously undescribed species, most of which remain recognized as valid today. Neither of these men was a zoologist, and both collected in regions not visited again by herpetological collectors until the present century. Confusion and doubt as to the origin of their collections have clouded the issue of the validity of certain names and the synonymy of others subsequently described. In the present paper the routes followed by the two pioneers and the sources of their materials are delineated for the first time. ACROSS THE GREAT DIVIDE: WARSZEWICZ IN WESTERN PANAMA Josef Warszewicz was born in Litwie (Wilno), Poland in 1812. He apparently studied some botany at the University of Krakow. He took part in the Polish Revolution against Russia of 1830-31 and rose to the rank of officer. After the defeat of the Polish insurgents he left Poland. From 1840-1844 he worked as a gardener in the Botanical Gardens in Berlin. There he came to the attention of a Belgian, Van Houtte de Gandawy, who owned a large garden in Santo Tomas (St. Thomas), now Matias de Galvéz, Guatemala. Warszewicz was sent to inventory the garden and to collect materials for Belgian gardens. He sailed from Europe, December 5, 1844, and was active in Guatemala by March 1, 1845. He added many local species to his employer’s gardens and in 1846 began work for himself, and forwarded living and dry plants, especially orchids, to Europe. In 1848 276 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. a 82° 30) = ==. _ 82°09) | | | ~ WARSZEWICZ'S ROUTE +++ +++" ae Sy DUNN AND DURYEA'S ROUTE--~—-— = re em hy) = we < { ’ oe 3 — \ ®& ss = + Re SS S| S LAGUNA DE CHIRIQUI 4 < | a ©... «*@ROBALO : ae CHIRIQUICITO 9°00 = =) i 2 x = / \ oN Qe / ecHIRIOU! GRANDE. Ja / ¢ ae iS f 4 aC. PANDO cee fi Ne | 2 -* S : ‘ -=— - f S . = Set Ss Wy—~_ ot s ~ C, HORQUETA 7 a i \ wooo’ ( VOLCAN CHIRIQUI a” %, / BOQUETE © = mr - PANAMA DOLEGA @ 4 % Z — 48°30 “ a ik ' \ 5 gi a i DAVID @ . \ ? Ve Sf [ fe A } \ S( S, T A i A LC > S A, ¥) 7 d \ R SC 5 ; Ficure 2. Map of western Panama, showing Warszewicz’ route divide and principal localities discussed in text. across the Continental Vor. XXXVIIT] SAVAGE: WARSZEWICZ AND GABB bo ~r ~I Warszewicz undertook a major trip through Central America. He traveled by land from Guatemala to San José, Costa Rica, where he was situated by February, 1848. On March 1, he climbed Volcan Irazu. Later he arrived in western Veragua (Chiriqui), Panama, where he climbed Volcan Chiriqui and crossed over to the Caribbean coast. Most of the amphibians and reptiles collected by Warszewicz were taken in western Panama. In 1851 he was again in the Chiriqui region and later that year he proceeded to South America, being in Guayaquil, Ecuador, at the end of the year. Warszewicz spent 1852 in South America, pri- marily in Pert and Bolivia. He is known to have visited Lima, Peru, and was at La Paz, Bolivia, on June 15, 1852. At the end of the year, December 28, he was at Huancabamba on the headwaters of the Rio Maranon, upper Amazon drainage, Departamento Piura, Peru. He returned to Germany in October 1853 and became Inspector of the Botanic Gardens in Krakow. He died there December 29, 1866. (Regal, 1867; Rouppert, 1927). A bust of Warszewicz was erected in the Univer- sity Botanical Garden in Krakow about 1880, where it still stands. Herpetological materials collected by Warszewicz were deposited at Berlin, Vienna, and Krakow. The last city had been made the capital of a small free state after the Napoleonic wars in 1815. In November 1846, it was annexed to the Austrian-Hungarian Empire, following a revolt in Poland. Through exchange, some specimens came to the museum at Munich and to the British Museum. The Central American specimens are all from Panama and were taken in 1848 and 1851. The former collection seems to have gone to Krakow and Vienna, the last to Berlin. The long residence of Warszewicz in Berlin prior to his American travels explains deposition of specimens there, probably as the result of long-time contacts. Apparently he loaned and gave some material to the Vienna Museum on his establishment in Krakéw in 1843, since the latter city was then part of the Austrian-Hungarian state. Fortunately Warszewicz’ route through the Chiriqui massif may now be traced with some accuracy (fig. 2). Information provided by Wagner (1863) and his map clearly define the route from David on the Pacific slope across the divide to the Laguna de Chiriqui. Wagner records that Warsze- wicz penetrated the interior of Chiriqui and traversed the great Cordillera to the Atlantic shore. Regel (1867) noted that Warszewicz climbed the 16,000 foot Volcan Chiriqui in 1848. Wagner (1863), using the same guides and carriers employed by Warszewicz, followed the same trails to the Chiriqui highlands. This route runs from David through Dolega and then up to Boquete (1158m.), from where there were two trails leading to the Caribbean shore. One trail skirted the east slope of Volcan Chiriqui and continued to Ranchos de Robalo, the other passed around the east slope of Cerro Horqueta and continued to the mouth of Cabbage Creek (Rio Guarmo) near present day Chiriqui Grande. Warszewicz certainly followed the Boquete-Robalo trail, since a branch from it leads to the top of Volcan Chiriqui (3478m.). Nevertheless, he may have returned via the other route. Emmett R. Dunn and Chester B. Duryea seem to have followed the 278 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47H Ser. latter trail from Chiriqui Grande to Boquete, in 1923 when they became the first herpetologists to re-collect several of Warszewicz’ species. HERPETOLOGICAL SPECIMENS COLLECTED BY WARSZEWICZ Most of the amphibians collected by Warszewicz were described by Oskar Schmidt (1857) and more extensively described and illustrated by him in 1858. The following are involved, with the Krakow Museum in the Department of Systematic Zoology, Jagellonian University (KM) numbers listed. These speci- mens were presented to the collection in 1870 and were examined by E. R. Dunn in 1928. Some of them are still extant. Location of other types and other Wars- zewicz material noted in preparation of this report is also indicated, but probably is not complete. Abbreviations for other collections are: Zoologisches Museum, Berlin (B); British Museum (Natural History), (BM); Zoologischen Museum, Hamburg (H); Zoologischen Staatssammlung in Mtinchen (M); Naturhistor- isches Museum Wien (W). An asterisk (*) indicates a new species. SPECIMENS COLLECTED BY WARSZEWICZ *Leiuperus sagittifer. New Granada (Colombia). *Txalus warschewitschii. KM 1006/1338; near Volcan Chiriqui, between 6000 and 7000 feet (4500-5250 feet = 1370-1600 m.). *Hyla pugnax. KM 1009/1339; Rio Chiriqui near Bocas del Toro. “Hyla splendens. KM 1008/1340 9 :Rio Chiriqui near Bocas del Toro. *Hyla molitor. KM 1010/1341, 24 6; W 16494, female designated as lectotype by Savage and Heyer (1969):Rio Chiriqui near Bocas del Toro. *Hyla molitor marmorata. KM 1010/1342 9 :Rio Chiriqui near Bocas del Toro. *Hylodes fitzingeri. KM 1012/1343; Mountains of New Granada (Panama), 4000 feet (3000 feet = 915 m.) ; now lost. *“Dendrobates speciosus. KM 1017/1345 nine specimens; W _ one specimen:trail between Bocas del Toro and Volcan Chiriqui, 5000-7000 feet (3777-5250 feet = 1150-1600 m.) ; now lost. *Dendrobates pumilio. KM 1018/1346:trail between Bocas del Toro and Volcan Chiriqui, 5000-7000 feet (3777-5250 feet = 1150-1600 m.) ; now lost. *Dendrobates lugubris. KM 1016/1347:trail between Bocas del Toro and Volcan Chiriqui, 5000-7000 feet (3777-5250 feet = 1150-1600 m.) ; now lost. Bufo margaritifer. Between Bolivia and Pert, 3000 feet (2250 feet = 685 m.). *Bufo pleuropterus. KM 1030/1348:between Bolivia and Pert, 3000 feet (2250 feet = 685 m.). *Bufo veraguensis. KM 1032/1350; New Granada, Provincia de Veragua. *Bufo simus. BM 95-9-14.6; H 1527; KM 1029/1351, 5 specimens (now lost); M 543/20; W 16521:Rio Chiriqui near Bocas del Toro. *Hylaemorphus dumerilii. KM 1014/1345:New Granada, Provincia de Chiriqui, 8000 feet (6000 feet = 1830 m.). *Hylaemorphus bibronit. KM 1015/1355; New Granada near Panama, 2000-3000 feet (1500- 2500 feet = 460-760 m.). *Phirix pachydermus. KM 1013/1356; Western New Granada near Buenaventura, 5000 feet (3777 feet =1150 m.) ; now lost. VoL. XXXVIII] SAVAGE: WARSZEWICZ AND GABB 279 Other specimens collected by Warszewicz. At Krakow: Basciliscus mitratus. KM 932/1317 America. Stenostoma albifrons. KM 962/1296 America. Cyclophis aestivus. KM 981/1270 America. Pelamis bicolor. KM 989/1304 Pacific Sea. Lacerta muralis viridis. KM 1019/1270 America. Bufo vulgaris. KM 1020 America. Phyllomedusa hypochondrica. KM 1024/1344 Guyana. Bufo chilensis. KM 1031/1349 Bolivia. At Berlin: *Rhinotyphlops albirostris. B 9529, 2 specimens; Veragua (Peters, 1857). *Anolis humilis. B 500; Veragua (Peters, 1863a). * Anolis intermedius. B 503; Veragua (Peters, 1863a). *Hyla sordida. B 3141; Veragua (Peters, 1863c). *Hyla punctariola. B 4918; Veragua (Peters, 1863c). *Strabomantis biporcatus. B 3222, 3330; Veragua (Peters, 1863b). Bufo haematiticus. B 3404; Veragua. Bufo typhonius. B 3442; Veragua. Most of the animals collected by Warszewicz are from what is today western Panama, but in his time constituted the Provincia de Veragua of the country of Nueva Granada (Colombia). Today the old Veragua comprises the Provincias de Veraguas, Chiriqui, and Bocas del Toro. In Warszewicz’ day, western Veragua was called Chiriqui and the Atlantic lowlands were called Bocas del Toro. Several corrections seem necessary in dealing with the data associated with his specimens. First, all altitudes listed are extremely high and well above the known distribu- tions for the species. As I have previously pointed out (Savage, 1968) 19th Century Polish feet contained the equivalent of only nine English inches. There- fore I have given the corrected elevations in parentheses above. Regel’s (1867) report of Warszewicz’ climbing 16,000 foot Volcan Chiriqui as previously cited shows the same point, since the mountain is 3478m. (11,311 feet) in height. Even these figures are out of the altitudinal range for several species, but since they were probably estimated, the differences are not extreme after the corrections have been made. Several forms described from Warszewicz’ materials by Oskar Schmidt have never been retaken in Central America but, because of the lack of details regard- ing his route and the inaccessibility of the area on the continental divide visited by him, herpetologists have assumed that these animals would ultimately be rediscovered in the field. Recently, I (Savage, 1969) demonstrated that one species, Bufo veraguensis, was based on a mislabeled Peruvian or Bolivian toad. At least three others, Hyla splendens, Hyla molitor, and Hyla molitor marmorata may similarly be removed from any list of Central American amphibians. Savage 280 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. 83°00 10°00” GABB’S ROUTE ...... *Cahuita Puerto Viejo - SA o . o gio Lo Amubric, (\ : . ES : 9°30 Oi ea fi eee FO Guabito ys C. Kamuk PANAMA COSTA RICA SSS es 0 es Ce ss Ss a GS aenarer e Soee . = 9° 00 Ficure 3. Map of southeastern Costa Rica, indicating area of Gabb’s collections and the route followed in his ascent of Cerro Utyum. Vor. XXXVIIT] SAVAGE: WARSZEWICZ AND GABB 281 and Heyer (1969) have shown that the latter two are totally unlike any Central American forms. Very likely they are also mislabeled South American frogs. The type of Hyla splendens appears to be a female of the genus Gastrotheca. Charles F. Walker, the leading student of this genus, informs me that the type is very similar to some Peruvian Gastrotheca species and unlike any Panamanian or Colombian form. GABB AND THE EXPLORATION OF THE TALAMANCA William More Gabb was born in Philadelphia on January 20, 1839. He is the subject of a biographical memoir of the National Academy of Science (Dall, 1909). Only details omitted from the memoir or matters directly related to his Central American experience are recounted here. Gabb was interested in geology and mineralogy and became associated with the California Geological Survey in 1862. As part of this work he spent the period 1862-1867 in California and was involved in the Survey’s study of Baja California in the latter year. In 1869-1870 Gabb was active in geological work in Santa Domingo. Gabb came to Costa Rica in February, 1873, to undertake a study of the geog- raphy, geology, resources, and climate of the southeastern section of the country, the Talamanca (fig. 3). During the 19 months of his contract, 17 were spent in the field (until August, 1874). He returned to the United States in 1876 whence he again visited Santa Domingo. Malaria apparently contacted in Costa Rica was inflamed in Santa Domingo and he ultimately succumbed to tuberculosis of the lungs after his final return to the United States in April, 1878. He died in Philadelphia May 30, 1878. Gabb’s fantastic activities during his Costa Rican stay are summarized in his reports (Gabb, 1875, 1877, 1913a, 1913b; Pittier, 1875, 1913). Most of his work was centered on the Valle de Talamanca, the region of the Rio Sixaola drainage. The upper portion of this area: The Valle de Rio Telire and the drainages of the Rio Urén, Rio Lari, Rio Coen, Rio Telire, and Rio Taberi, forms Alta Talamanca. The lower part of the valley from a line between Uatsi and the mouth of the Rio Yorkin to the coast is Baja Talamanca. The towering spires of the Cordillera de Talamanca border the Valle de Talamanca on the northwest. The Valle de Tala- manca was the original Spanish settlement in Costa Rica, where La Ciudad de Santiago de Talamanca was founded, near the present site of Suretka (60m.) on October 10, 1605. In Gabb’s day the central settlement of the area (it had the only church and was the home of Mr. John H. Lyon, an American, who had administrative responsibility for the district) was San Bernardo de Sipurio (70m.) between the Rio Suedi and Rio Urén about 3 miles above the mouth of the latter. This village and the Catholic church were destroyed by a flood of the Rio Lari in 1909. A new mission was established at Amubri (75m.) in 1910 and serves as the central settlement in the area today. In 1873-74 about 1240 people lived in the 282 CALIFORNIA ACADEMY OF SCIENCES [Proc, 47TH SER. region. Gabb was accompanied on most of his many trips through the area by two Costa Rican collectors, Jose Zeledén and Juan Cooper, both later famous naturalists in their own right, who collected most of the vertebrates. Gabb married an Indian girl, Victoria, and one son Guillermo was born to this marriage in 1874 or 1875. At least three grandchildren, Alfonso, Melania, and Francisco Gabb were alive in 1964 when I visited the Talamanca. Several great-grandchildren were also living including a Victoria Gabb, an exceedingly beautiful girl, who may have recalled the Indian beauty who married Gabb. Gabb visited almost every locality in the valley. As part of his fieldwork (Gabb, 1913b: 105-106, 114; 120-122, 127-128) he attempted to climb Pico Blanco (Cerro Kamuk) the highest peak (3554m.) in the southern Talamanca- Chiriqui range. Gabb tells it all—‘‘We followed hunter’s trails over a long, narrow, and very crooked ridge between the Urén and the Lari to a place called Bitsung- wo-ki, often scaling precipices, climbing around rocks, and in some parts scram- bling over bad places by means of ladders and bridges made of sticks placed there for this purpose. Beyond Bitsung-wo-ki, but two men had ever gone, and with one of them for a guide, we were forced to climb down to the Lavi River, and ascend the mountains on the other side, to avoid impassable rocks. At the end of seven (7) days, we found ourselves on the side of a peak, which we as- cended, made our observations, and returned.”’ His party consisted of 21 persons and subsisted mainly on pldtanos. They were on the peak June 13, 1873, after starting the ascent June 6. Gutiérrez (1960) has conclusively shown that Gabb, by detouring up the Rio Lari, actually ascended Cerro Utyum (3084m.) (Cerro Cruz del Obispo) rather than Kamuk. The route followed by Gabb, his altitude record for the peak, 9562 feet (2915m.), as well as his observations (1913b: 106) as pointed out by Gutiérrez (1960) and confirmed by Carballo (1960) who scaled Kamuk, sub- stantiate this conclusion. Gabb apparently returned to Alta Talamanca via the Rio Lari. I ascended the latter river in 1964 to a point approximately where Gabb crossed over the ridge from the Rio Urén. This place is 3 days hard hiking from Amubri and lies at 800m., near the juncture of the Rio Dipari and Rio Lari. In view of these data, none of Gabb’s animals should be listed from Pico Blanco (Cope 1875, 1876) but rather from Cerro Utyum. One of the principal supporters of Gabb’s explorations was the legendary Costa Rican entrepreneur Minor C. Keith, then manager of what became the Costa Rican Northern Railroad, that today connects Puerto Limon and San _ Jose. Keith began the planting of bananas along the rail lines, originally to keep the railroad hands busy and to provide food. Gradually bananas became the basis for the development of the United Fruit Company. The Compania Bananera began to exploit the Valle de Talamanca in 1916. Poor and thin soils led to re- duction of activity in 1922. The Valle was abandoned to local farmers in 1925. A railroad that connected with the United Fruit Company lines in the Bocas del Vor. XXXVIIT] SAVAGE: WARSZEWICZ AND GABB 283 Toro region of Panama, at Guabito, formerly extended up river past Suretka. The bridge across the Rio Sixaola above Suretka was washed out in 1925 and the rails abandoned. In 1964 the railroad still ran from Sixaola to Volio (Uatsi). A truck road connects Puerto Viejo and Cahuita to Fields where another truck road runs to beyond Suretka. A jeep trail runs north from this road to Pandora in the Valle de Estrella. HERPETOLOGICAL SPECIMENS IN THE GABB COLLECTIONS The herpetological materials from Gabb’s explorations were deposited at the United States National Museum (US) and reported on by E. D. Cope (1875, 1876) in a large monograph. Many examples served as types of new taxa as indicated below by an asterisk (*). Cope’s paper was originally published as a separate, with a limited letterpress run of 50 copies on November 26, 1876. The journal run (Cope, 1876) appeared early the next year. This original report on Gabb’s material has been reissued as a special number of the journal O’Bios and may be purchased from the Departamento de Biologia, Ciudad Universitaria, Costa Rica. Because the Gabb material is well known I have indicated catalog numbers only for type materials. Unless otherwise denoted all specimens are from Provincia de Limon, Canton de Limon in Costa Rica. SPECIMENS COLLECTED BY GABB Siphonops mexicanus. Holotype, US 29762; Paratype US 29763; Limon (described as new species Siphonops proximus Cope, 1878). Opheobatrachus vermicularis. One specimen from Cerro Utyum, 6000 feet (1830 m.); 2 ex- amples from lower country 20 miles (30 km.) from Coast. Oedipus moro ?. Eastern slope Cerro Utyum. *Cranopsis fastidiosus. Lectotype, US 32585; paratypes US 32584, 32586-87; Cerro Utyum, 2500 feet (760 m.). *Crepidius epioticus. Cerro Utyum, 5000 feet (1520 m.) (type lost), Savage and Kluge, 1961. *Ollotis coerulescens. Cerro Utyum, 3000-5000 feet (915-1520 m.) (type lost). “Bufo auritus. US 30676; east coast region (substitute name Bufo gabbi Taylor, 1952). Bufo valliceps. US 30592; eastern Costa Rica (described as new species, Bufo melanochloris Cope, 1878). Bufo agua. Eastern coast. Bufo haematiticus. Sipurio. Atelopus varius. Cerro Utyum and lower country. Dendrobates typographus. Low country about 10 miles (15 km.) inland. Dendrobates tinctorius. Lower country. *Dendrobates talamancae. Near Old Harbour on east coast (type lost). *Hyla gabbii. US 30658—59; near Sipurio. *Hyla uranochroa. US 30651; near Sipurio. *Hyla nigripes. US 30685-86; Cerro Utyum, 5000-7000 feet (1525-2135 m.). “Hyla elaeochroa. Lectotype, US 30689, paratypes US 30688, 30690; east foot of mountains near Sipurio. “Hyla punctariola pictipes. US 30652; Cerro Utyum, 5000-7000 feet (1525-2135 m.). 284 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. *Hyla punctariola monticola. US 30661, Cerro Utyum. *Phyllobates hylaeformis. US 30687; Cerro Utyum, 7000 feet (2135 m.). *Lithodytes podiciferus. US 30662, 30665-75 (US 30663 now at Harvard, US 30664 now at Michigan) ; Cerro Utyum, 5000-7000 feet (1525-2135 m.). *Lithodytes muricinus. Cerro Utyum (type lost). *Lithodytes habenatus. Cerro Utyum (type lost). *Lithodytes melanostictus. US 30608; Cerro Utyum, 7000 feet (2135 m.). *Lithodytes megalocephalus. US 32578; spur of Cerro Utyum, 6000 feet (1830 m.). *Lithodytes gulosus. US 32590; spur of Cerro Utyum, 6000 feet (1830 m.). *Hylodes cerasinus. US 32572; eastern slope of Cerro Utyum. Gnathophysia ocellaba. East side of the Cordillera. Ranula brevipalmata. Cerro Utyum. Mocoa assata. Old Harbour. *Mabuia alliacea. US 30619-20; from the low country. Mabuia cepedei. Below Sipurio. *Chalcidolepis metallicus. US 30568; Provincia de Alajuela, Fila de Aguacate. *Amiva gabbiana. US 32614-16; Old Harbour. Gerrhonotus fulvus. Summit of Cerro Utyum. Sphaerodactylus glaucus. Near Sipurio. Thecadactylus rapicaudus. North of Rio Estrella or North River. Anolis copei. Old Harbour (Puerto Viejo). Anolis trochilus. Talamanca. “Anolis pachypus. US 30683; slope of Cerro Utyum. *Anolis oxylophus. US 30556-57; Costa Rica. Anolis intermedius. Anolis capito. Old Harbour. Corythophanes cristatus. Sipurio. Iguana rhinophila. Low country. Basiliscus vittatus. Sipurio. *Basiliscus plumifrons. US 32622-6; Sipurio. *Niphosoma annulatum. US 32580. Boa imperator. Foot of mountains. *Leptognathus argus. US 30656; Sipurio. *Leptognathus pictiventris. US 30657; eastern Costa Rica. Leptognathus nebulata. Sibon annulatum. Old Harbour. Oxyrrhopus plumbeus. Low country Oxyrrhopus petola. Sipurio. *“Leptophis aeruginosus. US 30684; low country. *Leptophis saturatus. US 32563; Sipurio. Leptophis praestans. Sipurio. *Dendrophidium melanotropis. US 32597. Drymobius boddaerti. Talamanca. Herpetodryas carinatus. Low country. Spilotes corias. Talamanca. *Spilotes chrysobronchus. US 30623; coast region. Coniophanes fissidens. Sipurio and Old Harbour. Rhadinaea decorata. Sipurio. Erythrolamprus venustissimus. Sipurio. Xenodon angustirostris. Sipurio. VoL. XXXVIIT] SAVAGE: WARSZEWICZ AND GABB 28 OL Stenorhina ventralis. Old Harbour. *Contia pachyura. US 30618; Sipurio. *Catastoma psephotum. US 62972; Cerro Utyum, 5000-7000 feet (1525-2135 m.). Elaps circinalis. Talamanca. Teleurapsis schlegelii. Eastern Costa Rica, Old Harbour to 5000-6000 feet (1525-1830 m.). Bothriechis nigroviridis. Cerro Utyum. “Bothriopsis proboscideus. Sipurio (type lost). Bothrops atrox. Coast region. *Lachesis stenophrys. US 32479; Sipurio. Sphargis coriacea. Puerto Limon. Cinosternum leucostomum. Old Harbour and Sipurio. *Chelopus gabbii. US 45905. *Chelopus funerus. US 45900-01; 56134-35; Puerto Limon. Old Harbour, located on the coast between Punta Cahuita and the Boca de Sixaola, is now referred to as Puerto Viejo de Limon. A FINAL WORD At certain levels both Warszewicz and Gabb were successful in their quest. The modern observer who has been over some of the same ground can only marvel at the courageous determination, dedication, and curiosity of these scientific pioneers. In regions sparsely settled, without roads or other communication, dominated by primitive and rugged terrain, demanding climate and debilitating disease, they still prevailed against all odds to open a new and exciting world to those that followed. They could do no more. ACKNOWLEDGMENTS My great thanks go to Dr. Henryk Szarski, Department of Comparative Anatomy, Jagellonian University, Krakow, for data and advice regarding Wars- zewicz and his career. He also supplied the accompanying photograph of the Polish botanist. Norman J. Scott, Department of Biology, University of Con- necticut, Storrs, Connecticut, was my field companion during an arduous follow- ing of Gabb’s trails in the Talamancas and I appreciate his companionship and suggestions very much. Anthony J. Gaudin prepared the maps. This paper orig- inated while I was a John Simon Guggenheim Memorial Fellow 1963-1964 and I was encouraged in my studies in Costa Rica by the Organization for Tropical Studies. LITERATURE CITED CARBALLO GUTIERREZ, ENIO 1960. Viaje a los vertices Dudu y Kamuk, en la Cordillera de Talamanca. Informe Semestral Instituto Geografico de Costa Rica, Enero-Junio:1960, pp. 87-89. Corr, Epwarp D. 1875. On the Batrachia and Reptilia of Costa Rica. Journal Academy of Natural Sciences, Philadelphia (letterpress), ser 2, no. 8, pp. 93-157. 1876. On the Batrachia and Reptilia of Costa Rica. Journal Academy of Natural Sciences, Philadelphia, ser 2, no. 8, pp. 93-157. 286 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Serr. 1878. ‘Tenth contribution to the herpetology of tropical America. Proceedings of the American Philosophical Society, vol. 17, no. 100, pp. 85—98. 1893. Second addition to the knowledge of the Batrachia and Reptilia of Costa Rica. Proceedings of the American Philosophical Society, vol. 31, pp. 333-345. Dat, Wi11AM H. 1909. Biographical memoir of William More Gabb. 1839-1878. Biographical Memoirs National Academy of Science, vol. 6, pp. 347-361. Gasp, WILLIAM M. 1875. Informe sobre la exploracion de Talamanca verificada durante los Anos de 1873-4. Anales del Instituto Fisico-Geografico Nacional de Costa Rica, vol. 5, pp. 67-92. 1877. Aufnahme von Talamanca und der Kartographische Standpunkt von Costa Rica in 1877. Petermann’s (Geographischer) Mittheilungen, vol. 23, pp. 385-387, pl. 18. 1913a. Letter addressed by William M. Gabb to his Excellency General Don Tomas Guardia, President of the Republic of Costa Rica. Costa Rica-Panama Arbi- tration Documents, vol. 4, no. 581, pp. 95-97. 1913b. Report of the Talamanca Exploration made during 1873 and 1874 by W. M. Gabb. Costa Rica-Panama Arbitration Documents, vol. 4, no. 582, pp. 97-142. GUTIERREZ BRAUN, FEDERICO 1960. Ascensidn de Gabb al Kamuk o Pico Blanco. Informe Semestral Instituto Geo- grafico de Costa Rica, Enero-Junio 1960, pp. 11-16. PETERS, W. 1857. Vier neue Amerikanische Schlagen aus der familie der Typhlopineri. Monats- bericht Ko6niglich Preussischen Akademie der Wissenschaften zu Berlin, 1857, pp. 402-403. 1863a. Mittheilung uber eine neue Arten der Saurier-Gattung Anolis. Monatsbericht Koniglich Preussischen Akademie der Wissenschaften zu Berlin, 1863, pp. 135- 149. 1863b. Mittheilungen iiber eine neue Schlangen gattung, Styporhynchus, und _ ver- schiedene andere Amphibien des Zoologischen Museums. Monatsbericht Koniglich Preussischen Akademie der Wissenschaften zu Berlin, 1863, pp. 399-413. 1863c. Mittheilungen tber neue Batrachier. Monatsbericht Koniglich Preussischen Akademie der Wissenschaften zu Berlin, 1863, pp. 445-471. PITTIeER, ENRIQUE 1913. Introduction to the spanish translation of the report of Mr. William M. Gabb, by Professor H. Pittier. Costa Rica-Panama Arbitration Documents, vol. 4, no. 580, pp. 92-94. REGEL, EDUARD 1867. J.v. Warszewicz. Gartenflora, Erlangen, vol. 16, pp. 95—96. ROUPPERT, KAZIMIEZ, ED. 1927. J. Warszewicza. Ogroduictwo, vol. 23, pp. 1-32. SAVAGE, JAY M. 1968. The dendrobatid frogs of Central America. Copeia, 1968, no. 4, pp. 745-776. 1969. Clarification of the status of the toad, Bufo veraguensis O. Schmidt, 1857. Copeia, 1969, no. 1, pp. 178-179. SAVAGE, JAY M., anp W. Ronatp HEYER 1969. The tree-frogs (Family Hylidae) of Costa Rica: diagnosis and distribution. Revista de Biologia Tropical, vol. 16, no. 1, pp. 1-127. Vor. XXXVIIT] SAVAGE: WARSZEWICZ AND GABB 287 SAVAGE, JAY M., AND ARNOLD G. KLUGE 1961. Rediscovery of the strange Costa Rica toad, Crepidius epioticus Cope. Revista de Biologia Tropical, vol. 9, no. 1, pp. 39-51. ScHMIDT, O. 1857. Diagnosen neuer Frosche des zoologischen Cabinets zu Krakau. Sitzungsberichten der Kaiserlichen Akademie der Wissenschaften in Wien, Mathematisch-Natur- wissenschaftlichen Klasse, vol. 24, no. 1, pp. 10-15. 1858. Deliciae herpetologicae Musei Zoologica Cracoviensis. Beschreibung der im K. K. Museum zu Krakau befindlichen, von J. v. Warszewicz in Neu-Granada und Bolivia gesammelten Ungeschwanzten Batrachier. Denkschriften der Mathe- matisch-Naturwissenschaftlichen Klasse der Kaiserlichen Akademie der Wissen- schaften in Wien, vol. 14, pp. 237-258. WaGNER, Moritz 1853. Die Provinz Chiriqui (West-Veragua) in Mittel-Amerika. Petermann’s (Geo- graphischer) Mittheilungen, vol. 13, pp. 16-24. x - es : _ 7 g a ee ‘ mai = hee Gee : == © Chee Teed ws spin eatin th i = -_ 7 _ ae PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Festschrift for George Sprague Myers Vol. XXXVIII, No. 15, pp. 289-298. December 31, 1970 TELEOST HYBRIDIZATION STUDIES By Clark Hubbs The University of Texas at Austin 78712 “Traditionally, studies such as ours have been based on morphology, espe- cially the skeleton, which is the only complete organ system available for detailed comparisons with fossils. However, with the variety of both primitive and ad- vanced teleosts living today, we are most emphatically of the opinion that ap- proaches other than morphological ones would be exceedingly fruitful in the in- vestigation of teleostean interrelationships.” The above quotation from Greenwood e¢ al. (1966) clearly states George S. Myers’ philosophy that systematic studies are central to biology. Any difference or similarity between two groups of organisms can be of value in estimating the amount of divergence; therefore, all biologic investigations can provide direct or indirect taxonomic information. Similar or identical organisms should be used in order to obtain repeatable experimental results; therefore, all biological in- vestigations can be considered to be based on systematic research. At the present time, the classification of most major taxa is based on their gross anatomy. Although this is due primarily to tradition, there is a valid scien- tific basis. Most experimental* analyses cover such small fractions of the tax- onomic subdivisions that experiment-based classifications would have major gaps. Moreover, a typological concept has no place in modern systematics. * Hereafter the word experimental should be considered to equal all types of analyses that are not tradi- tional studies of museum specimens. [289] 290 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. When we contrast several related taxa we should compare the spectra of attri- butes in the diverse members of each taxon. Because of the difficulty of obtain- ing data and the short history of such studies, most experimental investigations contrast supposed typical representative members of the taxa to be compared ( = typology). In contrast, experimental studies are not burdened with traditional taxonomic relationships. The refreshing new viewpoint can challenge the validity of an un- supported traditional taxonomic conclusion. The resulting interaction between morphological taxonomists and experimentalists can provide a realistic arrange- ment of organisms approximating their phylogenetic relationships. The primary contribution of the taxonomist to this interaction may be to point out taxonomic problem groups so that the experimental biologist can concentrate his efforts efficiently. Among the host of problems faced by the ichthyologist concerned with taxo- nomic problems has been the separation of environmental and genetic factors. Taning (1952) and many others have shown that a single environmental variable can concurrently alter several morphologic entities regardless of genotype. Therefore, a morphologic comparison that emphasizes those attributes might produce a dichotomy between those fishes reared in warm water and those reared in cold water, despite their genetic affinities. Environment-related problems can further plague taxonomy because survival in similar environments tends to select for similar morphologic attributes. For example, fishes that live in rock crevices tend to be elongate. Existence in this environment seems also to be enhanced by small eyes and scales. Some may be blind or naked. Many also have anteriorly located or reduced pelvic fins and small gill slits. Theoretically, distortions of true relationships by convergent or parallel evolution can be resolved by the use of diverse attributes. Practically, one must be careful not to use apparently divergent characters that happen to have selective value in similar habitats. Use of “non-adaptive” characters for taxonomy should help resolve problems of this type, but can one be certain that any character is not adaptive? Johnson and Wicks (1964) have advocated the use of molecular biologic ( = electrophoretic) studies because they may provide the “ultimate” infor- mation on relationships. Studies of DNA hybridization seem to offer even more promise of approximating the degree of phylogenetic divergence. Even this “ul- timate” systematic tool may have potential weakness. Assume that we have two ancestral species of omnivorous fishes occupying estuaries. Both evolve into a freshwater herbivore and a saltwater carnivore. The DNA sequences in each species pair would diverge so that each saltwater type would have a sequence of codings favorable for survival in high salinity. Similarily each would have their DNA controlling their digestive enzymes designed to break down animal ma- Vor. XXXVIIT] HUBBS: TELEOST HYBRIDIZATION 291 terial. The freshwater representatives of each pair would have their nucleotide sequences designed to produce enzymes different from their sibling species, but the same as those of their more distantly related ecological counterpart. The above simplified model is undoubtedly extreme, but may indicate how depen- dence on a single analysis could be hazardous. It is also possible that this type of convergence would be missed despite the type of analysis used. In effect, we have returned to the premise that taxonomic conclusions should be based on the sum (or product?) of the biological studies available. Never- theless, each investigator should not attempt to carry out investigations in all areas, but should concentrate on those for which his aptitude, experience, and interest suit him. During recent years, I have used hybrid survival as an index of phylogenetic relationship. This approach has three major merits: 1) It neces- sitates minimal expenditures, 2) It measures genetic divergence, and 3) The results approximate those of classical morphological taxonomy. The general agreement between hybrid survival experiments and classical fish taxonomy (Hubbs, 1967) and the potential hazards in such tests (Hubbs and Drewry, 1960) combine to make such tests valuable contributions to, but not the ultimate answer for, problems of phylogenetic relationships. The exper- iments reported below relate to two levels of relationships: 1) Arrangement of fishes within the family Cyprinodontidae and 2) Arrangement of various fish families. MATERIALS AND METHODS The techniques of Strawn and Hubbs (1956) were used for removal and mix- ing of the gametes. Two modifications were used that reduced some of the ex- perimental difficulties. Most of our experiments have used gametes from “wild” fish, that is, the individuals were removed from natural populations when nearly ripe and taken to the laboratory for the experiments. This necessitated hurried field work to avoid having eggs shed or becoming overripe ( = stale) during transport. We have found that the gametes can be stripped and mixed in the field and then taken to the laboratory as they develop. Large numbers of ex- periments can be done in this manner if the trip is properly planned. We have used petri dishes for transportation of individual experiments. The ripe eggs attach to the surface of the basal unit and the top is held in place with rubber bands. A piece of tape on the edge of the basa] unit permits water circulation, another on the bottom is used as a lable. The sets of petri dishes are placed in styrofoam containers and the water changed when necessary. One still must be careful not to remain away from the laboratory for too long because careful examination of development is difficult in the field and newly hatched larvae can escape from the petri dishes. Keeping the transportation equipment cool prolongs the time that one can remain in the field. 292 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. We find that fertilization can be enhanced by use of mashed testes. Typi- cally, semen is removed from males by coelomic pressure. Some species have such small quantities of semen that milt is seldom extruded by this technique, but eggs can be fertilized by extracting and mashing the testes. The same tech- nique works with males whose sperm supply has been depleted in previous ex- periments. When pertinent, the sources of the stocks will be presented with the listing of the experiments. CROSSES OF CYPRINODONT FISHES Moenkhaus (1910), Newman (1908), Hubbs and Drewry (1960 and 1962), Archer (1966), Drewry (1967), and others have reported on many successful crosses among species of Fundulus and with species of related genera such as Adinia, Lucania, Rivulus, Jordanella, Crenichthys, and Cyprinodon. Unfor- tunately, the last two listed papers are in thesis form and have limited circulation. In general, the level of success parallels the estimate of phylogenetic similarity as determined by morphological taxonomy. Most species of Fundulus can be crossed with the others and the hybrids reared to mature size. Two previously unlisted crosses, F. seminolis 2 (Sumpter Lake, Florida) x F. cingulatus 4 (Green Cove, Florida) and F. seminolis 2 (Sumpter Lake, Florida) x F. heter- oclitus 6 (Matanzas Inlet, Florida) can be added to the extensive list of reared hybrids. Many authors have reported that hybrids between F. majalis or its near relative (race?) F. similis and other species of Fundulus will hatch if F. majalis type sperm is used and not if F. majalis type eggs are used. Six tests with F. zebrinus (Iraan, Texas) sperm and F. similis (9 mile pond, Texas) eggs had over 100 fertilized eggs fail to hatch. Drewry (1967) reported difficulties in crossing F. notatus or F. olivaceus with other species of Fundulus, but hybrids between them are easily reared (Thomerson, 1967). Drewry reported that only 1 of the 17 hybrids using F. olivaceus sperm (none available with F. notatus) hatched and it died shortly. The reciprocal experiments had 9 fertilized eggs of which 6 hatched but were not reared, indicating a difference between reciprocals that more recent results sup- port. Archer (1966) also failed to rear hybrids between Fundulus notatus or F. olivaceus and 2 other Fundulus species. Fundulus notatus 2 (Blanco R., Texas) has been crossed with F. kansae 6 (Colbert, Oklahoma) and F. grandis ¢ (Port Aransas, Texas) and one fish from 8 and 5 fertilized eggs respectively reared, but both were deformed. Fundulus olivaceus 2 (Scraper Park, Oklahoma) X F. kansae é (Colbert, Oklahoma) had 13 of 18 eggs hatch but the deformed larvae shortly died. The difficulty of rearing the hybrids supports Drewry’s observa- tions, but they can be reared. The reciprocal hybrids are much more difficult Vor. XXXVIIT] HUBBS: TELEOST HYBRIDIZATION 293 to rear. Fundulus kansae 2 (Colbert, Oklahoma) X F. olivaceus 6 (Scraper Park, Oklahoma) (twice); F. kansae 2 (Colbert, Oklahoma) X F. notatus ¢ (Blanco R., Texas) (12 times); F. kansae 2 (Miller Cr., Texas) X F. nota- tus 6 (Little Piney Cr., Texas), F. zebrinus 2 (Iraan, Texas) X F. notatus 4 (Onion Cr., Texas) (5 times); F. cingulatus 2 (Dog Lake, Florida) X F. oli- vaceus 6 (Baker, Florida), and F. similis 2 (Port Aransas, Texas) X F. nota- tus 6 (Blanco River, Texas) all failed to develop late embryos. The “greatest level of success” achieved in these 22 tests was 1 egg (F. kansae Colbert X F. notatus Blanco) that produced chromatophores but did not show indication of gastrulation. The low success and high frequency of abnormalities supports Drewry’s hypothesis that F. notatus and F. olivaceus are phylogenetically similar to each other but dissimilar to other species of Fundulus. Six of 7 attempts to cross F. notatus and F. olivaceus were successful but only 4 of 6 control tests. The previous intrageneric hybridization tests have not used the West Coast species, F. parvipinnis (Mission Bay, California, population). Failure of crosses with F. kansae (both reciprocals), F. grandis sperm, and eggs of F. cingulatus (3 tests, 2 populations), F. heteroclitus (2 tests), F. majalis, F. similis and F. olivaceus, indicates that this species is separate from other species now placed in Fundulus. Of course, only 10 failures may not be enough tests to insure valid results. The better results of hybridizing F. parvipinnis with Crenichthys baileyi (Hubbs, 1967) may indicate a common ancestry. Only Archer (1966) has previously reported hybridization tests with Jor- danella floridae. He reported that one set of Jordanella eggs crossed with Cy- prinodon variegatus sperm died as late embryos. Three tests with Cyprinodon females from Iraan, Texas, resulted in no fertilization, but 1 of 2 and 3 of 4 tests with female Fundulus zebrinus and Lucania parva respectively from the same locality produced late embryos, but none hatched. This indicates that Jor- danella is distinct from those 3 species. Drewry (1967) reported difficulty in rearing hybrids between Lucania and Fundulus; however, Archer (1966) reared several F. pulvereus X L. parva individuals to adult size. Apparently the hybrids did not exhibit sexual dimorphism. INTRAFAMILIAL HYBRIDIZATION The relationship of hybridization success to phylogenetic divergence of tele- ost families may be shown to have significant importance in the taxonomic ar- rangement of living fishes. In part this approach resurrects those of Moenkhaus (1910) and Hertwig (1936). The ability of producing late hybrid embryos in a series of cyprinodontid X atherinid crosses has been considered support of their close relationship (Rosen, 1964). One more combination can be added to the series already reported (Hubbs, 1967, and citations). Two of 15 eggs from F. notatus 2 (Denison Dam, Texas) exposed to Menidia audens 6 (U. Oklahoma 294 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Biological Station) sperm gastrulated. Both produced heads, one was attached to the yolk mass and the other was free. Black pigments covered the yolk mass on both. The one with the head free also had extensive orange pigmentation and remained alive until almost all of the yolk was expended. These results re- semble those of the reported series of cyprinodontid-atherinid hybrids. The fail- ure at gastrulation of a parallel experiment with Notemigonus crysoleucas sperm shows that fundulines will not hybridize with all fishes. Menidia audens has not previously been reported to have been tested in rearing experiments. The series (4) of controls done the same day all had 50 percent plus fertilization and hatching. Similar to most atherinids, the controls died a week later apparently due to starvation. Intrafamilial hybrids between M. audens 2 and Labidesthes sicculus é (Tishomingo, Oklahoma) (twice) had 25 percent—SO percent fertilization and all hatched and died with the maternal controls, showing that Menidia hybrids can be reared as far as the controls. Menidia audens has also been tested for hybridization survival with members of several other families. The tests with Notropis cornutus, Notemigonus cryso- leucas (3 times), Gasterosteus aculeatus, and Aphredoderus sayanus all termi- nated before gastrulation. Gastrulation and embryonic formation occurred in tests with a centrarchid and several percids. One egg gastrulated among 2 sets of M. audens eggs exposed to Lepomis macrochirus sperm. It died before pigmenta- tion. Another set of M. audens eggs exposed to Etheostoma radiosum (Blue River, Oklahoma) sperm produced 5 early embryos, of which only 2 developed pigmentation, the test with Percina caprodes males from the same locality failed. Seven reciprocal experiments were set up with percid eggs from Blue River fe- males, one E. spectabile and six E. radiosum. Six had some development and 58 of 192 gastrulated embryos produced pigmentation, but none showed any sign of circulatory development. Clearly these hybrids were more successful than most other interfamilial tests and approached that of the atherinid-cyprinodont tests. A series of other intrafamilial tests were done with Etheostoma or Hadrop- terus eggs. It is not surprising that all 7 tests with the ostariophysines, Moxo- stoma poecilurum, Notropis cornutus, Notropis umbratilis, Notemigonus cry- soleucas, and Opsopoedeus emiliae failed to gastrulate or that 3 crosses with the ‘‘black race” of Gasterosteus aculeatus (Chehalis, Washington) did not gas- trulate. Five of 17 tests with A phredoderus sayanus sperm had 1 or more eggs gastrulate. The males were from a stock obtained at Douglass, Texas. The suc- cessful combinations were essentially the same ones as the failures, indicating that the data are representative of the interfamilial combination. Two sets of £. asprigene (Douglass, Texas) eggs each had 2 eggs gastrulate and develop eye pigmentation. Two of them formed a heart that beat irregularly, but had no vis- ible cells in the tubes. Many erythrocytes were present on the yolk mass anterior to the head. Another embryo had no heart beat but died breaking the egg shell. UL VoL. XXXVIIT] HUBBS: TELEOST HYBRIDIZATION 29 Two sets of H. scierus (San Marcos and Pedro Creek, Texas) had 2 and 3 eggs gastrulate. Only 3 developed recognizable heads. Three additional embryos from an E. spectabile (San Marcos, Texas) X A. sayanus cross that developed pigmented eyes were sacrificed in an unsuccessful attempt to analyze the cy- tology. The putative hybrids were diploid but chromosome markers were not noted. Four of 11 sets of Etheostoma eggs exposed to Elassoma zonatum sperm had gastrulation. All but one involved E. spectabile eggs. The 2 successful sets from Shoal Creek, Missouri, females had 6 gastrulated eggs, 5 had eye pigmentation, and 1 a functional heart beat and flow. All died at the time the maternal con- trols hatched. The 2 successful tests with Blue River, Oklahoma, females (£. radiosum and E. spectabile), each had 1 embryo that developed eye pigmen- tation. The failure of 2 sets of G. aculeatus eggs exposed to A. sayanus sperm indi- cates that A. sayanus sperm will not fertilize all teleost eggs. DISCUSSION Most of the intrafamilial hybrid experiments substantiate those previously reported, i.e., teleost hybrids are relatively easily produced and if the parental morphology is similar the hybrids are easily reared. Drewry had evidence of a genetic block to the development of hybrids between F. notatus or F. olivaceus and other members of that genus. Because he had few tests (26 fertilized eggs), it might be possible that his results were due to chance. We had 26 fertilized eggs in the least studied reciprocal (F. olivaceus or F. notatus egg) and 22 tests with the reciprocal. Not only did the results confirm those of Drewry, they also showed a distinct difference between the reciprocals. Because most of our “standard Fundulus’’ in these tests were of the F. kansae—F. zebrinus type, it may be that the difference between reciprocals may relate only to those combi- nations. It is possible, however, that we have a second difference in reciprocal hybrid survival in funduline fishes. It is amply evident, however, that F. notatus and F. olivaceus are quite distinct from other species of Fundulus. Hybridization tests have clearly showed that gametes from Adinia xenica and Lucania parva are more compatible with gametes from typical Fundulus than are those of Fun- dulus notatus or F. olivaceus. Therefore, the genus should be expanded to in- clude the members of Lucania and Adinia, or F. notatus and F. olivaceus should be separated from the other Fundulus species and placed in the genus Zygonectes. The failure of 10 tests between Fundulus parvipinnis and 7 other species of Fundulus indicates that this species is quite distinct. The reasonable success of hybrids between F. parvipinnis and the Crenichthys-Empetrichthys complex suggests a possible relationship that makes biogeographic sense. The hybrids of all west coast Fundulines have earlier developmental blocks when crossed with an 296 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER. east-coast representative than when crossed with another western type. Perhaps the Fundulines had an east-west primary separation. The western types then diverged into a coastal (F. parvipinnis precursor) and a desert spring (Cre- nichthys, Empetrichthys) type. Because F. parvipinnis occupied a “typical Fundulus” environment, it retained “typical Fundulus” morphology, while Cre- nichthys and Empetrichthys occupied clear warm spring waters and became su- perficially distinct. Assuming that the cyprinodonts have lost spinous fin rays, it seems that hy- brids between soft-rayed and spiny-rayed fishes die at or before gastrulation. The consistent failures indicate a uniform distant relationship and perhaps a treelike phylogeny rather than a bushy type suggested by Greenwood et al. A treelike phylogeny is also indicated by hybrid development to late embryonic stages in almost all crosses between two spiny rayed types. If we placed A phredo- derus and Etheostoma in separate superorders than the hybrid success is incon- gruous; in contrast, if we follow Regan and make A phredoderus a relic of a group ancestral to other perciforms, the phylogeny would agree with the hybridization data. The success of most interfamilial spiny rayed hybrids is quite similar. Only two types of combinations usually survive to hatching. The cyprinodont- atherinid hybrids and hybrids among percids, serranids, and centrarchids. Re- cent classifications place each unit in a suborder. In the future, hybrid survival tests may aid in determining other family group relationships. ACKNOWLEDGMENTS The experiments reported in this paper were supported by NSF GB 6429. Many graduate students at the University of Texas at Austin aided in the col- lection of parental stocks. The stocks of Fundulus parvipinnis were obtained through the courtesy of Dr. Carl L. Hubbs at the University of California at San Diego. This paper is dedicated to Dr. George S. Myers because of his inspired teach- ing of systematic ichthyology. LITERATURE CITED ARCHER, JAMES DOUGLAS 1966. The behavior and characteristics of some hybrids of cyprinodont fishes (Atherini- formes). Unpublished M.S. Thesis, Cornell University, pp. i-vii + 1-70. Drewry, GEORGE EARL 1967. Studies of relationships within the family Cyprinodontidae. Unpublished Ph.D. Dissertation, The University of Texas, pp. i-vii + 1-134, text figs. 1-4. GREENWOOD, P. HumpHREY, Donn E. Rosen, STANLEY H. WEITZMAN, AND GEORGE S. MYERS 1966. Phyletic studies of teleostean fishes, with a provisional classification of living forms. Bulletin of the American Museum of Natural History, vol. 131, pp. 339- 456, text figures 1-9, plates 21-22, charts 1-32. VoL. XXXVIIT) HUBBS: TELEOST HYBRIDIZATION 297 HERTWIG, PAULA 1936. Artbastarde bei Tieren. Handbuch Verebungswissenschaft, vol. 2, pp. 1-140. Hupss, CLARK 1967. Analysis of phylogenetic relationships using hybridization techniques. Bulletin of the National Institute of Sciences of India, No. 34, pp. 48-59. Husps, CLARK, AND GEORGE E. DREWRY 1960. Survival of F; hybrids between cyprinodont fishes, with a discussion of the cor- relation between hybridization and phylogenetic relationship. Bulletin of the Institute of Marine Science, The University of Texas, vol. 6, pp. 81-91. 1962. Artificial hybridization of Crenichthys baileyi with related cyprinodont fishes. Texas Journal of Science, vol. 14, pp. 107-110. Jounson, Murray L., anp MERRILL WICKS 1964. Serum-protein electrophoresis in mammals: significance in the higher taxonomic categories. Jn: Leone, Charles A., Taxonomic biochemistry and _ serology, Ronald Press, New York, pp. 681-700, 7 text figures. MoeEnKHAUS, J. 1910. Cross fertilization among fishes. Proceedings of the Indiana Academy of Science, for 1910, pp. 353-393. Newman, H. H. 1908. The process of heredity as exhibited by the development of Fundulus hybrids. Journal of Experimental Zoology, vol. 5, pp. 503-561. Rosen, Donn ERIc 1964. The relationships and taxonomic position of the halfbeaks, killifishes, silversides, and their relatives. Bulletin of the American Museum of Natural History, vol. 127, pp. 219-267. STRAWN, Kirk, AND CLARK HUBBS 1956. Observations on stripping small fishes for experimental purposes. Copeia, 1956, pp. 114-116. TAninG, A VEDEL 1952. Experimental study of meristic characters in fish. Biological Reviews, vol. 27, pp. 169-193, text figures 1-10. THOMERSON, JAMIE E. 1967. Hybrids between the cyprinodontid fishes, Fundulus notatus and Fundulus oli- vaceus, in southern Illinois. Transactions of the Illinois Academy of Science, vol. 60, pp. 375-379. 7 - 7 - - > \ : © 7 = —— A - oe =. > eet 2 oe i [ = a a 4 ~@ 6 ¢ > 4 # v _ ies é _ j 7 ‘ in | = ' ay : 4 i : a 7 Be ine Awe < is Z - \@ Ve (te @ o= SJ) | an os, sat Ok a yea ee ag fe iee i ‘S. OS SPP i ae ‘am. ab hi ga / an 7 ay eee @,9S). 2-38 - 7 = 7 b (ee et ee ee Bra ' "ity vive ae ape Fig —_ bs ge a ae) “ Sage PROCEEDINGS care OF THE C1 oe CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Festschrift for George Sprague Myers Vol. XXXVIII, No. 16, pp. 299-340; 5 figs.; 4 tables. December 31, 1970 A REVISION OF THE FISHES OF THE GENUS NOTOTHENIA FROM THE NEW ZEALAND REGION, INCLUDING MACQUARIE ISLAND’ By Hugh H. DeWitt Marine Science Institute, University of South Florida, St. Petersburg, Florida 33701 INTRODUCTION While preparing a revision of the southern and Antarctic fishes of the genus Notothenia, it became evident that the taxonomy of the species found in the New Zealand region is confused. In the most recent review (Parrott, 1958), five species are identified as occurring there. Of these, only four are valid, only three of the four are found in the New Zealand region, and the nomenclature of the three is entirely confused. This should not reflect upon Parrott, for he fol- lowed Boulenger, Waite, Regan, and Norman. For these reasons it is timely to present new descriptions and a new key for the New Zealand species together with a Clarification of the nomenclatural confusion which has surrounded them. I include Macquarie Island in this paper because two of the three species of Notothenia recorded from there also occur in New Zealand waters. Further, Notothenia coriiceps is included in the key to the species because it is widely dis- tributed in the Southern Ocean, is known from the Kerguelen Islands, and even- tually also may be found at Macquarie Island; a description of it is not given. 1 Contribution number 15 from the Marine Science Institute of the University of South Florida. [299] 300 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. I have not included Notothenia cornucola in this revision because I do not believe it occurs at New Zealand. This species was early recorded from New Zealand waters (Gunther, 1860; p. 262; Hutton, 1872; p. 26; 1873; p. 262) and continues to be included in lists of New Zealand fishes although no specimens identified as NV. cornucola have been found for nearly 100 years. The most re- cent reference is Parrott (1958), who admits that its occurrence is doubtful, although he includes it in his key to the New Zealand species of Notothenia. Considering that specimens of V. cornucola are encountered most commonly in littoral and shallow inner sublittoral areas (for example, among and under the rocks of the beaches near Punta Arenas, Chile, at low tide), it seems likely that if the species actually occurred in the New Zealand region it would be well known there. Norman (1937b; p. 86) reviewed the evidence and concluded that it “. .. is very slender.” The specimen recorded by Giinther was probably mis- labeled, and Hutton’s 1872 record is probably based upon that of Giinther. Hutton’s 1873 record from the Chatham Islands was probably based upon specimens of N. angustata. This species has recently been collected there (Moreland, 1957) and I have seen the specimens (see the section on material examined under V. angustata). Further, Hutton’s (1873) statement that the upper lateral line “. . . extends to the end of the second dorsal, . . .” agrees best with my observations of NV. angustata rather than with N. magellanica, the species to which Norman believed Hutton referred. For these reasons I have included Hutton’s 1873 reference to NV. cornucola in the synonymy of N. an- gustata. I have not included the listings of NV. cornucola Richardson found in the lists and catalogues of New Zealand fishes because, in that form, they refer to a species which I believe does not occur in New Zealand. Several check lists of New Zealand fishes have been prepared at various times, some of which I have not seen. The most important are those by Gill (1893), which reviews in detail the earlier works, and by Phillipps (1927b) which refers to earlier lists. For lists that I have seen, I have included the references to Notothenia species in the synonymies according to my present interpretations of the names used. For example, the name Notothenia microlepi- dota is listed under that species even though during that period the name was used in reports on collections for specimens properly called NV. angustata. MUSEUM ABBREVIATIONS In preparing my descriptions I have utilized specimens from the collections of museums whose names are abbreviated in the lists of material examined as follows. BMNH: British Museum (Natural History), London. CM: Canterbury Museum, Christchurch, New Zealand. DM: Dominion Museum, Wellington, New Zealand. MACN: Museo Argentino de Ciencias Naturales, Buenos Aires. VoL. XXXVIII] DEWITT: NOTOTHENIA FROM NEW ZEALAND 301 MLP: Museo de La Plata, La Plata, Argentina. NMV: Naturhistorisches Museum, Vienna. PM: Muséum National d’Histoire Naturelle, Paris. SAM: South Australian Museum, Adelaide. SU: Division of Systematic Biology, Stanford University, Stanford, Cali- fornia. USC-Eltanin: material collected by the University of Southern California Antarctic Biological Research Program from the USNS Eltanin. USNM: United States National Museum, Washington, D.C. ZIL: Zoological Institute, Leningrad. ZMB: Zoologisches Museum, Humboldt-Universitat, Berlin. MEASUREMENTS AND COUNTS All measurements were made in a straight line with calipers, and are pre- sented in the descriptions as thousandths of the standard length unless otherwise specified. All were made on the left side unless there was a deformity or loss which necessitated using the right side. Lateral line and pectoral fin counts were usually made on both sides. Those measurements which are not usually made, or which have been made differently in the past, are defined in the following alphabetical list. Anal to Pelvic Distance: from base of pelvic spine to origin of anal fin. Body, Depth of: measured at origin of anal fin. Body, Width of: measured at thickest part of body above origin of anal fin. Dorsal Interspace: distance between base of last spine of first dorsal fin and first ray of second dorsal fin. Dorsal to Anal Distance: distance between origins of second dorsal and anal fins. Dorsal to Caudal Distance: distance between last ray of second dorsal fin and midbase of caudal fin. Head, Depth of: measured at vertical through cheeks. Head, Length of: measured from tip of snout (upper jaw) to posteriormost edge of opercular flap. Head, Width of: distance between cheeks. Pectoral Fin, Length of: measured from base of uppermost ray to tip of posteriormost extending ray. Pectoral to Pectoral Distance: distance between upper ends of bases of pectoral fins. Pelvic Fin, Length of: measured from base of pelvic spine to tip of posterior- most extending ray. Post Orbital Distance (Postorbital Part of Head): measured from posterior margin of orbit to posteriormost edge of opercular flap. Standard Length: measured from tip of upper lip to midbase of caudal fin. 302 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER. Upper Jaw, Length of: measured from tip of upper lip to posterior end of maxillary. The counts for the caudal fin include all the branched rays plus | additional ray above and below, i. e., the branched rays plus 2. The last ray elements in the second dorsal and anal fins are counted separately. The scales in a lateral longitudinal series are counted from the upper end of the base of the pectoral fin to the base of the caudal fin. Gill raker counts are given as follows: 6-9 + 0-1 + 12-17 = 18-26. This means that there are a total of 18-26 gill rakers, of which 6-9 are on the upper limb, none or 1 at the angle, and 12-17 on the lower limb. On each arch, except occasionally the fourth arch, there are 2 rows of gill rakers, one projecting anteriorly and the other posteriorly. These are called, respectively, the anterior and posterior series. The lateral lines and their counts as well as the terminology of the cephalic canals have already been described (DeWitt, 1962). GENUS NOTOTHENIA RICHARDSON A formal diagnosis of the genus will be presented elsewhere. The following characters serve to distinguish it from other genera of New Zealand marine fishes. The nostrils are tubular and single on each side; the New Zealand species have the hind margin of the tube extended into a flap. The gill mem- branes are joined to each other and to the isthmus, forming a free fold across the isthmus. The vomer and palatines are edentulous. Two dorsal fins are present, the first composed of 3-8 spines which are usually soft and flexible, the second long and composed of soft rays. The anal fin is similar to the soft dorsal fin. The pectoral fins have broad, almost vertical, slightly curved bases. The body is scaled; the head is nearly naked in the New Zealand species. The scales may be ctenoid or nonctenoid, with both types usually present. Two lateral lines are present on the body, one high near the bases of the dorsal fins, the other on the midside in the region of the caudal peduncle. In the New Zealand species the head is somewhat depressed, and the interorbital space and the top of the head are broad and flat. KEY TO THE SPECIES la, Lateral scales 78-99; middle lateral line 24-37; upper lateral line 61-75; 15-19 gill rakers on lower limb of first gill arch; total number of gill rakers on first arch DAES) eee ee igen So eee Se N. microlepidota, p. 325. 1b. Lateral scales 73 or less; middle lateral line 23 or less; upper lateral line 30-61; 8-15 gill rakers on lower limb of first gill arch; total number of gill rakers on first are. W5=23. oc nes te ee eee 2 Jan d(drom: sb). Rectoralsraysw2l—24 oe eee aes Ne POSSI Does 2s, Pectoral rays: 1O=U9) acne 3. 3a. (from 2b). Second dorsal fin with 35-41 rays; anal fin with 26-32 rays __ N. coriiceps. 3b. Second dorsal fin with 27-31 rays; anal fin with 22-26 rays __ 4. VoL. XXXVIII] DEWITT: NOTOTHENIA FROM NEW ZEALAND 303 4a. (from 3b). Upper lateral line with 36-48 tubular scales; total number of scales in upper and middle lateral lines 45-57; length of caudal peduncle 37.0-45.5 percent of head length; preoperculo-mandibular canal not connected with the temporal canal; dorsal surface of head without prominent ridges V. magellanica, p. 303. 4b. Upper lateral line with 45-61 tubular scales; total number of scales in upper and middle lateral lines 59-76; length of caudal peduncle 25.5-34.5 percent of head length; preoperculo-mandibular canal connected dorsally with the temporal canal; in larger specimens prominent ridges present on top of head extending from above each eye posteriorly onto temporal region ttt‘; SOXWN~«Y. an gsstata, p. 318. Notothenia magellanica (Forster). Gadus magellanicus Forster, in Bloch and Schneider, 1801: 10-11 (original description; type locality seas about Tierra del Fuego; no types preserved, description based upon notes taken from fresh specimens and an unpublished rough drawing) ; FORSTER, 1844: 361-362 (description); RICHARDSON, 1846: 61 (listed in footnote, see under Lota magellanica, below). Notothenia magellanica RicHARDSON, 1844: 9 (counts with reference to illustration: “Icon. ined. Bibl. Banks. fig. 178,” catalogued in British Museum (Natural History) in Banksian MSS. no. 6 & 7) ; Grr, 1862: 520 (listed). Lota magellanica RIcHARDSON, 1846: 61 (possibly a mistaken generic assignation”); GILL, 1862: 520 (listed). Notothenia magellanicus GUNTHER, 1860: 260 (listed). Notothenia magellanicuz DELFIN, 1899a: 21 (listed). Notothenia macrocephalus GUNTHER, 1860: 263 (original description; type locality Falkland Islands; type in British Museum); Grr, 1862: 520 (listed) ; CUNNINGHAM, 1871: 470 (color notes); PERUGIA, 1891: 618-619 (description) ; Smtr, 1897; 9-12, pl. 3, figs. 23-26 (description, scales) ; BOULENGER, 1900: 53 (listed). Notothenia maoriensis Haast, 1873: 276, pl. 16 (original description; type locality near Lyttleton Harbour, New Zealand; present location of type unknown, probably lost) ; Hutton, 1876: 212-213 (description); Hutton, 1890: 279 (listed); Grrr, 1893: 118 (listed) ; WartE, 1907: 29 (listed); Frost, 1928: legend for pl. 17, fig. 15 (otolith). Notothenia antarctica PreTERS, 1876: 837 (original description; type locality Accessible Bay, Kerguelen Island; type in Zoologisches Museum, Humboldt-Universitat, Berlin). Notothenia antarcticus STUDER, 1879: 131 (listed; color notes). Notothenia hassleriana STEINDACHNER, 1876: 69-70, pl. 6, left-hand figures (original descrip- tion; type localities Puerto Bueno and Port-Gallant, both in Strait of Magellan; types in Naturhistorisches Museum, Vienna); STEINDACHNER, 1898: 303 (listed). Notothenia arguta Hutton, 1879: 339 (original description; type locality Campbell Island; type in British Museum) ; Hutton, 1890: 280 (listed) ; Grrr, 1893: 118 (listed) ; WAITE, 1907: 30 (listed). Notothenia macrocephala GUNTHER, 1881: 20 (listed); VAMLANT, 1888: 27, pl. 3, figs. 2a-d (listed, illustrations) ; BOULENGER, 1902: 186 (listed) ; STEINDACHNER, 1903: 207 (listed) ; Dot1o, 1904: 86 (listed, distribution) ; LONNBERG, 1907: 10 (listed, color notes) ; REGAN, 1913: 277 (description, distribution) ; Hussaxor, 1914: 89 (listed with counts) ; WaAITE, 2In his description of Lota breviuscula, Richardson compares L. breviuscula with several other species, among which is “Lota magellanica of Forster.’’ In a footnote he lists the species and gives some data for each. Here Forster’s species is listed as Gadus magellanicus, with the following counts: B. 6; D. 5-31; A. 25; C. 14; P. 17; V. 6. These counts are identical with those given in Forster (in Bloch and Schneider, 1801: 11; 1844: 362) and Richardson (1844: 9) under Gadus magellanicus, except that Richardson does not give an anal fin count. It seems obvious that both Lota magellanica and Gadus magellanicus refer to the same fish, but the reason for the use of Lofa is unclear to me. 304 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. 1916: 66-69, pl. 3, fig. 2 (description, illustration) ; THompson, 1916: 431-433 (descrip- tion) ; REGAN, 1916: 378-379 (distribution) ; PHim1iipps, 1921: 123 (listed) ; THomPpson and ANDERTON, 1921: 94 (listed, synonymy); RENDAHL, 1925: 6 (listed); PHILLIPpPs, 1927a: 13 (listed) ; Puitiipps, 1927b: 44 (listed) ; Frost, 1928: 454-455, pl. 17, fig. 15 (otolith) ; NorMAN, 1937b: 88-90 (description, illustration, distribution) ; NoRMAN, 1938: 27 (distribution) ; OLIVER SCHNEIDER, 1943: 110 (listed, illustration) ; MacDonacH and Covas, 1944: 235-236 (description, distribution); Fowler, 1945: 128-129 (listed) ; Hart, 1946: 339 (pelagic young) ; Fowrer, 1951: 314 (key) ; ANDRIASHEV and TOKAREV, 1958: 199 (listed) ; ANDRIASHEV, 1959: 5 (vertebral count); BLANc, 1961: 124 (descrip- tion); Kenny and Haysom, 1962: 252 (habitat, food); SLAcK-SmiTH, 1962: 14 (color notes, habitat, food). MATERIAL EXAMINED. USNM 77329: Sandy Point (Punta Arenas), Strait of Magellan, 53 10’S., /0-554W. (1 183 °mm.)r USNM 88755: Municipal jetty (Port Stanley?), Falkland Islands (1; 193 mm.). USNM 88756: Mullet Creek, Falkland Islands, 51°44’S., 57°53’W. (2; 51.9 and 55.3 mm.). USNM_ 171000: Kainan Bay, Ross Sea, Antarctica, 78°14’S., 161°55’W. (22 90brana.) SU 59880: Macquarie Island (3; 48.0-169 mm.). SU 59882: Macquarie Island (2; 139 and 168 mm.). BMNH 1860.2.20.2: Falkland Islands (1, a skin; holotype of NV. macrocephala). BMNH 1886.11.18.28: Campbell Island, from Otago Museum, Dunedin, New Zealand (1; 150 mm.; type of NV. arguta). ZMB 21626: Deutsche Tiefsee-Expedition Station 123, 49°07’S., 08°40’E.; bottom depth 4418 m.; presumably taken at surface in a plankton net, 22 November, 1898 (1; 80.2 mm.). NMV 59926: Port Gallant (Puerto Gallant), 53°40’S., 71°58’W., field no. 1203a (1; 86.0 mm.; lectotype of NV. hassleriana). NMV 65389: Puerto Bueno, 50°59’S., 74°12’W.., field no. 1203b (1; 87.0 mm.; paralectotype of NV. hassleriana). MACN 1859: Punta Colnet (Cabo Colnett, 54°43’S., 64°20’W.), 17 fathoms (1; standard length not measured). MACN 2673a: Bahia Tethis (Tierra del Fuego), (1; 155 mm.). ZIL (no number): Transvaal Cove, Marion Island, about 2 meters (2; 189 and 216 mm.). ZIL (no number): Scotia Sea, 60°38’S., 44°08’W., bottom depth 287 m.; depth of capture 0-60 m.; gear Isaacs Kidd trawl; at Academician Knipowich Station 85 (1; 261 mm.). CM (no number): South Island, New Zealand, probably near Dunedin (1; 13/7 mim,)). I have also examined specimens deposited in New Zealand museums (all Vor. XXXVIII] DEWITT: NOTOTHENIA FROM NEW ZEALAND 305 uncatalogued) from the following localities. DM: Campbell Island, from Camp and Garden coves after tidal wave. CM: Campbell Island; Tucker Cove, Campbell Island; Penguin Harbour, Campbell Island; Perseverance Harbour, Campbell Island; Macquarie Island, 17 fathoms. DeEscripTION. Body evenly curved both dorsally and ventrally from head to base of caudal fin; compressed posteriorly, becoming broader and more rounded toward head; greatest depth of body at about origin of second dorsal fin; depth of body 208-282, its width 122-150; pectoral to pectoral distance 144-225; dorsal to anal distance 237-306. Caudal peduncle longer than deep, its length 107-135, its depth 93-102; dorsal to caudal distance 104-138. Head slightly shorter than average for genus, its length 280-320; its width, 146-248, about equal to its depth, 198-224. Vertebrae 16-18 + 28-30 = 45-47. Snout very bluntly rounded from dorsal view; from lateral view it rises steeply from tip of upper jaw to a point a little above and anterior to nostrils, where it becomes abruptly less steep; its length 82-102. Tubes of nostrils short, with posterior rim raised into a flap which may be folded over opening; placed 52-79 from tip of snout, 17-29 from orbit and 52—75 apart. Eyes placed high on sides of head, but below dorsal profile; diameter of orbit 58-96. Interorbital region very broad and flat, its least width 88-134; all of top of head, from pos- terior part of snout to occipital region, nearly straight and rising slightly pos- teriorly; length of postorbital part of head 141-176. Jaws short but wide, maxillary extending posteriorly to about vertical from pupil of eye; length of upper jaw 94-115. Teeth in each jaw in two almost uni- serial bands; those in outer bands much larger and more numerous than those of inner bands and extend full length of jaws; inner bands confined to anterior % or less of jaws. The numbers of teeth vary, for in some individuals the bands are almost entirely uniserial, whereas in others they may become essentially double for part of their length. Oral valves extend most of length of each jaw, the lower broadest; their exposed surfaces covered with coarse papillae, espe- cially close behind inner bands of teeth. Tongue fleshy and densely covered with short, slender papillae which may be covered by a mucous coating and appear as low rounded papillae. Anterior gill rakers of first gill arch nondentigerous, or occasionally with 1 to a few spines, the larger ones flattened, arranged 3-6 + 1 + 9-13 = 14-19. Posterior gill rakers of first arch dentigerous, arranged O—1 + 0-1 + 10-15 = 12—16. Gill rakers of remaining arches all dentigerous; 1—11 in posterior series of fourth arch. Branchiostegal rays 6; pseudobranchiae curved ventralward posteriorly. First dorsal fin 3—6, originating 306-343 from tip of snout, from just behind to just in advance of upper end of base of pectoral fin; lower than second dorsal fin, second or third spine longest, 67-99. Second dorsal fin 29-31, origi- 306 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. nating 396-437 from tip of snout, 25-65 behind base of last spine of first dorsal fin; length of sixth ray 125-171, of sixth from last ray 87-105. Membrane be- hind last spine of first dorsal fin may reach to base of first ray of second dorsal fin. Anal fin 22-26, originating 513-606 from tip of snout, below bases of rays 8-10 of second dorsal fin; length of sixth ray 103-134, of sixth from last ray 82-98. Caudal fin 14-16, its length 165-242; its posterior margin changes shape considerably with size, being deeply forked in very small individuals and becoming emarginate or even slightly rounded in larger specimens. Pectoral fins 16-18, their length 222-275, extending posteriorly to above bases of rays 1-8 of anal fin; width of their bases 81-88. In larger specimens (100 mm. or more) the upper rays are longest and cause the posterior margin to be obliquely truncate or slightly falciform; the lower posterior margin is rounded. Pelvic fins rather short, their length 166-216, third rays longest, not reaching posteriorly to base of anal fin; inserted 232-312 from origin of anal fin, not entirely to entirely in advance of bases of pectoral fins. Upper lateral line 36-46, separated from origin of second dorsal fin by 6-10 scale rows, ending below rays 3—6 from last of second dorsal fin; middle lateral line 5-14. The pores of the cephalic canals are small and often difficult to see, but are otherwise normal. Preoperculo-mandibular canals with 10—11 pores; infraorbital canals with 8—9 pores; supraorbital canals each with 4 pores and sharing a median coronal pore; temporal canals with 6 pores; supratemporal canal with 3—4 pores. Most scales on body nonctenoid, 47—64 in a lateral longitudinal series, 23-28 rows around caudal peduncle; ctenoid scales present in area of sides covered by appressed pectoral fins. These latter have a single row of weak teeth along the posterior margin which is vertical and straight and may be recessed into the scale. There may be also a few weak projections on other scales of the body. Scales extend onto base of caudal fin and exposed bases of pectoral fins. Medial bases of pectoral fins, including small portions of body posterior to bases, naked; a small scaleless area also present on exposed side just anterior to base of rays. Head nearly entirely naked; small patches of scales present behind eyes, on uppermost part of operculum, and at postero-lateral parts of top of head. Round fleshy papillae cover remainder of top of head, and are present around lower and posterior parts of eyes, on snout, opercles, and sometimes on skin covering posterior parts of maxillaries. The color patterns of preserved specimens seem to vary considerably. Most of the specimens examined show no striking patterns anywhere, being darker above (bluish-grey to warm brown) shading to paler ventrally. The vertical fins are dusky, with pigment on both rays and membrane in the dorsal and anal fins, but mainly on the rays in the caudal fin. The pectoral fins are more or less dusky, being darkest in the more recently caught specimens. However, the 2 Vor. XXXVIII] DEWITT: NOTOTHENIA FROM NEW ZEALAND 307 specimens from Marion Island in the collection of the Zoological Institute in Leningrad have a strikingly different coloration. Overall they are brown, darker above, lighter below, with small spots and mottlings, more or less distinct, on the upper parts of the body. There are very clear spots and vermiculations on the top and sides of the head, including, in the larger specimen, most of the snout and the upper medial part of the upper lip. Rather irregular spots and stripes are present on the dorsal and caudal fins, and there is faint spotting of the upper pectoral rays in the larger specimen. This spotted coloration is very similar to that found on most specimens of V. angustata. Norman (1937b) adds the following: ‘“. .. more or less distinct longitudinal stripes or series of spots on the sides; traces of oblique stripes below eye; . . . soft dorsal dusky, sometimes reticulated, and with a narrow pale margin. The young are more silvery, especially on the lower parts of head and body, and the fins are much paler.” Waite (1916) gives a good description of specimens from Macquarie Island: “The general color is olive grey, the lower parts yellow; the markings are black and somewhat irregular, but two oblique bands may be traced below the eye; a branch from the upper one crossing the lower part of the opercle; the rest of the upper parts and sides of the head bear irregular spots and lines; six or seven bands cross the back to below the lateral line, whence they break and form blotches alternating with the bands. The first dorsal is dark and clouded; the second has a dark intramarginal band and a white edge; diagonal bars cross the lower portion, and the clouding leaves lacunae in the membrane; the anal is sooty, but the tips of the rays are lighter; the other fins are also sooty but without markings.” In life the colors appear to be striking, as several authors have noted them. The back may be dark brown, dark grey-green, blue-grey, or rich golden-brown, passing to golden-yellow, cream, or reddish on the belly (the 189 mm. specimen in the Zoological Institute, Leningrad, was orange ventrally in life). The branchiostegal membranes may be bright orange-red or orange-yellow. The underparts of the head may be white, or the throat and jaws may be bright orange-red. The dorsal fins are blue-grey, the other fins grey (Cunningham, 1871; Lonnberg, 1907; Norman, 1937b; Studer, 1879). The existence of pelagic juveniles in this species, which have been collected some distance from land over great depths, explains satisfactorily the wide dis- tribution of the species and the apparent lack of differentiation between the many seemingly isolated populations. In their general coloration they resemble closely the pelagic young of NV. coriiceps and N. rossit, species which also have wide distributions. ANTARCTIC SPECIMENS. Among the Notothenia material which I have ex- amined are 2 large specimens captured well within the Antarctic Zone (Norman, 1938; Andriashev, 1965) which appear to belong with NV. magellanica. One, 308 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. USNM 171000, is from the Ross Sea and the other, in the Zoological Institute in Leningrad, is from the Scotia Sea. Both were collected from near the surface over fairly deep water. These 2 specimens thus present a problem with respect to both habitat and distribution. Information from the literature indicates that, except for the pelagic juveniles, V. magellanica is a near shore bottom fish, living among kelp, and that it can be captured with traps, hand lines and seines. Its distribution is primarily Subantarctic, extending into the edge of the Antarctic Zone only at Kerguelen and Macquarie islands. Further, very few species are known to inhabit both the Subantarctic and Antarctic zones. For these reasons the pelagic habit and high Antarctic localities of these 2 specimens suggest that they represent a different species. However, for nearly every character ex- amined they show no differences from Subantarctic material of V. magellanica, and it may be that the observed differences are products of their large size. Also, since V. magellanica is known to penetrate into the edge of the Antarctic Zone, it may prove to be one more species which inhabits both the Subantarctic and high Antarctic for at least part of the year. For the present, then, I shall consider these specimens as possible representatives of a differing population of NV. magellanica of unknown taxonomic rank. Table 1 presents the pertinent measurements and counts taken from the Antarctic specimens together with the ranges of the measurements expressed as thousandths of the Standard Length. Comparison of these data with those taken from the subantarctic material shows that the Antarctic specimens have smaller eyes, a shorter distance between the tip of the snout and the nostrils, a greater distance between the nostrils and the edge of the orbit, a wider interorbital space, a shorter upper jaw, a deeper body, a greater distance between the origins of the second dorsal and anal fins, shorter pectoral and pelvic fins, and more rows of scales about the caudal peduncle. Besides the above differences, the lowermost gill rakers in the anterior series of the first gill arch are dentigerous and appear similar to those of the posterior series; the caudal fin is distinctly emarginate and each lobe is pointed. Most striking, however, are the presence of ctenoid scales over most of the body. Those covered by, and just above and below, the appressed pectoral fins are strongly ctenoid, while those posteriorly on the sides of the body, anteriorly along the back, especially anterior to the first dorsal fin, and anterior to bases of pectoral fins are more weakly ctenoid. All of the scales on the belly, even anterior to the pelvic fins, are ctenoid. There are no obvious markings on the body or head. Top and sides of head and upper parts of body a dark grey-brown or bluish black; body shading to paler below, head becoming paler more abruptly along ventral edges of cheeks and opercles, and on lower jaw; the Scotia Sea specimen is very pale orange- pinkish below. First dorsal fin uniformly black; membranes of second dorsal Vor. XXXVIII] DEWITT: NOTOTHENIA FROM NEW ZEALAND 309 TaBLe 1. Measurements (in mm.) and counts from two Antarctic specimens of Notothenia magellanica, with ranges of measurements expressed as thousandths of the Standard Length. USNM ZIL Observation 171000 Specimen Range Measurements Standard Length (SL) 229 261 Length of head (HL) 64.3 68.8 253-281 Width of head (HW) 51.6 -- 215 Orbital diameter (O) 2: 10.1 39-53 Length of snout (Sn) 19.8 22.1 85-86 Snout to nostril distance (Sn-N) 11.1 10.5 40-48 Nostril to nostril distance (N-N) 16.3 18.4 71 Interorbital width (IO) S255 35.6 136-142 Length of postorbital part of head (PO) S/o = 162 Length of upper jaw (JL) Dee 24.0 92-93 Length of caudal peduncle (CPL) ell 34.9 121-134 Depth of caudal peduncle (CPD) DR Al 21.9 84-96 Dorsal to caudal distance (D-C) 28.6 — 125 Depth of body (BD) 72.6 70.1 269-317 Pectoral to pectoral distance (P-P) 50.2 = 219 Second dorsal to anal distance (D2.-A) 86.7 — 379 Snout to first dorsal (Sn-D:) 70.5 70.8 271-308 Snout to second dorsal (Sn-D») 91.7 99.9 383-400 Snout to anal (Sn-A) 126 141 540-550 Anal to pelvic distance (A-V) ales) 83.5 312-320 Length of caudal fin (CL) 46.7 48.5 186-204 Length of pectoral fin (PL) 49.9 57.0 218 Length of pelvic fin (VL) 35.8 39.8 152-156 Length of sixth ray of second dorsal fin 30.4 — 133 Length of sixth ray of anal fin 26.4 —- 115 Counts First dorsal fin (D1) 4 4 Second dorsal fin (Dz) 31 29 Anal fin (A) 25 25 Caudal fin (C) 16 16 Pectoral fin (P) 16 16 Lateral scales (LatSc) 52 53 Scale rows between lateral line and origin of second dorsal fin (LL-D:») 10 — Scales around caudal peduncle (ScArCP) 30 32 Upper lateral line (ULL) 37 44 & 45 Middle lateral line (MLL) 10 2 Branchiostegal rays (Br) 6 ~~ Preoperculo-mandibular pores 10 12 & 10 Infraorbital pores 8 9&8 Supraorbital pores 4 4 Temporal pores 6 6 Supratemporal pores 3 3 Anterior gill rakers, first arch 411+10=15 4+1+11=16 310 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. fin black, the rays pale hyaline, creating the effect of white stripes on a black field. Membranes of anal fin dusky basally, clear hyaline toward margin, the rays pale. Caudal fin dusky, especially along upper and lower edges of rays. Pectoral fins dusky along upper and posterior margins, paler centrally and below; pelvics dusky. DIstRIBUTION. Notothenia magellanica has been recorded from the Magel- lanic region; Kerguelan, Macquarie, Aukland, and Campbell islands; and from the South Island of New Zealand. In addition it is recorded here for the first time from Marion Island and 2 localities well within the Antarctic Zone. Except for pelagic juveniles and the 2 far southern records, the species appears to in- habit only very shallow water, as all records (where the information is given) state that specimens were secured by traps, hand lines, or seines. The Discovery obtained a few juveniles with dip nets and tow nets from or near the surface in open waters. Kenny and Haysom (1962) state that the species lives among kelp near the shore at Macquarie Island, and Forster (in Bloch and Schneider, 1801; 1844) states that about Tierra del Fuego it lives near the shore among sea weed. In the Magellanic region NV. magellanica appears to be restricted to the west coasts of Tierra del Fuego and Patagonia, and the Falkland Islands, a pattern similar to that of several other Subantarctic species. It is probable that adults everywhere are associated with rocky and protected areas near shore. Discussion. Although Norman (1937b) listed Gadus magellanicus Forster (in Bloch and Schneider, 1801) with a sign of interrogation under Notothenia macrocephala Giinther, he considered Forster’s description to be equally ap- plicable to N. macrocephala and N. microlepidota (non N. microlepidota of Hutton, but equals V. angustata of Hutton; see discussions under both species). His reasons for this position were that the unpublished drawing of the species by Forster is a rough sketch which, while definitely representing a Notothenia, is not of sufficient detail to identify the species, and that the anal fin is described as having 25 rays, a number common to both species. Through the courtesy of Mr. A. C. Wheeler of the British Museum (Natural History) and the Trustees of the British Museum I have been able to obtain a photograph of Forster’s drawing which is reproduced here (fig. 1). Although the drawing is obviously unfinished, it shows definitely that N. macrocephala is a synonym of Gadus magellanicus. The pectoral fin is drawn with an oblique posterior margin and with the upper rays longest, the snout is separated from the top of the head by an abrupt rounding above the nostrils, and the caudal fin is emarginate. These characters are diagnostic for the present species. Norman (1937b) also lists Notothenia porteri Delfin as a synonym of J. magellanica, but a careful reading of the description demonstrates that the name is a synonym of NV. angustata. A full discussion is presented under that species. Vor. XXXVIII] DEWITT: NOTOTHENIA FROM NEW ZEALAND 311 f° te i JOE, a prea Cdty, Ficure 1. Notothenia magellanica. Reproduction of J. G. A. Forster’s unpublished drawing of Gadus magellanicus, by permission of the Trustees of the British Museum (Natural History). A final problem has been the location and designation of the types of VV. maoriensis, N. arguta, N. hassleriana, and N. antarctica. It would appear that the type of V. maoriensis has been lost. In 1965 Miss M. Biichler (now Mrs. M. Darby), then Assistant Zoologist in the Canterbury Museum at Christchurch, New Zealand, informed me that an old register, dating back to the early part of this century or even into the last century, contains the following entry: “Notothenia maoriensis Haast, Trans. N. Z. Inst. vol. 5, p. 276 N. coriiceps Hutton, Cat. Fishes N. Z.: 32 (nec Richardson) Stuffed (Type lost, originally stuffed) .” Miss Biichler made a thorough search through the fish collection and catalogues and the above entry was the only positive result. Therefore it seems fairly certain that the type is no longer in existence. A number of fishes, some of which are types, were presented to the British Museum in the 1880’s by the Otago Museum in Dunedin, New Zealand. Among them is a specimen labelled as the type of Notothenia arguta. Its total length is about 179 mm., which is close to the length of 7% inches (equals 184 mm.) given by Hutton in the original description. Dr. D. R. Simmon of the Otago Museum informed me (letter dated 8 April 1964) that “although N. arguta is entered as a name in the register . . . there is no record of a specimen being held by this museum.” I therefore conclude that the British Museum specimen is indeed the type. Notothenia hassleriana was described from an unknown number of specimens collected at 2 localities in the Strait of Magellan. I have examined 2 specimens 312 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 2. Notothenia magellanica, from Steindachner, 1876. labeled as types, one each from the 2 localities. It is possible that these 2 specimens are all that Steindachner had, for Dr. P. Kahsbauer of the Naturhis- torisches Museum in Vienna indicated in letters dated 7 October and 7 Novem- ber, 1964, that these were all he could find. In any event, the specimen from Port Gallant (register number 59926) is very similar to the illustration pub- lished by Steindachner and reproduced here (fig. 2), and I designate this specimen as the lectotype. Notothenia antarctica was described from a single specimen, 35 cm. long, collected by Dr. Studer during the voyage of the SMS Gazelle from Accessible Bay, Kerguelen Island. Dr. C. Karrer of the Zoologisches Museum of Humboldt University in Berlin has written that a specimen identified as VV. antarctica of the proper size, from the above locality and collected by the Gazelle is in the fish collection there. Although it is not labeled as the type, it is undoubtedly the specimen Peters used for his description. Professor Kurt Deckert of the same museum had earlier written that although the register of the fish collection listed the type of NV. antarctica, he had been unable to find it. Notothenia rossii Richardson. Notothenia rossii RICHARDSON, 1844: 9-10, pl. 5, figs. 1 & 2 (original description and illustra- tion; type locality unknown, but probably the Kerguelen Islands (Regan, 1916); type lost); GUnrHeER, 1860: 263 (description) ; Norman, 1937a: 61, 64 (description, separa- tion from N. coriiceps); NorMAN, 1938: 25 (description, illustration, distribution) ; Branc, 1951: 495 (listed, food); Branc, 1954: 191 (listed); Brianc, 1958: 137 (listed, illustration) ; Branc, 1961: 123-124 (description) ; BELLIs1o, 1966: 69, foto 40 (listed, illustration) . Notothenia rossi REGAN, 1913: 240, 276-277 (description) ; ANDRIASHEV AND TOKAREV, 1958: 199 (juvenile listed). Macronotothen rossii GL, 1862: 521 (listed). Notothenia marmorata FiscHer, 1885: 53-55 (original description; type locality South Georgia, probably at about 54°31’S., 36°05’W.; types (2 specimens remain of original Vor. XXXVIII] DEWITT: NOTOTHENIA FROM NEW ZEALAND 313 TABLE 2. Measurements (in mm.) and counts from the types of Notothenia macrocephala, N. arguta and N. hassleriana. Abbreviations are as in table 1, with the addition of body width (BW), anterior gill rakers of first gill arch (AntGR) and longest pelvic rays (LongVR). Where two measurements or counts are given, the second is taken from the right side. N. macrocephala: N. hassleriana: N. hassleriana: N. arguta: BMNH 1860.2.20.2 NMV 59926 NMV 65389 BMNH 1886.11.18.28 Observation (Type) (Lectotype) (Paralectotype) (Type) SL = 86.0 87.0 150 HL 84 24.5 25.4 45.4 O 16 Sof 6.1 9.5 Sn — cd Toll 13.4 Sn-N —_ 4.9 4.5 8.7 N-N = 4.9 Si 9.9 IO 36 8.6 9.0 17.6 PO — 1g, 13.9 23.9 JL = 9.0 8.6 16.4 CPL 32 10.5 10.3 16.7 CPD — 8.3 8.4 14.7 BD = 21.6 RDS 38.7 BW — 10.5 WOES Depa P-P — 14.8 17.6 30.9 Sn-D, — Dies 27.4 49.0 Sn-D> = 36.7 36.3 63.8 Sn-A — 45.8 45.8 ies A-V = 26.7 24.6 40.5 CL — 18.4 18.0 29.4 12NL, — ZileO) 20.1 38.1 VL — 16.8 16.8 29.8 AntGR — 5+14+11=17 5+1+10=16 6+1+11=—18 D; ~~ 4 5 4 D2 Sil 30 Sil 30 A 24 24 24 24 (e — 16 16 16 1% ly) 16 &17 16 & 16 17 LongVR -—— 3 3 3 LatSc 49 51 54 50 LL-D, — 7 6 7 ScArCP — Dil 24 24 ULL 38 41 & 42 40 & 38 38 & 36 MLL 11 9& 11 10& 11 9 (?) & 10 3) in Hamburgischen Zoologischen Staatsinstituts und Zoologischen Museums, Ham- burg). Notothenia macrocephala marmorata LONNBERG, 1905: 34-36, 53 (description, spawning) ; LONNBERG, 1906: 94-95 (description, spawning, food). Notothenia coriiceps var. macquariensis WAITE, 1916: 64-66, pl. 5, fig. 3 (original description and illustration; type locality Macquarie Island; lectotype in South Australian Museum, Adelaide) ; REGAN, 1916: 378 (differentiation from N. coriiceps) ; NORMAN, 1937a: 60-61 (synonymized). 314 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER. Ficure 3. Notothenia rossi, from Waite, 1916. Notothenia rossti marmorata NYBELIN, 1947: 22-26 (differentiation from JN. r. rossii, de- scription, distribution) ; NyBELin, 1951: 23-27 (description, differentiation from JN. r. rossi, spawning); OLSEN, 1954: 373-382 (description, growth, food, habits); Ruup, 1954: 849 (oxygen capacity of blood); OLsren, 1955: 88 (biology compared to Chan- nichthyids) ; LApIGEs, WAHLERT, AND Mone, 1958: 165 (designation of lectotype) ; ANDRIASHEV, 1959: 5 (vertebrae). Notothenia rossii rossii ANDRIASHEV, 1959: 5 (vertebrae). MATERIAL EXAMINED. SU 67031: washed up on beach, Macquarie Island (1; 461 mm.; partly eaten and eviscerated). SAM (uncatalogued): Macquarie Island (1; 342 mm.; lectotype of NV. coriiceps macquariensis ) . The following material has been examined for purposes of comparison with the above Macquarie Island specimens. BMNH_ 1937.7.12.563-4: Jetty (probably Government Jetty, Grytviken), South Georgia (2; 129 & 143 mm.). USNM 107158: Stromness Harbour, South Georgia (1; 208 mm.). USNM 179080: King George Island, South Shetland Islands (1; 274 mm.). USC-Eltanin Station 671: South-west of South Georgia Island, 54°41’S., 38°38’W.; 220-320 m.; 10-foot Blake trawl (1; 432 mm.). DESCRIPTION Body less deep than \. magellanica, not becoming much deeper than head; ventral profile curves more evenly than dorsal profile, which rises most steeply in the snout; body compressed posteriorly, but becomes somewhat depressed anteriorly. Length of head 319-323, its width 234-292, its depth 204; depth of body 213, its width 138, dorsal to anal distance 235; pectoral to pectoral distance 219-228; length of caudal peduncle 103-104, its depth 85—91; dorsal to caudal distance 105-109. Vertebrae 20 + 31 = 51. Vor. XXXVIII] DEWITT: NOTOTHENIA FROM NEW ZEALAND 315 Snout rises steeply in a smooth curve from lateral view, its length 86-95. Nostrils short tubes with the posterior margin raised into a point, placed 53-63 from tip of snout, 24-25 from orbits, and 63-67 apart. The mouth appears larger than in V. magellanica, but the upper jaw extends posteriorly only under anterior edge of pupil; length of upper jaw 130-138; lower jaw projects slightly beyond upper jaw. Eyes directed laterally, placed just below dorsal profile of head; diameter of orbit 51-58. Interorbital region broad and almost flat, both from lateral and frontal views, its width 102-105. Length of postorbital part of head 174-186. Teeth in both jaws in 2 bands; outer band a single row of somewhat enlarged, evenly spaced, canine-like teeth extending almost full length of jaw, becoming slightly smaller anteriorly and absent near symphysis; inner band lies immedi- ately behind outer row, broad anteriorly, becoming narrow posteriorly, extending posteriorly only '% to % length of jaw (upper jaw) or as far as outer row (lower jaw). Tongue free anteriorly, fleshy, but not soft. Oral membranes extend most of length of jaws, papillose only along anterior edges. Larger gill rakers in anterior series of first gill arch flattened, nondentigerous, and not very elongate, arranged 6 + 0-1 + 13-14= 20. Posterior gill rakers of anterior gill arch dentigerous distally on anterior face, arranged 1 + 1+ 11 = 13 (SAM specimen); gill rakers of remaining arches similar. Branchiostegal rays 6; pseudobranchiae curved ventralward posteriorly. First dorsal fin 5—6, lower than second dorsal fin, length of longest spine 39-60; its origin 301-332 from tip of snout, from slightly behind to slightly in advance of upper end of base of pectoral fin. Second dorsal fin 32-33 (Waite, 1916, gives 33-34), its origin 406-451 from tip of snout, 31-51 from base of last ray of first dorsal fin; first ray short, heavy and unbranched but segmented; length of sixth ray 101-102, of sixth from last ray 70-86. Anal fin 27—28, its origin 558-562 from tip of snout, beneath bases of rays 8-10 of second dorsal fin; length of sixth ray 81—92, of sixth from last ray 69-77. Caudal fin 14-16, its length 154-173, its posterior margin truncate. Pectoral fins 22—23, their length 215—223, reaching posteriorly to above base of first ray of anal fin or not reaching to anal fin; middle portion of posterior edge truncate, upper and lower portions rounded; uppermost ray very short, about 48. Pelvic fins inserted 237-330 from origin of anal fin, entirely in ad- vance of bases of pectoral fins; their length 157-160, third, or third and fourth rays longest, not at all reaching to anal fin. Upper lateral line of body with 40-57 tubular scales, dipping slightly above upper end of base of pectoral fin, ending posteriorly below about fifth to ninth from last rays of second dorsal fin, and separated by about 6-7 scale rows from origin of second dorsal fin. Middle lateral line with 15-17 tubular scales, originating below or a little behind end of upper lateral line and extending a 316 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. short distance onto base of caudal fin. Cephalic lateral line system normal in pattern, but pores very small and difficult to see. Preoperculo-mandibular canals with 10 pores, not connected to temporal canals; infraorbital canals with 8—9 pores; supraorbital canals with 4 pores and sharing a coronal pore; temporal canals with 6 pores; supratemporal canal with 3 pores. Scales in lateral longitudinal series 55-57; 28-29 around caudal peduncle. Scales nonctenoid except for those in area of side of body covered by appressed pectoral fin and a little posteriorly which are weakly ctenoid; scales present everywhere on body except medial (posterior) base of pectoral fin and area im- mediately adjacent, and an arc along lateral base of pectoral fin; scales extend onto proximal part of caudal fin and onto lateral proximal part of pectoral fin; scales small on belly, ventral area anterior to pelvic fins and on back anterior to first dorsal fin. Scales absent on head except for about upper ” of cheeks behind eyes, about upper 4 of operculum, and 2 small patches on each side, one in front of the other, on posterolateral corners of top of head. Head only very slightly rugose, with small raised vermiculations, the most prominent radiating from eyes. Low ridges present, probably associated with parietal and pterotic bones. Color (in alcohol) of the SAM specimen is dark grey-brown with some blue above, becoming lighter, somewhat yellowish below. Second dorsal fin with dark longitudinal bands, 3 anteriorly, 2 posteriorly, rather irregular anteriorly. The SU specimen is brownish black above, lighter on belly. The second dorsal is marked with somewhat irregular brownish bands which extend posteroventrally in anterior part of fin and more or less parallel with back in posterior part of fin. Anal fin dusky except for a pale margin; caudal fin irregularly and indistinctly mottled. Two faint stripes on head, one extending along edge of upper jaw, the other extending from posteroventral edge of eye to angle of preopercular. Supspecies. Notothenia marmorata Fischer, described from South Georgia, has long been considered a synonym of J. rossi since comparison of material from Kerguelen and Macquarie islands with that from the region of the Scotia Sea has shown that the two populations are very similar. Nybelin (1947, 1951) was the first to call attention to differences between specimens from the two regions, and he resurrected the name “‘marmorata” as a subspecies of NV. rossit. Unfortunately, very little material has been reported from the Kerguelen- Macquarie region (Richardson, 1844; Waite, 1916; Blanc, 1951, 1954, 1961). The most reliable published information is that by Richardson; Waite’s 1916 paper contains numerous errors, and his methods of counting differ in some instances from those now in practice; the data given in the papers by Blanc seem to have been copied from reports on Antarctic material and cannot be used. For these reasons the data presented above, although obtained from only 2 specimens, are important additions to our knowledge of the species. Combining my observations with those of Richardson, it seems possible that Vor. XXXVIII] DEWITT: NOTOTHENIA FROM NEW ZEALAND 317 TaBLe 3. Measurements (in mm.) and counts from two specimens of Notothenia rossi from Macquarie Island. Abbreviations are as in table 2. Where two measurements or counts are given, the second is taken from the right side. Observation Lectotype SU 67031 Observation Lectotype SU 67031 SIL 342 461 A-V 98.1 152 HL 109 149 CL Soi 79.8 O 17.5 26.8 PL Bs 103 Sn S35) 39.7 VL 54.9 1253 Sn-N 21.6 24.4 AntGR 6+1+13 6+0+ 14 N-N 22.8 29.1 ——20 ==—/() IO 36.0 47.1 dD; 6 5 PO 59.5 85.7 D, 33 32 pe 44.4 63.4 A 28 27 CPI 35.1 48.1 Cc 16 14 CPD 31.0 39.3 Je 22 & 22 23 BD 72.3 — LongVR 3 3and3 &4 BW 47.1 -- LatSc 57 55 P-P 74.9 105 LL-D:; 7 6 Sn-D; 103 153 ScArCP 28 29 Sn-D> 139 208 ULL 57 & 54 40 Sn-A 191 259 MLL 17 &17 15 the Kerguelen and Macquarie material differ from the Scotia Sea material in having a longer snout (86-95 vs. 75-86), a broader interorbit (102-105 vs. 87-100), a longer upper jaw (130-138 vs. 106-123), a greater distance between the tip of the snout and the origin of the anal fin (558-562 vs. 472-547), fewer rays in the second dorsal fin (32-33 vs. 34-35), fewer vertebrae (51 vs. 52-53), and different coloration. It may be that the proportional differences are due to size, as the 2 Macquarie Island specimens examined are larger than nearly all of those seen from the Scotian region, but the counts and color differences seem to be reliable. In color, specimens from the Scotian region have the sides of the body covered by a series of irregular lines and blotches, with sometimes a dark arc at the base of each pectoral fin and spots on top of the head. The second dorsal is marked in much the same manner as in the Macquarie specimens, but the bands are much more distinct. For the above reasons I believe there is good evidence for following Nybelin in recognizing as subspecies two populations, one, V. rossi rossi, inhabiting the Kerguelen and Macquarie islands, and the other, V. rossii marmorata, inhabiting the islands of the Scotia Ridge system, including the South Shetland Islands. Discussion. In his original description of NV. coriiceps var. macquarienstis, Waite stated that the type was in the South Australian Museum, but he did not specifically designate either of the 2 specimens upon which his description was based. I therefore select the specimen from the South Australian Museum listed 318 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. under material examined as the lectotype. The second specimen, now presumably in the Australian Museum, Sydney, becomes a paralectotype. I do not know whether the lectotype is the specimen illustrated by Waite, but selecting the specimen in the South Australian Museum accords with Recommendation 74D of the International Code which suggests that a lectotype be selected from the material in the institution containing the largest number of types from the collection worked upon by the original author. Notothenia angustata Hutton. Notothenia coriiceps (non Richardson) Hutton, 1872: 26 (brief description) ; THomPson and ANDERTON, 1921: 94 (listed). Notothenia cornucola (non Richardson) Hurron, 1873: 262-263 (brief description). Notothenia angustata Hutton, 1875: 315-316 (original description; type locality Dunedin Harbour; type in Otago Museum); Hutton, 1876: 213 (an almost verbatim reprint of the previous paper; localities given as Dunedin and Bluff harbours) ; Hutton, 1879: 339 (listed, synonymy) ; Hurron, 1890: 279 (listed) ; Griz, 1893: 118 (listed) ; WatTE, 1907: 30 (listed). Notothenia parva Hutton, 1879: 339 (original description; type locality Auckland Islands; types in Dominion, Otago and British Museums); Hurron, 1890: 280 (listed) ; Git, 1893: 118 (listed) ; Waiter, 1907: 30 (listed). Notothenia porteri DeL¥IN, 1899b: 118-120 (original description; type locality Talcahuano, Chile; type (or types) possibly in the old natural history museum in Valparaiso, Chile). Notothenia microlepidota (non Hutton) BouLENGER, 1902: 185 (listed) ; WAITE, 1909: 590- 594 (description, illustration); Rrcan, 1913: 277-278 (description) ; WAITE, 1916: 69 (listed) ; ReGan, 1916: 379 (synonymy); THOMPSON AND ANDERTON, 1921: 94 (listed) ; MacDonacu, 1936: 428-429 (synonymy); Norman, 1937b: 90-91 (description, syn- onymy, distribution) ; FowLer, 1951: 314 (key) ; Moreranp, 1957: 34 (listed) ; Parrott, 1958: 110-111 (description, variation). Notothenia latifrons THompson, 1916: 434-435, pl. 3, fig. 1 (original description and illustra- tion; type locality Sandy Point (Punta Arenas), Strait of Magellan; holotype in U.S. National Museum). Notothenia macrocephala (non Ginther) FowLer, 1926: 283 (description). Notothenia patagonica MacDonacH, 1931: 100 (original description; type locality among the rock ledges of Bahia del Fondo, Golfo San Jorge, Santa Cruz (province), Patagonia (Argentina) ; holotype in Museo de La Plata); MacDonacu, 1934: 84-91, pl. 10, figs. 2 & 3, pl. 11, figs. 1 & 2, pl. 12 (description, illustrations, scales, systematics). MATERIAL EXAMINED. BMNH_ 1886.11.18.29: Auckland Islands; from the Otago Museum (1; 71.7 mm.; lectotype of NV. parva). BMNH 1886.11.18.30: Dunedin, New Zealand (1; 238 mm.). BMNH 1936.7.7.4: among the rocky ledges of Bahia del Fondo, Golfo San Jorge, Santa Cruz (province), Patagonia (Argentina) (1; 230 mm.; paratype of NV. patagonica). USNM 39670: New Zealand (1; 182 mm.). USNM 176391: Huiches Island, Chile (45°10’30”S., 73°33’W.) (1; 332 mm.). MLP 12—XII-30-1: same data as for BMNH 1936.7.7.4 (1; 315 mm.; Holo- type of NV. patagonica). Vot. XXXVIII] DEWITT: NOTOTHENIA FROM NEW ZEALAND 319 reese ss ee Ip ty 2? es ne ‘ Ficure 4. Notothenia angustata, from Waite, 1909. CM (uncatalogued): Ranui Cove, Auckland Island (2; 56.2 and 85.8 mm.). Selected measurements and counts were taken from the following specimens, all deposited in the Dominion Museum, New Zealand. 2864: Campbell Island (1); 2897: Oreti Beach, Southland (1); 3124: Campbell Island (1); 3332: outer Ranui Cove, Auckland Island (7); uncatalogued: Waitangi, Chatham Island, 43°36.2’S., 176°48.5°W. (3); uncatalogued: Glory Bay, Pit Island, Chatham Islands, shore, 43°47’S., 179°30’W. (1). Material in the Canterbury Museum (all uncatalogued) from the following localities was also examined. Tucker Point, Port Ross, Auckland Island, under stones (2); west coast of Campbell Island (1); Tucker Cove, Campbell Island, among kelp at low tide (1); Auckland Islands (2); Laurie Harbour, Auckland Island (1); Ranui Cove, Auckland Island (1). Description. Larger specimens more than usually compressed posteriorly ; caudal peduncle distinctly deeper than long. Smaller specimens more cylindrical; caudal peduncle may be longer than deep. In region of bases of pectoral and pelvic fins, body becomes broader and less deep; the head appears depressed and small, although its measured length is similar to those for other species. Dorsal and ventral profiles about equally convex, or the ventral profile, at least of head, may be a little more convex than dorsal profile. Length of head 302-345, its width 187-281, its depth 181-187; depth of body 197-259, its width 140-188; dorsal to anal distance 229-288, pectoral to pectoral distance 189-266; length of caudal peduncle 92-117, its depth 92-125; dorsal to caudal distance 89-125. Vertebrae 17-18 + 27-29 = 4446. Snout broad and flattened, its length 84-98, longer than diameter of orbit. In lateral view the snout appears short, but its breadth causes its measured length to be larger. A pair of ridges, through which the supraorbital canals extend, separate the medial and lateral parts of the snout. These ridges curve around 320 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. the posterior and medial sides of the nostrils and extend anteriorly to end at the edge of the groove behind the upper lip. In larger specimens the medial portion of the snout is flat and somewhat raised; in smaller specimens the ridges are little developed and the snout is more evenly rounded. The nasal tubes lie in shallow depressions, placed 52-66 from tip of snout, 17-30 from orbit, and 53-64 apart. The posterior half of each tube is raised into a pointed flap which can be used to constrict or close the nasal opening. Eyes rather small, diameter of orbit 45-82, placed entirely within upper half of side of head, either above or extending slightly below line between tip of snout and upper end of base of pectoral fin, not projecting into dorsal profile of head. Interorbital region very broad, its width 81-104. Ridges of supra- orbital canals, described above for snout, are continued through interorbital region on each side above eyes; these ridges are clearly visible on the small specimens. Medial portion of interorbital space flat and covered with elongate or finger-like papillae. Supraorbital ridges continue onto postorbital part of head as ridges of temporal canals, and extend to above operculum. Upper surface of head behind eyes almost flat, covered with papillae like those of interorbital region; posterior limit of papillae follows posterior line of head medially, but overlies post- temporal bone laterally. Length of postorbital part of head 167-198. Mouth broad, somewhat oblique, lower jaw projecting slightly; length of upper jaw 123-150, maxillary extending under anterior “4 to % of eye; width of jaws 176-180. Teeth all conical, arranged in 2 bands in each jaw. Outer bands uniserial, composed of enlarged, almost canine-like teeth; inner bands broader, especially anteriorly, composed of smaller and more slender teeth. Inner band of lower jaw extends only along anterior ’2 of jaw; that of upper jaw extends about full length of jaw. Outer bands of both jaws extend about full length of jaws, that of upper jaw being slightly longer than inner band. Gill rakers in anterior series of first gill arch short, blunt, somewhat flattened obliquely to long axis of arch and bearing teeth distally; arranged 5—7 + 0-1 + 11-15 = 17-22; in smaller specimens those near angle may be more elongate, bearing teeth along upper edges. Gill rakers of posterior series of first gill arch dentigerous and only slightly flattened at right angles to long axis of gill arch, arranged O—1 + 0-1 + 10-11 = 11-13; gill rakers of remaining arches similar. Branchiostegal rays 6. First dorsal fin 4—7, its origin 289-329 from tip of snout, above or slightly in advance of upper end of base of pectoral fin; second spine longest, its length 62-102. Second dorsal fin 27-30, its origin 400-457 from tip of snout and 17-75 from base of last spine of first dorsal fin; length of sixth ray 111-162, of sixth from last ray 97-122. Anal fin 22-26, its origin 505-577 from tip of snout, originating below bases of sixth to eighth rays of second dorsal fin; length Vor. XXXVIII] DEWITT: NOTOTHENIA FROM NEW ZEALAND 321 of sixth ray 96-138, of sixth from last ray 85-105. Caudal fin 14—16, its length 157-219, its posterior margin very slightly rounded, almost truncate. Pectoral fins 17-19, their length 193-240, not extending to origin of or reaching to above first four rays of anal fin, their posterior margins rounded, width of their bases 80-97. Pelvic fins placed 262—346 from origin of anal fin, entirely in advance of bases of pectoral fins, their length 174-217, third or fourth rays longest, not reaching posteriorly to origin of anal fin. Upper lateral line 45-61, terminating from below fourth from last ray to slightly behind posterior end of base of second dorsal fin, separated from origin of latter by 6~7 scale rows. Middle lateral line 9-18, extending a short distance onto base of caudal fin. Cephalic lateral line canals of normal pattern, the pores very small and difficult to see. Preoperculo-mandibular canals with 9-10 pores, connected to temporal canals; infraorbital canals with 8-10 pores; supraorbital canals each with 4 pores and sharing a median coronal pore; temporal canals with 5—6 pores; supratemporal canal normally with 3 pores, but in 1 specimen the canal is incomplete across the head and consists of a short tube on each side ex- tending dorso-medially from the temporal canals, each with a single pore at its end. Most scales on body ctenoid, 49-60 in a lateral longitudinal series, 27-31 rows around the caudal peduncle. Parrott (1958) records 61-69 scales in a lateral longitudinal series, but none of the specimens I have examined had counts that high; MacDonagh (1931) gives a lateral scale count of 68 for the holotype of N. patagonica, but I count only 60; Thompson (1916) records 67—73 lateral scales. Since Thompson’s counts were made along the lateral line, and above it, from the angle of the operculum to the base of the caudal fin, higher counts would be expected. It is probable that the other high counts were made in a similar manner. Nonctenoid scales present on belly and area anterior to bases of pectoral fins; a few may be found along bases of dorsal and anal fins. Scales extend onto basal parts of caudal fin and, except for a narrow naked crescent at bases of pectoral rays, onto lateral bases of pectoral fins. Scales absent directly in front of bases of pelvic fins, but medially they extend to area covered by fold of branchiostegal membrane across isthmus. Head almost entirely naked; a few scales, some of which may be ctenoid, present at posterolateral corners of head above temporal canals and on either side of supratemporal canal; a larger patch, some being ctenoid, present on upper part of operculum; a still larger patch, all nonctenoid, present on upper part of cheek behind eye; a few scattered scales may be present below eye. Ground color of BMNH specimen 1886.11.18.30 brown, somewhat lighter, perhaps originally white or yellowish, on belly. Head somewhat darker and greyish above. No prominent markings present on body. Sides of head with darker vermiculations creating a marbled appearance; these markings continued 322 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. onto lateral parts of snout, lips, lower jaw, and faintly onto branchiostegal rays. All vertical fins more or less uniformly brownish-dusky; second dorsal fin with indistinct and irregular darker brown markings on rays; anal fin with 1 or 2 series of darker markings on rays, tending to form horizontal lines. Rays of pec- toral fins with brown spots, arranged to form bars on left side, but irregular on right side; pelvic fins with faint marbling similar to that on sides of head. The USNM specimen from New Zealand is nearly entirely a uniform dark grey-brown. The larger of the 2 Canterbury Museum specimens has on the body irregular light areas over a dark background. The head is uniformly dark above and on the snout, but on each cheek is a series of 4 light lines, partially broken into spots, which radiate from the ventral and posterior parts of each eye; irregular light spots are present on the operculum. The vertical fins are generally dark; the second dorsal fin has 1 to 3 light spots along its rays creating horizontal lines, most distinct anteriorly and basally; the anal fin shows light areas, irregularly arranged anteriorly, horizontally arranged posteriorly; tips of rays of second dorsal and anal fins pale. The pectoral fins show only faint barring and spotting. The smaller specimen is essentially the same as the larger, but the light areas on the sides of the head are larger and less broken. Little has been recorded of life colors. Hutton (1875; p. 316) gives “Variable in color from dark olivaceous black to olive-green, slightly mottled with blackish on the back; lips speckled with white; axil of pectorals yellow; caudal and dorsal blackish.” In his description of NV. porteri Delfin (1899b; pp. 119-120) gives some color notes for the South American representatives of the species. The color of the iris is reddish yellow and the conjunctiva is green speckled with greenish yellow spots. The cheeks are described as hoary, with a scaled appearance due to the coloration, which probably refers to the dark vermiculations described above, which do sometimes look like scales. Most of the body is a greenish brown, with blackish overtones above, becoming paler ventrally; there also may be 1 or 2 longitudinal bands. Rays of pectoral fins with yellow spots, largest basally; the axil is yellow. Membranes of dorsal and anal fins dusky green, with spots of two shades of greenish yellow. Caudal fin greenish brown with a pale vertical band. DistriBuTiIon. I can find no essential differences between the New Zealand and South American material and I therefore concur with Norman (1937b; p. 91) that specimens from the two areas represent the same species. Such a broad distribution is not surprising when one considers the broad distributions of other closely related species (NV. coriiceps, N. rossii, and N. magellanica), all of which have characteristic pelagic young. Although no pelagic juveniles of this species have been found, it is probable that they do exist. Night-light fishing in the waters east of New Zealand might prove fruitful, for many pelagic juveniles of the other three species have been obtained in this manner. VoL. XXXVIII] DEWITT: NOTOTHENIA FROM NEW ZEALAND 323 Discussion. The use of the specific name “angustata’” for this species and the inclusion of Notothenia porteri in its synonyny represents a radical departure from the interpretations of all workers since Hutton’s time. My reasoning is as follows. Probably no one took the trouble to read Delfin’s description carefully, for the number of spines and rays in the dorsal and pectoral fins, and the color description, which have been abstracted and brought together above, clearly indicate that the species cannot be V. magellanica (D; 4-6; Dz 28-30; P 18-19 in NV. porteri vs. D; 3-6; Dz 29-31; P 16-18 in NV. magellanica). The realization that something was amiss in the interpretations of Hutton’s work by later authors came as a result of attempting to place the various early names applied to New Zealand nototheniids with the 5 species recognized from the area by Parrott (1958). Although it became clear that Hutton had himself confused species, certain important discrepancies were found between his descriptions of N. angustata and N. microlepidota and the species to which the names were applied. These include fin ray counts, scale counts, color, and shape of the caudal fin. I concluded that the name Notothenia angustata should apply to the species which has been called NV. microlepidota by all authors since Hutton’s time, and that the latter name should apply to the species which have been called NV. colbecki and N. filholi (see discussion under NV. microlepidota). The confusion can probably be traced back to the 1880’s, when a number of fishes were given to the British Museum by the Otago Museum, including some type material. Among these fishes is a specimen identified as VV. microlepidota which, although never labeled as such, was presumed to be the type (see Norman, 1937b; p. 89). I have examined this specimen (BMNH 1886.11.18.30) and it does belong with the species described here. Boulenger (1902; p. 185) was the first to apply the name N. microlepidota to the present species and, because he gave in the same paper an excellent description of the true V. microlepidota under the new name NV. colbecki, all later authors followed him. In an effort to obtain further and better evidence to support my belief that Hutton’s species had been confused, I wrote to the Otago Museum in Dunedin, New Zealand. Dr. D. R. Simmon of that institution was kind enough to locate the Notothenia material in the museum and to discover that the types of NV. angustata and N. microlepidota are probably there. Although there are no data which demonstrate that the Otago Museum specimens definitely are the types, the circumstantial evidence is very strong. In his 1876 paper, which redescribes both NV. angustata and N. microlepidota, Hutton stated that the types of both species were in the Otago Museum. Further, the lengths given by Hutton are close to those measured by myself on the stuffed specimens, my measurements being greater for both species (VV. angustata: WHutton’s length “about 14.5 inches,” equals 368 mm.; my measurement, TL = 407 mm.; NV. microlepidota: 324 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Hutton’s length ‘about 17 inches,” equals 432 mm.; my measurement 492 mm.). Mr. P. O’Brian, the preparator at the Otago Museum, stated that the process of stuffing tends to lengthen specimens slightly, and this may account for the ap- parent greater size of the stuffed specimens. There are other discrepancies be- tween the original accounts and my own data, the most serious being the pectoral count of 18 for N. microlepidota (1 counted 21 rays). This count is difficult to make on large specimens, however, because of thick investing skin, and I can only conclude that Hutton’s count is in error. The same may be said for my scale counts, which were made with difficulty because of the heavy lacquer with which the specimens are coated. Other differences between Hutton’s published accounts and my own observations include dorsal and anal fin ray counts. Whether one believes, as I do, that the specimens in the Otago Museum are the types, or because of the above discrepancies one believes that they are not, the interpretations of Hutton’s species by Boulenger, Waite, Norman, and Regan are untenable. Notothenia angustata is described as having “a bony ridge over each eye extending back to the posterior margin of the praeoperculum,” the “Caudal rounded,” the “Lips speckled with white,’ 19 rays in the pectoral fin and 52-58 scales in a lateral longitudinal series. The supraorbital ridges, rounded caudal fin and number of rays in the pectoral fin clearly distinguish it from NV. magellanica and demonstrate that it is the same as the V. microlepidota of the above authors. Hutton described NV. microlepidota as having 91 scales in a lateral longitudinal series, 12 scale rows between the origin of the second dorsal fin and the upper lateral line, and a truncate caudal fin. These characters are all incompatible with the species which has been called N. microlepidota and show its identity with V. colbecki and N. filholi (see discussion under N. micro- lepidota). Finally, the name ‘“‘microlepidota”’ certainly refers to the small and numerous scales implied by the high counts given in the original description, and which is most inappropriate if the conventional interpretation of the 2 species is accepted. Table 4 presents the data for the types of N. angustata and N. microlepidota, together with those for the types of other species synonymized with them. I have seen the holotype of NV. latifrons (USNM 76854), but it is not in good condition and I did little with it. Thompson’s description of the species seems good, and since the upper lateral line count of 51-56 is diagnostic for this species I again concur with Regan (1916) and Norman (1937b) that NV. latifrons belongs in the synonymy of N. angustata. Notothenia parva was described from 4 specimens ranging in size from 3 to 3% inches in length (equals 76-89 mm.). I have been able to locate three of these specimens, which are now deposited in 3 different institutions: the British Museum, the Dominion Museum, and the Otago Museum. Since the specimen in the British Museum is not mounted in gelatine or on a glass plate and is the VoL. XXXVIII] DEWITT: NOTOTHENIA FROM NEW ZEALAND WwW i) On most accessible, I designate it as the lectotype, the two others thereby becoming paralectotypes. The only remaining nomenclatural problem concerning this species is the identity of NV. maoriensis Haast, which has priority over the name “‘angustata.” However, it is not possible to determine without any doubt whether the original description applies to V. magellanica or to the present species. Characters which indicate an identity with \V. magellanica are the first dorsal fin with only 3 spines, the lack of any mention of supraorbital ridges on the head, and the dark coloration and lack of any speckling on the head. Characters which indicate an identity with V. angustata are scales present below the eye, the posterior end of the upper lateral line ending below the last ray of the second dorsal fin, and the shape of the pectoral fin as shown in the figure published with the original description. The illustration might constitute conclusive evidence except that it is a relatively crude drawing and contains some obvious errors which indicate it was not made with care. In the description the second dorsal fin is said to have 29 rays; the drawing shows only 27, and the membranes are drawn in a manner not found in any specimens belonging to Notothenia. The pelvic fins are drawn with a spine and 6 rays with the first ray longest; I have never ex- amined a specimen of Notothenia with 6 pelvic rays, and the third or fourth rays are longest, never the first. For these reasons I cannot trust the shape shown for the pectoral fin. Further, large specimens of V. magellanica have a number of low papillae below and behind the eyes, many of which are broad and flattened and appear similar to scales. It is possible that these were mistaken for scales by Haast. To conclude, there is enough doubt concerning the identity of NV. maoriensis to make me follow Regan and Norman in placing it with V. magellanica. Notothenia microlepidota Hutton. Notothenia microlepidota Hutton, 1875: 316 (original description; type locality Dunedin and Moeraki (45°23’S., 170°52’E.), New Zealand; holotype in Otago Museum, Dunedin) ; Hutton, 1876: 213 (virtual reprint of 1875 original description) ; Hutton, 1879: 339 (listed with counts); Hurron, 1890: 280 (listed); Grrr, 1893: 118 (listed); Warts, 1907: 30 (listed) ; Fow1er, 1945: 130 (listed). Nototoenia filholi SauvaGE, 1880: 228 (original description; type locality Campbeli Island; types in Museum National d’Histoire Naturelle, Paris) ; FrmHor, 1885: 345 (reprinting of Sauvage’s description). Notothenia filholi Dotxo, 1904: 127 (listed) ; VArLLANT, 1907: 22-23 (redescription of syn- types; correction of errors made by Sauvage); RecAan, 1913: 278 (description from Vaillant and Sauvage); Puirirrs, 1927a: 13 (listed); Puirtipps, 1927b: 44 (listed) ; Bianc AND HurEAv, 1962: 341-342 (disposition of syntypes). Notothenia colbecki BOULENGER, 1902: 185, pl. 16 (original description and illustration; type locality Campbell Island; types in British Museum); Warre, 1907: 30 (listed); WarrTe, 1909: 594-595 (description) ; REGAN, 1913: 278 (description) ; WAITE, 1916: 70 (listed) ; REGAN, 1916: 378 (distribution); RENDAHL, 1925: 6 (listed); PuHimtipps, 1927a: 13 [Proc. 4TH SER. CALIFORNIA ACADEMY OF SCIENCES 326 801 fifi GAS Tvs g's = — — OIT ILI SLE = W-us 7°68 ST [ers L9r ros = — — ZOT 971 Sze = E@eus e19 901 67 BZ ose Sr — = Weil T'16 1382 = tq-us v'6¢ Weise 9°01 ii 6°81 — PLT — 119 = 9°¢1 = d-d T'92 8°79 rs O'rT est = 6'+1 — r'6e VSP SOT = Md Z7'6¢ 9°9L eel (Pi rz — rSZ — 97S O19 VST — dd SST o'0¢ 6S 9°8 3'8 = vOL — 9'+Z 162 ol) = dado ZA Z'7r 6'6 ran | 6'¢1 = ei — Z'bZ OTE a = 1d9 7772 0'6¢ o8 SI Para LST = = 9're Zeb 76 = anti ose €19 cau 9°9T O'ST SYA 102 = Orr — ee = Od 761 Ore g's 78 06 VII 9°6 Sh 772 9°97 g's e9¢ OI Gel 8°02 wy gs g's oy — = 6'¢1 Cab Lv = N-N 0) 9°61 o¢ eS 9's a — — 6° ET eS LEG: os N-us 7 LT L0¢ 6S 18 Z'6 9°71 = = Gag $82 09 = us a 691 9's 39 89 76 0'6 — 771 anal Ls = O $79 SOT 872 ST¢ see Seb — aa! 7S £96 €7Z SIT TH (qnoqe) (qnoqe ) 961 Tee Poy, ZOT III IST 971 ber O¢z Ste LT Sre TES Ae: ess Boa Wo= Bisx ers ae a2 ae qs= Ga = = 2 pene Goa ree Ss ee Fee Gee Bae 8 ee es ee ee z Bis she Bro! oS Sie .S) SPS Bs gs a 2S a ve oot gs 5 es is a+ RES RSs crs SS £F 2s go B58 oR S See ae. See ees BOS ees Bee S Sse Bus So” < S Sos Ose SESS 2 SS ~ & $ EN, ABS, A 5 2 It if el yale ells S = a8 IS oe s ens ee he ee ae : eee Z 2 Z 2 Z * R 4 ae jap) q q ae sn 30] Kyonba asv SKDA Od} ayMIIpUL SaAN321f 0024 asaym “YASUOT Aof 7darxa ‘apis JysiA ayy WoAf UIYD] St PUOIAS AY] “UAAIS aD SJUNOI AO SJUAMAANSDAWM OM a4aY MY “7 31QD] U2 SD adAD SUONDIAaAQQHY “Way, {O SWKUOUKS SD PadsapisuoD SamDU Sutipaq suamads amos fo pup ejopidajort “N ‘e}E}sNBue vIUIYy}OJON fo Sadak ay. mor S]UNDD PUD ("Miu UL) SJuamaMspayyY “pb ATAVE$ VoL. XXXVIII] TABLE 4. (continued) (9d4}0}90[R1R q ) TLZ-OL'8° TT TO6T HNING °24929109 N (9d4}0}907T ) TZ-OL 8° TT 1061 HNING +74999/09 “N (9d4}0}90/P1k J ) bl-ZL 8 TT T06T HNING :74999]09 “N (9d4}0}99]R18 J ) blL-ZL 8 TT TO6T HNING +24999]09 “N (9d4}0}99]R1R J ) vL-CL 8 TT TO6T HN +24999199 “N (9d4}0}99'T) bsecV Wd oy *N (9d4}0}99[R1R J ) p8EcV Wd ours *N (od4 J) :DyopidajosIIM * Ny (adAjeieg ) pLloeol HNWG :pamMosDypd * Ny (ed4}0]07 ) T-Of-IIX-cl dT :pI1M0sDIDd * (904404997) 67 ST TL 988T HNING 2224040 -N (ada) [DIDISNIUD * AT uoT]BAIISq¢ DS}. 30.0 19.5 104 Shilss} 38.4 67.4 93.8 A-V 36.0 57.4 15.8 19.0 19.2 39.4 CL PL 28.7 Bde) lel 17.0 66.2 38.2 27.0 & 26.5 60.8 46.6 2 ine) = Se) 54.4 20.1 19.1 941418 23.6 N33 15.6 55.0 41.2 VL AntGR oO N 28 II == 73} = i) = 10) = 110) Di 27 27 27 27 24 14 23 about 14 29 29 29 29 about 27 24 24 about 24 — 16 28 25 Dz ine) N 14 20 & 20 23 14 20 2 14 1 20 23 14 21 20 & 20 20& 21 20 AAI 18 & 18 18 &17 18 & 17 3&4 19 Se 2&3 about 80 3&4 LongVR LatSce 84 10 41 74&72 32 & 32 88 11 39 67 & 67 34 & 34 89 96 10 45 98 10 40 about 87 about 90 51 60 59 52 DEWITT: NOTOTHENIA FROM NEW ZEALAND 11 43 75 &74 36 & 37 10 LL-D2 28 59 & 56 31 55 & 53 16 & 12 28 54 & 52 14 & 14 ScArGP ULL 71 35 & 32 73 72 70 about 30 about 30 about 71 52 &55 & 32 28 12 & 12 7, MLL 327 328 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. & ~ Ficure 5. Notothenia microlepidota. Lateral view and top of head of adult, and lateral view of young; from Boulenger, 1902. (listed) ; PuHitiipps, 1927b: 44 (listed); Norman, 1938: 27 (distribution); PArRrort, 1958: 112-113 (description). MATERIAL EXAMINED. PM A2384: Campbell Island (2; about 126 and 151 mm., not in good condition; paralectotype and lectotype, respectively, of N. filholi). BMNH 1901.11.8.70-71: Campbell Island (2; 196 and 331 mm.; paralectotype and lectotype, respectively, of V. colbecki). BMNH 1901.11.8.72—74: Campbell Island (3; 73.1-111 mm.; paralectotypes of NV. colbecki). DM 2734: Campbell Island (4; 111-141 mm.). The following New Zealand material was also examined, but not used for descriptive purposes. In the Dominion Museum: 1413, Tucker Cove, Campbell Island (1); 2084, off Big South Cape Island (1); 3123, off rocks in N. W. Bay, Campbell Island (1); 3333, outer Ranui Cove, Auckland Island (2). In the Canterbury Museum (uncatalogued): Campbell Island (3); Per- severance Harbour, Campbell Island, from throat of Shag (1); Perseverance Harbour, Campbell Island (1); Penguin Harbour, Campbell Island (1); Auck- land Islands (2). Description. Body fusiform, compressed throughout, including head (ex- cept in largest specimen); dorsal and ventral profiles nearly evenly curved throughout, a little more strongly so anteriorly, dorsal profile sometimes slightly Vot. XXXVIII] DEWITT: NOTOTHENIA FROM NEW ZEALAND 329 more convex than ventral profile; caudal peduncle distinctly longer than deep. Length of head 288-341, its width 126-247, its depth 164-182; depth of body 182-232, its width 115-190, pectoral to pectoral distance 138-254, dorsal to anal distance 196-254; length of caudal peduncle 123-137, its depth 79-91; dorsal to caudal distance 124-144. Vertebrae 18 + 27-28 = 45—46. Snout smoothly rounded from both lateral and dorsal views, rising from tip of upper jaw at about same angle as top of head; its length 81-100. Nostrils tubular, elliptic in cross section, each with its hind margin raised into a flap ending in a rounded point; nostrils placed 48—69 from tip of snout, 17-23 from orbit, and 52—63 apart. Eyes directed laterally, placed high on head, above a line between tip of snout and upper end of base of pectoral fin, but not protruding into dorsal profile of head; diameter of orbit 51-77. Interorbital space broad and nearly flat, only very slightly convex, its width 66-103. Length of postorbital part of head 154-185. Mouth oblique, lower jaw projecting slightly in front of upper jaw; length of upper jaw 104-128, maxillary extending posteriorly under first third of eye. Teeth in upper jaw may be described for convenience as being in 2 bands; outer band a uniserial row of enlarged, spaced (canine-like) teeth, extending only along anterior half of jaw; inner band composed of smaller, more closely spaced teeth, slightly broadened anteriorly, becoming a uniserial row posteriorly; inner teeth become slightly larger posterior to point where outer row ends. Teeth in lower jaw may be described as occurring in a single band, somewhat broadened an- teriorly, with outermost teeth largest, and becoming a uniserial row of enlarged teeth in posterior two-thirds of jaw, the teeth becoming smaller posteriorly. Oral valves extend nearly entire length of jaws; they may be covered with papillae or not. Tongue rounded and free anteriorly, with a slight depression in its upper surface, and covered with scattered low papillae. Gill rakers in anterior series of first gill arch slender and elongate, arranged 6—11 + O-1 + 15-19 = 24-30; those on lower limb near angle slightly flattened on ventral edge, those further below flattened dorsoventrally, those on upper limb more cylindrical; all bear a few to many teeth, those on lower limb near angle with fewest. Posterior gill rakers of first arch short and blunt, somewhat flattened dorsoventrally, and bearing teeth; arranged 1-3 + 1 + 14-15 = 17-19. Branchiostegal rays 6. First dorsal fin 6-8, its origin 295-338 from tip of snout, from above upper end to just in advance of bases of pectoral fins; its height relatively low, length of longest spine 86-110. Second dorsal fin 25-29, its origin 413-467 from tip of snout, 28-48 from base of last spine of first dorsal fin; highest anteriorly, length of sixth ray 105-130, of sixth from last ray 75-79. Anal fin 21-24, its origin 507-570 from tip of snout, below bases of rays five to seven of second dorsal 330 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. fin; highest anteriorly, length of sixth ray 88-111, of sixth from last ray 76-87. Caudal fin 14, its length 173-216, its posterior margin distinctly emarginate, almost forked. Although the sample counted is small, the apparent lack of variation in the number of principal rays may be due to the emarginate shape of the fin, in which the principal unbranched rays are nearly as long as the longest branched rays and form most of the upper and lower edges of the fin. Pectoral fins 20-21, their length 190-232, not reaching to, or extending as far as, above fourth ray of anal fin, the posterior margin rounded; width of their bases 66-88. Pelvic fins placed 251-314 from origin of anal fin, entirely in advance of bases of pectoral fins; their length 156-197, third ray longest, not reaching posteriorly to origin of anal fin. Upper lateral line with 61—75 tubular scales, ending below last few rays of second dorsal fin or extending a short distance posterior to it, separated from origin of second dorsal fin by 9-11 scale rows. Boulenger (1902; p. 185) gives a range of 59-71, but his counts are low in every case. Table 4 presents my counts from the same specimens (see under discussion), which range from 67—75. Middle lateral line with 24-37 tubular scales, originating below ninth to fifteenth rays of second dorsal fin, and extending onto base of caudal fin. Cephalic lateral-line canals normal in pattern except that preoperculo- mandibular canals are joined to temporal canals. The pores are small and diffi- cult to find. Preoperculo-mandibular canals with 9-10 (usually 9) pores, connected to temporal canals at areas of second pores of latter canals; infra- orbital canals with 9-11 (usually 10) pores; supraorbital canals each with 4 pores and sharing a median coronal pore; temporal canals with 6 pores; supra- temporal canal with 2—4 (usually 3) pores. Scales everywhere small, 84-98 in a lateral longitudinal series, with 37—45 rows around caudal peduncle; on body ctenoid except dorsally anterior to first dorsal fin, anterior to bases of pectoral fins, on ventral surface anterior to pelvic fins, and sometimes on lower sides of body between pelvic fins and anterior few rays of anal fin. A few nonctenoid scales may be found scattered among the ctenoid scales, especially at base of caudal fin, and the number of ctenae may be reduced to one. Scales extend onto basal parts of caudal fin and, except for a naked arc at bases of rays, onto exposed proximal portions of pectoral fins. Most of head naked; 2 patches of scales, some of which are ctenoid, present on each side at posterolateral corners of head, one just anterior to supratemporal canal, the other in triangle formed by temporal canal, supratemporal canal and very weak ridge of posttemporal bone. An elongate patch of nonctenoid scales present on uppermost part of operculum; a patch of similar scales present on upper and anterior part of cheek, extending ventrally and anteriorly in a nar- rowing arc around posterior and ventral margins of eye. Upper portions of head, including snout, lips, and naked parts of cheeks, as well as other parts in lesser Vot. XXXVIII] DEWITT: NOTOTHENIA FROM NEW ZEALAND 331 degree, covered with scattered and low papillae or ridges, the most marked being on top of head and anterior parts of lower lip and jaw. Ground color of body (in alcohol) uniformly brownish or greyish, becoming lighter ventrally; lower half of body may be somewhat silvery (this probably re- flects the method of initial fixation). Both dorsal fins dusky to deep brown; anal fin pale to brown; pectoral fins slightly brownish basally; pelvic fins a little dusky distally; caudal fin slightly dusky. Upper surface of head and tip of lower jaw dark, head otherwise becoming lighter ventrally; lower halves of operculum and cheek may be silvery. Indistinct and irregular dark areas may be present on top of head; a dark patch may be present behind eye at level of upper end of preopercular. Two dark lines may be present on lower parts of cheek, one extending from edge of upper jaw above end of maxillary posteriorly and ventrally towards lower margin of preopercular, the other ex- tending from ventral margin of eye towards angle of preopercular. A third line, extending from posteroventral edge of eye to upper end of preopercular, may be present, and the dark patch behind the eye mentioned above may repre- sent this line. Juvenile specimens are somewhat silvery in color and, although there are no striking color changes between the young and adults as seen in JN. rossit, the silvery color may indicate that the young of this species are also pelagic in habit. Little has been recorded of life colors. Hutton (1875) gives ‘“Purplish brown above, greyish below; throat, gill-membranes, axil of pectorals, and opercles yellowish.” Parrott (1958) notes that a specimen from Auckland Island was dark olive-green with dark red bands on the dorsal and ventral fins. DIstRIBUTION. Notothenia microlepidota is known only from the New Zealand region, including Macquarie Island. Its habits are apparently similar to those of NV. angustata, specimens having been captured primarily with hooks and lines close to shore. Discussion. Since the nomenclature and synonymy used for this species are totally different from those used by previous authors, some explanation of the present usage is desirable. My first suspicion that the names JV. filkoli and N. colbecki represented the same species was entertained upon reading the account by Filhol (1885; pp. 343-346) of his fishing efforts at Campbell Island. He stated that NV. filholi was the most common fish encountered there. Boulenger later described N. colbecki, also from Campbell Island, and it sub- sequently was found to be very common there, whereas N. filholi was never recorded again. Boulenger’s description is good and it was accompanied by an excellent figure; N. colbecki was therefore easily recognized by subsequent workers. Sauvage’s description, on the other hand, is not only brief, but con- 332 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER. tains a number of important errors (see Vaillant, 1907; p. 22, footnote), and no illustration was prepared. Thus JN. filholi was never again recognized, al- though the name continued to be included in keys and check lists because of the unusual counts which Sauvage had given. Vaillant (1907) corrected Sauvage’s errors, but his redescription and discussion has been disregarded. Regan (1913), apparently not knowing what to believe, gave the data of both Sauvage and Vaillant; later authors followed Sauvage. Through the courtesies of Dr. Maurice Blanc of the Museum National d'Histoire Naturelle, Paris, and of Mr. A. C. Wheeler and Dr. P. H. Green- wood of the British Museum (Natural History) I have been able to examine 2 syntypes of NV. filholi and 5 syntypes of NV. colbecki. Although the specimens of N. filholi are not in good condition, I was able to take some counts and measurements from them. These are presented in table 4 together with the more complete data from the specimens of NV. colbecki. There is no doubt that they all represent the same species, and the data from them have been incor- porated into the above description. I have related already the probable cause of the confusion attending Hutton’s species V. angustata and N. microlepidota (see discussion section under NV. angustata). Although there is some doubt whether the specimen in the Otago Museum thought to be the type of NV. microlepidota is indeed the type, since there are no records or catalogues dating back to the 1870’s, the original de- scription leaves no doubt that Hutton’s species is the same as both N. filholi and N. colbecki. The supposed type is now stuffed, and while its total length is somewhat greater than that recorded by Hutton, it is sufficiently near Hutton’s figure that the difference can be accounted for by the process of stuffing. The counts taken from the specimen are presented in table 4 for direct comparison with those from the types of \. filholi and N. colbecki, and show conclusively that the specimen, whether type or not, represents the same species as the others. A final matter is the selection of lectotypes from the type series of NV. filholi and N. colbecki. As the lectotype of the former name I choose the specimen of 151 mm. standard length (Paris Museum number A2384) listed above in the material examined. This is as nearly as I can tell the specimen which Vaillant used for his table (1907; p. 23) and is probably the specimen referred to by Sauvage in his original description when he gave a length of 350 mm. (corrected to 150 mm. by Vaillant, 1907; see also Blanc and Hureau, 1962; pp. 341-342). For the lectotype of the name NV. colbecki I choose the specimen of 331 mm. standard length (British Museum number 1901.11.8.70—71) listed above in the material examined. This is the largest of the specimens which remain of the type series, and is probably the specimen used for the figure of the adult published with the original description (there is some doubt because the legend for the plate states that the figure has been reduced Vor. XXXVIII] DEWITT: NOTOTHENIA FROM NEW ZEALAND WwW Ww Ww to %4, which would mean the specimen was a little over 500 mm. in standard length: if the above specimen was used for the illustration the reduction is about %). Boulenger also gives in his description a total length of 380 mm., which corresponds well with both Norman’s (1938; p. 27) and my measure- ments (385 and 388 mm., respectively) for the largest specimen in the series, and indicates that this specimen was considered as the type. Only 5 of the original 12 specimens of the type series remain, and they are undoubtedly the 5 specimens Boulenger used for his table of counts and measurements (Boulen- ger’s total lengths: 380, 230, 130, 120 and 85; my measurements: 388, 232, 130, 121 and 89). ACKNOWLEDGMENTS This paper is an extension of a portion of a doctoral dissertation submitted to the Department of Biological Sciences of Stanford University, and I take this opportunity to express my appreciation to Professor George S. Myers who, as my advisor, greatly aided me in formulating my dissertation topic and guided me through the initial stages of the work. The late Miss Margaret Storey of Stanford was very helpful in searching the literature and often gave me needed advice and encouragement. Much of the work was done at the University of Southern California where I participated in the Antarctic Marine Biological Research Program there, supported by the Antarctic Office of the National Science Foundation (grants G 19497, GA 238 and GA 448). Travel to and from the Antarctic research vessel Eltanin under these grants enabled me to visit museums in Argentina, New Zealand and Australia. Travel grants (GA 180 and GA 376) from the Antarctic Office enabled me to visit the British Museum in 1965 where I was able to examine the extensive Antarctic fish collection there and the Zoological Institute in Leningrad in 1966 where I was able to examine some of the recently collected Russian Antarctic material. I am much indebted to Dr. Jay M. Savage for leadership and advice during this period. The final writing was accomplished at the University of South Florida, St. Petersburg Campus. For the loan of specimens or for the innumerable courtesies shown me during my visits to museums in this and other countries, I give my most sincere thanks to the following persons: Drs. L. P. Schultz, E. A. Lachner, V. G. Springer, and S. H. Weitzman, and to Mr. R. Kanazawa of the United States National Museum; Dr. P. H. Greenwood, Mr. A. C. Wheeler, Dr. N. B. Marshall, and Miss V. duHeaume of the British Museum (Natural History); Drs. A. N. Svetovidov and A. P. Andriashev of the Zoological Institute in Leningrad; Professor K. Deckert and Dr. C. Karrer of the Zoologisches Museum, Humboldt- Universitit, Berlin; Professor M. Blanc of the Muséum National d’Histoire Naturelle, Paris; Dr. P. Kahsbauer of the Naturhistorisches Museum, Vienna; Drs. R. Lopez and N. Bellisio of the Museo Argentino de Ciencias Naturales, 334 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Buenos Aires; Dr. R. A. Ringuelet of the Museo de La Plata, La Plata; Mr. John Moreland of the Dominion Museum, Wellington; Mrs. Marie Darby (nee Biichler) of the Canterbury Museum, Christchurch; Dr. D. R. Simmon of the Otago Museum, Dunedin; and Mr. C. J. M. Glover of the South Australian Museum, Adelaide. I also thank the Trustees of the British Museum (Natural History) for permission to publish a reproduction of J. G. A. Forster’s manuscript illustration of Gadus magellanicus (fig. 1). Figure 3 was made possible through the courtesy of Dr. D. M. Cohen of the United States Bureau of Commercial Fisheries Ichthyological Laboratory, Washington, D. C. The staff in the Reference Department of the main library at the University of South Florida were very generous in their efforts to obtain for me publications not in their holdings. Finally, I thank Dr. Thomas L. Hopkins for critically reading portions of the manuscript. LITERATURE CITED ANDRIASHEV, A. P. 1959. On the number of vertebrae and some osteological characteristics of Antarctic fishes as shown by X-ray photographs. Voprosi Ikhtiologii, Moskva, no. 12, pp. 3-7 (in Russian). 1965. A general review of the Antarctic fish fauna. Monographiae Biologicae, vol. 15, pp. 491-550. ANDRIASHEV, A. P., AND A. K. TOKAREV. 1958. Ichthyofauna. The study of ichthyofauna and the goals of investigations. Trudy kompleksnoi antarkticheskoi Ekspeditsii. Akademiia Nauk S. S. S. R. (Opisanie Ekspeditsii d/e “Ob” 1955-1956), pp. 195-207 (in Russian). Betiisio, N. B. 1966. Fauna Marina Antartica. Republica Argentina, Secretaria de Marina, Servicio Hidrografia Naval, Publico H. 907, 93 pp. Branc, M. 1951. Poissons recueillis aux Iles Kerguelen par le Docteur Aretas. Bulletin du Muséum National d’Histoire Naturelle, Paris, ser. 2, vol. 23, pp. 493-496. 1954. Poissons recueillis aux Iles Kerguelen par P. Paulian (1951) et M. Angot (1952). Bulletin du Muséum National d’Histoire Naturelle, Paris, ser. 2, vol. 26, pp. 190-193. 1958. Sur quelques poissons des Iles Kerguelen rapportés par le Dr. Bourland. Bulletin du Muséum National d’Histoire Naturelle, ser. 2, vol. 30, pp. 134-138. 1961. Les poissons des terres australes et antarctiques francaises. Mémoires de l'Institut Scientifique de Madagascar, ser. F, vol. 4, pp. 109-159. Branc, M., ano J.-C. Hureavu. 1962. Catalogue des types de poissons de la famille des Nototheniidae, en collection au Muséum National d’Histoire Naturelle. Bulletin du Muséum National @Histoire Naturelle, Paris, ser. 2, vol. 34, pp. 339-342. BouLENGER, G. A. 1900. A list of the fishes collected by Mr. Rupert Vallentin in the Falkland Islands. The Annals and Magazine of Natural History, ser. 7, vol. 6, pp. 52-54. Vor. XXXVIII] DEWITT: NOTOTHENIA FROM NEW ZEALAND 335 1902. Pisces, pp. 174-189, pls. 11-18. Im Report on the Collections of Natural History made in the Antarctic Regions during the Voyage of the Southern Cross, London, British Museum (Natural History). CUNNINGHAM, R. O. 1871. Notes on the Reptiles, Amphibia, Fishes, Mollusca, and Crustacea obtained during the voyage of H.M.S. Nassau in the years 1866-69. Transactions of the Linnean Society of London, vol. 27, pp. 465-502. DetrFin, F. T. 1899a. Catalogo de los peces de Chile (continuacion). Revista Chilena de Historia Natural, vol. 3, pp. 15-24. 1899b. Descripcion de un nuevo Traquinido Chileno. Revista Chilena de Historia Natural, vol. 3, pp. 117-120. DeWt1rt, H. H. 1962. A new Antarctic nototheniid fish with notes on two recently described notothenii- forms. Copeia, 1962, pp. 826-833. Dotto, L. 1904. Poissons. Résultats du Voyage du S.Y. Belgica en 1897-1898-1899 sous le commandement de A. de Gelache de Gomery, Rapports Scientifiques, Zoologie, Anvers, 240 pp., 12 pls. FrrHo1, H. 1885. Poissons, pp. 343-346. In H. Filhol, Mission de l’le Campbell: Recherches zoologiques, botaniques et géologiques 4 l’Ile Campbell en Nouvelle Zélande. Recueil de Mémoirs, Rapports et Documents relatifs 4 Jl’observation du Passage de Venus sur le Soleil, vol. 3, pt. 2, no. 1. FIscHER, J. G. 1885. Ichthyologische und herpetologische Bemerkungen. I. Uber Fische von Sid- Georgien. Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten, Hamburg, vol. 2, pp. 49-65, pls. 1-2, figs. 1-4, 9. Forster, J. R. 1801. Gadus magellanicus, p. 10. In M. E. Bloch and J. G. Schneider, Systema Ichthyo- logiae Iconibus CX TIlustratum, Berolini. 1844. Descriptiones animalium quae in itinere ad maris australis terras per annos 1772 1773 et 1774 suscepto collegit obsevavit et delineavit . . . nunc demum editae auctoritate et impensis . . . Henrico Lichtenstein. Berlin, xiii + 424 =p ih isos Fow.er, H. W. 1926. Fishes from Florida, Brazil, Bolivia, Argentina, and Chile. Proceedings of the Academy of Natural Sciences of Philadelphia, vol. 78, pp. 249-285. 1945. Fishes of Chile. Systematic Catalog. El Imparcial, Santiago de Chile, 36 + 171 pp. 1951. Analysis of the fishes of Chile. Revista Chilena de Historia Natural, vol. 51-53, pp. 263-326. Frost, G. A. 1928. A comparative study of the otoliths of the neopterygian fishes. The Annals and Magazine of Natural History, ser. 10, vol. 1, pp. 451-456, 1 pl. Gut, T. N. 1862. Synopsis of the Notothenioids. Proceedings of the Academy of Natural Sciences of Philadelphia, 1861, pp. 512-522. 1893. A comparison of antipodal faunas. Memoirs of the National Academy of Sciences, Washington, vol. 6, pp. 89-124. 336 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. GUNTHER, A. 1860. Catalogue of the Acanthopterygian fishes in the collection of the British Museum. vol. 2. London, British Museum, xxi + 548 pp. 1881. An account of the zoological collections made during the survey of H.M.S. Alert in the Straits of Magellan and on the coast of Patagonia. III. Reptiles, Batrachians, and Fishes. Proceedings of the Zoological Society of London, 1881, pp. 18-22, 2 pls. Haast, J. 1873. Notes on some undescribed fishes of New Zealand. Transactions and Pro- ceedings of the New Zealand Institute, vol. 5, pp. 272-278, pl. 16. EVAR eel) fe 1946. Report on trawling surveys on the Patagonian continental shelf, compiled mainly from manuscripts left by the late E. R. Gunther, M. A. Discovery Reports, vol. 23, pp. 223-408, 1 pl. Hussaxor, L. 1914. Notes on a small collection of fishes from Patagonia and Tierra del Fuego. Bulletin of the American Museum of Natural History, vol. 33, pp. 85-94. Hutton, F. W. 1872. Catalogue with diagnoses of the species, pp. vii-xvi-+ 1-93. Jn F. W. Hutton and J. Hector, Fishes of New Zealand. Colonial Museum and Geological Survey Department, Wellington. 1873. Contributions to the ichthyology of New Zealand. Transactions and Proceedings of the New Zealand Institute, vol. 5, pp. 259-272, pls. 7-12, 15. 1875. Descriptions of new species of New Zealand fish. The Annals and Magazine of Natural History, vol. 16, pp. 313-317. 1876. Contributions to the ichthyology of New Zealand. Transactions and Proceed- ings of the New Zealand Institute, vol. 8, pp. 209-218. 1879. Notes on a collection from the Auckland Islands and Campbell Island. Trans- actions and Proceedings of the New Zealand Institute, vol. 11, pp. 337-343. 1890. List of the New Zealand fishes. Transactions and Proceedings of the New Zealand Institute, vol. 22, pp. 275-285. Kenny, R., anp N. Haysom. 1962. Ecology of rocky shore organisms at Macquarie Island. Pacific Science, vol. 16, pp. 245-263. LapIcEes, W., G. VON WAHLERT, AND E. Morr. 1958. Die Typen und Typoide der Fischsammlung des Hamburgischen Zoologischen Staatsinstituts und Zoologischen Museums. Mitteilungen aus dem Ham- burgischen Zoologischen Museum und Institut, vol. 56, pp. 155-167. LONNBERG, E. 1905. The fishes of the Swedish South Polar Expedition. Wissenschaftliche Ergebnisse der Schwedischen Sitidpolar-Expedition 1901-1903, vol. 5, pt. 6, 69 pp., 5 pls. 1906. Contributions to the fauna of South Georgia. I. Taxonomic and _ biological notes on vertebrates. Kungliga Svenska Vetenskapsakademiens Handlingar, new series, vol. 40, no. 5, 104 pp., 12 pls. 1907. Fische. Ergebnisse Hamburger Magalhaensischen Sammelreise, 1892/93, vol. 8, 16 pp., 1 pl. MacDonacu, E. 1931. Nota preliminar sobre Bovichthys argentinus y Notothenia patagonica n. spp. Notas Preliminares del Museo de La Plata, vol. 1, pp. 99-100. Vor. XXXVIII] DEWITT: NOTOTHENIA FROM NEW ZEALAND 337 1934. Nuevos conceptos sobre la distribucion geografica de los peces Argentinos. Revista del Museo de La Plata, vol. 34, pp. 21-170. 1936. Sobre algunos peces marinos. Notas del Museo de La Plata, Buenos Aires, vol. 1, Zoologia, no. 4, pp. 423-429. MacDonacH, E., AnD M. R. Covas. 1944. Peces Patagonicos y Fueguinos coleccionados por el Doctor Federico G. Lynch. Notas del Museo de La Plata, Buenos Aires, vol. 9, Zoologia, no. 76, pp. 229-241, 2 pls. MoreELanp, J. 1957. Report on the fishes, p. 34. Jn G. A. Knox, General account of the Chatham Islands 1954 Expedition. New Zealand Department of Scientific and Industrial Research, Bulletin 122. Norman, J. R. 1937a. Fishes. Banzare Reports, Ser. B (Zoology and Botany), vol. 2, pp. 49-88. 1937b. Coast fishes. Part II. The Patagonian region. Discovery Reports, vol. 16, pp. 1-150, 5 pls. 1938. Coast fishes. Part III. The Antarctic Zone. Discovery Reports, vol. 18, pp. 1-104, 1 pl. NYBELIN, O. 1947. Antarctic fishes. Scientific Results of the Norwegian Antarctic Expeditions 1927-1928 et Sqq., no. 26, 76 pp., 6 pls. 1951. Subantarctic and Antarctic fishes. Scientific Results of the “Brategg” Expedition 1947-48, no. 2, 32 pp. OLIVER SCHNEIDER, C. 1943. Catalogo de los peces marinos del litoral de Concepcién y Arauco. Boletin de la Sociedad de Biologia de Concepcion, Chile, vol. 17, pp. 75-126. OLsEN, S. 1954. “South Georgian Cod” (Notothenia rossit marmorata Fischer). Norsk Hvalfangst- tidende (The Norwegian Whaling Gazette), vol. 43, pp. 373-382. 1955. A contribution to the systematics and biology of Chaenichthyid fishes from South Georgia. Nytt magazin for Zoologi, vol. 3, pp. 79-93. Parrott, A. W. 1958. Fishes from the Auckland and Campbell Islands. Records of the Dominion Museum, Wellington, vol. 3, pp. 109-119. Peruci, A. 1891. Appunti sopra alcuni pesci Sud-Americani conservati nel Museo Civico di Storia Naturalle di Genova. Annali del Museo Civico di Storia Naturali di Genova, ser. 2, vol. 10, pp. 604-657. Peters, W. C. H. 1876. Ubersicht der wahrend der von 1874 bis 1876 unter dem commando des Hrn. Capitan z. S. Freihern von Schleinitz ausgefiihrten Reise S. M. S. Gazelle gesammelten und von der Kaiserlichen Admiralitat der Koniglichen Akademie der Wissenschaften iibersandten Fische. Monatsbericht der Koniglichen Akademie der Wissenschaften zu Berlin, Sitzung der physikalish-mathematischen Klasse, 1876, pp. 831-854. Puitiiers, W. J. 1921. Notes on the edible fishes of New Zealand, with a record of fishes exposed for sale in Wellington during 1918. The New Zealand Journal of Science and Technology, vol. 4, pp. 114-125. 338 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. 1927a. A check list of the fishes of New Zealand. Journal of the Pan-Pacific Research Institute, vol. 2, pp. 9-15. 1927b. Bibliography of New Zealand Fishes. New Zealand Marine Department, Fisheries Bulletin no. 1, 68 pp. REGAN, C. T. 1913. The Antarctic fishes of the Scottish National Antarctic Expedition. Transactions of the Royal Society of Edinburgh, vol. 49, pp. 229-292, 11 pls. 1916. Bibliographical notice. Antarctic and Subantarctic fishes. The Annals and Magazine of Natural History, ser. 8, vol. 18, pp. 377-379. RENDAHL, H. 1925. Papers from Dr. Th. Mortensen’s Pacific Expedition 1914-16. XXX. Fishes from New Zealand and the Auckland-Campbell Islands. Videnskabelige Meddelser fra Dansk Naturhistorisk Forening i Kobenhaven, vol. 81, pp. 1-14. RICHARDSON, J. 1844-1848. Ichthyology of the voyage of H.M.S. Erebus and Terror, under the com- mand of Captain Sir James Clark Ross, R.N., F.R.S., viii 139 pp., 60 pls. In J. Richardson and J. E. Gray (eds.), The Zoology of the Voyage of H.MS. Erebus and Terror, under the command of Captain Sir James Clark Ross, R.N., F.R.S., during the years 1839-1843, vol. 2. Ruup, J. T. 1954. Vertebrates without erythrocytes and blood pigment. Nature, London, vol. 173, pp. 848-850. SAUVAGE, H. E. 1880. Notice sur quelques poissons de l’Ile Campbell et de l’Indo-Chine. Bulletin de la Société Philomathique de Paris, vol. 4, pp. 228-233. SLACK-SMITH, R. J. 1962. A small collection of fish from Macquarie Island. Memoirs of the National Museum of Victoria, Melbourne, no. 25, pp. 13-16. Smitt, F. A. 1897. Poissons de l’Expedition Scientifique a la Terre de Feu. I. Nototheniae. Bihang Till Kongl. Svenska Vetenskaps-Academiens, Handlingar, Stockholm, vol. 235) pts 45) NOs 6.167 PPro) piss STEINDACHNER, F. 1875. Ichthyologische Beitrage (iii). Sitzungsberichte der K. Akademie der Wissen- schaften in Wien, I abt., vol. 72, pp. 29-96, 8 pls. 1898. Die Fische der Sammlung Plate. Zoologische Jahrbiicher, Supplement-Band 4, Fauna Chilensis, vol. 1, pp. 281-338, pls. 15-21. 1903. Die Fische der Sammlung Plate (Nachtrag). Zoologische Jahrbiicher, Sup- plement-Band 6, Fauna Chilensis, vol. 3, pt. 1, pp. 201-214. STUDER, T. 1879. Die Fauna von Kerguelensland. Archiv ftir Naturgeschichte, vol. 45, pp. 114-141. Tuompson, W. F. 1916. Fishes collected by the United States Bureau of Fisheries steamer Albatross during 1888, between Montevideo, Uruguay, and Tome, Chile, on the voyage through the Straits of Magellan. Proceedings of the United States National Museum, vol. 50, pp. 401-476, pls. 2-6. THomson, G. M., Aanp T. ANDERTON. 1921. History of the Portobello Marine Fish-Hatchery and Biological Station. Dominion of New Zealand, Board of Science and Art, Bulletin 2, 131 pp. Vor. XXXVIII] DEWITT: NOTOTHENIA FROM NEW ZEALAND 339 VAILLANT, L. 1888. Poissons. Mission Scientifique du Cap Horn, 1882-1883, vol. 6 (Zoologie), pt. 1, 35 pp., 4 pls. 1907. Poissons. Expedition Antarctique Francaise (1903-1905), Sciences Naturelles, Documents Scientifiques, Paris, 51 pp. Watte, E. R. 1907. A basic list of the fishes of New Zealand. Records of the Canterbury Museum, vol. 1, pp. 3-39. 1909. Vertebrata of the Subantarctic islands of New Zealand. Pisces, pp. 585-598. In C. Chilton (ed.), The Subantarctic Islands of New Zealand, Philosophical Institute of Canterbury, Wellington, vol. 2. 1916. Fishes. Australasian Antarctic Expedition, 1911-1914, Scientific Reports, Ser. C—Zoology and Botany, vol. 3, pt. 1, pp. 3-92, 5 pls., 2 maps. ' mace — La yy i ——. 7 y > f oes f ag vi i P a As j i a as : S _ 7 ‘ a ee 't 5 v Fw 1 “48 _ q -_ 7" - J , a’) : th ie Aas im na, ai gp aqin OP ee el eee af es | 2 Vien) ied 21a° ABATED rue My @ Wii (Ais 1 as ; : Peek i<4*; sod (ere haswre, Aritlis » Ra) ae © 340, ih Malem | 1H qeli’ lim © a oe ee oO | i re : _ .. 5 es a 7 s_% - PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Festschrift for George Sprague Myers Vol. XXXVIII, No. 17, pp. 341-346; 1 fig. December 31, 1970 HOW MANY RECENT FISHES ARE THERE? By Daniel M. Cohen Bureau of Commerical Fisheries Washington, D. C. It is a pleasure to dedicate this paper to Professor George S. Myers on the occasion of his 65th birthday. His interests in ichthyology have ranged widely and the topic of this paper seems especially appropriate, not least because he has been interested in this particular problem himself. Estimates of the number of species of Recent fishes in the current ichthyo- logical literature range from a low of 15,000-17,000 to a high of 40,000. Presented below is a brief list of some. SOME PAST ESTIMATES OF NUMBER OF RECENT FISH SPECIES Bailey (1960) gave 15,000 to 17,000, of which about 45 are Agnatha and about 575 are Chondrichthys. His estimate was apparently based on a group approach. Marshall (1965) mentioned that, “We know more than twenty thousand living kinds, but our inventory is by no means complete.’ He gave no basis for his estimate. Norman (1963) gave 25,000, with no mention of how the figure was reached. Myers (1958) stated there are, “. . . .33,000 or more living species of teleosts.”’ No mention of method of estimation was given. Schultz and Stern (1948) gave a figure of 40,000; however, Schultz (1965) later lowered his estimate to 32,000. No basis was given for either figure. [341] 342 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. PRIMARY le DIADROMOUS /seconoatt 0.6% y 8.1% ) Y y DEEP- BENTHIC 6.4% DEEP- PELAGIC 5.0% PGi 1.3% Ase “WARM oe J 39.9% 58. XN a MarR iné© SHALLOW. COLD Ficure 1. Percentages of Recent fish species living in various habitats. MY ESTIMATE OF NUMBER OF RECENT FISH SPECIES The wide range of figures suggests that a rational estimate, as opposed to an educated guess, is difficult for any one ichthyologist. With this in mind I com- piled a list of fish families and began to solicit estimates from specialists and to consult recent revisions. Seven years have gone by since the initiation of the project, and this seems an appropriate time and place to present the results of my canvas. For the several groups for which neither colleagues nor recent revisions could supply information, I was obliged to consult several large faunal works and interpret the results in what I hope was a judicious manner. Estimates are intended to be of the number of living species rather than described ones. Although approximately 75 to 100 species of Recent bony fishes are described each year (Zoological Record), we lack comparable information on how many species are placed in synonymy annually. The final results of the present survey are: Agnatha about 50; Chondrich- thyes 515 to 555; Osteichthyes 19,135 to 20,980. The figures given for bony Vor. XXXVIIT] COHEN: RECENT FISHES 343 fishes are two minimums rather than a maximum and a minimum. Most special- ists who volunteered a single figure gave it as a minimum. Many colleagues, however, gave a range. The first figure, 19,135, is the sum of single estimates and the lower figures of ranges; therefore, it represents a bare minimum. The second figure is the sum of single estimates and the upper figures of ranges; therefore, it is a combination of minimum and maximum estimates. I have attempted an ecological analysis of the data for Osteichthyes. The figures used for calculating percentages are averages of high and low estimates. (1) Primary freshwater (Myers, 1949) 6650. 33.1 percent. Approximately 6200 of this group belong to the Ostariophysi. (2) Secondary freshwater (Myers, 1949) 1625. 8.1 percent. Most of the species in this group belong to the families Cichlidae, Cyprinodontidae, and Poeciliidae. Total freshwater 8,275. 41.2 percent. If this astonishingly high percentage is valid it must be a reflection of the degree of isolation possible in the freshwater environment. (3) Diadromous (including Complementary of Myers, 1949) 115. 0.6 per- cent. As the systematics and life histories of tropical shore fishes become better known it seems likely that at least some species will be shifted from category 4 to this group. (4) Marine shore and continental shelf to depths of approximately 200 me- ters—warm water 8000. 39.9 percent. Perciform fishes and their derivatives are the major component of this category. Particularly important are percoid, blen- nioid, and gobioid fishes. Among nonperciforms, eels probably contribute the most species. (5) Marine shore and continental shelf to depths of approximately 200 me- ters—cold water 1130. 5.6 percent. A factor that may contribute to the sub- stantially smaller size of this fauna as compared with that of group 4 is the smaller area occupied. Also, much of the region has had long-term, unstable climatic conditions so that many of the species must be fairly recent in their present habitats. There is no doubt, however, that a high degree of endemism prevails. Important components of this group are Gadidae, Zoarcidae, northern blennioids, and scorpaeniform fishes. Total marine shore and continental shelf to 200 meters 9,130 45.5 percent. (6) Continental slope and deep sea benthic below 200 meters 1280. 6.4 per- cent. Important components of this group are Macruridae, and species of Brotu- lidae, Zoarcidae, Apodes, and Scorpaeniformes. Contrary to the opinion of Greenwood e¢ al. (1966), I do not believe that this group or group 8 contains a great number of unknown species. Fishes of these groups occupy a vast amount of space; however, conditions are relatively so stable and uniform that niches are correspondingly few. (7) Epipelagic (high seas) above 200 meters 255. 1.3 percent. Important 344 CALIFORNIA ACADEMY OF SCIENCES [PRroc. 4TH SER. groups in this category are Scombroidei and Synentognathi. These fishes are mostly mobile, living in an environment that offers few niches. The small number of species is scarcely remarkable. (8) Deep pelagic below 200 meters (including mesopelagic and bathy- pelagic) 1010. 5.0 percent. Clupeiform and myctophiform fishes are the chief constituents of this category. Probably more space is occupied by this group than by any other, yet the number of species is small. The environment is poor in niches and in energy; it is surprising that the fauna is not smaller. SOME CONCLUSIONS The number of species in any one of the 8 categories seems to be chiefly re- lated to the degree of isolation possible. Certainly tropical reefs, great river deltas, and major river drainages have contributed a great variety of habitats and ecological niches which are reflected in the high percentage of species found in freshwater and along tropical shores. The most important regions economically (though not necessarily in terms of biological productivity) are the cooler water shelf areas and the epipelagic, both regions with relatively few species. A final conclusion concerns the freshwater fishes. In view of the high per- centage of fishes found in freshwater and man’s increasing modification of this environment throughout the world, it is vital that research be drastically increased on the basic systematics of freshwater fishes while this is still possible. ACKNOWLEDGMENTS It is a pleasure indeed to acknowledge the cooperation that I have received from my colleagues. Whatever value this paper may have is due to their contri- butions. I thank R. Bailey, P. Banarescu, R. Behnke, F. Berry, J. Bohlke, R. Bolin, M. Bradbury, J. Briggs, W. Burgess, D. Caldwell, B. Collette, E. Cross- man, W. Davis, H. DeWitt, W. Eschmeyer, J. Garrick, R. Gibbs, W. Gosline, D. Greenfield, P. H. Greenwood, M. Grey, R. Haedrich, E. Herald, L. Knapp, E. Lachner, R. Lavenberg, N. B. Marshall, H. McCully, R. McDowall, G. Mead, A. G. K. Menon, G. Miller, G. S. Myers, T. Nalbant, N. Parin, J. Randall, W. Richards, L. Rivas, C. R. Robins, R. Rofen, D. Rosen, R. Rosenblatt, L. Schultz, W. B. Scott, V. Springer, R. Suttkus, A. N. Svetovidov, W. R. Taylor, J. Tyler, I. Trewavas, B. Walker, V. Walters, A. Wheeler, N. Wilimovsky, and L. Woods. LITERATURE CITED BaILey, R. M. 1960. Pisces (zoology), pp. 242-243 im McGraw-Hill Encyclopedia of Science and Tech- nology. Vol. 10. New York, McGraw-Hill. GREENWOOD, P. H. et al 1966. Phyletic studies of teleostean fishes with a provisional classification of living Vou. XXXVIIT] COHEN: RECENT FISHES 345 forms. Bulletin of the American Museum of Natural History, vol. 131, pp. 339-456. Marsuatt, N. B. 1965. The life of fishes. London, Weidenfeld and Nicolson. 402 pp. Norman, J. R. 1963. A history of fishes. 2nd edition by P. H. Greenwood. New York, Hill and Wang. xxxi + 398 pp. Myers, G. S. 1949. Salt-tolerance of fresh-water fish groups in relation to zoogeographical problems. Bijdragen tot de Dierkunde, vol. 28, pp. 315-322. 1958. Trends in the evolution of teleostean fishes. Stanford Ichthyological Bulletin, vol. 7, pp. 27-30. SCHULTZ, L. P. 1965. Fishes and how they live, Chapt. 1 in Wondrous world of fishes. Washington D. C., National Geographic Society. 367 pp. SCHULTZ, L. P. AND E. M. STERN 1948. The ways of fishes. New York, Van Nostrand. xii + 264 pp. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Festschrift for George Sprague Myers Vol. XXXVIII, No. 18, pp. 347-362; 8 figs.; 3 tables. December 31, 1970 DESCRIPTION OF A NEW SUBSPECIES OF RHABDOPHIS AURICULATA IN THE PHILIPPINES, WITH COMMENTS ON THE ZOOGEOGRAPHY OF MINDANAO ISLAND By Alan E. Leviton California Academy of Sciences, San Francisco, California 94118 During recent investigations on the snakes of the Philippine Islands, a study begun more than 15 years ago under the guidance of Professor George S. Myers, it became apparent that the island of Mindanao is inhabited by two clearly dis- tinguishable populations of the diminutive natricine snake, Rhabdophis auricu- lata. That the two populations of this species occupy different but contiguous parts of Mindanao make this discovery the more interesting, inasmuch as it helps shed further light on the paleogeography of the island. The type specimen of Rhabdophis auriculata, a species described by Giinther in 1858, was collected by Hugh Cuming, its locality being given simply as “Philippine Islands.” Miss Alice G. C. Grandison of the British Museum kindly examined Gunther’s type, compared it with the figure published by Boulenger (1893, pl. 17, fig. 1) [fig. 1], prepared a sketch | fig. 2] of the color pattern at the angle of the jaw, and responded that Boulenger’s figure could well have been based on the type specimen. With Miss Grandison’s sketch at hand, and after examining Boulenger’s figure with care, I do not doubt that Giinther’s type is based on a specimen drawn from the eastern Mindanao population. I have seen specimens from Davao Province that closely approximate the Gunther type, and [347] 348 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 1. Figure of Rhabdophis auriculata published by Boulenger (1893). Ficure 2. Sketch of scale and color pattern at angle of jaw on left side of holotype. VoL. XX XVIII) LEVITON: RHABDOPHIS AURICULATA 349 therefore, to insure stability of nomenclature, restrict the type locality of Tropidonotus auriculatus Giinther to Mt. Apo, Davao Province, Mindanao Island. Specimens of Rhabdophis auriculata from Zamboanga del Norte, Zamboanga del Sur, Misamis Occidental, and Bukidnon provinces of western Mindanao | fig. 3| differ from animals from the Davao-Agusan region in color pattern. These animals are referred to a new taxon, named in honor of Professor Myers, who has for many years concerned himself with the zoogeography of the Philip- pines, especially Mindanao Island: Rhabdophis auriculata myersi Leviton, new subspecies. (Figures 4-6.) Tropidonotus auriculatus, BOULENGER, 1893, Cat. Snakes British Mus., vol. 1, p. 261 (part; Mindanao [Pasanaca, Zamboanga City]). Natrix auriculata, Taytor, 1922, Philippine Jour. Sci., vol. 21, p. 294 (Basilan [Port Holland]) ; 1923, Philippine Jour. Sci., vol. 22, p. 542 (Basilan, Mindanao [Zamboanga City]). Diacnosis. Maxillary teeth 27-32, last 3 or 4 enlarged but not separated by diastemata from others; head short, distinct from neck; scales in 17 longi- tudinal rows on anterior third body, reducing to 15 at about level of 80th ventral plate; usually 3 labials bordering orbit; ventrals 143-162; subcaudals 80-93; hemipenes extending to end of 6th subcaudal plate, forked at end of 4th-mid 5th plate; sulcus forked; proximal portion opposite sulcus with longitudinal plicae, portion bordering sulcus spinose; 2 large basal spines present; distal forks uniformly spinose; ventrolateral black stripe extending forward to suture of 8th and 9th labials, continuing as diagonal stripe to corner of eye; lateral light stripe extending uninterrupted (rarely interrupted at angle of jaw by black spot on 8th upper labial) along side of body to anterior temporals. Measurements (of largest male and largest female, in mm.): 4 s-v 342, tail 138; 2 s-v 388, tail 165. Hototyrr. CAS-SU 23391, an adult male, taken near Masawan, approxi- mately 14 km. southeast of Buena Suerte, New Pinan, on west side of Dapitan Peak, Misamis Occidental Province, Mindanao Island, at an altitude of 4700 feet (1433 m.) on 13 April 1959 by Dr. Angel Alcala (figs. 4-5). ParaAtypEs (61). MINDANAO: Misamis OccIDENTAL PROVINCE: West side of Dapitan Peak: Masawan area, approximately 14 km. southeast of Buena Suerte, New Pinan, 26 March—20 April 1959 (CAS-SU 23372-23374, 23392- 23394, 23397 [elev. 4400 ft. (1340 m.)]; 23375 [elev. 4400-4500 ft. (1340- 1370 m.)]; 23376-23380, 23390 [elev. 4500 ft. (1370 m.)]; 23381-23382 [elev. 4700 ft. (1430 m.)]; 23383 [elev. 4300 ft. (1310 m.)]). Approximately 2 km. east of Masawan and 15 km. southeast of Buena Suerte, New Pinan, 5 April 1959 (CAS-SU 23384 [4700 ft. (130 m.)]). Approximately 12 km. east of Masawan, 6 April 1959 (CAS-SU 23387 [elev. 5500 ft. (1670 m.)]). Ap- 350 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. RHABDOPHIS AURICULATA @ R. a. auriculata @ R. a. myersi Ficure 3. Known distribution of Rhabdophis auriculata on Mindanao Island. proximately 2 km. southeast of Masawan and 16 km. southeast of Buena Suerte, New Pinan, 12 April 1959 (CAS-SU 23389 [elev. 4800 ft. (1460 m.)]). Ap- proximately 2—3 km. southeast of Masawan, 19 April 1959 (CAS-SU 23395— 23396 [elev. 5000-5200 ft. (1520-1580 m.)]). Ridge 11-12 km. southeast of Buena Suerte, New Pinan, 22 April 1959 (CAS-SU 23398 |elev. 4000 ft. (1220 m.)|). Bank of Dapitan River, 11-12 km. southeast of Buena Suerte, New Pian, 23 April 1959 (CAS-SU 23399 [elev. 4000 ft. (1220 m.)]). Bank of Dapitan River, approximately 13 km. southeast of Buena Suerte, 25 April 1959 (CAS-SU 23400 [elev. 4200 ft. (1280 m.)]). ZAMBOANGA DEL NoRTE PROv- INCE: West side of Dapitan Peak: Gumay area, approximately 6 km. south- east of Buena Suerte, New Pinan, 28-29 April 1959 and 4-8 May 1959 (CAS-SU 23401, 23403-23404, 23419 [elev. 2500 ft. (760 m.)], 23402 [elev. 2400 ft. (730 m.)]|, 23426-23427 [elev. 2300 ft. (700 m.)]). Dapitan River area, ap- proximately 1 km. southeast of Gumay and 7 km. southeast of Buena Suerte, New Pinan, 30 April-1 May 1959 (CAS-SU 23405-23413, 23415-23416 [elev. 2300 ft. (700 m.)], 23414 [elev. 2400 ft. (730 m.)]). Gumay Creek, approxi- mately 6 km. southeast of Buena Suerte, 2-6 May 1959 (CAS-SU 23417, 23422 VoL. XX XVIII] LEVITON: RHABDOPHIS AURICULATA 351 Ficure 4. Holotype of Rhabdophis auriculata myersi Leviton (dorsal view). [elev. 2300 ft. (700 m.) |). Bank of Dapitan River, approximately 2 km. south- east of Gumay and 8 km. southeast of Buena Suerte, New Pinan, 4 May 1959 (CAS-SU 23418 [elev. 2500 ft. (760 m.)|). Approximately 7 km. southeast of Buena Suerte, New Pifan, 5 May 1959 (CAS-SU 23420-23421 [elev. 2700— 2800 ft. (820-850 m.)]|). Approximately 10 km. southeast of Buena Suerte, New Pifian, 6 May 1959 (CAS-SU 23423-23425 |elev. 3500 ft. (1070 m.)]). Gumay, approximately 5-6 km. southeast of Buena Suerte, New Pinan, 26 March 1959 (CAS-SU 23371 [elev. 2500 ft. (760 m.)]). Gumay Creek, approx- imately 0.5 km. southwest of Gumay and 6 km. southeast of Buena Suerte, New Pinan, 8 May 1959 (CAS-SU 23428 [elev. 2400 ft. (730 m.) |). Mt. Malindang: Masawan, April-May 1956 (CAS-SU 19362-19364 [elev. 3500-4500 ft. (1067— 1372 m.)|). Between Masawan and Gandawan, April 1956 (CAS-SU 19535). Misamis OccIDENTAL AND ZAMBOANGA DEL NORTE PROVINCES BOUNDARY: West side of Dapitan Peak: Approximately 4 km. northwest of Masawan and 10 km. southeast of Buena Suerte, New Pinan, 9-11 April 1959 (CAS-SU 23385-23386 [elev. 3400 ft. (1040 m.)], 23388 [elev. 3500 ft. (1070 m.)]). 352 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 5. Holotype of Rhabdophis auriculata myersi Leviton (ventral view). ADDITIONAL MATERIAL EXAMINED (30). BASILAN: Abung-Abung, 5—25 October 1921 (CAS 60332-60339). Port Holland, 5-25 October 1921 (CAS 60468). BOHOL: Cantaub Sitio, Sierra Bullones, 30 April 1955 (CAS-SU 18895). MINDANAO: Buxipnon Province: Del Monte Plantation, 22 August 1940 (CAS-SU 12396). ZAMBOANGA DEL NoRTE PROVINCE: Catagan, 15 May 1906 (USNM 37390). Katipunan, 30 km. up the Dicayo River (FMNH 68905-68906). Miatan, Katipunan (FMNH 68916-68917). ZAMBOANGA DEL SUR Province: San Ramon, 10 July 1929 (FMNH 14948). Zamboanga City, 23 September—6 October 1920 (CAS 62023-62029). PHILIPPINE ISLANDS: (FMNH 68909, USNM 37415-37418). DESCRIPTION OF HOLOTYPE. Measurements (in mm.): snout-vent 318; tail 122; ventrals 157; subcaudals 82. Anal plate divided; 8 upper labials, 3—5 bordering orbit; nasal shield divided below nostril only; loreal 1; preocular 1; postoculars 3; temporals 2+ 3; dorsal scales reduce 17 (-4[89/91])15; hemipenes extend to end of 6th subcaudal plate, forked at middle of 5th plate; sulcus forked; forked portions spinose; half of proximal portion spinose on other VoLt. XXXVIII] LEVITON: RHABDOPHIS AURICULATA 353 TABLE 1. Comparison of ventral counts for samples of Rhabdophis auriculata. Locality Sex Mean+S.D.+S.E. Range Number Mindanao Zamboanga del Norte 3} 154.0) SEG S= OLS 150-160 29 Q Tgy4h (6) Se 25) ae (04! 150-162 33 Zamboanga del Sur 3 52) se AS) se 10) 149-146 5 2 153.5 153-154 2 Bukidnon fe) 148 — 1 Davao 3} 150 Se Bil Se OG 143-155 28 2 52-7 S= S5 == 0.7 146-160 28 Agusan 3 150.5 149-153 4 Q 150.5 147-154 4 Basilan 3 146.7 143-150 3 9 USP) a Shell SEP ss} 147-157 6 Bohol re 148 — 1 Luzon fe) 145 — 1 1+ Based on a large sample from the Buena Suerta area, on Mt. Dapitan, along the boundary between Misamis Occidental and Zamboanga del Norte provinces. side of sulcus; walls opposite sulcus with longitudinal plicae; 2 very large basal spines present on either side of sulcus. Color pattern as described in diagnosis; black ventrolateral stripe extending uninterrupted to meet lower postocular stripe; black midventral stripe originating on 24th ventral. VaRIATION. There is scarcely any variation in head and body scutellation beyond that already reported for ventral and subcaudal counts. Only two of TABLE 2. Comparison of subcaudal counts for samples of Rhabdophis auriculata. Locality Mindanao Zamboanga del Norte al Zamboanga del Sur Davao Agusan Basilan Bohol Luzon Sex WIG? a= SI Ds S= Slo. Range Number ) 86.32 341 0.6 80-92 25 Q 0.3) as 4s} ae 0)5) 82-93 28 g S574 2-4 Staal 82-89 5 Q 86.5 83-90 2 ry (Oe a= Bo) Se O48 71-86 22 Q SOLS Se 29) SE (Oa) 73-87 26 re 82.3 81-85 4 fe) 81.7 73-87 3 é 83.7 80-86 3 Q 834£19+08 80-86 5 ) 84 = 1 Q 78 — 1 354 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 6. Side view of head of CAS-SU 19362, Rhabdophis auriculata myersi (drawn by Marilyn Kramer). 87 specimens had 9 upper labials on both sides, one had 7 on both sides, one had 7 on one side, 8 on the other, and one had 9 and 8 labials. In most specimens there were 2 anterior temporals; however, in more than 7% of the sample the upper anterior temporal is divided by a vertical suture; 3 posterior temporals are usually present, only eight specimens have 2, and three have 2 on one side, 3 on the other. No variation was observed in numbers of preoculars, postoculars, or loreals. The nasal shield was carefully scrutinized under high magnification of a binocular stereoscopic microscope, and in no instance was I able to detect a suture in the shield above the nostril; a vertical suture was always present, extending from below the nostril to the lower border of the shield. Therefore, the nasal is only partially divided. The color pattern is remarkably stable, too. The dorsal ground color, in preserved specimens, is dark brown to black. There is a prominent middorsal light streak originating just behind the parietals and most distinct on the anterior fourth of the body. A lateral white stripe extends the length of the body along the outer edges of the ventrals to the neck, then angles forward onto the upper anterior temporals, ending just behind the postoculars [figure 6]. This stripe becomes narrower and less distinct posteriorly. It extends uninterrupted onto the side of the head, there being no squarish black spot on the neck behind the angle of the jaw as in the typical form | figure 7] connecting the dorsal ground color with the ventrolateral black stripe. This latter stripe, formed by black spots on the ventrals, extends forward, usually uninterrupted, onto the suture of the 8th and 9th lower labial and thence meets the lower postocular black bar which extends onto the 7th upper labial. Occasionally the stripe is interrupted for one scale width immediately behind the 8th lower labial. Otherwise it ex- tends posteriorly along the sides of the ventrals, broadening and usually coalesc- ing with the black middorsal stripe which originates on the 4th to 23rd ventral, extends posteriorly, becoming broad and joining the lateral black stripes. The Vou. XXXVIIT] LEVITON: RHABDOPHIS AURICULATA ios On UL Nea oe TER a, oe : os Ficure 7. Side view of head of CM 2592, Rhabdophis a. auriculata (drawn by Marilyn Kramer). venter is thus posteriorly almost entirely black except for a series of paramedial white spots, sometimes forming stripes. EcoLocicaL NoTES. The sample from Davao Province is large and is made up of both young and adult specimens. The smallest gravid female measured 299 mm. snout-vent length; young with umbilical scars still evident, though al- most completely closed, measured up to 200 mm., although most animals above 150 mm. in snout-vent length showed no visible signs of the scar. The young, defined here arbitrarily as those with some evidence of an umbilical scar or less than 200 mm. in snout-vent length, were collected mostly during the months of October and November 1946 by members of the Hoogstraal Expedition to Mindanao. The fact that young animals measuring from 104 to 200 mm. were collected during this period suggests that some egg laying takes place almost year round. However, 11 of the 19 young measured between 104 and 125 mm., suggesting further that there is probably a peak reproductive period during late spring or early summer. Gravid females were collected in June and July as well as during the October-November period previously mentioned. At least two speci- mens of R. a. myersi from northern Zamboanga, collected in April, were also gravid. Since most of the animals from the Zamboanga Peninsula were collected during April and early May, and those from Basilan in late October, this would add weight to my argument, based on the survey of the Davao sample, that a peak in egg laying takes place in late spring, possibly late May or June. Six of the sample measured less than 150 mm. snout-vent length, the smallest being a young female, 121 mm. from snout to vent. The rest of the sample divides up as follows: 150-199 mm. snout-vent, 19 specimens; 200-249 mm., 8; 250-299 mm., 13; 300-349 mm., 6; 350-399 mm., 6; and 400 and over, 1. Only amphibian remains were found in the gut, mostly frogs, though in two cases there were tadpoles, and in three, masses of gelatinous frog eggs. In three specimens of R. a. auriculata from Davao Province, the frogs were identified as Oreophryne, a small terrestrial species found in southern Mindanao living in 356 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. moss growing on logs or trees, under bark or in leaf axils (Inger, 1954, p. 447). Two specimens of R. a. myersi from near Buena Suerte had identifiable frog remains in the gut, one referable to Rana magna and the other to Ansonia muel- leri. It is not too surprising that these frogs are eaten by R. auriculata, consid- ering the diminutive size of the snake and of the frog, adults of which measure from 17.2—21.7 mm. (Inger, 1954, p. 447). The snakes for which we have adequate data are found at elevations from 2800 to about 6400 feet (850-1950 m.) in the mountains of Davao Province (Mt. McKinley and Mt. Apo regions). In like manner, specimens of R. a. myersi from the northern part of the Zamboanga Peninsula were taken largely between 2300 feet and 4500 feet (700-1370 m.). The type and those of the paratypic series for which data are available were taken from beneath rocks in dry river beds, while others were collected in brush or among other vegetation, usually near water courses, even if dry at that time of year (late September through November). A Basilan series of R. @. myersi collected by Taylor at Abung- Abung appear to come from near sea level. Also Taylor obtained a series from Zamboanga City, at or near sea level. However, the hills immediately behind Abung-Abung may in fact have been the source of these specimens. The Bunawan, Agusan Province, series collected by Taylor in 1921 was taken in the Agusan River valley, in the vicinity of water, usually from beneath leaves or logs at the edge of a small swamp (Taylor, 1922, p. 90). This locality lies below 500 feet. Since Taylor’s data seem quite reliable, the vertical distribution of R. auriculata is more than 600 feet, this being true for both nominal forms. Rhabdophis auriculata auriculata (Giinther). Tropidonotus auriculatus GUNTHER, 1858, Cat. Col. Snakes British Mus., p. 80 (type locality: Philippine Ids. [restricted here to Mt. Apo, Davao Province, Mindanao]; type in British Museum). Peters, 1861, Monatsb. Akad. wiss. Berlin, 1861, p. 687 (Samar). MULter, 1883, Verh. Naturf. Ges. Basel, vol. 7, p. 286. Borrrcrer, 1886, Ber. Senckenberg Naturf. Ges., p. 108; 1898, Kat. Rept. Samm]. Senckenberg Naturf. Gesell., Schlangen, p. 28 (Leyte). FiscHer, 1885, Jahrb. Hamburg wiss. Anst., vol. 2, p. 80 (Siid-Mindanao). BouULENGER, 1893, Cat. Snakes British Mus., vol. 1, p. 261, pl. 17, fig. 1 (part; “Philip- pines” [type specimen]). Natrix auriculata, GRIFFIN, 1911, Philippine Jour. Sci., sec. D, vol. 6, p. 257 (part; distribu- tion compiled). Taytor, 1922, Snakes Philippine Ids., p. 89, pl. 4, figs. 2-4 (Mindanao [Bunawan, Agusan], Samar). Dracnosis. See Rhabdophis a. myersi except as follows: ventrals 143-160; subcaudals 71-87; ventrolateral black stripe extends forward to side of neck, does not extend onto throat, does not meet dark lower postocular diagonal stripe; light lateral stripe usually interrupted on side of neck behind angle of jaw by squarish black spot joining black ventrolateral stripe to dark dorsal color. Measurements (of largest male and largest female, in mm.): ¢ s-v 352, tail 123: 29s-v 372, tail 140; Vor. XXXVIIT) LEVITON: RHABDOPHIS AURICULATA 3511 Taste 3. Variation in head scutellation in Rhabdophis auriculata auriculata. Preoculars 1-1°(42) 2-2(3) 2-1(2) 1 -2(3) Postoculars 3=—3(41) 3-2(2) 4-30) 4=-4@) 3=-4(1) 2-3(1) 2 —2(3) Upper labials 8—8(46) 7-—7(1) Temporals PEs 2 2612) 2a 3/2 ZO) 2k 2/2 1 34) tai / 2 SCD) ete 2 CL) ada) 22 3) 3 First number is count on right side, second number is for left side. 424 3 indicates anterior + posterior temporals for both sides; 2 +3/2-+ 2 are counts for right and left sides. Rance. Leyte?. Mindanao: Agusan Province (Bunawan); Cotabato Prov- ince (Burungkot) ; Davao Province (Mt. Apo [| Baclayan, Todaya], Mt. McKin- ley). Samar”. Luzon*. MATERIAL EXAMINED (86). LUZON: (CAS 15231). MINDANAO. AcusAN Province: Bunawan (CM 2590-2598). CoTABATO PROVINCE: Burungkot (FMNH 53345 [elev. 1500 ft. (460 m.)]). DAvao PROVINCE: Baganga River (USNM 34734-34735). Mt. Apo (FMNH 15016; 53322- 53323 [elev. 2800 ft. (850 m.)]; 53326 [elev. 5500 ft. (1670 m.)]; USNM 34713 [2 July 1904; elev. 4000 ft. (1220 m.)]; 34719 [3 July 1904; elev. 5000 ft. (1520 m.)]; 34712, 34714-34718, 34720-34727, 34770-34771 [June-July 1904; elev. 6000 ft. (1829 m.)]; 34728-34734). Todaya, Mt. Apo, November 1946 (FMNH 53316-53321, 53324-53325, 53329-53344 [elev. 2800 ft. (850 m.)]). Baclayan, Mt. Apo, 11 November 1946 (FMNH 53327-53328 |[elev. 6500 ft. (1980 m.)]). Mt. McKinley, 8 August-19 September 1946 (FMNH 53299-53314 [elev. 3000-6400 ft. (910-1950 m.)], 53315 [elev. 6300 ft. (1920 m.) ], 53346 [elev. 3400 ft. (1040 m.)]). VARIATION. Variation among the specimens examined is minimal. Ventral and subcaudal counts are summarized in tables 1 and 2. Variations in head scutellation are summarized in table 3. In color pattern in the typical form, variation is limited also. The light mid- dorsal stripe is present in 76 percent of the sample. However, its presence or ab- sence is not correlated with distribution, sex, or age group. The ventrolateral stripe is usually joined to the dorsal color by a vertical or almost vertical black bar on the neck behind the angle of the jaw in 83 percent of the sample. In 10 percent the vertical bar does not contact the dorsal pattern, being separated by about one-half of a scale width. In only four animals—one from Burungkot, Cotabato Province, one from Mt. McKinley, and two from Mt. Apo, Davao Prov- 2 Specimens from Leyte and Samar were reported by Boettger (1898, p. 28) and Peters (1861, p. 687) respectively. The specimens on which these records were based are probably extant, Peters’ at the Berlin Museum, Boettger’s at Senckenberg, and should be examined. I have not done so. The Luzon specimen is in the collection of the California Academy of Sciences, CAS 15231. Formerly it was in the Museo Santo Tomas and sent to the Academy around 1909 by Dr. J. C. Thompson, who said it probably came from Luzon. Indeed, the specimen may have originated on Luzon, but confirmation is needed. The specimen has an interrupted light lateral stripe and therefore is most similar to the eastern Mindanao population. 358 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. ince—is the bar absent. In most animals the ventrolateral stripe stops well short of the commissure of the mouth, being separated from the subpostocular blotch or stripe by at least 1 and usually 2 to 4 scales. Remarks. Rhabdophis auriculata is an unusually small natricine. I hesitate to venture an opinion on its relationships now, inasmuch as I haven’t seen several Bornean forms such as Rhabdophis sarawacensis (Giinther) with which, based on available descriptions, it seems to agree. Although R. auriculata myersi is readily separable from R. a. auriculata on the basis of color pattern alone, an examination of data for ventral and subcaudal counts is revealing (table 1). A comparison of the Zamboanga-Misamis sample with the eastern Mindanao samples does not indicate a statistically significant level of difference in these counts, yet inspection of the means and ranges, especially of subcaudal shields, suggests the populations do indeed differ, though there is a substantial overlap. Perhaps most interesting is the fact that in subcaudal counts both Basilan and Bohol samples agree with the western Mindanao population, and though this is not confirmed by similar close agreement of ventral counts, even here the counts are well within the range of the western Mindanao sample. Of course, in color pattern both Basilan and Bohol! samples are readily referrable to the typical subspecies. A further instructive comparison brings out the fact that the means and ranges of subcaudal counts for the Davao-Cotabato and Bunawan samples and the counts for the one specimen from Luzon are lower than for the typical form. Mindanao has long been of considerable interest to biogeographers concerned with the Philippine fauna and flora. Unquestionably it has served as a principal route for faunal and floral movements to the northern islands. Also it has had a complex post-Miocene history, which is still not completely unraveled. To the extent that we must deal with the matter here, it is sufficient to note that a substantial body of evidence, particularly in the form of coral reefs and numerous terraces, indicate clearly that, during the early and mid-Pleistocene, and possibly during the Pliocene, too, the island was divided into at least five and possibly seven smaller islands (fig. 8). These islands have been detailed by Dickerson (1928, pp. 85-87, fig. 19). Inger (1954, p. 453), in a paleographic map portray- ing the probable land areas during the lower Miocene based on Umbgrove (1938), suggests that even at that date Mindanao was probably not a single island, but rather a series of at least four island masses. The distribution of certain elements of the flora and fauna (see Merrill, 1928, p. 290; Cook, 1928, p. 269; Dickerson, 1928, p. 295) suggests at least a division between the Zamboanga Peninsula and the rest of Mindanao. Further, these authors suggest a greater faunal and floral similarity between the Cotabato-Agusan-Surigao regions than between Cotabato and Zamboanga. This probably indicates that the Pleistocene islands depicted by Dickerson, i.e. Cotabato, Agusan, and Surigao, were connected by emerging VoL. XXXVIIT) LEVITON: RHABDOPHIS AURICULATA 359 Sarangani Bay a4 Ficure 8. “Some probable Pleistocene islands in Mindanao,” from Dickerson (1928, p. 86, fig. 19). subaerial valleys before the Panguil-Illana Isthmus connecting the Zamboanga Peninsula with the Bukidnon-Lanao region emerged. In an earlier paper I pointed out that the populations of Cyclocorus nuchalis from eastern and western Mindanao differed significantly from one another (Leviton, 1967). This was a clue that suggested that in all probability other species of snakes might show similar patterns which could be related to the Plio-Pleistocene paleogeographic history of Mindanao. A careful study of Rhabdophis auriculata on Mindanao would seem to confirm this supposition. Populations of this species from Cotabato and Davao are most similar to the Surigao-Agusan populations I have seen and quite distinct from animals from the Zamboanga Peninsula. Samples of R. auriculata from Bukidnon and Lanao provinces, on the other hand, are indistinguishable from those from the Zambo- anga Peninsula and Basilan Island, raising the possibility that western Mindanao, 360 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser. composed of the ‘“‘Pleistocene islands” of Lano and Zamboanga, were joined to one another and separated from the eastern complex of islands until fairly late in the Pleistocene by a persistent Macajalar-Illana Bay seaway. Indeed, it may have been the persistence of this seaway that prevented any significant eastward movement of the Lanao freshwater fishes. Significantly, only one species of Puntius is known from eastern Mindanao, P. binotatus, a Bornean species. Of course the seaway no longer exists and certainly there has been sufficient time for faunal movements throughout the island. Terrestrial animals can move with greater freedom than those that must wait for stream-capture in order to go from one drainage basin to another, so it is not surprising to find, for example, a paucity of species of cyprinid fishes in eastern Mindanao, though many forms are known from Lake Lanao and several from the Zamboanga Pen- insula. On the other hand, many subspecies of frogs are widely distributed throughout Mindanao, Samar, and Leyte. This distribution is obviously a more recent development, probably a late Pleistocene phenomenon. Indeed, it is be- coming more apparent that several faunal movements and radiations can be identified in Mindanao and are correlated with its post-Oligocene geological history. Quite clearly, a more extensive and careful study of populations of animals and plants on Mindanao, especially the montane forms, should give us consider- able insight into the late geological history of that region. ACKNOWLEDGMENTS The writer wishes to express his appreciation for the loan of specimens to Mr. Neil B. Richmond and Dr. Clarence J. McCoy, Carnegie Museum; Dr. Robert F. Inger and Mr. Hymen Marx, Field Museum of Natural History; and Dr. James A. Peters, U.S. National Museum. The following abbreviations are used: CAS—California Academy of Sciences; CAS-SU—California Academy of Sciences-Stanford University (for specimens formerly housed at Stanford University and registered in the Stanford cata- logues) ; CM—Carnegie Museum; FMNH—Field Museum of Natural History: USN M— United States National Museum. LITERATURE CITED BOETTGER, OSKAR 1898. Katalog der Reptilien Sammlung in Museum der Senckenbergischen Naturfor- schenden Gesellschaft in Frankfurt am Main. 2. Teil. Schlangen. Frankfurt-am- Main, ix + 160 pp. BouLENGER, GEORGE A. 1893. Catalogue of the snakes in the British Museum (Natural History). Volume 1. London, xiii + 448 pp., 28 pls. Cooke, A. H. 1928. Land mollusks of the Philippines. In: Dickerson, Roy (ed.). Distribution of life in the Philippines. Monograph of the Bureau of Science, Manila, Philippines, no. 21, pp. 267-272. VoL. XXXVIIT) LEVITON: RHABDOPHIS AURICULATA 361 Dickerson, Roy 1928. Tertiary and Quarternary paleogeography of the Philippines. Monograph of the Bureau of Science, Manila, Philippines, no. 21, pp. 76-96. GUNTHER, ALBERT 1858. Catalogue of colubrine snakes in the collection of the British Museum. London, xvi + 281 pp. INGER, ROBERT F. 1954. Systematics and zoogeography of Philippine Amphibia. Fieldiana: Zoology, vol. 33, pp. 181-531. Leviton, ALAN E. 1967. Contribution to a review of Philippine snakes, IX. The snakes of the genus Cyclocorus. Philippine Journal of Science, vol. 94, pp. 519-533. MERRILL, Ermer D. 1928. Floral provinces and subprovinces of the Philippines. In: Dickerson, Roy (ed.). Review of the origins of the biologic provinces of the Philippines. Monograph of the Bureau of Science, Manila, Philippines, no. 21, pp. 289-294. PETERS, WILHELM C. 1861. Eine zweite Ubersicht (vergl. Monatsberichte 1859, s. 269) der von Hrn. F. Jagor auf Malacca, Java, Borneo und den Philippinen gesammelten und dem Kgl. zoologischen Museum iiberstandten Schlangen. Monatsberichte der Koniglich preussischen der Akademie der wissenschaften zu Berlin, 1861, pp. 683-691. Umecrove, H. J. F. 1938. Geologic history of the East Indies. Bulletin of the American Association of Petroleum Geologists, vol. 22, pp. 1-70, 1 pl. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Festschrift for George Sprague Myers Vol. XXXVIII, No. 19, pp. 363-382; 7 figs. December 31, 1970 A REINTERPRETATION OF THE TELEOSTEAN FISH ORDER GOBIESOCIFORMES By William A. Gosline University of Hawaii, Honolulu An intensive effort to interpret relationships among the old group “Jugulares” (Linnaeus, 1758, p. 249; Jordan, 1923, p. 228; etc.) led to a consideration of the Callionymidae and Draconettidae. For reasons dealt with below, the conclusion was reached that these two families (I do not agree with Davis, 1966, that they should be combined) are specialized derivatives of the notothenioid section of the perciform suborder Blennioidei (Gosline, 1968). Since, however, the Draconet- tidae and Callionymidae are morphologically well differentiated from the noto- thenioids, it appears best to remove them from the Perciformes entirely. Inves- tigation also suggested that the Gobiesocidae has evolved from the notothenioid section of the perciform suborder Blennioidei and in small part at least over the same route as the draconettids and callionymids. The Callionymidae, Draconet- tidae, and Gobiesocidae are therefore combined here in the order Gobiesociformes. The systematic position of the Callionymidae and Draconettidae has never been the subject of direct investigation. Various views concerning the relation- ships of these two families have, however, been suggested. Boulenger (1904, p. 708) included both the Callionymidae and Gobiesocidae in his Division Jugu- lares, and under his account of the Gobiesocidae stated: ‘The position of the ventral fins suggests, at first glance, affinity with the Callionymidae, and a com- [363] 364 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. parison of the skeletons of these two types has convinced me that they are really related to each other, though highly modified in different directions.”” (My own conclusions are essentially those of Boulenger.) Starks (1905, p. 302) in con- nection with his account of the gobiescoid Caularchus |=Gobiesox| maeandricus wrote: ‘The Callionymidae, however, possess some important characters not possessed by the Gobiesocidae, and these probably more than counterbalance the characters held in common.” Regan in 1913 (pp. 144, 145) placed the Calliony- midae and Draconettidae in the “Division Callionymiformes” of the perciform suborder Percoidei. He stated that the Callionymidae “‘may be related to the Pinguipedidae, but is much more specialized |a suggestion with which I also agree|. The Gobiesocidae differ in many characters of importance.” Referring again to the Callionymidae, Starks (1923, p. 267) said: ‘The osteology shows, however, that this family on account of several rather extraordinary and unique characters should be segregated in a suborder coordinate in value with the Batra- choid fishes.” Regan in 1929 also recognized the Callionymoidei as a perciform suborder. The most recent comment on the systematic position of the Draconet- tidae and Callionymidae is that of Briggs and Berry (1959, p. 125). They sum- marized as follows: ‘Considering the paucity of our knowledge about these two families and their relationships with other percomorph groups, we see no present need for setting them aside in a separate suborder. Their morphology is no more peculiar than that of several other families that are traditionally retained without subordinal recognition within the vast assemblage of the Percomorphi.” The best and most complete account of the anatomy of the Gobiesocidae re- mains that of Guitel (1889). However, Guitel draws no conclusions regarding gobiesocid relationships within the Acanthopterygii. Since the days of Starks (1905)! and Regan (1909) the family has generally been allocated to an order of its own. In his monograph of the family, Briggs (1955, p. 7) wrote: ““The Xeno- pterygii |=Gobiesociformes] seems to be most closely allied to the Haplodoci (batrachoids) but there is also some resemblance to the Callionymoidea. The order may be considered a highly specialized derivative of some still unknown primitive percomorph stock.” McAllister (1968, p. 165) also suggests a gobie- socid-batrachoidid relationship. Apparently on the assumption that such exists Greenwood et al. (1966, pp. 389, 397) have assigned the Gobiesociformes to the superorder Paracanthopterygii, thus separating the group superordinally from the callionymoids. Under the circumstances, it first seems advisable to discuss the possible rela- tionship between the gobiesocoid and batrachoid fishes. Though both groups hold certain characteristics in common, eé.g., the usually scaleless body, the flat- tened head, anterior pelvics, incomplete circumorbital series, etc., it is my pro- 1Starks (1905, p. 292) attributed the creation of ordinal status for the gobiesocids to Gill, but neither Briggs (1955, p. 7) nor I have been able to find where Gill recognized more than subordinal rank for this family. Vor. XXXVIIT] GOSLINE: GOBIESOCIFORMES 36 UL visional view that these similarities are the result of convergence. The batra- choid fishes differ from the Gobiesociformes, i.e., Callionymidae, Draconettidae, and Gobiesocidae, in the following features: In the batrachoid fishes the pelvic fins are fairly close together, small, and with 2 or 3 soft rays that are usually held out at an angle from the abdominal surface; in the Gobiesociformes the pelvic fins are wide apart, well developed (though highly specialized in the Gobiesocidae), and of 4 or 5 soft rays that are normally held flat against the body surface. In the batrachoids the upper hypu- rals have a peculiar intervertebral-like basal articulation with the rest of the caudal skeleton (Regan, 1912, fig. 2B); in the Gobiesociformes there is no such articulation. In the batrachoids the ascending process of the premaxillary has a movable basal articulation with the toothed portion, and a separate articular process of the premaxillary is well developed (Monod, 1960, fig. 49); in the Gobiesociformes the ascending and articular processes of the premaxillary have merged or fused and are firmly attached to the toothed portion. In the batra- choids there is no median ethmoid ossification; in the gobiesociform fishes a median ethmoid ossification is always present. Finally, the batrachoids have a peculiar gas bladder (Sgrensen, 1884); in the Gobiesociformes there is no gas bladder. With regard to the postulated derivation of the Gobiesociformes from the superfamily Notothenioidae (containing the parapercids [=mugiloidids], chei- marrichthyids, trichonotids, nototheniids, etc., see Gosline, 1968) of the perci- form suborder Blennioidei, the gobiesociform fishes have almost all of the diag- nostic notothenioid characteristics despite their high degree of specialization along other lines. Thus in the Gobiesociformes the head is always more or less flattened, some- times greatly so. The circumorbital ring of bones is incomplete. The medial tabulars are apparently lacking. There is a basisphenoid bone in Draconetta but not in the Callionymidae and Gobiesocidae. Flanges from the parasphenoid do not extend up in front of the prootics excluding the prootics from the internal cranial border of the orbit. (When, as in some gobiesocids and callionymids, the parasphenoid does have an upward expansion, this extends up between the mid- dle portion of the orbits, not in the form of a postorbital bar such as occurs for example, in the zoarcioid blennioids.) The pelvic fins are as noted above. (The Gobiesocidae, in which the pelvic fins form the anterior portion of the sucking disc, is the only group known to me in which such a disc extends well forward of the pectoral bases.) The pectoral actinosts are three or four in number. (The 3 broad plate-like actinosts of the Callionymidae are closely duplicated in such notothenioid families as the Nototheniidae.) The dorsal and anal rays are equal in number to the vertebrae between them. The caudal fin is rounded or brush- like, with fewer than 15 branched rays. Additional notothenioid resemblances of the Gobiesociformes are as follows. CALIFORNIA ACADEMY OF SCIENCES [Proc. 47H Serr. VoL. XXXVIIT] GOSLINE: GOBIESOCIFORMES 367 The ventral sucking disc of the Gobiesocidae would seem to be to some extent foreshadowed in the ridges on the flat ventral surface of the notothenioid family Cheimarrichthyidae. The notothenioid genera Prolatilus and Mugiloides are the only members of the suborder Blennioidei known to me with the draconettid supraoccipital crest and with the body musculature extending well forward over the top of the cranium. Seven branchiostegal rays, said to be present in some members of the Gobiesocidae (Briggs, 1955) also occur in a number of noto- thenioids, but rarely elsewhere among the Blennioidei. Finally, the opercular peculiarities of Draconetta (fig. 1A) are largely duplicated in the notothenioid Har pagifer (fig. 1B) and would seem to be foreshadowed in the more generalized notothenioid Parapercis (fig. 1C). The anatomical account of the draconettids, callionymids, and gobiesocids which follows is based primarily on alizarin-stained and dissected specimens from the following lots: Callionymidae: Callionymus flagris, 125 mm. in standard length (U.S. Na- tional Museum no. 71082); C. decoratus, 50 mm. (University of Hawaii no. 2073); and Pogonymus pogognathus, 24 mm., paratype (UH 1626). Draconettidae: Draconetta acanthopoma, 75 mm. (USNM 156956). Gobiesocidae: Gobiesox nigripinnis, 70 mm. (USNM 131163), and Tra- chelochismus pinnulatus, 55 mm., an exchange specimen from New Zealand in the UH collections. The external features of various other species of Callionymidae and Gobie- socidae in the U. S. National Museum were examined during tenure of a Smith- sonian Research Associateship. I wish to express my deep obligation to the members of the Fish Division of that institution for help and facilities during that time and for sending me on loan the specimens of Draconetta acanthopoma listed above. GENERAL FEATURES. The head and body of callionymids, draconettids, and gobiesocids are always scaleless, although Ochiai (1963, p. 66) finds “degenerate scales” partly surrounding the lateral line canal of the callionymid Diplogrammus goramensis. In callionymids the gill opening is a small hole; in Draconetta it is larger, but the gill membranes are broadly attached to the isthmus; and in the Gobiesocidae the gill membranes may be attached to or free from the isthmus (Briggs, 1955). The widely separate pelvic fin bases are entirely in front of the broad pectoral bases, which extend far down the sides; in some callionymids and i Ficure 1. Right suspensorium and opercular bones, external view, of A, Draconetta acanthopoma; B, Harpagifer bispinis; C, Parapercis cephalopunctata; D, Callionymus flagris ; and E, Gobiesox nigripinnis. ec, Ectopterygoid; hy, hyomandibular; io, interopercle; ms, mesopterygoid; mt, metapterygoid; op, opercle; pa, palatine; po, preopercle; and sy, sym- plectic. 368 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 2. Callionymus flagris. Sketch of right side of head to show lateral line canals (dashed lines) that are not enclosed in head bones. gobiesocids there is indeed a membrane extending from the innermost pelvic rays onto the outer surface of the pectoral fin. There is a short spinous dorsal in the Draconettidae and usually in the Callionymidae, but never in the Gobiesocidae. FIN sTRUCTURE. The Gobiesociformes show a transitional series from the usual percoid condition with spines and branched rays to that of the Gobiesocidae where the only spinous element is the flat outer pelvic plate and all the soft rays are simple. The loss of the spinous dorsal in this series has already been noted. As for soft rays, in the callionymid genera Verutia and Synchiropus all of the soft dorsal rays may be branched (Schultz, 1960, p. 399) and in large specimens of Draconetta acanthopoma all of the anal rays are branched, but elsewhere the dorsal and anal rays are mostly or all simple. In Draconetta and the callionymids examined, most of the pectoral and the 5 pelvic rays are branched. I count 8 branched caudal rays in Draconetta acanthopoma, 6 in Callionymus flagris. THE LATERAL LINE SYSTEM. Those portions of the lateralis system enclosed in head bones will be dealt with below. Here, only the peculiar extension of the lateralis system in the Callionymidae will be mentioned. Such extensions occur on both the head (fig. 2) and body. In Callionymus, the system includes such peculiar features as a commissure across the top of the caudal peduncle. On the head of the same genus the canals behind the frontals all lie superficial to the skull bones, extending across the surface of the pterotic and forming a complete supratemporal commissure that is not contained in extrascapulars. Again the preopercular canal, instead of running up within that bone, exits from its lower limb, passes out superficially across the preopercular spine, and then up over the flesh behind the preopercle (fig. 2). None of the peculiarities mentioned are found in either the Draconettidae or the Gobiesocidae, although in the Draconet- tidae there are membranous extensions of the lateralis system. VoL. XXXVIIT] GOSLINE: GOBIESOCIFORMES 369 NASAL APPARATUS. The nasal apparatus differs considerably among the gobiesociform fishes examined. It is most percoid-like in Gobiesox nigripinnis which has 2 nostrils, the anterior with a fringed flap and the posterior in a raised collar; these 2 nostrils lead into a nasal cavity, bordered mesially above by the nasal bone; the cavity contains a roundish nasal rosette. The nasal apparatus of Callionymus flagris is about the same except that there is only 1 nostril on each side. In Draconetta acanthopoma there are 2 tubular nostrils but no nasal bone; the nostrils lead into the two ends of a flattened, hollow, fleshy pad which seems to contain no specialized olfactory folds or lobes. THE CIRCUMORBITAL BONES. The circumorbital series in the Gobiesociformes is always reduced to the lacrimal bone. Behind the eye in Callionymus and Dra- conetta a membrane-enclosed canal exits from the main lateralis canal and extends downward. In Draconetta this canal is short, ending behind the eye; in Callio- nymus flagris it extends forward below the eye towards the base of the lacrimal bone (fig. 2) but fails to connect with the lacrimal-enclosed canal. Jaw stRucTuRE. The peculiarity of the upper jaw protrusion of Callionymus has been described by van Dobben (1935, pp. 47, 48) and by Kayser (1962). In most percoids, the maxillary heads twist on their axes extruding the premaxillary articular processes before them like a squeezed cake of soap (van Dobben, 1935, pp. 10-13). In Callionymus the maxillary heads and associated cartilages and ligaments of the two sides form a ring around the long ascending processes of the premaxillaries. The ascending processes of the premaxillaries are free to move in and out within this ring. Upper jaw protrusion is entirely produced by the lower- ing of the mandible with the associated downward movement of the lateral end of the maxillary. Anatomically Callionymus is peculiar in having no articular proc- esses on the premaxillaries lateral to their ascending processes. The gobiesocids also have premaxillaries without articular processes (Guitel, 1889, pl. 25, fig. 16, and Briggs, 1955, figs. 74-81). In Draconetta there are long, narrow, articular processes that are all but fused to the ascending processes. So far as I determine from preserved specimens, Draconetta and most gobiesocids use the same peculiar method of upper jaw protrusion that Callionymus does. In at least the gobiesocid genus Tomicodon, however, the upper jaw does not appear to be protrusile. GILL COVERS AND SUSPENSORIA. With the extreme flattening of the head region that has taken place in the Callionymidae, Draconettidae, and Gobiesoci- dae, the operculum becomes squashed, so to speak, into a horizontally elongate structure. In at least some members of all three families, backwardly projecting spines are developed, but they are formed in different ways. As already noted, the opercular apparatus of Draconetta (fig. 1A), with spi- nous opercles and subopercles, is essentially similar to that of the notothenioid Harpagifer (fig. 1B), although in Harpagifer an additional support for the oper- cle has been added by extending a vertical strut up to an abutment against the 370 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. cranium. How the spinous arrangement in Harpagifer and Draconetta might have originated is suggested by the basal notothenioid Parapercis (fig. 1C). In Parapercis the opercle ends in the not unusual point; the subopercle has two structurally different sections, an upper, flap-like ossified membrane and a lower rigid plate ending posteriorly in a few serrations. Disappearance of the upper membranous portion of the subopercle and development of the lower would provide essentially the configuration of gill cover spines found in Harpagifer and Draco- netta. Now, if instead of developing the lower portion of the subopercle of Para- percis, the upper flap-like portion were enlarged, the lower eliminated, and the preopercle developed backward as a strong spine, the configuration found in Cal- lionymus (fig. 1D) would result. To arrive at the gobiesocid-type opercle (fig. 1E), one could hypothesize a form of Callionymus in which the subopercle loses its association with the inter- opercle and swings back onto the end of the opercle where it may form a spine in gobiesocids. The changes in opercular structure just described are reflected in the inter- opercle. This bone, fairly long in Parapercis and longer in Draconetta, is pulled out into a long weakly ossified tendon in Callionymus. In Gobiesox the inter- opercle is wholly concealed by the preopercle and does not reach the subopercle at all but terminates in an abutment against the rear of the hyoid apparatus, as in the Blenniidae; the interopercle is, however, better developed in the more primitive Tvachelochismus, where it nearly reaches the subopercle. A last minor point about the gill cover structure of the Gobiesociformes should perhaps be made. In all three families those edges that are not rigid tend to have long, flexible bony fimbriae. The “squashing” of the opercle would also seem to have had an effect on the suspensoria of callionymids, draconettids, and gobiesocids. The preopercles of the callionymids (fig. 1D) and gobiesocids (fig. 1E) have been extruded backward, so to speak, and the hyomandibular, preopercle, and quadrate have come to form the three points of a triangle. Draconetta (fig. 1A), however, has retained the usual configuration with the hyomandibular, preopercle, and quadrate all more or less in line. There is however no separate metapterygoid in the Draconettidae, Callionymidae, or Gobiesocidae. The connection between the palatine and the posterior portion of the suspen- sorium has become rather tenuous. In Draconetta (fig. 1A) the palatine is at- tached to the quadrate by a long narrow strut composed of the ectopterygoid and mesopterygoid. In Callionymus (fig. 1D) these last two bones seem to have fused, but the strut is still present. In Gobiesox (fig. 1E) the palatine is only loosely connected with the rest of the suspensorium, the mesopterygoid is gone, and the minute ectopterygoid is only ligamentously attached to the palatine. THE HYOID APPARATUS AND GILL ARCHES. The hyoid apparatus is close to, Vot. XXXVIIT] GOSLINE: GOBIESOCIFORMES 371 Ficure 3. Hyoid and gill arches (1-5) in Gobiesox nigripinnis: A, hyoid arch and lower portions of the gill arches of the right side, from above; and B, the upper portions of the gill arches of the left side, from above. cb, Ceratobranchial; ch, ceratohyal; eb, epibranchial; eh, epihyal; gh, glossohyal; hb, hypobranchial; hh, hypohyal; ph, upper pharyngeal tooth plate. and firmly connected by the anterior basibranchial with, the gill arches in Callio- nymus; in Draconetta and Gobiesox the hyoid arch is well separated from and unconnected with the other gill arches. In Callionymus and Draconetta, a well developed glossohyal extends forward from the hypohyals; in Gobiesox (fig. 3) the glossohyal is a small sliver of bone completely contained in the interspace between the hypohyals of the two sides. In Draconetta and Callionymus there are 6 branchiostegal rays on each side, in the gobiesocids 5—7 (Briggs, 1955, p. 9). In Draconetta, Callionymus, and Gobiesox there are 2 anterior branchiostegals attached to the inner surface of the hyoid arch; the other 4 close to its lower rim. In Draconetta and Callionymus the first 2 are short; in Godiesox the first 3. In Draconetta 4 of the 6 branchiostegals are crowded back on the epihyal, in Callio- nymus 3, and in Gobiesox only 1 branchiostegal articulates with the epihyal. Among gobiesociform families the first spicular pharyngobranchial seems to have completely disappeared and there are never more than 2 sets of pharyngeal teeth on either side above. In Draconetta and Callionymus epihyal 2 extends up to the relatively small and narrow anterior tooth patch, while epihyals 3 and 4 articulate with the broader, posterior pharyngeal tooth patch”; in these two genera epihyals 3 and 4 are closely but movably attached to one another. In Gobiesox 2 Starks (1905, p. 302) stated that Callionymus had ‘‘three superior pharyngeals on each side’’ but in 1923 (p. 269) he describes 2 upper pharyngeals of the same shape as noted here. 372 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. Ficure 4. Cranium plus upper portion of pectoral girdle, right side, from above, of A, Gobiesox nigripinnis; B, Draconetta acanthopoma (only the upper surface of the rostral region is shown); and C, Callionymus flagris. Lateral line canals passing through cranial bones indicated by dashed lines. ca, Cartilage; cl, cleithrum; cv, cavity into which the ascend- ing processes of the premaxillaries extend; et, lateral extrascapular; ex, exoccipital; fr, frontal; le, lateral ethmoid; me, mesethmoid; pa, parietal; po, posttemporal; pt, pterotic; su, supra- cleithrum ; so, supraoccipital; st, sphenotic; su, supraoccipital crest; and vo, vomer. (fig. 3) there are no chondrified or ossified basibranchials, and the gill arches are not interconnected below. Above, there is only 1 small pharyngeal tooth plate on each side; epihyals 2, 3 and 4 articulate with it, and epihyals 3 and 4 are rigidly united to one another. SKULL. In Gobiesox (which lives under rocks in the tidal zone) the head is broad with small eyes in strong laterally placed bony sockets. In Callionymus and Draconetta the eyes are close together on the top of the head. These differ- ences are strongly reflected in the crania. Lateral line and associated skull bones. In the Gobiesocidae the forward por- tion of the supraorbital canal on each side commences near the snout rim and passes back through the paired nasal and frontal bones. Between the wide-set eyes there is a complete, bone-enclosed frontal commissure (fig. 4A). In the Dra- conettidae and Callionymidae the narrow interorbital region has doubtless caused the fusion of the 2 supraorbital canals into a single median canal between the eyes (fig. 4B, C). Furthermore, in Draconetta the frontals themselves have fused into a single median bone. However, in the two species of Callionymus examined the frontals appear to be only partially fused, and in the callionymid Pogonymus, VoL. XXXVIIT] GOSLINE: GOBIESOCIFORMES 3S which has a somewhat broader interorbital area, I believe I can see a suture com- pletely dividing the frontals. Anteriorly, the supraorbital canals of callionymids begin in the separate nasals as usual, but in Draconetta acanthopoma there are no nasal bones and the anterior median pore of the frontal canal is the anterior- most point in the supraorbital system. (In Draconetta oregona Davis, 1966, fig. 2, shows the supraorbital canals as separating ahead of the eyes and extending forward on each side to just behind the nostril. Perhaps these anterior extensions of the supraorbital system in D. oregona are represented by fine ridges of flesh running over the same areas in D. acanthopoma.) Behind the frontals the lateral line canals of Callionymus lie superficial to the skull bones, as previously noted (fig. 1). In Gobiesox and Draconetta the tem- poral canals pass backward from the frontals through what appears to be the sphenotic and pterotic (fig. 4A, B). Passage of the lateral line through the pterotic is normal in fishes, but a canal in the sphenotic is not. Possibly the “sphenotic”’ canal of Gobiesox and Draconetta extends through a dermosphenotic which has become fused to the sphenotic. In Draconetta the lateral line canal passes back from the pterotic into a lateral tabular, where it gives off the membranous, incom- plete supratemporal commissure, and then into the posttemporal, where it ends. In Gobiesox the lateral line canal ends in the pterotic; there is no tabular bone or posttemporal commissure. Ethmoid region of the skull. The peculiarities of the ethmoid region of the cranium of Callionymus (Starks, 1923, pp. 267-268) and of Draconetta can, I think, have developed through a pinching together of the broader, more normal ethmoid area of the Gobiesocidae. In the Gobiesocidae the ethmoid overlaps the vomer in the usual percoid fashion but lies behind the level of the lateral ethmoids (Guitel, 1889, pl. 25, fig. 1). In the narrower-snouted draconettids and callio- nymids the mesethmoid is completely separated from the vomer by cartilage and by the medial bases of the two lateral ethmoids which meet (fig. 4C) or nearly meet on the midline. In Draconetta (fig. 5) the mesethmoid is above and behind the lateral ethmoid bases, but in the callionymids it is entirely behind them. In both families the mesethmoid forms part of the orbital border. In the callionymid Pogonymus the ascending processes of the premaxillaries extend up and back over the rostral surface as usual; here the mesethmoid does not extend down into the interorbital space. But in Callionymus the ascending processes are more horizon- tal and their tips extend back into a medial rostral cavity; here the mesethmoid has been pushed down and back, as it were, into the infraorbital space (Starks, 1923, pl. 4, fig. 5). The same sort of thing seems to have happened in the chaeto- dontid percoids, as Starks (1926, p. 301, footnote 35) has noted. Upper surface of rear of skull. Major differences on the upper surface of the skull posteriorly have to do with the extent to which it is covered by the body musculature. In Callionymus the rear face of the skull drops away abruptly, and no musculature at all extends forward over its upper surface. The supraoccipital 374 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. WA TA MILLI PAD DY A Ficure 5. Draconetta acanthopoma. Lateral view of forward end of cranium with only the base of the lateral ethmoid indicated. Cartilage stippled. fr, Frontal; io, infraorbital fenestra; le, lateral ethmoid; me, mesethmoid; ra, parasphenoid; and vo, vomer. extends back from the skull as a flat superficial cap (fig. 4C) the bottom of which forms a surface for muscular attachment. In Gobiesox the rear face of the skull slopes more obliquely and two large lateral lobes of musculature extend forward nearly to the rear borders of the eyes. The musculature does not extend forward over the central portion of the skull and there is no median crest. The flat supra- occipital bone (fig. 4A) in Godiesox is pinched off into two parts by the overlap- ing parietals, but this is not true of at least certain other gobiesocid genera (see Guitel, 1889, pl. 25, fig. 1, and Starks, 1905, p. 283—-Starks’s Caularchus equals Gobiesox and his Gobiesox equals Sicyases according to Briggs, 1955). In Dra- conetta (fig. 4B) the musculature extends forward along either side of the midline to just behind the eye, and a median crest extends forward on the supraoccipital and even a short way on the fused frontals. Sphenoid region of the skull. In Draconetta a pleurosphenoid and small basisphenoid bone are present; the two bones are, however, widely separated, the basisphenoid ending posteriorly in the membrane lining the orbits posteriorly. In neither Callionymus nor Gobiesox are pleurosphenoids or basisphenoids present. As is true of all notothenioids, there is little upward extension of the para- sphenoid into a postorbital bar, and the prootic borders the orbit in all the Gobiesociformes. In Gobiesox, however, the parasphenoid is considerably ex- panded anteriorly, forming a broad shelf below and between the orbits; this ex- pansion is greater than that of the parasphenoid just behind the orbits. Otic and occipital regions of the skull. In none of the Gobiesociformes examined is there an expanded auditory bulla. In Draconetta there is a triangular VoL. XXXVIIT] GOSLINE: GOBIESOCIFORMES 375 Ficure 6. Draconetta acanthopoma. Pectoral girdle of right side, except postcleithra. Lateral line canal indicated by dashed lines. ac, Actinost; cl, cleithrum; co, coracoid; et, lateral extrascapular; li, ligament to intercalar; sc, scapula; sl, supracleithrum; and tm, post- temporal. intercalar on the lower surface of the cranium which serves for the attachment of the ligament from the short lower wing of the posttemporal. In Gobiesox and Callionymus there is neither an intercalar nor a lower wing to the posttemporal. The exoccipital condyles in Draconetta, Callionymus, and Gobiesox are widely separated from one another and indeed are practically or quite lateral to the basioccipital condyle. As Starks (1905, p. 293, footnote 1) has noted, this con- figuration of the occipital condyles is one frequently associated with a depressed body form. PECTORAL GIRDLE. In gobiesocids the supracleithrum and posttemporal bones are both present. The cleithrum and primary pectoral girdle extend up the sides of the body. From an articulation on the top of the cleithrum, the supracleithrum extends horizontally forward, and from the front of the supracleithrum the post- temporal extends horizontally inward to an articulation with the skull. The axes of the cleithrum, supracleithrum, and posttemporal thus lie primarily in three different planes (Guitel, 1889, pl. 24, fig. 3). In Draconetta the supracleithrum and posttemporal are present (fig. 6) but the supracleithrum and cleithrum have the same axes. Among callionymids Briggs and Berry (1959) and Ochiai (1963) state that a supracleithrum and supratemporal are both present, though the 376 CALIFORNIA ACADEMY OF SCIENCES [Proc, 47H SER. latter author shows only one of these two bones in his figures. Starks (1923, p. 268) says that the supracleithrum is absent in Callionymidae and I can find none in Callionymus flagris, C. decoratus, or Pogonymus. Judging from the position of the supracleithrum in Draconetta, it would seem to have become fused with the cleithrum in the callionymids investigated by me. Perhaps its loss as a separate element is variable in callionymids. In Draconetta there are 4 actinosts. The lower 3 are columnar, but the upper- most tapers to a basal point and has its entire upper edge contiguous with the scapula (fig. 6). In Callionymus, as in the Nototheniidae, there are only 3 acti- nosts, the uppermost of Draconetta having doubtless become fused with the scapula. In Gobiesox there are not only 4, more or less hourglass shaped, actinosts, but the scapula projects around the top of the uppermost in such a way as to resemble a fifth, as was noted by Starks (1930, p. 220; see also Guitel, 1889, pl. 24, fig. 10). It is very probably a scapular projection of similar sort that provides the uppermost fifth “‘actinost” of the batrachoid fishes. A further peculiarity of pectoral girdle structure unique among the Gobiesoc- idae is the specialization of the two postcleithra (see Starks, 1905; Guitel, 1889, pl. 24, fig. 3). Both of the postcleithra on either side are plate-like. The upper is vertically alined and has numerous fimbriae extending from its posterior sur- face; it appears to be only ligamentously attached to the main pectoral girdle. The lower extends inward from the side of the abdomen and, with its counterpart from the opposite side, supports the rear rim of the adhesive disc. I do not know of a similar specialization elsewhere in fishes, the postcleithra of Cheimarrichthys, for example, being quite normal. In Draconetta there is only a single, long, scimi- tar-like postcleithrum with the usual ligamentous attachment to the top of the cleithrum. Callionymus has an even longer, thinner postcleithral strut, but it is made up of 2 pieces closely bound together where they overlap. PELVIC GIRDLE. The pelvic girdle of the Gobiesociformes is short and broad, as in many notothenioids. The only peculiarity that I can find is in the flat, spatulate pelvic spine of Gobiesox, already mentioned. AXIAL AND CAUDAL SKELETONS. In Draconetta there are 7 abdominal and 16 caudal vertebrae, including the urostylar centrum. In the Callionymidae, so far as is known, there are 7 + 14 vertebrae. Briggs (1955, p. 9) gives the vertebral counts of Gobiesocidae as ranging from 25-54; in Gobiesox the count given by Starks (1905, p. 300) is 13 + 19. In all the Gobiesociformes the ribs start on the second vertebra. In Draco- ) netta and Callionymus there is only 1 pair of ribs per vertebra. These, in Draco- netta, extend out and up away from the abdominal cavity, which suggests that they are epipleurals. In Gobiesox the same set of ribs occurs, but from the third vertebra on there is another set of ribs extending lateroventrally from the lower surface of the main ribs about half way out along their length (Runyon, 1961, p. VoL. XXXVIIT]J GOSLINE: GOBIESOCIFORMES ay ~r 136 and fig. 27). These supplementary lower ribs are, despite their configuration, probably pleural ribs (but see Starks, 1905, p. 301). In the flattened nototheni- oid Bembrops there is only a single set of ribs, but these commence on the first, not the second vertebra (for the problem of whether a single set of ribs in acan- thopterans is pleural or epipleural, see Starks, 1923, p. 290). In the gobiesociform fishes examined there are no predorsal bones, and the first interneural extends down behind the second neural arch. In Draconetta the neural arch to interneural relationship is normal, but in Callionymus and Pogo- nymus the third and following vertebrae have V-shaped neural processes that extend out laterodorsally on either side of the interneurals. In Draconetta there are 2 separate hypurals in the caudal skeleton, the lower autogenous, and the upper fused to the urostylar centrum. In Callionymus and Gobiesox these 2 hypurals are fused into a single unit basally. In Draconetta and Callionymus there are 2 epurals, in Gobiesox none. Unlike Gobiesox, the penulti- mate vertebra of Callionymus and Draconetta has expanded, plate-like neural and hemal arches which are fused to the centrum. So far as the fishes examined are concerned, the characters described above may be grouped as follows. It should be noted, however, that the wider the spec- trum of variation within the group the less any definition based on one or a few species, such as those given below, can be expected to hold. Gobiesociformes.—Head and body scaleless. Circumorbital bones represented only by the lacrimal. Premaxillary with its articular process absent or merged with the ascending process (in Draconetta). Opercular apparatus with 1 or 2 backwardly projecting spines (except some Gobiesocidae). Metapterygoid absent. Ribs commencing on second vertebra. Gobiesocoidei—An abdominal adhesive disc. No spinous dorsal fin. None of the fin rays branched. Outer pelvic ray flattened and spatulate, followed by 4 segmented rays. Palatine separated by membrane from the ectopterygoid. No basibranchials. A single upper pharyngeal tooth plate on each side. Frontals separate. Mesethmoid not forming part of the orbital boundaries. Parasphenoid expanded below and between the orbits. Postcleithra expanded, platelike, the lower supporting the rear border of the adhesive disc. More than 10 abdominal vertebrae, more than 24 in all. Two sets of ribs from the third vertebra. Penultimate vertebra with its neural and hemal arches not expanded. No epurals. Gobiesocidae—Lateral line system limited to the head. A nasal bone on each side of head. Two nostrils on either side, which lead into a nasal sac contain- ing a well-developed olfactory rosette. A single spine, if any, on the opercular apparatus, formed by the subopercle. Gill openings not restricted to a small hole above or behind the opercle. Mesopterygoid absent. Supratemporal commissure lacking. No median supraoccipital or frontal crest. Pleurosphe- noid, basisphenoid, and intercalar absent. Posttemporal present. Four acti- 378 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. nosts. Two postcleithra. Neural arches normal. Hypurals fused into a single plate. Callionymoidei—No abdominal adhesive disc. A spinous dorsal fin present, except Draculo. At least 1 soft ray in each fin branched or divided to the base. Outer pelvic ray spinous, followed by 5 soft rays. Palatine firmly at- tached to the ectopterygoid. Basibranchials present. Two upper pharyngeal tooth plates on each side. Frontals fused or mostly so. Mesethmoid forming part of the orbital boundaries. Parasphenoid forming a narrow strut below and between the orbits. Postcleithral strut narrow. Seven abdominal verte- brae, fewer than 24 in all. A single set of ribs. Penultimate vertebra with its neural and hemal arches expanded and plate-like. Two epurals. Draconettidae.—Lateral line system limited to head. No nasal bone. Two nos- trils on each side of head; no nasal rosette. Two spines on the opercular apparatus, one on the opercle and one on the subopercle. Gill openings not restricted to a small hole above or behind the opercle. Mesopterygoid present. Supratemporal commissure incomplete. A low supraoccipital crest extending forward onto the rear of the frontals. Pleurosphenoid, basisphenoid, and intercalar present. Posttemporal present. Four actinosts. One postcleithrum. Neural arches normal. Two separate hypurals. Callionymidae.—Lateral line continued on body. A nasal bone on each side. One nostril leading into a nasal sac with a well developed olfactory rosette. Spine on the opercular apparatus single, formed by the preopercle. Gill openings restricted to a small hole above or behind the opercle. Mesopterygoid absent. Supratemporal commissure complete. No median crest on supraoccipital or frontals. Pleurosphenoid, basisphenoid, and intercalar absent. Posttemporal absent. Three actinosts. Two postcleithra. Neural arches of third and suc- ceeding vertebrae with V-shaped flanges. Hypurals fused into a single plate. Of the developments which characterize the Gobiesociformes as a whole, some are of a type that have repeatedly occurred in higher acanthopterans, e.g., the “simplification” of skull and fin ray structure. Perhaps the absence of scales and the loss of the circumorbital bones behind the lacrimal should be placed in the same category. In my opinion the definitive peculiarities held in common by the various members of the Gobiesociformes are those of the upper jaw, gill cover, and rib configuration. That the various members of the Gobiesociformes have diverged widely is obvious. In the first place, there is a most remarkable difference in habitat be- tween the callionymoids, which are mostly quiet water bottom fishes, and many gobiesocids. At least some of the latter, including the close relatives of one dis- sected here, live among the boulders of wave-washed rocky beaches. The way in which the gobiesocids have evolved from a proto-gobiesociform ancestor is suggested by the notothenioid Cheimarrichthys, which has the same VoL. XXXVIIT] GOSLINE: GOBIESOCIFORMES 379 Gobiesocidae Callionymidae Draconettidae GOBIESO|CIFORMES PERCIIFORMES NOTHENIOIDAE Ficure 7. Suggested gobiesociform relationships. sort of broad, flat head, small, wide-set eyes, and incipient adhesive organ on the abdomen as Gobiesox. However, the gobiesocids, in addition to having the pre- maxillary, opercular, and rib structure, etc., of all the gobiesociformes, which Cheimarrichthys does not have, have incorporated the postcleithra into the ad- hesive disc in a unique way. In short, the gobiesocids are much more highly specialized fishes than Cheimarrichthys. The callionymoids would seem to have diverged from their proto-gobiesoci- form ancestors in two principal respects. One is that the high-set eyes have left little room for the interorbital portion of the cranium. The frontals have not only fused, but their anterior portion appears to have been pinched off and replaced in part by the mesethmoid from the preorbital region. Second, there has been a reduction in the number of vertebrae. Between the draconettids and callionymids, the quite different opercular specializations of the two groups preclude the possibility of the one group having evolved directly from the other. In general, however, the draconettids have re- mained at a lower stage of specialization than the callionymids as indicated by the much lower degree of fusion in the draconettid skeleton. In my opinion then, the relationships between the three gobiesociform groups may be diagrammed as in fig. 7. REFERENCES BouLENceEr, G. A. 1904. Fishes (systematic account of the Teleostei). In “Cambridge Natural History,” vol. 7, pp. 541-727, figs. 325-440. 380 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser. Briccs, J. C., and F. H. Berry. 1955. A monograph of the clingfishes (Order Xenopterygii). Stanford Ichthyological Bulletin, vol. 6, pp. 1-224, 114 figs. Briccs, J. C., and F. H. Berry. 1959. The Draconettidae—A review of the family with the description of a new species. Copeia, 1959, pp. 123-133, 7 figs. Davis, W. P. 1966. A review of the dragonets (Pisces: Callionymidae) of the western North Atlantic. Bulletin of Marine Science, vol. 16, pp. 834-862, 10 figs. GOSLINE, W. A. 1968. The suborders of perciform fishes. Proceedings of the United States National Mu- seum, vol. 124, pp. 1-78. GREENWOOD, P. H., D. E. Rosen, S. H. Wetrzman, and G. S. MYEnrs. 1966. Phyletic studies of teleostean fishes, with a provisional classification of living forms. Bulletin of the American Museum of Natural History, vol. 131, pp. 339-456, pls. 21-23, text figs. 19. GUITEL, F. 1889. Recherches sur les lepadogasters. Archives de Zoologie Expérimentale et Génerale, ser. 2, vol. 6, pp. 423-627, pls. 24-37. Jorpan, D. S. 1923. A classification of fishes, including families and genera as far as known. Stanford University Publications, University Series, Biological Sciences, vol. 3, pp. 79-243. Kayser, H. 1962. Vergleichende Untersuchung tiber Vorstrecksmechanismen der Oberkiefer bei Fis- chen. Der Bau und die Funktion des Kiefer- und Kiemenapparates von Kno- chenfischen der Gattungen Ammodytes und Callionymus. Zoologische Beitrage, new ser., vol. 7, pp. 321-445. LINNAEUS, C. 1758. Systema Naturae, Ed. X. Vol. 1. Holmiae: 824 pp. McAttister, D. E. 1968. Evolution of branchiostegals and classification of teleostome fishes. National Mu- seum of Canada, Bulletin 221, xiv-+239 pp., 21 pls. Monon, T. 1960. A propos du pseudobrachium des Antennarius (Pisces, Lophiiformes). Bulletin de VInstitut Francais d’Afrique Noire, vol. 22, ser. A., pp. 620-698, 83 figs. Ocul, A. 1963. Two dragonet fishes obtained from the tidal zone of the Amami Islands. Bulletin of the Misaki Biological Institute, no. 4, pp. 63—74, 8 figs. REGANG Cale 1912. The classification of the teleostean fishes of the order Pediculati. Annals and Magazine of Natural History, ser. 8, vol. 9, pp. 278-289, 6 figs. 1913. The classification of the percoid fishes. Annals and Magazine of Natural History, ser. 8, vol. 12, pp. 111-145. 1929. Fishes. In “Encyclopaedia Brittanica,’ 14th ed., pp. 305-329. Runyon, S. 1961. Early development of the clingfish, Gobiesox strumosus Cope. Chesapeake Science, vol. 2, pp. 113-141, 33 figs. S@RENSEN, W. E. 1884. Om lydorganer hos fiske. Inaugural dissertation, Kj@benhavn: viii+-245 pp., 4 pls. VoL. XXXVIIT] GOSLINE: GOBIESOCIFORMES 381 Starks, E. C. 1905. The osteology of Caularchus maeandricus (Girard). Biological Bulletin, vol. 9, pp. 292-303, 2 figs. 1923. The osteology and relationships of the uranoscopoid fishes. Stanford University Publications, University Series, Biological Sciences, vol. 3, no. 3, pp. 259-290, 5 pls. 1926. Bones of the ethmoid region of the fish skull. Stanford University Publications, University Series, Biological Sciences, vol. 4, no. 3, pp. 139-338, 58 figs. 1930. The primary shoulder girdle of fishes. Stanford University Publications, Univer- sity Series, Biological Sciences, vol. 6, no. 2, pp. 149-239, 38 figs. VAN DosBen, W. H. 1937. Uber den Kiefermechanismus der Knochenfische. Archives Neerlandaises de Zoo- logie, vol. 2, pp. 1-72, 50 figs. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Festschrift for George Sprague Myers Vol. XX XVIII, No. 20, pp. 383-390 December 31, 1970 SCALE-EATING AMERICAN CHARACOID FISHES, WITH SPECIAL REFERENCE TO PROBOLODUS HETEROSTOMUS Ey Tyson R. Roberts Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138 INTRODUCTION Specialized scale-eaters have been discovered in three groups of American characoids. Kner (1860, p. 34) reported the stomach of a large specimen of Cato- prion full of scales and referred to them as ‘‘Raubfische.” Ladiges, observing this peculiar serrasalmid in an aquarium, saw one remove a row of scales from a specimen of Metynnis with one swipe of its teeth (reported by Géry, 1964, p. 460). Breder (1927, p. 127) reported substantial amounts of large scales in stomach contents of representatives of Roeboides occidentalis from eastern Pan- ama and identified some of the scales as coming from Ctenolucius, a pike-like characoid considerably larger than the specimens of Roeboides. Géry (1964, pp. 459-460) reported scale-eating in Exodon, Roeboides, and Roeboexodon, of the characid subfamily Characinae. In this paper the activity is verified for Cato- prion, Exodon, Roeboexodon, and two additional species of Roeboides and is re- ported for the first time in Probolodus heterostomus Eigenmann,' a member of the characid subfamily Tetragonopterinae. Although Géry supposed that scale-eating occurred in Catoprion, Exodon, Roeboexodon, and Roeboides only occasionally, in these genera and in Probolodus scales are definitely a major item in stomach contents, and eating scales is prob- 1 Myers (1942, p. 91), in recording specimens from the western end of the coastal plain of Rio, com- mented on their almost unbelievably strange dentition. The species also occurs in the rios Doce, Paraiba (formerly spelt Parahyba), and Ribeira. [383] 384 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. ably a significant factor in the mode of life of these fishes. Serrasalmus elongatus includes some scales in its diet but is primarily a fin-eater. Matthes (1961) re- ported that the African characoids Phago, Belonophago, and Eugnathichthys (family Ichthyboridae) are fin-eaters. There are reports in the literature on aquarium fishes that the eating of fins and scales occurs in Phago. Matthes (1964, pp. 65-66) reported scales in stomach contents of specimens of Belono- phago hutsebouti, Phago boulengeri, and (1961, p. 79) Eugnathichthys, but these fishes are primarily fin-eaters. The only other fresh-water scale-eaters of which I am aware are certain highly specialized African cichlids (see Fryer, Greenwood, and Trewavas, 1955). I have not searched thoroughly for accounts of marine scale-eating fishes. There probably are some; Springer and Woodburn (1960, p. 22) stated that annelids and fish scales (with no other fish remains) constituted the major portion of stomach contents of sea catfish (Galeichthys felis) taken in Tampa Bay. This paper was prepared at the Departamento de Zoologia of the Secretaria da Agricultura in Sao Paulo during a visit in April and May, 1969. All obser- vations are based on specimens in the Departamento’s collections. Measurements of fishes given in mm. refer to standard length. OBSERVATIONS Probolodus. MATERIAL EXAMINED. DZSP 7903, 40 specimens, 41-97 mm., from Rio Pa- raiba below représa de Santa Branca (state of Sao Paulo), col. 10-13 February 1962 by H. A. Britski; and DZSP 7904, 92 specimens, 38-71 mm., représa de Santa Branca, Rio Paraiba, collected 10-16 September 1963 by H. A. Britski and J. Rossi. STOMACH CONTENTS. Stomach contents were examined in 30 specimens rang- ing from 38 to 97 mm. Twenty of these were from the February, 1962 collection, and 10 from September, 1963. The stomachs contained food in all specimens. Scales were by far the major item encountered and occurred in all but 1 specimen. They were the only item present in about 50 percent. The number of scales in a stomach varied from 3 to 40, with a mean of about a dozen. Most of the scales were 3—5 mm. in diameter, substantially larger than Probolodus’ own scales. A white substance of loose consistency was present in large quantity in 5 specimens of the February, 1962, sample. Otherwise food items in the 2 samples were very similar. The following items were also encountered: small seeds (1 or 2), in 3 specimens; soil? (small quantities), 3 specimens; minute crustaceans (about 50), 1 specimen; insect larva (1), 1 specimen; hymenopteran (1), 1 specimen. The smallest specimens examined—38, 41, 47, and 49 mm.—have stomach contents similar to the others. DENTITION. The teeth of Probolodus have been described and figured by Eigenmann (1915, pp. 20-21, fig. 5). Probolodus has very few teeth and, as in VoL. XXXVIIT] ROBERTS: SCALE-EATING FISHES 385 many other characoids with highly specialized dentition, the number is constant or very nearly so. Basically there are 3 widely separated teeth on each pre- maxillary and 5 on each dentary. Often a tooth is missing, but this is due to loss or shedding to make way for a replacement tooth. There are usually either 3 or 4 teeth on each maxillary, but as few as 2 or as many as 5 were present on some specimens. Here, too, replacement affects the number present. All of the teeth are strictly tricuspid. The 3 cusps form a triangle with the enlarged median cusp at the anterior angle. The lateral cusps are equal in size and very small. The tooth base is moundlike and stout. The premaxillary teeth point out of the mouth. The first 3 dentary teeth also point out. Only the fourth and fifth dentary teeth lie inside the mouth. The enlarged fourth dentary tooth is situated internally to the third and slightly posterior to it, and the reduced fifth is directly behind the fourth. (Note— Eigenmann refers to one or more small teeth behind the fourth. In specimens I have examined there is only one. Perhaps the presence of additional teeth in an occasional specimen is a primitive or vestigial character.) The cusps of the ante- riormost premaxilliary and dentary teeth point almost straight ahead of the fish. The third dentary tooth, and to a lesser extent the third premaxillary tooth, pro- ject laterally from the mouth. The teeth are not juxtaposed but are separated from each other by a gap about equal to the diameter of a tooth base. When the mouth is closed the teeth of the upper and lower jaws interdigitate rather than truly oppose each other. Thus the first dentary tooth occupies the gap between the first and second premaxillary teeth, the second dentary tooth that between second and third premaxillary teeth, and the third dentary tooth that between the third premaxilliary and first maxillary teeth. The fourth and fifth dentary teeth do not oppose or interdigitate with other teeth and neither do the lowermost teeth on the maxillary. One can easily imagine how scales are firmly grasped by such teeth and then dislodged by the kind of tugging movements many characids make when feeding. The number, form, and arrangement of the teeth are the same in specimens from 38 to 97 mm. TooTH REPLACEMENT. Twenty specimens from the September, 1963, collec- tion were examined for signs of tooth replacement. In only 2 specimens were all of the premaxilliary and dentary teeth in functional position and firmly attached to the jawbones. In each of the remaining 18 from 1 to 4 teeth were in the proc- ess of replacement or had just come into functional position (teeth in the process of replacement can be detected immediately below the gum or making their way through it; teeth that have just come into functional position are recognizable as such because the cusps are unworn and very sharp, the bases are usually sur- rounded by soft, swollen tissue, and the attachment to the jaw is very loose). The data indicate that replacement occurs more frequently in lower jaw teeth than in upper, and that certain teeth are replaced with relatively high frequency. In all, 40 instances of tooth replacement in process and teeth newly in functional posi- 386 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER. tion were observed, 29 in the dentary and 11 in the premaxillary. No fewer than 9 instances involve the fourth dentary tooth. At the other extreme, the first pre- maxillary tooth is involved in only 1 instance. Judging from their appearance the teeth in the lower jaw receive more wear than those in the upper jaw. SEx. Almost all specimens in the February, 1962, collection have readily identifiable gonads. The others were sexed by the presence (males) or absence of tiny serrations on the anterior anal fin rays. The reliability of this method was checked in specimens-in which the sex of the gonads was obvious. Of the 40 specimens in the sample, 29 (72 percent) are females and 11 (28 percent) males. Females range from 40.5 to 97 mm. and average 68 mm., 12 mm. more than the males. Males range from 47 to 65 mm. and average 56 mm. The largest female is 32 mm. longer than the largest male. Combined biomass of females is slightly more than three times that of males. The 40.5 mm. specimen contained about 200 eggs, most .6—.7 mm. in diameter but a few somewhat smaller; a 73.5 mm. specimen contained about 2500, all about .7—.8 mm. in diameter. Serrasalmus elongatus. Stomach contents were examined in 7 specimens of S. elongatus Kner, 89-152 mm., from 3 Amazonian localities. Pieces of the rayed portion of fins and scales were present in every specimen; they were the only items encountered in 5 of the specimens. In all but 2 fin rays were by far the major item. One specimen had about 50 scales and only a few small bits of fin rays. The 152 mm. specimen (col- lected in Lago Jacupa, near Oriximina, state of Para, in February, 1967) contained 6 cichlid larvae of about 8 mm.; 13 fish? eggs of about 2 mm. in diameter; 2 large pieces of very hard, thick fin rays, perhaps from the caudal fin of a sorubim cat- fish; and 8 scales about 5-6 mm. in diameter. One specimen, with its stomach moderately full of fin rays and a few scales, had a small matted ball of fibrous plant material including 3 small seeds. All items encountered have been indicated ; noteworthy is the absence of pieces of meat. Stomach contents of several Pygo- centrus-type piranhas have been examined and when scales were encountered there were also bits of meat. Many piranhas feed to some extent on fins. S. elongatus is apparently a fin-eater which feeds to a certain extent on scales. Catoprion. Kner (1860, p. 34) found the stomach of a large Catoprion specimen full of scales. Gosline (1951, p. 54) examined the stomachs of 4 specimens and reported that ‘two were full of fish scales and two were empty except for a few fish scales; a small amount of unidentifiable debris was also found.” Géry (1964, p. 460) found scales in stomachs of specimens from Bolivia. In examining 4 specimens, 103-109 mm., from 3 Amazonian localities I find that their stomachs are more or less full of scales about 6-15 mm. in diameter. The only other items are a few bits of leaf from a higher plant (in 2 specimens) and a small ball of VoL. XXXVIIT] ROBERTS: SCALE-EATING FISHES 387 fibrous plant material, probably roots (in 1 specimen). Scales are thus the only item that has been encountered in substantial amounts in stomachs of Catoprion. The teeth in this genus are illustrated by Miller and Troschel (1845, pl. 2, fig) 5 )). Exodon. Géry (1964, p. 459) reported scales in stomach contents of Exodon from the Rio Araguaia. I examined 10 specimens, 36—59.5 mm., from the Rio Araguaia at Aruana and found from 4 to 15 scales, mostly 3-5 mm. in diameter, in every one. The only other item was small amounts of unidentifiable material in 2 speci- mens. Kner (1860, p. 47) found beetles in 2 specimens from the Rio Branco. The teeth of Exodon are figured by Miller and Troschel (1845, pl. 4, fig. la). Roeboexodon. This genus has hitherto been known only from a few specimens taken in French Guiana (Géry, 1959). In September, 1966 Heraldo A. Britski and P. E. Vanzolini collected 2 specimens (DZSP 4815, 41.5 and 45.5 mm.) from the Rio Araguaia near Aruana in the Brazilian state of Goias. The dentition of these specimens is identical with that in an alizarin preparation of a 29 mm. specimen from French Guiana (kindly sent to the Departamento de Zoologia by Géry) and they apparently represent the same species. The stomach contents of both specimens consist exclusively of scales from about 2.5 to 4 mm. in diameter. The 41.5 mm. specimen contained about 10 scales and the 45.5 mm. specimen about 20. The teeth of Roeboexodon are described and partially figured by Géry (1959, pp. 347-349, fig. 2). Roeboides. Naercio Menezes and I examined stomach contents in 9 specimens of Roebo- ides guatemalensis, 6 of R. myersi, and 25 of R. prognathus. In all 6 specimens of R. myersi (117-160 mm.) and in the 11 largest of R. prognathus (70-90 mm.) the stomachs are more or less filled with scales, to the exclusion of all else, those of R. myersi with from 15-35 scales mainly 6—9 mm. in diameter and those of R. prognathus with 40-150 scales 3-6 mm. in diameter. In 14 smaller examples of R. prognathus (41-68 mm.) scales predominate, but insects—Diptera, Hemip- tera (Notonectidae?), and a few Coleoptera—occur with high frequency. A 64 mm. specimen contained a fish larva. Our specimens of R. guatemalensis (72.5— 101 mm., from Gatun Lake, Panama Canal Zone, collected in November, 1965) have viscera heavily infested with nematodes and may not have been feeding normally. The stomachs are empty in 4 of them and the other five contain but little food, as follows: a few scales (in 4); shrimp (in 2); insect (in 1); and an unidentified, flocculent, white material (in 1). In very small specimens of Roeboides (20-30 mm.) the teeth can be recognized 388 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. as belonging to Roeboides because of their slightly hypertrophied bases, but they are all normal in position. Examination of stomach contents in a few specimens (unidentified to species) indicates that at these sizes they feed primarily on insects. Only at about 30-60 mm., depending on the species, is the transition made to the adult condition in which teeth with greatly hypertrophied bases pro- ject from the front of the jaws. DISCUSSION Stomach contents of fishes belonging to Catoprion, Probolodus, Exodon, Roe- boides, and Roeboexodon indicate that their diet consists mainly of scales. The teeth are so highly specialized in some of these fishes as to suggest that they could not survive in nature on the food that their non-scale-eating ancestors fed upon. The remarkable “twin spot” color pattern of Exodon and relatively small scales of Probolodus may have evolved after the scale-eating behavior to reduce intraspecific scale-eating. Breder (1927, p. 127) speculated that the small, thin, and very adherent nature of the scales of Roeboides occidentalis reduces auto- predation. Whereas Catoprion, Exodon, Roeboexodon, and Probolodus are mono- typic, Roeboides has speciated extensively. The relationships of the five genera, although not yet well understood, show that they represent at least three independent lines of evolution: 1. Catoprion is definitely a serrasalmid, and probably descended from Serrasalmus. Serrasal- mus elongatus includes scales in its diet but is primarily a fin-eater and does not appear to be closely related to Catoprion. 2. Eigenmann (1911; 1915) stated that Probolodus is very similar in general appearance to Astyanax fasciatus but placed it in his polyphyletic subfamily Aphyocharacinae (= Cheirodontinae). In my opinion Probolodus belongs in the Tetragonopterinae; it probably descended from Astyanax. It is certainly not related to either Chezrodon or A phyocharax. 3. Roeboides is closely related to Charax and Eucynopotamus. Géry (1959, pp. 404-405) suggested that Exodon was derived from Holobrycon and Roeboexodon from Exodon, and placed Roeboides and Charax in a different line. Naercio Menezes and I intend to study the osteology of these Characinae in an effort to clarify their relationships. We suspect that Eucynopotamus, apparently inter- mediate between Charax and Roeboides, is actually based on the young of Roe- boides, and note that Roeboexodon bears a strong superficial resemblance to Roeboides prognathus Boulenger. Perhaps the strange gymnotoid eel Oedemognathus exodon Myers is a scale- eater. According to Myers (1936, p. 115), in this apteronotid “the dentigerous portion of the premaxillaries is greatly expanded and bulbous, most of it not opposable to the lower jaw, and the upper part of it rising above the profile of the snout. The whole of this bulbous area is studded with many strong, slightly curved, conical teeth, placed irregularly and not very closely together. Most of the upper teeth therefore project forward, outward or upward, and are entirely VoL. XXXVIIT] ROBERTS: SCALE-EATING FISHES 389 outside the mouth. The lower teeth are similar to the upper ones in shape, and are numerous and arranged irregularly, but none is outside the mouth and all point in normal direction.” Oedemognathus is known only from the holotype, 202 mm. in total length, USNM 102040, and a 92 mm. specimen reported on and figured by Eigenmann and Allen (1942, pp. 325-326, pl. 15, figs. 2-3), CAS (I1UM) 15421, both from the Peruvian Amazon. ACKNOWLEDGMENTS I wish to thank Daniel M. Cohen of the U. S. Fish and Wildlife Service and Stanley H. Weitzman of the U. S. National Museum for critical comments on the manuscript. ADDENDUM Mr. William A. Bussing of the Departamento de Biologia, Universidad de Costa Rica, informs me that Sr. Carlos Leon, Administrador of the Parque Boli- var in San José, Costa Rica, observing 2 fish in an aquarium, saw one (Roeboides guatemalensis) butt the other (Astyanax sp.) with its snout and then catch the dislodged scales as they sank. Bussing has examined the viscera of about 100 representatives of R. gwatemalensis during fieldwork on the Atlantic and Pacific slopes of Costa Rica and found almost every specimen had scales and virtually nothing else in the stomach. A few contained small insect larvae and one a small fish. At the John G. Shedd Aquarium in Chicago, Mr. Emanuel Ledecky-Janecek, Curator of Exhibits, kindly responded to my request and placed a specimen of Leporinus (perhaps L. friderici) about 9 or 10 inches long in with a small tank- ful of fish belonging to Exodon paradoxus. Within a few moments we saw the latter agitatedly gang up to one side of the Leporinus victim and take turns making extremely rapid circular stabbing motions against its side, always strik- ing towards the free margin of the scales, and removing a single scale at about every other strike. The scales were swallowed directly. On only one occasion did a scale fall to the bottom of the tank and a moment later it too was devoured. In about 5 or 10 minutes 20 or 30 scales had been eaten. At the Steinhart Aquarium of the California Academy of Sciences I watched several fish belonging to Leporinus fasciatus determinedly nipping at fungus- infected sores on a specimen of Astronotus ocellatus. I am unsure, but think that the cichlid lost a few scales, although the Leporinus specimens seemed to confine their nipping to the sores. LITERATURE CITED BREDER, CHARLES M. 1927. The fishes of the Rio Chucunaque drainage, eastern Panama. Bulletin of the American Museum of Natural History, vol. 57, art. 3, pp. 91-176. 390 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. EIGENMANN, Car H. 1911. New characins in the collection of the Carnegie Museum. Annals of the Carnegie Museum, Pittsburgh, vol. 8, no. 3, pp. 164-181, pls. 4-9. 1915. The Cheirodontinae, a subfamily of minute characid fishes of South America. Memoirs of the Carnegie Museum, Pittsburgh, vol. 7, no. 1, pp. 1-99, pls. 1-17. EIGENMANN, Cart H., and W. R. ALLEN 1942. Fishes of western South America. University of Kentucky, pp. i-xv, 1-494, map. Fryer, Greorrroy, P. H. GREENwoop and E. TREWAVAS 1955. Scale-eating habits of African cichlid fishes. Nature, vol. 175, no. 4468, pp. 1089- 90. GErRY, JACQUES 1959. Contribution a l’étude des poissons characoides (Ostariophysi) (II.) Roeboexodon gen. n. de Guyane, redescription de R. guyanensis (Puyo, 1948) et relations probables avec les formes voisines. Bulletin du Muséum National d’Histoire Naturelle, second series, vol. 31, no. 4, pp. 345-352; and no. 5, pp. 403-409. 1964. Poissons characoides nouveaux ou non signalés de Vilha do Bananal. Vie et Milieu, suppl. 17 (Volume Jubilaire dédié a Georges Petit), pp. 447-471. GOSLINE, WILLIAM A. 1951. Notes on the characid fishes of the subfamily Serrasalminae. Proceedings of the California Academy of Sciences, fourth series, vol. 27, no. 2, pp. 17-61, pls. 1-3. Kner, RUDOLF 1860. Zur Familie der Characinen, IJ. Denkschriften der mathematisch-naturwissen- schaftlichen Classe der kaiserlichen Akademie der Wissenschaften, Wien, vol. 18, pp. 9-62, pls. 1-8. MattTues, HUBERT 1961. Feeding habits of some central African fishes. Nature, vol. 192, pp. 78-80. 1964. Les poissons du Lac Tumba et de la région d’Ikela. Annales Musée Royale de lAfrique Centrale, series in octavo, sciences zoologiques, no. 126, pp. 1-204, 2 maps, 1 chart, pls. 1-6. MULLER, JOHANNES, and F. H. TRosCHEL 1845. Horae Ichthyologicae. I & II, Die Familie der Characinen. Berlin, pp. 1-40, pls. 1-11. Myers, GEorGE 5S. 1936. A new genus of gymnotid eels from the Peruvian Amazon. Proceedings of the Biological Society of Washington, vol. 49, pp. 115-116. 1942. Studies on South American fresh-water fishes. I. Stanford Ichthyological Bulletin, vol. 2, no. 4, pp. 89-114. SPRINGER, Victor G., and K. D. WoopBuURN 1960. An ecological study of the fishes of the Tampa Bay area. Florida State Board of Conservation, Marine Laboratory (St. Petersburg), Professional Papers Series, no. 1, pp. 1-104. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Festschrift for George Sprague Myers Vol. XX XVIII, No. 20, pp. 383-390 December 31, 1970 SCALE-EATING AMERICAN CHARACOID FISHES, WITH SPECIAL REFERENCE TO PROBOLODUS HETEROSTOMUS ay Tyson R. Roberts Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138 INTRODUCTION Specialized scale-eaters have been discovered in three groups of American characoids. Kner (1860, p. 34) reported the stomach of a large specimen of Cato- prion full of scales and referred to them as ‘‘Raubfische.’’ Ladiges, observing this peculiar serrasalmid in an aquarium, saw one remove a row of scales from a specimen of Metynnis with one swipe of its teeth (reported by Géry, 1964, p. 460). Breder (1927, p. 127) reported substantial amounts of large scales in stomach contents of representatives of Roeboides occidentalis from eastern Pan- ama and identified some of the scales as coming from Ctenolucius, a pike-like characoid considerably larger than the specimens of Roeboides. Géry (1964, pp. 459-460) reported scale-eating in Exodon, Roeboides, and Roeboexodon, of the characid subfamily Characinae. In this paper the activity is verified for Cato- prion, Exodon, Roeboexodon, and two additional species of Roeboides and is re- ported for the first time in Probolodus heterostomus Eigenmann,' a member of the characid subfamily Tetragonopterinae. Although Géry supposed that scale-eating occurred in Catoprion, Exodon, Roeboexodon, and Roeboides only occasionally, in these genera and in Probolodus scales are definitely a major item in stomach contents, and eating scales is prob- 1 Myers (1942, p. 91), in recording specimens from the western end of the coastal plain of Rio, com- mented on their almost unbelievably strange dentition. The species also occurs in the rios Doce, Paraiba (formerly spelt Parahyba), and Ribeira. [383] 392 CALIFORNIA ACADEMY OF SCIENCES [PRroc. 4TH SER. anal spine has been overlooked in Naso, a genus thought to be alone among the acanthurids in having only two anal spines rather than the normal three. Naso, like Prionurus (including Xesurus), has fixed plates on the caudal peduncle rather than the folding spines typical of all other acanthurids. Naso and Paracanthurus are the only genera with 3 soft rays in the pelvic fin rather than 5. On the basis of Naso and Prionurus having fixed peduncular plates, Smith (1955: 169) recognized these 2 groups (with Naso divided into 5 genera on the basis of the number of peduncular plates and the snout horn development) as the family Nasidae distinct from the Acanthuridae. Randall (1955: 365-366), in a brief addendum to his revision of the surgeon fish genera, gave excellent rea- sons for not accepting this splitting of the family and of the genus Naso. Smith (1966: 635) subsequently recognized Naso (this time divided into 3 genera) as the subfamily Nasinae on the basis of its having 2 anal spines and 3 pelvic rays, in contrast to the 3 anal spines and 5 pelvic rays of the other acanthurids which he divided into 2 subfamilies on the basis of whether the caudal peduncle has folding spines (Acanthurinae) or fixed plates (Prionurinae). Such a system neglects Paracanthurus, which, in addition to 3 anal spines, has folding pedun- cular spines and only 3 pelvic rays. In Smith’s system, Paracanthurus would have to be recognized as an additional subfamily. By most contemporary stan- dards this system would seem to be far too finely split at the subfamilial and generic levels on the basis of somewhat superficial characters, even if all these characters were valid. The fact that Vaso has 3 anal spines like the other acanthurids is additional evidence that this genus should not be considered as a subfamily distinct from the other acanthurids. The anal spine that is supposedly absent in Naso is the first spine. This spine is substantially similar to the first spine in other acanthu- rids, except that its distal portion which would protrude through the skin is lost, leaving only the basal portion which acts as a complex locking device in basically the same manner as in other acanthurids. An initial survey of acanthurid osteology, which cannot be dealt with here, based on representatives of all of the genera recognized by Randall (1955), shows no features that warrant the recognition of subfamilies within the group. LOCKING MECHANISM OF THE DORSAL AND ANAL SPINES The locking mechanism in Acanthurus triostegus is osteologically representa- tive of all of the acanthurids except Vaso, and the descriptions and illustrations of the bony parts given here based on A. triostegus should apply well except in fine detail to the various species of Acanthurus, Ctenochaetus, Paracanthurus, Prio- nurus, and Zebrasoma. The musculature of the locking mechanism is described for A. triostegus, and although this has not been compared with the situation in the other genera that have 3 obvious anal spines, I suspect that, based on the shapes of the bony parts, it is similar in all of them. The bony parts of the Vor. XXXVIII] TYLER: SPINE LOCKING IN SURGEON FISHES 393 locking mechanism as described for Naso literatus are typical of that genus, and the few comments given here on the musculature probably will apply to all species of Naso. The dorsal and anal spines and their pterygial supports are more generalized in Acanthurus than in Naso, and are described first below. Acanthurus triostegus (Linnaeus). SPECIMENS EXAMINED. ANSP 109491, 7 specimens, 40.6—68.9 mm. stan- dard length, Caroline Islands, cleared and stained. ANSP 109490, 1, 69.0 mm., no locality, cleared and stained. ANSP 108288, 4 (out of 12), 83.6-97.0 mm., Seychelles Islands, alcohol. DoRSAL SPINES AND PTERYGIOPHORES. The first two are borne on the first basal pterygiophore, and the subsequent seven spines on their own individ- ual basal pterygiophores as well as distal pterygiophores. The first two spines ar- ticulate medially with the large flattened medial flange at the distal end of the basal pterygiophore, which projects above the level of the distal ends of the subse- quent pterygiophores, as well as ventrolaterally by their divergent bases with lat- eral flanges on the sides of the pterygiophore. The third and subsequent spines articulate at their less divergent bases with the regions of suturing between the distal pterygiophores anteroventral to their bases and the basal pterygiophores ventrally and posteroventrally. The first basal pterygiophore is held basally be- tween the exoccipitals and the dorsal half of the neural spine of the first vertebra. Each of the subsequent basal pterygiophores of the spiny dorsal fin is held between the neural spines of adjacent vertebrae, except that there is no pterygiophore between the third and fourth neural spines (fig. 1). This is true of all acanthurids, regardless of the number of dorsal spines, and is also true of zanclids. In the closely related siganids there is one basal pterygiophore of the spiny dorsal fin be- tween adjacent neural spines, except that there is no pterygiophore between the fifth and sixth neural spines. In siganids the first dorsal spine is normal, not modified into an acanthurid-like locking mechanism. However, the distal ends of the pterygiophores are laterally expanded into plates in much the same manner as explained below for acanthurids, especially Vaso. The basal region of the first spine is deeply concave, while that of the second spine has a complete foramen anteroposteriorly through which the medial flange of the first basal pterygiophore passes, the flange in this region having a hole to accommodate the extreme ventral end of the second spine, which is solid and without a medial suture (figs. 2-3). The third spine also has a complete antero- posterior foramen in its base through which passes the medial bridge formed by the posterodorsal process of the first distal pterygiophore and the anterodorsal process of the second basal pterygiophore. The fourth and subsequent dorsal spines usually have a complete foramen ventrally, although the ventromedial region may have a sutural mark medially. The bridges formed by the processes 394 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. supraoccipital , supracleithrum \st E abdominal vertebrae eee ig skpi uly juesinz0id \ ' ' ' Ficure 1. Lateral view of vertebral column and fin supports in Acanthurus triostegus, ANSP 109491, 46.2 mm. standard length. Bases of fin rays indicated in black; distal pteryg- iophores of fin rays not shown. VoL. XXXVIII] TYLER: SPINE LOCKING IN SURGEON FISHES 395 \ Qr nd spine 1st distal : pid spine 1st dorsal pterygiophore } haemal spine (40th vertebra) ist basal Spine i | pterygiophore | 2nd basal g._——_ base of 4th spine 2nd distal Pterygiophore pterygiophore fF 1st basal pterygiophore , ——2nd basal pterygiophore (7 neural spine (1st vertebra) 2nd distal pterygiophore a, base of 1st ray oo eae Sn = ss i aera 3rd spine fst anal spine \ ist distal pterygiophore Ficure 2. Lateral views of first three dorsal and anal spines and their supports in Acan- thurus triostegus, the spines only partially erected; composite drawings based on specimens from ANSP 109491. of the distal and basal pterygiophores, around which the foramina of the spines articulate, are slightly less well developed posteriorly in the series than anteriorly, the anterodorsal process of the basal pterygiophore especially tending to be of lesser length so that it does not quite contact the process of the distal pterygio- phore in front of it and thus fails to form a complete bridge. Distally the pterygiophores are expanded laterally to form a plate composed anteriorly of the expanded posterior half of the distal end of the basal pterygio- phore and posteriorly by the expanded anterior portion of the distal pterygio- phore. The tendinous insertions of the erector muscles of the spines (except for those of the first 2 spines) are accommodated by gaps between the composite plates, the gaps being between the posterior edges of the expanded portions of the distal pterygiophores and the anterior edges of the expanded portions of the basal pterygiophores (fig. 4). The depressor muscles are accommodated by similar gaps in the anterior third of the laterally expanded plates of each of the basal pterygiophores. The anterolateral edges of the basal pterygiophore plates tend to be prolonged anteriorly, partially enclosing the lateral surfaces of the insertion ends of the depressor muscles. The amount of bridging in these regions 396 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SEr. Ficure 3. Lateral view of first two dorsal spines in Acanthurus triostegus, the spines almost fully erected; composite drawing based on specimens from ANSP 109491. Vout. XXXVIII] TYLER: SPINE LOCKING IN SURGEON FISHES 397 Ist basal pterygiophore 2nd + 3rd basal pterygiophores distal pterygiophores Ficure 4. Dorsal view of first three basal and distal pterygiophores of dorsal fin of Acanthurus triostegus, after removal of the spines, ANSP 109491, 46.2 mm. standard length. may increase with increasing specimen size, but it probably never reaches the complete bridging found in Naso, as discussed later. As one of the major characteristics of the acanthurids, Gill stated (1884: 276; this statement often followed by others): “Interneurals with expanded buckler-like subcutaneous plates, which intervene between the spines and limit their erection forwards.” The plates he refers to obviously are the laterally ex- panded distal ends of the basal and distal pterygiophores, but these plates in no way interfere with the full erection of the spines, as explained below. Connective tissue in dried skeletal preparations or unnaturally inelastic membranes and ligaments in alcohol preserved specimens must have misled him. All of these spines can be erected at right angles to their pterygiophores. The first and second spines are bound together by a compact ligament, while the second and subsequent spines are bound together by more diffuse ligamentous connections spread out over most of the lengths of the spines. The erection or depression of one spine thus is in concert with the others. When the spines are fully erected at right angles to their pterygiophores, they tend to stay erected because of the frictional resistance of their ventral edges against the dorsal sur- faces of their pterygiophores caused by the down pull of the erector muscles, but the whole series of spines can be firmly locked in an erected position varying from full to any degree of partial erection by a catching action of the small first spine against grooved surfaces on the dorsomedial flange of the first basal pterygiophore. The locking mechanism probably develops very early in the larval stages, for the figures of 3 to 5 mm. long specimens of A. monroviae given by Abousso- 398 CALIFORNIA ACADEMY OF SCIENCES [Proc, 4TH SER. uan (1965: figs. 2-3) clearly indicate the grooved medial flange of the first basal pterygiophore, although the small first spine is not shown. The dorsomedial flange of the first basal pterygiophore is generally circular in outline as seen laterally, with the exception of the region just behind the mid- dle of the flange discussed below. The flange bears grooves running from its distal edge approximately towards the center. The deeply concave ventral sur- face of the first spine is relatively smooth and rides over the grooved peripheral surface of the medial flange of the first basal pterygiophore. The ventral arms, or extreme basal flanges, of the first spine rotate around a small flange projecting laterally from the lateral surface of the pterygiophore at the level of the horizon- tal base of the medial flange. The dorsal surface of this pterygiophore flange and the ventral edge of the ventral arm of the first spine which it supports are both knurled, the amount of knurling being highly variable in the specimens studied. As the first spine rotates in erection forward and downward it slides easily over the variously anteriorly to dorsally oriented grooves on the medial flange of the basal pterygiophore. When a downward and/or backward pressure is exerted on the first spine, it catches firmly on the grooves of the medial flange. Since the first and subsequent spines are ligamentously connected, the whole spiny dorsal fin is firmly held erect. Because the grooves on the medial flange are continuous from the region underlying the concave ventral surface of the first spine in its unerected position to its fully erected position at right angles to the pterygiophore, the first spine can be locked in any of the innumerable posi- tions between the two extremes. The first spine is unlocked from its erected position by relaxation of the pressure of the muscles (as discussed below) pulling it into intimate contact with the grooves on the medial flange of the basal pteryg- iophore, allowing it to slide upward and posteriorly without undue frictional resistance. The rotational course of the first spine is blocked anteriorly when it reaches its fully erected position at about a right angle to the pterygiophore by its anterior edge hitting against two bony obstacles on a prong-like portion of the pterygiophore anterior to the medial flange and separated from it by a deep vertical canal into which the head of the first spine rotates in erection. The extreme anterior end of the first spine hits the bottom of the canal at the same time that a slight indentation on the mid-dorsal (as seen when unerected) edge of the spine hits against the posterodorsal edge of the prong-like portion of the pterygiophore. The second spine rotates over the posterior half of the medial flange of the basal pterygiophore. The peripheral surface of the flange over which it slides is also grooved, although not so deeply or regularly as that portion over which the first spine slides. Nevertheless, downward pressure on the second spine serves to help lock the spiny dorsal fin in an erected position. The ventral arms (which are fully fused in the region of the foramen in the medial flange) of the second spine rotate around the oblique dorsal surface of a large flange running Vor. XXXVIII] TYLER: SPINE LOCKING IN SURGEON FISHES 399 vertically along most of the lateral surface of the first basal pterygiophore. The rotational course of the second spine is blocked anteriorly when it reaches a fully erected position at about a right angle (or a little more) to the pterygiophore by its anterior edge just at the dorsal end of the anteroposterior hole through its base hitting against an indented and thickened region on the medial flange. The medial flange is thicker beneath the position of the base of the unerected first dorsal spine than more posteriorly, and the lateral ridge so formed corresponds to an indented region on the flange just posterior to the ridge. The anterior edge of the second spine in full erection hits against this ridge and indentation, stop- ping any further rotational movement forward. The third and subsequent spines, by comparison, rotate simply over their articulations with the bridges formed by the posterodorsal processes of the distal pterygiophores and the anterodorsal processes of the basal pterygiophores, with- out any special mechanisms for locking them in an erected position, other than their ligamentous connections with the first 2 spines. MuscuLature. When the skin is removed from the upper part of the trunk below the dorsal spines, 3 main muscle groups are seen (fig. 5, top): the general epaxial muscle mass; the inclinators which originate on the undersurface of the skin and course posterodorsally to insert variously on the expanded plates of the pterygiophores and on the ventrolateral surfaces of the bases of the spines; and the supracarinales anterior, a band of longitudinal muscle running from the supraoccipital to the lateral surfaces of the anterodorsal prong-like portion of the first basal pterygiophore and of the base of the first spine. The epaxial muscle mass has nothing to do with the erection mechanism and is not further discussed. The inclinators of the first 2 spines are poorly developed in compari- son to those of the subsequent spines and of the soft rays, and they probably have little bearing on the locking mechanism. The band of muscle from the supraoccipital divides posteriorly into a deep segment attaching to the prong of the basal pterygiophore and a superficial segment attaching to the basal region of the first spine. The segment attached to the pterygiophore prong apparently is not associated with the locking mechanism, but the segment attached to the spine may help to unlock the spine prior to depression of the fin. When the first spine has been pulled firmly against the grooved surface of the pterygiophore flange it is locked in place, and a forward and/or upward directed pressure is necessary to disengage it from the grooves before it can slide without undue frictional resistance over the flange. Contraction of the longitudinal band of muscle between the supraoccipital and the first spine probably helps to disengage the spine by pulling it forward. However, the first anal spine has the same type of locking mechanism as the first dorsal spine, yet the anal spine does not have a band of muscle running forward from its base which, when contracted, would tend to disengage the spine from its locked position, nor is such a muscle present on the first dorsal and anal spines of Naso. I assume that the natural elasticity 400 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER. Ist basal pterygiophore 2nd basal pterygiophore 3rd spine 2nd_ spine ligament Ist spine supraoccipital 2nd distal pterygiophore 2nd_ basal pterygiophore Ist abdominal vertebra Ficure 5. Lateral views of musculature of first three dorsal spines in Acanthurus trioste- gus; above, superficial musculature after removal of skin; below, erector and depressor muscles as seen after removal of superficial muscles: composite drawings based on specimens from ANSP 108288; the representational fiber structures are diagrammatic. Legend: A, supracarinales anterior; B, inclinators; C, epaxial muscle mass; D, depressors; E, erectors. and resiliency of the various connective tissues between the first spine and the pterygiophore are strong enough to push or pull the spine the slight distance necessary to relieve the pressure on its concave ventral surface from the grooves on the medial flange of the pterygiophore. The relative smoothness of the ventral VoL. XXXVIII] TYLER: SPINE LOCKING IN SURGEON FISHES 401 surface of the spine perhaps facilitates the unlocking of the mechanism, for cer- tainly if it were deeply grooved like the medial flange a complex mechanism for unlocking the adpressed grooved surfaces would have to be present. The erector muscle of the first spine is the largest in the series (fig. 5, bot- tom). It originates over the middle of the posterior wall of the skull and over the lateral surfaces of the first vertebral neural spine and first pterygiophore. It inserts on the forward half of the lateral face of the base of the spine. In the upper third of its length, the muscle is partially divided into a forward and rear segment. Contraction of the muscle pulls the spine forward and downward over the medial flange. When the spine is erected it is probably the contraction of the rear segment of the upper third of the muscle that pulls the spine into close contact with the grooves of the medial flange. The muscle overlies much of the length of the large lateral flange running vertically on the pterygiophore, the head of this flange being the point around which the second spine pivots. The depressor muscle of the first spine originates under the upper regions of the erectors of the first and second spines. It is much smaller than any of the other depressor muscles in the series. It inserts through a band of connective tissue on a posteriorly directed process from the posterior edge of the basal flange of the first spine. The contraction of the depressor muscle simply rotates the first spine backward over the medial flange to its unerected position. The erector and depressor muscles of the second and subsequent spines re- quire no special comment. ANAL SPINES AND PTERYGIOPHORES. The 3 anal spines and the distal por- tions of the pterygiophores to which they articulate have much the same shapes and relationships as the first 3 dorsal spines do to their pterygiophores, as de- scribed briefly above. The ventral surface of the basal half of the first spine is deeply concave, fitting over a grooved medial flange of the first basal pterygio- phore; the second and third spines have complete anteroposterior foramina through their bases, that of the second spine fitting through the bridging of a hole in the posterior half of the medial flange of the first pterygiophore and that of the third spine through the bridge formed by the posteroventral process of the first distal pterygiophore and the anteroventral process of the second basal pterygiophore. The third anal spine is stouter than the third dorsal spine, and the distal ends of the pterygiophores are not expanded into large plate-like struc- tures such as are found on the pterygiophores of the spiny dorsal fin. The lock- ing mechanism of the first anal spine is like that of the first dorsal spine, and the rotational course of the first 2 spines is blocked when the spines are about at a right angle to the pterygiophore in the same manner as described for the spiny dorsal fin. The long shaft-like proximal portion of the first basal pterygiophore is firmly held by connective tissue against the anterior edge of the haemal spine of the tenth vertebra, the first of the caudal series. The proximal portion of the second 402 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. haemal spine - 10th vertebra Ist basal pterygiophore 2nd_ basal pterygiophore Ist distal pterygiophore pelvis 3rd_ spine Ist spine ligament 2nd spine Ficure 6. Lateral views of musculature of anal spines in Acanthurus triostegus; below, superficial musculature after removal of skin; above, erector and depressor muscles as seen after removal of epaxial muscles and inclinators; composite drawings based on specimens from ANSP 108288; the representational fiber structures are diagrammatic. Legend: A, infracarinales medialis; B, inclinators; C, hypaxial muscle mass; D, depressors; E, erectors. basal pterygiophore is held against the posterior edge of this haemal spine, while the first and second basal pterygiophores of the soft anal fin are held respectively to the anterior and posterior edges of the haemal spine of the second caudal vertebra. MuscuLature. Three main muscle groups are apparent when the skin is Vor. XXXVIII] TYLER: SPINE LOCKING IN SURGEON FISHES 403 removed from the trunk above the anal spines (fig. 6, bottom): the general hypaxial muscle mass; the inclinators; and a band of longitudinal muscle run- ning from the lateral surface of the pelvis just behind the level of the fin rays to the lateral surface of the anteroventrally directed medial prong-like portion of the first basal pterygiophore. The inclinators apparently are increased in number relative to those of the dorsal spines and are not associated with the anal spines on a one-to-one basis as they are with the dorsal spines. They originate on the undersurface of the skin and insert variously on the distal regions of the pteryg- iophores and on the lateral surfaces of the bases of the spines. The inclinators, like the hypaxial muscles, have no bearing on the locking mechanism. The band of muscle (infracarinales medialis) between the pelvis and first basal pterygio- phore becomes tendinous as it passes around the anus. This muscle in some ways corresponds to the one in the spiny dorsal fin which runs from the supraoccipital to the prong of the first basal pterygiophore and base of the first spine. How- ever, in the anal fin this muscle inserts only on the anteroventral prong of the first basal pterygiophore and does not have a superficial segment making con- tact with the first spine. It apparently has nothing to do with the locking mecha- nism. The erector and depressor muscles are shown at the top of figure 6. As with the first dorsal spine, the erector muscle of the first anal spine is the largest in the series, and its depressor is the smallest. The erectors and depressors of the subsequent spines are of about equal size. They rotate the spines in the same manner as described for the spiny dorsal fin. Naso literatus (Bloch and Schneider). SPECIMENS EXAMINED. ANSP 109497, 2, 111.2—208.5 mm., tropical western Pacific, cleared and stained. ANSP 109496, 1, 191.4 mm., no locality, cleared and stained. ANSP 108416, 1, 107.3 mm., Seychelles Islands, cleared and stained. ANSP 108272, 1, 168.3 mm., Seychelles Islands, alcohol; species undetermined, but not NV. literatus, used for muscle examination. DoRSAL SPINES AND PTERYGIOPHORES. There are 7 dorsal spines, the first of which is so distally aborted that it does not show externally, and the dorsal spine counts given in the literature are nearly always one less than in actuality (fig. 7). The second and subsequent dorsal spines are similar in their articula- tions to those described above for Acanthurus triostegus, except that the spines of Naso are stouter (and more heterocanth) and the distal pterygiophores to which the third and subsequent spines articulate are smaller, the latter difference being described below. The first dorsal spine of Naso differs from that of Acan- thurus and the other genera of acanthurids only in having lost its distal portion that would ordinarily protrude through the skin (fig. 8). The basal region of the spine (all that remains of it) is similar to that described above for A. ¢rio- stegus, including a deeply concave smooth ventral surface, ventral arms articu- 404 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER. supraoccipital supracleithrum i basioccipital Ist abdominal vertebrae | | | - ~ ! | ! Lateral view of vertebral column and fin supports in Naso literatus, ANSP 109497, 111.2 mm. standard length. Bases of fin rays indicated in black; distal pterygiophores of fin rays not shown. FIGURE 7. VoLt. XXXVIII] TYLER: SPINE LOCKING IN SURGEON FISHES 405 2nd spine 1st distal Pterygiophore 3rd spine 1st spine 2nd distal pterygiophore base of 4th Spine ee haemal spine (10 th vertebra) 2nd basal Pterygiophore x neural spine (1st vertebra) 1st basal pterygiophore et oh acal 2nd basal Pterygi pterygiophore ygiophore distal pterygiophore base of ist fin ray 2nd spine 3rd spine Ficure 8. Lateral views of first three dorsal and anal spines and their supports in Naso literatus, the spines only partially erected; composite drawings based on all of the cleared and stained materials listed. lating with lateral flanges on the first basal pterygiophore, and a posteriorly directed process from the posterior edge of the ventral arm to receive the inser- tion of the depressor muscle. A compact ligament connects the first spine at a groove along its dorsolateral surface to the anterolateral surface of the basal 406 CALIFORNIA ACADEMY OF SCIENCES [PRoc. 4TH SER. region of the second spine, and the second and subsequent spines are connected by more diffuse ligamentous connections, just as described for A. triostegus. The major differences between A. triostegus (and the other acanthurids with an externally visible first dorsal spine) and J. literatus (and the other species of Naso), other than the smaller size of the first dorsal spine and the somewhat stouter size and heterocanth arrangement of the other spines, are in the shapes of the pterygiophores supporting them. The medial flange of the first basal pterygiophore is much less deeply grooved in Naso than in other acanthurids, and the anterodorsal prong of the pterygiophore, which in other acanthurids is relatively laterally compressed, in Naso is expanded posterolaterally into a bridge which meets and fuses or sutures with the lateral flange on the pterygio- phore around which the ventral arms of the first spine rotate. The canal between the prong and the rest of the pterygiophore remains, and into this canal the first spine rotates when erected, just as described for A. triostegus. The rotational course of the first 2 spines is halted at about a right angle to the pterygiophore when the anterior edge of the base of second spine hits the indented region on the medial flange of the pterygiophore, and when, at the same time, the first spine can travel no farther in the canal between the anterodorsal prong and the medial flange. The second spine has a complete foramen anteroposteriorly through its base, the ventral bridge of the foramen being accommodated in a hole through the medial flange, as in A. tviostegus. The third and subsequent spines also have complete foramina, these articulating around the bridges formed by the postero- dorsal processes of the distal pterygiophores and the anterodorsal processes of the basal pterygiophores. The basal pterygiophores of the spiny dorsal fin in Naso otherwise differ from those described in A. triostegus mainly by the greater development of the ex- panded plates at their distal ends, and of the lesser involvement in this structure of the distal versus basal pterygiophores (fig. 8). In A. triostegus the distal pterygiophores have expanded anterolateral wings which contribute substantially to the plates, but in JV. literatus the plates are formed only by the basal pteryg- iophores, the distal pterygiophores remaining relatively small and medially placed in comparison to the width of the plates. Large foramina are present on each side of each expanded basal pterygiophore plate; these accommodate the depressor muscles of the second and subsequent spines, the erector muscles being accommodated in the gaps between the basal pterygiophore plates bordered medi- ally by the small distal pterygiophores (fig. 9). The locking mechanism in Vaso works just as described for A. triostegus, and, despite the less deep grooving on the medial flange in Naso, the apparatus seems to lock equally firmly. Muscutature. The muscles of Naso have been examined only cursorily, and in a species other than NV. literatus. But the major features of the bony struc- VoL. XXXVIII] TYLER: SPINE LOCKING IN SURGEON FISHES 407 2nd +3rd Ist basal pterygiophore basal pterygiophores Ist dorsal spine distal pterygiophores Ficure 9. Dorsal view of first three basal and distal pterygiophores of anal fin of Naso literatus, with the first spine in place at a position about halfway to full erection and the second and third spines removed, ANSP 108416, 107.3 mm. standard length. tures in the appropriate regions are similar in all species of Naso, and the muscu- lature probably follows suit. The longitudinal band of muscle (supracarinales anterior) between the supra- occipital and the anterodorsal prong of the first basal pterygiophore does not contact the first dorsal spine, its path being blocked by the posterolaterally ex- panded portion of the prong which forms a bridge to the lateral flange support- ing the ventral arm of the first spine. The erector and depressor muscles of the spines do not seem out of the ordinary, although the depressor muscles are even more deeply buried beneath the erectors than the condition described and shown above for A. triostegus. ANAL SPINES AND PTERYGIOPHORES. The 3 anal spines and associated re- gions of the pterygiophores are similar in their relationships to that described for the first 3 dorsal spines, except that there is no separate distal pterygiophore between the first and second basal pterygiophores, although each of the follow- ing soft rays has separate distal pterygiophores. The large proximal shaft-like portion of the first basal pterygiophore is 408 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. firmly held along the anterior edge of the haemal spine of the tenth vertebra, the first of the caudal series, and is curved much farther anteroventrally than in A. triostegus. The second basal pterygiophore is aborted proximally and squeezed in between the posteroventral region of the first basal pterygiophore of the spiny anal fin and the anterior edge of the slightly larger first basal pterygiophore of the soft anal fin. The second to sixth basal pterygiophores of the soft anal fin follow in series between the haemal spines of the first 2 caudal vertebrae. MuscutatureE. As in A. triostegus, the muscle (infracarinales medialis) between the pelvis and the first basal pterygiophore does not contact the first spine and become tendinous as it passes around the anus. The shapes and sizes of the erector and depressor muscles of the anal spines are not clear to me in the single specimen examined, but the depressor muscles seem even more deeply buried below the erectors than in the dorsal fin. MISCELLANEOUS NOTES ON THE AXIAL SKELETON. As indicated in figures 1 and 7, acanthurids have 3 epurals, the first in close association with the neural arch of the penultimate vertebra, and a well-developed paired uroneural. The hypurals (including the parhypural) number six in most acanthurids, as shown for A. triostegus, but in the several species of Naso investigated, the middle 4 elements are fused to each other and to the centrum, leaving only 2 separate elements. The haemal spines of the penultimate and antipenultimate vertebrae are autogenous in acanthurids. ACKNOWLEDGMENTS Mr. Richard Winterbottom, Queen’s University, Kingston, Ontario, Canada, generously advised on the musculature of the acanthurid locking mechanism, and Dr. Daniel M. Cohen, U.S. Fish and Wildlife Service, Washington, D.C., gave useful comments on the manuscript. The halftone illustrations are by the extremely competent Mrs. Mary H. Fuges. More generally, | am in the debt of Professor Myers for much good counsel and encouragement in the study of plectognath fishes and their relatives. This short paper in his honor is only an infinitesimal token of my deepest gratitude for his innumerable hours devoted to my education. LITERATURE CITED ABOUSSOUAN, A. 1965. Oeufs et larves de Téléostéens de [Ouest africain. I—Acanthurus monroviae Steind. Bulletin de Institute Francais d’Afrique Noire (Dakar), (A), vol. 27, no. 3, pp. 1183-1187, 3 figs. Donitz, W. 1867. Ueber die Gelenke an der Riicken- und Afterflosse der Teuthies C. Val. Archiv fur Anatomie, Physiologie, und wissenschaftliche Medizin (Leipzig), Jahrbuch, 1867, no. 1, pp. 210-220, pl. 7a. Git, THEopore N. 1884. Synopsis of the genera of the superfamily Teuthidoidea (families Teuthididae and VoL. XXXVIII] TYLER: SPINE LOCKING IN SURGEON FISHES 409 Siganidae). Proceedings of the United States National Museum (Washington, D.C.), vol. 7, no. 18, pp. 275-281. GUNTHER, ALBERT 1861. Catalogue of the acanthopterygian fishes in the collection of the British Museum, vol. 3, London, pp. 1-586. RANDALL, JOHN E. 1955. An analysis of the genera of surgeon fishes (family Acanthuridae). Pacific Science (Honolulu), vol. 9, no. 3, pp. 359-367. SMITH, Ji. L. B. 1955. East African unicorn fishes from Mozambique. South African Journal of Science (Johannesburg) vol. 51, no. 6, pp. 169-174, 2 pls. 1966. Fishes of the sub-family Nasinae with a synopsis of the Prionurinae. Ichthyolog- ical Bulletin (Department of Ichthyology, Rhodes University, Grahamstown), no. 32, pp. 635-682, 13 figs., pls. 103-104. SORENSEN, WILLIAM 1884. Om lydorganer hos fiske. En physiologisk og comparativ-anatomisk unders¢- gelse. University of Copenhagen, pp. 1-245, 4 pls. 1897. Some remarks on Dr. Thilo’s memoire on “Die Umbildungen an den Gliedmassen der Fische.” Morphologisches Jahrbuch (Leipzig), no. 25, pp. 170-189, 6 figs. THILO, OTTO 1896. Die Umbildungen an den Gliedmassen der Fische. Morphologisches Jahrbuch, no. 24, pp. 287-355, 7 figs., pls. 6-9. 1898. Erganzungen zu meiner Abhandlung “Die Umbildungen an den Gliedmassen der Fische.”” Morphologisches Jahrbuch, no. 26, pp. 81—90. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Festschrift for George Sprague Myers Vol. XXXVIII, No. 22, pp. 411-414; 1 fig. December 31, 1970 AMPUTATION AND REPLACEMENT OF MARGINAL SPINES IN CTENOID PERCOID SCALES By Howard McCully Divison of Systematic Biology, Stanford University, Stanford California 94305 In the ctenoid scales of most primitive percoid fishes the spines are found only on the free edge of the scale where it is not covered by another scale. Typically each spine with its base is a separate bone (a scalelet) fixed to the fiber layer of the scale. The fiber layer forms a flexible joint between adjacent scalelets. Cycloid scales in some genera, for example Rypticus and Grammistes, have homologous scalelets that lack spines; in other genera, for example Siniperca, the posterior fields lack scalelets that are homologous with the spines of ctenoid scales. Posterior growth in the scales that have spines is by increments of single scalelets. Except for rare and inconsistent specimens or species, new scalelets do not form radially to another unless at least the tip of the spine has been lost or amputated. When two spines outgrow, the one between them (the shorter and older spine) tends to lie flatter than it did and the tip is amputated by osteoclasis. Nearly always, solution pits can be seen on the end of the stump. Then a new scalelet will be laid down distal to the stump. Eventually the new one will grow until it extends beyond its neighbors and they in turn will be amputated and replaced. The fully grown scalelets with their spines stand erect or nearly so and hold [411] 412 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. up the connecting fiber layer and the overlying soft tissues of the epidermis and scale pocket. This forms a ridge usually in the arc of a circle that projects from the side of the fish. Except at the free edge, the posterior fields of such scales are covered with the amputated stumps of scalelets that once bore the spines that made the scale ctenoid. The figure shows all the steps in the life cycle of one of these marginal spines except for some intermediate sizes in the growth of the scalelet. It shows part of the free margin of the posterior field of a scale, 2.79 mm. long, from the flank of a pike-perch, Stizostedion canadense (Smith), SU-5673. It is unusual in that it shows the amputated tip of a scalelet still in position. Usually such a tip is lost soon after it is cut off. Most of the scales from this fish show 1 or 2 scalelets in this condition. It is the only fish I have yet encountered that showed any tips of amputated spines and the beginning of the new scalelets beneath them. The scalelet that has just been amputated is the marginal one that does not reach as far back as the others. The original length of this scalelet was 0.165 mm. The tip is 0.059 mm. long. The gap from which the bone was removed is 0.013 mm. wide. The primary ossification of the forming scalelet is wider than the old tip and lies beneath it. The new ossification measures 0.027 mm. in the anteroposterior diameter and is 0.045 mm. wide. Eventually the new scalelet will extend beyond the two beside it and they in their turn will be amputated and replaced. The specimen was cleaned of as much adhering soft tissue as possible, stained with Alizarin Red S., and mounted in air under a cover slip secured with a few drops of polyvinl chloride glue. The mount is so made that it dries under pres- sure and the free edge of the scale is held down. The gaps between the scalelets shrink on drying and are now narrower than they were in life. In spite of the cleaning, a layer or two of cells lies over the bony tissues in most of the figure. Williamson (1851) was the first to notice that the posterior field of perch, Perca, scales were made up of the bases of spines that were broken off. Baudelot (1873) saw that the perfect spines were only at the margins and that all spines not marginal were broken (brisée). He concluded, as had Williamson, that the spines were formed at the free edge. Neither one is able to explain how the scale- lets became broken and Bauelot says that more observations are needed. Hase (1911) studied perch, young of the year, and reached the mistaken conclusion that the posterior scalelets were formed near the nucleus and pushed out toward the margin. At the margin they then grew their spines. There was no clear ex- planation of what happened until I completed my doctoral research (McCully, 1961). From my examination of this and material from the Serranidae I conclude: 1. That the small size of the amputation gap means that only a few cells can be excreting the osteoclastic material. Vot. XXXVIII] McCULLY: MARGINAL SPINES IN CTENOID SCALES 413 Ficure 1. Posterior margin of scale from the flank of a pike-perch, Stizostedion cana- dense. (See text for explanation.) 2. Nearby cells must be protecting the bone that is not attacked. 3. The material removed from the bone may be redeposited nearby. 4. There is a regulating mechanism that can differentiate the excretory activities of a few selected cells from their neighbors. Another regulatory 414 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. change stops the unusual secretion and returns the cells, presumably, to their former state. 5. It is possible that migratory cells are the source of osteoclastic excretion. 6. There is, in a very small area, exposed to external observation and manipulation the whole of the sequence of bone growth and absorption. This type of scale offers a convenient way to test the action of drugs in an intact animal on any aspect of the physiological processes of bone growth and ab- sorption except for those peculiar to the replacement of cartilage. REFERENCES BAuDELOT, EMILE 1873. Recherches sur la structure et la développment des écailles des poissons osseux. Archives de Zoologie Expérimentale et Génerale, vol. 2, pp. 87-244 and pp. 429- 480, pls. v—xi. HAseE, ALBRECHT 1911. Die morphologische Entwicklung der Ctenoidschuppe. Anatomischer Anzeiger, 1911, vol. 40, pp. 337-356, 28 figs. McCutty, Howarp 1961. The comparative anatomy of the scales of the Serranid fishes. University Micro- films, Inc., Ann Arbor, Michigan, 248 pp. WILLIAMSON, WILLIAM CRAWFORD 1851. Investigations into the structure and development of the scales and bones of fishes. Philosophical Transactions of the Royal Society of London, 1851, pp. 643-702. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Festschrift for George Sprague Myers Vol. XXXVIII, No. 23, pp. 415-420; 3 figs.; 1 table. December 31, 1970 SIZE AND DISTRIBUTION OF PROTEINS IN ELASMOBRANCH PLASMA By Robert J. Heckly Naval Biomedical Research Laboratory, Naval Supply Center Oakland, California 94625 and Earl S. Herald Steinhart Aquarium, California Academy of Sciences Golden Gate Park, San Francisco, California 94118 As part of a project sponsored by the Office of Naval Research (N0014-67- C-0343) some members of the staff at Steinhart Aquarium embarked on an elec- trophoretic investigation of the classification value of elasmobranch plasma proteins. Samples from 11 of the 27 species available were sent to the Naval Biomedical Research Laboratory for ultracentrifuge analysis. The results show species and perhaps family differences not suspected from cellulose acetate and acrylimide gel electrophoresis. Similarities in blood constituents have been used by many investigators to show relationship between various animals. Genetic changes and natural selec- tion are believed to affect serum or plasma constituents less than the gross anat- omy or other such features. Thus, similarities in composition of blood or serum would indicate a relationship; certainly if they differed significantly it would indicate that the individuals are not closely related. Such classification of elasmo- branchs has been the subject of immunologic or electrophoretic analyses (Clem [415] 416 CALIFORNIA ACADEMY OF SCIENCES [PRroc. 4TH SER. 32 MINUTES | Ficure 1. Schlieren patterns obtained from plasma sample of 696 cm. adult male basking shark (Cetorhinus maximus). Photographs were taken at indicated times after rotor attained full speed (59,780 rpm). Numbers below major peaks in last two frames refer to observed sedimentation coefficients. and Small, 1967; Irisawa and Irisawa, 1954; Rasmussen and Rasmussen, 1967; Shuster and Goodman, 1968) but no comparative study of molecular size of serum proteins has been made. This report describes the results of some ultra- centrifugal analyses of blood plasma from several selected species. MATERIALS AND METHODS Blood was usually obtained by caudal puncture using Sequester-Sol (a dipotassiumethylenediaminetetracetate supplied by Cambridge Chemical Prod- ucts, Inc., Detroit) and for most samples, immediately centrifuged; the plasma was then removed and stored at 2° C. Analyses were made with a Spinco Model E Analytical ultracentrifuge fitted with schlieren optics using 12 mm. cells. The plasma were analyzed at the highest practical concentrations, either 1:2 or 1:4, VoL. XXXVIII] HECKLY & HERALD: ELASMOBRANCH PLASMA 417 Taste 1. Sedimentation coefficients of the principal plasma proteins. CM Total Total Common Name Scientific Name Sex Length Protein Component S Values Pacific lamprey, Lampetra tridentata F 49.5 34 8— 137° = Sevengill, Notorynchus maculatus EF 64.7 3 7 — 17 Horn shark, Heterodontus francisci M 75.7 5.3 4 7 — 17 Basking shark, Cetorhinus maximus M 696.0 = 4 7 — 18 Swell shark, Cephaloscyllium ventriosum F 94.0 3.4 45 7 — 17 Leopard shark, Triakis semifasciata F 119.3 2.75 As ee eel eli Dogfish, Squalus acanthias suckleyi F — — 4 7 — 17 Shovelnose guitarfish, Rhinobatos productus F 89.0 5.9 54 8 i2 iy Thornback ray, Platyrhinoidis triseriata F SZ 0.64 4 8 — 17 Pacific electric ray, Torpedo californica M 78.6 — 5 See ANDY = 7 Big skate, Raja binoculata (A) M 76.2 ~- 5 11 14 — Big skate, Raja binoculata (B) M 90.2 — 5 11 14 — so that minor constituents would be detected. In all instances the centrifuge was operated at 59,780 rpm. RESULTS Figure 1 shows a series of photographs taken during the sedimentation of serum from a basking shark. The first three frames clearly show the separation and sedimentation of the 18 S component. However, even after 48 minutes the 4 S component had not been completely resolved from the 7 S proteins. Since the area under the curve is proportional to the concentration of that component it is clear that the major protein in this serum had a high molecular weight. If the 7 S material is a globular protein, its molecular weight is probably in excess of 160,000. The results of ultracentrifuge analyses of plasma from a number of animals are summarized in figures 2 and 3 and the observed sedimentation coefficients are listed in table 1. The relative concentrations of the various components in each plasma can be estimated from the schlieren patterns. Except for the sevengill shark and thornback ray, the relative concentration of macroglobulin (17 S component) in shark plasma is higher than in mam- malian serum. As is evident in figure 2, the 7 S globulins in human serum have not been resolved from the albumin, but of course, continued centrifugation did separate the 7 S proteins from the albumin. Similarly, low molecular weight proteins were resolved from the 6 to 8 S components in the other plasma on pro- longed centrifugation indicating that there is at least a small amount of albumin-sized material in all of the elasmobranch plasma, even though this is not evident in the 32-minute frames shown in figures 2 and 3. In addition to indicating relative concentration, the schlieren patterns pro- vide a measure of homogeneity of the protein components in that if all mol- 418 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER. SWELL SHARK Ficure 2. Schlieren patterns obtained with various plasma 32 minutes after rotor at- tained full speed, 59,780 rpm. Numbers given below patterns indicate approximate sedimen- tation coefficients of components represented by peaks. VoL. XXXVIII] HECKLY & HERALD: ELASMOBRANCH PLASMA 419 IN) hy A Ws BIG SKATE (A) Ficure 3. Schlieren patterns obtained with various plasma 32 minutes after rotor at- tained full speed, 59,780 rpm. Numbers given below patterns indicate approximate sedimen- tation coefficients of components represented by peaks, 420 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. ecules are the same size the schlieren peak will be sharp, such as is demonstrated by the patterns for the skate and ray. A broad peak indicates either heteroge- neity of the component with respect to sedimentation velocity or a high diffusion coefficient. For instance, albumin, even though it is homogenous, would result in a rather broad peak after 32 minutes. The lamprey is unusual in that it appears to have a higher relative concen- tration of albumin (the 3.4 S component) than the elasmobranchs, and in this respect it is more like mammalian serum because, as is well known, albumin is the principal constituent in normal mammalian serum. In contrast, as is shown in figure 1, the principal protein in the basking shark was of the 7 S variety. Skates appear to lack 17 S protein in their plasma but they have high con- centrations of both the 10 and 14 S proteins. Although the patterns from dif- ferent skates do not yield identical patterns the variations that have been ob- served are shown in the last two photographs of figure 2. Since the presence or absence of a component is perhaps more significant than small differences in concentration, it is of interest that neither the lamprey nor the big skate have 17 S proteins and that they both have proteins in the 8 to 14 S range. Also of interest are the differences in the component S values between the guitarfish and the thornback. These two species were formerly placed in sep- arate families now combined in a single family (Rhinobatidae). It would seem logical that all family members would have similar sedimentation coefficients, yet in this case the shovelnose has a 12 S component which is strangely lacking in the thornback. Further study will be needed to evaluate this difference. As is shown in figures 2 and 3 and in table 1, the leopard shark, shovelnose, guitarfish, and electric ray have four distinct S components in their plasma, whereas all the other eight species studied have only three major components. With the exception of the guitarfish and thornback all species are represent- atives of individual families not closely related. LITERATURE CITED CieM, L. W., AND P. A. SMALL 1967. Phylogeny of immunoglobulin structure and function. I. Immunoglobulins of the lemon shark. Journal of Experimental Medicine, vol. 125, pp. 893-920. IrtsAwa, H., Anp A. F. IRISAWA 1954. Blood serum protein of the marine Elasmobranchii. Science, vol. 120, pp. 849-851. RASMUSSEN, L. E., anpD R. A. RASMUSSEN 1967. Comparative protein and enzyme profiles of the cerebrospinal fluid, extradural fluid, nervous tissue and sera of elasmobranchs. Chapt. 25, pp. 361-379. In Perry W. Gilbert (Ed.) Sharks, skates and rays. Baltimore, Johns Hop- kins Press, 624 pp. SHUSTER, J., AND J. W. GoopDMAN 1968. Phylogenetic studies of shark immunoglobulins. Nature, vol. 219, pp. 298-299, INDEX TO VOLUME XXXVIII FOURTH SERIES New names and principal reference in boldface type Ablepharus, 190 bivittatus, 190 bivittatus lindbergi, 170, 190 grayanus, 169, 190 pannonicus, 170, 190, 191 Acanthodactylus, 186 cantoris, 170, 186 Acanthogobius flavimanus, 207, 208, 209, 210, Blt 213, 237 Acanthogobius flavimanus in the San Fran- cisco Bay-Delta region of California, Explosive spread of the oriental goby, by Martin R. Brittan, John D. Hop- kirk, Jerrold D. Conners and Michael Martin, 207-214 (Acanthuridae), The dorsal and anal spine- locking apparatus of surgeon fishes, by James C. Tyler, 391-410 Acanthurus, 392, 393, 403 monroviae, 397 triostegus, 392, 393, 394, 395, 396, 397, 400, 402, 403, 406, 407, 408 Acerina, 255 Acestridium, 158 discus, 157, 158, 159 Acestridium discus Haseman, near Manaus, Brazil, Rediscovery of the loricariid catfish, by Robert L. Hassur, 157-162 Adinia, 292, 295 xenica, 295 Afghanistan, a checklist and key to the herpetofauna, The amphibians and reptiles of, by Alan E. Leviton and Steven C. Anderson, 163-206 Agama, 176 agilis, 167, 176 agrorensis, 168, 176 badakhshana, 168, 176 caucasica, 168, 176 erythrogastra, 168, 177 himalayana, 168, 177 himalayana himalayana, 177 lehmanni, 168, 177 megalonyx, 178 nupta, 168, 177 nuristanica, 168, 178 ruderata, 178 ruderata baluchiana, 178 ruderata megalonyx, 167, 178 scutellata, 181 tuberculata, 168, 178 versicolor, 178 Agamura, 182 femoralis, 166, 182, 205 persica, 166, 182 Agkistrodon, 201 halys, 172, 201 himalayanus, 172, 201 mokasen, 201 Ahaetulla prasina, 111, 114 Ailanthus, 2 Alcala, Angel C., see Brown, Walter C. Alosa sapidissima, 212 Alsophylax pipiens, 166, 182, 205 American characoid fishes, with special ref- erence to Probolodus heterostomus, Scale-eating, by Tyson R. Roberts, 383-390 Amiva gabbiana, 284 amphibians and reptiles of Afghanistan, a checklist and key to the herpetofauna, The, by Alan E. Leviton and Steven C. Anderson, 163-206 Amputation and replacement of marginal spines in ctenoid percoid scales, by Howard McCully, 411-414 spine-locking apparatus of fishes, (Acanthuridae), The and, by James C. Tyler, 391-410 Ancistrodon, 201, 202 halys, 201 himalayanus, 201 Anderson, Steven C., see Leviton, Alan E. Anguilla, 256 anal surgeon dorsal [421] 422 CALIFORNIA ACADEMY OF SCIENCES Anguis lumbricalis, 201 ventralis, 181 Annotated chronological bibliography of the publications of George Sprague M yers (to the end of 1969), 19-52 Anolis capito, 284 copel, 284 humilis, 279 intermedius, 279 oxylophus, 284 pachypus, 284 trochilus, 284 Ansonia mcegregori, 109 muelleri, 109, 356 Aphredoderus, 215, 226, 229, 232, 234, 235, 258, 259, 296 sayanus, 217, 229, 231, 235, 236, 239, 294, 295 Aphyocharax, 388 Aplopeltura boa, 114 Apogon, 215, 241, 244, 254, 255 astradorsatus, 217, 245 melanotaenia, 255 apogonid fishes, Some nerve patterns and their systematic significance in parac- anthopterygian, salmoniform, gobioid, and, by Warren C. Freihofer, 215-264 Aprionodon, 82 isodon, 67 Archolaemus, 267, 269, 270, 271 blax, 266, 267, 268, 270, 271 Argyropelecus, 234, 237 Argyropleura, 150 Astronotus ocellatus, 389 Astyanax, 388, 389 fasciatus, 388 Atelopus varius, 283 Aulopus, 256 Avocettina, 99 Barbourula, 117 busuangensis, 109 Basciliscus mitratus, 279 plumifrons, 284 vittatus, 284 [Proc. 4TH SER. Bassozetus, 236 Bathygobius, 237 lineatus, 217, 237, 249, 254 Bathylagus alascanus, 249 Batrachuperus, 174 mustersi, 165, 174 Batrachyperus mustersi, 174 Belonophago, 384 hutsebouti, 384 Bembrops, 377 bibliography of the publications of George Sprague Myers (to the end of 1969), Annotated chronological, 19-52 Blepharosteres grayanus, 190 Boa imperator, 284 johnii, 194 tatarica, 195 turcica, 194 Bohlke, James E., A new species of the doradid catfish genus Leptodoras, with comments on related forms, 53-62 Boiga, 195 angulata, 110, 115 cynodon, 115 dendrophila, 115 drapiezi, 115 philippina, 115 trigonata melanocephalus, 173, 195 Boreogadus, 227 saida, 217 Bothriechis nigroviridis, 285 Bothriopsis proboscideus, 285 Bothrops atrox, 285 Brachymeles, 116, 117, 121, 125, 128 bonitae, 112, 116, 117 cebuensis, 116, 117 elerae, 112, 116 gracilis, 112, 116, 117 hilong, 113, 116 pathfinderi, 112, 116 samarensis, 113, 116, 117 schadenbergi, 113, 116, 117 talinis, 113, 116 tridactylus, 110, 113, 116, 117 vermis, 116, 117 wrighti, 113, 116 Brazil, A new gymnotoid fish from the Rio Tocantins, by Maarten Korringa, 265— Dri VoL. XXXVIIT] Brazil, Rediscovery of the loricariid catfish, Acestridium discus Haseman, near Manaus, by Robert L. Hassur, 157- 162 Briggs, John C., Tropical shelf zoogeography, 131-138 Brittan, Martin R., John D. Hopkirk, Jer- rold D. Conners, and Michael Martin, Explosive spread of the oriental goby Acanthogobius flavimanus in the San Francisco Bay-Delta region of Cali- fornia, 207-214 Brotula, 252, 256, 258 clarkae, 217, 222, 223, 240 multibarbata, 222 Brotuloides emmalas, 217 Brown, Walter C., and Angel C. Alcala, The zoogeography of the herpetofauna of the Philippine Islands, a_ fringing archipelago, 105-130 Bufo, 174 agua, 283 andersonii, 165, 174 auritus, 283 biporcatus, 109 chilensis, 279 gabbi, 283 haematiticus, 279, 283 margaritifer, 278 melanochloris, 283 pleuropterus, 278 simus, 278 typhonius, 279 valliceps, 283 veraguensis, 278, 279 viridis, 165, 174 vulgaris, 174, 279 Bungarus caeruleus, 206 Bunopus, 182 tuberculatus, 166, 182 Calamaria, 111 bitorques, 114 gervaisi, 114 lumbricoidea, 114 palawanensis, 114 virgulata, 114 California, Explosive spread of the oriental goby Acanthogobius flavimanus in the INDEX 423 San Francisco Bay-Delta region of, by Martin R. Brittan, John D. Hopkirk, Jerrold D. Conners, and Michael Martin, 207-214 Callionymus, 368, 369, 370, 371, 372, 373, SE SIS, SUG, SUE decoratus, 367, 376 flagris, 366, 367, 368, 369, 372, 376 Calliophis calligaster, 115 Calliscyllium, 87, 89, 90, 93 venustum, 87, 89, 90 Calotes, 178 cristatellus, 112 marmoratus, 112 versicolor, 163, 167, 178 Carcharhinus, 67, 69, 80, 82 acronotus, 67 albimarginatus, 67 altimus, 67 amblyrhynchus, 67 borneensis, 67 cauta, 67 falciformis, 67 galapagensis, 67 leucas, 67 limbatus, 67 longimanus, 67 maculipinnis, 67 melanopterus, 67 menisorrah, 67 milberti, 67 obscurus, 67 pleurotaenia, 67 porosus, 67 remotus, 67 sorrah, 67 springeri, 67 tjutjot, 67 velox, 67 Catastoma psephotum, 285 catfish, Acestridium discus Haseman, near Manaus, Brazil, Rediscovery of the loricariid, by Robert L. Hassur, 157- 162 Catoprion, 383, 386, 387, 388 Catostomus occidentalis, 212 Caularchus, 364, 374 maeandricus, 364 Central America, On the trail of the golden 424 CALIFORNIA ACADEMY OF SCIENCES frog: with Warszewicz and Gabb in, by Jay M. Savage, 273-288 Centrotrachelus asmussi, 181 Cephaloscyllium ventriosum, 417 Cepola rubescens, 255 Ceramodactylus affinus, 186 Cerastes persicus, 202 Cetorhinus maximus, 416, 417 Chaenogaleus, 82 Chalcidolepis metallicus, 284 Chaperina fusca, 110 characid fish, Hysteronotus myersi, from Peru, A new species of glandulocau- dine, by Stanley H. Weitzman and Jamie E. Thomerson, 139-156 characoid fishes, with special reference to Probolodus heterostomus, Scale-eating American, by Tyson R. Roberts, 383- 390 Charax, 388 Chauliodus macouni, 237 Cheimarrichthys, 376, 378, 379 Cheirodon, 388 Chelopus funerus, 285 gabbil, 285 Chologaster, 258 papilliferus, 217, 229 Chrossomus neogaeus, 2 Chrysopelea paradisi, 111, 114 Cinosternum leucostomum, 285 Cleome, 2 Clevelandia ios, 211 Clupea pallasii, 250 Coelorhynchus scaphopsis, 217 Cohen, Daniel M., How many recent fishes are there?, 341-346 Coluber, 164, 195 aulicus, 197 bitorquatus, 198 blumenbachii, 199 collaris, 197 constrictor, 195 dione, 197 halys, 201 irregularis, 195 karelinii, 173, 195 lebetinus, 203 mucosus, 199 ravergieri, 173, 196 [Proc. 4TH Serr. rhodorhachis, 173, 196 schokari, 199 sibilans, 199 (Taphrometopon) lineolatus, 199 ventromaculatus, 173, 196 Compagno, L. J. V., Systematics of the genus Hemitriakis (Selachit: Carcharhini- dae), and related genera, 63-98 Coniophanes fissidens, 284 Conners, Jerrold D., see Brittan, Martin R. Contia bicolor, 197 pachyura, 285 Coregonus, 250 Cornufer, 104 Coronella taeniolata, 198 tessellata, 198 Corynopoma, 150 Corythophanes cristatus, 284 Cosymbotus platyurus, 111, 112 Cranopsis fastidiosus, 283 Crenichthys, 292, 295, 296 baileyi, 293 Crepidius epioticus, 283 Crossobamon, 182 eversmanni, 166, 182 lumsdeni, 166, 183 maynardi, 166, 183 Ctenochaetus, 392 ctenoid percoid scales, Amputation and re- placement of marginal spines in, by Howard McCully, 411-414 Ctenolucius, 383 Cursoria elegans, 194 Cyclocorus, 117, 118, 119, 128 lineatus, 114, 118 nuchalis, 359 Cyclophis aestivus, 279 persicus, 197 Cyprinodon, 292, 293 diabolis, 15 variegatus, 293 Cyrtodactylus, 164, 166, 183, 185 agusanensis, 112 annulatus, 112 caspius, 167, 183, 184 fedtschenkoi, 167, 184, 185 macularius, 185 VoL. XXXVIII) philippinicus, 112 pulchellus, 183 redimiculus, 112 russowii, 167, 184 scaber, 167, 184 watsoni, 167, 184 Dasia griffini, 113 olivaceum, 113 smaragdina, 111, 113 Dendrelaphis caudolineatus, 114 pictus, 111, 114 Dendrobates lugubris, 278 pumilio, 278 speciosus, 278 talamancae, 283 tinctorius, 283 typographus, 283 Dendrophidium melanotropis, 284 Description of a new subspecies of Rhab- dophis auriculata in the Philippines, with comments on the zoogeography of Mindanao Island, by Alan E. Levi- ton, 347-362 DeWitt, Hugh H., A revision of the fishes of the genus Notothenia from the New Zealand region, including Macquarie Island, 299-340 Dibamus argenteus, 112 Dicrolene, 237, 252, 256 intronigra, 217, 234, 235, 246 kanazawi, 217 Dinematichthys, 258 iluocoeteoides, 217, 222 Diplogrammus goramensis, 367 Dirrhizodon, 82 doradid catfish genus Leptodoras, with com- ments on related forms, A new species of, by James E. Bohlke, 53-62 Dormitator, 259 Dorosoma petenensis, 212 Dorsal and anal spine-locking apparatus of surgeon fishes (Acanthuridae), The, by James C. Tyler, 391-410 Draco bimaculatus, 112 INDEX 42 Un everetti, 112 mindanensis, 112 ornatus, 112 quadrisi, 112 rizali, 112 volans, 111, 112 Draconetta, 365, 367, 368, 369, 370, 371, 372, 373, 374, 375, 376, 377 acanthopoma, 366, 367, 368, 369, 372, RY APY LMSW is) oregona, 373 Draculo, 378 Drymobius boddaertii, 284 Dryocalamus subannulatus, 114 tristrigatus, 114 Dryophiops philippina, 114 Earle, Sylvia A., see Mead, Giles W. Echis, 202 carinatus, 172, 202 Eigenmannia, 269, 270, 271 conirostris, 271 macrops, 271 virescens, 270 Eirenis, 197 persica, 172, 197 Elaphe, 197 bairdii, 15 dione, 173, 197 erythrura, 111, 114 parreysli, 197 Elaps circinalis, 285 elasmobranch plasma, Size and distribution of proteins in, by Robert J. Heckly and Earl S. Herald, 415-420 Elassoma zonatum, 295 Eleginus, 227 gracilus, 217 Eleotris, 253 fuscus, 217, 253, 254 Emoia atrocostata, 111, 113 caeruleocauda, 113 ruficauda, 113 Empetrichthys, 295, 296 Enneacanthus, 2 Epigeichthys, 251 Epigonus robustus, 244 426 CALIFORNIA ACADEMY OF SCIENCES Eremias, 187 acutirostris, 171, 187 aporosceles, 169, 170, 187 aie 7 ile 137 fasciata, 170, 187 grammica, 171, 187 guttulata, 188 gcuttulata watsonana, 171, 188 intermedia, 171, 188 lineolata, 170, 188 (Mesalina) watsonana, 188 nigrocellata, 171, 188 persica, 189 regeli, 170, 189 (Rhabderemias) scripta, 189 (Scapteira) acutirostris, 187 (Scapteira) aporosceles, 187 (Scapteira) grammica, 187 scripta, 170, 189 velox, 189 velox persica, 171, 189 velox velox, 171, 189 Eridacnis, 87, 88, 89, 90, 91, 92, 93 alcocki, 90, 93 barbouri, 67, 72, 90, 93 radcliffei, 67, 90, 93 sinuans, 65, 67, 90, 93 Eristicophis, 202 macmahoni, 172, 202 Erythrolamprus venustissimus, 284 Eryx, 164, 171, 194 elegans, 171, 194 jaculus, 171, 195 jaculus czarewskul, 194 johnii, 172, 194, 195 miliaris, 171, 172, 194 persicus, 195 tataricus, 172, 195 Etheostoma, 294, 295, 296 asprigene, 294 radiosum, 294, 295 spectabile, 294, 295 Eublepharis, 185 hardwickii, 185 macularius, 165, 185 Eucynopotamus, 388 Eugaleus, 75 Eugnathichthys, 384 Eumeces, 191 blythianus, 169, 191, 192 [Proc. 4TH SEr. himalayanus, 193 schneideri, 169, 191, 192 schneideri princeps, 191 schneideri zarudnyi, 191 taeniolatus, 169, 192 Euprepes dissimilis, 192 Eurylepis taeniolatus, 192 Eutyx diagrammus, 217 Exodon, 383, 387, 388 paradoxus, 389 Explosive spread of the oriental goby Acanthogobius flavimanus in the San Francisco Bay-Delta region of Cali- fornia, by Martin R. Brittan, John D. Hopkirk, Jerrold D. Conners, and Michael Martin, 207-214 Farlowella, 157, 158, 159, 161 Frethofer, Warren C., Some nerve patterns and their systematic significance in paracantho pterygian, salmoniform, go- bioid, and apogonid fishes, 215-264 Fundulus, 2, 292, 293, 295, 296 cingulatus, 292, 293 grandis, 292, 293 heteroclitus, 292, 293 kansae, 292, 293, 295 majalis, 292, 293 notatus, 292, 293, 295 olivaceus, 292, 293, 295 parvipinnis, 293, 295, 296 pulvereus, 293 seminolis, 292 similis, 292, 293 zebrinus, 292, 293, 295 Furgaleus, 67, 80, 85 ventralis, 79 Gabb in Central America, On the trail of the golden frog: with Warszewics and, by Jay M. Savage, 273-288 Gadus, 227 magellanicus, 303, 310, 311, 334 marmoratus, 250 Galaxias, 250 Galeichthys felis, 384 Galeocerdo, 82 cuvier, 67 Vor. XXXVIIT] Galeorhinus, 75, 76, 80, 81, 83, 85, 86, 93 australis, 67, 76, 81, 83, 93 chilensis, 67, 76, 81, 83, 93 (Galeorhinus), 83 (Galeorhinus) galeus, 83 galeus, 67, 76, 81, 83, 93 (Hypogaleus) zanzibariensis, 83 hyugaensis, 83, 85 japanicus, 76, 83 molinae, 83 omanensis, 67, 83, 85, 93 vitaminicus, 83, 93 zanzibariensis, 83, 85 zyopterus, 64, 67, 71, 84 Galeus, 75 japanicus, 76, 93 Gammarus tigrinus, 214 Gasterosteus aculeatus, 210, 294, 295 Gastrotheca, 281 Gehyra mutilata, 111, 112 Gekko athymus, 112 gecko, 111, 112 mindorensis, 112 monarchus, 112 palawanensis, 112 tuberculosus, 185 George Sprague Myers, On the natural his- tory of, by Lionel A. Walford, 1-18 George Sprague Myers (to the end of 1969), Annotated chronological bibliography of the publications of, 19-52 Gephyrocharax, 150 Gerrhonotus fulvus, 284 Gillichthys mirabilis, 211 Glandulocauda, 150 glandulocaudine characid fish, Hysteronotus myersi, from Peru, A new species of, by Stanley H. Weitzman and Jamie E.. Thomerson, 139-156 Glauconia blanfordii, 201 Gnathophysia ocellaba, 284 Gobiesociformes, A reinterpretation of the teleostean fish order, by William A. Gosline, 363-382 Gobiesox, 364, 370, 371, 372, 373, 374, 375, 376, 377, 379 maeandricus, 364 nigripinnis, 366, 367, 371, 372 13, TO, Us Biky BB, INDEX 427 gobioid, and apogonid fishes, Some nerve patterns and their systematic signifi- cance in paracanthopterygian, sal- moniform, by Warren C. Frethofer, 215-264 Gonostoma elongatum, 237, 250 Gonyocephalus interruptus, 112 semperi, 112 sophiae, 112 Gonyosoma oxycephala, 114 Gosline, William A., A reinterpretation of the teleostean fish order Gobiesoci- formes, 363-382 Grammistes, 411 Gunnellichthys, 259 Gymnodactylus caspius, 183 eversmanni, 182 fedtschenkoi, 184 longipes, 184 persicus, 182 russowli, 184 watsoni, 184 gymnotoid fish from the Rio Tocantins, Brazil, A new, by Maarten Korringa, 265-272 Hadropterus, 294 scierus, 295 Halys himalayanus, 201 Harpagifer, 367, 369, 370 bispinus, 366 Harpodon, 256 Hassar, 53, 61 notospilus, 61 Hassur, Robert L., Rediscovery of the lori- cariid catfish, Acestridium discus Hase- man, near Manaus, Brazil, 157-162 Heckly, Robert J., and Earl S. Herald, Size and distribution of proteins in elas- mobranch plasma, 415-420 Hemidactylus, 185 flaviviridis, 166, 185 frenatus, 111, 112 garnoti, 112 luzonensis, 112 Hemigaleus, 80, 82 balfouri, 67 428 CALIFORNIA ACADEMY OF SCIENCES macrostoma, 67 microstoma, 67 pectoralis, 67 tengi, 67 Hemiphyllodactylus typus, 110, 111, 112 Hemipristis, 82 elongatus, 67 Hemitriakis, 63, 65, 67, 73, 75, 76, 79, 80, Si 22 88, 8B, Teeey, BS, So. Oil, Op. 93 jJapanica, 63, 64, 65, 68, 72, 75, 76, 77, 78, 79, 80, 83, 84, 92, 93 leucoperiptera, 63, 65, 75, 76, 78, 80, 87, 93 Hemitriakis (Selachit: Carcharhinidae), and related genera, Systematics of the genus, by L. J. V. Compagno, 63-98 Herald, Earl S., see Heckly, Robert J. Herpetodryas carinatus, 284 herpetofauna of the Philippine Islands, a fringing archipelago, The zoogeogra- phy of the, by Walter C. Brown and Angel C. Alcala, 105-130 Heterodon diadema, 198 Heterodontus francisci, 417 Heterogaleus, 82 Holobrycon, 388 Hologerrhum, 117, 118, 119, 128 philippinum, 114, 118 Hopkirk, John D., see Brittan, Martin R. Hoplostethus pacificus, 217 How many recent fishes are Daniel M. Cohen, 341-346 Hubbs, Clark, Teleost hybridization studies, 289-298 Hurria rychops, 114 hybridization studies, Hubbs, 289-298 Hydrosaurus pustulosus, 112 Hydrus piscator, 206 Hyla andersonii, 2, 3 elaeochroa, 283 gabbii, 283 molitor, 278, 279 molitor marmorata, 278, 279 there?, by Teleost, by Clark nigripes, 283 pugnax, 278 punctariola, 279 punctariola monticola, 284 [Proc. 4TH SER. punctariola pictipes, 283 sordida, 279 splendens, 278, 279, 281 uranochroa, 283 Hylaemorphus bibronii, 278 dumerilii, 278 Hylodes cerasinus, 284 fitzingeri, 278 Hypogaleus, 83, 85, 86, 93 (Hypogaleus), 83 hyugaensis, 64, 83, 85, 93 zanzibariensis, 64, 71, 84, 85, 93 Hypomesus olidus, 217 pretiosus, 217, 237, 246, 247 Hypopomus, 269, 270 brevirostris, 271 Hypoprion, 82 hemiodon, 67 macloti, 67 signata, 67 Hysterocarpus, 212 traski, 212 Hysteronotus, 139, 147, 150, 154 hesperus, 139, 144, 145, 146, 147, 149 150, 152, 154 megalostomus, 139, 145, 147, 148, 149 150, 151, 152, 153, 154 myersi, 139, 140, 141, 142, 143, 144 145, 146, 147, 149, 150, 151, 152, 153 Hysteronotus myersi, from Peru, A new spe- cies of glandulocaudine characid fish, by Stanley H. Weitzman, 139-156 ) , ’ Ichthyophis monochrous, 109 Ictalurus catus, 212 punctatus, 212 Iguana rhinophila, 284 Ilypnus gilberti, 211 Isogomphodon, 82 oxyrhynchus, 67 Ixalus warschewitschii, 278 Jordanella, 292, 293 floridae, 293 VoL. XXXVIITII Kalophrynus pleurostigma, 110 Kaloula baleata, 110 conjuncta, 110 picta, 110 rigida, 110 Korringa, Maarten, A new gymnotoid fish from the Rio Tocantins, Brazil, 265- 272 Labichthys, 99 Labidesthes sicculus, 294 Lacerta agama, 176 apoda, 182 aurata, 192 boskiana, 186 calotes, 178 grammica, 187 guttata, 179 mabouya, 192 muralis viridis, 279 mystacea, 180 pipiens, 205 varia, 193 variabilis, 187 velox, 189 Lachesis stenophrys, 285 Laemonema barbatulum, 217 Lamiopsis, 82 temmincki, 67 Lampanyctus, 241, 244, 256 leucopsarus, 215, 238, 243 Lampetra tridentata, 417 Lamprogrammis niger, 217 Landonia, 150 Lavinia exilicauda, 211 Leiuperus sagittifer, 278 Lepidodactylus aureolineatus, 112 christiani, 110, 112 herrei, 112 lugubris, 110, 112 naujanensis, 112 planicaudus, 112 Lepidogobius lepidus, 211 Lepomis macrochirus, 210, 294 Lepophidium, 222, 226, 252 prorates, 217, 218, 219 INDEX 429 Leporinus, 389 fasciatus, 389 friderici, 389 Leptobrachium hasselti, 109 Leptocharias, 67, 69, 80, 86 smithii, 67 Leptocottus armatus, 211 Leptodoras, 53, 59, 61 acipenserinus, 54, 59, 60, 61 juruensis, 53, 54, 58, 59, 60, 61 linnelli, 53, 54, 58, 59, 60, 61 myersi, 53, 54, 55, 56, 57, 59, 60, 61 Leptodoras, with comments on related forms, A new species of the doradid catfish genus, by James E. Bohlke, 53-62 Leptognathus argus, 284 nebulata, 284 pictiventris, 284 Leptophis aeruginosus, 284 praestans, 284 saturatus, 284 Leptotyphlops, 200 blanfordi, 171, 201 Leviton, Alan E., Description of a new sub- Species of Rhabdophis auriculata in the Philippines, with comments on the zoogeography of Mindanao Island, 347-362 Leviton, Alan E., and Steven C. Anderson, The amphibians and reptiles of Af- ghanistan, a checklist and key to the herpetofauna, 163-206 Lionurus gibber, 227 Liopeltis philippinus, 114 tricolor, 114 Lithodytes gulosus, 284 habenatus, 284 megalocephalus, 284 melanostictus, 284 muricinus, 284 podiciferus, 284 loricariid catfish, Acestridium discus Hase- man, near Manaus, Brazil, Rediscovery of the, by Robert L. Hassur, 157-162 Lota, 303 430 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. breviuscula, 303 Mabuia magellanica, 303 alliacea, 284 Loxodon, 67, 82 cepedei, 284 macrorhinus, 67 Mabuya, 192 Lucania, 292, 293, 295 aurata, 170, 192 parva, 213, 293, 295 bontocensis, 113 Luperosaurus, 117, 118, 128 dissimilis, 170, 192 amissus, 118 multicarinata, 111, 113 cumingi, 110, 112, 118 multifasciata, 111, 113 joloensis, 112, 118 Macquarie Island, A revision of the fishes megregori, 118 of the genus Notothenia from the New Lycodapus, 253 Zealand region, including, by Hugh H. Lycodon, 197 DeWitt, 299-340 aulicus, 111, 114 Macrobrachium brazilense, 146 dumerili, 114 Macronotothen rossii, 312 mulleri, 114 Macrurus, 227 striatus bicolor, 173, 197 Martin, Michael, see Brittan, Martin R. subcinctus, 114 Maticora intestinalis, 115 tesselatus, 114 McCully, Howard, Amputation and replace- Lygosoma (Leiolopisma) ment of marginal spines in ctenoid auriculatum, 113 percoid scales, 411-414 pulchellum, 113 Mead, Giles W., and Sylvia A. Earle, Notes quadrivittatum, 111, 113 on the natural history of snipe eels, rabori, 113 99-104 semperi, 113 Megrophrys monticola, 109 subvittatum, 113 Melamphaes, 217, 244, 250 vulcanium, 113 Menidia, 250, 294 zamboangensis, 113 audens, 293, 294 Lygosoma (Lygosoma) Merluccius, 252, 257 chalcides, 113 gayi, 217, 227, 235, 246 Lygosoma (Sphenomorphus), 113 productus, 217 acutum, 113 Metynnis, 383 arborens, 113 Micrixalus mariae, 109 atrigularis, 113 Microgadus, 227 coxi, 113 proximus, 217, 228 decipiens, 113 Mindanao Island, Description of a new sub- diwati, 113 species of Rhabdophis auriculata in fasciatum, 113 the Philippines, with comments on jagori, 113 the zoogeography of, by Alan E. luzonensis, 113 Leviton, 347-362 mindanensis, 113 Mocoa assata, 284 palawanensis, 113 Monomerepus, 217, 236, 252, 256 steerei, 113 Monomitopus, 236, 252, 256 stejnegeri, 113 Moxostoma poecilurum, 294 varigatum, 113 Musgiloides, 367 wrighti, 113 Mustelus, 67, 71, 79, 81, 86, $7, 88, 89, 91, 93 Lytorhynchus, 198 antarcticus, 67 maynardi, 173, 198 asterias, 67 ridgewayi, 173, 198 californicus, 67, 88 Mabouia blythiana, 191 canis, 67, 81, 88 Vor. XXXVIIIJ dorsalis, 67, 88 fasciatus, 67 griseus, 67, 88 henlei, 67, 72, 87, 88 higmani, 67, 88 kanekonis, 67 lenticulatus, 67 lunulatus, 67, 88 manazo, 67, 88 megalopterus, 88 mento, 67 mustelus, 67, 88 natalensis, 88 nigropunctatus, 88 norrisi, 67 schmitti, 67 Myersophis alpetris, 114 Naja, 200 lutescens, 200 naja, 115 oxiana, 172, 200 Naso, 391, 392, 393, 397, 403, 406, 407, 408 literatus, 393, 403, 404, 405, 406, 407 Natrix, 198 auriculata, 114, 349, 356 chrysarga, 114 dendrophiops, 114 lineata, 114 piscator, 206 spilogaster, 114 tessellata, 173, 198 tessellata tessellata, 198 torquatus, 198 Negaprion, 80, 82 acutidens, 67 brevirostris, 67 forsteri, 67 fronto, 67 Negogaleus, 82 Nemacheilus, 256 Nemichthys, 99, 102 scolopaceus, 101, 102 Neoscopelus, 244 Neotriakis, 87, 89, 90 sinuans, 87, 90 nerve patterns and their systematic signifi- in paracanthopterygian, sal- gobioid, cance moniform, and apogonid INDEX 431 fishes, Some, by Warren C. Frethofer, 215-264 new species of glandulocaudine charcid fish, Hysteronotus myersi, from Peru, A, by Stanley H. Weitzman and Jamie E. Thomerson, 139-156 new species of the doradid catfish genus Leptodoras, with comments on re- lated forms, by James E. Bohlke, 53-62 Zealand region, including Macquarie Island, A revision of the fishes of the genus Notothenia, by Hugh H. De- Witt, 299-340 Notemigonus crysoleucas, 294 Notes on the natural history of snipe eels, by Giles W. Mead and Sylvia A. Earle, 99-104 Notorynchus maculatus, 417 Notothenia, 299, 300, 302, 307, 310, 325 angustata, 300, 303, 307, 310, 318, 319, Be SE GIS, BAS. SA, Soll, so! antarctica, 303, 311, 312 antarcticus, 303 arguta, 303, 304, 311, 313 colbecki, 323, 324, 325, 326, 327, 328, 331, 332 corliceps, 299) 302; 307, 3115 3135318; 322 corliceps macquariensis, 313, 314, 317 cornucola, 300, 318 filholi, 323, 324, 325, 326, 327, 328, 331, 332 hassleriana, 303, 304, 311, 313 latifrons, 318, 324 macrocephala, 303, 304, 310, 313, 318 macrocephala marmorata, 313 macrocephalus, 303 magellanica, 300, 303, 307, 308, 309, 310, Bldeysi2, Siz, 315, 3222 323ho248 325 magellanicus, 303 magellanicuz, 303 maoriensis, 303, 311, 325 marmorata, 312, 316 microlepidota, 302, 310, 318, 323, 324, 325, 326, 327, 328, 331, 332 parva, 318, 324, 326, 327 patagonica, 318, 321, 326, 327 porteri, 310, 318, 322, 323 New 323) 432 CALIFORNIA ACADEMY OF SCIENCES rossi, 312 LOSsil,, GO2, 307, a2) SIA esiGs Sili7, 322 rossii marmorata, 314, 316, 317 rossii rossil, 314, 317 Notothenia from the New Zealand region, including Macquarie Island, A_ re- vision of the fishes of the genus, by Hugh H. DeWitt, 299-304 Nototoenia filholi, 325 Notropis cornutus, 294 cummingsi, 10 umbratilis, 294 Odontaspis taurus, 73 Oedemognathus, 389 exodon, 388 Oedipus moro, 283 Ogilbia, 222, 252, 258 ventralis, 220, 221 Oligodon, 198 ancorus, 114 arnensis, 206 maculatus, 115 modestum, 115 taeniolatus, 172, 198 vertebralis, 115 Ollotis coerulescens, 283 On the natural history of George Sprague Myers, by Lionel A. Walford, 1-18 On the trail of the golden frog: with Wars- zewicz and Gabb in Central America, by Jay M. Savage, 273-288 Oncorhynchus, 250 Ooeidozyga diminutiva, 109 laevis, 109, 111 Opheobatrachus vermicularis, 283 Ophichthys gomesii, 254 Ophiomorus, 193 brevipes, 169, 193 miliaris, 193 tridactylus, 169, 193 Ophiophagus hanna, 115 Ophisaurus, 181 apodus, 165, 182 Ophisops, 190 elegans, 190 jerdoni, 170, 190 [Proc. 4TH SER. Ophisthotropis alcalai, 115 typica, 115 Opsodoras, 53, 61 leporhinus, 61 linnelli, 61 notospilus, 61 Opsopoedeus emiliae, 294 Oreophryne, 355 annulata, 110 Orthodon microlepidotus, 211 Osmerus eperlanus, 255 Otosaurus cumingl, 113 Oxyrhabdium, 117, 118, 119, 128 leporinum, 115, 118 modestum, 115, 118 Oxyrrhopus petola, 284 plumbeus, 284 Palaemon macrodactylus, 213 paracanthopterygian, salmoniform, gobioid, and apogonid fishes, Some nerve pat- terns and their systematic significance in, by Warren C. Freihofer, 215-264 Paracanthurus, 392 Paragaleus, 82 Parapercis, 367, 370 cephalopunctata, 366 Parupeneus, 256 Parupygus, 270 savanensis, 271 Pelamis bicolor, 279 Pelophryne albotaeniata, 109 brevipes, 110 lighti, 110 Perca, 412 Percina caprodes, 294 percoid scales, Amputation and replacement of marginal spines in ctenoid, by Howard McCully, 411-414 Percopsis, 215, 229, 232, 234, 235, 236, 237, 238, 241, 244, 246, 251, 252, 254, 256, 257, 258, 260 omiscomaycus, 217, 230, 240, 241, 242 transmontana, 217 Periops parallelus schirazana, 200 VoL. XX XVIII] Perochirus, 117 ateles, 112 Peru, A new species of glandulocaudine characid fish, WHysteronotus myersi, from, by Stanley H. Weitzman and Jamie E. Thomerson, 139-156 Phago, 384 boulengeri, 384 Philautus acutirostris, 109 bimaculatus, 109 leitensis, 109 longicrus, 109 pictus, 109 schmackeri, 109 spinosus, 109 williamsi, 109 Philippine Islands, a fringing archipelago, The zoogeography of the herpetofauna of the, by Walter C. Brown and Angel C. Alcala, 105-130 Philippines, with comments on the zoogeog- raphy of Mindanao Island, Descrip- tion of a new subspecies of Rhab- dophis auriculata im the, by Alan E. Leviton, 347-362 Phirix pachydermus, 278 Phoxinus laevis, 250 Phrynocephalus, 179 clarkorum, 169, 179, 180 euptilopus, 169, 179 interscapularis, 169, 179 luteoguttatus, 168, 179 maculatus, 169, 180 mystaceus, 168, 180 ornatus, 169, 179, 180 raddei boettgeri, 181 reticulatus, 168 reticulatus boettgeri, 169, 181 scutellatus, 169, 181 tickelii, 181 Phyllobates hylaeformis, 284 Phyllomedusa hypochondrica, 279 Physiculus, 222, 226, 252, 258 talarae, 217, 224, 225, 238 plasma, Size and distribution of proteins in elasmobranch, by Robert J. Heckly and Earl S. Herald, 415-410 Platymantis, 104, 116, 117, 121, 128 cornutus, 109, 116 INDEX 433 corrugatus, 109, 116 dorsalis, 109, 116 guentheri, 109, 116 hazelae, 109, 116, 117 ingeri, 109, 116 polillensis, 116, 117 subterrestris, 109, 116 Platyrhinoidis triseriata, 417 Podarces (Eremias) intermedia, 188 (Scapteira) scripta, 189 Pogonichthys, 212 macrolepidotus, 211 Pogonymus, 372, 373, 376, 377 pogognathus, 367 Pomoxis, 210 Porichthys, 237, 251, 252, 258 margaritatus, 217, 232, 233, 236 Porogadus, 236, 252, 256 Prionace, 82 glauca, 67 Prionurus, 392 Probolodus, 383, 384, 388 heterostomus, 383 Probolodus heterostomus, Scale-eating Amer- ican characoid fishes, with special reference to, by Tyson R. Roberts, 383-390 Prolatilus, 367 Proscyllium, 88, 89, 90, 91, 92, 93 alcocki, 90 habereri, 67, 71, 72, 89, 90, 93 venustum, 90, 93 proteins in elasmobranch plasma, Size and distribution of, by Robert J. Heckly and Earl S. Herald, 415-420 Psammodynastes pulverulentus, 111, 115 Psammophis, 199 cerasogaster, 206 leithi, 173, 199, 206 lineolatus, 173, 199 schokari, 173, 199 Psammosaurus caspius, 194 Pseudoboa carinata, 202 Pseudocerastes, 202 bicornis, 203 persicus, 172, 202 persicus persicus, 202 Pseudocorynopoma, 150, 151, 154 doriae, 150, 151 434 CALIFORNIA ACADEMY OF SCIENCES Pseudogekko, 117, 118, 119, 128 brevipes, 112, 118 compressicorpus, 112, 118 Pseudorabdion ater, 115 mcnamatae, 115 montanum, 115 oxycephalum, 115 taylori, 115 Ptyas, 199 mucosus, 173, 199 Ptychozoon intermedia, 112 Puntius, 360 binotatus, 360 Pygocentrus, 386 Python reticulatus, 111, 114 Raja binoculata, 417 Rana, 175 bufo, 174 cancrivora, 109, 111 cyanophlyctis, 165, 175 erythraea, 109 everetti, 109 leytensis, 109 limnocharis, 109, 111 magna, 109, 356 microdisca, 109 nicobariensis, 109 ridibunda, 165, 175 ridibunda ridibunda, 175 sanguinea, 109 signata, 109 sternosignata, 165, 175 temporaria, 175 woodworthi, 109 Ranula brevipalmata, 284 Rediscovery of the loricariid catfish, Ace- stridium discus Haseman, near Ma- naus, Brazil, by Robert L. Hassur, 157-162 Reinter pretation of the teleostean fish order Gobiesociformes, A, by William A. Gosline, 363-382 Replacement of marginal spines in ctenoid percoid scales, Amputation and, by Howard McCully, 411-414 reptiles of Afghanistan, a checklist and key to the herpetofauna, The amphibians [Proc. 4TH Ser. and, by Alan E. Leviton and Steven C. Anderson, 163-206 revision of the fishes of the genus Noto- thenia from the New Zealand region, including Macquarie Island, A, by Hugh H. DeWitt, 299-340 Rhabdolichops, 270, 271 longicaudatus, 271 Rhabdophis auriculata, 347, 348, 349, 350, 353, 356, 357, 358, 359 auriculata auriculata, 355, 356, 358 auriculata myersi, 349, 351, 352, 354, S55 SO, SS sarawacensis, 358 Rhabdophis auriculata zm the Philippines, with comments on the zoogeography of Mindanao Island, Description of a new subspecies of, by Alan E. Leviton, 347-362 Rhacophorus appendiculatus, 109 emembranatus, 109 everetti, 109 leucomystax, 109, 111 lissobrachius, 109 pardalis, 109 surdus, 109 Rhadinaea decorata, 284 Rhinobatos productus, 417 Rhinotyphlops albirostris, 279 Rhizoprionodon, 67, 82 acutus, 67 lalandei, 67 longurio, 67 oligolinx, 67 porosus, 67 terraenovae, 67 Rhytiodus microlepis, 54 Rivulus, 292 peruanus, 146 Roberts, Tyson R., Scale-eating American characoid fishes, with special refer- ence to Probolodus heterostomus, 383— 390 Roccus, 212 saxatilis, 212 Roeboexodon, 383, 387, 388 Roeboides, 383, 387, 388 guatemalensis, 387, 389 VoL. XXXVIIT] myersi, 387 occidentalis, 383, 388 prognathus, 387, 388 Rypticus, 411 Salamandrella sinensis, 174 Salmo, 250 salmoniform, gobioid, and apogonid fishes, Some nerve patterns and their system- atic significance in paracanthoptery- gian, by Warren C. Freihofer, 215-264 San Francisco Bay-Delta region of Cali- fornia, Explosive spread of the orien- tal goby Acanthogobius flavimanus in the, by Martin R. Brittan, John D. Hopkirk, Jerrold D. Conners, and Michael Martin, 207-214 Sardinioides, 255 Savage, Jay M., On the trail of the golden frog: with Warszewicz and Gabb in Central America, 273-288 Scale-eating American characoid fishes, with special reference to Probolodus hetero- stomus, by Tyson R. Roberts, 383-390 scales, Amputation and replacement of mar- ginal spines in ctenoid percoid, by Howard McCully, 411-414 Scapteira acutirostris, 187 aporosceles, 187 lineolata, 188 Scincella, 193 himalayana, 170, 193 Scincus lateralis, 193 pavimentatus, 191 schneideri, 191 Scoliodon, 67, 82 laticaudus, 67 Scopelengys, 244, 254, 256 tristis, 217, 244, 248 Scorpaenichthys, 251 Scylliogaleus, 86, 87 quecketti, 65, 67 Scylliorhinus, 89 Scyllium, 89 (Proscyllium) habereri, 89 Sergestes, 102 (Sergestes) arcticus, 101, 102 INDEX 435 Serrasalmus, 388 elongatus, 384, 386, 388 Sibon annulatum, 284 Sibynophis bivittatus, 115 Sicyases, 374 Siniperca, 411 Siphonops mexicanus, 283 proximus, 283 Size and distribution of proteins in elasmo- branch plasma, by Robert J. Heckly and Earl S. Herald, 415-420 snipe eels, Notes on the natural history of, by Giles W. Mead and Sylvia A. Earle, 99-104 nerve patterns and their systematic significance in paracanthopterygian, salmoniform, gobioid, and apogonid fishes, by Warren C. Freihofer, 215- 264 Spalerosophis, 200 diadema, 172 diadema schirazana, 200 microlepis, 200 Sphaerodactylus glaucus, 284 Sphargis coriacea, 285 Sphenocephalus tridactylus, 193 Sphryaena argentea, 4 ensis, 4 idiastes, 4 Spilotes chrysobronchus, 284 corias, 284 spine-locking apparatus of surgeon fishes (Acanthuridae), The dorsal and anal, by James C. Tyler, 391-410 Spirinchus thaleichthys, 249 Squalus acanthias suckleyi, 417 Staurois natator, 109 Steatogenys, 269, 270 elegans, 271 Stegonotus miilleri, 115 Stellio agrorensis, 176 caucasicus, 176 Some erythrogaster, 177 himalayanus, 177 lehmanni, 177 spinipes, 181 436 CALIFORNIA ACADEMY OF SCIENCES Stenodactylus lumsdeni, 183 maynardi, 183 orientalis, 183 scaber, 184 scincus, 186 Stenorhina ventralis, 285 Stenostoma albifrons, 279 Sternopygus, 265, 269, 270, 271 macrurus, 269, 270 Stizostedion canadense, 412, 413 Strabomantis biporcatus, 279 surgeon fishes (Acanthuridae), The dorsal and anal spine-locking apparatus of, by James C. Tyler, 391-410 Synagrops, 244 bella, 244 Synchiropus, 368 Synodus, 256 Systematics of the genus Hemitriakis (Se- lachii: Carcharhinidae), and _ related genera, by L. J. V. Compagno, 63-98 Teleost hybridization Clark Hubbs, 289-398 teleostean fish order Gobiesociformes, A re- interpretation of the, by William A. Gosline, 363-382 Teleurapsis schlegelii, 285 Teratoscincus, 185 bedriagai, 166, 186 keyserlingi, 185 microlepis, 166, 186 scincus, 166, 186 Testudo, 175 graeca, 175 horsfieldii, 165, 176 Thaleichthys pacificus, 249 Thecadactylus rapicaudus, 284 Thomerson, Jamie E., see Weitzman, Stanley 181, Tomicodon, 369 Tomyris oxiana, 200 Torpedo californica, 417 Trachelochismus, 370 pinnulatus, 367 Trapelus megalonyx, 178 Triaenodon, 80, 82 obesus, 67, 80 Triakis, 63, 67, 81, 87, 88, 89, 90, 91, 93 studies, by [Proc. 4TH SrEr. acutipinna, 67, 88 attenuata, 87, 88, 91, 93 barbouri, 87, 90 fehlmanni, 67, 88, 90, 91, 93 habereri, 90 henlei, 87 maculata, 67, 81, 88 radcliffei, 90 scyllia, 67, 81, 87, 88 semifasciata, 67, 71, 81, 87, 88, 92, 417 sinuans, 90 venusta, 89, 90 Tridentiger trigonocephalus, 213 Trimeresurus flavomaculatus, 115 schultzei, 115 wagleri, 115 Tropical shelf zoogeography, by John C. Briggs, 131-138 Tropidonotus auriculatus, 349, 356 piscator, 206 Tropidophorus, 113 grayi, 113 leucospilos, 113 misaminus, 113 partelloi, 113 Tupinambis bengalensis, 193 Tyler, James C., The dorsal and anal spine- locking apparatus of surgeon fishes (Acanthuridae) , 391-410 Typhlops, 201 braminae, 111, 114 canlaonensis, 114 cumingi, 110, 114 dendrophis, 114 jagori, 114 longicauda, 114 luzonensis, 114 mindanensis, 114 nigricans, 200 ruber, 114 ruficauda, 114 rugosa, 114 vermicularis, 171, 201 Umbra, 2 Uromastyx, 181 asmussi, 167, 181 hardwickii, 167, 181 VoL. XXXVIII] Urophycis floridanus, 217 Varanus, 193 bengalensis, 169, 193 bengalensis bengalensis, 193 griseus, 169 griseus caspius, 194 griseus koniecznyi, 194 (Indovaranus) 193 (Psammosaurus) griseus caspius, 194 salvator, 112 Vipera, 203 lebetina, 172, 203 redi, 203 bengalensis bengalensis, Walford, Lionel A., On the natural history of George Sprague Myers, 1-18 Warszewicz and Gabb in Central America, On the trail of the golden frog: with, by Jay M. Savage, 273-288 Watasea sivicola, 217 Weitzman, Stanley H., and Jamie E. Thom- erson, A new species of glandulocau- dine characid fish, Hysteronotus myersi, from Peru, 139-156 INDEX 437 Xenochrophis piscator, 173, 200, 206 Xenodon angustirostris, 284 Xenopeltis unicolor, 114 Xesurus, 392 Xiphosoma annulatum, 284 Yerutia, 368 Zalieutes elater, 217 Zamenis rhodorachis, 196 Zaocys carinatus, 115 luzonensis, 115 Zebrasoma, 392 soogeography of Mindanao Island, Descrip- tion of a new subspecies of Rhab- dophis auriculata in the Philippines, with comments on the, by Alan E. Leviton, 347-362 zoogeography of the herpetofauna of the Philippine Islands, a fringing archi- pelago, The, by Walter C. Brown and Angel C. Alcala, 105-130 Zygonectes, 295 : uu LF 1 a ie es 7 hic he 4 4 2 ~~) ane yy i P y J r, , ¥ As me {) { is Mi ve. an nN at ath ua ay r ‘ae | ve : - } i ‘ie ! ‘ . La | | way ‘ i ; f == ue. x \ s] : ~ f | si i { by “S. as i “Lis 7 4 “ es aay ya ‘oy a ee at) nit ay 7 e 4 7 “ i + at 105 | { iad iA i) coe é. ” a is H Ae eee yr ie Oh ce i Vere erten ce Mae ee ait r eed cP Me CF aif! ats ihe Stiogstars as as Ae sr figs wited Tie ehh Mae 20e SOMME A Sepa Mi ee os pe pid pe ten PX, Se, een ah by Siletk eee A aed ed Neha ORT WA Ea NELS Oe OFA Ok SY manta " BRS MTA SANS Ora: My, ls =hyh ee te) we Hen Gas oe YO le gd het as 4 # bev, ee es red j Siar , wh wb KY ae Taselitace gun 0 Sisson’ ve Rohe . ia SH ead BY { a Ee kopbe Mt ES vhs Nets LHS Bowe ig