7 et ra arse area ee ye hid hes a ay tae hse a a ‘ Am St eee al DOC arn \ eons Bee alge , uth tan Rooms Vee Se Ra Vaan Mc alate avemen ets DORM v reel tet aig Stet Nn Te th the tetas * phy emit Eh a eS Elks tle oan GAG, sag Vat wn nie aa YO te ere SN as rae tea ren rae e ROL NT yee a ea Ve ete Oh ath Aas fae tar ag A Str seat 4S bh iR ae Sept BSE ead funds se syeges FF oe FRE ey POPE bak Sane & Fey PR ve edie Pept hve Se Os we 298 Roby it Were dete ; 1 GW aS St Eee ine ye 4 eee F 3 ze 2 Pat sequen , Ars da 5 Avie: “a . EP WS yh Bae See . 7 if : J uy 7 i 7 avr gs a: ii , 7 ; rs Warr ; mies ’ - rt é —. _ a ie oi " ‘ } “a Leas €% re oy Ae Pe =) . ; . a ca TA AN ee ; 7 1 : : * i. . att > 2 J 4 © . ‘ " — s vy io r] i : A Mi ve re SEK AAS ) oS we pal ee. ‘ he of ie ; iiooe Se 2 ae ‘ ¢ . ] eat ey 4 ‘qn oh fr +}: i leat 5 arr ay), Sa .: _ yo ty - : MPR Yoni ae Gi ih A ae PROCEEDINGS OF THE California Academy of Sciences Vol. 42 SAN FRANCISCO PUBLISHED BY THE ACADEMY 1979-1982 COMMITTEE ON PUBLICATIONS Tomio Iwamoto, Chairman and Editor Frank Almeda Daphne F. Dunn William N. Eschmeyer Frank H. Talbot (US ISSN 0068-547X) The California Academy of Sciences Golden Gate Park San Francisco, California 94118 PRINTED IN THE UNITED STATES OF AMERICA BY ALLEN PRESS, INC., LAWRENCE, KANSAS | ! eit. » 12, eS. . 14, CONTENTS OF VOLUME 42 WILDER, D. DEE. Systematics of the Nearctic Ptilodexia Brauer and Bergen- Stamm (Diptera: Tachinidae). Published’ March 2, 1979 _.._.. McCoskeEr, JOHN E. The snake eels (Pisces, Ophichthidae) of the Hawaiian Islands, with the description of two new species. Published March 2, CHEMSAK, JOHN A., AND E. G. LINSLEY. Review of the Rhinotragini of Mexico (Coleoptera: Cerambycidae). Published March 2, 1979 KAVANAUGH, DaAviD H. Studies on the Nebriini (Coleoptera: Carabidae), III. New Nearctic Nebria species and subspecies, nomenclatural notes, and lec- hee desivnations. Publushed December 22, 1979). Iwamoto, Tomio. Eastern Pacific macrourine grenadiers with seven branchi- ostegal rays (Pisces: Macrouridae). Published December 22, 1979 FRITZSCHE, RONALD A. Revision of the eastern Pacific Syngnathidae (Pisces: Syngnathiformes), including both Recent and fossil forms. Published July 2, OECD oct CAS NR Ral 7 ee ree Oe ean ee SUE NEE ent fee ee eee ene ee eee BRADBURY, MARGARET G. A revision of the fish genus Ogcocephalus with descriptions of new species from the western Atlantic Ocean (Ogcocephalidae; Panmitornes)bublished July 2; 19800 228" s eee Poss, STUART G., AND WILLIAM N. ESCHMEYER. Xenaploactis, a new genus for Prosopodasys asperrimus Gunther (Pisces: Aploactinidae), with descrip- imnisomiwo new species. Published July 2.1980)... ROBERTS, TYSON R. Sundasalangidae, a new family of minute freshwater salmoniform fishes from Southeast Asia. Published March 5, 198] ALMEDA, FRANK. New and reconsidered species of Miconia (Melastomata- ceae) from Costa Rica and Panama. Published March 5, 198] _------ Briccs, THOMAS S., AND DARRELL UBICK. Studies on cave harvestmen of the central Sierra Nevada with descriptions of new species of Banksula. BaneISHeGe UMC demos bee tee 2 oes = Slee esl PO en eg ene Le LINDBERG, DAviID R., AND JAMES H. MCLEAN. Tropical eastern Pacific lim- pets of the family Acmaeidae (Mollusca, Archaeogastropoda): generic criteria and descriptions of six new species from the mainland and the Galapagos istands senblished: June 24, 1981) <2. ee SMALL, GREGG, J. A review of the bathyal fish genus Antimora (Moridae: Gadioumes)ecubusned june 245 1981 U2: 225. ee ee eee FRICKE, RONALD. The kaianus-group of the genus Callionymus (Pisces: Callionymidae), with descriptions of six new species. Published October 26, [ORT un Set 5S ae aS 2 Rane Oa Wee Siemon Os EAM EO ENED S SOPME ECS re reese iat Rotu, Barry. Distribution, reproductive anatomy, and variation of Monade- nia troglodytes Hanna and Smith (Gastropoda: Pulmonata) with the proposal of anew subgenus. Published October 26, 1981) 2 [iii] 57-67 69-85 85-133 135-179 181-227 229-285 287-293 295-302 303-314 3 15-322 323-339 341-348 349-377 379-407 No. 17. No. 18. Index to Volume 42 JONES. RoBerT E. Food habits of smaller marine mammals from northern Cali- fornia. Published October 26, 1981 22 222 = Ss eee KAVANAUGH, Davip H. Studies on the Nebriini (Coleoptera: Carabidae), IV. Four new Nebria taxa from western North America. Published October IGUIOSL: 4 woe 1. Ro eee oe eee GryYGIER, MARK J. Dendrogaster (Crustacea: Ascothoracida) from California: sea-star parasites collected by the ALBATROSS. Published May 14, 1982 _______ ERWIN, TERRY L. Small terrestrial ground-beetles of Central America (Carab- idae: Bembidiina and Anillina). Published May 14, 1982 __--- == ROBERTS. TYSON R. The Bornean gastromyzontine fish genera Gastromyzon and Glaniopsis (Cypriniformes, Homalopteridae), with descriptions of new species..Published May l4, 1982 2 EE ZULLO, Victor A., AND RAs B. GURUSWAMI-NAIDU. Late Miocene balanid Cirripedia from the Basal Wilson Ranch beds (*‘Merced”’ Formation), Sonoma County, northern. California. Published May 14,1982 222 eee liv] 409-433 435-442 443-454 455-496 497-524 525-535 537-550 PROCEEDINGS OF THE ta a A MAD CALIFORNIA ACADEMY OF SCIENCES Vol. 42, No. 1, pp. 1-55; 113 figs. Wronrds Hole Ma: { March 2, 1979 SYSTEMATICS OF THE NEARCTIC PTILODEXIA BRAUER AND BERGENSTAMM (DIPTERA: TACHINIDAE)* By D. Dee Wilder Department of Entomology, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118 ABSTRACT: A revised classification of the Nearctic prosenine genus Ptilodexia Brauer and Bergenstamm (Dip- tera: Tachinidae) is presented. A total of 8,000 specimens and type material for nearly all species were studied. All seventeen previously described valid species of Ptilodexia are diagnosed and illustrated. Five new species, P. sabroskyi, P. pacifica, P. californica, P. westi, and P. maculata are described and illustrated. A key to the Nearctic species is presented. The following new synonymies are made: P. conjuncta (Wulp) (= Rhynchodexia simulans Wulp); P. contristans (Wulp) (= R. punctipennis Wulp); P. carolinensis Brauer and Bergenstamm (= P. neotibialis West, P. minor West); P. halone (Walker) (= P. hucketti West); P. harpasa (Walker) (= P. leucoptera West, Dinera robusta Curran); P. rufipennis (Macquart) (= Dexia cerata Walker, D. albifrons Walker, Rhynchodexia confusa West, R. translucipennis West, Rhynchodexia dubia Curran); P. major (Bigot) (= Dexiosoma fumipennis Bigot, Rhynchodexia fraterna Wulp, R. omissa Wulp); P. incerta West (= P. proxima West; Rhynchodexia elevata West). The biology of these parasitic flies is reviewed and possible host—parasite relationships are discussed. The taxonomic significance of numerous morphological characters in the genus and the subfamily is discussed. The phylogeny of Ptilodexia and its nearest relatives is discussed; six species groups are separated, and an evolu- tionary tree presented for these groups. The contemporary and historical zoogeography of the genus is discussed as it pertains to host and parasite distribution. A distribution map is presented for each species treated. INTRODUCTION Flies of the genus Prilodexia are large calyp- trate Diptera belonging to the Tachinidae, a fam- ily of exclusively parasitic flies. Ptilodexia adults are commonly collected on flowers during the summer months. The larvae parasitize the * Contribution from the Frost Entomological Museum, Pennsylvania State University. This research was supported by the Agricultural Experiment Station Project No. 2070, and constitues Contribution No. 5611 from that station. An earlier version of this paper was submitted to the Graduate School, Pennsylvania State University, as a thesis in partial fulfill- ment of the requirements for the degree of Doctor of Philos- ophy. (1 larvae of certain scarabaeid beetles. They are of economic interest because they are known par- asites of such pests as Phyllophaga spp., Po- pillia japonica Newman, and Macrodactylus subspinosus (Fabricius). The genus is distribut- ed throughout the New World. Although specimens of Ptilodexia are com- mon in collections, few are correctly identified. Sabrosky and Arnaud (1965) made no attempt to give synonymies or distributions in their cat- alog listing of 27 species names. The special problems leading to such confusion in Ptilodexia are (1) difficulty in associating the sexes, (2) a high degree of intraspecific variation, (3) an un- usually low degree of interspecific variation, and 2 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 1 (4) the lack of consistent traditional morpholog- ical characters. The purpose of this study is to revise the clas- sification of Ptilodexia on the basis of all avail- able material, to analyze the life history of its species, and to determine evolutionary and zoo- geographical trends which may apply also to other members of the family Tachinidae. MATERIALS This study was based on over 8,000 specimens borrowed from various museums. In addition, type-specimens for most of the known species and numerous representatives of other genera in the Prosenini were studied. Institutions loaning material used in this study were as follows: American Museum of Natural History (AMNH), Arizona State University (ASUT), British Museum (Natural History) (BMNH), California Academy of Sciences (CASC), University of California Berkeley (CISC), Canadian National Collection (CNCI), Cornell University (CUIC), University of Ne- braska (DEUN), Field Museum of Natural His- tory (FMNH), Florida State Collection (FSCA), Iowa State University (ISUI), Los Angeles County Museum of Natural History (LACM), Leningrad Museum of Natural History (LMNH), Museum of Comparative Zoology, Harvard Uni- versity (MCZC), Michigan State University (MSUC), Ohio State University (OSUC), Okla- homa State University (OSEC), Oregon State University (OSUO), Paul H. Arnaud, Jr., Col- lection (PHAC), Yale University (PMNH), Pur- due University (PURC), South Dakota State University (SDSU), University of Kansas (SEMC), Staten Island Institute of Science (SITS), University of Oklahoma (SMSH), Texas A & M University (TAMU), University of Ari- zona (UAIC), University of Alberta (UASM), University of California Davis (UCDC), Uni- versity of California Riverside (UCRC), Univer- sity of Idaho (UICM), University of Montreal (UMIC), University of Michigan (UMMZ), Utah State University (USUC), National Museum of Natural History (USNM), Vienna Museum of Natural History (VMNH), Washington State University (WSUC). METHODS The male genitalia of Prilodexia species are partially obscured on dry, pinned specimens. To examine them, the posterior half of the abdomen was removed, placed in a solution of 10 percent KOH, and heated until the structures were suf- ficiently softened to be dissected easily. They were later rinsed twice with water and twice with acetic acid, placed in glycerine, and ex- amined. They were stored in a microvial pinned beneath the insect. Illustrations of the genitalia were made using an ocular grid. The postabdomen was anchored to a small piece of soft wax on the bottom of the dish of glycerine to prevent it from drifting about. Drawings of the heads were made by project- ing photographic transparencies of them onto drawing paper. Manipulation of the projector provided images of uniform size. The image was then traced with a hard pencil; the details were filled in after thorough examination of the spec- imen with a dissecting microscope. All measurements were made using an ocular grid, calibrated by a stage micrometer. Ratios were calculated from these measurements. Because of the extreme intraspecific variabil- ity of Ptilodexia, a description which included all variation would be unwieldy and would be similar for each species. To make the descrip- tion more useful and manageable in size, only one specimen, the holotype, is described. A complete synonymy and list of citations are given for each species included in this study. New species are thoroughly described and di- agnosed. For previously described species, only a diagnosis is presented. Known information on each species is summarized and notes regarding types and nomenclature are presented. Intraspecific variation is discussed thoroughly for all new species. With previously described species, variation is discussed only where ‘it is necessary for species identification. Complete data from each specimen, including sex, locali- ty, collector, depository, and other information, have been recorded by Wilder (1976) and there- fore are not presented here. Wilder (1976) also gives complete redescriptions for all previously described species of Prilodexia. The scope of this revision has been limited for practical reasons. Inclusion of the southern Mexican species would have doubled the num- ber of species treated, and the material available for these species is wholly inadequate. All Nearctic species are treated herein. WILDER: NEARCTIC PTILODEXIA ACKNOWLEDGMENTS I extend my sincerest appreciation to Dr. C. W. Sabrosky, who initiated this study and with- out whose help this revision could not have been completed. Dr. Sabrosky also generously loaned types and was always available to answer ques- tions about Prilodexia and related tachinids. I thank Dr. K. C. Kim for his valuable assis- tance during this project. His enthusiasm and drive served as a constant inspiration. Persons and institutions loaning type material, for which I am grateful, are: Dr. R. W. Crosskey (BMNH), who was extremely generous in loan- ing 56 type-specimens for this project; Dr. R. Litchtenberg (VMNH); Dr. V. Richter (LMNH); Dr. L. L. Pechuman (CUIC) who kindly loaned me the types of 14 species for an extended pe- fiod of time; also J. C. Scott (MCZC), Dr. P. Wygodzinsky (AMNH), and Dr. F. C. Thomp- son (USNM) who helped in uncovering some important syntypes. I am indebted to Dr. P. H. Arnaud, Jr., and the Department of Entomology, California Academy of Sciences, for providing facilities and valuable assistance during my time on the West Coast. Thanks are also due to Dr. D. C. Rentz and D. H. Kavanaugh for their frequent assistance with many problems. I finally thank my husband, George Zelznak, for his unending encouragement and optimism throughout this study. BIOLOGY The larvae of Prilodexia flies parasitize and kill their scarabaeid larva hosts. The adult flies, however, feed on nectar and they spend consid- erable time probing at flowers, particularly com- posites. While feeding they become covered with pollen and probably act as pollinators. Ptilodexia conjuncta and P. agilis adults have been observed pollinating the flowers of dwarf mistletoe (Arceuthobium cyanocarpum). Adults of Ptilodexia have been collected with pollinia attached to the tarsi. Adults are collected in many environments. Members of some species are collected at the seashore, while others have been taken at alti- tudes as high as 3,000 m in the Sierra Nevada and the Rocky Mountains. These flies have been taken at UV light, Malaise, and other flight traps. But the most productive method of col- lecting seems to be sweeping flowers, especially composites such as Baccharis and Solidago. The occurrence of specimens of Ptilodexia is, as with most parasites, seasonal. At times, hundreds of individuals of one, two, or even three species may be collected simultaneously at one kind of flower. In other years the flies will be rare—perhaps representing the normal build- up and decline of a parasitic population. In some areas, such as Long Island, New York, and Riv- erside, California, specimens of certain species have been collected on the same dates every year for ten or more years. However, both these areas have been extensively surveyed regularly by specialists and may represent the actual sit- uation, which is not seen in other areas simply because of poor sampling. There is no information on the mating habits of Ptilodexia spp. Despite the numerous speci- mens collected, few have been pinned in copula. Males usually emerge before females, but the place and time of mating is unknown. The female carries hundreds of tiny larvae in her abdomen during larviposition. It is not known if she simply broadcasts them or if she places them directly on the soil. Neither is it known if the female is able to locate areas of host density, nor if the larva has the sensory capacity to find a host. If the larvae are depos- ited in a jar, they will wander along the sides for two to three days before dying. First-instar lar- vae of Prosena siberita survive a week or more in the soil (Clausen 1927). The larvae are presumably quite easy to rear if the host larvae can be kept alive under labo- ratory conditions. During the development of the parasite larva, a defensive response of the host causes a respiratory funnel to appear at the point of attachment to the host. This is a scler- otized funnel-shaped structure which encloses the caudal end of the larva and can be seen through the integument of the host. The larva feeds on the internal fluids and fat body of the host and finally leaves the host to pupate in the soil. By this time, the larva has ingested nearly all the contents of the host. Based on Davis’s (1919) data, Prilodexia lar- vae overwinter within the host. The pupal stage is quite short, cold temperatures are not re- quired to complete development. The host scarabaeid larvae, or white grubs, belong to the subfamily Melolonthinae, which 4 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 1 includes such common and economically impor- tant genera as Phyllophaga, Popillia, and Mac- rodactylus. The dynastinine scarabaeids of the genus Aphonus are also parasitized by these flies. Champlain and Knull (1944), and Peterson (1948) implicate Ptilodexia canescens as a par- asite of the cerambycids Saperda calcarata and Rhagium lineatum. Most species of Phyllophaga have a two- or three-year life cycle. Popillia japonica and some of the other hosts have one-year life cycles. The life cycle of Ptilodexia probably does not exceed one year. Early-instar white grubs are attacked in the fall and fed upon until the following spring or summer when the parasite pupates. Adult flies emerge shortly thereafter. According to Davis (1919), when infested host larvae are brought indoors in the fall, the Prilo- dexia larvae will continue their development within the host, pupate, and emerge as adults in the winter, without interruption. It is thus pos- sible that in areas with long seasons, two or more broods could develop. My samples from Texas, southern California, and some south- eastern states appear to support this. Two broods per year are possible only if there is an ample supply of grubs of the proper stage feed- ing in the soil. In these areas of bivoltinism, vari- ant populations in the species concerned are ap- parently more common than in areas where only one brood is possible. One can extrapolate to the tropical regions where even more genera- tions per year are possible, and the number of species and the variation among species is phe- nomenal. It is not known if any species of Ptilodexia is host specific, but some species are known to have more than one host. For example, P. car- olinensis can complete its life cycle either in Phyllophaga rugosa (fide Davis 1919, as Ptilo- dexia abdominalis) or in Popillia japonica. On the other hand, Ptilodexia maculata and P. prexaspes have restricted ranges and show little intraspecific variation—perhaps indicative of host specificity. Ptilodexia harpasa and one of its reported hosts, Macrodactylus subspinosus, are sympatric. Many questions remain unanswered regarding the relationship of host preference and specia- tion in Prilodexia. In some species, local aber- rant populations are found that only vaguely re- semble the typical population. The possibility exists that these aberrant populations have shift- ed to a host significantly different from the com- mon one. It is conceivable that such a shift could be an early step in speciation. Many factors involving the relationship of host and parasite presumably influence the ap- pearance of the adult fly. These factors include number of fly larvae per host, instar of parasi- tized host, rate of host development, and rate of parasitoid development. Specimens of P. caro- linensis developed in Phyllophaga grubs differ greatly from those developed in larvae of Popil- lia. These factors contribute to the extreme intra- specific variability in Ptilodexia, perhaps ulti- mately leading to speciation in the group. Care- fully controlled breeding experiments are needed to help understand the effects on the parasites of the host and host environment. TAXONOMIC CHARACTERS Most of the specific characters previously used by Prilodexia taxonomists are subtle, dif- ficult to see, and unstable, sometimes differing not only between individuals but also on each side of the same specimen. Of the characters traditionally used to separate species, many have been either stable within the genus (e.g., ‘‘arista plumose’’) or different within a species (e.g., color, wing venation). Most earlier work- ers lacked sufficient study material to recognize normal intraspecific variation. Certain charac- ters these early workers used were good, but they are more useful used in combination with certain other characters. Sexual dimorphism has caused problems in the taxonomy of Prilodexia. Abdominal color and color pattern, and leg color frequently differ between sexes; hence the two sexes of some species have been described under different names. I have freely used raw measurements in my descriptions, recognizing nonetheless that they are of limited use in this group. Proportional measurements are generally more useful, and I include them in my descriptions and diagnoses. The standards for these proportional measure- ments are head height and length of the first an- tennal segment—both measurements which are proportional to general size. This study is limited to dried adult flies, so structural characters are the only ones which form the basis of my classification. What follows WILDER: NEARCTIC PTILODEXIA is a brief discussion of the taxonomic impor- tance of the various physical characters and how their states are determined. The headings and general organization are similar to those used by Crosskey (1973a). It is hoped that other workers in the Tachinidae will adopt the same format, eventually bringing some order to the study of variation in the family. Body Color and Vestiture The general integumental color of Ptilodexia adults is a dull brown, although adults of certain more-advanced species may be black or testa- ceous, and teneral specimens are generally pal- er. The color of the scutellum compared to that of the rest of the notum sometimes is specifically useful (e.g., P. planifrons—P. contristans); how- ever, in adults of some species (P. rufipennis), it also varies intraspecifically. The color of the abdomen varies from reddish or testaceous with a dark longitudinal stripe, to concolorous black to testaceous. Although abdominal color may be of occasional taxonomic value, it almost always varies between males and females of the same species. In the female it is frequently concolor- ous, with the longitudinal stripe indistinct or ab- sent. General body color sometimes varies cli- nally, and in some species, smaller, darker populations exist in the northern parts of the range. Vestiture characters can aid in distinguishing members of different species. These characters seem to vary independently of integumental col- or. Facial tomentum varies from extremely heavy—totally obscuring the underlying integ- umental color—to fine and sparse. Occasionally there is a pattern or spot of color in this vestiture which can be distinct for a species (e.g., P. con- tristans, P. canescens). Facial tomentum may be dull (P. westi) or strongly shining (P. incerta). The color of the facial tomentum varies intra- specifically. The tomentum on the pleuron is of little taxo- nomic value, and that on the notum is only slightly more useful. The notal tomentum on adults of some species (e.g., P. westi, P. arida) is heavy, abundant, and almost flocculent, near- ly obscuring the integumental color, whereas on those of others (P. mathesoni) it is so fine that the notum appears polished. In members of oth- er species (P. conjuncta), the tomentum is flat- tened and shiny, giving the notum, or parts of it, a coppery hue. Usually notal tomentum is arranged in longitudinal stripes, but this striping varies among individuals and is of little diagnos- tic value. Notal tomentum also varies between the sexes, usually being heavier in the female. Tomentum on the mediotergite can be fine or heavy; in specimens of P. agilis and P. mathe- soni, however, it is absent. Abdominal tomentum varies more between species than between sexes. It is generally gray- ish and tessellate, although in some individuals it may be gold or brownish; and it may be shin- ing or dull. Only in adults of one species (P. mathesoni) has the tessellate pattern been re- placed by a more uniform distribution of tomen- tum, and even then, only in the males. In mem- bets of atew species (erg... 2. pacijica, P: ponderosa), the grayish tomentum is the only vestiture on the abdomen, but in most there are patterns of brown or gold tomentum which can aid in distinguishing species. In P. rufipennis adults the bases of the median marginal setae on the third and fourth abdominal tergites are sur- rounded with gold tomentum; adults of P. agilis have a marginal band of it on the third and fourth tergites; those of P. arida have brown tomentum overlying the longitudinal stripe. These characters of general color and vesti- ture show a greater degree of variation within Ptilodexia than within all other Nearctic Pro- senini, with the possible exception of the most closely related genus, Mochlosoma. When examining specimens for colors and patterns of tomentum, it is imperative that they be viewed from several different angles. Often a pattern can be distinguished only if it is seen obliquely. Chaetotaxy and Hairing Chaetotaxy is of minor taxonomic use in Ptilodexia, as it varies widely within species but little between them. Frequently, numbers and even the presence of setae vary from one side of the specimen to the other. This phenomenon is represented in the descriptions by separating the two states with a slash (1/0). Hairs differ from setae in being much smaller and finer. The taxonomically useful setae on the head are the oral vibrissae (Fig. 1). There is usually one pair (two in specimens of P. contristans). The size and spatial relationships between the vibrissae and the peristomal setae can aid in identifying adults in some species. In P. ruft- pennis adults the peristomals immediately below 6 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 1 Postorbital frontal vitta__~ (_ J seta frontal seta A gena Va ; arista Vk \ head vibrissa Nay lenatts —_| Parafrontal > profrons a . length iA epistome (i — A peristomal seta Wy parafacial — ih haustellum ) carina | U head height < je ninth tergite xo cercus surstylus aaa y ean) Co e;aculatory apodeme adeagus 3 Ficures 1-3. Structure of generalized Ptilodexia. Fig. 1. Head, anterolateral view. Fig. 2. Head, lateral view. Fig. 3. Genitalia, lateral view. the vibrissae are short, becoming longer with distance from the vibrissae; in P. conjuncta adults, on the other hand, the peristomals are subequal and nearly as long as the vibrissae. Numbers of peristomal setae vary between and within species, but considerable overlap be- tween species is common. The number and size of frontal setae vary, but these are even less reliable characters than are the number and size of the peristomal setae. The ocellar, postocellar, internal vertical, and external vertical setae show some intraspecific differences in size and number. I have described these differences, but they are too variable to be used diagnostically. The postorbital setae vary in length between species, but not as much as between the sexes. The hairs which are some- times inserted between them have minor signif- icance. In adults of some species the postorbit- als are long and closely spaced, while in those of others they are interspersed with fine setae half the length of the postorbitals, and in still others they are interspersed with tiny hairs. Another group of taxonomically useful hairs on the head are those immediately ventral to the postorbitals. Members of species such as P. planifrons and P. mathesoni are characterized as having two to four irregular rows of dark hairs between the postorbital setae and the yellow or white occipital hair. Members of other species have only one row of these dark hairs (P. cali- fornica), and those of others have none or just a few scattered hairs (P. maculata). This char- acter also shows much intraspecific variation and must be used cautiously. Perhaps the best diagnostic character in Prilo- dexia is the hairing of the parafacials (herein de- fined as the sides of the head bounded by the apex of the second antennal segment, the oral vibrissae, the frontal suture, and the anterior eye margin). The presence, size, distribution, and color of these hairs are extremely variable but species specific. There is slight variation in the characteristics of these hairs between males and females belonging to the same species; that is, the parafacial hairs of the female are slightly sparser, finer, and are not inserted as far ven- trally on the parafacial as they are on the male. These hairs may be absent (P. rufipennis); long, dark, and abundant (P. planifrons); sparse and pale (P. halone); minute and occurring only on the upper anterior parafacial (P. incerta); strong and concentrated at the lower edge of the eye (P. canescens); strongly inclined anteriad (P. contristans), or inclined ventrad (P. harpasa). Many other combinations exist. It appears that this character can be of diagnostic value even in the Tropics, where there are many undescribed species. I have illustrated the character state for parafacial hairs in every Nearctic species herein described or diagnosed. The nature of the para- facial hairs is also of importance in Mochloso- ma, where they are always present, but not in the other Nearctic Prosenini. Parafacial-hair characteristics appear fre- quently in the key to species. When the hairs are small and pale, specimens must be examined carefully from several angles; often it is the bas- es of the hairs rather than the hairs themselves which are visible. The parafrontal hairs are of much less taxo- nomic value. They are usually present, dark, and are either sparse or abundant. WILDER: NEARCTIC PTILODEXIA Thoracic chaetotaxy is of little diagnostic im- portance in Ptilodexia. The numbers of such se- tae as sternopleurals, notopleurals, and posta- lars are generally constant within the genus. Others such as posthumerals, presuturals, ac- rostichals, dorsocentrals, and scutellars vary somewhat between species, but they also show considerable intraspecific variation. The number of humeral setae and discal scutellars are more constant, but must be used in combination with other characters to aid identification. The length and density of hairs covering the Ptilodexia thorax vary between species. These, however, are difficult characters to divide into easily defined states. Propleural hairs do not oc- cur in Ptilodexia adults, but are present in mem- bers of several closely related genera. They have diagnostic value at the generic level. Another group of hairs on the thoraces of these flies is the infrasquamal setulae, small hairs inserted beneath the point of attachment of the squamae or calypters. In adults of some related genera in the Prosenini, these are always absent. In some Ptilodexia adults their absence may be a reliable specific character state (e.g., P. canescens, P. maculata), but in others, their absence carries less taxonomic importance. Six- ty percent of the specimens of P. rufipennis ex- amined had infrasquamal setulae, but they were present in only twenty percent of P. incerta specimens. This character is of equal value in both sexes. When using this character, one must realize that the ‘absence’ of infrasquamal setulae indicates absence on both sides of the body. Hairs and setae on the legs have little diag- nostic value; often the setal length reflects total body size more than any specific difference. The exception to this is the length of the antero- and posteroventral setae on the posterior leg of the male, which show species-level variation. These are difficult to measure, however, and have not been used in this revision. Other setae on the femora show some taxonomic potential, espe- cially the presence or absence of anterior setae on the posterior femora. Numbers of abdominal setae vary intraspecif- ically, but they usually vary around a certain number which can be defined for some species. The presence or absence of median marginal se- tae on the first syntergite can be a useful char- acter. The number of median discal and median mar- ginal setae on the third and fourth tergites is useful in separating members of some closely related species (e.g., P. californica and P. pa- cifica), while in others it shows considerable in- traspecific variation. The presence or absence of lateral discal setae on these tergites will sep- arate members of distantly related species. Length and density characters of abdominal hairs have about the same taxonomic value as those characters in thoracic hair. That is, they differ and seem to be constant among members of a species, but are difficult to separate into character states. Although hairing on the genitalia varies only slightly between species, the presence of strong setae on the ninth tergite (epandrium) is an ex- cellent diagnostic character in adults of P. con- tristans and P. westi. Head Head characters in Ptilodexia are of more use taxonomically than characters of any other part of the fly. Included are those of chaetotaxy, which have been discussed in the previous sec- tion. Drawings of the head, with the terms used in this paper, are presented in Figures 1-2. The head, in members of this genus, is wide and boxlike. The parafacials and genae are wide and covered with fine, dull-lustered tomentum. The genae are usually reddish, contrasting with the whitish parafacials and genal dilations. The velvety-appearing frontal vitta extends from the vertex to the frontal suture. The third antennal segment is rarely longer than twice the length of the second and bears an arista covered with long fine hairs. Between the antennae Is a raised ridge or carina which does not protrude beyond the antennae. The epistome may or may not pro- trude. Mouthparts are similar to those of other calyptrate flies with the mentum length from 0.3 to 0.8 times the head height. The width of the parafacial is of considerable diagnostic use in Prilodexia. Ptilodexia rufipen- nis adults have narrow parafacials, while in P. conjuncta adults they are quite wide. This char- acter is easier to evaluate in males than in fe- males, and it varies more among Ptilodexia adults than among Mochlosoma adults. The shape of the frontal vitta can be of use in this genus. Below the ocellar triangle and be- tween the eyes, the frontal vitta is usually oblit- erated (Fig. 4b), the parafrontals becoming con- 8 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 1 Figures 4-6. Comparison of specimens of Ptilodexia. Fig. 4. Comparison of adults of two generalized species of Ptilodexia showing variation in width of epistome, position of oral vibrissae, and width of frontal vitta; A. head, anterior view, with frontal vitta not obliterated and distance of oral vibrissae from oral margin greater than distance between oral vibrissae; B. head, anterior view, with frontal vitta obliterated and distance of oral vibrissae from oral margin less than dis- tance between oral vibrissae. Fig. 5. Comparison of antennae of adults of two generalized species of Prilodexia; A. antenna, showing length of plumosity on arista shorter than length of second antennal segment; B. antenna showing length of plu- mosity on arista longer than length of second antennal seg- ment. Fig. 6. Ptilodexia ponderosa (Curran), holotype, head of female, lateral view. tiguous. In members of some species (e.g., P. canescens, P. halone), however, the parafron- tals do not touch, and the frontal vitta is contin- uous from the antennal base to the ocellar tri- angle (Fig. 4a). This character varies to a similar degree in specimens of Mochlosoma. The size and shape of the carina are useful at the supraspecific level. Among Ptilodexia species, the carina is sometimes distinctly shaped (e.g., P. planifrons). Carina characters vary more among Mochlosoma than Ptilodexia adults, in general being wider and better devel- oped in individuals of the former genus. In mem- bers of several closely related genera such as Dinera and Hesperodinera, the carina is strong- ly developed, protruding from between the an- tennae, appearing almost bulbous, and visible from the lateral aspect. In Rhamphinina adults it is short, narrow, and strongly keeled. The ca- rina shows no sexual dimorphism in size and shape, and except in a few cases it is species specific. The oral vibrissae and adjoining areas hold characters of taxonomic value in Ptilodexia. These are the width of the depression between the bases of the oral vibrissae and the distance of the vibrissae from the epistome (sclerotized oral margin). Members of some species of Ptilo- dexia have this area wide and short; an example is P. rufipennis, in adults of which the vibrissae are far apart and close to the oral margin (Fig. 4b). The opposite state is shown in P. prexaspes adults, which are characterized by vibrissae that are far from the oral margin, but not far apart (Fig. 4a). In Rhamphinina specimens, the area is four times as high as wide. This character is diagnostic in a few species, but in others it varies intraspecifically. It shows no sexual dimor- phism. In evaluating this character, physical measurement is necessary; estimate is inade- quate. The invisible line connecting the vibrissae should pass through the center of their bases, and the sclerotized margin of the epistome should serve as the ventral boundary. Width is measured only at the vibrissae; height is mea- sured mesially. The epistome generally protrudes in Prtilo- dexia and Mochlosoma specimens, and al- though both genera show variation, in Ptilodexia adults it is sometimes species specific. In adults of species like P. prexaspes, P. canescens, and P. halone, the oral margin projects slightly if at all. As a result, the lower anterior portion of the head is vertical in profile (Fig. 71), and in some cases the anterior margin of the head protrudes further anteriad at the antennae than at the vi- brissae. Ptilodexia conjuncta adults show the opposite state, the epistome projecting strongly, as does the lower anterior portion of the face (Fig. 26). In other prosenines the character shows less intraspecific variation than it does among species of Ptilodexia. The length of the haustellum is an extremely valuable taxonomic character in Prilodexia (and Mochlosoma), and it is also the chief difference distinguishing Prilodexia from Mochlosoma specimens. In individuals of the former genus, the length of the haustellum varies from 0.3 to WILDER: NEARCTIC PTILODEXIA (0.9 times the head height, and the shape is broad and linear or slightly tapered; it is rigid in all individuals. Mochlosoma specimens have the haustellum much longer than the head height, and narrow and flexible. In other Prosenini this character serves to separate genera. In Prilo- dexia I have compared the length of the haus- tellum with the head height and used the result- ing ratio as a diagnostic character which varies consistently between species, little within species, and not at all between the sexes. When using this character in the key, actual measure- ments must be made; estimating the ratio is dif- ficult because a slender haustellum appears lon- ger than a broad one of the same length. There is intraspecific variation in the length of the haustellum among members of a few species. Among P. rufipennis (as well as P. arida, P. carolinensis, and P. pacifica) specimens, the haustellum length varies locally. The length can be short in members of one population and no- ticeably longer in those of another. The char- acter is still useful, though, since the variation remains within easily expressed values. The length and shape of the palpi vary slightly between members of different species of Ptilo- dexia. The length is expressed, in this paper, as a fraction of the haustellum length. Some Pro- senini, such as Prosena and Senostoma speci- mens, have short stubby palpi; and in Atelog- lossa adults they are completely absent. This character varies among Mochlosoma specimens much as it does among those of Prilodexia. In members of P. arida and P. prexaspes, the length of the palpi may nearly equal the length of the haustellum, while in those of P. obscura, it is rarely more than 0.3 times the haustellum length. The antennae possess some useful taxonomic characters: length, shape, and arista plumosity. The length of the third segment is herein ex- pressed in terms of its relationship to the rela- tively constant second segment. Measurement of the second segment is taken from a slightly anterodorsal aspect and is the longest dorsoven- tral length of the segment. Among Prosena and Senostoma adults, length of the third segment is approximately twice the length of the second; in those of most other Prosenini, it is considerably shorter. Among Mochlosoma species the length varies, the most usual state being the third segment equal to 1.4 to 1.5 the second. The same is true in Ptilodexia species, where this character can be used to separate adults of some species. Ptilodexia sabroskyi adults have a short third segment, subequal to or shorter than the second, whereas those of P. rufipennis have the third segment up to twice the length of the second. Specimens of P. obscura sometimes have the third antennal article broadened apically instead of slightly pointed as it is in members of most species. The length of the plumosity on the arista is an excellent diagnostic character in Ptilodexia. I have expressed it in relation to the length of the second antennal segment. The arista, including the plumosity, is measured at its greatest width (Fig. 5). In specimens of P. rufipennis and P. harpasa, two species with long third antennal segments, the length of the plumosity is greater than twice the length of the second antennal seg- ment, while in those of P. planifrons and P. prexaspes, the length of the plumosity is less than or equal to the length of the second seg- ment. This character is especially useful in sep- arating adults of the closely related P. califor- nica and P. sabroskyi. Thorax Most of the thoracic characters used in this revision have been discussed in the sections on vestiture and chaetotaxy, the remaining ones are those of the mediotergite, legs, and wings. Adults of Prilodexia have a typical calyptrate thorax with the mesothorax highly developed, and the prothorax and metathorax reduced. The scutellum is small; ventral to it is the bulging postscutellum, which distinguishes members of the family Tachinidae. The pleuron is typical of other calyptrate flies. The propleuron is bare, although the rest of the pleuron is beset with fine hairs and numerous groups of setae. The legs are long with extremely long tarsi. The wings are also long, the venation typical of ca- lyptrate flies. The mediotergite is the oval arched area ven- tral to the postscutellum. In members of some species the mediotergite is dorsally polished, although other parts of it may be tomentose. Care must be taken when observing this char- acter, since on adults of some species the me- diotergite has a thin layer of tomentum and still appears shiny. Leg color is a taxonomically useful character. In species where the color is similar in members 10 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 1 of both sexes, pale-colored legs are diagnostic. The color of the tarsi can also separate members of different species (e.g., P. halone and P. prex- aspes). Ptilodexia maculata specimens are dis- tinguished by distinct femoral patches which, although present on members of other species, are strikingly evident on those of P. maculata. Other species exhibit dimorphism in leg color, the males with dark legs, the females with pale legs (e.g., P. agilis, P. arida). Two other species, P. rufipennis and P. pacifica, have fe- males with pale legs and males with legs of var- ied color. Wing venation is useful in distinguishing members of some genera in this tribe (Nimio- glossa). Within Ptilodexia (and Mochlosoma), however, it is of dubious value. Wing color is constant within species and can be used diag- nostically. Adults of P. contristans have the wings distinctly darkened basally, while those of P. mathesoni have the entire wing darkened. The colors of the squamae, epaulet, and basi- costa also show slight differences between mem- bers of certain species, but they can be varied among those of others. Abdomen and Genitalia In Ptilodexia adults as in most Nearctic Pro- senini, the abdominal tergites meet ventrally, entirely obscuring the sternites. The first tergite is actually composed of two fused segments; the next three tergites—third, fourth, and fifth—are conspicuous. The sixth tergite is fairly broad, the edges not meeting ventrally (but embracing the fifth sternite); it and those remaining are withdrawn into the fifth tergite. The next two tergites are fused and become the seventh syn- tergite, which is fairly narrow in Ptilodexia members (not much wider than the epandrium), with its surface oriented posterodorsally, as is the epandrium (the ninth tergite). This pattern is similar in Mochlosoma members, but in other Prosenini it is different. In Prosena adults, for example, the ninth tergite appears to be fused with the seventh and eighth, and in members of Hesperodinera, the fused seventh and eighth tergites are exposed and greatly enlarged, the surface facing posterad, the epandrium forced beneath the abdomen. Abdominal color is varied intraspecifically but is still useful as a key characteristic distinguish- ing members of some species. In the key pre- sented herein, when the abdomen is described as reddish laterally, at least the second and third tergites (of the male—the character is not as consistently applicable to the female) have the integument reddish or rufotestaceous laterally. On specimens in which the abdomen is concol- orous dark brown or gray, there may be a slight rufescent cast along the margins of the tergites. This state should not be confused with the pre- vious one, in which the reddish color extends from the anterior to posterior margins of the ter- gites. In specimens of Ptilodexia, the external male genitalia (Fig. 3) have taxonomic value. The characters which vary slightly between mem- bers of different species are the shape of the ninth tergite, the shape of the surstyli, and the shape of the cerci. These characters are useful in distinguishing members of a few species, but sometimes vary more intra- than interspecifical- ly. Only in species with extraordinarily modified members (e.g., P. westi, P. rufipennis) can the external genitalia be called diagnostic, and even then, they must be dissected for characters to be examined properly. Often the genitalia of adults of Mochlosoma and Ptilodexia are iden- tical. The above-mentioned characters vary greatly between specimens belonging to differ- ent genera and are of excellent supraspecific group characters. Internal genitalia are generally not useful in separating members of species of Ptilodexia. The aedeagus is nearly identical in members of this genus and those of Mochlosoma. The ejac- ulatory apodeme, however, is useful in distin- guishing specimens of some species or species groups. Its shape can be distinct, as in P. con- tristans, P. planifrons, and P. rufipennis mem- bers; between many of the species, though, it does not vary. Female genitalia show no striking diagnostic differences, with the notable excep- tion of the surface sculpturing of the spermathe- cae which, with high-magnification studies, may reveal specific differences. The reproductive systems of both sexes of Ptilodexia have been described by Townsend (1938). The larvae of Ptilodexia have never been de- scribed, even though there is a figure of a mature larva and the puparium in Davis (1919). The na- ture of the cephalopharyngeal skeletons of first instar larvae (from the abdomen of gravid fe- males) has been used to separate species in some genera of tachinids (Archytas), but the character is of no use in Ptilodexia. Greene (1922) de- WILDER: NEARCTIC PTILODEXIA scribed the puparium of an unknown species of Ptilodexia (erroneously determined as P. tibi- alis). Of the useful diagnostic characters, none works to separate members of ail species from those of all others. Most of these characters are of high value in distinguishing members of the derived species, but when members of certain primitive species (P. carolinensis, P. major) are examined, they lose much of their value and more characters must be considered in making identifications. PHYLOGENY Present attempts to reconstruct the phylogeny of a genus or tribe in the Tachinidae are based on incomplete data and should be considered extremely tentative at best. Characters used at generic and tribal levels are so unstable that con- vergence, loss, and acquisition occur repeated- ly. Most of the species, and probably many of the genera, are unknown or poorly defined on a worldwide basis. Host relationships are largely unknown. For the phylogeny of Prilodexia, Neotropical species and representatives of closely related genera were carefully examined to infer apo- morphic and plesiomorphic states. Character matrices were then constructed and phylogenet- ic trees inferred. This method works well when trying to construct probable relationships in higher categories, but for relationships among species it is not adequate. This is becuase the characters distinguishing species are generally more unstable than those distinguishing families or tribes. Many specific characters can be lost or regained easily. Relationships within the Prosenini can be in- ferred only after examining members of the tribe on a world basis. I have not had the opportunity to do this. I have seen a few representatives of the North American genera, none of the exclu- sively Neotropical genera, one of an Australian genus, and one of Prosena, a worldwide genus. Most of the species in these genera can not yet be identified with existing keys. The characters used to infer relationships be- tween genera are facial carina, space between vibrissae, propleural hairs, and haustellum length. The form of the facial carina provides a good generic character, much as it does in the tribe Rutiliini (Crosskey 1973a). In members of Prosenini, the carina was lost once, although slight expression is common in members of some species of Prilodexia and Mochlosoma. The area between the vibrissae is another sta- ble generic character. In Ptilodexia and Moch- losoma adults, and in those of some of the other genera, this area is depressed and may be flat or slightly concave, while in members of the more primitive genera, it is slightly to strongly con- vex. In the primitive genera, the oral vibrissae are situated at or slightly above the oral margin, while in Ptilodexia, Mochlosoma, and Rham- phinina adults, they are inserted distinctly dor- sal to the epistome. In Arctophyto and Milada adults, their placement is intermediate between that in members of the primitive and the derived genera. Propleural hairs are lacking in members of Sentstoma and Prosena, two of the more prim- itive genera. They have been lost in Hespero- dinera adults and are never seen in those of the advanced genera. The haustellum is longer than the head height in specimens of Prosena, Prosenoides, Nimio- glossa, and Mochlosoma, but is much shorter in those of the other genera. Also, there is great variation in haustellum length among Ptilodexia species (from 0.3 to 0.9 times head height). The position of Ptilodexia within the Prosen- ini is advanced. Assuming an Oriental center of origin for the group, there is a wealth of forms (members of which have a broad, blunt, facial carina) in the Oriental and Palearctic regions. Some of these, such as Prosena and Dinera, are widespread. It was probably a form similar to Dinera, Milada, and Arctophyto which, along with its hosts, crossed the Bering Land Bridge during favorable conditions in the late Tertiary. Subsequent radiation before and during the Pleistocene must have been great, for Ptilodexia is the most diverse genus of prosenines in the western hemisphere. Dinera and Arctophyto-Milada remain on both sides of the Pacific with relatively few species. Other small genera which proably orig- inated from this complex are Ateloglossa, Dol- ichocodia, Prosenoides, and Hesperodinera, members of which have retained the inflated ca- rina, and Myoceropsis, Rhamphinina, and Nim- ioglossa, whose members have lost it. Ptilodexia and Mochlosoma, both large gen- era, probably had similar origins, Mochlosoma from a more restricted ancestral line. Although 12 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 1 the strong carina has been lost, it is expressed to a moderate degree in members of some species in both genera. It is my opinion that the only character which separates Ptilodexia from Mochlosoma adults, the haustellum shape and length, is a phylogenetically sound one, that is, its origin in Mochlosoma is monophyletic. Al- though radiation of both genera has been great, divergence between Mochlosoma and _ Ptilo- dexia members is only slight. Ateloglossa and Hesperodinera had their origins from a Dinera-like ancestor, members of the former having lost the palpi and the latter the propleural hairs, but both having retained the squarish head and inflated facial carina. Pro- senoides adults, on the other hand, bear a closer resemblance to Prosena specimens, and the two may be closely related. Both Nimioglossa and Rhamphinina had their origins early in the Prilo- dexia-Mochlosoma line, their members having diverged from the ancestral forms in having the area between the oral vibrissae depressed and the vibrissae placed considerably above the oral margin, as do representatives of Ptilodexia and Mochlosoma. Within Ptilodexia, characters indicating rela- tionship are difficult to determine. The ancestral and derived states of a few of these characters have been inferred. Small size and dark color seem to be primitive states within Prilodexia, while the derived states are large size and pale color. The primitive state of the length of the third antennal segment, the length of the plumosity on the arista, and length of the haustellum is an intermediate one, with the derived states being short and long. The presence of both parafacial hairs and infrasqua- mal setulae appear to be primitive within this genus. Primitively the oral margin is narrow and projecting, while the more advanced forms show it to be wide and not projecting. The primitive, typical shape of the ejaculatory apodeme can be seen in most species (Fig. 44). Members of some of the advanced species have this structure mod- ified in various ways (Fig. 29, 35), although those of others do not. The genus Ptilodexia in North America is comprised of six loosely knit groups. The first, and probably most primitive of these, is the agilis group, which consists of P. agilis, P. ob- scura, and P. mathesoni. The next group is the carolinensis group, with member species P. carolinensis, P. halone, P. prexaspes, and P. canescens. The harpasa group is composed of P. rufipennis, P. arida, P. harpasa, and P. pon- derosa. In the major group are P. major, P. incerta, P. maculata, and P. flavotessellata. The conjuncta group contains P. conjuncta, P. planifrons, P. contristans, and P. westi; and the californica group, P. californica, P. sabroskyi, and P. pacifica. The character states which segregate mem- bers of these groups are vague and difficult to define, but since the groups appear to have both a zoogeographical and morphological basis, they will be discussed. Their relationships to each other are somewhat less clear. Members of the agilis group are small, dark flies with little red color on the abdomen and a short haustellum. The legs of the females of P. agilis and P. mathesoni are pale, while those of P. obscura are dark. The mediotergite is pol- ished or shiny in members of all three species. This is probably the most primitive group of Ptilodexia; P. agilis members perhaps being similar to those of the prototype of the genus. Ptilodexia agilis is a widespread western form, occurring into central Texas; P. mathesoni, closely related, is a northern form found in New York, Michigan, and eastern Canada. Prilodexia obscura has a distribution from the Appalachi- ans to the Rocky Mountains and is nearly com- plimentary to that of P. agilis. Species included in the carolinensis group have members with a nonprojecting epistome and a nearly vertical anterior head profile. All adults have relatively short plumosity on the arista and the abdomen reddish laterally. Two of these species, P. halone and P. prexaspes, have limited east coast distributions. Ptilodexia ca- nescens occurs across the northern United States and Canada, while P. carolinensis is widely distributed from the east coast to the Rocky Mountains. Ptilodexia carolinensis is probably the oldest of the four, P. canescens, P. halone, and P. prexaspes being derived from it during the Pleistocene. The harpasa group is the most ambiguous of all, containing most of the Antillean and many Mexican species. Members of these species all have long antennae and long plumosity on the arista; some lack parafacial hairs. Ptilodexia ru- fipennis occurs from the east coast to the Rocky Mountains and across Canada; P. harposa is more restricted, P. arida is restricted to the Southwest and Mexico, while P. ponderosa is WILDER: NEARCTIC PTILODEXIA AGILIS CALIFORNICA CONJUNCTA — _— ie ee ee ee ee es pee oer os FIGURE 7. probably a West Indian species, with one record from Florida. Assuming that P. harpasa is the closest to the ancestor of the group, P. rufipen- nis became the most widespread and P. arida and P. ponderosa radiated in the southern lati- tudes. Ptilodexia major, the most primitive member of the next group shows slight similarities to specimens of P. harpasa. It ranges widely throughout the Midwest, Southwest, and Mex- ico. Ptilodexia incerta has an eastern distribu- tion almost exactly complementary to that of P. major, while P. maculata and P. flavotessellata are restricted in the Southwest and Midwest. These species all have members with short, pale, parafacial hairs. The next group, conjuncta, is probably de- rived directly from the ancestral agilis group and consists of only western species. Ptilodexia conjuncta, its most primitive member, ranges throughout the Rocky Mountains from Canada into Mexico and west to California. Prilodexia planifrons and P. contristans extend from the southwestern United States into Mexico; and P. westi, a close relative of P. contristans, is re- stricted to the extreme southern Midwest and the Southwest. OBSCURA CAROLINENSIS MAJOR HARPASA Inferred phylogeny of Prilodexia species groups. The last, or californica group, 1s related to the conjuncta group and probably had a similar or- igin. Its three species are confined to California and the West Coast, and members of these taxa show similarities only to members of the con- Juncta group. A graphic representation of the relationships of these species groups is given in Figure 7. Pos- sible events at the numbered branching points are as follows: 1. Major east-west split. Eastern group mem- bers with parafacial hairs extending low on the face: western group members with parafacial hairs high on the face. 2. Widening of face; lighter general color. Some females with yellow legs; lengthen- ing of plumosity on arista. Some advanced members radiating widely; radiation into Mexico and Antilles. 3. Lengthening of haustellum, increase in size, widening of face. Extensive radiation into Mexico. 4. Slight increase in size. Flattening of oral margin and anterior facial margin; short- ening of haustellum and plumosity on arista. 14 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 1 5. Shortening of haustellum. Radiation and isolation in southwestern United States and Mexican Pleistocene refugia. 6. Pleistocene isolation in California. De- crease in abundance of parafacial hairs and length of haustellum. ZOOGEOGRAPHY Although the dispersal powers of Diptera are relatively great, the distribution patterns seen in Ptilodexia seem to be dependent upon those of their hosts, the Scarabaeidae. Distribution of some Scarabaeidae are well known, and their possible histories have been discussed in several papers (Howden 1963; 1966). All statements made in this section are ten- tative. The patterns discussed are those of species of Prilodexia, but interpretations of those patterns are those which have been of- fered for some of the species of Scarabaeidae. No host specificity has been found, and it is only speculation that similar patterns of Prilodexia and their scarabaeid hosts are due to similar his- tories. Howden (1966) stated that North American species of Phyllophaga show a decline in num- bers from Georgia to Canada and from Texas or Arizona to Nebraska. This is true of Ptilodexia. He also stated that if certain areas of Texas were included with Arizona and New Mexico, there would be little overlap between the eastern and western faunas (approximate dividing line, 100th meridian). This holds true for the most part in Ptilodexia. However, a number of eastern species occur all the way into British Columbia in the northern parts of their ranges. As with Phyllophaga, many of the southern Arizona rec- ords represent the northern limits of Mexican species. When plotting the centers of distribution of species of Ptilodexia, it was noted that six species groups could be defined geographically. These were the same six groups which had been structurally and zoogeographically defined above. Although these groups show that the phylogeny presented herein has a zoogeograph- ical basis, they are not the best groupings for discussing zoogeography. I have categorized the species of Ptilodexia into six Zoogeographical groups, based on their complete distributions rather than centers of dis- tribution. The relationships of these categories may give insight into the historical zoogeogra- phy of the group. The first of these is an extreme northern pat- tern shown by P. canescens and P. mathesoni. The distribution is almost exclusively in areas which were previously glaciated. The range of P. mathesoni (Fig. 17) is restricted to Michigan, New York, and eastern Canada. Its ancestral and most closely related species, P. agilis, oc- cupies a large area from the Rocky Mountains west, extending eastward into Texas (Fig. 12). Ptilodexia canescens, from the carolinensis group, inhabits the northern United States and Canada from Newfoundland to British Colum- bia. In the East it extends southward only to the previous front of the Wisconsin glaciation, while in the West it extends southward into eastern Idaho, western Wyoming, and northern Utah (Fig. 67). The distribution of a species in pre- viously glaciated areas without representation south of the glacial front is fairly uncommon (Ross 1965). This deglaciated area may have of- fered considerable opportunity for expansion to certain Scarabaeidae and their Prilodexia para- sitoids. The next group has a widespread distribution, throughout the eastern United States into the plains states and, in some cases, even further west. Of these species, only P. harpasa (Fig. 82) lacks representation in the lower Midwest. The other species, P. carolinensis (Fig. 62), P. incerta (Fig. 103), P. obscura (Fig. 22), and P. rufipennis (Fig. 88) occur widely throughout the Midwest and the East. The remaining eastern distribution pattern is that of P. halone and P. prexaspes. Both of these species belong to the carolinensis group; they are closely related and complementary in distribution. Ptilodexia prexaspes occurs in Florida and along the Atlantic coast to Virginia (Fig. 77), while P. halone is found in Mississip- pi, Tennessee, and along the coast from Mary- land to New York (Fig. 72). It is possible that these relatively uncommon species are host-spe- cific parasites of some of the large, flightless scarabs found in the Southeast and discussed by Howden (1963). Two species, P. agilis and P. conjuncta, have large western ranges, the former from British Columbia to Texas and west to the Pacific coast (Fig. 12), the latter from Mexico to British Co- WILDER: NEARCTIC PTILODEXIA lumbia, west to the coast (excluding California), and eastward through Canada to Ontario (Fig. Daye Ptilodexia californica (Fig. 47), P. pacifica (Fig. 52), and P. sabroskyi (Fig. 57) have ranges which are restricted to the west coast of the United States. It is possible that some unique local populations are parasitic on the large flight- less genera of scarabs (such as Pleocoma) which survived in situ during the Pleistocene. The remaining distribution group is the most common in Ptilodexia, occurring in at least sev- en species. This is a southwestern distribution, with species which may have had Mexican re- fugia. Four of these are Mexican species whose ranges extend northward into the mountainous regions of Arizona and New Mexico, rarely into Utah and Idaho. These four are P. contristans (Fig. 37), P. planifrons (Fig. 32), P. maculata (Fig. 108), and P. arida (Fig. 93), all recently differentiated. It is possible that the ranges of many other Nearctic Mexican species also ex- tend into these areas, but specimens have not yet been taken by collectors. Two of this southwestern group, P. westi and P. flavotessellata, apparently do not range into Mexico. The former occurs broadly along the international boundary from central Arizona to eastern Texas and into Oklahoma and southern Kansas (Fig. 42), while P. flavotessellata occurs in northern New Mexico, Colorado, and Ne- braska (Fig. 113). The last species in the southwestern group is P. major. Its distribution is a combination of the ranges of the previous two groups, extending from Mexico (where it is widespread) into the mountains of Arizona, New Mexico, and Colo- rado and through Texas into the Plains in Ne- braska and Kansas (Fig. 98). Unlike those of Phyllophaga (Howden 1966), eastern species of Ptilodexia frequently occur from Georgia to southern Ontario; others range broadly across the northern part of the United States and Canada. I see this deviation as a re- sult of the vagility of these flies and the probable capability of developing in different hosts, fac- tors which may account for other deviations from typical scarabaeid distributions. Inferring the historical zoogeography of Prilo- dexia 1s extremely speculative. The genus, as we know it, probably evolved on this continent, its ancestor reaching the area via the Bering Land Bridge during the Tertiary. By the onset of the Pleistocene, most of the species were probably already established. Pleistocene cli- matic fluctuations must have affected the distri- bution patterns we see in the genus today. The eastern species in our fauna may have occupied southeastern Pleistocene refugia, most of them expanding westward in the north after the ice sheets retreated. Ptilodexia agilis seems to have been much more widespread at one time, one of the species derived from it being found only in the Northeast. While P. agilis may have had a wide refugium, P. conjuncta and P. californica perhaps survived the Pleistocene in Mexico and California, respectively, separated by the extensive desert barriers of the time. The three species endemic to California were prob- ably separated from the other species at a rela- tively early time, closely resembling each other considerably more than any other species. Their refugia were in central and southern California, and subsequent recolonization proceeded no further north than the southern limits of the ice sheet. The southwestern groups could have sur- vived the glacial periods in situ or in Mexican refugia. It appears that P. major was once a widespread species, extending well into Mexico before the Pleistocene and giving rise to many species there. The Southwest, including Texas and Arizona, has the largest number of endemics. In the warmer parts of the country such as these, more generations per year are possible, and evolution can proceed at a faster pace than in the north. This may account, in part, for the large number of endemics; it also helps explain the numerous divergent populations seen in California and Texas as well as the tremendous diversity of the genus in Mexico. Genus PTILODEXIA Brauer and Bergenstamm Ptilodexia BRAUER AND BERGENSTAMM, 1889:119 (Type- species, Ptilodexia carolinensis Brauer and Bergenstamm, 1889, by original designation.) Myoceropsis TOWNSEND, 1915:23 (Type-species, Rhynchio- dexia flavotessellata Walton by original designation.) Rhamphinina, authors, not Bigot. Rhynchiodexia, authors, not Bigot. Rhynchodexia, emend. Wulp, 1891. The genus Ptilodexia Brauer and Bergen- stamm is confined to the New World. It is best represented in the Neotropical region, as is the entire tribe Prosenini. 16 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 1 DiaGNosis.—Members of Prilodexia can be distinguished from those of all closely related species of Nearctic Prosenini by the following combination of character states: propleuron bare; facial carina sometimes well developed (but never broad, blunt, and separating the an- tennae); haustellum shorter than the head height; apical cell open or closed at wing margin; and infrasquamal setulae usually present. DESCRIPTION.—Color black to reddish, usu- ally with thorax dark and abdomen pale with dark longitudinal stripe. Head with face broad in profile, anterior margin usually vertical; epi- stomal margin somewhat projecting; frontal vitta strongly narrowed between eyes, often obliter- ated; facial tomentum heavy to sparse, dull to shining, color varied, but usually grayish; para- facial hairs varied, absent or present; carina long, not much deeper than width of third an- tennal segment; postocular setae long; one or two pairs of oral vibrissae; epistome generally projecting to some degree; haustellum rigid, var- ied, from 0.3 to 0.8 times head height; palpi long. Antennae with length of third segment varied from one to two times length of second; arista with length of plumosity more than length of second antennal segment. Thorax with propleu- ron bare; mesonotum strongly or weakly tomen- tose, usually indistinctly striped; three or four pairs of presutural and postsutural acrostichals: a tuft of small hairs on postalar wall. Wing length 2.5 times width; apical cell open or closed at wing margin; infrasquamal setulae present or absent. Legs dark, tibiae lighter in most cases, posterior tarsi very long, 1.5 times length of tib- ia; claws and pulvilli long. Abdomen broad, conical, tomentum usually in large irregular patches; numerous median discal and marginal setae on abdominal tergites: four abdominal ter- gites visible, lateral margins meeting ventrally. Genitalia slightly withdrawn, terminal, axis ver- tical; cerci and surstyli variously modified. Fe- male differs from male in following ways: frontal vitta wide with sides subparallel; eyes widely separated; profrons a little wider; frontal and peristomal setae not as abundant; vertex with few hairs or setae; postocular setae shorter and sparser; height of eves distinctly less; frontal orbital setae present (Fig. 6). Thorax with fewer setae and hairs and more heavily tomentose; thorax and legs frequently lighter in color with fewer and shorter major setae and hairs; tarsal claws and pulvilli much shorter. Abdomen broader, shorter, and much more heavily to- mentose; usually with fewer median discals and often lacking other setae; integumental color uniform brown or gray in many females, even when it is marked in males of the same species. Brauer and Bergenstamm erected the genus Ptilodexia in 1889 for the North American species P. carolinensis. There has been much confusion since that time regarding the limits of the genus. This is because several characters normally constant within tachinid genera vary among Ptilodexia species. These characters in- clude the presence or absence of parafacial hairs and infrasquamal setulae. Prior to 1889, Macquart and Walker described species belonging to Ptilodexia in the genus Dexia Meigen, while Bigot (1885) created the genus Rhamphinina for those species he de- scribed. Bigot thought that Rhamphinina, a neo- tropical genus, and Rhynchiodexia, one of his Australian genera, could be distinguished from one another by the presence or absence of a facial carina. Wulp (1891) considered this to be an inconsistent character. Wulp (1891) emended the name to Rhyncho- dexia, which he used for what we now call Ptilo- dexia. He felt that Rhynchodexia and Rham- phinina were congeneric and mistakenly placed one Mexican species, contristans, in Hystri- chodexia. West (1924; 1925), a North American worker, thought that the species with hairy parafacials belonged to Ptilodexia while those with bare parafacials were Rhynchodexia. Austen (1907) shared this opinion. Later, West (1950) agreed with Curran (1934) that the two were probably one and Rhynchodexia was the proper name for the complex. Reinhard (1943) stated that the name Ptilodexia was available for American species. Examination of species of Rhynchodexia, now Senostoma (Crosskey 1973a), shows that this genus differs from Ptilodexia in having a pronounced facial carina as well as numerous other differences which will be discussed later. Rhamphinina dubia, the type-species of the ge- nus, is not a Ptilodexia. The name Estheria tibialis Robineau-Desvo- idy is frequently used for species of Ptilodexia (Townsend 1921; Aldrich 1905; Austen 1907). The type of this species is lost, so we cannot know if E. tibialis belongs to Ptilodexia. How- ever, since a characteristic of Estheria is the WILDER: NEARCTIC PTILODEXIA presence in its members of a petiolate apical cell, and since this rarely occurs in Ptilodexia, I agree with previous workers who have chosen to reject the name E. tibialis. Key to the Nearctic Species of Ptilodexia la. lb. 2a. 2b. 3a. 3b. 4a. 4b. Parafacial hairs present, although very small and pale in some individuals; legs @ufemales varied in-color 92-22 2 2 Parafacial hairs absent; legs of females APEC OLOT: 2 2eee se ES toon ee | Infrasquamal setulae present ______-_------ 5 Infrasquamal setulae absent Parafacial hairs dark, long, present on most of parafacial (Fig. 66); length of plumosity on arista subequal to length of second antennal segment; face with traces or spots of brownish tomentum (northeastern U.S., trans-Canada, northern mountain states) oi ee canescens (Walker) Parafacial hairs pale and/or short, pres- ent only on upper anterior portion of parafacial; length of plumosity on arista at least 1.5 times length of second anten- nal segment (Fig. 5b); facial tomentum concolorous silvery gray or yellowish 4 Femora of members of both sexes brown or black (eastern U.S. to about 100th meridian) incerta (West) (in part) Femora of members of both sexes or- ange with definite black or brown patches on flexor surfaces, coxae also with dark patches (Arizona and New Mexico) maculata n.sp. Sa(2a). Flies pale colored; thorax, abdomen, Sb. 6a. 6b. Ta. 7b. and femora pale brown to orange, or width of depression between oral vibris- sae less than distance between oral vi- brissae and oral margin (Fig. 4a) Flies dark; thorax, abdomen, or legs brown or darker in color; width of depression between oral vibrissae equal to or greater than distance between oral vibrissae and oral margin (Fig. 4b) — 10 Integument of tarsi pale, concolorous with legs ventrally halone (Walker) Integument of tarsi brown or black -____- i Width of the depression between oral vibrissae greater than distance from vi- brissae to oral margin Width of depression between oral vibris- 8a. 8b. 9a. 9b. sae less than or equal to distance from vibrissae to oral margin (Fig. 4a) prexaspes (Walker) Parafacial hairs minute, confined to up- per anterior parafacial; haustellum length 0.6 times head height (Fig. 6) (southern Florida) ponderosa (Curran) Parafacial hairs long, scattered on para- facial; haustellum length no more than 0.5 times head height (southwestern WeSeandi Mexas) eee sh Se ae 9 Parafacial hairs dark, coarse and abun- dant (Fig. 61); femora or notum brown in many individuals; this color form un- common (Texas) _ carolinensis Bauer and Bergenstamm (in part) Parafacial hairs pale or light brown, fine, sparse (Fig. 112); femora and notum pale orange-brown (southwestern U.S. into Colorado and Nebraska) flavotessellata (Walton) 10a(Sb). Mediotergite polished immediately 10b. lla. Iii: beneath postscutellum; parafacial hairs distant from eye; haustellum 0.5 times head height or less (Fig.11); legs of fe- males yellow; species with members small, dark Mediotergite with at least a fine dusting of tomentum; parafacial hairs and haus- tellum varied Abdomen and thorax black, strongly shining; abdomen with tomentum evenly distributed; parafacial hairs black, coarse (Fig. 16); squamae of males dark brown (northcentral and eastern U.S.) _________ ee eis 2 ee mathesoni (Curran) Abdomen with tomentum in large irreg- ular patches; thorax with definite tomen- tose striping; parafacial hairs black, fine (Fig. 11); squamae of males white to pale brown (western U.S. and Texas) agilis Reinhard (in part) 12a(10b). Length of plumosity on arista less 12b. 13a. than or equal to 1.25 times length of sec- ond antennal segment (southwestern U.S. and Mexico) Length of plumosity on arista more than 1.25 times length of second antennal seg- ment (widespread ie ses 2s FFs 15 Abdomen and scutellum concolorous dark gray or brown, slightly lighter on ventral margins of tergites in a few in- 13b. 14a. 14b. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 1 dividuals: ninth tergite and parafacial hairs varied; haustellum length 0.7 to 0.8 times*head height (Fig. 31)! ---------- 14 Abdomen and scutellum distinctly red- dish laterally; ninth tergite with several strong setae (Fig. 33); parafacial hairs long, dark, occurring along parafacial ventrally to level of oral vibrissae; haus- tellum length 0.6 times head height (Fig. 36) contristans (Wulp) Parafacial hairs long, coarse, numerous, occurring on entire parafacial (Fig. 31) (Arizona, New Mexico into Mexico) _ eee tM ee planifrons (Wulp) Parafacial hairs fine, sparse, occurring only on anterior parafacial (Fig. 41) (Texas, Oklahoma, Kansas) westi n.sp. (in part) 1S5a(12b). Parafacial hairs short, pale or oth- IIb: 16a. 16b. 17a. 17b. 18a. 18b. 19a(17 ELrwise INCOMSPICUOUS: —- 2) = 2s 2 = 16 Parafacial hairs long, dark, although sparsely placed in some individuals _ 20 Mediotergite polished; legs of females yellowish; haustellum length no more than 0.5 times head height (Fig. 11); scu- tellum concolorous with rest of notum EU Sc le Pee agilis Reinhard (in part) Mediotergite tomentose to subshining; legs of females dark; haustellum length varied; scutellum lighter in color than rest of notum in most individuals —____- hz Parafacial hairs pale and, in most indi- viduals, small (Fig. 97) (west only to Ar- izona and Rocky Mountains) 18 Parafacial hairs light brown to black, short or medium in length (Pacific coast States) ee eae a 19 Infrasquamal setulae absent or greatly reduced in number in most individuals: parafacial hairs only on upper anterior parafacial (Fig. 102); length of palpi equal to or slightly less than half length of haustellum, broadened at tip (central and eastern U.S.)____ incerta (West) (in part) Infrasquamal setulae present in most in- dividuals; parafacial hairs extending ventrally on face to level of apex of an- tennae (Fig. 97); length of palpi greater than half length of haustellum, narrow (southwestern U.S. into Texas) 2 eI, ae tS major (Bigot) ). Third antennal segment subequal to 19b. 21b. 22b. 23b; or shorter than second segment; smallest distance between eyes of male subequal to width of frontal vitta at antennal base; female with parafacial hairs confined to area near antennae; facial tomentum dull (southern California) sabroskyi n.sp. Third antennal segment longer than sec- ond segment; smallest distance between eyes of male less than width of frontal vitta at base of antennae; females with parafacial hairs scattered, often along center of parafacial; facial tomentum shining (throughout California) californica n.sp. . Haustellum length greater than 0.6 times head height, thin, narrowed apically 21 . Haustellum length less than or equal to 0.6 times head height, broad and linear in: most individuals: .___-. = =e 24 . Length of plumosity on arista more than twice length of second antennal segment; haustellum length 0.65 to 0.7 times head height, narrow (Fig. 81) (northern and eastern U.S.) harpasa (Walker) Length of plumosity on arista less than or equal to twice length of second anten- nal segment; haustellum length varied (western U-S:) 22.2. === 22 . Parafacial hairs long, dark, and abun- dant, uniformly covering parafacial (Fig. 26); haustellum length 0.6 to 0.8 times head height, narrowed apically in most individuals; abdomen reddish laterally _ ie Sa ee ee conjuncta (Wulp) Parafacial hairs sparse (Fig. 41); haus- tellum length and abdomen varied Wit. we eae 23 . Length of haustellum more than 0.7 times head height, strongly narrowed apically; parafacial hairs sparse, located only on anterior portion of parafacial (Fig. 41); male abdomen dark brown or gray with little if any reddish color; ninth tergite with several strong setae (Fig. 38) (Texas and Oklahoma) Mee ee ee Sey ent westi n.sp. (in part) Length of haustellum less than 0.65 times head height, broad; not narrowed apically in most individuals; parafacial hairs occurring along center of parafacial (Fig. 46); male abdomen distinctly red- dish with longitudinal stripe; ninth ter- WILDER: NEARCTIC PTILODEXIA gite with hairs only (Pacific coast states) <<. californica n.sp. (in part) 24a(20b). Abdomen blackish with little if any orange coloration laterally; parafacial hairs inserted close to anterior margin of eye in most individuals; palpi very short, less than half length of haustellum (Fig. 21); wings of many males basally dark- ened obscura West . Abdomen, especially of male, with at least some red or orange coloration lat- erally; parafacial hairs not inserted close to anterior eye margin in most individu- als; palpi longer than half length of haus- tellum (Fig. 51); wings of males not ba- sally darkened . Femora of members of both sexes or- ange or marked with orange; abdominal tomentum concolorous (California)______ pacifica n.sp. Femora of members of both sexes brown or black (some specimens from Texas may have orange femora); abdominal to- mentum bicolored (widespread) —_____- 26 Epistomal angle of head not prominent (Fig. 61); females with 6 to 8 dorsal and lateral marginal setae on abdominal seg- ment III; males with | pair of dorsal mar- ginal setae on abdominal segment II; haustellum length 0.4 to 0.5 times head height (central and eastern U.S.) ________ Lae carolinensis Brauer and Bergenstamm (in part) . Epistomal angle of head prominent (Fig. 46); females with 10 to 12 dorsal and lat- eral marginal setae on abdominal seg- ment III; most males with 2 pairs of dor- sal marginal setae on abdominal segment II; haustellum length 0.5 to 0.65 times fread height: (western U:S:) -_2---__---____- californica n.sp. (in part) 27a(1b). Abdomen without any reddish or or- ange coloration laterally in most males; tip of abdomen and genitalia generally reddish yellow; facial tomentum strongly shining; length of plumosity on arista more than twice length of second anten- nal segment; face appearing narrow (Fig. 87); infrasquamal setulae absent in many individuals; femora of many males yel- low (central and eastern U.S. into New ple) Nm in jab} 25b. 26a. Mexico and British Columbia). rufipennis (Macquart) . Abdomen orange laterally in most males; tip of abdomen not noticeably lighter than rest of abdomen; facial tomentum rather dull; length of plumosity on arista at most equal to twice length of second antennal segment; face appearing broad (Fig. 92); infrasquamal setulae present; femora of males dark (Utah and southern Idaho south into Mexico) arida (Curran) The Nearctic Species of Prilodexia agilis Group Ptilodexia agilis Reinhard (Figures 8-12) Ptilodexia agilis REINHARD, 1943:22. SABROSKY AND AR- NAUD (1965:988). [HoLoTYPE, male, deposited in CNCI, labeled, College Station, Texas, 8 Oct. 1933, H. J. Rein- hard.] TAXONOMIC Notes.—Although I have not seen the holotype of P. agilis, members of this species are quite distinctive, and the original description is adequate to assure the identity of the specimens examined. Even though P. agilis was described from a disjunct population, the type-specimens are typical of the species. Type material is reported to be in excellent condition. DiaGNosis.—Ptilodexia agilis is a distinct species, members of which can be separated from their congeners by the following character states: size small; parafacial hairs fine, medium in length, inserted below apex of antennae or less than 0.25 mm from anterior margin of eye in only a few individuals; haustellum less than half head height; length of plumosity on the aris- ta 1.5 to 2.0 times length of second antennal segment; mediotergite polished; abdomen and scutellum entirely blackish; female with legs pale colored. MATERIAL EXAMINED.—Specimens exam- ined include 812 males and 488 females, data as listed by Wilder (1976). DISTRIBUTION.—Prtilodexia agilis ranges from Arizona and New Mexico north through Cali- fornia, the Great Basin, and the Rocky Moun- tains into Alberta and British Columbia. There are a few records from eastern Texas, where the topotypic population is found. BrioLoGcicAL Notes.—The flight period lasts from April to October, and adults can be col- 20 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 1 NORTH AMERICA Ficures 8-12. Ptilodexia agilis Reinhard. Fig. 8. Genitalia of male, posterior view. Fig. 9. Ejaculatory apodeme. Fig. 10. Genitalia of male, lateral view. Fig. 11. Head of male, lateral view. Fig. 12. Distribution of P. agilis. lected at any time during this period. July and August are the most frequent months of collec- tion, but local variation is common. In Califor- nia, for example, P. agilis adults are collected more frequently in September and October along the coast and in the south; but in the Sier- ra, northern California, and Oregon, June and July are the main periods of activity. This species inhabits both mountains and low- lands. Adults have been collected at elevations up to 2,600 m in Arizona, 3,800 m in California, above the 3,000-m level in Colorado, and fre- quently above 3,000 m elsewhere. Prtilodexia agilis adults have frequently been taken at low elevations in such areas as the San Joaquin Val- ley, the Great Basin, and eastern Texas. Spec- imens have been collected by UV light trap, Malaise trap, wind vane trap, and by sweeping foliage. Flowers visited include the Compositae Achil- lea Millefolium, Chrysothamnus viscidiflorus, Baccharis glutinosa, B. pilularis, Eriogonum nudum, Solidago trinervata, and Lepidospartum squamatum. Unidentified Compositae visited were Achillea sp., Baccharis sp., Solidago sp., Eriogonum sp., and Haplopappus sp. Other flowers from which P. agilis adults have been collected are Allium (Liliaceae) and Salix (Sal- icaceae). At two localities in Colorado, speci- mens were collected on Dwarf Mistletoe (Ar- ceuthobium cyanocarpum); and members of this species are believed to be pollinators of that plant. One specimen was collected in an emer- gence trap under a filbert tree in Oregon. There are no data on the life history of this insect. Its members are probably not host spe- cific, judging from the diverse assortment of habitats and wide geographical and temporal ranges of the species. Ptilodexia mathesoni (Curran) (Figures 13-17) Rhynchiodexia mathesoni CURRAN, 1931:93. West (1950:110); SABROSKY AND ARNAUD (1965:989). [LECTOTYPE (here des- ignated), male, deposited in CUIC, labeled, ** Douglas Lake, Mich., 24—VII-22’’/** Wing Slide, Cornell U., Lot. 919, Sub. 138, L. S. West’’/**d Holotype Rhynchiodexia mathesoni WILDER: NEARCTIC PTILODEXIA NORTH AMERICA Ficures 13-17. Ptilodexia mathesoni (Curran). Fig. 13. Genitalia of male, posterior view. Fig. 14. Ejaculatory apodeme. Fig. 15. Genitalia of male, lateral view. Fig. 16. Head of male, lateral view. Fig. 17. Distribution of P. mathesoni. Curran’’/** Holotype Cornell U. No. 1938’’/**Cornell U. Lot. 922, Sub. 40’’/*‘Lectotype Rhynchiodexia mathesoni Cur- ran designated by D. Wilder, 1976.’’] TAXONOMIC Notes.—Even though the label on the type-specimen reads ‘‘Holotype Rhyn- chiodexia mathesoni Curran,” this specimen is not a holotype. The author of the paper vali- dating the species made no mention of type ma- terial or of type-locality. The type label with Curran as author was put on the specimen at a later date; it was West’s “‘holotype,”’ not Cur- ran’s. The name P. mathesoni was proposed by West and validated by Curran in his 1931 key. I have designated West’s “‘holotype’’ as the lec- totype. It is a large specimen in excellent con- dition. There is one other specimen which I believe Curran had before him while writing his key. This specimen, also from Douglas Lake, Mich- igan, is deposited in AMNH and was collected on the same date as the specimen West labeled as holotype. I have designated this specimen as a paralectotype. It might be argued that this sec- ond specimen, because it is labeled with Curran as the author, was later sent to AMNH and not seen by Curran. However, the type labels of all five of the species validated in Curran’s key give Curran as the author. I believe that all these labels were changed at a later date, and since Curran makes no mention of material, I feel that these two identically labeled specimens are ac- tually syntypes. DiaGnosis.—Ptilodexia mathesoni is a dis- tinctive species, evidenced by the following combination of character states: body color black, shining; parafacial hairs long, black, in- serted ventral to apex of antennae or less than 0.25 mm from anterior margin of eye in only a few individuals; haustellum less than half head height in length; width of arista and plumosity 1.5 to 2 times length of second antennal seg- ment; mediotergite polished; wings dark in col- or; male abdomen with pollen evenly distribut- ed; female with pale-colored legs. MATERIAL EXAMINED.—Twenty-one males and eleven females were examined. 22 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 1 NORTH AMERICA Ficures 18-22. Ptilodexia obscura West. Fig. 18. Genitalia of male, posterior view. Fig. 19. Ejaculatory apodeme. Fig. 20. Genitalia of male, lateral view. Fig. 21. Head of male, lateral view. Fig. 22. Geographical distribution. DISTRIBUTION.—The species, although rec- ords are few, seems to range through the north- eastern United States and eastern Canada. There is one record from Victoria Beach, Man- itoba. Bro_oGicAL Notes.—The flight period is from April to October with the main period of activity between mid-June and mid-August. All records for months other than July and August are from Suffolk County, New York. There are no life-history data for P. mathe- sont. One male was collected on Solidago can- adensis. Ptilodexia obscura West (Figures 18-22) Ptilodexia obscura West, 1925:133. LEONARD (1928:822); CURRAN (1930:93); SABROSKY AND ARNAUD (1965:989). [HoLorype, female, deposited in SIIS, labeled, ** Wading River, L.I., June 29, 1917, W. T. Davis.’’] TAXONOMIC Notes.—West described P. ob- scura from three female specimens, one of which he designated holotype. The two para- types are so labeled and are deposited in CUIC. They closely resemble the holotype. There is a male specimen from Victoria Beach, Manitoba, deposited in AMNH, which bears a handwritten label reading, ““R. obscura West.” It appears that West recognized the male of the species at a later time, even though this particular male specimen differs considerably from the female type-series. West still used the generic name Ptilodexia for obscura in 1950, but because his concept of the genus changed, it is possible that he might have written ““R. obscura’’ instead of “P. obscura.’ It is doubtful that Curran labeled the specimen or ever looked at West’s types, since in his 1931 key, he describes P. obscura specimens as being over 12.5 mm long. DiaGnNosis.—Ptilodexia obscura is a fairly distinctive species. Its members may be sepa- rated from their congeners by the following combination of character states: face narrow, with parafacial hairs abundant, long, fine, dark, and inserted below lower edge of eye and close to its anterior edge; third antennal segment dark in most specimens, broadened apically; length of plumosity on arista 1.5 times length of second WILDER: NEARCTIC PTILODEXIA NORTH AMERICA Ficures 23-27. Ptilodexia conjuncta (Wulp). Fig. 23. Genitalia of male, posterior view. Fig. 24. Ejaculatory apodeme. Fig. 25. Genitalia of male, lateral view. Fig. 26. Head of male, lateral view. Fig. 27. Geographical distribution. segment; haustellum broad, length 0.55 to 0.6 times head height; palpi short, half haustellum length or less; notum covered with short, ap- pressed, grayish pollen, giving it a smooth, sub- shining appearance; legs of females brown; ab- dominal color, grossly appearing black, but actually rufescent laterally. MATERIAL EXAMINED.—One hundred forty- two males and 53 females of P. obscura were examined. DISTRIBUTION.—The range of P. obscura ex- tends from New Brunswick south into the Geor- gia Appalachians west into Arkansas, eastern Kansas and Nebraska, western South Dakota and Saskatchewan, and Manitoba. BIoLoGicAL Notes.—tThe flight period lasts from April to September with June and July the most common months of collection, a later av- erage seen only in Manitoba. Most specimens have been collected at low elevations, the ex- ceptions coming from 800—1,700 m in the Ap- palachians in Tennessee, North Carolina, and Virginia. Collecting methods yielding specimens of P. obscura include sweeping and Malaise trap. Specimens have been collected on the flowers of Solidago canadensis (Compositae), and Aruncus sp. and Spiraea latifolia, both Rosa- ceae. conjuncta Group Ptilodexia conjuncta (Wulp) (Figures 23-27) Rhynchodexia conjuncta WULP, 1891:228. ALDRICH (1905:499); GUIMARAES (1971:33). [LECTOTYPE (here designated), male, deposited in BMNH, labeled “‘Lectotype’’/**d°’/ “B.C.A. Dipt. II, Rhynchodexia conjuncta v.d.W.”’/**Cen- tral America. Pres. by F. D. Godman. O. Salvin. 1903- 172°*/*‘Ciudad, Mex., 8100 ft., Forrer’’/*“Lectotype Rhyn- chodexia conjuncta Wulp designated by D. Wilder, 1975.”’} Rhynchodexia simulans WULP, 1891:229. ALDRICH (1905:499); GUIMARAES (1971:34). [LECTOTYPE (here designated), male, deposited in BMNH, labeled, *‘Lectotype’’/"*N. So- nora, Mexico. Morrison’’/**3’’/**B. C. A. Dipt. II, Rhyn- chodexia simulans, v.d.W.’’/*‘Central America, pres. by F. D. Godman, O. Salvin. 1903—172’’/*‘Lectotype Rhyncho- dexia simulans Wulp, designated by D. Wilder, 1975.’"] NEw SYNONYMY Ptilodexia tibialis (partim): ALDRICH (1905:504). TAXONOMIC Notes.—Wulp described P. conjuncta from two male cotypes, one of which I have designated as lectotype. This specimen 24 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 1 is in good condition except for a few broken setae. Wulp described P. simulans and P. conjuncta in the same paper. The cotype (one of two) which I saw and designated lectotype is in good condition, although it has the dorsal setae and abdomen broken. Wulp realized that P. conjuncta and P. sim- ulans were very closely related. He separated members of each on the basis of size, curvature of the hind tibia, and some other minor char- acters. Size, of course, cannot be used effec- tively to separate these parasitic flies. A curved hind tibia is a character state which occurs fre- quently in members of many species of Prilo- dexia, especially in those of P. conjuncta, where it constitutes part of normal intraspecific varia- tion. DiAGNosis.—Ptilodexia conjuncta 1s a Vari- able species. Its members can be distinguished by the following: face wide; parafacial hairs long, fine, dark, abundant, inserted below level of lower edge of eye in some specimens and at least 0.12 mm from anterior edge of eye in all but a few specimens; haustellum long, ranging from 0.6 to 0.8 times head height, slender, nar- rowed apically in most specimens; carina fairly well developed; width of arista and plumosity 1.5 to 2 times length of second antennal seg- ment; oral margin distinctly protruding; scutel- lum and sides of abdomen reddish. MATERIAL EXAMINED.—Specimens exam- ined included 1,238 males and 629 females. DISTRIBUTION.—This species ranges from British Columbia and Alberta south through the Rocky Mountains into Mexico. There are scat- tered records from the plains states, the Great Basin, California, and Oregon. There are a few doubtful records from the eastern United States. BIOLOGICAL Notes.—The flight period ranges from February to October. Most of the activity occurs in July and August, earlier in the north- ern areas and later further south. In Arizona (and possibly Texas), P. conjuncta appears to have two broods, one in March and April, and another in August and September. Ptilodexia conjuncta adults are generally found in mountainous areas at altitudes from 1,500 to 2,750 m and in some areas up to 3,660 m. They are also, although less commonly, col- lected at low elevations in coastal as well as in- land areas. Label data indicate that they have been collected in meadow sweeps, in a meadow in spruce-fir zone (2,750 m), in pine-spruce-as- pen zone, and above timberline. Productive col- lecting methods for P. conjuncta specimens are UV light and Malaise trap. Specimens of P. conjuncta have been col- lected from many flowers. These include: Se- necio salignus, Heliopsis parvifolia, Cacalia de- composita, Encelia farinosa, Geraea canescens, Chrysothamnus greeni, Achillea Millefolium; and unidentified species of Encelia sp., Gutier- rezia sp., Solidago sp., Baccharis sp., Helian- thus sp., Senecio sp., Eriogonum sp., Achillea sp., Bigelovia sp., Aster sp., and Rudbeckia sp. (all Compositae). Other flowers visited include, Ceanothus fendleri and unidentified Ceanothus sp. (Rhamnaceae); Arctostaphylos sp. (Erica- ceae); Melilotus sp., and Dalea sp. (Fabaceae); Lippia wrightii (Verbenaceae); and Arceuthob- ium sp. (Loranthaceae). Ptilodexia conjuncta adults, along with those of P. agilis, have been observed pollinating Dwarf Mistletoe (Arceu- thobium cyanocarpum). Ptilodexia planifrons (Wulp) (Figures 28-32) Rhynchodexia planifrons WULP, 1891:234. ALDRICH (1905:499); GUIMARAES (1971:33). [HoLOTYPE, male, deposited in BMNH, labeled, ‘“‘Holotype’’/*‘Ciudad, Mexico, 8100 ft., Forrer’’/**3°’/*B. C. A. Dipt. Il, Rhynchodexia planifrons, v.d.W’’/*‘Central America pres. by F. D. Godman, O. Sal- vin. 1903-172.”"] Dexia harpasa (partim): ALDRICH (1925:114). (Misidentifica- tion). TAXONOMIC Notes.—The holotype of this species, deposited in BMNH, is in poor condi- tion, but is still recognizable. The facial band is obscured, the tibiae are quite light and distinctly curved (a frequently encountered anomaly in species of Prtilodexia), and the striping on the notum is more distinct. Labels on the specimen say only “Ciudad, Mexico, 8100 ft.,’> but in the description, the origin of this specimen is stated as *‘Ciudad in Durango, 8100 ft.”’ I follow Wulp’s original pub- lication in calling Durango the type-locality. DIAGNosIs.—Ptilodexia planifrons is a dis- tinctive species and its members can be identi- fied by the following character combination: face wide; presence of a contrasting tomentose diagonal band extending from antennal base to eye margin; parafacial hairs dark, coarse, abun- dant, inserted lower than level of oral vibrissae only in a few specimens; antenna with plumosity WILDER: NEARCTIC PTILODEXIA 28 5 NORTH AMERICA Ficures 28-32. Ptilodexia planifrons (Wulp). Fig. 28. Genitalia of male, posterior view. Fig. 29. Ejaculatory apodeme. Fig. 30. Genitalia of male, lateral view. Fig. 31. Head of male, lateral view. Fig. 32. Geographical distribution. on arista less than or equal to length of second antennal segment; carina long, slightly keeled and prominent; two pairs of oral vibrissae in most specimens; frontal vitta very wide at an- tennal base; haustellum long, 0.7 to 0.85 times head height, narrowed apically; scutellum and abdomen entirely dark gray or black. MATERIAL EXAMINED.—One hundred sev- enty-three males and 70 females of P. planifrons were examined. DISTRIBUTION.—This species ranges from Flagstaff south through the mountains of central and southeastern Arizona, through the central mountainous region of New Mexico, the west- ern tip of Texas, and south into Durango, Mex- ico. There is one record from Colorado; how- ever, no exact locality is given. BIOLOGICAL Notes.—The flight period lasts from August through October, with the majority of records from mid-August to mid-September. The earliest seasonal record is one specimen collected on 27 June (error?) from El Paso, Tex- as, in 1921, and the latest is a series of 24 females collected on 22 October 1964, in Cochise Coun- ty, Arizona. Ptilodexia planifrons adults generally occur in the mountains and have been collected at many elevations between 1,370 and 3,350 m. Flowers visited include Gutierrezia sarothrae (1,800- 2,440 m, Apache County, Arizona), Heliopsis parvifolia (2,590 m, Chiricahua Mts., Cochise County, Arizona), Solidago trinervata (Sierra Madre, 2,230 m), and Rudbeckia sp. in Chihua- hua, all Compositae. All except two of the flower-visiting flies were males (one female col- lected on Heliopsis sp. and another at Rudbeck- ia sp.). One specimen was collected at a light. Ptilodexia contristans (Wulp) (Figures 33-37) Hystrichodexia contristans WULP, 1891:221. [HOLOTYPE, male, deposited in BMNH, labeled, *‘Holotype’’/**Omil- teme, Guerrero, 8000 ft., July, H. H. Smith’’/**3"’/"'B. C. A. Dipt. II., Hystrichodexia contristans, v.d.W.’’/**Central America, Pres. by F. Godman, O. Salvin. 1903-172.’’] Rhynchodexia punctipennis WULP, 1891:233. ALDRICH (1905:499); GuIMARAES (1971:33). [LECTOTYPE (here des- ignated), male, deposited in BMNH, labeled, *‘Cotype’’/ ‘*Sierra de las Aguas Escondidas, Guerrero, 9500 ft., July. H. H. Smith’’/**3’’/"B. C. A. Dipt. Il. Rhynchodexia punc- tipennis, v.d.W.’’/*‘Central America. Pres. by F. D. God- man, O. Salvin. 1903-172’’/**Lectotype Rhynchodexia 26 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 1 Ficures 33-37. Ptilodexia contristans (Wulp). Fig. 33. Genitalia of male, posterior view. Fig. 34. Genitalia of male, lateral view. Fig. 35. Ejaculatory apodeme. Fig. 36. Head of male, lateral view. Fig. 37. Geographical distribution. punctipennis Wulp designated by D. Wilder 1975.”’] New SYNONYMY. Ptilodexia constrictans (Wulp): GUIMARAES (1971:33) [lapsus calamus |. TAXONOMIC Notes.—The holotype of this species is in fair condition except for a broken thorax and an abdomen which is oily, obscuring the tomentum patterns. Wulp described Rhynchodexia punctipennis and Hystrichodexia contristans in the same pa- per. He felt that the two genera differed by two superficial characters, the general body shape and the hairs of the abdomen. The lectotype of P. punctipennis, although in poor condition is certainly a teneral specimen of P. contristans. DIAGNosIs.—Specimens of P. contristans can be separated from those of related species by the following combination of characters: face very wide; parafacial hairs long, dark, abundant, occurring to, and in most specimens, also below level of lower eye margin; third antennal seg- ment broadened apically in some individuals; plumosity on arista short, its width subequal to length of second antennal segment; carina short but well developed; haustellum narrow, equal to 0.6 times head height; oral margin strongly pro- truding; scutellum and sides of abdomen reddish laterally, wing base distinctly darkened; ninth tergite with several strong setae. MATERIAL EXAMINED.—Ninety-one males and 20 females were examined. DISTRIBUTION.—This species ranges from the central and southeastern mountains of Arizona southward into the mountains in Durango, Ve- racruz, and Guerrero. There is one record from New Mexico. BIoLoGIcAL Notes.—The flight period is from mid-August to early September, with a few scattered records in July. The holotype was col- lected in July. All specimens were collected in mountainous areas. The lowest elevation indicated on labels is 2,130 m, in Veracruz, the highest, 2,900 m, in Guerrero. Specimens have been collected on the following Compositae: Heliopsis parvifolia, Ca- calia decomposita, Verbesina encelioides, He- WILDER: NEARCTIC PTILODEXIA NORTH AMERICA Figures 38-42. Prilodexia westi, n.sp. Fig. 38. Genitalia of male, posterior view. Fig. 39. Ejaculatory apodeme. Fig. 40. Genitalia of male, lateral view. Fig. 41. Head of male, lateral view. Fig. 42. Geographical distribution. lenium hoopesii, and unidentified Senecio sp., Cirsium sp., and Solidago sp. There are no as- sociated host data. Ptilodexia westi, new species (Figures 38-42) Type-LocaLity.—The holotype was collected at Imperial, Texas, 10 April 1954 by L. D. Beamer. TypPE-SPECIMENS.—The male holotype is deposited in CNCI, the allotype, from Las Cruces, New Mexico, in USNM. Complete data from these specimens and the 21 para- types are listed below. DIAGNosIs.—This is a distinctive species, its members easily separable from those of other species in the genus by the following combina- tion of character states: face wide, tomentum grayish, very heavy; parafacial hairs from short to medium in length, occurring only on anterior half of parafacial, below apex of antennae in a few individuals; length of haustellum 0.7 to 0.8 times head height, narrowed apically, length of plumosity on arista varied, from | to 1.25 times length of second antennai segment; abdomen heavily tomentose, in males integument dark gray with no median stripe, in females brown or rufous; ninth tergite of males with several strong setae; genitalia unlike those of adults of any oth- er species. DESCRIPTION.—Male: Total body length 10.5 mm, grayish. Head height 2.6 mm; profrons 0.25 of head length. Facial tomentum very heavy, dull, grayish white, obscuring integumental col- or; parafrontal hairs long, sparse, dark; parafa- cial hairs medium in length, dark, rather sparse, located along anterior half of parafacial ventrally to slightly beyond apex of antennae. Width of frontal vitta at antennal base 0.4 mm, at narrow- est part, 0.1 mm; carina strongly developed, slightly keeled, broad, not depressed below apex of antennae; facial cavities strongly darkened. Height of gena 0.25 times head height. One pair of strong ocellar setae, one pair of smaller post- ocellars and postverticals; one pair of strong and one pair of weak inner verticals; outer verticals subequal to postorbitals. Fifteen pairs of medi- um-length postorbital setae, some a little shorter 28 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 1 than others; ventral to postorbitals is an irreg- ular row of dark hairs merging into dense whitish occipital hair. Seven pairs of frontal setae; one pair of oral vibrissae 0.3 mm from oral margin, the shallow depression between them 0.4 mm wide; 9 additional oral setae, one above vibris- sae quite strong, peristomals gradually increas- ing in length with distance from vibrissae. Epi- stome not protruding, its width twice length of first two antennal segments. Haustellum narrow, length 0.7 times head height, narrowed apically; palpi testaceous, narrow, length 0.5 times haus- tellum length. Antennae dark testaceous, third segment brownish apically, arista and plumosity dark brown. Third segment of antenna 1.5 times length of second antennal segment. Thorax: Mesonotum dark brown with heavy gray tomen- tum; striping obscure. Integument of scutellum and postalar calli concolorous with rest of no- tum. Six humeral setae, 2 posthumerals and 1 presutural, 3 pairs of discal scutellars. Pleura brown, sutures lighter, tomentum grayish; infra- squamal setulae present; squamae whitish, wings light brown; epaulet light brown, basi- costa testaceous; mediotergite tomentose. Legs dark reddish brown, tibiae lighter apically. An- terior femur with 7 dorsal setae, 8 posterodor- sals, 13 posteroventrals; tibia with slightly dark- ened dorsal line. Midlle femur with 5 short anterior setae in 2 rows. Posterior femur with 8 anterodorsals, 5 anteroventrals, 6 posteroven- trals, and no anterior setae. Femora with color uniform, tibiae darkened on basal third, tarsi black. Abdomen dark brownish black with heavy gray tomentum which is slightly brownish around bases of median marginal setae on sec- ond tergite and along posterior margins of third tergite. First syntergite with | pair of lateral marginal setae. Second tergite with | pair of me- dian discals, | pair of median marginals, and | pair of lateral marginal setae. Third tergite with | pair of median discal setae, and 8 dorsal and lateral marginal setae. Genitalia: Eighth tergite with sparse small brown hairs. Ninth tergite long, covered with long dense brown hairs and many long, strong setae; lateral swelling absent. Surstylus small, narrow, lateral angle smoothly rounded, internal depression absent, mesal sur- face slightly convex with strong hairs; basally with strong setae, in profile, elongate, heart- shaped, inclined strongly in posteroventral di- rection, partially obscuring cerci. Cerci with lat- eral lobes angular, only slightly differentiated from arms; arms wide, height more than twice that of lateral lobes; arms extending ventrally well past apex of surstyli; in profile, smoothly but strongly incurved with no bulge, tips point- ed, facing in anteroventral direction. Ejacula- tory apodeme with stem distinct from bulb and about 1.3 times its length; bulb open on part of one side and top. Female: Similar to male except for usual sexual differences and the following. General color reddish brown. Width of frontal vitta at narrowest point 0.3 mm, at antennal base 0.4 mm, narrowest point near antennal base, strongly widened to vertex. Height of gena 0.25 times head height. Nine pairs of stout postorbital setae with smaller setae irregularly interspersed; six pairs of frontal setae. Notum brown with to- mentum heavy; six strong humeral setae; squa- mae whitish; pleura light reddish brown. Legs lighter in color than those of male, tibiae simi- larly marked. Anterior femur with 10 postero- ventrals, 8 posterodorsals, and 10 dorsal setae. Middle femur with 5 strong anterior setae in 2 rows, | anteroventral, and 5 posteroventrals. Posterior femur with 6 anterodorsals, 4 postero- ventrals, 5 anteroventrals, and no anterior se- tae. Abdomen brownish, lighter laterally, cov- ered with heavy grayish tomentum. First syntergite with | pair of lateral marginal setae. Second tergite with | pair of lateral marginals and | pair of median marginal setae. Third ter- gite with | pair of median discals and 10 dorsal and lateral marginals. VARIATION.—Ptilodexia westi adults are quite distinct, but even among the small number of specimens examined, a certain degree of varia- tion is seen. Total body length ranges from 10 to 12 mm. Facial tomentum is yellow or gray in a few specimens; parafacial hairs do not extend to below the antennal apex in most individuals, and the hairs are varied in length, from short to long. The carina is shorter and not visible be- yond the apices of the antennae in some speci- mens. There are two rows, instead of one, of dark hairs ventral to the postorbitals in a few adults. The haustellum length is varied, from 0.7 to 0.8 times the head height. The antennae are also varied, from rufotestaceous to light brown, with the third segment not darkened. The wings of some specimens are light in col- or. The femora are darkened apically on all ex- cept the dorsal surfaces in some individuals; this is especially noticeable in females, where ground color of the femora can be pale in color. WILDER: NEARCTIC PTILODEXIA NORTH AMERICA f. rr Ficures 43-47. Ptilodexia californica, n.sp. Fig. 43. Genitalia of male, posterior view. Fig. 44. Ejaculatory apodeme. Fig. 45. Genitalia of male, lateral view. Fig. 46. Head of male, lateral view. Fig. 47. Geographical distribution. In a few specimens, the abdomen is without brownish tomentum, and the integument is red- dish brown ventrally and laterally. The abdomen of many females is reddish with a vague narrow median stripe, and in most females, it has no median discal setae. MATERIAL EXAMINED.—Fifteen males and 8 females were examined from the following localities. Arizona: Pima Co.: 1g, Brown’s Canyon, Baboquivari Mts., 18 Aug. 1955, G. Butler (UAIC). Kansas: Clark Co.: 19, Sitka, 12 June 1960, W. Van Velzen (MSUC). New Mexico: Dona Ana Co.: 1°, Las Cruces, 21 Sep. (USNM). Oklahoma: Woodward Co.: 136, Range 1 mile NW Supply, 1 June 1961, D. Bryan, 7d d- 322,9 June, 25532 2, 15 June (OSEC). Texas: Brazos Co.: 13, College Station, 30 May 1936, student collector (TAMU). Pecos Co.: 16, Fort Stockton, 28 Sep. 1935 (TAMU); Id, Imperial, 10 Apr. 1954, L. Beamer (CNCI). Uvalde Co.: 16, Uvalde, 18 May 1914, Bishopp (USNM). Although few specimens were available for study, this species appears to be the most un- usual Prilodexia in the Nearctic region. Its members are, in fact, so dissimilar from those of other species that it may ultimately be as- signed to a new genus. BIoLoGIcAL Notes.—The flight period stretches from April to September, with too small a sample available to generalize on peak activity. There is no information available re- garding host, collecting techniques, or ecology of this species. DERIVATION OF NAME.—This species is named in honor of Dr. L. S. West, who did much of the early work with this genus in the United State. californica Group Ptilodexia californica, new species (Figures 43-47) Type-LocaALity.—Two miles (3.2 km) S of Luther Pass, Alpine County, California. Type-SPECIMENS.—The male holotype and the allotype are deposited in the collection of the California Academy of Sci- ences. The holotype bears the CASC type number 12571. Complete data from these specimens and the 52 male and 37 female paratypes are listed below. DiaGnosis.—Ptilodexia californica adults can usually be distinguished from those of other species in the genus using the following combi- nation of character states: facial tomentum shin- 30 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 1 ing, dark brown or pale hairs scattered sparsely along center of parafacial; haustellum length 0.5 to 0.65 times head height; third antennal seg- ment longer than second, and length of plumos- ity on arista equal to 1.4 times length of second antennal segment; frontal vitta obliterated at narrowest point; scutellum and abdomen of males reddish, legs of both sexes dark. DESCRIPTION.—Male: Total body length 13.5 mm, dark brown with abdomen reddish lat- erally. Head height 3.3 mm; profrons 0.3 times head length. Facial tomentum whitish, shining, obscuring integumental color; parafrontal hairs long, dark; parafacial hairs dark, rather long, scattered along middle of parafacial. Width of frontal vitta at antennal base 0.5 mm, at narrow- est point, obliterated; carina fairly well devel- oped, short; facial cavities brown. Height of gena 0.25 times head height. One pair of strong ocellar setae; 2 pairs of postocellars, | pair of postverticals, inner and outer verticals all sub- equal. Thirteen pairs of postorbitals, frequently interspersed with smaller hairs; between post- orbitals and the whitish occipital hair is an ir- regular row of dark setae. Twelve pairs of fron- tal setae; one pair of oral vibrissae 0.3 mm from oral margin, the depression between them 0.4 mm wide; 18 additional oral setae, at least one above vibrissae and peristomals subequal. Epi- stome strongly protruding, its width 1.6 times length of first two antennal segments. Haustel- lum long, of medium width, 0.6 times head height; palpi long, narrow, testaceous, length 0.5 times haustellum length. Antennae dark rufotes- taceous, third segment, arista and plumosity dark brown. Third segment of antenna 1.4 times length of second segment; length of plumosity on arista 1.4 times length of second antennal segment. Thorax: Mesonotum dark brown with fine, dull-gray pollen, striping indistinct; integ- ument of scutellum and postalar calli rufotesta- ceous. Six humeral setae, | posthumeral and 2 presuturals, 3 pairs of discal scutellars. Pleura dark brown with fine grayish tomentum; infra- squamal setulae present; squamae whitish, wings pale brown; epaulet dark brown, basi- costa testaceous; mediotergite tomentose, sub- shining. Legs dark brown, tibiae rufotestaceous darkened slightly basally and apically; tarsi blackish. Anterior femur with 9 dorsal setae, 8 posterodorsals, and 21 posteroventrals; tibia with dark dorsal line. Middle femur with 3/4 an- terior setae in 2 rows. Posterior femur with 11 anterodorsals, 8 anteroventrals, 5 posteroven- trals, and | anterior seta. Abdomen rufous with narrow, dark-brown median stripe; tomentum shining white, shining brown around median marginals and posterior median discals of sec- ond and third tergites. First syntergite with 1 strong pair of lateral marginal setae. Second ter- gite with 3 pairs of median discals, 2 pairs of median marginals, 2 pairs of lateral marginals, and 2 pairs of lateral discal setae. Third tergite with 2 pairs of median discals, 12 dorsal and lateral marginals, and 6 pairs of miscellaneous discal setae. Genitalia: Eighth tergite with sparse long hairs. Ninth tergite with long brown hairs, lateral swelling absent. Surstylus large with lateral angle fairly sharp, depression deep; surface strongly pitted. Cerci with height of lat- eral lobes greater than that of arms; lateral lobes slightly attenuated dorsally, mesal margins con- cave; in profile, arms strongly bulging. Ejacu- latory apodeme with bulb distinct from stem, which is twice height of bulb. Bulb open on one side and on top, where it is distinctly lipped. Female: Similar to male except for usual sexual differences and the following. Width of frontal vitta at narrowest point 0.4 mm, at antennal base 0.6 mm. Height of gena 0.3 times head height. Eleven pairs of short postorbital setae; 10 pairs of frontal setae. Legs colored similarly to those of male, tibiae not as pale. Anterior femur with 14 posteroventrals, 8 posterodorsals, and 6 dor- sal setae. Middle femur with 5 anterior setae in 2 rows. Posterior femur with no anterior setae, 5 anterodorsals, 5 posteroventrals, and 6 antero- ventrals. Abdomen dark reddish brown; median stripe broad and indistinct. First syntergite with 1 pair of lateral marginal setae. Second tergite with | pair of median marginals, | pair of median discals, and | pair of lateral marginal setae. Third tergite with 2 pairs of median discals, 2 pairs of lateral discals, and 12 dorsal and lateral marginal setae. VARIATION.—Ptilodexia californica adults show relatively little variation in many of the diagnostic characters of this group. However, they do seem to exhibit a distinctly greater than usual degree of variation in minor characters as well as in the genitalia. It is possible that later, more-detailed studies may uncover cryptic or incipient species within P. californica. There are also a few specimens which appear to be WILDER: NEARCTIC PTILODEXIA intermediate between P. californica and P. sa- broskyi, and between P. californica and P. pa- cifica. Total body length ranges from 11 to 16 mm. In a few individuals, facial tomentum has a yel- lowish cast and the carina is keeled. Most spec- imens have no small hairs interspersed between the postorbital setae. The haustellum is varied, with the length ranging from 0.5 to 0.65 times head height. Notal tomentum may be partially or wholly dull brown and the mediotergite can be fairly shiny. The abdomen is varied from the state shown in the holotype to dark reddish brown with the median stripe broad and indis- tinct. The genitalia are extremely varied. MATERIAL EXAMINED.—Fifty-three males and 39 females were examined from the following localities. California: Al- pine Co.: 16, 2 miles [3.2 km] S Luther Pass, 6 July 1959, at flower Achillea Millefolium, P. Arnaud (CASC); 1¢, Hope Valley, 9 July 1948, J. MacSwain (CISC). Fresno Co.: 12, Prather, 29 June 1956, R. Schuster (CISC); 22 2, Watts Val- ley, 22-23 June 1956, R. Schuster (CISC). Inyo Co.: 12, W Fork Coyote Creek, 2.1 miles [3.4 km] NE Coyote Lake, 37°14°N, 118°30°W, Sierra Nevada, 9,840 ft [ca. 3,000 m] sweeping, 16 Sep. 1969, S. Frommer (UCRC); 1d, near Mono Pass, 12,000 ft, [ca. 3,658 m] 19 Aug. 1956, C. MacNeill (CISC); 1¢ Panamint Mts., 28 May 1937, B. Brookman (CASC). Kern Co.: 16, 4 miles [ca. 6.4 km] N Muroc, 30 Apr. 1950, T. Leigh (CISC); 26 6, Short Canyon, 7 miles [ca. 11 km] NW Inyokern, 13 Apr. 1954, J. MacSwain (CISC); 1d, Walker Pass, 11 June 1962, G. Bohart (USUC). Los Angeles Co.: 36 6222, Oct., Coquillett (USNM); 52 2366, Crystal Lake, 29 June 1950, W. Bentnick, 13¢d¢1122, 9 July 1952, R. Wagner (CISC, LACM); 16, Little Rock, 10 May 1941, J. Wilcox (PHAC); 1¢12, 1 mile W Little Rock, 13 May 1956, J. MacSwain (CISC); 1d, 2 miles [ca. 3.2 km] W Pearblossom, 12 May 1956, E. Linsley (CISC); 12, Rock Creek, 21 July 1936, A. Basinger (CASC). Mariposa Co.: 16, Yosemite Val- ley, 26 June 1921, E. Van Dyke (CASC). Mono Co.: 1612, Sardine Creek, 27 June 1957, D. Flaherty (UCRC, UAIC); 13, Sonora Pass, 9,000—10,000 ft [ca. 2,740-3,050 m], 16 July 1951, P. Ashlock, 12, 10 Aug. 1960, C. Toschi (USNM, CISC). Nevada Co.: 12, Prosser Dam, 15 July 1966, D. Miller (UCDC); 1d, 7 miles [ca. 11 km] SE Truckee, 24 June 1954, G. Schaefers (CISC). Plumas Co.: 26 612 (Allotype), Mead- ow Valley, 10 June 1924, E. Van Dyke (CASC). Riverside Co.: 16, Hemet, 26 Apr. 1961, Ewart (UCRC); 1d, Hidden Valley, Joshua Tree National Monument, 18 May 1946, A. Melander (USNM); 1d, Pinon Flat, 24 Apr. 1950, C. MacNeill (CISC); 1d, Pinon Flat, San Jacinto Mts., 18 May 1939, E. Ross, 22 2, Eriogonum, 21 May 1940, H. Reynolds, 1d, 27 May 1939, B. Brookman (CISC, UCRC); 1d, Pinon Flat, San- ta Rosa Mts., 27 May 1941, D. Knull (OSUC). San Bernardino Co.: 16, Helendale, 16 May 1955, W. Richards (CNCI). San Diego Co.: 12, 16.5 miles [ca. 26.6 km] N of Carlsbad, Tim- berlake (UCRC). San Luis Obispo Co.: 16, Oso Flaco Lake, 19 July 1964, M. Irwin (UCRC). Santa Barbara Co.: 12, Jan- ama Beach, 9 July 1965, J. Powell (CISC); 12, Los Prietos, 23 June 1965, J. Powell (CISC). Santa Cruz Co.: 1d, Bear 31 Valley, Santa Cruz Mts., July 1913, F. Clark (CUIC). Siskiyou Co.: 266, Hebron Summit, 10 miles [ca. 16 km] S of Dorris, 5,300 ft [ca. 1,600 m] 27 July 1953, W. Gertsch (AMNH). Tehama Co.: 16322, Deer Creek, 6 July 1952, M. Cazier (AMNH). Ventura Co.: 12, Quatal Canyon, NW corner Ven- tura Co., Stanleya pinnata, 9 May 1959, J. Powell (CISC); 2366, Hungry Valley, 5 miles [ca. 8 km] S of Gorman, 6 May 1959, C. O’Brien (CISC). Nevada: Clark Co.: 12, Kyle Canyon, Charleston Mts., 5,200 ft [ca. 1,580 m] 4 June 1941, Timberlake (UCRC). Ormsby Co.: 12, 6 July, Baker (USNM). Oregon: Hood River Co.: 136, Hood River, 17 June, Childs (CISC). Jackson Co.: 12, Colestin, 31 July 1918, E. Van Duzee (CASC). Klamath Co.: 16; Eagle Ridge, Klamath Lake, 20 May 1924, C. Fox (CISC). Washington: Yakima Co.: 12, Signal Peak Ranger Stn., 4,000 ft [ca. 1,200 mJ], 15 July 1933, J. Wilcox (PHAC). BioLtoGcicAL Notes.—The flight period lasts from April to October with most of the activity from April to July. Adults have been collected at low as well as high elevations (up to 3,660 m). Flowers from which they have been collected are Eriogonum sp. (Polygonaceae) and Stanleya pinnata (Cruciferae). Ptilodexia pacifica, new species (Figures 48-52) Type-LocaLity.—Point Pinos, Pacific Grove, Monterey County, California. TyPE-SPECIMENS.—The holotype, a male, and the allotype are deposited in the collection of the California Academy of Sciences. The holotype bears the CASC type number 12572. Complete data from these specimens and the 62 male and 16 female paratypes are listed below. DiaGnosis.—Ptilodexia pacifica is a complex species, but specimens can generally be sepa- rated from those of other species in the genus by the following combination of character states: parafacial hairs long, dark, inserted along center of parafacial as low as ventral eye margin; haustellum length varied, from 0.4 to 0.55 times head height; legs of most adults either pale in color or marked with rufotestaceous; notum of most specimens covered with flat, shiny, brown- ish tomentum, giving it a coppery appearance; abdominal tomentum concolorous—either whit- ish or brownish in most individuals—white marked with brown in only a few. DESCRIPTION.—Male: Total body length 10.5 mm, dark brown, abdomen reddish. Head height 2.75 mm; profrons 0.25 times head length. Facial tomentum heavy, shining grayish, ob- scuring integumental color; parafrontal hairs long, dark, sparse; parafacial hairs long, dark, rather sparse, inserted along middle of parafacial to ventral margin of eye. Width of frontal vitta 32 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 1 FIGURES 48-52. NORTH AMERICA Ptilodexia pacifica, n.sp. Fig. 48. Genitalia of male, posterior view. Fig. 49. Ejaculatory apodeme. Fig. 50. Genitalia of male, lateral view. Fig. 51. Head of male, lateral view. Fig. 52. Geographical distribution. at base of antenna 0.4 mm, at narrowest part obliterated; carina low, broad, rounded; facial cavities dark brown. Height of gena 0.3 times head height. One pair of strong ocellar setae, 3 pairs of postocellars, 1 pair of postverticals, in- ner verticals, and outer verticals. Fifteen pairs of long postorbital setae, not interspersed with smaller hairs; between postorbitals and whitish occipital hairs are a few irregular dark setae. Thirteen pairs of frontal setae; 1 pair of strong oral vibrissae 0.2 mm from oral margin, the depression between them 0.4 mm wide. Thirteen additional oral setae; one above vibrissae and peristomals subequal to each other; oral margin protruding, its width equal to 2.2 times length of first two antennal segments. Haustellum of me- dium width, length 0.5 times head height; palpi long, rufotestaceous, the length 0.6 times haus- tellum length. Antennae rufotestaceous, third segment, arista and plumosity brown. Third seg- ment of antenna 1.5 times length of second seg- ment; length of plumosity on arista 1.75 times length of second antennal segment. Thorax: Mesonotum dark brown with tomentum appear- ing grayish from one angle and shiny brownish from others; striping indistinct. Integument of postalar calli, scutellum, and sides of notum ru- fotestaceous. Six humeral setae, 2 posthumerals and 2 presuturals, 2 pairs of discal scutellars. Pleura dark brown with grayish tomentum; in- frasquamal setulae present; squamae white, wings light brown; epaulet brown, basicosta ru- fotestaceous; mediotergite thinly tomentose, shining. Legs dark rufotestaceous, anterior and middle femora brownish with pale apical mark- ings; posterior femora with dark apicoventral patches; tibiae unmarked, tarsi dark brown. An- terior femur with 7 dorsal setae, 8 posterodor- sals, and 15 posteroventrals; tibia without dark dorsal line. Middle femur with 5 anterior setae in 2 rows. Posterior femur with 15 anterodorsals, 8 anteroventrals, 4 posteroventrals, and no an- terior setae. Abdomen rufotestaceous with a dark median stripe; tomentum whitish. First syntergite with | pair of lateral marginal setae. Second tergite with 2 pairs of median discals, 2 WILDER: NEARCTIC PTILODEXIA pairs of median marginals, 4 pairs of lateral mar- ginals, and 2 pairs of lateral discal setae. Third tergite with 2 pairs of median discals, 10 dorsal and lateral marginals, and 2 pairs of lateral discal setae. Genitalia: Eighth tergite with sparse, long, fine hair. Ninth tergite with long brown hair and a distinct lateral swelling. Surstylus with lateral angle sharp, internal depression deep. Cerci with lateral lobes longer than arms; arms reaching to ventral margin of surstylus; in profile, cerci bulging, tips directly ventrally. Ejaculatory apodeme with bulb and stem dis- tinct; stem twice as long as height of bulb, which is widely opened on one side and top; opposite side decumbent. Female: Similar to male except for usual sexual differences and the following: General color dark brown. Width of frontal vitta at narrowest point 0.4 mm, at antennal base 0.6 mm wide. Height of gena 0.35 times head height. Seven pairs of medium postorbital setae; 10 pairs of frontal setae. Notum dark brown with heavy brownish tomentum; six humeral setae. Legs rufotestaceous. Anterior femur with 11 posteroventrals, 4 posterodorsals, and 5 dorsal setae. Middle femur with 3 anterior setae in 2 rows, | anteroventral, and 4 posteroventrals, 4 anteroventrals, and no anterior setae. Abdomen dark brown, a little lighter laterally. Second ter- gite with 1 pair of median discals, | pair of me- dian marginals, 3 pairs of lateral marginals, and 1 pair of lateral discal setae. Third tergite with 1 pair of median discals, 10 dorsal and lateral marginals, and 2 pairs of lateral discal setae. Tip of abdomen reddish. VARIATION.—Ptilodexia pacifica is the most variable Nearctic speices of Ptilodexia. Several fairly distinct populations are present, each of which may eventually be recognized as a sepa- rate species. I fail to name these as species now because of the inadequate sample and numerous specimens which appear to be intermediates. The form to which the type-specimens belong has males and females with pale-colored legs. These specimens have the parafacial hairs rather sparse; are small in body size (7-13 mm); and the abdominal tomentum is uniformly whitish in most individuals. This form occurs from south- ern California along the coast into Monterey County. Members of the second form are larger; have parafacial hairs abundant; and only females have pale-colored legs, although the legs of some males are marked with orange or reddish. The 33 facial tomentum is brownish in some specimens, and the abdominal tomentum is so in most. These specimens are more hairy and much dark- er in color than those of the previous form. Many have one to two rows of dark hairs ventral to the postorbitals, and have dark squamae and wings. This form ranges from Oregon south along the California coast to Monterey County, where both this and the previous form are found. The third form is found in the Sierra Nevada from northern California south into Inyo Coun- ty. These specimens resemble those of the first form in size and facial hair; however, most in- dividuals are darker in color, and males and fe- males have dark-colored legs. The abdominal tomentum is concolorous, whitish. Specimens of the three forms have a number of character states in common which segregate them from specimens of other western species. It is for this reason and the lack of material that I have treated the complex as one species. It is hoped that further collecting, and ecological and life history studies will elucidate relationships and uncover a basis for the variability within this species. MATERIAL EXAMINED.—Two hundred twenty-six males and 115 females of P. pacifica were examined. Data from those specimens in the type-series are as follows: California: Monterey Co.: 353 312%, Point Pinos, Pacific Grove, 24 May, 1952, P. Arnaud, 46461229, 13 June 1959, 256 ¢529, 19 Aug. 1957 (PHAC). Data from the other specimens examined are recorded by Wilder (1976). DISTRIBUTION.—The species ranges from southern California north along the coast to northern Oregon. It is also found in the Central Valley and through the Sierra Nevada in Cali- fornia. BroLocicaAL Notes.—The flight period ranges from May to November with specimens occur- ring throughout that period with perhaps two activity peaks—one in late spring and early sum- mer, and the other in the fall. Specimens of P. pacifica have been collected on the following flowers belonging to the family Compositae: Baccharis pilularis, Chrysotham- nus viscidiflorus, and Achillea Millefolium, and undetermined Baccharis sp. and Solidago sp. Specimens have also been collected from Eri- ogonum elatum (Polygonaceae). They have also been collected while resting on the leaves of Rubus parviflorus and Umbellularia californica. This species inhabits both low and high (up to 34 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 1 NORTH AMERICA Figures 53-57. Ptilodexia sabroskyi, n.sp. Fig. 53. Genitalia of male, posterior view. Fig. 54. Ejaculatory apodeme. Fig. 55. Genitalia of male, lateral view. Fig. 56. Head of male, lateral view. Fig. 57. Geographical distribution. 3,660 m) elevations. Collecting methods used in- clude UV and white light, sweeping, flight trap, and light trap. Ptilodexia pacifica adults have been collected in meadows, near streams, on sand dunes, and on rocks. Ptilodexia sabroskyi, new species (Figures 53-57) Type-LocaLity.—San Clemente, Orange County, Califor- nia. TyPE-SPECIMENS.—The holotype, a male, was collected by G. Eickwort on 25 July 1961, and is deposited in MSUC. The allotype, deposited in LACM, is from near Wrightwood, 1,719 m, San Bernardino County, California. Complete data from these specimens and the 37 male and 7 female paratypes are listed below. DIAGNOsIS.—Specimens of P. sabroskyi are large, robust, and quite distinctive. They can be distinguished from those of other species of Ptilodexia by the following combination of char- acter states: parafacial hairs brown, rather short, concentrated near upper anterior edge of parafacial and scattered randomly elsewhere, lower ones often yellowish; length of plumosity on arista less than twice length of second anten- nal segment, the third antennal segment short, subequal to the second in length; smallest dis- tance between eyes greater than width of frontal vitta at antennal base; frontal vitta not obliter- ated at its narrowest point; length of haustellum varied from 0.5 to 0.6 times head height; legs of both sexes dark; notum light brown laterally in most individuals. DESCRIPTION.—Male: Total body length 17 mm, dark brown with abdomen reddish lat- erally. Head height 4 mm; profrons 0.3 times head length. Facial tomentum dull whitish, heavy, obscuring integumental color; parafron- tal hairs medium in length, dark, abundant; parafacial hairs brown, short, concentrated on upper anterior portion of face, short, pale hairs scattered elsewhere on parafacial. Width of frontal vitta at antennal base 0.6 mm, at narrow- est point 0.1 mm; carina well developed, broad; facial cavities dark gray. Height of gena 0.3 times head height. One pair of strong ocellar se- tae, two pairs of smaller postocellars, one pair WILDER: NEARCTIC PTILODEXIA each of postverticals, inner verticals, and outer verticals. Eighteen pairs of long postorbital se- tae, not interspersed with smaller hairs; ventral to postorbitals are no dark hairs, only yellowish occipital hair. One pair of frontal setae, | pair of long oral vibrissae 0.4 mm from oral margin, the depression between them 0.6 mm wide; 16 additional oral setae, one above vibrissae rather strong, peristomals subequal to each other, about 0.5 times vibrissae length. Epistome strongly protruding, its width 1.7 times length of first two antennal segments. Haustellum broad, blackish, length 0.5 times head height; palpi long, narrow, rufotestaceous, length 0.6 times haustellum length. Antennae dark rufotesta- ceous; third segment, arista, and plumosity dark brown. Third segment of antenna very short, 0.8 times length of second segment; length of plu- mosity on arista 1.5 times length of second an- tennal segment. Thorax: Mesonotum dark brown, rufescent laterally; tomentum heavy, grayish, striping distinct; integument of humeri, postalar calli, and scutellum rufescent. Six hu- meral setae, 2 posthumerals and 2 presuturals, 6 pairs of discal scutellars. Pleura dark brown, sutures lighter, tomentum heavy, grayish; infra- Ssquamal setulae present; squamae whitish, wings light brown, epaulet dark brown, basi- costa testaceous; mediotergite subshining to- mentose. Legs dark brown, tibiae rufous, dark- ened basally and slightly apically; tarsi blackish. Anterior femur with 12 dorsal setae, 11 postero- dorsals, and 18 posteroventrals; tibia with dark dorsal line. Middle femur with 4 strong anterior setae in 2 rows. Posterior femur with 13 antero- dorsals, 10 anteroventrals, 6 posteroventrals, and no anterior setae. Abdomen rufous with a narrow, dark brown, median stripe; tomentum white, heavy, patchy, brown around median marginal setae of second and third tergites. First syntergite with | pair of lateral marginal setae. Second tergite with 3 pairs of median discals, 2 pairs of median marginals, 3 pairs of lateral mar- ginals and 2 pairs of lateral discals. Third tergite with 3 pairs of median discals, 12 dorsal and lateral marginals, and 4 pairs of lateral discals. Genitalia: Eighth tergite with numerous small, fine hairs. Ninth tergite with long dark hairs; lateral swelling pronounced. Surstylus large with lateral angle sharp, internal depression rather shallow, basally strongly convex; in pro- file, inclined slightly posterad. Cerci with height of lateral lobes subequal to height of arms; in profile, lateral lobe with a strong dorsal hook pointing anteriad, arms bulging slightly ventral- ly, tips rounded. Ejaculatory apodeme with bulb distinct from long, thin stem, which ts 2.5 times height of bulb. Bulb large and somewhat flat- tened, open widely on one side and narrowly on top with a strong lip; opposite side strongly de- cumbent. Female: Similar to male except for usual sexual differences and the following. Width of frontal vitta at narrowest point 0.4 mm, at antennal base 0.6 mm. Height of gena 0.4 times head height. Nine pairs of short postor- bital setae irregularly interspersed with short hairs; 8 pairs of frontal setae. Legs same color as in male, anterior femur with 10 posteroven- trals, 8 posterodorsals, and 8 dorsal setae. Mid- dle femur with 3 anterior setae in 2 rows, 5 pos- teroventrals, and | anteroventral. Posterior femur with no anterior setae, 7 anterodorsals, 4 posteroventrals, and 4 anteroventrals. Abdomen dark reddish brown with a vague median stripe; tomentum heavy, white. First syntergite with | pair of lateral marginal setae. Second tergite with | pair of median discals, | pair of median marginals, and | pair of lateral marginal setae. Third tergite with | pair of median discals and 12 dorsal and lateral marginals. VARIATION.—P. sabroskyi exhibits relatively little variation. Total body length ranges from 12 to 18 mm. The scattered hairs on the parafacial are brown in some individuals, but are long in none. The third antennal segment is varied in length, at its longest being equal to the second segment. The number of discal scutellar setae is quite varied, as are the lengths of those setae. Abdominal color in males and females ranges from rufous to dark reddish brown, although few individuals exhibit the latter. There is no brown- ish tomentum on the abdomens of a few males and many females. MATERIAL EXAMINED.—Material examined included 38 males and 8 females. Data from these specimens are as fol- lows: California: Unknown: 16, Marina Beach, 17 June 1961, C. Philip (CASC); 1d, Pine Lake, southern California, John- son (USNM). Inyo Co.: 12, Wyman Canyon, White Mts., near stream, 21 July 1967, S. Frommer (UCRC). Los Angeles Co.: 16, Malibu Creek mouth, 17 July 1953 (LACM); 16, Claremont, Baker (LACM). Orange Co.: 16, Newport, 10 July 1916, D. Hall (USNM); 3¢¢, Newport Beach, 19 Aug. 1920, Timberlake (UCRC); 1¢, San Clemente, 25 July 1961, G. Eickwort (MSUC). Riverside Co.: 16, Coachella Valley. 7 Apr. 1928, R. Woglum (CISC); 1d, Palm Springs, Andreas 36 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 1 NORTH AMERICA Ficures 58-62. Ptilodexia carolinensis Brauer and Bergenstamm. Fig. 58. Genitalia of male, posterior view. Fig. 59. Ejaculatory apodeme. Fig. 60. Genitalia of male, lateral view. Fig. 61. Head of male, lateral view. Fig. 62. Geographical distribution. Canyon, 24 Apr. 1954, M. Wasbauer (CISC). San Bernardino Co.: 16, May, Coquillett (USNM); 1d, Barton Flats, 20 July 1950, A. Melander (USNM); 5d, Cajon, 11 May 1934, C. M. (UCRC); 3¢6, Colton, 26-28 May 1917, E. VanDuzee (CASC); 16, Deep Creek Public Camp, 15 June 1957, A. Menke (LACM); 16, South Fork Camp, San Bernardino Mts., 2 Sep. 1946, Timberlake (UCRC); 1¢, upper Santa Ana River, 6 July 1948, A. Melander, 1d, 9 July 1959, 1d, 16 July 1947, 1d, 18 July 1950, 1¢19, 22 July 1950, 1d, 4 Aug. 1946, 135, 10 Aug. 1949, 15, 29 Aug. 1946, 16, 15 Sep. 1946 (USNM); 1¢, Verdemont, 22 May 1954, A. Melander (USNM); 12, near Wrightwood, 1,719 m (LACM). San Diego Co.: 16, A. Moldenke (USNM); 16, 1 mile S of Del Mar, Eriogonum fasciculatum, 10 July 1963, P. Hurd (CISC); 1612, Dulzura, 14 June 1917 (AMNH); 12, 2.8 miles [ca. 4.5 km] SW of Poway, Los Penasquitos Creek, 400 ft [ca. 120 m], 25 July 1965, R. Somerby (UCRC); 12, Tecate Peak, 10 July 1963, J. Powell (CISC). Ventura Co.: 12, Oxnard Beaches, 13 June 1955, R. Erdmann, 1d, 16 July 1953, W. Gertsch, 1°, 25 July 1956, R. Erdmann (LACM, AMNH). DISTRIBUTION.—The range of P. sabroskyi is restricted to southern California and the White Mountains. BIOLOGICAL Notes.—The period of activity lasts from April to September. Adults have been collected near a stream and on Eriogonum fas- ciculatum (Polygonaceae). DERIVATION OF NAME.—Ptilodexia sabroskyi is named in honor of Dr. C. W. Sabrosky for his enormous contribution to the study of the Tach- inidae. carolinensis Group Ptilodexia carolinensis Brauer and Bergen- stamm (Figures 58-62) Ptilodexia carolinensis BRAUER AND BERGENSTAMM, 1899:119. BRAUER (1899:508); ALDRICH (1905:499); SABROSKY AND ARNAUD (1965:988); CoLe (1969:543). [LECTOTYPE (here designated), male, deposited in VMNH, labeled, **S. Car- olina’’/‘‘rufipennis Mq, carolinensis, Coll. Winthem, (two indistinguishable words)’’/*‘Carolinensis Type, det. Brauer. Bergenst.’’/*‘carolinensis Type Br Bgst’’/**Ptilodexia caro- linensis Br. Bgst.’’/*‘Lectotype Ptilodexia carolinensis des- ignated by D. Wilder 1976.’’] Estheria tibialis (partim): CoQUILLETT (1910:598); Town- SEND (1931:102). (misidentification) Ptilodexia abdominalis (partim): Davis (1919:84). (misidenti- fication) Ptilodexia neotibialis WEST, 1924:184. LEONARD (1928:822); WILDER: NEARCTIC PTILODEXIA CuRRAN (1930:93); SABROSKY AND ARNAUD (1965:989); CoLe (1969:543). [HoLoTYPE, male, deposited in MCZC, labeled, **Colebrook, Conn., Coll. W. M. Wheeler’’/Black square/**MCZ Type 26963’’/*‘Holotype Prilodexia neotibi- alis West.’’] NEw SYNONYMY Dexia harpasa (partim): ALDRICH (1925:114); JOHNSON (1925b:208). (misidentification) Rhynchodexia confusa (partim): JOHNSON (1925b:208). (mis- identification) Ptilodexia minor WEST, 1925:132. LEONARD (1928:822); WEST (1950:pl. I, fig. 3, pl. IV, fig. 8); SABROSKY AND ARNAUD (1965:989). [HoLoTyYPE, female, deposited in CUIC, labeled ‘‘Duck Lake, N.Y., 6-VIII-21"’/"*L. S. West Collector’’/ ‘Wing Slide, Cornell U., Lot 919, Sub 86, L. S. West’’/ ‘Holotype Prilodexia minor West’’/*“Holotype Cornell U., No. 1875’’/*‘Cornell U., Lot. 922, Sub. 43.’’] New Syn- ONYMY TAXxoNomic Notes.—Ptilodexia carolinensis has been confused routinely with other species by earlier workers. Neither Curran nor West had ever seen the type. West speaks of generic char- acters but never mentions P. carolinensis, the type-species of the genus, in any of his papers. Aldrich (1925) saw the types, matched them with his specimens of P. canescens and P. plan- ifrons, which he thought were Estheria tibialis. Following Austen (1907), he assigned this species to Dexia harpasa Walker. Townsend (1931) and Coquillett (1910) also felt that P. car- olinensis was a synonym of E. tibialis. Sabrosky and Arnaud (1965) considered P. carolinensis separate from P. harpasa. I have seen the type—there is only one male now, not the two which Aldrich (1925) report- ed—and it is in excellent condition. The speci- men has been designated lectotype. DiaGNosis.—Ptilodexia carolinensis is a di- verse species (or perhaps species group), its members lacking distinct characters. Specimens can be more easily assigned here by character- istics which they lack rather than those which they possess. Adults with rather broad, parallel-sided face, vibrissal angle protruding little or not at all; parafacial hairs long, fine, inserted along middle of parafacial; epistome protruding only slightly; third antennal segment 1.5 times length of sec- ond in most specimens; length of plumosity on arista varied from | to 1.5 times length of second antennal segment; haustellum length varied from 0.4 to 0.5 times head height; scutellum and sides of abdomen orange-brown in most adults; some specimens from Texas and South Dakota are to- tally orange-brown. 37 MATERIAL EXAMINED.—Two hundred fifty- six males and 133 females were examined. DISTRIBUTION.—This species ranges widely from Nova Scotia south to Georgia and west in- to Texas and the Rocky Mountains, with scat- tered records from Utah, Wyoming, and Sas- katchewan. BIOLOGICAL Notes.—The flight period is from July to September with the majority of specimens collected from mid-August to mid- September. Specimens have been collected as early as 30 May on Long Island and as late as 23 October in Virginia. In Texas, the insect seems to be double- brooded, some specimens being collected in April, May, and June, and others in September, October, and November. In material from other areas, such as New York, we find early-season and late-season specimens, but their numbers do not indicate a double brood. There may be a correlation between the brood and body color in the enigmatic Texas specimens. Unfortu- nately, the sample at hand, while giving weight to this possibility, is too small for any such con- clusions to be drawn. Specimens which have been brought into the lab complete their development and emerge much earlier than they would have in nature. Parasitized Japanese beetles, collected in Oc- tober, yielded adult Prilodexia carolinensis as early as 21 December. It is not known if these specimens were exposed to cold temperatures. It is feasible that in a warm climate two broods might occur. In the area where the above-men- tioned specimens were collected (Connecticut), the adult would not have emerged, under normal conditions, until the following summer. Nearly all of the specimens of this species were collected at low elevations. One was col- lected at 1,450 m in the southern Appalachians, and others in Colorado and Wyoming at 1,950 m and 2,060 m, respectively. A number of spec- imens have been collected by sweeping vegeta- tion, and some have been taken on Solidago sp. Perhaps coincidentally, Japanese beetle feeds on the foliage of Solidago juncea (Fleming 1972). Available host data indicate that this insect par- asitizes Popillia japonica in the Northeast and Phyllophaga sp. in Texas. Davis (1919:84) re- ports that adults (under the name of P. abdom- inalis were reared from the larvae of Phyllopha- ga rugosa collected in Manitoba. 38 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 1 NORTH AMERICA FiGures 63-67. Ptilodexia canescens (Walker). Fig. 63. Genitalia of male, posterior view. Fig. 64. Ejaculatory apo- deme. Fig. 65. Genitalia of male, lateral view. Fig. 66. Head of male, lateral view. Fig. 67. Geographical distribution. Ptilodexia canescens (Walker) (Figures 63-67) Dexia canescens WALKER, 1852:310. OSTEN SACKEN (1878:155); ALDRICH (1905:502); AUSTEN (1907:345); CHAMPLAIN AND KNULL (1944:214); PETERSON (1948:60); SABROSKY AND ARNAUD (1965:988). [HOLOTYPE, male, de- posited in BMNH, labeled *‘Dexia canescens, Walk.’’/ “Holotype’’/"‘U.S.’’/""United States. Ex. Coll. Saunders 68.4’’/Crosskey examination label, 1970] Ptilodexia harpasa (partim): Davis (1919:82); ALDRICH (1925:114); JOHNSON (1925b:208). (misidentification) Rhynchodexia confusa (partim): JOHNSON (1925b:208). (mis- identification) TAXONOMIC Notes.—The holotype has many dorsal setae broken and one antenna missing. The wings are in excellent condition. DIAGNosIs.—Adults of P. canescens are unique in the following combination of charac- ters: parafacial hairs long, coarse, denser near lower anterior edge of eye in most individuals. Oral margin in most adults not protruding; males with frontal stripe not obliterated at narrowest point; width of plumosity on arista less than or equal to length of second antennal segment: length of haustellum less than half head height; total absence of infrasquamal setulae; abdomen quite dark in most specimens, with a distinct median stripe in a few. MATERIAL EXAMINED.—One hundred forty- eight males and 68 females were examined. DISTRIBUTION.—This species ranges from Nova Scotia to Long Island and west to the Rocky Mountains. There are scattered records in British Columbia, Idaho, Wyoming, and Utah. BIOLOGICAL Notes.—tThe flight period is from June to August, with the majority of rec- ords in mid-July. Specimens have been collected as early as 24 April (Long Island, 1923) and as late as 8 September (Massachusetts, 1907) in nature. The majority of specimens were collected at low elevations; however, there are records from mountainous areas, the highest being 2,040 m in Wyoming. Specimens from Massachusetts and Utah have been collected on the flowers of par- snip. Specimens from State College, Pennsyl- vania, were taken at a UV light. In Kindred, in eastern North Dakota, a male and a female were WILDER: NEARCTIC PTILODEXIA 39 NORTH AMERICA Ficures 68-72. Ptilodexia halone (Walker). Fig. 68. Genitalia of male, posterior view. Fig. 69. Ejaculatory apodeme. Fig. 70. Genitalia of male, lateral view. Fig. 71. Head of male, lateral view. Fig. 72. Geographical distribution. collected on native prairie vegetation. Two males and six females have been reared and are associated with puparia. The only host data available indicate that this insect is a parasitoid of Phyllophaga sp. Ptilodexia halone (Walker) (Figures 68-72) Dexia halone WALKER, 1849:837. OSTEN SACKEN (1878:155); ALDRICH (1905:502); SABROSKY AND ARNAUD (1965:988). [LecTotyPE (here designated), male, desposited in BMNH, labeled, “Dexia halone Wlk.’’/**Lectotype’’/**Georgia, U.S.A., Ex. Coll. Abbot’’/**Georgia’’/Crosskey examina- tion label 1970/**Lectotype Dexia halone Walker designated by D. Wilder 1975.’’] Ptilodexia hucketti West, 1952:131. LEONARD (1928:822); CURRAN (1930:93); West (1950:pl. I, fig. 1); SABROSKY AND ARNAUD (1965:988) [HoLoTYPE, male, deposited in CUIC, labeled, “Riverhead, L.I., N.Y., 30 July 1922, H. C. Huck- ett’/"“Holotype Ptilodexia hucketti West, 3°’/**Holotype Cornell U. No. 1872°’/**Cornell U., Lot. 922, Sub. 44°’] New SYNONYMY TAXONOMIC Notes.—The lectotype and paralectotype are in poor condition but still eas- ily recognizable. The lectotype has all dorsal se- tae broken or absent, most tarsi and the left pos- terior leg missing, and the abdomen broken and glued. DIAGNosis.—Adults of Prilodexia halone can be distinguished from those of all other species of Ptilodexia by the following combination of characters: totally orange or orange-brown in color; parafacial hairs short and pale, inserted rather randomly on parafacial; oral margin not or only slightly protruding; area between oral vibrissae and oral margin higher than wide (length and width may be equal); length of haus- tellum 0.3 to 0.4 times head height; third anten- nal segment only slightly longer than second; length of plumosity on arista 1.4 times length of second segment; inner vertical setae strongly convergent; tarsi concolorous with femora and tibiae; presence of median marginal setae on first abdominal segment. MATERIAL EXAMINED.—Thirteen males and seven females were examined. DISTRIBUTION.—Scant available records show that P. halone ranges from southern Mississippi presumably through the Appalachians into Ten- nessee and along the East Coast from Maryland 40 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 1 NORTH AMERICA Ficures 73-77. Ptilodexia prexaspes (Walker). Fig. 73. Genitalia of male, posterior view. Fig. 74. Ejaculatory apodeme. Fig. 75. Genitalia of male, lateral view. Fig. 76. Head of male, lateral view. Fig. 77. Geographical distribution. to Albany, New York. There is one record from Georgia. Adults appear to be uncommon. BioLoGicAL Notes.—tThe flight period ranges from June to July in the southern part of its range and from mid-July to late August in the north. There are no ecological data associated with the material examined. Ptilodexia prexaspes (Walker) (Figures 73-77) Dexia prexaspes WALKER, 1849:837. OSTEN SACKEN (1878:155); BRAUER (1899:502); AUSTEN (1907:345); BRIMLEY (1938:368); SABROSKY AND ARNAUD (1965:989). [LECTOTYPE (here des- ignated), male, deposited in BMNH, labeled, **‘Dexia prex- aspes Walk.’’/*‘Lectotype’’/*‘Georgia, U.S.A., Ex. coll. Abbot’’/**Georgia’’/Crosskey examination label, 1970/ “Lectotype Dexia prexaspes Walker, designated by D. Wilder 1975.”’] TAXONOMIC Notes.—The lectotype has the right front and rear tarsi missing, most frontals, dorsal abdominal, and thoracic setae missing, and the haustellum withdrawn. DiaGnosis.—Adults of Ptilodexia prexaspes are unique in the following combination of char- acters: Rather light-colored flies, females uni- formly dark rufotestaceous except for darker tarsi; males may be partially brown dorsally, but with at least humeri and legs pale rufotesta- ceous; parafacial hairs dark, short in females, long and coarse in males, scattered on parafa- cial; third antennal segment 1.5 times length of second; length of plumosity on arista subequal to second antennal article; frontal vitta of males obliterated at some point between eyes; length of haustellum 0.3 to 0.4 times head height; oral margin protrudes slightly, if at all; area between oral vibrissae and oral margin distinctly higher than wide: legs of males and females light in color. MATERIAL EXAMINED.—Only six males and six females of P. prexaspes have been examined for this study. DISTRIBUTION.—This species appears to be localized and its members uncommon. The range extends from central Florida along the southern Atlantic coast to Cape Hatteras, North Carolina. BIOLOGICAL NoTES.—This insect appears to have two broods in Florida. The flight periods WILDER: NEARCTIC PTILODEXIA 41 NORTH AMERICA Ficures 78-82. Prtilodexia harpasa (Walker). Fig. 78. Genitalia of male, posterior view. Fig. 79. Ejaculatory apodeme. Fig. 80. Genitalia of male, lateral view. Fig. 81. Head of male, lateral view. Fig. 82. Geographical distribution. are from late March and April to June in Florida, and late August and September to October in Florida and North Carolina. Adults of P. prexaspes have been collected at a 15-watt UV light in March and June. In the fall they have been taken on Solidago micro- cephala and Eupatorium, both Compositae. One specimen was collected from a female asi- lid, Efferia aestuans (Linnaeus), which had pre- sumably taken it as prey. harpasa Group Ptilodexia harpasa (Walker) (Figures 78-82) Dexia harpasa WALKER, 1849:840. OSTEN SACKEN (1878:155); ALDRICH (1905:502); AUSTEN (1907:344); Davis (1919:82): WEST (1924: 186); ALDRICH (1925:114); JOHNSON (1925a:89): (1925b:208); (1927:203); LEONARD (1928:822); CURRAN (1930:93); TOWNSEND (1931:102); BRIMLEY (1938:368); PROCTER (1938:371); (1946:429); SABROSKY AND ARNAUD (1965:988). [HoLoTyPE, a female, deposited in BMNH, la- beled, “Dexia harpasa, Walk.’’/*‘Holotype’’/**40 3 30 1026°’/"*N. America. ex. Coll. Children 40.3.30.1026.°’/ Crosskey examination label 1970.] Ptilodexia leucoptera WEST, 1925:132. LEONARD (1928:822); CURRAN (1930:93); West (1950:pl. I, fig. 2); SABROSKY AND ARNAUD (1965:989). [HoLotyPE, female, deposited in CUIC, labeled, ‘Riverhead, L.I., N.Y., IX-17-13"’/* Wing Slide, Cornell U., Lot. 919, Sub. 115, L. S. West’’/**Ho- lotype Ptilodexia leucoptera West 2°’/**Holotype Cornell U., No. 1874°’/**Cornell U., Lot. 922, Sub. 42.”’] NEw Syn- ONYMY Dinera robusta CURRAN, 1930:93. West (1950:111); SABROs- KY AND ARNAUD (1965:989). [LECTOTYPE (here designat- ed), male, deposited in CUIC, labeled, **Poughkeepsie, N.Y., July 1906, N.Y.S. Coll.’’/** 3 Allotype Rhynchiodexia robusta Curran’’/**Allotype Cornell U., No. 1935°’/**Cor- nell U., Lot. 922, Sub. 3°’/*‘Prilodexia obscura West, det. H. J. Reinhard’’/**Lectotype Dinera robusta Curran des- ignated by D. Wilder 1976.""] NEw SYNONYMY Ptilodexia tibialis (partim): TOWNSEND (1931:102). TAXONOMIC Notes.—Ptilodexia harpasa was first described and named from one female by Walker in 1849. The holotype is in poor condi- tion, lacking posterior legs, right middle leg, and antennae (one antenna is glued to the double mount). All the setae from the dorsal surface of the thorax and abdomen are missing, and the coxal, sternopleural, and hypopleural areas of the left side are obliterated by the pin. The para- facial hairs have been rubbed off, but were ob- viously present at one time. Despite its condi- 42 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 1 tion, there is no question about the identity of the specimen. Austen (1907) placed the name P. harpasa in synonymy with P. tibialis, apparently without seeing the type of the latter (which had been lost). Aldrich (in Davis 1919) disputed this and restored validity to P. harpasa. In 1925, upon visiting the Vienna Natural History Museum, he placed P. carolinensis in synonymy with P. har- pasa, using for comparison specimens from USNM which were actually P. canescens and P. planifrons. At no time does Aldrich mention seeing the type of P. harpasa; he apparently based his conclusions solely on the word of Aus- ten. In 1931, Townsend reiterated this synony- my and restored the name P. tibialis as the valid one, again without looking at the latter type. Since that time, the names P. tibialis and P. harpasa have both been used for many species of Ptilodexia. Ptilodexia leucoptera was also described from a single female by West in 1925. The holotype, from Riverhead, Long Island, is in good condi- tion and is deposited in CUIC. Curran introduced the name Dinera robusta into the literature in his 1930 key. West (1950) claimed that he had never intended to describe robusta in the genus Dinera. This manuscript name was only tentative—until Curran validated it. The specific epithet, robusta, is totally mis- leading since this species is one of the smallest in the genus. West redescribed and illustrated Rhynchiodexia robusta in 1950, incorrectly des- ignating his earlier types as holotype and allo- type. Since Curran was the describer, the spec- imens he looked at (including West’s holotype and allotype) became a syntype series, and the only valid designation could be lectotype and paralectotype. I have designated West’s female “‘holotype”’ as paralectotype and his male ‘‘al- lotype’’ as lectotype. Other determined material in AMNH and CUIC which Curran probably used in developing his concept of the species have also been designated paralectotypes. DIAGNOsIS.—Ptilodexia harpasa adults are among the smallest, most slender flies in the ge- nus. They have parafacial hairs medium to long, fine, distributed sparsely along center of para- facial, in a few individuals inserted below ven- tral eye margin; haustellum length 0.65 to 0.7 times head height, slender; third antennal seg- ment long; length of plumosity on arista more than twice length of second antennal segment: abdomen and scutellum appear dark, but are ac- tually partially rufescent. MATERIAL EXAMINED.—Four hundred seven males and 465 females were examined, a sex ratio different from the usual in the genus. DISTRIBUTION.—Ptilodexia harpasa ranges from the southern Appalachians into northern Quebec and west into Ohio, Michigan, northern Illinois, Minnesota, North Dakota, and Mani- toba. BIoLoGicAL Notes.—tThe flight period lasts from May to October, the most active months being August and September. Ptilodexia har- pasa adults have been collected in Malaise traps, by net, on foliage, and as the prey of a crab spider. Flowers from which they have been collected are: Solidago canadensis, Solidago rugosa, Eupatorium coelestinum (Compositae); and Melilotus alba (Fabaceae). Several specimens have been reared and are pinned with puparia. Two specimens from On- tario bear the label, “reared from R. C. larvae.” This is perhaps an abbreviation for rose chafer, Macrodactylus subspinosus, a small scarab, whose range coincides with that of P. harpasa. Davis (1919) reports that adults have been reared from grubs (Phyllophaga?) from Con- necticut. Ptilodexia rufipennis (Macquart) (Figures 83-88) Dexia rufipennis MACQUART, 1842:244. OsTEN SACKEN (1878:155); BRAUER (1899:508); JOHNSON (1925b:209); AL- DRICH (1905:504); CURRAN (1930:93); BRIMLEY (1938:367); SABROSKY AND ARNAUD (1965:989). [HoLotyPeE, female, lost.] Dexia cerata WALKER, 1849:847. OSTEN SACKEN (1878:155); ALDRICH (1905:502); AUSTEN (1907:344); FATTIG (1944:3); SABROSKY AND ARNAUD (1965:989). [HoLoTyPE, female, deposited in BMNH, labeled, “Dexia cerata Walk.’’/**Ho- lotype’’/*"N. Amer.’’/**N. America. Pres. by the Entomo- logical Club, 44.12’’/Crosskey examination label, 1970.] New SYNONYMY Dexia albifrons WALKER, 1852:317. OSTEN SACKEN (1878: 155); ALDRICH (1905:501); AUSTEN (1907:345); SABROSKY AND ARNAUD (1965:988). [HoLoTYPE, male, deposited in BMNH, labeled, “Dexia albifrons Walk.’’/*‘Holotype’’/ “U.S."’/“United States Ex. Coll. Saunders. 68.4°’/*‘albi- frons’’/Crosskey examination label, 1970.] NEw SyYNony- MY Rhynchodexia confusa WesT, 1924:185. JOHNSON (1925b:208); LEONARD (1928:821); CURRAN (1930:93); WRAY (1938:91); REINHARD (1943:22); West (1950:109); SABROSKY AND AR- NAUD (1965:988). [HoLoTYPE, male, deposited in CUIC, labeled, ‘‘Selden, L.I., July 1, 1923, N.Y.’’/**Holotype WILDER: NEARCTIC PTILODEXIA 83 43 NORTH AMERICA t 500 Ficures 83-88. Prilodexia rufipennis (Macquart). Fig. 83. Genitalia of male, posterior view. Fig. 84. Ejaculatory apo- deme. Fig. 85. Ejaculatory apodeme. Fig. 86. Genitalia of male, lateral view. Fig. 87. Head of male, lateral view. Fig. 88. Geographical distribution. Rhynchodexia confusa West, 3°’/**Holotype, Cornell U. No. 1245’’/*Cornell U., Lot. 922, Sub. 35.’’] NEw Syn- ONYMY Rhynchodexia translucipennis West, 1925:135. LEONARD (1928:821); CURRAN (1930:93); West (1950:pl. I, fig. 6); Sa- BROSKY AND ARNAUD (1965:989). [HoLoTYPE, male, de- posited in CUIC, labeled, “Ithaca, N.Y., 18 July ‘21°7/""L. S. West Collector’’/**Wing Slide, Cornell U., Lot 919, Sub. 116, L. S. West’’/*Holotype, Rhynchodexia translucipen- nis West, 3°’/*Holotype Cornell U., No. 826°’/**Cornell U., Lot 922, Sub. 37.""] NEw SYNoNYMY Ptilodexia harpasa (partim): JOHNSON (1925b:209). (misiden- tification) Ptilodexia neotibialis (partim): JOHNSON (1925:208). (misiden- tification) Rhynchiodexia dubia CURRAN, 1930:93. West (1950:110); Sa- BROSKY AND ARNAUD (1965:988). [LECTOTYPE (here des- ignated), male, deposited in AMNH, labeled, “Sta. Study Insects, Tuxedo, N.Y., 28—VII-1928’’/*‘Collector C. H. Curran’’/*‘Lectotype Rhynchiodexia dubia Curran desig- nated by D. Wilder 1976.’"’] NEw SYNONYMY TAXONOMIC Notes.—The name P. rufipennis has been misused frequently in the literature. It also has a long list of synonyms. Part of the reason for this is clear—the species exhibits ex- ceptionally high variability. Females and males differ from each other in taxonomic characters which were used previously in the group to de- fine species. There is also strong clinal variation in males, which without being thoroughly sam- pled could lead workers to name certain popu- lations as separate species. Specimens also are commonly collected. These factors, combined with the failure of earlier workers to examine type material, have contributed to the taxonom- ic problems in this species. The name has been in general use since it was proposed in 1843. Brauer (1899) felt that this species was the same as P. carolinensis, and therefore placed the species in Ptilodexia. Sub- sequent workers used the name but failed to de- fine the species. This was understandable since the type was lost and the original description was short. Curran’s 1930 key characterized members of the species as having infrasquamal setulae ab- sent, parafacials haired, and the scutellum red- dish. Macquart’s description states that the fe- male from Nova Scotia, which he named Dexia 44 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 1 rufipennis, has, among other characteristics, the legs, basal antennal segments, and scutellum ‘“*testace.’’ He also describes the epistome as ‘*saillant’’ or protruding. The only species which ranges into Nova Scotia and whose members combine those characters with the others in Macquart’s original description is the one pres- ently being considered, specimens of which lack parafacial hairs. The holotype of Dexia cerata Walker, de- scribed as being from “North America,” fits Macquart’s description perfectly. It is in poor condition, with antennae, facial setae, and left middle and hind legs broken. The haustellum is withdrawn. Dexia albifrons Walker, another synonym of this species, is also from the *‘United States.” The male holotype is a dark, small specimen of this varied species. It is in poor condition, en- tirely covered with dust or mold and missing most setae. West used the genus Rhynchodexia for those species whose members lacked parafacial hairs. In describing Rhynchodexia confusa, he looked only at specimens from the northeastern United States and suspected that more than one species was involved. He also stated that this species had been represented in collections under the name P. harpasa, another Walker species de- scribed in the same paper as P. cerata. The de- scription of P. harpasa is much less character- istic of members of this species than is that of P. cerata, and it is difficult to understand why that name was being used instead of P. cerata. The holotype and allotype of P. confusa are both large specimens with light-colored legs. They are in excellent condition. West’s other synonym of P. rufipennis is Rhynchodexia translucipennis. This species was described from two unusual-looking specimens of P. rufipennis. The holotype (and paratype) has translucent, milky-white wings and a light, reddish-brown thorax. It is possible that these specimens are teneral; however, they lack the characteristic collapsed face normally associ- ated with teneral Ptilodexia. Tenerals do not generally have milky wings. Nevertheless, these specimens definitely belong in P. rufipennis. The type-specimens are in good condition. The name Rhynchiodexia dubia was intro- duced into the literature by C. H. Curran in 1931. Between 1921 and 1925, West had been working with tachinids of New York and adja- cent states and had placed type labels on certain specimens. His work was interrupted, and these specimens, with their manuscript names, were deposited in CUIC. Curran consulted this col- lection and used five of West’s manuscript names in his paper “‘Diptera from Tuxedo,”’ which was essentially a faunal list. Unfortu- nately, a key to adults of the species of Rhyn- chiodexia appears in this paper, making the names valid since they were accompanied by a statement of diagnostic characters. Of the five species described in this paper, Curran mentions actual material for only one— P. dubia. He does not mention West’s holotype (a female from Cimarron, Colorado), although he must have seen it to use the name. West’s holotype belongs to P. agilis Reinhard; the specimens which Curran mentions belong to P. rufipennis and are from New York. I have des- ignated the lectotype of P. dubia from the Cur- ran material for the following reasons. Curran, in mentioning these specimens, made them syn- types; he does not indicate that he saw West’s type, and there is a slight possibility that he did not. Also, West’s type is in poor condition, whereas Curran’s material is in good condition, and a lectotype designated from it would be sup- ported by a long series of paralectotypes. I have given West’s Colorado “‘holotype”’ the status of misassociated paralectotype. The lectotype and paralectotypes are in ex- cellent condition and are deposited in AMNH, FSCA, and CUIC. These specimens definitely belong to P. rufipennis (except West’s “‘holo- type’’). In Curran’s 1930 key, he separated P. rufi- pennis, as I have now defined it, into four species. The characters he used are: presence or absence of infrasquamals, color of male legs, color of scutellum, and presence or absence of parafacial hairs. All except the last are character states which vary among P. rufipennis adults. DIAGNosis.—Specimens of P. rufipennis are distinguished from those of all other species of Ptilodexia by the following combination of char- acters: face long and narrow, totally devoid of parafacial hairs ventral to second antennal seg- ment; antennae long, width of plumosity on aris- ta more than twice length of second antennal segment; abdomen grayish black or dark brown, reddish laterally in a few individuals; extreme tip of abdomen and genitalia yellowish; legs of females and often those of males pale in color; WILDER: NEARCTIC PTILODEXIA ejaculatory apodeme with bulb in shape of a shallow cup. VARIATION.—This species shows remarkable intraspecific variation. Northern specimens (those from Canada, Michigan, Wisconsin, North Dakota, and Montana) are dark in facial ground color; the antennae are dark in most adults; the legs of males are varied, ranging from light brown to almost black; and the length of the haustellum is almost half the head height. Specimens from Florida and Georgia into Texas are light in facial ground color; the antennae are rufotestaceous and the legs of males are pale, just as in females. These specimens have the haustellum short, about 0.3 times the head height. In intermediate geographical areas, there is a mixture of the two forms. It is not uncommon to have a series split into half dark individuals and half light. There is no striking difference in the genitalia of members of these two forms, and in the areas of intergradation, intermediates oc- cur. These intermediates are dark with short mouthparts or light with longer ones, and they are few in number. Many of the intermediate areas are represented by one or two specimens, and it is possible that a representative sample of these flies will show the presence of a smooth cline, northern areas having the largest percent- age of dark specimens, southern areas the larg- est proportion of light ones, and the intermediate areas showing a north-south gradient of light and dark. MATERIAL EXAMINED.—The sample of Prilo- dexia rufipennis includes 694 males and 612 fe- males. DISTRIBUTION.—This species ranges from Newfoundland to southern Florida, across the continent to British Columbia, southeastern Washington, and central Montana. There are scattered records in central Colorado, New Mexico, and Texas. BIOLOGICAL Notes.—The flight period lasts from May to November. However, most speci- mens have been collected in the months of July, August, and September. In the Midwest and Canada, records for the months of July and Au- gust predominate, while in the Northeast and Mid-Atlantic states most are collected during August and September. In areas of heavy col- lecting (Suffolk County, New York), specimens seem to occur with similar frequency throughout the season. The small sample from Florida 45 shows specimens collected in May and again in September, October, and November, perhaps indicating a bivoltine situation. In nearby Geor- gia the records show specimens taken from mid- May to mid-August with most occurring during June and July. Ptilodexia rufipennis is predominately a low- land species. A few specimens have been col- lected in mountainous areas. In New Mexico, specimens were collected at 2,440 m in the White Mountains; specimens from Georgia were taken up to 1,460 m in the southern Appalachi- ans; up to 1,890 m in North Carolina; and up to 1,830 m in Tennessee. Ptilodexia rufipennis adults have been col- lected on a number of different flowers, includ- ing the following Compositae: Solidago triner- vata, Achillea Millefolium, Eupatorium perfoliatum, Chrysanthemum leucanthemum, and unidentified Solidago sp., Aster sp., Cir- sium sp., and Anaphalis sp. They have also been noted visiting Melilotus alba (Fabaceae), Daucus carota (Umbelliferae), Ceanothus americanus, and unidentified Ceanothus sp. (Rhamnaceae), and Lonicera sp. (Caprifoli- aceae). Most of these flowers are yellow or white. One specimen was collected from a female asilid (Proctocanthus philadelphicus Macquart), which had presumably captured it as prey. Other means of collecting have been by net (sweeping foliage), UV light trap, aerial light, and Malaise trap. Ptilodexia rufipennis adults have been col- lected in a cranberry bog, marshy meadow, creek bank, maple-elm floodplain along stream, and in a maple—mixed-oak-hickory forest. Ptilodexia arida (Curran) (Figures 89-93) Rhynchiodexia arida CURRAN, 1931:93. West (1950:110); Sa- BROSKY AND ARNAUD (1965:988); CoLE (1969:543). [LEc- TOTYPE (here designated), a female, deposited in CUIC, labeled, ‘‘Pinnaleno Mts., Ft. Grant, Ariz.’’/**R. C. Shan- U., Lot. 919, Sub. 120, S. S. West’’/**2 Holotype Rhyn- chiodexia arida Curran’’/*‘Holotype, Cornell U., No. 1937°’/**Cornell U., Lot. 922, Sub. 39’’/**Lectotype Rhyn- chiodexia arida Curran designated by D. Wilder 1976.”’] TAXONOMIC Notes.—Curran introduced the name into the literature in his 1930 key. West, however, was the one to segregate and name the species. West's ““holotype,’’ a female, has been designated as lectotype, even though it bears a holotype label with Curran as author. A speci- 46 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 1 FIGURES 89-93. NORTH AMERICA Ptilodexia arida (Curran). Fig. 89. Genitalia of male, posterior view. Fig. 90. Ejaculatory apodeme. Fig. 91. Genitalia of male, lateral view. Fig. 92. Head of male, lateral view. Fig. 93. Geographical distribution. men from Manitoba has been designated as paralectotype. This specimen has an undated Curran determination label, and I think that he compared the specimen with West’s type, felt that they were the same, and consulted both when writing the 1930 key. West’s type is a female in poor condition, but the Manitoba male is in good condition. The Manitoba specimen is misassociated, actually belonging to P. major. DIAGNOSIS.—Ptilodexia arida adults are unique in the following combination of charac- ters: broad, square face, no parafacial hairs ven- tral to second antennal segment; plumosity on arista long, up to twice length of second antennal segment; legs of females pale in color; abdomen of males and many females reddish laterally. MATERIAL EXAMINED.—Sixty-four males and 33 females were examined. DISTRIBUTION.—Prtilodexia arida ranges from Veracruz in Mexico into the mountains of Ari- zona and Utah. There is one record from Jef- ferson County, Idaho. BIOLOGICAL Notes.—The flight period is from March to October, the majority of speci- mens being captured in July and August. There is only one March record and none for the months of April and May. One specimen from Morelos in Mexico was collected in November. This species inhabits a mountainous region. Specimens have all been collected in the moun- tains, some at elevations up to 2,560 m. One exception is a male collected in Jefferson Coun- ty, Idaho, an area of extensive lava fields. Other specimens have been collected by special de- vices, such as UV light and Malaise traps. One female was collected on Ceanothus (Rhamna- ceae), and four males were swept from Ipomoea (Convolvulaceae). There are five reared speci- mens; however, they lack data. They are pinned with their hosts, small scarabaeid larvae. Ptilodexia ponderosa (Curran) (Figure 6) Rhynchiodexia ponderosa CURRAN, 1930:93. West (1950:110); SABROSKY AND ARNAUD (1965:989). [LECTOTYPE, female, WILDER: NEARCTIC PTILODEXIA 47 Ficures 94-98. Ptilodexia major (Bigot). Fig. 94. Genitalia of male, posterior view. Fig. 95. Ejaculatory apodeme. Fig. 96. Genitalia of male, lateral view. Fig. 97. Head of male, lateral view. Fig. 98. Geographical distribution. deposited in CUIC, labeled, **Chokoloskee, Fla.’’/** Wing Slide, Cornell U. Lot. 919, Sub. 129, L. S. West’’/** 2 Ho- lotype Rhynchiodexia ponderosa Curran’’/**Holotype Cor- nell U. No. 1936’’/**Cornell U. Lot. 922, Sub. 38°*/**Lec- totype Rhynchiodexia ponderosa Curran.’’} TAXONOMIC NoTtes.—This species was orig- inally segregated and named by West, but the name was introduced into the literature by Cur- ran and dates from his 1930 key. The unique specimen was not designated as a holotype. Since Curran did not mention the number of specimens which he had, even though there is only one, it must be designated lectotype. The specimen is in excellent condition. DIAGNOosSIs.—Since this species is represented by one specimen, it is difficult to diagnose. The P. ponderosa specimen can be distinguished from members of all other related species by the following combination of characters. Size large, color light reddish brown; face wide; parafacial hairs small, brown, inserted only on anterior half of parafacial; vibrissal angle of head quite pro- nounced; epistome protrudes noticeably; third antennal segment long; length of plumosity 1.6 times length of second antennal segment; haus- tellum length 0.6 times head height; legs uni- formly testaceous. DISTRIBUTION.—The lectotype was collected in extreme southern Florida and almost certainly represents a West Indian species. It more closely resembles specimens from these islands (most of which have small, pale, parafacial hairs, long plumosity on the arista, and a me- dium-long haustellum) than those from the United States. It differs from the specimens I have seen from the islands in being much larger and having uniformly pale legs. major Group Ptilodexia major (Bigot) (Figures 94-98) Rhamphinina major BiGot 1888:265. WULP (1891:228); GiG- L1o-Tos (1894:60); BRAUER (1899:509); ALDRICH (1905:504); SABROSKY AND ARNAUD (1965:989); CoLe (1969:543); Gul- MARAES (1971:33). [LECTOTYPE (here designated), male, deposited in BMNH, labeled, **Lectotype’’/“Rhamphinina major Bigot’’/**Mexico. Ex. coll. Bigot. Pres. by G. H. Ver- rall, Oct. 1904. 1904-274"’/**Lectotype Rhamphinina major Bigot designated by D. Wilder 1975.’”] Rhynchiodexia tincticornis BIGOT, 1888:266. WULP (1891:228): BRAUER (1899:509); ALDRICH (1905:504); GUIMARAES 48 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 1 (1971:34). [Lectotype (here designated), male, deposited in BMNH, labeled, *‘Lectotype’’/““Rhynchiodexia tincti- cornis Bigot’’/**Mexico. Ex. coll. Bigot. Pres. by G. H. Verrall, Oct. 1904. 1904-274’’/Bigot’s original inked label/ Austen’s penciled note regarding Bigot’s label/*‘Lectotype Rhynchiodexia tincticornis Bigot designated by D. Wilder 1975.’”] Dexiosoma fumipennis BiGoT, 1888:270. WuLP (1891:230); BRAUER (1899:508); ALDRICH (1905:504); GUIMARAES (1971:34). [HoLotyPe, female, deposited in BMNH, la- beled, ‘‘Holotype’’/*‘Rhynchiodexia fumipennis’’/**Mexico. Ex. coll. Bigot. Pres. by G. H. Verrall, Oct. 1904. 1904—- 274°’/2 folded labels.| NEw SYNONYMY Rhynchodexia fraterna WULP, 1891:229. GIGLIo-Tos (1894:61); ALDRICH (1905:499); THOMPSON (1963:517); GUIMARAES (1971:33). [LEcToTYPE (here designated), male, deposited in BMNH, labeled, *‘Lectotype’’/**Cuernavaca, Morelos. June H. H. S.’’/*'3’’/"*B. C. A. Dipt. Il Rhynchodexia fra- terna, v.d.W.’’/*‘Central America Pres. by F. D. Godman, O. Salvin. 1903-172’’/**Lectotype Rhynchodexia fraterna Wulp, desig. by D. Wilder 1975.’’>] NEw SYNONYMY Rhynchodexia omissa WULP, 1891:235. ALDRICH (1905:499); GUIMARAES (1971:33). [LECTOTYPE (here designated), male, deposited in BMNH, labeled, *‘Lectotype’’/**N. Yu- catan, Gaumer’’/**d’’/**B. C. A. Dipt. II Rhynchodexia om- issa v.d.W.’’/**Central America Pres. by F. D. Godman, O. Salvin. 1903—172’’/**Lectotype Rhynchodexia omissa Wulp designated by D. Wilder 1975.’"] NEw SYNONYMY Ptilodexia harpasa (partim): Davis (1919:83). (misidentifica- tion) Rhynchiodexia arida (partim): CURRAN (1930:93). (misidenti- fication) TAXONOMIC Notes.—The confusion and syn- onymy of this species in the literature is under- standable. It is quite variable in color charac- teristics and size of its members. It also has a broad geographical range. Bigot proposed three names for P. major and Wulp two. The syntype series for P. major consists of five males. Two of these are misassociated, one resembling specimens of P. conjuncta but with a shorter haustellum, the other resembling spec- imens of P. scutellata but with much hairier parafacials. The other three are conspecific. The specimen chosen as a lectotype has the upper parafacials a little hairier than the others, and its facial tomentum has a slightly yellowish cast. The series is in poor condition, the lectotype having only one leg and one antenna; most of the setae are broken. Associated with the syntype series is a pen- ciled label in Austen’s handwriting. It mentions that Brauer (1899:509) felt the species belonged to Ptilodexia. He continued, however, saying that he (Austen) felt that the series was com- prised of representatives of three distinct gen- era, only one specimen belonging to Prilodexia. The specimen which Austen felt was a Prtilo- dexia is the misassociated type resembling spec- imens of P. conjuncta. His reasoning is not clear, but perhaps he failed to see the minute parafacial hairs on true P. major members and therefore felt it belonged in another genus. The type which resembles adults of P. scutellata could understandably be considered as repre- senting a different genus by a worker who be- lieved in restricted genera. Bigot stated the type-locality as Mexico and Washington Territory. All five syntypes have la- bels reading *‘Mexico.”’ Both of the cotypes of Rhynchiodexia tincti- cornis are in poor condition. The specimen cho- sen as lectotype is oily, the tomentum therefore obscured. Wulp (1891) redescribed the species and restricted the type-locality to Paso del Macho, possibly taking the name of the locality from another Bigot label. Since this locality is within the range of the species, the restriction is accepted. Wulp stated that he had a female from Costa Rica. It is possible, but unlikely, that the species ranges that far south. Brauer (1899) placed P. major in Ptilodexia and listed P. tinc- ticornis aS a synonym. Bigot described R. fraterna in 1891 from 21 males and 5 females. Of the four male specimens which I had the opportunity to see, one was misassociated, the other three belonged to P. major. Wulp himself stated after his description that three of his largest specimens ‘“‘agree fully with a typical example of Rhamphinina major.” and he then cited a few minor color differences. He also felt that some of the smaller specimens ‘bear a striking resemblance”’ to P. fumipennis adults. The lectotype, a male from Cuernavaca, is in good condition. Rhynchiodexia omissa was described from three male and two female cotypes. The male which I saw and designated as lectotype belongs to P. major. The specimen was collected in northern Yucatan. It is in excellent condition and is similar in color to the lectotype of P. tincticornis. Guimaraes (1971) was the first to place omissa in Ptilodexia. Another name must be considered while dis- cussing P. major. This is Rhamphinina dubia Bigot, the type-species of that genus. Bigot de- scribed R. dubia from two females labeled only ‘*Mexico.”’ The two specimens now in the Brit- ish Museum are, however, a male and a female. The male belongs to P. major and the female to Macrometopa calogaster, the latter placed by WILDER: NEARCTIC PTILODEXIA Austen. Townsend (1931:100) spoke of a female holotype; however, this cannot be considered a lectotype designation of the type-series accord- ing to Crosskey (in litt.). I have designated the female of the original material as the lectotype, giving the male the status of misassociated para- lectotype. Had the other specimen (d, P. major) been chosen as the lectotype, the name Rham- Phinina (1885) would have priority over Ptilo- dexia (1889) and our commonly used name would have to be changed. Macrometopa now becomes a synonym of Rhamphinina. DIAGNosis.—Ptilodexia major exhibits much variation throughout its range, and after a more thorough sample is available, may be found to consist of more than one species. It is most dif- ficult to diagnose, and specimens can be more easily identified by first eliminating other species. Thorax dark, the notum distinctly striped; parafacial hairs small, pale, inserted only on an- terior half of parafacial, ventrally on face to apex of antennae; they are light brown, medium length, and/or scattered on parafacial in few in- dividuals; haustellum length varied, from 0.55 to 0.65 times head height, in some small specimens it is slightly longer; palpi longer than half haus- tellum length in most individuals; length of plu- mosity on arista more than twice length of sec- ond antennal segment; abdomen reddish with dark median stripe in most males, entirely dark or dark with fourth tergite reddish in a few. VARIATION.—Ptilodexia major exhibits more variability than do most Ptilodexia. A summary of this variation is given by Wilder (1976). There are some specimens, notably those from Texas, New Mexico, Colorado, and Ne- braska, which seem to fall somewhere between specimens of P. incerta and P. major, two species which are, from available records, allo- patric. It is possible that these represent an as yet unrecognized species and that true P. major is a more southern species. Most of the char- acters involved, however, are vague and in other species are part of normal intraspecific varia- tion. This problem may be solved when a larger sample is available, thereby allowing the use of Statistical techniques. I have placed these spec- imens in P. major because it is with members of this species that they share the most character States. Most Nearctic Prilodexia do not have ranges which extend into neotropical Mexico (P. con- 49 juncta is an exception). Specimens of P. major from Chiapas, Campeche, and Yucatan, how- ever, differ very little from those from Arizona, at least in characters which easily separate adults or other species in the genus. I feel jus- tified in placing all these specimens in P. major, although when more Mexican material is avail- able for study, it may show that more than one species is involved. MATERIAL EXAMINED.—One hundred fifty- two males and 77 females of P. major were ex- amined. DISTRIBUTION.—This species ranges from Nebraska southward through Colorado, Kansas, Arizona, New Mexico, and Texas into southern Mexico. BIOLOGICAL Notes.—The flight period ranges from April to October, with the main activity in July and August in most parts of the range. Members of Texas populations exhibit two pe- riods of activity, one in April and May, and another in September and October. Specimens reared indoors appear to emerge much earlier, in December, February, and March. Specimens have been taken at both low and high elevations, the highest being 2,130 m in Coahuila, Mexico. Methods used to collect P. major adults are sweeping, UV light trap, Ma- laise trap, and rearing. Flowers from which P. major adults have been captured include: Se- necio sp., Baccharis glutinosa (Compositae); Lepidium thurberi (Cruciferae); Croton sp. (Eu- phorbiaceae); Medicago sativa, Dalea albiflora (Fabaceae); Marrubium vulgare (Labiatae); Gossypium sp. (Malvaceae), and Tamarix sp. (Tamaricaceae). Unlike other Prtilodexia, com- posites are not the group of flowers most fre- quently visited. Several specimens among the material exam- ined have been reared and are associated with skins of host larvae. Unfortunately, these skins cannot be identified. Host data on some speci- mens list the host as ‘May beetle’? and Phyl- lophaga farcta. David (1919) mentions Phylloph- aga grubs collected on 29 April, at Austin, Texas: Ptilodexia larvae were first observed from May 18 to June 18. In these cases, adult flies issued between June 11 and July 10, the puparium stage varying from 20-32 days. Grubs collected in New Mexico May 10 and 24 showed parasitism from May 20 to July 7, while larvae from the same locality collected on September 7 and confined in indoor cages showed parasitic larvae from October 7 to February 23. 50 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 1 Ficures 99-103. NORTH AMERICA Ptilodexia incerta (West). Fig. 99. Genitalia of male, posterior view. Fig. 100. Ejaculatory apodeme. Fig. 101. Genitalia of male, lateral view. Fig. 102. Head of male, lateral view. Fig. 103. Geographical distribution. Aldrich called these New Mexico and Texas specimens Prilodexia harpasa, but they are typ- ical of P. major adults. Davis also reports mem- bers of this species being reared from grubs of Aphonus pryiformis collected in Las Vegas and Caffrey, New Mexico. Ptilodexia incerta West (Figures 99-103) Ptilodexia incerta West, 1925:131. LEONARD (1928:822); CuRRAN (1930:93); SABROSKY AND ARNAUD (1965:989). [HoLotTyPe, male, deposited in CUIC, labeled, **River- head, L.I., N.Y. 6-30-21’’/**Holotype Ptilodexia incerta West d’’/**Holotype Cornell U. No. 1873’/*‘Cornell U. Lot. 922, Sub. 45’’] Ptilodexia proxima West, 1925:133. LEONARD (1928:822); CURRAN (1930:93); West (1950:pl. I, fig. 4); SABROSKY AND ARNAUD (1965:989). [HOLOTYPE, male, deposited in CUIC, labeled, “Riverhead, L.I., N.Y.’’/**Wm. T. Davis, Aug. 5, 1917°’/*‘Wing Slide, Cornell U. Lot. 919, Sub. 121, L. S. Cornell U. No. 1877°’/**Cornell U. Lot 922, Sub. 47’°] New SYNONYMY Rhynchodexia elevata WEST, 1925:135. LEONARD (1928:821); SABROSKY AND ARNAUD (1965:988). [HOLOTYPE, male, de- posited in CUIC, labeled, ‘Riverhead, L.I., N.Y., Aug. 20, 1922’’/*‘Wing Slide Cornell U. Lot. 919, Sub. 123, L. S. West’’/*‘Holotype Rhynchodexia elevata West 3°’/**Holo- type Cornell U. No. 1871°’/**Cornell U. Lot. 922, Sub. 36.”’| New SYNONYMY Ptilodexia harpasa (partim): JOHNSON (1925b:208). (misiden- tification) Rhynchiodexia levata West: CURRAN (1930:93). (lapsus cala- mus) TAXONOMIC Notes.—West described P. in- certa and its two synonyms in the same paper. The type-localities of all three are Riverhead, Long Island. None of his holotypes, all males, show any significant differences. The type of P. proxima is large, that of P. elevata is small, while the type of P. incerta is intermediate in size. West placed P. elevata in the genus Rhyn- chodexia because he thought that it lacked para- facial hairs. These hairs, however, are present and are similar in size, color, and number to those of the other two species. He also felt that the P. elevata adult differed in having the vi- brissae inserted far above the oral margin, another characteristic which is the same in the other two species. WILDER: NEARCTIC PTILODEXIA West stated no diagnostic difference between the P. incerta and P. proxima males, and the descriptions of the two are nearly identical. The types of these three species names are in excellent condition, with the possible exception of the holotype of P. incerta, which is dusty. They are all deposited in CUIC. The three are unquestionably conspecific. DiaGnosis.—Adults of P. incerta can be dis- tinguished from those of other species of the ge- nus overlapping in range by the following com- bination of character states: parafacial hairs tiny and pale (rarely brown), present only on ex- treme upper anterior portion of parafacial; length of third antennal segment about 1.5 times length of second; length of plumosity on arista twice length of second antennal segment. Haus- tellum length 0.5 to 0.6 times head height; palpi, broad in many individuals, equal to or less than half haustellum length. Thorax distinctly striped; in many specimens there are few or no infra- squamal setulae; in most abdomen reddish with a median longitudinal stripe. MATERIAL EXAMINED.—Specimens exam- ined include 105 males and 32 females. DisTRIBUTION.—This species occurs through- out the eastern U.S., west to about the 100th meridian. BioLoGicAL Notes.—The flight period lasts from May to September, with most specimens being collected during July. In certain areas, es- pecially in the Southeast, specimens have been taken in every month from 10 May to 26 Sep- tember. Associated data show that P. incerta adults have been collected while at blooms of Cirsium sp. and Solidago sp., both Compositae. They have also been taken on high prairie in Hennepin County, Minnesota. A male was taken from a phymatid, which had presumably captured it as prey. Records show that P. incerta adults have only been collected at low elevations. Ptilodexia maculata, new species (Figures 104-108) Type-LocaLity.—The holotype was collected 14 miles [ca. 22.5 km] north of Silver City, Grant County, New Mexico. TypPE-SPECIMENS.—The holotype, a male, is deposited in MSUC, the allotype, from Show-Low Arizona, in USNM. Complete data from these specimens and the 12 male and 10 female paratypes are listed below. DIAGNosIs.—Ptilodexia maculata is a distinc- tive species, its members differing from those of 51 its congeners by the following combination of character states. Reddish brown; face wide, an- teroventral angle not strongly protruding; para- facial hairs small, pale, occuring only along an- terior edge of parafacial; oral margin only slightly protruding; infrasquamal setulae lack- ing; haustellum length 0.4 to 0.5 times head height. The legs of males and females are pale, the femora with striking dark brown apicoven- tral patches. DESCRIPTION.—Male: Total body length 16 mm, brownish with median abdominal stripe. Head height 3.5 mm; profrons 0.3 times head length. Facial tomentum pale gold, shining, fine, not obscuring integumental color; parafrontal hairs long, dark, sparse; parafacial hairs small, pale, inserted only along anterior edge of para- facial and not ventral to antennal apex. Width of frontal vitta at base of antenna 0.55 mm, at narrowest part 0.25 mm; carina well developed, long, wide; facial cavities slightly darkened. Height of gena 0.35 times head height. One pair of ocellar setae, 2 pairs of postocellars subequal to ocellars, | pair of shorter postverticals, | pair of long inner verticals, and | pair of outer ver- ticals subequal to postorbitals. Fifteen pairs of postorbital setae not interspersed with smaller hairs, those closest to ocellar triangle long, be- coming smaller with distance from ocellar tri- angle, curved only slightly anterad: between postorbitals and the dense whitish occipital hair are a few scattered hairs. Twelve pairs of frontal setae; one pair of oral vibrissae 0.25 mm from oral margin, the depression between them 0.4 mm wide; 12 additional oral setae, 1 above vi- brissae fairly strong, peristomals subequal to each other. Epistome only slightly protruding, its width 1.3 times length of first two antennal segments. Haustellum broad, brown, 0.4 times head height; palpi testaceous, 0.7 times haustel- lum length. Antennae unique; second segment light brown, testaceous apically; third segment testaceous on basal half, light brown apically; arista and plumosity pale brown. Third segment of antenna slightly narrowed apically, 1.3 times length of second segment; length of plumosity on arista 1.5 times length of second antennal segment. Thorax: Mesonotum rufotestaceous, brown posteriorly, subshining; tomentum beige, striping indistinct. Integument of humeri and postalar calli rufotestaceous. Six pairs of strong humeral setae, | posthumeral, | presutural, and 2 pairs of discal scutellars. Pleura rufotestaceous 52 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 1 NORTH AMERICA Figures 104-108. Ptilodexia maculata, n.sp. Fig. 104. Genitalia of male, posterior view. Fig. 105. Ejaculatory apodeme. Fig. 106. Genitalia of male, lateral view. Fig. 107. Head of male, lateral view. Fig. 108. Geographical distribution. marked with brown, tomentum beige, translu- cent; infrasquamal setulae absent; squamae and wings pale testaceous; epaulet light brown, ba- sicosta testaceous, mediotergite tomentose, subshining. Legs rufotestaceous, tibiae dark- ened basally and apically, coxae and trochanters marked with dark brown; femora with dark brown, strongly contrasting ventral patches on apical half, bases of setae inserted on dark patches pale in color, pale setal bases on mac- ulae of anterior femur coalesced into pale stripe: tarsi black. Anterior femur with 12 dorsal setae, 8 posterodorsals, and 13 posteroventrals, tibia with a distinct dark dorsal line. Middle femur with 2 anterior setae in | row. Posterior femur with a row of 7 anterodorsal setae, 9 anteroven- trals, 6 posteroventrals, and | anterior seta; pos- terior tibia curved. Abdomen rufotestaceous with dark brown, narrow, median stripe; tomen- tum whitish, brown above median stripe. First syntergite with 3 pairs of lateral marginal setae. Second tergite with 3 pairs of median discals, 12 dorsal and lateral marginals, and 4 pairs of lat- eral discal setae. Ventral margins of tergite marked with brown. Genitalia: Eighth tergite with numerous fine, dark setae. Ninth tergite with many long, dark setae, lateral swelling small. Surstylus huge, lateral angle smoothly rounded, depression fairly deep, with only fine hairs; in profile, smoothly rounded with slight ventral swelling. Cerci elongate, lateral lobes laterally curved strongly anterad, mesal margins slightly concave; height of arms slightly greater than that of lobes, arms not reaching ventral margins of surstyli, in profile, slightly bulging, tips large, round. Ejaculatory apodeme with bulb and stem distinct, the latter slightly longer and curved. Bulb widely open on one side and top; opposite side strongly decumbent. Female: Similar to male except for usual sexual differ- ences and the following. Width of frontal vitta at narrowest point 0.4 mm, at antennal base 0.7 mm. Height of gena 0.35 times head height. Ten pairs of short postorbital setae, with 1 or 2 small hairs interspersed; 7/8 frontal setae. Anterior fe- mur with 9 posteroventrals, 7 posterodorsals, and 9 dorsal setae. Middle femur with 4 anterior setae in 2 rows, 4 posteroventrals, and no an- WILDER: NEARCTIC PTILODEXIA Ficures 109-113. Ptilodexia flavotessellata (Walton). Fig. NORTH AMERICA 109. Genitalia of male, posterior view. Fig. 110. Ejaculatory apodeme. Fig. 111. Genitalia of male, lateral view. Fig. 112. Head of male, lateral view. Fig. 113. Geographical distribution. teroventrals. Posterior femur with no anterior setae, 5 anterodorsals, 3 posteroventrals, and 3 anteroventrals. Abdomen rufous with vague dark-brown median stripe, pollen whitish, light brown around median marginal setae. First syn- tergite with | pair of strong lateral marginal se- tae. Second tergite with | pair of lateral margin- als, median discals, and median marginals. Third tergite with | pair of median discals and 5 dorsal and lateral marginal setae. VARIATION.—Intraspecific variation is slight. Total body length ranges from 10 to 16 mm. The parafacial hairs are brownish in some speci- mens, but they are small in all adults. The area between the oral vibrissae and the oral margin has the height equal to width in some specimens. The integument of the notum in some individuals is rufotestaceous striped with dark brown. All facial and body tomentum are shades of brown in some adults, in females, quite dark. MATERIAL EXAMINED.—Thirteen males and eleven females were examined from the following localities. Arizona: Cochise Co.: 1d, Garden Canyon, Huachuca Mts., 30 July 1949, W. Gertsch (AMNH); 16, Ramsey Canyon, 15 miles [ca. 24 km] S of Sierra Vista, 6,000 ft [ca. 1,830 m], 1 July 1964, Sternitzky (CNCI); 1d, Rucker Canyon, Chiricahua Mts., 22 Aug. 1965, G. Wallace (UCRC). Navajo Co.: 12, Show-Low, 24 July 1956, Butler (UAIC). Santa Cruz Co.: 12, Madera Canyon, 4,880 ft [ca. 1,490 m], 15 June 1965, D. Harrington, 12, 25 July, 1d, 6 Aug. 1961, at light, G. Nelson, 12, 11 Aug. 1965, D. Harrington, 2° 2, 12 Aug., 1d, 15 Aug., 1d, 17 Aug. 1972, D. Knull, 1¢12, 26 Aug. 1965, D. Harrington, 1d, 1 Sep., 16329, 7 Sep., 12, 15 Sep. (UCDC, OSUC). New Mexico: Catron Co.: 266, Mogollon Mts., 29 Aug. 1951 (CASC). Grant Co.: 16, 14 miles [ca. 22.5 km] N of Silver City, 8 July 1961, G. Eickwort (MSUC). Hidalgo Co.: 16, Guadalupe Canyon, 7 Aug. 1967, J. Smith (UCRC). BIoLoGIcAL Notes.—The flight period lasts from July to September. Adults of this species have been collected at lights. They inhabit mountainous areas and have been taken at ele- vations from 1,490 m to 1,830 m. DERIVATION OF NAME.—The name of this species refers to the striking femoral maculae on its members. Ptilodexia flavotessellata (Walton) (Figures 109-113) Rhynchiodexia flavotessellata WALTON, 1914:176. Town- SEND (1915:23); (1938:349); SABROSKY AND ARNAUD 54 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 1 (1965:988); CoLe (1969:543). [HoLotyPe, female, taken in copula and pinned with its mate, deposited in USNM, la- beled, *“‘Koehler, New Mexico, 8—-14’’/**Webster No. 7707°’/**W. R. Walton collector.”’] TAXONOMIC Notes.—Walton stated that he described this species from four specimens tak- en at Eagle Tail Mountain, Colfax County, New Mexico. He named a female, pinned in copula with its mate, the holotype. These specimens bear a label which reads only, **Koehler, New Mexico”’ as the locality; however, since Walton himself was the collector, his restriction is ac- cepted. There are more specimens than the de- scription records with identical labels, so the three males which he indirectly makes paratypes cannot be segregated. The type material is in ex- cellent condition. Townsend (1915) erected the genus Myocer- opsis for P. flavotessellata. He believed that specimens of Ptilodexia had the haustellum lon- ger than 0.75 times the head height and that P. flavotessellata, its members with the haustellum length 0.5 times the head height, differed suffi- ciently to be segregated generically. D1aGNosis.—Specimens of P. flavotessellata can be distinguished from those of other North American Ptilodexia by the following combi- nation of character states: general color pale or- ange-brown, tarsi darker; parafacial hairs pale to brown, of medium length, sparse, scattered randomly on parafacial; third antennal segment not much longer than second; length of plumos- ity 1.25 to 1.5 times length of second antennal segment; haustellum light brown in color, nar- row, about 0.5 times head height; area between oral vibrissae and oral margin wider than high; epistome protrudes only slightly. MATERIAL EXAMINED.—Five male and three female specimens were examined. DISTRIBUTION.—Ptilodexia flavotessellata ranges from northeastern New Mexico north into Colorado and Nebraska. BIOLOGICAL Notes.—The collection dates on the specimens examined range from 22 June in Boulder, Colorado, to 9 September in Nebraska. There are no associated ecological data. LITERATURE CITED ALpRICH, J. M. 1905. A catalogue of North American Dip- tera. Smithson. Misc. Collect. 46(2): 1-680. 1924. Notes on some types of American muscoid Diptera in the collection of the Vienna Natural History Mu- seum. Ann. Entomol. Soc. Am. 17:209-218. 1925. Notes on some types of American muscoid Diptera in the collection of the Vienna Natural History Mu- seum (cont.). Ann Entomol. Soc. Am. 18:107—130. AUSTEN, E. E. 1907. The synonymy and general position of certain species of Muscidae (sens. /at.) in the collection of the British Museum, described by the late Francis Walker. Ann. Mag. Nat. Hist., Ser. 7, 19:326-347. Bicot, J. M. F. 1885. Diagnoses de trois genres nouveaux de Diptéres du groupe des Dexiaires. Ann. Soc. Entomol. France (6)5:xi-xi1. 1889. Dipteres nouveaux ou peu connus 34? partie, XLII, Diagnoses de nouvelles especes. Ann. Soc. Entomol. France (6)8:253-270. BLANCHARD, E. E. 1966. Dipteros parasitos de escarabeo- ideos argentinos. Rev. Invest. Agric., Ser. 5, 3(11):175—229. BRAUER, F. 1899. Beitrage zur Kenntniss der Muscaria Schi- zometopa. Bemerkungen zu den original exemplaren der von Bigot, Macquart und Robineau-Desvoidy, beschreiben Muscaria Schizometopa aus der Sammlung des Herrn G. H. Verrall, III. Sitzunsber. K. Akad. Wiss. Wien, Math.- Naturwiss. KI]. 1(108):495—529. AND J. E. VON BERGENSTAMM. 1889. Die Zweifltigler des Kaiserlichen museums zu Wien. IV. Vorarbeiten zu ein- er monographie der Muscaria Schizometopa. Pars I. Denkschr. K. Akad. Wiss. Wien 56(1):69—-180. BRIMLEY, C. S. 1938. Insects of North Carolina. North Car- olina Dep. Agric., Div. Entomol., Raleigh. 491 pp. CHAMPLAIN, A. B., AND J. N. KNULL. 1944. Notes on Penn- sylvania Diptera. Entomol. News 34(7):211-215. CLAUSEN, C. P., J. L. KING, AND C. TERANISHI. 1927. The parasites of Popillia japonica in Japan and Chosen (Korea) and their introduction into the United States. U.S. Dep. Agric. Agric. Res. Serv. Tech. Bull. 1429. 56 pp. Cote, F. R. 1969. The flies of western North America. Univ. Calif. Press, Berkeley and Los Angeles. 693 pp. CoguILLeTT, D. W. 1910. The type species of the North American genera of Diptera. Proc. U.S. Natl. Mus. 37:499— 647. CrosskEY, R. W. 1973a. A revisionary classification of the Rutiliini (Diptera: Tachinidae), with keys to the described species. Bull. Bnt. Mus. (Nat. Hist.) Entomol. Suppl. 19: 1-167. 1973b. A conspectus of the Tachinidae (Diptera) of Australia, including keys to the supraspecific taxa and taxo- nomic and host catalogues. Bull. Brit. Mus. (Nat. Hist.) Entomol. Suppl. 21:1—221. CurRRAN, C. H. 1930. Report on the Diptera collected at the Station for the Study of Insects, Harriman Interstate Park, N.Y. Bull. Am. Mus. Nat. Hist. 61:21-115. . 1934. The Diptera of Kartabo, Bartica district, Brit- ish Guiana with descriptions of new species from other Brit- ish Guiana localities. Bull. Am. Mus. Nat. Hist. 66:287—532. Davis, J. J. 1919. Contributions to a knowledge of the natural enemies of Phyllophaga. Ill. Nat. Hist. Surv. Bull. 8(5):53- 138. Fattic, P. W. 1944. Phyllophaga of Georgia. Emory Univ. Mus. Bull. Georgia, no. 2, 32 pp. FLEMING, W. E. 1963. The Japanese Beetle in the United States. U.S. Dep. Agric. Agric. Res. Serv. Agric. Handb. 236. 30 pp. GiGLio-Tos, E. 1894. Ditteri del Messico. Pt. 3. 76 pp. To- rino. GREENE, C. T. 1922. An illustrated synopsis of the puparia of 100 muscoid flies (Diptera). Proc. U.S. Natl. Mus. 60(10): 1-39. GuIMaAraes, J. H. 1971. Family Tachinidae. Pages 1-333 in WILDER: NEARCTIC PTILODEXIA A catalogue of the Diptera of the Americas south of the United States. 104. Museum of Zoology, University of Sao Paulo, Brazil. Howpen, H. F. 1963. Speculation on some beetles, barriers and climates during the Pleistocene and pre-Pleistocene pe- riods in some nonglaciated portions of North America. Syst. Zool. 12:178-201. . 1966. Some possible effects of the Pleistocene on the distributions of North American Scarabaeidae. Can. Ento- mol. 98:1177—1190. JOHNSON, C. W. 1925a. Diptera of the Harris collection. Proc. Bost. Soc. Nat. Hist. 38(2):57-99. . 1925b. Fauna of New England, List of the Diptera or two-winged flies. Occas. Pap. Bost. Soc. Nat. Hist. 7(15): 1-326. . 1927. The insect fauna with reference to the flora and other biological features. Jn Procter, W., Biological survey of the Mount Desert Region. Part I. Wistar Institute Press, Philadelphia. LEONARD, M. D. 1928. A list of the insects of New York. Cornell Univ. Agric. Exp. Stn. Mem. 101:1-1121. Macquart, J. 1843. Dipteres exotiques nouveaux ou peu con- nus. Mém. Soc. Sci. Agric. Arts Lille 1842:162—460. 1846. Dipteres exotiques nouveaux ou peu connus. Supplement. Mem. Soc. Sci. Agric. Arts Lille (1845), 1844: 133-364. OsTEN SACKEN, C. R. 1878. Catalogue of the described Dip- tera of North America (ed. 2). Smithson. Misc. Collect. 16(2): 1-276. PETERSON, L. O. T. 1948. Some aspects of Poplar Borer (Saperda calcarata Say) (Cerambycidae) infestations under parkbelt conditions. Ann. Rep. Entomol. Soc. Ont. 78:56— 61. Procter, W. 1938. The insect fauna with references to meth- ods of capture, food plants, the flora and other biological features. In Procter, W., Biological survey of the Mount Desert Region. Part VI. Wistar Institute Press, Philadel- phia. 496 pp. 1946. The insect fauna with references to methods of capture, food plants, the flora and other biological fea- tures. Jn Procter, W., Biological survey of the Mount Des- ert Region. Part VII. Wistar Institute Press, Philadelphia. 566 pp. REINHARD, H. J. 1943. New North American Muscoidea. J. Kansas Entomol. Soc. 16:14—23. RoBINEAU-DeEsvoipy, J. B. 1830. Essai sur les Myodaires. Mem. Acad. Sci. Inst. Fr., Ser. 2, 2:1-813. Ross, H. H. 1965. Pleistocene events and insects. Pages 583-— 595 in Wright, H. E., Jr., and D. G. Frey, editors. The Quaternary of the U.S. Princeton University Press, Prince- ton, N.J. SABROSKY, C. W., AND P. H. ARNAUD, JR. 1965. Family Tachinidae. Pages 961-1108 in Stone, A., C. W. Sabrosky, W. W. Wirth, R. H. Foote, and J. R. Coulson. Catalogue of the Diptera of America North of Mexico. U.S. Dep. Agric. Agric. Res. Serv. Agric. Handb. 276. STEYSKAL, G. C. 1976. The terminology of the bristles on the upper back of the head in the higher Diptera. J. Kansas Entomol. Soc. 49(2): 155-159. THompson, W. R. 1963. The tachinids of Trinidad. II. Echy- nomyiines, dexiines and allies. Can. J. Zool. 41:335-576. TOWNSEND, C. H. T. 1915. Proposal of new muscoid genera for old species. Proc. Biol. Soc. Wash. 28:19-23. . 1921. Notes on American oestromuscoid types. Rev. Entomol. 1:65—104. . 1938. Manual of myiology in twelve parts. Pt. 7: Oes- troid generic diagnoses and data (Gymnosomatini to Seno- stomatini), 434 pp. Sao Paulo. WALKER, F. 1837. Descriptions, etc. of the insects collected by Cpt. P. P. King, R.N.F.R.S., in the survey of the Straits of Magellan, Diptera. Trans. Linn. Soc. London. 17:331- 359. 1849. List of specimens of dipterous insects in the collection of the British Museum. London. 4:689-1172. . 1852. Diptera. Pages 157-252, 253-414 in Saunders, W. W., ed. Insecta Saundersiana. Vol. 1. London. WALTON, W. R. 1914. Report on some parasitic and preda- ceous Diptera from northeastern New Mexico. Proc. U.S. Natl. Mus. 48:171-186. West, L. S. 1924. New northeastern Dexiini. Psyche 31:184— 193. . 1925. New Phasiidae and Tachinidae from New York State. J. N.Y. Entomol. Soc. 33:121-135. 1950. The status of Rhynchiodexia robusta Curran together with a consideration of certain cephalic and other characters useful in muscoid taxonomy. Pap. Mich. Acad. Sci. Arts Lett. 34:109-117. WivperR, D. D. 1976. Systematics of the Nearctic Ptilodexia Brauer and Bergenstamm (Diptera: Tachinidae). Ph.D. dis- sertation. Pennsylvania State University. 390 pp. WILLISTON, S. W. 1896. On the Diptera of St. Vincent (West Indies). Trans. Entomol. Soc. London; 253-446. Wray, D. L. 1938. Insects of North Carolina. North Carolina Dep. Agric. Third Supp., 181 pp. Raleigh, N.C. Wu tp, F. M. VAN DER. 1891. Family Muscidae. Pages 225— 248 in F. D. Godman, O. Salvin, editors. Biologia Centrali- Americana. Zoologia. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF S Vol. 42, No. 2, pp. 57-67, 6 figs., 2 tables. F=5 poe, IENCES RARY MAR 13 197¢ 1979 Msen. Sods. eS if 2. THE SNAKE EELS (PISCES, OPHICHTHIDAE) OF THE HAWATIAN ISLANDS, By WITH THE DESCRIPTION OF TWO NEW SPECIES John E. McCosker Steinhart Aquarium, California Academy of Sciences, San Francisco, California 94118 ABSTRACT: The ophichthid eels of the Hawaiian Islands (including Johnston and the Leeward islands) are reviewed; included are species new to Hawaii and extralimital records of species previously considered to be endemic. A key to species identification is provided. Two new species captured in deepwater traps off Oahu are described: Muraenichthys puhioilo, subfamily Myrophinae, captured at 275 m, and Ophichthus kunaloa, subfamily Ophichthinae, captured at 350 m. Data concerning the following species are provided: Schismorhyn- chus labialis, Muraenichthys cookei, M. macropterus, Apterichtus flavicaudus, Ichthyapus vulturis, Phaenomonas cooperae, Callechelys luteus, Myrichthys maculosus, M. bleekeri, Cirrhimuraena playfairii, Brachysomophis sau- ropsis, B. henshawi, Phyllophichthus xenodontus, Ophichthus polyophthalmus and O. erabo. Differences in ver- tebral number of populations of Myrichthys maculosus are discussed and the eastern Pacific nominal species M. xysturus (Jordan & Gilbert), M. tigrinus Girard and M. pantostigmius Jordan & McGregor are placed in its synonymy. The endemism (5 of the 15 species) of the Hawaiian ophichthid fauna and the problems of populations and species differences are discussed. INTRODUCTION The snake eels, family Ophichthidae, of the Hawaiian Islands (including Johnston and the Leeward islands) were first treated by Jordan and Evermann (1905) and subsequently re- viewed by Gosline (1951) and Gosline and Brock (1960). Recent collections by the George Van- derbilt Foundation, John E. Randall of the Bish- op Museum, and Thomas A. Clarke of the Uni- versity of Hawaii have added important additional specimens. The Hawaiian Archipela- go contains a particularly interesting eel fauna in terms of its abundance and the range of dis- tributional conditions which exist, including species that are endemic to the islands as well as those that are distributed eastward to Austra- lia and the Red Sea. This, while recognizing the dispersal mechanism allowed by the leptoceph- alus larva, provides an intriguing study for ma- rine zoogeographers. Those considerations, as well as the recent capture of other Hawaiian ophichthids and two apparently undescribed deepwater species, have prompted this review. METHODS All measurements are straight-line (point to point). Standard length, trunk length, and tail length were read on a 300-mm ruler with 0.5-mm gradations and were recorded to the nearest 0.5 mm. All other measurements were made with dial calipers and were recorded to the nearest 0.1 mm. Head length was measured from the snout tip to the posterodorsal margin of the gill opening; trunk length was taken from the end of the head to mid-anus; body depth does not in- clude the fin. Vertebrae (which include the last centrum) were counted from radiographs. Comparisons are based in part on specimens [57] 58 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 2 extralimital to the Hawaiian Islands when ma- terial was insufficient. Partial synonymies are provided where applicable to the Hawaiian Is- lands taxa and where new synonymies are pro- posed. Specimens utilized in this study are deposited in the following institutions: Australian Mu- seum, Sydney (AMS); Academy of Natural Sci- ences of Philadelphia (ANSP); British Museum (Natural History) (BMNH); Bernice P. Bishop Museum (BPBM); California Academy of Sci- ences (CAS), now including the George Van- derbilt Foundation (GVF) and the Stanford Uni- versity collections (SU); Hawaii Institute of Marine Biology (HIMB); Los Angeles County Museum of Natural History (LACM); Scripps Institution of Oceanography (SIO); University of Hawaii (UH); and the National Museum of Natural History (USNM). KEY TO THE OPHICHTHID EELS OF JOHNSTON AND THE HAWAIIAN ISLANDS la. Caudal fin rays conspicuous, confluent with dorsal and anal; tail tip flexible; gill openings mid-lateral, a constricted open- ing; pectoral fin absent in Hawaiian species. Subfamily MYROPHINAE ____- 2 Ib. Tail tip a hard or fleshy, finless point; gill openings mid-lateral to entirely ventral, unconstricted; pectoral fin present in some species. Subfamily OPHICHTHI- NAE 2a. A prominent median toothed groove on ventral side of snout, bordered by der- mal folds, extending forward to anterior nostrils; anterior nostrils elongated tubes equalito.eyeunlencth sss eee ae SY eee Schismorhynchus labialis 2b. Ventral side of snout without a promi- nent groove bordered by dermal folds; anterior nostrils less than eye in length 3a. Teeth absent on vomer, absent or embedded on intermaxillary, those on maxillary and dentary minute or villi- form; dorsal fin origin (DFO) behind anus Schultzidia johnstonensis Teeth present on intermaxillary, maxil- lary, dentary, and vomer; DFO either before or behind anus Posterior nostril entirely outside of mouth; teeth on maxillary, dentary, and 3b. 4a. 4b. ley 8a. 8b. 9a. 9b. 10a. 10b. lla. I 1b. . DFO above or behind anus . Snout blunt; DFO above or slightly be- vomer in broad bands; snout bluntly rounded] === Muraenichthys schultzei Posterior nostril inside mouth, covered externally by a flap; teeth uniserial or biserial, not in broad bands; snout either blunt or acute _...... ae 5 . DFO anterior to anus, about midway to gill openings Muraenichthys puhioilo n.sp. fOFetaNws = Muraenichthys cookei . Snout acute; DFO slightly behind ants: 22-33) ) >. Muraenichthys gymnotus . Body entire finless; coloration either uni- form or darker dorsally, without large spots or saddles At least a minute, short, dorsal fin pres- ent; coloration variable, either uniform, banded, or spotted, or somewhat darker dorsally. nn 9 Posterior nostril opening outside mouth, with a flap; anterior nostril tubular; body extremely elongate; head 15-20 times in i) omen ee? Crikey Apterichtus flavicaudus Posterior nostril opening inside mouth; anterior nostril flush with snout; body moderately elongate; head 11-12 times in TL Ichthyapus vulturis Only fin a short dorsal originating just behind occiput and ending in anterior trunk region; body extremely elongate, the depth 120-150 times in TL ___________- Phaenomonas cooperae Dorsal and anal fins present, the dorsal extending nearly to the tail tip; body moderately to extremely elongate, but the depth less than 120 times in TL — 10 Dorsal fin orgin (DFO) on nape; pectoral fins absent; gill openings inferior, con- verging forward; coloration pale to tan, overlain with small dark spots BERD dat ee eae Callechelys luteus DFO behind nape, either on head or slightly behind gill openings; pectoral fins present; coloration either uniform, spotted, or banded ___ aaa 11 DFO well in advance of gill openings; teeth molariform or granular; pectoral fins broad-based, short and rounded _ 12 DFO before, above, or behind gill open- ings; teeth pointed; pectoral fin base re- McCOSKER: HAWATIAN SNAKE EELS 12a. 12b. 3a. 13b. 14a. 14b. 15a. 15b. 16a. 16b. lak 17b. 18a. 18b. 19a. stricted, opposite upper half of gill open- ings and longer than broad __- 13 Coloration consists of several longitudi- nal series of dark spots along sides and dorsal surface _____ Myrichthys maculosus Coloration consists of about 30 dark sad- dles reaching approximately to the lat- Baling. <22 2 8). Myrichthys bleekeri DFO well ahead of gill openings; edge of upper lip fringed with a conspicuous row of barbels Cirrhimuraena playfairii DFO above or behind gill openings; up- per lip either naked or fringed 14 Postorbital region with a conspicuous transverse depression; lips fringed; ca- nine teeth in jaws and on vomer; color- 2. 0) 15 Dorsolateral profile on head even; lips entire; jaw and vomerine teeth not ex- cessively developed; coloration uniform, spotted. or banded 2... -... 2.2.5 16 Dorsal fine pale; snout contained about Isstumes in head lenpth 22... 2.2.2 Brachysomophis sauropsis Dorsal fin dark with a white border; snout contained about 10 times in head eneth 2... 2. Brachysomophis henshawi Conspicuous leaflike appendages on an- terior nostrils; head and trunk equal to or greater than tail (coloration uniform; vomerine teeth absent) Phyllophichthus xenodontus No leaflike appendages on anterior nos- trils; head and trunk equal to or less than ees cine Pn! Bite ante ual Ci i Vomerine teeth absent or 1-3; head and body coloration light to tan, overlain with a series of black saddles ______________ aol Leiuranus semicinctus A series of teeth on the vomer: color- ation uniform or spotted, not as above ir eh Oe See Od 18 DFO above pectoral tips; pectoral fin elongate, attenuate; coloration uniform, darker dorsally Perse Ophichthus kunaloa n.sp. DFO above gill openings; pectoral fin rounded; coloration not uniform, mark- SBS POMC Mle 0. ee. ee 19 Head and body overlain with numerous ocellated spots; those on body in 3 reg- ular alternating rows, the spots separat- 59 ed by pale interspaces; vertebrae of | SVo(s/ei 0002100 2s Sa, coe es See ee ee ees ae wee Ophichthus polyophthalmus Head and body overlain with numerous dark spots; those on body in 2 irregular rows, the spots about equal in size to their interspaces; vertebrae of 6 speci- mens 152-155... = Ophichthus erabo 19b. Muraenichthys puhioilo, new species (Figures 1-2) Holotype: CAS 29115 (originally SIO 70-32), 355 mm total length, captured in a benthic shrimp trap set overnight at 275 m depth, N of Barber’s Point, Oahu, Hawaiian Islands, by Thomas A. Clarke on 28 October 1969. COUNTS AND MEASUREMENTS (in mm).—The description of this new species is based on the holotype and only known specimen. Total length 355 mm; head length 37.3; trunk length 120; tail length 198; body depth at gill openings 15; body width at gill openings 8; body depth at anus 12: body width at anus 7.5; origin of dorsal fin 86; snout length 8.2; upper jaw length 10.7; eye di- ameter 3.2; interorbital distance 4.4. Total ver- tebrae 160; preanal vertebrae 62. GENERAL DESCRIPTION.—Body elongate, depth at gill openings 23.7 in total length, taper- ing and laterally compressed posteriorly. Head and trunk 2.26 and head 9.5 in total length. Snout moderately acute; lower jaw included, its tip reaching the posterior edge of anterior nos- trils. Anterior nostrils tubular, slightly shorter than eye diameter. Posterior nostril entirely in- side upper lip, before anterior margin of the eye, opening inward, appearing externally as a flap. Eye less than twice in fleshy interorbital dis- tance. Interorbital region flat. Rictus of jaw slightly behind posterior margin of eye. Median fins low, except near tail tip and an- terior portion of anal fin. Dorsal fin arises closer to gill openings than to anus. Paired fins absent. Median fins confluent with caudal. Head pores minute. Single temporal and in- terorbital pores present. Five pores along man- dible, three overlying preopercle. Lateral-line pores difficult to discern posteriorly; 10 pores before gill opening; approximately 140 pores along left side, 65 before the anus. Last lateral- line pore occurs about a head length before tail tip. Teeth slender, small and conical, uniserial throughout. The pattern of dentition is nearly 60 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 2 CHCl Oe Cw Oe 0 Cie SOP TS CeO wer eee te er ey Neth elt a ta oP tee rea FiGures | and 2. Fig. 1. Left lateral view of holotype of Muraenichthys puhioilo McCosker, new species, CAS 29115, 355 mm TL. Fig. 2. Head region of holotype of Muraenichthys puhioilo McCosker, new species. identical to that of Muraenichthys chilensis (cf. McCosker 1970, fig. 4), except that the new species lacks the anteriormost intermaxillary tooth. None is extremely elongate although the anteriormost vomerine tooth is slightly larger than all jaw teeth. Four intermaxillary teeth, forming an inverted v, are followed by 14 uni- serial vomerine teeth. The maxillary teeth are equal in size, 15 right and 17 left. Teeth of lower jaw separated at symphysis, about 17 on each side. Body coloration in isopropyl alcohol uniform tan, although the belly and lateral-line are slight- ly darker. Median fins pale except for the pos- terior portion (slightly longer than head length) of anal fin which is dark. (The functional signif- icance of this highly contrasting fin coloration is not known.) Eyes dark blue. ETYMOLOGY.—From the Hawaiian puhi oilo, small eels about as large in diameter as a finger, here considered a noun in apposition. Eels, par- ticularly puhi oilo, were highly esteemed as food by ancient Hawaiians. Mary Kawena Pukui (1902) wrote that “the eel was a fish of which chiefs were fond .. . so much prized by those of Koolau, Maui. . . that they said only beloved guests were served with eels . . . for eels were considered choicer than wives.” REMARKS.—This individual was captured at a depth far greater than that of any previously McCOSKER: HAWAIIAN SNAKE EELS known Muraenichthys. The nearly 20 species of the genus are generally known from water shal- lower than 50 meters. This specimen was reported by Clarke (1972:312), on the basis of my erroneous iden- tification, as Muraenichthys macropterus Bleeker. I have subsequently examined a radio- graph of Bleeker’s type-specimen (BM 1867.11.28.303) and found it to possess 130 ver- tebrae, with 22 before the dorsal fin origin and 47 before the anal fin origin. Bleeker’s type was from Ambon; a series from Palau (CAS 41186) had 127-132 vertebrae (« = 129.8, n = 5). My examination of more than 100 specimens of M. macropterus from throughout Oceania found them to differ from the new species in having fewer vertebrae, biserial vomerine dentition (be- coming uniserial posteriorly), uniform fin col- oration, and in occupying shallower water. The new species differs from all other species of Muraenichthys, subgenus Scolecenchelys, on the basis of its uniserial dentition, anterior dor- sal fin location, coloration, and vertebral num- ber. In my review of Muraenichthys (McCosker 1970), I followed Schultz (1953) in considering M. breviceps Gunther to be a probable synonym of M. macropterus. | have subsequently rec- ognized M. breviceps as a distinct species and include M. devisi Fowler, M. ogilbyi Fowler, and Aotea acus Phillipps in its synonymy (McCosker and Allen, ms.). I also examined the other known synonyms of M. macropterus, M. owstoni Jordan and Snyder from Japan and Echidna uniformis Seale from Guam, and de- termined that they are M. macropterus. The ho- lotype of M. owstoni (SU 6472) has 131 verte- brae. Ophichthus kunaloa, new species (Figures 3-4) Holotype: CAS 29136 (originally SIO 70-33), 440 mm total length, captured in a benthic shrimp trap set overnight at 350-m depth, SE of Barber’s Point, Oahu, Hawaiian Islands, by Thomas A. Clarke on 31 December 1969. Paratypes: Collected with the holotype. BPBM 21086, 473 mm total length. USNM 218274, only the head and anterior trunk region of a partially eaten specimen. COUNTS AND MEASUREMENTS (in mm).—The condition of the holotype is parenthetically fol- lowed by that of the intact paratype. Total length 440 mm (473 mm); head length 42.0 (45.7); trunk length 135 (144); tail length 263 (283); body depth 61 at gill openings 16 (15); body width at gill open- ings 13.7 (12.5); origin of dorsal fin 58 (68); left pectoral fin length 20.0 (19.5); left pectoral fin base 4.6 (4.5); snout length 8.6 (7.7); upper jaw length 17.2 (20.0); gill opening height 5.6 (5.5): eye diameter 7.0 (8.0); interorbital distance 6.9 (6.9). Total vertebrae 185 (181); preanal verte- brae 66 (67). GENERAL DESCRIPTION.—Body elongate, depth at gill openings 10.3-10.5 in total length, tapering posteriorly. Tail slender, laterally com- pressed posteriorly. Head and trunk 2.49 and head 10.3-10.5 in total length. Snout rounded; lower jaw included, its tip in advance of anterior nostril base. Anterior nostril tubular; posterior nostril at edge of lip, entirely outside of mouth, covered anteriorly by a small flap. Eye large; its center lies slightly behind midpoint of upper jaw. Head broad; interorbital area flat. Median fins low, lying partially within a groove. Dorsal fin arises before end of pectoral. Median fins disappear within a flabby groove before the tail tip. Pectoral fins elongate, the dorsalmost rays tapering posteriorly. Caudal tip naked. Head pores minute, difficult to locate on pre- served specimens. Single temporal and interor- bital pores. Five pores along mandible, two overlying preopercle. Lateral-line pores be- gin above second preopercular pore. Lateral- line pores difficult to discern, about 64 before anal opening. Teeth small, conical; biserial in jaws, the out- er row smaller and closer set. Vomerine teeth biserial anteriorly, followed by a uniserial row of about 10 teeth. Intermaxillary tooth patch surrounded anterolaterally by a ring of 6-8 teeth. Body coloration in isopropyl alcohol tan, be- coming lighter along chin, snout, throat, lower third of trunk region, below the lateral-line of the trunk region, along the dorsal midline, and at the tail tip. A black smudge exists along the lower edge of the tail, about “% head length from the tail tip. Fins pale. Lateral-line pores lie with- in minute white dots. Eyes blue. EtyMoLoGcy.—Named kunaloa in reference to Kuna Loa, the Long Eel, of ancient Hawaiian legends. It is said that the sixth great deed of Maui the Wonder Boy was to behead Kuna Loa after the treacherous eel had assaulted the fair maiden Hina (see Colum 1937). The legend as- 62 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 2 Ficures 3 and 4. Fig. 3. Left lateral view of holotype of Ophichthus kunaloa McCosker, new species, CAS 29136, 440 mm TL. Fig. 4. Head region of holotype of Ophichthus kunaloa McCosker, new species. serts that from the cut tail evolved the common conger eel and from the blood which fell into the fresh and salt water came all of the other Hawai- ian eels. This, clearly, was the first attempt at a phylogenetic interpretation of Hawaiian an- guilliforms. REMARKS.—The depth of capture of the new species is remarkable in that mest benthic ophichthids live shallower than 100 meters. It is entirely likely that future deepwater collections in outlying areas might reveal this or a closely related species. In his key to the Hawaiian ophichthids, Gos- line (1951:309) mentioned a new species of oph- ichthine which possessed characters similar to the new species, that was ‘‘probably from mod- erately deep water,’ and had been killed by the Mauna Loa lava flow of 1950. This specimen was not mentioned in further publications, and neither Gosline (in litt.), John E. Randall of the Bishop Museum (BPBM), nor Leighton Taylor of the University of Hawaii (UH) have been able to locate it after the majority of the UH fish collection had been transferred to the BPBM. The closest relatives to the new species appear to be those congeners which also possess large eyes, similar dentition, posterior nostrils along the lip (rather than opening into the mouth) and preceded by a flap, two rather than three pre- opercular pores, and a plain coloration. Those species of Ophichthus, mostly within the sub- genus Coecilophis Kaup (cf. McCosker 1977), share a preference for moderate-depth sand or mud substrates. Ophichthus kunaloa is partic- ularly similar to O. urolophus (Temminck and Schlegel), an oriental species, which differs in its proportionately shorter tail and much deeper body, and to the eastern Pacific O. pacifici Gunther, a species with a comparatively longer head, deeper body, and white spots along the lateral-line. The following comments comprise new rec- ords and systematic information concerning the Hawaiian ophichthid fauna. SUBFAMILY MYROPHINAE Schismorhynchus labialis (Seale) Muraenichthys labialis Seale, 1917:79 (type-locality, Arno Atoll, Marshall Islands). Leptenchelys labialis: Schultz 1953:80. Schismorhynchus labialis: McCosker 1970:509. REMARKS.—This wide-ranging species has been collected from the Society Islands (BPBM 12016), Marshall Islands, Johnston Island, and Easter Island, but has not been captured in Ha- wail. McCOSKER: HAWAIIAN SNAKE EELS Muraenichthys cookei Fowler Muraenichthys cookei Fowler, 1928:41 (type-locality, Oahu). REMARKS.—I concur with Gosline (1951), that M. cookei, a Hawaiian endemic, is closely re- lated to but distinct from M. laticaudata. A comprehensive discussion of specific differences exists in Gosline (1955:469-470). SUBFAMILY OPHICHTHINAE Apterichtus flavicaudus (Snyder) Sphagebranchus flavicaudus Snyder, 1904:516 (type-locality, between Maui and Lanai). Caecula (Sphagebranchus) flavicauda: Gosline 1951:311. Verma flavicauda: Bohlke 1968:3. Apterichtus flavicaudus: Bohlke and McCosker 1975:4. REMARKS.—This species, previously consid- ered a Hawaiian endemic, has now been col- lected at several South Pacific locations. I com- pared all of Snyder’s specimens with specimens from Midway Island (SIO 68-487) and Rapa Is- land (BPBM 12306), and found them to differ only in vertebral number. Six Hawaiian and Midway specimens had 155-166 (¥ = 158.7) ver- tebrae, whereas six Rapa specimens had 163- 166 (x¥ = 164). Ichthyapus vulturis (Weber and de Beaufort) Sphagebranchus vulturis Weber and de Beaufort, 1916:319 (type-locality, Nasi besar Island, Sumatra). Caecula (Sphagebranchus) platyrhyncha Gosline, 1951:312 (type-locality, Oahu, Hawaiian Islands). REMARKS.—Randall and McCosker (1975) synonymized Caecula platyrhyncha with Sphagebranchus vulturis after comparing Hawaiian specimens with the holotype. Varia- tion exists in the number of preopercular pores (either 3 or 4) of Hawaiian specimens, but the pore number of specimens from other localities was found to be constant. Phaenomonas cooperae Palmer Phaenomonas cooperae Palmer, 1970:219 (type-locality, Gil- bert Islands). REMARKS.—This unmistakable elongate species is known from Hawaii on the basis of a specimen (HIMB 68-52) dredged from a depth of 60 m, offshore from Keehi Lagoon, Oahu, in 1968. The specimen has 262 total vertebrae, 169 preanal, and falls within the range of meristic and morphometric variation of its Indo-West- Pacific conspecifics (McCosker 1975). 63 Callechelys luteus Snyder Callechelys luteus Snyder, 1904:517 (type-locality, near the southern coast of Molokai). REMARKS.—This elongate species, unique in dorsal fin condition and coloration, is known only from the Hawaiian Islands and Midway Is- land (SIO 68-497). Its closest relative appears to be the eastern Pacific endemic C. galapagen- sis McCosker and Rosenblatt. A radiograph of the holotype of C. luteus (USNM 50864) evi- denced 216 vertebrae, with 123 before the anal fin origin. Myrichthys maculosus (Cuvier) Muraena maculosa Cuvier, 1817:232 (type-locality, European Seas?). Pisoodonophis magnifica Abbott, 1861:476 (type-locality, Hawaiian Islands). Ophichthus stypurus Smith and Swain, 1882:120 (type-locali- ty, Johnston Island). REMARKS.—This common species, perhaps better than any other Hawaiian ophichthid, de- picts the isolation of the Hawaiian and Johnston population as evidenced by vertebral number. I am unable except by vertebral number to sep- arate the Hawaiian, Midway, and Johnston specimens from those from the Red Sea, Indian Ocean, Oceania, and the eastern tropical Pacific (see Table 1). For this and related faunal studies I have examined many living and dead speci- mens of Myrichthys as well as the types of most of the nominal species. Until now, I have fol- lowed the conventional view that the eastern Pacific nominal species M. xysturus (Jordan and Gilbert) (which includes M. tigrinus Girard and M. pantostigmius Jordan and McGregor) differs from M. maculosus, although the only signifi- cant difference is that of vertebral number. In the absence of any apparent morphological dif- ferences, I am resigned to recognize these pop- ulations at a subspecific level, and therefore rec- ognize this ophichthid as a trans-Pacific species. The differences in M. maculosus population vertebral numbers is thus parallel to the condi- tion of several trans-Pacific species of muraenid eels (Rosenblatt et al. 1972; McCosker and Ro- senblatt 1975). I suspect that as data are gath- ered concerning the duration of larval life, dis- tance of larval transport, and the critical period of leptocephalus transformation, a more lucid view of Myrichthys taxonomy will be achieved. I have examined the holotypes of M. magni- 64 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 2 TABLE 1. Myrichthys maculosus (CUVIER): VERTEBRAL DATA FOR EASTERN PACIFIC AND INDO-PACIFIC POPULA- TIONS. All counts made from radiographs of adults. mean range n Eastern Pacific’ 153.1 149-157 19 Revillagigedos and Clipperton Is. 163.1 161-168 10 Hawaii, Johnston, and Midway 178.5 174-182 16 Western Pacific? 192.8 190-199 8 Red Sea 194.5 194-195 2 ' Specimens from the Gulf of California, Tres Manias Is- lands, Cocos Island, Panama, and the Galapagos Islands. * Specimens from Wake, Palau, Philippines, RyuKyus, and the Line Islands. ficus and M. stypurus and found them to be con- specific. The holotype of M. stypurus is aberrant in that its tail had been severed and subsequent- ly healed. Myrichthys bleekeri Gosline Ophisurus fasciatus var. semicinctus Bleeker, 1864:64 [a homonym of Ophisurus semicinctus Lay and Bennett, 1839:66] (type-locality, Indonesia). Myrichthys bleekeri Gosline, 1951:314 [a substitute name for Ophisurus fasciatus var. semicinctus Bleeker, 1864, preoc- cupied]. REMARKS.—This species 1s known from John- ston Island and the Indo-West-Pacific, but not from Hawaii. Cirrhimuraena playfairii (Gunther) Ophichthys playfairii Giinther, 1870:76 (type-locality, Zanzi- bar). Microdonophis macgregori Jenkins, 1903:422 (type-locality, Lahaina, Maui). Jenkinsiella macgregori: Jordan and Evermann, 1905:82. Cirrhimuraena playfairii: Barnard, 1925:205. Cirrhimuraena macgregori: Gosline, 1951:315. REMARKS.—Smith (1962) synonymized the Hawaiian species Microdonophis macgregori with the wide-ranging Indo-Pacific species Cir- rhimuraena playfairii. He based this action on his comparison of four specimens from Aldabra with published descriptions by Gosline (1951) of Hawaiian specimens and by Weed and Howarth (1961) of specimens from Ceylon. I have ex- amined the type-specimens from Hawaii and compared them with material from Midway (SIO 68-497) and the Marquesas (BPBM 12304), and concur with Smith. The holotype of M. mac- gregori (USNM 50721) has 180 vertebrae; two Marquesan specimens have 186 and 187 verte- brae. Brachysomophis sauropsis Schultz Brachysomophis sauropsis Schultz, 1943:18 (type-locality, Samoa). REMARKS.—Not known from Hawaii. I have been unable to examine Gosline’s (1955:443) specimen of B. sauropsis from Johnston Island. However, based on his description, I would agree that B. sauropsis and B. henshawi are dis- tinct species. Brachysomophis henshawi Jordan and Snyder Brachysomophis henshawi Jordan and Snyder, 1904:940 (type-locality, Honolulu). REMARKS.—Apparently, this species is a Hawaiian endemic. A comprehensive revision of Brachysomophis is necessary, however, be- fore assumptions concerning its species can be made. Phyllophichthus xenodontus Gosline Phyllophichthus xenodontus Gosline, 1951:316 (type-locality, Oahu). REMARKS.—Phyllophichthus is currently rec- ognized to contain a single wide-ranging species, known from the western Indian Ocean, through- out Oceania, and Hawaii (McCosker and Allen, MS). Four specimens from Johnston and Oahu islands had 168-175 vertebrae (¥ = 170.8). Ophichthus polyophthalmus Bleeker (Figure 5) Ophichthys polyophthalmus Bleeker, 1864:43 (type-locality, Ambon). Microdonophis polyophthalmus: Jordan and Snyder 1907:207. Ophichthus polyophthalmus: Fowler 1927:5. REMARKS.—The markedly spotted Hawaiian eels placed by recent authors in either Micro- donophis or Ophichthus is a composite. Oph- ichthus polyophthalmus (Fig. 5), a medium- length adult ophichthid with ocellated spots on the head and body, was reported by Fowler (1927) on the basis of a Kahoolawe (BPBM 3700) dredge specimen. Fowler (1928) subsequently recorded eleven Honolulu market specimens (the largest, 1,039 mm) which possessed dark spots; he considered them to also be O. poly- ophthalmus and recognized the nominal species Microdonophis fowleri Jordan and Evermann McCOSKER: HAWAIIAN SNAKE EELS 65 FIGURE 5. and Ophichthus garretti Gunther to be syn- onyms which differed only in coloration. Oph- ichthus garretti is a valid and distinctly different species. My comparison of numerous specimens of the large form with solid dark spots (Ophich- thus erabo, Fig. 6) with the medium-length eel with ocellated spots indicated that they are in fact separate species differing in coloration and vertebral number. Too few specimens of O. polyophthalmus were available for a proper morphological comparison, although O. erabo appears to possess a proportionately longer tail. I herein report a second Hawaiian specimen of O. polyophthalmus, collected in 1968 by hook and line from Nanakuli, Oahu (BPBM 11981, 399 mm SL). The specimen has 148 vertebrae, 75 before the anal opening. Ophichthus erabo (Jordan and Snyder) (Figure 6) Microdonophis erabo Jordan and Snyder, 1901:780 (type-lo- cality, Misaki, Japan). Microdonophis fowleri Jordan and Evermann, 1904: 164 (type- locality, Honolulu). Ophichthus erabo: McCosker 1977:81. REMARKS.—The majority of Hawaiian rec- ords of O. polyophthalmus are based on speci- mens of O. erabo. I have compared the Japa- nese holotype (SU 6477) and cotypes (SU 6667 Adult specimen of Ophichthus polyophthalmus, taken from Bleeker (1864). and 6744) of O. erabo with specimens from Ha- wail (SU 8407; SU 8466: BPBM 12510; and USNM 50613, the holotype of M. fowleri) and Taiwan (CAS 15600), and found them not to dif- fer. The vertebral range of six specimens was 152-155 (@ = 154). The holotype of O. erabo has 155 vertebrae and the holotype of M. fowleri has 152. DISCUSSION The endemism of the Hawaiian shorefish fau- na has been recognized by numerous authors and most recently summarized by Randall (1976). He calculated that 29 percent of the 442 reef and shorefish species were endemic at the specific or subspecific level. He elucidated the situation of the problematical species such as Acanthurus triostegus, wherein an identifiable difference in coloration exists between the Hawaiian and extralimital populations, but a consensus of opinion concerning the biological significance of that difference has not been reached. A similar problem exists with the Hawaiian ophichthids. Of the 15 ophichthids present at Hawaii, five species are endemic. The new species described herein, Ophichthus kunaloa and Muraenichthys puhioilo, are known only from a single deep- water collection and are therefore of little value FIGURE 6. Adult specimen of Ophichthus erabo, taken from Jordan and Snyder (1901). 66 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 2 TABLE 2. DISTRIBUTION OF HAWAIIAN AND JOHNSTON ISLAND OPHICHTHIDS. Schultzidia johnstonensis Schismorhynchus labialis Muraenichthys schultzei Muraenichthys cookei Muraenichthys gymnotus Muraenichthys puhioilo Ichthyapus vulturis Apterichtus flavicaudus Callechelys luteus Myrichthys maculosus Myrichthys bleekeri Cirrhimuraena playfairii Phyllophichthus xenodontus Phaenomonas cooperae Leiuranus semicinctus Brachysomophis sauropsis Brachysomophis henshawi Ophichthus erabo Ophichthus polyophthalmus Ophichthus kunaloa Indo-West Hawaiian Leewards Johnston Pacific Xx x X x x x X X X X x x X X x X X x Xx x Xx x x x Xx x rK ms Xx x Xx X x Xx X Xx Xx Xx mR X x Xx X x X Xx Xx to a zoogeographic analysis. The endemics, Mu- raenichthys cookei and Brachysomophis hen- shawi, are perhaps no more different than are several of the other ophichthids discussed here from their Indo-West-Pacific “‘conspecifics.”’ Callechelys luteus is the only Hawaiian ophichthid endemic distinctly different at the specific level from all known congeners. It is most closely related to C. galapagensis, another insular endemic (McCosker and Rosenblatt 1972). In my analysis of Hawaiian ophichthids, I have been able to recognize the Hawaiian pop- ulations of several species (viz., Myrichthys maculosus, Phaenomonas cooperae, Phylloph- ichthus xenodontus, Ichthyapus vulturis, and probably several others) to be distinct from oth- er Indo-Pacific populations of their conspecifics on the basis of vertebral differences. Yet I am hesitant to assign them specific rank. It should be recognized, however, that the absolute dif- ferences in ophichthid vertebral numbers appear to be less when calculated on a percentage basis. For example, the difference of eight vertebrae between the Hawaiian specimen of Phaenomon- as cooperae and the Gilbert Island holotype is only three percent, which is less than a differ- ence of one vertebra for most perciform fishes. As Randall (1976:49-S0) has discussed, these differences are real and apparently indicate lim- ited gene flow with other insular populations. Whether the Hawaiian forms are in fact distinct biological species, incipient species, or what- ever taxon a systematist deems them, awaits the discovery of biological data concerning lepto- cephalus transport, longevity, and gene flow. Untaxing the taxonomy of the Hawaiian ophichthids, initiated by Maui the Wonder Boy and continued by Jordan, Evermann, and Gos- line, remains a challenge. ACKNOWLEDGMENTS This study depended primarily upon recent specimens collected by J. E. Randall (BPBM), the George Vanderbilt Foundation (CAS), and the 1968 STYX Expedition of the Scripps Insti- tution of Oceanography. Other specimens and assistance were provided by the following indi- viduals: J. Paxton and D. Hoese (AMS); J. Bohlke (ANSP); A. Wheeler (BMNH): W. Esch- meyer (CAS); R. Lavenberg (LACM); R. Ro- senblatt and J. Copp (SIO); V. Springer and N. Gramblin (USNM); and L. Taylor of the Wai- kiki Aquarium, Hawaii. J. Gordon and C. Ruark assisted with the preparation of radiographs, M. Giles prepared the photographs, and K. Smith prepared Figures 1-4. I am very grateful to all of the above-mentioned individuals for their as- sistance, and in particular, to Thomas A. Clarke for providing the new species and to William A. Gosline for his pioneering studies of apodal fish- es. McCOSKER: HAWAITAN SNAKE EELS LITERATURE CITED ApsoTt, C. C. 1861. Description of new species of apodal fishes in the museum of the Academy of Natural Sciences of Philadelphia. Proc. Acad. Nat. Sci. Phila. 12:475-479. BARNARD, K. H. 1925. A monograph of the marine fishes of South Africa. Part I. Ann. S. Afr. Mus. 21:1-418. BLEEKER, P. 1864. Atlas ichthyologique des Indes Orientales Neerlandaises. Vol. 4. Amsterdam. 132 pp. BOHLKE, J. E. 1968. A new species of the ophichthid eel genus Verma from the West Atlantic, with comments on related species. Not. Nat. (Phila.), no. 415. 12 pp. , AND J. E. McCosker. 1975. The status of the ophichthid eel genera Caecula Vahl and Sphagebranchus Bloch, and the description of a new genus and species from fresh waters in Brazil. Proc. Acad. Nat. Sci. Phila. 127(1): 1-11. CLARKE, T. A. 1972. Collections and submarine observations of deep benthic fishes and decapod crustacea in Hawaii. Pac. Sci. 26(3):310-317. Cou, P. 1937. Legends of Hawaii. Yale Univ. Press, New Haven. 220 pp. Cuvier, G. 1817. Le regne animal. Vol. 2. Paris. 532 pp. Fow er, H. W. 1927. Fishes of the tropical central Pacific. Bernice P. Bishop Mus., Bull. 38. 32 pp. . 1928. The fishes of Oceania. Bernice P. Bishop Mus., Mem. 10. 540 pp. GosLINE, W. A. 1951. The osteology and classification of the ophichthid eels of the Hawaiian Islands. Pac. Sci. 5(4):298- 320. 1955. The inshore fish fauna of Johnston Island, a Central Pacific atoll. Pac. Sci. 9(4):442—-480. , AND V. E. Brock. 1960. Handbook of Hawaiian fish- es. Univ. Hawaii Press, Honolulu. 372 pp. GUNTHER, A. C. 1870. Catalogue of the fishes in the British Museum. Vol. 8. London. 549 pp. JENKINS, O. P. 1904. Report on collections of fishes made in the Hawaiian Islands, with descriptions of new species. U.S. Bur. Fish. Bull. 22:417-511. JORDAN, D. S., AND B. W. EVERMANN. 1905. The aquatic resources of the Hawaiian Islands. Part I. The shore fishes. U.S. Fish. Comm. Bull. 23(1):1-574. , AND J. O. SNYDER. 1901. A review of the apodal fishes or eels of Japan, with descriptions of 19 new species. Proc. U.S. Natl. Mus. 23(1239):837-890. . 1904. Notes on collections of fishes from Oahu Island and Laysan Island, with descriptions of four new species. Proc. U.S. Natl. Mus. 27:939-948. . 1907. Notes on fishes of Hawaii, with descriptions of new species. U.S. Bur. Fish. Bull. 26:207-218. Lay, G. T., AND E. T. BENNETT. 1839. The zoology of Cap- tain Beechey’s voyage. Fishes. London. Pp. 41-75. McCosker, J. E. 1970. A review of the eel genera Lep- tenchelys and Muraenichthys, with the description of a new genus, Schismorhynchus, and a new species, Muraen- ichthys chilensis. Pac. Sci. 24(4):506-516. 1975. The eel genus Phaenomonas (Pisces, Ophichthidae). Pac. Sci. 29(4):361-363. . 1977. The osteology, classification, and relationships of the eel family Ophichthidae. Proc. Calif. Acad. Sci., Ser. 4, 41(1):1-123. , AND R. H. ROSENBLATT. 1972. Eastern Pacific snake-eels of the genus Callechelys (Apodes: Ophichthi- dae). Trans. San Diego Soc. Nat. Hist. 17(2):15-24. , AND . 1975. The moray eels (Pisces: Muraen- idae) of the Galapagos Islands, with new records and syn- onymies of extralimital species. Proc. Calif. Acad. Sci., Ser. 4, 40(13):417-427. PALMER, G. 1970. New records, and one new species, of teleost fishes from the Gilbert Islands. Bull. Brit. Mus. (Nat. Hist.) Zool. 19(6):211-234. Pukul, M. K. 1902. Ka hana kuhikuhi no ka lawai’a ana. Ka Nupepa Kuokoa, newspaper article, 23 May. (Not seen; reference from: Tircoms, M. 1972. Native use of fish in Hawaii. Univ. Hawaii Press, Honolulu. 175 pp.) RANDALL, J. E. 1976. The endemic shore fishes of the Hawaiian Islands, Lord Howe Island and Easter Island. Colloque Commerson 1973, ORSTOM Trav. et Doc. no. 47:49-73. , AND J. E. McCosKer. 1975. The eels of Easter Island with a description of a new moray. Los Angeles County Mus. Contrib. Sci., no. 264. 32 pp. ROSENBLATT, R. H., J. E. MCCoskeR, AND I. RUBINOFF. 1972. Indo-West-Pacific fishes from the Gulf of Chiriqui, Panama. Los Angeles County Mus. Contrib. Sci., no. 234. 18 pp. SCHULTZ, L. P. 1943. Fishes of the Phoenix and Samoan islands. U.S. Natl. Mus. Bull. 180. 316 pp. 1953. Fishes of the Marshall and Marianas islands. Families from Asymmetrontidae through Siganidae. U.S. Natl. Mus. Bull. 202, 1. 685 pp. (With collaborators.) SEALE, A. 1917. New species of apodal fishes. Bull. Mus. Comp. Zool. Cambridge 61:79-94. SMITH, J. L. B. 1962. Sand-dwelling eels of the western In- dian Ocean and the Red Sea. Rhodes Univ. Ichthyol. Bull. 24:447_466. SMITH, R., AND J. SWAIN. 1882. Notes on a collection of fishes from Johnston’s Island including descriptions of five new species. Proc. U.S. Natl. Mus. 5:119-143. SNYDER, J. O. 1904. A catalogue of the shore fishes collected by the steamer “‘Albatross”’ about the Hawaiian Islands in 1902. U.S. Bur. Fish. Bull. 22:513-538. WEBER, M., AND L. F. DE BEAUFORT. 1916. The fishes of the Indo-Australian Archipelago. Vol. 3. Leiden. 455 pp. WEED, W. A. III, AND J. N. HowARTH. 1961. Range exten- sions of the ophichthid eels Phyllophichthus xenodontus and Cirrhimuraena macgregori. Copeia 1961(3):357. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIEN Vol. 42, No. 3, pp. 69-85; 7 figs. Marine Bistogical Laberator) LIBRARY ‘JAN 8 1960 Woods Hole, Mass. December 22, 1979 REVIEW OF THE RHINOTRAGINI OF MEXICO (COLEOPTERA: CERAMBYCIDAE) John A. Chemsak and E. G. Linsley University of California, Berkeley, CA 94720 ABSTRACT: The Rhinotragini known to occur in Mexico are reviewed. Keys are provided for the determination of the seven genera and also for species in the genera. Distribution maps are included for the more common species and most of the new taxa are illustrated. New species include: Tomopterus exilis, Acyphoderes amoena, A. fulgida, A. prolixa, and A. parva. New synonymies and one new combination are proposed. INTRODUCTION The Rhinotragini comprise a large group of Neotropical Cerambycidae which is particularly abundant and diverse in South America. The group is of special interest because of the re- markable degree of mimicry exhibited by most species. This is evident in modifications in form, coloration, and behavior, with the various taxa utilizing a wide range of models, particularly bees, wasps, and “‘protected”’ beetles of other families. Frequently, the mimicry is expressed in strong sexual dimorphism and dichromatism ‘with males and females utilizing different models. The number of species diminish as the group extends northward from South America, and only one is thus far known to occur in the United States. As presently defined, the Mexican fauna consists of 22 species in 7 genera. Some of the species appear to be restricted to Mexico, while others range into Central America to Costa Rica or Panama. One is also found in South America. Tribe RHINOTRAGINI Lacordaire Rhinotragides LACORDAIRE, 1869:497. Rhinotraginae BATES, 1873:21. Rhinotragini LINSLEY, 1963:164. [69] Body more or less elongate. Head prolonged below eyes to form a muzzle; antennae usually shorter than body in both sexes, outer segments more or less serrated; eyes finely faceted, with lower lobes very large, nearly contiguous in front in males; palpi short, last segment subcy- lindrical or ovate-cylindrical, apex truncate; mandibles short. Pronotum cylindrical or round- ed, without a lateral spine or tubercle; proster- num with intercoxal process narrow, level, an- terior coxae usually obliquely exserted, cavities feebly angulate externally; intermediate coxal cavities open externally; episterna of metatho- rax very large, triangular, and in front broad and narrowly separated from coxae. Elytra nearly always modified, attenuated, subulate, or squamiform. KEY TO THE MEXICAN GENERA OF RHINOTRAGINI Ie Elytra short, not extended over abdo- TC Tage ee Beene = Bee aries oe Sh 2 Elytra longer, extended at least partially OVE adOMeN 9 4 2(1). Elytra narrowing posteriorly, apices de- hiscent and narrowly rounded ___-__ 3 70 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 3 Elytra not narrowing posteriorly, apices obliquely truncate _-- Tomopterus Pronotum with disk punctate, calluses, if present, small, rounded; posterior tibiae with very dense brush of long pubescence Epimelitta Pronotum with three longitudinal, gla- brous calluses on disk; posterior tibi- ae with apical brush small, the pubes- cence short Bromiades Pronotum uniformly punctate or with rounded calluses on disk; femora 3(2). 4(1). moderately clavate = ee ae 6 Pronotum usually with three distinct, longitudinal calluses on disk; anterior and intermediate femora often very Strongly clavate = 5 Antennae short, serrate apically; ante- rior and intermediate femora very Strongly clavate .....-... _- Acyphoderes Antennae slender, elongate, filiform api- cally; anterior and intermediate fem- ora narrowly clavate ___ Stenopseutes Elytra with disk vitreous transparent; pronotum about as long as broad or slightly longer == == Odontocera Elytra with disk shining, not transpar- ent; pronotum narrow, distinctly longer than broad Ommata 5(4). 6(4). Genus Tomopterus Audinet-Serville Tomopterus AUDINET-SERVILLE, 1833:544; NEWMAN 1840:21; WHITE 1855:176; THOMSON 1860:166, 168; 1864:163, 416; LACORDAIRE 1869:509; BATES 1870:329; 1880:44. This genus may be recognized by the short, rather squat form, short, subserrate antennae, rounded pronotum, short, apically truncate ely- tra, and arcuate posterior femora. TYPE-SPECIES: Tomopterus staphylinus Au- dinet-Serville, 1833 (monotypic). The species of Tomopterus appear to mimic eumenine vespids. Two species occur in Mexi- co. Tomopterus vespoides White Tomopterus vespoides WHITE, 1855:176, pl. 5, fig. 8; BATES 1880:44; 1885:291; FisHER 1930:17. MALE: Integument black, antennae reddish distally, apices of hind femora and tibiae pale, elytra usually narrowly reddish along lateral margins and with a reddish median stripe ex- tending from humeri arcuately to middle of apex. Head with eyes contiguous on front; an- tennae with segments from fifth expanded, sub- serrate. Pronotum almost as broad as elytra: disk coarsely, contiguously punctate; apex and base with bands of yellow, appressed pubes- cence. Scutellum elongate, triangular, nonpu- bescent. Elytra moderately coarsely, densely punctate, sparsely pubescent. Abdomen with sternites margined with yellowish, appressed pubescence; parameres prominent. Length, 8- 10 mm. FEMALE: Form more robust. Head with eyes separated on front by about width of antennal scape. Scutellum yellow-pubescent. Elytra black medially, remainder reddish. Abdomen bulbous, pointed apically. Length, 8-10 mm. TYPE-LOCALITY: Guatemala. RANGE: Tamaulipas, Mexico to Panama. FLOWER RECORDS (Costa Rica): Bixa, For- steronia, Byrsonima, Paullinia, Cordia. New REeEcorps: | female, 10 miles [16 km] N Cuidad Vic- toria, Tamaulipas, 20 Aug. 1941 (H. S. Dybas); 1 female, 9 miles [ca. 14 km] S Tuxpan, Veracruz, 11 June 1961 (Mich- ener and Ordway); 1 female, Tehuantepec, Oaxaca, 8 July 1962. Tomopterus exilis, new species (Figure 1) FEMALE: Form small, rather slender; integu- ment black, antennae brownish; elytra narrowly pale at base, each side with a broad, oblique, pale vitta extending from behind humerus to apex; anterior and intermediate legs brownish, posterior femora pale over basal one-half; pu- bescence silvery, appressed. Head with front rather short, moderately coarsely, densely punctate; eyes separated on front by little more than diameter of antennal scape; separated on vertex by about twice diameter of scape; anten- nae clavate, enlarging from fifth segment, outer segments broader than long, with small apical poriferous areas, third segment longer than first, fourth shorter than first, fifth longer than fourth, basal segments with a few, long, erect setae be- neath. Pronotum as long as broad, cylindrical; disk coarsely, deeply reticulate-punctate; apex narrowly impressed, base rather broadly im- pressed; apex and base with narrow transverse silvery bands of appressed pubescence, each side with a narrower band behind middle ex- tending from sides to coxal cavities; long, pale, erect hairs numerous; prosternum densely punc- tate, sparsely clothed with long, erect hairs: CHEMSAK & LINSLEY: RHINOTRAGINI OF MEXICO a mesosternum with a broad pubescent band over epimeron; metasternum sparsely punctate, with a long erect seta rising from each puncture, a narrow, oblique, pubescent band present pos- teriorly. Scutellum densely silver-pubescent. Elytra broader than long, densely, deeply, con- tiguously punctate; pubescence long, erect; pale vittae in form of a broad V; apices obliquely truncate. Legs slender; femora enlarged over apical one-half, hind pair arcuate, extending al- most to apex of abdomen; tibiae slender. Ab- domen only slightly enlarged posteriorly; punc- tures and pubescence fine, sternites more densely pubescent along posterior margins; last sternite rounded at apex. Length, 8 mm. Holotype, female (National Museum of Natural History) from Cancun, Quintana Roo, Mexico, 25 Apr. 1974 (D. Pletsch). The more slender form, cylindrical rather than transverse pronotum, and longer antennae will separate this species from T. vespoides White. Genus Epimelitta Bates Charis NEWMAN, 1840:21; THOMSON 1864:163; LACORDAIRE 1869:507 (name preoccupied). Type-species: Charis eu- phrosyne Newman, 1840, Thomson designation, 1864). Epimelitta BATES, 1870:330; 1873:123. Charisia CHAMPION, 1892:161 (new name for Charis New- man, 1840, preoccupied). The Mexican species in this genus may be rec- ognized by the short elytra, lack of longitudinal calluses on the pronotum, and densely tufted posterior tibiae. TYPE-SPECIES: Epimelitta meliponica Bates, 1870 (by present designation). Most species of Epimelitta mimic meliponid bees. The group is dominantly South American with only two species presently known from Mexico. Epimelitta nigerrima (Bates) Charisia nigerrima BATES, 1892:160, pl.6, fig. 8. Epimelitta nigerrima,; AURIVILLIUS 1912:284. FEMALE: Integument black, antennae reddish brown. Antennae short, serrate from fifth seg- ment. Pronotum densely clothed with long erect pubescence: disk callused behind middle. Elytra strongly dehiscent, densely punctate. Legs short, femora and tibiae with dense tufts of dark pubescence. Length, 11 mm. TYPE-LOCALITY: Atoyac, Veracruz, Mexico. Only the holotype of this species is known. 71 FiGure 1. Dorsal view of pronotum and body of Tomop- terus exilis Chemsak and Linsley, female. Epimelitta aureopilis Fisher Epimelitta (?) aureopilis FISHER, 1953:16. MALE: Form slender, elongate; head, pro- thorax, part of metasternum and often bases of hind femora black, antennae reddish brown, elytra testaceous except for narrow black lateral and sutural margins, legs and abdomen partially reddish, posterior tarsi yellow. Head small, front golden pubescent; antennae slender, mod- erately serrate. Pronotum densely, coarsely punctate; apex and base with a broad band of appressed golden pubescence, basal band reach- ing to about middle at center. Elytra dehiscent, sparsely pubescent. Legs with posterior pair very elongate, femora feebly clavate, tibiae with 72 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 3 a dense tuft of long reddish-orange pubescence. Abdomen elongate, slender, last sternite barely impressed. Length, 16-21 mm. FEMALE: Form and coloration similar. Ab- domen with last sternite narrowly rounded at apex. Length, 18-23 mm. TYPE-LOCALITY: Tolome, Veracruz, Mexico. This species differs greatly in facies from oth- er known Epimelitta. Its long, slender form with very long posterior legs, make it especially dis- tinctive. However, except for these character- istics, aureopilis is structurally similar to other species within the genus. Fisher (1953) tentatively assigned aureopilis to Epimelitta because of the difficulty in delim- iting genera of Rhinotragini. We have retained it in this genus since it is beyond the intent of this paper to attempt to clarify the generic clas- sification of this primarily South American group. New Recorps: 3 females, 3 males, Cotaxtla Exp. Sta., Cotaxtla, Veracruz, 1 Aug. 1962 (D. H. Janzen); | male, Ixmal, Yucatan (G. F. Gaumer); | female, 30 miles [48 km] NE Tehuantepec, Oaxaca, 8 July 1955 (D. Giuliani). Genus Bromiades Thomson Bromiades THOMSON, 1864:165; LACORDAIRE 1869:506: BaTEs 1873:120; Zayas 1975:130. The tricallused pronotum, strongly serrate an- tennae, and short, tapered elytra distinguish this genus. TYPE-SPECIES: Odontocera brachyptera Chevrolat, 1838 (by original designation). Bromiades brachyptera (Chevrolat) Odontocera brachyptera CHEVROLAT, 1838:285; JACQUELIN DUVAL in SAGRA 1857:269, pl. 10, fig. 9. Bromiades brachyptera; THOMSON 1864:165; BATES 1873:120; FISHER 1930:13; LINsLEY 1935:85, pl. 2, fig. 3; ZAYAS 19563115 S975 iSilesplasldautigera: Bromiades meridionalis FisHER, 1930:14 (type-locality, Ca- bima, Panama). NEw SYNONYMY. MALE: Integument black, shining, antennal segments four to usually eight, yellowish basal- ly; elytra broadly testaceous medially; femora usually with basal one-half testaceous; anterior and intermediate tibiae basally and posterior pair except apically, testaceous. Pronotum densely clothed with golden, recumbent pubes- cence between calluses. Abdomen not modified apically. Length, 15-17 mm. FEMALE: Form and coloration similar to male, abdomen broader. Antennae occasionally al- most all black. Length, 16-18 mm. TYPE-LOCALITY: Cuba. RANGE: Central Mexico to Colombia and Cuba. Host PLANTS: Andira inermis (W. Wright) H-B-Ke ex D:€; (Cuba) FLOWER RECORDs. Costa Rica: Cordia, Ca- searia, Forsteronia, Coccoloba, asclepiad vine. Mexico: Spondias. Color variation is apparent in the antennae and legs. Specimens from Panama have domi- nantly black antennae and orange posterior fem- ora and tibial brushes. This type of color vari- ation occurs in numerous species throughout the tribe. We have seen no additional specimens from Mexico since those reported by Linsley (1935) from Bejucos. Genus Acyphoderes Audinet-Serville Acyphoderes AUDINET-SERVILLE, 1833:549; WHITE 1855:194; THOMSON 1860:179; 1864:165; LACORDAIRE 1869:505; BaTEs 1873:117; 1880:43. This genus may be recognized by the dorsal callosities of the pronotum. The antennae are distally serrate and usually extend to about the middle of the elytra. The elytra are subulate or elongate, narrowed posteriorly and extend to the abdomen. The anterior and intermediate femora are usually strongly clavate. Males of most species have the last abdominal sternite modi- fied in varying degrees. TYPE-SPECIES: Acyphoderes aurulenta Kirby, 1818 (Thomson designation, 1864). Acyphoderes, with ten species, is the largest group of Rhinotragini found in Mexico. Two other species, velutinus Bates, 1885, and ves- piventris Bates, 1880, described originally from Guatemala, probably also occur in Mexico. Key to the Mexican species of Acyphoderes l. Pronotum black, with discal callosities shining, punctures sparse or absent _ callosities opaque, densely punctate tn 2(1). Pronotum with transverse or longitudi- nal bands of dense, appressed pubes- Ceénce.... 5... 2 eee 3 CHEMSAK & LINSLEY: RHINOTRAGINI OF MEXICO without basal and apical transverse PUDESCENE DanGs <.-8 =. =-f et 4 3(2). Pronotum densely pubescent along api- cal and basal margins, dorsal calluses usually punctate, lateral pair rounded, punctures very dense, confluent. Length, 13-20 mm. Sinaloa to Oaxaca And VELACKUZ 8 81S. ae cribricollis Pronotum densely pubescent except on calluses, calluses shining, elongate, not punctate, punctures obscured. Length, 14-18 mm. Sinaloa to Chia- (OGG). St a ee | ae amoena Elytra subulate, extending only to an- terior margin of second abdominal segment; abdomen strongly inflated toward apex. Length, 17-21 mm. Nayarit to Oaxaca and Veracruz ______ __. 5 ee, a oe sexualis Elytra more gradually tapering, extend- ing over first three abdominal seg- ments; abdomen narrowed toward apex. Length, 14-18 mm. Veracruz to Msiar Rica ee Fy fulgida Elytra with apices rounded, as broad as or broader than diameter of antennal 4(2). SCh)E Elytra with apices truncate to emargin- ate, narrower than diameter of anten- HAIRS CAD Cee See eee gee eS 28 7. Pronotum deeply impressed at base and apex, with bands of dense appressed pubescence in impressions; elytra black around scutellum. Length, 20- PSMmMioiC OUMA ys 2 ee prolixa Pronotum not deeply and broadly im- pressed at base and apex, without transverse bands of dense pubes- cence; elytra pale around scutellum. Length, 11-18 mm. Baja California _ oe ee delicata Pronotum with apical and basal trans- verse bands of dense, appressed pu- bescence, dorsal calluses moderately el SEC sk © ee ree 8 Pronotum rather uniformly pubescent, without transverse bands, dorsal cal- luses prominently elevated. Length, 15-20 mm. Colima to Oaxaca and Ve- Tel CHUN ee SE a acutipennis Elytra 2.5 or more times longer than basal width, extending at least to third abdominal segment 6(5). 7G): 8(7). Elytra short, 2.0 times as long as basal width, extending only to second ab- dominal segment. Length, 10-18 mm. Campeche to Honduras ____ yucateca Legs with hind femora pedunculate; ab- domen abruptly inflated toward apex; elytra with two oblique, usually brownish, scabrous bands at base. Length, 11-16 mm. Sinaloa to Nay- LITT prereset ee NE SS parva Legs with hind femora gradually en- larged; abdomen gradually expanded toward apex; elytra punctate at base, with a triangular yellowish area around scutellum. Length, 13-22 mm. Sonora and Tamaulipas to Costa Rica Beet 2 OA PR eget ine begs SUAVIS 9(8). Acyphoderes cribricollis Bates Acyphoderes cribricollis BATES, 1892:160, pl. 6, fig. 6; LiNs- LEY 1935:84; CHEMSAK 1967:74 (lectotype). MALE: Form slender, elongate; integument black: antennae reddish brown, usually infus- cated apically; elytra testaceous, shining, mar- gins dark, legs pale, hind pair with femoral club and apices of tibiae brownish; abdomen with first two segments yellowish, usually infuscated apically, other segments usually reddish. Head with front densely pubescent, eyes separated by less than diameter of antennal scape. Pronotum longer than broad; apex and base broadly im- pressed: disk with median callus punctate, lat- eral pair more elevated anteriorly, glabrous, rounded; punctures dense, opaque; apical and basal margins with broad bands of dense, ap- pressed, golden pubescence, bands joined lat- erally and usually extended longitudinally on disk on each side of middle; pro- and mesoster- num pubescent; anterior one-half of metaster- num and posterior end of episternum densely pubescent. Elytra extending to posterior margin of third abdominal segment; base and disk fine- ly, sparsely punctate, margins densely punctate; apices rounded, about as broad as antennal scape. Legs with femora clavate, anterior and intermediate pairs moderately clavate. Abdo- men slender, slightly enlarged toward apex: last sternite deeply, broadly excavated, margins prominent, elevated apically. Length, 14-20 mm. FEMALE: Form similar. Head with widely separated on front. Abdomen with last sternite not modified. Length, 13-20 mm. eyes 74 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 1000 FOOT CONTOUR BONNE’S EQUAL-AREA PROJECTION FIGURE 2. TYPE-LOCALITY: Ventanas, Durango. FLOWER RECORDS: Croton, Xanthoxylum, Jatropha. The dense pubescence, rounded, glabrous cal- luses, and dense, opaque punctation of the pronotum distinguish this species. New Recorps: See Figure 2. Acyphoderes amoena, new species (Figure 3) MALE: Form moderate sized; integument black; antennae reddish brown; elytra shining, yellow-brown, margins black, base often with two short dark vittae extending back from inside of humeri; legs orange-brown, posterior femora often dark over clavate portion; abdomen usu- ally pale and dark. Head with eyes separated on front by less than diameter of antennal scape; front and vertex densely clothed with appressed golden pubescence. Pronotum longer than broad, sides impressed before and behind mid- dle; disk uneven, dorsal! calluses glabrous, ele- vated, sides with smaller calluses before middle, pubescence between calluses dense, appressed, golden, obscuring surface, long, erect hairs rath- Ww Known distribution of Acyphoderes cribricollis Bates. er sparse; prosternum densely pubescent; meso- sternum densely clothed with appressed golden pubescence; metasternum densely pubescent except for rectangular areas near base, epister- num with dense pubescent patches posteriorly and anteriorly on upper margin. Elytra elongate, extending to fourth abdominal segment; lateral margins darkened, narrowly punctate, sutural margins elevated, narrowly dark; disk shining, very sparsely punctate except on base; apices rounded, broad. Legs with anterior and inter- mediate femora moderately clavate; posterior femora gradually clavate; posterior tibiae dense- ly pubescent over apical one-half. Abdomen not apically enlarged; sternites with small patches of dense pubescence at sides of apical margins; last sternite broadly excavated, margins elevat- ed toward apex, angles blunt. Length, 14-18 mm. FEMALE: Form similar. Head with eyes widely separated on front. Abdomen with last sternite broadly rounded at apex. Length 16-18 mm. Holotype, male (California Academy of Sciences) from 3 miles [ca. 5 km] E Villa Union, Sinaloa, Mexico, 24 July 1972, CHEMSAK & LINSLEY: RHINOTRAGINI OF MEXICO FiGuRE 3. Acyphoderes amoena Chemsak and Linsley, male. on Jatropha curcas flowers (J. A. and M. A. Chemsak, A. E. and M. M. Michelbacher). Paratypes as follow: 1 male, same data as type: 1 female, 16 miles [ca. 26 km] NE San Blas, Nayarit, 21 July 1963 (R. Westcott); | male, 14 miles [ca. 23 km] NW Tehuantepec, Oaxaca, 26 June 1961, on flowers of Croton (Univ. Kans. Mex. Exped.); 1 male, 23 miles [ca. 37 km] S Matias Romero, Oaxaca, 14 Aug. 1963 (Parker and Stange); 1 female, Playa La Ventosa, 3.5 miles [ca. 5.6 km} 76 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 3 NE Salina Cruz, Oaxaca, 28 July 1970 (Fisher, Sullivan); 1 female, 5 miles [ca. 8 km] NW Escuinapa, Sinaloa, 25 July 1971 (Fisher). We are also assigning to this species five spec- imens from various localities in Mexico which differ in having the antennae black, with seg- ments 8-11 all or partially yellow. In most of these, the two basal black vittae of the elytra are very distinct and the pubescence is more silvery than golden. Structurally they are iden- tical with the typical forms. Records are: 1 female, Tecolopa, Colima, 31 July 1954 (Ca- zier, Gertsch, Bradts); 1 female, 10 miles [ca. 16 km] W Col- ima, | Aug. 1954 (Cazier, Gertsch, Bradts); 1 male, Manzan- illo, Colima, 12 July 1956 (R. and K. Dreisbach); 1 male, Acapulco, Guerrero, 30 July 1933 (M. A. Embury); | male, 31 miles [ca. 50 km] SE Comitan, Chiapas, 18 June 1965 (Burke, Meyer, Schaffner). Acyphoderes sexualis Linsley Acyphoderes sexualis LINSLEY, 1934:349; 1935:84, pl. 2, figs. [Pela Ma_Le: Form rather elongate; integument black, antennae with basal segments infuscated beneath, reddish above, outer segments pale an- nulate basally; elytra with disk pale vitreous; abdomen with first segment reddish at middle; posterior legs with femora narrowly pale basal- ly, tibiae yellowish at basal one-half. Head with eyes separated on front by about diameter of third antennal segment. Pronotum slightly broader than long, sides impressed behind mid- dle, base narrower than apex; disk with three longitudinal, sparsely punctate, shining calluses; punctures between calluses fine, dense; pubes- cence fine, moderately dense, appressed, long erect hairs numerous; mesosternum with epi- meron densely pubescent; metasternum with a broad, densely pubescent band anteriorly and a narrow band along posterior margin. Elytra sub- ulate, extending to anterior portion of second abdominal segment; lateral margins broadly black and densely punctate, sutural margins nar- rowly black, impunctate; base scabrous, with two oblique black vittae behind scutellum; disk vitreous, almost impunctate. Legs with anterior and intermediate femora strongly clavate; pos- terior tibiae with a dense brush of hairs over apical one-half. Abdomen strongly inflated to- ward apex; sternites with first three segments margined at sides and apices with dense pubes- cence; last sternite with a large circular plate, sides prominently elevated; claspers large. Length, 20-21 mm. FEMALE: Form similar. Head with eyes widely separated on front. Pronotum with dorsal calluses less prominent. Abdomen with last ster- nite excavated but lacking a plate. Length, 17 mm. TYPE-LOCALITY: Bejucos, Temascaltepec, Mexico. FLOWER RECORDS: Acacia, Spondias. New Recorps: | male, Arroyo Santiago, near Jesus Maria, Nayarit, 5 July 1955 (B. Malkin); 1 male, Mexcala, Guerrero, 29 June 1951 (H. Evans); 2 males, 20 miles [32 km] S Matias Romero, Oaxaca, 25 June 1961, on flowers of Acacia (Univ. Kans. Mex. Exped.); | female, 8 miles [ca. 13 km] NE Ca- temaco, Veracruz, 3 July 1971 (Clark, Murray, Hart, Schaff- ner). Acyphoderes fulgida, new species (Figure 4) MALE: Form moderate sized, tapering; integ- ument black, shining; antennae often brownish toward apex; elytra with disk testaceous, vitre- ous; hind femora usually testaceous over basal one-half except at extreme base. Head with eyes contiguous on front; inner margins of front ele- vated, nonpubescent. Pronotum longer than broad, sides impressed behind middle; disk with dorsal calluses elongate, barely punctate, shin- ing; sides with oblique calluses on anterior one- half; punctures between calluses fine, dense: pubescence fine, pale, appressed, denser on sides, long, pale, erect hairs numerous; proster- num densely punctate at middle, densely clothed with long, pale, erect hairs; mesosternum with a dense, pale, pubescent patch on epimeron; metasternum densely pubescent over apical one- fourth, episternum with a small pubescent patch at end. Elytra elongate, extending to fourth ab- dominal segment; lateral margins broadly black at base, narrowly black toward apex, sutural margins more broadly black at base; punctures dense on dark margins, disk very sparsely punc- tate; apices rounded, broader than antennal scape. Legs with anterior and intermediate fem- ora strongly clavate, clavae large; posterior fem- ora gradually clavate; posterior tibiae with a moderately dense brush of dark hairs over apical one-half. Abdomen not enlarged apically; ster- nites one to three densely clothed with pale pu- bescence along lateral and apical margins and with a double row at middle; last sternite shal- lowly excavated with a large blunt tubercle on CHEMSAK & LINSLEY: RHINOTRAGINI OF MEXICO 77 FiGures 4-5. Acyphoderes parva Chemsak and Linsley, male. each side at base, claspers densely pubescent. Length, 16-18 mm. FEMALE: Form similar. Head with eyes sep- arated on front by more than diameter of anten- nal scape. Pronotum very sparsely punctate be- tween calluses. Abdomen with last sternite narrowly rounded at apex. Length, 14-18 mm. Holotype, male; allotype (California Academy of Sciences) from La Pacifica, 4 km NW Canas, Guanacaste Prov., Costa Rica, 25 May 1974, on Casearia nitida flowers (P. Opler). Six paratypes (3 males, 3 females) same locality, all collected by P. Opler, 25 May 1974, 2-4 June 1973, on asclepid vine, 3 June 1971, on Trigonia floribunda; | female, 2 miles [ca. 3.2 km] NW Canas, 18-21 May 1974 (E. Giesbert); 2 females, Playa Hermosa, Guanacaste, 6-10 June 1974 (Giesbert). Dorsal view of pronotum and body of (/eft) Acyphoderes fulgida Chemsak and Linsley, male, and (right) Two additional specimens (1 male, 1 female) from Salto Eyiplantla, near Catemaco, Veracruz, Mexico, 17 Aug. 1976 (J. Hafernik and R. Garrison) are also assigned to this species. In addition to the marked sexual dimorphism, this species varies slightly in coloration. Males occasionally have the posterior femora yellow and brownish. Some of the females have the femora and tibiae pale brownish except for the apical one-half of the hind tibiae. The elongate elytra, punctation of the prono- tum, and nonbulbous abdomen will separate ful- gida from sexualis. The structure of the male copulatory apparatus is also strikingly different in the two species. 78 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 3 Acyphoderes prolixa, new species MALE: Form very elongate; head reddish; an- tennae brownish; pronotum black; dorsal callus- es reddish; elytra testaceous, margins brownish, base with a triangular black spot around scutel- lum; thoracic sterna black; legs pale, tibiae dark on dorsal edge, femora black internally on an- terior portion of club; abdomen pale basally, reddish toward apex, sternites variably infus- cated at bases. Head with eyes separated on front by about diameter of antennal scape; inner margins of front densely pubescent; antennae with segments from fourth opaque. Pronotum slightly longer than broad, sides rounded; apex and base broadly impressed; disk convex, dorsal calluses moderately elevated; calluses punctate, other punctures fine, dense; apex and base with densely pubescent bands; prosternum densely pubescent; mesosternum with epimeron densely clothed with pale appressed pubescence; meta- sternum densely pubescent anteriorly, epister- num with a dense pubescent patch posteriorly. Elytra narrow, extending at least to middle of third abdominal segment; lateral margins brown- ish, densely punctate, sutural margins elevated, pale brownish; disk shining, testaceous, very sparsely punctate; apices rounded, about as wide as antennal scape. Legs with anterior and intermediate femora strongly clavate; posterior femora petiolate, clavae moderate. Abdomen elongate, not enlarged apically; sternites densely punctate, moderately densely pubescent; fourth sternite shallowly impressed at apex; last ster- nite broadly, deeply impressed, margins elevat- ed basally, not forming angle at apex. Length, 21-25 mm. FEMALE: Form more robust, less elongate. Head with eyes widely separated on front. Pronotum reddish, black on apical and basal margins; metepisternum partially reddish. Ab- domen enlarged apically; last sternite not im- pressed. Length, 20-24 mm. Holotype, male, allotype (American Museum of Natural His- tory) and 6 paratypes (3 males, 3 females) from 10 miles [ca. 16 km] W Colima, Colima, Mexico, | Aug. 1954 (M. Cazier, W. Gertsch, Bradts). This species may be recognized by the elon- gate form and black triangular patch around the scutellum. The coloration, more petiolate pos- terior femora, and broadly excavated, margined last abdominal sternite of the males will separate prolixa from suavis. Acyphoderes delicata Horn Acyphoderes delicatus HORN, 1894:400; HAMILTON 1896: 168; LINSLEY 1942:54. MALE: Form slender, gradually tapering; in- tegument reddish; antennae orange-brown; ely- tra shining pale brownish, margins darker; un- derside partially infuscated; legs pale, femora with clavate portions brownish, hind tibiae broadly brown at apices. Head with eyes sepa- rated on front by more than diameter of antennal scape. Pronotum as long as broad, broadly rounded; disk opaque, median callus narrow, elongate, lateral pair broader, arcuate, densely punctate; pubescence fine, uniform, dense, short, erect; mesosternum with sides densely pubescent; metasternum densely pubescent over anterior one-half, episternum with a dense pubescent patch posteriorly. Elytra extending to fourth abdominal segment; base narrowly yel- lowish, sparsely punctate, margins densely punctate, disk sparsely punctate; apices round- ed, much broader than antennal scape. Legs with anterior and intermediate femora clavate, clavae small; posterior tibiae densely pubescent at apical one-half. Abdomen slender, not en- larged apically; last sternite very shallowly im- pressed at apex, sides not margined. Length, 11- 18 mm. FEMALE: Not seen. TYPE-LOCALITY: El Taste, Baja California Sur. This species is distinctively characterized by the inflated, opaque pronotum and apically slen- der abdomen. New REcorDs: 2 males, Santa Victoria Trail: La Burrera— La Laguna ridge, 1,200-1,350 m, Baja California Sur, 28 Aug. and 2 Sep. 1977 (R. L. Westcott). Acyphoderes acutipennis Thomson Acyphoderes acutipennis THOMSON, 1860:179; BATES, 1880:43; 1885:290. MALE: Form elongate, strongly tapered; in- tegument brownish; calluses of pronotum often yellowish; elytra yellowish brown. Head with eyes separated on front by about diameter of antennal scape; pubescence sparse. Pronotum longer than broad, sides impressed behind mid- dle; disk with calluses prominently elevated, punctate, subopaque; pubescence moderately dense, pale, appressed, rather uniform; meso- sternum with a dense patch of appressed pubes- CHEMSAK & LINSLEY: RHINOTRAGINI OF MEXICO cence at sides; metasternum with a small pu- bescent patch on posterior part of episternum. Elytra elongate, strongly tapered, extending to anterior margin of third abdominal segment; lat- eral margins broadly brownish, densely punc- tate, sutural margins narrowly brownish; base finely, densely asperate punctate; disk finely, sparsely punctate, each puncture bearing a short erect seta; apices truncate, narrower than di- ameter of antennal scape. Legs with anterior and intermediate femora strongly clavate. Abdomen gradually inflated toward apex; last sternite shal- lowly impressed at apex, sides barely margined. Length, 15—20 mm. FEMALE: Form similar. Head with eyes more widely separated on front. Abdomen more bul- bous, last sternite not impressed. Length, 16-20 mm. TYPE-LOCALITY: Mexico. The prominent, often pale calluses of the pronotum and rather elongate, apically narrow elytra make this species distinctive. New records: 4 males, 2 females, Temascal, Oaxaca, 3 June 1964, 1 Nov. 1963 (D. Janzen); 1 male, Tezonapa, Veracruz, July 1906; 1 male, Colima, Colima (Conradt). Acyphoderes yucateca (Bates), new combination Odontocera yucateca BATES, 1892:159, pl. 6, fig. 7; CHEMSAK 1967:78 (lectotype). MALE: Form moderately elongate, strongly tapered; integument light to dark reddish brown; elytra with disk testaceous. Head with eyes sep- arated on front by about diameter of antennal scape. Pronotum longer than broad, sides lightly impressed behind middle, base slightly narrower than apex; disk with elongate calluses punctate, opaque; punctation dense, shallow, scabrous: base and apex with narrow bands of dense, ap- pressed pubescence, long erect hairs rather sparse; meso- and metasternum with small, densely pubescent patches at sides of posterior margins. Elytra strongly tapered, extending to anterior margin of second abdominal segment: lateral margins dark from behind humeri to api- ces, densely punctate, sutural margins narrowly dark: impunctate; base moderately densely punctate, punctures sparse toward apex, each bearing a short seta: humeri and often a V- shaped basal vitta dark; apices truncate, nar- rower than diameter of antennal scape. Legs with anterior and intermediate femora strongly clavate. Abdomen moderately inflated toward We apex; two basal segments yellowish; last sternite shallowly impressed, sides lightly margined; last tergite densely clothed with appressed pubes- cence. Length, 10-18 mm. FEMALE: Form similar. Head with eyes more widely separated on front. Abdomen more in- flated toward apex; last sternite unmodified. Length, 12-18 mm. TYPE-LOCALITY: Temax, Yucatan, Mexico. The pubescent bands and less elevated callus- es of the pronotum will distinguish this species from acutipennis. New REcorbs: 4 males, Yucatan (G. F. Gaumer); 3 males, 3 females, Piste, Yucatan, 6 and 26 Sep. 1967 (E. Welling); 1 male, Yaxche, Yucatan, 10 Sep. 1964 (J. and D. Pallister); 1 male, | female, X-Can, Quintana Roo, 25 June 1967, 26 Sep. 1967 (Welling); | male, Hwy. 180, Quintana Roo, 23 Mar. 1964 (E. L. Mockford); 1 male, 26 miles [ca. 42 km] S Yucatan line, Campeche, 30 June 1963 (Porter); | male, Campeche, Campeche, 30 June 1966 (McFadden); 1 male, El Zapotal, 2 miles [ca. 3.2 km] S Tuxtla Gutiérrez, Chiapas, 9 July 1957 (J. Chemsak); | female, Puerto Castilla, Honduras, Sep. 1945 (Crowell). Acyphoderes parva, new species (Figure 5) MALE: Form rather small, short; integument pale brownish; head, pronotum, underside, and abdomen variously infuscated; elytra testa- ceous, dark margined, and with two dark, oblique basal bands. Head with eyes separated on front by a little more than diameter of anten- nal scape; inner margins of front densely pubes- cent. Pronotum longer than broad, sides slightly impressed behind middle; disk convex, dorsal calluses rather prominent, punctate; surface densely punctate, opaque; apical and basal mar- gins with narrow pubescent bands; sternum with a narrow, densely pubescent band at sides; metasternum densely pubescent, with a pubes- cent patch at end of episternum. Elytra extend- ing to anterior margin of third abdominal seg- ment; base finely scabrous, densely punctate; lateral margins densely punctate, disk finely, sparsely punctate; apices truncate, narrower than antennal scape. Legs with anterior and in- termediate femora strongly clavate, clavae large; hind femora petiolate, clavae short; hind tibiae arcuate. Abdomen enlarged toward apex: two basal segments with lateral patches of dense pubescence; last sternite broadly excavated, margins elevated at apex forming obtuse angles. Length 11-15 mm. 80 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 3 FEMALE: Form similar. Head with eyes more widely separated on front. Pronotum with dorsal calluses less prominent. Abdomen bulbous, last sternite not impressed. Length, 12-16 mm. Holotype, male; allotype (California Academy of Sciences) from 5 miles [ca. 8 km] N Mazatlan, Sinaloa, Mexico, | Aug. 1972, on flowers of Buddleia wrightii (J. A. and M. A. Chem- sak), 9-15 Aug. 1970, on B. wrightii (J. A. Chemsak). Para- types as follow: 1 female, 5 miles [ca. 8 km] N Mazatlan, 9- 15 Aug. 1970, on Buddleia (J. A. Chemsak); 10 males, Ma- zatlan, 28 Nov. 1962 (T. Gantenbein); | female, Acaponeta, Nayarit, 19 Nov. 1955 (E. C. Bay); 2 males, 15 miles [ca.24 km] NW Acaponeta, 19 Oct. 1964 (A. E. Michelbacher); 1 male, 15 miles [ca. 24 km] S Acaponeta, 20 Aug. 1964 (Burke and Apperson); | female, Teacapan, Sinaloa, 29 June 1956 (R. and K. Dreisbach); 2 females, Tuxpan, Nayarit, 20 Aug. 1964 (Burke and Apperson); | female, 60 miles [ca. 97 km] N Tepic, Nayant, 15 Aug. 1957 (J. A. Chemsak). This species averages smaller in size than most other Mexican Acyphoderes. It can be sep- arated from suavis by the different coloration, more inflated abdomen, and petiolate posterior femora. Additionally, the excavation of the last abdominal sternite of males is deeper and the margins more highly elevated. The coloration of parva varies from brownish to black and the basal oblique dark bands of the elytra are often vague. Acyphoderes suavis Bates Acyphoderes suavis BATES, 1885:290, pl. 20, fig. 20. MALE: Form elongate; integument reddish; antennae with segments 6-8 usually infuscated; elytra pale brownish with a pale triangular basal area; underside variously infuscated: front and middle femora basally and dorsally dark, hind femora dark at middle; hind tibiae dark annulate on apical one-half. Head with eyes separated on front by slightly more than diameter of antennal scape. Pronotum longer than broad, sides lightly impressed behind middle; apex and base nar- rowly impressed; dorsal calluses moderately el- evated, opaque; apex and base with a band of dense, appressed pubescence; mesosternum with epimeron densely pubescent; metasternum with a densely pubescent patch at posterior edge of episternum. Elytra extending to third abdom- inal segment; lateral margins lightly infuscated, densely punctate; disk finely, sparsely punctate; apices truncate, as broad as or narrower than antennal scape. Legs with anterior and inter- mediate femora strongly clavate, posterior pair gradually clavate. Abdomen gradually enlarged apically; last sternite impressed at apex, sides moderately produced apically. Length, 13-24 mm. FEMALE: Form similar. Abdomen more bul- bous, last sternite not impressed. Length, 15—22 mm. TYPE-LOCALITY: Cordoba, Mexico. FLOWER RECORDs: Buddleia, Jatropha (Mex- ico); Casearia, Baltimora (Costa Rica). This species may be readily recognized by the color and pubescent bands of the pronotum. In flight, adults greatly resemble and behave like those of Polistes instabilis Sauss. New Recorps: See Figure 6. Known from Mexico to Costa Rica. Genus Stenopseutes Bates Stenopseutes BATES, 1873:130; 1880:44. This genus may be recognized by the slender antennae, longitudinally tricallose pronotum, re- duced prosternal process, and slender, feebly clavate femora. TYPE-SPECIES: Sfenopseutes aeger Bates, 1873 (monotypic). Two species are known, one from Mexico. Stenopseutes sericinus Bates Stenopseutes sericinus BATES, 1880:44. FEMALE: Integument testaceous, antennae brownish. Pronotum densely clothed with ap- pressed pubescence. Elytra vitreous with a tri- angular patch of appressed pubescence at base: margins narrowly dark. Legs yellowish, femora feebly clavate. Length, 14 mm. TYPE-LOCALITY: Mexico. No new material of this species has been seen. The above characterization was based upon a 35-mm color transparency of a specimen in the British Museum (Natural History). Genus Odontocera Audinet-Serville Odontocera AUDINET-SERVILLE, 1833:546; WHITE 1855:187; THOMSON 1860:176; 1864:417; LACORDAIRE 1869:503; BATES 1873:36; 1880:42; LINSLEY 1963:165. The primary characteristic of this genus in Mexico is the vitreous disk of the elytra. Odon- tocera differs from Ommata by this character and also by the less elongate pronotum. Acy- phoderes differs by the tricallused pronotum and usually more strongly clavate femora. CHEMSAK & LINSLEY: RHINOTRAGINI OF MEXICO TYPE-SPECIES: Odontocera vitrea Audinet- Serville (Thomson designation, 1864). Key to the Mexican species of Odontocera L. Pronotum margined apically and basally by bands of dense, appressed pubes- Ge(Ca. . wees eee eee Ss 2 ee 2 Pronotum lacking dense pubescent bands, disk usually with a longitudinal median callus. Length, 11-16 mm. Mexico to Nicaragua clara 2(1). Elytra more than 3 times longer than basal width, erect hairs at base short: antennae extending beyond middle of elytra. Length, 11-20 mm. Southern Arizona to Chiapas and Veracruz ___ ae. e 3 ee ener ee ee eee aurocincta Elytra less than 3 times longer than bas- al width, erect hairs at base long; an- tennae not reaching to middle of ely- tra. Length, 12-13 mm. Tamaulipas to WiETAGHUIZ 2-82. ES et fuscicornis Odontocera fuscicornis Bates Odontocera fuscicornis BATES, 1885:289. MALE: Form slender, rather short; integument black; antennae reddish; legs brownish, femora pale at bases; elytra yellowish, vitreous, nar- rowly margined by black, apices black, base with two dark humeral vittae; abdomen with first two segments partially pale. Head with front short, eyes contiguous; antennae short, segments from sixth expanded at apices, basal segments with a few long, erect hairs beneath. Pronotum moderately coarsely, subreticulately punctate; base deeply impressed, apical and basal pubescent bands silvery, long, erect hairs numerous. Elytra less than 3 times longer than basal width; punctures on vitreous areas sparse: pubescence at base long, erect. Legs with fem- ora clavate; posterior tibiae moderately pubes- cent. Abdomen with last sternite impressed for most of its length, sides feebly elevated. Length, 12-13 mm. FEMALE: Form and coloration similar. Head with eyes well separated on front. Abdomen with last sternite not modified. Length, 13 mm. TYPE-LOCALITY: Cordoba, Mexico. The smaller size, shorter elytra and antennae will separate this species from aurocincta. New ReEcorps: 2 males, | female, 22 miles [ca. 35 km] N El Limon, Tamaulipas, 9 June 1951 (P. D. Hurd). 81 Odontocera aurocincta Bates Odontocera aurocincta BATES, 1873:42. MALE: Form elongate, slender, integument black; antennae pale with basal segments often infuscated; elytra pale, shining, base often black, lateral margins black to beyond middle: legs black or with only clavate portions of fem- ora black; abdomen with segments one and two mostly pale. Head with front moderate, eyes subcontiguous; antennae slender, extending a little beyond middle of elytra, segments from sixth slightly produced apically. Pronotum long- er than broad, sides rounded; disk coarsely, sub- confluently punctate; apex and base with a band of dense, silvery or golden, appressed pubes- cence, bands extended along sides and united: long erect hairs numerous; mesosternum dense- ly pubescent on epimeron: pubescent bands on anterior and posterior margins of metasternum and on episternum. Elytra more than 3 times longer than basal width; margins densely punc- tate, pale portion of disk rather sparsely, sepa- rately punctate; pubescence sparse, short, erect over basal half. Legs slender, femora clavate. Abdomen with last three sternites densely punc- tate, last sternite feebly impressed over most of its length, sides margined but not elevated. Length, 11-19 mm. FEMALE: Form similar; head and pronotum often reddish; antennae usually pale; legs pale, with femoral clavae black; abdomen usually mostly yellowish or reddish; pubescence golden. Head with eyes widely separated on front. Ab- domen somewhat inflated toward apex; last ster- nite rounded at apex. Length, 12-20 mm. TYPE-LOCALITY: Tehuantepec, Mexico. This species 1s easily separated by the elon- gate form and pubescent bands of the pronotum. Two distinct subspecies can be recognized. Odontocera aurocincta aurocincta Bates Odontocera aurocincta BATES, 1873:42; 1880:43. Odontocera aurocincta aurocincta;, LINSLEY 1961:163. Odontocera aurocincta var. nigroapicalis FISHER, 1947:52; LINSLEY 1961:164 (type-locality, La Gloria, Cardel, Vera- cruz). Males with pubescence golden; antennae usu- ally all pale; elytra without basal black bands; legs with femoral clubs black. Females with head and pronotum black. TYPE-LOCALITY: Tehuantepec. 82 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 3 {000 FOOT CONTOUR BONNE’S EQUAL-AREA PROJECTION FIGURE 6. RANGE: State of Nayarit to Chiapas, north to Veracruz (Fig. 7). FLIGHT PERIOD: June to August. Adults have been taken on flowers of Acacia. Odontocera aurocincta arizonensis Linsley Odontocera aurocincta arizonensis LINSLEY, 1961:163; 1963: 165. Males with silvery pubescence; antennae with basal segments infuscated; legs usually all black; elytra black basally. Females with head and pronotum reddish. TYPE-LOCALITY: Box Canyon, Santa Rita Mts., Arizona. RANGE: southern Arizona to Sinaloa (Fig. 7). FLIGHT PERIOD: July to September. Adults frequent the flowers of Acacia, Bud- dleia, Croton, and Jatropha. Although the models have not been specifi- cally determined, it is clear that the two sexes of this subspecies mimic different insects. Fe- males greatly resemble and fly like vespids, while males appear more like ichneumonids in the field. Known distribution of Acyphoderes suavis Bates in Mexico. Odontocera clara Bates Odontocera clara BATES, 1873:38; 1874:222; 1880:43, pl. 5, fig. 4. Odontocera carinicollis LINSLEY, 1934:348; 1935:84, pl. 2, fig. 6 (type-locality: Bejucos, Temascaltepec, Mexico). NEw SYNONYMY. MALE: Form slender, elongate; integument black; pronotum often reddish; elytra pale vit- reous except for narrow black margins; legs oc- casionally pale; antennae often with outer seg- ments pale annulate. Head with front moderately long, eyes subcontiguous; antennae slender, ex- tending beyond middle of elytra, segments from sixth slightly produced apically. Pronotum long- er than broad, sides rounded; disk rather finely, irregularly punctate; middle usually with a lon- gitudinal, slightly elevated callus which extends over most of the length; pubescence fine, erect, with appressed pubescence at sides; metaster- num densely pubescent. Elytra more than 3% times longer than basal width; base and black margins densely punctate, pale disk sparsely punctate; pubescence sparse. Legs slender; femora moderately clavate. Abdomen densely CHEMSAK & LINSLEY: RHINOTRAGINI OF MEXICO SCALE 400 RILOMETERS 1000 FOOT CONTOUR BONNE’S EQUAL-AREA PROJECTION FIGURE 7. Linsley (open circles). pubescent; last sternite shallowly impressed at apex, sides barely elevated. Length, 11-14 mm. FEMALE: Form similar. Pronotum a little broader. Abdomen with last sternite narrowly rounded at apex. Length, 14-16 mm. TYPE-LOCALITY: Chontales, Nicaragua. This species varies in having the pronotum reddish or black. The elongate, median callus of the pronotum characterizes it. New ReEcorps: | male, | female, 6 miles [ca. 10 km] N La Ventosa, Oaxaca, 19 July 1963 (W. A. Foster); 1 female, Su- chiapa, Chiapas, 17 July 1957 (J. A. Chemsak); | female, 5.5 miles [ca. 8.9 km] NW Acala, Chiapas, 23 June 1965 (Burke, Meyer, Schaffner); 1 female, 30 miles [ca. 48 km] NE Te- huantepec, Oaxaca, 8 July 1955 (D. Giuliani); | female, 4.4 miles [ca. 7.1 km] E Cuernavaca, Morelos, 6-8 July 1974 (Clark, Murray, Ashe, Schaffner). Genus Ommata White Ommata Wuite, 1855:194; THOMSON 1864: 166; LACORDAIRE 1869:502: Bates 1870:319; 1873:26; 1880:42; Zasciw 1966:875; 1970:37. This is probably the most difficult genus of Rhinotragini to define. It has been divided into Known distribution of Acyphoderes aurocincta aurocincta Bates (closed circles), and A. aurocincta arizonensis a number of subgenera, but a study of the entire group will be necessary to correctly determine the systematic position of the Mexican Ommata. In Mexico, the genus may be recognized by the usually long antennae, non-callused, elongate pronotum and usually non-vitreous elytra. TyYPE-sPECIES: Ommata elegans White, 1855 (monotypic). Three species are presently known from Mex- ico. Key to the Mexican species of Ommata 1. Pronotum elongate, cylindrical, not broad- ly impressed at base and apex: elytra subglabrous, with fine erect hairs arising frommeach. puncte) 2 a eee 2 Pronotum about as long as broad, broadly impressed at base and apex; elytra densely clothed with appressed, golden pubescence. Length, 11 mm. Veracruz to Panama sallaei 2. Integument, especially pronotum and un- derside, metallic bluish to greenish: pos- 84 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 3 terior tibiae densely clothed with sub- erect, black setae. Length, 11-13 mm. Veracruz t0 Oaxaca. 22-2 cyanea Integument non-metallic, abdomen often reddish; posterior tibiae sparsely pubes- cent. Length, 7-10 mm. Morelos to Gua- temala championella Ommata championella Bates Ommata (Eclipta) championella BATES, 1880:42; Zajciw 1970:38. Ommata (Ommata) rubriventris LINSLEY, 1934:347; 1935:84; ZAJCIW 1966:91 (type-locality: Tejupilco, Temascaltepec, Mexico). NEW SYNONYMY. MALE: Integument black; antennae with outer segments basally yellow annulate; elytra with disk longitudinally pale brownish; femora yel- lowish basally; abdomen often reddish. Head with antennae enlarged from seventh segment. Pronotum narrow, shallowly reticulate-punc- tate. Elytra moderately coarsely, densely punc- tate, punctures becoming denser toward apex. Abdomen elongate; last sternite impressed at apex. Length, 7-9 mm. FEMALE: Form more robust. Prothorax often reddish. Abdomen broader, reddish; last sternite narrowly rounded at apex. Length, 8-10 mm. TYPE-LOCALITY: Calderas, Guatemala. RANGE: Morelos, Mexico to Guatemala. The small, slender form, punctation of the pronotum, and paler longitudinal stripes of the elytra will separate this species from other known Mexican Ommata. New ReEcorpbs: | female, 7 miles [ca. 11 km] SW Yautepec, Morelos, 2 July 1961, 3,500 ft [ca. 1,067 m] (Univ. Kansas Mex. Exped.); 1 male, | female, 3 miles [ca. 5 km] SE La Trinitaria, Chiapas, 18, 19 June 1965 (Burke, Meyer, Schaff- ner); | female, Las Margaritas, Chiapas, 18 June 1965 (Burke, Meyer, Schaffner); | female, 7 miles [ca. 11 km] SW Teopisca, Chiapas, 23 May 1969 (J. M. Campbell). Ommata cyanea Bates Ommata cyanea BATES, 1885:288. Ommata (Eclipta) cyanea; ZAsiciw, 1970:38. MALE: Integument greenish or bluish metallic; antennae bluish, yellow annulate from fourth segment; legs bluish; elytra brownish down mid- dle. Head with antennae as long as elytra, seg- ments from sixth apically expanded. Pronotum cylindrical; coarsely punctate at middle; sparse- ly pubescent; strongly narrowed toward apex; apices truncate. Legs with posterior femora elongate, feebly clavate. Abdomen with last sternite deeply impressed. Length, 11 mm. FEMALE: Form similar. Antennae black; basal segments densely pubescent. Abdomen broader: last sternite narrowly rounded at apex. Length, 12-13 mm. TYPE-LOCALITY: Cordova, Mexico. The metallic coloration makes this species distinctive. New Recorps: | female, 5 miles [ca. 8 km] S Lake Cate- maco, Veracruz, 6 July 1961 (D. H. Janzen); 1 female, | male, Temescal, Oaxaca, 6 July 1965 (G. Nelson and family). Ommata sallaei Bates Ommata sallaei BATES, 1885:289, pl. 20, fig. 15. FEMALE: Integument yellowish; mandibles, eyes, antennae basally and at apices of outer segments, apical and basal margins and rounded median spot on pronotum, elytra except for tri- angular basal area, apices and dorsal edges of femora and bands on clubs of hind femora, and parts of underside black. Antennae slender, seg- ments feebly produced at apices. Eyes widely separated on front. Pronotum coarsely punctate, sparsely pubescent. Elytra opaque, densely punctate and pubescent, apices truncate. Length, 11 mm. TYPE-LOCALITY: Cordova, Veracruz. This species is distinctive among Mexican Ommata in its color and opaque elytra. It is also known from Cerro Campana in Panama. ACKNOWLEDGMENTS These studies were carried out in conjunction with a National Science Foundation-sponsored study on North American Cerambycidae through Grant DEB 76-23849 AOI. The authorities of the following institutions and individuals are grate- fully acknowledged for the loan of specimens: American Museum of Natural History, New York; California Academy of Sciences, San Francisco; Canadian National Collection, Otta- wa; Cornell University, Ithaca; Essig Museum of Entomology, Berkeley; Field Museum of Natural History, Chicago; Los Angeles County Museum of Natural History; Museum of Com- parative Zoology, Cambridge; Texas A & M University, College Station; United States Na- tional Museum of Natural History, Washington, D.C.; University of Arizona, Tucson; Univer- sity of California, Davis; University of Kansas, CHEMSAK & LINSLEY: RHINOTRAGINI OF MEXICO Lawrence; E. Giesbert, D. Marqua, G. Nelson, and R. Westcott. Celeste Green prepared the illustrations and Kathleen Sorenson the maps. LITERATURE CITED AUDINET-SERVILLE, J. G. 1833. Nouvelle classification de la famille des longicornes. Ann. Soc. Entomol. France. 2:528- SW/3 AURIVILLIUS, C. Berlin. Bates, H. W. 1870. Contributions to an insect fauna of the Amazon Valley (Coleoptera, Cerambycidae). Trans. Ento- mol. Soc. London. 1870:243-335, 391444. . 1873. Notes on the longicorn Coleoptera of Tropical America. Ann. Mag. Nat. Hist. (4) 11:21-45, 117-133. 1874. Supplement to the longicorn Coleoptera of Chontales, Nicaragua. Trans. Entomol. Soc. London. 1874 (2):219-235. 1879-1885. Longicornia. Biologia-Centrali-Ameri- cana, Insecta, Coleoptera. S:iii-xii, 1-436, pls. 1-25. 1892. Additions to the Longicornia of Mexico and Central America with remarks on some of the previously recorded species. Trans. Entomol. Soc. London. 1892 (2): 143-183, 3 pls. CHAMPION, G. C. 1892. [Footnote proposal of Charisia to replace Charis Newman]. Trans. Entomol. Soc. London. 1892:161. CHEMSAK, JOHN A. 1967. Lectotype designations of Cer- ambycidae in the British Museum (Natural History). J. Kan- sas Entomol. Soc. 40:73-81. CHEVROLAT, L. A. A. 1838. decouverts par M. Lanier dans I’intérieur de l’ile de Cuba. Rev. Zool. 1838:279-286. FisHER, W. S. 1930. Notes on the rhinotragine beetles of the family Cerambycidae, with descriptions of new species. Proc. U.S. Natl. Mus. 77:1-20. . 1947. New cerambycid beetles belonging to the tribe Rhinotragini. Proc. U.S. Natl. Mus. 97:47-57. . 1953. New cerambycid beetles belonging to the tribe Rhinotragini. Pan-Pac. Entomol. 29: 14-17. HAMILTON, J, in LENG AND HAMILTON, 1896. Synopsis of the Cerambycidae of North America. Part 3, The Lamiinae. Trans. Am. Entomol. Soc. 23:101-178. 1912. Coleopterorum catalogus 39: 1-574. Insectes coleopteres inedits, 85 Horn, G. H. 1894. The Coleoptera of Baja California. Proc. Calif. Acad. Sci. (2) 4:302-449. JACQUELIN DU VAL, P. N. C. in SAaGRA, 1857. Insectes. Ordre des coleoptéres. Histoire. . .de ile de Cuba. 7:137— 328. KirBy, W. 1818. A century of insects, including several new genera described from his cabinet. Trans. Linn. Soc. Lon- don 12:375-453. LACORDAIRE, J. T. 1869. Histoire naturelle des insectes. Genera des coléopteres ou exposé méthodique et critique de tous les genres proposés jusqu’ici dans cet ordre dinsects. 8:1-552. Linstey, E. G. 1934. A new genus and several new species of Neotropical rhinotragine beetles. Rev. Entomol. 4: 345— Soir 1935. Studies in the Longicornia of Mexico. Trans. Am. Entomol. Soc. 61:67—102, | pl. . 1942. Contributions toward a knowledge of the insect fauna of Lower California. No. 2. Coleoptera: Cerambyci- dae. Proc. Calif. Acad. Sci. (4) 24:21-96, pls. 4-5. 1961. A new rhinotragine cerambycid from Arizona and Sonora. Entomol. News 72: 163-164. . 1963. The Cerambycidae of North America. Part IV. Taxonomy and classification of the subfamily Cerambycin- ae, tribes Elaphidionini through Rhinotragini. Univ. Calif. Publ. Entomol. 21:165, 52 figs. NewMan, E. 1840. Entomological notes. The Entomologist 1: 1-16; 2:17-32. THOMSON, J. 1860. Essai d'une classification de la famille des cerambycides et matériaux pour servir a une monogra- phie de cette famille. Pp. 1-404, illus. Paris. 1864. Systema cerambycidarum ou expose de tous les genres compris dans la famille des cerambycides et fam- illes limitrophes. Mem. Soc. Roy. Sci. Liege. 19: 1-540. Wuite, A. 1855. Catalogue of coleopterous insects in the collection of the British Museum. Pt. VIII, Longicornia II, pp. 175-412, pls. 5-10. London. Zaiciw, D. 1966. Estudos do genero “Ommata™’ White, 1855. II: Subgenero **Ommata’’. Rev. Brasil. Biol. 26:87- 92, | fig. 1970. Estudos do genero Ommata White, 1855. V: Notas sobre 0 subgenero Eclipta Bat., 1873. Atas Soc. Biol. Rio de Janeiro 13:37—40. ZAYAS, F. DE. 1956. El genero Essostrutha Thoms., adicion de una especie nueva. Mem. Soc. Cubana Hist. Nat. 23:105-114, 1 pl. 1975. Revision de la familia Cerambycidae. Acad. Cien. Cuba, Inst. Zool. 443 pp., 35 figs. ho A) en WS NTE ry as | ke Oe) Marine Biological Labbraicr, LIBRARY scriNtEs 0 Woods Hole, Mass. ; PROCEEDING OF THE CALIFORNIA ACADEMY Vol. 42, No. 4, pp. 87-133, 70 figs. STUDIES ON THE NEBRIINI (COLEOPTERA: CARABIDAE), III.! NEW NEARCTIC NEBRIA SPECIES AND SUBSPECIES, NOMENCLATURAL NOTES, AND LECTOTYPE DESIGNATIONS? By David H. Kavanaugh Department of Entomology, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118 ABSTRACT: The purpose of this paper, third of a series on the nebriine carabid beetles, is to upgrade the status of nomenclature for Nearctic members of genus Nebria in preparation for subsequent papers on classi- fication, phylogeny, zoogeography, and natural history of members of the Nearctic fauna. Names are provided for five undescribed species [Nebria carri (type-locality—Dollarhide Summit, Idaho), N. darlingtoni (type-lo- cality—South Fork American River, California), N. gouleti (type-locality—Rattlesnake Creek, Washington), N. lituyae (type-locality—Mount Blunt, Lituya Bay, Alaska), and N. navajo (type-locality—19 miles SW of Kayenta, Arizona)]. Twenty-three subspecies are also described as new [Nebria acuta quileute, N. arkansana edwardsi, N. arkansana oowah, N. arkansana uinta, N. fragilis teewinot, N. gebleri cascadensis, N. gebleri fragariae, N. gebleri siskiyouensis, N. gyllenhali lassenensis, N. gyllenhali lindrothi, N. kincaidi balli, N. lacustris bellorum, N. meanyi lamarckensis, N. meanyi sylvatica, N. nivalis gaspesiana, N. obliqua chuskae, N. sahlbergii modoc, N. sahlbergii triad, N. schwarzi beverlianna, N. spatulata sierrae, N. trifaria utahensis, N. vandykei wyeast, and N. zioni oasis|. Diagnosis and comment on geographical distribution are provided for each new taxon. Lectotypes are designated for Carabus gyllenhali Schonherr, C. nivalis Paykull, Helobia castanipes Kirby, Nebria bifaria Mannerheim, N. eschscholtzii Menetries, N. gregaria Fischer von Waldheim, N. hudsonica LeConte, N. ingens Horn, N. livida LeConte, N. mannerheimii Fischer von Waldheim, N. metallica Fischer von Waldheim, N. mollis Motschulsky, N. moesta LeConte, N. obliqua LeConte, N. sahlbergii Fischer von Waldheim, N. suturalis LeConte, N. trifaria LeConte, N. vandykei Banninger, N. violacea Motschulsky, and N. viridis Horn. Nebria longula LeConte and N. obtusa LeConte are recognized as junior synonyms of N. suturalis LeConte and N. obliqua LeConte, respectively. Nebria rathvoni LeConte is reduced in status to a subspecies of N. gebleri Dejean. Emendations or restrictions of type-localities are provided for Nebria catenata Casey, N. eschscholtzii Menetries, N. expansa Casey, N. hudsonica LeConte, N. incerta Casey, N. ingens Horn, N. longula LeConte, N. obliqua LeConte, N. oregona Casey, N. ovipennis LeConte, and N. tenuipes Casey. Additional notes on type-specimens and nomenclature are provided for other species group names. INTRODUCTION ‘ Papers which serve as contributions I and II in this series My interest in beetles of the genus Nebria are, respectively, Kavanaugh (1971) and Kavanaugh and Mar- _—_ Latreille began in 1967. While collecting carabid Bebo e772). beetles in the Rocky Mountains of Colorado, I ° Data presented here have been extracted from the author's EOnEoteatedodiiveouneprcterced haunts unpublished Doctoral Dissertation, submitted to the Univer- y P eee sity of Alberta, Edmonton, September 1978 (Kavanaugh namely, riparian, nival, and peri-nival habitats 1978). This specialization provided frequent encov [87] 88 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 4 ters with members of various Nebria species because, in these habitats, few insect groups ap- proach Nebria in their conspicuous abundance. With the aid of Lindroth’s (1961) treatment of the genus for Canada and Alaska, I was able to identify most of my material. It soon became evident that Nebria species formed a biologi- cally important and interesting element of the montane fauna. Species diversity and population densities in suitable microhabitats were impres- sively high, and I began to recognize patterns of altitudinal zonation among the resident species. Observations made on collecting trips to mon- tane areas in Utah and California suggested that similar patterns of Nebria species diversity and habitat distribution existed in these and other areas in western North America. The opportunity to begin serious study of Nearctic Nebria appeared in July 1970, when I enrolled at the University of Alberta, Edmon- ton. Although Nearctic Nebria species were at that time already among the taxonomically bet- ter-known carabid groups, I hoped to build on my previous interest and experience through ad- ditional studies which seemed both feasible and potentially rewarding scientifically. My plan was to obtain data with which to further define the structural and distributional limits of Nearctic species and explore the evolutionary relation- ships among them. Initial goals were the for- mulation of an improved infrageneric classifi- cation of the New World species and generation of hypotheses concerning the historical devel- opment of diversity and distributional patterns observed in the fauna. Nebria, in the broadest sense, comprises a group of over 500 nominal taxa which, in aggre- gate, demonstrate Holarctic distribution. The Palaearctic component of the genus is more di- verse, both in species and major lineages, than the Nearctic. Consequently, all hypotheses on relationships among Nearctic species and on historical development of the Nearctic fauna would be highly speculative in the absence of a working knowledge of the Palaearctic fauna. Fortunately, as work toward the initial project goals progressed, new sources of taxonomic data became available as by-products of field work and efforts to obtain comparative material on loan. These additional resources have per- mitted expansion of the project to include cov- erage of the Palaearctic Nebria fauna, compar- isons with members of other nebriine geriera and related carabid tribes, and the use of additional character systems in study of the Nearctic Ne- bria fauna. These other systems include external structure of immature stage individuals, life his- tory features (especially life cycle timing), hab- itat preferences, and adult internal structure. Various facets of the expanded project have progressed at different rates, and some of these are yet incomplete or barely begun. This paper represents a first report on the Nearctic Nebria fauna, specifically on nomenclature relating to same. My purpose is twofold. First, I provide names for five species and 23 subspecies not previously described. These names are needed immediately for use in several other manuscripts and by several other workers. To this end, data and discussion presented for each name are lim- ited to little more than the minimum required by the International Code of Zoological Nomencla- ture for availability. My second purpose is to provide additional notes on nomenclature for previously described Nearctic species, including emendations and/or restrictions of type-locali- ties, notes on type-specimens, and designations of lectotypes. In a subsequent paper (manuscript in prepa- ration) I will present additional information on Nearctic Nebria taxa, including those presented here as new. Each taxon will be more fully de- scribed and illustrated. A classification and key for identification of adults will be provided, as will results of comparative studies on structure, distribution, and natural history. A review of the known fossil record of Nearctic Nebria will also be included. MATERIALS This study is based on examination of over 66,000 Nearctic and 5,800 Palaearctic adult Ne- bria specimens. Approximately 35,000 speci- mens were borrowed from various institutional and private collections in North America, Eu- rope, and Asia. Following is a list of abbrevia- tions used in the text which refer to various col- lections from which specimens were received. Most, but not all, correspond to abbreviations proposed by Arnett and Samuelson (1969). Where appropriate, names of curators or assis- tants who sent specimens are also included. AHab—A. Habu, National Institute of Agricultural Sciences, Tokyo, Japan. KAVANAUGH: NEW NEARCTIC NEBRIA ALar—A. Larochelle, College Bourget, Rigaud, Québec JOP 1P0. AMor—A. Morgan and A. V. Morgan, University of Water- loo, Waterloo, Ontario N2L 3G1. AMNH—American Museum of Natural History, New York, New York 10024; L. H. Herman, Jr. ANSP—Academy of Natural Sciences, Philadelphia, Penn- sylvania 19103; W. W. Moss. BFCa—B. F. Carr, 24 Dalrymple Green NW, Calgary, Al- berta T3A 1Y2. BMNH—British Museum (Natural History), London, En- gland; P. M. Hammond, H. K. Kenward, M. E. Bacchus. BRot—B. Rotger, CR., Immaculate Heart of Mary Church, P.O. Box 451, Pagosa Springs, Colorado 81147. CArm—C. Armin, 191 West Palm Avenue, Reedley, Califor- nia 93654. CAS—California Academy of Sciences, San Francisco, Cali- fornia 94118; H. B. Leech. CDA—California State Department of Food and Agriculture, Sacramento, California 95814; F. G. Andrews. CNC—Canadian National Collection of Insects, Biosystema- tics Research Institute, Ottawa, Ontario KIA 0C6; R. de Ruette, A. Smetana. CSU—Colorado State University, Fort Collins, Colorado 80521; T. O. Thatcher. CUB—University of Colorado, Boulder, Colorado 80302; H. Rodeck. CUIC—Cornell University, Ithaca, New York 14850; L. L. Pechuman. DBUM—Université de Montreal, Montreal, Quebec H3C 3J7; M. Coulloudon. DENH—University of New Hampshire, Durham, New Hampshire 03824; W. J. Morse. DEUN—University of Nebraska, Lincoln, Nebraska 68503; B. C. Ratcliffe. DHKa—D. H. Kavanaugh, California Academy of Sciences, San Francisco, California 94118. DJLa—D. J. Larson, Department of Biology, Memorial Uni- versity of Newfoundland, St. John’s, Newfoundland A1C 5S7. DMan—D. Mann, University of Washington, Seattle, Wash- ington 98195. DRWh—D. R. Whitehead, Systematic Entomology Labora- tory, U.S.D.A., % United States National Museum, Wash- ington D.C. 20560. DZEC—Montana State University, Bozeman, Montana 59715; N. L. Anderson. EAMa—E. A. Martinko, University of Kansas, Lawrence, Kansas 66045. EDNC—North Carolina Department of Agriculture, Raleigh, North Carolina 27602; J. F. Greene. EMUS—Utah State University, Logan, Utah 84332; W. J. Hanson. ETHZ—Eidgenéssische Technische Hochschule Entomolo- gisches Institut, Ziirich, Switzerland; W. Sauter. FMNH—Field Museum of Natural History, Chicago, Illinois 60605; H. Dybas. GASh—G. A. Shook, 1209 West Hays, Boise, Idaho 83702. GRNo—G. R. Noonan, Milwaukee Public Museum, Milwau- kee, Wisconsin 53233. HGou—H. Goulet, Biosystematics Research Institute, Otta- wa, Ontario K1A 0C6. 89 HoKn—R. L. Hoffman and L. Knight, Radford College, Rad- ford, Virginia 24141. HNHM—Hungarian Natural History Museum, Budapest, Hungary; S. Horvatoviich. ICCM—Carnegie Museum, Pittsburg, Pennsylvania 15213; G. E. Wallace. INHS—Illinois Natural History Survey, Urbana, Illinois 61803; M. W. Sanderson. ILar—I. La Rivers, University of Nevada, Reno, Nevada 89507. ISUI—Iowa State University, Ames, Iowa 50010; R. Miller. IUIC—Indiana University, Bloomington, Indiana 47401; F. N. Young. JBel—J. Belicek, 9345-98A Street, Edmonton, Alberta T6E 3N2. JKus—J. Kuster, University of Alberta, Edmonton, Alberta T6G 2E3. JNeg—J. Negre, 9 Boulevard de Lesseps, Versailles, France. JSch—J. Schuh, 4039 Shasta Way, Klamath Falls, Oregon 97601. JSpe—J. Spence, University of Alberta, Edmonton, Alberta T6G 2E3. JVMa—J. V. Matthews, Jr., Geological Survey of Canada, Ottawa, Ontario KIA OE8. KMFe—K. M. Fender, 911 Ashwood Avenue, McMinnville, Oregon 97128. KSUC—Kansas State University, Manhattan, Kansas 66502; H. D. Blocker. KUSM—University of Kansas, Snow Museum, Lawrence, Kansas 66045; G. Byers. LACM—Los Angeles County Museum of Natural History, Los Angeles, California 90007; C. L. Hogue. LEMC—Macdonald College, Lyman Entomological Museum, Ste. Anne de Bellevue, Québec J7V 3N8. LRus—L. Russell, 828 NW 27th Street, Corvallis, Oregon 97330. MCZ—Museum of Comparative Zoology, Harvard Universi- ty, Cambridge, Massachusetts 02138; J. F. Lawrence. MGF—Museum G. Frey, Entomologisches Institut, Tutzing bei Miinchen, West Germany; M. Wirmli. MHNG—Museum d°’Histoire Naturelle, Geneva, Switzer- land; C. Besuchet. MNHP—Muséum National d’Histoire Naturelle, Paris, France; A. Bons, H. Perrin. MLLu—M. L. Luff, University of Newcastle, Newcastle- upon-Tyne, England. MSU—AMichigan State University, East Lansing, Michigan 48823; R. L. Fischer. NMB—Naturhistorisches Museum Basel, Basel, Switzerland; W. Wittmer. NMDo—N. M. Downie, 505 Lingle Terrace, Lafayette, In- diana 47901. NMPC—National Museum in Prague, Prague, Czechoslova- kia; Z. Mlynar. NRSS—Naturhistoriska Riksmuseet, Stockholm, Sweden; T. Nyholm. NSDA—Nevada State Department of Agriculture, Reno, Ne- vada 89504; R. C. Bechtel. ODA—Oregon State Department of Agriculture, Salem, Or- egon 97310; R. L. Westcott. OSEC—Oklahoma State University, Stillwater, Oklahoma 74074: W. A. Drew. 90 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 4 OSUC—Ohio State University, Columbus, Ohio 43210; C. A. Triplehorn. OSUO—Oregon State University, Corvallis, Oregon 97331; P. Oman. PADA—Pennsylvania State Department of Agriculture, Har- risburg, Pennsylvania 17120; T. J. Henry. PMCh—P. M. Choate, Jr., University of Florida, Gainesville, Florida 32601. PMNH—Peabody Museum of Natural History, Yale Univer- sity, New Haven, Connecticut 06520; C. L. Remington, K. W. Brown. PSMi—P. S. Miliotis, Depot Street, Dunstable, Massachu- setts 01827. PSUC—Pennsylvania State University, University Park, Pennsylvania 16802; K. C. Kim. PUCA—Pacific Union College, Angwin, California 94508; L. E. Eighme. PURC—Purdue University, Lafayette, Indiana 47907; A. Pro- vonsha. RCGr—R. C. Graves, 627 Crestview, Bowling Green, Ohio 43402. RCra—R. Crawford, University of Washington, Seattle, Washington 98195. RDav—R. Davidson, University of Vermont, Burlington, Vermont 05401. RFre—R. Freitag, Lakehead University, Thunder Bay, On- tario P7B SE1. ROM—Royal Ontario Museum, Toronto, Ontario MSS 2C6; G. B. Wiggins. RPPa—R. P. Papp, Bernice P. Bishop Museum, Honolulu, Hawaii 96818. RTBe—R. T. Bell, University of Vermont, Burlington, Ver- mont 05401. SFVS—California State University, Northridge, California 91324; P. F. Bellinger. SDSU—South Dakota State University, Brookings, South Dakota 57006; E. U. Balsbaugh, Jr. SJSU—San Jose State University, San Jose, California 95114; J. G. Edwards. TCBa—T. C. Barr, Jr., University of Kentucky, Lexington, Kentucky 40506. TNak—T. Nakane, National Science Museum, Tokyo, Japan. TMZM—Troms¢g Museum, Tromsg, Norway; J. H. Ander- sen. UAFA—University of Arkansas, Fayetteville, Arkansas 72701; R. T. Allen. UASM—University of Alberta, Strickland Museum, Edmon- ton, Alberta T6G 2E3; G. E. Ball. UBC—University of British Columbia, Spencer Museum, Vancouver, British Columbia V6T 1W5; G. G. E. Scudder. UCB— University of California, Essig Museum of Entomol- ogy, Berkeley, California 94720; J. A. Chemsak. UCD—University of California, Davis, California 95616; R. O. Schuster. UCR—University of California, Riverside, California 92502; S. Frommer. UIMI—University of Idaho, Moscow, Idaho 83843; W. F. Barr. UMHF—Universitetets Zoologiska Museum Entomologiska Avdelningen, Helsingfors, Finland; H. Silfverberg. UMMZ—University of Michigan, Ann Arbor, Michigan 48104; R. D. Alexander. UMRM—University of Missouri, Columbia, Missouri 65201; W.R. Enns. UMTF—University of Turku, Turku (Abo), Finland; H. Hip- pa. USNM—United States National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560; T. L. Erwin. UWBM— University of Washington, Burke Museum, Seattle, Washington 98105; M. H. Hatch, S. Rohwer. UWEM—University of Wisconsin, Madison, Wisconsin 53706; L. J. Bayer. UWLW—University of Wyoming, Laramie, Wyoming 83070; R. J. Lavigne. VMKi—-’V. M. Kirk, Northern Grain Insects Research Lab- oratories, Brookings, South Dakota 57006. WSU—Washington State University, Pullman, Washington 99163; W. J. Turner. ZILR—Zoological Institute, Academy of Science, Leningrad, U.S.S.R.; O. L. Kryzhanovskij, V. G. Shilenkov. ZMKD—Universitetets Zoologiske Museum, Kgbenhavn, Denmark; O. Martin. ZMLS—Zoological Institute, University of Lund, Lund, Swe- den; C. H. Lindroth. ZMUM—Zoological Museum, Moscow University, Moscow, U.S.S.R.; S. Keleinikova. ZSBS—Zoologische Sammlung des Bayerischen Staates, Munchen 19, West Germany; G. Scherer. I have been able to study the type-specimens of all 85 previously described Nearctic nominal taxa through the cooperation of many of the cu- rators listed above. The following three minor nomenclatural problems must remain unsolved, but none, I think, will require further attention. Nebria melsheimeri Sturm (1826:173) and Ne- bria eschscholtzii Sturm (1826:173) (not of Meé- nétries) must be considered nomina nuda be- cause these names appear only in Sturm’s catalog to his collection, unaccompanied by either description or statement of locality; and no type-specimens exist. The problem with and current status of the name Nebria elias Mot- schulsky is discussed with a treatment of Nebria gyllenhali (Schonherr) below. Where necessary, lectotypes have been cho- sen and so labelled, and their formal designa- tions appear here. I have also labelled holotypes for previously described species names where prior labels were unclear or lacking. Holotypes designated for taxa described here as new (with the exception of the holotype of N. trifaria utah- ensis n. ssp.) have been deposited in the type collection of the California Academy of Sci- ences. METHODS A detailed discussion of preparative and pro- cedural methods used in this project will be in- KAVANAUGH: NEW NEARCTIC NEBRIA cluded in a subsequent paper (Kavanaugh, manuscript in preparation). I present here com- ment on only those methods which relate spe- cifically to data and results presented in this pa- per. DISSECTING TECHNIQUES.—Characters used in diagnoses of many taxa named here include form of the hindwings and genitalia of adults. Successful examination of these structures re- quired some dissection. Specimens were pre- pared for dissection by soaking them for 5 or 10 minutes in boiling water. Addition of a very small amount of liquid detergent hastened the relaxation process. Specimens collected in ethyl acetate fumes were relaxed easily, but material collected in potassium cyanide fumes or in ethyl alcohol required a brief (two to five minute) treatment in warm potassium hydroxide (10% solution). A general impression of the size (but not shape) of the hindwings was obtained by simply lifting the left elytron to expose the wing. For a study of venation and wing shape, the left hindwing was torn free at the wing base using fine forceps, then spread and mounted in alcohol on a slide. Permanent mounts (in Euparal me- dium) were made for representatives of all taxa studied; however, most hindwings were later dried, glued to cards, and pinned with their re- spective specimens. The apex and shaft of the median lobe and parameres in male specimens or the coxostyli in female specimens were quickly examined in re- laxed specimens by simple eversion of these structures using fine forceps. For study of the base of the median lobe in males or the sper- matheca, bursa copulatrix, and other internal structures in females, the genitalia were extract- ed as a unit by tearing the membranous connec- tions between eighth tergum and sternum and the ‘‘ring sclerite’’ (in males) or the valvifers, paraprocts, and proctiger (in females). Genitalia were next treated for 5 to 10 minutes in hot (10%) potassium hydroxide solution to remove nonsclerotized tissue, rinsed in water, further dissected as needed, then transferred to and ex- amined in glycerine. After genitalic specimens were studied and drawn, they were placed in polyethylene microvials in a drop of glycerine and pinned with their respective specimens. SEx DETERMINATION.—Determination of the sex of individuals was made simply by reference 91 to form of the front tarsi. Adult males have the basal three or four tarsomeres dilated, laterally expanded, and have pads of adhesive setae on ventral surfaces of the basal two or three tar- someres. Females have all tarsomeres slender and without ventral pads of adhesive setae. MEASUREMENTS.—The only mensural char- acter used in this study is ‘‘standardized body length.’’ This expression refers to the sum of three measurements (Fig. 1A): length of head, measured along midline from apical margin of clypeus to a point opposite posterior margin of eye; length of pronotum, measured along mid- line from apical to basal margin; and length of elytron, measured along midline from apex of scutellum to a point opposite elytral apex. While this measure of relative size avoids error due to variation in extension or retraction of body parts among specimens, it consistently underesti- mates the apparent total body length of speci- mens (such as could be measured from the most anterior to the most posterior point) by from 12 to 18 percent. Range in standardized body length in my sam- ple for each taxon was established by visual se- lection of smallest and largest specimens for each sex. All measurements were made with the aid of a Leitz stereoscopic dissecting micro- scope at a magnification of 16 diameters, using a calibrated ocular grid with a scale interval of 0.1 mm. ILLUSTRATIONS AND MaAps.—Line drawings illustrating structural characters were made with the aid of an ocular grid mounted in a Leitz stereoscopic dissecting microscope. Unless oth- erwise noted, scale lines which accompany il- lustrations equal 1.0 mm. All drawings of homologous structures are consistent in scale and aspect to facilitate comparisons. In some drawings, uniform sparse stippling is used to in- dicate membranous areas. Where only part of a structure is illustrated (such as the basal region of an elytron), a wavy line serves as the artificial margin of the drawing. Maps are presented to illustrate known distri- butions of taxa. Symbols used denote approxi- mate locations of samples (see taxon descrip- tions for respective lists of sample localities). (Written permission has been obtained for use of Goode Base Map No. 202 [or parts thereof] [copyright by the University of Chicago, De- partment of Geography].) 92 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 4 FORMAT FOR PRESENTATION OF NEW TAXA.— Included for each new name are: a synonymy (i.e., list of names which have been misapplied in the literature to members of this taxon, in- cluding all known literature records); designa- tions and listings of type-specimens and their places of deposition; designation of type-locali- ty; the diagnostic combination (i.e., those char- acter states which, in combination, distinguish members of the taxon from members of other taxa); derivation of the taxon name; a brief syn- opsis of geographical distribution; and a list of localities (including months of collection and place of deposition) for paratypes. Several comments are required on the format and data content of the locality lists. For con- venience, all data (including countries, states or provinces, counties, and localities) are arranged alphabetically. All records are from specimens I have personally studied. Records solely from the literature are not included here. The geo- graphical subdivisions ‘‘County’’ or *‘Comté”’ are used where possible as an aid in organizing the data. Provinces and states for which these subdivisions are not used include: Alberta, Brit- ish Columbia, Manitoba, Saskatchewan, Yukon Territory, and Alaska. All national parks are treated as counties; and all localities within park boundaries are listed under the park rather than under their appropriate counties. Because alti- tude data are often important in defining the hab- itat ranges of Nebria taxa, these data are re- corded here where known. No attempt has been made, however, to present altitude data not ac- tually recorded on the specimen labels. Where necessary, altitude values have been converted to their metric equivalents. Only the months in which specimens were ac- tually collected at a given locality have been not- ed here. This level of precision adequately de- scribes the temporal distributions of most samples. The addition of year and day data would have expanded the size of this paper greatly but contributed little. I have also omitted the names of collectors in order to conserve space. Collections in which specimens are de- posited are noted for each locality, but holdings for each collection are not itemized. Specimens with illegible or essentially mean- ingless locality labels are grouped with speci- mens bearing no locality labels under the head- ing “‘SPECIMENS WITHOUT LOCALITY DATA.” Specimens listed under the heading ** DOUBTFUL RECORDS’’ are of two types: (1) those labelled as from localities outside the known and prob- able ranges of the taxon; and (2) those labelled as from localities within the known geographical range of a taxon but from areas where I am cer- tain no suitable habitat for the beetles exists. For some specimens in category (2), however, where appropriate habitat can be found within reasonable proximity of the stated locality, I have chosen to add the word “‘area’’ after the entry. Although imprecise, this convention ac- commodates the common practice among some collectors of simply recording the nearest city or town on specimen labels rather than more precise locality data. CRITERIA FOR RANKING TAXA.—AII available evidence supports the universality of sexual re- production among nebriine species. Because ap- parently all Nebria species are bisexual, I ac- cept as appropriate Mayr’s (1969:26) proposed definition of the biological species (as amended by Whitehead 1972:139), namely, that species are populations or groups of populations through which gene flow actually or potentially exists, but which are reproductively [intrinsically] iso- lated from all other such populations [or groups of populations]. As noted by Whitehead (1972:139) and Larson (1975:251), the species so defined is a category with a nonarbitrary, objec- tive basis; but this applies only to the extent that interspecific hybridization is excluded or non- introgressive. Success in application of this species definition to the recognition of contem- porary species and, therefore, the ‘‘objective”’ bases of taxa recognized depend on the degree to which reproductive isolation can be demon- strated or inferred where it exists and excluded where it does not. Direct, unequivocal evidence for this intrinsic isolation or lack of same can seldom, if ever, be provided, even from breeding tests. Only criteria by which indirect evidence can be evaluated are generally useful. The following working criteria were used in recognizing species. Two sympatric or parapa- tric forms were considered separate species if all individuals (of one or both sexes) of one differ from all members (of the same or both sexes) of the other in one or more structural characters (other than color or size). In practice, members of sympatric or parapatric Nebria species gen- erally differ in several characters and therefore easily fulfill this criterion. Two allopatric forms were considered separate species if differences KAVANAUGH: NEW NEARCTIC NEBRIA between members of each are roughly equiva- lent to differences between members of two closely related sympatric species AND if mem- bers of geographically intermediate populations of either fail to exhibit intermediate character states for the differentiating characters. Use of the above criteria required the following as- sumption: that continuity or discontinuity in characters of external structure, internal geni- talic structure, geographical and habitat distri- bution, and life history provided by the study of population samples (comprised of individuals and groups of same) is evidence of reproductive continuity or isolation, respectively, among nat- ural populations. Clearly, recognition of allo- patric species is more arbitrary than distinguish- ing sympatric or parapatric species; but hypotheses about relationship must be drawn from available evidence, and some estimate of these relationships is preferable to none. I accept the subspecies definition of Edwards (1956b:230) in slightly altered form; namely, that subspecies are populations or groups of popu- lations whose members are recognizably differ- ent from members of other conspecific popula- tions and would interbreed with the latter 1F they occurred sympatrically and synchronously (un- der natural conditions) BUT are distinctly iso- lated from them during their mating periods. THE SUBSPECIES CATEGORY.—I frequently used the subspecies category in ranking allopat- ric Nebria populations and population groups because patterns of geographical variation are often such that discontinuities in structural and other characters correlate well with discontinu- ities in habitat or physiography (see also Ball 1966; and Ball and Negre 1972). Other workers (Erwin 1970; Madge 1967; Noonan 1973; and Whitehead 1976) have avoided use of the sub- species category in their studies of lowland, mainland groups. Recognition of the usefulness of a subspecies concept appears to depend on one’s particular perspective; more specifically, on the distribution of habitats or areas occupied by the organisms one studies. Where gaps be- tween areas of suitable habitat are broad and clear, such as with predominantly montane groups like Nebria, correlated discontinuity in variation is more confidently recognized and suggestive of active, effective barriers to gene flow. Because names for subspecies are accorded the same rights and strictures as species names and compete with the latter for priority under the present International Code of Zoological Nomenclature, the description of new subspe- cies (and resultant introduction of new names into the literature) must be justified. I suggest that recognition of subspecies as defined above is justified because they represent incipient species. They are at present independent evo- lutionary units whose members are already dif- ferentiated from members of other conspecific populations. Because these units are generally of zoogeographic and historical significance, the formal subspecies trinomen is as welcome a ‘‘shorthand notation’? (Larson 1975:252) as the species binomen or any other formal name. In addition to the question of usefulness is that of practicality, recognizing inherent difficulties in the formulation and application of a subspe- cies concept. Use of the subspecies category has been warmly debated for decades (see for ex- ample, Brown and Wilson 1954; Edwards 1954, 1956a, 1956b; Inger 1961; and Wilson and Brown 1953). Without question, great variation in sub- species concepts has existed among taxono- mists, so that taxa recognized by various work- ers are seldom equivalent or even comparable units. The inability of taxonomists to formulate and apply a common subspecies concept to their studies is unfortunate but understandable in light of evolutionary theory, which predicts a spec- trum of intermediate stages in the process of speciation. Assignment of formal subspecific rank to populations at one particular stage in the process is complicated by the potential for al- most infinite variety in evolutionary rate and direction of change. I agree with Edwards (1954, 1956a, 1956b) that the subspecies as defined above is a relatively objective (“‘comparatively but not absolutely nonarbitrary’’ [Simpson 1961:116]) category. Extensive barriers to gene flow between sub- species can, as suggested earlier, be recognized in some instances through detailed studies of habitat and geographical distributions and life histories. These may in fact be easier to dem- onstrate than intrinsic isolating mechanisms (i.e., reproductive isolation) between separate species. Arbitrariness arises in decisions on ‘*how different’> members of allopatric or al- lochronous populations must be for recognition as either distinct species or conspecific subspe- cies. Working criteria I used for recognizing sub- 94 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 4 species were as follows. Two populations or groups of populations were considered separate subspecies if (1) their respective geographical ranges are allopatric and separated by recogniz- able environmental barriers, (2) differences be- tween members of each are constant but less distinctive than differences between members of two closely related sympatric species in one or more characters of structure, color, or size, AND (3) the geographical pattern of variation in dis- tinguishing characters is nonclinal or distinctly step-clinal. In general, all members of a subspe- cies can be recognized on structural characters alone. However, in some instances, where two subspecies were recognized as distinct by a step- clinal pattern of variation, all members of one subspecies may be distinguished only from all members of the most proximate populations of the other. A good example of this type of vari- ation pattern is seen for Nebria lacustris Casey (see below; details to be presented elsewhere [Kavanaugh, manuscript in preparation]). The following assumption was required for use of the above criteria: that the amount of phenotypic divergence between members of two allopatric populations is a measure not only of the com- pleteness and duration of their extrinsic isolation but also of the probability that reproductive (in- trinsic) isolation has evolved between them. Where this assumption is invalid, the resultant error in ranking (i.e., recognizing as subspecies taxa which, in fact, conform to the biological species definition) is, in my opinion, less objec- tionable than its opposite. If two de facto species are ranked as subspecies, close relation- ship between the taxa is merely overempha- sized, and species-group names are still provid- ed and protected by priority. Only changes in status are required if actual relationships are correctly determined later. New NEArcTIC NEBRIA SPECIES AND SUBSPECIES The order of presentation of new taxa in this section follows a new classification of Nearctic Nebria to be presented elsewhere (Kavanaugh, manuscript in preparation). Nebria gouleti, new species (Figures 2, 13, 51) HOLOTYPE, a male, in CAS, labelled: ‘‘U.S., Wash., Asotin Co., Hwy. 129, 10 mi. sw. Antone, Rattlesnake Cr., 3000’, 25 August 1973 DHKavanaugh Family’’/ *‘D. H. Kavanaugh Col- lection’ [orange label]/ *‘Holotype Nebria gouleti Kavanaugh det. D. H. Kavanaugh 1976” [red label]/ ‘‘California Academy of Sciences Type No. 12504.’ ALLoTyYPE (same data as ho- lotype) also in CAS. In total, 514 PARATYPEs (227 males and 288 females) are deposited in the following collections: AMNH, ANSP, BFCa, CAS, CUIC, DHKa, DJLa, LRus, MCZ, MSU, OSUO, PURC, RTBe, SJSC, UASM, UIMI, UMMZ, USNM, UWBM, and WSU. Type-LocaLity.—Rattlesnake Creek (10 miles [ca. 16 km] sw of Antone), Asotin County, Washington. DIAGNOSTIC COMBINATION.—Head uniform- ly dark, without pale spot(s) on vertex, antennal scape moderately long, symmetrically ellipsoid (Fig. 2); pronotum distinctly cordate, midlateral seta present (Fig. 13), without longitudinal tu- bercle medial to midlateral seta; elytra dull, mi- crosculpture deeply impressed, meshes isodia- metric or very slightly transverse, fifth elytral interval without setae; metepisternum impunc- tate; hind coxa bi- or trisetose basally, tarso- meres of hind tarsus glabrous dorsally; second visible abdominal sternum with patch of setae medially (between hind coxae), third to fifth vis- ible sterna each with two to six pairs of posterior paramedial setae (Fig. 37). DERIVATION OF TAXON NAME.—I take great pleasure in naming this species in honor of my good friend and frequent field companion, Henri Goulet, from whom I have learned to observe and appreciate carabid beetles while they are still alive! GEOGRAPHICAL DISTRIBUTION.—Figure 51; restricted to portions of the Columbia Plateau region in Washington, Oregon, and Idaho; the lower Columbia River valley; and the mountains of northern and central Idaho. I have studied specimens from the following localities. United States of America IDAHO: Adams County, New Meadows ([{1,170 m]) [June] (1; UIMI); Bonner County, Sandpoint [Sep.] (1; CAS); Clear- water County, Canyon Ranger Station [Sep.] (1; UIMI); Cus- ter County, Lower Stanley (10 miles [ca. 16 km] E on Salmon River [1,800 m]) [Aug.] (1; DHKa), Salmon River (at Bayhorse Creek [1,620 m]) [Aug.] (45; DHKa); Idaho County, Fenn Ranger Station (Selway Fork Clearwater River) [Aug.] (3; UIMI), Lochsa River (at Canyon Creek [520 m]) [July] (2; DHKa), Lowell (39 miles [ca. 63 km] NE on Lochsa River [850 m]) [July] (1; DHKa), Meadow Creek (1.5 miles [ca. 2.4 km] s of Selway Falls) [Sep.] (1; CAS), Orogrande Creek [Aug.] (3; UWBM), Selway Falls [July] (1; UWBM): Kootenai Coun- ty, Coeur d’Alene [June—July] (5; CAS, MCZ, USNM), Hay- den Lake [Aug.] (9; AMNH, MCZ, UIMI, USNM); Latah County, Kendrick [Sep.] (2; UIMI), Moscow (1; USNM); Lemhi County, Salmon (and 21 miles [ca. 34 km] N) [July— Aug.] (2; UASM, UIMI); Nez Perce County, Lenore [May] (2; UIMI), Lewiston (and 2 and 3 miles [ca. 3.2 and 4.8 km] KAVANAUGH: NEW NEARCTIC NEBRIA E on Clearwater River [300 m—380 m]) [May—Aug., Oct.] (52; DHKa, UIMI), Waha [Aug.] (5; CAS, MCZ, PURC, USNM), Waha Lake [July] (4; CAS, MCZ); Shoshone County, Avery [Sep.] (2; UWBM), Clarkia [June] (1; UIMI), St. Joe River (17 miles [ca. 27 km] E of St. Maries) [July] (10; CAS); Valley County, Dagger Falls [July] (2; BFCa), Zena Creek [June] (1; UIMI); County unknown, Bungalow [Aug.] (15; UWBM). OREGON: Baker County, East Fork Eagle Creek [July] (2; UWBM), Halfway [July] (1; BFCa); Hood River County, Hood River [Sep.] (2; MCZ); Multnomah County, Portland [July] (3; CUIC); Umatilla County, Cottonwood Creek [June] (3; USNM, UWBM), Kamela [June] (1; CAS), Milton (Walla Walla River) [May-June] (4; UWBM), s of Kooskooskie (Washington) [June] (2; UWBM); Union County, Elgin (and 0.1 mile [0.16 km] s at Grande Ronde River [1,130 m]) [May, Aug.] (2; DHKa), Hilgard Junction State Park [Aug.] (3; RTBe), La Grande (1 mile [ca. 1.6 km] NE on Grande Ronde River [1,070 m]) [May] (5; DHKa); Wallowa County, Minam (0.2 miles [ca. 0.3 km] se at junction of Minam and Wallowa rivers [790 m]) [May] (5; DHKa). WASHINGTON: Asotin Coun- ty, Antone (10 miles [ca. 16 km] sw at Rattlesnake Creek [910 m]) [Aug.] (21; DHKa), Asotin (Asotin Creek [370 m]) [May— June] (18; ANSP, CAS, DHKa), Buford Creek (3.6 miles [ca. 5.8 km] s of Grande Ronde River [760 m]) [May] (8; DHKa), Grande Ronde River (at Highway 129 [550 m]) [May, Sep.] (2; DHKa, SJSC); Chelan County, Leavenworth [July] (20; ANSP, CAS, MCZ, PURC, USNM, UWBM), Peshastin Creek [July] (1; UASM), Tumwater Canyon ([610 m]) [May] (2; LRus); Columbia County, Dayton (Touchet River [580 m]) [May, July] (5; DKHa, WSU); Franklin County, Palouse Falls State Park [May] (2; LRus); Klickitat County, Trout Lake {July} (1; UWBM), White Salmon [Aug.] (2; UWBM); Ska- mania County, Underwood (15 miles [ca. 24 km] w on Colum- bia River [90 m]) [May] (4; DHKa); Spokane County, Spokane Falls (1; CAS); Walla Walla County, Burbank [May] (1; ANSP), Coppei [May] (1; UAFA), Coppei Creek [May] (1; UIMI), Kooskooskie [June, Aug.] (33; UIMI, UWBM), Touchet River (near Waitsburg) [July] (98; DJLa), Walla Wal- la (and 2 miles [ca. 3.2 km] s [320 m], Mill Creek) [May—Sep.] (139; CAS, CUIC, DHKa, MCZ, OSUO, PURC, UIMI, UMMZ, UWBM, WSU): Whitman County, Almota [May] (2; MCZ, MSU), Wawawai (2.8 miles [ca. 4.5 km] Nw at Snake River [300 m]) [May] (11; DHKa), Wilma [Apr.—May] (6: ANSP, UWBM). Doubtful Records CANADA—ALBERTA: Longview (Highwood River) [July] (7; CAS). UNirED STATES OF AMERICA—CALIFORNIA: (2; USNM). Nebria lacustris bellorum, new subspecies (Figures 14, 52) Nebria lacustris; BELL 1955:265 (in part). Ho.orype, a male, in CAS, labelled: “‘U.S., Tenn., Great Smoky Mts. N. P., Chimneys Picnic Area, W. Prong Little Pigeon R., 3000’, 24May73 DHKavanaugh & HGoulet’’/ **D. H. Kavanaugh Collection” [orange label]/ **Holotype Nebria lacustris bellorum Kavanaugh det. D. H. Kavanaugh 1976” [red label]/ ‘‘California Academy of Sciences Type No. 12506... ALLoTYPE (same data as holotype) also in CAS. In total, 276 PARATYPES (139 males and 137 females) are depos- ited in the following collections: CAS, CUIC, DHKa, HoKn, MCZ, RTBe, TCBa, UAFA, UASM, USNM. Type-LocaLity.—West Prong Little Pigeon River (at Chimneys Picnic Area), Great Smoky Mountains National Park, Tennessee. DIAGNOSTIC COMBINATION.—Head uniform- ly dark, without pale spot(s) on vertex; prono- tum (Fig. 14) distinctly cordate, midlateral seta present, without longitudinal tubercle medial to midlateral seta; elytra very shiny or faintly iri- descent, microsculpture consists of thin, trans- verse lines without (or with only scattered) meshes, striae very deeply impressed, intervals markedly convex, fifth interval without setae; metepisternum impunctate; tarsomeres of hind tarsus glabrous dorsally; second visible abdom- inal sternum glabrous medially, third to fifth vis- ible sterna each with single pair of posterior par- amedial setae; specimen from locality in southeastern United States, south of French Broad River (North Carolina or Tennessee) (Fig. 52): DERIVATION OF TAXON NAME.—I take plea- sure in naming this subspecies in honor of Ross and Joyce Bell (Burlington, Vermont), who have long been interested in the Nebria of eastern North America, and who have generously pro- vided encouragement and information for this study. GEOGRAPHICAL DISTRIBUTION.—Figure 52; restricted to that portion of the southern Appa- lachian region including Great Smoky Moun- tains National Park and adjacent mountain ranges. I have studied specimens from the fol- lowing localities. United States of America NortH CAROLINA: Great Smoky Mountains National Park [June] (1; MCZ), Indian Gap [Sep.] (1; USNM), Kephard Prong Oconaluftee River [Aug.] (14; DHKa, TCBa), New- found Gap ([{1,520 m—1,580 m]) [Aug.] (1; MCZ), Oconaluftee River (11 miles [ca. 18 km] se of Newfound Gap on Highway 441 [SSO m], at Smokemont [840 m] and 2.3 miles [ca. 3.7 km] Nw of Smokemont [610 m]) [May, Sep.] (61; DHKa, USNM), Smokemont Loop Trail ({760 m—910 m]) [July] (18; UAFA); Haywood County, Beach Gap (7 miles [ca. 11 km] s of Sun- burst at Middle Prong West Fork Pigeon River [1,520 m]) {[June, Aug.] (4; HoKn), Bubbling Spring Branch ([{1,520 m]) [July] (10; DHKa, TCBa), Graveyard Fields (Blue Ridge Park- way at Yellowstone Prong Pigeon River [1,540 m]) [May] (1; DHKa), Mount Sterling ({910 m]) [Oct.] (7; CAS, CUIC), Re- treat [May] (6; USNM):; Swain County, Bryson City (Deep Creek [610 m]) [Aug.] (2; MCZ). TENNESSEE: Great Smoky Mountains National Park, Alum Cave Creek ([{1,220 m—1,250 m]) [May] (4; DHKa), Chimneys Camp ((910 m]) [May, Sep.] (18; CAS, CUIC, USNM), Newfound Gap ({1,520 m—i,580 m]) [Aug.] (2; MCZ, UASM), Tennessee Branch Bradley Creek ([980 m]) [May] (15; USNM), Walker Prong Little Pi 96 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 4 geon River [Aug.] (1; DHKa), West Prong Little Pigeon River ({1,040 m—1,220 m]), Chimneys Picnic Area [910 m], (3 miles [ca. 4.8 km] s of Gatlinburg [520 m], 3.7 miles [ca. 6.0 km] N of Newfound Gap [1,220 m]) [May, July, Sep.] (109; DHKa, MCZ, UAFA, USNM); Sevier County, Gatlinburg [June] (1; DHKa). Doubtful Records UNITED STATES OF AMERICA—COLORADO: Jefferson County, Golden (Clear Creek) [June] (1; USNM). LoutsIAna: St. Tammany Parish, Highway 90 (10 miles [ca. 16 km] w of Mississippi border) [June] (1; CAS). Nebria nivalis gaspesiana, new subspecies (Figures 15, 53) Nebria suturalis; LENG (not LeConte). —LENG 1920:47 (in part). —LINDROTH 1955a:39. Nebria nivalis bifaria MANNERHEIM. —LINDROTH 1955a:39 (in part). Nebria nivalis; auctorum. —LAROCHELLE 1972:62 (in part); 1976:48; 1977:12 (in part). —LINDROTH 1961:81 (in part); 1971:1459. Nebria bifaria; LINDROTH 1961:81 (in part). HOLoryPe, a male, in CAS, labelled: *‘Can., Que., Penin- sule de Gaspe, Mt. Albert, Ruisseau du Diable, 3200’—3500', 11-12 June 73 DHKavanaugh & HGoulet’’/ *‘D. H. Kavan- augh Collection’’ [orange label]/ ‘Holotype Nebria nivalis gaspesiana Kavanaugh det. D. H. Kavanaugh 1976” [red la- bel]/ “‘California Academy of Sciences Type No. 12510.” AL- LOTYPE (same data as holotype) also in CAS. In total, 287 PARATYPES (119 males and 168 females) are deposited in the following collections: ALar, CAS, CNC, DBUM, DHKa, MCZ, RTBe, UASM, USNM. Type-LocALity.—Ruisseau du Diable (980 m—1,070 m), Mont Albert, Parc de La Gaspésie, Comté de Gaspeé-Ouest, Quebec. DIAGNOSTIC COMBINATION.—Head uniform- ly dark, without pale spot(s) on vertex; prono- tum (Fig. 15) distinctly cordate, midlateral seta present, with small longitudinal tubercle medial to midlateral seta; fifth elytral interval with one to four setae; metepisternum impunctate; tar- someres of hind tarsus glabrous dorsally; spec- imen from locality in Maine (Mount Katahdin), Gaspé Peninsula, Island of Newfoundland, or Labrador (west only to east shore of Ungava Bay) (Fig. 53). DERIVATION OF TAXON NAME.—This subspe- cies is named for the Gaspé Peninsula, on which the type-locality is found. GEOGRAPHICAL DISTRIBUTION.—Figure 53; disjunct; restricted to the western part of the Island of Newfoundland, Mount Katahdin (Maine), the high mountains of the Gaspé Pen- insula, and coastal Labrador (northwest to the eastern shore of Ungava Bay). I have studied specimens from the following localities. Canada NEWFOUNDLAND: Island of Newfoundland, Little Codroy River [July] (4; MCZ, USNM), St. John Bay (Doctors Brook) [July] (5; CNC, ZMLS), South Branch (Grand Codroy River) [July] (4; MCZ), Spruce Brook (1; MCZ). QUEBEC: Comté de Bonaventure [July] (1; ALar); Comte de Gaspé-Ouest, Parc de La Gaspésie (Mont Albert at Ruisseau du Diable [980 m— 1,070 m]) [June-July] (261; ALar, CNC, DBUM, DHKa, RTBe): Territoire du Nouveau-Québec, Riviere Abloviak [July] (14; DBUM). United States of America MAINE: Piscataquis County, Baxter State Park (Mount Ka- tahdin [910 m—1,580 m]) [June—Aug.] (10; CAS, MCZ, USNM). Nebria gyllenhali lassenensis, new subspecies (Figures 3, 54) HOoLotyPE, a male, in CAS, labelled: *‘U.S.A., Calif., Las- sen Volcanic N. P., Cascade Range, s. slope Mt. Lassen, Emerald Lk., 2450m, 22Aug.74 D.H.Kavanaugh Coll.’’/ **D. H. Kavanaugh Collection’ [orange label]/ *‘Holotype Nebria gyllenhali lassenensis Kavanaugh det. D. H. Kavanaugh 1976” [red label]/ “California Academy of Sciences Type No. 12511.’° ALLOTYPE (same data as holotype) also in CAS. In total, 34 PARATYPES (18 males and 16 females) are deposited in the following collections: CAS, DHKa, FMNH, USNM. Type-LocaLity.—Emerald Lake, Mount Lassen (south slope), Lassen Volcanic National Park, California. DIAGNOSTIC COMBINATION.—Head, prono- tum, and elytra without metallic reflection; head uniformly dark, without pale spot(s) on vertex, moderate in size and convexity, with genae and occiput not broadened, eyes full-sized; antennal scape (Fig. 3) short, straight, symmetrically ovoid or anteriorly more convex; penultimate labial palpomere trisetose; pronotum small, slightly narrow, moderately cordate (lateral margin moderately arcuate at middle), slight- ly dull to moderately shiny, microsculpture moderately to faintly impressed, meshes isodi- ametric, alveolae flat, midlateral seta present, without longitudinal tubercle medial to midlat- eral seta; elytral intervals flat, third interval con- tinuous or faintly catenate (setiferous pores slightly to moderately foveate), fifth interval without setae; hindwing full-sized; metepister- num impunctate; tarsomeres of hind tarsus gla- brous dorsally; second visible abdominal ster- num glabrous medially, third to fifth visible sterna each with single pair of posterior para- medial setae; specimen from locality in northern Sierra Nevada (California) or Cascade Range south of Columbia River (Fig. 54). DERIVATION OF TAXON NAME.—This subspe- KAVANAUGH: NEW NEARCTIC NEBRIA cies is named for Mount Lassen, on the south slope of which the type-locality is found. GEOGRAPHICAL DISTRIBUTION.—Figure 54; restricted to the southern Cascade Range (south of the lower Columbia River valley) and the northern Sierra Nevada (south to Sonora Pass). I have studied specimens from the following lo- calities. United States of America CALIFORNIA: Lassen Volcanic National Park, Emerald Lake ([2,450 m]) [Aug., Oct.] (6; CAS, DHKa, FMNH), Little Hot Springs Area (East Sulphur Creek [2,440 m-—2,530 m]) [Aug.] (6; CAS, DHKa), Mount Lassen (3 miles [ca. 4.8 km] se) [July] (1; USNM), Ridge Lake [Aug.] (16; CAS); Tuol- umne County, Sonora Pass [Aug.] (3; CAS). OREGON: Des- chutes County, Todd Lake (and | mile [ca. 1.6 km] s) [Aug.] (4; JSch). Nebria gyllenhali lindrothi, new subspecies (Figures 4, 54) Nebria sahlbergi: WICKHAM 1902:232. Nebria gyllenhali; auctorum. —ARMIN 1963:90. HOoLorypPe, a male, in CAS, labelled: **U.S., Wyo., Albany Co., Medicine Bow Mts., Snowy Range, Brooklyn Lk, 10500’, 11July72 DHKavanaugh’’/ *‘D. H. Kavanaugh Collection” {orange label]/ *‘Holotype Nebria gyllenhali lindrothi Kavan- augh det. D. H. Kavanaugh 1976” [red label]/ *‘California Academy of Sciences Type No. 12512... ALLOTYPE (same data as holotype) also in CAS. In total, 1579 PARATYPES (795 males and 784 females) are deposited in the following collec- tions: AMNH, ANSP, CArm, CAS, CDA, CNC, CUB, CUIC, DHKa, DRWh, EAMa, FMNH, ICCM, KSUC, MCZ, MSU, PADA, PMCh, PMNH, PURC, RCGr, RDav, RTBe, SDSU, UAFA, UASM, USNM, UWBM, UWL, ZMLS. TypeE-LocaLity.—Brooklyn Lake (3,200 m), Albany Coun- ty, Wyoming. NoTeES ON NOMENCLATURE AND TYPES.— This subspecies is the form noted by Lindroth (1969b:1109) as an undescribed species related to N. gyllenhali. DIAGNOSTIC COMBINATION.—Head, prono- tum, and elytra without metallic reflection; head uniformly dark, without pale spot(s) on vertex, moderate in size and convexity, with genae and occiput not broadened, eyes full-sized; antennal scape (Fig. 4) long, straight, slightly arcuate, moderately or markedly narrowed basally; pen- ultimate labial palpomere trisetose; pronotum small, slightly narrow, moderately cordate (lat- eral margin moderately arcuate at middle), slightly dull to moderately shiny, microsculpture moderately to faintly impressed, meshes isodi- ametric, alveolae flat, midlateral seta present, without longitudinal tubercle medial to midlat- 97 eral seta; elytral intervals flat, third interval con- tinuous or faintly catenate (setiferous pores slightly to moderately foveate), fifth interval without setae; hindwing reduced in size and ve- nation pattern; metepisternum impunctate; tar- someres of hind tarsus glabrous dorsally; second visible abdominal sternum glabrous medially, third to fifth visible sterna each with single pair of posterior paramedial setae; specimen from lo- cality in southern Rocky Mountains—in Medi- cine Bow Mountains or Sierra Madre (Wyo- ming), or Uinta Mountains (Utah), or further south (Fig. 54). DERIVATION OF TAXON NAME.—I am hon- ored to name this subspecies after Carl H. Lind- roth, a friend whose knowledge of and studies on Holarctic Carabidae, and Nebria in particu- lar, have been a guiding light in this study, and who first called my attention to the distinctive- ness of members of this taxon. I deeply regret that Carl did not live to see the completion of this study, to which he contributed so much. GEOGRAPHICAL DISTRIBUTION.—Figure 54; restricted to the southern Rocky Mountains, from northern New Mexico north to the Medi- cine Bow Mountains and Sierra Madre of south- ern Wyoming, west to the Uinta Mountains of eastern Utah. I have studied specimens from the following localities. United States of America CoLorRAbo: (2; KUSM, PURC); Boulder County, ([3,810 m]) [July] (1; CUB), Arapaho Pass ([2,740 m—3,350 m]) [Aug. ] (1; RTBe), Arapaho Peak ([3,510 m]) [Aug.] (3; CArm), Bea- ver Creek ({2,530 m]) [July] (1; CArm), Fourth of July Mines ({3,430 m]) [Aug.] (2; CArm), Jenny Lake ([{3,200 m]) [July] (2: CArm), Middle St. Vrain Creek ({3,600 m]) [July] (1; CArm), Niwot Ridge ([{3,050 m]) [July] (S; CArm, CUB), Rain- bow Lakes ([3,350 m]) [Aug.] (3: CNC), Rollins Pass ([(3,410 m-—3,540 m]) [Aug.] (2; CUB, DHKa), South Fork Middle Boulder Creek ({2,900 m—3,960 m]) [Aug.] (1; CArm); Clear Creek County, Leavenworth Valley (Argentine Road [3,660 m-—3,960 m]) (2; USNM), Loveland Pass ({3,660 m—3,840 m]) [July] (42; CAS, RTBe, UWBM), Mount Evans ([3,600 m— 4,330 m], Summit Lake [3,900 m—3,960 m]) [July—Aug.] (76; CAS, CNC, DHKa, FMNH, MCZ, PURC, RCGr, UASM); Conejos County, Cumbres Pass ([3,050 m]) [June] (1; CAS): Custer County, Ophir Creek (7.3 miles [ca. 11.7 km] sw of Fairview [3,140 m]) [June] (9; DHKa), St. Charles River (1.9 miles [ca. 3.1 km] sw of San Isabel [2,710 m—2,830 m]) [June] (14; DHKa), Wet Mountains ((3,470 m]) [July] (2; USNM), Dolores County, Lizard Head Pass ({3,050 m]) [Sep.] (1; CAS), Rico area ({2,590 m-3,050 m]) [July] (10; ANSP, MCZ, USNM);: El Paso County, Pikes Peak ({3,960 m]) [July] (8: AMNH, CAS), South Cheyenne Creek (at Gold Camp Road (2,530 m—2,650 m]) [June] (1; DHKa); Garfield County, Trap- pers Lake ([{2,530 m—2,650 m]) [June] (49; DHKa): Gilpin 98 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 4 County, Central City ({2,620 m]) [July] (1; CAS), Corona ({3,350 m]) [Sep.] (7; CAS, CUIC), South Boulder Creek (at Rollinsville and 2.5 miles [ca. 4.0 km] w [2,440 m]) [July— Aug.] (5; CArm, DHKa, UAFA); Grand County, Ptarmigan Peak (20; MCZ, ZMLS); Gunnison County, Gothic ({2,930 m]) [July] (2; USNM), Mexican Cut Lakes (1 mile [ca. 1.6 km] Nw of Elko Park [3,440 m]) [Aug.] (1; PMNH); Huerfano County, Apishapa Pass [June] (3; CUB), Bear Lake [June] (13; DHKa, EAMa), Blue Lake [June] (10; DHKa, EAMa), Cucharas Pass ({3,050 m] and 1.5 miles [ca. 2.4 km] N) [June] (109; DHKa, EAMa), Cucharas River (near Blue Lake [3,196 m]) [June] (5; DHKa); Jackson County, Cameron Pass ({3,140 m]) [Aug.] (1; UASM), Rabbit Ears Pass ({2,740 m]) [June— Aug.] (8: CAS, USNM); La Plata County, Hermosa Creek Trail (NW of Hermosa [2,960 m]) [Aug.] (9; UASM); Larimer County, Browns Lake Trail [Aug.] (9; RTBe), Crown Point [July] (21; RTBe), Zimmerman Lake [Aug.] (3; RTBe); Los Animas County, Apishapa Pass (2.1 miles [ca. 3.4 km] SE (3,230 m]) [June] (9; DHKa), Bear Creek (5 miles [ca. 8 km] s of Cucharas Pass [2,800 m]) [June] (83; DHKa); Mesa Coun- ty [Aug.] (5; PMCh, RDav), Grand Mesa ([3,050 m]) [July— Aug.] (6; CAS, CUB, USNM), Skyway ([3,050 m]) [June] (2; DHKa, FMNH):; Mineral County, Pagosa Springs (20 miles [ca. 32 km] N) [June] (1; DRWh), Wolf Creek Pass ({3,250 m— 3,660 m] and 2 miles [ca. 3.2 km] w, Thunder Mountain [3,570 m]) [June—Aug.] (96; CAS, CDA, CNC, DHKa, EAMa, UASM, USNM); Ouray County, American Flats ({3,660 m]) [July] (4; AMNH), Ouray area [June—July] (4; AMNH, CAS); Park County, Middle Fork North Platte River (near Hoosier Pass [3,540 m]) [July] (1; DHKa), Mount Lincoln ({3,350 m}) [Aug.] (1: UASM): Rio Grande County, Rio Grande River (1 mile [ca. 1.6 km] s of South Fork [2,400 m]) [Aug.] (29; DHKa, EAMa), Summitville ({3,050 m—3,350 m]) [Aug.] (7; CNC, MCZ); Rocky Mountain National Park ([{3,200 m—3,510 m]) [June—Aug.] (11; CAS, CNC, UASM), Fall River Pass ([3,600 m]) [July] (S$; CUB), Longs Peak ({3,840 m], Chasm Lake (3,810 m]) [July—Aug.] (14; CArm), Milner Pass ({3,280 m]) [July] (1; AMNH), Ouzel Falls ({3,050 m]) [Aug.] (1; CArm), Sandbeach Lake ([3,050 m]) [June] (7; CArm), Trail Ridge Road ({3,660 m—3,960 m]) [Aug.] (2; CUB, USNM); Routt County, Buffalo Pass (Summit Lake [3,140 m]) (4; CAS), Wal- ton Creek (above Dumont Lake [2,900 m—2,960 m]) [Aug.] (5; DHKa); San Juan County, Silverton ([(3,660 mJ) [July] (1; USNM); Summit County, Quandary Peak (south slope [3,410 m-—3,960 m]) [Aug.] (35; DHKa); Teller County ([3,410 m)]) {[June] (1; PADA), Phantom Canyon ([{2,960 m]) [June] (1; CAS); County unknown, “‘Hot Springs”’ (1; ICCM), **King’s Lake” ({3,350 m]) [Aug.] (2; CArm), La Plata Mountains ({3,200 m—3,660 m]) [July—Aug.] (4; DHKa, USNM), **Lara- mie Co.” (2; ICCM), Medicine Bow Range ({2,440 m-—3,660 m]) [July] (3; MCZ), Mount Baldy (summit [3,750 m]) [July] (1; CArm), San Juan Mountains ((3,660 m]) [July] (1; USNM), “foothills of Rocky Mts.’ (1: PMNH). New Mexico: (9; ANSP, ICCM, KUSM, MCZ, USNM); Mora County, Pecos Baldy (2; AMNH); Rio Arriba County, Gregorio Lake ({2,790 m]) [June] (S; DHKa), Rio Puerco (at Rio Puerco Campground (2,350 m—2,440 m]) [June] (3; DHKa); San Miguel County, Beaver Creek (Sw of Rincon Montoso [2,620 m]) [Aug.] (15; UASM), Cowles area ({2,440 m], Spirit Lake Trail) [June] (4; AMNH, UASM), Las Vegas area ([2,130 m]) [July] (1; KUSM), Willow Creek (near Torrero [2,740 m]) [June] (8; UASM); Santa Fe County, Lake Katherine ((3,600 m]) [June] (30; DHKa), Lake Peak (5; AMNH), Nambe Divide ({3,350 m]) [July] (2; ANSP), Santa Fe Ski Basin (5 miles [ca. 8 km] Nw) (1: AMNH); Taos County, Red River (2.2 miles [ca. 3.5 km] [2,610 m], 4.1 miles [ca. 6.6 km] [2,580 m], and 5 miles [ca. 8 km] w of Red River, 8 miles [ca. 13 km] s of Red River [2,870 m]) [June] (40; AMNH, DHKa), Tres Ritos [July] (4; CAS), Wheeler Peak ((3,050 m—3,640 m], Nw slope, Middle Fork Lake [3,340 m], Middle Fork Red River [3,260 m—3,690 m]) [June] (172; AMNH, DHKa). Utau: Duchesne County, Garfield Basin (stream below Tungsten Lake [3,450 m]) [Sep.] (3; USNM); Grand County, Mill Creek (at Oowah Lake [2,680 m]) [Aug.] (12; DHKa); San Juan County ({2,800 m]) [Aug.] (1; FMNH), Dark Canyon ([{2,960 m], NE slope [3,240 m]) {[Aug.] (30; UASM), Geyser Pass ([3,050 m], N slope [3,120 m], E slope [3,220 m], at Blue Lake [2,960 m]) [July—Aug.] (79; AMNH, UASM), Mill Creek Valley ({2,900 m]) [Aug.] (2; UASM). Wyominc: Albany County, Brooklyn Lake ({3,200 m]) [July] (41; DHKa), Centennial area ({3,050 m]) [Aug.] (22; MSU, SDSU), Douglas Creek (1 mile [ca. 1.6 km] sseE of Keystone [2,680 m]) [July] (32; DHKa), Little Brooklyn Lake ({3,120 m]) [July] (113; DHKa), Medicine Bow Peak ({3,350 m—3,660 m]) [July—Aug.] (3; FMNH, KSUC, UWL), Mirror Lake ({3,290 m]) [July] (37; USNM), Nash Fork Little Laramie River (7.5 miles [ca. 12.1 km] Nw of Centennial {2,990 m]) [July] (1; DHKa), Snowy Range Pass ([3,200 m]) [June, Aug.] (33; RTBe, UASM); Carbon County, Elk Moun- tain (35.0 miles [ca. 56.3 km] NE of Saratoga [3,400 m]) [July] (32; USNM), Hidden Treasure Gulch (11.5 miles [ca. 18.5 km] wsw of Encampment [2,870 m]) [Julyj (2; DHKa), Lake Marie ({3,230 m] and 8 miles [ca. 13 km] sw [2,740 m]) [July] (7; DHKa), North French Creek (4 miles [ca. 6.4 km] w of Mirror Lake [3,050 m]) [July] (3; DHKa), Silver Lake ({3,170 m]) [July] (32; USNM). Doubtful Records UNITED STATES OF AMERICA—MASSACHUSETTS: (1; UMRM). Nebria acuta quileute, new subspecies (Figures 16, 38, 46, 55) Nebria acuta LINDROTH 1961:71 (in part). HoLoryPe, a male, in CAS, labelled: “‘U.S., Wash., Clym- pic N. P., Olympic Mts., Boulder Cr. at Olympic Hot Springs, 2000’, 20July71 DHKavanaugh & EAMartinko’’/ “‘D. H. Ka- vanaugh Collection’’ [orange label]/ *‘Holotype Nebria acuta quileute Kavanaugh det. D. H. Kavanaugh 1976” [red label]/ “California Academy of Sciences Type No. 12494.” ALLo- TYPE (same data as holotype) also in CAS. In total, 135 ParRa- TYPES (81 males and 54 females) are deposited in the following collections: CAS, CNC, DHKa, LACM, MCZ, UASM, UIMI, USNM, UWBM. Type-Locatity.—Boulder Creek (at Olympic Hot Springs), Olympic National Park, Washington. DIAGNosTIC COMBINATION.—Pronotum (Fig. 16) moderately cordate, basal sinuation of lateral margin moderately short and deep, basal angle rectangular or slightly acute, midlateral seta ab- sent; elytra slightly shiny, microsculpture mod- erately impressed, meshes isodiametric (broken) or slightly transverse, elytral silhouette subrect- angular, not narrowed basally, humeral angle not rounded or obtuse, humeral carina markedly KAVANAUGH: NEW NEARCTIC NEBRIA developed and projected anterolaterally; hind coxa bi- or plurisetose basally; third to fifth vis- ible abdominal sterna each with two or more pairs of posterior paramedial setae; male with apex of median lobe as in Figure 38; female without anterodorsal sclerite in bursa copulatrix (Fig. 46); specimen from locality on Olympic Peninsula, Washington (Fig. 55). DERIVATION OF TAXON NAME.—This subspe- cies is named for the people native to the area including the type-locality, the Quileute tribe. GEOGRAPHICAL DISTRIBUTION.—Figure 55; restricted to the Olympic Peninsula, Washing- ton. I have studied specimens from the following localities. United States of America WASHINGTON: Olympic National Park [Aug.] (5; LACM, MCZ), Dosewallips River (at Mascott Campground) [July] (10; UWBM), Eagle Point ({1,830 m]) [July] (1; CNC), Olympic Hot Springs (Boulder Creek [610 m—780 m]) [May—Aug.] (120; CAS, CNC, DHKa, UIMI, USNM, UWBM), Sol Duc Hot Springs [June] (1; CAS). Nebria sahlbergii modoc, new subspecies (Figures 17, 56) HoLortyPE, a male, in CAS, labelled: **U.S., Calif., Modoc Co., Warner Mts., 4 mi. e. New Pine Creek, Pine Cr., 5700’, 22May71 DHKavanaugh & EAMartinko’’/ *‘D. H. Kavanaugh Collection’ [orange label]/ *‘Holotype Nebria sahlbergii mo- doc Kavanaugh det. D. H. Kavanaugh 1976” [red label]/ ‘‘California Academy of Sciences Type No. 12513... ALLo- TYPE (same data as holotype) also in CAS. In total, 15 PARA- TYPES (10 males and 5 females) are deposited in the following collections: CAS, DHKa, UASM, USNM. Type-LocaLity.—Pine Creek (4 miles [ca. 6.4 km] E of New Pine Creek), Modoc County, California. DIAGNOSTIC COMBINATION.—Pronotum (Fig. 17) size moderate, basal sinuation of lateral mar- gin long and deep, basal angles rectangular, not denticulate, lateral explanation broad through- out its length, midlateral seta absent; proepister- num faintly punctulate anteriorly; elytra slightly short and narrowed basally, elytral silhouette subrectangular, humeral angle not markedly rounded or obtuse, humeral carina moderately developed, slightly projected anterolaterally; specimen from locality in Warner Mountains (northeastern California or southcentral Oregon) (Fig. 56). DERIVATION OF TAXON NAME.—This subspe- cies is named for the county in which the type- locality is found, as well as for the people native to the region, the Modoc tribe. GEOGRAPHICAL DISTRIBUTION.—Figure 56; 99 restricted to the Warner Mountains of north- eastern California and southcentral Oregon. I have studied specimens from the following lo- calities. United States of America CALIFORNIA: Modoc County, Cedar Pass [June] (6; USNM), New Pine Creek (4 miles [ca. 6.4 km] £ at Pine Creek [1,740 m]) [May] (11; DHKa). Nebria sahlbergii triad, new subspecies (Figures 18, 56) HoLoryPe, a male, in CAS, labelled: **U.S.A., Calif., Trin- ity Co., Trinity Alps, S. F. Salmon R. at Big Flat Cmpgd., 1490m, 18July75 D.H. & B.A. Kavanaugh Stop #75-3677/ **D. H. Kavanaugh Collection” [orange label]/ *‘Holotype Nebria sahlbergii triad Kavanaugh det. D. H. Kavanaugh 1976” [red label]/ *‘California Academy of Sciences Type No. 12514.” ALLOTYPE (same data as holotype) also in CAS. In total, 212 PARATYPES (98 males and 114 females) are deposited in the following collections: CAS, DHKa, FMNH, MCZ, PUCA, UASM, USNM. Type-LocaLity.—South Fork Salmon River (at Big Flat Campground), Trinity County, California. DIAGNOSTIC COMBINATION.—Pronotum (Fig. 18) large, broad, basal sinuation of lateral margin moderate in length and depth, basal angles rect- angular, not denticulate, lateral explanation broad throughout its length, midlateral seta ab- sent; proepisternum smooth anteriorly: elytra moderate in length and basal width, elytral sil- houette subrectangular, humeral angle not markedly rounded or obtuse, humeral carina moderately developed, slightly projected an- terolaterally; specimen from locality in Klamath Mountains system of the Coast Ranges (north- western California or southwestern Oregon) (Fig. 56). DERIVATION OF TAXON NAME.—This subspe- cies is named for the Trinity Alps, in which the type-locality is found. I use a synonym of ‘trin- ity’ for this purpose. GEOGRAPHICAL DISTRIBUTION.—Figure 56; restricted to the Klamath Mountains system of northwestern California, probably also in south- western Oregon. I have studied specimens from the following localities. United States of America CALIFORNIA: Siskiyou County, Gulick Creek (at road to Josephine Lake [1,550 m]) [Aug.] (1; CAS); Trinity County, Boulder Creek (at Goldfield Campground [1,070 m]) [July] (56: CAS, DHKa), Coffee Creek (at Coffee Creek Ranch [1,070 m]) [July] (32; CAS, DHKa), Morris Meadows ([{1,340 m]) [Aug.] (1; PUCA), South Fork Salmon River (at Big Flat Campground [1,490 m]) [June—Aug.] (124: CAS, DHKa, FMNH, MCZ, UASM, USNM). 100 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 4 Nebria lituyae, new species (Figures 29, 57) Ho.ortyPe, a male, in CAS, labelled: **4 June 1977 3356’ under stones, summit Mt. Blunt, 2 mi. s. Lituya Bay, SE Alaska D. Mann coll. 102’’/ *‘Holotype Nebria lituyae Kavan- augh det. D. H. Kavanaugh 1979°° [red label]/ “‘California Academy of Sciences Type No. 13460.°° ALLOTYPE (same data as holotype) also in CAS. In total, 21 PARATYPEs (11 males and 10 females) are deposited in the following collec- tions: CAS, DHKa, DMan, UASM, USNM. Type-LocaLiry.—Mt. Blunt, 2 miles [ca. 3.2 km] s of Li- tuya Bay, Alaska. DIAGNOSTIC COMBINATION.—Head width moderate or slightly broadened, eyes moderate- ly convex, each epilobe of mentum with apical tooth short or moderately long; pronotum with lateral explanation broad throughout its length, basal angles rectangular, basal sinuation of lat- eral margin moderate in length and depth, mid- lateral seta absent; prosternal intercoxal process (Fig. 29) lanceolate, completely margined (even at apex); elytra slightly elongate, narrow, with or without faint metallic violet reflection, elytral silhouette subovoid, slightly to moderately nar- rowed basally, humeral angle not markedly rounded or obtuse, humeral carina slightly de- veloped and projected anterolaterally; hind coxa bi- or plurisetose basally; third to fifth visible abdominal sterna each with two or more pairs of posterior paramedial setae; specimen from lo- cality in southeastern Alaska (mainland) (Fig. Sys DERIVATION OF TAXON NAME.—This species is named for Lituya Bay, near which the type- locality is found. GEOGRAPHICAL DISTRIBUTION.—Figure 57; known only from southeastern Alaska, near Li- tuya Bay. I have studied specimens from the following locality. United States of America ALASKA: Mount Blunt (2 miles [ca. 3.2 km] s of Lituya Bay [720 m-—760 m]) [June] (23; CAS, DHKa, DMan, UASM, USNM). Nebria arkansana edwardsi, new subspecies (Figures 30, 33, 39, 47, 58) Nebria sahlbergi, auctorum.—EDWARDS 1975:48. —MANK 1934:74. Nebria arkansana; LINDROTH 1961:70 (in part). HOLoryPE, a male, in CAS, labelled: **U.S., Mont., Glacier N. P., Livingston Range, Logan Pass Area, ne. slope Mt. Clements, 7100’, 17July72 DHKavanaugh & JGEdwards’’/ “D. H. Kavanaugh Collection’? [orange label]/ *“Holotype Nebria arkansana edwardsi Kavanaugh det. D. H. Kavanaugh 1976”? [red label]/ *‘California Academy of Sciences Type No. 12495.°° ALLoTYPE (same data as holotype) also in CAS. In total, 1872 PARATYPES (1060 males and 812 females) are de- posited in the following collections: AMNH, ANSP, BFCa, CAS, CNC, CUIC, DBUM, DHKa, DJLa, FMNH, INHS, JSch, JVMa, KUSM, LRus, MCZ, MSU, OSUO, PMNH, PURC, ROM, RTBe, SJSC, UAFA, UASM, UBC, UIMI, USNM, UWBM. Type-LocaALity.—Logan Pass, Glacier National Park, Montana. NoTes ON NOMENCLATURE AND TyYPES.— Lindroth (1961:70) designated the valley of the Upper San Juan River, Archuleta County, Col- orado, as type-locality for N. arkansana Casey. As a result, the nominal subspecies of N. ar- kansana is one restricted to the southern Rocky Mountains (southcentral Wyoming and further south). The name presented here is for the widely distributed subspecies left unnamed by Lindroth’s choice of type-locality. DIAGNOSTIC COMBINATION.—Head moderate in size and relative width; pronotum with lateral explanation broad throughout its length, basal angles rectangular or slightly acute, basal sin- uation of lateral margin moderately deep, mid- lateral seta absent; elytra slightly to moderately dull, microsculpture with meshes isodiametric or slightly transverse, elytral silhouette rectan- gular or subrectangular, broad basally, basal margination (Fig. 30) straight or slightly con- cave, merged smoothly with lateral margination, humeral angle not markedly rounded or obtuse, humeral carina absent or very slightly devel- oped; hindwing with full reflexed apex distal to stigma (Fig. 33); hind coxa bi- or plurisetose ba- sally; third to fifth visible abdominal sterna each with two or more pairs of posterior paramedial setae; male with median lobe (Fig. 39) broad, with apex short and narrow; female with pos- terodorsal sclerite of bursa copulatrix as in Fig- ure 47; specimen from locality in Rocky Moun- tain region, but only north or west of Utah and southern Wyoming (Fig. 58). DERIVATION OF TAXON NAME.—I take great pleasure in naming this subspecies in honor of J. Gordon Edwards, whose keen interest in life at high altitude has fostered my own. GEOGRAPHICAL DISTRIBUTION.—Figure 58; widespread, distributed in Rocky Mountain re- gion from southern Yukon Territory south to northeastern Nevada, Idaho, and northern Wy- oming. I have studied specimens from the fol- lowing localities. KAVANAUGH: NEW NEARCTIC NEBRIA Canada ALBERTA: Banff National Park, Banff [Aug.—Sep.] (3: CUIC, UASM), Boom Lake ([1,830 m]) [July] (1; CNC), Bow Lake [Aug.] (1; UASM), Bow River ([1,830 m]) [Aug.] (6; CNC), Johnson Creek Trail (at Johnson Creek Campground) [July] (1; ROM), Lake Agnes [Aug.] (1; PMNH), Lake Louise [June—Sep.] (16; AMNH, CAS, CNC, MCZ, UBC), Ptarmigan Mountain ({[2,590 m]) [Aug.] (13; CNC), Spray Lake [July] (1; JVMa), Sunset Pass ([{1,950 m]) [Aug.] (4; CNC); Jasper Na- tional Park [Aug.] (2; AMNH), Amethyst Lake (and Ame- thyst Lake Trail [1,710 m]) [July] (10; DHKa, UASM), Angel Glacier [Sep.] (1; RTBe), Hardisty Creek (1 mile [ca. 1.6 km] s on Highway 93) [July] (1; RFre), Jasper [Aug.] (3; USNM), Miette Hot Springs [Aug.] (1; RTBe), Miette River (at Mead- ow Creed [1,100 m]) [July—Aug.] (22; DHKa, UASM), Mount Edith Cavell ({1,360 m] and below Angel Glacier) [July] (3; DHKa, UAFA), Portal Creek Trail ({1,950 m]) [July] (3: USNM); Waterton Lakes National Park [July—Aug.] (5; UASM, UWBM), Cameron Lake (creek below lake) [July] (7; USNM), Red Rock Canyon [July] (2; CAS), Waterton Park (Cameron Creek) [July] (36; CNC, MCZ, UASM): Other lo- calities, Bellevue (2 miles [ca. 3.2 km] w at Gold Creek [1,370 m]) [July] (1; DHKa), Blackstone Gap [Aug.] (2; CNC), Blair- more (1; USNM), Blakiston Brook [July] (10; UASM), Car- bondale River (near Lost Creek) [Sep.] (1; DJLa), Cline River (at Highway 11) [Aug.] (2; DHKa), Gap [Aug.] (1; CAS), Gorge Creek [July] (2; UASM), Happy Valley (Pincher Creek) [Aug.] (11; CAS, UASM), Highwood Pass ([2,380 m]) [July] (15; CNC), Highwood River [Aug.] (1; CAS), Marmot Creek (10 miles [ca. 16 km] sw of Kananaskis Forest Experiment Station [1,520 m]) [Aug.] (10; CNC), Mill Creek Ranger Sta- tion [Aug.] (2; CAS), Pincher Creek ([{1,620 m]) [July] (3: DJLa), Ram River area [July] (2; BFCa), Sheep River (4.5 miles [ca. 7.2 km] above Gorge Creek) [Aug.] (7; DHKa), West Branch Castle River ([{1,340 m]) [Aug.] (6; DJLa). Brit- IsH CoL_uMBIA: Glacier National Park, Asulkan Glacier area [Aug.] (1; CNC), Glacier [July—Aug.] (6; CAS, CNC, PURC), Rogers Pass ({1,370 m]) [Aug.] (2; ANSP, CAS): Kootenay National Park [Aug.] (3; CNC), Kimpton Creek ([{1,220 m]) [July] (2; CNC), Kindersley Pass ({2,130 m]) [July] (4; CNC), Kootenay River (at Highway 1B) [Aug.] (1; UASM), Marble Canyon [July] (1; CNC), Sinclair Creek ([1,830 m] and at Highway 13) [June—Aug.] (25; CAS, CNC, UASM, UBC), Vermillion Creek [July] (4: CAS, MSU, UBC); Yoho National Park [July] (3; CAS), Amiskwi River (and at Amiskwi Falls [1,830 m]) [Aug.] (34; CNC), Field (and 3 miles [ca. 4.8 km] E on Kicking Horse River) [June, Aug.] (4: CAS, USNM), Lake Oesa ([2,130 m]) [July] (18; CNC), Lake O'Hara [July] (10; CAS, CNC, UBC), Otterhead River ({1,680 m]) [Aug.— Sep.] (29; CNC, DHKa), Yoho River ({1,400 m], 4 to 12 miles [ca. 6.4 toca. 19 km] £ of Field, at Takakkaw Falls [1,520 m]) [July—Aug.] (25; CNC, DHKa, UASM); Other localities, Bo- sewell [Aug.] (1; UASM), Cabin Creek (Ne of Fernie, near Michel) [July] (13; CNC, UASM), Creston (8 miles [ca. 13 km] w) [June] (3; CNC), Crowsnest Pass (2 miles [ca. 3.2 km] w [1,280 m] and 5 miles [ca. 8 km] w on Elk River [1,220 m]) [July] (S51; DHKa), Edgewood [July—Aug.] (3: CAS), Elk Creek (£ of Fernie) [July] (4; UASM), Elk River (10.7 miles {[ca. 17.2 km] w of Fernie) [July] (3; ROM), Erickson (Arrow Creek) [Aug.] (12; CAS, UBC), Fernie (Coal Creek [1,680 m], Fairy Creek, Lizard Creek) [July—Aug.] (63; BFCa, CAS, CNC, DBUM, FMNH, MSU, UBC), Flathead [June] (1; CUIC), Fort Steele [June, Sep.] (2; CAS, UBC), Fortysixmile Camp (at Hope Trail) [July] (4; CAS, CNC, UBC), Hope 101 Mountains [July—Sep.] (6; CNC, INHS, MCZ), Hosmer (Elk River) [July] (1; CAS), Lorna [June] (2: CAS, UBC), Manning Provincial Park (Nicomen Ridge) [July] (10; CAS, CNC, MSU, UBC), Michel (5; UASM), Mount Copeland ([1,980 m]) [Aug.] (21; CNC), Radium Hot Springs [Aug.] (6; CAS, CNC, UBC), Slocan [Aug.] (2; BFCa), South Fork Creek [Aug.] (1; UWBM), Summit Creek (near Creston) [Aug.] (5; UASM), Tetsa River (Alaska Highway mile 378 and mile 390) [Aug.] (3; DHKa, MSU), Vernon [July] (1; CAS), Whipsaw Creek (at Hope Trail [1,460 m]) [July] (1; UBC), Wyndel [Sep.] (1; CNC). YUKON Territory: Rancheria (Swift River) [Aug.] (6: CNC, UASM). United States of America IDAHO: Blaine County, Alturas Lake (at Alturas Inlet Campground [2,130 m]) [Aug.] (11; DHKa), Cherry Creek (0.1 miles [ca. 0.2 km] Nw of Highway 93 [2,230 m]) [Aug.] (6; DHKa), Dollarhide Summit (2 miles [ca. 3.2 km] £ at North Fork Warm Springs Creek [2,350 m—2,410 m]) [Aug.] (9; DHKa), Galena [July] (1: BFCa), Galena Summit (2.4 miles {[ca. 3.9 km] NE [2,320 m]) [Aug.] (12; DHKa), Hyndman Creek [July] (22; UWBM), Ketchum (2 miles [ca. 3.2 km] NNW at Big Wood Creek [1,830 m]) [Aug.] (28; DHKa), North Fork Warm Springs Creek (10 miles [ca. 16 km] w of Ketchum [1,890 m]) [Aug.] (92; DHKa), Sun Valley [Sep.] (2; JSch): Camas County, Carrie Creek (32 miles [ca. 51 km] [2,440 m] and 36 miles [ca. 58 km] [2,100 m] Ese of Ketchum) [Aug.] (106; DHKa), South Fork Boise River (14 miles [ca. 23 km] E of Featherville at Skeleton Creek [1.550 m], 34 miles [ca. 55 km] £ of Featherville at Bear Creek [1,830 m]) [Aug.] (157: DHKa); Cassia County, Rock Creek (15 miles [ca. 24 km] s of Hansen, at Rock Creek Ranger Station) [June-July] (6: UIMI): Custer County, Bayhorse Creek (7 miles [ca. 11 km] Nw of Highway 93 [2,440 m]) [Aug.] (3: DHKa), Fourth of July Creek (3 miles [ca. 4.8 km] © of Highway 93 [2,230 m]) [Aug.] (S8; DHKa), Stanley [July] (1; UWBM), Trail Creek Summit (2 miles [ca. 3.2 km] NE at Summit Creek [2,320 m]) {Aug.] (9; DHKa); Fremont County, Island Park [July] (3; BFCa): Shoshone County, Avery (7 miles [ca. 11 km] £) [July] (1; UIMI); Valley County, Dagger Falls [July] (2; BFCa), Middle Fork Salmon River (near Boundary Creek) [Aug.] (7; RTBe); County unknown, Twin Creek Forest Camp ([{1,520 m]) [July] (S57; UWBM). MoNTANA: Carbon County, Quad Creek (17 miles [ca. 27 km] sw of Red Lodge [3,050 m]) [July] (66: DHKa): Fergus County, East Fork Rock Creek ([{1.710 m]) [July] (42; DHKa); Flathead County, Marias Pass (10 miles [ca. 16 km] w of summit [1,370 m]) [July] (11; LRus), South Fork Flathead River (at Hungry Horse Reservoir [1,220 m]) [July] (3; LRus); Gallatin County, Fairy Lakes [Aug.] (1; CUIC); Glacier National Park [July—Aug.] (6; CUIC), Baring Creek (at St. Mary Lake) [Aug.] (3; UWBM), Grinnel Falls ({1,860 m]) [Aug.] (1; SJSC), Grinnel Glacier ([{1,860 m—1,890 m]) [Aug.] (21; DHKa, SJSC, USNM), Grinnel Lake (at inlet {1,540 m]) [Aug.] (1; DHKa), Lake McDonald (and at Sprague Creek Campground) [July—Aug.] (4; CAS, RTBe), Lake Sher- burne (at Canyon Creek) [July] (1; CAS), Logan Pass ({2,030 m—2,130 m] and 5 miles [ca. 8 km] w [1,770 m]) [July—Sep.] (16; DHKa, JSch, RTBe, UASM), Morning Eagle Falls ({ 1.680 m]) [July] (1; SJSC), Mount Clements (NE slope [2,160 m}) [July] (S; DHKa), Piegan Pass ({2,410 m]) [Aug.] (1; SJSC). St. Mary [Aug.] (8: SJSC, UASM); Madison County, Beaver Creek ([{1,920 m]) [Aug.] (1: KUSM), Ennis [July] (2; UIMI): Mineral County, Haugan [Aug.] (14; KUSM), Lookout Pass ({1,280 m]) [July] (1; LRus), Saltese [July, Sep.] (3; MSU, 102 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 4 SJSC); Sweetgrass County, Big Timber Creek (at Half Moon Campground [2,230 m—2,290 m]) [July] (1; DHKa). NeEvapa: Elko County, Lamoille Creek (at Lower Lamoille Camp- ground [1,920 m]) [Aug.] (35; DHKa), Thomas Creek (at Thomas Creek Campground [2,320 m—2,380 m]) [Aug.] (130; DHKa). OREGON: Baker County, Cornucopia [June—July] (35; BFCa, UWBM); Union County, Elgin (and 9 miles [ca. 14 km] Nw at Little Phillips Creek [1,300 m]) [May, Aug.] (6: DHKa), Langdon Lake area (se of summit) [Aug.] (2; UWBM), Tollgate-Elgin Road [June] (3; UWBM); Wallowa County [Aug.] (1; JSch), Aneroid Lake ({2,320 m]) [July] (2; FMNH), East Fork Eagle Creek [July] (9; UWBM), Indian Crossing (Imnaha River) [Aug.] (2; MSU, OSUO), Lostine River ({1,310 m], at French Campground, at Two Pan Camp- ground [1,710 m—1,980 m]) [Aug.] (112; DHKa, USNM, UWBM), Lost Lake [Aug.] (1; UWBM), Wallowa Lake [July— Aug.] (10; DHKa, UWBM), Wallowa River (above inlet to Wallowa Lake) [May] (1; DHKa). WASHINGTON: Walla Walla County, Walla Walla [Aug.] (1; UWBM). Wyominc: Big Horn County, Granite Pass (8 miles [ca. 13 km] sw on Granite Creek [2,380 m]) [July] (5: DHKa), Shell Creek (11 miles [ca. 18 km] sw of Granite Pass [2,130 m]) [July] (14; DHKa), West Tensleep Creek ({3,140 m—3,200 m]) [July] (17; DHKa); John- son County, Clear Creek (12 miles [ca. 19 km] w of Buffalo {2,190 m]) [July] (1; DHKa), South Fork Clear Creek (16 miles [ca. 26 km] w of Buffalo [2,350 m], at South Fork Campground (2,350 m]) [July—Aug.] (6; DHKa, FMNH); Park County, Fox Creek (at Highway 212) [Aug.] (1; CAS); Sheridan County, Little Tongue River (13 miles [ca. 21 km] wsw of Dayton (2,380 m]) [July] (22; DHKa); Washakie County, Tensleep (6 miles [ca. 10 km] [1,550 m] and 11 miles [ca. 18 km] [1,890 m] NE on Tensleep Creek) [July] (4; DHKa); Yellowstone Na- tional Park [Aug.] (25; USNM); County unknown, Bighorn Reserve (1; MCZ). Nebria arkansana oowah, new subspecies (Figures 40, 48, 58) HOLoryPe, a male, in CAS, labelled: “‘U.S., Utah, Grand Co., LaSal Mts., Mill Cr. at Oowah Lk., 8800’, 2August71 DHKavanaugh & EAMartinko’’/ *‘D. H. Kavanaugh Collec- tion” [orange label]/ ‘‘ Holotype Nebria arkansana oowah Ka- vanaugh det. D. H. Kavanaugh 1976” [red label]/ “California Academy of Sciences Type No. 12496.’> ALLoOTYPE (same data as holotype) also in CAS. In total, 75 PARATYPES (54 males and 21 females) are deposited in the following collec- tions: CAS, DHKa, MCZ, UASM, USNM. Type-LocaLity.—Mill Creek (at Oowah Lake), Grand County, Utah. DIAGNOSTIC COMBINATION.—Head moderate in size and relative width; pronotum with lateral explanation broad throughout its length, basal angles rectangular or slightly acute, basal sin- uation of lateral margin moderately deep, mid- lateral seta absent; elytra with humeral angle not markedly rounded or obtuse, humeral carina moderately developed, elytral silhouette sub- rectangular, broad basally, hindwing full-sized; hind coxa bi- or plurisetose basally; third to fifth visible abdominal sterna each with two or more pairs of posterior paramedial setae; male with median lobe broad, apex as in Figure 40; female with posterodorsal sclerite of bursa copulatrix as in Figure 48; specimen from locality in La Sal Mountains (eastern Utah) (Fig. 58). DERIVATION OF TAXON NAME.—This subspe- cies is named for the small, sheltered lake at the type-locality. GEOGRAPHICAL DISTRIBUTION.—Figure 58: restricted to the La Sal Mountains of eastern Utah. I have studied specimens from the follow- ing localities. United States of America Urau: Grand County, Mill Creek (at Oowah Lake [2,680 m]) [Aug.] (74; CAS, DHKa, MCZ, USNM); San Juan Coun- ty, Mill Creek Valley ({2,900 m]) [Aug.] (3; UASM). Nebria arkansana uinta, new subspecies (Figures 35, 41, 58) HOLoryPE, a male, in CAS, labelled: *‘U.S., Utah, Summit Co., Uinta Mts., Hwy. 150, Lost Lk., 9800’, 14Aug71 DHKa- vanaugh’’/ *“‘D. H. Kavanaugh Collection” [orange label]/ “Holotype Nebria arkansana uinta Kavanaugh det. D. H. Kavanaugh 1976” [red label]/ *‘California Academy of Sci- ences Type No. 12497.”’ ALLoTYPE (same data as holotype) also in CAS. In total, 93 PARATYPES (44 males and 49 females) are deposited in the following collections: CAS, DHKa, MCZ, UASM, USNM. Type-Locatity.—Lost Lake, Summit County, Utah. DIAGNOSTIC COMBINATION.—Head moderate in size and relative width; pronotum with lateral explanation broad throughout its length, basal angles rectangular or slightly acute, basal sin- uation of lateral margin moderately deep, mid- lateral seta absent; elytra with humeral angle not markedly rounded or obtuse, humeral carina moderately developed; elytral silhouette sub- rectangular or subovoid, slightly narrowed ba- sally; hindwing reduced in length and width (Fig. 35); hind coxa bi- or plurisetose basally; third to fifth visible abdominal sterna each with two or more pairs of posterior paramedial setae; male with median lobe as in Figure 41; specimen from locality in Uinta Mountains or in Wasatch Mountains at or north of Weber River, Utah (Fig. 58). DERIVATION OF TAXON NAME.—This subspe- cies is named for the Uinta Mountains, in which the type-locality is found. GEOGRAPHICAL DISTRIBUTION.—Figure 58; restricted to the Uinta Mountains and the Wa- satch Mountains at or north of Weber River (northern Utah). I have studied specimens from the following localities. KAVANAUGH: NEW NEARCTIC NEBRIA United States of America UrAu: Cache County, Logan River (13.7 miles [ca. 22 km] E of Logan at Cottonwood Creek [1,740 m]) [Aug.] (65; CAS, DHKa, MCZ, UASM): Duchesne County, Painter Basin ({3,470 m]) [Sep.] (1; USNM); Summit County, Lost Lake ({2,990 m]) [Aug.] (7; DHKa), Upper Provo River (20.8 miles [ca. 33 km] £ of Kamas [2,600 m], at Highway 150) [Aug.] (6: DHKa, USNM); Uintah County, Big Brush Creek (7 miles [ca. 11 km] wnw of Highway 44 [2,620 m]) [Aug.] (14; DHKa); Weber County, Ogden (6 miles [ca. 10 km] E on Og- den River [1,490 m]) [Aug.] (2; DHKa). Nebria fragilis teewinot, new subspecies (Figures 35, 42, 59) HoLortyPe, a male, in CAS, labelled: “‘U.S., Wyo., Teton N. P., Teton Mts., Mt. Teewinot, stream on se. slope, 7100'— 9000’, 3Aug.73 DHKavanaugh Family’’/ *‘D. H. Kavanaugh Collection’’ [orange label]/ ‘‘ Holotype Nebria fragilis teewinot Kavanaugh det. D. H. Kavanaugh 1976” [red label]/ **Cali- fornia Academy of Sciences Type No. 12500." ALLOTYPE (same data as holotype) also in CAS. In total, 377 PARATYPES (219 males and 158 females) are deposited in the following collections: CAS, DHKa, KUSM, MCZ, SJSC, UASM, USNM. Type-LocALity.—Mount Teewinot (southeast slope), Grand Teton National Park, Wyoming. DIAGNOSTIC COMBINATION.—Head moderate in size and relative width; pronotum with lateral explanation broad throughout its length, basal angles rectangular or slightly acute, basal sin- uation of lateral margin moderately deep, mid- lateral seta absent; elytra slightly shiny, micro- sculpture faintly impressed, with meshes irregularly isodiametric or slightly transverse, with or without faint metallic green or violet re- flection, elytral silhouette subrectangular or sub- ovoid, slightly narrowed basally, humeral angle not markedly rounded or obtuse, humeral carina moderately developed; hindwing reduced in length and width (Fig. 35); hind coxa bi- or plu- risetose basally; third to fifth visible abdominal sterna each with two or more pairs of posterior paramedial setae; male with median lobe as in Figure 42; specimen from locality in westcentral Wyoming (Fig. 59). DERIVATION OF TAXON NAME.—This subspe- cies is named for the type-locality. GEOGRAPHICAL DISTRIBUTION.—Figure 59; restricted to mountain ranges of the Rocky Mountain system in westcentral Wyoming. I have studied specimens from the following lo- calities. United States of America Wyomina: Grand Teton National Park, Mica Lake ([2,970 m]) [July] (1; DHKa), Mount Teewinot (se slope [2,160 m— 2,740 m]) [Aug.] (79; DHKa), South Fork Cascade Canyon 103 ((3,080 mJ) [July] (1; SJSC), South Fork Garnet Canyon ({2,820 m]) [July] (1; SJSC), Timberline Lake area [July] (1; DHKa); Lincoln County, White Creek (20 miles [ca. 32 km] se of Alpine Junction [1,950 m—2,190 m]) [Aug.] (40; DHKa), Wolf Creek (16.1 miles [ca. 26 km] sw of Hoback Junction [1,770 m—-1,780 m]) [July—Aug.] (137; CAS, DHKa, MCZ, UASM, USNM); Sublette County, Green River Lakes (west shores [2,440 m—2,590 m]) [July] ($3; DHKa), Hoback River (14 miles [ca. 23 km] sw of Highway 187/189 [2,440 m}) [July] (11; DHKa); Teton County, Alaska Basin (near Sunset Lake [2,940 m]) [July] (1; SJSC), Granite Creek (at Granite Falls {2,160 m]) [July] (37; DHKa), Togwotee Pass ({2,960 m] and 3.7 miles [ca. 6.0 km] wNw at Blackrock Creek [2,770 m]) [Aug.] (20; DHKa, KUSM). Nebria zioni oasis, new subspecies (Figures 5, 60) HoLorypPe, a male, in CAS, labelled: **U.S., Utah, Wash- ington Co., Pine Valley Mts., Leeds Cr. at Oak Grove Cmpegd., 6300'-6500', 20June71 DHKavanaugh & EAMartin- ko’’/ **D. H. Kavanaugh Collection” [orange label]/ *‘Holo- type Nebria zioni oasis Kavanaugh det. D. H. Kavanaugh 1976” [red label]/ **California Academy of Sciences Type No. 12518... ALLOTYPE (same data as holotype) also in CAS. In total, 46 PARATYPES (18 males and 28 females) are deposited in the following collections: CAS, DHKa, UASM, USNM. Type-LocaLity.—Leeds Creek at Oak Grove Camp- ground, Washington County, Utah. DIAGNOSTIC COMBINATION.—Body and ap- pendages dark (piceus or black); antennal scape (Fig. 5) medium length, straight, almost cylin- drical (slightly narrowed basally); each epilobe of mentum with tooth moderately long; prono- tum with lateral explanation moderately broad throughout its length, basal angles rectangular or slightly acute and projected posteriorly, lat- eral margin without subapical sinuation but with basal sinuation moderately long and deep, mid- lateral seta absent; elytra not vaulted, slightly flattened, elytral silhouette ovoid, markedly nar- rowed basally, humeral angle markedly rounded and obtuse, humeral carina markedly devel- oped, fifth and seventh elytral intervals without setiferous pores; hind coxa bi- or plurisetose basally; third to fifth visible abdominal sterna each with two or more pairs of posterior para- medial setae; specimen from locality in Pine Valley Mountains (Utah) (Fig. 60). DERIVATION OF TAXON NAME.—This species is named for the oasislike nature of the type- locality—a spot of green, as it were, in an oth- erwise dry land. GEOGRAPHICAL DISTRIBUTION.—Figure 60; restricted to the Pine Valley Mountains of south- western Utah. I have studied specimens from the following locality. 104 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 4 United States of America UTAH: Washington County, Leeds Creek (at Oak Grove Campground [1,920 m—1,980 m]) [June] (46; CAS, DHKa, UASM, USNM). Nebria obliqua chuskae, new subspecies (Figures 19, 34, 61) HoLortypPe, a male, in CAS, labelled: *“‘USA Arizona Apache Co., sw. Red Rock Chuska Mts. Wagon Wheel Camp Gr. aspen-fir forest 2260m. Loc. 11 July 11-12, 1976°7/ ‘““SOUTHWESTERN USA EXP.—1976 G.E.Ball, J.M. Campbell, P.M.Hammond’’/ *‘Holotype Nebria obliqua chus- kae Kavanaugh det. D. H. Kavanaugh 1979’ [red label]/ “California Academy of Sciences Type No. 13461.’’ ALLo- TYPE (same data as holotype) also in CAS. In total, 16 PARA- TYPES (10 males and 6 females) are deposited in the following collections: CAS, MCZ, UASM, USNM. Type-LocaLity.—Lukachukai Creek (at Wagon Wheel Campground), Apache County, Arizona. DIAGNOSTIC COMBINATION.—Mandible markedly explanate laterally, broad; tooth of mentum bifid, with notch V-shaped; pronotum (Fig. 19) relatively long and narrow, moderately cordate, moderately narrowed basally, widest anterior to middle, slightly shiny (microsculp- ture faintly or moderately impressed, meshes isodiametric), basal margin not or only slightly sinuate laterally, lateral bead narrow, obliterat- ed basally in most specimens, basal foveae broad, smooth or faintly rugulose or punctulate, basal angles obtuse, basal sinuation of lateral margin absent, lateral explanation moderately wide and more broadened posteriorly only, mid- lateral seta absent; elytral silhouette subovoid, slightly narrowed basally, lateral margins slight- ly rounded; hindwing (Fig. 34) reduced in length; hind coxa bi- or plurisetose basally; third to fifth visible abdominal sterna each with two or more pairs of posterior paramedial setae; specimen from locality in Chuska Mountains, Arizona (Fig. 61). DERIVATION OF TAXON NAME.—This subspe- cies is named for the Chuska Mountains, in which the type-locality is found. GEOGRAPHICAL DISTRIBUTION.—Figure 61; restricted to the Chuska Mountains, Arizona. I have studied specimens from the following lo- cality. United States of America ARIZONA: Apache County, Lukachukai Creek (at Wagon Wheel Campground [2,260 m—2,320 m]) [July—Aug.] (18; CAS, MCZ, UASM, USNM). Nebria darlingtoni, new species (Figures 20, 62) HOLOTYPE, a male, in CAS, labelled: ‘‘U.S.A., Calif., El Dorado Co., Sierra Nevada, S. F. American River., 3 mi. w. of Riverton, 910m, 16July75 D. H. & B.A. Kavanaugh Stop #75-29"'/ **D. H. Kavanaugh Collection’? [orange label]/ ‘‘Holotype Nebria darlingtoni Kavanaugh det. D. H. Kavan- augh 1976” [red label]/ **California Academy of Sciences Type No. 12499.’ ALLOTYPE (same data as holotype) also in CAS. In total, 170 PARATYPES (105 males and 65 females) are de- posited in the following collections: AMNH, CAS, DHKa, ICCM, KSUC, KUSM, MCZ, UASM, USNM. Type-LocaALitry.—South Fork American River (3 miles [ca. 4.8 km] w of Riverton), El] Dorado County, California. DIAGNOSTIC COMBINATION.—Body black, legs dark brown or piceus; microsculpture on pronotum and elytra very deeply impressed, meshes isodiametric, alveolae convex; prono- tum (Fig. 20) with lateral explanation narrow at middle, broadened posteriorly, basal angles rectangular or slightly acute, basal sinuation of lateral margin moderately short and deep, mid- lateral seta absent; elytra with humeral carina slightly developed, slightly projected anteriorly, elytral silhouette ovoid, markedly narrowed ba- sally, humeral angles moderately rounded and obtuse; hind coxa bi- or plurisetose basally; third to fifth visible abdominal sterna each with two or more pairs of posterior paramedial setae; specimen from Sierra Nevada, California (Fig. 62). DERIVATION OF TAXON NAME.—I take great pleasure in naming this species in honor of Philip J. Darlington, Jr., biogeographer, student of carabid evolution, and friend, whose work and personal encouragement have provided inspira- tion in this study. GEOGRAPHICAL DISTRIBUTION.—Figure 62; apparently restricted to the canyon of the South Fork of the American River, Sierra Nevada, California. I have studied specimens from the following localities. United States of America CALIFORNIA: El Dorado County [Aug.] (9; CAS, ICCM, KSUC, MCZ, USNM), Kyburz (2 miles [ca. 3.2 km] E£ on South Fork American River [1,340 m]) [July] (5; CAS, DHKa), Pacific House [Sep.] (5; CAS), Riverton (South Fork American River [980 m], 3 miles [ca. 4.8 km] w at Bridal Falls Picnic Area [910 m]) [July—Aug.] (151; AMNH, CAS, DHKa, KUSM, MCZ, UASM), White Hall [June] (2; UASM). Nebria navajo, new species (Figures 43, 63) HOLotyPe, a male, in CAS, labelled: ‘‘Kayenta Ariz. 19 mi. sw VI-18-33"'/ **Navajo Co. Alt. 6500 ft.”’/ **Ansel F Hall Exp. 1933°’/ ‘Holotype Nebria navajo Kavanaugh det. D. H. Kavanaugh 1976” [red label]/ *‘California Academy of Sci- ences Type No. 12509.’’ One male PARATYPE (Same data as holotype) also in CAS. Type-Locatity.—Kayenta (19 miles [ca. 31 km] sw, Na- vajo County, Arizona. KAVANAUGH: NEW NEARCTIC NEBRIA DIAGNOSTIC COMBINATION.—Body pale tan- orange, legs pale yellow; microsculpture on pronotum and elytra moderately impressed, meshes isodiametric, alveolae flat; pronotum with lateral explanation narrow at middle, broadened posteriorly, basal angles rectangular or slightly acute, basal sinuation of lateral mar- gin moderately short and deep, midlateral seta absent; elytra with humeral carina slightly de- veloped, slightly projected anteriorly, elytral sil- houette ovoid, markedly narrowed basally, hu- meral angles moderately rounded and obtuse; hindwing short, narrow; hind coxa bi- or plu- risetose basally; apex of fourth tarsomere of hind tarsus with ventrolateral lobe long; third to fifth visible abdominal sterna each with two or more pairs of posterior paramedial setae; male with apex of median lobe as in Figure 43; spec- imen from Arizona (Fig. 63). DERIVATION OF TAXON NAME.—This species is named for the people native to the region in- cluding the type-locality. GEOGRAPHICAL DISTRIBUTION.—Figure 63; known only from the type-locality in northeast- ern Arizona. I have studied specimens from the following locality. United States of America ARIZONA: Navajo County, Kayenta (19 miles [ca. 31 km] sw [1,980 m]) [June] (2; CAS). Nebria gebleri cascadensis, new subspecies (Figures 21, 64) Nebria gebleri; auctorum. —HATcCH 1939:118 (in part). —LINDROTH 1961:83 (in part). —VAN Dyke 1925:119 (in part); 1926:10 (in part). HOoLorype, a male, in CAS, labelled: **U.S., Wash., Mt. Rainier N. P., Paradise R. above Narada Falls, 4580’—4800’, 8Aug72 DHKavanaugh & HGoulet’’/ *‘D. H. Kavanaugh Col- lection’’ [orange label]/ ‘‘ Holotype Nebria gebleri cascadensis Kavanaugh det. D. H. Kavanaugh 1976”” [red label]/ **Cali- fornia Academy of Sciences Type No. 12502... ALLOTYPE (same data as holotype) also in CAS. In total, 1696 PARATYPES (944 males and 752 females) are deposited in the following collections: AMNH, ANSP, BFCa, CAS, CNC, CUIC, DHKa, DJLa, FMNH, INHS, JSch, KSUC, KUSM, LRus, MCZ, MSU, NMDo, ODA, OSUC, OSUO, PURC, ROM, RTBe, SDSU, SJSC, UAFA, UASM, UBC, UCR, UIMI, UMMZ, USNM, UWBM, UWEM, WSU. Type-LocaLity.—Paradise River (above Narada Falls), Mount Rainier National Park, Washington. DIAGNOSTIC COMBINATION.—Head moderate in size, dark, with a pair of pale paramedial spots on vertex; pronotum (Fig. 21) markedly cordate, with or without faint violet metallic reflection, sinuation of lateral margin very deep, midlateral 105 seta present, basolateral seta present; elytra moderately convex, with distinct violet or brassy metallic reflection, elytral silhouette sub- ovoid, slightly narrowed basally, basal margin- ation moderately concave, humeral angle round- ly obtuse, humeral carina moderately to markedly developed, fifth elytral interval without setae (in few individuals with one to three setae), not (or in few individuals moderately) catenate, striae moderately impressed; specimen from locality in Cascade Range of central Oregon to southern British Columbia (Fig. 64). DERIVATION OF TAXON NAME.—This subspe- cies is named for the Cascade Mountain Range, in which members of this taxon are widely dis- tributed. GEOGRAPHICAL DISTRIBUTION.—Figure 64; restricted to the Cascade Range (from central Oregon to southern British Columbia), the ad- jacent mountain ranges west of the Okanagan lowland in southwestern British Columbia, and to Vancouver Island (not yet recorded from the Olympic Peninsula, Washington, although I pre- dict that it will be found to occur there). I have studied specimens from the following localities. Canada BritisH Co_tuMBIA: Copper Mountain [Aug.] (4; UBC), Cultus Lake Provincial Park (Entrance Bay Campground) [June] (4; ROM), Garibaldi (Cheakamus River) [Sep.] (2: JSpe), Garibaldi Provincial Park (Diamond Head Trail [1,070 m]) [July] (3; CNC), Hope [June-July] (8; BFCa, IHNS), Hunter Creek (at Restmore Lodge) [July] (3; CAS), Manning Provincial Park (Blackwell Peak [910 m], Lightning Lake, Nicomen Ridge, Skagit River) [May—Sep.] (35; CAS, CNC, DHKa, KUSM, MCZ, MSU, RTBe, UBC, UMMZ), Midday Valley (near Merritt) [Aug.] (1; INHS), Nicolum River (at Hope Trail) [July] (2; UBC), North Bend [July] (19; USNM), Skagit River (at Hope Trail [760 m]) [July] (3; UBC), Snass Creek (at Warburton Trail [760 m]) [July] (S; UBC), Spious Creek [May] (1; CAS), Vancouver area (2; ANSP), Whipsaw Creek (at Hope Trail [1,220 m]) [May, July] (4; UASM, UBC): Vancouver Island, Port Alberni (Robertson Creek Spawning Channel) [June] (1; ROM). United States of America OREGON: Clackamas County, Austin Hot Springs [July] (1; OSUO), Brightwood (Salmon River) [July] (8; DHKa), Mount Hood (s slope below Timberline Lodge [1,190 m]) [Aug.] (2; DHKa), Rhododendron ({490 m] and 1 mile [ca. 1.6 km] SE at Zig Zag River [600 m]) [July—Aug.] (55; CAS, DHKa, SJSC, UWBM); Deschutes County, Elk Lake [July] (1; UWBM), Upper Deschutes River [May] (1; UWBM): Hood River Coun- ty, Hood River [May, July] (8; CAS, USNM), Mount Hood (Cloud Cap, Hood River Meadows Ski Area [1,710 m], Sand Creek, Tillie Jane Creek, Umbrella Falls) [June—Sep.] (70; CAS, DHKa, MCZ, NMDo, ODA, UCR, UIMI, UWBM): Marion County, Detroit ([460 m] and 5 miles [ca. 8.0 km] N) [July—Aug.] (3; LRus, UWBM); Multnomah County, Horsetail Falls ({120 m]) [May-July] (87; DHKa, JSch, ODA, OSUO) WASHINGTON: (32; ANSP, CAS, INHS, MCZ, OSUC, 106 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 4 USNM); Chelan County, Fish Lake [July] (1; UWBM), Leav- enworth [July] (8; ANSP, KUSM, MCZ, SDSU, USNM), Tumwater Canyon ((610 m]) [May] (1; LRus); Cowlitz County, Ariel [May] (1; UWBM], Cougar [Sep.] (2; OSUO), Hoffstadt Creek (at Toutle River) [July] (2; MSU, OSUO), North Fork Toutle River (14 miles [ca. 23 km] E£ of Toutle (300 m]) [Aug.] (33; CAS, DHKa), Toutle River [July] (11; UWBM); King County, Baring [July] (2; FMNH), Green River Gorge (Green River) [May, Aug.] (5; UWBM), Green- water River (at Greenwater Campground) [June] (1; UWBM), North Bend [July] (17; CAS, CUIC, MCZ), Renton (Cedar River) [May] (1; UWBM), Skykomish [July, Sep.] (2; UASM, UWBM), Snoqualmie [June] (4; UIMI), Stevens Pass (and 4 miles [ca. 6.4 km] w) [June, Aug.] (5; LRus, UASM), Welling- ton [July] (10; CAS, MCZ, USNM, UWEM), White River (5 miles [ca. 8 km] w of Greenwater [1,490 m], Slippery Creek) [July—Aug.] (42; DHKa, UWBM); Kittitas County, Blewett Pass (Niger Creek) [July] (2; USNM), Easton (22; CAS, INHS, MCZ, USNM), Snoqualmie Pass [July] (4; CAS), Teanaway Ridge ([910 m]) [July] (10; LRus); Klickitat County, Klickitat Valley [July] (1; MCZ); Lewis County, Lake Creek (3 miles [ca. 4.8 km] NE of Packwood [370 m]) [July] (5; DHKa), White Pass (and 1.5 miles [ca. 2.4 km] w at Millridge Creek [1,280 m]) [June-July] (42; DHKa, SJSC, USNM); Mount Rainier National Park ({760 m—1,830 m]) [June—Sep.] (30; ANSP, CAS, CUIC, KSUC, MCZ, OSUO, SJSC, UIMI, UWBM), Bear Flat ([{1,520 m—1,830 m]) [July] (1; CAS), Carbon River [June] (4; UWBM), Cayuse Pass [1,400 m] [June] (4; USNM), Fish Creek (at West End Road) [June] (1; ROM), Fryingpan Creek ([{1,160 m—1,190 m]) [July—Aug.] (38; CNC, DHKa, SJSC), Horse Creek (near Long- mire) [July] (15; CAS), Ipsut Creek Campground {[Aug.] (1; OSUO), Kautz Creek (at Nisqually Riv- er) [Sep.] (2; UWBM), Longmire ({760 m]) [June—Aug.] (25; CAS, SJSC, USNM), Narada Falls ({1,370 m—1,460 m]) [July—Aug.] (118; DHKa, USNM, UWBM), Nisqually River (0.1 miles [ca. 0.2 km] above Paradise River [980 m], toe of Nisqually Glacier [1,220 m—1,520 m]) [July—Aug.] (7; AMNH, CAS, CUIC, DHKa, MCZ), North Fork Puyallup River (be- low toe of Puyallup Glacier [1,100 m—1,650 m]) [July—Aug. ] (22; CNC, DHKa, USNM), Ohanapecosh [June, Aug.] (18; DJLa, LRus, SJSC, UIMI, USNM), Paradise area (below Slu- skin Falls, Paradise Park [1,220 m—1,830 m], Paradise River {1,400 m—1,680 mJ], Paradise Valley) [June—Sep.] (88; CAS, CUIC, DHKa, FMNH, MCZ, USNM, UWBM), St. Andrews Creek (at West End Road [1,160 m]) [Aug.] (5; DHKa), Silver Creek [Aug.] (10; UWBM), Sluskin Falls [July] (8; UWBM), Stevens Creek ({1,190 m—1,220 m]) [June-July] (4; DHKa, USNM), Sunbeam Falls ({1,220 m]) [May] (1: CNC), Sunrise area [July] (2; LRus), Tahoma Creek ({910 m]) [July] (1; USNM), West End Road [July] (1; CAS), White River ({1,190 m] and at White River Campground [1,520 m]) [July—Aug.] (12; FMNH, SJSC, USNM, UWBM), Van Trump Creek (above Christine Falls [1,220 m—1,280 m]) [July—Aug.] (62: DHKa, UWBM); Pierce County, Goat Creek (at Nisqually River [610 m}]) [July] (130; DHKa), Fairfax [Oct.] (1; FMNH), Poch Creek (at Carbon River) [Aug.] (S; UCB, UWBM), Ten- as Creek (5 miles [ca. 8.0 km] £ of Ashford [610 m]) [July] (1; DHKa), White River (at Silver Springs Campground [790 m]) (June, Aug.] (15; DHKa, USNM); Skamania County, Carson {Aug.] (6; UIMI), Council Lake (se on Forest Road 123 [1,260 m]) [Aug.] (4; UASM), Mount St. Helens (Maratta Creek) {July} (3; DHKa), North Fork Cipsus River [June] (14; UWBM), North Fork Toutle River (at Spirit Lake Lodge [940 m]) [Aug.] (31; CAS, DHKa), Spirit Lake [July, Sep.] (3; DHKa, UWBM), Stabler [June] (1; OSUO), Wind River (8 miles [ca. 13 km] N of Carson [340 m]) [May, July] (14; DHKa); Snohomish County, Darrington (Clear Creek Falls) [May] (5; UWBM), Glacier Peak [July—Aug.] (2; CAS, UWBM); Whatcom County, Glacier (North Fork Nooksack River) [July] (13; CAS, LRus), Lookout Creek (4 miles [ca. 6.4 km] £ of Glacier [380 m]) [Aug.] (18; CAS, DHKa), Mount Baker (Austin Pass [1,520 m], Bagley Creek [670 m], Razor Hone Creek [1,680 m]) [July—Sep.] (125; CAS, DHKa, JSpe, LRus, UCR, UWBM), North Fork Nooksack River (4 miles [ca. 6.4 km] E of Glacier [380 m], Silver Fir Campground [610 m]) [Aug.] (3; CAS, DHKa), Shuksan ({760 m]) [Aug.] (3; LRus), Shuksan Creek [Aug.] (12; UWBM), Skagit River (at Newhalem) [July] (2; UWBM); Yakima County, American River (at Hell’s Crossing [1,040 m]) [July—Aug.] (17; DHKa, ICCM, KUSM, MCZ, UIMI, UMMZ, UWBM), Bumping River (0.8 miles [ca. 1.3 km] sw of Highway 410 [1,010 m]) [July—Sep.] (92; CAS, DHKa, ICCM, MCZ, MSU, PURC, UAFA, USNM, WSU), Dog Lake ([{1,310 m]) [July] (25; DHKa), Mount Adams (Bird Creek [910 m—2,440 m], Chap- arral Creek [880 m], Klickitat River) [June—July, Sep.] (82; AMNH, ANSP, CAS, CUIC, FMNH, ICCM, MCZ, UASM, UIMI, USNM, UWBM, WSU), Naches River [July] (2; USNM), Tampico [July] (1; USNM), Tieton Dam [Aug.] (2; AMNH, WSU), Wildcat Creek (at Tieton River [880 m]) [July] (3; DHKa, OSUO), Yakima area (11; JSch, MCZ, USNM, WSU), Yakima Indian Reservation ({910 m]) [July] (1; CAS). Doubtful Records UNITED STATES OF AMERICA—COLORADO: Leavenworth Valley (1; ANSP). WASHINGTON: Pullman (1; ICCM). Specimens Without Locality Data: (4; CDA, MCZ, MSU, UWBM). Nebria gebleri fragariae, new subspecies (Figures 22, 44, 49, 64) HoLorype, a male, in CAS, labelled: *‘U.S., Ore., Grant Co., Blue Mts., Strawberry Cr., 5800’ 28 May 1971, DHKa- vanaugh & EAMartinko’’/ *‘D. H. Kavanaugh Collection” [orange label]/ **Holotype Nebria gebleri fragariae Kavanaugh det. D. H. Kavanaugh 1979” [red label]/ *‘ California Academy of Sciences Type No. 12501.’’ ALLoTYPE (same data as ho- lotype) also in CAS. In total, 27 PARATYPES (21 males and 6 females) are deposited in the following collections: CAS, DHKa, UASM, USNM. Type-LocaLity.—Strawberry Creek (1,770 m), Grant County, Oregon. DIAGNOSTIC COMBINATION.—Size moderate, standardized body length male less than 10.5 mm, female less than 11.0 mm; head moderate in size, dark, with a pair of pale paramedial spots on vertex; pronotum (Fig. 22) moderately cor- date, with or without faint green metallic reflec- tion, midlateral seta present, basolateral seta present; elytra with distinct green metallic re- flection, elytral silhouette subrectangular, broad basally; male with apex of median lobe as in Figure 44; female with anterodorsal sclerite in KAVANAUGH: NEW NEARCTIC NEBRIA bursa copulatrix (Fig. 49); specimen from local- ity in Strawberry Range, Blue Mountains, east- ern Oregon (Fig. 64). DERIVATION OF TAXON NAME.—This subspe- cies is named for the Strawberry Range of the Blue Mountains, in which the type-locality is found. The subspecific epithet is a form of the scientific name for the strawberry genus, Fra- garia Linnaeus. GEOGRAPHICAL DISTRIBUTION.—Figure 64; restricted to the Strawberry Range of the Blue Mountains of eastern Oregon. I have studied specimens from the following locality. United States of America OrEGON: Grant County, Strawberry Creek ({1,770 m]) [May] (29: CAS, DHKa, UASM, USNM). Nebria gebleri siskiyouensis, new subspecies (Figures 23, 64) HoLoryPeE, a male, in CAS, labelled: *‘U.S.A., Calif., Trin- ity Co., Trinity Alps, S. F. Salmon R. at Big Flat Cmpgd., 1490m, 18July75 D. H. & B. A. Kavanaugh Stop #75-36°7/ “—D. H. Kavanaugh Collection’ [orange label]/ “‘Holotype Nebria gebleri siskiyouensis Kavanaugh det. D. H. Kavan- augh 1976” [red label]/ **California Academy of Sciences Type No. 12503.’ ALLOTYPE (same data as holotype) also in CAS. In total, 33 PARATYPES (19 males and 14 females) are depos- ited in the following collections: CAS, DHKa, FMNH, UASM, USNM. Type-LocaLity.—South Fork Salmon River (at Big Flat Campground), Trinity County, California. DIAGNOSTIC COMBINATION.—Head dark, with a pair of pale paramedial spots on vertex; prono- tum (Fig. 23) with apical angles moderately nar- row, midlateral seta present, basolateral seta present; elytra without metallic reflection, ely- tral silhouette subovoid, relatively short and broad; hindwing full-sized; specimen from lo- cality in Klamath Mountains system of Coast Ranges in northwestern California or south- western Oregon (Fig. 64). DERIVATION OF TAXON NAME.—This subspe- cies is named for the Siskiyou Mountains, in which members of this taxon were first collect- ed. GEOGRAPHICAL DISTRIBUTION.—Figure 64; restricted to the Klamath Mountains system of the Coast Ranges in northwestern California and southwestern Oregon. I have studied specimens from the following localities. United States of America CALIFORNIA: Trinity County, Boulder Creek (at Goldfield Campground [1,070 m]) [July] (2; CAS, DHKa), South Fork 107 Salmon River (at Big Flat Campground [1,490 m]) [June-July] (30; CAS, DHKa, FMNH, UASM, USNM), Taylor Lake Road (branch of Taylor Creek [1,750 m]) [Aug.] (1; CAS). OREGON: Jackson County, Little Applegate River ({700 m}) [Aug.] (2; CAS, FMNH). Nebria carri, new species (Figures 10, 24, 65) HOLotyPe, a male, in CAS, labelled: *‘U.S., Ida., Blaine Co., Sawtooth Mts., Smoky Range, 2 mi. e. Dollarhide Sum- mit, N. F. Warm Springs Creek, 7700’-7900', 21 Aug.73 DHKavanaugh Family’’/ *‘D. H. Kavanaugh Collection’’ [or- ange label]/ *‘Holotype Nebria carri Kavanaugh det. D. H. Kavanaugh 1976” [red label]/ ‘‘California Academy of Sci- ences Type No. 12498.°° ALLoTYPE (same data as holotype) also in CAS. In total, 152 PARATypPEs (63 males and 89 fe- males) are deposited in the following collections: BFCa, CAS, DHKa, FMNH, GASh, MCZ, UASM, USNM. Type-LocaLity.—Dollarhide Summit, Blaine County, Ida- ho. DIAGNOSTIC COMBINATION.—Head dark, with a pair of pale paramedial spots on vertex; pen- ultimate labial palpomere bisetose; mentum (Fig. 10) with M, setae absent; pronotum (Fig. 24) with midlateral seta present, basolateral seta absent; specimen from locality east of Columbia Basin (Fig. 65). DERIVATION OF TAXON NAME.—I take plea- sure in naming this species in honor of John and Betty Carr (Calgary, Alberta), who first collect- ed specimens of same, and who have provided help and encouragement to me and to other stu- dents during our respective studies. GEOGRAPHICAL DISTRIBUTION.—Figure 65; known at present only from ranges of the Saw- tooth Mountains system and from the Seven Devils Mountains of southcentral and western Idaho, respectively. I have studied specimens from the following localities. United States of America IpaHo: Adams County, Towsley Springs (9 miles [ca. 14 km] NNE of Cuprum [2,070 m]) [July—Aug.] (4; BFCa, DHKa); Blaine County, Dollarhide Summit (1.0 to 2.0 miles [ca. 1.6 to 3.2 km] £ on North Fork Warm Springs Creek [2,350 m- 2,500 m]) [Aug.] (138; CAS, DHKa, MCZ, UASM, USNM), Twin Lakes Trail (w of Petit Lake) [July] (2; BFCa); Camas County, Carrie Creek (32 miles [ca. 51 km] wsw of Ketchum [2,400 m]) [Aug.] (9; DHKa); Elmore County, Trinity Lakes [Aug.] (1; GASh). Nebria kincaidi balli, new subspecies (Figures 11, 31, 65) Nebria kincaidi; auctorum. —BLACKWELDER AND BLACK- WELDER 1948:2 (in part). —ERWIN AND BALL 1972:85 (in part). —Hatcu 1939:121 (in part). —LINDROTH 1961:88 (in part). —VAN Dyke 1919:8 (in part). 108 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 4 Nebria columbiana; auctorum (not Casey). —HATCH 1939:121 (in part). —VAN Dyke 1919:8 (in part). HoLortype, a male, in CAS, labelled: *“U.S., Wash., Mt. Rainier N. P., Paradise R. above Narada Falls, 4580’—4800’, 8Aug72 DHKavanaugh & HGoulet’’/ **D. H. Kavanaugh Col- lection’’ [orange label]/ ‘‘Holotype Nebria kincaidi balli Ka- vanaugh det. D. H. Kavanaugh 1976” [red label]/ *‘California Academy of Sciences Type No. 12505.’ ALLOTYPE (same data as holotype) also in CAS. In total, 366 PARATYPES (186 males and 177 females) are deposited in the following collec- tions: AMNH, CAS, CNC, CUIC, DHKa, FMMH, KUSM, LACM, LRus, MCZ, MSU, ODA, OSUO, SJSC, UASM, USNM, UWBM, WSU. Type-LocaLity.—Paradise River (above Narada Falls), Mount Rainier National Park, Washington. DIAGNOSTIC COMBINATION.—Head dark, with a pair of pale paramedial spots on vertex; pen- ultimate labial palpomere trisetose; mentum (Fig. 11) with M, setae present; pronotum with midlateral seta present, basolateral seta absent; elytra with basal margination (Fig. 31) moder- ately long, smooth or faintly wavy, slightly to moderately concave; specimen from Cascade Range of central Washington to northern Oregon (Fig. 65). DERIVATION OF TAXON NAME.—I take great pleasure in naming this subspecies in honor of George E. Ball, my mentor and friend. GEOGRAPHICAL DISTRIBUTION.—Figure 65; restricted to the Cascade Range of central Washington and northern Oregon. Present known range extends from Mount Rainier (Washington) south to Mount Hood (Oregon). I have studied specimens from the following lo- calities. United States of America OREGON: Hood River County, Mount Hood ({910—1,830 m], Cloud Cap Road at Tillie Jane Road, Hood River Meadows Ski Area [1,620 m—1,710 m], Tillie Jane Creek [1,830 m], Um- brella Falls) [June—Sep.] (67; AMNH, CAS, CUIC, DHKa, MCZ, MSU, ODA, USNM, UWBM, WSU). WASHINGTON: Mount Ranier National Park [July—-Sep.] (4; CUIC, SJSC, WSU), Cayuse Pass ({1,400 m]) [June] (1; USNM), Fryingpan Creek ([{1,160 m—1,190 m]) [July—Aug.] (2; CNC, DHKa), In- dian Henry’s ({1,370 m—1,520 m]) [July] (1; UASM), Louise Lake [July] (5; UASM), Mowich Lake [July] (4; OSUO, UWBM), Mystic Lake [July] (1; WSU), Nisqually River ({1,220 m]) [May] (4; CNC), Paradise area (above Narada Falls {1,520 m], Paradise Park [1,520 m—1,830 m], Paradise River {1,460 m—1,830 m]) [June—Aug.] (265; CAS, CNC, CUIC, DHKa, FMNH, KUSM, LRus, MCZ, OSUO, UASM, USNM, UWBM), Ricksecker Point [Sep.] (1; UWBM), Slu- skin Falls [July] (1; UWBM), Snow Lake ([{1,520 m]) [Aug.] (1; CNC), Spray Park ({1,980 m]) [Aug.] (1; LRus), Sunrise area [July] (2; CAS, LRus), Van Trump Creek (above Chris- tine Falls [1,220 m—1,280 m]) [July] (7; DHKa), Van Trump Park ([{1,830 m]) [Aug.] (1; CNC), West End Drive [July] (1; CAS); Pierce County, Coplay Lake [May] (1; UWBM); Ska- mania County, Spirit Lake ({910 m—1,770 m]) [Aug.] (1; OSUO). Doubtful Records UNITED STATES OF AMERICA—OREGON: Deschutes Coun- ty, Bend [July] (8; LACM). Nebria spatulata sierrae, new subspecies (Figures 6, 66) Nebria spatulata; ERWIN AND BALL 1972:83 (in part). HOLOTYPE, a male, in CAS, labelled: ““U.S.A., Calif., Mono Co., Sierra Nevada Mts., ne. face White Mt., along stream in cirque above Big Horn Lake, 3290m—3480m, 21 July 74 D.H.Kavanaugh Collector’/ **D. H. Kavanaugh Collec- tion’ [orange label]/ **Holotype Nebria spatulata sierrae Ka- vanaugh det. D. H. Kavanaugh 1976” [red label]/ **California Academy of Sciences Type No. 12516..° ALLOTYPE (same data as holotype) also in CAS. In total, 58 PARATYPEsS (28 males and 30 females) are deposited in the following collec- tions: CAS, DHKa, MCZ, RPPa, UASM, UCB, USNM. Type-LocaLity.—White Mountain (east slope, above Big Horn Lake), Mono County, California. DIAGNOSTIC COMBINATION.—Size moderate: standardized body length male less than 12.0 mm, female less than 12.4 mm; head with a pair of pale paramedial spots (or pair fused to form a single median spot), genae and occiput not swollen, eyes moderate in size and convexity; antennal scape (Fig. 6) moderate in length, slightly arcuate, markedly narrowed basally; pronotum with midlateral seta present, basolat- eral seta present; elytra rufous or piceus, very dull, with microsculpture deeply impressed, meshes isodiametric, without metallic reflec- tion, elytral silhouette markedly ovoid, nar- rowed basally; hindwing vestigial; specimen from locality on Eastern Divide or more north- ern portion of Sierra Nevada (Fig. 66). DERIVATION OF TAXON NAME.—This subspe- cies is named for the Sierra Nevada, California, to which its geographical range is restricted. GEOGRAPHICAL DISTRIBUTION.—Figure 66; restricted to the Eastern Divide of the Sierra Nevada in California; presently known from as far north as Sonora Pass (Tuolumne County) and as far south as Big Pine Creek (Inyo County). I have studied specimens from the following lo- calities. United States of America CALIFORNIA: Inyo County, Big Pine Creek (at Finger Lake {3,290 m]) [Aug.] (4: CAS); Mono County, H. M. Hall Natural Area ((3,350 m]) [July—Sep.] (9; RPPa), Saddlebag Lake [Sep.] (3; CAS), White Mountain (NE slope in cirque above Big Horn Lake [3,290 m—3,480 m]) [July] (13; DHKa); Tuolumne Coun- ty, Sonora Pass ({2,740 m—3,350 m]) [July] (3; UCB, USNM); Yosemite National Park, Maclure Creek Basin ((3,290 m— KAVANAUGH: NEW NEARCTIC NEBRIA 3,350 m]) [Sep.] (1; DHKa), Mount Lyell ({3,350 m]) [July— Aug.] (23; CAS, MCZ, UASM), Tioga Pass ({3,050 m]) [Aug. ] (2; CAS). Doubtful Records UNITED STATES OF AMERICA—CALIFORNIA: San Bernar- dino County [July] (2; CAS). Nebria meanyi lamarckensis, new subspecies (Figures 25, 32, 67) HoLotyPe, a male, in CAS, labelled: “‘U.S., Calif., Inyo Co., Sierra Nevada, Lamarck Cr. above Upper Lamarck Lk., 10700’—11000’ 6July71 DHKavanaugh & EAMartinko’’/ **D. H. Kavanaugh Collection”’ [orange label]/ ‘Holotype Nebria meanyi lamarckensis Kavanaugh det. D. H. Kavanaugh 1976” {red label]/ *‘California Academy of Sciences Type No. 12507... ALLOTYPE (same data as holotype) also in CAS. In total, 21 PARATYPES (6 males and 15 females) are deposited in the following collections: CAS, DHKa, UASM. Type-LocaLity.— Lamarck Creek (above Upper Lamarck Lake), Inyo County, California. DIAGNOSTIC COMBINATION.—Size moderate, standardized body length male less than 12.0 mm, female 9.5 to 12.3 mm; head dark, with a pair of pale paramedial spots on vertex; prono- tum (Fig. 25) markedly cordate, basal angles acute, projected posteriorly, midlateral seta present, basolateral seta present; elytra with faint or moderate violet, blue, or green metallic reflection, elytral silhouette subrectangular, broad basally, basal margination (Fig. 32) straight or slightly concave; specimen from lo- cality in southern Sierra Nevada (California) (Fig. 67). DERIVATION OF TAXON NAME.—The name for this subspecies refers to the type-locality, Lamarck Creek, which in turn refers to Jean Baptiste Pierre Antoine Lamarck, the noted French naturalist and pioneer evolutionist. GEOGRAPHICAL DISTRIBUTION.—Figure 67; restricted to the east slope of the southern Sierra Nevada. I have studied specimens from the fol- lowing localities. United States of America CALIFORNIA: Inyo County, Big Pine Creek (below Finger Lake [3,140 m]) [Aug.] (2; CAS), Lamarck Creek (above Up- per Lamarck Lake [3,260 m—3,350 m]) [July] (21: CAS, DHKa, UASM). Nebria meanyi sylvatica, new subspecies (Figures 26, 67) Nebria meanyi; HATCH 1939:118 (in part). Hototype, a male, in CAS, labelled: ““‘U.S.A., Wash., Olympic N. P., Olympic Mts., Boulder Cr. at Olympic Hot Springs, 610m, 15SAug.74 D. H. Kavanaugh’’/ *‘D. H. Kavan- augh Collection’? [orange label]/ *‘Holotype Nebria meanyi 109 sylvatica Kavanaugh det. D. H. Kavanaugh 1976” [red label]/ “California Academy of Sciences Type No. 12508.°° ALLo- TYPE (same data as holotype) also in CAS. In total, 177 Para- TYPES (83 males and 94 females) are deposited in the following collections: CAS, CNC, CUIC, DHKa, ICCM, JSch, UASM, USNM, UWBM. Type-LocaLity.—Boulder Creek (at Olympic Hot Springs), Olympic National Park, Washington. DIAGNOSTIC COMBINATION.—Head moderate in size and width, dark, with a pair of pale par- amedial spots on vertex; pronotum (Fig. 26) markedly cordate, basal sinuation of lateral mar- gin very short and deep, midlateral seta present, basolateral seta present; elytra moderately con- vex, elytral silhouette subrectangular to sub- ovoid, moderately broad basally, basal margin- ation slightly short, humeral angles faintly or moderately obtuse, humeral carina present, slightly to markedly developed, elytral intervals markedly convex, fifth interval with three to six setae, markedly catenate, striae deeply im- pressed; specimen from locality on Vancouver Island (British Columbia) or Olympic Peninsula (Washington) (Fig. 67). DERIVATION OF TAXON NAME.—This subspe- cies name is derived from the Latin word, syl- vaticus, meaning “‘of the forest’’—a reference to the deep forest habitat occupied by taxon members. GEOGRAPHICAL DISTRIBUTION.—Figure 67; restricted to Vancouver Island, British Colum- bia, and the Olympic Peninsula, Washington. I have studied specimens from the following lo- calities. Canada BritisH CoLuMBIA: Vancouver Island, Alberni (38 miles [ca. 61 km] w) [May] (1; CNC). United States of America WASHINGTON: Olympic National Park [Aug.] (11; ICCM, MCZ, UASM, USNM), Elwha River (at Elwha Campground) [Aug.] (1; CUIC), Dosewallips River (at Muscott Camp- ground) [July] (3; UWBM), Olympic Hot Springs (Boulder Creek [670 m—760 m]) [May—Aug.] (160; CAS, CNC, DHKa, JSch, UWBM), Sol Duc Hot Springs [July] (3; JSch, UWBM). Nebria vandykei wyeast, new subspecies (Figures 7, 68) Nebria vandykei; auctorum. —ERWIN AND BALL 1972:87 (in part). —Hatcu 1939:118 (in part). —LINDROTH 1961:86 (in part). Nebria trifaria vandykei BANNINGER. —HaTcuH 1953:56 (in part). HoLotyPE, a male, in CAS, labelled: *‘U.S.A., Ore., Clack- amas Co., Cascade Range, s. slope Mt. Hood, headwaters Salmon R. near Timberline Lodge, 1830m—1950m, 3 Aug.74 110 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 4 D. H. Kavanaugh’’/ *‘D. H. Kavanaugh Collection” [orange label]/ ‘‘ Holotype Nebria vandykei wyeast Kavanaugh det. D. H. Kavanaugh 1976” [red label]/ ‘*California Academy of Sci- ences Type No. 12517."’ ALLoTyPE (same data as holotype) also in CAS. In total, 186 PARATYPES (104 males and 82 fe- males) are deposited in the following collections: CAS, DHKa, MCZ, OSUO, UASM, USNM, UWBM. Type-Locatity.—Salmon River headwaters, Mount Hood, Clackamas County, Oregon. DIAGNOSTIC COMBINATION.—Size very large, standardized body length male greater than 12.0 mm, female greater than 12.5 mm; head dark, with a pair of pale paramedial spots on vertex; antennal scape (Fig. 7) narrowed basally, straight or slightly arcuate; pronotum with mid- lateral seta present, basolateral seta present; elytra without metallic reflection, elytral silhou- ette markedly ovoid; specimen from locality in Cascade Range south of Columbia River (Fig. 68). DERIVATION OF TAXON NAME.—For this sub- species, I have selected that name given to Mount Hood by local native peoples—Wy’ east, which means ‘*The Mountain.”’ GEOGRAPHICAL DISTRIBUTION.—Figure 68; restricted to the Cascade Range south of the Columbia River; present known range from Mount Hood south to the Three Sisters area (Oregon). I have studied specimens from the fol- lowing localities. United States of America OREGON: Deschutes County, Middle Sister Peak (E slope at North Fork Squaw Creek [1,950 m—2,190 m]) [Aug.] (19: CAS, DHKa); Hood River County, Mount Hood ([{1,740 m], Hood River Meadows Ski Area [1,920 m—1,950 m], headwaters of Salmon River near Timberline Lodge [1,830 m—1,950 m], Sand Creek) [June—Aug.] (130; CAS, DHKa, UWBM): Marion County, Mount Jefferson (N slope [1,860 m—1,920 m], s end of Jefferson Park [1,830 m], Russell Creek [1,680 m—1,740 m], West Branch Whitewater River [1,830 m—1,890 m]) [Aug.] (39; CAS, DHKa, OSUO). Nebria schwarzi beverlianna, new subspecies (Figures 1, 27, 36, 69) HOLoryPE, a male, in CAS, labelled: *‘U.S., Wyo., Sub- lette Co., Gros Ventre Mts., Hwy. 187/189, 8 mi. nw. Bon- durant, Hoback R., 6900’, 1-2 August 73 DHKavanaugh Fam- ily’’/ **D. H. Kavanaugh Collection’ [orange label]/ *‘Holotype Nebria schwarzi beverlianna Kavanaugh det. D. H. Kavan- augh 1976” [red label]/ ‘California Academy of Sciences Type No. 12515.”’ ALLoType (same data as holotype) also in CAS. In total, 254 PARATYPES (133 males and 121 females) are de- posited in the following collections: CAS, DHKa, FMNH, MCZ, ROM, UASM, USNM. Type-Locatity.—Hoback River (8 miles [ca. 13 km] Nw of Bondurant), Sublette County, Wyoming. DIAGNOSTIC COMBINATION.—Figure 1; size very large, standardized body length male great- er than 12.0 mm, female greater than 12.5 mm; head moderate in size and width, dark, with a pair of pale paramedial spots on vertex; prono- tum (Fig. 27) slightly flattened, only slightly cor- date, broad basally, midlateral seta present, ba- solateral seta present; elytra with marked violet, blue, or green metallic reflection, elytral silhou- ette subrectangular, broad basally; hind coxa (Fig. 36) with four to six basal and one (in very few individuals two or three) apical seta(-ae); specimen from locality in western Wyoming (Fig. 69). DERIVATION OF TAXON NAME.—I take great pleasure in naming this subspecies in honor of my wife, Beverly Ann Kavanaugh, in grateful acknowledgment of her constant support through the course of this study. GEOGRAPHICAL DISTRIBUTION.—Figure 69; at present known only from Hoback River at the south slope of the Gros Ventre Mountains. I have seen specimens from the following locality. United States of America WYOMING: Sublette County, Bondurant (8 miles [ca. 13 km] NW at Hoback River [2,100 m]) [July—Aug.] (255; CAS, DHKa, FMNH, MCZ, UASM, USNM). Specimens Without Locality Data: (1; ROM). Nebria trifaria utahensis, new subspecies (Figures 8, 28, 45, 50, 70) HOLOTYPE, a male, in CNC, labelled: *‘Lonesome Beaver, 7500', Henry Mts., Utah VII 20-22, 68 H. F. Howden’’/ “Holotype Nebria trifaria utahensis Kavanaugh det. D. H. Kavanaugh 1976” [red label]. ALLOTYPE (same data as holo- type) also in CNC. In total, 13 PARATYPES (9 males and 4 females) are deposited in the following collections: CAS, CNC, UASM. Type-LocaLtity.—Lonesome Beaver, Henry Mountains, Garfield County, Utah. DIAGNOSTIC COMBINATION.—Head moderate in size and width, dark, with a pair of pale par- amedial spots on vertex; antennal scape (Fig. 8) slightly arcuate or straight, slightly to markedly narrowed basally; pronotum (Fig. 28) with basal sinuation of lateral margin moderately deep, midlateral seta present, basolateral seta present; elytra without metallic reflection, microsculp- ture deeply impressed, meshes isodiametric, elytral silhouette subovoid; hindwing reduced in width and length, truncate distal to stigma; male with apex of median lobe as in Figure 45; female with posterodorsal sclerite in bursa copulatrix KAVANAUGH: NEW NEARCTIC NEBRIA as in Figure 50; specimen from Henry Moun- tains (Utah) (Fig. 70). DERIVATION OF TAXON NAME.—This subspe- cies is named for the State of Utah, in which its geographical range is restricted. GEOGRAPHICAL DISTRIBUTION.—Figure 70; restricted to the Henry Mountains of southcen- tral Utah. I have studied specimens from the following localities. United States of America Uran: Garfield County, Hanksville (24 miles [ca. 39 km] s [2,290 m]) [July] (3; CNC), Lonesome Beaver ([2,290 m]) [July] (12; CAS, CNC, UASM). NOMENCLATURAL NOTES AND LECTOTYPE DESIGNATIONS Carabus gyllenhali SCHONHERR, 1806:196. [=Nebria gyllenhali (Schonherr).] LEcTo- TYPE (here designated), a male, in NRSS, la- belled: **d’’/ [triangular blue label]/ **Lecto- type Carabus gyllenhali Schonherr designated by D. H. Kavanaugh 1976”’ [red label]. Five paralectotypes (same data as lectotype) also in NRSS. The name Carabus rufescens Strgm (1768:331) did not appear in the literature between the date of its original description and 1880, when Schgy- en (1880:183) revived its use (Carl Lindroth, per- sonal communication). Jeannel (1937:4), how- ever, accepted N. rufescens as the senior synonym of N. gyllenhali; and, following him, a number of authors have subsequently applied the name “‘rufescens’’ to this species (Blair 1950:220; Friden 1971:30; Greenslade 1968:41; Horvatovitch 1973:252; Luff 1972:174; Luff and Davies 1972:47; Nakane 1963:18, 1971:180; and Ueno 1953:59, 1955:47). The combination N. rufescens gyllenhali has sporadically appeared (Jeanne 1966:12; and Jeannel 1937:4, 1941:19) as has the unfortunate combination N. gyllenhali rufescens, where ‘‘rufescens’’ referred various- ly to a subspecies, race, variety, or aberration of N. gyllenhali (Andrewes 1939:161; Banninger 1925:345, 1960:261; Csiki 1927:359; Munster 1933:267; and Reitter 1908:93). The name given to this particular species is of considerable importance. The accumulated lit- erature on this species is very extensive, and its members are among the most broadly distrib- uted and commonly collected of all carabid bee- tles. Both names have been so often used in the literature that a decision for or against either will 11] cause some confusion, but it is still more im- portant to reach some decision which will pro- vide for consistency in all future usage. Lindroth (1939:59, 1954:121, 1961:78) and Banninger (1949:144) have refused to recognize N. rufes- cens as the valid name for this species. For the reasons they have cited, because no type-spec- imen has been found or perhaps ever existed, and because Strgm’s original description ‘‘can- not be interpreted (not even as a Nebria)’’ (Carl Lindroth, personal communication), I concur that the name N. gyllenhali (Schonherr) should be conserved as the valid name for the species. Any future use of the epithet rufescens should be discouraged; and the name Carabus rufes- cens Strgm should be listed as a nomen dubium. Carabus nivalis PAYKULL, 1790:52. [=Nebria nivalis (Paykull).] LEcTrotyPe (here designat- ed), a male, in NRSS, labelled: ‘‘Mus. Payk.’’/ **Riksmuseum Stockholm’”’ [green la- bel]/ *‘Lectotype Carabus nivalis Paykull des- ignated by D. H. Kavanaugh 1976’ [red la- bel]. The specimen chosen as lectotype of Carabus nivalis is one of three specimens so identified in NRSS from the Paykull Collection. The other two specimens are actually Nebria gyllenhali gyllenhali (Schonherr) adults. Because all three specimens bear identical labels and have tradi- tionally been considered syntypes, the two N. gyllenhali specimens could perhaps be consid- ered paralectotypes. Paykull (1798:119) subse- quently described a ‘‘variety B”’ of C. nivalis, which is clearly synonymous with N. gyllenhali gyllenhali and was based on specimens from a different locality and collector. The original de- scription of C. nivalis does not fit the two het- ero-specific specimens, but the description of “variety B’’ does so. I conclude that these spec- imens are probably not part of Paykull’s original type-series of C. nivalis; but they may be the specimens he described later as “‘variety B.” Helobia castanipes KirBy, 1837:20. [=Nebria gyllenhali castanipes (Kirby).] LECTOTYPE (here designated), a female, in BMNH, la- belled: ‘“‘Type’’ [red-trimmed disk]/ **70°’/ **N. Amer.”’ [pale blue disk with **5699"° on underside]/ ‘“‘apparently Type of castanipes Kby. Lindroth 1952’’/ ‘‘Lectotype Helobia castanipes Kirby designated by D. H. Kavan- augh 1976” [red label]. 112 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 4 The lectotype bears no label in Kirby’s hand. However, according to P. M. Hammond (per- sonal communication), the number ‘*5699”’ on the underside of one label refers to an entry in manuscript catalogs of the Coleoptera in BMNH. An original label in Kirby’s hand is pasted in the catalog at this entry and reads **73. 1. 2?/ castanipes.’’ The fate of the second of two specimens mentioned by Kirby in his orig- inal description is unknown. Nebria arkansana CASEY, 1913:52. Lindroth (1961:70) was certainly correct in denying the validity of Casey’s records for Ar- kansas and Indiana for this species; and his choice for the new type-locality (‘Valley of the Upper San Juan River,’ Archuleta County, ‘*Colorado’’) is quite reasonable. The specimens in Casey’s series are labelled ‘‘Ind’’ and ‘'S. Ark’’:; and it is at least possible that the latter refers to the South Arkansas River (a quite suit- able locality for the species, in Chaffee County, Colorado) rather than to the State of Arkansas as supposed by Casey. It is also possible that ‘‘Ind’’ refers to the ‘‘Indian Territory,’ al- though general use of that name for the region was abandoned long before 1913. Nebria bifaria MANNERHEIM, 1853:120. [=Ne- bria nivalis nivalis (Paykull).] LECTOTYPE (here designated), a male, in ZILR, labelled: ‘*38°’/ ‘‘Nebria carbonaria Esch. Insel. Paul’’/ ‘‘Lectotype ?’’ [red label]/ *‘Lectotype Nebria bifaria Mannerheim designated by D. H. Ka- vanaugh 1976” [red label]. Nebria catenata CASEY, 1913:49. [=Nebria tri- faria catenata Casey.] TypeE-LOCALITY.— Colorado; restricted by Erwin and Ball (1972:97) to San Juan Mountains, and here further restricted to Wolf Creek Pass, Mineral County, Colorado. Nebria diversa LECONTE, 1863a:2 [as a replace- ment name for N. livida LeConte]. LEctTo- TYPE (same as for N. livida LeConte; see be- low for data). Nebria elias MOTSCHULSKY, 1865:274. The type- specimen of N. elias has been completely de- stroyed and only its pin and labels remain (in ZMUM). Motschulsky’s original description is too vague to permit assurance that N. elias and N. gyllenhali castanipes are synony- mous. The occurrence of the latter in Alaska (the type area of N. elias) is restricted and local, which casts further uncertainty as to appropriate application of the name. I there- fore refrain from designation of a neotype and list N. elias as a nomen dubium, but, for con- venience, place it as a synonym of N. gyllen- hali castanipes, following Lindroth (1961:78). Nebria eschscholtzii MENETRIES, 1844:55. LEc- TOTYPE (here designated), a female, in ZILR, labelled: [gold-coated square]/ ‘‘California”’ [pink label]/ ‘‘Eschscholtzii Menet. Californ.’’/ ‘‘Syntypus’”’ [red label]/ ‘‘Lectotype Nebria eschscholtzii Menetries designated by D. H. Kavanaugh 1976” [red label]. One paralecto- type also in ZILR. Type-LocaLiry.—Califor- nia; here restricted to South Fork of American River, 3 miles [ca. 4.8 km] w of Riverton, El Dorado County, California. Nebria expansa Casey, 1913:56. [=Nebria la- custris Casey, 1913:56.] TypE-LOCALITY.— Indiana; here restricted to Turkey Run State Park, Parke County, Indiana. The name N. lacustris Casey has priority over N. expansa by precedence of position as well as by action of first reviser (Bell 1955:265; see also Lindroth 1961:77). In his original descrip- tion, Casey recorded N. expansa from **Texas and Indiana,’ based on his study of two fe- males. Bell (1955:267) and Lindroth (1961:77, 1975:112) accepted the record from Texas. In my judgment, this record must be erroneous. Acting on my suggestion, Lindroth (1975:147) amended his lectotype designation (1975:112) and selected the Casey specimen labelled “*L.,”” interpreted by Casey as from Indiana, instead of the ‘‘Texas”’ specimen. Individuals recognizable as the ‘‘expansa’’ form of N. lacustris have often been collected at the restricted type-local- ity chosen. Nebria fusiformis VAN DyKE, 1926:11. [=Nebria spatulata spatulata Van Dyke.] Because this name appeared unaccompanied by any description or indication, it should be considered a nomen nudum. It is clear to me, however, from the text in which the name ap- peared, that Van Dyke was referring to his N. spatulata, the original description of which ap- peared the previous year. KAVANAUGH: NEW NEARCTIC NEBRIA Nebria gebleri DEJEAN, 1831:573. HOLOTYPE, a female, in MHNP, labelled: ‘‘Sitka’’/ ‘‘Ge- bleri Eschs’’/ ‘‘Ex Musaeo Mniszech’’/ **‘Ho- lotype Nebria gebleri Dejean det. D. H. Ka- vanaugh 1976” [red label]. It is clear from his original description that the specimen of N. gebleri seen by Dejean was at that time complete. In his review of the Dejean types, Lindroth (1955b:12) noted that the head and prothorax of the type-specimen had been lost. In my study of the type (in 1976), I found it to be complete again! Unfortunately, the parts replaced are not only from a different specimen but also from one representing a different species [namely, Nebria picicornis (Fabricius)]. There is no doubt, however, that the pterothor- ax and abdomen (through which the pin passes) are the recognizable remains of the valid type- specimen. Nebria gregaria FISCHER VON WALDHEIM, 1821:72. LECTOTYPE (here designated), a male, in ZMUM, labelled: ‘‘47. gregaria Fisch.’’/ ‘‘Lectotype Nebria gregaria Fischer v. Wald. designated by D. H. Kavanaugh 1976” [red label]. Fischer (1821:73) indicated that his descrip- tion of N. gregaria was based on material in his own and Eschscholtz’s collections. I have been unable to locate any Eschscholtz specimens of N. gregaria from among those in UMHF. Ap- parently, the single Fischer specimen (ZMUM) chosen as lectotype is the only survivor from the type-series. Nebria hudsonica LECONTE, 1863b:3. LECTO- TYPE (here designated), a male, in MCZ, la- belled: ‘‘Saskatchewan’’/ ‘‘Type 643” [red la- belj/~ N* hudsonica LeC.”’?/ “Lectotype Nebria hudsonica LeConte designated by D. H. Kavanaugh 1976” [red label]. Typr-Lo- CALITY.—Saskatchewan, Hudson’s Bay Ter- ritory; here restricted to North Saskatchewan River at Rocky Mountain House, Alberta. In 1863, the area now occupied by the Prov- ince of Saskatchewan was part of Hudson’s Bay Territory (or the Northwest Territories), and at least up to that date, the name *‘Saskatchewan’”’ was apparently applied only to the Saskatche- wan River system. Therefore, LeConte’s use of the name as a type-locality probably refers to the river system rather than the province. Only those parts of the system in Alberta are within 113 the continuous range of N. hudsonica, although I have seen two specimens from localities in the Province of Saskatchewan. My selection of the restricted type-locality reflects these facts and considerations. Nebria incerta CASEY, 1913:53. [=Nebria obli- qua LeConte.] Type-LocaLity.—Colorado; here restricted to North Fork of South Platte Canyon at Santa Maria, Park County, Colo- rado. This restricted type-locality is the same as for N. obliqua. Nebria ingens Horn, 1870:98. LECTOTYPE (here designated), a female, in MCZ, labelled: ““Cala.’’/ “‘Type No. 1026’’ [red label]/ **Ne- bria ingens Horn’’/ ‘‘Lectotype Nebria ingens Horn designated by D. H. Kavanaugh 1976” [red label]. One female paralectotype, also in MCZ, labelled: ‘‘Cala.’’/ *“Type 8127 [red la- bel]/ “*N. ingens Horn.” Type-LocALity.— Sierra Nevada Mountains east of Visalia, Cal- ifornia; here restricted to Franklin Lakes, Tu- lare County, California. In his original description, Horn mentioned two specimens, a male and a female. The MCZ specimen seen by Lindroth (1961:87), his ‘‘al- lotype,’’ is a female. The other specimen, from the Horn Collection (formerly at ANSP, now at MCZ), is also a female; so Horn did not actually see a male of this species. The lectotype chosen is the specimen from the Horn Collection. Nebria livida LECONTE, 1859:84 [preoccupied by Carabus lividus Linnaeus, 1758:414; =Ne- bria diversa LeConte]. LECTOTYPE (here des- ignated), a male, in MCZ, labelled: [blue disk]/ ‘Type 642° [red label]/ ‘‘N. diversa LeC. || livida LeC.’’/ ‘‘Lectotype Nebria livida LeConte designated by D. H. Kavanaugh 1976’ [red label]. One male paralectotype (same data as lectotype) also in MCZ. Nebria longula LECONTE, 1878:478. [=Nebria suturalis LeConte—NEW SYNONYMYyY.]| TYPE- LocaLiry.—Colorado; here restricted to Longs Peak, Rocky Mountain National Park, Colorado. To date, all Colorado records for N. suturalis have been from the Front Range of the Rocky Mountains. The restricted type-locality is a pro- tected wilderness area in that range. 114 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 4 Nebria mannerheimii FISCHER VON WALDHEIM, 1828:253. LecToType (here designated), a male, in ZMUM, labelled: ‘‘Sitcha’’/ ‘‘48. mannerheimii Fisch.’’/ ‘“‘Lectotype Nebria mannerheimii Fischer v. Wald. designated by D. H. Kavanaugh 1976” [red label]. One male paralectotype in UMHF. Both specimens that I recognize as comprising the type-series of N. mannerheimii were prob- ably those collected by Eschscholtz (Fischer von Waldheim 1828:253). The specimen chosen as lectotype is from the Fischer Collection (ZMUM). The paralectotype is from the Man- nerheim Collection (UMHF) and bears an **Eschsch.”’ label. Nebria metallica FISCHER VON WALDHEIM, 1821:71. Lectotype (here designated), a male, in ZMUM, labelled: *‘Unalaschka. F.’’/ “75 metallicay Fischs’?/ “Lectotype Nebria metallica Fischer v. Wald. designated by D. H. Kavanaugh 1976” [red label]. One male and one female paralectotype are in UMHF. Fischer's description of N. metallica was based on material in his own and Eschscholtz’s collections (Fischer von Waldheim 1821:72). Specimens from the latter, identified as such, are now in UMHEF as part of the Mannerheim Collection. I have chosen the Fischer Collection specimen as lectotype, the two Eschscholtz specimens as paralectotypes. A fourth speci- men, in the Fischer Collection (ZMUM), bears a ‘‘Kadjak’’ [=Kodiak] label and is thus exclud- ed from the type-series. Nebria moesta LECONTE, 1850:209. [=Nebria gyllenhali castanipes (Kirby).] LECTOTYPE (here designated), a male, in MCZ, labelled: {light green disk]/ ‘Type 645” [red label]/ *“N. moesta LeC.’’/ ‘‘N. castanipes Kby.’’/ ‘‘Lec- totype Nebria moesta LeConte designated by D. H. Kavanaugh 1976” [red label]. One male and two female paralectotypes (same data as lectotype) also in MCZ. Nebria mollis MOTSCHULSKY, 1865:274. [=Ne- bria_ nivalis nivalis (Paykull).] LECTOTYPE (here designated), sex undetermined (speci- men incomplete, badly damaged), in ZMUM, labelled: ‘‘A. borealis’’ [green label]/ *‘Nebria mollis Motsch. Am. arc. cont.’’ [green label]/ [red rectangle]/ ‘‘Lectotype Nebria mollis Motschulsky designated by D. H. Kavanaugh 1976”° [red label]. In his original description, Motschulsky pre- sented the species epithet as “‘molbis.’’ I have found no published statement by Motschulsky that ‘‘molbis’> was a misspelling. However, the type-specimen is labelled ‘‘mollis’’; from as ear- ly as 1868, every subsequent citation of the name has used the latter spelling. This suggests early recognition, possibly initiated (privately) by Motschulsky himself, of an original error. The first clear indication of an intended emen- dation is that of Csiki (1927:389). Nebria muhlenbergii STURM, 1826:173 [here emended to Nebria muehlenbergii Sturm]. Because Sturm never selected type-speci- mens, none exists for this name. Furthermore, no description ever accompanied the use of this name. However, Sturm (1843:16) subsequently cited N. muehlenbergii as a synonym of N. pal- lipes, thereby providing the only clue to his ap- plication of the name to a taxon. The name should be treated as a nomen nudum. Nebria obliqua LECONTE, 1866c:363. LECTO- TYPE (here designated), a male, in MCZ, la- belled: “‘Col.’’/ ‘“‘Type 646’’ [red label]/ *‘Ne- bria obliqua Lec. Lewis Col.’’/ “‘obliqua 2’’/ ‘‘Lectotype Nebria obliqua LeConte desig- nated by D. H. Kavanaugh 1976” [red label]. One female paralectotype (same data as lec- totype) also in MCZ. Type-LocALity.—Col- orado; here restricted to North Fork of South Platte Canyon at Santa Maria, Park County, Colorado. The type-locality of N. obliqua, as originally stated, was Colorado. LeConte (1878:478) sub- sequently listed ‘‘North Fork of South Platte Canon (7,000 to 8,000 ft.) as a locality for the species, and Lindroth (1961:73) accepted this as the type-locality. I here formally restrict the type-locality to that area. Nebria obtusa LECONTE, 1878:478. [=Nebria obliqua LeConte—NEw SYNonyYMYyY.] My interpretation of LeConte’s original de- scription of N. obtusa is that the name is based on a single specimen (see LeConte 1878:479, lines 7-8). I therefore consider the type-speci- men (in MCZ) to be a holotype. The synonymy noted here was actually suggested by LeConte KAVANAUGH: NEW NEARCTIC NEBRIA (1878:479) and is supported by abundant data (Kavanaugh, manuscript in preparation). Nebria oregona CASEY, 1913:52. [=Nebria man- nerheimii Fischer von Waldheim.] Type-Lo- CALITY.—Clackamas County, Oregon; here restricted to Zigzag River at Rhododendron, Clackamas County, Oregon. Nebria ovipennis LECONTE, 1878:477. Type-Lo- CALITY.—Sierra Nevada, California; here re- stricted to Mount Conness (east slope, above Greenstone Lake), Mono County, California. The holotype of N. ovipennis (in MCZ) is rep- resentative of populations north of Mount Lyell, Yosemite National Park, but not south of that point in the Sierra Nevada. The restricted type- locality chosen is therefore both more precise and appropriate for the form represented by the type-specimen. Nebria pallipes SAy, 1823:78. Lindroth (1969a:1149, and in Lindroth and Freitag 1969:326) discussed the problem of es- tablishing the actual publication date of Say’s paper in which the original description of N. pallipes appeared. His comments, however, support the date cited above, rather than 1825, the year Lindroth (1961:76) cited. Nebria rathvoni LECONTE, 1853:400. [=Nebria gebleri rathvoni LeConte—NEwW STATUS.] HOLOTYPE, a male, in MCZ, labelled: **6456 ft. Lake Tahoe, Cal. May 24, 1879°’/ [light green square]/ ‘*79.’’ [red-tipped label]/ ““N. rathvoni LeC.’’/ *‘Type 7403’’ [red label]/ ‘*Holotype Nebria rathvoni LeConte det. D. H. Kavanaugh 1976” [red label]. Type-Lo- CALITY.—Sacramento, California; here emended to Truckee River at Truckee, Ne- vada County, California. There is no reason to doubt that the specimen recognized here as the holotype of N. rathvoni is actually the specimen seen by LeConte, al- though locality and date data on one label indi- cate otherwise. It is the only specimen of this taxon in the LeConte Collection (MCZ). Fur- thermore, the troublesome specimen label itself appears to be relatively new, and I suspect that it has been more recently added to the original type-specimen and thereby represents a misla- belling. 115 LeConte’s type-locality, Sacramento, is well outside the habitat range of the taxon: so the type was surely not collected there (although Sacramento was probably the closest important town to the true collection site. Truckee, the emended type-locality, is about at the midpoint of both the habitat and geographical ranges of the taxon. Nebria sahlbergii FISCHER VON WALDHEIM, 1828:254. LecTrorype (here designated), a male, in UMHF, labelled: ‘*¢”’/ ‘‘Eschsch.’’/ ““Sitka’’/ ‘‘Lectotype Nebria sahlbergii Fi- scher v. Wald. designated by D. H. Kavan- augh 1976” [red label]/ ‘“‘Lectotype Nebria violacea Motsch. designated by D. H. Kavan- augh 1976”° [red label]. Fischer's description of N. sahlbergii was based on a study of Eschscholtz material (Fi- scher von Waldheim 1828:254). I have located only two specimens probably representing the original type-series—one in the Fischer Collec- tion (ZMUM), the other in the Mannerheim Col- lection (UMHF). I have chosen the UMHF specimen as lectotype because, first, the original description better fits this specimen, and, sec- ond, the Fischer (ZMUM) specimen is actually a representative of N. gyllenhali castanipes rather than N. sahlbergii as the latter name was applied by Lindroth (1961:68). The Fischer specimen must be considered a paralectotype of N. sahlbergii in spite of its identity. In fact, the strange misuse of the name “‘sahlbergii’’ by nu- merous North American authors in referring to N. gyllenhali castanipes (Kirby) (e.g., Casey 1913:51; Hatch 1939:120; Horn 1870:102; Le- Conte 1853:400, 1878:479) can only now be ex- plained—that is, if these authors assumed that the Fischer specimen was the type of N. sahl- bergii. | choose not to follow their interpretation in order to preserve the name Helobia castan- ipes Kirby (see above), which would become an objective junior synonym of N. sahlbergii if the Fischer specimen were designated as lectotype. Nebria suturalis LECONTE, 1850:209. LeEcTo- TYPE (here designated), a male, in MCZ, la- belled: [light green disk]/ **Type 650” [red la- bel]/ ‘‘suturalis 3’’/ ‘‘Lectotype Nebria suturalis LeConte designated by D. H. Ka- vanaugh 1976” [red label]. Two female para- lectotypes (same data as lectotype) also in MCZ. 116 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 4 Nebria tenuipes CASEY, 1913:51. [=Nebria esch- scholtzii Ménétriés.] TypeE-LOCcALITy.—Ala- meda County, California; here restricted to Niles Canyon, Alameda County, California. Nebria texana Casey, 1913:54. [=Nebria obli- qua LeConte.] TypeE-LOcALITy.—‘‘Texas.”’ The holotype of N. texana (in USNM) bears the label ‘‘Tex,’’ interpreted by Casey as refer- ring to the State of Texas (a highly improbable record). It is more probable that the specimen was collected in Colorado, perhaps at Texas Creek, a small tributary of the Arkansas River, Fremont County, Colorado. However, I prefer not to amend or restrict the type-locality at pres- ent, pending further field search in the moun- tainous regions of western Texas. Nebria trifaria LECONTE, 1878:478. LECTOTYPE (here designated), a female, in MCZ, labelled: **9500 ft. Amer. Fork Canon Utah, Aug. 2-3, 1877 7 Type 651° [red labell/ “N-. tnfania LeC.”’/ ‘‘Lectotype Nebria trifaria LeConte designated by D. H. Kavanaugh 1976”’ [red label]. Nebria vandykei BANNINGER, 1928:5. LEcTO- TYPE (here designated), a male, in ETHZ, la- belled: ‘‘Paradise Val. Mt. Rainier Wash. VII- (83192077) Cole by EC] Van Dyke7/ +l. Bublstdstass i 2) BS?) “FG: Basizs bil: OP!’’/ *‘Nebria trifaria LeC.’’/ ‘‘Type Nebria Van Dykei 10.1927”° [red-trimmed label]/ ‘‘Lectotype Nebria vandykei Banninger des- ignated by D. H. Kavanaugh 1976’’ [red la- bel]. Two paralectotypes also in ETHZ. In his original description, Banninger men- tioned four specimens in the type-series, but only three of these are accounted for at present (W. Sauter, personal communication). Nebria violacea MOTSCHULSKY, 1850:73. [=Ne- bria sahlbergii sahlbergii Fischer von Wal- dheim.] LeEctorype (here designated), same specimen as lectotype of Nebria sahlbergii Fischer von Waldheim (see above). Mannerheim apparently obtained and exam- ined the Eschscholtz specimen of N. sahlbergii now in UMHF, studied the specimen identified as N. sahlbergii in Fischer’s Collection (ZMUM), and then noted (Mannerheim 1843:189) that two ‘varieties’ existed. He called the form repre- sented by the Eschscholtz specimen in his pos- session ‘“‘var. B.”’ Motschulsky (1850:73), refer- ring to Mannerheim’s brief description of variety ‘*B,’’ named this form N. violacea. Conse- quently, the specimen now in UMHF appears to be the specimen upon which both N. violacea and N. sahlbergii are based. I have therefore designated and so labelled the specimen as lec- totype for both names. Nebria viridis HORN, 1870:101. [=Nebria frigida R. F. Sahlberg.] LEcTotype (here designat- ed), a male, in MCZ, labelled: ‘‘R. [super- script ‘““M’’] A.’’/ ‘‘Type No. 1027-”’ [red la- bel]/ *‘N. viridis Horn’’/ ‘‘Lectotype Nebria viridis Horn designated by D. H. Kavanaugh 1976” [red label]. One male paralectotype (same data as lectotype) also in MCZ. In Horn’s original description, only two spec- imens are mentioned. There are now three spec- imens in MCZ (one from the Horn Collection, two from the LeConte Collection). I have des- ignated the Horn specimen as lectotype and the first LeConte specimen as the paralectotype. The second LeConte specimen (labelled ** viridis 2°’) has no type status. All three specimens bear identical first labels; but LeConte may have re- ceived his second specimen directly from Ulke (see Horn 1870:14), his first specimen from Horn. ACKNOWLEDGMENTS I gratefully acknowledge the assistance of those curators and other individuals listed above (under Materials and Methods) who loaned ma- terial in their care to me for study. In addition, the following individuals made type-material available to me on loan or provided critical in- formation on type-specimens: T. L. Erwin (USNM), P. M. Hammond (BMNH), H. Hippa (UMTF), S. Keleinikova (ZMUM), O. L. Kry- zhanovski and V. G. Schilenkov (ZILR), J. F. Lawrence and J. Scott (MCZ), H. B. Leech (CAS), O. Martin (ZMKD), T. Nyholm (NRSS), H. Perrin (MHNP), S. Rohwer (UWBM), W. Sauter (ETHZ), G. Scherer (ZSBS)? and ir Silfverberg (UMHF). T. L. Erwin (USNM), M. H. Hatch (UWBM), J. F. Lawrence (MCZ), H. B. Leech (CAS), P. Oman (OSU), Ey ere chuman (CUIC), A. Provonsha (PURC), and A. Smetana (CNC) were helpful hosts during my visits to study material in their respective insti- tutions. I thank also the many colleagues and KAVANAUGH: NEW NEARCTIC NEBRIA friends who assisted me in the field by providing companionship, physical labor, and other re- sources in support of the project, and my col- leagues at both the University of Alberta and the California Academy of Sciences for their en- couragement and assistance. Special thanks are due to.P. H. Armaud; Jr:, G. E. Ball, T. L. Er- win, H. Goulet, G. R. Noonan, and D. R. White- head for their continued interest in the project and their willingness to share with me their own observations and experiences. This study was supported by the National Re- search Council of Canada with funds from Grant A-1399 to G. E. Ball. Additional financial sup- port was obtained from the Boreal Institute for Northern Studies (University of Alberta), through a Grant In Aid of Research (1973-74) for fieldwork in Alaska, and from the California Academy of Sciences. LITERATURE CITED ANDREWES, H. E. 1939. The generic names of the British Carabidae, with a check list of the British species. The Ge- neric Names of British Insects, 6:153-192. ARMIN, C. 1963. A study of the family Carabidae (Coleop- tera) in Boulder County, Colorado. Unpublished Ph.D Dis- sertation. Department of Biology, University of Colorado, Boulder, Colorado. 476 pp. ARNETT, R. H., AND G. A. SAMUELSON. 1969. Directory of Coleoptera collections of North America (Canada through Panama). Purdue University Press, Lafayette, Indiana. vil + 123 pp. BALL, G. E. 1966. The taxonomy of the subgenus Scaphi- notus Dejean with particular reference to the subspecies of Scaphinotus petersi Roeschke. Transactions of the Ameri- can Entomological Society, 92:687-722. , AND J. NEGRE. 1972. The taxonomy of the Nearctic species of the genus Calathus Bonelli (Coleoptera: Carab- idae: Agonini). Transactions of the American Entomologi- cal Society, 98:412-533. BANNINGER, M. 1925. Neunter Beitrag zur Kenntnis der Car- abinae: die Nebriini. Entomologische Mitteilungen, 14:180— 195, 256-281, 329-343. . 1928. Uber die Nebriini. 13. Beitrag zur Kenntnis der Carabinae. Koleopterologische Rundschau, 14:1—7. Ento- mologicae, 11:143-242. . 1949. Ueber Carabinae (Col.). Erganzungen und Be- richtigungen III, mit Bemerkungen zu R. Jeannels neuer Einteilung der Carabiden. Mitteilungen der Minchner En- tomologischen Gesellschaft, 35-39: 127-157. . 1960. Die schweizerischen Arten der Gattung Nebria Latr. (Col. Carab.). Mitteilungen der Schweizerischen En- tomologischen Gesellschaft, 32:337—-356. BELL, R. T. 1955. Species of the pallipes-group of Nebria in the eastern United States. Proceedings of the Entomo- logical Society of Washington, 57:265—267. BLACKWELDER, R. E., AND R. M. BLACKWELDER. 1948. Fifth supplement to the Leng Catalogue of Coleoptera of 117 America, north of Mexico. John D. Sherman, Mount Ver- non, New York. 87 pp. BLairR, K. G. 1950. Nebria nivalis Paykull (Col., Carabidae) in Scotland. Entomologist’s Monthly Magazine, Ser. 4, 86:220-222. Brown, W. L., JrR., AND E. O. WiLson. 1954. The case against the trinomen. Systematic Zoology, 3:174-176. Casey, T. L. 1913. Studies in the Cicindelidae and Carabidae of America. Memoirs on the Coleoptera, 4:1—192. Csiki, E. 1927. Pars 92. Carabidae: Carabinae II. Pp. 315- 622 in W. Junk and S. Schenkling, editors, Coleopterorum Catalogus 1. W. Junk, Berlin. 345 pp. + 648 pp. [1926-33]. DeJEAN, P. F. M. A. 1831. Species general des Coléoptéres de la Collection de le Comte Dejean. 5. Mequignon-Marvis, Paris. viii + 883 pp. Epwarps, J. G. 1954. A new approach to infraspecific cat- egories. Systematic Zoology, 3:1—20. . 1956a. Clarification of certain aspects of infraspecific systematics. Systematic Zoology, 5:92-94. . 1956b. What should we mean by subspecies? Turtox News, 34:200-202, 230-231. . 1975. The Carabidae of Glacier National Park, Mon- tana. The Coleopterists Bulletin, 29:47-58. ERwin, T. L. 1970. A reclassification of bombardier beetles and a taxonomic revision of the North and Middle American species (Carabidae: Brachinida). Quaestiones Entomologi- cae, 6:4-215. , AND G. E. BALL. 1972. Classification of the ovipen- nis and trifaria groups of Nebria Latreille (Coleoptera: Ca- rabidae: Nebriini). Proceedings of the Biological Society of Washington, 85:77—108. FISCHER VON WALDHEIM, G. 1820-22. perii russici. I. Moscow. viii + 210 pp. 1825-28. Entomographia imperii russici. III. Mos- cow. vii + 315 pp. FRIDEN, A. 1971. Beetle fauna on the borders of some Scan- dinavian glaciers. Norsk Entomologisk Tidsskrift, 18:29-32. GREENSLADE, P. J. M. 1968. Habitat and altitude distribution of Carabidae (Coleoptera) in Argyll, Scotland. Transactions of the Royal Entomological Society of London, 120:39-54. Hatcn, M. H. 1939. A key to the species of Nebria of north- western North America. The Pan-Pacific Entomologist, 15:1 17=121: 1953. The beetles of the Pacific Northwest. Part I: Introduction and Adephaga. University of Washington Press, Seattle, Publication No. 16. vii + 340 pp. Horn, G. H. 1870. Descriptive catalogue of the species of Nebria and Pelophila of the United States. Transactions of the American Entomological Society, 3:97—-10S. Horvatovicn, S. 1973. Ergebnisse der zoologischen For- schungen von Dr. Z. Kaszab in der Mongolei. 306. Nebria nivalis changaica ssp. n. aus der Mongolei (Coleoptera: Carabidae). Folia Entomologica Hungarica, Series nova, 26:251-25S. INGER, R. F. 1961. Problems in the application of the sub- species concept in vertebrate taxonomy. Pp. 262—285 in W. F. Blair, editor, Vertebrate speciation. Conference on ver- tebrate speciation, University of Texas, 1958. University of Texas Press, Austin. xvi + 642 pp. JEANNE, C. 1966. Carabiques de la Peninsule Iberique. (3' note). Actes de la Société Linnéenne de Bordeaux, 103:3- 18. JEANNEL, R. 1937. Notes sur les Carbiques (Premiere note). Revue Frangaise d Entomologie, 4:1—23. Entomographia im- 118 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 4 . 1941. Faune de France, 39. Coléopteres Carabiques. Premiere partie. P. Lechevalier et Fils, Paris, 571 pp. KAVANAUGH, D. H. 1971. A new species of Nebria Latreille from Utah. Journal of the Kansas Entomological Society, 44:40-46. . 1978. The Nearctic species of Nebria Latreille (Co- leoptera: Carabidae: Nebriini): classification, phylogeny, zoogeography, and natural history. Unpublished Ph.D. Dis- sertation. Department of Entomology, University of Alber- ta. xlvili + 1041 pp. , AND E. A. MARTINKO. 1972. Notes on distribution and a peculiar behavior pattern in Nebria purpurata Le- Conte (Coleoptera: Carabidae). The Coleopterists Bulletin, 26:147-149. Kirpy, W. 1837. Part 4. The insects. [xxxix + 325 pp.] In J. Richardson, Fauna Boreali-Americana; or the zoology of the northern parts of British America. Josiah Fletcher, Nor- wich and London. LAROCHELLE, A. 1972. Notes sur les périodes d’accouplement de quelques Carabidae du Québec. Le Naturaliste Cana- dien, 99:61-63. 1976. Dates of egg-production in some carabid bee- tles (Coleoptera: Carabidae) from Québec. Cordulia, 2:47- 49. 1977. A list of teneral carabid beetles (Coleoptera: Carabidae) from Québec and their dates of capture. Cor- dulia, 3:109-113. Larson, D. J. 1975. The predaceous water beetles (Coleop- tera: Dytiscidae) of Alberta: systematics, natural history and distribution. Quaestiones Entomologicae, 11:245-498. LeConte, J. L. 1850. General remarks upon the Coleoptera of Lake Superior. Pp. 201-242 in J. L. R. Agassiz, Lake Superior, Natural History 4. Gould, Kendall, and Lincoln, Boston. x + 248 pp. + 8 plates. . 1853. Notes on the classification of the Carabidae of the United States. Transactions of the American Philosoph- ical Society (new series), 10:363-403. 1859. Catalogue of the Coleoptera of Fort Tejon, California. Proceedings of the Academy of Natural Sciences of Philadelphia, 11:69-90. . 1863a—66a. List of the Coleoptera of North America. Smithsonian Miscellaneous Collections, No. 140, 6:1-49 (1863), 50-70 (1866). . 1863b-66b. New species of North American Coleop- tera, Part 1, Smithsonian Miscellaneous Collections, No. 167, 6: 1-86 (1863), 87-168 (1866). 1866c. Additions to the coleopterous fauna of the United States. I. Proceedings of the Academy of Natural Sciences of Philadelphia, 1866:361—-394. 1878. The Coleoptera of the alpine regions of the Rocky Mountains. Bulletin of the United States Geological and Geographical Survey of the Territories, 4:447-480. LENG, C. W. 1920. Catalogue of the Coleoptera of America, North America. John D. Sherman, Mount Vernon, New York. x + 470 pp. LinpRrotH, C. H. 1939. Zur Systematik fennoskandischer Carabiden. 2. Nebria nivalis Payk. und gyllenhali Schonh. Entomologisk Tidskrift, 60:54—-62. . 1954. Random notes on North American Carabidae. Bulletin of the Museum of Comparative Zoology, 111:117— 161. . 1955a. The carabid beetles of Newfoundland includ- ing the French islands St. Pierre and Miquelon. Opuscula Entomologica, Supplement 12. 160 pp. + 9 plates. . 1955b. Dejean’s types of North American Carabidae. Opuscula Entomologica, 20:10-34. . 1961. The ground-beetles of Canada and Alaska, Part 2. Opuscula Entomologica, Supplement 20. Pp. 1-200. 1969a. The theory of glacial refugia in Scandinavia. Comments on present opinions. Notulae Entomologicae, 49:178-192. 1969b. The ground-beetles of Canada and Alaska, Part 6. Opuscula Entomologica, Supplement 35. Pp. 945— 1192. . 1971. On the occurrence of a continental element in the ground-beetle fauna of eastern Canada (Coleoptera: Ca- rabidae). The Canadian Entomologist, 103:1455—1462. . 1975. Designation of holotypes and lectotypes among ground beetles (Coleoptera, Carabidae) described by Thom- as L. Casey. The Coleopterists Bulletin, 29: 109-147. , AND R. FREITAG. 1969. North American ground-bee- tles (Coleoptera, Carabidae, excluding Cicindelinae) de- scribed by Thomas Say: designation of lectotypes and neo- types. Psyche, 76:326-361. LINNAEUS, C. 1758. Systema naturae. Tenth edition. Part 1. Stockholm. 824 pp. Lurr, M. L. 1972. The larvae of the British Carabidae (Co- leoptera) II. Nebriini. The Entomologist, 105:161-179. , AND L. Davies. 1972. Ecological observations on some Carabidae (Col.) from St. Kilda, with notes on other beetles new to the island. The Entomologist’s Monthly Magazine, 108:46—-S1. MapceE, R. B. 1967. A revision of the genus Lebia Latreille in America north of Mexico (Coleoptera, Carabidae). Quaestiones Entomologicae, 3:139-244. MANK, E. W. 1934. The Coleoptera of Glacier Park, Mon- tana. The Canadian Entomologist, 66:73-81. MANNERHEIM, C. G. 1843. Beitrag zur Kaefer-Fauna der Aleutischen Inseln, der Insel Sitkha und Neu-Californiens. Bulletin de la Société Impériale des Naturalistes de Mos- cou, 16:175—314. 1853. Dritter Nachtrag zur Kaefer-Fauna der nord- amerikanischen Laender des russischen Reiches. Bulletin de la Société Impériale des Naturalistes de Moscou, 26:96— 273. Mayr, E. 1969. Principles of systematic zoology. McGraw- Hill Book Co., New York. xi + 428 pp. MENETRIES, M. 1844. Sur un envoi d’Insectes de la cote N. O. d’Amérique. Bulletin de la Classe Physico-Mathema- tique de Académie Imperiale des Sciences de Saint-Pe- tersbourg, 2:50-63. MorTscHULSKY, V. 1850. Die Kaefer Russlands. I. Insecta Carabica. Gautier, Moscow. vii + 91 pp. . 1865. Enumération des nouvelles especes de Coleop- teres. IV. Bulletin de la Société Impériale des Naturalistes de Moscou, 38:227-380. Munster, T. 1933. Tillaeg og Bemaekninger til Norges Ko- leopterfauna. III. Norsk Entomologisk Tidsskrift, 3:267— 278. NAKANE, T. 1963. Carabidae. Pp. 5-52 and plates 3-26 in Iconographia insectorum Japonicorum colore naturali edita. II. (Coleoptera). Hokurykan, Tokyo. 1971. Notes on the Coleoptera from the alpine zone of the Daisetsu Mountains. Pp. 179-183 in Faunal Survey of the Mt. Daisetsu area, JIBP Main area—XII. Annual Report of JIBP—CTS for Fiscal Year 1970. Noonan, G. R. 1973. The anisodactylines (Insecta: Coleop- KAVANAUGH: NEW NEARCTIC NEBRIA tera: Carabidae: Harpalini): classification, evolution, and zoogeography. Quaestiones Entomologicae, 9:266—480. PAYKULL, G. 1790. Monographia Caraborum Sueciae. J. Ed- man, Uppsala. 138 pp. 1798. Fauna Suecica. Insecta. I. Uppsala. x + 234 pp. REITTER, E. 1908. Fauna Germanica. Die Kafer des deutsch- en Reiches. I. Stuttgart. viii + 248 pp. Say, T. 1823. Descriptions of insects of the families of Car- abici and Hydrocanthari of Latreille inhabiting United States. Transactions of the American Philosophical Society, new series, 2:1—109. SCHONHERR, C. J. 1806. Synonymia insectorum, oder: Ver- such einer Synonymie aller bisher bekannten Insecten; nach Fabricii Systema Eleutheratorum geordnet. I. H. A. Nords- trom, Stockholm. xii + 293 pp. SCH@YEN, W. M. 1880. Coleopterologiske notitser. Ento- mologisk Tidskrift (Stockholm), 1:177—185. Simpson, G. G. 1961. Principles of animal taxonomy. Colum- bia University Press, New York. xii + 247 pp. StrgM, H. 1768. Beskrivelse over Norske insecter. II. Det Kongelige Norske Videnskabers Selskab Skrifter, 4:313- Bile Sturm, J. 1826. Catalog meiner Insecten-Sammlung, erster Theil, Kafer. Niirnberg. 207 pp. + 4 plates. 1843. Catalog der Kafersammlung von Jacob Sturm. Niirnberg. xiii + 386 pp. + 6 plates. 119 UENO, S.-I. 1953. Carabidae. Shin Konchu (Tokyo), 6:52— 60. 1955. Two new species of the genus Nebria. The Entomological Review of Japan, 6:45—50. VAN Dyke, E. C. 1919. The distribution of insects in western North America. Annals of the Entomological Society of America, 12:1-12. . 1925. Studies of western North American Carabinae with descriptions of new species. The Pan-Pacific Ento- mologist, 1:111—125. . 1926. Certain peculiarities of the coleopterous insect fauna of the Pacific Northwest. Annals of the Entomological Society of America, 19:1-12. WHITEHEAD, D. R. 1972. Classification, phylogeny, and zoo- geography of Schizogenius Putzeys (Coleoptera: Carabi- dae: Scaritini). Quaestiones Entomologicae, 8:131—348. 1976. Classification and evolution of Rhinochenus Lucas (Coleoptera: Curculionidae: Cryptorhynchinae), and Quaternary Middle American zoogeography. Quaestiones Entomologicae, 12:118—201. WICKHAM, H. F. 1902. A catalogue of the Coleoptera of Col- orado. Bulletin from the Laboratories of Natural History of the State University of Iowa, 5:217-310. WILson, E. O., AND W. L. BRown, Jr. 1953. The subspecies concept and its taxonomic application. Systematic Zoology, 2:97—111. 120 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 4 Ficure 1. Nebria schwarzi beverlianna, new subspecies: adult male, dorsal aspect (approximately 6 times actual size). (Illustration by C. L. Mullinex.) KAVANAUGH: NEW NEARCTIC NEBRIA 121 Figures I-11. Fig. 1. Components of ‘*standardized body length’? measurement: HL = head length; PL = pronotal length; EL = elytral length. Figs. 2-8. Right antennal scape, dorsal aspect. 2. Nebria gouleti n.sp. (Underwood, Washington). 3. Nebria gyllenhali lassenensis n.ssp. (a. Mount Lassen, California; b. Todd Lake, Oregon). 4. Nebria gyllenhali lindrothi n.ssp. (a. Brooklyn Lake, Wyoming; b. Rio Puerco, New Mexico). 5. Nebria zioni oasis n.ssp. (Pine Valley Mountains, Utah). 6. Nebria spatulata sierrae n.ssp. (Big Horn Lake, California). 7. Nebria vandykei wyeast n.ssp. (Mount Hood, Oregon). 8. Nebria trifaria utahensis n.ssp. (Henry Mountains, Utah). Figs. 9-11. Mentum, ventral aspect. 9. General form and chaetotaxy; M, through M, are designations for specific pairs of setae. 10. Nebria carri n.sp. (Dollarhide Summit, Idaho). 11. Nebria kincaidi balli n.ssp. (Mount Hood, Oregon). All scale lines equal 1.0 mm. 122 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 4 am ati b _pti Figures 12-16. Fig. 12. Pronotum, general form and associated terms (a. dorsal aspect; b. left lateral aspect; c. cross-sec- tional aspect); abbreviation code: aa = apical angle; am = apical margination; ati = anterior transverse impression; ba = basal angle; bf = basal fovea; bls = basolateral seta; /b = lateral bead (= lateral margination); /e = lateral explanation; /g = lateral groove; mli = median longitudinal impression; mls = midlateral seta; pep = proepipleuron; pti = posterior transverse impres- sion. Figs. 13-16. Pronotum, dorsal aspect. 13. Nebria gouleti n.sp. (Salmon River, Idaho). 14. Nebria lacustris bellorum n.ssp. (Great Balsam Mountains, North Carolina). 15. Nebria nivalis gaspesiana n.ssp. (Mont Albert, Québec). 16. Nebria acuta quileute n.ssp. (Olympic Hot Springs, Washington). Scale line equals 1.0 mm. KAVANAUGH: NEW NEARCTIC NEBRIA 123 Ficures 17-24. Pronotum, dorsal aspect. 17. Nebria sahlbergii modoc n.ssp. (Warner Mountains, California). 18. Nebria sahlbergii triad n.ssp. (Trinity Alps, California). 19. Nebria obliqua chuskae n.ssp. (Chuska Mountains, Arizona). 20. Nebria darlingtoni n.sp. (Kyburz, California). 21. Nebria gebleri cascadensis n.ssp. (Glacier, Washington). 22. Nebria gebleri fra- gariae n.ssp. (Strawberry Mountains, Oregon). 23. Nebria gebleri siskiyouensis n.ssp. (Trinity Alps, California). 24. Nebria carri n.sp. (Dollarhide Summit, Idaho). Scale line equals 1.0 mm. 124 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 4 FiGures 25-29. Figs. 25-28. Pronotum, dorsal aspect. 25. Nebria meanyi lamarckensis n.ssp. (Upper Lamarck Lake, Cali- fornia). 26. Nebria meanyi sylvatica n.ssp. (Olympic Hot Springs, Washington). 27. Nebria schwarzi beverlianna n.ssp. (Hoback River, Wyoming). 28. Nebria trifaria utahensis n.ssp. (Henry Mountains, Utah). Fig. 29. Prosternal intercoxal process, Nebria lituyae n.sp. (Mount Blunt, Lituya Bay, Alaska). All scale lines equal 1.0 mm. KAVANAUGH: NEW NEARCTIC NEBRIA Ficures 30-36. Figs. 30-32. Basal region of left elytron, dorsal aspect. 30. Nebria arkansana edwardsi n.ssp. (a. Rancheria Yukon Territory; b. Gorge Creek, Alberta). 31. Nebria kincaidi balli n.ssp. (Mount Rainier, Washington). 32. Nebria meanyi lamarckensis n.ssp. (Upper Lamarck Lake, California). Figs. 33-35. Left hindwing. 33. Full-sized wing, Nebria arkansana edwardsi n.ssp. (Logan Pass, Montana). 34. Wing reduced in length, Nebria obliqua chuskae n.ssp. (Chuska Mountains, Arizona). 35. Wing reduced in length and width, Nebria arkansana uinta n.ssp. (Logan River, Utah). Fig. 36. Right hind coxa, ventral aspect, Nebria schwarzi beverlianna n.ssp. (Hoback River, Wyoming). All scale lines equal 1.0 mm. 126 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 4 Ficures 37-40. Fig. 37. Abdominal venter, form and associated terms (a. ventral aspect; b. left lateral aspect); abbreviation code: AS = anal sternum; HC = hind coxa; HF = hind femur; LP = lateral pit; MTE = metepisternum; MTS = metaster- num; PPMP = posterior paramedial puncture and associated seta; VS/ to VSS = first to fifth visible abdominal sterna. Figs. 38-40. Median lobe of male genitalia. 38. Nebria acuta quileute n.ssp. (Olympic Hot Springs, Washington) (a. left lateral aspect; b. ventral aspect, apical region only). 39. Nebria arkansana edwardsi n.ssp. (Yoho National Park, British Columbia) (a. left lateral aspect; b. dorsal aspect, apical region only; c. cross section at middle of shaft). 40. Nebria arkansana oowah n.ssp. (La Sal Mountains, Utah) (left lateral aspect). All scale lines equal 1.0 mm. KAVANAUGH: NEW NEARCTIC NEBRIA 127 ne = 41 42 43 44 Ficures 41-45. Median lobe of male genitalia, left lateral aspect. 41. Nebria arkansana uinta n.ssp. (Logan River, Utah). 42. Nebria fragilis teewinot n.ssp. (Mount Teewinot, Wyoming). 43. Nebria navajo n.sp. (19 miles [ca. 31 km] sw of Kayenta, Arizona). 44. Nebria gebleri fragariae n.ssp. (Strawberry Mountains, Oregon). 45. Nebria trifaria utahensis n.ssp. (Henry Mountains, Utah). Scale line equals 1.0 mm. 128 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 4 ee — See ese FiGures 46-50. Bursa copulatrix (a. dorsal aspect; b. left lateral aspect; c. bursal sclerite enlarged, dorsal aspect; d. bursal sclerite enlarged, left lateral aspect); in a. or b., stippled area denotes shape and location of distinct sclerite or moderately to markedly sclerotized, but less clearly defined, area; in c. or d., stippled area denotes weakly sclerotized membrane, sclerite is not stippled. 46. Nebria acuta quileute n.ssp. (Olympic Hot Springs, Washington). 47. Nebria arkansana edwardsi n.ssp. (Amiskwi Falls, British Columbia). 48. Nebria arkansana oowah n.ssp. (La Sal Mountains, Utah). 49. Nebria gebleri fragariae n.ssp. (Strawberry Mountains, Oregon). 50. Nebria trifaria utahensis n.ssp. (Henry Mountains, Utah). Scale line **x’’ equals 1.0 mm and applies to all figures ‘‘a’’ and ‘‘b.’’ Scale line “‘y’’ equals 0.10 mm and applies to all figures ‘“*c’’ and “‘d.” KAVANAUGH: NEW NEARCTIC NEBRIA 129 Figures 51-52. Geographical distribution maps. 51. Nebria gouleti n.sp. 52. Nebria lacustris Casey [N. lacustris lacus- tris = solid circles; N. lacustris bellorum n.ssp. = solid squares]. All scale lines equal 500 km. 130 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 4 FiGures 53-54. Geographical distribution maps. 53. Nebria nivalis (Paykull) [N. nivalis nivalis = solid triangles; N. nivalis gaspesiana n.ssp. = solid circles]. 54. Nebria gyllenhali (Schénherr) [N. gyllenhali castanipes (Kirby) = solid circles; N. gyllenhali lassenensis n.ssp. = solid squares; N. gyllenhali lindrothi n.ssp. = solid triangles]. All scale lines equal 500 km. KAVANAUGH: NEW NEARCTIC NEBRIA 131 FiGures 55-59. Geographical distribution maps. 55. Nebria acuta Lindroth [N. acuta acuta = solid circles; N. acuta quileute n.ssp. = solid triangles]. 56. Nebria sahlbergii Fischer von Waldheim [Nebria sahlbergii sahlbergii = solid circles; N. sahlbergii modoc n.ssp. = solid triangles: N. sahlbergii triad n.ssp. = solid square]. 57. Nebria lituyae n.sp. 58. Nebria arkansana Casey [N. arkansana arkansana = solid triangles; N. arkansana edwardsi n.ssp. = solid circles; N. arkansana oowah n.ssp. = open square; N. arkansana uinta = open triangles]. 59. Nebria fragilis Casey [N. fragilis fragilis = solid circles; N. fragilis teewinot n.ssp. = solid triangles]. All scale lines equal 500 km. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. Ficures 60-65. Geographical distribution maps. 60. Nebria zioni Van Dyke [N. zioni zioni = solid triangles; N. zioni oasis n.ssp. = solid circle]. 61. Nebria obliqua LeConte [N. obliqua obliqua = solid circles; N. obliqua chuskae n.ssp. = solid triangle]. 62. Nebria darlintoni n.sp. 63. Nebria navajo n.sp. 64. Nebria gebleri Dejean [N. gebleri gebleri = solid squares; N. gebleri cascadensis n.ssp. = solid triangles; N. gebleri fragariae n.ssp. = open square; N. gebleri rathvoni LeConte solid circles; N. gebleri siskiyouensis = open triangles]. 65. Nebria carri n.sp. [solid squares] and Nebria kincaidi Schwarz [N. kincaidi kincaidi = solid circles; N. kincaidi balli n.ssp. = solid triangles]. All scale lines equal 500 km. KAVANAUGH: NEW NEARCTIC NEBRIA 133 FiGures 66-70. Geographical distribution maps. 66. Nebria spatulata Van Dyke [N. spatulata spatulata = solid circles; N. spatulata sierrae n.ssp. = solid triangles]. 67. Nebria meanyi Van Dyke [N. meanyi meanyi = solid circles; N. meanyi lamarckensis n.ssp. = solid square; N. meanyi sylvatica n.ssp. vandykei = solid circles; N. vandykei wyeast n.ssp. solid circles; N. schwarzi beverlianna n.ssp. = N. trifaria catenata Casey = solid triangles; N. trifaria utahensis n.ssp. solid triangles]. 68. Nebria vandykei Banninger [N. vandyket solid triangles]. 69. Nebria schwarzi Van Dyke [N. schwarzi schwarzi = solid triangle]. 70. Nebria trifaria LeConte [N. trifaria trifaria = solid circles: solid square]. All scale lines equal 500 km. . PROCEEDINGS OF THE CALIFORNIA ACADEMY O Vol. 42, No. 5, pp. 135-179 Mating Bislogical Laboraior) | | SCIENCBRARY GAN8 1980 December 22, 1979 Woods -flole, Mags. EASTERN PACIFIC MACROURINE GRENADIERS WITH SEVEN BRANCHIOSTEGAL RAYS (PISCES: MACROURIDAE) By Tomio Iwamoto Department of Ichthyology, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118 ABSTRACT: Seventeen species representing eight genera of macrourine grenadiers with seven branchiostegal rays are treated as part of the eastern Pacific fauna. Among the eight genera, Nezumia is the most diverse, with 10 species represented. Nezumia ventralis new species, is described from two Galapagos specimens; the high pelvic fin ray count of 15 contrasts this species with all other eastern Pacific Nezumia. Two apparently disjunct populations of N. loricata are given subspecific recognition: subspecies /oricata from the Galapagos and sub- species atomos (new) from central Chile. Echinomacrurus, Hymenocephalus, Paracetonurus, Ventrifossa, Mal- acocephalus, Mataeocephalus, and Mesobius are each represented by only one species—the first four genera listed are first recorded from the eastern Pacific. Echinomacrurus occidentalis (a second species in the genus) is newly described from a single specimen taken off Peru in 4,334 m. Macrurus fragilis Garman, 1899, is tentatively aligned with members of Paracetonurus. Ventrifossa is recognized as consisting of three subgenera: Ventrifossa, Lucigadus, and Sokodara (new). Only subgenus Lucigadus is represented in the eastern Pacific. INTRODUCTION The large grenadier subfamily Macrourinae is divisible into two distinct groups based on bran- chiostegal ray counts. Those macrourines with six branchiostegal rays, typified by Macrourus Bloch and Coryphaenoides Gunnerus, form one group; those with seven branchiostegal rays and One monotypic genus (Pseudonezumia Okamu- ra) with seven or eight form a second group. This second group may be further divided by species differences in the development and lo- cation of the periproct region (the specialized area surrounding the anal and urogenital open- ings) and the development of ventral light or- gans. Hymenocephalus is the most distinctive member of this second group and forms a phy- logenetic line well removed from the others. Another line of related genera, characterized by the members having a broad periproct situated close to the origin of the anal fin, includes such diverse genera as Echinomacrurus Roule, Par- acetonurus Marshall, Cetonurus Gunther, Trachonurus Gunther, and Sphagemacrurus Fowler. These genera contrast with Nezumia Jordan, Malacocephalus Gunther, Ventrifossa Gilbert and Hubbs, Kumba Marshall, and Pseu- donezumia Okamura, members of which have a smaller periproct removed by some distance from the origin of the anal fin and preceded by an anteriorly extended light organ of various size. Mataeocephalus Berg falls somewhere be- tween these two divisions in that members of one species (M. adjustus) have a periproct that is well removed from the anal fin origin, and they also have a distinct dermal window of the light organ extended forward of the periproct, while members of the other species apparently have a periproct situated close before the ana! [135] 136 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 5 fin, and their light organ lacks a distinct dermal window. The 16 species here considered constitute about 40 percent of the total (approximately 40 spp.) macrourid fauna of the eastern Pacific. The genus Coelorinchus from this region (6 spp.) was reviewed in a previous paper (Iwamoto 1979). A review of the large complex of ma- crourines with six branchiostegal rays and the trachyrincines (2 spp.) from the eastern Pacific is currently underway. A summary of past stud- ies of the Macrouridae from eastern Pacific waters is given in Iwamoto and Stein (1974). In addition, the recent works of Hubbs and Iwa- moto (1977), Chirichigno and Iwamoto (1976), Shcherbachev et al. (1979), Ojeda and Camus (1977), and Parin et al. (1973, 1976) should be consulted. METHODS Methods for making counts and measure- ments generally follow procedures described by Hubbs and Lagler (1958) and modified for ma- crourids by Iwamoto (1970, 1978). Six or seven branchiostegal rays are found in all but one species of macrourid. The exception is Pseu- donezumia Okamura, 1971a, which has seven or eight (personal communications, Osamu Oka- mura, September 1978). Four of the rays are at- tached laterally on the epihyal and ceratohyal; the remaining two or three (or probably four in Pseudonezumia; condition not known) are at- tached anteriorly and medially on the cerato- hyal—thus only the latter group of rays need be counted to determine the total number. How- ever, these anteriormost rays (i.e., those closest to the isthmus) are generally small and slender, making them difficult to see without probing or dissection. Synonymies are limited to primary synonyms and other combinations. Materials for this study are based primarily on collections made by the ANTON BRUUN in 1966 and the TE VEGA in 1968; most of these are deposited in the ichthyological collection of the California Academy of Sciences (CAS). Other sources have been extensively used, and abbreviations for the depositories are as follows: AMNH, American Museum of Nat- ural History, New York; BMNH, British Mu- seum of Natural History, London; CAS-SU, Natural History Museum, Stanford University, now housed at CAS; FAKU, Faculty of Agri- culture, Kyoto University, Maizuru; FSFRL, ventral striae . ventral striae smooth fragile bones eas \ : \ \ ventral striae FiGureE |. Diagrammatic ventral (a) and lateral (b) views of a hypothetical representative of the genus Hymenocepha- lus, showing diagnostic features of the genus. Far Seas Fisheries Research Laboratory, Shi- mizu; IMARPE, Instituto del Mar, Callao; LACM, Natural History Museum of Los An- geles County, Los Angeles; MCZ, Museum of Comparative Zoology, Harvard University, Cambridge; SIO, Scripps Institution of Ocean- ography, La Jolla; UMML, School of Marine and Atmospheric Science, University of Miami, Miami; USNM, National Museum of Natural History, Washington, D.C. KEY TO THE ADULTS OF EASTERN PACIFIC MACROURINE SPECIES WITH SEVEN BRANCHIOSTEGAL RAYS la. Ventral striae (fine black transverse lines, most readily visible under magni- fication) present over gular membranes, isthmus, and parts of chest and abdomen (Fig. 1). Anus situated immediately be- fore anal fin and preceded by a small, IWAMOTO: EASTERN PACIFIC MACROURIDAE smooth large teeth in one row dermal windows b scales on branchiostegal rays FiGurRE 2. Malacocephalus laevis. (a) Lateral view show- ing smooth leading edge of second spinous dorsal ray and enlarged teeth of lower jaw. (b) Ventral view showing scales on branchiostegal rays and location of anus and light organ structures. raised lenslike structure; this structure connected by a thin (often obscure) black median line to similar structure on chest. Head bones extremely fragile, some almost membranous. Gill-rakers numerous, more than 20 (total count) on first arch. Second spinous dorsal ray smooth _____ Hymenocephalus sp. (p. 140) Ib. No ventral striae. Anus removed by a short to moderate distance from anal fin; lenslike structure, if present, far re- moved from anal fin origin and usually located in a shallow fossa. Head bones relatively strong. Gill-rakers fewer than 20 (total) on first arch. Second spinous dorsal ray smooth or serrated Z 2a. Spinous second ray of first dorsal fin with smooth leading edge (Fig. 2a) 3 2b. Spinous second ray of first dorsal fin with serrated leading edge ___._.________- 2) seme Cloviniil oats ote ral oi) :| ie an oe it ee ee Mesobius berryi Hubbs and Iwamoto (p. 141) broad naked margin al dermal window b FiGureE 3. Diagrammatic ventral views of abdomen of grenadiers showing periproct location (a) abutting or close to (separated by a few scale rows in some individuals) origin of anal fin, and (b) periproct far removed from origin of anal fin. 3b. 4a. 4b. ae Chin barbel present, well developed ____ 4 Teeth large, in a single row on lower jaw (Fig. 2a); scales on branchiostegal mem- braneChig. 2b) See see eee eee __ Malacocephalus laevis (Lowe) (p. 149) Teeth small, in a narrow band on lower jaw; no scales on branchiostegal mem- LOU REN VM aabendiach vive MEF, taal eich Dy le i de ala Periproct close to or abutting (Fig. 3a) origin of anal fin; anus usually closer to origin of anal fin than to insertion of pel- POAT ee oe. eee. Sa a: ae ee 6 . Periproct far removed from origin of anal fin (Fig. 3b); anus usually closer to in- 138 FIGURE 4. 6a. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 5 Ventrolateral views of (a) Ventrifossa nigro- maculata and (b) Nezumia latirostrata comparing relative lengths of first dorsal fin, anterior extent of gill openings (ar- rows), and relative positions of pelvic and anal fins. sertion of pelvic fins than to origin of YT NS Veen epee eg re eae 2c dn encgee co Oey em 8 Head massive, globose; snout broadly rounded. Scales on body distinctly non- imbricate (not overlapping), the exposed field of each scale separated by a fleshy border from the exposed fields of adja- CONUS Cale =esn= 9s: Aen eee ear re Echinomacrurus occidentalis n. sp. (p. 143) 6b. Head relatively slender, angular; snout ae pointed. Body scales distinctly imbri- cate; exposed fields of adjacent scales not distinctly separated from each other Byeanflesnyy DOGdER 2. 62222 eute seers ee. 7 Head bones stout; head ridges strongly armed with coarse scutelike scales. Mouth small, notably inferior, and dis- tinctly U-shaped. Scales densely cov- ered With spmmules ee fb: 8a. 8b. 9a. 9b. 10a. 10b. Mataeocephalus tenuicauda (Garman) (p. 145) Head bones fragile; head ridges naked or with unmodified scales. Mouth moderate in size, not notably inferior, and more normally shaped. Scales with few or no spinules on exposed field] >a Paracetonurus fragilis (Garman) (p. 147) Height of first dorsal fin much greater than length of head. Opercular opening extends far forward to vertical through hind edge of maxillae (Fig. 4a) _ Ventrifossa (Lucigadus) nigromaculata (McCulloch) (p. 153) Height of first dorsal fin about equal to or less than length of head. Anteriormost extent of opercular opening much pos- teriad of vertical through hind edge of maxillae (Fig.-4b) 1) =e 9 Second ray of first dorsal fin with 0 to 4 weak denticles on leading edge (Fig. Sa). Scales thin, highly deciduous with few or no spinules on exposed fields of body scales aia Oech Nezumia liolepis (Gilbert) (p. 157) Second ray of first dorsal fin with nu- merous prominent denticulations on leading edge (Fig. 5b). Scales relatively adherent, with spinules densely covering exposed fields...._.....4.) a Species bathypelagic. Gill filaments short; length about half diameter of eye lens. Outer pelvic ray prolonged, 70- 160% HL. Body scales small, bearing 1- 15 slender, erect spinules that render body surface velvety; 11-14 scale rows below origin of second dorsal fin. Color black to brownish black Nezumia parini Hubbs and Iwamoto (p. 176) Species benthopelagic. Gill filaments moderate to long; length about equal to or greater than diameter of eye lens. Outer pelvic ray usually less than 70% HL. Body scales small to moderate; spi- nules on scales few to numerous, mod- erately to greatly inclined, conical, lan- ceolate, or shield shaped, usually rough in texture; fewer than 11 scale rows be- low origin of second dorsal fin. Color IWAMOTO: EASTERN PACIFIC MACROURIDAE FIGURE 5. 4 b Comparison of denticulations on leading edge of second spinous dorsal rays of (a) Nezumia liolepis and (b) Nezumia loricata. lla. 11b. various shades of blue, violet, brown, or Mandibular rami usually completely na- ked; pores of lateralis system large and prominent on mandibular rami and along ventral border of suborbital region (Fig. 6a). Mouth relatively large, length upper jaw usually 30-36% HL. Outer gill-slit relatively wide, 17-22% HL Most of mandibular rami scaled, al- though anterior end naked in some; pores of cephalic lateralis system small, relatively inconspicuous (Fig. 6b). Mouth relatively small, length upper jaws 23- 32% HL. Length outer gill-slit 12-16% HL 12a. 12b. I3a. 14a. 14b. 139 Patches of small scales on base of low- ermost branchiostegal rays (Fig. 6a). Spinules on body scales relatively broad, lanceolate, arranged in a some- what quincunx pattern, rows not dis- cretely parallel or slightly convergent and ridgelike. Chin barbel long, 20-25% HL. Rays of pelvic fin 10-11. Orbit di- ameter 25=297o) Milly ses eee __ Nezumia stelgidolepis (Gilbert) (p. 161) Few isolated scales or no scales on bran- chiostegal rays. Spinules on body scales narrow, usually conical, arranged in dis- crete parallel or slightly convergent rows. Length of chin barbel 10-16% HL. Rays of pelvic fin 11-12. Orbit diameter 5s 7 oa lies ile ee NS grt oe ewe, 4, wk a | __.. Nezumia pulchella (Pequeno) (p. 159) Gill-rakers on inner side of first (outer- most) arch 9 or fewer (total count), 5—6 (rarely 7) on lower limb __ Nezumia convergens (Garman) (p. 171) . Gill-rakers on inner side of first arch 9 or more (total), 8 or 9 on lower limb ____ Pelvic fin rays 9. First dorsal with prom- inent black blotch distally, pale basally. Interorbital region narrow, width 17- Shere Nezumia orbitalis (Garman) (p. 167) Pelvic fin rays 9-12. First dorsal fin usu- ally uniformly blackish. Interorbital width moderate, 17-26% (usually more thans2 (7a) vb os a ee 15 FiGURE 6. Ventrolateral views of (a) Nezumia stelgidolepis and (b) Nezumia pudens comparing pore development (slightly exaggerated) and squamation of head in the two species. 140 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 5 . Snout relatively blunt, high. Suborbital region smoothly and almost completely scaled; scales along ridges not especially stout and coarse (Fig. 6b). Barbel long, 19-25% HL Nezumia pudens Gilbert and Thompson (p. 163) . Snout pointed, low. Suborbital region often with ventral surfaces partially na- ked; scales along ridges stout, coarse. Barbel shorter, less than 17% HL _.. 16 . Pelvic fin rays 9-10. First dorsal fin rays II,8—11 (usually 9-10) Length barbel S— 1 2/om § | eee neneee en.” Ss Ce ee __ Nezumia latirostrata (Garman) (p. 168) Pelvic fin rays 10-12. First dorsal fin rays II,9-12 (usually 10-12). Length bar- bela O St Ponta lego" aS Ue cee a oes Ie Be Nezumia loricata (Garman) (p. 174) 16b. Hymenocephalus Giglioli Hymenocephalus GiGLioLi, 1882:199 (type-species Hymeno- cephalus italicus Giglioli, 1884, by monotypy). Mystaconurus GUNTHER, 1887:124 (as subgenus of Macrurus) (type-species Hymenocephalus italicus Giglioli, 1884, by subsequent designation of Jordan and Evermann 1898:2580). Hymenogadus GILBERT AND Huss, 1920:521 (as subgenus) (type-species Hymenocephalus gracilis Gilbert and Hubbs, 1920, by original designation). —Okamura 1970a:58 (rec- ognized as genus). DIAGNOsIs.—Macrourine grenadiers with anus immediately before anal fin. Striae, consisting of fine parallel black lines over silvery pigment, on most ventral areas of body. Two lenslike dermal windows of light organ on midventral line; one immediately before anus and one on chest be- fore pelvic fin bases; dermal window connected by black ridge along midventral wall of abdom- inal cavity. Head large, cavernous; bones thin, head covering membranous. Mouth large, sub- terminal. Gill openings and outer gill-slit rela- tively wide. Gill-rakers numerous, more than 15 on lower limb of outer arch except in slender, cylindrical-bodied species (subgenera Hymeno- gadus and Spicomacrurus). Second dorsal spine smooth or weakly denticulate (in Hymenoga- dus). Small species, usually less than 200 mm TL. (Adapted after Iwamoto 1970:374-375.) REMARKS.—Okamura (1970a) elevated the subgenus Hymenogadus to full generic status, including in it H. gracilis Gilbert and Hubbs, 1920, H. tenuis Gilbert and Hubbs, 1917, and H. kuronumai Kamohara, 1938. The three species are obviously closely related and form a distinct group readily distinguished from most other Hymenocephalus (sensu lato), but it is not the differences between the species groups in Hymenocephalus but the similarities among them that are most striking. Used in its widest sense, the genus Hymenocephalus encompasses a group of species that is notably distinct and distantly removed from other groups of ma- crourine grenadiers. By recognizing both HAy- menocephalus and Hymenogadus, differences are emphasized and the obvious close relation- ship of the two groups is masked. This is re- grettable and entails a needless proliferation of names. Recognition of Hymenogadus (with gra- cilis and tenuis) and Spicomacrurus (with ku- ronumali) as subgenera within Hymenocephalus is to me a more practicable and meaningful treat- ment of the species groups involved. Hymenocephalus sp. DIAGNosIs.—Refer to generic diagnosis. COUNTS AND MEASUREMENTS.—I1D. II,8; gill-rakers on outer arch about 20. The following in millimeters: estimated HL 25; estimated TL 170; horizontal orbit diameter 10.3; length upper jaw 15.0; width suborbital 3.8; orbit to angle of preopercle 12.3; length barbel 3.7. REMARKS.—The single specimen of this genus from eastern Pacific waters was in such poor condition when examined in June 1975, that an adequate description could not be prepared. The head was damaged and had become separated from the trunk, and the paired fins were gone; but there was no question as to its genus because of the presence of striae on the gular membrane and along the abdomen; the presence of two lenslike light organs on the chest and before the anus; the number, shape, and coloration of the gill-rakers; and the shape of the preopercle ridge—these combined features uniquely char- acterize the genus Hymenocephalus. This specimen represents the first record of the genus from eastern Pacific waters. The genus is common throughout the warm-water regions of the Atlantic, Indian, and most of the central and western Pacific oceans. That only a single specimen has been collected—and that a fair- sized adult taken in a midwater haul—suggests that the species is not a regular inhabitant of eastern Pacific waters. The specimen may rep- resent a stray or an expatriate, possibly from populations far to the westward. IWAMOTO: EASTERN PACIFIC MACROURIDAE MATERIAL EXAMINED.—USNM 149049 (1 specimen, est. 170 mm TL); Peru, off Aguja Pt., 5°57’30’S, 81°50’ W; vertical haul, 732-0 m over bottom depth of 4,023 m; ALBATROSS sta. 4655, 12 Nov. 1904. Mesobius Hubbs and Iwamoto Mesobius HUBBS AND IWAMOTO, 1977:235 (type-species Me- sobius berryi Hubbs and Iwamoto, 1977, by original desig- nation). DIAGNOosIS.—Bathypelagic. Periproct region broad, somewhat raised, close to anal fin origin (removed from origin by 24 scale rows in some individuals); anus centrically located within periproct. Large light organ abutting rectum, but no anterior extensions between pelvic fin bases or on chest. Abdomen short, distance isthmus to anal fin origin 1.0-1.5 of orbit diameter in adults. Chin barbel absent. Dentition in both jaws consist of narrow bands of small teeth. Scales of head elongate, each bearing 1-3 rows of spinules that form low, sharp ridges with 2— 9 spinules per row. Head and body laterally compressed. Swim bladder greatly reduced, bearing 2 retia and 2 gas glands. Postlarvae and prejuveniles pass through a polka-dotted (*‘phal- acromacrurus’’) stage; adults mostly black. (Adapted from original description.) REMARKS.—Since publication of the genus description, Dr. Carl L. Hubbs and I have re- ceived information concerning additional speci- mens of the genus. Dr. Nikolai V. Parin has written (to Hubbs, 25 Nov. 1977) that represen- tatives of the genus have been collected by Rus- sian vessels in the Atlantic and Indian oceans between latitudes 31° and 37°S. These specimens are presumably those recently reported by Shcherbachev et al. (1979) who recorded M. berryi from the Indian Ocean, and M. antipo- dum from the Indian Ocean and the Atlantic Ocean off the southwestern tip of Africa. Unlike the type-specimens of both species, the Russian specimens were captured in bottom trawls. The largest M. antipodum recorded by Shcherba- chev et al. was more than 661 mm in total length and 136 mm in head length, and their six spec- imens ranged 104-136 mm HL and 465+ to 661+ mm TL. Their five specimens of M. berryi ranged 64.5—75.5 mm HL and 330+ to 412+ mm TL. The M. antipodum specimens are consid- erably larger than any of the M. antipodum and M. berryi Dr. Hubbs and I had examined (the largest M. berryi we had was 392 mm in total length and 70 mm in head length; the holotype 141 of M. antipodum was 390 mm long, with an in- complete tail, and 75 mm in head length). Dr. Christine Karrer has also informed us (personal communication, Oct. 1977) of having examined specimens of Mesobius in the Institut fiir See- fischerei (ISH), Hamburg, that were captured in the Atlantic by the WALTHER HERwIG. Four of these (ISH 1816/68; 88-107 mm HL) were bor- rowed through the kindness of Dr. M. Stehman and were identified as M. antipodum. Takao Arai (1979) recently reported the collection of 12 specimens of M. antipodum off New Zealand, the largest of which was 128 mm in head length. It thus appears that M. antipodum attains a much larger size than M. berryi, and that spec- ulation (Hubbs and Iwamoto 1977:246) as to the probable distribution of M. antipodum extend- ing widely across the Southern Hemisphere is supported. Mesobius berryi Hubbs and Iwamoto (Figure 7a) Mesobius berryi HUBBS AND IWAMOTO, 1977:236-244, figs. 1-8, 10A (original description; holotype and 18 paratypes from eastern North Pacific). DiaGNosis.—A species of Mesobius with 12- 17 pyloric caeca; mesial gill-rakers on first arch 10-13 ( = 11.8), on second arch 10-12 (¢ = 11.0). Length posterior nostril less than 4 into least suborbital width. Differentiated squama- tion of posttemporal region not extending pos- teriad of vertical through origin of pectoral fin. Outer margin of gill cover not notably incised at subopercle. REMARKS.—General features of the fish can be seen in Figure 7a. This species has been ad- equately described in the original description, and in the recent reports by Arai (1979) and Shcherbachev et al. (1979). Mesobius berryi and Nezumia parini are the only eastern Pacific species of macrourid normally living bathype- lagically as adults (adults of other species make excursions into bathypelagic depths, but their normal habitat is near bottom, i.e., benthope- lagic; see Marshall and Merrett (1977) and Mer- rett (1978) for recent discussions on this sub- ject). The unique squamation of the head of Mesobius berryi is unlike that of any other species encountered in the eastern Pacific and makes adults of the species easily recognizable. The specific differences originally reported be- tween M. berryi from the North Pacific and M. antipodum from the South Pacific have generally 142 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 5 FIGURE 7. been confirmed by Arai (1979) upon his exami- nation of 12 specimens of M. antipodum cap- tured off the New Zealand shelf and by Shcher- bachev et al. (1979) from examination of specimens from the Indian Ocean and the south- western Atlantic. Meristic features show the best differences (see Table 1), with the pyloric caeca count showing no overlap (12-17 in eight specimens of M. berryi, 30-43 in more than six specimens of M. antipodum) (data from Hubbs and Iwamoto (1977), Shcherbachev et al. (1979), and Arai (1979)); the last author gave only the range of counts in his 12 specimens of M. an- tipodum). Echinomacrurus Roule Echinomacrurus ROULE, 1916:22 (type-species Echinomacru- rus mollis Roule, 1916, by monotypy). DIAGNOosIs.—Macrourine grenadiers with anus located within a broad naked area that lies im- mediately adjacent to anal fin origin; head mas- sive, swollen by the expansive cephalic lateral- (a) Mesobius berryi Hubbs and Iwamoto. (Adapted from fig. 1 in Hubbs and Iwamoto 1977.) (b) Echinomacrurus occidentalis [wamoto, new species. Reconstruction of holotype, USNM 135612, collected by the ALBATROss off Peru in 4,334 m. line canals; second spinous ray of first dorsal fin serrated along leading edge; scales nonimbri- cate, with slender, erect spinules; scales along dorsal fins not especially enlarged (as in Ceto- nurus), no scales on gular and branchiostegal membranes; swim bladder reduced or absent. (See Marshall (1973:599) for additional charac- ters.) REMARKS.—The genus Echinomacrurus has hitherto been known only from the eastern North Atlantic and the western Indian Ocean (Marshall 1973). The eastern Pacific specimen here reported is the first Pacific and, at 4,334 m, the shallowest record of the genus. Other spec- imens of Echinomacrurus have been taken at depths from 5,000 to 5,413 m, and except for one Indian Ocean specimen, all were taken in nets fished at the bottom—Marshall (1973) thus considers E. mollis bathypelagic to benthope- lagic in habit. The genus is closely related to Cetonurus Vaillant, 1888; the two agree in most diagnostic characters given above but not in squamation IWAMOTO: EASTERN PACIFIC MACROURIDAE TABLE 1. COMPARISON OF SELECTED COUNTS IN MESO- BIUS BERRYI AND M. ANTIPODUM. Data from Hubbs and Iwamoto (1977), Arai (1979), and Shcherbachev et al. (1979). Total gill-rakers on first arch range x S.D. n M. berryi 10-13 ile 7/ 0.82 19 M. antipodum 12-15 13.9 0.85 31 Total gill-rakers on second arch range x S.D. n M. berryi 10-12 EZ 0.62 20 M. antipodum 12-16 13.9 0.89 31 Pectoral fin rays range X S.D. n M. berryi 12-14 12.9 0.63 33 M. antipodum 13-16 14.5 0.68 31 Pelvic fin rays range x S.D. n M. berryi 7-9 Tel 0.57 35 M. antipodum 6-7 6.9 0.35 30 and swim-bladder characters. Cetonurus has imbricate scales over most of the body and no- tably enlarged scales along the base of the sec- ond dorsal fin; it also has scales over the lower branchiostegal rays and on the gular membrane, and a well-developed swim bladder. Echinomacrurus occidentalis new species (Figure 7b) DIAGNOosIs.—An Echinomacrurus with 12 pelvic fin rays. Orbits about 20 percent HL. Eleven gill-rakers on mesial side of second arch. Interspace between first and second dorsal fins 31 percent HL. Counts.—1D. ca. II,9; 1P. i18/118; 2P. ca. 12/ 12. Gill-rakers on first arch 1 + 9; on second arch 2 + 9. Pyloric caeca 6. MEASUREMENTS (all in mm; measurements preceded by ca. are estimates).—Total length ca. 400; head length ca. 80; snout length ca. 27; preoral length ca. 23; horizontal orbit diameter 16; least postorbital length 39; orbit to angle of preopercle 36; suborbital width 10; length upper jaw 22.5; length barbel 7; length outer gill-slit 9; preanal length ca. 110; length snout to anus ca. 143 102; outer pelvic to anal 29; isthmus to anal ca. 54; greatest body depth ca. 62; depth over anal origin ca. 53; 1D.—2D. interspace 31. DESCRIPTION.—Head deep, massive, presum- ably swollen in appearance when fresh; snout high, broad; nostrils high, located at about level of dorsal margin of orbits; interorbital space broad, convex. Mouth rather small, subinferior, but upper jaw extends near to vertical through hind edge of orbit; ascending limb of premaxilla high (height about 0.9 of ramus length), inclined forward. Interopercle mostly covered by, and closely adhered to, preopercle. A distinct notch in outline of gill cover formed by posteroventral border of subopercle. Scales small, nonimbricate, almost in mosaic pattern on parts of head, more widely spaced on body. Thin, erect spinules cover most scales, giving shagreenlike feel to skin. Scales every- where cover exposed surfaces of head and body except on fins, lips, parts of subopercle, wide margin surrounding anus, small areas behind pectoral and dorsal fins, and gill membranes. Shoulder girdle beneath gill cover naked; border between naked and scaled areas of shoulder sharply demarcated. Periproct large, raised, im- mediately adjacent to anal fin and occupying al- most half distance between insertion of pelvic fins and origin of anal fin; anus slightly protrud- ing. Premaxillary teeth small, conical, recurved, in a narrow, tapered band of about four rows deep anteriorly, tapering to a single row poste- riorly; band extends about three-fourths length of premaxillary ramus; outer series of teeth slightly enlarged. Mandibular dentition about the same as that of premaxillary, but without enlarged outer teeth. Gill-rakers short, spiny, tubercular. Outer gill- slit relatively long; about 8 rudimentary rakers on outer side of first arch, 1 + 9 on inner side, these armed with slender conical spines. Pyloric caeca 6, large, thick. Specimen a male with 2 well-developed testes. Linings of gill, buccal, and peritoneal cavities black. Stomach black. REMARKS.—It is with some reluctance that | describe this single eastern Pacific specimen of Echinomacrurus as a new species, because of its extremely poor condition. It is unlikely, how- ever, that any additional material of this rare species will be soon forthcoming, and there is little doubt that the specimen represents an un- described taxon. The pelvic fin ray count of 12, 144 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 5 which I have confirmed on the right fin by stain- ing, is distinctly higher in the eastern Pacific specimen than the 9-10 reported for E. mollis by Nybelin (1957) and Marshall (1973:599). The larger orbit (20 percent HL cf. 10-15 percent), which goes into the distance orbit to angle of preopercle about 2.2 times (compared with 2.5 or more in E. mollis), the slightly more rays of the first dorsal fin (II,9 in E. occidentalis, 11,10 or II,11 in E. mollis), and the somewhat higher gill-raker count on the second arch (11 compared with 9-10 for E. mollis) are other noteworthy differences. Additionally, comparison of the ho- lotype of E. occidentalis with a specimen of E. mollis from the Swedish Deep-Sea Expedition of 1947-48 (reported on by Nybelin (1957) and borrowed through the courtesy of Dr. Huben- dick of the Natural History Museum, Goteborg) has revealed that the scales on the abdomen of the new species are smaller, with about seven scale rows separating the periproct from the pel- vic fin bases as compared with three at the most separating the two regions in E. mollis. The pel- vic girdle of E. occidentalis is much shorter than that of E. mollis—its length from anterior point to base of pelvic fin goes about 2.5 into distance orbit to angle of preopercle, as compared with about 2.0 for the same measurement in E. mol- lis. The new species is otherwise closely similar to its congener, for which Nybelin (1957) gives an excellent description and a photograph of two freshly caught specimens. DISTRIBUTION.—Known from only the holo- type taken off the northern coast of Peru in 4,334 m. MATERIAL EXAMINED.—Holotype: USNM 135612 (1, ca. 80 mm HL, ca. 400 mm TL); off Peru, 8°30’S, 85°36’ W, 4,334 m, ALBATROSS Sta. 4658, 14 Nov. 1904. Mataeocephalus Berg Coelocephalus GILBERT AND CRAMER, 1897:422 (non Agas- siz, 1843) (type-species Coelocephalus acipenserinus Gil- bert and Cramer, 1897, by monotypy). Mataeocephalus BERG, 1898:43 (replacement name for Coe- locephalus Gilbert and Cramer, 1897, preoccupied). DIAGNosIs.—Macrourine grenadiers with anus located within a moderately broad to broad na- ked area (periproct) whose posterior edge is im- mediately adjacent to or close to anal fin origin; anus usually closer to origin of anal fin than to insertion of pelvic fins. Spinous ray of first dor- sal fin with a serrated leading edge or serrations obsolete (in M. adjustus). Snout produced, dor- soventrally depressed, with a stout, two-pronged scute. Mouth small (less than 30 percent HL), inferior; suborbital ridge stout, sharply angular in cross section. Premaxillary and mandibular teeth in broad bands which are usually confined to anterior portion of jaws and which usually fall well short of posterior corners of mouth open- ing. Gill openings restricted; gill membranes broadly connected to isthmus. Outer gill-rakers on first arch rudimentary or absent. COMPARISONS.—Mataeocephalus appears su- perficially close to Coelorinchus Giorna but is immediately distinguished from that genus by the branchiostegal ray count of 7 (6 in Coelorin- chus), the strongly serrated spinous ray in first dorsal fin (except in M. adjustus with serrations obsolete; cf. smooth or, rarely, with few distal teeth in Coelorinchus), and the two-pronged ter- minal snout scute (one- or three-pronged in Coe- lorinchus). The genus Mataeocephalus is clos- est to Nezumia but differs in: (1) anus closer to anal fin origin than to pelvic fin insertion (anus generally closer to pelvic fin insertion in Nezu- mia); (2) premaxillary and mandibular teeth in short, broad bands except in M. adjustus (cf. narrow tapering bands that extend posteriorly to rictus except in N. burragei (Gilbert, 1905)); (3) generally longer snout, and smaller, more infe- rior mouth; and (4) outer rakers of first arch ru- dimentary or absent (cf. tubercular rakers pres- ent in Nezumia species). Members of the genus are unlikely to be confused with any other ma- crourine grenadier with the combination of sev- en branchiostegal rays and periproct adjacent or close to anal fin origin because of their small, inferior mouth, much produced snout, and sharp, angular suborbital ridge. REMARKS.—Mataeocephalus includes a small group of about five closely related species con- fined to the tropical waters of the Pacific and Indian oceans. The genus is not known from the Atlantic Ocean. Mataeocephalus adjustus (Smith and Rad- cliffe, 1912) from the Philippines appears to be the most primitive member judged by the follow- ing characters: relatively large mouth that is not strongly U-shaped; dentition in both jaws ex- tending posteriad in a tapered band; head and snout not dorsoventrally depressed to the extent found in the other members; ventral surfaces of snout and suborbital regions completely scaled; terminal snout scute rather small; and periproct area relatively small. Most of these supposedly IWAMOTO: EASTERN PACIFIC MACROURIDAE FIGure 8. 8468, collected by the ARCTURUS, 96 km south of Cocos Island. Scale bar under otolith represents 5 mm; that under tail, 25 mm. primitive features of M. adjustus cloud the oth- erwise strong differences between the genera Mataeocephalus and Nezumia, but viewing the group as a whole, and on the basis of the diag- nosis given above, recognition of each as dis- tinct genera seems Justified. Mataeocephalus tenuicauda (Garman) (Figure 8) Macrurus tenuicauda GARMAN, 1899:216-217, pl. 49, fig. 1 (original description; type-locality Gulf of Panama, 838 m, ALBATROSS Sta. 3384). Mataeocephalus tenuicauda: G1LBERT AND Huss, 1916:146 (name only). DIAGNOsISs.—A species of Mataeocephalus with 8 (9 in three fins of 18 specimens) pelvic fin rays; 22-26 pectoral fin rays. Chin barbel about 4-6 percent HL. Upper jaw 19-28 percent HL. No small naked fossa (anterior dermal window of light organ) anterior to periproct. DESCRIPTION.—General features of fish seen in Figure 8. Head shallow, depressed; greatest width of head about equal to or more than great- est depth of head. Dorsal and ventral surfaces of head sharply demarcated by a strong ridge si ra PATE Mataeocephalus tenuicauda (Garman). Composite drawing from specimens catalogued AMNH 8451, 8467, and running from snout tip posteriad to preopercle, but not connected to preopercular ridge. Mouth small, inferior, protrusible, U-shaped. Periproct region large, situated almost midway between origin of anal fin and insertion of pelvic fins, but slightly closer to former. No separate dermal window of light organ apparent in specimens examined. Swim bladder large, with two long, slender, uncoiled retia and two small, flattened gas glands. Males with large drumming muscles on each side of anterior end of swim bladder. Pyloric caeca simple, short, thick; 16 to 21 in five specimens. Intestine with two major loops, the first extending posterodorsally from pylorus, the second extending anteroventrally along dex- tral wall of abdomen, passing anteriad and si- nistrally around front of stomach and caecal mass. Opercular openings restricted dorsally and ventrally; gill membranes broadly connect- ed and attached to isthmus with no free posterior fold. A fleshy ridge on shoulder girdle along pos- terodorsal margin of gill cavity. Outer gill-rakers of first arch rudimentary, platelike, 3 or 4 in number. Dorsal fin with thornlike spinous first ray 146 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 5 upper lower FIGURE 9. View of upper and lower jaws showing shape of tooth bands in (a) Mataeocephalus tenuicauda (Garman), specimen 48 mm HL and (6) Nezumia latirostrata (Garman), specimen 40.5 mm HL. closely appressed to spinous second ray, the lat- ter with widely spaced teeth along leading edge. Outer ray of pelvic fins prolonged, extending well beyond anal fin origin. Scales on body with slender, conical spinules arranged in sharp, ridgelike rows. Scales on head variously developed; those on ridges more coarsely developed with spinule rows arranged in a Stellate pattern in some and a broad poste- riorly radiating pattern in others; those on top of head generally with low, longitudinal spinule rows that diverge slightly posteriorly. Ventral surfaces of head naked except for a small patch of scales at anteroventral corner of preopercle and along leading edge of snout where large, coarse, spinous, nonimbricate scales overlap slightly onto ventral surfaces. Tip of snout armed with two closely appressed, conical scutes. Suborbital region covered dorsally with two distinct rows of large, coarse, nonimbricate, strongly adherent scales. Supraorbital and su- pranarial ridges coarsely scaled. A prominent lunate naked groove dorsally along each side of anterior snout margin. Premaxillary teeth in broad, short, cardiform bands truncated at posterior ends (Fig. 9a). Mandibular teeth in a broad, short band with tapered ends; tooth band falling well short of lateral corners of mouth. Coloration in alcohol. All specimens exam- ined have lost most of their body scales. Overall color brown to swarthy, often with a violet tinge on trunk and tail. Abdominal region blackish. Naked ventral surfaces of head pale to dusky. Fins dusky to blackish. Oral, branchial, and peritoneal cavities blackish. Counts (from 18 specimens unless otherwise indicated).—1D. II,8—10; 1P. 22-26 ( = 23.76); 2P. 8-9 (9 rays in 3 of 36 fins). Gill-rakers on first and second arch 0-1 + 6-7 (6-8 total; x = 7.18); on second arch 0-1 + 5-7 (6-8 total; x = 7.29). Scales below first dorsal 7-11 (6 speci- mens); below midbase of first dorsal 6% to 9 (6 specimens); below second dorsal 8% to 11 (9 specimens); over distance equal to predorsal length of head 37-48 (4 specimens). MEASUREMENTS (from 18 specimens unless otherwise indicated).—Total length 131+ to 303 mm; head length 30.0-65.3 mm. The following in percent of head length: postrostral length of head 60-66 (% = 61.8; S.D. = 1.56); snout length 36-42 (« = 39.9; S.D. = 1.46); preoral length 35—43 (¢ = 38.6; S.D. = 2.31); internasal width 19-22 (€ = 20.5; S.D. = 0.80); orbit di- ameter 26-30 (¢ = 27.2; S.D. = 1.48); interor- bital width 19-22 (¢ = 20.3; S.D. = 0.93); post- orbital length 29-36 (¢ = 32.7; S.D. = 1.51); orbit to angle of preopercle 28-33 (¢ = 30.0; S.D. = 1.25); suborbital width 13-16 ( = 14.6; S.D. = 0.96); upper jaw length 19-28 (¢ = 22.4; S.D. = 2.07); barbel length 3.7-5.7 (« = 4.7; S.D. = 0.61); length outer gill slit 8.5-—12.7 (@ = 10.4; S.D. = 1.08; n= 13); preanal length 122- 138 (¢ = 129.5); outer 2P. to A. 21-29 @ = 25.9; S.D. = 2.16; n = 15); greatest body depth 42- 56@ = 52.2; S.D. = 3.4720 = 14) 102) ein- terspace 19-29 (x = 25.1); height 1D. 52-63 @ = 58.5; n = 8); length 1P. 37-41 @ = 38.7; n = 12); length 2P. 37-49 @ = 43.2). COMPARISONS AND RELATIONSHIPS.—Matae- ocephalus tenuicauda closely resembles the Hawaiian endemic M. acipenserinus (Gilbert and Cramer, 1897) but differs in having a longer upper jaw (22-28 percent HL vs. about 19-21.5), a somewhat longer chin barbel (3.7—5.7 percent HL vs. 2.64.0), and in lacking a small naked fossa anterior to the periproct. M. nigrescens (Smith and Radcliffe, 1912) from the Philippines also closely resembles both M. tenuicauda and M. acipenserinus, and the three are considered as close allies by Gilbert and Hubbs (1920:564). My cursory examination of five paratypes of M. nigrescens (USNM_ 149310; 149311 [2 speci- mens]; 149312: 149313) revealed minimal differ- ences between nigrescens and tenuicauda. Morphometric features of the two are indistin- IWAMOTO: EASTERN PACIFIC MACROURIDAE guishable. Scale-row counts, however, show no- table separation; nigrescens specimens had about 7/2 rows between the origin of the second dorsal fin and 5’ below the midbase of the first dorsal fin, whereas tenuicauda specimens had 8% to 11 and 6% to 9, respectively. Mataeocephalus adjustus (Smith and Radcliffe, 1912) is readily distinguished from tenuicauda, nigrescens, and acipenserinus by the following characters: fewer pelvic rays (7 cf. 8 or 9), fewer pectoral rays (19 or fewer cf. 21-26), bands of teeth in both jaws extending posteriad about to end of rictus. Because of the many features dis- tinguishing M. adjustus on the one hand and M. tenuicauda, M. nigrescens, and M. acipenseri- nus on the other, a wide phylogenetic diver- gence between the two groups Is suggested. The lack of specimens of M. microstomus (Regan, 1908) from the Indian Ocean and the paucity of information in the original description preclude an adequate comparison of this species with its congeners. DISTRIBUTION.—Mainland Pacific coasts of Panama and Ecuador between latitudes 6°36'N and 3°15’S; in the Galapagos; and south of Co- cos Island. S1zE.—To at least 65 mm HL and 303 mm TL. MATERIAL EXAMINED (55 specimens, 7 localities).—Pana- ma: USNM 148879 (2, 57-59 mm HL, 202-253 mm TL), 6°36'N, 81°45’W, 581 fms (1,063 m), ALBATROSS sta. 4621, 21 Oct. 1904. Ecuador: CAS 38325 (1, 34 HL, 155 TL), 3°15’S, 80°5S'W, 945-960 m, ANTON BRUUN cr. 18B, sta. 770 (field no. LWK66-120), 10 Sep. 1966. Cocos Island (60 miles [96 km] S of): AMNH 8451 (7, 35-64 HL), AMNH 8467 (6 spec- imens), AMNH 8468 (23 specimens), AMNH 8469 (5 speci- mens), AMNH 8470 (7 specimens), all from ARCTURUS Sta. 74, May 1925. Galapagos: CAS-SU 25239 (1, 55 HL, 230 TL), 0°29'S, 89°54'30’W, 392 fms [717 m], ALBATROSS Sta. 2818, 15 Apr. 1888.—USNM 135340 (1, 43 HL, 185+ TL), 0°36'30'S, 89°19'00"W, 634 fms [1,159 m], ALBATROSS sta. 2808, 4 Apr. 1888.—CAS 42075. (2, 30-31 HL, 158-131+ TL), 1°06’S, 89°22’W, 700-800 m, TE VEGA cr. 19, sta. 102, 12 Apr. 1968. Paracetonurus Marshall Paracetonurus MARSHALL, 1973:615 (type-species Macrurus parvipes Smith and Radcliffe, 1912, by original designation). DIAGNOsSIS.—A macrourine grenadier with anus and urogenital openings within a broad, black periproct immediately preceding anal fin origin. Spinous ray of first dorsal fin with a ser- rated leading edge. Snout relatively high, broad; suborbital region relatively deep, without a strong spinous ridge. Scales flanking base of sec- ond dorsal fin not enlarged. (Adapted from Mar- shall 1973.) 147 REMARKS.—Marshall (1973:615) erected this genus to include Macrurus flagellicauda Koe- foed, 1927, M. parvipes Smith and Radcliffe, 1912, and Lionurus cetonuropsis Gilbert and Hubbs, 1916—three species that are closely re- lated to Cetonurus Giinther, 1887, but which are distinguished by their somewhat less-inflated head and their lack of enlarged scales along the base of the second dorsal fin. The monotypic genus Kumba Marshall, 1973, is also closely re- lated to Paracetonurus but differs primarily in lacking serrations on the second spinous ray of the first dorsal fin. I have included Macrurus fragilis Garman, 1899, in Paracetonurus for reasons given in the description of that species. By doing so, the def- inition of the genus is expanded and its contrast with the genera Cetonurus and Kumba is less- ened. I recognize, however, that a detailed study comparing features of P. fragilis and the three other species of the genus may necessitate a fur- ther rearrangement of the taxon, but the material available does not allow such a study at this time. Paracetonurus fragilis (Garman) (Figure 10) Macrurus fragilis GARMAN, 1899:203-204, pl. 46, fig. 1 (orig- inal description; ALBATROSS specimens from off Panama and Colombia, 3,058—3,334 m). Lionurus (Lionurus) fragilis: GILBERT AND Huss 1916:146 (listed). Sphagemacrurus fragilis: MARSHALL 1973:623 (listed). DIAGNOSIS.—A species of Paracetonurus with 8-9 pelvic rays. Nasal rostrum extremely fragile; head covering thin, almost membranous. Counts.—1D. II,6-9; 1P. 19-21; 2P. 8-9. Gill-rakers on inner side of first arch 1-2 + 8- 10 (10-11 total); on inner side of second arch I- 2 + 8-9 (9-11 total). MEASUREMENTS (from three specimens; most measurements are estimates).—Head length about 26-35 mm; total length 175+ to 200 mm. The following in percent of head length: snout length about 28-40; orbit diameter about 22-23; interorbital width 24-27; orbit to angle of pre- opercle 33-44; suborbital width 14-18; length upper jaw 31-39; length barbel 19-24; length outer gill-slit 17-20; greatest body depth 66-69. DESCRIPTION.—Head moderately wide, trunk short, tail very long and straplike. Head bones thin, fragile, particularly nasal rostrum, which most often is bent to side giving appearance of 148 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 5 Figure 10. Paracetonurus fragilis (Garman). (a) Diagrammatic reconstruction from paratype MCZ 28585 and UMML uncat. (b) SIO 55-265, snout reconstructed. (c) Stomach and intestines of UMML uncat. (d) Retia and gas glands of UMML uncat. Scale rule for a and b equals 25 mm. blunt snout. Snout relatively high and wide. In- terorbital region broad, width usually greater than diameter of orbit. Suborbital region deep, without a strongly angular ridge. Mouth mod- erately large, upper jaws extend to vertical through hind margin of orbits or slightly beyond. Barbel moderately long, slender, length equal to or slightly greater than length outer gill-slit. Gill Openings extend forward to below posterior third of orbit. Gill membranes little restricted (probably with a moderate free fold posteriorly over isthmus). Gill-rakers tubercular, gill lamel- lae short. Pyloric caeca short, thick, 10-15 in UMML specimen. Intestine short, only two loops from pyloric caeca to anus. Esophagus and caecal portion of stomach black; pyloric portion of stomach grayish brown. Rectum black, remainder of intestine and pyloric caeca pallid. Retia long, well developed, 2; gas glands 2. First dorsal fin short, base low; second spi- nous ray weakly serrated and produced beyond segmented rays. Second dorsal fin rudimentary, originating posterior to vertical through origin of anal fin. Pectoral and pelvic fins small; the latter far forward, origin below preopercle angle, with outer ray slightly produced and extended past origin of anal fin. Snout and probably most of anterior and ven- tral parts of head naked and covered with thin black integument. (I could find no scale pockets on the head of specimens I examined, but Gar- man’s (1899:pl. 46, fig. 1) figure suggests that there are some over the gill covers.) Garman (1899:204) describes the scales as “‘small, thin, deciduous, with concentric striae; five scales in a row from the lateral line to the base of the first dorsal or thirty-one in a series from this dorsal to the anal.”’ Garman’s figure of a scale shows a single point on the posterior edge, but an oth- erwise spinuleless exposed field. Teeth short, slender, sharp, in roughly two rows in premaxillae, in narrow band in mandi- bles; no enlarged series. Coloration in alcohol. Head and abdominal region black, remainder of body brownish. Oral, branchial, and peritoneal cavities black. Gill arches and rakers blackish but lamellae pallid. DISTRIBUTION.—The species is known only from the tropical eastern Pacific where it is found in waters of considerable depths. Speci- mens were obtained in three trawls fished on bottom in 3,058—3,334 m, and a single specimen was taken in an open midwater trawl fished to a depth of 1,335 m. COMPARISONS AND REMARKS.—As the name implies, members of this species are extremely fragile. The few specimens available for exami- IWAMOTO: EASTERN PACIFIC MACROURIDAE nation were in such poor condition that an ad- equate description and illustration could not be prepared. The thin, flexible rostrum is often damaged and folded to the side (as is the rostrum on the specimen illustrated by Garman 1899:pl. 46, fig. 1), but when intact, the rostrum is mod- erately long and pointed. The fragile head, and particularly the rostrum, is highly reminiscent of the condition obtaining in specimens of Hymen- ocephalus, but members of that genus and P. fragilis differ in many other fundamental fea- tures. The small adult size of P. fragilis is in- dicated by the ripe condition of a 200-mm-TL female paratype (MCZ 28585), whose ovaries contained eggs as large as 1.3 mm in diameter. This species is quite unusual and may repre- sent a genus distinct from Paracetonurus, but it is placed in that taxon out of convenience, because an adequate study comparing it with other related forms could not be made. It differs from other members of Paracetonurus in having a less inflated head; thin, membranous, and mostly naked head covering (cf. moderately thick, completely scaled head covering); more pelvic fin rays (8-9 cf. 6-7); and a larger mouth (upper jaw extends posteriad to below hind third of orbits, cf. below middle third of orbits). The species is also fairly close to members of the genus Sphagemacrurus Fowler (in which Mar- shall (1973) has placed the species), but it differs in lacking a strong spinous suborbital ridge, a high first dorsal fin base, and a short blunt snout—all characteristic of Sphagemacrurus. The origin of the vent is also more anteriorly placed in species of Sphagemacrurus (below anterior third of first dorsal fin compared with below hind edge of first dorsal fin in P. fragilis). MATERIAL EXAMINED (7 specimens, 4 localities) —Pana- ma: MCZ 28586 (holotype, about 230 mm TL); 6°17'N, 82°0S'W, 3,058 m, beam trawl, ALBATROSS sta. 3360, 24 Feb. 1890. —UMML uncat. (1, about 26 HL, 175+ TL); 6°53'N, 79°27'W, 3,193 m, R/V PiLtsBury sta. 526, 5 May 1967. Colombia: MCZ 28585 (3 paratypes, about 29-35 HL, 190+ to 200 TL) and USNM 57857 (1, est. 26 HL, est. 175 TL); 2°35'N, 83°53'W, 3,334 m, beam trawl, ALBATROSS sta. 3374, 3 Mar. 1890. Eastern Pacific: SIO 55-265 (1, 117 TL); 00°02’S, 100°23'W, 0-1,335 m, 3-m midwater trawl, R/V HoriZon sta. ET(b)-H-65. Malacocephalus Gunther Malacocephalus GUNTHER, 1862:396 (as subgenus of Macru- rus) (type-species Macrourus laevis Lowe, 1843, by mono- typy). DIAGNOSIS.—A macrourine grenadier with 149 anus remote from anal fin and closer to pelvic fins; periproct large. Two large dermal windows of light organ, the anterior one in a bean-shaped depression situated between bases of pelvic fins, the posterior one in a shallow circular depres- sion close before the anus. Teeth large, widely spaced, in single row in lower jaw; usually larger posteriorly. Teeth in upper jaw in two rows or in narrow band; outer series distinctly spaced and enlarged. Pyloric caeca numerous (50-100 or more), multiply branched. Lowermost three or four branchiostegal rays scaled. Mouth large, upper jaw usually more than 45 percent of head length. No strongly developed scutelike scales on head ridges. REMARKS.—Malacocephalus comprises a close-knit group of about six species, one of which is undescribed (see Iwamoto 1970:410). Relationships of the genus lie closest to Ventri- fossa Gilbert and Hubbs, 1920, a taxon repre- sented in the eastern Pacific by a single member (of the subgenus Lucigadus), although the genus is common in most other warm-water areas. Members of the genus Malacocephalus are con- fined to upper-slope waters of tropical and warm-temperate regions. Three of the six species (laevis (Lowe, 1843), nipponensis Gil- bert and Hubbs, 1916, and hawaiiensis Gilbert, 1905) are closely related and may eventually prove to represent one widely distributed species. Okamura (1970a:69) has, in fact, syn- onymized M. nipponensis with M. laevis. Malacocephalus laevis (Lowe) (Figure 11) Macrourus laevis Lowe, 1843:92 (original description; off Madeira). Malacocephalus laevis: GUNTHER 1862:397-398. —HuBBs, FOLLETT, AND DEMPSTER 1979:14 (list; first record from eastern Pacific). Macrurus (Malacocephalus) laevis: GUNTHER 1887:148, pl. 38, fig. b. See Marshall 1973:653 for extensive synonymy. Malacocephalus sp.: HUBBS AND IWAMOTO 1977:243 (1 spec. from California). DIAGNosIs.—A species of Malacocephalus with spinous second ray of first dorsal fin smooth; teeth in upper jaw in two distinct rows; pectoral fin rays 19-20; upper jaw less than 50 percent of head length. DESCRIPTION OF EASTERN PACIFIC SPECI- MEN.—General features of fish seen in Figure 11. Head compressed and deep. Ridges not sharp or coarsely scaled; head contours smooth- ly rounded. Snout narrow, pointed (in somewhat 150 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 5 FiGurReE 11. shriveled eastern Pacific specimen; probably rounded in life). Gill openings wide, extending anteroventrally to below hind margin of orbit. Gill membranes loosely connected to isthmus with a posterior free fold. Interopercle broadly exposed and scaled ventrally. Periproct region large, long-ovate, anus and urogenital opening at broader posterior end of region; anterior der- mal window of light organ in deep, lunate to bean-shaped fossa between bases of pelvic fins and separated from periproct by a broad isthmus of scales. Several partially digested euphausids in stomach. Pyloric caeca fragile, thin, branched, difficult to count, but more than 50 terminal dig- its. Scales uniformly small and finely spinulated. Snout and suborbital region completely scaled, without prominent naked areas. Coarse, scute- like scales completely absent; ridges of head all Malacocephalus laevis (Lowe). SIO 79-344, 58 mm HL, from off Santa Barbara County, California, in 229 m. smoothly rounded. Lower jaw uniformly scaled. Lowermost branchiostegal rays scaled; gular membrane apparently without scales (but scales here normally highly deciduous and may have been present in life). Color gray-brown overall; violet over abdom- inal region, shoulder girdle, and opercle; and blackish over chest, on branchiostegal mem- brane, and on gular region (violet regions prob- ably not as prominent in undenuded and fresh specimens). Peritoneum brownish black. Oral valves and lower lips peppered with melano- phores, remainder of oral cavity pale. First dor- sal fin, pectoral fin, pelvic fin, and anterior por- tion of anal fin brownish black; second dorsal fin and most of anal fin pale or dusky. Counts.—1D. II,11; 1P. 20/19; 2P. 9/8. Gill- rakers on first arch 3 + 9; on second arch 3 + 7. Scales below first dorsal approximately 16. IWAMOTO: EASTERN PACIFIC MACROURIDAE MEASUREMENTS.— [I otal length 335 mm; head length 58 mm. The following in percent of head length: postrostral length of head 78; snout length 26; preoral length 17; internasal width 17.5; orbit diameter 36; interorbital width 27; postorbital length of head 41; distance orbit to angle of preopercle 43; suborbital width 12; up- per jaw length 48; barbel length 17; length first gill-slit 26; preanal length 123; outer pelvic ray to anal origin 28; anus to anal origin 17; greatest body depth 85; interspace between 1D. and 2D. 48. COMPARISONS AND RELATIONSHIPS.—I have tentatively identified this single eastern Pacific specimen of Malacocephalus as M. laevis. All morphometric and meristic characters examined fell within the range of variation enumerated in a previous paper (Iwamoto 1970) for specimens I examined from the Atlantic Ocean. However, the snout length and interorbital width in the Pacific specimen were at the lower limits of the range for these characters in Atlantic specimens, and the orbit diameter was at the upper limit. Close comparison of the eastern Pacific speci- men with others from the Atlantic Ocean showed only one difference—scales were absent on the gular membrane whereas most Atlantic specimens had a small patch of scales there. The deciduous nature of the scales in this area, how- ever, may account for their absence in the some- what denuded eastern Pacific specimen. Com- parison of SIO 79-344 with specimens of M. hawatiensis (USNM 51618, holotype; CAS-SU 8522, 3 paratypes) showed slightly lower values in relative snout length (26 percent head length cf. 28-29), preoral length (17 cf. 19-21), inter- nasal width (17.5 cf. 21-25), interorbital width (27 cf. 31-35), length snout to anus (111 cf. 117- 120), and 1D.-2D. interspace (48 cf. 57-75). Similar comparison of proportional measure- ments with specimens of M. nipponensis (FAKU 13316, 13318, 13321, and 13878) showed lower values in the eastern Pacific specimen in snout length (26 percent HL cf. 28-31), preoral length (17 cf. 20-21), interorbital width (27 cf. 30-35), and barbel length (17 cf. 21-24), but a greater value for the orbit diameter (36 cf. 29- 34). The gill-raker count of 12 was also slightly higher than the 10-11 of the M. nipponensis specimens. Whether or not these slight differ- ences are meaningful in delimiting species is as yet conjectural. Adequate series of each popu- lation should be compared before definitive 151 statements are made regarding the number of valid species in the genus. Until then it seems wisest to retain the established names. REMARKS.—Malacocephalus laevis is listed by Hubbs, Follett, and Dempster (1979:14) in their “‘List of the fishes of California’ on the basis of the present specimen. Mr. Eiichi Fujii has informed me (in litt., 6 Mar. 1979) of having collected specimens of the species from sea- mounts off the coast of Baja California while aboard the KAryo MARU in January and Feb- ruary of 1979. MATERIAL EXAMINED.—SIO 79-344 (1, 57.8 mm HL, 335 mm TL), California, Santa Barbara County, between Gav- iota and Point Conception, 229 m, otter trawl, 10 Dec. 1968. Ventrifossa Gilbert and Hubbs Ventrifossa GILBERT AND Huss, 1920:553 (type-species Cor- yphaenoides garmani Jordan and Gilbert, 1904, by original designation). DIAGNOSIS.—A genus of macrourine grena- dier with anus removed from anal fin origin and closer to pelvic fin insertion; anus situated pos- teriorly in an oval-shaped area of naked black skin (the periproct) that extends forward to a small fossa (anterior dermal window) between pelvic fin bases. An often-inconspicuous lenslike posterior dermal window in front of anus. Sec- ond spinous ray of first dorsal fin slightly pro- duced and finely toothed along leading edge (ex- cept in V. atherodon). Jaw teeth small, in narrow to moderately broad bands in upper jaw, outer series slightly enlarged in most species; lower-jaw teeth small, none enlarged, in one to several irregular series laterally. Branchiostegal membranes naked (in subgenus Ventrifossa) or usually with patches of scales (in subgenus Lu- cigadus) along exposed lowermost branchioste- gal rays. No sharp, coarsely scaled, angular ridges on head; suborbital region with flat to gently rounded contours—scales here not mod- ified to form stout shelves or ridges (as in Ne- zumia). Snout angular to rounded in profile, either without a spiny terminal scute or with a small unilateral one; supranarial ridge without modified scutelike scales; lateral angles of snout without spinous tubercular scale at tip. Body and head scales generally small, densely cov- ered with fine, slender, conical or shield-shaped spinules. REMARKS.—Ventrifossa as here diagnosed comprises three major groups which I treat as subgenera: Ventrifossa, Lucigadus, and Soko- 152 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 5 dara (new). In describing Ventrifossa, Gilbert and Hubbs (1920:543) recognized four subgen- era, three of which were monotypic: Atherodus (with Optonurus atherodon Gilbert and Cramer, 1897), Lucigadella (with Macrourus nigromar- ginatus Smith and Radcliffe, 1912), and Lugi- gadus (with Macrourus lucifer Smith and Rad- cliffe, 1912). Atherodus was distinguished from the other subgenera on the basis of dentition (teeth of lower jaw in two series, arrowhead- shaped canines on upper jaw), scale spinules (few and short), orbit size (3 in head), and dorsal spine (without denticulations). My studies of the genus have shown that all but the last character are nondiagnostic for the subgenus in that the character states are shared with other members of the genus, or that they are part of a graduated character spectrum within the genus. The ab- sence of denticulations on the dorsal spine ap- pears to be the sole character distinguishing V. atherodon from other members of the genus. The sister-group relationship of V. atherodon and V. ctenomelas (Gilbert and Cramer, 1897) is strongly suggested in other shared specializa- tions and in the common occurrence of the two species in the Hawaiian Islands, where V. ath- erodon is apparently endemic (V. ctenomelas is also found in the western Pacific). Characters that Gilbert and Hubbs (1920) used to separate V. lucifer into a distinct subgenus, Lucigadus, include the ventral light organ (*‘a conspicuous pearly body in a sheath’’), mouth angle (“highly oblique’’), and spinules on scales (‘in quincunx order’’). My reexamination of the type-specimens of V. lucifer has shown that the ‘*sheath’’ in which the ‘‘pearly body” lies is an artifact of preservation, the ‘‘sheath’’ having formed by the ripping midventrally of the thin abdominal wall below the light organ. The pearly color of the organ is from the reflective layer surrounding the organ. The light organ in V. /u- cifer is not notably different from that of other closely related members of the genus, including V. nigromarginata. The arrangement of spinules on the scales appears to show no meaningful relationships in this genus nor in the closely re- lated Nezumia—the character is useful, how- ever, at the species level. The notably oblique mouth of V. lucifer represents one extreme in a graded series that includes—from mouth little oblique to mouth notably oblique—Macrourus nigromarginatus, Macrurus fasciatus Weber, 1913, Macruroplus ori Smith, 1968, Macrourus nigromaculatus McCulloch, 1907, and Macrou- rus lucifer. 1 consider these five species as rep- resentatives of the subgenus Lucigadus Gilbert and Hubbs. The subgenera of Ventrifossa as | recognize them can be characterized in the following man- ner: Subgenus Ventrifossa Gilbert and Hubbs, 1920:553 (type-species Coryphaenoides garmani Jordan and Gilbert, 1904). 1. Mandibular teeth in one to three irregular series laterally. Premaxillary teeth in a narrow band with outer series slightly to prominently en- larged; tooth band extends posteriad be- yond maxillary process. 3. Mouth slightly oblique, moderate to large, upper jaw length 42—53 percent head length. 4. Tip of snout with a blackish spot, or entire leading edge black. . Branchiostegal and gular membranes com- pletely naked. 6. Gill-rakers 13-20 total on inner series of out- er (first) arch. 7. Pores of cephalic lateral line system small and inconspicuous. 8. Pyloric caeca numerous, 40-70 in distal count, usually branched near base. 9. Snout without spinous tubercular scute at tip. 10. Abdominal vertebrae 11-12. 11. Ventral aspects of body not appearing to have shifted notably forward. 12. Infraorbital shelf not exceedingly narrow anteriorly. i) ‘n Included species: V. atherodon (Gilbert and Cramer, 1897), V. ctenomelas (Gilbert and Cra- mer, 1897), V. divergens Gilbert and Hubbs, 1920, V. garmani (Jordan and Gilbert, 1904), V. macropogon Marshall, 1973, V. mucocephalus Marshall, 1973, V. petersonii Alcock, 1891, and one or two undescribed speces (specimens in the CAS collection). Subgenus Lucigadus Gilbert and Hubbs (type- species Macrourus lucifer Smith and Rad- cliffes 1912): 1. Mandibular teeth in several irregular series laterally or in a narrow to moderately wide band. IWAMOTO: EASTERN PACIFIC MACROURIDAE 2. Premaxillary teeth in a narrow to moderate- ly wide band with outer series slightly en- larged; teeth not present posteriad of hind margin of maxillary process. 3. Mouth slightly to greatly oblique, moderate in size, 33-45 percent HL. 4. Tip and leading edge of snout without dis- tinct pigmentation. 5. Lowermost of branchiostegal rays scaled. 6. Gill-rakers 7—16 total on inner series of outer (first) arch. 7. Pores of cephalic lateral line system prom- inent in most species. 8. Pyloric caeca 30-55, generally unbranched. 9. Snout tip without stout, spinous, scutelike scale. 10. Abdominal vertebrae 10-11. 11. Ventral aspects of body appearing to have migrated far forward resulting in anal origin below first dorsal; pelvic origin below oper- culum; gill membranes united below orbits; snout high, rounded; base of first dorsal high. 12. Infraorbital shelf not especially narrow an- teriorly. Included species: V. fasciata (Weber and de Beaufort, 1929), V. lucifer (Smith and Radcliffe, 1912), V. nigromarginata (Smith and Radcliffe, 1912), V. nigromaculata (McCulloch, 1907), and V. ori (Smith, 1968). Subgenus Sokodara Iwamoto, new subgenus (type-species Coryphaenoides misakius Jordan and Gilbert, 1904). 1. Mandibular teeth in a narrow band of two to three irregular series laterally. 2. Premaxillary teeth in a narrow band; outer series scarcely enlarged. 3. Mouth slightly oblique, large, 35-42 percent head length. 4. Tip of snout blackish. 5. Branchiostegal and gular membranes com- pletely naked. 6. Gill-rakers 14-16 total on inner series of out- er (first) arch. 7. Pores of cephalic lateral line system small and inconspicuous. 8. Pyloric caeca 54-65 in distal count, branched near base. 9. Snout with a small, unilateral, spinous scute at tip. 10. Abdominal vertebrae 14. 11. Ventral aspects of body not appearing to have shifted notably forward. 12. Infraorbital shelf greatly narrowed anterior- ly. Included species: V. misakia and one or two undescribed species (specimens in CAS and USNM collections). DISTRIBUTION.—The genus Ventrifossa is represented in the eastern Pacific by one wide- ranging, Southern Hemisphere species, V. (Lu- cigadus) nigromaculata. That representatives of the more diverse subgenus Ventrifossa are en- tirely absent from this region forms a striking parallel with a similar situation in the eastern Atlantic, where the genus is entirely unknown (except off Cape Point, South Africa)—this de- spite the presence in the western Atlantic of two species of subgenus Ventrifossa (V. macropo- gon and V. mucocephalus) and one of subgenus Lucigadus (species cf. V. ori). Ventrifossa is known in most other warm-water regions of the Pacific and Indian oceans. Ventrifossa nigromaculata (McCulloch) (Figures 12a, 18d) Macrourus nigromaculatus MCCULLOCH, 1907:346—348, pl. 63, figs. 1, la (original description; holotype and 4 para- types; 56 km E of Sydney, Australia, in 1,463 m). Lionurus nigromaculatus: MCCULLOCH 1919:32 (listed), pl. 11, fig. 114a. ?Macruroplus nigromaculatus: SMITH 1949: 135 (bref de- scription; southern Africa). Nezumia nigromaculata: MAKUSHOK 1967:207 (name). Macruroplus potronus Pequeno, 1971:290-291, fig. 15 (origi- nal description; holotype only, off Chile, 34°58’S, 72°36'W, in 200 m). DIAGNosIs.—A species of Ventrifossa, sub- genus Lucigadus, with 13-15 pelvic fin rays. Total gill-rakers on first arch 12-16; on second arch 12-14. Upper jaw length 39-45 percent HL. Scales below first dorsal fin about 16-20; below second dorsal fin 10-12. First dorsal fin with a prominent black blotch. DESCRIPTION.—General features of the fish are best seen in Figure 12a. Gill membranes unite across isthmus at a point below middle of orbits and slightly ahead of posterior end of maxillae. Head ridges virtually nonexistent; all surfaces smoothly rounded; suborbital region almost flat. Pores on head well developed but not especially prominent. Pyloric caeca well developed, numerous. rather large; unbranched except at very base. 154 PROCEEDI NGS OF TH E CALIFORNIA ACADEMY OF SC IENCES, V. , Vol. 42, No. 5 NAD ‘eo je \\\ AN AY \\\ NIN ° QW WW As KQQ0 \ IWAMOTO: EASTERN PACIFIC MACROURIDAE Configuration of intestinal tract a rather simple ‘S’ pattern. Gas glands a flattened kidney shape; mesial surfaces of the two glands closely abut- ting; retia attached mesially at about middle of each gland. Retia slender, short, uncoiled. Spinules on body scales very small, slender, conical, greatly reclined, and arranged in more or less parallel horizontal rows in middle part of vertically elongate exposed field. Spinule rows number 11-12 on larger scales of CAS speci- mens. Scales present on lowermost three or four branchiostegal rays and also densely cover ex- posed ventral surfaces of interopercle. Anterior dermal window of light organ large, situated between pelvic fin bases, and connected to periproct by isthmus of naked black skin. Posterior dermal window relatively prominent in most specimens examined, discernible as a discrete translucent area lying immediately an- teriad of anus and occupying most of anterior portion of periproct. Fins well developed. First dorsal fin large; height about equal to or greater than length of head. Anal fin high along almost entire length. Pelvic fins broad but not especially long; outer ray slightly produced, extending slightly beyond anal fin origin. Coloration in alcohol. Ground color tawny to grayish. Trunk region with violet tinge becoming bluish over abdomen and blackish around peri- proct and pelvic fins. First dorsal fin with a prominent black blotch on distal one-half to two- thirds; blotch not extending to margin of fin. Anal rays lightly punctate anteriorly and black- ish along distal margins. Median fins blackish posteriorly towards tail tip. Pelvic fins blackish or covered with bold punctations. Gill covers and gill membranes blackish or densely punc- tate. Oral cavity generally pale with few scat- tered small areas blackish or densely punctate. Outer wall of gill cavity black along first gill-slit and around outer margins, but pale otherwise. Gill-rakers and gill-arches blackish, but gill fil- aments pale. Lips dusky to somewhat blackish. Counts (from 13 specimens).—1ID. II,10-11: IP. 19-23; 2P. 13-15 (one specimen with 12 on left fin, 13 on right fin). Total gill-rakers on first arch 12-16; on second arch 12-14. Scales below first dorsal about 16-20; below second dorsal 10-12; over distance equal to predorsal length 42-45. Pyloric caeca 30, 32, and about 57 in three specimens. MEASUREMENTS.—Iwelve specimens exam- ined ranged 17-50 mm HL; 94+ to 339 mm TL. The following are in percent of head length: postrostral length 73-81 (t= 77.2; S.D. = 2.57); snout length 25-30 (¥ = 28.1; S.D. = 1.55); ven- tral length of snout 18-22 = 19.1; S.D. 1.38); orbit diameter 40-47 (* = 42.9; S.D. 2.71); interorbital width 20-26 (¢ = ee Se DE 2.07); suborbital width 12-18 (¢ = 15.7; S.D. 2.14); length upper jaw 39-45 (¢ = 7 Se, 1.80): length barbel 18-26 (usually 20-26); Mee outer gill-slit 23-30 (¢ = 26.0; S.D. = 1.68); preanal length 140-158; outer pelvic ray to anal origin 41-54; anus to anal origin 20-30; greatest body depth 92-112 (usually over 100); depth over anal origin 75-105; 1D.—2D. interspace 32- 58; height 1D. 101-134; length 1P. 63-73; length 2P. 44-58. REMARKS.—Macruroplus potronus Pequeno, 1971, was described from a single specimen tak- en off Chile in 200 m. The specimen was ex- amined and photographed by Dr. William N. Eschmeyer in 1974. Examination of his photo- graphs and the original description leaves no doubt as to the conspecificity of that specimen with those I have examined from the eastern Pacific and called Ventrifossa nigromaculata. Should these eastern Pacific specimens prove distinct from western Pacific specimens now re- ferred to this species, the name potronus re- mains available. Ventrifossa nigromaculata is a distinctive species widely distributed in temperate waters of the South Pacific. Although considerable variation was found in several characters in the specimens examined, the material available did not suggest a difference in populations from op- posite sides of the Pacific. Atlantic Ocean spec- <_< Ficure 12. (a) Ventrifossa nigromaculata (McCulloch). Specimen CAS 41668, 35 mm HL, from off Chile in 750 m. (6) Nezumia liolepis (Gilbert), CAS-SU 21402, 55 mm HL, from off Santa Cruz Island, California in 1,397-1,629 m. Enlarged views of scales from interorbital region and region just below anterior end of second dorsal fin. (c) Nezumia pulchella (Pequeno), CAS 28763, 46 mm HL, from off Peru in 272 m. (d) Nezumia stelgidolepis (Gilbert), CAS 33109, 56 mm HL, off Pescadero Point, California, in 439 m. 156 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 5 imens previously referred to this species are ap- parently distinct and may represent an undescribed species. DISTRIBUTION.—Australia, New Zealand, and Chile, in 230-1,463 m. S1z—E.—To about 50 mm in head length and about 340 mm total length. MATERIAL EXAMINED (13 specimens, 8 localities) —Chile: USNM uncat. (2, 20-35 mm HL, 140-246 mm TL), NW of Valparaiso, ca. 400 m; M/V Ruiz I; 11 Feb. 1966. —CAS 41669 (1, 33 HL, 240 TL), 32°08.5’S, 71°43’W, 960 m, ANTON BRUUN cr. 18A, sta. 703 (field no. LWK66—-47), 12 Aug. 1966. —CAS 41668 (1, 35 HL, 262 TL), 34°06.5’S, 72°18.5’'W, 750 m, ANTON BrRUUN cr. 18A, sta. 687 (field no. LWK66-25), 5 Aug. 1966. New Zealand: BMNH 87.12.7.118 (1, 17 HL, 94+ TL), CHALLENGER Sta. 166, 275 fms (503 m). —LACM 11336-3 (1, 38 HL, 280 TL), 41°35’S, 175°00’E, 256-490 m, ELTANIN Sta. 1848, 19-20 Dec. 1961. —LACM 10968-12 (5S, 25-37 HL, 192- 283 TL), 44°00’S, 178°06’W, 230-421 m, ELTANIN sta. 1398, 29 Nov. 1964. —FAKU 42147 (1, 41 HL, 300 TL), Karyo MARU Sta. 29, July 1968. —FSFRL B3052 (1, 50 HL, 339 TL), 44°20.5'S, 179°17.5'W, 750 m, Katyo MARu sta. 33, 15 July 1968. Nezumia Jordan Nezumia JORDAN, 1904, in Jordan and Starks: 620 (type- species Nezumia condylura Jordan and Gilbert, 1904, by original designation). ?Macruroplus BLEEKER, 1874:369 (type-species Macrourus serratus Lowe, 1843, by monotypy) (nomen nudum; see Poll 1953:238). Lionurus:auct. (non Gunther, 1887). DIAGNOSIS.—A macrourine grenadier with anus removed from anal fin origin (closer to pel- Vic fin insertion in most species) and situated in an oval-shaped area of naked black skin (the periproct). A small fossa (anterior dermal win- dow of ventral light organ), round to teardrop shaped, forming anterior point of periproct in most species, detached from periproct in some species. Teeth small, in narrow to broad bands in both jaws; outer series of teeth in upper jaw of most species uniformly spaced and slightly enlarged. Teeth on premaxillary bone do not arise posteriad of maxillary process. Mouth moderate to small, upper jaw length less than 40 percent of head length in most species. Snout pointed or bluntly rounded, with stout, spiny, tuberclelike scales at tip and lateral angles, prominent in almost all species. Barbel present. Suborbital shelf covered with two (in most species) or more rows of stout, deeply embed- ded (except in N. liolepis), nonimbricate, spi- nous scales; these scales form a prominent edge or crest that runs longitudinally from lateral an- gle of snout to preopercle bone and divides sub- orbital region into upper and lower halves. Body scales covered with needlelike to shield-shaped spinules (spinules almost obsolescent in N. lio- lepis). Snout and suborbital areas with some na- ked areas along ventral margins in most species, almost entirely naked ventrally in these areas in several species. Total gill-raker number on me- sial side of outer (first) arch less than 12 in all but a few species. Color various shades of black, brown, blue, or violet; some species with last two colors have silvery reflections on body. Py- loric caeca unbranched and fewer than 30 in dis- tal count in most species, as many as 60 in the few species with branched caeca. Retia two (oc- casional individuals may have four), slender, uncoiled; gas glands globular, sometimes some- what flattened. REMARKS.—The genus Nezumia is a diverse group of slightly more than 40 species of which ten are known and confined to the eastern Pa- cific. Most of these ten species have restricted distributions, but N. stelgidolepis is known from southern Peru to southern British Columbia, and N. convergens is common in waters from north- ern Peru to Costa Rica and is here recorded from the Islas Tres Maria and the Gulf of California. The distribution of six species (/atirostrata, lio- lepis, orbitalis, parini, pudens, pulchella) each spans fewer than 30 degrees of latitude. N. lor- icata is known only from isolated captures in the Galapagos and off central Chile. Although most other genera are poorly rep- resented in the eastern Pacific, the region has proved rich in Nezumia species. The ten species here recorded rank the fauna among the largest found in broadly comparable geographical areas. Based on my examination of specimens and from the literature, a breakdown of the number of Nezumia species by area is as follows: Eastern Pacific—10 spp. Atlantic—14 spp. Eastern—10 spp. Western—9 spp. Japan (8) + Philippines (4)—10 spp. Hawali—7 spp. Western Indian Ocean— ?4 spp. Central Indian Ocean— 4 spp. The few species recorded from the Philippines is surprising, as the area is otherwise exceed- ingly rich in grenadier species, especially of the genus Coelorinchus. Despite the extensive col- IWAMOTO: EASTERN PACIFIC MACROURIDAE lecting by the ALBATROSS in the early part of the century, the area remains poorly known in terms of its deepwater fauna, and doubtless, other species of Nezumia will subsequently be found there. The South China Sea and the Indo-Aus- tralian Archipelago likewise remain relative un- knowns, and more deepwater collecting is badly needed in these biologically rich areas. Indian Ocean grenadiers have not been comprehen- sively reviewed by recent workers; the generic allocations of many species are still question- able. Members of the genus are of small to mod- erate size; only a few attain lengths of more than 340 mm in total length (only N. stelgidolepis in the eastern Pacific, but other species in the At- lantic and western Pacific). The larger species tend to have a larger, more terminal mouth and stronger, longer teeth—these features being suggestive of a predatory habit on larger, more active prey. The smaller species with their smaller, more inferior mouth, and longer, more pointed snout most likely feed on small prey captured on or in the bottom substratum. Ne- zumia parini is an enigma, however, in that it is strictly bathypelagic (the only member of the genus to be so), yet it retains a physiognomy much like that of its bottom-dwelling congeners (see Hubbs and Iwamoto (1977) for additional discussion of this peculiar species). The genus is primarily one of upper- and mid- dle-slope fishes, with few species ranging deeper than 2,000 m. Of the eastern Pacific species (Table 2), N. convergens not only has the sec- ond broadest horizontal range, but also the greatest vertical range, and it is probably the deepest-living species. In contrast, N. stelgi- dolepis, which has the broadest horizontal range of the eastern Pacific species, has only a mod- erate depth range and has been captured at the shallowest depth. The sole bathypelagic member of the genus, N. parini, has been captured in midwater nets fished primarily in depths of 1,000 m or greater. In the type-series, 14 of 23 captures of the fish were made in nets fished to an estimated depth of 1,000 m or greater; in seven, the nets were fished to depths estimated at between 675 and 940 m; and only in two were the nets fished shal- lower (estimated depths of 420 and 455 m) (Hubbs and Iwamoto 1977). Genera most closely related to Nezumia are Ventrifossa Gilbert and Hubbs, 1920, and Mal- 157 TABLE 2. COMPARISON OF CAPTURE DEPTHS OF NINE SPECIES OF NEZUMIA FROM THE EASTERN PACIFIC OCEAN. Capture depths (m) Species min max. difference stelgidolepis 227 909 632 pulchella 272 735 463 orbitalis 523 800 277 ventralis 549 717 168 pudens 580 1,238 758 loricata 600 1,480 880 convergens 600 1,865 1,265 latirostrata 605 1,400 795 liolepis 682 1,629 947 acocephalus Gunther, 1887; the three constitute a close-knit unit which I (Iwamoto 1972) have called the tribe Malacocephalini. The unifying characters of this tribe are the presence in the members of seven branchiostegal rays, a well- developed periproct region that is remote from the anal fin origin, and the presence of one or two dermal windows of the light organ anterior to the anus. Nezumia liolepis (Gilbert) (Figures 126, 18c) Macrurus (Lionurus) liolepis GILBERT, 1890:117 (original de- scription, off southern California, ALBATROSS sta. 2980, in 1,103 m). Lionurus liolepis: GOODE AND BEAN 1896:409 (occurrence). Macrurus liolepis: GARMAN 1899:199-200 (description; rec- ords from ALBATROSS sta. 3418, 3424, 3436). Macrurus barbiger GARMAN, 1899:197, pl. 45, figs. 2-2b (orig- inal description; off Islas Tres Marias, Mexico, ALBATROSS sta. 3424, in 676 fm [1,236 m)). Lionurus (Lionurus) barbiger: GILBERT AND Huss 1916:146 (listed). Lionurus (Lionurus) liolepis: GILBERT AND Husss 1916:146 (listed). Nezumia liolepis: FitCH AND LAVENBERG 1968: 142 (listed). Ventrifossa barbiger: MARSHALL 1973:654 (listed). DiaGNosis.—A Nezumia with 10-11 (rarely 12) pelvic fin rays. Few serrations on second spinous ray of first dorsal fin. Thin, deciduous scales on body with few or no spinules on ex- posed field. Small ventral light organ. Upper jaw 31-40 percent HL. DESCRIPTION.—General features of fish seen in Figure 12b. Head moderately compressed and deep. Suborbital ridge rounded; shallow, obtuse angle formed by dorsal and ventral surfaces. In- terorbital region shallowly concave; width about equal to or less than orbit diameter. Snout nar- row, bluntly pointed, protruding slightly beyond 158 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 5 rather large, almost lateral mouth. Jaws only slightly restricted by lip-folds at posterior angle. Opercular openings wide, extending above a horizontal through dorsal edge of orbit; gill membranes loosely attached to isthmus. Nape high, strongly arched in large adults. First dorsal fin base low. Periproct region about equidistant from pelvic fin insertion and anal fin origin; ven- tral light organ poorly developed, inconspicuous without dissection. Gill-rakers small, tubercular. Gill filaments long; length of longest more than half diameter of orbit, about equal to or longer than least width of suborbital region. Pyloric caeca slender; rather short, less than diameter of orbits; biramously branched, each main stem branching one or two times. Fin rays all relatively slender and delicate. Second spinous ray of dorsal fin slightly pro- longed; leading edge smooth except for a few small denticles near distal end. Outer ray of pel- vic fin slender; filamentous tip extends slightly beyond origin of anal fin. Overall coloration grayish brown tinged with violet; surface over abdomen much darker, over operculum blackish, but anteriorly over rest of head and especially snout paler. Lips dark brown to blackish; fins dusky to blackish. Gill membranes black. Outer-wall lining of gill cham- ber blackish; inner-wall lining grayish to pale. Lining of oral cavity blackish. Scales highly deciduous; few specimens with any scales remaining. A sizable patch remaining dorsally on head and snout of CAS-SU 2545, 50 mm HL. Larger of these scales with 5—7 rows, each with 2-3 long conical spinules (Fig. 12b). Scales over dorsal surface of snout either with one to several low ridges that are sometimes armed with one or a few short spinules, or scales completely lack ridges and spinules. Dorsal sur- face of snout lacks scales along leading edge and along snout ridges. Ventrally, snout entirely na- ked back onto suborbital region to vertical through end of maxillary, but a narrow, thin wedge of small, thin, cycloid scales extends for- ward to about level of anterior border of orbits. Rami of lower jaw with small, fine, spinuleless scales located mainly about median axis of each ramus. Counts (from 38 specimens).—ID. II,8-11 (total 10-13; x = 11.87; S.D. = 0.75); 1P. 20-25 @= 2212? n= 61; S.D, = 1.23); 2P. 10-11 (rarely 12). Gill-rakers on first arch 1-3 + 7-10 (total 9-12, usually 10-11); on second arch 1-3 + 7-10 (9-12 total). Scales below first dor- sal 8-10; below second dorsal 7-10; over dis- tance equal to predorsal length of head 39-44 (3 specimens). Pyloric caeca 25-37 (6 speci- mens). MEASUREMENTS (from 38 specimens).—Total length 114-290 mm; head length 26-63 mm. The following in percent of head length [range (x; n: S.D.)]: postrostral length of head 73-81 (77.68: 34; 1.77); snout length 24—28 (25.95; 36; 1.12); preoral length 13-23 (19.12; 34; 2.00); internasal width 17-23 (19.00; 26; 1.50); orbit diameter 25— 31 (27.18; 38; 1.50); interorbital width 21- 27 (23.28; 37; 1.25); postorbital length of head 43-56 (50.17; 35; 3.42); distance orbit to angle of preopercle 36—44 (39.39; 37; 1.88); suborbital width 11-15 (12.92; 37; 0.96); upper jaw length (28) 31-40 (34.12; 38; 2.31); barbel length 10-20 (14.68; 37; 2.59); length first gill-slit 14-23 (17.38; 37; 1.65); preanal length 119-142 (131.52; 27; 5.81); outer pelvic ray to anal origin 32—57 (42.26; 26; 6.24); anus to anal origin 13-27 (19.76; 24; 4.66); greatest body depth 60-82 (70.89; 29; 4.86); 1D.-2D. interspace 28-48 (34.63; 36; 6.15); 1D. height 58-73 (62.267; 21; 4.27); IP. length 43-53 (49.15; 24; 3.30); 2P. length 42-55 (48.03; 25; 6.13). COMPARISONS AND RELATIONSHIPS.—Ne- zumia liolepis belongs with that group of Ne- zumia species characterized by (1) a moderate- sized mouth, (2) a relatively deep, compressed head, (3) arather blunt, high, and usually weakly armed snout, and (4) relatively wide gill open- ings. This group includes such geographically separated species as N. stelgidolepis (Gilbert). N. atlantica (Parr, 1946), N. africana (lwamo- to, 1970), N. bubonis Iwamoto, 1974, N. dara (Gilbert and Hubbs, 1916), N. burragei (Gilbert, 1905), N. hebetata (Gilbert, 1905), N. holocen- trus (Gilbert and Cramer, 1897), N. kamoharai Okamura, 1970, and N. macronema (Smith and Radcliffe, 1912). N. liolepis is readily distin- guished from these species in having the com- bination of (1) few serrations on the dorsal spine, (2) reduced spinulation on the scales of the head and body, (3) extensive naked areas on the dor- sal and ventral surfaces of the snout and sub- orbital region, (4) a distinctive shape of the oper- culum, and (5) a relatively posteriad position of the anus. REMARKS.—Garman (1899:197) described Macrurus barbiger from a 10-inch (25-cm) spec- imen taken off the state of Guerrero, Mexico. IWAMOTO: EASTERN PACIFIC MACROURIDAE He contrasted the species with N. liolepis, re- porting that barbiger has ‘‘the head more round, the cheeks more convex, the snout narrower, and the barbel longer, besides which differences there are others in the fins and the colors.”’ I compared the holotype of M. barbiger (MCZ) 28597) with specimens Garman identified as “Macrurus liolepis’’ and found that these dif- ferences do not hold up. There is little doubt that the M. barbiger holotype and the **M. lio- lepis’’ specimens are conspecific. DISTRIBUTION.—The species is known from off Monterey Bay, California (36°49’20°N), to south of Guerrero, Mexico (17°24'N), including the Gulf of California south of Guaymas (Gar- man 1899:200, ALBATROSS sta. 3436). Capture depths have ranged from 768-823 m (CAS 26638) to 1.655 m (ALBATROSS Sta. 3436). MATERIAL EXAMINED (40 specimens from 9 localities).— California (north to south): CAS-SU 5351 (4, 28-52 mm HL), off Monterey Bay, 36°49'20’N, 122°12'30’W, 834 m, ALBaA- TROSS sta. 3126, 13 Mar. 1889. —CAS-SU 21402 (5, 53-63), off Santa Cruz Islands, 1,397—1,629 m, ALBATROSS sta. 4428, 14 Apr. 1904. —USNM 44271, syntypes (2, 45-50), £ of Santa Cruz Islands, 33°49'45"N, 119°24'30’W, 1,103 m, ALBATROSS sta. 2980, 12 Feb. 1889. —CAS 26638 (4, 41-56), off San Ma- teo Point, 33°15’30’N, 117°38'W, 768-823 m, N. B. SCOFIELD sta. 53B59, 23 June 1953. —CAS-SU 2545 (6, 27-50), off San Diego, 32°49'N, 117°27'30"W, 656 m, ALBATROSS Sta. 2936, 4 Feb. 1889. —USNM 77495 (3, 33-33), off Point Loma, San Diego, 940-989 m, ALBATROSS sta. 4333, 9 Mar. 1904. Mexico: MCZ 28597 (holotype of Macrurus barbiger Gar- man, 1899, 54 mm HL), off Islas Tres Marias, 21°15'N, 106°23’W, 1,236 m, ALBATROSS sta. 3424, 18 Apr. 1891. — CAS 40230 (10, 38-47), off Jalisco, 19°43.5'N, 105°35.5'W, 700-900 m, TE VEGA cr. 19, sta. 17, 10 July 1968. —CAS 40231 (5S, 51-58), off Guerrero, 17°24'N, 101°31’W, 940-1,000 m, TE VEGA cr. 19, sta. 19, 12 July, 1968. Nezumia pulchella (Pequeno, 1971) (Figures 12c, 185) Macruroplus pulchellus PEQUENO, 1971:293-294, fig. 17 (orig- inal description; off Chile; type-locality 25°26’S, 70°37'W, 374-424 m). DIAGNosIs.—A species of Nezumia with 11- 12 pelvic fin rays. Gill-rakers on first arch 9-11 total; on second arch 8-10. Spinules on body scales conical, in distinct, slightly convergent rows. Ventral surfaces of snout, suborbital re- gion, mandibular rami, and gular and branchio- stegal membranes usually naked, but occasion- ally with few scattered scales. Barbel 9-16 percent of HL; upper jaw 30-34 percent HL. DESCRIPTION.—General features of fish seen in Figure 12c. Head moderately compressed, trunk deep in large adults, less so in smaller in- 159 dividuals. Suborbital ridge prominent in study material, accentuated by shrunken lower portion of region; suborbital ridge of fresh specimens probably not as pronounced. Other head ridges rather smoothly rounded without reinforcing spiny scutelike scales. Snout narrow and short, relatively blunt; armed with small spiny scutes at tip and at lateral angles. Mouth moderately large, lateral, unrestricted by lip folds at angle. Posterodorsal corner of opercle slightly pro- duced. Interopercle broadly exposed posteriorly and ventrally. Gill openings wide, membranes broadly connected over isthmus with a moder- ately broad free fold. Anterior dermal window of light organ small, in shallow fossa between bases of pelvic fins and slightly removed from periproct. Gill filaments long; length of longest about equal to diameter of pupil, greater than width of suborbital region. Pyloric caeca well developed, but fairly short (about 0.6 into orbit diameter), branched only at bases; 23-31 in nine specimens. Scales of body with distinct, slightly conver- gent rows of small conical spinules on exposed fields. Scales generally lacking on gular mem- brane, mandibles, lower part of snout and sub- orbital region, and branchiostegal membrane, but one specimen (CAS 38323) with two small scales on right gill membrane near base of fifth branchiostegal ray. Small areas dorsally behind leading edge of snout naked. Stout, spiny scales at tip ard lateral angles of snout and in two rows along dorsal surface of suborbital region, but scales otherwise not modified. Sensory pores along ventral margins of suborbital region and preopercle, and along inner margins of mandi- bles large and prominent. Teeth in both jaws in rather narrow band; teeth small except for outer premaxillary series, which is composed of enlarged, spaced, sharp, conical teeth. Coloration in alcohol medium brown overall with bluish tinge over abdomen and blackish ventrally on trunk. Gill membranes black; fins blackish. Lining of buccal cavity pale except for grayish oral valves. Peritoneal cavity pale but peppered with small melanophores. Counts (from 16 specimens).—ID. II,10-11 (9 in one specimen); IP. 19-28 (* = 24.52: n = 31; S.D. = 2.06); 2P. 11-12. Mesial gill-rakers on first arch 1-3 + 7-9 (9-11 total); on second arch 1-2 + 7-9 (8-10 total). Scales below first dorsal 642-82; below second dorsal 52-742; 160 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 5 below midbase of first dorsal 42-6; over dis- tance equal to predorsal length 35-44 (usually 37-40). MEASUREMENTS (from 15 specimens, 22-mm- HL specimen from SIO 65-675 excluded).—To- tal length 137+ to 286+ mm; head length 29-61 mm. The following in percent of head length [range (x; n: S.D.)]: postrostral length of head 79-82 (80.47; 15; 1.06); snout length 21-26 (23.59; 15; 1.38); preoral length 16-19 (17.61; 15; 0.62): internasal width 15-21 (18.47; 15; 1.42): orbit diameter 30-34 (32.29; 15; 1.47); interor- bital width 20-25 (22.79; 15; 1.31); postorbital length 40-46 (43.34; 15; 1.88); orbit to angle of preopercle 34-38 (36.05; 15; 1.30); suborbital width 11-13 (11.85; 15; 0.66); upper jaw length 30-34 (32.15; 15; 1.29); length barbel 9-16 @ils73= 15; 3:38); length first gill-slit 17—23 (19.95; 15; 1.54); preanal length 117-150 (131.93; 15; 7.51); greatest body depth 68-88 (79.80; 15; 5.43); 1D.-2D. interspace 31-56 (41.83; 15; 7.40); height first dorsal fin 65-76 (69.30; 10; 4.00); length pectoral fin 51-70 (61.53; 15; 5.17); length pelvic fin 39-53 (45.87; 15; 3.48). COMPARISONS AND RELATIONSHIPS.—Ne- zumia pulchella is closely related to N. stelgi- dolepis but differs primarily in having 11-12 pelvic fin rays rather than the 9-10 of N. stelgidolepis. Scale spinules also show notable differences between the two species. In N. pul- chella the spinules are all conical and in distinct rows that converge slightly towards the midline, whereas in stelgidolepis the spinules are conical to lanceolate and arranged in a more quincunx pattern. Retia and gas glands of pulchella are also much larger and stouter, the barbel is short- er, and the branchiostegal rays are naked except for occasional scattered scales (as opposed to short rows of deciduous scales along the bases of the lowermost branchiostegal rays in stelgi- dolepis). Nezumia pulchella is readily distinguishable from N. pudens in having extensive naked areas on the ventral surface of the head and on the man- dibles (as compared with surfaces mostly scaled in pudens), fewer scale rows below the first and second dorsal fins (62 to 8% vs. 10% to 12 be- low the first dorsal; 5%2 to 7% vs. 8% to 10% below the second dorsal, fewer pyloric caeca (23-31 vs. 37-44), and a shorter barbel (9-16 vs. 20-25). S$1zE.—To at least 286 mm in total length (in a specimen, IMARPE uncat., 61 mm in head length). A specimen in the type-series measured 67 mm HL and 272 mm TL (Pequeno 1971:294, table). DISTRIBUTION.—Northern Peru (7°49’S) to central Chile (about 33°S) in 250—960 m. MATERIAL EXAMINED (16 specimens from 8 localities).— Peru: CAS 38323 (3, 54-57 mm HL, 207+ to 270 mm TL), 7°49'S, 80°38’ W, 605-735 m, ANTON BRUUN cr. 18B, sta. 754 (field no. LWK66—93), 5 Sep. 1966. —CAS 28761 (1, 43 HL, 187 TL) and IMARPE uncat. (1, 61 HL, 286+ TL), 13°49.4’S, 76°46.9'W, 570 m, 24 Jan. 1972. —CAS 28763 (1, 46 HL, 226 TL), 13°53.3’S, 76°42.0’W, 272 m, 25 Jan. 1972. —CAS 28764 (1, 35 HL, 164 TL), 16°31.0’S, 73°12.2'W, 510 m, 1 Feb. 1972. —IMARPE uncat. (1, 34 HL, 161 TL), 18°07’S, 71°02.5’W, 28 Jan. 1972. Chile: CAS 38320 (6, 26-39 HL, 135-185 TL), 23°41'S, 70°34’W, 250-400 m, ANTON BRUUN cr. 18A, sta. 717 (field no. LWK66—63), 17 Aug. 1966. —CAS 38318 (1, 38 HL, 158 TL), 32°08.5'S, 71°43’W, 960 m, ANTON BRUUN cr. 18A, sta. 703 (field no. LWK66-47), 12 Aug. 1966. —SIO 65-— 675 (1, 22 HL, 104 TL), about 29 km off Valparaiso Harbor (ca. 33°N), 22-23 Dec. 1965. Nezumia stelgidolepis (Gilbert) (Figures 12d, 13, 187) Macrurus stelgidolepis GILBERT, 1890:116 (orginal descrip- tion; off Pt. Conception, California, ALBATROSS sta. 2960, in 488 m). Macrurus gracillicauda GARMAN, 1899:206—207, pl. H, fig. 1 (original description; Gulf of Panama, ALBATROSS sta. 3384 and 3385, in 837 and 523 m). Lionurus (Nezumia) stelgidolepis: GILBERT AND HUBBS 1916:145 (listed). Lionurus stelgidolepis: BARNHART 1936:24 (brief description). Nezumia stelgidolepis: ROEDEL 1951:509, fig. 183 (16 records off California). Nezumia gracillicauda: MARSHALL AND IWAMOTO in MAR- SHALL 1973:626 (listed). DIAGNOSIS.—A species of Nezumia with 9— 10 pelvic fin rays. Gill-rakers on first arch 8-12 total; on second arch 8-11 total. Spinules on body scales conical to narrowly lanceolate. Ven- tral surfaces of snout, most of suborbital region, and anterior half or more of mandibular rami naked. Lowermost branchiostegal rays scaled. Barbel 15-26 percent HL; upper jaw 31-37 per- cent. DESCRIPTION.—General features of fish seen in Figures 12d and 13. Head moderately com- pressed and deep; ridges of head not strongly developed, usually somewhat rounded; subor- bital region relatively flat. Snout narrow, bluntly pointed (more pointed in young), protruding slightly beyond rather large mouth, which is only slightly restricted at posterior angle by lip folds. Posterodorsal corner of opercle somewhat IWAMOTO: EASTERN PACIFIC MACROURIDAE 161 Gr ; MN H S pA ag FiGureE 13. Nezumia stelgidolepis (Gilbert). (a) CAS 44197, 14.7 mm HL, 73 mm TL (fins and head region somewhat structed. (c) CAS 40022, 53 mm HL, 255 mm TL. 162 flaplike, posterior margin of gill cover steeply oblique along opercle, gently rounded along sub- opercle, connecting to broadly exposed (and scaled) interopercle. Shape of gill cover similar to that in N. liolepis. Gill openings wide; gill membranes rather narrowly connected over isthmus, with free posterior fold. Nape region slightly arched. Anus within lenticular area of black naked skin, situated slightly behind bases of pelvic fins, but closer to these than to anal fin. Anterior dermal window of light organ small, in shallow fossa between pelvic fin bases. Gill filaments well developed; length longest filament equal to or longer than least suborbital width and about equal to pupil diameter. Pyloric caeca numerous, branched; counts highly variable, in eight specimens 24, 34, 36, 38, 38, 40, 43, and 58. Periproct region relatively large, close be- hind pelvic fin bases; anterior window of light organ small, circular, lying between inner edges of pelvic fin bases. Scales densely covered with long, narrowly lanceolate to conical spinules arranged in short, slightly convergent rows. Ventral surfaces of snout and most of suborbital region, and anterior half to two-thirds of lower jaw naked. Sensory pores rather prominent over these naked areas. Short rows of small deciduous scales along bas- es of one or more branchiostegal ray. Coloration of fresh specimen (CAS 40022) swarthy overall with blackish predominant over ventral aspects of abdomen, chest, head, and gill covers. Bluish tinge over blackish abdominal walls. Fins dusky to blackish; first dorsal slightly paler basally. Lining of buccal cavity pale, but oral valves gray. Pharyngeal region dark gray; gill-rakers gray but gill filaments pale. In juve- niles (CAS 41196 and 41197), oral cavity densely peppered with small melanophores. Dentition in broad bands in both jaws, nar- rowly tapered posterolaterally. Outer series of upper jaw teeth slightly enlarged and evenly spaced. Four juvenile specimens presumably of this species were captured in midwaters. The largest of these (LACM 30611-16) measured 19 mm in HL, 106 mm in TL, and was taken in an Isaacs- Kidd midwater trawl off the northwestern coast of Baja California between the surface and about 600 m, over a bottom depth of 1,600 to 1,100 fms (2,962—2,012 m). This specimen was dark colored, superficially resembling N. parini, but the coarse scale spinules, the large pores on the PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 5 head and mandibles, the distinct lateral line, the number of pelvic fin rays, and the large gill fil- aments distinguish the species as N. stelgido- lepis. The three other specimens (CAS 41196 and 41197) captured in midwaters were too small to positively identify without more comparative material, but the general appearance (see Fig. 13c) and counts (of dorsal and pelvic fin rays, and gill-rakers) also suggest N. stelgidolepis as the correct identification. Proportional measure- ments of these juveniles were generally diver- gent from those of the adults, indicating sub- stantial allometric growth in such features as preoral length of snout, orbit diameter, inter- obital width, postorbital length of head, distance orbit to angle of preopercle, upper jaw length, and barbel length. The smallest examined specimen (USNM 57861) captured in a bottom trawl measured 39 mm in head length and 176 mm in total length. Counts (from 33 specimens).—1D. II,9-10 (8 in one specimen, 11 in two specimens); IP. 20-26 (& = 21.00; n = 59; S.D. = 3:30): 2P2 9— 10 (8 in one specimen, 11 in two others). Gill- rakers on first arch 10-11 total (8 in one, 9 in one, 12 in three specimens); on second arch 9— 11 total. Scales below first dorsal 8—9 (rarely 10); below second dorsal 7-8 (9 in one); below mid- base of first dorsal 6-7; over distance equal to predorsal length of head 34-47, usually between 36 and 42. MEASUREMENTS (from 38 specimens, juve- niles excluded).—Total length 176-405+ mm; head length 39-96 mm. The following in percent of head length [range (x; n; S.D.)]: postrostral length of head 73-82 (78.60; 35; 1.80); snout length 22—28 (24.51; 36; 1.34); preoral length 12- 18 (14.86; 36; 1.28); internasal width 14-19 (16.46; 30; 1.19); orbit diameter 26—32 (28.06; 38; 1.68); interorbital width 20-26 (23.46; 38; 1.55); postorbital length 43-52 (46.56; 33; 2.17); orbit to angle of preopercle 35—45 (39.81; 37; 2.01); suborbital width 11-16 (12.84; 37; 1.01); upper jaw length 31-37 (34.93; 38; 1.49); length barbel 15-26 (20.98; 36; 2.72); length first gill-slit 12-18 (16.37; 36; 1.54); preanal length 119-157 (135.95; 37; 9.51); greatest body depth 70-90 (77.68; 33; 5.77); 1D.-2D. interspace 26-58 (45.92; 38; 11.19); height first dorsal fin 48-68 (60.73; 29; 4.26); length pectoral fin 44-56 (49.69; 35; 3.23); length pelvic fin 35—49 (43.48; 34; 3.61). COMPARISONS AND RELATIONSHIPS.—In the eastern Pacific, Nezumia stelgidolepis is most IWAMOTO: EASTERN PACIFIC MACROURIDAE closely related to N. liolepis and N. pulchella, but is readily distinguished from these two by differences in scale spinulation, squamation pat- tern, pelvic ray count, and other features noted in the key. Nezumia stelgidolepis is represen- tative of that group of Nezumia spp. character- ized by relatively blunt snout, large mouth, broadly unrestricted gill openings, deep body, and large size. Some of the species in this group (e.g., Nezumia atlantica (Parr, 1946), N. afri- cana (Iwamoto, 1970)) have been treated in oth- er works (Parr 1946; Iwamoto 1970; Marshall 1973) as members of the genus Ventrifossa. S1zE.—Nezumia stelgidolepis is the largest eastern Pacific member of the genus, attaining a total length of at least 445 mm (see Roedel 1951:509). Nezumia atlantica, a close relative from the western Atlantic, attains a comparable size, probably exceeded in the genus only by the peculiar species N. bubonis Iwamoto, 1974, from the Hawaiian Islands and the western At- lantic. DIsTRIBUTION.—British Columbia (off Van- couver Island) to southern Peru (18°10.0'S), in 277-909 m (Makushok (1967:table 18) gives the depth distribution as **(61—91)379-909""). MATERIAL EXAMINED (excluding material previously listed in Iwamoto and Stein 1974:49-S0).—California: CAS 23396 (1, 63 mm HL, 310 mm TL), off Ft. Bragg in 475-494 m. — CAS 33110 (1, 43 HL, 243 TL), off Bodega Bay, 494-585 m, 23 Feb. 1965. —CAS 31509 (1, 73 HL, 365+ TL), off Point San Simeon, 530 m, N. B. SCOFIELD sta. 54BB71, 25 Oct. 1954. —CAS 40022 (1, 53 HL, 255 TL), off Santa Barbara County, 34°18.4’N, 120°14.4’'W, 439-443 m, COMMANDO, 10 Aug. 1977. —USNM 87579 (1, 68 HL), ALBATROoss. —USNM 127072 (1, 39 HL, 280+ TL), off southern California, ALBA- TROSS. Mexico: LACM 30611-16 (1, 19 HL, 106 TL), Guadalupe I., 28°57'15"N, 118°05'19°W, est. fishing depth 0-600 m, bot- tom depth 2,926—2,012 m, IKMT, VELERO sta. 12494, 21 Nov. 1968. —AMNH 12902 (4, 55-70 HL, 250-305+ TL), San Cris- tobal Bay, 27°07'08"gN, 114°33'10”W, 519 m, ALBATROSS sta. 5675, 15 Mar. 1911. Galapagos: USNM 135605 (1, 77 HL, 346 TL), ALBATROSS. Ecuador: CAS 44196 (1, 14 HL, 67+ TL), 4°14’S, 81°26’W, 0-1,830 m, IKMT, ANTON BRUUN cr. 18B, sta. 756B (field no. LWK66-113), 8 Sep. 1966. Peru: LACM 33883 (1, 66 HL, 303 TL), 6°42’S, 80°59’05’W, 780 m, sta. SNP1-25, 22 Jan. 1974. —LACM 33886 (1, 41 HL, 168 TL), 7°44'05"S, 80°30'05"W, 750-760 m, sta. SNP1-28, 23 Jan. 1974. —CAS 38324 (7, 48-66 HL, 180+ to 280 TL), 7°49'S, 80°38’W, 605-735 m, ANTON BRUUN cr. 18B, sta. 754 (field no. LWK66-93), 5 Sep. 1966. —CAS 41197 (2, 13.5- 14.7 HL, 66-73 TL), 11°53’S, 78°05'W, 0-1,125 m, IKMT, ANTON BRUUN cr. 16, sta. 656-0, 14 June 1966. —IMARPE uncat. (1, 74 HL, 290+ TL), 18°10’S, 71°29’W, 610 m, 23 Aug. 1972. 163 Nezumia pudens Gilbert and Thompson (Figures 6b, 14c, 18h) Nezumia pudens GILBERT AND THOMPSON, 1916:472-473, pl. 5, fig. 2 (types from ALBATROsS sta. 2791 off Lota, Chile, 38°08'S, 75°53'W, 1,240 m). Lionurus (Nezumia) pudens: GILBERT AND Huss 1916:146 (name only). Macruroplus pudens: FOWLER 1944:48, fig. (list). DIAGNOSIS.—A species of Nezumia with 10- 11 pelvic fin rays. Gill-rakers on first and second arches 9-10 total. Spinules on body scales slen- der, sharp, conical, in dense cluster on exposed fields. Head surface fully scaled except for small area along ventromedian line of snout and over gill membranes. Barbel 20-25 percent of HL; upper jaw 30-39 percent HL. DESCRIPTION.—General features of fish seen in Figure 14a. Head rather compressed, surfaces smoothly rounded. Suborbital ridge low, round- ed. Snout short, blunt narrow; blunt scutes at terminal and lateral angles not especially spiny or set off from adjacent scales. Mouth subter- minal, rather short; posterior corners somewhat restricted by lip folds; lips thick, papillaceous, especially in large adults. Barbel rather thick throughout, not tapering to filamentous tip. In- teropercle broadly exposed posteriorly and ven- trally. Posterodorsal corner of opercle angular, without broad tablike extension. Gill mem- branes rather broadly connected over isthmus, with a narrow free fold. Anus located in middle third of distance between pelvic fin base and anal fin origin. Anterior dermal window small, round, lying between or slightly in advance of line connecting insertion of pelvic fins. Gill fil- aments well developed but relatively short; length of longest less than diameter of eye lens, about 0.6 into least suborbital width. Pyloric caeca slender, numerous, 37, 39, and 44 in three specimens, branched two or three times at base; length of longest about equal to interorbital width. Exposed fields on body scales densely cov- ered with relatively erect, slender, stiletto- shaped spinules arranged in quincunx pattern or in close, strongly convergent rows. Almost all of head uniformly covered with small, relatively adherent scales (Fig. 6b). Small area along ven- tral midline and ventral margin of snout naked. Scales along dorsal half of suborbital region heavier, larger, more adherent, and in two dis- tinct rows. Mandibles and exposed surfaces of 164 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 5 < ey x a PES RERERY BEE REPL) a FiGurE 14. (a) Nezumia pudens (Gilbert and Thompson). CAS 38310, 50 mm HL, from off Chile in 750 m. (b) Nezumia ventralis Hubbs and Iwamoto, new species. Holotype, CAS-SU 68361, 26.4 mm HL, taken off the Galapagos in 549 m. (c) Nezumia orbitalis (Garman), LACM 33885-2, 48 mm HL, off Peru in 800 m. IWAMOTO: EASTERN PACIFIC MACROURIDAE interopercle covered with small scales, but gill membranes naked. Fins moderate in size. First spinous ray of dorsal fin slender, sharp, spikelike; second spi- nous ray slightly prolonged, armed along leading edge with slender, sharp spikes spaced suffi- ciently apart so that spikes do not overlap. Outer ray of pelvic fins with filamentous tip extending well beyond origin of anal fin. First dorsal fin placed rather far posteriad; origin about on same vertical as anus and about one eye-lens diameter behind vertical through origin of pectoral fins. Jaw teeth in rather broad, coarse bands that narrow posteriorly. Outer series of upper jaw slightly enlarged, particularly along anterior bor- der. Coloration in alcohol medium brown with ab- dominal and cheek area blackish tinged with vi- olet. Gill membranes black; naked margins of snout, nostrils, lower jaw, opercular bones, and rim of orbit blackish or darkly swarthy. Fins all blackish; distal and anterior portions of mem- branes of first dorsal fin more intensely blackish. Lining of mouth and gullet blackish; that of gill cavity blackish along external margins but pale towards inner portions; that of abdominal cavity black to swarthy. Counts (from 7 specimens).—1D. II,9-11; IP. 19-24; 2P. 10-11. Medial gill-rakers on first arch 1-2 + 7-9, usually 2 + 8 (9-10 total); on second arch 1-2 + 8-9 (9-10 total). Scale rows (from 4 specimens) below first dorsal 10% to 12; below second dorsal 8% to 10/2; below midbase of first dorsal 72 to 8’; over distance equal to predorsal length 43-46. MEASUREMENTS (from 6 specimens; 22-mm- HL specimen excluded).—Total length 165—400 mm; head length 32-68 mm. The following in percent of head length: postrostral length of head 74-76; snout length 28-30; preoral length 18-22; internasal width 19-23; orbit diameter 31-35; interorbital width 22-23; postorbital length of head 40-48; distance orbit to angle of preopercle 37-43; suborbital width 1S—16; upper jaw length 30-39; length barbel 20-25; length first gill-slit 14-18; preanal length 143-158; greatest body depth 73-81; 1D.—2D. interspace 34-43; height first dorsal fin 80-93; length pec- toral fin 56-64; length pelvic fin 44-58. COMPARISONS AND RELATIONSHIPS.—Ne- zumia pudens is readily distinguished from its most closely related eastern Pacific congeners N. stelgidolepis, N. pulchella, and N. liolepis by: (1) ventral surface of snout, suborbital re- 165 gion, and mandibular rami scaled (vs. mostly naked in these other species); (2) mucous pores in these areas not prominent (vs. prominent); (3) posterodorsal angle of opercle not produced into a short tab or flap (vs. a distinct flap or tab); (4) gill filaments relatively short, length less than diameter of eye lens (greater than eye lens in others); (5) first dorsal fin originating more pos- teriad than others; and (6) buccal cavity com- pletely blackish (mostly pale in stelgidolepis and pulchella, but blackish in liolepis). Its relation- ship to other Pacific species of Nezumia is un- certain and probably distant. Nezumia pudens bears some resemblance in general physiognomy of the head and body to N. sclerorhynchus (Valenciennes, 1838), and N. suilla Marshall and Iwamoto, 1973, from the Atlantic, but the higher pelvic fin ray count and the more complete squamation on the ventral aspects of the head in N. pudens are distinguish- ing. S1zeE.—To at least 68 mm HL and 400 mm TL. DISTRIBUTION.—Known only off central Chile between latitudes 32°S and 38°S, in 580- 1,238 m. MATERIAL EXAMINED (7 specimens, all from Chile).— USNM uncat. (formerly CAS 38317) (1, 39 mm HL, 240 mm TL), 32°08.5'S, 71°43'W, 960 m, ANTON BRUUN cr. I8A, Sta. 703 (field no. LWK66—-47), 12 Aug. 1966. —CAS 38315 (2, 22- 39 HL, 106+ to 234 TL), 32°17’S, 71°39.5'W, 580 m, ANTON Bruun cr. 18A, sta. 702 (field no. LWK66—44), 11 Aug. 1966. —CAS 38310 (2, 50-68 HL, 267-400 TL), 34°06.5’S, 72°18.5'W, 750 m, ANTON BruUN cr. 18A, sta. 687 (field no. LWK66-25), 5 Aug. 1966. —USNM 76860 (holotype, 32 HL, 170 TL) and CAS-SU 22727 (paraiype, 34 HL, 165 TL), 38°08'S, 75°53’W, 1,238 m, ALBATROSS sta. 2791, 14 Feb. 1888. Nezumia ventralis Hubbs and Iwamoto, new species (Figure 145) DiaGNosis.—A Nezumia with 15 pelvic fin rays and 24-27 pectoral fin rays. Mandibular rami, ventral surfaces of snout, and most of ven- tral surfaces of suborbital region naked; mucous pores over these surfaces large. Scales small, about 9% below origin of second dorsal fin; spi- nules on body scales slender, conical, greatly reclined, in parallel to subparallel rows. DESCRIPTION OF HoLotype.—General fea- tures of fish seen in Figure 14b. Head and trunk moderately compressed and deep. Suborbital region divided into an upper portion covered with two longitudinal rows of stout scales and 166 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 5 a lower portion with almost no scales; the sur- faces of the two halves meet along the suborbital ridge at an obtuse angle. Opercular opening moderately restricted, extends ventrally to be- low a point anteriad of preopercular ridge. Gill membranes restricted, without a free fold over isthmus. Trunk short, distance isthmus to anal fin origin less than postrostral length of head. Periproct moderate in size, anus much closer to insertion of pelvic fins than to origin of anal fin. Ventral light organ well developed; a large, oval dermal window forms anterior extension of peri- proct; dermal window extends forward almost to a line connecting pelvic fin insertions. Gill filaments long, well developed; longest filament about 3.5-4.0 mm, about as long as diameter of eye lens. About 23 slender pyloric caeca. Teeth in moderately broad bands in both jaws; bands narrower posteriorly on jaws. Outer se- ries of teeth in upper jaw slightly enlarged. Scales of body covered with slender, conical, reclined spinules arranged in parallel to subpar- allel ridgelike rows that give a somewhat striated appearance to body surfaces. Four to five rows of spinules on larger body scales. Spinules on head scales more erect than those on body scales, and spinules often arranged in divergent rows. Scales behind bases of paired fins, behind first dorsal fin, and beneath outer margin of gill covers without spinules. Scales dorsally on sub- orbital region stout, adherent, and covered with coarse spinules. Scales at tip and lateral angles of snout large, tuberclelike, and coarsely spined. Exposed posterior tip of interopercle scaled. Mandibular rami, ventral surfaces of snout, and most of ventral surface of suborbital region na- ked; mucous pores large and prominent in these areas. There appear to be some scale pockets along lowermost branchiostegal rays of holo- type, but condition of specimen left doubts as to their actual presence (the ‘‘pockets’’ may simply be torn skin). Fins generally well developed. First dorsal fin high, length almost as long as head length (long- est ray broken at tip in holotype and paratype). Leading edge of spinous second ray armed with sharp, widely spaced denticles. Outer ray of pel- vic fin slightly prolonged, extends beyond origin of anal fin. Pelvic and anal fins originate rela- tively far forward; pelvic fin origin below sub- opercle, anal fin origin below hind third of first dorsal fin base. Holotype too badly faded for accurate de- scription of coloration. See color description of paratype. DESCRIPTION OF PARATYPE.—The single paratype is in poor condition and falling apart. The snout region is destroyed, and the posterior half of the tail has broken off and almost com- pletely disintegrated. The fins and scales are in- tact, however, and most of the diagnostic fea- tures of the species could be examined. The specimen is notably larger than the holotype (postrostral length of head 31.0 mm compared with 20.8 mm) and the two exhibit some onto- genetic differences. Thus, the spinules on scales are more numerous than in the holotype, and the larger spinules on body scales tend to be lanceolate rather than conical. Spinule rows on body scales are also more numerous, six to eight on the larger scales of paratype compared with four to five in comparable scales of holotype. The paratype had 29 pyloric caeca compared with the approximate count of 23 obtained in the holotype (I did not enlarge the original slit in the abdominal wall of the holotype for fear of dam- aging the specimen further). Two spinulated scales were present at the base of the left fourth branchiostegal ray, but none were on the op- posite member. Ground color of paratype dark brown with violet tinge over abdomen. Fin membranes, gill membranes, and naked surfaces of snout, sub- orbital region, and mandibular rami chocolate- brown. Gill cavity brown on inner wall and along periphery of outer wall but pallid everywhere else. Gill filaments and gill arches pale, the tu- bercular rakers dusky. COUNTS (paratype counts, if different, in brackets).—1D. II,10; 1P. 24 (left), 27 (right) [25, 25]; 2P. 15. Mesial gill-rakers on first arch 2 + 9 (2 + 10], on second arch 2 + 8 [2 + 9]. Scales below first dorsal about 12 [about 13]; below second dorsal 92; below midbase of first dorsal about 8 [about 914]; lateral line scales over dis- tance equal to predorsal length 41 [not taken in paratype]. Pyloric caeca about 23 [29]. MEASUREMENTS OF HoLotrypPe.—Total length 142 mm, head length 26.4 mm. The following in percent of head length: postrostral length of head 79.0; snout length 23.8; preoral length 20.4; internasal width 19.7; orbit diameter 36.7; inter- orbital width 22.0; postorbital length of head 38.2; distance orbit to angle of preopercle 34.0; IWAMOTO: EASTERN PACIFIC MACROURIDAE suborbital width 12.1; upper jaw length 27.2; barbel length 14.0; length first (outer) gill-slit 15.9; preanal length 129; isthmus to anus 47.4; outer pelvic ray to anal origin 34.0; greatest body depth 68; depth over anal origin 57; 1D.- 2D. interspace 34; length pectoral fin about 53; length outer pelvic ray about 53. COMPARISONS AND RELATIONSHIPS.—Ne- zumia ventralis appears most closely related to two Atlantic species, N. sclerorhynchus (Valen- ciennes, 1838) and N. suilla Marshall and Iwa- moto, 1973. The combination of general head and body shape, the presence of broad naked areas on the ventral surfaces of the snout and suborbital region, the naked mandibular rami, the large pores on the head, the general color- ation. and the placement and size of the fins sug- gest this relationship. The new species is readily distinguished from the two by its high pelvic fin ray count of 15, compared with 7 in suilla and usually 9 (rarely 7, 8, or 10) in sclerorhynchus. N. propinqua (Gilbert and Cramer, 1897) is the only other species of the genus having a pelvic fin ray count as high as that of N. ventralis, but N. ventralis lacks the distinct black blotch on the tip of the first dorsal fin that is so prominent in propinqua, and it has more pectoral fin rays (23 or fewer in propinqua), a shallower body, a more anteriorly placed pelvic fin, and fewer scale rows below the second dorsal fin. DISTRIBUTION.—The species is known from only two specimens taken by the ALBATROSS in the Galapagos Islands. ETYMOLOGY.—From the Latin ventralis, adj., of or belonging to the belly—in reference to the ventral light organ. REMARKS.—The paratype was first examined (circa 1920) by Dr. Carl L. Hubbs, who recog- nized the distinctness of the specimen and gave it the manuscript name of **Lionurus ventralis.” Because of other commitments, Dr. Hubbs was unable to formally describe the species at the time and consequently returned the specimen to the National Museum along with other macrou- rids collected by the ALBATROSS in the eastern Pacific in 1888 and 1904. MATERIAL EXAMINED.—Holotype: CAS-SU 68361, 26.4 mm HL, 142 mm TL, Galapagos Islands, off Hood (Espanola) I., 1°30'30’S, 89°39'W, 549 m, ALBATROSS sta. 4642, 7 Nov. 1904. Paratype: USNM 135345, 1 badly deteriorated specimen, 167 Galapagos Islands, 00°29’S, 89°54'30’W, 717 m, ALBATROSS sta. 2818, 15 Apr. 1888. Nezumia orbitalis (Garman) (Figure 14c) Macrurus orbitalis GARMAN, 1899:207—208, pl. 47, figs. 1-1b (original description; Gulf of Panama, ALBATROSS sta. 3385, in 523 m). Lionurus (Nezumia) orbitalis: GILBERT AND Huszss 1916:146 (listed). Nezumia orbitalis: MAKUSHOK 1967:table 18. DIAGNOsIS.—A species of Nezumia with 9 pelvic fin rays. Gill-rakers on first arch 1-2 + 8- 10 (10-11 total); on second arch 1-3 + 7-9 (9- 11 total). Spinules on body scales slender, con- ical to narrowly lanceolate; no enlarged middle row of spinules on scales. Barbel length 6-12 percent HL; upper jaw length 27-32 percent HL. First dorsal fin with distinct black tip. DESCRIPTION.—General features of fish seen in Figure 14c. A petite species probably not ex- ceeding 200 mm in total length. Head about 21- 25 percent of total length in specimens with complete tails. Snout pointed; head ridges rela- tively well developed with spinous tubercles at terminal and lateral points of snout. Dorsal pro- file arched high over nape to give distinctive humpbacked appearance. Gill membranes broadly attached across isthmus, with only a narrow posterior free fold. Anterior dermal win- dow of light organ small, circular, situated in a shallow fossa between bases of pelvic fins: anus close behind. Spinous second ray of first dorsal fin armed along leading edge with smail, closely spaced teeth. Outer pelvic ray slightly prolonged, ex- tends past origin of anal fin. Pelvic fin origin forward of pectoral fin origin, which in turn is slightly forward of first dorsal fin origin. Body scales with slender, conical to narrowly lanceolate spinules densely covering exposed fields, disposed in as many as 15 rows in large specimens, with spinule rows converging to- wards middle of field; no enlarged middle row of spinules. Most ventral surfaces of snout and suborbital region naked. Lower jaw surfaces na- ked except for a line of loose scales along mid- line of each ramus. Coloration. Garman (1899:208) has given a good color description from the types: **Black- ish over the body cavity; reddish brown over the muscular portions of the caudal section; lighter to translucent on the head and snout; en- 168 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 5 tire surface tinted with grayish, from the spi- nules of the scales. Fins light; first dorsal and pectorals tipped with black. Orbit surrounded by a narrow border of black.’’ The peritoneal lining of the lectotype is silvery with bold punc- tations. Dentition of jaws consists of small teeth in moderately wide bands with outer series of up- per jaw slightly enlarged. Counts (from 15 specimens).—1D. II,9-10 (11 in one specimen); 1P. 20-22 ( = 20.8); 2P. 9. Gill-rakers on first arch 1-2 + 8-10 (9-11 to- tal; x = 10.3); on second arch 1-3 + 7-9 (9-11 total; x = 10.2). Scales below first dorsal 7-9; below second dorsal 62 to 7; below midbase of first dorsal 4’ to 52; lateral-line scales over distance equal to predorsal length of head 36- 45. Pyloric caeca 17, 17, and 23 in three speci- mens. MEASUREMENTS (from 15 specimens).—Total length 115-186 mm; head length 27-41 mm. The following in percent of head length [range (x; 7; S.D.)]: postrostral length of head 73-78 (76.0; 14; 1.47); snout length 25-30 (27.8; 15; 1.27); preoral length 20-25 (22.9; 15; 1.81); internasal width 18-22 (19.9; 15; 0.97); orbit diameter 29- 34 (31.9; 15; 1.28); interorbital width 17-19 (17.7; 15; 0.72); postorbital length 39-45 (41.5; 13; 2.11); orbit to angle of preopercle 33-40 (34.5; 15; 1.96); suborbital width 11-14 (12.4; 15; 0.83); upper jaw length 27-32 (28.5; 15; 1.55); length barbel 6-12 (9.0; 13; 1.59); length outer gill-slit 14-18 (15.9; 15; 1.33); preanal length 119-146 (123.8; 15; 12.42); outer pelvic ray to anal origin 28-41 (33.9; 15; 5.19); anus to anal origin 14-29 (21.5; 15; 4.81); greatest body depth 63-82 (72.2; 15; 6.45); 1D.—2D. interspace 34-57 (44.1; 15; 7.35); height first dorsal fin 75-90 (80.4; 10; 6.15); length pectoral fin 48-78 (55.6; 12; 7.35); length pelvic fin 41-58 (49.2; 11; 4.49). RELATIONSHIPS AND COMPARISONS.—WNe- zumia orbitalis is closely related to N. latiros- trata and the two have narrow, coincident, geo- graphic ranges, although N. orbitalis is usually found at shallower levels than N. latirostrata (the two species were represented together in only one haul, SNP1-28, off Peru in 800 m, the deepest record for N. orbitalis). Meristic and morphological features of the two species are close, and the general appearance is similar. Color differences are the most noticeable. The first dorsal fin of N. orbitalis is distinctly pallid basally and black tipped distally, whereas in N. latirostrata the fin is uniformly blackish. The head of N. orbitalis is paler than that of N. la- tirostrata, and the prominent black orbital ring of the former is essentially lacking in the latter. The area ventrally on the naris appears smudged with black in N. /atirostrata but is scarcely pig- mented in N. orbitalis. Silvery reflections per- sist on the preopercle of N. orbitalis specimens but not in N. latirostrata specimens, and the belly denuded of scales has a more bluish cast in N. orbitalis. Nezumia latirostrata is generally a much darker fish. The low pelvic fin ray count, the number of gill-rakers, and the short barbel together serve to distinguish N. orbitalis from other eastern Pacific members of the genus. The species appears close to the widespread Atlantic species N. aequalis (Gunther), the two sharing many meristic, morphometric, and col- oration features. Even the humpbacked appear- ance of N. orbitalis is shared with one popula- tion of N. aequalis—that population being coincidentally also one from the eastern side of an ocean (the Gulf of Guinea) (see Marshall and Iwamoto in Marshall 1973:641). N. orbitalis is readily distinguished from N. aequalis, how- ever, by its somewhat shorter snout length (20- 25 percent HL cf. 25-32 in aequalis), its rela- tively longer head length compared with the to- tal length (head length 21-25 percent of total length, cf. 15-18), and its smaller adult size (probably less than 200 mm TL, as judged from the 175-mm length of the ripe female lectotype, cf. more than 270 mm TL in aequalis [Marshall and Iwamoto in Marshall 1973:643]). DISTRIBUTION.—Gulf of Panama to Peru, in 523-800 m. S1zZE.—To at least 41 mm HL and 186 TL. MATERIAL EXAMINED (15 specimens, 4 localities) —Pana- ma: MCZ 28578 (lectotype, 41 mm HL, 175+ mm TL), MCZ 50956 (5 paralectotypes, formerly MCZ 28578, 28-35 HL, 115+ to 153 TL), and USNM 57856 (2 paralectotypes, 28-34 HL, 129+ to 130+ TL), all from Gulf of Panama, 7°32'36’N, 79°16'W, 286 fms (523 m), ALBATROSS sta. 3385, 8 Mar. 1891. Peru: LACM 33885 (2, 40-40 HL, 174+ to 186 TL), 7°46’S, 80°31'W, 800 m, sta. SNP1-27, 23 Jan. 1974. —LACM 33886 (4, 27-39 HL, 118-178 TL), 7°44'05"S, 80°30'05"W, 750-760 m, sta. SNP1-28, 23 Jan. 1974. —IMARPE (1, 40 HL, 175+ TL), 13°57.2'S, 76°42.0’'W, 580-600 m, 7 Aug. 1972. Nezumia latirostrata (Garman) (Figures 4b, 15a, 18e) Macrurus latirostratus GARMAN, 1899:211—212, pl. 48, fig. 2 (original description; types from ALBATROSS Sta. 3354, 3384 and 3394 in Gulf of Panama, 322-511 fms [589-935 m)}). IWAMOTO: EASTERN PACIFIC MACROURIDAE Ficure 15. convergens (Garman), CAS 36816, 44 mm HL, from off Ecuador in 850—1,400 m. (c) Nezumia loricata atomos new subspecies. Holotype, CAS 38312, 51 mm HL, 297+ TL, from off Chile in 1,170—1,480 m. Lionurus (Nezumia) latirostratus: GILBERT AND HUuBBS 1916:146 (name only). Nezumia latirostrata: MAKUSHOK 1967:table 18. DIAGNosIs.—A species of Nezumia with 9- 10 pelvic fin rays. Gill-rakers on first and second arch 9-11 total (7-9 on lower limb). Barbel length 5-10 percent of HL; upper jaw length 27- 169 med vies Oars Ss S SS (a) Nezumia latirostrata (Garman), CAS 38330, 40 mm HL, from off Ecuador in 700-1,000 m. (b) Nezumia 32. Spinules on body scales lanceolate, arranged in indistinct rows without enlarged middle row. Scales below first dorsal 72 to 9; below second dorsal 6 to 7%. First dorsal fin uniformly black- ish. DESCRIPTION.—General features of fish seen in Figure 15a. Head and trunk moderately com- 170 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 5 pressed, tail greatly so. Head ridges prominent, suborbital ridge strongly angular. Mouth subter- minal; rictus extends to below anterior edge of eye lens; maxilla extends to below middle of eye. Terminal and lateral angles of snout armed with small but stout, spiny tubercular scales. In- terorbital region slightly concave, width about 60 percent of orbit diameter. Gill membranes broadly attached over isthmus (Fig. 4b), with a narrow posterior free fold; gill opening extends anteroventrally slightly anteriad of a vertical through posterior edge of preopercle. Body moderately deep in adults, generally more slen- der in the young; greatest body depth more than postrostral length of head in all specimens larger than about 30 mm HL. Light organ externally represented by a small, black, oblate, naked fos- sa between bases of pelvic fins and anterior to anus. Anus situated notably closer to insertion of pelvic fins than to ongin of anal fin. Gill fil- aments short; length of longest slightly shorter than diameter of eye lens. First dorsal fin of moderate height, length of longest ray less than head length; second spi- nous ray slightly produced into a thin filament; leading edge of ray armed with small, closely spaced teeth; in some specimens, tip of each tooth overlaps base of adjacent tooth. Paired fins of moderate size; outer ray of pelvic fin pro- duced, extends posteriad well beyond origin of anal fin. Body scales moderately deciduous, as judged from generally denuded state of specimens ex- amined. Spinules on body scales distinctly lan- ceolate, densely covering exposed fields, ar- ranged in indistinct longitudinal rows that converge towards middle of field. Scales absent on ventral surface of snout, but surface copi- ously dotted with dark sensory papillae. Sub- orbital region below ridge with a wedge-shaped cluster of small scales. Mandibles with small loose scales along midline of each ramus but naked at anterior end. Teeth small, in bands of moderate width in both jaws. Outer series of teeth in upper jaw scarcely enlarged. Coloration in alcohol swarthy. Bluish tinge over trunk and violet tinge over head and tail pronounced in denuded specimens. All fins dark brown to brownish black. Lining of oral cavity dusky, that of gill cavities black except for pale areas ventrally and along medial wall. Peritoneal cavity pale but peppered with large melano- phores. Counts (from 57 specimens).—1D. II,9-10 (11,8 in two specimens, II,11 in two others); 1P. 1823 @ = 20.65; n = 105; S-D! = T0222. 9 — 10. Gill-rakers on first and second arch 1-3 + 7— 9 (total 9-11). Scales below first dorsal 7/2 to 9 (rarely 7); below second dorsal 6 to 72; lateral- line scales over distance equal to predorsal length of head 32-38 ( = 35.88; n = 34;S.D. = 1.5). Pyloric caeca 17-25 @ = 21.23; n = 30; S.Dy = 2:4). MEASUREMENTS (from 56 specimens).—Total length 128-231 mm; head length 26-48 mm. The following in percent of head length [range (x; n;S.D.)]: postrostral length of head 71-77 (73.53; 56; 1.78); snout length 27-34 (29.97; 56; 4.36); preoral length 23-31 (26.16; 56; 1.78); internasal width 20—26 (23.39; 55; 1.23); orbit diameter 30— 36 (33.78; 55; 1.46); interorbital width 17-25 (20.84; 56; 1.57); postorbital length of head 36— 43 (38.94; 54; 1.52); orbit to angle of preopercle 33-39 (35.25; 55; 1.36); suborbital width 13-17 (15.18; 56; 1.11); upper jaw length 27-32 (29.55; 56; 1.30); length barbel 5-10 (8.19; 49; 1.57); length first gill-slit 11-16 (13.59; 54; 1.13); pre- anal length 123-157 (140.93; 56; 7.57); outer pel- vic ray to anal fin origin 27-48 (37.14; 55; 4.59); anus to anal fin origin 16-33 (22.27; 52; 3.78); greatest body depth 66-96 (77.22; 56; 11.01); 1D.-2D. interspace 34-55 (one specimen with 72) (45.41; 56; 6.30); height first dorsal fin 75—98 (83.78; 18; 6.10); length pectoral fin 52-68 (59.32; 50; 4.96); length pelvic fin 41-67 (55.57; 49; 6.92). COMPARISONS AND RELATIONSHIPS.—Ne- zumia latirostrata and N. convergens are closely related and specimens of the two are dif- ficult to differentiate. Chief differences between the two species lie in (1) gill-raker counts, (2) barbel length, (3) scales below first dorsal, (4) scales over distance equal to predorsal length, and (5) body length. Nezumia latirostrata gen- erally has 7-9 gill-rakers on the lower limb of the first and second arches, whereas N. conver- gens has S—7. Barbel lengths contrast at 5—10 percent HL for N. /atirostrata and 8—20 percent (12-18 in most) for N. convergens. Scale rows below the origin of the first dorsal fin are less numerous in latirostrata (74% to 9) than in con- vergens (82 to 10, a few individuals with 11). The number of anterior lateral-line scales over IWAMOTO: EASTERN PACIFIC MACROURIDAE a distance equal to the predorsal length is gen- erally lower in Jatirostrata (32-38 in most spec- imens compared with 36-42 in convergens), but there is considerable overlap with convergens in this feature. The body is somewhat shorter in latirostrata; the ratio HL:TL is between 1:4.5 and 1:5.5 (fewer than 12 percent of the speci- mens with an intact tail had a ratio of more than 1:5.2), whereas in convergens it is 1:5.3 to 1:5.8. Nezumia latirostrata also closely resembles N. loricata. The principal differences between the two rest in the pelvic fin ray counts (9-10 in latirostrata, 10-12 in loricata); barbel length (5— 10 percent HL vs. 10-17); number of scale rows below midbase of first dorsal fin (usually 4% to 5% vs. 64% to 7/2) and below the origin of the second dorsal fin (6 to 7/2 vs. 8-9); and the ratio of head length to body length (1:4.8 to 1:5.5 vs. feSaietorl<6:1): Nezumia latirostrata is readily distinguished from other eastern Pacific species of Nezumia by characters given in the key. Nezumia lati- rostrata is distinguishable from the Hawaiian species of the genus by a combination of char- acters that includes snout shape (blunter and shorter in holocentrus (Gilbert and Cramer, 1897); burragei (Gilbert, 1905); hebetata (Gil- bert, 1905); and ectenes (Gilbert and Cramer, 1897)) and pelvic fin ray count (more in propin- qua (Gilbert and Cramer, 1897) and obliquata (Gilbert, 1905)). Of the Japanese species report- ed by Okamura (1970a), all but four (proxima (Smith and Radcliffe, 1912, in Radcliffe), tomi- yamai (Okamura, 1963), burragei (Gilbert, 1905), and dara (Gilbert and Hubbs, 1916)) can be differentiated from /atirostrata by the num- ber of pelvic fin rays, but N. tomiyamai, N. bur- ragei, and N. dara have more scale rows below the origin of the second dorsal fin than does N. latirostrata, and N. proxima has a longer barbel (less than two times into orbit diameter com- pared with three or more times in /atirostrata). Most Atlantic members of the genus (see Mar- shall and Iwamoto in Marshall 1973:624-649) are distinguishable from N. latirostrata by a combination of pelvic fin ray count, barbel length, and shape of scale spinules. The western Atlantic species N. cyrano Marshall and Iwa- moto, 1973, is ‘probably most closely related to N. latirostratus’’ (ibid.:629), but the two differ in snout length (longer in cyrano, 32-36 percent HL, as opposed to 27-34, most specimens below 171 33, in latirostrata), upper jaw length (24-28 in cyrano Vs. 27-32 in latirostrata), and length of first gill-slit (6-10 vs. 11-16). DISTRIBUTION.—Panama (SE of Punta Mala) to northern Peru (SE of Lobos de Afuera) in 605— 1,400 m. S1zE.—To at least 48 mm HL and more than 234 mm TL. MATERIAL EXAMINED (182 specimens from 11 localities).— Panama: USNM 57855 (2 syntypes, 39-41 mm HL, 198+ to 205 mm TL), 7°21'N, 79°35'W, 935 m, ALBATROSS sta. 3394, 10 Mar. 1891. —GCRL 14259 (2, 47-48 HL, 215-198+ TL), 7°18'N, 79°38'W, 732-805 m, CANOPUS sta. 1285, 24 Mar. 1974. —GCRL 14260 (1, 41 HL, 167+ TL), 7°13'N, 79°18'W, 805-841 m, CANopus sta. 1291, 12 Mar. 1974. —CAS-SU 25246 (2, 44-46 HL, 195+ to 245 TL), ‘‘between Galapagos and Panama ALBATROSS 1888” (no other data). Ecuador: CAS-SU 24091 (8, 28-41 HL) and CAS-SU 25223 (9, 38-42 HL), 00°37'S, 81°00'W, 733 m, ALBATROSS sta. 2792, 2 Mar. 1888. —CAS 38328 (1, 42 HL, 175+ TL), 2°20’S, 81°16’W, 850-1,400 m, TE VEGA cr. 19, sta. 84, 4 Aug. 1968. —CAS 38330 (49, 25-45 HL, 133-215 TL), 2°25’S, 81°10’ W, 700-1,000 m, TE VEGA cr. 19, sta. 148, 1 Sep. 1968. —CAS 40826 (60, 38-45 HL, 190-234 TL), 3°15'S, 80°55’ W, 945-960 m, ANTON BRUUN cr. 18B, sta. 770 (LWK66-120), 10 Sep. 1966. Peru: USNM 118019 (1, 42 HL, 215+ TL), 5°47’S, 81°24’W, 1,030 m, ALBATROSS sta. 4653, 12 Nov. 1904. —LACM 33885 (2, 37-38 HL, 175-192 TL), 7°46’S, 80°31’'W, 800 m, sta. SNP1-27, 23 Jan. 1974. —CAS 38321 (35, 16-46 HL, 81-205 TL) and CAS 38322 (10, 27-46 HL, 100+ to 203 TL), 7°49’S, 80°38'W, 605-735 m, ANTON BRUUN cr. 18B, sta. 754 (LWK66-93), 5 Sep. 1966. In addition, the following ALBATROSS specimens were ex- amined but no measurements or counts were taken from them: MCZ 28570 (6 syntypes), sta. 3354; MCZ 28571 (10 syntypes), sta. 3384; and MCZ 28572 (3 syntypes), sta. 3394. Nezumia convergens (Garman) (Figures 156, 18f) Macrurus convergens GARMAN, 1899:210-211, pl. 48, fig. 1 (Gulf of Panama, 695-1,020 fms [1,271—1,865 m], ALBaA- TROSS sta. 3353, 3357, and 3393). Macrurus cuspidatus GARMAN, 1899:209-210 (Gulf of Cali- fornia, 27°34’N, 110°53'40’W, 905 fms [1,655 m], ALBA- TROSS sta. 3436). Macrurus trichiurus GARMAN, 1899:215 (Gulf of Panama, 555 fms [1,014 m], ALBATROSS sta. 3358). Lionurus (Nezumia) convergens: GILBERT AND HUBBS 1916:146 (listed). Lionurus (Nezumia) cuspidatus: GILBERT AND HUBBS 1916: 146 (listed). Lionurus (Nezumia) trichiurus: GILBERT AND HuBBs 1916:146 (listed). Nezumia convergens: MAKUSHOK 1967:table 18. Nezumia cuspidata: MAKUSHOK 1967:table 18. Nezumia trichiura: MAKUSHOK 1967:table 18. Sphagemacrurus trichiurus: MARSHALL 1973:623 (listed). DIAGNOosIs.—A species of Nezumia with pel- 172 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 5 vic fin rays 10-11. Gill-rakers on first arch 7-9 (S—7 on lower limb); on second arch 8-9. Spi- nules on body scales slender, conical to narrow- ly lanceolate, in discrete longitudinal rows, mid- dle row often enlarged. Barbel length 8-20 percent HL (usually 2.5—3.0 in orbit); upper jaw length 26-32. First dorsal fin uniformly blackish. DESCRIPTION.—General features of fish seen in Figure 15b. Head slightly compressed, ridges of head generally well developed; suborbital ridge prominent. Mouth subterminal, upper jaw extends posteriad to below middle of eye. Snout moderately pointed; terminal and lateral angles with stout tubercles armed with radiating rows of small spinules. Interorbital region flat to slightly concave, narrow, somewhat more than half orbit diameter. Gill membranes broadly at- tached over isthmus, with only a narrow pos- terior free fold; gill openings extend anteroven- trally to beneath posteroventral angle of preopercular ridge (Fig. 4b). Body slender; greatest body depth (at origin of first dorsal fin) less than postrostral length of head; 7-8 in total length in large adults. Light organ not well de- veloped externally; no scaleless fossa between pelvic fins. Anus within middle third of distance between pelvic fin insertion and anal fin origin. Gill filaments rather short, length of longest shorter than diameter of eye lens. Second spinous ray of first dorsal fin longer than postrostral length of head; leading edge armed with rather closely spaced, nonoverlap- ping slender, sharp teeth. Pectoral and pelvic fins originate about on same vertical, which is anterior to origin of first dorsal fin. Outer pelvic ray in adults extends slightly past anal fin origin. Second dorsal fin rudimentary throughout. Body scales somewhat deciduous. Slender, sharp, conical spinules uniformly cover exposed fields, aligned in slightly convergent longitudinal rows; spinules in middle row usually larger than those of lateral rows; 10-12 rows of spinules on scales of large adults. Almost all of ventral snout surface and part of anteroventral portion of sub- orbital region without scales. Mandibular rami covered posteriorly with narrow band of small, loose scales; mandibles generally naked ante- riorly, but liberally pocketed there with small pores. Dentition composed of small teeth in broad bands in both jaws; outer series of teeth in upper jaw not notably enlarged. Coloration in alcohol generally brownish black with hint of violet (especially on tail); bluish tinge over abdomen. Fins uniformly black or brownish black. Oral cavity dusky; peritoneal cavity flesh colored, often lightly peppered with large melanophores. Counts (from 80 specimens). —1ID. II,9-10 (rarely 8 or 11); 1P. 18-22 @ = 20.18; n = 142; S.D. = 1.02); 2P. 10-11 (rarely 9). Gill-rakers on first and second arch 1-2 + 5-7 (total 7-9: x = 8.04; n = 80; S.D. = 0.56). Scales below first dorsal 8'2—-10 (rarely 11); below second dor- sal 52-9 (usually 62-9); lateral line scales over distance equal to predorsal length of head 33-44 (usually 36-42). Pyloric caeca 21—32 (usually 25— 30)@ = 26.973 n= 3524S: Da2 ss MEASUREMENTS (from 80 specimens, 25 lo- calities, 17.5-51.6 mm HL, 97-300 mm TL), ex- pressed as percentage of head length [range (x; n; S.D.)]: postrostral length of head 69-76 (72.60; 79; 1.43); snout length 27-34 (30.46; 80; 1.37); preoral length 22-31 (26.99; 78; 1.59); in- ternasal width 17—23 (20.50; 70; 1.31); orbit di- ameter 31-37 (33.74; 80; 1.31); interorbital width 16-23 (19.73; 79; 1.29); postorbital length 34—40 (37.78; 78; 1.59); orbit to angle of preopercle 32— 40 (35.17; 77; 1.72); suborbital width 12-17 (14.41; 77; 0.96); upper jaw length 26-32 (28.83; 78; 7.37); length barbel 8-20 (14.08; 74; 2.64); length first gill-slit 11-19 (13.22; 76; 1.36); pre- anal length 133-181 (150.13; 73; 8.47); distance outer pelvic ray to anal origin 27-55 (40.11; 75; 5.24): vent to anal origin 14—28 (20.54; 66; 3.11); greatest body depth 60-78 (71.21; 64; 5.23); 1D.- 2D. interspace 28-68 (45.74; 75; 9.00); height first dorsal fin 68-93 (79.68; 53; 5.13); length pectoral fin 46-60 (52.30; 64; 3.52); length pelvic fin 36-61 (46.03; 59; 4.85). RELATIONSHIPS AND COMPARISONS.—In the eastern Pacific, Nezuwmia convergens appears closest to N. latirostrata (Garman), N. loricata (Garman), and N. orbitalis (Garman), but it is readily distinguished from these three by its few- er gill-rakers on the first arch (9 or fewer total, 5-6 [rarely 7] on lower limb, as compared with 9 or more total, 7-11 on lower limb). Spinules on body scales are generally more slender and conical than those of N. latirostrata and N. lor- icata, and they are arranged in discrete parallel rows with the middle row often enlarged (this compares with spinules in a somewhat quincunx arrangement or in rows that converge strongly toward the midline in the other two species). In addition, the body is somewhat more slender IWAMOTO: EASTERN PACIFIC MACROURIDAE than that of the other three species (greatest depth less than 75 percent of head length in most adults of N. convergens compared with more than 75 percent in most adults of other species), the anus is usually more posteriorly placed, often closer to the anal fin origin than to the insertion of the pelvic fins (always closer to the pelvic fin insertion in the others), and the ante- rior dermal window of the light organ is poorly defined externally (well defined in others). The barbel is slightly longer than that of N. latiros- trata, and the first dorsal fin is uniformly black- ish, not black blotched as in N. orbitalis. Nezumia convergens seems closely related to the western Atlantic species N. suilla Marshall and Iwamoto, 1973, with which it shares a close similarity in general physiognomy, coloration, scale spinulation, and extent of naked areas on the ventral surfaces of the head. The lower pel- vic fin ray count (7) and broader scale spinules in that species, nonetheless readily distinguish it from N. convergens. REMARKS ON SYNONYMY.—In my studies of the grenadiers described by Garman (1899), I have encountered three instances where he de- scribed different specimens of a single species- group taxon as different species. In the present instance, the nominal species concerned are Macrurus cuspidatus, M. convergens, and M. trichiurus. As first revisor, I have chosen to es- tablish the name M. convergens over the other two, primarily because the type-series for the species is large and representative, while M. cuspidatus and M. trichiurus were each de- scribed from single specimens, the last from a juvenile. The holotype of M. cuspidatus agrees well with the many well-preserved type-specimens of M. convergens except for its much deeper body. The entire trunk area of the holotype, in fact, appears to be abnormally swollen, as if it had been over-injected with preservative. If this is true, the deeper body is of no consequence. The Gulf of California capture of M. cuspidatus rep- resents the northernmost record of the species and the only record from the Gulf. The holotype of M. trichiurus is a young spec- imen (17.5 mm HL, 111 mm TL) slightly dam- aged ventrally. It agrees in all salient features with small specimens of N. convergens I have examined. Although Garman (1899:215) gave a pelvic fin ray count of 8 for the species, I count- ed 10 on both fins of the holotype. The gill-raker 173 count on that specimen was | + 6 on the mesial side of both the first and the second arches. DISTRIBUTION.—Nezumia convergens is one of the most numerous and widely distributed grenadier in the eastern Pacific Ocean. The species has the greatest latitudinal range of any Nezumia in the eastern Pacific except N. stel- gidolepis, although present collections show a broad gap between the Islas Tres Marias and Costa Rica. This gap is of little significance, however, in that deep trawling along the Central American coast between the Gulf of Panama and the Gulf of California has been extremely lim- ited. One Gulf of California record: others from off northern Mexico, Costa Rica, Panama, Co- cos Ridge, Galapagos, Ecuador, Peru, and Chile (to 34°53.5'S). Capture depths from 600 m to 1,865 m. S1zE.—To at least 50 mm HL and 300 mm TL. MATERIAL EXAMINED.—Mexico: MCZ 28575 (1, 52 mm HL; holotype of Macrurus cuspidatus Garman, 1899), Gulf of California off Guaymas, Sonora, 27°34'’N, 110°53’40’W, 1,655 m, ALBATROSS Sta. 3436, 22 Apr. 1891. —LACM 31124-4 (3, 43-45 HL, 250-270 TL), N of Islas Tres Marias, 21°52'30'N, 106°47'36"W, 800-550 m, VELERO IV sta. 13770, 21 Jan. 1970. Costa Rica: LACM 33588 (4, 28-SO HL, 161-295 TL), 9°45'18"N, 85°52'24"W, 1,865—1,372 m, VELERO IV sta. 18932, 12-13 May 1973. Panama: MCZ 28574 (1 damaged syntype), 7°15'N, 79°36'W, 1,865 m, ALBATROSS sta. 3393, 10 Mar. 1891. — GCRL 14258 (1, 26 HL, 132+ TL), 7°13'N, 79°18'W, 805-841 m, CANOPUS sta. 1291, 12 Mar. 1974. —MCZ 28573 (3 syn- types, 31-42 HL, 170+ to 220+ TL), 7°06'15’N, 80°34'W, 1,271 m, ALBATROSS sta. 3353, 23 Feb. 1891. —MCZ 28556 (1, 17.5 HL, 111 TL; holotype of Macrurus trichiurus (Gar- man, 1899), 6°30'N, 81°44’W, 1,006 m, ALBATROSS sta. 3358, 24 Feb. 1891. Cocos Island: AMNH 3471 (3, 29-43 HL), 96 kms of Cocos, ARCTURUS Sta. 74, May 1925. Galapagos: CAS 36814 (1, 47 HL, 248 TL), 27 km sseE of Isla San Cristobal, 1°06’S, 89°22'W, 700-800 m, TE VEGA cr. 19, sta. 102, 12 Aug. 1968. Ecuador: CAS 36817 (1, 47 HL, 265 TL), off Gulf of Gua- yaquil, 2°10’S, 81°13’W, 800-1,000 m, TE VEGA cr. 19, sta. 144, 31 Aug. 1968. —CAS 36816 (8, 19-44 HL, 101-252 TL), 2°20'S, 81°16’ W, 850-1,400 m, TE VEGA cr. 19, sta. 84, 4 Aug. 1968. —CAS 36815 (2, 22-30 HL, 108-175 TL), 2°25’S, 81°10’W, 700-1,000 m, TE VEGA cr. 19, sta. 148, 1 Sep. 1968. —CAS 36812 (35, 23-48 HL, 134—230+ TL), 3°15’S, 80°55’ W, ANTON BruUuN cr. 18B, sta. 770 (field no. LWK66-120). Peru: CAS 36813 (3, 40-43 HL, 205-230 TL), 4°10’S, 81°27'W, 1,815-1,860 m, ANTON BRUUN cr. 18B, sta. 766 (field no. LWK66-115), 9 Sep. 1966. —CAS 28760 (1, 21 HL, 97 TL), 17°08'05"S, 73°28'04"W, 860 m, 27 Jan. 1972. —IM- ARPE (1, 37 HL, 206+ TL), 18°17.3’S, 71°11'W, 600 m, 28 Jan. 1972. —CAS 28762 (1, 33 HL, 177 TL), 18°19’S, 71°12’ W, 810 m, 28 Jan. 1972. —IMARPE (1, 32 HL) (bottle label illeg- ible), 800 m, 1972. Chile: SIO 72-184 (3, 35-39 HL, 182—205+ TL), 18°40.4’S, 70°36.0'W, 768-967 m, THOMAS WASHINGTON, field no. 174 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 5 TABLE 3. RANGE, MEAN (x), AND STANDARD DEVIATION (S.D.) oF SELECTED MEASUREMENTS (VALUES IN PERCENT- AGE OF HEAD LENGTH) AND COUNTS OF Two SUBSPECIES OF NEZUMIA LORICATA (GARMAN). Sub- Character species n Range % S.D 1D. rays (total) loricata 11 12-14 12.73 0.8 atomos 9 11-13 11.78 0.8 IP. rays loricata 23 20-24 21.48 1.1 atomos 18 21-25 22.22 1.0 2P. rays loricata 24 «10-11 10.92 0.3 atomos 18 11-12 11.28 0.5 GR I (total) loricata 12 9-12 10.58 0.9 atomos 9 11-14 12.44 0.9 GR II (total) loricata 12 10-12 10.83 0.7 atomos N= 13 2256 n 057 Postrostral len. loricata 10 70-75 72.60 1.4 of head atomos 9 67-72 69.56 1.3 Snout length loricata 120 29=32 0/5 etal atomos 9 32-37 33.89 1.4 Internasal loricata 11 20-24 22.00 1.2 length atomos 922-251 23-56) 0 Interorbital loricata 12 19-24 21.00 1.4 width atomos 9: 22-26 23578 1-6 Orbit to loricata 12 32-36 33.92 1.0 preopercle atomos 9 3438 35.89 1.2 MV72-II-27, 7 May 1972. —CAS 36807 (6, 27-50 HL, 150+ to 287 TL), 24°29.5'S, 70°40’W, 950 m, ANTON BRUUN cr. 18A, sta. 714 (field no. LWK66—60), 16 Aug. 1966. —CAS 36810 (7, 34-41 HL, 184+ to 220 TL), 32°08.5'S, 71°43'W, 960 m, ANTON BRUUN cr. 18A, sta. 703 (field no. LWK66—-47), 12 Aug. 1966. —CAS 36809 (3, 36-42 HL, 218-220 TL), 32°17’S, 71°39.5'W, 580 m, ANTON Bruun cr. 18A, sta. 702 (field no. LWK66-44), 11 Aug. 1966. —CAS 36805 (28, 29-50 HL, 162- 300+ TL), 33°39’S, 72°09.5'W, 1,170-1,480 m, ANTON BRUUN cr. 18A, sta. 699 (field no. LWK66-41), 10 Aug. 1966. —CAS 36811 (2, 35-39 HL, 208-224 TL), 34°06.5'S, 72°18.5'W, 750 m, ANTON BRUUN cr. 18A, sta. 687 (field no. LWK66-25), 5 Aug. 1966. —CAS 36808 (8, 28-39 HL, 148+ to 203 TL), 34°53.5'S, 72°44'W, 780-925 m, ANTON BRUUN cr. 18A, sta. 698 (field no. LWK66—40), 9-10 Aug. 1966. Nezumia loricata (Garman) (Figures 15c, 16, 18g; Table 3) Macrurus loricatus GARMAN, 1899:208-209, pl. 47, figs. 2-2b (original description; specimens from ALBATROSS sta. 3409 and 3410, off Galapagos Islands, 327-331 fms). Lionurus (Nezumia) loricatus: GILBERT AND Huss 1916: 146 (listed). Nezumia loricata: MAKUSHOK 1967:table 18. DIAGNOsIS.—A species of Nezumia with 10- 12 (usually 11) pelvic fin rays. Gill-rakers on first arch 11—14 (8-11 on lower limb); on second arch 10-13 (8-11 on lower limb). Spinules on body scales moderately to broadly lanceolate; no en- larged middle row of spinules on scales. Barbel length 10-17 percent HL; upper jaw length 29— 33. First dorsal fin uniformly blackish. DESCRIPTION.—General features of fish seen in Figure 15c. Body rather deep, especially in adults; greatest depth at origin of first dorsal fin significantly greater than postrostral length of head. Head length varies from 17 to 20 percent of total length. Rictus extends to below middle of orbit; maxillary extends to vertical slightly behind middle of orbit. Tubercular scutelike scales at tip and lateral angles of snout spiny and stout; the terminal scute bifid. Interorbital space flat to slightly concave, relatively narrow, 1.3- 1.7 into orbit. Gill membranes broadly attached across isthmus with little, if any, free fold. Gill openings extend anteroventrally to a point slightly behind vertical through posterior margin of orbits. Dermal window of light organ small, roundish, situated between pelvic fin bases. Anus much closer to origin of pelvic fin than to origin of anal fin. Gill filaments short, length of longest less than half diameter of orbit. Second spinous ray of dorsal fin stout, armed on leading edge with sharp, pointed, reclined teeth; tip of ray slightly produced. Paired fins moderate in size; outer ray of pelvic fin extends beyond first three or four anal rays, other pelvic rays barely (or do not) reach anal fin origin. Body scales relatively adherent (compared with those of N. convergens and N. latirostra- ta). Spinules on body scales dagger-shaped to narrowly triangular, densely cover exposed fields, arranged in longitudinal series that con- verge towards middle of field. Scales absent on ventral surface of snout and posteriorly along part of suborbital region; naked areas dotted with sensory papillae, arranged in discrete rows and clusters in most specimens. Mandibular rami covered with small, thin, rather deciduous scales. Teeth small, in bands of moderate width in both jaws. Outer series of teeth in upper jaw slightly enlarged. Color in alcohol swarthy to medium brown. Ventral aspects of trunk and head much darker. All fins blackish. Oral cavity dusky. Peritoneal lining blackish and densely punctate in Chilean specimens (subspecies atomos), but more flesh colored with scattered large punctations in TE VEGA Galapagos specimens (subspecies /orica- jal). GEOGRAPHIC VARIATION.—I here recognize two subspecies of Nezumia loricata based on IWAMOTO: EASTERN PACIFIC MACROURIDAE differences in certain counts, measurements, and structural features enumerated and dis- cussed below. Key to the Subspecies of Nezumia loricata la. Terminal snout scute composed of two distinctly separated halves. Gill-rakers on first arch 9-12 total. Postrostral length of head 70-75% of head length ( = 72.6); snout length 26-32% (x = 30.8). Head length 17.6-20.0% of total length (¢ = 1920) Galapagos Islands: __.--.--._------_.-+-- Ree ee ee ee N. loricata loricata 1b. Each half of terminal snout scute closely adpressed, without a deep, median gap between them. Gill-rakers on first arch 11-14 total. Postrostral length of head 67- 71% HL (& = 69.6); snout length 32-37 (¢ = 33.9). Head length 16.6—-18.2% of to- fimtenethi(x,— 17.5). Off Chile__...__-____ Bee: N. loricata atomos, new subspecies The few specimens of restricted size repre- senting the populations off Chile and the Gala- pagos Islands limit comparisons, but there ap- pear to be sufficient differences in the study material to warrant formal recognition of the two populations. Frequency distributions of counts and measurements showing notable differences between the subspecies are enumerated in Table 3. Chief among the other differences is the shape of the terminal snout scute (Fig. 16)—in Gala- pagos specimens, the scute is composed of two coarsely spined halves medially split almost to the base, whereas in Chilean specimens, the scute is rather finely spined with the two halves closely adjoined medially, with little gap be- tween. The tubercular scales along the leading edge of the snout and at the lateral angles are also more coarsely spined in the Galapagos specimens, and the Chilean specimens have a broader double row of thickened scales along the suborbital region. The two populations of N. loricata are widely disjunct, and apparently, no material from in- tervening areas has been collected despite nu- merous trawl hauls at appropriate depths along the coasts of Ecuador, Peru, and Chile. The subspecific name atomos is derived from the Greek word meaning indivisible or uncut, in reference to the relatively undivided terminal snout scute of this subspecies. Counts (of both subspecies combined).—1D. 175 FIGURE 16. Nezumia loricata (Garman). Dorsal views of terminal snout scute in two subspecies: (a) N. loricata lori- cata; (b) N. loricata atomos. I1,9-12; 1P. 20-25; 2P. 10-12. Gill-rakers on first arch 1-3 + 8-11; on second arch 2 + 8-11. Scales below first dorsal 9 to 1242; below second dorsal 7/2 to 9; below midbase of first dorsal 6% to 7/2; over distance equal to predorsal length of head 35—41. Pyloric caeca 20-32 (¢ = 25.9, n = 9). MEASUREMENTS (of both subspecies com- bined, other than those given in Table 3).—Total length 175+ to 297+ mm, head length 34-52. The following in percent of head length: preoral length 25-30; orbit diameter 31-36; postorbital length of head 36—40; suborbital width 14-17; upper jaw length 29-33; length pelvic fin 44-68; length barbel 10-17; length outer gill slit 14-17; preanal length 140-157; base outer pelvic ray to anal origin 31-44; anus to anal origin 16-28; greatest body depth 67-87; height first dorsal fin 80-101; length pectoral fin 48-63. COMPARISONS AND RELATIONSHIPS.—Ne- zumia loricata appears most closely related to N. latirostrata, but the two are readily distin- guished by differences given in the key and in the description of N. latirostrata. Nezumia lor- icata may be distinguished from another close relative, N. convergens, by differences in gill- raker numbers, spinules on body scales, relative body depth, and placement of anus (see com- parisons in description of N. convergens). Other eastern Pacific species of the genus are differ- entiated from N. loricata by characters given in the key. DISTRIBUTION.—Galapagos (subspecies /ori- cata) and central Chile (subspecies atomos). S1zeE.—To at least 52 mm HL and 300 mm TL. MATERIAL EXAMINED.—Galapagos (subspecies /oricata): MCZ 28577 (4 syntypes, 45-55 mm HL, 215+ to 250 mm TL), and USNM 57859 (1 syntype. 47 HL, 240+ TL), 00°19’N, 90°34’ W, 331 fms (605 m), ALBATROSS Sta. 3410, 3 Apr. 1891. —MCZ 28576 (1 syntype, 48 HL, 260 TL), 00°18'40’N, 90°34’ W, 327 fms (598 m), ALBATROSS Sta. 3409, 3 Apr, 1881. 176 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 5 FIGURE 17. —USNM 135603 (remains of about 6 specimens, mostly dis- integrated), 00°29’S, 89°54'30’W, 392 fms (717 m), ALBATROSS sta. 2818, 15 Apr. 1888. —CAS 38329 (7, 34-50 HL, 175+ to 250 TL), 1°06’S, 89°22’W, 700-800 m, TE VEGA cr. 19, sta. 102, 12 Aug. 1968. —CAS-SU 25246 (2, 44-46 HL, poor con- dition), captured “‘between Galapagos and Panama, ALBA- TROSS 1888.” Chile (subspecies atomos): HoLotyPe, CAS 38312 (51 mm HL, 297+ mm TL) and 8 ParATyPeEs, CAS 43427 (47-51 HL, 270-290 TL), 33°39’S, 72°09’'W, 1,170-1,480 m, ANTON BRUUN cr. 18A (field no. LWK66-41), 10 Aug. 1966. Nezumia parini Hubbs and Iwamoto (Figures 17, 18a) Nezumia parini HUBBS AND IwAMoToO, 1977:246—250, fig. 8, 11, table 4. DIAGNOsIS.—A bathypelagic species of Ne- zumia with 11-12 pelvic fin rays. First and sec- ond gill arches each with 10-13 rakers. Small, circular body scales each with 1-15 long, erect, needlelike spinules. Scale rows below second dorsal about 11-14. Barbel length 7-13 percent HL; upper jaw length 31-36 percent HL; outer pelvic fin ray length 71-160 percent HL. Overall color dark brown to black. REMARKS.—General features of the fish can be seen in Figure 17. This distinctive species, described in detail in the original description, is the only Nezumia with bathypelagic adults, al- though in other species (e.g., N. stelgidolepis) the early-life stages are probably bathypelagic. The relationships of N. parini are obscure, but probably lie closest to the convergens-latiros- trata-loricata-orbitalis complex of species. The dense, fine, erect spinules on the small scales give a distinctive furry texture and feel to body surfaces. SIZE.—Nezumia parini is a small species at- p SARA atetannetoiecensearertmarrommemneer Rig meee Wueded eH~x S Ss < Nezumia parini Hubbs and Iwamoto, 1977. Paratype, CAS 29414, 35 mm HL, 216 mm TL, from off Peru. taining a head length of 35 mm and a total length of 216 mm. DISTRIBUTION.—Midwaters of the equatorial eastern Pacific and off the western shores of the Americas between Panama (7°30'N) and north- ern Chile (20°00’S). MATERIAL EXAMINED.—See original description. In addi- tion, one specimen recently discovered in the CAS-SU col- lections filed under unidentified Gadidae: CAS-SU 57651 (for- merly New York Zoological Society cat. no. 28704), (1, 35.6 mm HL, 202 mm TL), off Colombia, 4°45’N, 78°02’W, meter net fished over bottom depth of 500 fms (914 m), ZACA sta. 233, 3 Apr. 1938. ACKNOWLEDGMENTS I thank the many persons who helped in dif- ferent aspects of this study. The following helped in the loan of specimens and/or provided information and assistance during museum vis- its: T. Arai (Nippon Luther Shingaku Daigaku, Tokyo); N. Chirichigno-F. (IMARPE); D. M. Cohen (National Marine Fisheries Service, Washington, D.C.); C. E. Dawson (GCRL); D. Hubendick (Natural History Museum, Gote- borg); C. L. Hubbs (deceased), R. H. Rosenblatt and assistants (SIO); C. Karrer (formerly ISH); R. J. Lavenberg and assistants (LACM); R. Lee (University of Alaska); K. Leim and K. Hartel (MCZ); T. McLellan (formerly MCZ and Mas- sachusetts Institute of Technology); C. R. Rob- ins (UMML): C. L. Smith and G. Nelson (AMNH); V. G. Springer and assistants (USNM); M. Stehman (ISH) W. N. Eschmeyer examined and photo- graphed the holotype of Macruroplus potronus, critically reviewed the manuscript, and provided useful advice during the study. L. J. Dempster assisted in literature searches, reviewed the IWAMOTO: EASTERN PACIFIC MACROURIDAE = FiGuRE 18. Otoliths (left saggita) from (a) Nezumia parini, (b) N. pulchella, (c) N. liolepis, (d) Ventrifossa nigromacu- lata, (e) Nezumia latirostrata, (f) N. convergens, (g) N. lor- icata atomos, (h) N. pudens, and (i) N. stelgidolepis. Increments on scale bars equal 1 mm. manuscript, and advised on grammatical, no- menclatural, and editorial matters. O. Okamura reviewed the manuscript, offered advice and in- formation, and provided many profitable hours of discussions on the Macrouridae. K. P. Smith rendered the fine illustrations of Mataeocepha- lus tenuicauda (Fig. 8) and Nezumia ventralis (Fig. 14b). Curatorial and technical assistance was provided by J. E. Gordon, T. Keating, B. 177 Powell, W. C. Ruark, and P. Sonoda. My sin- cerest thanks to all. This study was supported in part by a grant from the National Science Foundation (BMS 075—03153), Tomio Iwamoto, principal investi- gator. LITERATURE CITED AGASSIZ, JOHN Louis R. 1843. Bericht iiber die fossilen Fische des Old Red Sandstone, in Auftrage der britischen Versammlung, im Jahre 1842, erstattet. Biblioth. Univ. Ge- neve, 1843, 43:353-369. ALCOCK, ALFRED W. 1891. On the deep-sea fishes collected by the ‘Investigator’ in 1890-91. Ann. Mag. Nat. Hist., ser. 6, 8:16—-34; 119-138, pls. vii—viii. ARAI, TAKAO. 1979. Additional information on a rare ma- crourid fish, Mesobius antipodum, from New Zealand. Japn. J. Ichthyol. 25(4):286—390, figs. 1-3. BARNHART, PERCY SPENCER. 1936. Marine fishes of southern California. Univ. Calif. Press, Berkeley. 209 pp. BERG, CARLOS. 1898. Substitucion de nombres genéricos. Comm. Mus. Nac. Buenos Aires 1:41-43. BLEEKER, PIETER. 1874. Typi nonnulli generici piscium neg- lecti. Versl. Akad. Amsterdam, ser. 2, 8:367-371. CHIRICHIGNO F., NORMA, AND TOMIO IWAMOTO. 1977. Cor- yphaenoides delsolari,a new species of macrourid fish from the Pacific coast of South America. Proc. Biol. Soc. Wash. 89(45):519-528. FitcH, JOHN E., AND ROBERT J. LAVENBERG. 1968. Deep- water teleostean fishes of California. Univ. Calif. Press, Berkeley. 155 pp. Fow Ler, HENRY W. 1944. Fishes of Chile. Systematic cat- alog. Apartado Rev. Chil. Hist. Nat. Santiago. GARMAN, SAMUEL. 1899. Reports on an exploration off the west coasts of Mexico, Central and South America, and off the Galapagos Islands, in charge of Alexander Agassiz, by the U.S. Fish Commission steamer ‘“‘Albatross’’ during 1891. Lieut.-Commander Z. L. Tanner, U.S.N., command- ing. Part 26, The Fishes. Mem. Mus. Comp. Zool. Harv. Coll. 24:1-431, pls. 1-84, A-N. GIGLIOLI, ENRICO H. 1882. New deep-sea fish from the Med- iterranean. Nature (London), 27(1882): 198-199. , AND A. IsseEL. 1884. Pelagos, saggi sulla vita et sul prodotti del mare. Genova. 433 pp. GILBERT, CHARLES H. 1890. A preliminary report on the fish- es collected by the steamer *‘Albatross”’ on the Pacific coast of North America during the year 1889, with descriptions of twelve new genera and ninety-two new species. Proc. U.S. Natl. Mus. 13(797):49-126. 1905. The deep-sea fishes of the Hawaiian Islands. Part 2, section 2, pp. 576-713, figs. 230-276, pls. 66-101, in Jordan, David Starr, and Barton Warren Evermann, The aquatic resources of the Hawaiian Islands. Bull. U.S. Fish Comm. (1903)22. , AND FRANK CRAMER. 1897. Report on the fishes dredged in deep water near the Hawaiian Islands, with de- scriptions and figures of twenty-three new species. Proc. U.S. Natl. Mus. 19(1114):403-435, pls. 36-48. , AND CARL L. Husss. 1916. Report on the Japanese macrouroid fishes collected by the United States fisheries steamer *‘Albatross”’ in 1906, with a synopsis of the genera. Proc. U.S. Natl. Mus. 51(2149): 135-214, pls. 8-11. . AND 1917. Description of Hymenocephalus 178 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 5 tenuis, a new macrouroid fish from the Hawaiian Islands. Proc. U.S. Natl. Mus. 54(2231):173-175. , AND . 1920. The macrourid fishes of the Phil- ippine Islands and the East Indies. U.S. Natl. Mus. Bull. 100, 1(pt. 7):369-588. , AND WILLIAM F. THOMPSON, 1916. Family Macrou- ridae, pp. 471-476, pls. 5-6, in Thompson, William F., 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. Proc. U.S. Natl. Mus. 50(2133):401- 476, pls. 2-6. Goope, G. BROWN, AND TARLETON H. BEAN. 1896. Oceanic ichthyology, a treatise on the deep-sea and pelagic fishes of the world .... Smithson. Contnb. Knowl. 1895[1896], 30(981), 1(text): 1-553; 2(atlas): 1-26, pls. 1-123. [Also is- sued as U.S. Natl. Mus. Spec. Bull. 2, and Mem. Mus. Comp. Zool. Harv. Coll. 22.]. GUNTHER, ALBERT. 1862. Catalogue of the fishes of the Bnt- ish Museum 4: 1-534. 1887. Report on the deep-sea fishes collected by H.M.S. CHALLENGER during the years 1873-76. Rep. Sci. Res. CHALLENGER 22(Zool.):pt. 1(text): 1-335, pt. 2(plates): pls. 1-73. Husss, Cart L., W. I. FOLLETT, AND LILLIAN J. DEMp- STER. 1979. List of the fishes of California. Occas. Pap. Calif. Acad. Sci. 133:1-S1. , AND ToMIO IWAMOTO. 1977. A new genus (Meso- bius), and three new bathypelagic species of Macrouridae (Pisces, Gadiformes) from the Pacific Ocean. Proc. Calif. Acad. Sci., ser. 4, 41(7):233-251, figs. 1-11. . AND KARL F. LAGLER. 1958. Fishes of the Great Lakes region. Rev. ed. Cranbrook Inst. Sci. Bull. 26:1-213. Iwamoto, Tomio. 1970. The R/V Pillsbury Deep-Sea Bio- logical Expedition to the Gulf of Guinea, 1964-65. 19. Mac- rourid fishes of the Gulf of Guinea. Stud. Trop. Oceanogr. Miami 4(pt. 2):316—-431, figs. 1-27. . 1972. Macrourid fishes of the tribe Malacocephalini (Macrouridae: Gadiformes). Ph.D. Thesis, Univ. Miami. . 1974. Nezumia (Kuronezumia) bubonis, a new sub- genus and species of grenadier (Macrouridae: Pisces) from Hawaii and the western North Atlantic. Proc. Calif. Acad. Sci., ser. 4, 3%22):507—516, figs. 1-3. . 1978. Eastern Pacific macrourids of the genus Coe- lorinchus Giorna (Pisces:Gadiformes), with description of a new species from Chile. Proc. Calif. Acad. Sci., ser. 4, 41(12):307—337, figs. 1-20. . AND DavipD L. STEIN. 1974. A systematic review of the rattail fishes (Macrouridae: Gadiformes) from Oregon and adjacent waters. Occas. Pap. Calif. Acad. Sci. no. 111:1-79. JORDAN, DAVID STARR, AND BARTON WARREN EVERMANN. 1898. The fishes of North and Middle America. Bull. U.S. Natl. Mus. 47(3):2183-3134. , AND CHARLES HENRY GILBERT. 1904. Macrouridae. Pp. 602-621 in Jordan, David Starr, and Edwin C. Starks, List of fishes dredged by the steamer Albatross off the coast of Japan in the summer of 1900, with descriptions of new species and a review of the Japanese Macrouridae. Bull. U.S. Fish Comm. (1902) 22:577-630, pls. 1-8. , AND EDwIn C. Starks. 1904. See Jordan and Gil- bert, 1904. KAMOHARA, TOSHUI. 1938. On the offshore bottom-fishes of Prov. Tosa, Shikoku, Japan. Tokyo. 86 pp. KOEFOED, EINAR. 1927. Fishes from the sea-bottom. Report on Scientific Results of the ‘‘Michael Sars’’ North Atlantic Deep-Sea Expedition. 1910, 4(Pt. 1):1-147, pls. 1-6. Lowe, RICHARD T. 1843. Notices of fishes newly observed or discovered in Madeira during the years 1840, 1841 and 1842. Proc. Zool. Soc. Lond. 11:91. MAKUSHOK, M. 1967. Whiptails (family Macrouridae or Cor- yphaenoididae auct.). Chapter IV, pp. 200-227 in V. G. Kort (chief ed.), Biology of the Pacific Ocean. Book III. Fishes of the open waters. Moscow. (Transl. from Russian.) U.S. Naval Oceanogr. Off., Transl. 528, Wash., D.C. MARSHALL, NORMAN B. 1973. Family Macrouridae. Fishes of the western North Atlantic. Mem. Sears Found. Mar. Res., no. I(pt. 6):496—665. , AND Tomio Iwamoto. 1973. Genus Nezumia. Pp. 624-649, figs. 39-47 in Marshall, Norman B. Family Ma- crouridae. Fishes of the western North Atlantic. Mem. Sears Found. Mar. Res., no. I(pt. 6):496—665. , AND NIGEL MERRETT. 1977. The existence of a ben- thopelagic fauna in the deep-sea. Pp. 483-497 in Martin, Angel, (ed.), A voyage of discovery: George Deacon 70th Anniversary Volume. Pergamon Press, Oxford. McCuLLocnH, ALLAN R. 1907. The results of deep sea in- vestigation in the Tasman Sea. II. The expedition of the ‘“Woy Woy.”’ |. Fishes and crustaceans from eight hundred fathoms. Rec. Aust. Mus. Syd. 6:345-355, pls. 63-65. . 1919. Check-list of the fish and fish-like animals of New South Wales, Parts 1 and 2. Australian Museum Syd- ney. 58 pp. 24 pls. MERRETT, NIGEL R. 1978. On the identity and pelagic oc- currence of larval and juvenile stages of rattail fishes (Fam- ily Macrouridae) from 60°N, 20°W and 53°N, 20°W. Deep- Sea Res. 25:147-160. NYBELIN, ORVAR. 1957. Deep-sea bottomfishes. Rep. Swed. Deep-Sea Exped. 1947-48, 2(Zool.)(20):247-345, pls. 1-7. OJEDA R., FEDERICO P., AND JUAN Camus Y. 1977. Mor- fometria y nicho trofico de Coelorhynchus patagoniae Gil- bert y Thompson (Pisces: Macrouridae). Bol. Mus. Nac. Hist. Nat. Chile 35:99-104. OKAMURA, OSAMU. 1963. Two new and one rare macrourid fishes of the genera Coelorhynchus and Lionurus, found in the Japanese waters. Bull. Misaki Mar. Biol. Inst., Kyoto Univ. 4:21-35. 1970a. Fauna Japonica. Macrourina (Pisces). Aca- demic Press of Japan, Tokyo. 216 pp., 64 pls. 1970b. Studies on the macrouroid fishes of Japan. Morphology, ecology and phylogeny. Rep. Usa Mar. Biol. Sta. 171(1-2): 1-179. ParRIN, N. V., V. E. BECKER, O. D. BORODULINA, AND V. M. TcHUvAssov. 1973. Deep-sea pelagic fishes of the southeastern Pacific Ocean [in Russian, English summary]. Tr. Inst. Okeanol., Akad. Nauk SSSR 94:71-159. , G. N. PoKHILskaya, Y. I. SAZONOV, AND B. I. FE- DORYAKO. 1977. Rare and poorly know midwater fishes from the central and eastern Pacific Ocean [in Russian, En- glish summary]. Tr. Inst. Okeanol., Akad. Nauk SSSR 104:206-236. PARR, ALBERT E. 1946. The Macrouridae of the western North Atlantic and Central American seas. Bull. Bing. Oceanogr. Coll. 10(1): 1-99. PEQUENO-R., GERMAN. 1971. Sinopsis de Macrouriformes de Chile (Pisces, Teleostomi). Bol. Mus. Nac. Hist. Nat. Santiago 32:269-298. PoLL, MAx. 1953. Poissons III. Téléostéens malacoptery- IWAMOTO: EASTERN PACIFIC MACROURIDAE giens. Rés. Sci. Expéd. Océonogr. Belge eaux cOotieres Africaines de |’ Atlant. Sud (1948-1949) 4(2): 1-258. RADCLIFFE, Lewis. 1912. See Smith and Radcliffe, 1912. REGAN, C. TATE. 1908. The Percy Sladen Trust Expedition to the Indian Ocean in 1905 under the leadership of Mr. J. Stanley Gardiner. No. XIV.—Report on the marine fishes collected by Mr. J. Stanley Gardiner in the Indian Ocean. Pp. 217-255, pls. 23-32. ROEDEL, PHIL M. 1951. Noteworthy southern California rec- ords of four species of marine fishes. Calif. Fish Game 37(4):509-S10. RouLe, Louis. 1916. Notice préliminaire sur quelques es- peces nouvelles ou rares des Poissons provenant des croi- sieres de S.A.S. le Prince de Monaco. Bull. Inst. Océanogr. Monaco 320: 1-32. SHCHERBACHEV, YU. N., YU. I. SAZONOV, AND A. S. Pio- TROVSKIJ. 1979. Occurrence of Trachonurus villosus (Gun- ther) and Mesobius (Hubbs et Iwamoto) (Macrouridae, Os- teichthyes) in the Indian Ocean. [In Russian.] Akad. Nauk SSSR [Problems in Ichthyology] 19:20-28, figs. 1-2, tables 1-2. SmitH, HuGH M., AND Lewis RADCLIFFE. 1912. Jn Rad- 179 cliffe, Lewis. Descriptions of a new family, two new genera, and twenty-nine new species of anacanthine fishes from the Philippine Islands and contiguous waters. Proc. U.S. Natl. Mus. 43(1924): 105-140, pls. 22-31. SMITH, J. L. B. 1949. The sea fishes of southern Africa. Central News Agency, Cape Town, 550 pp. (Also 1953, 1961, and 1965 editions.) . 1968. New and interesting fishes from deepish water off Durban, Natal and southern Mozambique. Oceanogr. Res. Inst. Invest. Rep. 19:1-30, pls. 1-6. VAILLANT, LEON L. 1888. Expédition scientifiques du Tra- vailleur et du Talisman pendant les années 1880, 1881, 1882, 1883. Poissons. Paris. Pp. 1406, pls. 1-28. VALENCIENNES, ACHILLE. 1838. Ichthyologie des iles Ca- naries. Pp. 1-109, pls. 1-27, in Philip Barker Webb and Sabin Berthelot, Histoire Naturelle des iles Canaries, Zool., tome 2, pt. 2. Bethune, Pans. WEBER, MAx. 1913. Die Fische der S1BOGA-Expedition. Si- boga Exped. 57:1-719, pls. 1-12. , AND L. F. DE BEAUFORT. 1929. The fishes of the Indo-Australian Archipelago, vol. 5. E. J. Brill, Leiden. 458 pp. sre ee SI Oe oe PAC IMC SY AINa Tin Re 3 A rerd vine Bere Wie ery AC POSIT, YORU Sem 7 = — - = - _ a i oo ar iN CASS fay © : - - ~~" 7 — a ==> (Se Ga = S a : a = = a=) BS) 4 oT ae * : tome inerne Bigtogical bdiperdttery LIBRARY PROCEEDINGS STE JUL 18 1880 CALIFORNIA ACADEMY OF SCIENCES Woods’ HOle, Mass: Vol. 42, No. 6, pp. 181-227, 17 figs. July 2, 1980 REVISION OF THE EASTERN PACIFIC SYNGNATHIDAE (PISCES:SYNGNATHIFORMES), INCLUDING BOTH RECENT AND FOSSIL FORMS By Ronald A. Fritzsche Department of Biology, The University of Mississippi, University, Mississippi 38677 Abstract: The marine and estuarine eastern Pacific Syngnathidae comprises 17 extant and 3 extinct species. Diagnostic characters for species and genera, including meristic and morphometric characters, and osteological features, are given. All species both living and fossil are diagnosed and described. The recognized species and their ranges are: Hippocampus ingens Girard [=H. hildebrandi] (San Francisco Bay, California, south to Pucusana, Peru, including the Gulf of California); Doryrhamphus melanopleura (Bleek- er) (Indo-Pacific; in the eastern Pacific from Bahia Magdalena, Baja California, south to Isla la Plata, Ecuador, including the Gulf of California, Galapagos Islands, and Clipperton Island); Doryrhamphus paulus n.sp. (Islas Revillagigedo, Mexico); Leptonotus blainvilleanus (Eydoux and Gervais) [=S. acicularis| (Hornitos, Chile, to Golfo Nuevo, Argentina); Bryx arctus (Jenkins and Evermann) (Tomales Bay, California, south to Mazatlan, Mexico, including the Gulf of California); Bryx heraldi n.sp. (Islas Juan Fernandez and Isla San Felix, Chile); Bryx coccineus (Herald) (Bahia Banderas, Mexico, south to Punta Aguja, Peru, and the Galapagos Islands); Bryx veleronis Herald (Galapagos Islands; Islas Revillagigedo; Isla Murcielago and Isla del Cano, Costa Rica; and Islas San Jose and Canal de Afuera, Panama); Bryx clarionensis n.sp. (Isla Clarion, Mexico); Syngnathus auliscus (Swain) (Santa Barbara Channel, California, south to Paita, Peru, including the Gulf of California); Syngnathus carinatus (Gilbert) (confined to upper Gulf of California); Syngnathus exilis (Osburn and Nichols) (Half Moon Bay, California, to Bahia Magdalena, Baja California, and Isla Guadalupe, Mexico); Syngnathus californiensis Storer (Bodega Bay, California, south to Bahia Santa Maria, Baja California); Syngnathus ma- crobrachium n.sp. (Tumbes, Peru, south to Puerto Montt, Chile); Syngnathus euchrous n.sp. (Redondo Beach, California, to Punta Eugenia, Baja California); Syngnathus leptorhynchus Girard [=S. griseolineatus, S. bar- barae| (southeastern Alaska south to Bahia Santa Maria, Baja California); Syngnathus insulae n.sp. (Isla Guadalupe, Mexico). Studies of growth and variation show that S. leptorhynchus is highly variable with each population distinct. Marked seasonal variation, when combined with growth data, indicates that individuals probably live for one year or less. The fossil pipefishes of Southern California are all only known from the Miocene. Hipposyngnathus imporcitor n.sp. from the upper Modelo Formation is most closely related to two species from the Oligocene of Europe. Syngnathus emeritus n.sp. is known only from the Puente Formation. Syngnathus avus Jordan and Gilbert is known from the lower Modelo Formation. A comparison between inferred relationships of the various species and their geographical distribution sug- gests that the evolution of the eastern Pacific Syngnathus is a result of at least two separate invasions. Doryr- hamphus melanopleura invaded the eastern Pacific by crossing the East Pacific Barrier and gave rise to Doryr- hamphus paulus. Leptonotus blainvilleanus is related to other species of Leptonotus in New Zealand and southern Australia and was probably derived from an ancestor in these areas via waif dispersal. The reduction and loss of elements of the branchial skeleton is useful in characterizing urophorine genera and may be of general use when relationships within the family Syngnathidae are studied in more detail. 181 182 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 6 INTRODUCTION Pipefishes of the family Syngnathidae inhabit most temperate and tropical seas. In the eastern Pacific, pipefishes occur from southeastern Alaska to Tierra del Fuego. They are primarily marine or euryhaline, but some species are con- fined to fresh water. Even though the family Syngnathidae con- tains the seahorses, of general interest to aquar- ists, the family has, in general, been poorly stud- ied. There are a number of undescribed species and the intrafamilial relationships are poorly understood, due, in part, to the great variability in meristic and morphometric characters. A revision of the Syngnathidae was published by Duncker (1915), but the American species were not included. Ginsburg (1937) and Herald (1940-1965) have been the major contributors to the taxonomy of the American syngnathids. To date, a definitive treatment of the eastern Pacific syngnathids has not appeared. Detailed osteological work on adult syngnath- ids is limited to that of Jungersen (1910), Rauther (1925), and Banister (1967). Jungersen’s study included the genera Hippocampus, Syngnathus, and Nerophis, two of which occur in the eastern Pacific. Rauther also studied the osteology of Syngnathus, as did Banister. The osteology of the other four genera of eastern Pacific syng- nathids has not been treated. It is generally rec- ognized that for the study of higher taxa, osteo- logical characters are a good indicator of phylogeny because of their conservative nature. The goal of this study has been to characterize the eastern Pacific genera and species of Syng- nathidae, both fossil and recent, and to examine intraspecific variation. In addition, a preliminary examination of the osteology of a few of the syngnathid genera was made to help in under- standing the relationships between at least a few of the genera, and as a test of the relationships proposed by Herald (1959) based on the method of brood-pouch closure. MATERIALS AND METHODS Specimens that have contributed to the data are listed in the Material Examined section for each species. In those lists, the number of spec- imens is given, followed by the size range (SL in mm) enclosed in parentheses. If the size range was not determined or the specimens are dam- aged, the range of standard length is not given. Abbreviations for listed collections ares AMS, the Australian Museum, Sydney; BC, Univer- sity of British Columbia: BMNH, British Mu- seum (Natural History); BOC, Bingham Ocean- ographic Collection, Yale University; CAS, California Academy of Sciences; EMBCh, Es- tacion de Biologia Marina, Chile; GCRL, Gulf Coast Research Laboratory Museum; HSU, Humboldt State University; IMARPE, Instituto del Mar, Peru; LACM, Natural History Museum of Los Angeles County; MCZ, Museum of Com- parative Zoology, Harvard University; MNHN, Muséum National d’Histoire Naturelle, Paris; MNMH P, Museo Nacional de Historia Natural, Santiago, Chile; SCCWRP, Southern California Coastal Water Research Project; SIO, Scripps Institution of Oceanography Marine Vertebrates Collection; SU, Stanford University (now housed at CAS); UCLA, Department of Zoolo- gy, University of California, Los Angeles; UMMZ, University of Michigan Museum of Zo- ology, USNM, National Museum of Natural History, Smithsonian Institution; UW, Univer- sity of Washington. Measurements were made with dial calipers to the nearest 0.1 mm for lengths less than 17 cm; an ocular micrometer was used for mea- surements less than 2 mm. Measurements great- er than 17 cm were made to the nearest mm with a centimeter rule. The principle characters used in identifying syngnathids are illustrated in Figures 1 and 2 (based on a generalized syngnathid). The ter- minology used for the various ridges on the body is that of Herald (1943). The condition of the lateral trunk ridge (Fig. 2) is important in characterizing syngnathid gen- era. It is subcontinuous with the superior tail ridge in Bryx and Syngnathus, but continuous in Leptonotus. In Hippocampus and Doryrham- phus the lateral trunk ridge is continuous with the inferior tail ridge. Clausen (1956) has shown that these ridge patterns are not wholly consis- tent within species, however, they are of value when used with other characters. Scutella (Fig. 1), small oval plates interpolated between the larger dermal plates, may be pres- ent or absent. Their size is important in species determinations and is therefore included in the species descriptions. Size is given as a compar- ison of an individual scutellum with an adjacent plate. FRITZSCHE: REVISION OF EASTERN PACIFIC SYNGNATHIDAE 183 TLL Patt FiGureE 1. Characters used in syngnathid identification (modified after Herald 1943). (A) Characters in the region of the head: a. snout ridge; b. supraorbital ridge; c. supraocci- pital, nuchal, and prenuchal ridges; d. opercular ridge; e. pec- toral cover plate ridges; f. ventral trunk ridge. (B) Characters in the region of the dorsal fin: g. superior trunk ridge; h. superior tail ridge; 7. lateral trunk ridge; /. inferior trunk ridge; k. anus; /. anal fin; m. inferior tail ridge; n. scutellum. A ring is defined as one unit in the series of dermal plates which form definite bands around the body. Pouch protecting plates are not illustrated. They are merely ventral extensions of the infe- rior trunk or tail ridge which support the brood pouch and protect the eggs. The pectoral cover plate is defined as the plate covering the base of the pectoral fin. The methods of making counts and measure- ments are those of Hubbs and Lagler (1958), with the following exceptions: Number of trunk and tail rings: The ring bearing the pectoral fins is counted as the first trunk ring; the ring bearing the anus is the last trunk ring. The ring bearing the anal fin is the first tail ring. If the anus and the anal fin are borne on the same ring, then that ring is the first tail ring and the preceeding ring is the last trunk ring. Number of dorsal and anal fin rays: last two rays are counted as two. Number of rings covered by dorsal fin: fractions c FiGuRE 2. Configuration of ridges above the anus (modi- fied after Herald 1943). (A) Lateral trunk ridge subcontinuous with superior tail ridge. (B) Lateral trunk ridge continuous with superior tail ridge. (C) Lateral trunk ridge continuous with inferior tail ridge. of a ring estimated to the nearest tenth of a ring. Number of lateral-line papillae: range of the number of lateral-line papillae per dermal plate. For Hippocampus the standard length is the distance from the tip of the coronet, with the head held perpendicular to the trunk, to the tip of the tail, with the tail held straight (Ginsburg 1937). In species diagnoses the modal range of a par- ticular meristic character is given in parentheses after the range of that character. Osteology was studied from trypsin-digested and alizarin-stained specimens (Taylor 1967) listed below. Osteological nomenclature follows Jungersen (1910) and Banister (1967). Bones were drawn with the aid of a camera lucida. Diagnoses are based on adults unless other- wise noted. 184 ’ PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 6 Statistical procedures follow the text of Dixon and Massey (1957) except that the regression analyses are based on the Bartlett regression analysis as given by Hoese (1971). Unless oth- erwise specified, differences are considered sig- nificant at P < .05. CLEARED AND STAINED MATERIAL EXAMINED.—Heraldia nocturna, SIO 75-54, 1, Sydney, Australia; Maroubra perser- rata, SIO 75-53, 1, Sydney, Australia; Choeroichthys brachy- soma SIO 73-196, 1, Timor Sea; Doryrhamphus melanopleu- ra, SIO 65-343, 1, Gulf of California; Dunckerocampus dactylophorus, CAS uncat., 1, unknown; Dentirostrum jans- si, CAS 14148, 3 paratypes, Thailand; Syngnathoides biacu- leatus, SIO 61-693, 1, 10°N, 103°50’E; Leptonotus blainvil- leanus, USNM 176569, 1, Isla Chiloé, Chile; Leptonotus blainvilleanus, USNM 176564, 1, Bahia Lin, Chile; Parasyng- nathus elucens, CAS 13696, 1, Virgin Islands; Ichthyocampus belcheri, CAS uncat., 2, Philippine Is.; Micrognathus brevi- rostris, SIO 73-196, 4, Timor Sea; Penetopteryx taenio- cephalus, SIO 66-587, 1, Great Tulear Reef; Syngnathus cari- natus, S. Guevarra pers coll., 1, Gulf of California; S. acus, SIO 73-310, 1, Yugoslavia; S. auliscus, SIO 68-168, 1, Sonora, Mexico; S. californiensis, S1O H47-180, 1, Santa Cruz Island; S. leptorhynchus, E. B. Brothers pers. coll., 1, Mission Bay; S. pelagicus, SIO 65-358, 1, western Atlantic; Bryx veleronis, SIO 71-52, 1, Panama; B. clarionensis, SIO 74-116, 1, para- type, Clarion Is.; B. dunckeri, SIO 70-376, 1, Panama; B. arctus, SIO H52-218, 1, Baja California; B. coccineus, USNM 220972, 1, Galapagos Is.; Cosmocampus brachycephalus, CAS 24025, 2, Panama; Corythoichthys flavofasciatus, R. Nolan pers. coll., 1, Eniwetak; Corythoichythys sp., SIO 73- 206, 1, Timor Sea, Pseudophallus starksi, USNM 208371, 2, Panama; P. elcapitanensis, USNM 208369, 1, Panama; Hip- pocampus kuda, S1O-60-250, 1, Hawaii; H. ingens, NMFS uncat., 1, eastern Pacific; Phyllopteryx foliatus, S1O 73-361, 1, Australia. SYSTEMATICS Family Syngnathidae Bonaparte, 1838 Type-Genus: Syngnathus Linnaeus, 1758. DIAGNOsIS.—Syngnathiforms with body en- cased in armor formed of dermal plates arranged in rings; pelvic and spinous dorsal fins absent; dorsal and pectoral fins moderately developed to absent; caudal and anal fins weakly developed to absent (tail often prehensile when caudal fin absent); opercular slit reduced to a small open- ing at dorsoposterior margin of opercle, four complete gill arches bearing lobate gills; pseu- dobranchiae present; supracleithra and post- cleithra absent; ribs absent; teeth lacking on jaws, but premaxillae and dentaries may bear odontoid processes (Dawson and Fritzsche 1975); pharyngobranchial tooth plates present or absent; upper part of cleithrum forms part of the external armor; posttemporals suturally united to neurocranium; I—3 branchiostegals; no basisphenoid; no metapterygoid; eggs incubated by males in a special area under the trunk or tail, which may or may not be developed into a pouch; no pyloric sphincter; no distinct stom- ach; right kidney only present, aglomerular; pre- dorsals reduced to 2-3 nuchal plates. The family is usually divided into two groups: Gastrophori—those which develop the brood pouch under the abdomen; Urophori—those with the brood pouch under the tail. Key to Genera of Eastern Pacific Syngnathidae la. Caudal fin absent; tail prehensile; head at right angle to main body axis________________ Ne eet aE WA 9, Hippocampus Rafinesque 1b. Caudal fin present; tail never prehensile; head in line with main body axis __________ 2 2a. Brood pouch under abdomen 2b. Brood pouch under tail 3a Trunk rings more numerous than tail Doryrhamphus Kaup 3b. Trunk rings fewer than tail rings __- Oostethus Hubbs 4a. Lateral trunk ridge continuous with su- perior tail ridge; brood pouch without protecting plates; mature females with deep, compressed trunk === Leptonotus Kaup 4b. Lateral trunk ridge subcontinuous with superior tail ridge; brood pouch with pro- tecting plates; mature females with sub- cylindrical trunk 5a. Dorsal rays 23 or fewer; snout short, con- tained 2.3—4.0 times in head; anal fin pres- ent or absent; small, never longer than 124 smmsS, 17 eee Bryx Herald 5b. Dorsal rays 26 or more; snout longer, con- tained 1.6—2.8 times in head; anal fin pres- ent; larger, most species reach 200 mm or more Sh..2 aes Syngnathus Linnaeus Records of Doubtful Validity Regan (1908) included Oostethus brachyurus (Bleeker) and Syngnathus spicifer Ruppell in his list of the fishes occurring at Tehuantepec, Mex- ico. Herald (1940) repeated these records but doubted their validity. Herald (1943) indicated that there was considerable doubt that these two species were collected at Tehuantepec, because he had been able to confirm that Regan’s spec- imens had come from an animal dealer. Duncker (1915) had also realized that the specimens had FRITZSCHE: REVISION OF EASTERN PACIFIC SYNGNATHIDAE 185 come from an animal dealer, but the records have persisted. The animal dealer had received specimens from both Mexico and the Philip- pines. O. brachyurus and S. spicifer occur in the Philippines, and it seems most probable that the specimens were collected there. There is, however, a valid record of Oostethus from the eastern Pacific. McCosker and Dawson (1975) reported a single individual of the eury- haline Atlantic Oostethus lineatus collected on the Pacific side of Panama in 1971 and concluded that the specimen represents a transit of the Pan- ama Canal. Dermatostethus punctipinnis Gill (1863) was described from four specimens collected at San Diego. Presently there are three type-specimens at the USNM (lectotype, here designated as USNM 8128; paralectotypes here designated as USNM 214484). Much confusion has been gen- erated by these specimens. The types are quite large (302-344 mm SL), have a very flexible ‘“‘neck,’’ and dark spotting on the dorsal fin. Herald (1940, 1941) referred D. punctipinnis to the synonymy of Syngnathus californiensis be- cause the latter is a large species with counts like that of the type material of D. punctipinnis. Miller and Lea (1972:212) suggested that D. punctipinnis might be a valid species, but did not include it in their key to California species. I have examined the types of D. punctipinnis and have compared them to specimens of S. acus from Europe and have been unable to find significant differences. I therefore relegate D. punctipinnis to the synonymy of S. acus. Syngnathus acus has been recorded from the Indo-Pacific (Weber and De Beaufort 1922), and my examination of a specimen from that region (Pakoi, China) suggests the existence of a dis- tinct acus-like species. But the putative Califor- nia material is typical of acus and could not rep- resent trans-Pacific migration. Dumeril (1870) described Syngnathus bairdi- anus from a locality given as ‘‘Cote du Mexique, voisine de la Californie.’’ The locality as listed in the catalog of the MNHN is ‘‘Mexique pres la Californie, Lucas 1867.’ I examined the two types (lectotype, here designated as MNHN 6112; paralectotype, here designated as MNHN 2756) and found them to be indistinguishable from S. pelagicus Linnaeus. I therefore relegate S. bairdianus to the synonymy of S. pelagicus. S. pelagicus inhabits the Sargasso Sea and ap- parently can be transported over long distances. It has been recorded from Tierra del Fuego (Fowler 1944) and from New Zealand (Weber and De Beaufort 1922). Histrio histro, another inhabitant of the Sargasso Sea, is widely distrib- uted and has been recorded from the Galapagos Islands (Schultz 1957). It is therefore remotely possible that S. pelagicus could have been col- lected near the coast of Mexico. None of the above species will be treated fur- ther because of their doubtful standing as mem- bers of the eastern Pacific fauna. Hippocampus Rafinesque Hippocampus RAFINESQUE, 1810:18 (type-species by mono- typy, H. heptagonus Rafinesque [=Syngnathus hippocam- pus Linnaeus]); DUNCKER 1912:237 (diagnosis); 1915:115 (diagnosis); GINSBURG 1937:525 (diagnosis; discussion). Farlapiscus WHITLEY, 1931:313 (type-species by original des- ignation, Hippocampus breviceps Peters). Hippohystrix WHITLEY, 1940:44 (type-species by original des- ignation, Hippocampus spinosissimus Weber). DIAGNosis.—Urophori characterized by a prehensile tail; absence of caudal fin, scutella, brood pouch protecting plates, and basibranchi- als; head at right angle to trunk; brood pouch sealed along midline except for small anterior opening; occiput raised to form coronet; dorsal fin base raised. DIsTRIBUTION.—Marine; world-wide in trop- ical and subtropical seas. Hippocampus ingens Girard (Figures 3 & 7C) Hippocampus ingens GIRARD, 1859:342 (original description; San Diego, California) (lectotype here designated as USNM 982): JORDAN AND GILBERT 1880:23 (San Diego); 1881:453 (San Diego); JORDAN AND Jouy 1881:1 (California); Jor- DAN AND GILBERT 1882:69 (San Diego); 1883:386 (descrip- tion; range); EVERMANN AND JENKINS 1891:127, 135 (Guay- mas, Mexico); GILBERT 1891:450 (ALBATROSS sta. 2795); EIGENMANN AND EIGENMANN 1892:144 (San Diego); VAIL- LANT 1894: 70 (Gulf of California); JORDAN 1895:417 (de- scription; Mazatlan, Mexico); JORDAN AND EVERMANN 1896:776 (synonymy; description); GILBERT AND STARKS 1904:57 (Panama Bay); GILL 1905:807 (range); STARKS AND Morris 1907:186 (range); OSBURN AND NICHOLS 1916:155 (Concepcion Bay); NICHOLS AND MurpuHy 1922:506 (Peru); MEEK AND HILDEBRAND 1923:256 (description; synony- my); BREDER 1928:23 (Cape San Lucas, Concepcion Bay); ULREY AND GREELEY 1928:41 (synonymy; range); ULREY 1929:6 (Lower California, Gulf of California); 1932:77 (Baja California); GINSBURG 1937:534 (range; synonymy; descrip- tion); ATz 1937:62 (size); FOWLER 1944:496 (range): KoepcKE 1962:200 (references; range); CLEMENS AND NowELL 1963:262 (off Mexico; in stomachs of fish); HUBBS AND HINTON 1963:12 (California record; range); CHIRI- CHIGNO-F. 1963:8, 34 (Peru; range); CASTRO-AGUIRRE ET AL. 1970:132 (common in Gulf of California); MILLER AND 186 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 6 FIGURE 3. 1860 at San Francisco, California, by Alexander Agassiz (not examined). Hippocampus ingens, MCZ 35292, collected in LEA 1972:89 (description; range); CHIRICHIGNO-F. 1974: 86, 337 (Peru; in key; range). Hippocampus gracilis GiLL, 1863:282 (original description; Cape San Lucas, Baja California). Hippocampus ecuadorensis FOWLER, 1921:446 (original de- scription; Bahia, Ecuador). Hippocampus hildebrandi GiNSBURG, 1933:562 (original de- scription; Chame Point, Panama); holotype USNM 82063): 1937:579 (description; range). DIAGNOsIS.—A Hippocampus with 17—22 (19) dorsal rays; 1S—17 pectoral rays; 4—S anal rays; 11-12 (11) trunk rings; 37-41 (39) tail rings; dor- sal fin on 1.2—2 + 1.5—4 rings; total rings cov- ered by dorsal fin 3—4; brood pouch on 5-8 rings; head 4.2—5.8 in SL; dorsal fin base 1.8—2.9 in head; snout 1.8—2.5 in head; maximum size ex- amined 247 mm SL; smallest mature male 54 mm, said to attain 12” (297 mm) (Miller and Lea 1972), which is approximately the size of my largest specimen if the head is included in the measurement. DESCRIPTION.—Body ridges typically well de- veloped with a blunt recurved tubercle at center of each plate. First, fourth, seventh, and tenth trunk rings usually with larger and better devel- oped tubercles. Sixth, tenth, fourteenth, and eighteenth tail rings may have strongly devel- oped tubercles, but are usually more obsoles- cent than trunk tubercles. Coronet with five ra- dially arranged tubercles. Males with the more weakly developed tubercles. Tubercles in both sexes generally become obsolete with growth. Strong nuchal ridge, without spines. Opercle with faint radiating striae. No prominent snout ridge. Internasal spine blunt. Prominent broad- based supraopercular spine directed laterally. Lateral trunk ridge and superior tail ridges over- lap for one ring. Trunk rings octangular under dorsal fin, heptangular anteriorly. Trunk mod- erately compressed. First tail ring heptangular, remainder quadrangular. Superior trunk and tail ridges overlap for two to three rings. Dermal flaps, when present, on enlarged tubercles and head ridges, consisting of a stout base with nu- merous filamentous outgrowths. Coloration in life. Red, yellow, or green. One specimen captured at La Paz, Baja California, had undersides and tips of tubercles yellowish, most of body mottled with dark brown to black, and covered with many small dark spots as well as smaller white ones. The white spots tend to coalesce into longitudinal streaks; yellowish col- oration more pronounced on underside of tail. White bands around body every six or seven rings. Dorsal fin with distinct dark submarginal band. Pectoral fins hyaline. Coloration in alcohol. Enlarged tubercles usu- ally whitish, often with white ring around body at that point. Ground color dark brown with small white papillae often forming streaks and reticulations, and radiating lines around orbit. Ventral surface of tail without white markings. Dorsal fin with a dark band near margin. Median abdominal ridge often dark brown in males. However, color variable; some specimens may be uniform light tan. HaBITAT.—Collections of H. ingens are un- common. Some have been made in shallow FRITZSCHE: REVISION OF EASTERN PACIFIC SYNGNATHIDAE 187 water, but most specimens have been captured at depth with dredges or trawls, or at the surface in the open ocean. Dredge and trawl collections have usually been made at depths of 10 m or more. Juveniles (ca. 40 mm) and larger individ- uals (115 mm) are not uncommonly taken at the surface. Alverson (1963) studied the food items in stomachs of eastern Pacific yellowfin tuna, Thunnus albacares, and found H. ingens in 18 stomachs out of the 2846 he examined. Judging from the small displacement volumes of these fish, they were all juveniles. One 34-mm indi- vidual (SIO 71-186) was taken from the gut of a bluefin tuna, Thunnus thynnus. The habitat is not yet precisely known. H. ingens appears to spend much of its life in the open sea. COMPARISONS.—Ginsburg (1933) described H. hildebrandi from Chame Point, Panama, as differing from H. ingens in having lower, broad- er tubercles. My examination of 38 specimens from the eastern Pacific has shown that all spec- imens are referrable to H. ingens. | have ex- amined the types of H. ingens and H. hilde- brandi and conclude that the types of H. hildebrandi are juveniles of H. ingens. Since H. ingens is the only species of sea- horse in the eastern Pacific, it is easily identifi- able. The closest relative of H. ingens is pos- sibly H. reidi Ginsburg from the western Atlantic, from which it differs in number of dor- sal rays (17-22 vs 15-19) and in number of tail rings (36-41 vs 34-37). The relationship of H. ingens to seahorses of the Indo-Pacific is im- possible to determine because knowledge of the genus Hippocampus is very incomplete. RANGE.—San Francisco Bay, California, south to Pucusana, Peru, including the Gulf of California. Infrequently taken north of central Baja California. During periods of unusually warm water, H. ingens may enter California waters. MATERIAL EXAMINED.—California: San Diego, USNM 982, 1(167), lectotype; USNM 214485, 2(125—146), paralecto- types; Point Loma, SIO 63-1085, 1(195). Baja California: Laguna Guerrero Negro, SIO 61-10, 3(155— 185). Bahia Sebastian Vizcaino, SIO 63-1046, 1(29). Bahia Magdalena, SIO 62-713, 1(108); SIO 64-73, 1(27): SIO 60-305, 1(55). Punta Hughes, SIO 64-45, 1(109). Gulf of California: La Paz, SIO 74-81, 1(200). Punta Man- gles, SIO 65-335, 1(122). Isla Santa Inéz, SIO 65-306, 1(99). San Felipe, SIO 67-1, 1(168). Mazatlan South: Boca Teacapan, SIO 60-90, 1(118). Nay- arit, SIO 60-89, 1(92). Bahia de Banderas, SIO 62-29, 1(148). Acapulco, UCLA WS52-119, 1(70). Golfo de Tehuantepec, SIO 73-258, 5(116—-136); SIO 68-16, 1(47); SIO 63-1031, 1(1i5); SIO 72-123, 1(80). Guatemala, SIO 63-623, 1(70); UCLA W56-273, 1(120). Costa Rica, UCLA WS54-139, 1(114). Panama, SIO 7}- 260, 1(115); SIO 71-186, 1(36); USNM 82063, 1(66), holotype of H. hildebrandi; USNM 82037, 1(44), paratype of H. hil- debrandi; USNM 82039, 1(47), paratype of H. hildebrandi. Galapagos Islands: BC 56-440, 2(238-247); SIO 54-174, 1(128). Isla Santa Cruz, SIO H51-388, 1(89). Doryrhamphus Kaup Doryrhamphus Kaup, 1853:233 (nomen nudum); 1856:54 (type-species by monotypy, D. excisus Kaup; Red Sea); DUMERIL 1870:585 (description); JORDAN AND EVERMANN 1896:773 (in part; description); DUNCKER 1912:231 (descrip- tion; synonymy); 1915:244 (description; synonymy); Jor- DAN, EVERMANN, AND CLARK 1930:243 (synonymy): HER- ALD 1953:244 (description; synonymy). Pristidoryrhamphus FOWLER, 1944:158 (type-species by orig- inal designation, P. jacksoni Fowler =Doryrhamphus ne- grosensis Herre). DIAGNOos!Is.—Gastrophori with lateral trunk ridge continuous with inferior tail ridge; inferior trunk and tail ridges discontinuous; superior trunk and tail ridges discontinuous; trunk rings more numerous than tail rings; snout ridge strongly serrate, markedly so in mature males; each dermal plate armed with strong retrorse spine; no pouch-protecting plates; caudal fin large and brightly colored; branchial skeleton with all elements present; scutella present; two nuchal plates; three infraorbitals; strongly de- veloped opercular ridge; 14-19 trunk rings; 10- 17 tail rings; 21-29 dorsal rays; 4 anal rays; 10 caudal rays. RANGE.—Four or five species ranging throughout the tropical Indo-Pacific among coral and rocky reefs. DiscUSsSION.—Kaup first published the name Doryrhamphus in 1853, but his reference to D. excisus Hemprich and Ehrenberg cannot be tak- en to be an indication as defined by the /nter- national Code of Zoological Nomenclature Art. l6a(v). The species D. excisus was an unpub- lished manuscript name in 1853. The require- ments of the Code were not fulfilled until Kaup (1856) published descriptions of Doryrhamphus and D. excisus. Key to Eastern Pacific Species of Doryrhamphus la. Trunk rings 16-18 (usually 17) tail rings 14-17 (usually 15); head 4.0-4.4 in SL _ Se eee ee ee melanopleura (Bleeker) Tropical Indo-Pacific 1b. Trunk rings 16-17 (usually 16); tail rings 188 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 6 FiGureE 4. Anterior part of body of four species of eastern Pacific Syngnathidae. (A) Leptonotus blainvilleanus, a 168-mm- SL female, SIO 72-168. (B) Doryrhamphus paulus, holotype, SIO 72-67. (C) Doryrhamphus melanopleura, a 52-mm-SL male, SIO 65-343. (D) Syngnathus auliscus, a 110-mm-SL female, SIO 65-181. 13-15 (usually 14); head 3.4—4.0 in SL _ CRIT TES a 5 Eee paulus n.sp. Islas Revillagigedo, Mexico Doryrhamphus melanopleura (Bleeker) (Figure 4C) Syngnathus melanopleura BLEEKER, 1858:464 (original de- scription; ‘‘Kokos-eilanden’’). Doryrhamphus californiensis GULL, 1863:284 (original descrip- tion; Cabo San Lucas; holotype SU 19255); JORDAN AND EVERMANN 1896:773 (description; range); DUNCKER 1915:62 (as “‘species dubia’); ULREY 1929: 6 (Cape San Lucas); JORDAN, EVERMANN, AND CLARK 1930:243 (range); ULREY 1932:77 (Cabo San Lucas); NICHOLS AND MuRPHY 1944:239 (Panama); FOWLER 1944:496 (range). Doryichthys californiensis: GUNTHER 1870:186 (new combi- nation; description; range). SE FRITZSCHE: REVISION OF EASTERN PACIFIC SYNGNATHIDAE 189 Microphis extensus SNYDER 1911:525 (original description; Naha, Okinawa; holotype USNM 98266). Doryrhamphus melanopleura: HERALD 1940:59 (in key; syn- onymy; range); 1953:246 (description; subspecies; range). DiaAGnosis.—A Doryrhamphus with 22-27 (24-25) dorsal rays; 19-23 pectoral rays; 16-18 (17) trunk rings; 14-17 (15) tail rings; dorsal cov- ering 3-5 + 2-4 rings; total rings covered by dorsal fin 6—-7.5; brood pouch covering 17-19 rings; head 4.0-4.4 in SL; dorsal fin base 1.6— 1.9 in head; snout 2.0-2.5 in head; maximum known size 69 mm SL; smallest mature male 31 mm SL. DESCRIPTION.—A sharply ridged and strongly spined fish. Head with prominent prenuchal, nu- chal, and occipital crests. Snout ridge strongly serrated, markedly so in mature males. Pectoral cover plate with well-developed superior and in- ferior ridges. Strong spination on maxillae and along dorsal edge of infraorbitals. Ventral edge of quadrate occasionally with strong serrations. Each ring bears a single posteriorly directed spine on each ridge, becoming obsolete along ventral ridges. Scutella large. No dermal flaps. Brood pouch abdominal, without protecting plates. Eggs about 0.5 mm in greatest diameter, arranged in three layers, four across. Males with eggs and young collected from March through August. Fins all large and well developed. Caudal a little longer than snout. Dorsal base about equal to snout and orbit combined. Anal conspicuous, its length about half orbital diameter. Pectoral fins broad based. Coloration in life. Essentially the same as col- oration in alcohol except that caudal has a white border, two yellowish spots near base, and an oval orange area in middle part of fin. Coloration in alcohol. Uniform dusky with darker streak from tip of snout to pectoral base. Fins colorless except for caudal, which is dis- tinctively marked with two light basal spots and one larger median spot. Individual fin rays often lined with melanophores on each side. Young less than 20 mm SL have a banded color pattern with eight brown bands on a lighter background. HABITAT.—In crevices in rocks, under over- hangs, and among coral heads. RANGE.—In eastern Pacific from Bahia Mag- dalena, Baja California, south to Isla la Plata, Ecuador (UMML), and at the Galapagos Islands and Clipperton Island. In the Gulf of California north to Isla Angel de la Guarda (UCLA W60- 6, not examined) in the west to Punta Guillermo, Sonora (UCLA WS1I-11 not examined), in the east. Widely distributed throughout Indo-Pacif- ic. COMPARISONS.—D. melanopleura is an inter- esting and complex species. It has been divided into several subspecies (Herald 1953) that are probably not valid. A detailed study of the pop- ulations is needed before much can be said about possible relationships. It is related to D. paulus from which it can be distinguished by the char- acters given in the description of that species. MATERIAL EXAMINED.—Mexico: Bahia Magdalena, SIO 64- 54, 3(55-62). Cabo San Lucas, SIO 61—227, 10(18-56). Los Frailes, SIO 61-249, 14(32-64). Bahia de los Muertos, SIO 74- 90, 3(32-46). Isla San José, SIO 65-265, 5(48-55). Isla Santa Cruz, SIO 65-343, 6(19-61). Isla Carmen, SIO 65-302, 15(45- 62). Isla San Ignacio de Farallon, SIO 59-228, 3(40—42). Lobos Rock, SIO 61-280, 20(16—-53). Bahia de Banderas, SIO 62-29, 2(29-45). Costa Rica: Isla del Cano, LACM 32548, 36(33-55). Panama: Taboguilla Island, SIO 67-34, 21(16—-48). Galapagos Islands: Plaza Island, SIO 64-1015, 3(53-63); BC 54-392, 1(64). Comparative material from Indo-West Pacific: Hawaii, CAS 20402, 1(54). Eniwetak, R. S. Nolan personal collection, 3(39- 45). Guam, CAS 15835, 2(44-44). Australia, AMS IA-2713, 1(38); AMS 110740, 2(31-39). Japan, CAS 14512, 2(68-69). Doryrhamphus paulus n.sp. (Figure 4B) Doryrhamphus melanopleura pleurotaenia: (in part) HERALD 1953:248 (size; meristic data). Doryrhamphus melanopleura: Ricker 1959:2 (Socorro Island). DIAGNosis.—A dwarf species of Doryrham- phus with 23-26 dorsal rays; 20-21 pectoral rays; 16-17 (16) trunk rings; 13-15 (14) tail rings; 29-31 (30) total rings; dorsal on 3—4.2 + 2.5-4 rings; total rings covered by dorsal fin 6.5—7.5; brood pouch on 14-17 rings; head 3.4—4.0 in SL; dorsal fin base 1.7—2.2 in head; snout 2.1—2.7 in head; smallest mature male 24.5 mm SL; maxi- mum known size 38.9 mm SL. DESCRIPTION.—Strongly ridged and spined. Occipital, nuchal, and prenuchal ridges low but sharply defined. Snout ridge serrate in males, with 4—5 spines on anterior tip and three or so larger isolated spines further posterior; females and juveniles with or without obvious snout- ridge serrations. Pectoral cover plate with su- perior and inferior ridges. Each body plate with a sharp retrorse spine. No dermal flaps. Only one brooding male known, collected in March. Fins well developed. Caudal shorter than 190 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 6 snout. Dorsal fin base equal to combined snout and orbit. Pectoral fins broad based but short, only extending posteriorly to junction of first and second trunk rings. Coloration. Essentially same as that given for eastern Pacific populations of D. melanopleura. HABITAT AND RANGE.—Three to 17 m, among rock reefs in Islas Revillagigedo, Baja California. COMPARISONS.—D. paulus is most closely re- lated to, and was probably derived from, D. melanopleura, which is widespread in the Indo- Pacific but is not represented at the Islas Revil- lagigedo. Although D. melanopleura ranges throughout the Indo-Pacific and is quite vari- able, the number of trunk rings is fairly stable at 18. Some populations have modal counts of 17 or 19 rings; however, 16 trunk rings have nev- er been found in D. melanopleura. Doryrhamphus paulus can be distinguished from D. melanopleura by the characters given in the key. D. excisus from the Red Sea also has 16 trunk rings, but differs from D. paulus in having 10- 11 tail rings rather than 13-15, and 18-20 dorsal rays rather than 23-26. ETYMOLOGY.—From the Latin paulus, little. MATERIAL EXAMINED.—Holotype: SIO 72-67, a 32.7-mm- SL mature male collected at a depth of 6-12 m with Chemfish, ca. 100 m se of ‘“‘Humpback Cove,’’ Isla Socorro, Islas Re- villagigedo, Mexico, 19 Feb. 1972, by D. Diener and party. Paratypes. Mexico. Islas Revillagigedo: SU 67255, 24.5 mm SL, mature male, 8 m, rotenone, “‘Grayson’s Cove,” Isla So- corro, 11 Mar. 1940, by Vernon Brock. SU 36442, 1(31), same data as SU 67255. CAS 13699, 9(20-34) and LACM 31781-2, 3(22-32), ‘“‘Grayson’s Cove,”’ ca. 200 m N of *‘Old Man of the Rocks,” Isla Socorro, R/V SEARCHER sta. 52, 8-12 m, rote- none, 14 Feb. 1971. LACM 317821-12, 5(26-32), s of Cape Henslow, Isla Socorro, R/V ;SEARCHER sta. 53, 13-17 m, ro- tenone, 15 Feb. 1971. GCRIe 15753, 1(27), Braithwaite Bay, Isla Socorro, 3 m, rotenone, 13 Feb. 1956, by George Lindsay. SIO 72-67, 6(16-26), collected with holotype. SIO 74-155, 2(29-39), Sulfur Bay, Isla Clarion, 10-17 m, rotenone, 11 Dec. 1974, by Robert Kiwala. Leptonotus Kaup / Leptonotus KAupP, 1853:232 (type-species by monotypy, Syngnathus blainvilleanus Eydoux and Gervais, 1837); 1856:46; DUNCKER 1912:235; 1915:88. Acmonotus PHILLIPI, 1896:382 (type-species by original des- ignation, Acmonotus chilensis Philippi [=S. blainvilleanus Eydoux and Gervais]). Novacampus WHITLEY, 1955:110 [type-species by original designation, Syngnathus norae (Waite)). DIAGNos!Is.—Urophori without pouch-pro- tecting plates; trunk compressed and much ex- panded in females; lateral ridge system of the ascending pattern (Fig. 2B); all elements of bran- chial skeleton present, none reduced; dorsal fin usually located on two or more trunk rings; 10 caudal rays; opercular ridge weak or absent; most body ridges smooth and reduced; two branchiostegals. DISTRIBUTION.—Approximately five species known only from south temperate seas; South America, South Australia, Tasmania, and New Zealand. Leptonotus blainvilleanus (Eydoux and Gervais) (Figure 4A) Syngnathus blainvilleanus EyYDOUX AND GERVAIS, 1837:79 (original description; ‘‘Mare Indicum’’; holotype MNHN 6050); GUICHENOT 1848:348 (description). Syngnathus acicularis JENYNS, 1842:147 (original description; Valaparaiso, Chile; holotype BMNH 1917.7.14.28). Leptonotus Blainvillei: KAUP 1853:232 (range); 1856:16 (de- scription; range). Syngnathus blainvillianus; GUNTHER 1870:162 (description; range); STEINDACHNER 1898:331 (Chile); THOMPSON 1816:423 (Patagonia). Hemithylacus Petersi DUMERIL, 1870:600 (original descrip- tion; Puerto Montt, Chile). Acmonotus chilensis PHILIPPI, 1896:382 (original description; Pelluhue, Chile). Leptonotus blainvillianus: ABBOTT 1899:338 (references; range); DUNCKER 1915:88 (synonymy; description; range); NORMAN 1937:40 (Patagonia); HERALD 1940:59 (synony- my; range); 1942:132 (diagnosed in key); FOWLER 1944:496 (range); MANN 1954:189 (description; range); DE BUEN 1963:89 (synonymy; description); KOEPCKE 1962:200 (range); CHIRICHIGNO-F. 1974:339 (in keys; range). Syngnathus blainvillei: DELFIN 1901:43. Siphostoma blainvilliana: EVERMANN AND RADCLIFFE 1917:53. Leptonotus blaenvillianus: SiccARD1 1954:211—242 (brood pouch; variation). Leptonotus blainvilleanus: DUMERIL 1870(2):581 (description; habitat); VAILLANT 1888:16 (Orange Bay, Patagonia; col- oration); HERALD 1965:364 (common name). DIAGNosiIs.—A Leptonotus with 34-41 (35- 37) dorsal rays; 12-14 pectoral rays; 2-3 anal rays; 18—20 (19) trunk rings; 48—52 (50) tail rings; 67-70 total rings; dorsal on 0.5—2 + 7-8.2 rings; total rings covered by dorsal fin 7.3—10; brood pouch on 10-14 rings; head 7.0-9.0 in SL; dorsal fin base 1.0-1.6 in head; snout 1.8—2.2 in head; maximum known size 217 mm SL (Duncker 1915); smallest mature male 108 mm SL. See Table 1 for meristic variation. DESCRIPTION.—Ridges on head and body mostly obsolete. Nuchal and prenuchal ridges low and smooth. Opercles strongly convex; opercular ridge reduced to basal one-fourth of FRITZSCHE: REVISION OF EASTERN PACIFIC SYNGNATHIDAE TABLE i. Trunk rings Tail rings Locality 18 19 20 48 49 50 Chile: Antofagasta 1 12 1 1 3 6 Valparaiso — — 2 — — ly Concepcion — 1 — — — — Coquimbo — 2 — — — 1 Puerto Montt 1 4 2 1 2 4 Isla Chiloé — 9 3 4 4 4 Totals 2 28 8 6 2) 16 x = 1951 x = 49.6 Argentina*: Golfo Nuevo (females) 27 4 — — 4 16 (males) R) 34 3 5 12 20 Totals 30 38 3 5 16 36 x = 18.6 xX = 49.8 ““Mare Indicum’’** — 1** — — — — + Holotype of S. acicularis. * Data from Siccardi 1954. ** Holotype of L. blainvilleanus. opercle, striations faint. Snout ridge smooth, low, and reaches posteriorly to interorbit. Su- praorbital ridges smooth, one-half orbit diameter in length. Pectoral cover plate without ridges. Trunk of mature females compressed and ex- panded dorsoventrally, with sharp dorsal and ventral borders. Scutella small, indistinct. Plates with reduced ridgelets. Entire body very fleshy over plates. Lateral line papillae 2—4 per plate. No dermal flaps. Brood pouch without lateral protecting plates. Males brooding eggs have been collected in Jan- uary and March. Dorsal fin high, its height equal to width of two trunk rings. Coloration in alcohol. Juveniles with alternat- ing pattern of dark and light bands along length of body. Darker bands about four rings in width, light bands only one-half a ring in width. Caudal dark brown with a light border. Snout and in- terorbit darker than remainder of head. Fins col- orless. Adult females sometimes entirely light tan with dark brown venter and dorsum on trunk, and with posterior half of tail dark brown. Usu- ally the head is a dark olive-brown and the trunk is dark brown with small, dark-bordered ocelli. 191] MERISTIC VARIATION IN Leptonotus blainvilleanus. Dorsal rays 52 34 35 36 37 38 39 40 41 — 2 6 1 4 1 — — — ae = al = ie Le a4 aS “ie 2=. te = 1 ate = _ = = 1 =s _ = = —_ = = = = — 1 3 3 — _ — — — 2 3 4 2 — — — — 3 8 5 13 6 = —_ == 50.5 1 — 1 1 8 14 5 1 1 2 — — 7 di 13 8 1 1 3 — 1 8 15 27 13 2 2 x = 37.8 Dorsal fin may have a few melanophores along the ray margins. Adult males yellowish brown with dark brown area surrounding the nares; ocelli may develop on the first few tail rings. HaBITAT.—Kelp beds along the open coast; occasionally into brackish water (Fischer 1963). REMARKS.—Meristic data from the Golfo Nuevo, Argentina, population of L. blainvil- leanus (Table 1) (from Siccardi 1954) show a difference in mean number of trunk rings be- tween males and females from Golfo Nuevo (P < .005). The lower number of trunk rings in the Golfo Nuevo females contributes to the significant dif- ference between the Chilean and Argentinean populations (P < .005). However, the dorsal ray count also differs significantly between these two regions. Siccardi (1954) studied a large number of specimens of L. blainvilleanus from both coasts of southern South America and was able to pro- vide some data on morphometric variation. She found that the depth of the trunk in males re- mains fairly constant during growth, from 112- 140 mm, so that the standard length/trunk depth ratio was 22.4 in 112-mm fish and 26.7 in 140- mm fish. Also, the maximum depth of the trunk 192 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 6 in females was attained by a standard length of 140-150 mm. This was correlated with reaching sexual maturity. The trunk in males became rel- atively shorter and the tail longer with growth, whereas relative lengths of the trunk and tail in females remained constant with growth. COMPARISONS.—Because the genus Lepton- otus has not received systematic treatment, it can only be said that L. blainvilleanus seems to be most closely related to the southern Austra- lian species L. semistriatus. L. blainvilleanus can be distinguished from L. semistriatus by the snout length (1.8—2.2 in head rather than 1.6— 1.7) and by the dorsal fin placement (on 0.5—2 + 7-8.2 rings rather than 3—4 + 7). Leptonotus blainvilleanus has long been con- sidered to be a member of the New Zealand fau- na (Waite 1909). However, a comparison of the holotype of L. blainvilleanus with specimens of Leptonotus from New Zealand reveals that L. blainvilleanus and examined New Zealand species of Leptonotus are distinct. The New Zealand specimens of Leptonotus are referable to L. elevatus (Hutton) and L. norae (Waite). Mr. A. Wheeler (BMNH) examined the ho- lotype of Syngnathus acicularis Jenyns at my request. His counts and description of lateral ridge pattern confirm the fact that the holotype of S. acicularis is conspecific with L. blainvil- leanus. RANGE.—Hornitos, Chile, to Golfo Nuevo, Argentina. MATERIAL EXAMINED.—‘‘Mare Indicum,’> MNHN 6050, holotype. Chile: Hornitos, SIO 72-168, 1(168). Antofagasta, EBMCh 1520-28, 9(86-207); GCRL 12466, 4(117-215). Coquimbo, MNMHP 5302, 1(169); MNMHP 5305, 1(246); Valparaiso, EBMCh 222, 1(190). Concepci6n, MNMHP 5574, 1(126). Ba- hia Lin, USNM 176564, 4(75-121). Puerto Montt, USNM 205179, 3(104-129). Isla de Chiloé, USNM 176569, 4(108— 142); MNMHP 5304, 3(84-173); MNMHP 5303, 5(141-162). No collection data, CAS (Indiana label), 1(184). Argentina: Patagonia, CAS 36440, 1(130). Bryx Herald Bryx HERALD, 1940:52 (type-species by original designation Bryx veleronis Herald); 1959:468 (subgenus of Syngnathus). Microsyngnathus HERALD, 1959:468 (subgenus of Syngna- thus, type-species by original designation Syngnathus dunckeri Metzelaar). D1AGNosis.—Urophori characterized by a very short snout; small size (generally less than 100 mm); first epibranchial reduced or not; sec- ond hypobranchials and epibranchials reduced; frontals not reaching anteriorly past middle of lateral ethmoids; ossified epaxialis tendons pres- ent or absent; one infraorbital; two nuchal plates; pouch protecting plates present; dermal flaps present or absent; 14-17 trunk rings; 27—40 tail rings; 19-31 dorsal rays; 10—14 pectoral rays; anal fin present or absent; 10 caudal rays; 10-19 rings covered by brood pouch; 0-2 + 3-7 rings covered by dorsal fin. DISTRIBUTION.—Ten tropical species, and one species at Islas Juan Fernandez and Isla San Félix. DiscussIon.—Bryx is closely related to Cos- mocampus, and the two were probably derived from a common ancestor. Bryx was erected by Herald in 1940 for the sole reception of B. veleronis. Later, Herald (1959) transferred B. veleronis to Syngnathus and retained Bryx as a subgenus. In the same paper Microsyngnathus was erected as a sub- genus, with Syngnathus dunckeri as type-spe- cies, and S. arctus and §. coccineus included therein. §S. hildebrandi Herald, 1965, S. ran- dalli Herald, 1965, and S. banneri Herald and Randall, 1972, were described as being related to, or tentatively referred to, the subgenus Mi- crosyngnathus species in Herald (1965) and Her- ald and Randall, 1972. The species of Bryx and Microsyngnathus (and Micrognathus balli Fow- ler, 1925, and Syngnathus darrosanus Dawson and Randall, 1975) are separated from other syngnathids and united by shared characters given in the diagnosis above. This combination of characters defines a related lineage of syng- nathids worthy of generic recognition. The type- species of Microsyngnathus further agrees with Bryx veleronis in lacking an anal fin: Micro- syngnathus Herald, 1959, thus becomes a junior synonym of Bryx Herald, 1940. The subgenus Simocampus is proposed for those species of Bryx possessing an anal fin. Key to Eastern Pacific Species of Bryx la. Anal fin present (subgenus Simocampus) Ib. Anal fin absent (subgenus Bryx) 2a. Head shorter than length of dorsal fin baS@r 2. heraldi n.sp. Islas Juan Fernandez and Isla San Felix 2b. Head longer than length of dorsal fin base 2.2... eee ee 3 3a. Dorsal fin usually entirely on tail rings, FRITZSCHE: REVISION OF EASTERN PACIFIC SYNGNATHIDAE 193 TABLE 2. FREQUENCY DISTRIBUTIONS OF TRUNK AND TAIL RINGS, AND DORSAL FIN RAYS IN EASTERN PACIFIC SPECIES OF Bryx. Trunk rings Tail rings Dorsal fin rays [4a SOS a5 4S SOS AOS B40 4) oats 19e 20) 21225 2324595; 26 27) 28 heraldi — 18* — — — — 2 10 6 — — — — ~—~ —~ ~— 10* 8 ~—~ ~ ~ ~ — arctus 1 31* 20 — — — 3 5) HQ Sikes Te) i EO Ie) 2 —- — — — = coccineus ee lilies 1— — — 1 SNe GP? | eee veleronis eG * nny De tam ATOR eS —= — | IF i} 3h ek 2 clarionensis — 9* — — — 3 6 — — — — — — — =—- = — ~~ — 1 =2 «4F 2 * Primary type. except in some southern California spec- imens with the dorsal on a fraction of a trunk ring; 18—23 dorsal rays; double row Oijaatkespotsvon tink, 2.2. 222.3 ces arctus (Jenkins and Evermann) Tomales Bay, California, and San Felipe, Gulf of California, to Mazatlan, Mexico 3b. Dorsal fin always on at least a fraction of last trunk ring; 18—20 dorsal rays; no dou- ble row of dark spots on trunk. coccineus (Herald) Banderas Bay, Mexico, to Punta Aguja, Peru, and Gala- pagos Islands 4a. Snout longer than postorbital length _____ clarionensis n.sp. Clarion Island, Revillagigedo Islands, Mexico 4b. Snout shorter than postorbital length ___ veleronis Herald Galapagos and Revillagigedo Islands, and coasts of Panama and Costa Rica Simocampus n.subgen. TYPE-SPECIES.—Siphostoma arctum Jenkins and Evermann. DIAGNosis.—A Bryx with anal fin. TABLE 3. INCLUDED SPEcIES.—B. arctus (Jenkins and Evermann):; B. coccineus (Herald); B. balli (Fowler); B. banneri (Herald and Randall); B. hildebrandi (Herald); and B. darrosanus (Daw- son and Randall). ETYMOLOGY.—From the Greek simos, pug- nosed, and campos, sea-creature. Bryx (Simocampus) heraldi n.sp. (Figure SC) DiaGNosis.—A Bryx with 22-23 (22) dorsal rays; 11 pectoral rays; 3 anal rays; 15 trunk rings; 36—38 (37) tail rings; 52—54 total rings; dor- sal on 0-1 + 5.2-6 rings; 17-18 rings covered by brood pouch; head 10.3—12.3 in SL; dorsal fin base 0.8-1.0 in head; snout 2.4—2.8 in head; maximum known size 99.8 mm SL; smallest mature male 70 mm SL. See Tables 2 and 3 for additional counts and measurements. DESCRIPTION.—AII ridges of head and body moderately developed and smooth. Prenuchal and nuchal ridges elevated and easily discern- ible. Opercular ridge extends posteriorly about half length of opercle. Snout ridge moderately developed, extending posteriorly to interorbit. Ridges of pectoral cover plate obsolete. Trunk and tail ridges low but easily visible. Superior NUMBER OF SPECIMENS (N) AND STANDARD LENGTH RANGE, TOGETHER WITH RANGE AND MEAN (x) OF SE- LECTED CHARACTERS EXPRESSED IN THOUSANDTHS OF STANDARD LENGTH IN Bryx. Head Snout Snout Body Dorsal base Pectoral length length depth depth length fin length Species SL Range mG Range OF oF ff snosinp VC Cum Cam 0G 6l 81 yt Syl Sil 1A sBuL [Ie “snyIDUSUAG AO SAIOAdS DIAIOVd NUALSVY NI SAVY NI TvSuod ANV ‘SONIY ssull yUNIy, JIV], GNV ANAY] JO SNOILLNAIISIG AONANOAN “pf ATAV FRITZSCHE: REVISION OF EASTERN PACIFIC SYNGNATHIDAE 201 TABLE 5S. NUMBER OF SPECIMENS (N) AND STANDARD LENGTH RANGE, TOGETHER WITH RANGE AND MEAN (%) OF SE- LECTED CHARACTERS EXPRESSED IN THOUSANDTHS OF STANDARD LENGTH IN Syngnathus. Head Snout length length Species SL Range x Range ¢ auliscus 46-134 98-136 113 44-56 48 carinatus 116-230 113-141 128 65-89 76 exilis 75-238 102-148 121 50-88 68 californiensis 78-382 111-143 120 55-93 69 macrobrachium 92-190 112-144 137 60-82 72 euchrous euchrous 111-249 100-132 114 47-78 55 e. ollotropis 116-244 93-108 103 44-S7 49 leptorhynchus 57-245 104-148 122 48-87 65 insulae 88-180 107-117 111 46-57 52 1928:42 (synonymy; range); ULREY 1929:6 (Lower Califor- nia); JORDAN, EVERMANN, AND CLARK 1930:240 (range); ULREY 1932:77 (Golfo de California); BARNHART 1936:34 (description; range); HERALD 1940:60, 63 (key; synonymy; range); CASTRO-AGUIRRE ET AL. 1970:131 (not common in Gulf of California); MILLER AND LEA 1972:89 (description; range); CHIRICHIGNO-F. 1974:351 (range). Singnathus auliscus; VAILLANT 1894:70 (Gulf of California). Siphostoma sinaloae JORDAN AND STARKS 1896:268 (original description; Mazatlan; holotype SU 2945). Syngnathus tweedlei MEEK AND HILDEBRAND, 1923:259 (orig- inal description; Chame Point, Panama; holotype USNM 82088). DiAGNosis.—A Syngnathus with 26-33 (27- 30) dorsal rays; 10-14 pectoral rays; 2-3 anal rays; 14-16 (15) trunk rings; 34-39 (36-37) tail rings; 0.2—1.5 + 5—7.2 rings covered by dorsal fin; total rings covered by dorsal fin 6—7.9; 15— 21 pouch rings; head 7.1—10.1 in SL; dorsal fin base 0.8-1.6 in head; snout 2.0-2.8 in head; maximum known size 178 mm SL; smallest sex- ually mature male 71 mm SL. See Tables 4 and 5 for additional counts and measurements. DESCRIPTION.—AII ridges of body smooth, rounded, low but distinct. Prenuchal and nuchal ridges low but distinct. Opercles strongly con- vex, with median ridge reaching half the length. Snout ridge running full length of snout to inter- orbital. Head not raised at orbit, so that snout ridge almost horizontal. Supraorbital ridge orig- inating just anterior to nostrils and ending above anterior edge of opercle. Pectoral cover plate with one obsolete ridge. Lateral trunk and su- perior tail ridges overlap for one ring. Slightly acute ventral trunk ridge. Three lateral line pa- pillae per plate. Dermal flaps rarely present. Plates protecting brood pouch well developed, often as deep as overlying tail rings. Eggs 0.6— Snout Body Dorsal-base Pectoral-fin depth depth length length Range ~—> — 2 1 6 4 7 3 1 1 1 — Puget Sound —- —- —> ~—> > — 2 3 8 7 4 2 2 — — — Coos Bay —- -—- -—|- -—|- -—|- —- —- = 5 4 16 14 10 8 2 — Humboldt Bay —- —- FS Fe eee a 7 9 12 17 14 12 8 2 Tomales Bay —- -—- —- —- —- —- = 1 4 9 11 3 4 1 2 — San Francisco Bay —- -—- —- —- —>— — 1 9 31 55 61* 60) 25 8 1 — Morro Bay - -—- —> — 3 5 10 10 6 5) a —- —- — — — Pt. Mugu Lagoon —- —- — — 2 2 72 5 2 1 — —- -—- =—- => = Santa Barbara —- —- —|- —|- —|- —- —>— — ———— 2 SS eS Newport Bay —- =—- — => 3 5 6 6 7 3 1 —- -—- —- —> — Mission Bay 2 12 On Oe Same) 37 18 6 1 — —- -—- —- —- — San Diego Bay —- — 5 3 8 6 2 [-e- == — = —- —- FeO Oe Estero de Punta Banda 2 22 7 5 30 — SO — — — —- — > > Bahia San Quintin —- — — 4 14 7 8 3 2 — 1 —- —- —- —>— — Scammon Lagoon — 1 1 — 1 — 3 1 — = = aa eS EES Total 4 Is 38 Si) 87 7 Ws 58 85 98 113 99. S56 30 = 14 Z ti Seo 323 * Holotype of S. barbarae. + Holotype of S. abbotti. ** Syntypes of S. brevirostris. Coloration in life. Mostly browns, purples, and greens, with varying amounts of mottling, spots, etc. Coloration in alcohol. Coloration highly vari- able, mainly shades of brown. Great inter- and intra-population variability. Highly mottled, blotched, and striped. Dorsal fin always with some brownish spots at base and fin rays bor- dered with melanophores. HaBITAT.—Ordinarily inhabiting eelgrass in bays and occasionally near shore outside. VARIATION AND GROWTH.—Variation: Syng- nathus leptorhynchus shows extreme variation in meristic characters (see Table 7) approaching the North Atlantic species S. typhle in this re- gard. Duncker (1908) statistically analyzed sev- eral meristic characters of S. typhle populations from the Baltic Sea, Plymouth, and Naples and FRITZSCHE: REVISION OF EASTERN PACIFIC SYNGNATHIDAE showed a definite clinal pattern in the variation of numbers of trunk rings and pectoral rays. Other meristic features showed populational dif- ferences, but in a mosaic pattern of different se- quences. Meristic variation in S. leptorhynchus is mo- saic, and no single character allows or justifies recognition of more than one species or subspe- cies. Herald (1940, 1941) and others have used the same meristic characters herein analyzed in support of recognizing northern and southern species or subspecies. A mosaic pattern of vari- ation is not limited to Syngnathus species, e.g., Rosenblatt (1963) reported a similar pattern for populations of the Central American tripterygiid Enneanectes sexmaculatus. Each bay along the west coast of North Amer- ica appears, on the basis of available evidence, to support a distinctive population of S. lepto- rhynchus. Meristic features of each population are not stable, but do fluctuate with changes in environmental temperature. To underscore this point, C. L. Hubbs ana- lyzed three collections from the northwest cor- ner of Mission Bay on 22-24 April 1923. All three collections were made with exactly the same method (seine) and at the same location in the bay. For analysis, all specimens 97 mm SL or less are considered to be winter reared, and in view of the demonstrated growth (see below) it may be presumed that all specimens longer than 97 mm SL were at least mostly reared in warmer water. The summary of meristic data (Table 8) compiled for these collections indi- cates that the winter-reared young have higher means in all meristic characters studied than do the larger (adult) fish. The means for these two size groups are all significantly different (P < .00S). Although Mission Bay has been highly modi- fied in recent years, the data are indicative of changes in meristic characters as a result of en- vironmental temperature. Similar seasonal me- ristic variation has been documented by Broth- ers (1975) for the goby Quietula y-cauda—using samples taken throughout the year, he deter- mined that the mean vertebral number of the postlarvae is inversely correlated with seasonal changes in environmental temperature. Each local population of Syngnathus lepto- rhynchus is therefore distinctive and also highly modified by local environmental conditions. S. SEASONAL VARIATION IN MERISTIC CHARACTERS OF Syngnathus leptorhynchus. TABLE 8. Tail rings Trunk rings SD 37 38 39 40 36 SD 18 19 17 34 10 0.51 17.8 Young Adults 1.02 38.4 110 15 0.57 7/5) 111 53 54 Dorsal rays Total rings SD SD 54 53 7.80 1.55 1.47 c 31.6 111 aN 10.75 1.04 129 Wil7 SA) OM S59 41 39 18 4 13 al Young Adults 3} 214 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 6 ——1 FiGur_E 13. leptorhynchus may be a useful indicator species for the effects of pollution. Growth: On 16 June 1974, a male S. lepto- rhynchus (165 mm SL) gave birth to 85 young in the laboratory of the National Marine Fish- eries Service, La Jolla. The young were imme- diately separated from their father and kept alive until a failure in the seawater system on 6 Sep. The young were fed Artemia nauplii and fresh- caught plankton. An overabundance of food was always present. Samples of the young were re- moved frequently during the period from 16 June to 6 Sep. and preserved in 10% formalin for later analysis. Figure 14 represents the measured growth of the young for the period. The data have been analyzed and fitted to a Laird-Gompertz equa- tion (Laird, Barton, and Tyler 1968) calculated after the methods of Zweifel and Lasker (1976). The Laird-Gompertz equation is preferred when analyzing growth for periods less than one year. On 16 Aug. development of the brood pouch had already begun in a 76-mm-SL individual. Four males (80-93 mm SL), each with a fully developed brood pouch, were found on 24 Aug. By the termination of the experiment on 6 Sep., the largest specimen was a fully-mature, 114-mm male. That the ranges of meristic variation for the adults do not completely subtend the ranges for the young together with rapid growth to sexual maturity strongly indicate that individuals of S. leptorhynchus (at least from Mission Bay) prob- ably live for only one year. This is in agreement with the data presented by Brown (1972) for S. louisianae, S. scovelli, S. floridae, and Micro- gnathus crinigerus for Cedar Key, Florida. Based on length-frequency analyses, Brown estimated a life span of perhaps one year. COMPARISONS.—A highly variable species that is similar to the Japanese S. schlegeli. In fact, it is rather difficult to separate members of northern populations of /eptorhynchus from Syngnathus leptorhynchus at time of emergence from brood pouch. 16.3 mm SL. those of schlegeli. However, S. schlegeli has a longer snout and longer fins, and the ridges of the body are better developed. S. insulae, also closely related to S. leptorhynchus, may be dis- tinguished by the characters given in the account of that species. RANGE.—Southeastern Alaska south to Bahia Santa Maria, southern Baja California. MATERIAL EXAMINED.—Alaska: UW 14486, 1610, 1(212). British Columbia: Vancouver, BC 53-253a, 28(60—206); SIO 62-206, 2(137-192). Washington: Puget Sound, CAS 10935, 3(125—140); UW 4927, 3(136-151); UW 5296, 2(146-245); UW 4258, 3(86-130); UW 4918, 4(139-167); SIO 74-73, 19(73-215). Oregon: Tillamook, UMMZ 93951, 33(49-235). Coos Bay, UMMZ 93956, 63(35—235). California: Klamath Lagoon, CAS 45070, 1(216). Humboldt Bay, HSU OT62-7, 44(136-207); HSU OT61-18, 14(121-218); HSU OT64-5, 12(110-210). Mendocino County, SU 20515, 5(165-214). Tomales Bay, SIO 75-459, 37(54—234); UCLA W55-198, 35(61-207); CAS 19860, 2(211-293); CAS 26207, 2(104-252). Drakes Bay, CAS 7518, 1(130). San Pablo Bay, CAS 45072, 33(59-272). San Francisco Bay, SU 4190, 24(115— 11179); UW 159); USNM 971, 1(228), holotype of S. arundinaceus; UMMZ 64416, 1(215); UMMZ 6044, 1(151); CAS ALBATROSS Biological Survey of San Francisco Bay*, D5826, 1(131); H5159, 2(139-165); DS5816, 1(129); H5311, 1(119); H5013, 4(111-184); D5847, 1(92); H5003, 1(87—121); H5318, 2(110- 128); H5185, 1(61); DS848, 2(101-118); H5312, 2(66—-94); H5004, 2(77-110); H5308, 1(67); H5230, 1(62); H5248, 1(64):; HS259, 1(61); H5310, 1(80); H5316, 2(69—98); H5309, 2(106— 120); H5250, 7(51-123); H5009, 9(33—109); H5257, 1(86); H5261, 2(41—43); H5345, 13(25—133); H5313, 6(69-93): H5251, 6(73—-125); H5010, 2(33-66); H5005, 3(44-69); H5275, 3(38- 70); H5315, 5(63—113); H5252, 5(62—95); H5317, 4(76—-103): H5311, 8(75—90); H5274, 8(67-77); H5007, 11(28—108); H5314, 13(67—105); H5272, 17(64-146); H5276, 25(49-130); H5006, 76(27-116). Elkhorn Slough, SIO 62-510, 3(77-103). Del Monte, SIO 48-372, 2(92-196). Morro Bay, SIO H50-191, 1(243); UCLA W50-196, 50(98-236); SU 48888, 10(162-217). Point Mugu, CAS 45076, from E. S. Herald personal collec- tion, 4(151-181). Santa Barbara, USNM 31253, type of S. barbarae. San Pedro, SIO H48-180, 1(257). Newport Bay, UCLA WS0-171, 50(99-278); UCLA WS50-172, 3(101-132):; SIO H46-33, 5(215—280). San Juan Capistrano, SIO H52-109, 1(101). Mission Bay, SIO H45-8, 50(72-198); SIO H46-10, * Station data may be found in Anonymous 1921. FRITZSCHE: REVISION OF EASTERN PACIFIC SYNGNATHIDAE 215 120 fe) 10 20 30 DAYS AFTER BIRTH 40 50 60 70 80 Figure 14. Growth of Syngnathus leptorhynchus. Standard length plotted against time in days. The Laird-Gompertz Equa- tion fitted to these data is included. 3(116-125). San Diego Bay, USNM 969, 3, types of S. bre- virostris; SIO 64-337, 25(100-158). Baja California: Estero de Punta Banda, SIO H45-15, 19(104—158). Bahia San Quintin, SIO 60-453, 128(69-245): SIO 60-450, 6: SU 15251, 10(123-299). Laguna Scammon, SIO H52-140, 1(95). Bahia Santa Maria, SIO H50-149; SIO 60-294, 1(94). Syngnathus insulae n.sp. (Figure 12D) DiaGNosis.—A Syngnathus with 31-34 (32- 33) dorsal rays; 10-12 pectoral rays; 2-4 anal rays; 17-19 (18) trunk rings; 40-42 (41-42) tail rings; 58—60 total rings; dorsal on 0.2-1.2 + 6.5- 8 rings; 19 rings covered by brood pouch; head 8.6—9.3 in SL; dorsal fin base 1.0 in head; snout 1.7-2.3 in head; maximum known size 204 mm SL; smallest sexually mature male 125 mm SL. See Tables 4 and 5 for additional measurements. DESCRIPTION.—AIl ridges of head and body poorly developed, smooth. Prenuchal and nu- chal crests obsolete. Opercular ridge reduced to short basal portion. Ridges of pectoral cover plate low. Trunk and tail ridges low, rounded but distinguishable. Superior tail ridge not reaching beyond first tail ring, only overlapping lateral trunk ridge for one-half ring. Scutella quite distinct and about half as deep as adjoining plate. Lateral line consisting of S—6 papillae per plate. No dermal flaps. Protecting plates of brood pouch well devel- oped, each about as deep as adjacent tail ring; edges of brood pouch folds entire. Eggs 1.2—1.3 mm in greatest diameter, arranged in one layer two across. Males with eggs and young have been collected in April. Fins normally developed. Caudal fin as long as last three tail rings. Pectoral fin reaches pos- teriorly to end of first trunk ring. Young. About 12 mm SL at birth. About five dark bands posterior to vent. Body sprinkled with small round melanophores, especially in the area of the bands. Snout short, mouth dis- tinctly upturned. Coloration in life. From C. L. Hubbs’s field notes for SIO H46-143: The greener specimen has a dark streak running downward and backward from eye. In this specimen the streak ex- tending upward and backward from eye is less distinct. The streak running forward from eye to chin is conspicuous pos- teriorly but interrupted anteriorly. The redder specimen has a dark spot just behind and below eye, but no definite streaks radiating from eye. Coloration in alcohol. A light tan to tan fish without any distinguishing pattern. The caudal fin may be lighter at its border. Darker speci- mens may have a sprinkling of small melano- phores bordering rays. HaBITAT.—Occasionally in floating Sargas- 216 sum, but much more often taken at depths of 20-34 m in beds of Sargassum and algae. COMPARISONS.—Appears most closely related to S. leptorhynchus. The two are very close in meristic characters. §. leptorhynchus has the longer snout and postorbital. S. insulae differs from leptorhynchus in the broader interorbital, smaller orbit, and less robust body. RANGE.—Endemic to Isla Guadalupe, an oceanic island well off the coast of Baja Cali- fornia. ETYMOLOGY.—From the Latin insulae, gen- itive of insula (island). MATERIAL EXAMINED.—Holotype: SIO 63-169, a 160-mm- SL mature male collected at Caleta Malpomene, a cove at the south end of Isla Guadalupe, Mexico, from a depth of 22-50 m using a try-net on 22 Apr. 1963, by C. L. Hubbs and party aboard the R/V Horizon. Paratypes: All taken from Isla Guadalupe. SIO 63-169, 32(77-204), same data as holotype. SIO H46-143, 2(98—98.4), Caleta Melpomene, under nightlight, 6 Dec. 1946, by C. L. Hubbs aboard R/V E. W. Scripps. SIO H46-144, 1(118), Cal- eta Melpomene, 3-6 m, dredge, 7 Dec. 1946, by C. L. Hubbs aboard R/V E. W. Scripps. GCRL 15756, 2(93-117), Caleta Melpomene, under nightlight, 27 Jan. 1950, by C. L. Hubbs and party aboard R/V Orca. SIO HS0-31, 1(99), Caleta Mel- pomene, 3-7 m, dredge, 29 Jan. 1950, by C. L. Hubbs and party aboard R/V Orca. SIO 54-214, 2(51—87), “% mile [200 m] “off Barracks’’ (Northeast Anchorage), under nightlight, 13 Nov. 1964, by C. L. Hubbs and party. SIO 63-188, 1(116), just inside s side Punta Amarillo on rocky bottom, 15 m, Chemfish and scuBA, 26 Apr. 1963, by J. R. Stewart and party. FossIL SYNGNATHIDAE OF SOUTHERN CALIFORNIA The following discussion of the fossil pipe- fishes of southern California is based on a large collection in the Section of Vertebrate Paleon- tology at the Natural History Museum of Los Angeles County (LACM), including the collec- tions formerly housed at the California Institute of Technology (CIT). The fossil syngnathids from southern Califor- nia are all of Miocene to possibly Pliocene age. Most specimens have been collected in diato- mites and diatomaceous siltstones of the upper Modelo and Puente formations, although a num- ber have been collected from argillaceous diat- omites of the lower Modelo Formation. Locali- ties are given by number, and more precise data can be obtained by referring to the Locality Catalogue in the Vertebrate Paleontology Sec- tion (LACM). These formations have provided the bulk of material used in the descriptions of California PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 6 fossil fish (Jordan and Gilbert 1919; David 1943). Since most of the fishes contained in these de- posits are pelagic to bathypelagic, previous au- thors have inferred that during the Miocene this area was a large basin with depths ranging from 200-500 m (David 1943) to 1000 m (Crane 1966). Materials and Methods The anal and pectoral fins of fossil pipefishes are usually not visible. Since the anal fin marks the first tail ring, it is impossible to make precise counts of trunk and tail rings. However, the number of plates in a ring around the trunk is usually seven and there are four around the tail. When these plates could be seen, accurate counts were made. When such counts were not determinable, the trunk rings were counted as ending at the first ring under the dorsal fin or the posterior end of the median ventral ridge (in Hipposyngnathus). The count of dorsal fin rays in fossil pipefishes is difficult because of frequent splitting of the two ray-halves. However, with proper precau- tion, a fairly accurate count is often possible. Accurate counts of the rings and dorsal rays are of particular importance in syngnathid tax- onomy, as these counts provide the primary ba- sis for classification of living forms. Correct generic allotment of fossil syngnathid species is based primarily on the author’s knowledge of the general appearance of syng- nathid genera. The characters often used for placement of syngnathids into a genus (i.e., lat- eral ridge pattern, placement and position of brood pouch) are very rarely preserved in fos- silized pipefishes. This section on fossil pipefishes is intended only to describe those forms known to occur in deposits along the eastern Pacific and is not meant to be a critical review or stratigraphic analysis of them. Subfamily Hipposyngnathinae nom. nov. Eogastrophinae JERZMANSKA 1968:436 (“‘genre typique: Hip- posyngnathus Danil’chenko 1960°’). Since the subfamily name was incorrectly pro- posed as Eogastrophinae (the sole-included ge- nus is Hipposyngnathus), and since subfamilial ranking is obviously called for, I propose the name Hipposyngnathinae, with Hipposyngna- thus as the type-genus. Jerzmanka diagnosed the subfamily adequate- FRITZSCHE: REVISION OF EASTERN PACIFIC SYNGNATHIDAE 217 Ficure 15. Holotypes of three species of California fossil syngnathids. (A) Hipposyngnathus imporcitor, LACM 12055. (B) Syngnathus avus, CAS Geology Dept. No. 58372. (C) Syngnathus emeritus, LACM 27445. iy except that she incorrectly stated that scutella are absent. Hipposyngnathus Danil’chenko Hipposyngnathus DANIL’CHENKO, 1960:87 (type-species by original designation H. convexus Danil’chenko); JERZMAN- SKA 1968:436. DIAGNosIs.—Lateral protecting plates of _ brood pouch absent, however, a very pro- nounced median ventral trunk ridge probably in- dicates position of brood area. Dorsal, pectoral, and caudal fins present. Dorsal mainly on the tail. Ridges of body generally moderately de- veloped and rugose, but without spines or ser- rations. Three described species from the Oli- gocene and Miocene. DISTRIBUTION.—Oligocene of the Caucasus and Carpathian Mountains of southeastern Eu- rope. Miocene of southern California. Hipposyngnathus imporcitor n.sp. (Figure 15A) Syngnathus avus (in part, not of Jordan and Gilbert, 1919): Davip 1943:70 (description). Holotype: LACM 12055, an almost-complete specimen from the upper Modelo Formation of the northeast slope of Santa Monica Mountains, Los Angeles County, California (Locality LACM (CIT) 326). REFERRED MATERIAL (all from California)—Upper Modelo Formation, NE slope Santa Monica Mountains, Los Angeles County, 24 identifiable specimens, 33 fragments, LACM loc. no. (CIT)326: LACM cat. nos. 10060-10061, 10063, 10114, 10134, 10143, 10145, 12054, 25341-25351; and LACM loc. no. 1267: LACM cat. nos. 13362-13373, 13451-13465, 13469- 13473, 13475, 13477-13478, 25789-25791. Near Lompoc, LACM loc. no. 6589, 10 fragmentary specimens: LACM cat. nos. 12160-12169. Near Emery Park, Alhambra, Los Angeles County, LACM loc. nos. 1031, 1033, 14 specimens: LACM cat. nos. 1056-1058, 1349, 12459-12464, 25668. Dixie Canyon, Santa Barbara County, LACM loc. no. (CIT)329, 6 speci- mens: LACM cat. nos. 25357-25359, 25363-25365. D1aGNosis.—A Hipposyngnathus from the upper Modelo Formation of southern California with about 22 dorsal rays; 20-22 trunk rings; 42- 44 tail rings; snout 1.5-1.9 in head; orbit 7.8— 12.0 in head; pouch rings equal to trunk rings; pectoral fin present but rays not countable; pouch ridge as long as adjacent trunk. Maximum size known 242 mm SL. 218 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 6 DESCRIPTION.—Body ridges moderately de- veloped and rugose. Plates rhomboid-ovate with 10-13 ridgelets on either side of median ridge, large; each trunk plate about as wide as orbit. Opercular ridge smooth, about half as long as opercle. Snout without distinct ridge. Scutella small. Two infraorbitals. Vertebrae as in Syng- nathus. Dorsal high, about 1.5 times higher than adjacent tail. Caudal normally developed. DiscUSsION.—H. imporcitor is the most com- mon syngnathid in the upper Modelo Formation. The extinct Hipposyngnathus was previously known from two European Oligocene species. Hipposyngnathus imporcitor can be easily dis- tinguished from its congeners H. neriticus and H. convexus by the characters given in the di- agnosis. The genus Hipposyngnathus is unusual in having a very exaggerated ventral trunk ridge. Jerzmanska (1968) suggested that the males alone had this ridge and that it was the site of egg deposition. She also speculated that pouch flaps were present on either side of the ridge. However, the extant genus Maroubra also has an enlarged ventral trunk ridge, although not so well developed as in Hipposyngnathus, and its eggs are not protected by flaps. It seems im- probable that Hipposyngnathus had brood- pouch flaps. The Gastrophori are typically spiny with the exception of Nerophis and Entelurus. Hippo- syngnathus is most like Nerophis in general body form. PALEOECOLOGY.—Crane (1966) has discussed the paleoecology of Chauliodus eximius, a chau- liodontid from the diatomite of the upper Mo- delo. Most of his specimens were from the same localities as those where Hipposyngnathus was collected. The upper Modelo is approximately 15 million years old (Crane 1966). David (1943) discussed the paleoecology and climate of the upper Sisquoc Formation and the deposits at Lompoc. She concluded that the Lompoc deposits were younger and that the fishes found there were neritic and pelagic, and those of the upper Modelo more bathypelagic. Her conclusion that the fish fauna of the upper Modelo indicated a basin depth of 200-500 m is probably in error. Recent evidence (Crane 1966) supports the view that the basin was probably at least 1000 m in depth. With the above evidence in mind, it seems most probable that H. imporcitor was a pelagic species, which may have accompanied floating algae, much as do the Recent species Syngna- thus pelagicus and S. californiensis. This con- clusion is supported by the evidence presented by Danil’chenko (1960) for the fishes of the Mai- kop basin. He showed that this basin was anoxic below a depth of 300 m, therefore H. convexus may be assumed to be a pelagic species. ETYMOLOGY.—From the Latin Jmporcitor— ‘‘a diety that presides over the drawing of fur- rows,’ in reference to the greatly expanded trunk ridge. To be treated as a noun in apposi- tion and as of masculine gender. Syngnathus Linnaeus Syngnathus emeritus n.sp. (Figure 15C) Holotype: LACM 27445, an almost complete specimen from the Puente Formation near San Dimas, Los Angeles County, California (Locality LACM 7153). REFERRED MATERIAL.—Puente Formation near San Dimas, Los Angeles County, California, LACM loc. nos. 5153, 7046, 14 specimens and fragments: LACM 26146-26148, 27431- 27434, 27437-27440, 27446-27448. DIAGNosis.—A Syngnathus with 43-47 dor- sal rays; 10 caudal rays; pectoral rays not de- tectable; 14-15 trunk rings; 51 tail rings; snout 1.5-1.8 in head; orbit 5.8-8.7 in head; brood pouch not seen; largest specimen examined 181 mm SL. DESCRIPTION.—Ridges of body smooth and generally not accentuated. Osteology basically the same as in extant species of Syngnathus. Plates small, width of trunk plate less than orbit diameter. Dorsal about as high as depth of ad- jacent body. COMPARISONS.—S. emeritus is unusual in having 15 trunk rings combined with 51 tail rings. These characters and the number of dorsal rays distinguish it from S. avus. The relationships of S. emeritus perhaps lie with the extant species S. leptorhynchus and S. auliscus. It is possible that S. emeritus was on the ancestral line leading to one or both of these species. ETYMOLOGY.—From the Latin emeritus— honorably retired. PALEOECOLOGY.—S. emeritus is found in the diatomaceous shale of the Puente Formation. Its habits probably were much like those of S. avus, i.e., living in relatively shallow water among marine vegetation. FRITZSCHE: REVISION OF EASTERN PACIFIC SYNGNATHIDAE 219 Syngnathus avus (Jordan and Gilbert) (Figure 15B) Syngnathus avus JORDAN AND GILBERT, 1919:34 (Miocene chalk near Titus Avenue, Bairdstown, California, U.C. loc. no. 3525; type-specimen by original designation, SU XLIV, B of original description): JORDAN 1921:259 (reconstruc- tion); Davip 1943:70 (in part; description; synonymy). Syngnathus californiensis avus: HERALD 1941:69 (description; discussion). Holotype: CAS Geology Dept. no. 58372 (SU no. XLIV, B of original description), all fossil fish types and other Jordan paleoichthyological material is now in the Geology Depart- ment, California Academy of Sciences. Paratypes: CAS Geology Dept. no. 58373 (SU no. V, also marked 115, A of original description); CAS Geology Dept. no. 58374 (SU no. 115A, C of original description). REFERRED MATERIAL.—Lower Modelo Formation, Santa Monica Mountains, Los Angeles County, California, LACM loc. no. 1681, 15 specimens: LACM cat. nos. 11598-11605, 26438-26444. DiaGnosis.—A Syngnathus with 36 dorsal rays; 10 caudal rays; 18 trunk rings; 43 tail rings; dorsal on 0.5 + 7.5 rings; pouch covering 22 rings; head 6.4—7.0 in SL; snout 1.7—1.8 in head; dorsal fin base 1.3 in head; maximum known size 225 mm SL. DESCRIPTION.—Generally as given by Jordan and Gilbert (1919). Osteology of neurocranium same as that described for extant species of Syngnathus (Jungersen 1910). Opercles without median ridge. DiscUSSION.—The type material is now in such poor condition that the characters as given by Jordan and Gilbert (1919) cannot accurately be determined. However, specimen No. 58373 (paratype) does have a brood pouch with pro- tecting plates developed; they are not missing as stated by Herald (1941). Syngnathus avus is a rather common member of the fossil fauna of the argillaceous diatomites of the lower Modelo Formation. The original field notes indicate that one slab measuring 4 by 4 ft [1.2 x 1.2 m] contained 42 specimens. It must be pointed out that Figure 2 on Plate XXIX of Jordan and Gilbert (1919) is not of the type of S. avus. In fact, Jordan and Gilbert mis- takenly included a figure of a fossil pipefish from Austria rather than the correct figure of S. avus. This mistake was pointed out by Duncker (1923) and was subsequently corrected by Jordan (1926, 1927). DISTRIBUTION AND RELATIONSHIPS The eastern Pacific syngnathids are generally distributed, with some exceptions, in commonly recognized zoogeographic provinces. Most of the pipefishes are found in bays and sloughs where the water temperature may be higher than along the adjacent open coast. As a result their distribution often extends past the higher-lati- tude, colder-water boundary established for the open-coast fauna. This kind of distribution is shown by other bay-dwelling fishes such as the blennies and gobies (distributions in Miller and Lea 1972). For example, Hypsoblennius gentilis ranges from the Gulf of California to Monterey, Ilypnus gilberti ranges from the Gulf of Califor- nia to Tomales Bay, Clevelandia ios ranges from the Gulf of California to British Columbia, and Quietula y-cauda ranges from the Gulf of Cali- fornia to Morro Bay. Syngnathus leptorhynchus ranges from the outer coast of Baja California to southern Alaska. Thus, these bay-dwelling fishes cross the Point Conception faunal bound- ary (Briggs 1974). There are more species of Syngnathus in the Northern Hemisphere than in the Southern Hemisphere. The close similarity of Japanese Syngnathus schlegeli to generalized members of two distinct eastern Pacific lineages supports the hypothesis that the eastern Pacific Syngnathus are derivable from two separate invasions from the northwest. An alternative hypothesis is in- vasion of Japan from the eastern Pacific fol- lowed by a reinvasion of the eastern Pacific. However, the first hypothesis appears to be the most parsimonious. One lineage consists of S. carinatus, S. macrobrachium, and S. folletti. The other consists of S$. leptorhynchus, S. eu- chrous, and S. insulae. The related S. califor- niensis and S. exilis occupy a somewhat isolated position in the genus and may be the products of another radiation. The problem of defining these lineages and relating them to S. schlegeli is one of not being able to determine plesiomorphic and apomor- phic character states at the species level within the genus Syngnathus. Thus, overall similari- ties, including meristic characters and strength of ridge development, become the most impor- tant criteria used. The carinatus-macrobrachium-folletti lineage is characterized by a relatively long snout; large, well-developed fins; large orbit; and rather well- developed ridges. The presence of carinatus in the upper Gulf of California and macrobrachium and folletti in South America parallels the dis- 220 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 6 tribution given by Chen (1975) for the exsul-spi- norbis and capensis complexes of the subgenus Sebastomus. Chen speculated that these lin- eages probably shared a common ancestor that was able to both invade the Gulf and cross the tropics using the same climatic event. He was, however, unable to rule out multiple crossings. The lack of any close relative of S. carinatus along the outer coast of Baja California or Cal- ifornia is puzzling and quite different from the Sebastomus example given above. It is possible that the carinatus-macrobrachium-folletti \in- eage may have had a southern origin, although more evidence is needed in support of this hy- pothesis. The leptorhynchus-euchrous-insulae lineage is characterized by the medium-length snout, which is round in cross section, and an almost straight snout ridge which does not rise much anterior to the orbit. The restriction of this group essentially to Baja California northward is inter- preted to mean that, assuming a more northern origin, this lineage is much more recent than the preceding one. It is apparent that S. /Jeptorhynchus has given rise to the Guadalupe Island endemic S. insulae, probably as a result of waif dispersal via the California Current. Syngnathus leptorhynchus and S. euchrous probably share a common ancestor. The isola- tion of a population of S. euchrous in Turtle Bay has produced the subspecies S. e. ollotropis. Turtle Bay is unusually warm and the Macro- cystis pyrifera found there has a different growth form than that along the coast (Neushul 1971). A zone of intergradation between S. e. euchrous and S. e. ollotropis exists at Puerto Malarrimo. Syngnathus exilis and §S. californiensis are characterized by an elongate, compressed snout, and rounded ridges. It is possible that they are related to the carinatus-macrobrach- ium-folletti lineage, but their relationships can- not be determined at this time. The relationships of the fossil S. avus and S. emeritus are difficult to determine. It seems highly probable that S. avus is related to the generalized S. leptorhynchus based mainly on meristic characters. The low trunk ring and high tail ring counts of S. emeritus are unlike those seen in living eastern Pacific Syngnathus spp. Syngnathus auliscus is the only truly tropical member of the genus Syngnathus in the eastern Pacific. The characteristic spination of young S. auliscus can also be seen in young S. fuscus from the northwestern Atlantic. This spination has also been reported (D’ Ancona 1933) for the Mediterannean S. agassizi and S$. phlegon. Spi- nation in young Syngnathus may be indicative of a distinct lineage within the genus. A system- atic study of the juveniles of Syngnathus spp. is definitely called for. Syngnathus auliscus is not uncommonly taken in southern California. It is apparent that this species is little affected by the tropical-temper- ate transition between about Sebastian Vizcaino Bay and Magdalena Bay. This distribution pat- tern is shown by other fishes such as Paralabrax maculatofasciatus (distribution in Miller and Lea 1972). However, in San Diego Bay the warm-water effluent produced by the San Diego Gas and Electric power plant provides an ideal environment for tropical species. As a result there is a large population of §. auliscus in San Diego Bay, and the typical California bay species S. leptorhynchus is now less abundant. The genus Bryx is a circumtropical genus with only B. arctus and B. heraldi having temperate populations. There are eight species in the New World: one, B. balli, is in Hawai, B. banneri is in the Indo-West Pacific, and B. darrosanus is in the Indian Ocean. The two subgenera, Bryx and Simocampus, were present before the clo- sure of the Panamanian seaway in the late Plio- cene to early Pleistocene (Woodring 1966), since the eastern Pacific and western Atlantic now have members of both subgenera. The subgenus Bryx is composed of four species, two in the Atlantic and two in the Pa- cific Ocean. This subgenus is presumably de- rived from the subgenus Simocampus and is dis- tinguished from it by the absence of the anal fin. The two species in each ocean are more closely related to each other than either is to the species in the other ocean. In fact the radiations in the two oceans have paralleled each other. The pri- mary difference between the two species is the relative length of the snout. Without a knowledge of the plesiomorphic condition with regard to meristic characters, it is impossible to speculate on the relationships within the subgenus Simocampus. However, the Juan Fernandez endemic B. heraldi was probably derived from B. coccineus or their im- mediate common ancestor. This is unusual since FRITZSCHE: REVISION OF EASTERN PACIFIC SYNGNATHIDAE the ichthyofauna on the Chilean islands of Juan Fernandez and San Félix has a strong Indo-Pa- cific component (McCosker 1970, 1971). The eastern Pacific B. arctus and B. coccineus are more closely related to each other than either is to members of the genus in other oceans. Doryrhamphus melanopleura is an Indo-Pa- cific species that has apparently invaded the eastern Pacific by crossing the East Pacific Bar- rier, since there are no members of this genus in the Atlantic, even though suitable habitat would seem to be available. The eastern Pacific populations of D. melanopleura differ in color- ation from their Indo-West Pacific conspecifics. The Islas Revillagigedo endemic D. paulus has the same color pattern as the eastern Pacific D. melanopleura and was probably derived from mainland populations of D. melanopleura. The genus Leptonotus is a southern cold-tem- perate group with species in South Australia (Munro 1958), New Zealand, and southern South America. This genus is represented in South America by L. blainvilleanus. The mech- anism by which cold-temperate organisms have achieved a distribution spanning the South Pa- cific Ocean is currently under debate. Rosen (1974) has argued that the distribution of at least the freshwater galaxiids covers most of the com- ponents of the original Gondwanaland mass. Thus, the distribution of southern cold-temper- ate organisms reflects the break-up of Gon- dwanaland. However, the possibility of chance dispersal probably utilizing the eastward-flowing South Pacific current seems to be more likely for most marine fishes. This has been docu- mented by McCosker (1970, 1971) for Paraper- cis and Muraenichthys. McCosker pointed out that most of the groups showing this particular distribution pattern are derived from Indo-West Pacific groups dispersing southward to southern Australia and New Zealand, and the subsequent movement eastward of cold-adapted forms across the cold-temperate South Pacific Ocean to Chile. Eschmeyer and Poss (1976) have shown that with some cooling during the Pleis- tocene, a number of near-surface sea mounts, reefs, and islands would have been available to eastward-moving temperate biota in the south- ern Pacific Ocean. Although they were dealing with the scorpionfish genus Maxillicosta, the effects of Pleistocene or earlier cooling would also facilitate dispersal of Leptonotus. The abil- 22] ity of marine fishes to cross oceanic barriers has been well documented (Rosenblatt et al. 1972). Pipefishes (e.g., S. pelagicus) are known to be borne great distances by the currents. It is there- fore likely that the marine L. blainvilleanus was derived from an ancestor from the South Aus- tralia-New Zealand region by chance dispersal of waifs to South America. The relationships of Hippocampus must await a revisionary study of the genus Hippocampus. Our knowledge of the genus Hippocampus is so poor that it is now impossible to determine the exact relationship of H. ingens to other mem- bers of the genus. Since the genus is circum- tropical, it is at least pre-Tethyan in origin. Hipposyngnathus has become extinct since the Tertiary. Because of our almost complete ignorance of the relationships between genera of living syngnathids, it is impossible to relate Hip- posyngnathus to living genera. The reason or reasons for the extinction of this genus remain unknown. BRANCHIAL SKELETON Previous investigators (Duncker 1910, 1912; and Herald 1959) have used the position and de- velopment of the male brood pouch as the di- agnostic character when defining genera and subfamilies. The method of brood pouch closure was used by Herald (1959) in constructing a phy- logeny of the Syngnathidae. The few osteological studies which have in- cluded the Syngnathidae have generally been based on only one or a few of the genera (Jun- gersen 1910; Banister 1967). These studies were therefore not adequately comparative and pro- vided little information on intrafamilial relation- ships. In order better to characterize the genera and subfamilies within the Syngnathidae, I have be- gun a study of the osteology of the syngnathid genera. This study is also designed to be a test of the hypotheses of relationships presented by previous authors who studied the brood pouch. Although the present investigation is still prelim- inary, data can be presented at this time. Because the syngnathids are such highly spe- cialized fishes, many aspects of their osteology (e.g., caudal skeleton) are so reduced that they can provide little phylogenetic information. However, the branchial skeleton does seem to provide useful information. All the genera ex- in) nN Nm PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 6 TABLE 9. SYNGNATHID BRANCHIAL FORMULAE. Genus Bl B2 B3 H1 H2 El E2 E3 12 13. Remarks GASTROPHORI: Heraldia (1)* + aa + + + + + + + fe Maroubra (1) + + = + + + + of ot st Choeroichthys (1) + + - + + 4 + ae + ae Dentirostrum (1) + + ~ + = + + - + + Doryrhamphus (1) + + — + + + + at + + Dunckerocampus (1) ae + = + ae + + - + + Syngnathioides (1) + + — + = at a 1 Nerophis (1)** + + = 4 te - UROPHORI: Leptonotus (1) + ar _ + 3F + + + + at Parasyngnathus (1) + + = + + ar a + + + 2 Ichthyocampus (1) f + = + + +f + + + = Micrognathus (1) a + = + 4 3+ af + + + 3 Syngnathus (7)+ ao + - —/r + 4 + + te + ““Corythoichthys”’ (Atl.) (1) + + - + r + 4 Su at ae Bryx (Bryx) (3) + + - - r +/r r a + + Bryx (Simocampus) (2) + +f = + r +/r r + ate + Pseudophallus (2) + + - = + + + = 4 4 Hippocampus (2) = ~ = 4 + + + + + + Phyllopteryx (1) ~ — = + 9 ae a 4 Corythoichthys (Pac.) (2) + ate Penetopteryx (1) te + - + + + + dt dt ae + = present. — = absent. = reduced: * Number in parenthesis represents number of species examined. ** Data from Jungersen (1910). + Only Syngnathus acus has reduced H1. 1 12&3 with tooth plates. 2 with additional I element. 3 [2&3 with tooth plates. 4 12&3 with tooth plates. amined so far (Table 9) have a full complement of ceratobranchials (i.e., five pairs). The re- maining elements in the branchial skeleton are variously lost or reduced within the family and possibly along distinct lineages within the fam- ily. The Urophori in particular show reduction and loss of every element except the cerato- branchials. The structure of the branchial skel- eton is constant within a genus. Although the information provided by the branchial skeleton is based on characters undergoing reduction and not on uniquely derived characters, it is reason- able to use these characters when defining re- lationships. It seems quite probable that an ele- ment which is lost from such a highly evolved structure would not be regained. The presence of well-developed first hypo- branchials and reduced second hypobranchials and epibranchials clearly distinguish the genus Bryx from Syngnathus (Fig. 16). Bryx was in- cluded by Herald (1959) as a subgenus of Syng- nathus. The loss of all elements except the first basi- branchial and second hypobranchials in the Pa- cific species usually referred to the genus Cor- ythoichthys is clear evidence that the Atlantic and Pacific species should be placed in separate genera (Fig. 17). In fact, Dawson (1977) provi- sionally referred Atlantic species of Corythoich- thys to the catchall genus Syngnathus and more recently placed them in a new genus, Cosmo- campus (Dawson 1979). Herald’s hypothesis that Corythoichthys is ancestral to, or on the lineage leading to, Syngnathus is clearly erro- neous. However, Atlantic species of Cosmo- campus could very well be on the lineage leading to Bryx. Herald’s hypothesis that Leptonotus and Mi- FRITZSCHE: REVISION OF EASTERN PACIFIC SYNGNATHIDAE 723 FIGURE 16. Dorsal view of the branchial skeleton of several syngnathids with the epibranchials and infrapharyngobranchials shown from their ventral aspect. (A) Parasyngnathus elucens, CAS 13696. (B) Bryx arctus, SIO H52-218. (C) Syngnathus cali- forniensis, SIO H47-180. Abbreviations: B—basibranchial; H—hypobranchial; C—ceratobranchial; E—epibranchial; I—infra- pharynogobranchial. crognathus are derivable from Parasyngnathus can neither be supported nor refuted on the basis of their branchial skeletons. Parasyngnathus has a primitive branchial skeleton (Fig. 16). The retention of the upper pharyngeal tooth plates in Micrognathus is the only major difference between it and Parasyngnathus. One interesting point is the extreme reduction of the branchial skeleton in the gastrophorine genus Syngnathoides and in the urophorine gen- era Phyllopteryx and Hippocampus (Fig. 17). These three genera are presently placed in sep- arate subfamilies because of differences in the position and development of the brood pouch. However, in addition to the reduction of bran- chial elements, all three genera have the head at an angle to the main body axis and a prehensile tail without a caudal fin. This evidence may in- dicate that the three genera are much more closely related than has previously been recog- nized. The amount of variation in the branchial skel- eton is much greater in the Urophori than in the Gastrophori. The Gastrophorine genus Heraldia is the only syngnathid genus with three basi- branchials, and it might well be the most prim- itive of the syngnathid genera. The genus Nerophis is usually considered to be the most primitive genus in the family. How- ever, the species referred to this genus have the most specialized branchial skeleton, in terms of reduction, in the Gastrophori (Table 9). FiGurE 17. Branchial skeletons of four syngnathids. (A) Syngnathoides biaculeatus, SIO 60-693. (B) Phyllopteryx fol- iatus, SIO 73-361. (C) Cosmocampus brachycephalus, CAS 24025. (D) Corythoichthys intestinalis, SIO 73-206. 224 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 6 The evidence as is cited above indicates that the structure and development of the branchial skeleton in the Syngnathidae is of great utility in determining relationships. The fact that rela- tionships as determined by the branchial skele- ton transcend recognized subfamilial boundaries is a strong indication that the within-family re- lationships of this group are far from being understood. ACKNOWLEDGMENTS I thank the following individuals for loan of material and/or providing information on speci- mens: John Paxton, AMS; Alex Peden, BC; AI- wyne Wheeler, BMNH; Keith Thompson, BOC; William N. Eschmeyer, Tomio Iwamoto, and Pearl Sonoda, CAS; Fernando Balbontin, EMBCh; C. E. Dawson, GCRL; Robert Behrs- tock, HSU; Norma Chirichigno-F., IMARPE; Robert J. Lavenberg, Camm Swift, and Shelton Applegate, LACM; Myvanwy Dick, MCZ; M. L. Bauchot, MNHN; James Allen, SCCWRP; Leslie Knapp, SOSC; W. R. Taylor and Susan Karnella, USNM; Robert R. Miller, UMMZ; J. Moreland, NMNZ. I thank the following individuals for material aid in the completion of this work: Donald Dockins (deceased), Joseph Copp, and Jan Pul- sifer provided curatorial aid; James Zweifel pro- vided the Laird-Gompertz analysis; John Butler raised the pipefish used in the study of growth; Myvanwy Dick supplied the figure of Hippo- campus ingens collected at San Francisco; fel- low graduate students who discussed various aspects of this problem with me and provided insights from their own research. Dr. Robert Lavenberg provided assistance in preparing the manuscript for publication. I am particularly indebted to Dr. Richard H. Rosenblatt who made this study possible through his aid and advice, and to Dr. Carl L. Hubbs (deceased), who generously provided data collected on pipefishes for over 30 years and for his advice, support, and encouragement. And finally, I wish to thank my wife, Edith, for her support and encouragement throughout the course of this research. Without her I am sure this work would never have been complet- ed. This study is based on a dissertation submit- ted to the University of California, San Diego, in 1976, in partial fulfillment of the requirements for the Ph.D. degree. LITERATURE CITED ANONYMOUS. 1921. Dredging and hydrographic records of the U.S. Fisheries Steamer ‘‘Albatross’’ 1911-1920. Ap- pendix 3, Rep. U.S. Comm. Fish. 1920, Doc. 897, 190 p. ABBoTT, J. F. 1899. The marine fishes of Peru. Proc. Acad. Nat. Sci. Phila. 1899:324—364. ALVERSON, F. G. 1963. The food of yellowfin and skipjack tunas in the eastern tropical Pacific Ocean. Inter-Amer. Trop. Tuna Comm. Bull. 7:293-396. Atz, J. W. 1937. Neptune’s knight: the seahorse. Bull. N.Y. Zool. Soc. 40(2):57-6S. Ayres, W. O. 1854. Description of Syngnathus griseo-linea- tus. Pacific 3(51):202. BANISTER, K. E. 1967. The anatomy and classification of the order Gasterosteiformes (Pisces). Ph.D. Dissertation. Uni- versity of Newcastle upon Tyne, England. 211 p. BARNHART, P. S. 1936. Marine fishes of southern California. Univ. Calif. Press, Berkeley. 209 p. Berry, F. H., AND H. C. PERKINS. 1965. Survey of pelagic fishes of the California Current area. U.S. Fish Wildl. Serv. Fish. Bull. 65(3):625-682. BLEEKER, P. 1858. Vijfde bijdrage tot de kennis der ichth- yologische fauna van de Kokos-eilanden. Nat. Tijdsch. Neder. Indie. 15:457—-468. BONAPARTE, C. L. J. L. 1838. Synopsis vertebratorum sys- tematis. Nuov. Ann. Sci. Nat. 2:105—133. BREDER, C. M., JR. 1928. Scientific results of the second oceanographic expedition of the *‘Pawnee’’ 1926. Nematog- nathi, Apodes, Isospondyli, Synentognathi, and Thoracos- traci from Panama to Lower California with a generic anal- ysis of the Exocoetidae. Bull. Bing. Oceanogr. Exped. 2(2):1—25. Briccs, J. C. 1974. Marine Zoogeography. McGraw-Hill, New York. 475 p. Brown, J. 1972. A comparative life history study of four species of pipefishes (family Syngnathidae) in Florida. Ph.D. Dissertation. University of Florida, Gainesville. 163 p. BROTHERS, E. B. 1975. The comparative ecology and behav- ior of three sympatric California gobies. Ph.D. Dissertation. University of California, San Diego. 370 p. CASTRO-AGUIRRE, J. L., J. ARVIZU-MARTINEZ, AND J. PAEZ- BARRERA. 1970. Contribucion al conocimiento de los peces del Golfo de California. Rev. Soc. Méx. Hist. Nat. 31:107— 181. CHEN, L. 1975. The rockfishes, genus Sebastes (Scorpaeni- dae), of the Gulf of California, including three new species, with a discussion of their origin. Proc. Calif. Acad. Sci., Ser. 4, 40(6): 109-141. CHIRICHIGNO-F., N. 1963. Estudios de la fauna ictiologia de los esteros y parte baja de los rios de departmento de Tumbes (Peru). Servicio de Pesqueria Peru, Serie de Di- vulgacion Cientifica, No. 22. 87 p. 1974. Clave para identificar los peces marinos del Peru. Inf. Inst. Mar Peru-Callao, No. 44: 1-387. CLEMENS, H. B. AND J. C. NOWELL. 1963. Fishes collected in the eastern Pacific during tuna cruises, 1952 through 1959. Calif. Fish Game 49(4):240-264. CRANE, J. M., JR. 1966. Late Tertiary radiation of viperfishes (Chauliodontidae) based on a comparison of recent and Miocene species. Los Angeles Cty. Mus. Contrib. Sci., no. 115:1-29. D’ Ancona, U. 1933. Family Syngnathidae. /n Fauna e Flora del Golfo di Napoli, 38. Monografia: Uova, larve, e stadi FRITZSCHE: REVISION OF EASTERN PACIFIC SYNGNATHIDAE 225 giovanili di teleostei (Monografia eleborata con l’uso del materiale raccolto e seriato da Salvatore Lo Bianco). Stn. Zool. Napoli, pt. 2:284—299. [Translation of the Israel Pro- gram for Scientific Translation 1969] DANIL’CHENKO, P. G. 1960. Kostistye ryby maikopskich otlozenii kavkaza. Tr. Paleontol. Inst. Akad. Nauk SSSR 78:3-207. Davip, L. R. 1943. Miocene fishes of southern California. Geol. Soc. Am. Spec. Pap. no. 43:1-143. Dawson, C. E. 1977. Review of the pipefish genus Cory- thoichthys with description of three new species. Copeia 1977(2):295-338. . 1979. Notes on western Atlantic pipefishes with de- scription on Syngnathus caribbaeus n.sp. and Cosmocam- pus n.gen. Proc. Biol. Soc. Wash. 92(4):671-676. , AND R. A. FRITZSCHE. 1975. Odontoid processes in pipefish jaws. Nature 257:390. , AND J. E. RANDALL. 1975. Notes on Indo-Pacific pipefishes (Pisces: Syngnathidae) with description of two new species. Proc. Biol. Soc. Wash. 88(25):263-280. Dawson, E. Y. 1944. The marine algae of the Gulf of Cali- fornia. Allan Hancock Pac. Exped. 3(10): 189-464. DEBUEN, F. 1963. Peces Chilenos Beloniformes, Syngnath- iformes y Gobiidae. Bol. Soc. Biol. Concepcion 35- 36(1960-61):82-101. DELFIN, F. T. 1901. Catalogo de los peces de Chile. Rev. Chilena Hist. Nat. 3(1899) and 4(1900). 133 p. Dixon, W. J., AND F. J. MAsseEy, JR. 1957. Introduction to statistical analysis. 2nd ed. McGraw-Hill, New York. 488 p. DuMErIL, A. H. A. 1870. Histoire naturelle des poissons ou ichthyologie générale. Vol. 2. Paris. 624 p. DuNcKER, G. 1908. Syngnathiden-Studien. I. Variation und Modifikation bei Siphonostoma typhle L. Mitt. Naturh. Mus. (Jahrb. Wiss. Anst.) Hamburg 25:1-115. 1912. Die Gattungen der Syngnathidae. Ibid. 29(2):219-240. 1915. Revision der Syngnathidae. Erster teil. Ibid. 32:9-120. . 1923. Warnung. Zool. Anz. 56:238-239. EIGENMANN, C. H., AND R. S. EIGENMANN. 1892. A cata- logue of the fishes of the Pacific coast of America, north of Cerros Island. Ann. N.Y. Acad. Sci. 6:349-358. ESCHMEYER, W. N., AND S. G. Poss. 1976. Review of the scorpionfish genus Maxillicosta (Pisces:Scorpaenidae), with a description of three new species from the Australian- New Zealand region. Bull. Mar. Sci. 26(4):433-449. EVERMANN, B. W., AND O. P. JENKINS. 1891. Report upon a collection of fishes made at Guaymas, Sonora, Mexico, with descriptions of new species. Proc. U.S. Natl. Mus. 1891:121-165. , AND H. B. Latimer. 1910. Ona collection of fishes from the Olympic Peninsula, together with notes on other west coast species. Proc. Biol. Soc. Wash. 23:131-139. , AND L. RADCLIFFE. 1917. The fishes of the west coast of Peru and the Titicaca Basin. Bull. U.S. Natl. Mus. 95: 1-166. Eypoux, F., AND P. GERVAIS. 1837. Voyage de “‘La Favor- ite.’ Poissons. Echeneis sexdecim-lamellata et Syngnathus blainvilleanus. Mag. Zool. 7:1-4. FEDER, H. M., C. H. TURNER, AND C. LIMBAUGH. 1974. Observations on fishes associated with kelp beds in south- ern California. Calif. Dep. Fish Game Fish. Bull. no. 160:1- 144. FISCHER, W. 1963. Die Fische des brackwassergebiets Lenga bei Concepcion (Chile). Int. Rev. Gesamten Hydrobiol. 48:419-S11. Fow er, H. W. 1921. Notes on hemibranchiate and lopho- branchiate fishes. Proc. Acad. Nat. Sci. Phila. 73:437-448. . 1944a. Fishes obtained in the New Hebrides by Dr. Edward L. Jackson. Proc. Acad. Nat. Sci. Phila. 96:155— 199. . 1944b. Results of the Fifth George Vanderbilt Ex- pedition (1941). The fishes. Monogr. Acad. Nat. Sci. Phila. no. 6:57-S83. GILBERT, C. H. 1891. Scientific results of explorations by the U.S. Fish Commission Steamer Albatross. No. XIX.—A supplementary list of fishes collected at the Galapagos Is- lands and Panama, with descriptions of one new genus and three new species. Proc. U.S. Natl. Mus. 1890, 13:449-455. 1892. Scientific results of explorations by the U.S. Fish Commission Steamer Albatross. No. XXII.—Descrip- tions of thirty-four new species of fishes collected in 1888 and 1889, principally among the Santa Barbara Islands and in the Gulf of California. Proc. U.S. Natl. Mus. 14:539-566. , AND C. C. Starks. 1904. The fishes of Panama Bay. Mem. Calif. Acad. Sci. 4:1-304. GILL, T. N. 1863. Synopsis of the species of lophobranchiate fishes of western North America. Proc. Acad. Nat. Sci. Phila. 1862:282-284. . 1905. The life-history of the sea-horses (Hippocam- pids). Proc. U.S. Natl. Mus. 28:805-814. GiInsBuRG, I. 1933. Descriptions of five new species of sea- horses. J. Wash. Acad. Sci. 23(12):560-563. . 1937. Review of the seahorses (Hippocampus) found on the coasts of the American continents and of Europe. Proc. U.S. Natl. Mus. 83:497-594. GIRARD, C. F. 1854. Observations upon a collection of fishes made on the Pacific Coast of the United States by Lieut. W. P. Trowbridge, U.S.A., for the museum of the Smith- sonian Institution. Proc. Acad. Nat. Sci. Phila. (for 1854), 7(4): 142-156. 1859. Fishes. Jn General Report upon the Zoology of the several Pacific railroad routes, 1857. U.S. Senate Miscell. Doc. no. 78, 1859, 33 Congr., 2 Sess. 400 p. GUICHENoT, A. 1848. Fauna Chilena. Peces. In Gay, C., Historia fisica y politica de Chile . . . , Paris, (“en casa del Autor’’) Zoologia 2:137—372. GUNTHER, A. 1870. Catalog of the fishes in the British Mu- seum. Vol. 8. Taylor and Francis, London. 549 p. HERALD, E. S. 1940. A key to the pipefishes of the Pacific American coasts with descriptions of new genera and species. Allan Hancock Pac. Exped. 9(3):51—-64. . 1941. A systematic analysis of variation in the west- ern American pipefish, Syngnathus californiensis. Stanford Ichthyol. Bull. 2(3):49-73. _ 1943. Studies on the classification and relationships of the American pipefishes. Ph.D. Dissertation. Stanford University. 339 p. 1953. Family Syngnathidae:Pipefishes. Pp. 231-278 in Leonard P. Schultz et al. Fishes of the Marshall and Marianas Islands. Bull. U.S. Natl. Mus. 202, 1. _ 1959. From pipefish to seahorse—a study of phylo- genetic relationships. Proc. Calif. Acad. Sci. ser. 4, 29(13):465—473. 1965. Studies on the Atlantic American pipefishes with descriptions of new species. Proc. Calif. Acad. Sci. ser. 4, 32(12):363-375. HILDEBRAND, S. F. 1946. A descriptive catalog of the shore fishes of Peru. Bull. U.S. Natl. Mus. 189:1—530. 226 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 6 HoeseE, D. E. 1971. A revision of the eastern Pacific species of the gobiid fish genus Gobiosoma, with a discussion of relationships of the genus. Ph.D. Dissertation. University of California, San Diego. 213 p. Husss, C. L. 1916. Notes on the marine fishes of California. Univ. Calif. Publ. Zool. 16:153—-169. . 1912. Notes on the pipe-fishes of California. Occas. Pap. Mus. Zool. Univ. Mich. no. 100:1-4. , AND K. F. LAGLER. 1958. Fishes of the Great Lakes region. Bull. Cranbrook Inst. Sci. 26:1-213. , AND S. D. HINTON. 1963. The giant sea horse re- turns. Pac. Discov. 16(5):12—-15. JENKINS, O. P., AND B. W. EVERMANN. 1888. Descriptions of eighteen new species of fishes from the Gulf of California. Proc. U.S. Natl. Mus. 1:137-158. JENYNS, L. 1842. Fish. Jn Darwin, C., ed., The zoology of H.M.S. ‘‘Beagle,’’ during the years 1832-1836. Part 4. 172 p. JERZMANSKA, A. 1968. Ichtyofaune des couches a ménilite (flysch des Karpathes). Acta Palaeontol. Pol. 13(3):379-488. JORDAN, D.S. 1895. The fishes of Sinaloa. Proc. Calif. Acad. Sci. ser. 2, 5:377-514. 1921. The Miocene shore-fishes of California. Sci. Mon. 13(5):459-563. . 1926. Ein ungliickliches Versehen. Zool. Anz. 6:221— 224. . 1927. A confusing error. Copeia 1927(161):190—-191. , AND B. W. EVERMANN. 1896. The fishes of North and Middle America. Part 1. Bull. U.S. Natl. Mus. 47:1- 954. , AND . 1927. New genera and species of North American fishes. Proc. Calif. Acad. Sci. ser. 4, 16(5):501— 507. , AND H. W. CrLark. 1930. Checklist of the fishes and fish-like vertebrates of North and Middle Amer- ica north of the northern boundary of Venezuela. Rep. U.S. Comm. Fish. (1928) pt. 2: 1-670. , AND C. H. GILBERT. 1880. Notes on a collection of fishes from San Diego, California. Proc. U.S. Natl. Mus. 3:23-34. , AND 1881. List of the fishes of the Pacific Coast of the United States, with a table showing the distri- bution of the species. Proc. U.S. Natl. Mus. 3:452-458. , AND . 1882. Notes on the fishes of the Pacific Coast of the United States. Proc. U.S. Natl. Mus. 4:29-70. , AND 1883. Synopsis of the fishes of North America. Bull. U.S. Natl. Mus. 16:1-1018. , AND J. Z. GILBERT. 1919. Fossil fishes of southern California. II. Fossil fishes of the Miocene (Monterey) for- mations of southern California. Stanford Univ. Publ. (Univ. Ser.): 13-60. , AND P. L. Jouy. 1882. Checklist of duplicates of fishes from the Pacific Coast of North America, distributed by the Smithsonian Institution in behalf of the United States National Museum. Proc. U.S. Natl. Mus. 4:1-18. , AND E. S. Starks. 1896. Description of a new species of pipefish (Siphostoma sinaloae) from Mazatlan. Proc. Calif. Acad. Sci. ser. 2, 6:268. JUNGERSEN, H. F. E. 1910. Structure of Aulostomidae, Syng- nathidae, and Solenostomidae. K. Danske Vidensk. Skrift. Naturv., ser. 7, 8(5):268—384. Kaup, J. J. 1853. Ubersicht der Lophobranchier. Arch. Na- turges. 19(1):226—234. . 1856. Catalogue of the lophobranchiate fishes in the collection of the British Museum. London. 76 p. KERMODE, F. 1929. Report of the Provincial Museum of Nat- ural History for the year 1928. [British Columbia] F1—F27. 1930. Report of the Provincial Museum of Natural History for the year 1929. [British Columbia] F1—F26. KoepckE, H. W. 1962. Lista de los peces marinas conocidos del Pert con datos de su distribucion geografica. Pt. 3. Biota 4(33): 193-204. LairD, A. K., A. D. BARON, AND S. A. TYLER. 1968. Growth and time: An interpretation of allometry. Growth 32:347-354. LINNAEuS, C. 1758. Systema naturae. Regnum animale. 10th ed. Guiliemi Engelman, Lipsae. 824 p. LoBeELL, M. J. 1942. Some observations on the Peruvian coastal current. Trans. Am. Geophys. Union, pt. II:332- 336. MANN, F. G. 1954. La vida de los peces en aguas chilenas. Ministerio de Agricultura, Univ. of Chile, Santiago. 342 p. McCoske_er, J. E. 1970. A review of the eel genera Lep- tenchelys and Muraenichthys, with the description of a new genus, Schismorhynchus, and a new species, Murae- nichthys chilensis. Pac. Sci. 24(4):506—-516. —. 1971. A new” species of Parapercis (Pisces: Mugiloididae) from the Juan Fernandez Islands. Co- peia 1971(4):682-686. , AND C. E. DAwson. 1975. Biotic passage through the Panama Canal, with particular reference to fishes. Mar. Biol. 30:343-351. MEEK, S., AND S. F. HILDEBRAND. 1923. The marine fishes of Panama, pt. 1. Field Mus. Nat. Hist. Publ. 215, zool. ser. 15: 1-330. MILLER, D. J. AND R. N. LEA. 1972. Guide to the coastal marine fishes of California. Calif. Dep. Fish Game Fish Bull. no. 157. 235 p. Morrow, J. E. 1957. Shore and pelagic fishes from Peru, with new records and the description of a new species of Sphoeroides. Bull. Bing. Oceanog. Coll. 26(2):5—S5. Munro, I. S. R. 1958. Syngnathiformes. Pp. 81—96 in Hand- book of Australian fishes. Fish. Newsl. 17(2-S). NEUSHUL, M. 1971. The species of Macrocystis with partic- ular reference to those of North and South America. Pp. 211-222 in North, W. J., ed. The biology of giant kelp beds (Macrocystis) in California. Beihefte zur Nova Hedwigia 32. NicHo_s, J. Y., AND R. C. MurpHy. 1922. On a collection of marine fishes from Peru. Bull. Am. Mus. Natl. Hist. 46:501—-516. , AND 1944. A collection of fishes from the Panama Bight, Pacific Ocean. Bull. Am. Mus. Nat. Hist. 83(4):217-260. NorMAN, J. R. 1937. Coast fishes. Part II. The Patagonian region (including the Straits of Magellan and the Falkland Islands). Discovery Rep. 16:1—150. OsBuRNn, R. C., AND J. T. NicHoLs. 1916. Shore fishes col- lected by the *‘Albatross’’ expedition in Lower California with descriptions of new species. Bull. Am. Mus. Nat. Hist. 35(16): 139-181. PHILIPPI, R. A. 1896. Peces nuevos de Chile. Anal. Univ. Chile 93:375-390. RAFINESQUE, C. S. 1810. Caratteri de alcuni nuovi generi e nuove specie di animali e plante della Sicilia, con varie os- servazioni sopra i medisimi. Palermo. 105 p. RAUTHER, M. 1925. Die syngnathiden des Golfes von Nea- pel. Publ. Stn. Zool. Napoli 36:1-365. FRITZSCHE: REVISION OF EASTERN PACIFIC SYNGNATHIDAE 227 REGAN, C. T. 1908. Pisces. Biologia Centrali-Americana, 1906-1908. 203 p. RICKER, K. E. 1959. Fishes collected from the Revillagigedo Islands during the 1954-1958 cruises of the “‘Marijena.”’ Univ. Brit. Col. Inst. Fish. Mus. Contrib. no. 4:1-10. Rosen, D. E. 1974. Phylogeny and zoogeography of salmon- iform fishes and relationships of Lepidogalaxias salaman- droides. Bull. Am. Mus. Nat. Hist. 153(2):267—325S. ROSENBLATT, R. H. 1963. Some aspects of speciation in ma- rine shore fishes. Pp. 171-180 in Speciation in the sea. Sys- tematics Assoc. Publ. no. 5. , J. E. McCoskER, ANDI. RUBINOFF. 1972. Indo-west Pacific fishes from the Gulf of Chiriqui, Panama. Los An- geles Cty. Mus. Contrib. Sci. no. 234:1-18. SCHULTZ, L. P. 1957. The frogfishes of the family Anten- nariidae. Proc. U.S. Natl. Mus. 107:47-105. Siccarpi, E. M. 1954. Consideraciones sobre el modo de reproduccion de ‘‘Leptonotus blaenvillianus’’ Eydoux and Gervais, 1837 (Pisces: Syngnath.). Comun. Inst. Nac. In- vest. Cien. Nat. Cien. Zool. 2(14):211-242. SNYDER, J. O. 1911. Descriptions of new genera and species from Japan and the Riu Kiu Islands. Proc. U.S. Natl. Mus. 40:525-549. Starks, E. C. 1911. Results of an ichthyological survey about the San Juan Islands, Washington. Ann. Carnegie Mus. 7:162-213. , AND E. L. Morris. 1907. The marine fishes of south- ern California. Univ. Calif. Publ. Zool. 3:159-252. SFEINDACHNER, F. 1898. Die Fische der Sammlung Piate. Fauna Chilensis. Zool. Jahrb. (Suppl.) 4:28 1-337. Storer, D. H. 1848. Description of a new species of Syng- nathus, brought from the western coast of California by Capt. Phelps. Proc. Boston Soc. Nat. Hist. 1845, 2(10):73. Swain, J. 1882. A review of the Syngnathinae of the United States, with a description of one new species. Proc. U.S. Natl. Mus. 5:307-315. , AND S. E. MEEK. 1884. Notes on the pipefishes of Key West, Florida, with description of Siphostoma Mc- Kayi, a new species. Proc. U.S. Natl. Mus. 7:237—239. TayLor, W. R. 1967. An enzyme method of clearing and staining small vertebrates. Proc. U.S. Natl. Mus. 122:1-17. THompson, 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. Proc. U.S. Natl. Mus. 50:401-476. Urey, A. B. 1929. A check-list of fishes of southern Cali- fornia and Lower California. J. Pan-Pac. Res. Inst. 4(4):2- Ml 1932. Lista de los peces de las costas de la Baja California. Ann. Inst. Biol. 3(1):75—80. , AND P. O. GREELEY. 1928. A list of the marine fishes (Teleostei) of southern California with their distribution. Bull. South. Calif. Acad. Sci. 27(1):1—53. VAILLANT, L. L. 1883. Remarques sur les affinités naturelles des familles composant le sous-ordre des poissons mala- coptérygiens abdominaux. Ann. Sci. Nat. (Zool.), sér. 6, 15(7): 1-13. . 1888. Mission scientifique du Cap Horn, 1882-1883. VI. Zool., Poissons. Paris. 35 p. 1894. Sur une collection de poissons recueillie en Basse-Californie et dans le Golfe par M. Léon Diguet. Bull. Soc. Philom. Paris, sér. 3, 6:69-75. Waite, E. R. 1909. Subantarctic islands of New Zealand. Art. XXV. Vertebrata of the subantarctic islands of New Zealand. Pisces:585—598. WEBER, M. AND L. F. DE BEAUFORT. 1922. The fishes of the Indo-Australian Archipelago. IV. Leiden. 410 p. WHITLEY, G. P. 1931. New names for Australian fishes. Aust. Zool. 6(4):310-334. 1940. Illustrations of some Australian fishes. Aust. Zool. 9(4):397—428. . 1955. Sidelights on New Zealand ichthyology. Aust. Zool. 12(2):110-119. WooprRING, W. P. 1966. The Panama land bridge as a sea barrier. Am. Phil. Soc. 110(6):425—453. ZWEIFEL, J. R. AND R. LASKER. 1976. Prehatch and post- hatch growth of fishes—A general model. Fish. Bull., U.S. 74(3):609-621. CALIFORNIA ACADEMY OF SCIENCES Golden Gate Park San Francisco, California 94118 Vol. 42, No. 7, pp. 229-285; 30 figs., 7 tables tuarRARy PROCEEDING: JUL 18 1980 OF THE CALIFORNIA ACADEMY SCIENCES ''<, Mass. A REVISION OF THE FISH GENUS OGCOCEPHALUS WITH DESCRIPTIONS OF NEW SPECIES FROM THE WESTERN ATLANTIC OCEAN (OGCOCEPHALIDAE; LOPHITFORMES) By Margaret G. Bradbury’ Department of Biological Sciences, San Francisco State University, San Francisco, California 94132 ABSTRACT. The New World genus Ogcocephalus comprises twelve species. Two species are island endemics in the eastern Pacific, Ogcocephalus darwini Hubbs in the Galapagos archipelago and Ogcocephalus porrectus Garman off Cocos Island, but the remaining ten are western Atlantic species. Five of the Atlantic species are described as new: Ogcocephalus pantostictus and Ogcocephalus declivirostris from the northern and western Gulf of Mexico, Ogcocephalus rostellum from the Atlantic coast of the southeastern United States, Ogcocephalus corniger also from the Atlantic coast of the southeastern United States but ranging into the eastern Gulf of Mexico, and Ogcocephalus pumilus from the Caribbean and coasts of the Guianas. Ogcocephalus parvus Longley and Hildebrand has a wider range than formerly known, from the coast of the southeastern United States and eastern Gulf of Mexico through the Caribbean to the Atlantic coast of South America. Ogcocephalus vespertilio (Linnaeus) has a more restricted range than formerly thought, the coast of Brazil from the mouth of the Amazon to the mouth of the Rio de la Plata. Ogcocephalus notatus (Cuvier and Valenciennes) also has a southerly distribution in the western Atlantic, the coast of northern South America from Colombia to northern Brazil. The variable species Ogcocephalus nasutus (Cuvier and Valenciennes) appears to be allopatric with O. vespertilio (Linnaeus), which it most resembles; O. nasutus ranges from the mouth of the Amazon through the Caribbean to the Bahamas and southeastern Florida. Ogcocephalus cubifrons (Richardson) ranges from the Bahamas and the coast of the southeastern United States into the eastern Gulf of Mexico to at least Pensacola, Florida, and Campeche Banks. The name Ogcocephalus radiatus (Mitchill) is placed in the synonymy of Ogcocephalus cub- ifrons. Illustrated keys, photographs, diagnoses, and distribution maps are provided. F Marine Bldiopica) A abara cy ty | July 2, 1980 INTRODUCTION Ogcocephalids from the western Atlantic Ocean have become available in large numbers during the last two or more decades, much of the new material a handsome dividend from the work of exploratory vessels of the U.S. National Marine Fisheries Service sampling new fishing 1 Research Associate, Department of Ichthyology, Califor- nia Academy of Sciences. grounds. Prior to this exploratory period, spec- imens of ogcocephalids were uncommon in col- lections except for those captured inshore by swimmers or divers, particularly off Florida and off islands in the Caribbean. Few in number and little studied, these specimens were difficult to assign names to because variation in characters then used in diagnoses was not understood. Variation in the length of the prominent rostrum was a frequent source of error. The information developed in this study is that some species may be diagnosed by their long rostrums at all sizes, [229] 230 but the rostrum in other species varies allo- metrically so that juveniles have long rostrums, but adults have short ones. Analysis of morphometric data, including data about the rostrum, reveals that the genus Ogco- cephalus is represented in the western Atlantic by ten species, five of which are described here as new. More recently, new material from the eastern Pacific has also become available. Study of this material confirms that two derivative species are present in the eastern Pacific (Gar- man 1899; Hubbs 1958). Studies on the remaining genera of Ogcoce- phalidae of the Atlantic Ocean are underway currently. Discussions of the zoogeography of the group and intrafamilial relationships of all the Atlantic species will be published with the results of those studies. ACKNOWLEDGMENTS This study has been pursued intermittently since the late 1950’s. During this time I became indebted to many colleagues to whom it is a pleasure to extend my grateful thanks at last. I deeply appreciate the encouragement given me by the late Loren P. Woods, who first suggested this problem and made available the specimens, skeletal material, and photographs of ogcoce- phalids in the Field Museum of Natural History; he also read an early version of the manuscript and devoted a great deal of time to assisting me. My dear friend, the late Marion Grey, gave valu- able bibliographic assistance as well as the en- thusiastic encouragement that so often charac- terized her relations with her colleagues. Pearl M. Sonoda sent specimens on each of a number of occasions and cheerfully gave much other as- sistance. Hymen Marx discussed ideas with zest and humor during one memorable Chicago sum- mer. George S. Myers and the late Rolf L. Bolin, who were my professors at Stanford University, gave valuable help; Prof. Bolin gave skillful crit- icism of an early version of this report. With characteristic generosity, Giles W. Mead kindly made possible a study trip to museums in the eastern United States and gave support and en- couragement in various ways. The late Carl L. Hubbs gave advice about taking counts and measurements and graciously examined for me a Linnaean type-specimen at Uppsala. He along with W. I. Follett and Lillian Dempster helped with the coining of names. Photographs and radiographs were generously PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 7 provided by the following: Harvey R. Bullis, Jr., Bruce B. Collette, William N. Eschmeyer, Da- vid W. Greenfield, Ake Holm, Robert J. Lav- enberg, Garnett W. Link, Jr., Robert N. Lea, Giles W. Mead, and Loren P. Woods. For their valuable time, I am grateful to the following peo- ple who loaned specimens or made arrange- ments for me to work at their institutions: Wil- liam W. Anderson, Richard T. Barber, Frederick H. Berry, James E. Bohlke, the late Anton Brunn, Harvey R. Bullis, Jr., Daniel M. Cohen, Bruce B. Collette, C. E. Dawson, Myvanwy M. Dick, Robert H. Gibbs, Carter R. Gilbert, David W. Greenfield, Jena Guibé, Carl L. Hubbs, Rob- ert K. Johnson, Robert J. Lavenberg, Giles W. Mead, George C. Miller, Martin A. Moe, Jr., C. Richard Robins, Richard H. Rosenblatt, Jay M. Savage, Leonard P. Schultz, Robert L. Shipp, Victor G. Springer, Camm C. Swift, Ethelwynn Trewavas, Vladimir Walters, and Loren P. Woods. I thank Lillian J. Dempster and W. I. Follett for clarifications on nomenclatural ques- tions. This investigation was supported by Public Health Service Fellowship No. 8543 from the Division of General Medical Sciences, Public Health Service, and by National Science Foun- dation grant GB-4777 from the Division of Bio- logical and Medical Sciences. Support from these agencies was essential for the completion of this work, and I am deeply appreciative. SPECIAL TERMINOLOGY The depressed anterior portion of the fish is called the disk and consists of the cranium, opercular apparatus, and the trunk back to the pectoral axillae. The lateral margins of the disk are formed by the opercular bones, which are very elongate relative to the cranium. The cra- nium is of ordinary proportions relative to the standard length, so the large total size of the disk is due primarily to the elongate opercular series, which sweeps backward from the crani- um and envelops the trunk on either side to form the disk. The (usually) conspicuous protuberance from the front of the head dorsal to the nasal openings is called the rostrum. Appearing as a median horn in most species, it varies in relative length from short to very long. It may bear cirri at its distal end, but its function remains unknown at this time. No skull bones are involved in the BRADBURY: FISH GENUS OGCOCEPHALUS OUTSIDE MARG/N OF LATERAL ETHMO/D & ' ty x -x\t * re * X-2 ~— = E D C FIGURE 1. Ki See 231 2 ESCA 4 A XW SERIES Lateralis system of Ogcocephalus cubifrons, 275 mm SL. Each lateral-line scale indicated by pair of opposing U-shaped symbols representing flaps of skin associated with such scales. Canals indicated by dashed line. (a) Lateral view showing lateral line beginning at A; portion of supraorbital series also shown, beginning at B. (b) Oblique view of head showing cheek series, A to C, premaxillary series, and subopercular series, C to E. structure of the rostrum; it is formed solely of modified scales. Just beneath the rostrum is a cavity lined with scaleless skin into which the angling lure or il- licium may be drawn. This cavity is here called the illicial cavity. The somewhat bulbous struc- ture seen within the illicial cavity is the bait of the angling apparatus, or esca (Fig. 1b), a mov- able glandular appendage attached to the distal end of the illicium. The illicium is a bony rod, quite short, but when extended, can be seen to be covered with scaleless skin like the skin which lines the illicial cavity; the skin of the illicium is thrown into accordionlike folds when the illicium is retracted. In members of the Ogcocephalidae the scales are highly modified structures of three main types (Bradbury 1967: fig. 5). The simplest type is a pyramid- or cone-shaped scale called a tu- bercle, typically with a spine at its apex, al- though there may be several. The kind of scale that characterizes Ogcocephalus, called a buck- ler, is larger and more complex; bucklers are cone-shaped, too, but covered with small spi- nules rather granular in appearance and ar- ranged in rows that radiate out from the apex of the buckler. These granular spinules fill the spaces between bucklers. The morphological re- lationship between tubercles and bucklers is not clear; the spinules of bucklers often resemble tubercles, suggesting that the buckler may be a compound structure consisting of one enlarged tubercle overlain with small tubercles arranged in the radiating pattern, but the matter cannot be clarified until developmental studies of the scales are undertaken. Finally, besides tubercles and bucklers, spe- cially shaped lateral-line scales occur. With ap- propriate preparations it can be shown that lat- eral-line scales are bowl- or dish-shaped (Bradbury 1967: fig. 5), but under ordinary view- ing conditions their shapes are difficult to dis- cern. However, they are often made conspicu- ous because of associated flaps or fringes of epidermis; when epidermal decorations are ab- sent, lateral-line scales may often be distin- guished because of the large single neuromast lying in the center of the cup of each scale. METHODS CouNTS.—Vertebral counts were made from radiographs and include the hypural plate. Scales of the lateral-line system can generally be seen in most species of Ogcocephalidae, al- 232 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 7 POSTERIOR SWELLING OF THE MANDIBLE MEDIAL RIM ok: OF ye SUBOPERCLE fe ’ TERMINAL PAIR OF A SUBOPERCLE MAJOR DISK BUCKLERS FIGURE 2. Ventral view of Ogcocephalus cubifrons, 275 mm SL, showing disk margin. (a) Length of disk margin is distance between two points indicated by the two arrows. (b) Position of subopercle relative to disk margin. though the canals are difficult to discern as a rule. Thus it may be difficult to follow and count lateral-line scales when they are small and spaced far apart on the surface of the disk where they tend to become lost in the rough integu- ment, but elsewhere on the body it is generally possible to see the scales and follow the lines of the canals by using adequate magnification. In this paper I have used four series of counts of scales of the lateral-line system as taxonomic characters. 1. Lateral line (Fig. la). The lateral line begins just posterior to the eye at A. Care must be tak- en not to include scales of the supraorbital series which begins at B. The line of scales continues back across the dorsal surface of the disk and suddenly descends ventrally just posterior to the gill pore, continuing along the lateral sides of the tail and on to the base of the caudal fin. 2. Subopercular series (Fig. 1b). This portion of the cephalic lateral-line system branches from the preoperculomandibular canal at C and con- tinues posteriorly along the ventral side of the subopercle to E. A secondary branch leaves the subopercular series at D and runs up onto the dorsal side of the disk for a short distance. Care must be taken to avoid including the first scale of this branch series in the subopercular latéral- line scale count. The subopercular lateral-line scale count was always taken as the count be- tween C and D because the posterior section (between D and E) does not vary from a count of 3. Thus, the subopercular lateral-line scale count in the figure is 6. 3. Cheek series (Fig. 1b). I have called the section of lateral line that extends in a sigmoid curve from A to C the cheek series, although its topographic position suggests that it consists of lateral-line scales from the infraorbital canal plus the opercular section of the preoperculomandib- ular canal. The cheek series excludes scales of the branch extending from B towards the eye. 4. Premaxillary series (Fig. 1b). The premax- illary series is a short section of infraorbital ca- nal just above the upper lip. MEASUREMENTS.—The highly specialized body form of these fishes requires some modi- fication of standard measuring procedures as pointed out by Hubbs (1958). I have followed his procedures in large part, but made adjust- ments that permit comparative measurements of members of other genera. Measurements were made either with sharp- pointed steel calipers and a steel rule, or with dial calipers, and were recorded to the nearest 0.1 mm; measurements of approximately 15 mm or less were taken viewed through a low-power dissecting microscope. All measurements in- volving the upper or lower jaws were taken from the anteriormost median point of the lips without pressing the calipers into the lip tissue. All mea- surements involving bucklers were taken to the base of the buckler and not to its apex, unless otherwise stated. Standard length is the distance from the upper lip to the base of the caudal rays. The latter point is difficult to establish in large specimens in which the integument is thick and stiff; the cal- ipers were pressed firmly back against the swell- ing formed by the bases of the rays. Predorsal distance is the distance from the center of the upper lip to the base of the first dorsal ray; preanal distance is the distance from the center of the lower lip to the base of the first anal ray. The distance from the jaw to the anus is the distance between the center of the lower lip and the center of the anal opening. The point of articulation of the mandible with the suspensorium is visible externally as a pro- nounced swelling at the posterior end of the BRADBURY: FISH GENUS OGCOCEPHALUS OUTLINE OF ILLIC/IAL FOSSA FIRST PAIR OF MAJOR BUCKLERS 233 FIRST PAIR OF b MAJOR BUCKLERS POINT OF GREAT- EST CONCAVITY OF THE CRANIUM APEX OF ANGLE OF SUBORBITAL POSTERIOR SWELLING OF THE MAND/BLE BUCKLERS FIGURE 3. Views of head of Ogcocephalus cubifrons, 275 mm SL, showing specific locations of points used in taking measurements of head. Head depth is taken with one needle of calipers set between bucklers of first major pair (a) and (b), with other needle on apex of swelling of mandible (b). Width of cranium is taken between points of greatest concavity just posterior to orbits (b); also shown by arrows in (a). Suborbital width is least distance between orbit and apex of angle formed by suborbital bucklers (5). mandible; the measurement of the disk margin was taken with one point of the calipers pressed against the front of this swelling and the other point pressed against the posterior end of the subopercle (arrows, Fig. 2a). Care must be tak- en in determining the posterior end of the sub- opercle in specimens with a thick stiff integu- ment, but the tip of the subopercle can be felt with the caliper point just posterior to the ter- minal pair of large bucklers at the angle of the disk. This measurement of the disk margin was substituted for head length, which proved to be an awkward measurement in some species. Trunk width is the ‘“‘width behind P, [pectoral] fins’’ (Hubbs 1958) which ‘‘is taken where the body margin reaches a maximum curvature near the solid part of the urosome, behind the disk.”’ Length of rostrum was taken as the distance from the dorsal rim of the posterior nostril to the distal tip of the terminal spine of the rostrum. The nostril landmark was chosen, in spite of being off-center, because it is in precisely the same relative position in all species. Although the rostrum appears to be highly variable in length in Ogcocephalus, results of this study show that some of the variation results from al- lometric growth patterns. When this allometry is borne in mind, the length of the rostrum proves a useful distinguishing feature in these fishes. Width of cranium (Fig. 3a) corresponds to ‘skull prominence width’’ of Hubbs (1958). It is the distance taken between the ‘“‘points of greatest concavity’? which lie just posterior to the orbits (Fig. 3a, arrows, and 3b). Head depth is the oblique distance from a median point on the roof of the cranium between the first pair of major bucklers (Fig. 3a) to the apex of the cone- shaped swelling formed by the mandible where it articulates with the suspensorium (Fig. 3b). The first pair of major bucklers invariably occurs just posterior to a noticeably soft spot on the dorsal surface of the cranium, which is actually the fossa into which the illicium retracts (Fig. 3a). Although off-centered, this measurement of head depth taken as described is a useful, re- peatable measurement. The eye measurement is the greatest diameter of the bony orbit; since there are no circumorbital bones, the lateral pro- cess of the sphenotic bone forms al! there is of a posterior bony margin of the orbit. The rear needle of the calipers was always pressed against this bone. Suborbital width is the least distance between the margin of the cornea (well marked by its rim of tubercles) and the apex of the sharp angle formed by the suborbital buck- lers just above the angle of the lips (Fig. 3b). Lateral ethmoid width is the least distance be- tween the outside margins of the lateral ethmoid bones, which are the bones that form the sides of the illicial cavity (Fig. 1b). Mouth width is the greatest overall width, i.e., the distance between the posterior tips of the maxillaries. Taking this measurement and the following one require some care because the maxillary tips are usually concealed beneath the suborbital integument. 234 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 7 The length of the jaw is taken from the median point of the upper lip to the posterior tip of the maxillary. The snout length measurement is the least distance from the anterior bony margin of the orbit to a point on the median line just above the upper lip and exactly at the base of a strip of scaleless skin lying between the two armored halves of the snout. Length of dorsal and length of anal are the lengths of these fins from the base of the first ray to the tip of the longest ray with the fin depressed. The patterns of dentition in species of Ogco- cephalus show little variation from the pattern described for O. darwini by Hubbs (1958). Brad- bury (1967) described the dentition in the genus and compared it with patterns in other genera with the use of alizarin-stained and cleared spec- imens. Since there are no readily discernible dis- tinctions in the dentitions of species of Ogco- cephalus that are taxonomically useful, features of dentition are omitted from this report. As is often the case with endeavors of this kind, sampling was not random, and specimens were studied as they came available regardless of their size or the locality from which they came. Usually habitat differences were not known. Meristic differences proved to be less useful for discriminating among populations than body proportions and color patterns, and the latter were used repeatedly in diagnosing specimens. It was convenient to express body proportions as ratios, and these ratios are used in the species diagnoses. However, for system- atic comparisons, proportions are also given in terms of percent of standard length in tabular format. Counts of pectoral fin rays and the var- ious lateral-line scale series were made on both right and left sides, and counts from both sides are included in the tables; most of the tables present these data in right-and-left pairs. Abbreviations for fish collections are as fol- lows: AMNH, American Museum of Natural History; ANSP, Academy of Natural Sciences of Philadelphia; BMNH, British Museum (Nat- ural History): CAS, California Academy of Sci- ences; FMNH, Field Museum of Natural His- tory; FSBC, Florida State Board of Conservation Marine Laboratory; GCRL, Gulf Coast Re- search Laboratory Museum; LACM, Los An- geles County Museum; MCZ, Museum of Com- parative Zoology, Harvard University; MNHN, Muséum National d’Histoire Naturelle, Paris; SIO, Scripps Institution of Oceanography; SU, Stanford University (this collection is now housed in the California Academy of Sciences and therefore listed as CAS-SU); UF, Florida State Museum, University of Florida; UMML, Marine Laboratory, University of Miami; USNM, U.S. National Museum of Natural His- tory, Smithsonian Institution; ZMUC, Univer- sitetets Zoologiske Museum, Copenhagen. Ogcocephalus Fischer Ogcocephalus FISCHER, 1813:78 [type-species Lophius ves- pertilio Linnaeus, by subsequent designation of Jordan and Evermann 1896:511]; BRADBURY 1967. Malthe Cuvier, 1816:311 (part) [type-species Malthe vesper- tilio Cuvier and Valenciennes (non Linnaeus), 1837, by sub- sequent designation of Gill 1878:232]. Malthea (or Malthaea) CUVIER AND VALENCIENNES, 1837:440 [emended spelling of Malthe and therefore taking the same type-species]. Oncocephalus JORDAN, 1895:506 [unjustified emendation of Ogcocephalus Fischer]. Onchocephalus GiLL, in GOODE AND BEAN, 1896:498 [unjus- tified emendation of Ogcocephalus Fischer]. Of the two unjustified emendations of Ogco- cephalus which have appeared in the literature, at least one had the objective of correcting an alleged improper transliteration from the Greek. Jordan and Evermann (1898:2736) state with re- gard to the original spelling of Ogcocephalus, ‘‘properly written Oncocephalus, but Fischer chose the above monstrous spelling.’’ The sec- ond emended spelling may have been introduced for the same reason, but in fact no reason was put forward. In any event, the original spelling of the name is a correct original spelling, and the name stands as Ogcocephalus Fischer (see also Briggs 1961). DIAGNOs!Is.—A genus of Ogcocephalidae with gill rakers in the form of oval pads which are studded with teeth resembling the fine teeth elsewhere in the buccal cavity (Bradbury 1967; fig. 7). Gills 22, none on the first arch; complete holobranchs on the second and third arches, a hemibranch on the fourth arch. Scales on ventral surface of tail consisting of numerous, tiny, closely spaced tubercles; if any bucklers occur, they are scattered or may occur ina single short median row, but in any case, never are arranged in two longitudinal rows that cover the entire ventral surface as in Zalieutes or Malthopsis. Esca distinctly trilobed, with a median dorsal, somewhat leaf-shaped, lobe and two ventral lobes arranged symmetrically below the median lobe (Bradbury 1967: fig. 3). BRADBURY: FISH GENUS OGCOCEPHALUS 235 DESCRIPTION.—Body form. Body depressed vA iol ledees alien anteriorly, forming a disk which is subtriangular in outline. Cranium well elevated above general Slit ee pele eae lbs nal allen al slope of disk, proportions between width of cra- Bi nium and head depth showing little variation Scam Mileabeslieil dst agi mlehisa yeh within genus (width of cranium into head depth Fa = Tipe keel lie te id 1.42.1). Also, there is little variation in length of snout and width of suborbital relative to width SSeouIO Ls Me Ale sess Hl pallens | of cranium (suborbital width into width of cra- Seam omee en a el WI nium 1.0—1.9; snout length into width of cranium Sass 1.42.4). Rostrum variable in length, from a rel- Seine fl | atively small knob to a long median horn. Gill K fe pores opening dorsally, lying in the axillae a = Brea ialele or tLe 1 short distance in advance of rear margin of disk. S SRl ip prmeme7y yd i No membranes connecting pectoral peduncles & a with trunk as occur in Halieutichthys. Ventral Soa tee el ol sl surface of disk flat. 2 ellie =. ber gas ott Tail stout and muscular, relatively wide at its o 2 5. | | Joell a junction with disk, tapering to a narrow caudal cS js jst | | | base. Ventral surface of tai! flattened so that a eee. cross section through anterior half of tail would = s S she Pe ee [ieee be nearly a half circle. Predorsal distance 1.4— 2) 2 rs 1.7 in standard length; preanal distance 1.2—-1.4 s eas = Se | : . : Stel in standard length. Anus slightly in front of a < Elag oma ONAAR] = | line drawn between posterior tips of pectoral z g re elbows when pressed against trunk; distance Peet | er eee re | from jaw to anus 1.6—1.9 in standard length. 6 se|scane pron jo] ' Head Features. Eyes directed laterally and a 6 a ie little anteriorly and downward. Cornea much 5 Sapa Oe JOAN NOK | | smaller than orbit, skin extending from cornea z around eyeball studded with small tubercles. E eae | Depending upon its: condition at time of pres- 5 =elro-++ enn o~m | ervation, iris shows a more or less pronounced a pupillary operculum (except in O. notatus) on g Soe wee ec its ventral margin, giving pupil an approximate ce EE RP I eee ree eee kidney shape. Sometimes a less noticeable dor- és ne | a Mad sal operculum (again excepting O. notatus) ex- a seil[rm[—m[amtramnaveon tends downward towards its ventral mate so that < pupil is dumbbell-shaped; presumably these Gy We Oh col Sale ae ig ana opercula regulate pupil size, but I know of no pte hs ano te experiments that demonstrate this function. aula | | iP | Anterior nostril round, the aperture encircled Seale eae Wal el Halas | by arim of skin which often forms a short tube; posterior nostril slitlike, larger than anterior Z|RRAPSRSREESRS nostril. Jaws very protractile, mouth moderate to small, usually semicircular, its width DAES)? a 2 in length of disk margin except in O. notatus, pers 2 ine Ei Ey E 2 which has a relatively wide mouth and in which S = S = = & § = ~ S = = the ratio is 1.8—2.5. In normal position, posterior SSSSISSESSES S ends of maxillaries concealed beneath the sub- orbital bucklers. Upper jaw terminal or subter- minal relative to snout; lower jaw included; gape oblique. 236 TABLE 2. Ogcocephalus. Dorsal fin 1 2 3 4 5 notatus — 1 — 35 17 darwini —_ ae 1 16 1 porrectus —_— a= 11 22 _ vespertilio — — 3 26 8 nasutus — 1 1 57 11 cubifrons —_ — 3 23 10 pantostictus — — 3 25 11 rostellum — — 5 20 1 corniger — — 7 30 1 parvus — 1 1 64 3 declivirostris —_— — 1 39 13 pumilus 1 — 9 37 3 Fins. Dorsal fin small, placed far posteriorly on tail to rear of level of anus; dorsal rays usu- ally 4-5, unbranched. Anal fin ‘‘like a narrow lappet’’ (Hubbs 1958) or a prop for the tail, usu- ally fleshy, particularly along anterior edge and tips of first two rays. Apparently this fin lies twisted on its side while fish rests on the sub- strate. Typically the anal fin has 4 unbranched rays, but O. pumilus frequently has only 3 (Ta- ble 2). Pectorals sturdy, variable in shape from relatively long and narrow to broad and fan shaped; pectoral rays with skin especially fleshy on distal ventral surfaces in 9 of the 12 species, this fleshy tissue not developed on ventral sur- face of pectorals in very small specimens as a rule. Pectoral rays 10-15, unbranched (Table 3). PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 7 FREQUENCY DISTRIBUTIONS FOR DORSAL AND ANAL FIN RAY COUNTS AND VERTEBRAL COUNTS IN SPECIES OF Anal fin Vertebrae 6 3 4 18 19 20 21 2 5 50 6 11 — = — 2 16 — 14 3 _ — — 33 2 14 — — — — 37 3 15 — — — 4 66 3 30 — _— = 1 35 — 5 14 1 — — 37 — 2 12 3 — 1 25 — 5 17 2 — 1 37 3 16 — — — 2 67 3 9 1 _— — 4 49 1 14 28 — — 28 21 — 14 — — Pelvics also sturdy, narrow at their base for more than half their length, then abruptly flared distally with tips of rays very fleshy in most species, so much so in adults of some species that fin resembles a small paddle. Pelvic fin count always I,5; rays unbranched. Caudal fin with rounded margin and 9 rays, at least 6 of which are branched; formula either 1,7,i or 1,6,i1. Teeth. Teeth conical, retrorse, villiform, in bands on jaws, in broad plates on tongue, and in smaller patches on prevomer and palatines. Gill rakers in the form of oval plates, variable in size but always bearing teeth similar to those on jaws and in buccal cavity. Patches of slightly larger teeth on second and third pharyngeobran- chials, these patches always well developed FREQUENCY DISTRIBUTION OF PAIRS OF PECTORAL FIN RAY COUNTS IN SPECIES OF Ogcocephalus. Paired pectoral ray counts TABLE 3. +6 tt tt +3 a) notatus —_ — _— _ — darwini — —_— —_— — — porrectus —_ ~ — —_— — vespertilio — _ — -- a nasutus (South Amer.) — — 1 — 3 nasutus (Cent. Amer.) — — 1 1 17 nasutus (W. Indies) — — — 1 24 cubifrons — — — 1 24 pantostictus oo — I 3 17 rostellum — — — 1 2 corniger 2 5 25 2 3 parvus (northern) 23 6 12 — — parvus (southern) ~ — 11 3 1 declivirostris 2 4 41 2 2 pumilus — l 30 8 9 8 | Ne nuh nvn hn Le ce) iG Se 43 4 other x Do) Sars inte B 13.4 a age 8 6hagnle ae 14.6 SF ee za 14.0 4 5 28 4 4 — 14.0 200 6 se ee ae 13.1 yi Sisk. Gate eg ee = 12.1 ieee! * at 12.3 AA ei) Mabie, ean HA 12.7 21 — 1 — — — 1225 32 3 3 — — — 13.0 ae Cle (see cs 11.0 ee eee ee a 10.4 6 = i ee = 11.2 a ee ee em * 11.0 — = aS — a= = 11.3 BRADBURY: FISH GENUS OGCOCEPHALUS FREQUENCY DISTRIBUTION OF LATERAL-LINE SCALE COUNTS IN SPECIES OF Ogcocephalus. Counts were attempted on both right and left sides of specimens, but counts which were not repeated within two subsequent attempts were discarded. TABLE 4. Lateral-line scale counts 39 16 15 54 26 53 notatus darwini nN porrectus 31.8 62 vespertilio nasutus 30.8 19 17 nN nN 114 32.5 14 10 iN pantostictus cubifrons rostellum corniger N 71 16 10 73 44 37 parvus 16 16 59 declivirostris pumilus nN 10 10 nN 34 Nm Los) ~) even in species in which gill rakers are smallest (Bradbury 1967). Integument. Most species with dermal cirri, but these variable among individuals of the same species; also, cirri are probably fragile and most likely do not preserve uniformly well, so their variability is difficult to assess. In any event, when they appear in preserved materials, they may occur singly or in clusters or thick fringes, most often in association with lateral-line sys- tem, particularly on face, disk margin, and lat- eral sides of tail. Scales consisting of close-set tubercles and bucklers, their bases sometimes slightly overlapping, forming a heavy armor. Large bucklers in dense groups over cranium and face and around disk margins, where they may be ankylosed to underlying bone. Heavy bucklers also on dorsal surface of trunk and tail where they form a median band which varies in width, extending in some species down sides of tail to lateral line. All species with large bucklers scattered elsewhere over dorsal surface of disk and pectoral peduncles with tubercles and small bucklers interspersed among them. Tubercles always invest skin covering surface of eyeballs between cornea and edges of bony orbits. Ven- tral surface of disk invariably covered with fine tubercles so that skin looks and feels like sha- green. Ventral surface of tail similarly covered, but there may be a short median line or a scat- tering of small bucklers. Dorsal and pectoral fins may or may not have minute tubercles running out along rays, but the fleshy anal and pelvic fins always have fine tubercles covering at least basal 14 or %4 of fin, except in O. notatus in which fins are nearly naked. Caudal fin with thick bands of small bucklers and tubercles run- ning out along rays, but in alternating fashion so that rays 1, 3, 5, etc., have their thickest cov- ering on one side of the fin while alternate rays have their thickest covering on opposite side, permitting fin to be tightly folded in spite of its dense armor. Color in preservative. Most species with markings on ‘‘wings’’ of disk and in axillae in the form of clusters of spots. Less commonly, tracts of spots or reticula occur along lateral walls of tail, and in Ogcocephalus porrectus and O. darwini the disk and tail markings have co- alesced into a pair of prominent stripes. Color of dorsal and pectoral fins variable from species to species, but color of anal and pelvic fins gen- erally the same as color of ventral surface of PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 7 FREQUENCY DISTRIBUTION OF PAIRS OF CHEEK LATERAL-LINE SCALE COUNTS IN SPECIES OF Ogcocephalus. Paired cheek lateral-line scale counts 238 TABLE 5. N ; f 4 notatus 43 = — — darwini 24 1 4 1S porrectus 19 — _- 18 vespertilio 34 — — —_ nasutus (South Amer.) 38 _- — — nasutus (Cent. Amer.) 20 = — 4 nasutus (W. Indies) 37 — — 8 cubifrons 50 — 3 31 pantostictus 39 —- 1 2 rostellum 25 — — 19 corniger 26 -= 1 22 parvus (northern) 35 1 6 24 parvus (southern) 15 — 1 11 declivirostris 42 — — 36 pumilus 32 — — 23 body. Ventral surface of body usually much pal- e~ than ground color of dorsal surface. Dorsal and pectoral fins may have spotted or reticulate pattern, but anal and pelvics seldom have any discrete spotting or banding. Caudal fin mark- ings nearly constant throughout genus, consist- ing of a dark basal third about the same intensity as ground color of dorsal surface of body, a pale middle third, and a dark distal third. Spots sometimes superimposed on this basic pattern in a few species. Color in life. Observations of life colors in these fishes suggest that, as a group, they TABLE 6. cephalus. 5 10 10 ) tN = ASonuan tA wr vy a are remarkably colorful if one considers how drab they become in preservative. In most cases the underparts are some shade of red: reddish orange, or crimson, or even purplish red, with fins, lips, and buccal membranes likely to be col- ored a shade of red as well. One relatively deep- water form, O. pumilus, is pinkish colored when freshly caught. Life colors are known from too few species, however, to be able to generalize yet about life colors in these fishes. DISTRIBUTION.—Ogcocephalus is confined to the West Indian—Panamanian fauna, with ten species in the western Atlantic Ocean and two FREQUENCY DISTRIBUTION OF PAIRS OF SUBOPERCULAR LATERAL-LINE SCALE COUNTS IN SPECIES OF Ogco- Paired subopercular lateral-line scale counts N $ 3 3 é + t 5 3 3 3 to te dt other x notatus 4 —- —- => —S — 3 4 3 8 19 3 — — $,5,5,7%9 8.5 darwini 21 l 1 — i 3 4 2 l 1Lo—> => SS = g 6.5 porrectus 31 — — I) 8 I 1 —- ~— ~ ~~ ~~ — — 6.1 vespertilio 0 — — —- —- — — 4 6 8 18 5 —- — 75.5, 10, 10 8.6 nasutus (South Amer.) a 2 3 1 7 2 9 6 4 — — $,8,5,5 8.0 nasutus (Cent. Amer.) a I l 2 4 2 3 3 l 2— — $5.9 7.6 nasutus (W. Indies) i 4 6 610 7 2 2 l l 1 — $,8 We. cubifrons 57 AG = - -—- = 8,355, 10 7.5 pantostictus 84 — —- — —- =— — — 3 Sie? 7 3 l $.osa eats 9.1 rostellum a 3 9 12 he 1 —- — — = 35.85 6.6 corniger 33 ————— 6 12 10 3 l = => SS SS SS = 3 6.2 parvus (northern) 35 3 10 15 4 1 — ee 3 5.4 parvus (southern) i) 2 8 2 3s SS ee Sa — 5.1 declivirostris 38 — = 2 8 vy il} 4 20 SS Se iS 3,3 6.7 pumilus 30 I 4 Sis 4 —- — — — 5.8 BRADBURY: FISH GENUS OGCOCEPHALUS 239 TABLE 7. MORPHOMETRIC PROPORTIONS FOR SPECIES OF Ogcocephalus. Body measurements are given as thousandths of standard length. Length Jaw to anus Snout to dorsal of disk margin Width of urosome N Range x N Range x N Range x N Range 5 notatus 29 ~«.549-.796 .585 22 .633-.680 .657 50 .365-.438 .403 36 .191-.318 .268 darwini 28 .547—.836 .703 28 .589-.668 .639 29 .328-.456 .385 28 .155—.244 .210 porrectus 31) AS 716 -585e Ole -615—.699) 647" 935) 419=/539) 4508 35) 1892277) 243 vespertilio 37. —-.542-.617 .586 37 ~=.635-.733 .670 46 .414-.490 .456 37 .180-.282 .231 nasutus (South Amer.) 11 .520-.618 .564 14 .611-.682 .643 40 .419-.506 .464 40 .185-.271 .236 nasutus (Cent. Amer.) 14 .535-.625 .571 25. —-.415-.482 446 7 = .207-.241 = .221 nasutus (W. Indies) 8 .550-.588 .566 9 .626-.684 .656 40 .415-.508 .465 26 .160-.262 .228 pantostictus 5) 929=574) £558 5 .646-.671 .657 48 .420-.483 .441 25 .196-.281 .247 cubifrons 16 = 543—:62)1 See 1G) ee G1S— 6847 96538) 54) S96 48284385) Si 169= 2945 237 rostellum 20 =.528-.608 .551 20 .624-.686 .651 71 .405-.493 .448 19 .211-.282 .249 corniger 32. -.545-.641 S91 33 .641-.732 .680 37 .388-.481 .441 33 .192-.258 .228 parvus (northern) 18 .480-.652 .603 17 .646-.704 .674 40 .452-.524 .486 29 .183-.284 .241 parvus (southern) 7 .610-.636 = .621 8 .644-.691 .666 15 .453-.501 .479 13 .199-.280 .245 declivirostris 17 .540-.601 .564 16 .605-.679 .635 56 .390-.447 .419 29 .193-.342 .264 pumilus 29 .523-.588 .547 29 .611-.844 .635 49 .398-.463 .429 29 .204-.264 .235 Head depth Width of cranium Length of rostrum Width of mouth N Range x N Range x N Range x N Range x notatus 49 .198—.260 .223 53 .121-.185 .146 50 .0S0-.195 .079 53 .172-.213 .196 darwini 291 96= 29 1223) 29) 033155 123, 29) > 20883181 128) 929) 106=21387- = 2 porrectus B52 0/25 Oe ees SOO 207 = slain) 32) eelSi— 164) 149 ee 35127193) S9 vespertilio 35224255259) 46) 2104157, © 2140) 46 3169=3293) 218" es) 1384=9165) 152 nasutus (South Amer.) 41. .213-.282 .234 41 .127-.172 .142 39 .079-.230 .135 41 .134-.177 .149 nasutus (Cent. Amer.) 24 .212-.262 .240 25 .110-.150 .135 25 .068-.258 .136 24 .122-.162 .144 nasutus (W. Indies) 345 7-208=262) 229" 40) 18—176"" 1383" 37 1083—229" 1827 34" 2120=5164" 137 pantostictus 36 )=. .200-.250 36.223 48—Siw«119-.155 = 1360S 47s «067—.170 =.104 «40S 130-.168 .142 cubifrons Aji 206-2 58u8 225) 52) 19= 177, 4 Sl O75=.224) I 4 097214 ei, rostellum SSeS — 254.2045 eal) O5—.165)) 2130) 969) 2053—5123) Fe 0735 525 100-3147 2 ils corniger S728 — 290 25 38) L217) 13659 36) a189— 267822855 37, 180-1835 eal 52 parvus (northern) Ales 8-5 Olen Omen 4 le S6—"81 800m loan 40 eee 07415 2a Onn [eee 09914928 parvus (southern) 150 -241=2270) 2253) 15) 2133=.158 +2146) 15S (094-136 118 S15 {095=120) ~-106 declivirostris SOn lS = 244 e285 56) 2107= 1570" 2183) S67 029=.131 92077 S60 -i7= Si 2133 pumilus AD 2 OS — 24455 2225649) ee 123= 155) 21379 49)” 21B5— 219) 1845 49) 1255S 44 Width Depth of Length of jaw Width of eye of interorbital caudal peduncle N Range x N Range % N Range x N Range x notatus 52087124) 538) 2072=.112) 2086) 53) 2046—.090) 061 52) 20690947082 darwini 28 .081-.110 .090 29 .081-.103 .090 29 .050-.080 .064 28 .066-.090 .079 porrectus 35. .087-.117 .098 35 .078-.105 .088 35 .055-.090 .064 34 .065-.083 .076 vespertilio 46 .095-.113 .106 46 .076-.106 .095 46 .060-.093 .071 46 .069-.094 .080 nasutus (South Amer.) 41 .091-.121 .101 40 .079-.118 .091 41 .054-.074 .064 40 .070-.090 .081 nasutus (Cent. Amer.) 25 .096-.112 .104 25 .070-.110 .094 25 .056-.077 .064 8 .081-.089 — .085 nasutus (W. Indies) 40 .087-.115 .098 40 .076-.130 .088 40 .053-.097 .067 35 .079-.122 .089 pantostictus 48 .085-.110 .096 43 .068-.106 .087 48 .054-.080 .066 47 .077-.096 .086 cubifrons 53 .069-.100 .081 36 .074-.116 .091 53 .062-.092 .075 43 .078-.105 .091 rostellum 71 .070-.107 .080 69 .073-.109 .085 71 .044-.083 .059 59 .055-.088 .074 corniger 38 =-.093-.125 .104 38 8 .082-.124 .101 38 .044-.082 .064 38 .061-.087 .076 parvus (northern) 41 .082-.102 .091 41 .098-.134 .116 41 .042-.076 .056 40 .075-.102 .083 parvus (southern) Si 0710842078 15 2091=. 116 2103915. 055=.073 062) 5) 0780952087, declivirostris 56 ©.080-.108 .092 56 .081-.107 .095 56 .039-.070 .050 55 .077-.100 .086 pumilus 49 .070-.111 .093 49 .094-.128 .106 49 .037-.054 .044 49 .070-.088 .079 240 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 7 TABLE 7. Cont. Width Width between Length Length of suborbital lateral ethmoids Length of snout of dorsal fin of anal fin N Range x N Range x N Range x N Range x N Range i notatus 35. .067-.112 .087 36 .086-.110 .096 36 .068-.097 .080 48 .105-.219 .146 48 .114-.180 .150 darwini 29 .065-.096 .084 29 .057-.088 .071 29 .059-.083 .066 26 .087-.121 .105 25 .144-.191 .162 porrectus 34 .085-.103 .093 34 .073-.092 .080 34 .049-.079 .070 33 .087-.121 .102 34 .150-.190 .171 vespertilio 46 .089-.114 .103 37 .064-.084 .077 37 .065-.082 .074 34 .099-.152 .123 35 .147-.183 .167 nasutus (South Amer.) 31 .084-.123 .097 32 .069-.100 .083 31 .063-.084 .073 31 .081-.147 .108 31 .114-.178 .146 nasutus (Cent. Amer.) 8 .099-.116 .104 21 .073-.096 .085 7 .076-.100 .083 7 .095-.119 .109 7 .140-.172 .156 nasutus (W. Indies) 40 .087-.117 .103 34 .074-.103 .083 34 .065-.083 .074 25 .089-.153 .111 30 .123-.193 .160 pantostictus 31 .084-.112 .097 25 .076-.095 .084 24 .065-.084 .074 41 .088-.160 .120 36 .133-.191 .164 cubifrons 35 .085-.110 .096 29 .072—.096 .079 28 .061-.097 .073 41 .097-.159 .126 29 .152—.204 .170 rostellum 38 .070-.097 .085 44 .065-.099 .076 20 .057-.077 .065 19 .088-.141 .103 17 .122-.169 .147 corniger 33 .088-.112 .101 33 .069-.110 .081 33 .064-.084 .073 37 .072-.125 .091 38 .124—.177 .150 parvus (northern) 29 .110-.136 .120 29 .082-.117 .097 29 .078-.098 .089 36 .109-.175 .145 39 .132-.177 .158 parvus (southern) 13 .091-.110 .101 13 .079-.089 .084 13 .071-.083 .079 14 .120-.176 .143 15 .143-.202 .165 declivirostris 34 .069-.105 .084 34 .068-.100 .083 34 .058-.084 .070 53 .114~.186 .150 54 .141-.187 .166 .078-.099 .087 29 .069-.094 .078 29 .060-.075 .067 43 .096-.192 .132 44 .108-.178 .150 pumilus 29 in the eastern Pacific. In the Atlantic the genus occurs along the coast of the United States from Cape Hatteras south to the Gulf of Mexico, West Indies, the coasts of Central America and South America to Uruguay. In the Pacific the genus is known from the vicinity of Cocos Is- land, the Galapagos Islands, and Peru (but not from south of the Gulf of Panama as stated by Lundy (1956) or from off Cabo Corrientes as stated by Hubbs (1958); of course, it is possible, even likely, that Ogcocephalus will be found eventually at these localities, but I know of no such records to date. I suspect the error is owed simply to a misreading of coordinates on the map of the eastern Pacific Ocean concerning the locality of the type-series of Ogcocephalus por- rectus (Garman, 1899). Relative to other genera in the family, Ogco- cephalus has the shallowest bathymetric range. Although data on depth distribution are still scanty for some species, some generalizations may be attempted. Four species appear to occur most commonly at depths shallower than 50 m; these species are O. cubifrons, O. pantostictus, O. vespertilio, and O. darwini, with cubifrons often recorded from depths of 10 m or less. Two species that have their depth distribution cen- tered at 50-60 m are O. nasutus and O. notatus, and three that have their depth distributions cen- tered at 60-100 m are O. parvus, O. pumilus, and QO. declivirostris. Ogcocephalus porrectus has been taken only between 100 and 170 m, but the species has been taken from few localities. Finally, there are two species, O. rostellum and O. corniger, that have relatively wide bathy- metric ranges from 30 to 250 m. Thus, for the genus as a whole, most specimens have been taken at depths shallower than 100 m, but some have been taken as deep as 200 m, and a few individuals have been taken from depths over 350 m. Key to Species of Ogcocephalus Few species were found to be unique in the characters studied, and constructing a key proved difficult. This key will permit identifi- cation of most specimens of juvenile and adult specimens of Ogcocephalus, but some couplets utilize characters that do not discriminate com- pletely between members of the couplet. In these cases, species intermediate between mem- BRADBURY: FISH GENUS OGCOCEPHALUS bers of a couplet are carried from there through both sides of the key. la. 1b. 2a. 2b. 3a. 3b. 4a. 4b. Sa. Distribution: western Atlantic Ocean _ 2 Distribution: eastern Pacific Ocean_._.__ 13 Pectoral rays 10/10 to 11/11 or 11/12 3% of specimens belonging to species in- cluded here have pectoral rays 12/12) _ Pectoral rays 12/12 to 15/15 (2% of spec- imens belonging to species included here have pectoral rays 11/12 or 11/11) Rostrum very long, length 0.9-1.6 in head depth, 1.6—3.1 in length of disk PCT Se ee eee En a oe een 4 Rostrum moderate to short, length 1.7- 4.0 in head depth, 3.2-8.5 in length of etes Kem Lee Ws Pe hs ee 5 Anal rays 3-4. Body pale tan or gray without markings except dark spots oc- casionally present dorsally on either side of disk. No fleshy pads on ventral sur- face of pectoral rays; fin membrane rel- atively thin and translucent (Fig. 4a). A small species reaching 70 mm SL. Known from northern Bahamas through Puerto Rico and Lesser Antilles to Sur- inam. One record from Honduras _______- Ogcocephalus pumilus new species (Fig. 26), p. 272 Anal rays 4. Dorsal body surface brown- ish, evenly covered with small whitish spots (which are the tips of bucklers). Distal ends of pectoral rays with thick- ened fleshy pads on their ventral sur- faces; fin membrane thick and opaque (Fig. 4b). A moderate-sized species reaching 140 mm SL. Known from east- ern Cuba, eastern Gulf of Mexico, and eastern coast of United States as far north as Cape Lookout, North Carolina Ogcocephalus corniger new species (Fig. 26), p. 274 Rostrum short, usually tilted upward; never sloping downward. Body surface rough, contours craggy with prominent bucklers. Head raised relatively high above disk, head depth 3.1—4.2 in stan- dard length. Mouth small, its width 2.0- 2.8 in head depth. Distal ends of pectoral 241 FIGURE 4. (a) Ventral view of pectoral fin of type found in O. notatus, O. pumilus, and O. declivirostris. Skin rela- tively thin, membranes translucent, and articulations of fin rays visible through skin. (b) Ventral view of pectoral fin of type found in all other species. Skin thick, spongy along the rays, especially distally where small pads form. Membranes thick and elastic. (These features often undeveloped in small individuals.) 5b. 6a. 6b. rays with thickened fleshy pads on ven- tral surfaces; fin membrane thick and opaque (Fig. 4b). A small species reach- ing 85 mm SL. Western Atlantic from South Carolina to Recife, Brazil, except absent from western Gulf of Mexico ___ Ogcocephalus parvus (Fig. 28), p. 276 Rostrum horizontal or sloping down- ward relative to long axis of body. Buck- lers giving body a roughened surface, but not craggy contours. Head relatively low to the disk, head depth 4.3-5.1 in standard length. Mouth width 1.4—-1.9 in head depth. Distal ends of pectoral rays without fleshy pads; fin membrane thin and translucent (Fig. 4a). A moderate- sized species reaching 140 mm SL. Known only from northern and western Gulf of Mexico Ogcocephalus declivirostris new species (Fig. 24), p. 269 Mouth wide, its width 1.8—2.5 in length of disk margin, 1.0-1.4 in head depth. Lateral-line scales in premaxillary series 4/4, in cheek series usually 10/10 (Fig. 1b). Pupil oval; no opercula. Atlantic coast of South America from Caribbean coast of Venezuela to mouth of Rio Plata in Uruguay Ogcocephalus notatus (Fig. 5), p. 243 Mouth width 2.6 or more in length of 242 TE 7b. 8a. 8b. 9a. 9b. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 7 disk margin, 1.4 or more in head depth. Lateral-line scales in premaxillary series 3/3, in cheek series 8/8 to 9/9 or 9/10, rarely 10/10 (Fig. 16). Pupil kidney- or dumbbell-shaped because of presence of opercula (flaps of iris tissue) encroaching OMG PU pile. 22 Se a Rostrum very long, length 1.4—3.1 in length of disk margin, 0.8-1.7 in head Gepthe. 5 OO |e © 30 _ a oO ° + ! oO e.e? 5 fo) oo ae ‘ = a Se | fe) oD So 20 “eo - 1 t = = oF . | a. 2. ° | °— roste/lum a ee Be °* | | o - cub/frons fe) ¢ | =: | o° | | | Te) 20 30 40 50 60 70 80 90 100 Length of disk margin in mm. FiGureE 20. Comparison between Ogcocephalus rostellum and O. cubifrons to show that depth of head is greatest in O. cubifrons relative to length of disk margin. (Table 1) and distinguished by the combined fea- tures of a bright pattern of dark spots on the pale pectoral fins, a small mouth (Fig. 6), a wide interorbital distance, and a thick, massive head on which, in large individuals, is perched a small, knoblike rostrum, because the rostrum is characterized by ontogenetic allometry: very long in juveniles, very short in large adults (Fig. 22): In general appearance, O. cubifrons most closely resembles nasutus, rostellum, and pan- tostictus, especially the latter, which is also large and heavy. O. cubifrons is distinguished from pantostictus by its smaller mouth (Fig. 19) and lower count of lateral-line scales in the sub- opercular series (usually 7-8 in cubifrons, mod- ally 9 in pantostictus, Table 6). O. cubifrons is distinguished from both pantostictus and nasu- tus by the lower number of lateral-line scales in the cheek series (modally 8 in cubifrons, 9 in pantostictus and nasutus, Table 5), and is fur- ther distinguished from nasutus by having a shorter jaw and wider interdrbital (Figs. 14 and 15) and in having allometric growth of the ros- trum (Fig. 22). Although the ranges of cubifrons and rostel- lum overlap off the southeastern United States, the two evidently seldom co-occur, cubifrons being taken in shallower water than rostellum. O. rostellum has a more flattened disk as ex- pressed by the head height (Fig. 20), a less mas- sive head as expressed by the rostrum measure- ment (Fig. 9), and a smaller caudal peduncle (Fig. 21); it is a smaller species than cubifrons, reaching only 160 mm SL. Ogcocephalus parvus and corniger occur off both coasts of Florida and off the Carolinas and Yucatan, as does cubifrons, but parvus and cor- niger are taken in deeper water than cubifrons. Both parvus and corniger have fewer pectoral fin rays than cubifrons (usually 12-13 in cubi- frons, 10-11 in parvus and corniger, Table 3) and differing color patterns; parvus, a small species reaching only 85 mm SL, has blotches of varying tones as the color pattern on the dor- sal surface of the body, but it lacks the promi- nent clusters of dark circles seen on the body and pectoral fins in cubifrons. O. corniger dif- fers from cubifrons in having close-set pale spots on a dark background evenly covering the entire dorsal surface of the body. Ogcocephalus declivirostris is restricted to the northwestern Gulf of Mexico and is evi- dently allopatric with cubifrons. O. decliviros- 262 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 7 Depth of caudal peduncle in mm. e — roste//um 20 30 40 o-— cubifrons 50 60 70 Length of disk margin in mm. FIGURE 21. greatest in O. cubifrons relative to length of disk margin. tris 1s distinguished, in any case, in having fewer pectoral fin rays (modally 11 in declivirostris, usually 12-13 in cubifrons, Table 3); decliviros- tris lacks the fleshy ventral pads on the distal tips of the pectoral fin rays as well as the spotted pattern of the pectoral fins so prominent in cub- ifrons (although occasionally declivirostris has a few very pale spots on the pectorals). Finally, declivirostris differs from cubifrons in having its rostrum jutting horizontally or downwards in- stead of upwards as in cubifrons. DESCRIPTION.—Counts and measurements from 95 specimens 29.9 to 224.0 mm SL (Table 1 Counts. Counts given in Tables 2-6. Two counts for cubifrons lie in the middle of the range for the genus; pectoral fin ray count is generally 12-13 in cubifrons, which is the mid- dle of the generic range of 10-15, and the sub- opercular lateral-line scale count in cubifrons is usually 7-8, which is the middle of the generic range (excluding extremes) of 5—10. The lateral- line scale count, although showing wide varia- tion, is also near the middle of the range for the Comparison between Ogcocephalus rostellum and O. cubifrons to show that depth of caudal peduncle is genus (Table 4). However, cubifrons agrees with pantostictus, rostellum, and declivirostris in having a high modal vertebral count of 20, whereas the more usual count for the genus is 19. Proportions. Proportions expressed as ratios given as the mean followed by the range in pa- rentheses. Length of disk margin intermediate within range of variation for the genus, 2.3(2.0—2.5) in SL. Width of tail also intermediate, varying from thin in smallest specimens to thick and broad in large specimens, 1.8(1.5—2.4) in length of disk margin; caudal peduncle thick and deep, its depth 2.5(2.2-2.9) in head depth. Rostrum long, conical, and upturned in small specimens, be- coming relatively small and knoblike in large ones (Fig. 22); length of rostrum 4.3(1.8—6.1) in length of disk margin. Aperture of illicial cavity subtriangular in small specimens, much higher than wide, becoming merely oval or round in large ones. Cranium rising steeply above the disk, its contours becoming massive and boxlike in adults; head depth 2.0(1.7-2.2) in length of BRADBURY: FISH GENUS OGCOCEPHALUS 135 mm 304 mm FiGuRE 22. Heads of four examples of Ogcocephalus cub- ifrons showing allometric growth of rostrum. disk margin. Width of cranium 3.1(2.6—3.6) in length of disk margin. The following in width of cranium: eye 1.6(1.4—1.9), lateral ethmoid width 1.8(1.5—2.0). Interorbital very wide, slightly concave, 1.9(1.5-2.2) in width of cranium, 3.0(2.3-3.4) in head depth. Mouth small, its width 1.9(1.6—2.2) in head depth, 3.8(3.2—4.4) in length of disk margin; jaw 2.8(2.5—3.2) in head depth. Upper lip fleshy, of an even width for its entire length; lower lip fleshy, usually thickened medially although no median lobe is developed. Dorsal fin 3.4(2.7—4.8) in length of disk margin; anal fin 2.6(2.2—3.1) in length of disk margin. Integument. Dermal cirri generally well de- veloped on lateral sides of tail, less so on disk margin; cirri also frequently scattered thickly on dorsal surface of body, particularly in half- grown specimens. Large bucklers prominent, arranged as in generic description. Tubercles relatively sparse on fins, occurring at bases of dorsal and pectoral fins with only a few scattered along fin rays. Anal and ventrals with tubercles at bases and running out on fin rays for a third to a half their length. Caudal as in generic de- scription. Skin covering eyeballs bearing tiny tubercles with rim of cornea beaded by a row of small bucklers. Pectoral membranes thick, opaque, the ventral surface of ray tips with well- developed fleshy pads. Color in preservative. Ground color of dorsal surface of body light tan to dark brown or gray with black spots sprinkled on face, on skin cov- ering eyeballs, on shoulders and lateral sides of tail, in the axillae, on pectoral fins, and usually on lateral margins of disk. Spotted pattern on shoulders does not form a reticulum as it does in O. vespertilio. Shoulder spots often with pale centers in small specimens. Ground color of pectoral fins and sides of tail usually pale with dark spots so close together that the background appears as a reticulum; even in specimens so dark as to obscure most of the spotted pattern of body, the pattern on pectoral fins remains prominent. Pectoral fins with a broad dark brown or black border in medium-sized and small specimens. Lips pale; rostrum with a dif- fuse black subterminal ring; iris black, or golden with black spots, in which case color pattern of face extends over eyes to margin of pupil. Ven- tral surface of body uniformly pale or creamy. Ventral and anal fins the same ground color as ventral surface of body, ventrals often with a pale patch distally in dark specimens, anal fin sometimes dark or at least tipped with brown or black in small specimens. Ventral pads of pec- toral fin rays white or pale. Dorsal fin spotted or sometimes with merely vague blotches. Caudal fin as in generic description but generally with spotted pattern of tail carried back and super- imposed on caudal’s basic pattern. Color in life. | have not seen this species alive. The following is a description by Longley (Longley and Hildebrand 1941), ‘*The young are dark gray on the under side; older fish coppery red, with much individual variation in color on the dorsal side, usually lighter or darker brown, variegated with reddish orange or yellow-orange patches, disappearing in preserved specimens; the chief patches behind the eyes, the two some- times meeting across the nape to form a great U-shaped mark open to the rear; usually an ad- ditional one above and behind the branchial ap- erture; and one about base of soft dorsal fin. Even at a standard length of 50 mm. there is some, and in fish of medium and large size much spotting with light-bordered dark spots in a long, narrow tract from eye to gill opening. Here the spots usually are less crowded than elsewhere, and more nearly circular. On the sides of the head, margin of the disk, and sides of the trunk, the pattern is less open, being principally a net- work of light lines on the dark ground. Small fish have pectorals and caudal yellow proximally and dark brown distally, with a netted pattern appearing first at the base of these fins and ex- tending gradually to their tips, the yellow ad- vancing before it at the expense of the darker brown, which gives way to the lighter color on which the marbling spreads.” 264 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 7 A photograph in Walls (1975:263) shows a specimen (his O. radiatus) with margins of pec- torals bright yellow; the size of the fish is not given, but the small relative size of the rostrum indicates that the specimen is probably a large adult. A smaller specimen is shown in color in Herald (1972). In this specimen the yellow color lies just proximal to the dark brown fin margins about as Longley described (Longley and Hil- debrand 1941). Both photographs show the ‘‘netted’’ pattern on the base of the pectorals. DISTRIBUTION (Fig. 17).—Northern Bahamas, coast of the southeastern United States from the vicinity of Cape Lookout, North Carolina, around the Florida peninsula to Pensacola, Flor- ida, and south to Campeche Banks, Yucatan. Bathymetric range: shore to 68 m. MATERIAL EXAMINED.—BMNH 1855.9.19.848, the holo- type, said to be from Labrador, but this undoubtedly an error (see p. 260). For following material, numbers in parentheses are numbers of specimens. For data of OREGON stations (= O), refer to Springer and Bullis (1956). United States: North Carolina: UF 24912 (1), near Cape Lookout Bight, Carteret Co. South Carolina: UF 24903 (1), 31°58’N, 79°43'W, about 77.3 km se Edisto Beach, Beaufort Co. Georgia: UF 25931 (1), off Brunswick. Florida: Dade County: FMNH 46128 (1), inshore reef between Miami and Angelfish Creek; FMNH 46854 (1), Triumph Reef off Elliot Key; USNM 89710-89711 (2), Biscayne Bay near Cocoanut Grove, Miami. Monroe County: ANSP 90667 (1), E end Grassy Key, Florida Bay; CAS 18585 (1), near Key West; FMNH 7020 (1), Key West; FMNH 21645-21647 (3), near Marathon; FMNH 64123 (1), O-1004; FSBC 342 (4), FSBC 381 (1), FSBC 410 (1) all from 24°41'—24°52'N, 82°00’— 82°3S'W: FSBC 626 (1) and FSBC 762 (1), both from 24°43'10"—24°52'00"N, 82°00'-82°35'W: UF 4071 (1), Marque- sas Keys; UF 7022 (1), Big Pine Key; UMML 1201 (10), 24°45’—24°50'N, 82°10'-82°30'W; UMML 4786 (1), 5.3 km sw of Greyhound Key; USNM 57243 (1), Matecumbe Key; USNM 84035 (1), 24°36’N, 81°48’W; USNM 117003 (7), Tor- tugas; USNM 117006 (3), channel w of White Shoal, Tortugas; USNM 117170 (2), Tortugas; USNM 126488 (2), Key West. Collier County: FSBC 1157 (1), Everglades. Lee County: CAS-SU 5841 (1), Captiva Pass; CAS-SU 36404 (6), Sanibel I.; FMNH 510, 512, and 513 (3), Captiva Pass; FMNH 38584— 38586 (3) and FMNH 38587 (1), Boca Grande; USNM 76025 (1), Boca Grande Pass; USNM 86159 (1), Boca Grande. Hills- borough County: USNM 143160 (2), Port Tampa. Pinellas County: FSBC 61 (1), Blind Pass, St. Petersburg Beach; FSBC 878 (1), John’s Pass, Madeira Beach; FSBC 1369 (1), due w of Don-ce-Sar; USNM 158543 (1), Bay at Clearwater. Levy County: the following all from Cedar Key: ANSP 90668 (2); CAS-SU 99 (5); UF 229 (1); UF 753 (1); UF 916 (1); UF 919 (1); UF 1522 (1); UF 1612 (1); UF 3169 (1); UF 3374 (1); UF 7832 (1); UF (1), R-I-650-4; UF (1), R-I-651-3; UF (2), R-I- 2351-1; UF (1), R-II-551-6; UF (1), R-II-2051-4; UF (1), R-III- 451-1; UF (1), R-III-451-4; UF (2), R-IV-551-1; UF (1), R-IV- 551-7; UF (2), R-VI-650-3; UF (4), R-VI-750-2; UF (4), R-VI- 1750-4; UF (1), R-VII-150-5; UF (4), R-VII-150-6; UF (6), R- VIII-2650-1; UF (4), R-VIII-2650-1; UF (2), R-VIII-2650-2: UF (1), R-IX-1250-1; UF (1), R-IX-1250-1; UF (2), R-X-850- 7; UF (1), R-X-2250-5-5; UF (1), R-XI-750-6; UF (1), R-XI- 750-7; UF (1), R-XII-1050-1; UF (2); R-XII-1050-2; USNM 88499 (1); USNM 164928 (1). Franklin County: CAS 23914 (1), Apalachicola Bay. Okaloosa County: UF 5586 (1), UF 5587 (1), UF 5596 (1), all from Choctawatchee Bay at Destin; UF 2730 (1), Pensacola. Florida Straits: UF 28649 (1), SILVER Bay sta. 144, 27°44'N, 82°46’ W, 7.3 m. Bahamas: ANSP 103627 (1), GEORGE M. Bowers sta. 659, w side Eleuthra I.; ANSP (3), Chaplin Program sta. 513B, Hatchet Bay, Eleuthra I.; ANSP (2), Chaplin Program sta. 526, Hatchet Bay, 4.8 km offshore, Eleuthra I.; UMML 4719 (1), Exuma Sound adjacent to Eleuthra I., 7.3 m. Mexico: Campeche Banks: FMNH 46745 (5), O-436; FMNH 64107 (8), O-713-716; FMNH 64134 (3), O-710; UF 28647 (1), SILVER Bay sta. 362, 20°00'N, 91°43'W, 44 m; UF 28648 (1), O-417. Yucatan: USNM 148497 (1). Undetermined material. Seven specimens from the Bahama Islands are intermediate in color pattern between cubifrons and nasutus, and also cannot be diagnosed with certainty on the basis of morphometric characters. These specimens are: ANSP 72335 (1), Hog I.; ANSP 90670-90671 (2), s shore of Hog I.; and FMNH 64139 (4), Grand Bahama I. Ogcocephalus pantostictus new species (Figure 23) Ogcocephalus vespertilio (non Linnaeus): Woops 1942:192 {off Corpus Christi, Texas]. Ogcocephalus cubifrons (non Richardson): HILDEBRAND 1954:321 [off Texas]. Ogcocephalus radiatus: ANONYMOUS 1976:131 and 427 [Mex- ican coast of Gulf of Mexico; color photograph; key]; HOESE AND Moore 1977:144, fig. 101 [Nw Gulf of Mexico; key; partial description; range]. DIAGNOSIS AND COMPARISONS.—A very large Ogcocephalus (Table 1) with the entire dorsal surface of the body covered with a polkadotlike pattern of dark spots (sometimes dim in museum specimens). Resembling cubifrons in many fea- tures, it has allometric growth of the rostrum: a relatively long rostrum in small specimens, but short and sometimes merely knoblike in large specimens. O. pantostictus is distinguished from cubifrons by its wider mouth (Fig. 19; also expressed by the jaw measurement, Fig. 14) and in having higher modal counts of lateral-line scales in the cheek series (9 in pantostictus, 8 in cubifrons, Table 5) and in the subopercular series (9 in pantostictus, 8 in cubifrons, Table 6). With its range restricted to the northern and western Gulf of Mexico, pantostictus is largely sympatric with declivirostris, but the two are distinguished by the pectoral fin ray count (usu- ally 12-13 in pantostictus and 11 in decliviros- tris, Table 3) and the lateral-line scale count in the subopercular series (modally 9 in pantostic- tus, 7 in declivirostris, Table 6), O. pantostictus BRADBURY: FISH GENUS OGCOCEPHALUS FIGURE 23. Ogcocephalus pantostictus new species, all from Gulf of Mexico. (A) and (D) Paratype 77.0 mm SL, CAS-SU 40559, Galveston, Texas. (B) Paratype 96.0 mm SL, FMNH 64111, off Mobile Bay, Alabama. (C) and (E) Holotype 226.0 mm SL, FMNH 50765, off Pensacola, Florida. (F) Paratype 187.0 mm SL, CAS-SU 40559, showing variation of rostrum resembling O. cubifrons. Compare with (G), O. cubifrons (same specimen as in Fig. 18C). also has the ventral surfaces of the pectoral fin rays thickened with fleshy pads which are not present in declivirostris. The species corniger and parvus have ranges slightly overlapping that of pantostictus in the northern Gulf of Mexico, but both are separable from pantostictus by their lower pectoral fin ray counts (usually 12-13 in pantostictus and 10-11 in corniger and parvus, Table 3). O. parvus is a far smaller species than pantostictus, reaching only about 85 mm SL. O. corniger has a much longer rostrum than does pantostictus (Fig. 9) and has a distinctive color pattern of pale spots on a darker background distributed evenly over the dorsal surface of the body in contrast to the darkish circles on a lighter background found in pantostictus. DESCRIPTION.—Counts and measurements from 65 specimens, 38.6 to 261.0 mm SL (Table ie Counts. Counts given in Tables 2-6; in the following description, counts for the holotype given in brackets. Pectoral fin ray counts in pan- tostictus in middle of range for genus, 12—13 in a range of 10-15 [12,12]. Subopercular lateral- line scale count, modally 9 [10,10] in pantostic- tus, high relative to range of 5—10 (excluding ex- tremes) for genus; only two other species have as high a count: vespertilio usually has 9 scales in subopercular series, notatus usually 10. The count for cheek lateral-line scales nears high end of range, too, with count in pantostictus usually 8 or 9 [9,10]; in this character pantostictus agrees with nasutus, and only vespertilio and 266 notatus have higher counts in cheek series. Lat- eral-line scale count [35,36] varies widely, but mean is highest for genus. O. pantostictus re- sembles cubifrons, rostellum, and declivirostris in vertebral count of 19-21, modally 20, in con- trast to remaining eight species which nearly al- ways have 19, occasionally 18, vertebrae. [D.4; A.4.] Proportions. Proportions expressed as ratios given as the mean followed by the range in pa- rentheses and the proportion for the holotype in brackets. Length of disk margin intermediate within range of variation for the genus, 2.3(2.1- 2.4)[2.4] in SL. Tail moderately thick and broad in large specimens, relatively narrow in small ones, its width 1.8(1.5—2.0)[1.7] in length of disk margin; caudal peduncle thick and deep, its depth 2.6(2.3—2.9)[2.5] in depth of head. Onto- genetic allometry in growth of rostrum resem- bles that in cubifrons; however, I have not seen specimens of pantostictus less than 38.6 mm in standard length and therefore do not know whether or not the remarkably long rostrum seen in very small cubifrons (30 mm SL) is also characteristic of small pantostictus. Rostrum cone-shaped in small examples, becoming rela- tively small and finger-shaped in large ones. Length of rostrum 4.3(2.6—6.3)[5.0] in length of disk margin. Aperture of illicial cavity subtrian- gular in small specimens, higher than wide, be- coming oval in adults. Cranium rising steeply above disk, becoming heavy and massive in large specimens; head depth 2.0(1.8—2.1)[2.1] in length of disk margin. Width of cranium 3.2(2.9- 3.7)[3.4] in length of disk margin. The following in width of cranium: eye 1.6(1.4—1.8)[1.8], lat- eral ethmoid width 1.6(1.4—1.8)[1.6]. Interorbital slightly concave, wide, 2.0(1.8—2.5)[1.9] in width of cranium, 3.3(2.9—4.1)[2.9] in head depth. Mouth moderate, its width 1.6(1.4—1.8)[1.6] in head depth, 3.1(2.8—3.6)[3.3] in disk saat 2.3(2.1-2.7)[2.5] in head depth. Upper| lip fairly fleshy, of an even width along its entire length; lower lip fleshy, usually thickened medially al- though no median lobe is developed. Dorsal fin 3.7(2.9-4.8)[4.8] in length of disk margin; anal fin 2.7(2.3-3.2)[3.2] in length of disk margin. Integument. Dermal cirri often present on disk margin and lateral sides of tail. Large buck- lers prominent, arranged as in generic descrip- tion. Tubercles relatively sparse on fins, occur- ring at very base in dorsal and anal fins and on about proximal third of anal and ventrals. Cau- PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 7 dal as in generic description. Skin covering eye- balls bearing tubercles with rim around cornea usually beaded by a row of small bucklers. Pec- toral membranes thick, opaque; ventral surfaces of ray tips with well-developed fleshy pads. Color in preservative. Dorsally the ground color of body gray or tan sprinkled fairly evenly with round dark spots over entire tail and disk, including face and skin covering eyeballs. Mark- ings on shoulders, sides of tail, and in axillae generally darkest and showing greatest contrast with background color. Ventral surface uniform- ly pale, usually paler than ground color of dorsal body surface. Lips pale; rostrum with a diffuse dark subterminal ring; iris golden with black spots, or entirely black. Pectorals with same spotted pattern as dorsal surface of body. In small or medium-sized spec- imens, spotted pattern on pectorals grades into a dark border. In many individuals spots close together so that background appears as a light- colored reticulum. Ventrals and anal same color as ventral surface of body; anal sometimes dusky near tip in specimens under about 100 mm SL. Dorsal fin usually spotted, especially when spotted pattern of dorsal body surface relatively intense. Color in life. Unknown. DISTRIBUTION (Fig. 17).—Coasts of northern and western Gulf of Mexico from approximately longitude 87°W to latitude 25°N. Bathymetric range: from limited data available, about 9- 31m: NAME.—The name pantostictus means all spotted, in reference to the distinctive over-all spotted color pattern. MATERIAL EXAMINED.—HOLOTYPE: FMNH 50765, Santa Rosa Sound, Pensacola, Florida; a male 226.0 mm SL. PARATYPES. Numbers in parentheses are numbers of speci- mens. United States: Florida: ANSP 71329 (1), Pensacola; MCZ 45080 (1), OREGON sta. 1648, 30°11'N, 86°55'W, 31 m. Alabama: FMNH 64111 (1), OREGON sta. 882, 30°10'N, 87°50'W, 10 m. Mississippi: CAS 23934 (1), off Horn I.; FMNH 21636 (1), Horn I.; FMNH 46734 (1), outside Petit Bois I.; GCRL 562 (1), outside Petit Bois I.; GCRL 563 (1), sw of Ship I.; GCRL 564 (1), off Horn I.; GCRL 565 (1), off w end of Horn I.; UF uncat. (1), St-veR Bay sta. 167, 30°12'N, 88°40’ W, 7.3 m. Louisiana: FMNH 50997 (2): GCRL 566 (1), s of Grand Isle, 17 m; USNM 188755 (1), PELICAN Sta. 66-7, 28°59’N, 91°48’W. Texas: CAS 23932 (1); USNM 94443 (1); USNM 118610 (9). Vicinity of Galveston: CAS-SU 40559 (10); USNM_ 118603 (1); USNM 118604 (1); USNM 118606 (1); USNM 120071 (1); USNM 176982 (1); USNM 188803 (2), PELICAN sta. 56-5, 29°10.5'N, 94°50.5'W; USNM 188805 (1), PELICAN sta. 62-2, 29°19’N, 94°31.5'W. Vicinity of Freeport: MCZ 45079 (1), USNM 117804 (14), USNM 118605 (1), USNM 118611 (1). Southern Texas: USNM 120070 (1), BRADBURY: FISH GENUS OGCOCEPHALUS 67 FiGuRE 24. (A) and (B) Ogcocephalus rostellum new species, holotype, 133.0 mm SL, USNM 188809, off Atlantic coast of Florida, 29°10’N, 80°19’W. (C) and (D) Ogcocephalus declivirostris new species, holotype, 129.5 mm SL, FMNH 64136, Texas coast of Gulf of Mexico, 28°02'N, 96°03’W. (E) Paratype of O. declivirostris, 107.5 mm SL, FMNH 61577, Texas coast of Gulf of Mexico, 28°02'N, 96°04’W, showing pattern of dim spots sometimes seen on dorsal surface of body. Aransas Pass; FMNH 37728-37729 (2), Port Aransas; FMNH 40298 (1), within 80 km of Corpus Christi; ANSP 90675 (1), Brownsville Ship Channel, Pt. Isabel; USNM 188800 (1), Pt. Isabel. Mexico: USNM 188775 (1) and USNM 188783 (1), both from off Huts Bayon, approximately 24°30'N, 6-18 m. Material without exact localities: USNM 188756 (1), PELI- CAN sta. 37; USNM 188758 (1), PELICAN sta. 104-2; USNM 188769 (1), MC 9; USNM 188798 (1), 8.1 km offshore. Ogcocephalus rostellum new species (Figure 24A —B) Malthaea nasuta (non Cuvier and Valenciennes): DEKAY 1842:166, pl. 28, fig. 89 [synonymy; description; range, compiled]. DIAGNOSIS AND COMPARISONS.—A moderate- sized Ogcocephalus with a very abbreviated ros- trum resembling the shelflike condition seen in notatus. (In the smallest specimens of both ros- tellum and notatus, 40 mm SL or less, a short median horn is developed on the rostrum.) O. rostellum is distinguished from notatus by its narrower mouth (Fig. 6) and lower number of scales in the cheek lateral-line series (modal number 8 in rostellum, 10 in notatus, Table 5) and in the premaxillary series (3 in rostellum, 4 in notatus). In general appearance and color pattern, ros- tellum most closely resembles cubifrons; the ranges of the two species overlap along the At- 268 lantic coast of the southeastern United States, but rostellum is ordinarily taken from deeper water than cubifrons. The two are distinguished by some differences in body proportions: ros- tellum has a relatively more flattened disk as expressed by the head height measurement (Fig. 20) and a less massive head as expressed by the rostrum measurement (Fig. 10); also it has a nar- rower caudal peduncle than cubifrons (Fig. 21). O. rostellum is a smaller species than cubifrons, reaching 160.0 mm in standard length compared to 230.0 mm in cubifrons. Finally, the color pat- tern of the pectoral fin helps discriminate be- tween rostellum and cubifrons; often there are no dark spots on the pectoral fins in rostellum compared to the bright pattern of dark spots on a light background covering the pectoral fins in cubifrons, but when spots do occur on the pec- torals in rostellum, they are generally dim and also less numerous than in cubifrons and re- stricted to the basal portions of the fins. Ogcocephalus rostellum has its range adjacent to that of nasutus but is a smaller species than nasutus (Table 1) and lacks the long rostrum seen in nasutus (Fig. 10); rostellum differs fur- ther from nasutus in having a shorter jaw (Fig. 16). The range of rostellum overlaps the ranges of parvus and corniger; all occur at moderate depths on the continental shelf off the south- eastern United States. O. rostellum may be sep- arated from parvus and corniger by the pectoral fin ray count (usually 13 in rostellum, usually 10-11 in parvus and corniger, Table 3). O. ros- tellum is further distinguished from corniger in lacking a long rostrum (Fig. 9), and is further distinguished from parvus in its flatter disk with less elevated cranium (head depth 4.3-5.4 into SL in rostellum, 3.1-4.2 in parvus). DESCRIPTION.—Counts and measurements from 69 specimens, 33.3 to 154.2 mm SL (Table 1). Counts. Counts given in Tables 2-6; in the following description, counts for holotype given in brackets. Pectoral fin ray count usually 13 [13, 13], range 11-15, about intermediate within range of variation for genus. Count of lateral- line scales in subopercular series moderately low for genus, usually 6 or 7 [5,6], range 5-9. Count of cheek series of lateral-line scales 8 [8,8], the most common count for the genus; range 7-9. Lateral-line scale count, 21-32 [26, PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 7 25]. Mean for each of the counts of series of lateral-line scales in rostellum lower than in the closely similar cubifrons. O. rostellum agrees with cubifrons, pantostictus, and declivirostris in having a modal count of 20 vertebrae, com- pared to other species in genus in which modal count is 19. [D. 4; A. 4.] Proportions. Proportions expressed as ratios given as the mean followed by the range in pa- rentheses and the proportion for the holotype in brackets. Length of disk margin intermediate relative to range of variation for genus, 2.2(2.0—2.4)[2.4] in SL. Tail moderate in width, 1.8(1.6—2.1)[1.6] in length of disk margin; caudal peduncle slender, 2.7(2.4-2.9)[2.7] in head depth. Rostrum pro- duced into a short cone in small specimens, be- coming more shelflike in large specimens with the median horn extremely abbreviated, rostrum 6.1(3.7-8.0)[7.1] in length of disk margin. Ap- erture of illicial cavity almost diamond-shaped. Cranium well demarcated from disk, but head less deep than in other species in the genus, head depth 2.2(2.0—2.4)[2.2] in length of disk margin. Cranium relatively narrow, its width 3.5(2.7— 3.9)[3.6] in length of disk margin (one specimen had this ratio 4.4). The following in width of cranium: eye 1.5(1.3—1.7)[1.5], lateral ethmoid width 1.6(1.5—1.8)[1.8]. Interorbital space slight- ly concave, its width moderate, 2.2(1.9—2.7)[2.4] in width of cranium, 3.4(2.8—4.1)[3.9] in head depth. Mouth small, its width 1.7(1.5—2.0)[1.8] in head depth, 3.8(3.1—4.4)[4.0] in length of disk margin; jaw 2.6(2.3—2.8)[2.7] in head depth. Up- per lip fleshy, of an even width for its entire length; lower lip fleshy, very thickened medial- ly. Dorsal fin 4.3(3.2-4.9)[4.5] in length of disk margin; anal fin 3.0(2.5—3.5)[3.4] in length of disk margin. Integument. Dermal cirri sometimes present but never conspicuous. Large bucklers promi- nent, arranged as in generic description. In more than half the specimens studied, the ventral sur- face of tail with low bucklers making surface markedly coarse relative to smooth shagreenlike texture of ventral surface of disk. All fins with at least a few tubercles; dorsal with only a few tubercles at base, but pectoral, ventral, and anal fins with tubercles at their bases and running out along rays for a third to three-fourths their length. Caudal as in generic description. Skin covering eyeballs bearing tiny tubercles with rim BRADBURY: FISH GENUS OGCOCEPHALUS around cornea beaded by a row of small buck- lers. Pectoral membranes thick, opaque, ventral surfaces of ends of rays with fleshy pads. Color in preservative. Ground color of dorsal surface of body light tan to dark brown with round dark spots (occasionally rings) in an elon- gate cluster on either shoulder and in a band down either side of tail, each continuous with a cluster in axillae. Sometimes the same spotted pattern occurs on face. Ventral surface uniform- ly pale; lips pale. The small rostrum sometimes with a dusky tip, but no dark subterminal ring except in a few small specimens; iris golden or black. Basal portions of pectoral fins pale, oc- casionally with a few faint spots, darkening dis- tally so that frequently a distinct black border is formed. Ventrals and anal the same pale ground color as ventral surface of body, anal with a dusky spot distally in a few small speci- mens. Dorsal dusky with faint splotches or sim- ply pale. About one-half the specimens at hand have no dark markings on caudal, but remainder show pattern common for genus. Color in life. The following description from three color transparencies of a freshly killed specimen captured in winter off North Carolina; one transparency shows a dorsal, one a lateral, and one a ventral view. Ground color of dorsal surface of body me- dium brown with whitish patches where skin evidently abraded from protruding tops of buck- lers. Darker mottlings of pigment in axillae. Fine black lines radiating from apices of large buck- lers. An elongate cluster of round dark spots on either shoulder; among these spots are sprinkled a few smaller, intensely orange spots, about four on each side (cast shadows on the photographs may be obscuring other orange spots). Pectoral and caudal fins with mottled brownish basal por- tions fading to translucent white with pale tan striations on middle portions and terminating in broad, bright orangy-red borders blending to pure dark red and finally to narrow black distal margins. Face the same brownish ground color as dor- sal surface of body. Lips bright orangy red; iris a rosy-brass color with tan spots in a circle around pupil. Ventral operculum well extended in example and the same rosy-brass color as rest of iris. Ventral surface of body with a transparent wash of pinkish orange over a pale gray surface. 269 DISTRIBUTION (Fig. 17).—Atlantic coast of the United States from Cape Hatteras, North Carolina, to the Florida Keys. (One specimen from Jamaica, USNM 160732, 150.0 mm SL, has the general shape of the head and the small mouth that characterize rostellum, but it differs from rostellum in other body proportions and has no diagnostic color markings; I am unable to assign this specimen.) Bathymetric range: 28— 228 m. NAME.—The name rostellum is the diminu- tive of the word rostrum, referring to the partic- ularly small rostrum in this species. MATERIAL EXAMINED.—Numbers in parentheses are num- bers of specimens. For data from ComBaTt stations (=C) and SILVER Bay stations (=SB), refer to Bullis and Thompson (1965); for data from PELICAN stations (=P), refer to Anderson (1956). HoLotype. USNM 188809, C-336, Atlantic coast of Florida; a male 129.5 mm SL. PARATYPES. North Carolina: FMNH 64338 (1), C-384; USNM 72288 (1), Cape Lookout; USNM 111848 (1), Beaufort; USNM 132093 (1), ALBATROSS sta. 2417, 33°18'30’N, 77°07'W, 174 m; USNM 148317 (2), ALBATROSS sta. 2600, 34°39'40"N, 75°35'30"W, 159m; USNM 148320 (3), ALBATROSS sta. 2602, 34°38'30"N, 75°33'30"W, 227 m; USNM 151919 (1), ALBATROSS III sta. 35, 34°45.5-48’N, 75°28’-33'W, 77-88 m: USNM 151962 (1), ArBatross III, 31-B, sta. 14, 33°26’N, 77°03.5—06'W, 59 m: USNM 151971 (1), ALBATROss III, 31-D, sta. 3, 34°11'N, 76°06’W, 82-119 m. South Carolina: CAS 23925 (3), SB-1782; FMNH 64100 (1), Bowers sta. 50, 31°50'N, 79°45’'W, 46 m; USNM 84510 (4), ALBATROSS sta. 2312, 32°54’N, 77°53'30"W, 161 m. Georgia: ANSP 103634 (1), P-178-10; CAS-SU 62117 (2), SB-1768; FMNH 66385 (15), Bowers sta. 32, 30°14’N, 80°16’W, 73 m; UF 24189 (1), Brunswick; USNM 149959 (2); USNM_ 188810 (1), P-198-5. Florida: FMNH 66388 (2), SB-3442: MCZ 32940 (1), inlet near New Smyrna; MCZ 45088 (3), SB-3022; MCZ 45089 (1), SB- 2775; MCZ 45090 (1), P-168-2; MCZ 45091 (1), Cape Canaveral Pier; MCZ 45092 (1), SB-2771; MCZ 45093 (1), P-169-3: USNM 18031 (1), St. Augustine; USNM 38026 (1), 3.2 kms of Mosquito Inlet, Hillsborough River; USNM 131628 (2), ALBATROSS Sta. 2641, 25°11'30"N, 80°10’W, 110 m; USNM 188790 (1), P-208-8: USNM 188794 (1), P-171-2; USNM 188806 (L), P-169-8; USNM 188807 (1), C-334; USNM 188813 (2), C-336 (same haul as holotype). Florida Straits: CAS-SU 62123 (1), SB-2039; UF 24190 (1), C-455. Florida Keys: USNM 117004 (2), Tortugas; USNM 117005 (1), Tortugas. OTHER: USNM 83785-83786 (3), ALBATROSS sta. 2311 and 2318, 32°5S5'N, 77°54'W (South Carolina) and 24°25'45’N, 81°46'00’W (Florida Keys), 145 and 82 m. Ogcocephalus declivirostris new species (Figure 24C-E) Ogcocephalus sp.: HILDEBRAND 1954:322 [off Texas; partial description; stomach contents]; WALLS 1975:120 [N Gulf of Mexico; called Ogcocephalus sp. B; partial description; figs.]; HOESE AND Moore 1977:144, fig. 100 [Nw Gulf of Mexico; key; partial description; range]. 270 DIAGNOSIS AND COMPARISONS.—An Ogco- cephalus with rostrum generally sloping down- ward with respect to the long axis of the body instead of being horizontal or tilted upward. O. declivirostris differs from all its congeners ex- cept pumilus and notatus in lacking the devel- opment of fleshy pads on the ventral surfaces of the pectoral and pelvic rays. It may be separated from the long-nosed species, pumilus, corniger, and vespertilio, by the length of the rostrum (length of rostrum into length of disk margin 3.4— 8.4 in declivirostris, 1.5—3.0 in the long-nosed species, Fig. 9) and from notatus by the width of the mouth (width of mouth into length of disk margin 2.8—3.7 in declivirostris, 1.8-—2.5 in no- tatus). It is distinguished from all species except pumilus, parvus, and corniger by its low pec- toral ray count (usually 11 in declivirostris, 12 or more in other species except pumilus, par- vus, and corniger, Table 3). O. declivirostris is sympatric with O. pantostictus, both being species of the northern and western coasts of the Gulf of Mexico; besides the lower pectoral fin ray count in declivirostris, just mentioned, declivirostris has a lower count for the sub- opercular lateral-line series than pantostictus (modally 7 in declivirostris, 9 in pantostictus). The range of parvus overlaps with that of de- clivirostris in the northern Gulf of Mexico; de- clivirostris is distinguished from parvus by its flatter head (head depth into SL 4.3-5.1 in de- clivirostris, 3.1—4.2 in parvus) and larger mouth (width of mouth into head depth 1.4—-1.9 in de- clivirostris, 2.0—2.8 in parvus). DESCRIPTION.—Counts and measurements from 99 specimens, 39.6 to 137.0 mm SL (Table 1). Counts. Counts given in Tables 2-6; in follow- ing description, counts for holotype given in brackets. In declivirostris all counts except ver- tebral count at low end of range of variation within genus. Pectoral fin ray count usually 11 [11,11]; range 9-12. Subopercular lateral-line scale count usually 8 [8,9]; range 7-9. Mean for lateral-line scale count 25 [25,27]; range 22-28. Vertebral count most frequently 20; range 18— 202 (ID 252-Ae 3] Proportions. Proportions expressed as ratios given by the mean followed by the range in pa- rentheses and the proportion for the holotype in brackets. Length of disk margin intermediate relative to range of variation for genus, 2.4(2.3—2.6)[2.4] in PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 7 standard length. Tail stout and broad, its width 1.6(1.3—2.0)[1.3] in length of disk margin, taper- ing slightly but rounding off more abruptly at caudal fin than in other species. Caudal peduncle thick, its depth 2.5(2.3—2.8)[2.5] in head depth. Rostrum a rod-shaped protuberance with a nar- row base, generally pointing downward but sometimes horizontal relative to long axis of body; never tilted upward. Length of rostrum 5.7(3.4-8.4)[7.0] in length of disk margin; the series at hand shows that during ontogeny the rostrum becomes relatively shorter. Aperture of illicial cavity approximately oval to subtriangu- lar, wider than high in medium- and large-sized specimens, higher than wide in specimens less than about 65 mm SL. Cranium when viewed from front rising in a gradual curve above disk, head depth 1.9(1.8-2.1)[1.9] in length of disk margin. Width of cranium 3.2(2.7—3.7)[3.3] in length of disk margin. The following in width of cranium: eye 1.4(1.1—1.6)[1.6], lateral ethmoid width 1.6(1.3—1.9)[1.6]. Interorbital space con- cave, narrow, its width 2.7(2. 1—3.4)[3.2] in width of cranium, 4.5(3.3—5.7)[5.7] in head depth. Mouth average size, its width 1.6(1.4—1.9)[1.5] in head depth, 3.1(2.8—3.7)[2.8] in length of disk margin; length of jaw 2.4(2.1—2.7)[2.1] in head depth. Upper lip moderately fleshy, of an even width for its entire length; lower lip thickened medially, the thickening becoming a pronounced lobe in largest specimens. Dorsal fin relatively large, its length 2.8(2.4—3.4)[2.8] in length of disk margin; anal fin also large, its length 2.5(2.1- 2.9)[2.6] in length of disk margin. Integument. Dermal cirri well developed on lateral sides of tail and disk, more so than in other species of Ogcocephalus. Cirri present on tip of rostrum in small specimens. Large buck- lers arranged as described in generic descrip- tion. Dorsal fin devoid of tubercles, anal fin with only a few at base. Pectoral fins with tubercles at base on both dorsal and ventral surfaces, a few tubercles extending out along anterior and posterior rays. Pelvics with tubercles present on basal quarter of ventral surface but none on dor- sal surface. Caudal as in generic description. Skin covering eyeballs bearing tiny tubercles with rim around cornea beaded by a row of small bucklers in small specimens, disappearing in large specimens. Pectoral membranes relatively thin and translucent; no fleshy pads on ventral tips of rays. Color in preservative. Ground color of dorsal BRADBURY: FISH GENUS OGCOCEPHALUS 271 @ parvus © pumilus ° declivirostris FIGURE 25. from Recife, Brazil (Giinther 1880). surface of body gray or tan, apices of bucklers paler than ground color. Dim rounded spots sometimes clustered on shoulders, face, in ax- illae, and along sides of tail, occurring mostly in small specimens. A few small specimens have these spots covering entire disk dorsally. Ven- tral surface of body uniformly pale; lips pale, occasionally with dusky splotches. Rostrum with a diffuse black tip in small specimens, this mark disappearing in larger specimens. Iris black or golden, sometimes golden with a ring of dim spots circling pupil. Pectoral fins dusky, sometimes darkened towards tips, but no pro- nounced dark borders except in some small in- dividuals. Pelvics and anal the same pale ground color as ventral surface of body, anal sometimes with a dusky tip in small specimens. Dorsal dusky with inconspicuous splotches. Caudal as described in generic description except for a few specimens that have the caudal uniformly dusky. Color in life. Unknown. DISTRIBUTION (Fig. 25).—Northern and west- ern coasts of Gulf of Mexico from longitude 86°W to latitude 26°N. One record from Florida Straits. Bathymetric range: 3.5—180 m except for Florida Straits record (388 m). NAME.—The name Ogcocephalus decliviros- tris means an Ogcocephalus with rostrum slop- ing downward. Localities for study material of Ogcocephalus parvus, O. pumilus, and O. declivirostris. O. parvus also known MATERIAL EXAMINED.—HOLOTYPE: FMNH 64136, off Texas, 28°02'N, 96°03’W, 40 m; a male 129.3 mm SL. PARATYPES. Numbers in parentheses are numbers of spec- imens. For data from OREGON stations (=O), refer to Springer and Bullis (1956) and Bullis and Thompson (1965). Gulf coast of Florida: ANSP 103628 (1), O-331; FMNH 61579 (1), 0-858; FMNH 64120 (2), O-944. Florida Straits: UMML 3897 (1), O-1550. Louisiana: CAS 23930 (1), O-2700; CAS 23931 (1), O-2704; CAS-SU 62119 (3), PELICAN sta. 84-3, 28°14’N, 91°41'W, 55 m; FMNH 46735 (1), 0-72; FMNH 46736 (3), O-75; FMNH 46737 (3), O-283-284; GCRL 557 (2) and GCRL 559 (1), s of Grand Isle, 37 m; GCRL 558 (2) and GCRL 560 (1), s of Grand Isle, 27 m; MCZ 45072 (3), PELICAN sta. 74-3, 29°04'N, 88°44.5'W, 110m; MCZ 45073 (1), PELICAN sta. 72-6, 29°11.5'N, 88°52'W, 73 m; MCZ 45074 (1), PELICAN sta. 69-6, 28°48'N, 89°S51’W, 55 m; USNM 131166 (21), Morgan City, 3.7-9.1 m; USNM 188761 (1), sw of Eugene I. Beacon, Atchafalaya Bay; USNM 188764 (1), PELICAN sta. 85-4, 28°11'N, 91°24.5'W, 86 m; USNM 188765 (1), PELICAN sta. 85-3, 28°09’N, 91°27'W, 91 m; USNM 188767 (1), PELICAN sta. 81-8, 28°50’N, 91°29.5'W, 18 m; USNM 188771 (1) and USNM 188774 (1), ssw of Eugene I. Beacon, Atchafalaya Bay; USNM 188777 (1), PELICAN sta. 86-3, 28°28'N, 91°13.5’W, 40 m; USNM 188784 (2), PELICAN sta. 34, 28°33’N, 90°55'W, 26 m; USNM 188792 (2), PELICAN sta. 84-1, 28°22.5'N, 91°44.5'W, 59 m; USNM 188796 (1), PELICAN sta. 94-1, 28°27'N, 92°14’W, 53 m; USNM 188799 (1), PELICAN sta. 93-8, 28°33'N, 92°15.5’W, 44 m; USNM 188801 (2), PEL- ICAN sta. 93-3, 28°55’N, 92°15.5'W, 26 m. Texas: CAS-SU 62140 (30), Galveston; FMNH 61576 (1), 27°04'N, 96°23’W, 55 m: FMNH 61577 (2), 28°02'N, 96°04’W, 40 m; FMNH 61578 (1), 28°02'N, 96°04’ W, 37 m; FMNH 61580 (1), 27°22'N, 96°34'W, 49 m; FMNH 61582 (1), 27°30'N, 96°28'W, 73 m; FMNH 61583 (1), 27°39'N, 96°28’ W, 68 m; FMNH 64135 (1), 27°49'N, 96°31'W, 40 m; FMNH 64137 (1), 27°34'N, 96°43’W, 42 m: FMNH 64138 (2), off Mustang I., 46-48 m; USNM 272 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 7 FIGURE 26. 118606 (23), Galveston; USNM 171763 (2), near Brownsville: USNM 188776 (1), StLveER BAy sta. 200, 27°59'N, 95°15'W, 79-84 m. Material without exact locality: USNM 188766 (1), Madame Canaia. Ogcocephalus pumilus new species (Figure 26D-E) DIAGNOSIS AND COMPARISONS.—A long-nosed Ogcocephalus distinctive for its small size and lack of pigment markings. Ogcocephalus pumi- lus is the smallest member of the genus; maxi- mum standard lengths for specimens seen in this study are all under 70 mm (Table 1). O. pumilus differs from all its congeners except decliviros- tris and notatus in lacking the fleshy ventral pads on the tips of the rays of the pectoral and pelvic fins. Ogcocephalus pumilus is sympatric with par- (A) Paratype of Ogcocephalus corniger new species, 61.0 mm SL, UF 24183, from w of Andros I., Bahamas, 24°25'N, 79°13'W, showing color pattern of juvenile. (B) and (C) Ogcocephalus corniger new species, holotype, 124.5 mm SL, USNM 188808, Gulf of Mexico, 29°00’—-10'N, 85°01’W. (D) and (E) Ogcocephalus pumilus new species, holotype, 57.5 mm SL, USNM 188812, Surinam, 7°05'N, 54°08’ W. vus and nasutus, but as it does not occur on the coasts of the southeastern United States and Gulf of Mexico, its range is less extensive than that of parvus. It is distinguished from parvus by its long rostrum (length of rostrum into length of disk margin 2.0-3.0 in pumilus, 3.2—5.0 in parvus) and shorter disk margin (length of disk margin into SL 2.2-2.7 in pumilus, 1.9-2.2 in parvus) as well as its pale color contrasted with the mottlings and spotted patterns seen in par- vus. O. pumilus is also distinguished from most examples of nasutus by its lack of color pattern as well as its diminutive size (nasutus being among the largest species in the genus). Also, many specimens of pumilus may be separated from nasutus by the pectoral fin ray count (mod- ally 11 in pumilus, 12 or 13 in nasutus, Table 3). BRADBURY: FISH GENUS OGCOCEPHALUS Off the northern coast of South America, the range of pumilus overlaps that of notatus. The two species are separable on the basis of the pectoral fin ray count (11-12 in pumilus, 13-14, rarely 12, in notatus) and subopercular lateral- line scale count (range 5—7, modally 6, in pumi- lus, 7-10, modaliy 9, in notatus). O. pumilus also has a markedly smaller mouth than notatus (width of mouth into length of disk margin 2.6— 3.4 in pumilus, 1.8—2.5 in notatus). DESCRIPTION.—Counts and measurements from 63 specimens, 25.8 to 66.7 mm SL (Table 1). Counts. Counts given in Tables 2-6. In fol- lowing description, counts for holotype given in brackets. All counts in pumilus low compared to counts for genus as a whole. Pectoral fin ray count most often 11 [11,11]; range 10-12. Sub- opercular lateral-line scale count most often 6 [6,7]; range 4—7. Cheek lateral-line scale count most often 8 [8,8]; range 8—9. Mean for lateral- line scale count 21 [21,21]; range 19-24. Anal fin ray count 3 more than half the time [3], remark- able because count normally 4 throughout fam- ily. [D. 4.] Vertebral counts on 14 specimens all 19. Proportions. Proportions expressed as ratios given as mean followed by range in parentheses and proportion for holotype in brackets. Length of disk margin 2.3(2.2—2.5)[2.3] in SL. Tail moderately wide, its width 1.8(1.6—2.1)[1.8] in length of disk margin. Caudal peduncle 2.8(2.5—3.2)[2.6] in head depth. The grotesque rostrum thin and long, its length 2.4(2.0- 3.0)(2.4] in length of disk margin, its long axis about horizontal with body axis rather than tilt- ed upward. Aperture of illicial cavity subtrian- gular, higher than wide. Head depth 1.9(1.7- 2.1)[2.1] in length of disk margin. Width of cra- nium 3.1(2.7—3.5)[3.3] in length of disk margin. The following in width of cranium: eye 1.3(1.1- 1.5)[1.3]; lateral ethmoid width 1.7(1.5—1.9)[1.6]. Interorbital space slightly concave, narrow, its width 3.1(2.4—3.7)[2.8] in width of cranium, 5.1(3.8-6.2)[4.4] in head depth. Mouth moder- ate-sized, its width 1.5(1.4-1.7)[1.7] in head depth, 3.0(2.7—3.4)[3.2] in length of disk margin; length of jaw 2.4(2.1-2.6)[2.4] in head depth. Upper lip moderately fleshy, of an even width for its entire length; lower lip fleshy, no median lobe. Dorsal and anal fins about average in pro- portion to body size, length of dorsal fin 3.3(2.3- 273 4.3)[3.9] in length of disk margin; length of anal fin 2.9(2.4-3.7)[3.0] in length of disk margin. Integument. Dermal cirri usually present on lateral sides of tail and disk margin. No cirri on tip of rostrum. Large bucklers arranged as de- scribed for genus. Dorsal fin devoid of tubercles, anal fin with only a few near base. Pectorals and pelvics with tubercles at base and running out on the fins for 4 to % their length. Caudal as in generic description. Skin covering eyeballs bearing tiny tubercles with rim around cornea often beaded by a row of small bucklers. Pec- toral membranes thin, translucent; pectoral rays without fleshy pads on their ventral tips. Color in preservative. Dorsal surface of body white to pale gray or grayish brown; in the latter case, apices of bucklers paler than ground color. Pigment markings usually absent; if present, they are dusky and ill-defined spots, occurring on shoulders, lateral sides of tail, and (very in- frequently) face. Ventral surface of body uni- formly pale; lips also pale. Rostrum with a dusky subterminal ring. Iris black or silvery. Pectorals colorless in white examples, dusky in gray ones; in the latter, the gray color may shade to a dark- er tone towards tips of pectorals. Pelvics and anal the same pale whitish color as ventral sur- face of body; anal sometimes with a dusky tip. Dorsal colorless in white specimens, dusky in others, sometimes with a faintly darker stripe or two. Caudal colorless to dusky in white exam- ples; gray or brownish individuals with caudal as in generic description. Color in life. From a color transparency taken shortly after capture: body pale bluish gray, ventral surface of body paler than dorsal sur- face, the whole suffused with a wash of trans- lucent pinkish orange. Rostrum a deeper orange. Distal thirds of pectorals and caudal deep or- ange. DISTRIBUTION (Fig. 25).—Known from the Bahamas just north of Cuba, from Puerto Rico, the Leeward Islands, the western Caribbean, and the northern coast of South America to Sur- inam. Bathymetric range: 35-348 m. NAME.—The name pumilus, which is derived from a word meaning dwarf, is given in refer- ence to the diminutive size and grotesque ap- pearance of this species. MATERIAL EXAMINED.—Numbers in parentheses are num- bers of specimens. For data from OREGON (=O) gnd SILVER 274 Bay (=SB) stations, refer to Bullis and Thompson (1965). Ho.otyPe: USNM 188812, Surinam, O-2018, 57.5 mm SL. PARATYPES. Northern Bahamas: USNM 188759 (2), SB- 2455. Puerto Rico: FMNH 64843 (2) and FMNH 64844 (10), O-2626; FMNH 64845 (2), O-2627; MCZ 45085 (3), O-2668; MCZ 45087 (8), O-2626. Virgin Islands: CAS 23920 (1), O- 2618; FMNH 64841 (4), O-2616; USNM 108353 (1), 18°25'30"N, 63°32'36”W. Leeward Islands: FMNH 64846 (5), O-2631; FMNH 64848 (2), O-2633; FMNH 64891 (33), O-2231; UMML 7488 (14), O-2632; USNM 190467 (1), O-2632. Hon- duras: CAS 23921 (1), O-1865. Guyana: ANSP 103632 (3), O- 2231; ANSP 103633 (1), O-2262; CAS 23919 (1), O-2249; CAS- SU 62122 (3), O-2345; CAS-SU 62125 (1), O-2259; FMNH 64893 (3), O-2249; FMNH 65965 (1), O-2001; FMNH 65966 (1), O-2234; FMNH 65967 (3), O-2248; FMNH 65968 (1), O- 2250; FMNH 65969 (3), O-2261; FMNH 65970 (4), O-2344; MCZ 45086 (3), O-2261; USNM 185984 (3), 8°33'N, 58°46’W. Surinam: FMNH 64899 (2), O-2330. Ogcocephalus corniger new species (Figure 26A—-C) Ogcocephalus vespertilio (non Linnaeus): LONGLEY AND HIL- DEBRAND 1941:311, 314 [Tortugas, Florida; key; description]. Ogcocephalus sp.: LONGLEY AND HILDEBRAND 1941:311, 315 [Tortugas, Florida; key; partial description by Hildebrand]; WALLS 1975:118 [northern Gulf of Mexico; partial descrip- tion; figs.; called Ogcocephalus sp. A; color photograph]. Oncocephalus vespertilio (non Linnaeus): FOWLER 1952:14 {off southern Florida; partial description]. Probable references: Lophius nostratus MITCHILL 1818:325 [“‘Straits of Bahamas”’; a misspelling of Lophius rostratus Shaw]. Malthe vespertilio (non Linnaeus): UHLER AND LUGGER 1876:77 [southern Chesapeake Bay; partial description]. Oncocephalus vespertilio (non Linnaeus): GARMAN 1896:87 {off Key West, Florida; partial description]. Ogcocephalus vespertilio (non Linnaeus): H. M. SMITH 1907:401, figs. 185-186 [off Beaufort, North Carolina; par- tial description; figures the same as Jordan and Evermann 1898:pl. 392, figs. 958 and 958b (legend: Ogcocephalus ves- pertilio)). DIAGNOSIS AND COMPARISONS.—A long-nosed Ogcocephalus distinctive for its dorsal color pattern, which consists of small pale round spots close together and evenly distributed over the darker background color of the dorsal surface of the body (specimens under about 65 mm SL lack this pattern; see sections on color below). The long rostrum in corniger separates it from all other species of Ogcocephalus except ves- pertilio, pumilus, long-nosed morphs of nasutus, and small examples of cubifrons less than 50 mm SL (Fig. 9). The range of corniger (coasts of southeastern United States and eastern Gulf of Mexico) is widely separated from that of ves- pertilio (coast of Brazil) and, at this time, ap- pears to be separated from that of pumilus (northern Bahamas, Caribbean, and coast of South America to Surinam). Besides separate PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 7 ranges, other aspects separate these long-nosed species. O. corniger differs from vespertilio in its lower number of pectoral rays (10-12 in cor- niger, 13-15 in vespertilio) and lower number of subopercular lateral-line scales (usually 5-7 in corniger, usually 8-9 in vespertilio, Table 6); it differs from pumilus in its larger body size (pumilus is less than 70 mm in SL) and in having fleshy pads on the ventral tips of the pectoral rays which pumilus lacks. The range of corniger overlaps the ranges of nasutus and cubifrons, but long-nosed forms of nasutus and cubifrons may be distinguished from corniger by the pectoral fin ray count (modally 11 in corniger, usually 12 or 13 in na- sutus and cubifrons, Table 3). O. corniger dif- fers further from nasutus in the number of cheek lateral-line scales (usually 8 in corniger, modally 9 in nasutus, Table 5) and from cubifrons by its longer jaw (length of jaw into width of cranium 1.2-1.5 in corniger, 1.6—2.0 in cubifrons). Two short-nosed species of Ogcocephalus, parvus and rostellum, occur within the range of corniger but are separable from corniger not only by the proportions of the rostrum but also by color pattern—neither ever has the pattern of small round pale spots on a darker back- ground seen in corniger. Other characters that separate corniger from parvus include the smoother shagreenlike quality of the integument in corniger compared to the rough, lumpy sur- face of the integument in parvus, and the larger mouth in corniger (width of mouth into head depth 1.5-2.0 in corniger, 2.0-2.8 in parvus). An additional character separating corniger from rostellum is the pectoral fin ray count (modally 11 in corniger, modally 13 in rostellum, Table 3). DESCRIPTION.—Counts and measurements from 57 specimens 26.2 to 134.0 mm SL (Table }): Counts. Counts given in Tables 2-6. In the following description, counts for holotype given in brackets. Counts in corniger low compared to those of genus as a whole. Pectoral fin ray count usually 11 [11,11]; range 10-12. Suboper- cular lateral-line scale count usually 6 or 7 [6,6]; range 4-8. Cheek lateral-line scale count 8 [8,8] (the commonest count for genus); range 7-9. Mean for lateral-line scale count nearly 21 [20,20]; range 18-24. Majority of sample of cor- niger had 19 vertebrae, the most usual vertebral count for the genus. [D. 3; A. 4.] BRADBURY: FISH GENUS OGCOCEPHALUS Proportions. Proportions expressed as ratios given as mean followed by range in parentheses and proportion for holotype in brackets. Length of disk margin intermediate relative to range of variation in genus, 2.3(2.1—2.5)[2.4] in SL. Tail narrow to moderately wide, the width 2.0(1.7—2.4)[1.7] in length of disk margin, taper- ing to a relatively thin caudal peduncle; depth of caudal peduncle 3.3(2.8—4.1)[3.3] in head depth. Rostrum with a wide base but becoming slender distally, upturned and very long, the length 2.0(1.7—2.3)[2.1] in length of disk margin. Aperture of illicial cavity triangular, always higher than wide. Cranium, when viewed from front, rising steeply above disk; head relatively deep, its depth 1.8(1.6—1.9)[1.7] in disk margin. Width of cranium 3.3(2.9-3.5)[3.2] in length of disk margin. Eye comparably large, 1.3(1.1- 1.7)[1.5] in width of cranium. Lateral ethmoid width 1.7(1.5—2.0)[1.7] in width of cranium. In- terorbital space flat to moderately convex, its width 2.1(1.7—2.5)[1.9] in width of cranium, 3.9(3. 1-4.6)[3.6] in head depth (except one spec- imen, which has these proportions 2.9 and 5.7, respectively). Mouth average size, its width 1.6(1.5—2.0)[1.5] in head depth, 2.9(2.6—3.7)[2.6] in length of disk margin; length of jaw 2.4(2.2- 2.6)[2.5] in head depth. Upper lip fleshy, of an even width for its entire length; lower lip also fleshy, thickly swollen medially. Dorsal fin small compared to other species in genus, its length 4.8(3.7—5.8)[S.8] in length of disk margin; anal fin also small, its length 3.0(2.5—3.5)[3.0] in length of disk margin. Integument. Dermal cirri absent, or at least inconspicuous, except near tip of rostrum, where fairly long cirri occur in subterminal ring. Large bucklers arranged as in generic descrip- tion. Dorsal fin devoid of tubercles; anal fin with tiny tubercles covering proximal third or half of fin. Pectorals and pelvics with tiny tubercles at base on both dorsal and ventral surfaces, run- ning out along fin rays for a third to half their length. Caudal as in generic description. Skin covering eyeballs bearing fine tubercles with rim around cornea beaded by a row of slightly larger ones. Pectoral membranes thick, opaque, ven- tral surfaces of the ray tips with well-developed fleshy pads (pads not developed in small speci- mens). Color in preservative. Dorsal surface of body uniformly covered with small pale spots on a darker ground color, the ground color generally 275 FIGURE 27. corniger. Localities for study material of Ogcocephalus brown or gray, becoming darker on shoulders in many examples; sometimes spots so close to- gether that ground color may appear as a retic- ulum. The pale spots are unpigmented apices of bucklers. Specimens under about 65 mm SL generally lack this color pattern and are uni- formly pale or dusky, but occasionally, dark rings occur clustered on shoulders. Ventral sur- face of body uniformly pale; lips pale. Rostrum with a diffuse dark subterminal ring, incomplete below. Iris black or golden, or golden with spots, but never with radiating lines. Pectorals pale to dusky proximally with broad dark brown or black margins; ray tips white, ventral pads of rays white. Pelvic and anal fins the same ground color as ventral surface of body; pelvics some- times with darkened or blotched tips in dark specimens; anal most often with distal third very dark to black. Dorsal dusky, often with a pale diagonal stripe or two. Caudal as in generic de- scription, the border always dark to black. Color in life. The following description from three color transparencies made of a freshly killed specimen captured in winter off North Carolina. One transparency shows a dorsal view of the fish and one a ventral view; the remaining one shows a lateral view of the head. 276 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 7 Ground color of dorsal surface of body choc- olate brown with fine, close-set pearl-gray spots completely covering the surface, including face and skin covering eyeballs. Even iris of eye has the spotted pattern, with a brassy highlight around pupil. Lips bright red-orange, with a translucent wash of red-orange pigment over lower face and lower edge of disk, where it be- comes continuous with the same red-orange wash over ventral surface of body, including pelvic and anal fins. This red-orange coloration on ventral surface is more intense on chin, on a ring encircling anus, and on tail including anal fin, which also has a black tip. Pectoral fins with translucent red-orange color basally, the rays becoming deep red-orange distally and the fins each with a broad blackish margin. Caudal dark basally, becoming pale red-orange in middle portion with rays a darker shade than mem- branes, and black again distally in a broad mar- gin. Longley (Longley and Hildebrand 1941) stat- ed for examples from off Tortugas (his Ogco- cephalus vespertilio) that the pectorals and cau- dal were, “‘buff-yellow over basal half, passing through maroon-purple to prune purple at the tips, the contrast between the darker outer third and the yellow basal two-thirds being sharper on the under side of the pectorals; belly, maxillary membranes and buccal cavity coppery red.”’ Walls (1975) presents a photograph of an ex- ample from the Gulf of Mexico (his Ogcoceph- alus sp. A) which shows the pectorals red-or- ange basally, succeeded by bright yellow, which gives way to more red-orange, which in turn gives way to black borders. DISTRIBUTION (Fig. 27).—Known from the Atlantic coast of the United States from Cape Lookout, North Carolina, to Florida, Campeche Banks, and the northeastern Gulf of Mexico to Louisiana; one record from off Great Inagua I., Bahamas. Bathymetric range: 29-230 m. NAME.—This form is named corniger, mean- ing horn-bearer, in reference to the long, up- turned rostrum. MATERIAL EXAMINED.—Numbers in parentheses are num- bers of specimens. For data from early OREGON stations (= O), refer to Springer and Bullis (1956); for data from later OREGON stations and for ComBat (=C) and SILver Bay (= SB) stations, refer to Bullis and Thompson (1965). HOLOTYPE: USNM 188808, Gulf coast of Florida, O-729-730; a male 124.5 mm SL. PARATYPES. North Carolina: CAS 23927 (1), SB-2930: USNM 152030 (1). Atlantic coast of Florida: ANSP 103626 (1), C-101; CAS-SU 62124 (1), C-497; CAS-SU 62128 (1), SB-1931; USNM 161375 (1), Palm Beach. Bahamas: FMNH 66387 (1), SB-3496; UF 24183 (1), SB-2470. Florida Straits: CAS 23926 (1), SB-437; UF 24182 (1), SB-2362. Florida Keys: FMNH 64124 (6), O-1020; USNM 37853 (1), ALBATROSS sta. 2316, 24°25'30"N, 81°47'45"W, 91 m; USNM 74089 (1), s of Key West; USNM 117001 (1) and 117002 (3), Tortugas. Gulf coast of Florida: FMNH 64103 (1), O-603; FMNH 64104 (1), O-727- 728; FMNH 64109 (1), O-732; FMNH 64113 (1), O-897; FMNH 64116 (2), O-917; FMNH 64119 (1), O-936; FMNH 64122 (2), O-945; 64126 (4), O-1021; FMNH 64129 (3), O-1022; MCZ 45070 (1), O-326; USNM 134087 (1), 28°47'30’N, 84°37'00"W. Alabama: FMNH 46744 (1), O-265; MCZ 45071 (1), O-2838; USNM 159159 (1), 29°47'N, 87°17'W. Louisiana: USNM 188793 (1), SB-175. Yucatan: USNM 188804 (1), SB- 404. Ogcocephalus parvus Longley and Hildebrand (Figure 28) Malthe vespertilio (non Linnaeus): GUNTHER 1880:7 [Pernam- buco (=Recife), Brazil]. Ogcocephalus parvus LONGLEY AND HILDEBRAND, 1940:283, fig. 28 [s of Tortugas, Florida; holotype USNM 109313]; 1941:311 and 314 [Tortugas, Florida; key]; ERDMAN 1956:338 [Joyuda, Puerto Rico; partial description]; BRIGGS 1958:301 [Florida, compiled]; MOE ET AL. 1966:120 [Flori- da; list of museum specimens]; BRADBURY 1967:417 [listed]; STARKS 1968:33 [Alligator Reef, Florida; composition of reef fauna]; WALLS 1975:116 [N Gulf of Mexico; partial description]. Oncocephalus parvus: FOWLER 1945:336 [Key West, Florida; listed]; 1952:15 [Sombrero Key Light, Florida; description of color]. Probable reference: Ogcocephalus vespertilio: DAHLBERG 1975:45, fig. 93 [coast of Georgia; partial description; range, compiled]. DIAGNOSIS AND COMPARISONS.—An Ogco- cephalus with exceptionally angular, craggy body contours and large, prominent bucklers. O. parvus differs from all its congeners except cubifrons in having a very small mouth relative to head depth (width of mouth into head depth 2.0-2.8 in parvus, 1.0—2.0 in all others except cubifrons). O. parvus may be separated from cubifrons by its narrower interorbital space (in- terorbital width into head depth 3.6—6.6 in par- vus, 2.3-3.4 in cubifrons). It may be distin- guished from all but corniger, declivirostris, and pumilus by its low pectoral fin ray count (usually 10-11 in parvus, 12 or more in all others except corniger, declivirostris, and pumilus). O. par- vus is distinguished from corniger and pumilus by its short rostrum (length of rostrum into length of disk margin 3.2—5.1 in parvus, 1.7-3.0 in corniger and pumilus) and from declivirostris BRADBURY: FISH GENUS OGCOCEPHALUS FIGURE 28. Right: FMNH 65957, Guyana, 64.0 mm SL. by the fleshy ventral pads developed on the dis- tal ends of the pectoral fin rays (absent in decli- virostris ). DESCRIPTION.—Counts and measurements from 69 specimens, 27.3 to 84.7 mm SL (Table 1). Because specimens in the northern part of the range (coasts of southeastern United States and eastern Gulf of Mexico) differed in pigment pattern from those in the southern part (Carib- bean Sea and Atlantic coast of South America), the two groups were analyzed separately with the expectation that samples would accumulate that were large enough to differentiate the groups well. However, too few specimens from the southern part of the range were obtained, so only the color patterns and counts are treated separately in the following description. Data for body proportions were combined for the two groups. Counts. Counts given in Tables 2-6. Counts for parvus the lowest found in genus. Pectoral fin ray count usually 10 in northern parvus and Ogcocephalus parvus Longley and Hildebrand. Left and center: FMNH 46742, Gulf of Mexico, 75.0 mm SL. 11 in southern parvus; range 10-12 for species. Subopercular lateral-line count usually 5 or 6; range 4-8. Cheek lateral-line scale count usually 8: range 7-9. Mean count for lateral-line scales 17; range 15-19. Vertebral count modally 19; range 18-20 (all vertebral counts are from ho- lotype and paratypes). Proportions. Proportions expressed as ratios given as mean followed by range in parentheses. Relative length of disk margin longest for the genus, its length 2.1(1.9-2.2) in SL. Tail mod- erately wide, its width 2.0(1.7-2.5) in length of disk margin. Caudal peduncle 3.1(2.5—4.0) in head depth. Rostrum variously finger- to cone- shaped, but always short, its length 4.2(3.2—-5.1) in length of disk margin. Aperture of illicial cav- ity oval or subcircular. Cranium rising steeply above disk, head depth 1.8(1.6—2.0) in length of disk margin. Width of cranium 3.2(2.6—3.6) in length of disk margin. The following in width of cranium: eye 1.4(1.1-1.6); lateral ethmoid width 1.7(1.4-1.9). Interorbital space narrow, con- 278 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 7 cave, its width 2.7(2.1—4.1) in width of cranium, 4.8(3.6—6.6) in head depth. Mouth small and nar- row, its width 2.3(2.0-2.8) in head depth, 4.1(3.4—5.2) in length of disk margin; jaw short, its length 3.1(2.8-3.5) in head depth. Upper lip fleshy, of an even width for its entire length; lower lip with a fleshy median lobe. Dorsal fin average size, its length 3.4(2.4-4.3) in length of disk margin; anal fin short, its length 3.1(2.4— 3.6) in length of disk margin. Integument. Dermal cirri usually sparse or absent, but a few examples have cirri well de- veloped over dorsal surface of body. Most spec- imens from southern population have subter- minal rostral cirri. Large bucklers arranged as in generic description. Dorsal fin devoid of tu- bercles; anal fin with only a few tiny ones at base. Pectorals and pelvics with tiny tubercles covering basal third or fourth of their length. Caudal as in generic description. Skin covering eyeballs bearing tiny tubercles with rim around cornea beaded by a row of small bucklers. Pec- toral membranes thick, opaque; ventral surfaces of ray tips with well-developed fleshy pads. Color in preservative. Northern examples: ground color of dorsal surface of body pale tan to medium brown. Markings variable but irreg- ular or rounded dark brown spots, with or with- out whitish margins, clustered on shoulders, face, in axillae, and sometimes along lateral sides of tail. Occasionally some of these mark- ings are rings instead of spots. Dark splotches, sometimes indistinct, may occur elsewhere on dorsal surface. A few specimens with minute dark spots scattered over dorsal surface in ad- dition to other markings. Ventral surface of body uniformly creamy pale, sometimes becom- ing dusky on anterior part of head, particularly on chin, in which case two distinct white patches may occur, one on either side of mandible. Pec- torals pale to nearly white with a broad black margin extending through all but the three short- est rays; ray tips white, ventral pads white. Pel- vics and anal same pale ground color as ventral surface of body; anal often with a dusky or dark spot distally. Dorsal colorless or a little dusky basally. Caudal colorless in about half the spec- imens at hand, otherwise as in generic descrip- tion. Iris black or golden, or golden with dark spots. Southern examples: ground color of dorsal surface of body usually a much darker brown than in northern specimens (this difference does not show well in Fig. 28 because photographed specimens were chosen to show pigment mark- ings contrasted as well as possible with ground color; fish on right in Fig. 28 is relatively pale). Minute but very distinct black dots on lateral portions of disk and on top of cranium, continu- ing posteriorly along dorsal surface of tail. Large, sharply defined brown or black spots and rings on shoulders and face and also in axillae, extending from these on to sides of tail. A few specimens with an absolutely pigmentless white patch a little larger than top of cranium in area and located in middorsal region just posterior to cranium. Ventral surface of body, fins, and eyes as in northern examples. Color in life. From an aquarium specimen newly captured in Gulf of Mexico off Mobile, Alabama, kindly shown me by Dr. Robert L. Shipp and his students at the University of South Alabama: ground color of face and dorsal surface of body dark brown, the large irregularly shaped spots on shoulders and axillae lavender separated by cream-colored reticula in approx- imately the pattern seen on the right in Figure 28; smaller pepperlike spots on tail and disk seen in figure are black in living specimen. Down face on either side from eye to angle of jaw a cascade of orange spots having dark brown rims and sep- arated by white-colored reticula. Lips orange. Iris gold flecked with orange. Pectorals a trans- lucent orange with broad black margins. Caudal dark brown basally, white in middle third, or- ange on distal third. Several Kodachrome transparencies made from specimens immediately after capture have been available. From specimens from off the southeastern United States, the following notes are offered: ground color of dorsal surface of body bright tan suffused with red-orange in large patches; dark spots arranged as described for preserved specimens. Dorsal fin red-orange. Pectorals with a red-orange stripe just proximal to black margin; ray tips white. From a specimen newly captured from off coast of Guyana: dorsal surface of body very dark, the markings difficult to discern except those on shoulders (the usual cluster of irregular spots). Skin surrounding gill pores tan. Tips of many bucklers faintly red-orange. Pectorals with red-orange stripe just proximal to black margins; ray tips white. Ventral surface of body, includ- BRADBURY: FISH GENUS OGCOCEPHALUS ing pectoral peduncles and bases of pectoral fins, entirely bright red-orange. Anal fin red-or- ange with white stripe through second quarter. Lips red-orange, corresponding to an observa- tion by Erdman (1956) of a specimen from Puer- to Rico with ‘‘bright red lips.’’ A rectangularly shaped white patch on either side of mandible and an oblong white patch in median line about halfway between chin and bases of pelvics. DISTRIBUTION (Fig. 25).—Known from the Atlantic coast of the United States from Cape Hatteras south to the eastern Gulf of Mexico, the Caribbean Sea, and the coast of South America to Recife, Brazil. Bathymetric range: 29-126 m. MATERIAL EXAMINED.—Numbers in parentheses are num- bers of specimens. For data from early OREGON stations (= O), refer to Springer and Bullis (1956); for data from later OREGON stations as well as COMBAT (=C) and SILVER BAy (=SB) stations, refer to Bullis and Thompson (1965). Northern examples North Carolina: CAS 23922 (1), SB-2927; CAS 23924 (1), SB-3339; FMNH 64336 (1), C-384; UF 24186 (1), C-385. South Carolina: MCZ 45082 (1), C-166. Georgia: FMNH 66383 (9), BOwERS sta. 32, 30°14’N, 80°16’W, 73 m; MCZ 45083 (1), PELICAN Sta. 178-15, 31°20'N, 80°17’ W, 38 m; UMML 114 (1), between Jacksonville, Florida, and Brunswick, Georgia. At- lantic coast of Florida: ANSP 103631 (1), SB-2721. Florida Straits: CAS-SU 62118 (2), C-457; UF 24187 (2), SB-2363; UF 24188 (1), SB-2382. Florida Keys: FMNH 64125 (6), O-1020; USNM 109313 (holotype) and USNM 109314 (12 paratypes), Tortugas, 134-201 m. Gulf coast of Florida: FMNH 46742 (1), 29°04'N, 84°23.5'W, 37 m; FMNH 46743 (2), O-35; FMNH 64105 (1), O-729-730; FMNH 64108 (2), O-732; FMNH 64112 (1), and FMNH 64114 (2), O-897; FMNH 64115 (1), O-916; FMNH 64117 (3), O-917; FMNH 64118 (2), O-936; FMNH 64127 (3) and FMNH 64128 (3), O-1021; FMNH 64130 (1), O-1022; GCRL (1), O-35; MCZ 45084 (1), O-1024; USNM 188795 (1), PELICAN sta. 153-3, 29°24’N, 85°54’W, 37 m. Al- abama: USNM 188778 (1), PELICAN sta. 137-2, 29°36'N, 87°29'W, 66 m. Yucatan: CAS 23923 (1), SB-438; FMNH 46741 (1), O-222. Southern examples Puerto Rico: USNM 164504 (1), Joyuda. Honduras: ANSP 103630 (1), O-1874. Guyana: CAS-SU 62121 (1), O-2000; FMNH 64892 (2), O-2247; FMNH 64895 (3), O-2261; 65954 (1), O-2245; FMNH 65955 (1), O-2232; FMNH 65956 (1), O- 2257; FMNH 65957 (4), O-2000; FMNH 65958 (1), 1999; MCZ 48081 (1), O-2262; USNM 188768 (2), O-2000. Brazil: BMNH 79.5.14.527 (1), CHALLENGER Sta. 122, off Pernambuco (=Re- cife). Ogcocephalus darwini Hubbs (Figure 29 [left]) Ogcocephalus darwini Huss, 1958:161 [Isla Isabella, Gala- pagos Islands; holotype SIO H51-214; photographs]; BRAD- BURY 1967:417 [listed]; MCALLISTER 1968:161 [Tagus Cove, Albemarle (=Isabella) I., Galapagos Islands; branchioste- gals described]. 279 DIAGNOSIS AND COMPARISONS.—An QOvco- cephalus distinguished by the unique character of its smooth integument, likened to shagreen by Hubbs in the original description (1958); the bucklers are obscured by a covering of fine spi- nules in contrast to the prominent, coarse buck- lers of other species. O. darwini is also distin- guished from all its congeners except the single other eastern Pacific species, O. porrectus, by a pair of solid dark stripes on the dorsal surface of the disk which continue on to the lateral sides of the tail; all Atlantic species of Ogcocephalus either are spotted on the dorsal surface of the body or else lack markings altogether. Differ- ences between darwini and porrectus include a shorter disk margin in darwini (Fig. 9) and a higher modal pectoral fin ray count (15 in dar- wini, 14 in porrectus; Table 3). DESCRIPTION.—A detailed description and comparison with the two other eastern Pacific ogcocephalines is given in the original descrip- tion of Ogcocephalus darwini (Hubbs 1958), but in order to keep the presentations of species in this revision strictly parallel for ease of refer- ence, a description is given again below, includ- ing some new observations. Counts and mea- surements from 30 specimens 35.1 to 166.1 mm SL (Table 1). Counts. Counts given in Tables 2-6. O. dar- wini has the highest mean for pectoral fin ray count of any in genus; range 14-15 (but Hubbs (1958) gives 14-16; however, his sample includ- ed three specimens I have not seen). O. darwini most often has 6—7 subopercular lateral-line scales but shows considerable variation in count; range 4—9. Cheek lateral-line count usu- ally 8, as for most species in genus; range 6-9. Range for lateral-line scale count 19-30, which is about center of distribution for genus. Modal vertebral count 19; range 19-20. Proportions. Proportions expressed as ratios given as mean followed by range in parentheses. Disk margin shorter on the average than in any other species of Ogcocephalus, its length 2.6(2.3-2.9) in SL. Tail thin, its width 2.0(1.7— 2.4) in length of disk margin, tapering evenly to caudal fin; caudal peduncle of moderate depth, 2.7(2.4—3.1) in head depth. Rostrum thick and blunt, moderately long, its length 3.1(2.8—3.6) in disk margin. Aperture of illicial cavity subtrian- gular, higher than wide or the two dimensions about equal. Cranium, when viewed from front, 280 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 7 FIGURE 29. Ogcocephalus darwini Hubbs, left, top and bottom, paratype 128.5 mm SL, CAS-SU 17112. Ogcocephalus porrectus (Garman), right, top and bottom, paratype 108.0 mm SL, MCZ 28733. rising steeply above disk, head depth 1.8(1.6— 2.1) in length of disk margin. Width of cranium 3.2(2.9-3.5) in length of disk margin. The follow- ing in width of cranium: eye 1.3(1.3—1.5), lateral ethmoid width 1.6(1.4—-1.8). Interorbital space wide, concave, the width 1.8(1.6—2.1) in width of cranium, 3.2(2.9-3.6) in head depth. Mouth average in width, the width 1.7(1.4—2.0) in head depth, 3.1(2.7-3.4) in length of disk margin; length of jaw 2.4(2.0-2.6) in head depth. Upper lip moderately fleshy, of an even width for its entire length; lower lip fleshy, no median lobe. Dorsal fin average size, its length 3.8(3.0—4.3) in length of disk margin; anal fin comparatively long, 2.4(2.1—2.6) in length of disk margin. Integument. Dermal cirri present or absent. Large bucklers present but inconspicuous be- cause of their covering of skin embedded with BRADBURY: FISH GENUS OGCOCEPHALUS E Eo 5 Q x = £ lo) “OO Ow ‘= Se DD soRe ye Oo A os : = O- Garwini (je Mm a © — porrectus [e) ° — 50 60 70 80 90 100 Standard length in mm. 110 120 130 140 I50 160 FiGure 30. Comparison between Ogcocephalus darwini and O. porrectus to show that length of disk margin is greatest in O. porrectus relative to standard length. fine spinules (except small specimens 35 mm SL or less have perfectly visible bucklers not yet covered by the fine-grained, shagreenlike integ- ument); bucklers arranged as in generic descrip- tion. All fins with fine tubercles at their bases and extending out variable distances along fin rays. Caudal as in generic description. Skin cov- ering eyeballs bearing tiny, densely set tuber- cles, with rim around cornea beaded by a row of somewhat larger ones. Pectoral membranes thick, opaque; ventral surfaces of ray tips with well-developed fleshy pads. Color in preservative. Ground color of dorsal surface tan to brown, the conspicuous markings a pair of dark stripes, one on each side, origi- nating behind eyes and extending posteriorly over disk and on to lateral walls of tail where they are sometimes interrupted to form a series of blotches. Top of head dark, from whence a median dusky stripe extends posteriorly, be- coming wider and more intense around dorsal fin. Face also dark, marked by a narrow light stripe on either side extending obliquely from eye to lip just anterior to corner of mouth, a pattern very similar to that observed in Ogco- cephalus porrectus. Ventral surface uniformly pale except for chin, which is sometimes dusky; the pale shade extends up sides of tail to the lateral line or even a little above. Rostrum the same dark shade as face and head. Iris black or gold, or black with golden spots. Dorsal surfaces of pectorals pale basally, grading to black dis- tally, the ventral pads near tips of rays pale or white. Pelvics and anal the same ground color as ventral surface of body, anal sometimes dusky or black on distal third. Dorsal fin dusky to dark, sometimes blotched. Caudal fin pale basally with distal third dark except in one spec- imen, which displays color pattern described for genus. Color in life. Hubbs (1958) states for one spec- imen which retained some color, “‘upper parts, purplish gray; rather blue-gray on tubercles and in an irregular blotch near each side of the disk; the two dark streaks, reddish brown; under- parts, bright rose-red, becoming white or whit- ish on the lower (but not the upper) surface of the pelvics, on the outer tip of the anal, and, weakly, on the lower border of the caudal, also on the esca (but not the red stem) of the illicium; pectoral rays pink-gray, encroached by the wid- ening, blackish interradial streaks; the fin be- coming almost solidly blackish inside the narrow red outer border.” I observed two live specimens in August 1968 which had been freshly taken from coral rubble in 25 m of water off Isla Isabela by pipe dredge. In these the dorsal surfaces of disk and tail me- dium brown, the two longitudinal stripes dark cocoa brown. Ventral surfaces of body creamy white, as were esca and tops of eyeballs, all con- trasting sharply with brown head and brown ros- trum. Lips cherry-red; striking white blotches along sides of disk posterior to mouth. Pelvics creamy white tipped with brown on dorsal sur- faces. Pectorals brown on dorsal surfaces, grad- 282 ing to rich dark shade distally, creamy white on ventral surfaces. DISTRIBUTION.—Galapagos Islands. Bathy- metric range: 3.5—73.5 m. MATERIAL EXAMINED.—Numbers in parentheses are num- bers of specimens. All material from the Galapagos Islands. Isla Isabela: SIO H51-214 (holotype) and SIO 54-175 (1), Cale- ta Tagus; SIO H50-18 (1), Punto Moreno, shallow water near shore; SIO 55-16 (1), Punto Moreno, under 9 m; SIO H50-132 (1), Bahia Elizabeth, 5.5 m; SIO 54-199 (1) and SIO 58-116 (1), Bahia Elizabeth; SIO 57-20 (1), Bahia de Banks, approx. 00°01'S, 91°29'W; CAS-SU 14977 (1), Bahia de Banks, under 77 m; SIO HS1-51 (1), just outside Caleta Webb; SIO 57-111 (1), Caleta Webb; CAS 39904 (13), TE VEGA cr. 17, sta. 91, 00°15'22"S, 91°22'26”W, Canal Bolivar; SIO 56-60 (1), 4.6—-5.6 m; SIO 58-39 (1), w side I. Isabela or E side I. Fernandina. Isla Fernandina: SIO H53-196 (1), Punta Mangle. Isla Santa Cruz: USC (1), Allan Hancock Pacific Exped. 1935, sta. 345- 35, 00°24'50"S, 90°21'40’W, 55 m. Other material: SIO 57-166 (1), exact locality unknown; CAS-SU 17112 (1), exact locality unknown; CAS-SU 46654 (1), from stomach of shark (Gyropleurodus quoyi) taken in Caleta Tagus. Ogcocephalus porrectus (Garman) (Figure 29 [right]) Oncocephalus porrectus GARMAN, 1899:86 [5°32'45"N, 86°54'30"W; lectotype MCZ 28733]. Ogcocephalus porrectus: HuBBS 1958:161 [redescription of type-series; photographs; selection of lectotype]; BRAD- BURY 1967:417 [listed]. DIAGNOSIS AND COMPARISONS.—One of only two species of Ogcocephalus known from the eastern Pacific Ocean (the other is O. darwini). O. porrectus and O. darwini are morphologi- cally distinguishable by the nature of the squa- mation, shagreenlike and relatively smooth in darwini but rough with prominent spiny buck- lers in porrectus. Other differences include the relatively longer disk margin in porrectus (Fig. 9) and lower modal number of pectoral rays (14 in porrectus, 15 in darwini, Table 3). Although a smaller species than nasutus (Ta- ble 1), porrectus most resembles nasutus in body proportions and quality of the squamation. However, the color pattern in porrectus, con- sisting of a longitudinal stripe on each side of the body, is unknown in nasutus or any other Atlantic species. The relationship in color pat- tern between Atlantic and Pacific species in this: the clusters of spots or reticulations where found in tracts in Atlantic species are represented in porrectus (and in darwini) by solid stripes. DESCRIPTION.—Counts and measurements from 35 specimens 25.6 to 138.5 mm SL (Table 1). Counts. Pectoral fin ray count relatively high PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 7 with modal number 14 in range of 10 to 15 for genus (Table 3). Lateral-line scale counts below average, however, with lateral-line count always under 30 (Table 4) and subopercular lateral-line count modally 6 (Table 6). Cheek lateral-line scale count usually 8 as in most species in the genus (Table 5) and vertebral count usually 19 (Table 2). Proportions. Proportions expressed as ratios given as mean followed by range in parentheses. Disk outline subtriangular, length of disk margin 2.2(2.1-2.4) in SL. Tail thin to moderately wide, its width 2.0(1.6—2.4) in length of disk margin, tapering evenly to caudal fin; caudal peduncle of average thickness, its depth 3.0(2.6—3.4) in head depth. Rostrum moderately long with a thick base, its length 3.0(2.8-3.4) in length of disk margin; seen from side, rostrum arches slightly with its distal end pointing downwards except that terminal buckler is turned abruptly upwards (Fig. 29). Aperture of illicial cavity sub- triangular in outline, higher than wide. Cranium, when viewed from front, rising steeply above disk; head depth 2.0(1.9-2.2) in length of disk margin. Width of cranium 3.2(2.3—3.9) in length of disk margin. The following in width of cra- nium: eye 1.6(1.3—-2.0), lateral ethmoid width 1.7(1.5—1.9). Interorbital space flat, not convex as in vespertilio, of average width, the width 2.1(1.8-2.5) in width of cranium, 3.6(3.0—4.0) in head depth. Mouth moderate, its width 1.6(1.4— 1.8) in head depth, 3.2(2.8—3.6) in length of disk margin; length of jaw 2.3(2.1—2.5) in head depth. Upper lip moderately fleshy, of an even width for its entire length; lower lip fleshy, thickened medially. Dorsal fin small, its length 4.4(3.4—4.8) in length of disk margin; anal fin relatively long, its length 2.6(2.4—3.0) in length of disk margin. Integument. Dermal cirri inconspicuous, usu- ally present on disk margin, chin, lateral sides of tail near lateral line, and on sides of large bucklers on dorsal surface of body. Large buck- lers prominent, with coarse spines bristling from apex of each; bucklers arranged as in generic description. All fins except dorsal with fine tu- bercles running out for 4 to % length of fin rays; dorsal fin with none or only a few prickles on anterior edge of first ray. Caudal as in generic description. Skin covering eyeballs bearing fine tubercles with rim around cornea beaded by a row of small but prominent bucklers. Pectoral membranes thick, opaque; ventral surfaces of ray tips with well-developed fleshy pads. Color in preservative. Freshest material has BRADBURY: FISH GENUS OGCOCEPHALUS dorsal surface of body uniformly brown except for darker brown longitudinal stripes, one on each side as in darwini, beginning anteriorly on dorsal surface of disk behind eyes as an elongate ‘*shoulder’’ blotch which then narrows poste- riorly and trails back along sides of tail where sometimes interrupted in one or two places. Garman’s four specimens faded (Garman 1899), but holotype (Hubbs 1958) shows the longitu- dinal markings, and original description refers to markings. Markings on faces of fresh specimens also re- semble markings in darwini; suborbital space (between eye and mouth) dark except for a markedly pale stripe descending from eye to corner of mouth. Lips the same creamy-white shade as ventral surface of body; pelvics and ventral surfaces of pectorals also creamy white. Dorsal surface of pectorals dusky, grading dis- tally to black tips. Anal often tipped with black, especially in small specimens. Dorsal dusky, occasionally blotched with dark pigment. Iris golden with very dark spots in a ring around the eccentrically shaped pupil (Garman described the iris as having “‘radiating bars of brown’’). Caudal as in generic description. Color in life. Two 35-mm color slides provided by Dr. Robert Lea show that, as in other species of Ogcocephalus, O. porrectus has considerable bright-reddish coloring. One slide shows entire ventral surface of body to be orange-red except for ventral surfaces of pelvic and anal fins, which are whitish. The other slide, a dorsal view of body, shows skin around gill openings to be reddish and fin rays nearest sides of pectoral and caudal fins to have their tips scarlet. DISTRIBUTION.—Vicinity of Cocos Island from depths of 88—146 m. Hubbs (1958) erred in stating that the material described by Garman (1899) from ALBATROSS station 3368 was from “south of the Gulf of Panama, in the vicinity of Cabo Corrientes, Colombia.’’ The coordinates for this station (5°32’45”N and 86°54’30’W, cor- rectly quoted by Prof. Hubbs) designate a lo- cality near Cocos Island, not the coast of Co- lombia. But this small error may foreshadow things to come. A specimen from Peru, USNM 200363 (data given below) agrees fairly well with O. porrectus in squamation, color pattern, and morphometric characters. Whether a population of batfishes that properly may be assigned to O. porrectus occurs along the coast of western South America is a question that awaits future work. 283 MATERIAL EXAMINED.—Numbers in parentheses a n- bers of specimens. Cocos Island: MCZ 28733 (lectotyp: d MCZ 41594 (3 paralectotypes) both from ALBATROSS 3368, 5°32'45"N, 86°54'30"W, 121 m. The following from R/V SEARCHER cruise 72-4: LACM 32263 (23), 5°33'32’N, 87°04'44"W, 110 m; LACM 32264 (6), 5°33'30’N, 87°05'50’W, 137-146 m; LACM 32268 (1), 5°31'10”N, 87°01'58”W, 88-91 m; LACM 32269 (1), 5°28’30"N, 87°04'00”W, 119-121 m. Peru: USNM 200363 (1), Caleta Cruz (Tumbes), 37-55 m. LITERATURE CITED ANDERSON, W. W. 1956. January to April distribution of the common shrimp on the south Atlantic continental shelf. U.S. Fish Wildl. Serv., Spec. Sci. Rep. Fish. 171:1-14. ANONYMOUS. 1976. Catalogo de peces marinos Mexicanos. Secretaria de Pesca Instituto Nacional de Pesca. 462 p. BEEBE, W., AND J. TEE VAN. 1928. The fishes of Port-au- Prince, Haiti, with a summary of the known species of ma- rine fish of the island of Haiti and Santo Domingo. Zoolog- ica 10(1):1-279, figs. Biocn, M. E. 1787. Ichthyologie, ou Histoire naturelle, gé- nérale et particuliére des poissons. Pt. 4 (de la Garde ed.). Berlin. 134 p. BOHLKE, J. E., AND C. C. G. CHAPLIN. 1968. Fishes of the Bahamas and adjacent tropical waters. Livingston Publ. Co., Wynnewood. 771 p. BRADBURY, M. G. 1967. The genera of batfishes (Ogcoce- phalidae). Copeia 1967(2):399—422. BREDER, C. M. 1929. Field book of marine fishes of the At- lantic coast from Labrador to Texas. G. P. Putman Sons. 332 p., figs. 1949. On the relationship of social behavior to pig- mentation in tropical shore fishes. Bull. Am. Mus. Nat. Hist. 94:87-106, 8 pls. Briccs, J. C. 1958. A list of Florida fishes and their distn- bution. Bull. Florida St. Mus. Biol. Sci. 2(8):223-318. 1961. Emendated generic names in Berg’s classifi- cation of fishes. Copeia 1961(2): 161-166. BROWNE, P. 1756. The civil and natural history of Jamaica, vol. 8. 509 p., 49 pls. BuLuis, H. R., JR., AND J. R. THOMPSON. 1965. Collections by the exploratory fishing vessels Oregon, Silver Bay, Com- bat, and Pelican made during 1956-1960 in the southwest- ern North Atlantic. U.S. Fish Wildl. Serv., Spec. Sci. Rep. Fish. 510:1—130. CARVALHO, J. P. 1943. Nota preliminar sobre a fauna ictiolo- gica do litoral sul do Estado de Sao Paulo. Bol. Industria Animal 150:27-80. CASTELNAU, F. DE. 1855. Animaux nouveaux ou rares re- cueillis pendant |’expédition dans les parties centrales de 1’ Amérique du Sud, de Rio de Janeiro a Lima, et de Lima au Para. Poissons. Paris. 112 p., 50 pls. CERVIGON M., F. 1966. Los peches marinos de Venezuela. Tomo II (Monografia No. 12). Estacion de Investigaciones Marinas de Margarita Fundacion La Salle de Ciencias Natu- rales. Caracas. Pp. 449-951. Cuvier, G. 1816. Le regne animal distribué d’apres son or- ganisation, . . . Poissons, vol. 2. Paris. 532 p. 1829. Le régne animal distribué d’apres son organi- sation, . . . Ed. 2. Poissons, vol. 2. 406 p. , AND A. VALENCIENNES. 1837. Histoire naturelle des poissons, vol. 12. Pp. i-xxiv, 1-507. Paris. DAHL, G. 1971. Los peces del norte de Colombia. Inst. De- sarrollo de los Recursos Nat. Renov., Bogota. 392 p. 284 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 7 DAHLBERG, M. D. 1975. Guide to coastal fishes of Georgia and nearby states. Univ. Georgia Press. 186 p. Dekay, J. E. 1842. Zoology of New York. Pt. 4, Fishes. Albany, N.Y. 415 p. (plates in a separate volume). DELSMAN, H. C. 1941. Pisces. Mem. Mus. Roy. Hist. Nat. Belgique, sec. 2, fasc. 21, Résultats scientifiques des Croi- sieres du Navire-Ecole Belge 3(3):65—77. ERDMAN, D. S. 1956. Recent fish records from Puerto Rico. Bull. Mar. Sci. Gulf Caribb. 6(4):315—340. EVERMANN, B. W., AND W. C. KENDALL. 1900. Check-list of the fishes of Florida. Rep. U.S. Comm. Fish Fish. 25:35— 103. FISCHER, G. 1813. Zoognosia, tabulis synopticis illustrata. Ed. 3. Vol. 1. Moscow. 465 p. Fow er, H. W. 1906. Some cold-blooded vertebrates of the Florida Keys. Proc. Acad. Nat. Sci. Phila. 58:77-113. 1915. Cold-blooded vertebrates from Florida, the West Indies, Costa Rica, and eastern Brazil. Proc. Acad. Nat. Sci. Phila. 67:244-269. 1926. Fishes from Florida, Brazil, Bolivia, Argen- tina, and Chile. Proc. Acad. Nat. Sci. Phila. 78:249-285. . 1941. A list of fishes known from the coast of Brazil. Arquiv. Zool. Sao Paulo 3(6):115—184. . 1945. A study of the fishes of the southern Piedmont and coastal plain. Acad. Nat. Sci. Phila. Monogr. 7:1—408. 1947. Notes on Bahama fishes obtained by Mr. Charles G. Chaplin in 1947, with descriptions of two new species. Not. Nat. 199. 14 p. 1952. Fishes from deep water off southern Florida. Not. Nat. 246. 16 p. GARMAN, S. 1896. Report on the fishes collected by the Ba- hama expedition of the State University of lowa, under Pro- fessor C. C. Nutting, in 1893. Bull. Lab. Nat. Sci. St. Univ. Iowa 4:77-93. . 1899. Reports on an exploration off the west coasts of Mexico, Central and South America, and off the Gala- pagos Islands, by the U.S. Fish Commission steamer *‘Al- batross”’ during 1891. XXVI. The Fishes. Mem. Mus. Comp. Zool. Harvard 24:1—431, 97 pls. GILL, T. 1862. Catalogue of the fishes of the eastern coast of North America, from Greenland to Georgia. Proc. Acad. Nat. Sci. Phila. 1861(1862)(Suppl.): 1-63. 1873. Catalogue of the fishes of the east coast of North America. Smithson. Misc. Collect. 283. 50 p. . 1878. Note on the Maltheidae. Proc. U.S. Natl. Mus. 1:231—232. 1883. Supplementary note on the Pediculati. Proc. U.S. Natl. Mus. 5:551-556. GoopE, G. B., AND T. H. BEAN. 1879. Catalogue of a col- lection of fishes sent from Pensacola, Florida, and vicinity, by Mr. Silas Stearns, with descriptions of six new species. Proc. U.S. Natl. Mus. 2:121-156. , AND 1882. A list of the species of fishes re- corded as occurring in the Gulf of Mexico. Proc. U.S. Natl. Mus. 5:234-240. , AND 1896. Oceanic Ichthyology. Special Bull. U.S. Natl. Mus. 2:1-555, atlas with 123 pls. GUNTER, G., R. H. WiLLiaAmMs, C. C. DAvis, AND F. G. W. SMITH. 1948. Catastrophic mass mortality of marine ani- mals and coincident phytoplankton bloom on the west coast of Florida, November 1946 to August 1947. Ecol. Monogr. 18(3):309-324, GUNTHER, A. 1861. Catalogue of the acanthopterygian fishes in the collection of the British Museum. British Museum (Natural History), London. 586 p. 1880. Report on the shore fishes. Report on the sci- entific results of the voyage of H.M.S. Challenger during the years 1873-76. Zoology I(pt. 6): 18-82. HENSHALL, J. A. 1891. Report upon a collection of fishes made in southern Florida during 1889. Bull. U.S. Fish Comm. 1889, 9:371—-389. 1895. Notes on fishes collected in Florida in 1892. Bull. U.S. Fish Comm. 1894, 14:209-221. HERALD, E. S. 1972. Fishes of North America. Doubleday, New York. 256 p. Herre, A. W. C. T. 1942. Notes on a collection of fishes from Antigua and Barbados, British West Indies. Stanford Univ. Publ. Univ. Ser. Biol. Ser. 7(2):287-305. HILDEBRAND, H. H. 1954. A study of the brown shrimp (Penaeus aztecus Ives) grounds in the western Gulf of Mex- ico. Publ. Inst. Mar. Sci. Univ. Texas 3(2):233-366. 1955. A study of the fauna of the pink shrimp (Pe- naeus duorarum) grounds in the Gulf of Campeche. Publ. Inst. Mar. Sci. Univ. Texas 4(1): 169-232. HoeseE, H. D., AND R. H. Moore. 1977. Fishes of the Gulf of Mexico, Texas, Louisiana, and adjacent waters. Texas A&M Univ. Press. 327 p. Hoi, A. 1957. Specimina Linnaeana i Uppsala bevarade Zoologiska samlingar fran Linneéstid. [In Swedish with sum- mary.] Uppsala Univ. Arsskr. 1957(6): 1-68. Husss, C. L. 1958. Ogcocephalus darwini, a new batfish endemic at the Galapagos Islands. Copeia 1958(3): 161-170. JORDAN, D. S. 188Sa. A catalogue of the fishes known to inhabit the waters of North America, north of the Tropic of Cancer, with notes on the species discovered in 1883 and 1884. Rep. U.S. Comm. Fish Fish. 1884:789-973. . 1885b. List of fishes collected at Key West, Florida, with notes and descriptions. Proc. U.S. Natl. Mus. 7:103— 150. . 1885c. List of the fishes from Egmont Key, Florida, in the Museum of Yale College, with descriptions of two new species. Proc. Acad. Nat. Sci. Phila. 1884:42—46. 1895. The fishes of Sinaloa. Stanford Univ. Publ., Contrib. Biol. Hopkins Lab. Biol. 1:377—514. (Published simultaneously in Proc. Calif. Acad. Sci., ser. 2, 5:377—514.) 1899. A manual of the vertebrate animals of the northern United States including the district north and east of the Ozark mountains, south of the Laurentian hills, north of the southern boundary of Virginia, and east of the Mis- souri river, inclusive of marine species. Ed. 8. Chicago. 397 p. , AND B. W. EVERMANN. 1896. A check-list of the fishes and fish-like vertebrates of North and Middle Amer- ica. Rep. Comm. Fish Fish. 1895, Append. 5:207—584. , AND 1898. The fishes of North and Middle America. Bull. U.S. Natl. Mus. 47(pt. 3):2183-3136 and (pt. 4):i-ci, 3137-3313, pls. I-cccxcl. 5 , AND H. W. CLarK. 1930. Checklist of the fishes and fish-like vertebrates of North and Middle Amer- ica north of the northern boundary of Venezuela and Co- lombia. Rep. U.S. Comm. Fish. 1928 (pt. 2): 1-670. , AND C. H. GILBERT. 1882. Synopsis of the fishes of North America. Bull. U.S. Natl. Mus. 16:1—1018. , AND J. SWAIN. 1885. Notes on fishes collected by David S. Jordan at Cedar Keys, Florida. Proc. U.S. Natl. Mus. 7:230-234. BRADBURY: FISH GENUS OGCOCEPHALUS LINNAEUS, C. 1758. Systema naturae. Vol. 1. Ed. 10. Hol- miae. 824 p. LoNGLEY, W. H., AND S. H. HILDEBRAND. 1940. New gen- era and species of fishes from Tortugas, Florida. Pap. Tor- tugas Lab. 32(Carnegie Inst. Washington Publ. 517):223- 285. , AND 1941. Systematic catalog of the fishes of Tortugas, Florida, with observations on color, habits, and local distribution. Pap. Tortugas Lab. 34 (Carnegie Inst. Washington Publ. 535):1—331. LONNBERG, A. J. E. 1896. Linnean type-specimens of birds, reptiles, batrachians, and fishes in the Zoological Museum of the R. University in Upsala. K. Sven. Vetenskapakad. Handl. 22(4)(1):1-45. Lowe (McCoNNELL), R. H. 1962. The fishes of the British Guiana continental shelf, Atlantic coast of South America, with notes on their natural history. J. Linn. Soc. London Zool. 44(301):669-700. Lunpby, W. E. 1956. Galapagos produces the *‘Thing.’” Nat. Hist. 65:468—469. LUTKEN, C. 1866. Ichthyologiske notiser. II. Om arterne af slaegten Malthaea og saerligt om M. notata (truncata). Vidensk. Medd. Naturhist. Foren. Kjgbenhavn 1865:205— 223. MARCGRAVE, G. 1648. Historiae rerum naturalium Brasiliae. Batavia and Amsterdam. 293 p. plus an index of 7 unnum- bered pages. MARTIN S., F. 1956. Ictiologia del archipiélago de Los Ro- ques. Pp. 87-144 in El archipiélago de Los Roques y La Orchila por la Sociedad de Ciencias Naturales La Salle. Caracas. MCALLISTER, D. E. 1968. Evolution of branchiostegals and associated opercular, gular, and hyoid bones and the clas- sification of telestome fishes, living and fossil. Bull. Natl. Mus. Canada 221 (biol. ser. 77):1—239. MEEK, S. E., AND S. F. HILDEBRAND. 1928. The marine fishes of Panama. Field Mus. Nat. Hist. Publ. 249, zool. ser. 15:709-1045. MENEZES, N. 1964. Sobre ogcocephalideos das costas do Brazil (Pisces, Ogcocephalidae). Papeis Avulsos do Depar- tamento de Zoologia 16(16):153-171. MIRANDO RiBEIRO, A. DE. 1915. Fauna Brasiliense—Peixes. Physoclisti. Pt. 5. Arch. Mus. Nac. Rio de Janeiro 17:1- 679. . 1918. Summario. Fauna Brasiliense—Peixes. Physo- clisti. Pt. 5. Arch. Mus. Nac. Rio de Janeiro 21:1—227. MitcuHiLt, S. L. 1818. Dr. Mitchill’s memoir on the fishes of New-York. Am. Monthly Mag. Critical Rev. 2(5):321-328. Mog, M. A., Jr., P. C. HEEMSTRA, J. E. TYLER, AND H. WAHLQUIST. 1966. An annotated listing of the fish refer- ence collection at the Florida Board of Conservation Marine Laboratory. Florida Bd. Conserv. Mar. Lab. Spec. Sci. Rep. 10, 121 p. [mimeo.]. , AND G. T. MARTIN. 1965. Fishes taken in monthly trawl samples offshore of Pinellas County, Florida, with new additions to the fish fauna of the Tampa Bay area. Tulane Stud. Zool. 12(4):129-151. 285 PaRRA, A. 1787. Peces y crustaceos de la Isla de Cuba. Ha- vanna. [Not seen; pages 11-13 in a volume of pages hand- copied from Parra by Senorita Poey in the Library of the California Academy of Sciences were used.] Puyo, J. 1936. Contribution a l'étude ichthyologique de la Guyane frangaise—Peches et Pecheries. Bull. Soc. Hist. Nat. Toulouse 70. 258 p. . 1949. Faune de l’Empire frangais. 12. Poissons de la Guyane frangaise. Office de la Recherche Scientifique outre-Mer. Paris. 280 p. RANDALL, J. E. 1967. Food habits of reef fishes of the West Indies. Stud. Trop. Oceanogr. Miami 5:665—847. . 1968. Caribbean Reef Fishes. T.F.H. Publ., New Jer- sey. 318 p. Reip, G. K. 1954.. An ecological study of the Gulf of Mexico fishes in the vicinity of Cedar Key, Florida. Bull. Mar. Sci. Gulf Caribb. 4(1): 1-94. RICHARDSON, J. 1836. Fauna Boreali-Americana; or the zo- ology of the northern parts of British America. Pt. 3. The Fish. London. 327 p., pls. 74-97. ROSENTHAL, F. 1822. Ichthyotomische Tafeln. Section 2, Pt. 4. Berlin. SCARABINO, S. M. DE. 1974. Sobre la presencia de ‘‘pez mur- cielago’’ Ogcocephalus vespertilio (Linné) (Pisces, Ogco- cephalidae) en la boca de la Plata. Rev. Inst. Invest. Pes- queras 2(3):306—313. SesBA, A. 1734. Locupletissimi rerum naturalium thesauri ac- curata descriptio et iconibus artificiosissimis expressio, per universam physicis historiam .... Vol. 1. Amstelaedami. SHAW, G. 1804. General zoology 5(2):251—463. SmitH, H. M. 1907. The fishes of North Carolina. North Carolina Geol. Econ. Surv. 2:1-453. Soto, A. DE. 1922. A batfish from the Amazon. Copeia (108):51. SPRINGER, S., AND H. R. BuLLis, JR. 1956. Collections by the Oregon in the Gulf of Mexico. U.S. Fish Wildl. Serv. Spec. Sci. Rep. Fish. 196:1—134. SPRINGER, V. G., AND A. J. MCERLEAN. 1962. Seasonality of fishes on a south Florida shore. Bull. Mar. Sci. Gulf Caribb. 12(1):39-60. , AND K. D. WoopsBurn. 1960. An ecological study of the fishes of the Tampa Bay area. Florida State Bd. Cons. Mar. Lab., Prof. Pap. Ser. 1. 104 p. STarK, W. A., II. 1968. A list of fishes of Alligator Reef, Florida, with comments on the nature of the Florida reef fish fauna. Undersea Biol. 1(1):4—40. Storer, D. H. 1846. A synopsis of the fishes of North Amer- ica. Mem. Am. Acad. Arts Sci. Boston 2(7):44-298. Storey, M., AND E. W. GupDGER. 1936. Mortality of fishes due to cold at Sanibel Island, Florida, 1886-1936. Ecology 17(4):640-648. UHLER, P. R., AND O. LuUGGER. 1876. List of fishes of Mary- land. Rep. Comm. Fish Maryland 1876:67-176. WALLs, J. G. 1975. Fishes of the northern Gulf of Mexico. T.F.H. Publ., New Jersey. 432 p. Woops, L. P. 1942. Rare fishes from the coast of Texas. Copeia 1942(3):191. CALIFORNIA ACADEMY OF SCIENCES Golden Gate Park San Francisco, California 94118 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF; CIENCES 1 8 1960 Vol. 42, No. 8, pp. 287-293, 5 figs. rr 8 ime ee arine Bledagical:taboraicy, LIBRARY XENAPLOACTIS, A NEW GENUS FOR PROSOPODASYS ASPERRIMUS GUNTHER (PISCES: APLOACTINIDAE), WITH DESCRIPTIONS OF TWO NEW SPECIES By Stuart G. Poss Division of Biological Sciences and Museum of Zoology, University of Michigan, Ann Arbor, Michigan 48109 and William N. Eschmeyer California Academy of Sciences, Golden Gate Park, San Francisco, California 94118 ABSTRACT. A new genus, Xenaploactis, is created for Prosopodasys asperrimus Gunther, 1860, which is re- described. X. anopta from Luzon Island in the Philippines and X. cautes from the Andaman Sea and the Gulf of Thailand are described as new. These species exhibit differences in the configuration of ridges between the eyes, body depth, head pores, and other features. Species of Xenaploactis differ from those of other genera of the Aploactinidae by the presence of several features in combination: 3 anterior dorsal fin spines forming a separate fin, rather sharp head spines, a markedly upturned mouth, and a body densely covered with modified pointed scales. INTRODUCTION Gunther (1860) described Prosopodasys as- perrimus, assigning it to a genus otherwise com- posed of tetrarogid scorpaenoids. Because of this and because the species remains known from only the holotype, Prosopodasys asperri- mus has not been generally recognized as an aploactinid scorpaenoid. Gunther described the head and body as ‘“covered with small prickles,’ and this caused Poss and Eschmeyer (1978) to suspect that this species was an aploactinid. Other features men- tioned in Gunther’s brief description—absence of palatine teeth, presence of one spine and three soft rays in the pelvic fins, and presence of flexible anal spines—tended to support this view. More recent examination of the holotype confirms this suspicion. Species assigned to Prosopodasys, a name originally proposed by Cantor (1849:1026) as a replacement name for Apistus Cuvier and Val- enciennes, have been placed into a number of genera. Prosopodasys asperrimus does not be- long in any of these nor does it belong in any existing genus of aploactinid. Two undescribed and closely related species have been discovered among specimens in the collections of the National Museum of Natural History (USNM) and the California Academy of Sciences (CAS). Together with Prosopodasys asperrimus, they form a natural cluster quite distinct from other aploactinids. Provided below [287] 288 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 8 are descriptions of these species and a new ge- nus to contain them. METHODS The methods used in this study follow those of Eschmeyer (1969) as modified by Poss and Eschmeyer (1978). Spines and rays are difficult to distinguish and median fin-ray counts were checked against radiographs. The last fin ray in the dorsal and anal fins is double, borne on a single pterygiophore, and counted as one ray. Xenaploactis new genus TYPE-SPECIES.—Prosopodasys asperrimus Giinther, 1860. DiaGNosis.—Aploactinid fishes with spinous dorsal fin in 2 parts; the first 3 dorsal fin spines inserting on cranium and forming a separate fin, widely separate from the 4th spine of the second part of the spinous dorsal fin; rather pungent head spines, including 2 prominent preorbital spines and strong spine on lateral face of third infraorbital bone; head and body densely cov- ered with modified scales; mouth strongly up- turned. Maxillary with prominent angular point at end of anteroventral margin, with ridge on lateral face. Dorsal III, X, 8—9 (last double) or III, XI, 8—9 (last double). Anal I, 9-10 (last dou- ble). Pelvic I, 3. Pectoral 13-14. Vertebrae 27- 28. Branchiostegal rays 6. Four dorsal spines and associated pterygiophores anterior to third neural spine. Upper posterior margin of oper- cular bone very close to base of dorsal fin. Teeth on jaws and vomer, none on palatines. DESCRIPTION.—(See also species descriptions below.) Dorsal fin in 2 parts, originating on cra- nium above posterior border of eye; first 3 spines close together, widely separate from 4th spine, which originates just anterior to end of head; 2nd spine longest, all spines rather weak, flexible, without cirri. Dorsal fin membrane of second part of fin notably incised, dorsal spines free from fin membrane at about midlength; de- gree of fin incision decreases posteriorly. Pec- toral fin rays all unbranched, longest 5—7 from above, reaching just past anus. Head covered with modified scales, scales ab- sent in interorbit and behind eye. Mouth strong- ly upturned. Movable lachrymal bone (infraor- bital 1) bladelike, with 3 spines; Ist small, directed anteriorly, followed by 2 prominent sharp spines, 2nd spine about equal in length to 3rd, directed downward; 3rd spine directed down and slightly back. Third infraorbital with prominent spine on ventral margin of bone pro- jecting ventrally and laterally; lateral face of bone with prominent ridge and 2 blunt spines. Infraorbital (suborbital) stay appearing as a strong ridged bar. One postorbital bone (infraor- bital 4 or 5). Interorbit with prominent ridges. Nasal bones tubed, without spines. Preopercle with 5 rather strong spines, uppermost largest, diminishing in size ventrally. Upper arm of pre- opercle forming strong ridge. Opercle with 2 weak ridges, lower ending in small blunt spine on opercular margin. Opercular flap extending nearly to base of dorsal fin between spines 4 and 5. Interopercle forming spinous projection on posterior opercular margin. Posterior dorsal border of cleithrum ending in small, poorly de- fined blunt spine. Angular bone prominent, strongly jutting ventrally with mouth closed. Maxillae reaching anterior border of eye, with ridged dorsal border, ridge near ventral border. Maxillary cirrus minute. Pelvic fin origin slightly in advance of lower- most pectoral fin rays. Pelvic fin membrane not adnate to body. Caudal fin rounded, with 18 total fin-ray elements (9 upper and 9 lower), all un- branched. Caudal skeleton with parahypural, Ist and 2nd hypurals fused; 3rd and 4th hypurals fused; Sth hypural small, autogenous; Ist preural neural spine long; 2 epurals. Seven upper and 7 lower fin-ray elements are attached to the hy- purals, 2 (procurrent) rays are free above and below. ETYMOLOGY.—The generic name Xenaploac- tis is derived from the Greek xenos (stranger) + Aploactis (a related genus). Xenaploactis is feminine. Key to the Species of Xenaploactis la. Interorbit with ridges nearly parallel (Fig. 4). Body depth less than % of standard length. Dorsal fin III, 1X, 8-9 (based on limited material; some variation to be ex- pected) ___ X.. cautes (Figs. 4 lower and 5) 1b. Interorbit with ridges divergent anterior- ly, convergent over middle of orbit, di- vergent posteriorly (Figs. 2 and 4 upper). Body depth equal to or greater than 4% of standard length. Dorsal fin III, X, 8-9 (based on limited material; some variation tobe expected). 2) aa 2 2a. Second infraorbital bone with 1 or 2 POSS & ESCHMEYER: NEW APLOACTINID GENUS AND SPECIES 289 FIGURE 1. spines. Pore of infraorbital lateral line ca- nal at second infraorbital bone as simple obscure pore. Body depth greater than 4 of standard length. No fingerlike cirri above uppermost preopercular lateral line pores. Dorsal fin III, X, 9. Anal I, 10 (based on limited material; some variation WMEPCECROCEICO) so a2 X. asperrima (Figs. 1 and 2) 2b. Second infraorbital bone without spines. Pore of infraorbital lateral line canal at second infraorbital bone as prominent elongate slit. Body depth equal to 4% of standard length. Fingerlike cirri present above uppermost preopercular lateral line pores. Dorsal III, X, 8. Anal I, 9 (based on limited material; some variation to be EN DEC IS 1) Ieee aR ee Sees oe eee Xenaploactis asperrima (Gunther) (Figures 1 and 2) Prosopodasys asperrimus GUNTHER, 1860:140-141 (original description; type-locality East Indies). MATERIAL.—Holotype: BMNH 1979.5.5:1 (39.9 mm SL). East Indies, Sir E. Belcher, no other data. Counts.—Dorsal fin III, X, 9 (last double). Anal fin I, 10 (last double). Pectoral fin 13 (left), 14 (right). Pelvic fin I, 3. Lateral line scales 10 (left), 11 (right). Vertebrae 27. DESCRIPTION.—(See also generic diagnosis above.) Body notably elevated behind head, Lateral view of holotype of Prosopodasys asperrimus (=Xenaploactis asperrima) (BMNH 1979.5.5:1, 39.9 mm SL). Specimen formerly dried, somewhat distorted. body depth more than % of standard length. Body densely covered with modified scales which form spinous points, best developed on upper back behind head. Lateral line with 10-11 tubed scales, each with 2 small laterally pro- jecting spinules, best developed anteriorly, last scale extending over base of caudal fin. Gill rak- ers short, difficult to count, total 8-10; 3 on up- per arch, S—7 on lower arch. No modified scales on snout. Lachrymal bone (infraorbital 1) with 2 large spines, first notably curved. A small spine in front at base of first spine, a small spine at base of second spine. Second infraorbital bone with a small double or single spine, with obscure small circular lateral iine pore. Third infraorbital bone with a large spine directed out and down; a strong ridge attached to preopercle. Mouth very strongly upturned, nearly vertical. Interorbit with prominent ridges, divergent an- teriorly, convergent over middle of interorbit, divergent posteriorly (Fig. 2). Postocular spine appearing as a sculptured ridge, weakly con- nected to supraorbital ridge. Parietal spine lumplike. Pterotic spine as a strong, slightly curved ridge. Posttemporal spine well ossified, sculptured, appearing as a ridged lump, followed by small bladelike supracleithral spine. Dorsal posterior border of cleithrum appearing as a marked ridge, ending in a blunt, poorly defined spine. Preopercular lateral line pores simple, no fingerlike cirri above uppermost pores. Ventral surface of dentary without distinct cirri. Color in life unknown. Color of head and body 290 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 8 FiGuRE 2. Dorsal view of head of holotype of Prosopo- dasys asperrimus (=Xenaploactis asperrima). in preservative light brown, fins slightly darker and possibly speckled in life. Measurements in millimeters as follows (per- cent standard length in parentheses): standard length 39.9; head 13.2 (34); snout 3.5 (9); orbit 3.0 (7); interorbital width 2.2 (5); jaw 6.0 (15); postorbit 7.0 (17); body depth 14.6 (37); predor- sal 6.2 (15); anal fin 14.0 (35); caudal fin 9.5 (24); pectoral fin 9.2 (23); pelvic fin 4.5 (11); Ist dorsal spine 3.2 (8); 2nd dorsal spine 4.1 (10); 3rd dorsal spine 3.1 (8); 4th dorsal spine 1.8 (4): Sth dorsal spine 2.7 (7); penultimate dorsal spine 3.1 (8); last dorsal spine 3.5 (9); anal spine 2.1 (5); least depth of caudal peduncle 4.2 (10); snout to base of 2nd dorsal spine 7.2 (18); snout to base of 3rd dorsal spine 7.8 (19); snout to base of 4th dorsal spine 11.5 (29); snout to base of Sth dorsal spine 14.6 (37); width of Ist dorsal spine at midlength 0.2 (1); incision of dorsal fin membrane at 4th dorsal spine (from tip to membrane) 1.8 (4). DISTRIBUTION.—Known only from the holo- type from the ‘‘East Indies.”’ Xenaploactis anopta, new species (Figures 3 and 4 upper) No literature applies to this species. MATERIAL.—Holotype: CAS 32633 (37.0 mm SL). Philip- pines, Luzon I., Zambales, 4 km w of Calguaguin Cove, 64— 81 m, 0835-0910 hrs, J. E. Norton, 9 June 1966. Counts.—Dorsal fin III, X, 8 (last double). Anal fin I, 9 (last double). Pectoral fin 13 (left), 14 (right). Pelvic fin I, 3. Lateral line scales 10 (left), 11 (right). Vertebrae 27. DESCRIPTION.—(See also generic description above.) Body somewhat elevated behind head, body depth % of standard length. Body densely covered with modified scales which form spi- nous points, best developed anteriorly; lateral line with 10-11 tubes, each with 2 small, later- ally projecting spinules which are best devel- oped anteriorly; last scale extending over base of caudal fin. Gill rakers short, difficult to count, total 10,°3 on upper arch, 7 on lower arch. Few modified scales on snout, none on inter- orbit. Movable lachrymal bone (infraorbital one) IO mm. FIGURE 3. — Lateral view of holotype of Xenaploactis anopta (CAS 32633, 37.0 mm SL). POSS & ESCHMEYER: NEW APLOACTINID GENUS AND SPECIES 291 bladelike with 3 spines: Ist small, directed to- ward premaxilla; followed by 2 large, sharp spines, 2nd about equal in length to 3rd, di- rected downward; 3rd spine directed down and slightly back. Spine on second infraor- bital bone absent; a large elongate lateral line pore present. Mouth strongly upturned. Inter- orbit with prominent ridges, divergent anterior- ly, convergent over middle of interorbit, strong- ly divergent posteriorly (Fig. 4 upper). Postocular spine and pterotic spine ridgelike. Posttemporal spine a large well-ossified lump ending in blunt spine, followed by blunt supracleithral spine. Dorsal posterior border of cleithrum ending in a small, poorly defined blunt spine. Preopercular lateral line pores opening as small tubes, finger- like cirri above uppermost pores. In ventral view, surface of dentary with 5 small fingerlike cirri along outer margin; 5 pairs of similar cirri anteriorly, between dentaries. Color in life unknown. Color of head and body in preservative (Fig. 3) brown, with scattered black specks. Fins darker, possibly speckled in life; caudal with vertical bands. Measurements in millimeters as follows (per- cent standard length in parentheses): standard length 37.0; head 13.0 (35); snout 3.7 (10); orbit 3.3 (9); interorbital width 2.5 (6); jaw 5.3 (14): postorbital 7.3 (20); body depth 12.3 (33); pre- dorsal 6.4 (17); anal fin 14.1 (38); caudal fin 9.2 (25); pectoral fin 8.5 (23); pelvic fin 5.2 (14); Ist dorsal spine 1.9 (5); 2nd dorsal spine 3.5 (9); 3rd dorsal spine 2.4 (6); 4th dorsal spine 1.5 (4); Sth dorsal spine 2.2 (6); penultimate dorsal spine 2.2 (6); last dorsal spine 2.3 (6); anal spine 1.9 (5); least depth of caudal peduncle 4.1 (11); snout to 2nd dorsal spine 6.8 (18); snout to 3rd dorsal spine 8.4 (23); snout to 4th dorsal spine 12.9 (35); snout to Sth dorsal spine 13.7 (37); width of Ist - dorsal spine at midlength 0.3 (1); incision of dor- sal fin membrane at 4th dorsal spine (from tip to membrane) 1.5 (4). ETYMOLOGY.—The species-group name is de- rived from the Greek anoptos (unseen). DISTRIBUTION.—Known only from the type- locality in the Philippines at 64-81 m. Xenaploactis cautes, new species (Figures 4 lower and 5) No literature applies to this species. MATERIAL.—Holotype: CAS 16105 (28.0 mm SL). Gulf of Thailand, 12°19'15"N, 100°43'40’E, 28.6 km from Goh Chuang, 33 m, muddy sand bottom, MV STRANGER, 16-ft (4.9- FIGURE 4. tis anopta (upper) and X. cautes (lower). Dorsal view of head of holotypes of Xenaploac- m) otter trawl, George Vanderbilt Foundation sta. 60-449, GVF reg. no. 2724, Scripps locality 60-185 C.N. 633f.5-9a, 0117-0202 hrs, 13 Dec. 1960. Paratype: USNM 221143 (24.3). Andaman Sea, 14°07’N, 97°05’E, 69-73 m, International In- dian Ocean Expedition, ANTON BRUUN cruise 1, sta. 38, Gulf of Mexico shrimp trawl, 30 Mar. 1963. Counts.—Dorsal fin III, XI, 8-9 (last dou- ble). Anal fin I, 10 (last double). Pectoral fin 14. Pelvic fin I, 3. Lateral line scales 9-10. Verte- brae 27-28. DESCRIPTION.—(See also generic description above.) Body not notably elevated behind head, body depth less than 3 of standard length. Body covered with modified pointed scales. Lateral line with 9-10 tubed scales, each with 2 small laterally projecting spinules which are best de- veloped anteriorly, last scale extending over base of caudal fin. Gill rakers short, difficult to count, total 10-12, 3-4 on upper arch, 6-8 on lower arch. Many modified scales on snout. Lachrymal bone (infraorbital 1) with 3 spines; Ist, of mod- erate size, points mostly forward, continuous 292 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 8 10 mm FIGURE 5. with ridge at base of larger 2nd spine; 2nd spine about equal in length to 3rd, directed downward, slightly curved; 3rd spine points mostly back. Second infraorbital bone with 2 spinous points, 1 above other, with obscure circular lateral line pore. Interorbit with nearly parallel ridges, stronger posteriorly (Fig. 4 /ower). Postocular spines as slightly curved ridges, meeting at mid- line of interorbit, connected to interorbital ridges. Pterotic spine ridgelike. Posttemporal spine ridgelike, followed by blunt supracleithral spine. Cleithrum ending in small blunt spine. In ventral view, surface of dentary with 5 tiny fin- gerlike cirri along outer margin, 5 pairs of similar cirri anteriorly between dentaries. Color in life unknown. Color in preservative pale, probably strongly faded. Head and body without scattered specks. Fins not darker than body. Measurements in millimeters as follows (ho- lotype first, percent standard length in parenthe- ses): standard length 28.0, 24.3; head 9.9, 9.4 (35, 39); snout 2.4, 2.4 (9, 10); orbit 2.4, 2.4 (9, 10); interorbital width 1.8, 2.1 (6, 9); jaw 3.6, 4.2 (13, 17); postorbit 4.7, 4.9 (17, 20); body depth 7.8, 7.4 (28, 30); predorsal 5.0, 4.2 (18, 17); anal fin 11.1, 10.8 (40, 44); caudal fin 6.7, 6.2 (24, 26); pectoral fin 6.2, 5.5 (22, 23); pelvic fin 3.0, 3.4 (11, 14); Ist dorsal spine 1.7, 1.7 (6, 7); 2nd dor- sal spine 2.9, 2.6 (10, 11); 3rd dorsal spine 1.7, 1.6 (6, 6); 4th dorsal spine 1.3, 1.3 (4, 5); Sth dorsal spine 1.7, 1.5 (6, 6); penultimate dorsal spine 2.2, 1.6 (8, 6); last dorsal spine 2.3, 1.4 (8, Lateral view of holotype of Xenaploactis cautes (CAS 16105, 28.0 mm SL). 6); anal spine 1.8, 2.0 (6, 8); width between in- terorbital ridges 0.7, 0.8 (3, 3); least depth of caudal peduncle 3.3, 2.7 (12, 11); snout to 2nd dorsal spine 5.6, 5.2 (20, 21); snout to 3rd dorsal spine 6.0, 5.4 (21, 22); snout to 4th dorsal spine 8.9, 8.4 (32, 34); snout to Sth dorsal spine 10.3, 8.5 (37, 35); width of Ist dorsal spine at mid- length 0.2, 0.2 (1, 1); incision of fin membrane at 4th dorsal spine (from tip to membrane) 1.3, 1.3 (4, 5). ETYMOLOGY .—The species-group name is de- rived from the Latin cautes (a rough, pointed rock) and is to be treated as a noun in apposi- tion. DISTRIBUTION.—Known only from the type material from the Andaman Sea and Gulf of Thailand. This species appears to inhabit muddy sand bottom at depths of 33-79 m. ACKNOWLEDGMENTS We thank a number of friends and colleagues who assisted us in the course of this study: Al- wyne Wheeler and Mary Connolly (BMNH) for giving us the opportunity to examine the holo- type of Prosopodasys asperrimus,; Leslie Knapp, of the Smithsonian Oceanographic Sorting Cen- ter, for sending us the second specimen of Xe- naploactis cautes; James Gordon, Betty Powell, William Ruark, and Pearl Sonoda (all of CAS) for their continous help. Tomio Iwamoto and Lillian Dempster (CAS) reviewed the manu- script and offered critical suggestions. We also wish to thank Ellie Koon and Joanne Zupan, of POSS & ESCHMEYER: NEW APLOACTINID GENUS AND SPECIES 393 the University of Michigan, for their assistance. GUNTHER, A. 1860. Catalogue of the acanthopterygian fishes The drawings were made by Beth Meinhard in the British Museum. Vol. 2, Squamipinnes, Cirrhitidae, Triglidae, Trachinidae, Sciaenidae, Polynemidae, Trichiur- (CAS). idae, Scombridae, Carangidae, Xiphiidae. London. xxi + 548 p. LITERATURE CITED Poss, S. G., AND W. N. ESCHMEYER. 1978. Two new Aus- CANTOR, T. 1849. Catalogue of Malayan fishes. J. R. Asiatic tralian velvetfishes, genus Paraploactis (Scorpaeniformes: Soc. Bengal 18(2): 983-1443, 14 pls. Aploactinidae), with a revision of the genus and comments EscCHMEYER, W. N. 1969. A systematic review of the scor- on the genera and species of the Aploactinidae. Proc. Calif. pionfishes of the Atlantic Ocean (Pisces: Scorpaenidae). Acad. Sci. 41(18):401-426, 14 figs., 6 tables. Occas. Pap. Calif. Acad. Sci. 79. 130 p. CALIFORNIA ACADEMY OF SCIENCES Golden Gate Park San Francisco, California 94118 ee a eM Voy Vol. 42, No. 9, pp. 295-302, 6 figs. PROCEEDINGS ° OF THE ha A we | CALIFORNIA ACADEMY OF SCIENCES SUNDASALANGIDAE, A NEW FAMILY OF MINUTE FRESHWATER SALMONIFORM FISHES FROM SOUTHEAST ASIA By Tyson R. Roberts California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118 ABSTRACT: Sundasalangidae, a new family of minute freshwater salmoniform fishes related to the Salangidae or East Asian icefishes, is based upon a new genus and two new species recently discovered in Southeast Asia, Sundasalanx praecox and S. microps, both described in the present paper. With males and females sexually ripe at standard lengths of only 14.9 mm, S. praecox is the smallest known adult salmoniform and is among the smallest of all adult vertebrates. The new family differs from all other teleosts including Salangidae in the following features of its skeletal anatomy, which is largely cartilaginous: the two halves of the pectoral girdle are united to each other by a median scapulocoracoid cartilage; in branchial arches 1-3 the basibranchials and hypobranchials of either side are represented by a single cartilaginous element; and each half of the pelvic girdle is provided with a pair of rod-shaped parapelvic cartilages. Despite these and many other differences, Sundasalangidae and Salangidae are clearly closely related. They agree with each other but differ from all other known teleosts in having the jaw suspension with bilaterally paired palatohyomandibuloquadrate cartilages. March 5, 1981 INTRODUCTION Among the freshwater fishes recently collect- ed by the author in Southeast Asia are two sam- ples, one from far up the mainstream of the Ka- puas River in Kalimantan (Indonesian Borneo) and the other from a creek draining into the Tale Sap in peninsular Thailand, of minute transpar- ent teleosts with a skeleton that is almost en- tirely cartilaginous (Figs. 1-2). Superficially re- sembling drawings of early or mid-metamorphic leptocephalus larvae of Elops (cf. Gehringer 1959:figs. 10-11), closer study of the fishes in these samples reveals that they are sexually ma- ture, represent two very distinct species, and are actually salmoniforms most closely related to the East Asian icefishes of the family Salan- gidae. Yet they differ so markedly from Salan- gidae, and in certain respects from all other known teleosts, that a new family is proposed for them. METHODS AND MATERIALS; ACKNOWLEDGMENTS Observations on the largely cartilaginous skel- etal anatomy of Salangidae and Sundasalangidae have been made on specimens prepared by means of the newly developed alcian blue—aliz- arin technique for counterstaining cartilage and bone in whole, cleared specimens of small ver- tebrates (Dingerkus and Uhler 1977). I am grate- ful to Robert Drewes of the California Academy of Sciences for preparing specimens of both species of Sundasalangidae and of all of the genera and nearly half of the species of Salang- idae, and also to William N. Eschmeyer, Cu- rator of Fishes of the California Academy of Sciences, for making the specimens of Salang- idae available. A complete list of these skeletal preparations will be presented in an exten- sively illustrated account of the skeletal anat- omy of Salangidae and Sundasalangidae now [295] 296 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 9 TERA ARE fir falls dab js ity RERAA BAS CSR Se Saal ‘ net (a) Sundasalanx praecox, based mainly on 17.0-mm paratype, MCZ 54390; (b) Sundasalanx microps, based mainly on 17.0-mm paratype, CAS 44220. * FIGURE 1. ROBERTS: SUNDASALANGIDAE—NEW FISH FAMILY being prepared (Roberts, ms). I also thank Kar- sten E. Hartel and William Fink for loaning the specimens of Sundasalanx praecox which had been deposited by me in the Museum of Com- parative Zoology at Harvard. My fieldwork in Thailand and Indonesia which led to the discov- ery of Sundasalanx was supported or aided by the following institutions: Museum of Compar- ative Zoology; Kasetsart University College of Fisheries; National Research Council, Thailand; Indonesian National Research Council (LIPI); and Smithsonian Tropical Research Institute (STRI). I particularly thank Supap Monkolprasit of Kasetsart and Ira Rubinoff of STRI for facil- itating my travel and fieldwork. The preparation of this paper was supported by grant DEB77- 24574 in the Systematic Biology Program of the National Science Foundation. SUNDASALANGIDAE, new family TYPE-GENUS: Sundasalanx, new genus. Sundasalangids differ from all other teleosts, so far as known, in having a pectoral girdle with a median cartilaginous scapulocoracoid and a single pair (one on each side) of fan-shaped, ““externalized’’ radial cartilages (Fig. 3) which form the peduncular portion of the pedunculated pectoral fins; a pair of rod-shaped parapelvic cartilages (Fig. 2) which apparently serve to anchor each half of the pelvic girdle to the free ventral end of a myotomal muscle; and branchial arches with hypobranchial elements 1-3 absent as separate elements, evidently fused to basi- branchials 1-3 (Fig. 4). In Salangidae the scap- ulocoracoids are paired, separate elements, and the two halves of the pectoral girdle are separate from each other; each pectoral fin is supported by three or more radial cartilages; parapelvic cartilages are absent; and the first three bran- chial arches have separate basibranchial and hy- pobranchial elements. Members of the Sundasalangidae agree with most Salangidae but apparently differ from all other teleosts in having a jaw suspension con- sisting of a single cartilaginous element or pal- atohyomandibuloquadrate (Fig. 5); they agree with Salangidae but apparently differ from adults of all other known teleosts (Nelson 1960:61) in having well-developed separate fourth hypobranchials (Fig. 4). Sundasalangids agree with the Salangidae but differ from adults of most other teleosts in having pedunculate pectoral fins; a scaleless body; no symplectics; 297 no circumorbital bones; myotomal muscles fail- ing to meet at ventral midline of body; and max- illary bones with distal two-thirds curved in- wards underneath the head so that the portion of the maxillary toothrow they bear projects medially rather than ventrally when the mouth is closed. In addition to the unique characters associated with their pectoral and pelvic girdles and gill arches, which have been described above, Sundasalangidae differ from Salangidae in their much smaller size (the smallest salangids are over 35 mm in standard length [SL] when sexually mature); olfactory organs each with a single nasal opening instead of two openings; interopercle absent; pectoral fin without seg- mented bony rays; adipose fin absent; sexually mature males without enlarged or otherwise modifed anal fin, or a row of large pored scales on the base of the anal fin (present in all salan- gids); pelvic fins five-rayed (seven-rayed in all salangids); myotomes <-shaped (3-shaped in salangids); and vertebrae only 37-43 (48-77 in Salangidae). Sundasalangidae comprises two species, Sundasalanx praecox and §. microps, described below. Sundasalanx, new genus TyYPE-SPECIES: Sundasalanx praecox, new species. Minute, transparent, freshwater fishes with a largely cartilaginous skeleton; a single large na- sal opening on each side of snout; body bilat- erally compressed, scaleless; branchiostegal rays four (usually three in Salangidae); pectoral fins pedunculate, without bony rays; pelvic fins midabdominal, with five rays; myotomal mus- culature of opposite sides widely separated ven- trally; each half of pelvic girdle anchored to free ventral end of a myotomal muscle by a pair of parapelvic cartilages; a median membranous keel extending on abdomen from pelvic fins to vent; gut a simple straight tube with no differ- entiated stomach; vent immediately anterior to anal fin origin; no secondary sexual dimorphism or dichromatism; vertebrae (including hypural centrum as one) 37-43. Dentition (Figs. 4-5): Premaxillary and max- illary with a single row of minute conical teeth; lower jaw with two rows of minute conical teeth, inner row curved inwards away from outer row; fifth ceratobranchials with 0-10 conical teeth. Roof of mouth and upper pharyngeal elements edentulous. Median fins: Dorsal and anal fins originating 298 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 9 Ny ‘ Nal i i | | \i | : Al) ae SS VB, 3 (\ NYU 5 \NIGZ ae wh \ Ze 2 \ Nia é ENS Ke f\ \ \ \ P \ \ \ \ q f = . \ 7 \ \ y \ : ‘ \SX SS SS yd — NS Nuss ‘ ~ Y 4 Ne vA S17 >} — ™ Sia ml bho / \ { rs if SO 4 £ Vea | f <—<— LE — FiGuRE 2. (a) Sundasalanx praecox, MCZ 54390, 17.0 mm: (b) Sundasalanx micr te ddd ebb lUll ROBERTS: SUNDASALANGIDAE—NEW FISH FAMILY on posterior third of body; origin of anal fin on a vertical through or slightly posterior to middle of dorsal fin base. Dorsal fin with three simple and 8-11 branched rays (last ray, counted as one, split to base); anal fin with 3—4 simple rays and 12-17 branched rays (last ray split to base). Adipose fin absent. Caudal fin moderately forked, with 10 + 9 principal rays and 9-12 + 7-11 procurrent rays. ETYMOLOGY.—Sunda (=Sundaland, the con- tinental landmass of Southeast Asia connected to the Asian mainland by the isthmus of Kra) plus Salanx (Greek, masc.), type-genus of the family Salangidae. Sundasalanx praecox, new species (Figures la, 2a, 4a, Sa, 6a) MATERIAL.—Holotype: MCZ 47129, 17.2-mm male with well-developed testes, from Khlong Falamee, a swift, muddy creek 1-2 m deep and 3-5 m wide with hard-packed mud bottom flowing into inner lake of Tale Sap, at about 2 km W of Pak Payoon on the isthmus of Kra, southern Thailand; ny- lon flyscreen pushnet; 20 June 1970. Paratypes: MCZ 54390, 119: 14.9-18.3 mm, same collection data as holotype; nine utilized for alcian blue—alizarin prepa- rations. DIAGNOSIS.—S. praecox is distinguished from S. microps, its only congener, by its much larger eyes: horizontal diameter of eye measured in 10 specimens including smallest and largest speci- men of each species, 4.3—5.0% of SL in S. prae- cox vs. 2.7—3.2% in S. microps. In S. praecox eyeballs separated from each other by a distance about equal to transverse diameter of their pig- mented portion, while in S. microps distance separating them equal to at least twice trans- verse diameter. In S$. praecox head deeper, more compressed, and nasal septum much nar- rower (Figs. 1, 6). Maxillary teeth about 15-19 vs. about 30 (Fig. 5). Palatohyomandibuloquad- rate cartilage entire vs. palatine separate from hyomandibuloquadrate (Fig. 5). Fifth cerato- branchial with about 8—10 large conical teeth vs. 0-3 small conical teeth (Fig. 4). First gill arch with | + 9 well-developed gill rakers vs. 0 + I- 2 rudimentary gill rakers; all ceratobranchials with well-developed gill rakers vs. gill rakers greatly reduced in size, and ceratobranchials moderately elongate vs. slender and very elon- gate (Fig. 4). Posterior parapelvic cartilage orig- inating dorsal to anterior parapelvic cartilage and extending further into myotomal muscle mass instead of lying parallel with anterior par- apelvic cartilage (Fig. 2). No midventral row of 299 large, round pigment spots on abdomen and postpelvic abdominal keel corresponding in number to myotomal muscles (present in S. #i- crops) (Fig. 1). Anal fin with 12-15 branched rays vs. 14-17. Caudal peduncle more slender (Figs. 1, 2). Vertebral centra more elongate (Fig. 2.) Total vertebrae (excluding basioccipital half centrum but counting upturned hypural centrum as one) 37(n = 2) or 38(7) vs. 41(2), 42(4) or 43(1). Sex: The 120 S. praecox from Khlong Fala- mee were all caught in a segment of the creek less than 100 m long and presumably represent a random sample from a single breeding popu- lation. The sample includes 32 males 14.9-18.3 mm SL with well-developed testes, 19 females 14.9-17.3 mm with well-developed eggs, and 68 specimens 15.2-17.9 mm of undetermined sex in which the gonads are inactive or relatively undeveloped. The testes extend nearly the entire length of the abdomen dorsolaterally to the gut. Obscured anteriorly by the liver, the testes are otherwise readily visible through the transparent ventral body wall and translucent ventral por- tion of the myotomal muscles (Fig. la); they are uniformly divided throughout their length into obliquely aligned divisions or partitions, of which there are about five or six per myotome. Similarly partitioned testes, present in some minute or small freshwater African Clupeidae (personal observation with Peter Whitehead) have not been observed in Salangidae. The larg- est specimen in the entire sample is a male of 18.3 mm with very well developed testes; the smallest male, 14.9 mm, has testes almost as well developed. In females the ovaries also ex- tend virtually the entire length of the abdomen and lie dorsolateral to the gut. Eggs in varying stages of development are present. In the ripest ovaries observed, the largest eggs, 0.20—0.25 mm in diameter, are aligned in a single row of about 25 eggs in each ovary. The smallest fe- male, 14.9 mm, has ovaries with well-developed eggs. Food: Gut contents, either whole macro- scopic animals or fragments of them, readily ob- served through the transparent body and gut walls, are present in 34 (28%) of the 120 speci- mens. In the other 86 (72%), the guts appear to be entirely empty. Items ingested consist exclu- sively of animals, almost all apparently either aquatic insect larvae or segmented vermiform organisms (also aquatic insects?). None of the 300 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 9 FiGuRE 3. Sundasalanx microps, CAS 44220, 17.0 mm, pectoral girdle and fin (ventral view). PT, SCL, CL = post- temporal, supracleithrum, cleithrum; R, F = pectoral radial, frayed margin of fin; SCO = scapulocoracoid (secondary pec- toral girdle and frayed margin of fin of right side omitted). guts contain plant material, sediment, or signif- icant amounts of nonidentifiable debris. ETYMOLOGY.—Latin praecox, too early ripe, premature. | 1mm | FIGURE 4. Sundasalanx microps, new species (Figures 1b, 2b, 3, 4b, 5b, 6b) MATERIAL.—Holotype: Museum of Zoology, Bogor, Indo- nesia, 3000, 17.0 mm, near shore of mainstream Kapuas River at Kampong Nibung, about 100 km NE of Sintang and 7 km NE of Selimbau, Kalimantan, Indonesia, lat. 0°39’N, long. 112°10.5'E; current moderate, water muddy, 26°C, pH 5.5-6, bottom soft mud, depth to 1 m; nylon flyscreen seine; 5-6 July 1976. Paratypes: Museum of Zoology, Bogor, Indonesia, 3001, and CAS 44220, 34:14.6-19.9 mm, same collection data as holotype, seven utilized for alcian blue—alizarin preparations. DIAGNOSIS.—S. microps is distinguished from S. praecox, its only known congener, in the di- agnosis of that species given above. Sex and food: The gonads in the sample of 35 S. microps are not well developed, and I have been unable to distinguish males and females. The guts of nearly all appear to be empty; a few contain unidentified fragments, but no whole in- sect larvae or other animals, plant material, or sediment. ETYMoOLoGy.—Greek mikros, small, little, and ops, eye. Sundasalanx species undetermined Vaillant (1893:110-112, pl. 2, fig. 4) described some delicate little fishes collected in the Ka- Ventral parts of gill arches, dorsal view (gill rakers of right side omitted): (a) Sundasalanx praecox, MCZ 54390, 17.2 mm; (b) Sundasalanx microps, CAS 44220, 17.2 mm. B, H, C = basibranchial, hypobranchial, ceratobranchial (see text for explanation ). ROBERTS: SUNDASALANGIDAE—NEW FISH FAMILY 301 1mm FIGURE 5. SO Jaws, jaw suspension, and gill cover, lateral view: (a) Sundasalanx praecox, MCZ 54390, 17.1 mm; (b) Sun- dasalanx microps, MCZ 44220, 17.0 mm. P, MX, LJ = premaxillary, maxillary, lower jaw; PHQ = palatohyomandibuloquad- rate; PL, HQ = palatine, hyomandibuloquadrate; OP, SO = opercle, subopercle. puas River by Chaper in 1890-91 which he hes- itantly identified as young needlefish (?Belone caudimaculata). Subsequent to completing the manuscript of this paper, I visited the Muséum National d’Histoire Naturelle in Paris, where Chaper’s Kapuas collection is deposited, and examined this material. The specimens (MNHN 91-596, 27:18.8—23.4 mm), although not in the best state of preservation, are clearly Sundasa- lanx but do not agree well with my diagnoses of either S. microps or S. praecox. They agree with S. microps rather than S. praecox in their relatively large size, vertebral counts of about 42-45, modally 43 (determined by counting the myotomes), and number of branched anal fin rays, about 15—18. On the other hand, the eyes seem to be much larger than in §. microps and possibly even larger than in S. praecox, and the number of maxillary teeth fewer than in S. mi- crops and S. praecox. In nearly all of the spec- imens, the eyes are ruptured or their shape so distorted that horizontal diameter cannot be measured accurately. In a 22.5-mm specimen in which the eye is intact and nearly normal in shape, its horizontal diameter is 5.0% of SL (2.7-3.2% in S. microps, 4.3—5.0% in S. prae- cox). The eyeballs are so large that they nearly meet in the middle of the head, whereas in both S. microps and §. praecox they are consider- ably further apart. The number of teeth on the maxillary bone is only 10-14 (n = 3) vs. about 30 in S. microps and about 15-19 in S. praecox. A row of melanophores is visible on the side of the body external to the free ventral ends of the myotomal muscles, but not on the ventral mid- line of the abdomen and postpelvic abdominal keel (the latter present in S. microps but not in S. praecox). I was unable to detect testes or eggs in any of the specimens, although many of 1mm FiGure 6. Anterior end of cranium, dorsal view: (a) Sun- dasalanx praecox, MCZ 54390, 17.2 mm; (b) Sundasalanx microps, CAS 44220, 17.0 mm. EP, NS = ethmoid plate, na- sal septum; ACF, TC = anterior cranial fontanel, tectum cra- nil. 302 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 9 them have ruptured abdomens. The precise lo- cality where they were collected is unknown, but judging from Vaillant’s remarks (1893:60) they must have been taken in or near the main- stream of the Kapuas River somewhere between its confluence with the Sebruang River and Sem- itau, or in the same general area as S. microps, the type-locality of which is less than 40 km upriver from Semitau. So far as I have been able to find out, no additional material of Sundasa- lanx has been reported upon or collected by any- one else. ' _» DISCUSSION Alizarin is a specific stain for bone but does not always stain bone in an early~stage of de- velopment or poorly calcified bone. Alcian blue usually stains cartilage, but apparently also sometimes stains uncalcified bone or bone in an early stage of development. In some instances structures which are clearly bone or cartilage fail to be noticeably stained by either alizarin or alcian blue. Thus, it is not possible to state in every instance which skeletal elements in Sun- dasalangidae and Salangidae are cartilage and which are bone based merely on their staining reaction to alcian blue and alizarin. In Sunda- salangidae the only skeletal elements deeply stained with alizarin are vertebral centra, distal ends of neural and haemal spines, caudal fin rays, hypural fan, teeth, and bony toothplates on fifth ceratobranchials. Maxillary bone, tooth- bearing portion of the lower jaw, secondary pec- toral fin girdle (posttemporal, supracleithrum and cleithrum) and pelvic fin rays are weakly stained with alizarin. Gill rakers, branchiostegal rays, and subopercle are either weakly stained with alcian blue or are not stained at all, in which case they may be difficult to observe even with transmitted light. In alcian blue—alizarin prepa- rations of adult Salangidae, the distribution of elements stained by alcian blue and alizarin is basically similar to that in Sundasalangidae, but alizarin is taken up more extensively. Sundasalanx praecox is among the smallest of adult vertebrates and is the smallest known sal- moniform fish. Sundasalanx microps is the smallest of more than 250 fish species present in the Kapuas River (personal observations), thus providing an excellent example of survival of a peripheral-division fish family in the midst of a rich primary-division freshwater ichthyofauna by evolution of minute body size and a wholly freshwater life history (for further discussion see Roberts 1978:20-21). Salangidae are anadro- mous and freshwater fishes inhabiting coastal waters and rivers of East Asia from North Viet- nam northwards to Korea, Vladivostok, and Sakhalin. They are unknown from Thailand and Borneo. The basic references to systematics and geographical distribution of Salangidae are Fang (1934) and Wakiya and Takahasi (1937). LITERATURE CITED DINGERKUS, G., AND L. D. UHLER. 1977. Enzyme clearing of alcian blue stained whole small vertebrates for demon- stration of cartilage. Stain Tech. 52(4):229-232, 3 figs. FANG, P. W. 1934. Study on the fishes referring to Salangidae of China. Sinensis 4(9):231—268, 7 figs., 6 tables. GEHRINGER, J. W. 1959. Early development and metamor- phosis of the ten-pounder, Elops saurus Linnaeus. U.S. Fish Wildl. Serv. Fish. Bull. 59 (155):619-647, 32 figs., 14 tables. NELSON, G. J. 1970. Gill arches of some teleostean fishes of the families Salangidae and Argentinidae. Jpn. J. Ichthyol. 17(2):61-66, 2 figs. Roserts, T. R. 1978. An ichthyological survey of the Fly River in Papua New Guinea with descriptions of new species. Smithson. Contrib. Zool. 281, 72 pp., 32 figs. VAILLANT, L. 1893. Contribution a |’étude de la faune ichth- yologique de Borneo. Nouv. Arch. Mus. Hist. Nat., ser. 3, 5:23-114, 2 pls. WaKIYA, Y., AND N. TAKAHASI. 1937. Study on fishes of the family Salangidae. J. Coll. Agric. Tokyo Univ. 14(4):265—295, 3 figs., pls. 16-21, 2 tables. CALIFORNIA ACADEMY OF SCIENCES Golden Gate Park San Francisco, California 94118 PROCEEDINGS [Maring Bitor'c OF THE : CALIFORNIA ACADEMY OF SCIENCES Vol. 42, No. 10, pp. 303-314, 4 figs. r Y ACO: MAR 19 1931 . March 5, 1981. rf. #rIidsSo. é fee A AE RE ET i Waingds Ho 3 YV¥UeVIUO 7? te 6 - —— ore aE ES NEW AND RECONSIDERED SPECIES OF MICONIA (MELASTOMATACEAE) FROM COSTA RICA AND PANAMA By Frank Almeda Department of Botany, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118 ABSTRACT: Diagnoses, descriptions, and discussions are presented for five new Central American species of Miconia (M. chiriquiensis and M. coloradensis from Panama, and M. confertiflora, M. grandidentata, and M. longibracteata from Costa Rica). Diagnostic illustrations are provided for four of the novelties, and the new name Miconia concinna is proposed for a Panamanian endemic heretofore known as Topobea micrantha. INTRODUCTION In the four decades since Standley (1938) at- tempted to summarize knowledge of the Costa Rican Melastomataceae, increased botanical ex- ploration in Costa Rica and adjacent Panama has resulted in the collection of much new material of the montane species of Miconia. Field and herbarium study of this genus for a comprehen- sive treatment of Melastomataceae for Flora Costaricensis reveals the need for nomenclatur- al changes, emended species descriptions, and recognition of several undescribed taxa. This paper places some of this new information on record now to make the names available to other researchers prior to completion of the floristic treatment. Miconia chiriquiensis Almeda, sp. nov. (Figure 1) Sect. Cremanium. Frutex vel arbor parva 2- 4(-10) m. Ramuli teretes vel obscure rotundato- quadrangulati sicut folia primum paulo furfu- racei mox glabrati. Petioli (5-)7-18(-25) mm; lamina 3.7-7.5(-10) x 1.5-4.6 cm chartacea el- liptica apice caudato-acuminato basi acuta margine obscure serrulato. Panicula 4-6.5 cm longa glabra laxe multiflora; flores 5-meri, pedi- cellis 0.5-1 mm longis, bracteolis 0.5-1.5 x 0.5 mm caducis. Hypanthium (ad torum) 1 X 1mm, lobis interioribus 0.5-1 mm altis late deltoideis apice obtuso, dentibus exterioribus acutis ca. 0.5 mm eminentibus. Petala 1.5 X 1.5 mm sub- orbicularia glabra. Stamina isomorphica gla- bra; filamenta 1.5 mm longa; antherarum thecae 0.75 x 0.5 mm paulo cuneatae poro 0.5 mm diam. ventraliter inclinato, connectivo non pro- longato. Stylus 0.7-1 x 0.5 mm glaber; stigma subcapitatum; ovarium omnino inferum apice glabro. Shrub or tree 2—4(—10) m tall. Internodes and distal branches + terete, essentially glabrous at maturity, but vegetative buds and young leaves commonly beset with a brownish furfuraceous indument. Leaves chartaceous, distantly ciliate- serrulate, 3.7—7.5(-10) cm long and 1.5—4.6 cm wide, elliptic, bluntly caudate-acuminate apical- ly and acute basally, glabrous at maturity, [303] 304 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, vol. 42, No. ALMEDA: NEW SPECIES OF MICONIA 3-nerved or 3-plinerved (excluding inconspic- uous submarginal pair) punctate and with a con- spicuous network of secondaries below; petioles (5—7)7—18(—25) mm long and | mm broad. Inflo- rescence a laxly branched suberect to + pendant panicle 4-6.5 cm long; rachis + quadrate, gla- brous throughout; bracteoles sessile, early-de- ciduous, 0.5—1.5 mm long and 0.5 mm wide, lin- ear-oblong, entire. Pedicels 0.5—1 mm long. Hypanthia (at anthesis) campanulate, | mm long to the torus, glabrous. Calyx lobes (on fruiting hypanthia) persistent, glabrous, semicircular or depressed-triangular with entire to irregularly, + hyaline margins, 0.5—1 mm long and about | mm wide; calyx teeth persistent, subulate, + appressed to and shorter than mature calyx lobes. Petals 5, erect to antrorsely spreading, glabrous, + concave, white or greenish white, suborbicular, entire, 1.5 mm long and wide. Sta- mens 10, isomorphic, incurved toward central axis of flower at anthesis; filaments glabrous, subulate, white to translucent, mostly 1.5 mm long and 0.5 mm wide basally; anthers about 0.75 mm long and 0.5 mm wide distally, white, + infundibuliform to obliquely cuneate in profile view, shallowly emarginate dorsally, the pore oblong and ventrally inclined; connective sim- ple, inconspicuous and lacking prolongations or appendages. Ovary inferior. Style straight, gla- brous, 0.7—1 mm long; stigma subcapitate. Berry globose, 2—3 mm long to the torus and 2—3.5 mm in diameter. Seeds + pyriform, beige, papillate, mostly 0.7—1 mm long. TypPes.—Panama. Chiriqui: about 8 km w of Cerro Punta, vicinity of Las Nubes, elevation 6100-6400 ft [1859-1951 m], il Feb. 1978, Almeda & Nakai 3535 (holotype: CAS!; iso- types: C!, DUKE!, F!, MO!, PMA!, US!). ADDITIONAL SPECIMENS EXAMINED.—Panama. Chiriqui: Vicinity of Las Nubes, 2.7 miles [4.3 km] Nw of Rio Chiriqui Viejo, w of Cerro Punta, Croat 22392 (CAS, DUKE, MO), Croat 22425 (MO, US); Bajo Chorro, Boquete, Davidson 181 (US), Davidson 390 (MO). DIsTRIBUTION.—A little-collected cloud-for- est species apparently restricted to Chiriqui province in western Panama at elevations of 1850-2200 m. Available specimens, all of which 305 were collected in February, are in flower and fruit. Miconia chiriquiensis is apparently rare and occurs in a region which continues to yield new and narrowly endemic taxa. Diagnostic features include the ciliate-serrulate, elliptic leaves that are caudate-acuminate apically, short (0.5-1 mm), linear-oblong, early-deciduous bracteoles, concave, suborbicular petals, geniculate fila- ments, and minute (0.75 x 0.5 mm), 4-celled an- thers. In his account of Miconia for the Melasto- mataceae of Panama, Gleason (1958) referred this entity to M. rubens (Sw.) Naud. Study of his description and examination of selected cited specimens indicate that Gleason also confused M. chiriquiensis with the taxon treated here as M. concinna. The latter differs markedly from M. chiriquiensis by virtue of its epiphytic habit, trichotomously branched, corymbiform panicle, and distinctive androecial morphology. In foliar size and shape the new species bears a strong resemblance to M. rubens, which is known only from Jamaica and Venezuela. Miconia rubens does differ conspicuously, however, by the somewhat swollen nodes, ferrugineous pubes- cence on distal nodes and juvenile foliage, mar- ginally fimbriate bracteoles, dioecious floral condition, peltate stigma, and cuneate, apically truncate anthers. Although M. chiriquiensis re- sembles M. rubens most closely in the totality of its vegetative characters, it is difficult to pres- ent meaningful speculation regarding the origin and exact relationships of these taxa. Aside from the possibility of evolutionary convergence, the most logical alternative hypothesis is that M. rubens is a Close relative and possibly dioecious derivative of M. chiriquiensis. Miconia coloradensis Almeda, sp. nov. (Figure 2) Sect. Amblyarrhena. Herba | m alta (fide col- lectore). Ramuli glabri primum obscure subquadrangulati demum teretes. Petioli 3- 7.5 x 1.5-3 cm; lamina 13.5-20.5 X 9.8-I17 cm <_— FIGURE 1. Miconia chiriquiensis Almeda. A, habit, x2; B, representative leaves, lower surface (left) and upper surface (right), x1; C, seeds, x12; D, mature berry, <8; E, stamens, ventral view (left) and lateral view (right) x ca. 10; F, petal, x ca. 8; G, fully expanded flower showing natural posture of petals and stamens (left), floral bud and pedicellar bracteoles (right), x9. (A-G from Almeda & Nakai 3535.) 306 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, vol. 42, No. 10 chartacea vel subcoriacea denticulata cordata apice acuminato vel cuspidato, 9-I]-nervata, supra glabra et paulo reticulato-bullata, subtus in superficie pilis stellatis obsita. Panicula 7-19 cm longa multiflora; flores (4—)5-meri, 1.5-5 mm pedicellati, bracteolis 3-5 mm longis obovatis vel spathulatis valde caducis. Hypanthium (ad torum) 2.5 X 2 mm, lobis interioribus | mm altis rotundatis vel truncatis, dentibus exterioribus crassis ca. 0.5 mm eminentibus. Petala 3-5.5 X 2-4 mm obovata. Stamina isomorphica glabra; filamenta 2.5 mm longa; antherarum thecae 2.5 x 1 mm anguste obovatae poro 0.50.75 mm diam. paulo ventraliter inclinato; connec- tivum nec prolongatum nec appendiculatum. Stylus 5 x 0.5 mm glaber; stigma subcapita- tum. Herb to | m tall (according to collectors). Cauline internodes glabrous, subquadrangular to + terete. Leaves + bullate above, chartaceous to subcoriaceous, denticulate, 13.5—20.5 cm long and 9.8-17 cm wide, cordate, acuminate to cus- pidate apically, 9-11-nerved with a prominulous network of secondary and tertiary nerves below, glabrous and green above, red to purple and moderately beset with sessile stellate trichomes on and between primary and secondary nerves below; petioles 3—7.5 cm long and 1.5-3 cm broad, each petiole bearing an abaxial, humplike protuberance proximal to nodal junction. Inflo- rescence a laxly branched terminal panicle 7-19 cm long; rachis + rounded to subquadrangular, moderately stellate pubescent; bracteoles ses- sile, early deciduous, 3-6 mm long and 1|.5-3 mm wide, obovate to spatulate, erose to dentic- ulate, + enveloping young buds and pedicels, glabrous above, sparsely to moderately stellate below. Pedicels 1.5—5 mm long, beset with ses- sile stellate trichomes. Hypanthia (at anthesis) narrowly campanulate, mostly 2.5 mm long to the torus, moderately to sparsely stellate pubes- cent. Calyx lobes (on fruiting hypanthia) persis- tent, glabrous, erect, broadly semicircular but + truncate apically, entire to somewhat erose, | mm long and about 2 mm wide; calyx teeth persistent, bluntly subulate or knoblike, ap- pressed to and shorter than the calyx lobes on mature berries. Petals mostly 5, but 4 in some flowers, reportedly pink, obovate, entire, round- ed to irregularly emarginate at apex, 3—5.5 mm long and 2-4 mm wide. Stamens 10, but 9 in 4- merous flowers, isomorphic; filaments straight, subulate, thickened and somewhat fleshy, at least basally, 2.5 mm long; anthers 2.5 mm long and mostly | mm wide distally, yellow, obovoid, border of apical pore + emarginate ventrally but truncate dorsally; connective inconspicuous and without prolongations or appendages. Ovary in- ferior. Style straight, 5 mm long, glandular-cil- iate basally, otherwise glabrous; stigma subcap- itate. Berry + globose, 4.5-5.5 mm long to the torus and 5—5.5 mm in diameter. Seeds galei- form, nitid, smooth to obscurely papillate, 0.5 mm long. Types.—Panama. Bocas del Toro/Chiriqui border: Cerro Colorado along intersection of Bocas Road with main ridge road, 15.4 km from Chami along ridge road, elevation 1400- 1700 m, 24 Oct. 1977, Folsom 6143 (holotype: CAS!; isotype: MO). ADDITIONAL SPECIMENS EXAMINED.—Panama. Chiriqui: Cerro Colorado, Bocas Road, Folsom & Collins 1748 (CAS, MO). DISTRIBUTION.—Known only from Cerro Colorado in western Panama at an elevation of 1400-1700 m. Flowering and fruiting specimens have been collected in February and October. This distinctive species is characterized by a lax, elongate inflorescence, truncate calyx lobes, nitid, galeiform seeds and cordate, den- ticulate leaves clothed abaxially with sessile, stellate trichomes. In the few collections avail- able for study, the inflorescence is consistently terminal but superficially appears to diverge from a lateral position because of the overtop- ping effect created by lengthening of proximal axillary shoots. The characteristic number of floral parts has been difficult to determine on the basis of material at hand. The inflorescence on the holotype has both 4-merous flowers with nine stamens and 5-merous flowers with ten sta- mens, but the significance of this variation can- not be properly assessed without a more exten- sive series of specimens. The relationships of this species are unclear. Assuming sect. Amblyarrhena represents a nat- ural grouping, it is tempting to suggest that M. coloradensis diverged from the Andean stock that gave rise to M. andreana Cogn. of Colom- bia and M. gibba Markgraf of Ecuador. The for- mer differs from M. coloradensis in having long- er leaves (2-3.5 dm), a furfuraceous pubescence, linear-oblong to subulate bracteoles, and a com- pact inflorescence with ultimate units consisting of congested glomerules. Aside from striking differences in inflorescence size and structure, ALMEDA: NEW SPECIES OF MICONIA 307 FiGureE 2. Miconia coloradensis Almeda. A, habit, x ca. %; B, stamens, ventral view (left) and 3% lateral view (right), x ca. 6; C, mature berry, x3%, D, stellate trichomes, x ca. 35. E, petal, x7; F, seeds, 16. (A-F from the holotype.) the new species approaches M. gibba in overall foliar length and shape. Petioles of the latter are shorter (1.5—2 cm), and the leaves are 7-nerved, caducously furfuraceous below, and bigibbous dorsally at the base. Miconia concinna Almeda, nom. nov. Topobea micrantha Pittier, J. Wash. Acad. Sci. 14:451. 1924. Nec Miconia micrantha Cogn. (Bull. Torrey Bot. Club 23:16. 1896) nec M. micrantha Pilger (Verh. Bot. Ver. Brand. 47:173. 1905; M. wittii Ule, nom. nov., Notizbl. Bot. Gart. Berl. 6:367. 1915) nec M. micrantha Pittier (Bol. Soc. Venez. Cienc. Nat. 11:27. 1947; M. tabayensis Wurdack, nom. nov., Phytologia 21:359. 1971). Study of the holotype and recently collected material of this entity reveals some inaccuracies in Pittier’s incomplete Latin diagnosis. It seems appropriate, therefore, to present the following emended species description. 308 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, vol. 42, No. 10 Epiphytic shrub with lax arching branches to 2 m long. Cauline internodes terete, the distal branches glabrous and + vernicose on drying. Leaves of a pair isomorphic to anisomorphic, chartaceous, ciliate-serrulate (the trichomes mostly 1-2 mm long), 1.7-4.7 cm long and 1.4— 3 cm wide, elliptic to elliptic-ovate acuminate apically and acute basally, sparsely pubescent to glabrate at maturity but usually lepidote to brown-punctate below, sparsely brown furfura- ceous to glabrous above, the pubescence usually tardily deciduous and imparting a brown-punc- tate appearance, 3-nerved with a network of sec- ondaries mostly 1-2 mm apart; petioles 5—20 mm long, | mm broad. Inflorescence a pendant, con- gested, trichotomously branched corymbiform panicle 1-1.5 cm long (shorter than foliage leaves borne at the node initiating the inflores- cence) borne on a stout peduncle 2—5 mm long; bracteoles sessile, foliaceous, persisting on the infructescence, oblanceolate to narrowly spat- ulate, 3—6(—10) mm long, 1-2 mm wide, glabrous to sparsely furfuraceous. Pedicels terete, gla- brous, mostly 0.5 mm long. Hypanthia (at an- thesis) glabrous, campanulate, |—1.5 mm long to the torus and 1—1.5 mm broad. Calyx lobes (on fruiting hypanthia) + ascending, persistent, semicircular but varying to rounded-deltoid, en- tire to minutely lacerate, | mm long and 1.5 mm wide at base between sinuses; calyx teeth per- sistent, triangular, 0.5 mm long. Petals 5, + erect to antrorsely spreading, glabrous, + con- cave, white but sometimes yellowish on drying, suborbicular, entire, 2—-2.5 mm long and wide. Stamens 10, isomorphic; filaments glabrous, subulate, distally geniculate, 2—2.5 mm long; an- thers 0.5 mm long, + cuneate, broadly flared and terminated by a + ovoid ventrally inclined pore; connective thickened and prolonged (0.5 x 0.5 mm) below thecae, truncate to shal- lowly bilobed in ventral view, dilated dorsally into a blunt projecting appendage in profile view. Ovary ca. 4/5 inferior. Style straight, glabrous, 3 mm long; stigma + clavate to subcapitate. Ber- ry reportedly black at maturity, 2-2.5 mm long to the torus, 2.5-3 mm in diameter. Seeds + pyriform, smooth and nitid, mostly 0.75 mm long. SPECIMENS EXAMINED.—Panama. Chiriqui: humid forests on precipitous slopes of Cerro de la Horqueta, Pittier 3276 (US, holotype of T. micrantha); s slopes of Cerro Horqueta N of Boquete, Wilbur, Teeri & Foster 13490 (CAS, DUKE); Cerro Pando, on continental divide and Panama/Costa Rica border, ca. 16 km Nw of El Hato del Volcan, Mori & Bolten 7291 (CAS), Mori & Bolten 7301 (CAS). DISTRIBUTION.—Apparently a localized cloud- forest epiphyte endemic to Chiriqui province in western Panama at elevations of 2000-2500 m. Flowering and fruiting specimens have been col- lected in January, March, and July. Miconia concinna is recognized by its lax, arching branches, adaxial furfuraceous indu- ment on juvenile foliage, large, persistent, folia- ceous floral bracts, and pendant, few-flowered corymbiform inflorescences that are markedly shorter than subtending foliage leaves. Pittier’s (1924) initial placement of this species in Topobea is difficult to understand, since his discussion makes note of several features which made this decision questionable. In choosing this course, Pittier was apparently impressed by the presence of conspicuous foliaceous floral bracteoles and by the position of the inflores- cence, which he erroneously described as axil- lary. In describing this species as a tree, it also seems likely that Pittier mistook its habit for that of its host. Label Information for all recently gathered material indicates that this species is an epiphyte, and until noted otherwise, it seems advisable to accept this habital description as characteristic of the species. On the basis of limited material, Standley (1938) misinterpreted M. concinna to be con- specific with the Costa Rican endemic described here as M. longibracteata. This confusion was compounded when he referred specimens of these two taxa to M. myrtillifolia Naud., a species of Andean Colombia and Venezuela which differs in having quadrate branchlets, a longer (2-4 cm), erect panicle, and very different oblong anthers with diminutive apical pores and unprolonged connectives. More recently, Glea- son (1958) included M. concinna in his miscon- strued concept of M. rubens (Sw.) Naud. The latter, a dioecious species, is known only from Jamaica and Venezuela and differs most notably by its elongate multiflowered inflorescence, in- conspicuous bracteoles, + swollen nodes, and different anther morphology. In foliar shape and floral details, M. concinna and M. longibracteata are more similar to each other than to any other species of the genus, and there is little doubt that they were derived from common ancestral stock. Available collections suggest that these two species are allopatric, with the range of M. concinna lying south of ALMEDA: NEW SPECIES OF MICONIA that of M. longibracteata. The prevailingly gla- brous leaves, early-deciduous floral bracteoles, and laxly branched, elongate panicle of M. lon- gibracteata serve to separate these species most readily. Miconia confertiflora Almeda, sp. nov. (Figure 3) Sect. Chaenopleura. Frutex epiphyticus ca. 2 m altus. Ramuli sulcato-quadrangulati sicut folia inflorescentia plerumque glabri. Petioli 3- 16(-20) mm; lamina 1.7-7 Xx 1-3.4 cm elliptica, elliptico-ovata vel obovata apice acuta vel acu- minata basi acuta, 3(-—5)-nervata, chartacea et serrulata. Panicula corymbiformis, pedunculo plus minusve 1.5 cm longo; flores 5-meri breviter (1 mm) pedicellati, bracteolis 1.5-2.5(-4) mm longis valde caducis. Hypanthium (ad torum) 1.5 x 1 mm, lobis interioribus 0.5-1 mm altis late deltoideis vel rotundatis, dentibus exteriori- bus acuminatis 0.5-1.5 mm longis. Petala 1- 1.5 x 0.5-l1 mm ovata apice acuto vel paulo un- cinato. Stamina isomorphica glabra; filamenta 1-1.5 mm longa; antherarum thecae 0.75-I xX 0.25-0.50 mm anguste oblongae, connectivum nec prolongatum nec appendiculatum. Stylus 2 X 0.5 mm glaber; stigma truncatum non ex- pansum. Epiphytic shrub to 2 m tall, distal branches quadrangular with carinate to narrowly alate an- gles, entirely glabrous but bearing a pair of + pustulate setiform appendages at opposing nodal faces. Leaves chartaceous, glabrous, |.7—7 cm long and 1-3.4 cm wide, basally entire but dis- tally serrulate, elliptic but sometimes varying to elliptic-ovate or obovate, acute to acuminate apically and acute basally, 3(—5)-nerved with a conspicuous network of secondary nerves, dark green above, pale green and occasionally punc- tate below; petioles 3—16(—20) mm long and about | mm broad. Inflorescence a multiflow- ered corymbiform panicle with flowers borne in congested terminal glomerules; rachis glabrous, quadrangular, mostly less than 1.5 cm long; bracteoles sessile, glabrous, early-deciduous, 1.5—2.5(—4) mm long and 0.5—1 mm wide, linear- subulate. Pedicels 1 mm long. Hypanthia (at an- thesis) campanulate, glabrous, 1.5 mm long to the torus. Calyx lobes (on fruiting hypanthia) persistent, erect, broadly deltoid to + rounded, entire with conspicuous hyaline margins, 0.5-1 mm long and | mm wide; calyx teeth persistent, 309 subulate to setiform, equaling or commonly ex- ceeding calyx lobes on mature berries. Petals 5, glabrous, erect and + concave at anthesis, white but tinged with red externally, narrowly to broadly ovate, entire, acute to bluntly uncinate apically, 1-1.5 mm long and 0.5-1 mm wide. Stamens 10, isomorphic, erect to slightly in- curved at anthesis; filaments white, + translu- cent, subulate, 1-1.5 mm long; anthers 0.75-1 mm long and 0.25—0.50 mm wide basally, gla- brous, white, narrowly oblong but + obovoid in profile view, distally rounded with a subtermi- nal, oblong ventrally inclined cleftlike pore, the margins of which often form a + elevated hya- line border; connective thickened, not conspic- uously dilated or prolonged dorsally but com- monly + prolonged ventrally below thecae. Ovary wholly inferior. Style straight, 2 mm long; stigma truncate. Berry purple at maturity, + glo- bose, 3—3.5 mm long to the torus and 3.5 mm in diameter. Seeds narrowly ovoid with an en- larged, + flattened lateral raphe, densely papil- late on the convex surface, mostly 2 mm long and | mm broad. Types.—Costa Rica. San José: about 18 km N of San Isidro de Coronado off C.R. #216 on lower w slopes of Volcan Ir- azu, elevation 1700-1800 m, 5 July 1977, Almeda et al. 2908 (holotype: CAS!; isotypes: CR!, F!, MO!, US!). ADDITIONAL SPECIMENS EXAMINED.—Costa Rica. Here- dia: slopes NE of Cerro Chompipe about 16 km NNE of San Rafael, Wilbur, Almeda & Daniel 22249 (CAS, DUKE); Cerro Zurqui, NE of San Isidro, Standley & Valerio 50542 (US), Standley & Valerio 50644 (US); saddle area between Cerro Chompipe and se flank of Volcan Barba off secondary road N of C.R. Hwy #113 connecting with Calle Gallito, Baker, Utley & Utley 232 (CAS, DUKE). San Jose: 5 km NE of Cas- cajal, Almeda & Nakai 3611 (CAS); 3-6 km beyond Las Nubes in vicinity of Cascajal, Almeda 2636 (CAS); about 7 km by road NE of Cascajal and 14 km NE of San Isidro de Coronado, Wilbur 19816 (DUKE); 3-5 km ne of Cascajal in vicinity of Rio Cascajal, Wilbur 24420 (DUKE); about 3 km NE of Cascajal and 9 km NE of San Isidro de Coronado, Wilbur 19787 (DUKE). DISTRIBUTION.—A local cloud-forest epi- phyte apparently endemic to the Cordillera Cen- tral of Costa Rica at elevations of 1600-2400 m. Flowering and/or fruiting specimens have been collected in February, March, July, October, and December. Field observations and label information of known collections indicate that this species is an obligate epiphyte. Extirpation of moist forests within the limited range of this taxon poses a real threat to its survival. Fortunately, some in- dividuals persist as inhabitants of remnant pas- 310 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, vol. 42, No. 10 W j Wy 4 \) tau Vd STR | YY ig /2 = a » . es i SI Pr, BD AcGa Ns } Tewwe, nl \\ We ESS ec) BR UD. co FiGurE 3. Miconia confertiflora Almeda. A, habit, x4; B, representative leaves, upper surface (left) and lower surface (right), x34; C, fully expanded flower showing petals and stamens, <9; D, petal, x 16; E, stamens, ventral view (left) and lateral view (right), «13; F, mature berries with persistent calyx lobes and calyx teeth, x3; G, seeds, x6. (A-—G from Almeda et al. 2908.) ture trees long after surrounding vegetation has inconspicuous anthers are also noteworthy in been cut away. having subterminal, oblong pores, the margins This new species is readily separated from of which are elevated into a low, continuous, congeners by its quadrangular branchlets, dis- hyaline border. This feature becomes distorted tally serrulate leaves, congested corymbiform with pressing and drying and is best observed in inflorescence, glabrous hypanthia, prominent — pickled or hydrated material. calyx teeth, and narrowly ovoid seeds that are Miconia confertiflora superficially resembles densely papillate on the convex side. The small, MM. chionophylla Naud. of sect. Chaenopleura, ALMEDA: NEW SPECIES OF MICONIA which ranges from Andean Colombia to Bolivia. The latter differs in its procumbent or scandent habit, puberulent branchlets and petioles, small- er leaves (1-1.5 x 0.7-1.3 cm), 4-merous flow- ers, and capitate stigma. The congested inflo- rescence of M. confertiflora is also reminiscent of that found in M. parvifolia Cogn. (of sect. Cremanium), another high-elevation Colombian species easily separated by its dwarf shrubby habit (2-3 dm tall), smaller, revolute leaves (8- 15 mm long), ovate, apically truncate petals, and copious stellate pubescence on distal branchlets. Miconia grandidentata Almeda, sp. nov. Sect. Chaenopleura. Frutex epiphyticus ca. 1 m altus. Ramuli sulcato-quadrangulati sicut folia novella primum modice vel dense pilis stel- latis induti mox glabrati. Petioli 4-15(-19) mm; lamina 3-6 xX 1.8-3.5 cm elliptica vel elliptico- ovata apice acuto vel acuminato basi acuta, 3(-5)-nervata, chartacea obscure distanterque serrulata. Panicula 3-5.3 cm longa multiflora; flores 5-meri breviter (1-2 mm) pedicellati, brac- teolis conspicuis (1-)3-6 mm longis persistenti- bus. Hypanthium (ad torum) 2-2.5 x 2 mm, lob- is interioribus 0.5 mm altis late deltoideis vel rotundatis, dentibus exterioribus subulatis 2 mm longis. Petala 1.5—2 X 1 mm plus minusve ovata apice uncinato. Stamina isomorphica glabra; filamenta 2 mm longa; antherarum thecae 0.75— 1 x 0.5 mm oblongae, rectae vel paulo curvatae poro ventraliter inclinato, connectivo ad basim dorsaliter dente hebeti truncato glabro ornato. Stylus 2 x 0.5 mm glaber; stigma truncatum non expansum. Epiphytic shrub to | m tall, distal branchlets moderately to densely stellate, quadrangular with carinate to narrowly alate angles. Older branches somewhat corky on drying, cracking and excorticating in age. Leaves chartaceous, entire but obscurely serrulate distally, 3-6 cm long and 1.8-3.5 cm wide, elliptic to elliptic- ovate, acute to acuminate apically and acute ba- sally, glabrous at maturity but clothed with brown stellate trichomes when young, 3(-—5)- nerved with a conspicuous network of second- ary nerves below; petioles 4—15(-19) mm long and 1-2 mm broad. Inflorescence an erect, ter- minal thyrse mostly 3—5.3 cm from base to apex; the rachis prominently quadrangular; bracteoles sessile, essentially glabrous, persistent, gradu- ally reduced in size upward, (1-)3-6 mm long 311 and 0.5—2 mm wide, narrowly lanceolate to sub- ulate and + concave adaxially. Pedicels !—1.5 (-2) mm long caducously stellate pubescent, each pedicel commonly subtended by three bracteoles. Hypanthia (at anthesis) + globose, 2-2.5 mm long to the torus, beset with sessile stellate trichomes (these early-deciduous and generally not present on fruiting hypanthia) or persistent only as remnant arms of stellate tri- chomes, which superficially resemble puncti- form glands. Calyx lobes (on fruiting hypanthia) persistent, sparsely stellate pubescent, erect to somewhat incurved, broadly deltoid to + round- ed, entire to bluntly undulate with hyaline mar- gins, 0.5 mm long and | mm wide; calyx teeth persistent, subulate, 2 mm long and 1.5 mm wide at base, markedly exceeding calyx lobes on ma- ture berries. Petals 5, glabrous, erect and + con- cave at anthesis, white, + ovate in outline, en- tire, but bluntly uncinate apically, 1.5-2 mm long and mostly | mm wide. Stamens 10, iso- morphic, erect but + incurved and exceeding the style; filaments white to translucent, subu- late, 2 mm long; anthers 0.75—1 mm long and mostly 0.5 mm wide basally, glabrous, white, oblong to rhomboid in profile view, but broad- ened distally to an oval or oblong, ventrally in- clined apical pore; connective thickened and prolonged dorsally at the base into a deflexed + truncate appendage. Ovary wholly inferior. Style straight, 2 mm long; stigma truncate. Berry pink when young but deep purple at maturity, globose, mostly 4 mm long to the torus and 4 mm in diameter. Seeds cuneate and conspicu- ously angled, reddish brown, vernicose, 1.5— 1.75 mm long. Type.—Costa Rica. San José: About 5 km NE of Cascajal, elevation 5400 ft [1646 m], 17 Feb. 1978, Almeda & Nakai 3627 (holotype: CAS!). ° ADDITIONAL SPECIMENS EXAMINED.—Costa Rica. Heredia: pastured slopes above Rio Para Blanco on lower slopes of Cerro Zurqui about 7 km ne of San Josecito, Wilbur & Luteyn 18634 (DUKE). San José: slopes and thickets at Alto La Pal- ma about 15 km ina straight line NE of San José, Wilbur 20340 (CAS, DUKE). DISTRIBUTION.—A rare cloud-forest epiphyte known only from the south-facing slopes of the Cordillera Central of Costa Rica at elevations of 1600-1800 m. Flowering material has been col- lected in February, July, and December. Initial study of this species led me to interpret it as an atypical large-leafed variant of M. con- fertiflora, a species that grows in the same gen- 312 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, vol. 42, No. 10 eral area. Subsequent field observations of ad- ditional flowering and fruiting material indicate that this is a distinctive taxon worthy of specific rank. Miconia confertiflora and M. grandiden- tata share an epiphytic habit, quadrangular branchlets, and elliptic to elliptic-ovate, distally serrulate leaves having a conspicuous network of secondary nerves below. Miconia grandiden- tata differs consistently in several diagnostic characters. It lacks the nodal, setiform append- ages so characteristic of distal branchlets in M. confertiflora. It also differs in having stellate pubescence on branchlets and young hypanthia, an elongate thyrsoid panicle, persistent floral bracteoles, longer calyx teeth, and angulate, vernicose seeds that lack a well-defined pro- longed lateral raphe. The stamens of M. gran- didentata also provide several distinguishing features. Hydrated anthers, which are oblong to rhomboid in profile view, are broadened distally to an oval or oblong, ventrally inclined terminal pore, and the connective is thickened and pro- longed dorsally into a deflexed caudiform ap- pendage. Miconia longibracteata Almeda, sp. nov. (Figure 4) Sect. Chaenopleura. Frutex 1-3 m altus. Ra- muli glabri obscure quadrangulati demum te- retes. Petioli 7-16(-27) x 1 mm; lamina (1.3-) 2.5-4.2 x 1.1-2.8 cm chartacea elliptica, ellip- tico-obovata aliquando suborbicularia apice acuta vel acuminata basi acuta vel obtusa, tri- nervata supra primum sparse ferrugineo-furfu- racea mox glabrata, subtus primum sparse vel modice lepidota mox glabrata. Panicula 3.5-8 cm longa multiflora; flores 5-meri, pedicelli (1-) 2.4 mm longi, bracteolis 4-11(-18) mm longis oblongis vel anguste spatulatis usque ad an- thesim persistentibus. Hypanthium (ad torum) 1-1.5 x 1-1.5 mm, lobis interioribus 1-].5 mm altis rotundatis vel deltoideis, dentibus exteriori- bus acutis 0.5 mm longis. Petala 2-2.5 x 2mm suborbicularia glabra. Stamina isomorphica glabra; filamenta 2.5 mm longa; antherarum thecae 0.5 X 0.25 mm apice late biporosae, connective sub loculis 0.5 mm prolongato dor- saliter ad basim (0.5 mm) hebeti-tuberculato. Stylus 2-2.5 * 0.5 mm _ glaber; stigma plus minusve clavatum. Shrub 1-3 m tall. Cauline internodes glabrous, glossy black and obscurely quadrangular when ~by a broad, + ‘connective markedly thickened and prolonged young, becoming brown and terete with age. Leaves firmly chartaceous, ciliate-serrulate (the trichomes mostly 0.5—1 mm long), (1.3—)2.5—4.2 cm long and 1.1—2.8 cm wide, elliptic, elliptic- obovate or sometimes varying to suborbicular, acuminate to acute apically and acute to obtuse basally, glabrous at maturity but caducously lep- idote below and glabrous to sparsely brown fur- furaceous above when young, 3-nerved, the sec- ondaries conspicuous below and mostly 2 mm apart; petioles 7—16(-—27) mm long and | mm broad. Inflorescence an erect, laxly branched, elongate panicle 3.5-8 cm long, exceeding fo- liage leaves borne at the node initiating the in- florescence; rachis glabrous, quadrate to + rounded; bracteoles sessile, foliaceous, decidu- ous following anthesis and mostly absent on the infructescence, linear-oblong to narrowly spat- ulate, + concave to navicular, 4-11(-18) mm long, 0.5—3 mm wide, glabrous above and below. Pedicels terete, glabrous, (l1—)2—-4 mm long. Hy- panthia (at anthesis) glabrous, campanulate, |— 1.5 mm long to the torus and 1-1.5 mm broad distally. Calyx lobes (on fruiting hypanthia) erect to ascending, persistent, semicircular but varying to + deltoid, entire, |—-1.5 mm long and 1-1.5 mm wide basally between sinuses; calyx teeth persistent, triangular, 0.5 mm long. Petals 5, antrorsely spreading, glabrous, + concave, white to yellowish white, suborbicular, entire, 2—2.5 mm long, 2 mm wide. Stamens 10, iso- morphic; filaments glabrous, subulate, white, distally incurved, 2.5 mm long; anthers about 0.5 mm long or less, 0.25 mm wide distally, white, + cuneate, flared distally and terminated ovoid, ventrally inclined pore: (0.5 x 0.5 mm) below thecae, dilated dorsally into a blunt knobby protuberance. Ovary totally inferior. Style straight, glabrous, 2—2.5 mm long; stigma + clavate. Berry deep purple to purple- black at maturity, 3.5-5S mm long to the torus, 3—4(-5) mm in diameter. Seeds + pyriform, ob- scurely muriculate, mostly | mm long. TypPes.—Costa Rica. Alajuela: wooded slopes of Volcan Poas about 12 km w of Varablanca, elevation ca. 2400 m, 21 Jan. 1968, Wilbur & Stone 9845 (holotype: DUKE!; isotypes: CAS!, US!). ADDITIONAL SPECIMENS EXAMINED.—Costa Rica. Alajue- la: Volcan Poas, forest between crater and cold lake, Davidse & Pohl 1169 (US); Volcan Poas, cloud forest along road 2.5- 3.5 miles [4.0-5.6 km] w of Poasito, Webster, Miller & Miller 12242 (DUKE, US); se slope of Volcan Poas, Hatheway 1388 ALMEDA: NEW SPECIES OF MICONIA 313 = SSF i , ees £ Vv 4 } i FiGuRE 4. Miconia longibracteata Almeda. A, habit, x ca. %; B, seeds, x14; C, petal, x9; D, representative leaves, upper surface (left) and lower surface (right), x1; E, mature berry, x ca. 4; F, stamens, lateral view (left) and 34 ventral view (right), x10: G, stamens, dorsal view (left) and ventral view of anther and prolonged connective (right), x 10. (A & D from Schnell 727: B, C, E-G from Wilbur & Stone 9845.) 314 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, vol. 42, No. 10 (DS, US). Cartago: El Empalme, Schnell 727 (US—2 sheets); Talamanca Range, Panamerican Highway, Carlson 3611 (US); El Canon (Carretera Panamericana), O. Jimenez 42 (US); La Chonta, Schnell 725 (US); Panamerican Highway about 24 km w of Villa Mills, Wilbur & Stone 8800 (DUKE); s of El Em- palme, A. Jimenez 1979 (US). Cartago/San José border: 6 km Nw of Dos Amigos, Wilbur & Luteyn 18321 (DUKE); 5 km Nw of Ojo de Agua or 19 km sE of El Empalme, Wilbur 28674 (DUKE); 5 km se of Trinidad along Carretera Interamericana, Wilbur 27780 (DUKE). DISTRIBUTION.—Endemic to Costa Rica where it is presently known from the slopes of Volcan Poas and the Cordillera de Talamanca at eleva- tions of 2000-3000 m. Available collections in- dicate that flowering and fruiting occurs sporad- ically throughout the year. Among Costa Rican taxa of Miconia, this species is recognized by the combination of large foliaceous floral bracteoles, prevailingly glabrous foliage, suborbicular petals, and pecu- liar androecial morphology (see Fig. 4F). Sta- minal posture and morphology are best observed in pickled or boiled material hydrated with the aid of a wetting agent. Materials so treated show short, compressed, anther thecae terminating in a broad apical pore; the connective which is conspicuously thickened and prolonged below the thecae is narrowly triangular in ventral view, knobby and somewhat angulate in profile view, and + horseshoe shaped in dorsal view. In many respects the elaborate connective is reminiscent of a pedestal providing anchorage and support for the anther. This distinctive anther morphol- ogy appears to represent an extreme in the evo- lutionary line that has led to great reduction in anther size. The very broad apical pores and pronounced geniculation of the filaments consis- tently bring anthers to an incurved position within the flower and may contribute to preva- lent self-pollination. In size and shape of floral bracteoles and an- thers, M. longibracteata is most similar to M. concinna (also treated herein). The characters distinguishing these species are enumerated in the discussion under M. concinna. Miconia lon- gibracteata also resembles M. superposita Wur- dack in vegetative aspect. The latter, a Colom- bian species, differs in having squamulose pubescence on juvenile branchlets, shorter brac- teoles (3.7 x 0.8 mm), and 4-celled anthers with the dorsal loculus of each theca overlapping but distal to the ventral one. ACKNOWLEDGMENTS I thank Terry Bell for preparing the line draw- ings, Dr. J. J. Wurdack for review of the manu- script, the Museo Nacional de Costa Rica and the Organization for Tropical Studies for logis- tical support, and curators of the following her- baria for special loans or use of their facilities: CR, DUKE, F, MO, US. This study was sup- ported by U.S. National Science Foundation Grants DEB 76-83040 and DEB 78-25620. LITERATURE CITED GLEASON, H. A. 1958. Melastomataceae. Flora of Panama. Ann. Missouri Bot. Gard. 45:203-304. Pitter, H. 1924. New or little known Melastomataceae from Venezuela and Panama. II. J. Wash. Acad. Sci. 14:447-451. STANDLEY, P. C. 1938. Flora of Costa Rica. Field Mus. Nat. Hist., Bot. Ser. 18(3):783-1133. CALIFORNIA ACADEMY OF SCIENCES Golden Gate Park San Francisco, California 94118 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENGESS = *’ Vol. 42, No. 11, pp. 315-322, 5 figs. Marine Bislogical Laborator; JUL 2 0 1981 June 24, 1981 Waeds Hole, dass. ; STUDIES ON CAVE HARVESTMEN OF THE CENTRAL SIERRA NEVADA WITH DESCRIPTIONS OF NEW SPECIES OF BANKSULA By Thomas S. Briggs Research Associate, Department of Entomology, California Academy of Sciences, San Francisco, California 94118 and Darrell Ubick Biology Department, San Jose State University, San Jose, California 95114 ABSTRACT: New ecological and biogeographic information on Sierra Nevada caye harvestmen in Banksula was obtained while environmental impact and mitigation work was being done for the Army Corps of Engineers New Melones Dam project. Isolation appears to be the principal factor leading to speciation in Banksula, but the distribution of species in the vicinity of the New Melones Reservoir is not readily explained. Four new species of Banksula are described: B. rudolphi, B. martinorum, B. grubbsi, and B. elliotti. INTRODUCTION Troglobitic organisms are scarce in California caves, possibly due to the relatively small size and geologic youth of these habitats. Laniatorid harvestmen of the genus Banksula are distinc- tive because they are relatively abundant obli- gate cavernicoles of the Calaveras Formation of the Sierra Nevada. Intensive collecting by bio- speleological investigators contracted by the Army Corps of Engineers, the Fish and Wildlife Service, and the Office of Endangered Species has yielded numerous new records and four new species of Banksula. Project teams worked pri- marily in the vicinity of the New Melones Res- ervoir site on the Stanislaus River, Calaveras and Tuolumne counties, where several caves are threatened by completion of the New Melones Dam. Biological surveys were begun in May 1975 when a mine tunnel in limestone was se- lected for transplanting biota, including Bank- sula grahami and B. melones, from McLean's Cave, the largest of the threatened caves. As additional workers transplanted animal and plant life from McLean’s Cave and studied other nearby caves, some distributional, behavioral, and ecological information were obtained which allow us to present some biogeographic discus- sion of Banksula. Briggs (1974) reviewed and expanded the ge- nus Banksula and described four species with functional eyes. He showed that the single pre- viously known species, Banksula californica (Banks), lacked corneas and retinae. His con- tention that all Banksula are confined to caves is supported by all subsequent investigations. Only Banksula melones, which has relatively (315] 316 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 11 well-developed eyes, has been collected near or at the cave twilight zone. Sitalcina sierra Briggs and Hom is the only laniatorid harvestman found in epigean habitats adjacent to cave lo- calities. METHODS AND DEPOSITION The structures measured and morphological terminology used in this paper are generally those described by Briggs (1968) for laniatorid harvestmen, with one modification that appears in this paper and in Briggs (1974): we substitute ‘*second endites’’ for that portion of the second coxae previously referred to as the maxillary processes. Another structure, the labial process, first described by Briggs (1971), is located be- tween the second endites. Unless otherwise noted, all specimens record- ed in this paper were preserved in ethyl alcohol and are deposited in the collection of the Cali- fornia Academy of Sciences. Key to the Species of Banksula la. Operculum small, held almost entirely be- tween mesal margins of fourth coxae; males with apex of aedeagus not enclosed in sheath, velum on dorsal plate smooth (californica 2toup)) 2282s 2 Ib. Operculum large, with posterior margin well behind mesal extreme of posterior margin of fourth coxae; males with apex of aedeagus enclosed in sheath, velum on dorsal plate in transverse folds (melones 2a. Retinae entire; eye tubercle obliquely truncate in lateral view __ B. galilei Briggs 2b. Retinae absent or incomplete; if retinae present, eye tubercle a rounded cone _ 3 3a. Small spines or tubercles between prin- cipal dorsal spines on palpal femur; cor- neas absent B. californica (Banks) 3b. No small spines or tubercles between principal dorsal spines on palpal femur; corneas present or absent 4a. Proximal ventral spinose tubercle on pal- pal tibia equal to ventral spinose tubercle on patella; retinae absent; corneas pres- Chit OA SENG eee ments 0 DA See te Ae 5 4b. Proximal ventral spinose tubercle on pal- pal tibia always smaller than ventral spi- nose tubercle on patella; retinae incom- plete or absent; corneas present 5a. Submarginal row of tubercles on eighth tergite and last sternite’ >see etree Fe eee B. rudolphi new species 5b. No submarginal rows of tubercles on ter- gites or sternites; __..2 = ae a) eee B. martinorum new species 6a. Eye tubercle a rounded cone; small spur ectal to proximal ventral spine on palpal femur; incomplete retinae usually pres- B. tuolumne Briggs 6b. Eye tubercle obliquely truncate in lateral view; no spur ectal to proximal ventral spine on palpal femur; retinae absent ____ £2. 3 ee B. grubbsi new species 7a. First legs longer than 4 mm; retinal di- ameter greater than 0.03 mm; aedeagus and enclosing sheath fold anterior to re- curved prongs of dorsal plate __- poss, Ce B. melones Briggs 7b. First legs shorter than 4 mm; retinal di- ameter less than 0.03 mm or missing; ae- deagus and enclosing sheath held within bifurcate dorsal plate 8a. Eye tubercle and elevated area behind without tubercles; retinae present ______ est 3d Snes can) B. grahami Briggs 8b. Eye tubercle and elevated area behind tu- berculate; retinae reduced or missing -_-- oct) ORES bt a B. elliotti new species Banksula rudolphi new species (Figure 2) HoLotypPe.—Male. Body length 1.67 mm; scute length 1.40 mm; scute width 1.14 mm; eye tubercle length 0.23 mm; eye tubercle width 0.28 mm; operculum length 0.18 mm; operculum width 0.21 mm. Palp: trochanter 0.19 mm, femur 0.79 mm, patella 0.44 mm, tibia 0.65 mm, tarsus 0.44 mm. Leg II: trochanter 0.19 mm, femur 1.42 mm, patella 0.44 mm, tibia 1.26 mm, metatarsus 1.02 mm, tarsus 1.58 mm. Scute with segmentation delineated by tuber- cles, eye tubercle and area behind tuberculate. Eye tubercle a rounded cone without retinae (some individuals also without corneas). Ter- gites with row of tubercles at margin, eighth ter- gite also with medial row of tubercles. Posterior BRIGGS AND UBICK: CAVE HARVESTMEN PLACER CO. EL DORADO CO. californica galilei grahami r melones tuolumne elliotti e grubbsi martini rudolphi OCODOC¢@E RO FIGuRE 1. TUOLUMNE CO. 317 i ‘ @ Bs my v @ ales Transplant Mine Ws +\f 6 --~-7}, 7 , Vf tee eN) lr air - ‘ji é « 4 ) (SIE a iF j ae . H >> 2 Columbia § ee (is ny Cz Map showing counties in the central Sierra Nevada where Banksula have been collected. The inset shows the region along the Stanislaus River where environmental impact and mitigation work on Banksula has been concentrated. sternite with two rows of tubercles. Anal plate with medial tubercles. Second endites with con- cave ectal margin. Labial processes rounded. Operculum with narrow, rounded anterior; pos- terior margin adjacent to mesal extreme of pos- terior margin of fourth coxae. Palpal femur with six prominent proximal dor- sal spines (some individuals with only five spines) and one small distal dorsal spine; no se- tae or tubercles between dorsal spines; ectal spur at base of first ventral spine small and rounded; venter with numerous small tubercles. Proximal ventral tubercle on palpal tibia re- duced, spine from tubercle absent. Ventral spine on palpal patella arising from a reduced tubercle. Tarsal formula 4-6-5-6. Body concolorous yellow-orange. Penis typical of californica group (see Briggs 1974:11—12). ALLOTYPE.—Female. Slightly smaller than holotype. Proximal ventral spinose tubercle on palpal tibia present, equal in size to robust ven- tral spinose tubercle on palpal patella. Opercu- lum with truncate apex (rounded in some indi- viduals); posterior margin behind mesal extreme of posterior margin of fourth coxae. TYPE-SPECIMENS.—Holotype, allotype, 21 paratypes (9 d, 9 2, 3 juv.): Chrome Cave, near Jackson, Amador County, California, 5 Apr. 1979, D. C. Rudolph, S. Winterath, and B. Martin. Paratypes, 2 d, 2 2, 3 juv.: same locality, 21 Sep. 1980, T. S. Briggs and D. Ubick. Paratype, | ¢: same locality, 24 Jan. 1981, T. S. Briggs and D. Ubick. ETyMOLOGY.—This species is named for biospeleologist D. Craig Rudolph. EcoLoGIcAL NotEes.—Chrome Cave is a small cave situated in about 0.02 km? of limestone. The outcrop is surrounded by serpentine which also forms much of the cave’s walls. The hu- midity is relatively high and a mean temperature of 18 C was recorded on 24 January 1981. Bank- 318 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 11 FIGURE 2. Lateral view of left palpus of female allotype. sula rudolphi is found throughout the upper re- gions of the cave, predominantly on the under- sides of rocks. Associated with Banksula is a rich assortment of predators, of which spiders (Araneae) are most abundant. We recorded the following species: Archoleptoneta schusteri Gertsch, Liocranoides sp., Trogloneta para- doxa Gertsch, and Usofila (Telema) sp. Of the other cavernicoles encountered, the most inter- esting were specimens of Prokoenenia sp. (Pal- pigradida). Banksula martinorum new species (Figure 3) Ho.Lotype.—Male. Body length 1.75 mm; scute length 1.49 mm; scute width 1.23 mm; eye tubercle length 0.26 mm; eye tubercle width 0.33 mm; operculum length 0.18 mm; operculum width 0.19 mm. Palp: trochanter 0.19 mm, femur 0.88 mm, patella 0.53 mm, tibia 0.65 mm, tarsus 0.42 mm. Leg II: trochanter 0.21 mm, femur 1.81 mm, patella 0.39 mm, tibia 1.67 mm, metatarsus 1.23 mm, tarsus 2.37 mm. Scute with segmentation delineated by small tubercles; eye tubercle and area behind slightly tuberculate. Eye tubercle subconical, without Banksula rudolphi Briggs and Ubick, new species. a. Lateral view of body and left palpus of male holotype; b. retinae or corneas. Tergites with submarginal row of tubercles. Second endites large, ectal margin slightly concave. Labial processes with acute anterior margin. Operculum with rounded anterior, posterior margin slightly behind mesal extreme of posterior margin of fourth coxae. First coxae with two prominent spinose tuber- cles. Anal plate smooth. Palpal femur with five prominent proximal dorsal spines and two slightly smaller distal dor- sal spines; no setae or tubercles between dorsal spines; ectal spur at base adjacent to second ventral spine. Proximal ventral spinose tubercle on palpal tibia equal to ventral spinose tubercle on patella. Tarsal formula 4-6-5-6. Body concolorous pale yellow. Penis typical of californica group. ALLOTYPE.—Female. Similar to male. Ty PE-SPECIMENS.—Holotype, allotype, 3 paratypes (1 9, 2 juv.): Heater Cave, 8 km N Columbia, Calaveras County, Cal- ifornia, 15 Mar. 1979, D. C. Rudolph, B. Martin, and S. Win- terath. ETYMOLOGY.—This species is named for arachnologist Barbara Martin and for Army Corps of Engineers environmental planner Rob- ert Martin. BRIGGS AND UBICK: CAVE HARVESTMEN 319 FIGURE 3. Banksula grubbsi new species (Figure 4) HoLotyPeE.—Male. Body length 1.49 mm; scute length 1.26 mm; scute width 1.23 mm; eye tubercle length 0.25 mm; eye tubercle width 0.35 mm; operculum length 0.18 mm; operculum width 0.21 mm. Palp: trochanter 0.19 mm, femur 0.70 mm, patella 0.39 mm, tibia 0.53 mm, tarsus 0.65 mm. Leg II: trochanter 0.18 mm, femur 1.49 mm, patella 0.44 mm, tibia 1.23 mm, metatarsus 0.96 mm, tarsus 1.49 mm. Scute with segmentation delineated by small tubercles, area behind eye tubercle tuberculate. Eye tubercle obliquely truncate, declining pos- teriorly; entire surface tuberculate. Eyes with small corneas and no retinae. Tergites with sub- marginal row of tubercles. Second endites se- tose, with acute ectal invagination behind mid- point. Labial processes large, rounded. Small operculum with rounded anterior, margin adja- cent to fourth coxae, posterior margin slightly behind mesal extreme of posterior margin of fourth coxae. Chelicerae setose, only slightly tuberculate. Palpal femur with four prominent proximal dorsal spines and three reduced distal dorsal spines; no setae or tubercles between proximal dorsal spines; ectal spur at base replaced by a broad, low tubercle; ventral margin straight. Palpal tibia without significant proximal ventral spine. Tarsal formula 4-6-5-6. Body concolorous light yellow. Penis typical of californica group, aedeagal velum held in smooth dorsal plate. Banksula martinorum Briggs and Ubick, new species. Lateral view of body and left palpus of male holotype. TyYPE-SPECIMEN.—Holotype: Black Chasm Cave, near Vol- cano, Amador County, California, 19 Feb. 1978, A. G. Grubbs. FEMALE.—Unknown. ETYMOLOGY.—This species is named for biospeleologist Andrew G. Grubbs. Banksula elliotti new species (Figure 5) HoLotyPeE.—Male. Body length 1.61 mm; scute length 1.23 mm; scute width 1.28 mm; eye tubercle length 0.26 mm; eye tubercle width 0.35 mm; operculum length 0.30 mm; operculum width 0.32 mm. Palp: trochanter 0.23 mm, femur 0.63 mm, patella 0.39 mm, tibia 0.53 mm, tarsus 0.39 mm. Leg II: trochanter 0.16 mm, femur 1.14 mm, patella 0.35 mm, tibia 0.96 mm, metatarsus 0.72 mm, tarsus 1.14 mm. Scute with segmentation delineated by tuber- cles, eye tubercle and area behind tuberculate. Eye tubercle rounded, slightly conical, with small corneas and without retinae. Tergites with row of tubercles at margin. Second endites se- tose, mesally broad and rounded; labial pro- cesses spatulate. Operculum large, extending posterior to hind coxae. Hind sternite and anal plate with row of tubercles. All coxae tubercu- late. Chelicerae with strongly tuberculate anterior margin. Palpal femur with six prominent proximal dor- sal spines and four reduced distal dorsal spines; no setae or tubercles between proximal dorsal spines; ectal spur well developed at base of 320 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 11 Ficure 4. Banksula grubbsi Briggs and Ubick, new species. Lateral view of body and left palpus of male holotype. proximal ventral spine, ventral surface tuber- culate. Tarsal formula 4-6-5-6. Body concolorous yellow-orange. Penis with folds at base of dorsal plate; apex of aedeagus enclosed in rounded sheath and held within distal bifurcation of dorsal plate. ALLOTYPE.—Female. Similar to male. TYPE-SPECIMENS.—Holotype, allotype, 4 paratypes (2 d, 1 2, 1 juv.): Pinnacle Point Cave, near Camp Nine Power House, 9.6 km N Columbia, Tuolumne County, California, 8 Feb. 1979, D. C. Rudolph, B. Martin, and S. Winterath. Para- types, 2 gd, 2 juv.: same locality, 20 Jan. 1978, W. Elliott, A. Grubbs, and S. Winterath. Paratypes, 2 d, 1 2: same locality, 1 May 1977, A. Grubbs, N. Boice, M. McEachern and J. Davis. Paratypes, | 2: Digger Pine Cave, 6.5 km N Columbia, Calaveras County, 22 Mar. 1979, D. Rudolph, B. Martin, S. Winterath, W. Elliott, and J. Reddell. Paratypes, 3 5, 5 9: Rabbit Hole Cave, 6.5 km N Columbia, Calaveras County, 22 Mar. 1979, D. Rudolph, B. Martin, and S. Winterath. Para- types, | d, 1 2: Grapevine Gulch Cave, 6.5 km N Columbia, Calaveras County, 22 Nov. 1979, D. Rudolph, B. Martin, and S. Winterath. Paratypes, | ¢, 4 2: same locality, 27 Apr. 1977, B. Hawson, A. Grubbs, J. Munthe, and M. McEachern. ETyMOLOGY.—This species is named for biospeleologist William R. Elliott. VARIATIONS.—Specimens taken in Pinnacle Point Cave are without retinae, although half of the specimens taken in the remaining caves have retinal pigment in their eyes. The specimen from Digger Pine Cave has a relatively more conical eye tubercle. Some specimens have only three distal dorsal spines on the palpal femur. Banksula grahami Briggs Banksula grahami BricGs 1974:7. Recorps.—Calaveras County: Moaning Cave, near Valle- citos, 22 Aug. 1963, 22 Dec. 1968, 6 Dec. 1977, R. Graham, T. Briggs, W. Elliott, A. Grubbs, and S. Winterath; Linda’s Cave, 6 km w Columbia, 16 May 1977, A. Grubbs, N. Boice, and D. Broussard; Carlow’s Cave, 6 km w Columbia, 16 May 1977, A. Grubbs, N. Boice, and D. Broussard. Tuolumne County: Experimental Mine Cave, 3 km N Columbia, 25 Jun. 1975, R. Lem; Mine tunnel along road to Experimental Mine, 2.5 km N Columbia, 25 Jun. 1975, T. Briggs (identification only); Snell’s Cave, 3 km N Columbia, 26 Feb. 1978, 14 Feb. 1978, 2 Apr. 1979, S. Winterath, D. Rudolph, and J. Reddell; Crystal Palace Cave, 5 km N Columbia, 4 Nov. 1967, 21 Dec. 1977, 4 Feb. 1979, 25 Mar. 1979, T. Briggs, V. Lee, D. Ru- dolph, S. Winterath, A. Grubbs, W. Elliott, B. Martin, and J. Reddell; Porcupine Cave, 5 km N Columbia, 4 Feb. 1979, 28 Feb. 1979, D. Rudolph, S. Winterath, and B. Martin; mine on ridge, 4.5 km N Columbia, 22 Feb. 1979, D. Rudolph, S. Win- terath, and B. Martin; Banksula Cave, 6 km N Columbia, 21 Mar. 1979, D. Rudolph, B. Martin, S. Winterath, and W. El- liott; McLean’s Cave, 4.5 km N Columbia, 13 May 1967, 14 Dec. 1977, 18 Dec. 1977, 6 Mar. 1979, 27 Mar. 1979, 2 Apr. 1979, K. Hom, T. Briggs, W. Elliott, A. Grubbs, S. Winterath, D. Rudolph, and B. Martin; Transplant Mine, 3 km N Colum- bia, 17 Apr. 1979, D. Rudolph, S. Winterath, and E. vanIngen. Notes.—The migration of Banksula grahami into two of the mine tunnels recorded above is the only known example of probable interstitial movement by a species of Banksula. Banksula melones Briggs Banksula melones BriGGs 1974:8. Recorps.—Calaveras County: Cave of Skulls, 5 km Nw Columbia, 16 Apr. 1977, 29 Mar. 1979, T. Briggs, D. Rudolph, B. Martin, S. Winterath, W. Elliott, and J. Reddell; Quail (Gerritt’s) Cave, 5 km Nw Columbia, 3 Jul. 1975, 17 Apr. 1977, 30 Mar. 1979, W. Rauscher, D. Cowan, B. Martin, and S. Winterath; Barren Cave, 5 km Nw Columbia, 30 Mar. 1979, D. Rudolph, B. Martin, S. Winterath, W. Elliott, and J. Red- dell; Beta Cave, 5 km Nw Columbia, 7 May 1977, 29 Mar. 1979, A. Grubbs, D. Broussard, S. Winterath, D. Rudolph, W. Elliott, J. Reddell, and B. Martin; Poison Oak Cave, 5 km NW Columbia, 29 Mar. 1979, D. Rudolph, B. Martin, S. Win- terath, W. Elliott, and J. Reddell; Coral Cave, 5 km N Colum- bia, 24 Feb. 1978, 22 Mar. 1979, A. Grubbs, D. Rudolph, B. Martin, S. Winterath, W. Elliott, and J. Reddell; Bryden’s Cave, 5 km Nw Columbia, 29 May 1977, A. Grubbs and B. Hopkins; Cone Cave, 5 km Nw Columbia, 30 Mar. 1979, D. Rudolph, B. Martin, S. Winterath, W. Elliott, and J. Reddell; Eagle View Cave No. 2, 5 km Nw Columbia, 29 Mar. 1979, D. Rudolph, B. Martin, S. Winterath, W. Elliott, and J. Red- BRIGGS AND UBICK: CAVE HARVESTMEN FIGURE 5. dell; Lost Piton Cave, 6.5 km w Columbia, 6 May 1977, 26 Mar. 1979, A. Grubbs, N. Boice, D. Broussard, S. Winterath, W. Elliott, and J. Reddell; Bone Cave, 6 km w Columbia, 20 Apr. 1980, T. Briggs and D. Ubick. Tuolumne County: Quarry (McNamee’s) Cave, 3 km Nw Columbia, 24 May 1969, G. Leung, W. Rauscher, and T. Briggs; Gate Pit Cave, 3 km Nw Columbia, 1 Apr. 1979, D. Rudolph; Mine Cave, 3 km Nw Columbia, 1 Apr. 1979, D. Rudolph, B. Martin, and S. Win- terath; Scorpion Cave, 4.5 km N Columbia, 8 Feb. 1979, 25 Mar. 1979, D. Rudolph, S. Winterath, D. Cowan, and T. Briggs; Vulture Cave, 4.5 km N Columbia, 10 Feb. 1979, 17 Feb. 1979, 22 Feb. 1979, 19 Mar. 1979, S. Winterath, D. Ru- dolph, B. Martin, and W. Elliott; McLean’s Cave, 4.5 km n Columbia, 13 May 1967, 17 June 1967, 24 June 1967, 14 Dec. 1977, 18 Dec. 1977, 6 Mar. 1979, 27 Mar. 1979, K. Hom, T. Briggs, V. Lee, W. Elliott, A. Grubbs, S. Winterath, D. Ru- dolph, and B. Martin; Transplant Mine, 3 km N Columbia, 17 Apr. 1979, D. Rudolph, S. Winterath, and E. vanIngen. Notes.—Some of the specimens collected in Vulture Cave and Bone Cave were found under rocks in the twilight zone. All other collections were made in permanently dark regions of caves. ECOLOGICAL REVIEW Several ecological studies on Banksula (Briggs _ 1975; Elliott 1978; Rudolph 1979) have been con- ducted in McLean’s Cave at the confluence of the Middle Fork and the South Fork of the Stan- islaus River (approximate elevation, 300 m). This cave is one of the largest in the Calaveras limestone of the central Sierra Nevada and con- tains the only sympatric populations of Bank- sula species. Banksula melones and B. grahami are mixed in habitats near the base of a broad talus cone formed by debris gradually moving through two small entrances to the lowest levels of the cave. Because there is no flowing water in the cave, the talus cone and deep-penetrating roots are important food sources for the inhab- itants. The cave temperature ranges from 14 C Banksula elliotti Briggs and Ubick, new species. Lateral view of body and left palpus of male holotype. to 16 C and the humidity from 82% to 97% (El- liott 1978). Specimens of Banksula melones and B. grahami were found under rocks or wander- ing on the floor or walls. They were rarely cap- tured in baited pitfall traps (Briggs 1975; Elliott 1978). Although biased by more intensive winter collecting, records from McLean’s and other caves suggest greatest activity in winter and spring. At least 30 species of arthropods, most of which were listed by Elliott (1978), coexist with Banksula melones and B. grahami in McLean’s Cave. Elliott (1978) identified some species, but many have not yet been identified. His feeding experiments showed that captive Banksula, kept in McLean’s Cave, will eat live Collembola but not equally abundant Psocoptera. Rudolph (1979) maintained eight immature Banksula in McLean’s Cave using Collembola as food; three molted within 43 days. One adult and two im- mature individuals maintained at the same time without food survived. Rudolph (1979) searched the mine tunnel into which Briggs and Elliott transplanted Banksula melones, B. grahami, and miscellaneous arthro- pods from McLean’s Cave. The transplanted population, which included a few hundred Banksula, appeared to be reproducing. The fu- ture of this transplant may depend on how well conditions in the mine duplicate conditions in McLean’s Cave. If they both flourish, the trans- planted Banksula species may demonstrate that their sympatry in a small cave is stable. BIOGEOGRAPHY The distribution of Banksula species is similar to the invertebrate troglobite distributions re- ported from caves in the Appalachian Mountains 322 of the eastern United States in that these cave species tend to be bounded within ‘‘karst is- lands,’’ within which subterranean dispersal and genetic communication readily occurs (Barr 1967; Culver et al. 1973). Such karst-island spe- ciation exists for Banksula in the Sierra Nevada if the limestone outcrops are widely separated. Thus, the species B. galilei Briggs, B. califor- nica (Banks), B. rudolphi new species, B. grubbsi new species, and B. tuolumne Briggs . occupy karst that is separated by many kilo-, meters of nonporous rock (Fig. 1). The isolation’ of these cave species is, therefore, more com- plete than that of the species in the Appalachian caves. The area of the karst in which B. galilei, B. californica, and B. rudolphi are found is 0.02—0.1 km? and is considerably smaller than the smallest karst island (10 km?) studied in the eastern United States (Culver et al. 1973). Fur- thermore, quarrying shows that these small out- crops have small volume and little internal water flow. All known Banksula species are allopatric ex- cept for B. melones and B. grahami in Mc- Lean’s Cave. This suggests that sympatric species compete, and the sympatry in McLean’s Cave is a result of a recent invasion. About half of the 70 or so Calaveras Formation caves care- fully checked yielded a species of Banksula. Thirty-one of these are located in the karst of the Stanislaus River. Eighteen apparently hab- itable caves in this region did not contain Bank- sula (Rudolph 1979). The absence of harvest- ment in caves located in karst outside of the Stanislaus River region may be due to an uneven distribution of their epigean ancestors. The distribution of the four species of Bank- sula (B. melones, B. grahami, B. martinorum, and B. elliotti) that occupy the karst of the Stanis- laus region is not readily explained. Volcanic rock divides Coyote Creek (west of the Stanis- laus River) from the Stanislaus River. It forms an irregular barrier between B. elliotti and B. melones, and separates B. melones from B. gra- hami southeast of the Stanislaus River. Several continuous bands of amphibolite divide the re- maining limestone regions into parallel lenses in which allopatric populations of B. melones and PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 11 B. grahami occupy most of the available cav- erns. These populations seem to split into east- ern and western clusters that do not correlate well with geology or water systems. The karst that contains the easternmost population of B. elliotti is isolated by one kilometer of metavol- canic rock from the karst that contains the rest of B »elliotti and B. martinorum. Coyote Creek, ‘the much. larger Stanislaus River, and the south “fork of the Stanislaus that branches to the east ‘have no apparent. effect on the distribution of species of Banksula. The four Stanislaus species show increasing cavernicolous specialization in relative eye loss, from B. melones with the larg- est eyes, to B. elliotti with missing retinae, and B.- martinorum with complete eye loss. One might infer that adaptive radiation began with a B. melones-like ancestor and progressed to B. elliotti. If correct, one must explain how the in- creasingly troglobitic species B. grahami and B. elliotti were able to cross geologic and drainage barriers. Further studies on the harvestmen of the Stanislaus River region are needed to explain this unexpected distribution of species. It is re- grettable that this portion of the river will soon be inundated by the New Melones Reservoir and some of the harvestmen caves will be lost. LITERATURE CITED Barr, T. C., Jn. 1967. Observations on the ecology of caves. Am. Nat. 101(922):475—491. BricGs, T. S. 1968. Phalangids of the laniatorid genus Sital- cina (Phalangodidae:Opiliones). Proc. Calif. Acad. Sci. 36(1):3-7. . 1971. Relict harvestmen from the Pacific Northwest. Pan-Pac. Entomol. 47(3): 168-169. . 1974. Phalangodidae from caves in the Sierra Nevada (California) with a redescription of the type genus. Occas. Pap. Calif. Acad. Sci. 108:1-15. 1975. Biological transplant project, New Melones Lake, California—final report. Serial #DACWO05-75-P-1845, U.S. Army Corps of Eng., Sacramento Dist., Calif. Cutver, D., J. R. HOLSINGER, AND R. BARoopDy. 1973. To- ward a predictive cave biogeography: the Greenbrier Valley as a case study. Evolution 27(4):689-695. Etutiotr, W. R. 1978. Final report on the New Melones cave harvestman transplant. Contract #DACW05-78-C-0007, U.S. Army Corps of Eng., Sacramento Dist., Calif. Rupocpn, D. C. 1979. Final report on the status of the Me- lones cave harvestman in the Stanislaus River drainage. Contract #14-16-0009-79-009, U.S. Fish Wildl. Serv., Wash. D.C. CALIFORNIA ACADEMY OF SCIENCES Golden Gate Park San Francisco, California 94118 Vol. 42, No. 12, pp. 323-339, 34 figs. PROCEEDINGS OF THE... CALIFORNIA ACADE VOF 51 : EE I i Laberatory | { IGE SE ONG Woodgs Nole, Maes. | TROPICAL EASTERN PACIFIC LIMPETS OF THE FAMILY ACMAEIDAE (MOLLUSCA, ARCHAEOGASTROPODA): GENERIC CRITERIA AND DESCRIPTIONS OF SIX NEW SPECIES FROM THE MAINLAND AND THE GALAPAGOS ISLANDS By David R. Lindberg* Center for Coastal Marine Studies, University of California, Santa Cruz, California 95064 and James H. McLean Section of Malacology, Natural History Museum of Los Angeles County, Los Angeles, California 90007 ABSTRACT: ters. Lottia, previously considered monotypic, is expanded to include Panamic species with a secondary gill (branchial cordon) formerly assigned to Scurria. Scurria has a similar gill, but the shell structure differs. The new species Notoacmea ubiquita from Mexico and N. pumila from Ecuador are small-shelled allopatric species with radular teeth modified for feeding on coralline algae. Two new species of Notoacmea (N. rothi and N. immaculata), endemic to the Galapagos Islands, constitute a species pair differing chiefly in radular features: the radular teeth of N. immaculata are adapted for feeding on calcareous algae; those of N. rothi for noncal- careous algae. A pair of endemic new species of Lottia from the Galapagos Islands (L. mimica and L. smitht) also differ mainly in radular characters. Lottia mimica is a noncalcareous-alga feeder and L. smithi is a calcar- eous-alga feeder. These four endemic species are the principal acmaeid limpets of the Galapagos. Two mainland We define genera on conservative shell structure characters and on qualitative radular charac- 24, 1981 species, Notoacmea filosa and Lottia mesoleuca, are known only sporadically from the Galapagos Islands. INTRODUCTION The last comprehensive, illustrated review of the Acmaeidae of the tropical eastern Pacific was given by McLean (1971). At the time of preparation of that account, two of the species * Research Associate, Department of Invertebrate Zoology, California Academy of Sciences, San Francisco, California 94118. described herein—Notoacmea ubiquita and Lottia mimica—were recognized as new but were not described. The generic placement of these two species was puzzling because they had previously unknown combinations of radular, shell, and gill characters. Further study of generic relationships has now provided a basis for the convincing allocation of these species. Although a full review of generic [323] 324 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 12 criteria in the family is beyond the scope of this paper, we include some discussion pertaining to the Panamic species. A major distinction has become apparent between Lottia Sowerby, 1834, and Scurria Gray, 1847, two genera having a secondary gill (accessory gill lappets on the mantle margin). These two genera are redefined here, the name Lottia thus being made available for use for some tropical species previously con- sidered to belong to Scurria. A closer examination of the acmaeids of the Galapagos Islands has resulted in the recogni- tion of four new endemic species, representing two species pairs wherein the principal differ- ences are in radular tooth morphology. The shell characters of each pair are insufficiently distinct to permit reliable identification by shell alone. Radular characters in the Acmaeidae have been found by all workers to be species-specific. In no species has ontogenetic or situs variation in radulae been found. Similar shell morphologies have been reported, however, in both conge- neric and noncongeneric species of Acmaeidae (McLean 1966; Lindberg 1979). We therefore consider each radular morphotype to represent a separate species. Because the shell characters of each pair are insufficiently distinct to permit reliable identification, both species are dis- cussed in a combined discussion section follow- ing their formal descriptions. Abbreviations are as follows: AHF, Allan Hancock Foundation, University of Southern California, Los Angeles (collection on loan to LACM); AMNH, Department of Invertebrates, American Museum of Natural History, New York; ANSP, Department of Malacology, Acad- emy of Natural Sciences, Philadelphia; CAS, Department of Invertebrate Zoology, California Academy of Sciences, San Francisco; LACM, Section of Malacology, Natural History Mu- seum of Los Angeles County, Los Angeles; MCZ, Museum of Comparative Zoology, Har- vard University, Cambridge; SU, Stanford Uni- versity, Stanford (collection on loan to CAS); USNM, Division of Mollusks, U.S. National Museum of Natural History, Washington, D.C. GENERIC CRITERIA FOR THE PANAMIC ACMAEIDAE Generic assignments in McLean’s (1971) re- view were based on shell sculpture, presence or absence of the secondary gill, and whether mar- ginal radular teeth are represented by two pairs of fully developed teeth, a single pair of rudi- mentary teeth (uncini), or are absent altogether. We now realize that a system based on these three characters alone is not adequate. Christiaens (1975) proposed a generic classi- fication of the Acmaeidae in which tooth shape and configuration were especially emphasized. He felt some genera had three pairs of lateral teeth and some two, the latter group having a bicuspid second lateral tooth. We maintain that all acmaeids have three pairs of lateral teeth. We fail to see how the third lateral tooth can be interpreted as part of the second, because in all acmaeid radulae we have examined, we find that the ventral plates of the radular ribbon have three lateral plate components, one correspond- ing to each lateral tooth. We believe that the reduction of the outermost tooth that occurs in some species is a result of dietary specialization. Relation of diet to tooth shape was discussed by McLean (1966), and we are now aware of similar tooth shape and configuration in species of di- verse genera. We do not consider lateral tooth shape to be useful as a generic character. We continue to maintain full generic separa- tion of species groups in which the marginal teeth have three possible expressions: (1) two pairs of fully functional marginals (Patelloida Quoy and Gaimard, 1834); (2) a single pair of marginal remnants or uncini (Collisella Dall, 1871; Lottia Sowerby, 1834; Scurria Gray, 1847); and (3) no marginals or uncini (Acmaea Eschscholtz, 1833; Notoacmea Iredale, 1915; Problacmaea Golikov and Kussakin, 1972; Rhodopetala Dall, 1921; and Tectura Gray, 1847). We therefore disagree with Christiaens’s ranking of Notoacmea as a subgenus of Colli- sella. Recent work by Lindberg (1976, 1978) has em- ployed shell structure characters first used for patellacean limpets by MacClintock (1967). We believe that the relationships suggested by shell structure are conservative and are basic to a modern classification of the family. We are now inclined to define genera using shell structure, branchial characters, radula bas- al plate structure, and the three possibilities for marginal teeth listed above, recognizing that shell sculpture characters are convergent in all genera and that lateral tooth shape is likewise LINDBERG & McLEAN: TROPICAL EASTERN PACIFIC ACMAEIDAE 325 Ficures 1-4. Ventral views of preserved specimens of Loftia and Scurria species showing secondary gill in relation to head. Ficure 1. Lottia gigantea, Isla de Guadalupe, Mexico (LACM 55618). FiGureE 2. Lottia mesoleuca, Bahia Tenacatita, Jalisco, Mexico (LACM 66-55). FiGuRE 3. Lottia mimica new species, paratype, Academy Bay, Isla Santa Cruz, Galapagos Islands, Ecuador (LACM 1926). FiGure 4. Scurria scurra, Punta El Lacho, Santiago Province, Chile (LACM 75-32). convergent and widely variable interspecifically (although not intraspecifically).! Acmaea and Notoacmea differ in lateral plate morphology. In Acmaea the lateral plates are similar in size and shape and are arranged in a posteriorly diverging V-configuration. In No- toacmea the lateral plates are unequal in size and shape—the first and second lateral plates tend to lie in the same line and the third lateral plates are always lateral and slightly posterior to the second lateral plates. 'In his discussion of Notoacmea fascicularis (Menke, 1851) McLean (1971:327) alluded to two different radular types within that species, suggesting that a “complex involv- ing more than one species’ was a possibility. Lindberg will report separately on the two species of the N. fascicularis complex. Acmaea and Tectura also differ in lateral plate morphology. In both genera dentition consists of three pairs of equal-sized and equal-shaped lateral teeth; however, in Tectura the lateral plates that support these teeth are complex and similar in shape and position to those found in the genus Collisella. All species of Acmaea are known to feed on coralline algae and have blunt, equal-sized teeth. Some of the tropical eastern Pacific and Carib- bean species of Notoacmea are now known to have teeth similarly blunt and of equal size. These species may also be coralline alga feeders. Three of the four new species of Notoacmea described in this paper (N. ubiquita, N. pumila, and N. immaculata) have blunt equal-sized lat- eral teeth. The other new species of Notoac- mea, N. rothi, has the outermost lateral tooth 326 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 12 greatly reduced and the first two pairs more elongate, which is the pattern characteristic of most temperate and tropical species of Notoac- mea. Similar modification of the lateral teeth for feeding on coralline algae is known in some trop- ical species of Collisella. Eastern Pacific species of this uncinate genus with lateral teeth so mod- ified are: C. atrata (Carpenter, 1857), C. discors (Philippi, 1849), C. mitella (Menke, 1847), and C. pediculus (Philippi, 1846). Two generic names have been used for ac- maeid limpets in which there is a secondary gill (branchial cordon) in addition to the normal ac- maeid ctenidium: Lottia Sowerby, 1834 (type- species L. gigantea Sowerby, 1834), and Scur- ria Gray, 1847 (type-species Patella scurra Lesson, 1830). Lottia has usually been consid- ered monotypic, with the single Californian species L. gigantea. It has been diagnosed (Dall 1871) as having a secondary gill incomplete or interrupted in front of the head, whereas in Scurria the gill is complete or continuous. The radular dentition in both genera consists of three pairs of lateral teeth and one pair of uncini. We have examined the secondary gill in living and preserved specimens of L. gigantea and find that many specimens have a greatly reduced but distinct gill in front of the head (Fig. 1). The secondary gill of a tropical eastern Pacific species usually assigned to Scurria, S. mesoleu- ca (Menke, 1851), is normally much less prom- inent in front of the head than along the sides (Fig. 2). The secondary gill of Lottia mimica, new species (Fig. 3), is also much reduced in front of the head. The secondary gill of Scurria scurra (Fig. 4) is complete over the head, but it is also somewhat reduced in prominence in this region. We therefore do not regard the reduction of the secondary gill near the head as a useful generic character. MacClintock (1967) found that Scurria in the Peruvian faunal province differ in shell structure from other eastern Pacific species with the sec- ondary gill. The Peruvian Scurria species are in MacClintock’s shell structure ‘‘group 3,”’ whereas Lottia gigantea and the two species placed by McLean (1971) in Scurria (S. meso- leuca and S. stipulata (Reeve, 1855)) are in shell structure “‘group |’ (along with most other species of Collisella and Notoacmea). Because we believe that shell structure is more conser- vative than branchial characters, and we place even less emphasis on shell sculpture and col- oration, we infer that the Panamit acmaeids pre- viously assigned to Scurria are more closely re- lated to L. gigantea than to Scurria. The two Panamic species plus L. mimica and L. smithi described in this paper are therefore assigned to Lottia. Lottia is redefined to include uncinate species in shell structure “‘group 1,’’ with a sec- ondary gill that is usually reduced but not nec- essarily absent over the head. New SPECIES OF ACMAEIDAE FROM THE TROPICAL EASTERN PACIFIC Notoacmea ubiquita new species (Figures 5-7, 23, 29) There are two situs forms of this species, a laterally compressed form and an oval form. A description for each follows. Shell (oval form) (Figs. 5, 6): Relatively small (maximum length 12 mm), profile of medium height; apex anterior to center; all slopes con- vex; large shells frequently with a flattened area posterior to apex; sides of shell somewhat par- allel. Sculpture of rounded radial ribs, with weaker secondary ribs beginning below apex; ribs extending slightly beyond the shell edge, crenulating the aperture; concentric sculpture of well-defined but nearly microscopic, sharply raised ridges. Exterior translucent white with red-brown markings on early shell, the markings becoming darker, reticulate, and limited to rib- interspaces with growth; ribs white, overlain with dark brown radial markings that may be concentrated into lateral rays. Interior margin white with dark markings that correspond to ex- terior interspaces; intermediate area white; cen- tral area with yellow stain, exterior markings visible through shell. Shell (compressed form) (Fig. 7): Lateral pro- file high, ends raised relative to sides; all slopes convex; sOme specimens compressed in early stage, changing abruptly to oval form. Sculpture and coloration as in oval form. Radula (Figs. 23, 29): First pair of lateral teeth closely set at anterior edge of ribbon segment, medial edges convex, lateral edges straight to slightly concave, cusps rounded, blunt; second pair of lateral teeth posterior and lateral to first pair, medial and lateral edges convex, cusps rounded, blunt; third lateral teeth lateral to sec- LINDBERG & McLEAN: TROPICAL EASTERN PACIFIC ACMAEIDAE vs) tw ~— Ficures 5-10. FiGure 5. Notoacmea ubiquita new species. Holotype, LACM 1917. Santiago Peninsula, Colima, Mexico. ~ Length 11.7 mm. FiGure 6. Notoacmea ubiquita. LACM 54773. Guaymas, Sonora, Mexico. Length 6.9 mm. FiGurRe 7. Notoacmea ubiquita. LACM 54773. Guaymas, Sonora, Mexico. Length 5.4 mm. FiGure 8. Notoacmea pumila new species. Holotype, LACM 1919. Punta Ancon, Ecuador. Length 4.7 mm. FIGURE 9. Notoacmea pumila. Paratype, LACM 1920. Punta Ancon, Ecuador. Length 3.6 mm. FiGurE 10. Notoacmea pumila. LACM 72-17. Bahia Jobo, Costa Rica. Length 5.6 mm. ond pair, medial edges concave, lateral edges Straight, extending to edges of ventral plates, cusps blunt. Marginal teeth lacking. First lateral plates overlap anterior ribbon segment, postero- lateral edges concave; second lateral plates ir- regular, posterior edges convex; third lateral plates lobate with lateral lobes extending to edges of ventral plates; second and third lateral plates separated by a partial suture. Ventral plates closely set with both anterior and poste- rior processes. Animal: Pigmentation lacking, snout with oral lappets. Ho.LotyPe DIMENSIONS.—Length 11.7, width 8.1, height 3.6 mm. Type-LocaALity.—Mekxico: Colima; Manza- nillo, Santiago Peninsula, Playa Las Hadas (19°05'57’"N, 103°19'36”W) (LACM 63-10), inter- 328 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 12 tidal zone to 5 m. Leg. J. H. McLean and C. Tenney, 21-24 Mar. 1963, 3 specimens. TypeE-MATERIAL.—Holotype (oval form), LACM 1917; 2 paratypes, LACM 1918 (both oval and compressed forms). The holotype is the largest specimen examined. DISTRIBUTION.—Baja California Sur, Mexico, from Punta Pequena (26°14’N) (LACM 71-6) to Bahia Magdalena and Cabo San Lucas, north in the Gulf of California to Puertecitos (30°25’N) (LACM 65-34) and Guaymas, south to Bahia Tangola Tangola, Oaxaca (15°45'N) (AHF 215). MATERIAL EXAMINED.—58 lots, approxi- mately 350 specimens, 3 radula preparations. ETYMOLOGY.—The name is based on the Lat- in adverb uwbique (everywhere). The species is indeed ubiquitous—shells, at least, have been recovered from sediment residues taken by divers from nearly all LACM station localities throughout the range. Discussion.—Although Notoacmea ubiquita has been known for many years, it was not de- scribed earlier because of uncertainties about its generic position. Its shell shape and sculpture suggested Collisella, and its lack of uncini and subtidal habitat suggested Acmaea. We are now satisfied to place it in Notoacmea because of the basal plate configuration. Although most species of Notoacmea are finely ribbed, the rather more prominent ribbing of N. ubiquita is not as strong as occurs in many species of Col- lisella. Notoacmea ubiquita is the only Panamic ac- maeid with an oval form and a laterally com- pressed form. Notoacmea ubiquita has broader, stronger ribs than any other tropical species of Notoacmea. On the basis of shell characters, it is more similar to some of the Panamic Colli- sella, which differ in having an uncinate radula. The fine brown concentric markings of the early stages do not occur in any other species. Col- lisella turveri (Hertlein and Strong, 1951) differs in having broader, more projecting ribs. Colli- sella acutapex (Berry, 1960) has a higher shell profile with sharper, more prominent ribbing, and its pattern of brown lines is more coalescing. Collisella mitella (Menke, 1847) also has white ribs, but its ribs are more numerous and the in- terspaces are dark colored. Patelloida semiru- bida (Dall, 1914) has sharper radial and concen- tric sculpture, with red rather than brown markings; its radula is also markedly different, having two pairs of marginal teeth per ribbon segment. Large lots show a complete series of possible shell shapes between the elevated narrow forms with raised ends and the low oval forms. Some shells have an early compressed phase, with lat- er growth stages like the oval form; some rela- tively large shells are angulate at the sides, giv- ing the shell a flat-topped appearance. Color variation is relatively minor; one color variant characteristic of specimens from Jalisco is or- dinary in early stages, changing to solid maroon at later stages. Largest shells seen are from Ja- lisco and Colima; specimens from localities in the Gulf of California attain about two-thirds the size of southern specimens. Notoacmea ubiquita has features in common with two more northern species, Collisella triangularis (Carpenter, 1864) and Tectura ro- sacea (Carpenter, 1864), both of which differ in lacking the radial ribbing. All three species have laterally compressed forms, are primarily sub- tidal, and have equal-sized lateral teeth adapted for feeding on calcareous algae. In C. triangu- laris the compressed form predominates, where- as in T. rosacea the oval form is more abundant, but in both species the compressed forms occur on branching coralline algae and the oval forms occur on crustose coralline algae, and all inter- mediate conditions are known. Although we have not directly observed the compressed form of N. ubiquita on branching coralline algae, it probably so occurs, judging from its ability to change from compressed to oval during growth, which implies a change of situs. Notoacmea pumila new species (Figures 8-10, 24, 30) Shell (Figs. 8-10): Small (maximum length 7 mm); profile medium-high; apex anterior to cen- ter; anterior slope straight to convex, lateral slopes convex; usually encrusted with coralline algae. Sculpture of fine, sharp radial ribs origi- nating below apex, secondary ribs arising in the interspaces, not reaching thickness of primary ribs. Aperture oval, not crenulate. Color pattern independent of ribbing: most frequently white near apex, gray at margin, with 6 to 10 white rays in a stellate pattern, some rays not reaching margin; some specimens with fine brown lines bordering white rays and fine brown lines that LINDBERG & McLEAN: TROPICAL EASTERN PACIFIC ACMAEIDAE 329 produce a concentric network. Interior translu- cent white, showing the exterior pattern. Raduia (Figs. 24, 30): First pair of lateral teeth closely set at anterior edge of ribbon segment, medial edges convex, lateral edges concave, cusps rounded. Second pair of lateral teeth pos- terior to first pair, medial edges convex, lateral edges straight, cusps rounded. Third pair of lat- eral teeth positioned posterior and lateral to sec- ond pair, medial edges convex, lateral edges straight to slightly concave. Third laterals broader than second laterals, with lateral exten- sions to edges of ventral plates; cusps rounded. Marginal teeth lacking. First lateral plates irreg- ular, anterior portions overlapping anterior rib- bon segments; second lateral plates elongate, ovoid; third lateral plates triangular, with con- vex posterior edge. Ventral plates with strong anterior and posterior processes. Lateral por- tions with strong sutures parallel to edges. Animal; Pigmentation lacking, oral fringe sim- ple. HoLotyPe DIMENSIONS.—Length 4.7, width 3.3, height 2.0 mm. Type-LocALiry.—Ecuador: Santa Elena Peninsula; Punta Ancon, north and south sides (2°20'S, 80°54’W), intertidal zone. Leg. J. H. McLean and D. Shasky, 6-7 Mar. 1970 (LACM 70-11, 70-12), 72 LACM specimens, 12 Shasky specimens. TyPE-MATERIAL.—Holotype, LACM 1919, paratypes, LACM 1920; paratypes have also been deposited in the collections of CAS and USNM, and in Shasky collection (Redlands, California). DISTRIBUTION.—El Velero, Nicaragua (12°01'N) (LACM 74-86), south to Ecuador (type-locality). There are numerous dead spec- ~imens from Bahia Salinas, Costa Rica (11°02'N, 85°45’'W) (LACM 72-17, 72-19); two specimens only from Panama at San Carlos (8°29'N, 79°57'W) (LACM 75-55), and a number of lo- calities in Ecuador collected by D. Shasky. MATERIAL EXAMINED.—19 lots, approxi- mately 200 specimens, 3 radula preparations. EtTyMOoLoGy.—The name is a Latin adjective, pumilus, meaning small or dwarfish—fitting for this, the smallest tropical eastern Pacific mem- ber of the family. DiscUSSION.—Notoacmea pumila could be confused only with two other relatively small forms, N. ubiquita new species and Patelloida semirubida. It differs from the first in having much sharper ribbing and not being compressed. Although both N. pumila and P. semirubida have fine sharp radial ribs, N. pumila lacks the sharp concentric sculpture and pink markings of P. semirubida. The radula of N. pumila is similar to that of two new species described herein, N. ubiquita and N. immaculata. It differs from both by hav- ing a complete rather than partial suture be- tween the second and third lateral plates and having strong ventral plate sutures parallel to the lateral edges. The ventral plates of N. pum- ila have anterior and posterior processes which N. immaculata lacks, and the third lateral plates are triangular rather than biformed as in N. im- maculata and N. ubiquita. The radula of N. pumila differs from that of P. semirubida by lacking marginal teeth. Large lots show similar color patterns both in the material from Costa Rica and from stations in Ecuador. A small percentage of specimens change with growth from dark rayed to solid dark (see Fig. 9); fewer specimens are rayed only with brown linear markings and fine brown reticulate markings. Shell proportions vary only slightly. Notoacmea pumila undoubtedly feeds on cor- alline algae—the lateral teeth are blunt and of equal size. Living specimens have been collect- ed in the low intertidal zone and the species probably also occurs in the immediate subtidal zone on coralline-encrusted rocks. Notoacmea rothi new species (Figures 11-13, 25, 31) Shell (Figs. 11-13): Size medium (maximum length 20 mm), height medium; apex anterior to center; all slopes convex, aperture ovoid. Sculp- ture of unequal riblets and concentric growth lines; one to three secondary riblets between each two primary riblets. Exterior dark gray with scattered white markings; apical pattern tessellate; white markings often aligned in lateral rays that define a broad, dark posterior ray. In- terior margin broad, dark, streaked with white corresponding to exterior pattern; intermediate area blue-white; central area blue-white with brown stain. Radula (Figs. 25, 31): First pair of lateral teeth closely set at anterior edge of ribbon segment, medial edges convex, lateral edges slightly con- 330 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 12 Ficures 11-16. Notoacmea from the Galapagos Islands. FiGure 11. Notoacmea rothi new species. Holotype, LACM 1921. Wreck Bay, Isla San Cristobal. Length 16.8 mm. FiGure 12. Notoacmea rothi. LACM 54774. East side, Isla Fernandina. Length 11.5 mm. FiGure 13. Notoacmea rothi. LACM 54777. Bahia Cartago, Isla Isabela. Length 8.8 mm. FiGure 14. Notoacmea immaculata new species. Holotype, LACM 1923. Isla Baltra. Length 5.6 mm. FiGureE 15. Notoacmea immaculata. AHF 173-34. Isla Baltra. Length 7.8 mm. Figure 16. Notoacmea filosa (Carpenter, 1865). LACM 54775. Isla San Cristobal. Length 15.5 mm. cave, rounding to pointed cusps. Second pair of — First lateral plates subrectangular; second lat- lateral teeth lateral to first pair, both edges con- _ eral plates rounded, separated from third lateral vex, broad, with pronounced pointed cusps; plates by partial suture; third lateral plates bi- third lateral teeth lateral to second pair, re- formed, posterior section rounded, lateral sec- duced, medial edges convex, lateral edges tion pointed, extending to lateral edges of ven- straight, cusps angular. Marginal teeth lacking. tral plates. Ventral plates closely set, LINDBERG & McLEAN: TROPICAL EASTERN PACIFIC ACMAEIDAE 331 subrectangular, with weak anterolateral exten- sions. Animal: Body pigmentation lacking; snout with oral lappets. HoLortyPe DIMENSIONS.—Length 16.8, width 14.0, height 4.8 mm. TypeE-LocaLity.—Ecuador: Galapagos Is- lands; Isla San Cristobal, Wreck Bay (0°54’S, 89°36’W). Leg. J. DeRoy, 12 May 1968, 4 spec- imens. TyPE-MATERIAL.—Holotype, LACM 1921 (shell and radular slide), 1 paratype, LACM 1922; paratypes also deposited in the collections of CAS and USNM. DisTRIBUTION.—Galapagos Islands; Isla Fer- nandina (ANSP 152554), Isla Isabela (SU 239), Isla Rabida (LACM 71-69), Isla Bartolomé (AMNH 163290), Isla Santa Cruz (ANSP 154889), Isla Baltra (AMNH 163263), Isla Santa Maria (CAS 23025), Isla Santa Fé (AHF 48-33), Isla Espanola (USNM 102359), Isla San Cristo- bal (MCZ 205068). MATERIAL EXAMINED.—40 lots, 435 speci- mens, 9 radula preparations. ETYMOLOGY.—We are pleased to name the species in honor of Barry Roth of the California Academy of Sciences in recognition of his work in molluscan systematics. Notoacmea immaculata new species (Figures 14, 15, 26, 32) Shell (Figs. 14-15): Small (maximum length 12 mm), thin, diaphanous, height medium. Apex anterior to center, anteriorly directed; all slopes convex. Aperture ovoid; sides straight, sides elevated. Sculpture of faint gray, broad riblets and concentric growth lines. Exterior light gray, mottled with yellow-brown, brown, and white; darker markings concentrated into broad pos- terior ray bordered with white. Interior margin broad, dull, marked with exterior pattern; inter- mediate area translucent, glossy white; exterior pattern readily visible through shell; central area glossy, translucent, marked with sparse yellow streaks, central stain lacking. Radula (Figs. 26, 32): First pair of lateral teeth closely set at anterior edge of ribbon segment, medial edges convex, lateral edges straight to slightly concave, tapering to rounded cusps; second pair of lateral teeth posterior and slightly lateral to first pair, both edges convex, tapering to rounded cusps. First and second lateral teeth approximately equal in width. Third pair of lat- eral teeth posterior and lateral to second pair, medial edges convex, lateral edges elongate, straight to slightly concave, extending to edges of ventral plates, cusps rounded. Marginal teeth lacking. First lateral plates ovoid; second lateral plates rounded posteriorly, separated from third lateral plates by partial suture; third lateral plates lobate. Ventral plates closely set, subrect- angular with strong anterior sutures. Animal: Body pigmentation lacking; snout with oral lappets. Ho.LotyPe DIMENSIONS.—Length 5.6, width 4.3, height 1.4 mm. Type-LocaLiry.—Ecuador: Galapagos Is- lands; Isla Baltra (0°26'S, 90°17'W), Caleta del Norte, 0-3 m. Leg. ANTON Bruun, cr. 18B, sta. 791, 21 Sep. 1966, 1 specimen. TypeE-MATERIAL.—Holotype, LACM 1923 (shell and radula slide), 1 paratype, CAS 15920 (shell and radula slide). Paratype from Isla Santa Cruz, Academy Bay. DIsTRIBUTION.—Galapagos Islands; Isla Fer- nandina (LACM 62-196), Isla Isabela (LACM 71-70), Isla Bartolomé (AMNH_ 163290), Isla Santa Cruz (ANSP 154889), Isla Baltra (LACM 66-206), Isla San Cristobal (ANSP 153328). MATERIAL EXAMINED.—1]4 lots, 63 speci- mens, 5 radula preparations. EtyMoOLoGy.—The name is a Latin adjective, immaculatus (unstained), referring to the lack of a central stain in the area within the myo- stracum. DiscuUssIoN.—The radular difference that is the chief basis of the separation of the two species is unmistakable and qualitative: in N. rothi the third lateral teeth are reduced (Fig. 25) and in N. immaculata the third lateral teeth are large (Fig. 26). The lateral plate morphologies are correspondingly different. In N. rothi the second and third lateral plates are approximately equal in size, and the lateral edges of the third lateral plates form small pointed projections. In N. immaculata the third lateral plates are larger than the second lateral plates and the lateral pro- jections are rounded. The shells of N. rothi and N. immaculata have similar overall proportions and sculpture. The color pattern consists of radiating and scat- tered whitish tessellations, with the greatest concentration of white tessellations in two lat- ero-posterior rays, the posterior area between 332 the two rays having the least amount of tessel- late flecking so that it may appear to be a single, uniformly dark posterior ray. Those specimens confirmed on radular examination to have the tooth pattern of N. rothi have the dark gray- green ground color predominating, whereas those identified as N. immaculata have a light gray or white ground color. The largest speci- mens examined have proven to be N. rothi; the largest specimen verified as N. immaculata is 12 mm in length. The large specimens of N. rothi have a dark interior stain, which is generally lacking in N. immaculata. One small, stunted specimen verified as N. immaculata (LACM 71- 48) shows a slight trace of brown interior stain. It is possible that the brown stain is indicative of the attainment of size rather than a species- specific character. Too few specimens verified as N. immaculata are available to enable us to be certain that any shell characters may be used as proof of identity. Of the mainland acmaeid species, the No- toacmea rothiimmaculata complex resembles Notoacmea filosa (Carpenter, 1865) (Fig. 6), which has similar shell characters. They differ from N. filosa in the following ways: N. rothi- immaculata has a profile of medium height; N. filosa has a low profile. In N. rothi-immaculata the interspaces are broader than the riblets; in N. filosa the riblets are more numerous and the interspaces approximately equal in width to the riblets. Notoacmea filosa has a color pattern of radiating dark and lighter rays, often interrupt- ed, but not tessellated in circular or oval pat- terns. The tessellate markings are characteristic of N. rothi-immaculata. The dark posterior ray of N. rothi-immaculata is not a feature of N. filosa. Although the configuration of the lateral teeth of N. filosa has little in common with that of N. immaculata, there is a similarity between N. fi- losa and N. rothi. However, the shape of the second lateral teeth differs: in N. filosa the sec- ond lateral teeth are triangular; in N. rothi the second lateral teeth are broad with convex edges. No detailed observations on the habitat of either species are available to us. We know from collection data on museum specimens that N. rothi occurs intertidally. Specimens are relative- ly free of encrustations except for some coralline algae and spirorbid worm tubes. The edges of PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 12 the apertures are smooth and oval, not molded to fit a habitual site of attachment, suggesting that the normal habitat is likely to be on the undersides of stones in tidepools. Station data for the holotype of N. immaculata indicate a depth of 0 to 3 m. The absence of specimens in the intertidal collections of J. DeRoy suggests that N. immaculata is essentially a subtidal species. The elongated teeth of N. rothi are similar to those of such temperate species as Collisella pelta (Rathke, 1833), Notoacmea persona (Rathke, 1833), and Lottia gigantea. All have pointed cusps on the first and second laterals and reduced third laterals. These temperate species are known to feed upon sessile diatoms and noncalcareous algae in the middle and high intertidal zones, so we infer that N. rothi does also. The short blunt teeth of N. immaculata are similar to those of species known to feed on cor- alline algae. The presumed subtidal occurrence of N. immaculata is in accordance with the abundant subtidal occurrence of coralline algae. Lottia mimica new species (Figures 17-19, 27, 33) Shell (Figs. 17-19): Size medium (maximum length 25 mm), height medium; apex anterior to center; all slopes convex. Aperture ovoid, lat- eral edges somewhat parallel. Sculpture of raised angular ribs with one or two secondary ribs between each pair of primary ribs; ribs ex- tending slightly, crenulating the margin. Exte- rior gray-brown with white radial markings that may or may not correspond to ribs. Apex white, with fine, dark radial lines typically concentrat- ed in rays. Interior margin dark, with white markings corresponding to exterior color pat- tern; intermediate area blue-white; central area stained with dark brown; apical region white; exterior markings visible through shell. Radula (Figs. 27, 33): First pair of lateral teeth closely set at anterior edge of ribbon segment, medial edges convex, lateral edges straight, cusps pointed. Second pair of lateral teeth po- sitioned posterior to and slightly lateral to first pair, medial edges convex, lateral edges slightly convex, cusps pointed. Third lateral teeth lateral to second pair, medial edges strongly convex, lateral edges concave, cusps pointed. All lateral teeth approximately equal in width. Marginal LINDBERG & McLEAN: TROPICAL EASTERN PACIFIC ACMAEIDAE 333 FiGures 17-22. 22 New species of Lottia from the Galapagos Islands. All from Academy Bay, Isla Santa Cruz. FiGure 17. Lottia mimica new species. Holotype, LACM 1925. Length 16.2 mm. FiGure 18. Lottia mimica. Paratype, LACM 1926. Length 14.8 mm. FiGure 19. Lottia mimica. Paratype, LACM 1926. Length 9.6 mm. FiGureE 20. Lottia smithi new species. Holotype, LACM 1927. Length 12.4 mm. FiGure 21. Lottia smithi. Paratype, LACM 1928. Length 12.9 mm. FiGure 22. Lottia smithi. Paratype, LACM 1928. Length 7.5 mm. teeth small, narrow, extending over ventral plates in vicinity of third pair of lateral teeth. First lateral plates square, slightly overlapping anterior ribbon segments; second lateral plates rounded, separated from third lateral plates by partial suture; third lateral plates irregular, with prominent lateral extensions extending to edges of ventral plates. Ventral plates closely set, with broad, rounded anterior process, lateral edges concave, posterior process weak. Animal: Base of every second or third mantle tentacle with dark red-brown pigmentation; 334 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 12 snout with oral lappets; secondary gill complete but reduced in front of head, composed of lap- pets (approximately 14 per mm); every other lappet reduced, less than one-half the size of the larger ones. HoLotyPeE DIMENSIONS.—Length 16.2, width 12.4, height 4.2 mm. Type-LocaLiry.—Ecuador: Galapagos Is- lands; Isla Santa Cruz (0°38’S, 90°23’W), Acad- emy Bay, Coamano Island, intertidal zone. Leg. J. DeRoy, Oct. 1967. TypPE-MATERIAL.—Holotype, LACM 1925 (shell and radula slide), 17 paratypes, LACM 1926, paratypes also deposited in the collections of CAS and USNM. Paratypes collected at sev- eral stations in Academy Bay by J. DeRoy be- tween 1967 and 1969. DISTRIBUTION.—Galapagos Islands; Isla Fer- nandina (AMNH 163363), Isla Isabela (AHF 74- 33), Isla Bartolomé (AMNH 163290), Isla Santa Fé (AMNH 163362), Isla Espanola (AHF 359- 35), and Isla San Cristobal (CAS 23103). MATERIAL EXAMINED.—33 lots, 198 speci- mens, 23 radula preparations. EtTyMOLOGY.—The name mimica is a Latin adjective, imitative, indicative of the difficulty of distinguishing the two members of the Lottia pair by their external appearance. Lottia smithi new species (Figures 20-22, 28, 34) Shell (Figs. 20-22): Size medium (maximum length 25 mm), height medium; apex positioned in anterior third of shell; all slopes convex. Ap- erture ovoid. Sculpture of rounded primary ribs, secondary ribs of equal strength but beginning below apex. Primary and secondary ribs white, interspaces brown; apex white, with fine dark radial lines gathered into rays; interior margin dull yellow with irregular brown markings that correspond to exterior interspaces; intermediate area and central areas white; interior of myo- stracum bordered by yellow-brown halo. Exte- rior markings visible through shell. Radula (Figs. 28, 34): First pair of lateral teeth closely set at anterior edge of ribbon segment, both edges convex, rounding to blunt cusps; second pair of lateral teeth posterior and slightly lateral to first pair, both edges convex, rounding to blunt cusps. Third pair of lateral teeth posi- tioned lateral and posterior to second pair, both edges convex, rounding to blunt cusps. Marginal teeth small, narrow, overlapping ventral plates just anterior of third pair of lateral teeth. First lateral plates ovoid, slightly overlapping anterior ribbon segment; second lateral plates distinctly smaller than other lateral plates, medial edges rounded, separated from third lateral plates by a partial suture. Posterior edge of third lateral plates concave, with lateral extensions termi- nating in strongly hooked edges. Ventral plates closely set with strong posterior process; ante- rior process also present. Lateral edges in vicin- ity of marginal teeth concave; anterior sutures parallel with anterior edges of ventral plates. Animal: Mantle tentacle pigmentation some- times present; snout with oral lappets; second- ary gill complete, but reduced in front of head, composed of lappets (approximately 11 per mm). HoLotyPe DIMENSIONS.—Length 12.4 mm, width 9.7 mm, height 4.5 mm. Type-LocaLity.—Ecuador: Galapagos Is- lands; Isla Santa Cruz (0°38'S, 90°23’W), Acad- emy Bay, Punta Nunez, intertidal zone. Leg. J. DeRoy, 13° Octs1969: Type-MATERIAL.—Holotype, LACM 1927 (shell and radula slide), 9 paratypes, LACM 1928; paratypes also deposited in the collections of CAS and USNM. All type-material from Isla Santa Cruz, Academy Bay. DISTRIBUTION.—Galapagos Islands; Isla Fer- nandina (LACM 72-196), Isla Isabela (CAS 27221), Isla Bartolomé (AMNH_ 163290), Isla Santa Cruz (LACM 28839), Isla Santa Maria (ANSP. 153370), Isla San Cristobal (ANSP 153328). MATERIAL EXAMINED.—15 lots, 70 speci- mens, 14 radula preparations. ETyMoLoGy.—We are pleased to name this species in honor of the late Allyn G. Smith of the California Academy of Sciences in recogni- tion of his work with eastern Pacific mollusks, including those of the Galapagos Islands. Discuss1on.—The radular difference that separates L. mimica and L. smithi is readily ap- parent. In L. mimica the lateral teeth are point- ed distally; in L. smithi they are rounded. The third lateral teeth of L. mimica are of the same width as the second; in L. smithi the third lateral teeth are much broader than the second. Lottia mimica lacks the strong posterior process on the ventral plates that is present in L. smithi. In addition to radular difference, there is LINDBERG & McLEAN: TROPICAL EASTERN PACIFIC ACMAEIDAE 335 FiGurReEs 23-28. a 28 ha? Vag! ab Radular dentition. Figure 23. Notoacmea ubiquita new species. FIGURE 24. Notoacmea pumila new species. FIGURE 25. Notoacmea rothi new species. FIGURE 26. Notoacmea immaculata new species. FIGURE 27. Lottia mimica new species. FIGURE 28. Lottia smithi new species. Anterior towards top of page. another significant anatomical difference: in L. mimica the mantle tentacles are darkly pig- mented (Fig. 3) in much the same way as in L. gigantea (Fig. 1). This pigmentation is weakly developed or entirely lacking in L. smithi. This distinction could prove useful in future field studies because it provides a reliable, nonfatal method of species determination. Shells of L. mimica and L. smithi are essen- tially indistinguishable and are characterized by moderately strong radial ribs, variable in the number reaching the margin. Some specimens of both species have relatively few primary and secondary ribs, and these ribs project, crenulat- ing the margin (Figs. 18, 21). In others the sec- ondary ribs are more numerous and the primary ribs less prominent. In these specimens the ribs project only slightly and the shell margin is rel- atively even (Figs. 17, 19, 20, 22). The normal pattern on the apical region of juvenile shells is identical in both species (Figs. 19-22). The api- cal tip is dark colored with a pattern of thin, dark lines, concentrated in six bundles in the 1, 3,5, 7,9, and 11 o’clock positions. In most spec- imens this pattern changes abruptly to one in which rib surfaces are lighter colored and the interspaces darker, often showing concentric variations in intensity. In a tew specimens of both species (Figs. 19, 22), the juvenile pattern changes to a solid 6-rayed pattern in the adult. The holotype of L. mimica is unusual; in the early stage it is uniformly dark, changing abrupt- ly to a rayed pattern in which the lighter rays do not necessarily correspond to the ribs. Four specimens of the type lot of L. mimica, includ- ing the holotype, are predominantly dark col- ored; none in the type lot of L. smithi may be so described. The range of variability of L. mim- ica is therefore somewhat broader than that of L. smithi. The peculiar markings in the juvenile shell of L. mimica—smithi as well as its particular adult pattern are unlike those of any other acmaeid species. The extreme specimens with few ribs are similar to Collisella pediculus, although that species has fewer, more prominent ribs. Colli- 336 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 12 On 29 Gh 31 Lac = 33 FiGuRES 29-34. 30 Ge 32 Go: = 34 Radular basal plates. FIGURE 29. Notoacmea ubiquita new species. FiGURE 30. Notoacmea pumila new species. FIGURE 31. Notoacmea rothi new species. FiGURE 32. Notoacmea immaculata new species. FIGURE 33. Lottia mimica new species. FIGURE 34. Lottia smithi new species. Anterior towards top of page. sella mitella has a greater number of ribs and has a more uniform color pattern, with lighter ribs and darker interspaces. There is no particular similarity to the large, dark brown shells of Lottia gigantea or the blue- green shells of L. mesoleuca and L. stipulata, but that is not surprising in view of the lack of consistent shell characters within all acmaeid genera. We have no information about the habitat of either L. mimica or L. smithi. The original col- lecting information supplied by J. DeRoy indi- cates that they were collected on rocks exposed to heavy surf. All of the shells in both type lots were heavily encrusted with coralline algae, as are those of the Panamic species of Collisella that live under exposed surf conditions, such as C. pediculus. Limpets in this habitat have an LINDBERG & McLEAN: TROPICAL EASTERN PACIFIC ACMAEIDAE 337 habitual site of attachment with the shell edge molded to fit the home site. The margins of L. mimica and L. smithi are sufficiently irregular to suggest that they conform to specific sites. The short, blunt, equal-sized radular teeth of L. smithi are well adapted for feeding on cor- alline algae. The shape of the lateral teeth, par- ticularly the expansion of the third laterals, is similar to that found in other coralline-feeding Collisella of the tropical eastern Pacific. The lat- eral teeth of L. mimica, however, are not those of a coralline feeder, nor are they like those of most of the diatom-feeding species of Notoac- mea or Collisella, in which the third lateral teeth are reduced. The medium length teeth with pointed cusps, all equally large, are like those of many of the Peruvian Scurria and the Cali- fornian Notoacmea insessa (Hinds, 1842). The last species is known to feed on the stipes of brown algae. The teeth of L. mimica are most likely adapted for feeding on some of the fleshy, encrusting, but noncalcareous algae. Limpets of the L. mimica—smithi type are present in late Pleistocene deposits on Isla San Salvador (Hertlein and Strong 1939). The range of variation in the fossil specimens is similar to that seen in Recent specimens of both L. mimica and L. smithi. However, specific identifications are not possible from the shells alone. ACMAEIDAE OF THE GALAPAGOS ISLANDS The Galapagos Islands, or Archipiélago de Colon, are located approximately 800 km west of Cabo San Lorenzo, Ecuador. The fifteen is- lands and numerous islets extend from 1°40'N to 1°36’S and from 89°17'W to 90°01'W. The is- lands have been the subject of numerous sci- entific explorations (see Slevin 1959) and are re- - nowned for their unique fauna and flora. The marine molluscan fauna comprises tropical east- ern Pacific elements, endemics, and a few forms from the Indo-Pacific faunal region (Emerson 1978). Some nine different taxa of acmaeid limpets have been reported from the Galapagos Islands since 1855 (Reeve 1855; Carpenter 1864; Wim- mer 1880; Stearns 1893; Pilsbry and Vanatta 1902; Dall 1909; Schwengel 1938; Hertlein and Strong 1955; Keen 1958; McLean 1971). None of the species reported have been recognized as endemics, having been considered instead as vagrants from the Californian, Panamic, and Pe- ruvian molluscan faunal provinces. Many of the records have been based on beachworn shells or small shells dredged dead and in poor condition. Such specimens are difficult to refer to known species and could hardly be recognized as dis- tinct new species. After examining a number of museum collections, we are able to confirm the presence of only two previously reported species at the Galapagos Islands, Notoacmea filosa and Lottia mesoleuca. Specimens of L. mesoleuca from the Gala- pagos were originally misidentified as the Aus- tralian species Patelloida striata Quoy and Gai- mard, 1834, and were so treated by Reeve (1855:pl. 33, fig. 99) (as *‘Patella striata’’), Car- penter (1864) (as “‘Acmaea striata Reeve’’), and Stearns (1893) (as ‘‘Acmaea striata Reeve’’). Occurrences of L. mesoleuca at the Galapagos are rare. It is represented in the collections ex- amined by three specimens: two from Isla Genovesa (AHF 782-38) and a single beach specimen from the *‘Galapagos’’ (ANSP 39189). Isla Genovesa is the most northeastern of the islands, and it is conceivable that the species may be established there and not elsewhere in the archipelago. Notoacmea filosa is also rare in collections. It is represented by only two museum lots, one from Isla Santa Cruz (AMNH 177320) and one from Isla San Cristobal (LACM 54775) (Fig. 16) collected in 1929. Further information about the occurrence of these two species at the Galapa- gos Islands is desirable. Considering that recent collecting efforts have not produced these species, we only provisionally list them in the Galapagos Islands fauna. We consider all other records of Acmaeidae from the Galapagos Islands to be misidentifica- tions of the four new species described herein. Because separation of members of the species pairs is based on radular characters, it is not possible to list species-specific synonymies for each member. In both species pairs, additional characters segregate with the radular morphotypes. In the Notoacmea siblings, there are differences in shell size and coloration associated with the rad- ular types. In the Lottia pair, there are no sig- nificant differences in shell size, sculpture, or coloration; however, there are differences in mantle pigmentation and secondary gill mor- phology. We interpret these separate character 338 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 12 states as evidence against a simple radular poly- morphism and instead recognize the pairs as separate species. There is also evidence that the habitat differs in the Notoacmea pair but not in the Lottia pair. Whether speciation of the two pairs of sibling species was sympatric or allopatric is unknown. However, we hope that future workers will pur- sue this problem in the field and the laboratory. Electrophoretic analysis could prove produc- tive. ACKNOWLEDGMENTS We are indebted to Jacqueline DeRoy of Isla Santa Cruz, Galapagos Islands, who furnished preserved specimens of the new species from those islands. Donald R. Shasky of Redlands, California, loaned pertinent specimens from his collection. Courtesies were extended to Lind- berg on his museum visits by George M. Davis (ANSP), William K. Emerson (AMNH), Welton L. Lee (CAS), Joseph Rosewater and Kathy Lamb (USNM), Barry Roth (CAS), and Ruth Turner (MCZ). Barry Roth also made available the late Leo G. Hertlein’s unpublished notes on the Galapagos molluscan fauna. Bertram C. Draper, Los Angeles, made the prints of shell specimens, and Sally Walker, University of Cal- ifornia, Santa Cruz, prepared the line drawing. We thank Eugene V. Coan, Myra Keen, and Barry Roth for reading the manuscript and of- fering helpful suggestions. RESUMEN La definicion de género esta basada en las ca- racteristicas conservativas de la estructura de la concha y también en las caracteristicas cualita- tivas de la radula. La Lofttia, que anteriormente se habia considerado monotipica, se extende a incluir las especies panamicas con la agalla se- cundaria (un cordon branquial), que anterior- mente se habia atribuido a la Scurria. La Scur- ria tiene la agalla semejante, pero la estructura de la concha se diferencia. Las nuevas especies Notoacmea ubiquita de México y N. pumila de Ecuador tienen las conchas pequenas y son es- pecies alopatricas con los dientes radulares que estan modificados para alimentarse de la alga coralina. Dos nuevas especies de Notoacmea (N. rothi y N. immaculata), que son endémicas a las Islas Galapagos, constituyen un par de es- pecies que se diferencian principalmente por los rasgos de la radula: los dientes radulares de N. immaculata estan adoptados para alimentarse de la alga calcarea; los dientes radulares de N. rothi, para alimentarse de la alga no calcarea. Un par de nuevas especies de Lottia endémicas a las Islas Galapagos (L. mimica y L. smithi) también se diferencian principalmente por las caracteristicas radulares. La Lottia mimica se alimenta de la alga y es no calcarea; y la L. smithi se alimenta de alga y es calcarea. Estas cuatro es- pecies endémicas son las principales lapas ac- maeidas de las Islas Galapagos. Dos especies de la tierra firme, Notoacmea filosa y Lottia me- soleuca, se han observado no mas esporadica- mente en las Islas Galapagos. LITERATURE CITED CARPENTER, P. P. 1864. Supplementary report on the present state of our knowledge with regard to the Mollusca of the west coast of North America. Rep. Brit. Assoc. Adv. Sci. 1864:517-686. CHRISTIAENS, J. 1975. Revision provisoire des mollusques marins récents de la famille des Acmaeidae. Inf. Soc. Belge Malacol. 4(4):3-20. Dati, W. H. 1871. On the limpets; with special reference to the species of the west coast of America, and to a more natural classification of the group. Am. J. Conchol. 6(3):227-282. . 1909. Report on a collection of shells from Peru, with a summary of the littoral marine Mollusca of the Peruvian zoological province. Proc. U.S. Natl. Mus. 37:147-294. Emerson, W. K. 1978. Mollusks with Indo-Pacific faunal affinities in the eastern Pacific Ocean. Nautilus 92:91—96. HERTLEIN, L. G., AND A. M. StrRoNG. 1939. Marine Pleis- tocene mollusks from the Galapagos Islands. Proc. Calif. Acad. Sci., ser. 4, 23:367—380. , AND . 1955. Marine mollusks collected at the Galapagos Islands during the voyages of the Velero III, 1931-1932. Pp. 111-115 in Essays in the Natural Sciences in Honor of Capt. Allan Hancock, Univ. So. Calif., Los Angeles, Calif. KEEN, A. M. 1958. Sea shells of tropical west America. Ist ed. Stanford Univ., Stanford, Calif. 624 pp. LINDBERG, D. R. 1976. Cenozoic phylogeny and zoogeog- raphy of the Acmaeidae in the eastern Pacific. Ann. Rep. West. Soc. Malacol. 9:15—16. 1978. On the taxonomic affinities of Collisella ed- mitchelli, a late Pleistocene limpet from San Nicolas Island, California. Bull. So. Calif. Acad. Sci. 77:65-70. . 1979. Variation in the limpet Collisella ochracea and the northeastern Pacific distribution of Notoacmea testu- dinalis (Acmaeidae). Nautilus 93:50—56. MacCLintock, C. 1967. Shell structure of patelloid and bel- lerophontoid gastropods (Mollusca). Peabody Mus. Nat. Hist. Yale Univ. Bull. 22:1—140. McLean, J. H. 1966. West American prosobranch Gastro- poda: Superfamilies Patellacea, Pleurtomariacea, and Fissu- rellacea. Ph.D. dissertation, Stanford Univ., Stanford, Cal- if. 255 pp. LINDBERG & McLEAN: TROPICAL EASTERN PACIFIC ACMAEIDAE 339 . 1971. Family Acmaeidae, in Keen, A. M., Sea shells of tropical west America. 2nd ed. Stanford Univ. Press, Stanford, Calif. 1064 pp. . 1973. Family Acmaeidae, in Marincovich, L., Inter- tidal marine mollusks of Iquique, Chile. Nat. Hist. Mus. Los Angeles Co. Sci. Bull. 16:1—49. Pitssry, H. A., AND E. G. VANATTA. 1902. Papers from the Hopkins Stanford Galapagos Expedition, 1898-99, no. 13, marine Mollusca. Proc. Washington Acad. Sci. 4:549-560. REEVE, L. 1855. Conchologia iconica: or, illustrations of the shells of molluscous animals. Vol. 8. Monograph of the ge- nus Patella. London. Pages not numbered, 41 pls. SCHWENGEL, J. S. 1938. Zoological results of the George Vanderbilt South Pacific Expedition, 1937. Part I. Galapa- gos Mollusca. Proc. Acad. Nat. Sci. Philadelphia 90: 1-3 SLEVIN, J. R. 1959. The Galapagos Islands: A history of their exploration. Occas. Pap. Calif. Acad. Sci. 25:1—150. STEARNS, R. E. C. 1893. Report on the mollusk fauna of the Galapagos Islands with descriptions of new species. Proc. U.S. Natl. Mus. 16:353—450. Wimmer, A. 1880. Zur Conchylien-Fauna der Galapagos-In- seln. Sitzungsber. K. Akad. Wiss. Wien Math.-Natwiss. KI. 80(Abt. 1)(10)(1879):465—5 14. CALIFORNIA ACADEMY OF SCIENCES Golden Gate Park San Francisco, California 94118 oe) = : = a oo - ~~ a 7 . se _ he oe ee an ie | | ; _ — —_ F » ane : = a - we $- : by - ; 7 r 7 . wy é ee PTE BATH), F011 D)». . - a ioe 10665 s. rid ; . t r ‘ \ a a, = Panui! 4 Dan hi, hie ( “ieee ms Seid y+) 7 Gareth ey = et a> = au PROCEEDINGS Vol. 42, No. 13, pp. 341-348, 5 figs. A REVIEW OF THE BATHYAL FISH GENUS ANTIMORA (MORIDAE: GADIFORMES) By Gregg J. Small* Systematics Laboratory, National Marine Fisheries Service, NOAA, National Museum of Natural History, Washington, DC 20560 ABSTRACT: There are two valid species of the genus Antimora: A. microlepis Bean in the North Pacific, and A. rostrata (Gunther) from the southeastern Pacific, Southern Ocean, and Atlantic Ocean. Junior synonyms of A. rostrata include: A. australis Barnard, A. meadi Pequeno, A. rhina Garman, and Haloporphyrus viola Goode and Bean. Antimora microlepis has 90 to 103 gill filaments on the first arch; A. rostrata has 76 to 90. Differences in the regression equations of gill filament length on standard length, and of head length on standard length between fish from several geographic areas are shown. Other characters examined include numbers of vertebrae, fin rays, gill rakers, and scale rows; and morphometric ratios, using lengths of eye, snout, predorsal, first dorsal fin ray, maxillary, and gill rakers, and width of interorbital. INTRODUCTION Fishes of the benthopelagic morid genus An- timora are widely distributed in the world oceans, ranging in depth between 402 and 2905 m (Grey 1956), and are very abundant in some regions (Wenner and Musick 1977). They appear to be most common on the continental slopes of subarctic, subantarctic, and temperate regions, but are generally rare in the subtropics and trop- ics, although they are apparently common in the vicinity of the Hawaiian and Galapagos islands. The genus Antimora is distinguished from other members of the family Moridae by the combination of a pronounced pointed snout, a pelvic fin with six rays, a well-developed mental barbel, a long-based second dorsal fin with more than SO rays, and a deep indentation in the out- * Present address: Northwest and Alaska Fisheries Center, National Marine Fisheries Service, NOAA, 2725 Montlake Boulevard East, Seattle, Washington 98112. line of the anal fin (Svetovidov 1948). There is little information on food habits because speci- mens brought to the surface routinely evert their stomachs. Sedberry and Musick (1978) found only 10 specimens with intact stomachs in nu- merous deep-water trawlings. Individuals with ripe eggs are unknown, and specimens smaller than 100 mm are rare. Males longer than 325 mm are uncommon, although females are often long- er than 600 mm. Within its depth range, there is a segregation by sex and also by size of individ- uals (Iwamoto 1975; Wenner and Musick 1977). Available information on the taxonomy and dis- tribution of the genus was summarized by [wa- moto (1975). Although this study shows that only Antimora rostrata, occurring in all areas except the North Pacific, and A. microlepis, occurring only in the North Pacific, are valid species of the genus Antimora, the following species and their as- sociated localities have been previously pro- posed: Haloporphyrus rostrata Gunther, 1878, [341] 342 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 13 CIFIC: | lepericen el NORTH ATLANTIC i | | + tt ii | i: nr += ———— toed t SOUTHEAST PACIFIC SOUTHERN OCEAN | | | SOUTHERN OCEAN + — +4- —— = leo ~® FiGure |. Heavy black lines divide regions. from subantarctic seas; Haloporphyrus viola Goode and Bean, 1878, from the temperate western North Atlantic; Antimora microlepis Bean, 1890, from the temperate eastern North Pacific; Antimora rhina Garman, 1899, from the Gulf of Panama; Antimora australis Barnard, 1925, from off the Cape of Good Hope; and An- timora meadi Pequeno, 1970, from off the coast of Chile. No comparison has ever been made of popu- lations represented by these six names. The closest approach was that of Schroeder (1940) who examined limited material of the first four nominal species (he apparently overlooked A. australis) and suggested that they be referred to the same species. The object of the present pa- per is to resolve the number of valid species of Antimora using measurements, counts, and the known species distributions. Biological and distributional information are useful in clarifying the taxonomy of Antimora. It has been shown that in the western North Atlantic Antimora becomes larger with increas- ing depth but does not reproduce within the area World localities from which specimens of Antimora spp. were studied. Stars = A. microlepis, dots = A. rostrata. of its greatest known abundance (Wenner and Musick 1977). Migrations may be a regular part of the life history of the western North Atlantic population. The mobility of Antimora has been confirmed by Cohen (1977) who has shown that these fish are able swimmers. Perhaps there is a single interbreeding North Atlantic population. If Antimora from other regions have the same swimming ability, then there may be single pop- ulations in the southeastern Pacific, the southern oceans, and the northern Pacific. Although data for this study were originally segregated on the basis of six geographical regions (see Materials), characters of Antimora in the western and east- ern North Pacific were found to be similar as were those from the western and eastern North Atlantic. A preliminary analysis indicated that the four geographical groupings mentioned above provide a more appropriate basis for com- parison (Fig. 1). METHODS Measurements and counts were made accord- ing to Hubbs and Lagler (1970) and include: SMALL: FISH GENUS ANTIMORA standard length, head length, snout length, pre- dorsal length, eye diameter, upper jaw length, interorbital distance, first dorsal fin ray length, number of scales along the lateral line, and num- ber of scale rows between lateral line and dorsal origin. Numbers of vertebrae (including the hy- pural plate), anal rays, and dorsal rays were read from radiographs. Numbers of gill filaments and gill rakers on the first gill arch, lengths of the longest gill raker and gill filament (measured from base to tip, Fig. 2), length of longest gill raker at the gill angle, and dorsal fin ray length (from the anterior base to the tip of the first ray) were compared. Bro- ken and otherwise damaged rays were not mea- sured. Coloration, often used in early descriptions, was not recorded due to color changes which occur in preserved specimens. However, an at- tempt was made to recheck other characters presented in original descriptions. MATERIALS A total of 449 specimens were examined (refer to figures for length summary). WESTERN NortTH PaciFic (between 33°N to 48°N and 135°E to 145°E; 6 specimens): U.S. National Museum (USNM): 161494 (1); 160607 (1); 160606 (1); 149228 (1); 117886 (2). EASTERN NortH PacliFic (between 18°N and 56°N; 96 spec- imens): California Academy of Sciences (CAS): 3883 (2); 37559 (1); 34354 (2); 34353 (2); 32308 (2); 27525 (1); 26226 (1); uncat. (3); CAS-SU 5276 (5); 77 (1). Museum of Comparative Zoology (MCZ): 28250 (1). Scripps Institution of Oceanogra- phy (SIO): 70-249 (22); 70-247 (2); 68-443 (1); 59-265 (1). Uni- versity of Washington (UW): 19309 (2); 19235 (1); 19228 (7); 19139 (4); 18492 (8); 18201 (2) 18190 (2); 17180 (7); 19149 (1); 18493 (5). USNM: 45361 (2 syntypes of Antimora microlepis),; 54573 (1); 54364 (1); 53876 (3); 48562 (1); 47238 (1); 47237 (1). WESTERN NorTH ATLANTIC (between 27°N and 59°N; 117 specimens): Institut fiir Seefischerei, Hamburg (ISH): 79/73 (2). MCZ: 53949 (1); 38282 (3); 38073 (3); 37633 (1); 37619 (2); 37595 (1); 37520 (1); 37585 (2). University of Maine, Darling Center (UMDC): 313-1 (1); 306-2 (1). USNM: 21837-8 (2 syn- types of Haloporphyrus viola); uncat. (35); 31725 (1). x-ray counts only: 143250 (1); 45872 (1); 45845 (1); 45808 (1); 36163 (1); 38142 (1); 38068 (1); 38064 (3); 38019 (1); 35595 (1); 35566 (1); 33446 (4); 33443 (7); 33340 (3); 33337 (5); 33014 (1); 31768 (1); 28612 (1); 28611 (1); 28610 (1); 28609 (1); 28608 (1); 24746 (1). Virginia Institute of Marine Sciences (VIMS): 3458 (2); 3243 (1); 1471 (7); 1460 (3); 872 (4); 870 (1); uncat. (3). EASTERN NorTH ATLANTIC (between 50°N and 60°N; 6 specimens): ISH: 112/74 (1); 111/74 (1); 146/74 (2); 745/74 (2). SOUTHEASTERN PAaciFIc (between 0°S to 56°S and 70°W to 91°W; 123 specimens): USNM: uncat. (120). MCZ: 28610 (1) and 28611 (2) (3 syntypes of Antimora rhina). SOUTHERN OCEANS (specimens from southern Atlantic, In- dian, and Pacific oceans, excluding southeastern Pacific spec- imens; 101 specimens): ISH: 2191/68 (2); 1250/66 (1); 1241/66 (2); 1142/66 (11); 1129/66 (1); 361/71 (2); 344/71 (1); 286/71 (1); 343 FiGure 2. Medial view of first gill arch of left side to il- lustrate location of gill filament measurement. (A) Antimora microlepis, UW 17180, off Columbia River, 46°N, 124°W, 310 mm SL; (B) Antimora rostrata, LACM 10985-5, southwest of New Zealand, 56°19’S, 158°29’E, 330 mm SL. Drawn by Kei- ko Hiratsuka Moore. 152/67 (8); 151/67 (3); 150/67 (1); 33/76 (5); WH32/76 (5). Los Angeles County Museum (LACM): 10033 (12); 10032 (8); 10985-5 (3); uncat. (9). University of Florida (UF-TABL): 503 (1). UMDC: uncat. station numbers 01343 (1); 01342 (1); 00198 (1); 00165 (1); 00157 (1); 00152 (1). USNM: 188827 (1 syntype of Antimora australis); 188829 (3); 188823 (5); 188822 (2); un- cat. (3). British Museum Natural History (BMNH): 1887.12.- 17.36 (holotype of Haloporphyrus rostratus, x-ray only). RESULTS Antimora may be divided into two distinct species instead of the six described, based on the number of gill filaments and secondarily on the ratio of gill filament length to standard length. Other measurements can be used but with less distinct separation of species. Counts.—North Pacific specimens possess 90-103 gill filaments on the first gill arch, com- pared with 76-90 in specimens from the other 2 ) Z al vt S > na ss O Z eB 3) N f, ie) > = ia a < Ss) < S Z a2 fe) Be s < ) w Be = rs fe) Ze) S) Z a a a S) fe) om a 344 I I I v Vv + 10r hu I t c tC OOb-ISE I IT Cc 4 I I I REE |i a4 we OS CE WG SE ve Ot € € OSt-l0€ |! ke! Il v I (i qu We EEE NE VO GC € Gane CC Sec (G = Az OO0E-1ST v I I | II I € (GG As (GAY ve t OSc—-10C vy OUP [PU I I v As 6G WG (he {s 4 Creer. OOc-ISI v v v 4 I I I I CCG (306 v by C oo OSI—O01 901 SOL Ol €0I ZOI IOI O01 66 86 46 96 S6 6 £6 7 16 06 68 88 48 98 S8 8 £8 78 18 08 6L 8L LL YL (ww) TS susWe]Y [Is Jo loquny ‘(p) SNVAJIO NYFHLNOS GNV “(€) OIdIOVg LSVWAHLNOS§ *(Z) OILNVILY HLYON AHL WOds DIDAISO4 DAOUNUY GNV “([) OISIOVG HLYON WOU sidajosonu DAOWIUY YOA dNOUH HLONAT GUVGNVIG Ad YaaWON LNAWW II THY AO AONANOAY AHL JO NOSIMYVdWOD ‘[ ATAVE SMALL: FISH GENUS ANTIMORA Gill filament length (mm) 100 200 300 345 400 500 Standard length (mm) FIGURE 3. Regression of gill filament length on standard length in Antimora from four divisions of the world oceans: North Pacific, squares; southeast Pacific, Southern Ocean, and North Atlantic, dots. North Pacific: y = 0.03x — 0.10, n = 51. All other oceans (areas): y = 6.02x — 0.70, n = 230. regions (Table 1). Number of gill filaments ap- pears to be constant over the size range of spec- imens in all populations. Only slight differences between samples were found from the four geographical areas in total vertebral number, number of gill rakers on the first arch, and anal and dorsal fin rays (Table 2). Although the size and number of scales have been used in several of the species descriptions previously mentioned, they are not useful char- acters for the differentiation of these species. TABLE 2. Specimens of Antimora are very fragile and on capture the scales and scale pockets do not re- main intact for use as a reliable character. MEASUREMENTS.—Specimens collected from the eastern and western North Pacific north of latitude 10°N have a gill filament length relative- ly greater than that in fish caught elsewhere. Above the size range of approximately 150 mm standard length, the length of the filaments dis- tinctly separate North Pacific fishes from all oth- er groups. Least square regression lines were fit SUMMARY OF SELECTED COUNTS AND LENGTH PROPORTIONS IN Antimora FROM THE FOUR GEOGRAPHICAL REGIONS SHOWN IN FiGurReE | (lengths presented as ratio of standard length to size of part). N Pacific SE Pacific N Atlantic S Ocean Character x n SD G n SD X n SD x n SD Snout length ile) 7A 0.86 11.8 105 1.33 125] ee Cee LG 2G oS 153 Predorsal length 3.9) 169 1.47 3357 108 0.20 Bh) aki st ly/ 3.9 94 0.20 Maxillary length eu 70 0.36 6.9 110 = 0.50 UP They BIS 7.4 94 0.49 First dorsal fin ray length S29 47 1.43 Tail 94 1.58 55! 63 1.45 6.1 54 1.40 Eye diameter 15.0 66 1.20 1533 96 134 6{ ON GOs ull 42 16.2.- 49 1.36 Interorbital width 17.6 38 1.45 18.6 71 1.61 1525) 282s) HIRS6 18.3 32 1.64 Longest gill raker length 73.4 23 14.07 = 0 - 76.9 1 = 103.0 11 16.00 Total number of vertebrae 591 47 0.86 58.8 76 0.96 59.8 64 1.26 59.6 31 1.02 Total number of gill rakers 16:5) 38 1.93 16.2 74 1.29 166 46 e250 16.0 79 1.54 Total number of anal fin rays 40.6 41 1.40 39.3 56 1.39 41.9 67 1.56 40.0 28 1.70 Total number of dorsal finrays 52.4 40 1.15 Sl 59 Vey Seep 7 GY 1.45 Sou 32 1.45 346 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 13 e 110 e eee b ee e e oO 108 North Pacific oT o e e = 90@® oo e E 90 o e ee = oo op a ie og ee e e & aleao c goof e 5 80 00 op’oo ee e ee All other oceans (areas) 12) og ee ee @ e a oo ee a6 Oo ot e 70 oqgp oe e e oop e e ce ee eee on @ @ ee a e ee BE o,eee “e 0 Aee ee Oo e e e@ ee SB e e e epee o @ e 40 e ee ee 30 fer eee Bs ee ee 20 100 200 300 400 500 Standard Length (mm) FIGURE 4. Regression of head length on standard length in Antimora from four divisions of the world oceans: North Pacific, squares; North Atlantic, southeast Pacific, and Southern Ocean, dots. North Pacific: y = 0.26x + 2.22, n = 98. All other oceans (areas): y = 0.23x + 2.9432, n = 277. to the data, and the regression of gill filament length on standard length was determined fol- lowing the methods of Zar (1974) (Fig. 3). Head length against standard length (Fig. 4) also separates North Pacific specimens from all other Antimora. This difference is most marked in fish larger than 200 mm standard length. Bean (1890), Garman (1899), Barnard (1925), and Pequeno (1970) noted head length to total length or standard length proportions in original descriptions with no apparent differences among the described species. Comparisons of the length of snout, predorsal distance, maxillary, first dorsal fin ray, eye di- ameter, longest gill raker, and width of interor- bital are summarized in Table 2. None of the measurements serve unequivocally to separate the North Pacific specimens from other Anti- mora. Fish from all other areas do show some differences: southeast Pacific, North Atlantic, and Southern Ocean populations may have slightly different lengths of first dorsal fin ray; North and southeast Pacific fish appear to have larger eyes than do other Antimora; and North Atlantic specimens have larger interorbital widths. Gill raker length may also show some differences, however, data for this character are incomplete. Within the range of A. rostrata, there are local differences in some morphometric characters, but these differences are not consistent through- out the range of size or geography. For example, the first dorsal fin ray to standard length ratio is higher in North Atlantic specimens than those from other areas. This longer fin ray is most pro- nounced in specimens in the 200-400 mm range. North Atlantic fish also have a shorter snout and wider interorbital distance over certain seg- ments of their size range as compared to speci- mens from other regions. Although only one specimen from low latitudes in the mid-Atlantic was examined in this study, there may be con- tact between North Atlantic and South Atlantic populations of Antimora, as specimens have been taken in the Bahamas and the Gulf of Guinea (personal communication, Daniel M. SMALL: FISH GENUS ANTIMORA FIGURE 5. 347 (A) Antimora rostrata, USNM 218479, SL 346 mm, male, western North Atlantic, 36°39’N, 74°28’W, 1530-1610 m; (B) Antimora microlepis CAS 32308, SL 371 mm, male, off California. Drawn by Keiko Hiratsuka Moore. Cohen, Systematics Laboratory, National Ma- rine Fisheries Service, NOAA, Washington, DG;). CONCLUSIONS The species can be characterized as follows: Antimora microlepis Bean, 1890 (Figure 5B) Antimora microlepis BEAN, 1890:38 (type-locality: 51°23’N, 130°34'W, ALBATROSS Sta. 2860, off Cape St. James, Queen Charlotte Islands, 876 fms [1602 m], 13 Aug. 1888). CHARACTERS.—Gill filaments on first gill arch 90-103; gill filaments relatively long, regression equation of gill filament length on standard length y = 0.03x — 0.10; head length relatively long, regression equation of head length on stan- RANGE.—Eastern and western North Pacific Ocean, north of latitude 10°N. Antimora rostrata (Glinther, 1878) (Figure 5A) Haloporphyrus rostratus GUNTHER, 1878:18 (type-locality: ‘midway between the Cape of Good Hope and Kerguelen Island; east of the mouth of the Rio de la Plata,’ CHAL- LENGER sta. 146, 1375 fms [2515 m], and sta. 320, 600 fms [1097 m}). Haloporphyrus viola GOODE AND BEAN, 1878:257—260 (type- locality: ‘‘outer edge of Le Have Bank, at a depth of four or five hundred fathoms”’ [approximately 43°N, 64°W)). Antimora rhina GARMAN, 1899:185—186 (type-locality: Gulf of Panama, ALBATROSS sta. 3353, 7°06'15"N, 80°34’ W, 695 fms [1271 m], sta. 3393, 7°15'N, 79°36’W, 1020 fms [1865 m]). ss ot. CHaracters.—Gill filaments on first gill arch = o RANGE i - 348 Antimora australis BARNARD, 1925:499 (type-locality: ‘‘off Cape Point, 475-900 fathoms’’ [869-1646 m]). Antimora meadi PEQUENO, 1970:14—16 (type-locality: ANTON BRUUN cruise 13, between 34°06'S, 72°26’W, and 34°12’S, --»72°25'W, in 1400-1475*m, 3 Feb. 1966). filaments relatively short, regression equation of gill filament length on standard Jengthey ='0.02x — 0.70; head length relatively ny short, regression equation of head length on standard length y = 0.23x + 2.9432. Alboééans except the North Pacific N. wT o ACKNOWLEDGMENTS I am thankful for the advice and support given me by Daniel M. Cohen of the Systematics Lab- oratory, National Marine Fisheries Service (NMEBS), for initially suggesting this study, and his valuable guidance throughout the course of this work. Tomio Iwamoto (CAS) recommended that I examine gill filament length, arranged for the loan of specimens from the California Acad- emy of Sciences, and reviewed the manuscript. Bruce B. Collette (Systematists Laboratory, NMFS) and Hugh H. DeWitt (University of Maine, Orono) also reviewed the manuscript and made valuable comments. Special thanks are due Charles A. Wenner and John A. Musick for allowing me to read their then-unpublished manuscript on the life history of North Atlantic Antimora and examine spec- imens in their collection at the Virginia Institute of Marine Science. The Allan Hancock Foun- dation and Los Angeles County Museum pro- vided Antarctic material, William Fink (MCZ) provided important materials from the North Atlantic as did Alfred Post (ISH). Richard Ro- senblatt (SIO), Arthur Welander (UW), Carter Gilbert (UF), Jean Dunn (NMFS, Seattle Lab- oratory), and the National Museum of Natural History were all instrumental in providing ad- ditional materials. Keiko H. Moore prepared the illustrations and figures. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 13 LITERATURE CITED BARNARD, K.H. 1925. Descriptions of new species of marine fishes from South Africa. Ann. Mag. Nat. Hist. ser. 9, 15:498-504. BEAN, T. H. 1890. Scientific results of explorations by the U.S. Fish Commission Steamer Albatross. No. XI—New fishes collected off the coast of Alaska and adjacent region southward. Proc. U.S. Natl. Mus. 13:37-45. CoHEN, D. M. 1977. Swimming performance of the gadoid fish Antimora rostrata at 2400 meters. Deep-Sea Res. 24:275-277. GARMAN, S. 1899. Reports on an exploration off the west coasts of Mexico, Central and South America, and off the Galapagos Islands, in charge of Alexander Agassiz, by the U.S. Fish Commission Steamer ‘*Albatross** during 1891, Lieut-Commander Z. L. Tanner, U.S.N. Commanding. Part 26, The Fishes. Mem. Mus. Comp. Zool. Harvard 24:1-431. GoopeE, G. B., AND T. H. BEAN. 1878. Descriptions of two gadoid fishes, Phycis chesteri and Haloporphyrus viola from the deep-sea fauna of the northwestern Atlantic. Proc. U.S. Natl. Mus. 1:256-260. Grey, M. 1956. The distribution of fishes found below a depth of 2000 meters. Fieldiana Zool. 36:74—336. GUNTHER, A. 1878. Preliminary notices of deep sea fishes collected during the voyage of H.M.S. Challenger. Ann. Mag. Nat. Hist. ser. 5, 2:17—28. Husss, C. L., AND K. F. LAGLER. 1970. Fishes of the Great Lakes region. 4th ed. Ann Arbor: Univ. Michigan Press. 213 pp. Iwamoto, T. 1975. The abyssal fish Antimora rostrata (Gtn- ther). Comp. Biochem. Physiol. 52B:7-11. PEQUENO-R., G. 1970. Antimora meadi n.sp. en Chile. Not. Men. Mus. Nac. Hist. Nat. (Santiago) 15:14—-16. SCHROEDER, W. C. 1940. Some deep sea fishes from the North Atlantic. Copeia 1940:231-238. SEDBERRY, G. R., AND J. A. Musick. 1978. Feeding strate- gies of some demersal fishes of the continental slope and rise off the Mid-Atlantic coast of the USA. Mar. Biol. (N.Y.) 44:357-375. Svetovipov, A. N. 1948. Treskoobraznye. Faune SSSR, Ryby 9(4), 221 pp. Akad. Nauk SSSR. (Transl. from Rus- sian: 1962, Gadiformes, Fauna USSR, Fishes. 304 pp. U.S. Dep. Comm., Natl. Tech. Inf. Serv., Springfield, Virginia; OTS 63-11071.) WENNER, C. A., AND J. A. Musick. 1977. Biology of the morid fish, Antimora rostrata, in the western North Atlan- tic. J. Fish. Res. Bd. Canada 34:2362-2368. ZAR, J. H. 1974. Biostatistical analysis. Englewood Cliffs, New Jersey: Prentice Hall Inc. 620 pp. CALIFORNIA ACADEMY OF SCIENCES Golden Gate Park San Francisco, California 94118 PROCEEDING OF THE CALIFORNIA ACADEMY O Vol. 42, No. 14, pp. 349-377; 18 figs., 2 tables es eee ee Marine Bislagical Le ouralory pis ARY | | Noy 19 1981 | = fhiess October 26, 1981 THE KAIANUS-GROUP OF THE GENUS CALLIONYMUS (PISCES: CALLIONYMIDAE), WITH DESCRIPTIONS OF SIX NEW SPECIES By Ronald Fricke Saalestrasse 3 a, D-3300 Braunschweig, Federal Republic of Germany ABSTRACT: The kaianus-group of the genus Callionymus, subgenus Callionymus, is revised. The recognized species and their ranges are: Callionymus kaianus Gunther, 1880 (Kai Islands, eastern Indonesia); Callionymus moretonensis Johnson, 1971 (northern half of Australia, New Ireland, New Caledonia); Callionymus whiteheadi n.sp. (southwestern Indonesia); Callionymus guentheri n.sp. (Philippine Islands); Callionymus formosanus n.sp. (Formosa Strait, Taiwan); Callionymus sokonumeri Kamohara, 1936 (southern Japan); Callionymus altipinnis n.sp. (South China Sea: China coast); Callionymus ochiaii n.sp. (southern Japan); Callionymus regani Nakabo, 1979 (Saya de Malha Bank, western-central Indian Ocean); Callionymus kotthausi nom. nov. (for Callionymus indicus (Kotthaus, 1977), a secondary homonym; India); Callionymus africanus (Kotthaus, 1977) (east Africa); Callionymus bentuviai n.sp. (southern Red Sea); Callionymus carebares Alcock, 1890 (northern Indian Ocean). INTRODUCTION The dragonets of the family Callionymidae are a group of benthic marine fishes (except for two euryhaline species which enter and even live in freshwater rivers). About 130 species are known. The two largest genera, Callionymus and Synchiropus, are nearly circumtropical in warm and temperate seas, but some species also live in cold waters; e.g., Callionymus lyra and C. maculatus of the northern Atlantic follow the warm Gulf Stream to Iceland and northern Nor- way. Callionymids usually live on sand or mud bottoms, sometimes also on coral sand bottom in coral reefs, or among seaweed, from very shallow waters and even tide pools down to about 800 m. Callionymus is the largest genus of the family, comprising about 75 species. Fricke (1980) dis- tinguished three subgenera (Callionymus, Cal- liurichthys, and Spinicapitichthys) which differ principally in the shape of the preopercular spine. The subgenus Callionymus is the largest, comprising about 55 species which can be ar- ranged into various species groups. The kaia- nus-group is one of the larger species groups and contains deepwater mud-bottom species of the Indian and western Pacific oceans. Seven of the species included in the kaianus- group have been described. Callionymus kaia- nus Giinther, 1880, was originally described from Kai Islands, west of New Guinea, and later re- corded from India, Indonesia, Saya de Malha Bank (western-central Indian Ocean), Zanzibar, Arabian Sea, Japan, Pescadore Islands (near Taiwan), Gulf of Tonkin, and the coast of China. Johnson (1971) described a new subspecies, Callionymus kaianus moretonensis, from north- eastern Australia. Callionymus carebares Al- [349] 350 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 14 cock, 1890, was described from the Bay of Ben- gal; C. sokonumeri Kamohara, 1936, from Japan; and C. regani Nakabo, 1979, from the Saya de Malha Bank. Under the generic name Diplogrammus, Kotthaus (1977) described two further species, D. africanus and D. indicus. I found several closely allied new species, which are described and compared in the pres- ent paper, from examination of specimens (most previously identified as Callionymus kaianus) in the Australian Museum, Sydney (AMS); British Museum (Natural History), London (BMNH); California Academy of Sciences, San Francisco (CAS); Faculty of Agriculture of Kyoto Univer- sity, Kyoto (FAKU); Fish Collection, Hebrew University of Jerusalem (HUJF); Institut Royal des Sciences Naturelles, Brussels (IRSN); Ma- rine Science Laboratory, Chinese University of Hong Kong (MSL); Staatliches Naturhisto- risches Museum, Braunschweig (NMB); Zoo- logisches Institut und Zoologisches Museum der Universitat Hamburg (ZIM); and Museum ftr Naturkunde, Zoologisches Museum, East Berlin (ZMB). METHODS Methods used are the same as those in my previous papers (especially Fricke 1980; Fricke 198 1a). The preopercular spine formula, explained by Fricke (198la), is calculated by the following formula: a es, Cc where a is the number of antrorse spines at the base, b is the number of points or serrae at the dorsal edge, c is the number of points or serrae at the ventral edge, and d is | and reflects the main tip of the spine. The formula treats sim- plified /eft spines. Right spines have to be treat- ed as left (e.g., the number of antrorse spines at the base, a, is always on the left side of the formula). The pectoral fin base is divided in two by a membrane connecting it with the fifth pelvic fin ray. The formula a/b is used where a is the pec- toral fin base length above the connecting mem- brane, b is the corresponding length below. THE CALLIONYMUS KAIANUS-GROUP The Callionymus kaianus-group, including the new species described in this paper, com- prises thirteen deepwater mud-bottom species in the subgenus Callionymus (see Fricke 1980:59) distributed in the Indian and west Pa- cific oceans (Figs. | & 2): Callionymus kaianus, C. moretonensis, C. whiteheadi, C. guentheri, C. sokonumeri, C. formosanus, C. altipinnis, C. ochiaii, C. regani, C. kotthausi, C. africanus, C. bentuviai, and C. carebares. The group is characterized by the presence in its members of large eyes; dorsal and anal fin formulae D IV + vill,l, A (vil, l—)vili,l; one or two unbranched median caudal fin rays which are often filamen- tous; a characteristic shape of the preopercular spine (see Fig. 3); and (usually) a characteristic black blotch on the third membrane of the first dorsal fin. Species of the Adianus-group are similar to the deepwater mud-bottom species groups of the genus Synchiropus (e.g., phaeton-group, alti- velis-group), agreeing with them in some aspects of body shape and even color markings (black spot on third membrane of first dorsal fin, etc.). The Synchiropus species groups are easily dis- tinguished from the kaianus-group by the shape of the preopercular spine and by generic differ- ences between Callionymus and Synchiropus discussed in detail in my revision of the genus Synchiropus (Fricke 1981b). Characters and distribution of the species of the Aadianus-group are compared in Tables 1-2. Further distinguishing features (not compared in the tables) include other proportions, the pre- opercular spine shape, and body color pattern. Key to the Species of the Callionymus kaianus-Group la. Head in SL 2.7-3.1; branchial opening very broad, same size as or larger than pupil; upper edge of preopercular spine with 2 large curved points, but without a small antrorse point C. carebares Ib. Head in SL 3.4—4.6; branchial opening small, about “3-2 of pupil; upper edge of preopercular spine with 1 small an- trorse and | or 2 large curved points _ 2 2a. D, and anal fins very high, males with convex distal) margins === 33 2b. D, and anal fins relatively low, with Straight distal margins... 3a. First spine of D, filamentous 3b. First spine of D, not filamentous — 5 FRICKE: KAIANUS-GROUP OF GENUS CALLIONYMUS 4a. 4b. Sa. _@ ochiaii oO moretonensis a Kalanus v africanus FiGure I. D, relatively low, Ist ray about 1.0, Sth ray about 0.9 in head length; preoper- cular spine with 2 curved points (and 1 small antrorse point) at its upper side; black blotch on 3rd membrane of D, large, nearly covering entire membrane; anal fin with a distal black streak on each membrane; distal %4 of anal fin dark brown, tips of rays white _.-- ons al ae C. formosanus D, high, Ist ray about 1.1, 5th ray about 0.8 in head; preopercular spine with 1 large curved point (and | small antrorse point) at its upper side; black blotch on 3rd membrane of D, small, distal in po- sition; anal fin without a distal black streak on each membrane; distal margin of anal fin black, tips of fin rays also black C. altipinnis Main tip of preopercular spine long and slender; distal half of anal fin black, tips of rays white; distal margin of caudal fin regular; black blotch on 3rd membrane of D, relatively large, central in position (not reaching distal margin) ecarebares 351 Geographical distribution of species of the Aaianus-group of the genus Callionvmus. 5b. Main tip of preopercular spine short; 6a. 6b. Hae 7b. 8a. 8b. 9a. anal fin with a distal black streak on each membrane, distal 24 of membranes and tips of fin rays brown; distal margin of caudal fin irregular; black blotch on 3rd membrane of D, very small, extremely distal in position C. sokonumeri Caudal fin convex, without filaments _ 7 Caudal fin convex or slightly pointed, with 1 or 2 filaments Anal fin with a small distal yellow margin or colorless; sides of body with a row of large indistinct brownish blotches _______- BOS A We a Ree rs oe A! C. regani Anal fin with a broad dark brown or black margin; sides of body with a row of small distinct black blotches Main tip of preopercular spine long and slender; lower part of caudal fin colorless MeN 2 Rati le he anh eles C. kotthausi Main tip of preopercular spine short; lower part of caudal fin with a broad Black Sticdkes Se ee ee cee eee 9 Pectoral fin base with a large dark brown 352 9b. 10a. 10b. lla. Ib. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol.No. 14 aregani — v kotthausi m bentuvial | o sokonumer! FIGURE 2. area; back marbled with olive-green; operculum with large white spots; lower part of caudal fin black distally; D, with a basal row of dark blotches Pectoral fin base with 2 black streaks; back yellowish brown, with white blotches surrounded by semicircular dark brown bands; operculum with small black spots; lower part of caudal fin with a broad, curved black band, but color- less distally; D, without a basal row of black spots _.______ _C.. africanus (female) First spine of D, with a long filament First spine of D, without a filament — 13 Main tip of preopercular spine short, largest point on its dorsal side with a bas- al hook; D, colorless, with transverse white lines; sides of body with a row of distinct black blotches - C. kaianus Main tip of preopercular spine long and slender, largest point on its dorsal side without a basal hook; D, with rows of v guenther! _ aaltipinnis o whitehead! e formosanus 12a. 12b. liSae 13b. Geographical distribution of species of the kaianus-group of the genus Callionymus. blotches; sides of body with a row of in- distinct brownish biotches Anal fin with a dark margin; D, with 2 rows of white blotches and a darkish dis- tal margin; main tip of preopercular spine about 2.5—3.0 times as long as larg- est point at its dorsal side; body with light blotches edged with semicircular darkslines "= 2 a C. moretonensis Anal fin pale, without a dark margin; D, with a basal and 2-3 more distal rows of dark spots; main tip of preopercular spine about 1.0—1.5 times as long as larg- est point at its dorsal side; body with minute blackish spots forming rings and blotches C. ochiaii Second membrane of D, deeply incised; caudal fin with 2 long filaments which are nearly twice as long as rest of fin; D, with 2 or more black blotches, largest reaching from Ist to 4th spine; D, with vertical dark streaks _.___ -C.. bentuviai Second membrane of D, not incised; caudal fin with | or 2 relatively short fil- FRICKE: KAIANUS-GROUP OF GENUS CALLIONYMUS aments which are not longer than rest of fin; D, with a black blotch on 3rd mem- brane (rarely also with an additional black blotch distally on the same mem- brane), occasionally reaching to 2nd spine; D, with rows of dark and/or light blotches (but without vertical dark streaks) 14a. Lateral line in area behind eye with a long branch running downwards; D, with a large black blotch basally on 3rd mem- brane, 1 or 2 branches of which reach 2nd membrane; pectoral fin base with 2 dark streaks; anal fin with a distal black margin (usually including tips of fin rays); caudal fin without 2 median trans- verse black lines; D, with 1 basal, | me- dian, and 1| distal row of light spots and 2 median rows of black spots. tot C. africanus (male) Lateral line without a branch in the post- orbital area; D, with a relatively small distal black blotch on 3rd membrane; pectoral fin base with a dark area; distal half of anal fin black, tips of fin rays white; caudal fin with 2 median trans- verse black lines; D, with a basal and a median row of black blotches. 14b. Callionymus carebares Alcock, 1890 (Figures 4—5) Callionvmus carebares ALCOCK, 1890:209 (‘‘off Madras coast, 98-102 fms’’); 1898:73; 1899:pl. 20, fig. 4; REGAN 1906:329 (Sea of Oman, 98-180 fms [179-329 m]); SmitH 1963:555, pl. 84K (after Alcock). MATERIAL EXAMINED.—Syntypes: BMNH 1890.11.28. 18- 24,2 6,5 2, ‘Investigator,’ off Ganjam Coast, India. Other specimens: BMNH 1903.5.14.34, 1 spec., 39.0 mm SL, J. W. Townsend, Karachi. BMNH 1903.9.24.2-4, 3 spec., J. W. Townsend, Iranian Mekran coast, Gulf of Oman (25°19'N, 58°21'E), 98 fms (179 m). BMNH 1904.5.25.218- 220, 3 spec., J. W. Townsend, Sea of Oman, 180 fms (329 m). BMNH 1939.5.24.1384, | spec., John Murray Exped., 23 Nov. 1933, Gulf of Oman, 193 m. BMNH 1939.5.24.1385—-1409, 24 spec., John Murray Exped., Arabian Sea, 135-183 m. BMNH 1939.5.24.1410-1421, 15 spec., John Murray Exped., Gulf of Aden, 220 m. IRSN 1797, 2 2, M. Frank, 4 Apr. 1894, Gulf of Bengal. FMNH 5740, 1 2, J. W. Townsend, 1906, Sea of Oman (Dr. D. J. Stewart, FMNH, kindly examined the spec- men). D1AGNosis.—A Callionyvmus of the kaianus- group with a very large branchial opening (same size as, or larger than, pupil), an unusually large 353 2y S yee TZ sans FiGuRE 3. Preopercular spines of species of the Adianus- group. The main characteristics are the straight ventral side of the spine, the strong straight antrorse point, the upcurved main tip, and the antrorse spine in combination with two (rare- ly one) large curved points on the dorsal side. (A) Left pre- opercular spine of Callionvmus kaianus. (B) Left preopercu- lar spine of C. guentheri. (C) Left preopercular spine of C. africanus. (D) Left preopercular spine of C. moretonensis. (E) Left preopercular spine of the most primitive species of the kaianus-group, C. carebares (without an antrorse point on the dorsal side). head (head length in SL 2.7—3.1), and two large curved points, but without a small antrorse spine at dorsal edge of preopercular spine. DESCRIPTION.—Counts and measurements (see also Table 1): D, IV; D. vuii,1; A vin,t; P, li—ii, 14—16,111; P, 1,5; C t1,2—3,0-11,2—4, 101. Body elongate and depressed. Head very large, depressed, 2.7—3.1 in SL. Eye large, 2.7— 3.5 in head. Pupil large, about 2.5 in eye. Bran- chial opening very large, same size as pupil or larger. Occipital region with a bony plate and two low bony protuberances. 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Specimen 1, female, 85.8 mm SL: (A) lateral view; (B) dorsal view; (C) ventral view; (D) left preopercular spine; (£) right preopercular spine. Specimen 2, female, 86.2 mm SL: (F) right preopercular spine. 5 base (formula: | 1; see Fig. 4D-F). Lat- eral line reaching from hind margin of eye to base of caudal fin; the line of the opposite side is interconnected by a transverse branch across the occipital region. Caudal peduncle length 5.8- 6.6 in SL, minimal caudal peduncle depth 21.0- 22 6nsS 1. First spine of first dorsal fin nearly as long as first ray of second dorsal fin, filamentous only in females. Distal margin of second dorsal fin straight. Anal fin beginning on vertical through second ray of second dorsal fin. Distal margin of caudal fin slightly convex, the two median rays elongate but usually not filamentous. Outer edge of pelvic fin convex; longest pelvic fin ray reaching to base of first anal fin ray. Pectoral fin reaching nearly to fifth ray of second dorsal fin when laid back. Color in alcohol. Head and body dark gray, belly white. Eye darkish. First dorsal fin in male monochromatic dark, in female nearly colorless, with a large distal dark blotch reaching from sec- ond to fourth spine. Second dorsal fin colorless, distal margin darkish. Distal one-third of caudal fin dark. Distal two-thirds of anal fin black, anal fin base colorless. Pelvic fin colorless, pectoral fin with few dark spots. DISTRIBUTION.—Northern parts of Indian Ocean: Gulf of Aden, Gulf of Oman, coast of India, Arabian Sea (see Fig. 1); 135-330 m on muddy bottoms. DiscUssiOn.—Callionvmus carebares seems to be the most primitive member of the kaianus- group based on the shapes of its preopercular spine (no antrorse spine at its dorsal side) and caudal fin (often no median unbranched ray FRICKE: KAIANUS-GROUP OF GENUS CALLIONYMUS 7 Ficure 5. First dorsal fin in Callionvmus carebares. (A) BMNH 1890.11.28.18, male, 95.0 mm SL, syntype, Ganjam coast, India. (B) BMNH 1890.11.28.19, female, 86.2 mm SL, syntype, Ganjam coast, India. present). It is also unique, however, in its very large head and its extremely large branchial opening (which is porelike and very small in oth- er callionymid fishes). Therefore, it seems to belong to another evolutionary branch in the Kaianus-group, and I assign it to a subgroup of its own. Juvenile specimens, which have a smaller head and a smaller branchial opening, are more similar to the other species of the kaianus-group. Callionymus kaianus Gunther, 1880 (Figure 6) Callionvmus kaianus GUNTHER, 1880:44, pl. 19, fig. B (Kai Is., 129 fms [236 m]); DE BEAUFORT 1951:66-67, fig. 12 (after Gunther); SMitH 1963:553, pl. 84/ (in part: after Giin- ther); SUWARDJI 1965:308—310 (Kai Is., 180-290 m). MATERIAL EXAMINED.—Holotype: BMNH_ 1879.5.14.565, 1 3d, 128.6 mm SL, Challenger Exped., Kai Is. DIAGNosiIs.—A Callionymus of the kaianus- group with a small branchial opening, a short head (about 4.6 in SL), preopercular spine with a small antrorse and two large curved points on dorsal side, straight distal margin of secend dor- sal fin, two short caudal fin filaments, long fila- mentous first spine of first dorsal fin, short main tip of preopercular spine, and pale anal fin. DESCRIPTION.—Counts and measurements (see also Table 2): D, IV; D, viti,1; A viii,1; P, ivig—teniesP, 1,5; C i,2,1,3-iil. Body elongate and depressed. Head de- pressed, about 4.6 in SL. Eye large, 2.35 in head. Pupil relatively small, 3.3 in eye. Bran- chial opening of normal size, about two times in pupil. Occipital region with two low bony pro- tuberances. Preopercular spine 1.45 in eye di- ameter, with a relatively short, slightly upcurved main tip, a small antrorse and two large curved points (the larger with a small additional basal point) on its dorsal side and a large antrorse spine at its base (formula: 1———1; see Fig. 6B). Lateral line reaching from area behind eye to end of third branched caudal fin ray (seen B a 2mm Ficure 6. Callionvmus kaianus, BMNH 1879.5.14.565, holotype, male, 128.6 mm SL, Kai Islands: (A) lateral view; (B) left preopercular spine. 358 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 14 gv * (PA Z 5 Sot csioe) i 2mm D 2mm FiGureE 7. Callionvmus moretonensis, NMB 37074, 69.4 mm SL, New Caledonia: (A) lateral view; (B) dorsal view; (C) left preopercular spine; (2) right preopercular spine. from above); the line of one side is intercon- nected to its opposite member by a transverse branch across the occipital region and one across the dorsal side of caudal peduncle. Cau- dal peduncle length 5.2 in SL, minimal caudal peduncle depth 23.8 in SL. First spine of first dorsal fin long and filamen- tous. Distal margin of second dorsal fin straight. Anal fin beginning on the vertical through mid- base of second membrane of second dorsal fin. Distal margin of caudal fin pointed, the two un- branched median rays filamentous (filaments rel- atively short). Distal margin of pelvic fin con- vex; longest pelvic fin ray only reaching the anal papilla. Pectoral fin reaching to second anal fin ray when laid back. Color in alcohol. Head and body light brown, back with some lighter blotches. Belly white, thorax brownish. Eye light gray. Pectoral fin base with a large dark blotch. A row of dark blotches in groups along body side. Operculum with some dark spots, head with indistinct brownish blotches and lines. First dorsal fin light, with a black blotch on third membrane, a FRICKE: KAIANUS-GROUP OF GENUS CALLIONYMUS basal branch of which reaches nearly to second spine. Second dorsal fin colorless, with white lines in characteristic arrangement (see Fig. 6A). Anal fin colorless, with few yellow pigment on the membranes between the five posterior rays. Upper and median parts of caudal fin with dark spots and blotches; lower part with a broad curved dark band. Distal part of pelvic fin dark; pectoral fin colorless. DIsTRIBUTION.—Kai Islands (west of New Guinea), at three different localities (Fig. 1); 180-290 m, muddy bottom. Discussion.—C. kaianus differs from C. moretonensis of northern Australia, the species geographically nearest, by its shorter main tip of preopercular spine, and by various color markings (e.g., pale anal fin without a black dis- tal margin); it differs from C. whiteheadi by the presence of filaments in the first dorsal and cau- dal fins, the barbed largest point on dorsal side of preopercular spine, and various color mark- ings. Specimens referred to C. kaianus from the Gulf of Thailand seem to belong to another species. Callionymus moretonensis Johnson, 1971 (Figure 7) Callionymus calauropomus: (nec Richardson, 1844) PETERS, 1876:841 (New Ireland); JoRDAN AND SEALE 1905:415 (after Peters); FOWLER 1928:422 (after Peters); MUNRO 1958:253 (after Peters). Callionymus kaianus moretonensis JOHNSON, 1971: 1085113, figs. 1-2 (Ss Queensland); 1973:217—230 (biology). MATERIAL EXAMINED.—Holotype: AMS 115608-001, 1 spec., 158.3 mm SL, C. R. Johnson, 1 Aug. 1969, E of Cape Moreton, Queensland, 68-72 fms (124-132 m). Paratypes: CAS 24764, | spec., 131.2 mm SL; CAS 24765, 1 spec., 134.0 mm SL; CAS 24766, 1 spec., 141.7 mm SL; CAS 24767, 1 spec., 164.9 mm SL; all with same data as holotype. Other specimens: BMNH 1892.1.14.26-27, 2 spec., 52.4— $8.2 mm SL, Mr. Walker, Holothuria Banks (Nw Australia). ZMB 9399, 1 6, 1 juv., 35.0-81.9 mm SL, R/V GAZELLE, “shortly before the year 1876,°° New Ireland. NMB 37074, 1 spec., 69.4 mm SL, P. Fourmanoir, 1979, Havannah, s New Caledonia, 150 m. DiaGnosis.—A Callionymus of the kaianus- group with a small branchial opening; short head (3.44.7 in SL); preopercular spine with a small antrorse and two large curved points on its dor- sal side and a long, slender main tip; second dorsal fin with a nearly straight distal margin; caudal fin with two short median filaments; first dorsal fin with a relatively long, filamentous first spine; anal fin with a dark distal margin; and second dorsal fin with rows of white blotches and a dark distal margin. DESCRIPTION.—Counts and measurements (see also Table 2): D, FV; D, viii,1; A vii, 1—viii,1; Body elongate and depressed. Head de- pressed, about 3.4—4.7 in SL. Eye large, 2.0-2.4 in head. Pupil relatively small, 2.9-3.0 in eye. Branchial opening of normal size, about 2-3 times in pupil. Occipital region with two low bony protuberances. Preopercular spine about 1.1-1.2 in eye diameter, with a long slightly up- curved main tip, a small antrorse and two large curved points at its dorsal side, and a_large an- trorse spine at its base (formula: | 1; see Fig. 7C, D). Lateral line reaching from area be- hind eye to end of third branched caudal fin ray (counted from above); the line of the opposite side is interconnected by a transverse branch across the occipital region and one across the dorsal side of the caudal peduncle. Caudal pe- duncle length 5.8—6.5 in SL, minimal caudal pe- duncle depth 23.0—30.0 in SL. First spine of first dorsal fin relatively long and filamentous (in adults). Distal margin of sec- ond dorsal fin straight. Anal fin beginning on a vertical through second ray of second dorsal fin. Distal margin of caudal fin convex in small spec- imens (somewhat pointed in adults), with two short median filaments. Distal margin of pelvic fin convex; longest pelvic fin ray reaching nearly to midbase of first membrane of anal fin. Pec- toral fin reaching to fourth ray of second dorsal fin. Color in alcohol. Head and body light brown, ventral side of body and belly whitish. Thorax white. Back with white spots bordered by semi- circular black lines. Eye grayish. A row of ir- regular brownish spots along sides of body. Head with small white spots. Some dark spots at upper part of pectoral fin base. First dorsal fin light brown, with a large black blotch on third membrane, a basal branch of which reaches to second spine, and some smaller white blotches surrounding it. Second dorsal fin with two or three rows of white spots and a darkish distal margin. Anal fin colorless, with a black distal border. Pelvic fin with small dark spots on distal parts of fourth and fifth rays. Pectoral fin col- orless or with three vertical rows of darkish spots. Caudal fin whitish, with some dark spots forming a broad curved band in the upper part, a broad curved black band in the lower part. 360 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 14 DIsTRIBUTION.—Northern half of Australia (Holothuria Banks to southern Queensland), New Caledonia, and New Ireland (see Fig. 1); 101-150 m, mud bottom. Discussion.—The differences between C. kaianus and C. moretonensis are discussed in the description of the former species. Callio- nymus moretonensis differs from C. whiteheadi by the presence of filaments in first dorsal and caudal fins, by the shape of the main tip of the preopercular spine, and by color markings. The record of Synchiropus calauropomus (Richardson, 1844) from New Ireland (Peters 1876) is based on two specimens of Callionymus moretonensis. Synchiropus calauropomus does not occur in that area. Johnson (1971) originally described C. more- tonensis as a subspecies of C. kaianus, but the differences are sufficient to regard C. moreton- ensis aS a distinct species. Callionymus whiteheadi new species (Figure 8) ?Callionvmus kaianus: (non Gunther, 1880) WEBER 1913: (Madura Sea, 7°2.6’S, 115°23.6’E, 100 m). MATERIAL EXAMINED.—Holotype: BMNH_ 1980.6.20.1, 112.3 mm SL, P. J. P. Whitehead, 14 July 1979, off Bali (8°50'S, 114°14’E), 110-220 m. Paratype: BMNH 1980.11.25.2, 1 spec., 105.5 mm SL, P. J. P. Whitehead, summer 1979, se coast of Java (near type-locality). DIAGNOsISs.—A Callionymus of the kaianus- group with a small branchial opening; short head (3.9-4.2 in SL); preopercular spine with a small antrorse and one or two large curved points, and a short main tip; second dorsal fin with a nearly straight distal margin; caudal fin with a convex distal margin, but without filaments; first dorsal fin without a filament and with a basal black blotch on third membrane; light brown cheeks with few dark spots; anal fin with a black distal margin; second dorsal fin with a basal row of dark spots; and sides of body with a row of dis- tinct black blotches. DESCRIPTION.—Counts and measurements (see also Table 1): D, IV; D, viii,1; A viii,1; P, i-11, 15-17, ii; P, 1,5; C ii,3,i—i1,2—3, iii. Body elongate and depressed. Head de- pressed, 3.9-4.2 in SL. Eye large, 2.2—2.3 in head. Pupil relatively small, about 3.2 in eye. Branchial opening of normal size, about 3 times in pupil. Occipital region with two low bony pro- tuberances. Preopercular spine 1.7—1.9 in eye diameter, with a short, slightly upcurved main tip, a small antrorse and one or two large curved points at its dorsal side, and a large antrorse spine at its base (formula: 1 lessee #Pig: 8B). Lateral line reaching from area behind eye to third branched caudal fin ray (counted from above); the line of the opposite side is intercon- nected by a transverse branch across the occip- ital region and another across the dorsal side of caudal peduncle. Caudal peduncle length 5.5— 6.1 in SL, minimal caudal peduncle depth 25.5— 26.3 nS: First spine of first dorsal fin somewhat longer than first ray of second dorsal fin, but not fila- mentous. Distal margin of second dorsal fin nearly straight. Anal fin beginning on the vertical through second ray of second dorsal fin. Distal margin of caudal fin convex; no median fila- ments. Distal margin of pelvic fin convex; long- est pelvic ray only reaching to anal papilla when laid back. Pectoral fin reaching to midbase of second membrane of second dorsal fin when laid back. Color in alcohol. Head and body dark olive- green; lower surface of body white. Back mar- bled with brown. Thorax and belly white. A dark area at upper part of pectoral fin base. Some large whitish spots bordered by black on oper- culum. Head with dark brown spots and lines. Eye dorsally black, ventrally dark blue. A row of irregular black blotches along sides of body. First dorsal fin olive-green; first spine marbled alternating black and white; a large black blotch basally on third membrane, a basal branch of which reaches to second spine. Second dorsal fin with two rows of indistinct darkish blotches and a basal row of distinct blackish blotches. Anal fin white, with a broad distal black margin. Caudal fin rays in upper part of caudal fin with black blotches; lower part of caudal fin blackish distally. Distal two-thirds of pelvic fin darkish; upper half of pectoral fin with four vertical rows of black spots. DISTRIBUTION.—Bali and southeastern Java, possibly also Madura Sea (see Fig. 2); 110-220 m, mud bottom. Discussion.—The differences between C. whiteheadi, C. kaianus, and C. moretonensis have been discussed in the descriptions of the last two species. C. whiteheadi differs from C. regani by the shape of the preopercular spine and by various color markings (e.g., black bor- der of anal fin; shape and position of black blotch in first dorsal fin; color patterns of second FRICKE: KAJANUS-GROUP OF GENUS CALLIONYMUS 361 B Pi 2mm FIGuRE 8. opercular spine. dorsal and caudal fins; body color pattern, etc.). It differs from C. guentheri in lacking caudal fin filaments, in the different shape of the preoper- cular spine, and in various color markings. ETYMOLOGY.—This new species is named af- ter Dr. Peter J. P. Whitehead, British Museum (Natural History), who collected the type-spec- imens and allowed me to examine them. Callionymus regani Nakabo, 1979 (Figure 9) Callionvmus kaianus: (non Gunther, 1880) REGAN 1908:248 (Saya de Malha Bank, 123 fms [225 m]); SmiTH 1963 (part):553 (after Regan). , Callionvmus regani NAKABO, 1979:231-234, fig. 1, table 1 (Saya de Malha Bank). MATERIAL EXAMINED.—BMNH 1908.3.23.263, 1 d, 101.3 mm SL, Gardiner-Expedition, Saya de Malha Bank (western- central Indian Ocean), “‘over 123 fms” (225 m). DiAaGNosis.—A Callionymus of the kaianus- group with a small branchial opening; short head (about 4.1 in SL); preopercular spine with a small antrorse and two large curved points at dorsal side, and a short main tip; second dorsal fin with a nearly straight distal margin; caudal fin with a convex distal margin but without fil- Callionymus whiteheadi, holotype, BMNH 1980.6.20.1, 112.3 mm SL, off Bali: (A) lateral view; (B) left pre- aments; first dorsal fin without a filament and with a distal black blotch on third membrane surrounded by white lines; dark brown cheeks with characteristic light blotches; colorless anal fin without a distal black margin; second dorsal fin with rows of white spots and lines; and sides of body with few large indistinct brownish blotches. ; DESCRIPTION.—Counts and proportions (see Table 1): D, IV; D, viii,1; A viti,1; P, i-ii,17- 19,0-i; P, 1,5; C 0-ii,3,i—ii1,2—3,ii-1. Body elongate and depressed. Head de- pressed, 4.1 in SL. Eye large, about 2.7 in head. Pupil relatively small, 3.4 in eye diameter. Bran- chial opening of normal size, about 3 in pupil. Occipital region with two low bony protuber- ances. Preopercular spine 1.4 in eye diameter, with a short, slightly upcurved main tip, a small antrorse and two larger curved points at its dor- sal side, and a large antrorse spine at its base (formula: 1 1; see Fig. 98). Lateral line reaching from area behind eye to end of third branched caudal fin ray (seen from above); the line of the opposite side is interconnected by a transverse branch across the occipital region and another across the caudal peduncle. Caudal 362 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 14 B ay 2mm FiGureE 9. Callionvmus regani, BMNH 1908.2.23.263, male, preopercular spine. peduncle length 6.25 in SL; minimal depth of caudal peduncle 22.5 in SL. First spine of first dorsal fin somewhat longer than first ray of second dorsal fin, but not fila- mentous. Distal margin of second dorsal fin nearly straight. Anal fin beginning on vertical through second ray of second dorsal fin. Distal margin of caudal fin nearly convex; no median filaments. Distal margin of pelvic fin convex; longest pelvic fin ray reaching to base of first anal fin ray when laid back. Pectoral fin reaching to midbase of third membrane of second dorsal fin when laid back. Color in alcohol. Head and body brown. Belly white, thorax light brown. Three to four large indistinct darkish areas along sides of body. Head brown; males with many light blotches, females with few light blotches. Occipital region with dark spots. Eye posteriorly gray, anteriorly yellowish. A large dark brown blotch at pectoral fin base. First dorsal fin pale; about four hori- zontal dark lines on first and second membranes. 101.3 mm SL, Saya de Malha Bank: (A) lateral view; (B) left A black blotch distally on third membrane, oc- casionally reaching to posterior part of second membrane; distal edge of third membrane also black. Remaining parts of third and fourth mem- branes covered with curved white lines. Second dorsal fin mostly colorless, with rows of white blotches and/or lines. Anal fin pale; distal margin yellowish. Caudal fin pale, occasionally with two darkish blotches at upper edge; lower part sometimes dusky. Distal parts of fourth and fifth rays of pelvic fin darkish; pectoral fin colorless. DISTRIBUTION.—Saya de Malha Bank, west- ern-central Indian Ocean (see Fig. 2); 126- 225 m. DiscUSSION.—The differences from Callio- nymus whiteheadi were discussed in the descrip- tion of that species. Callionymus regani differs from C. africanus by the absence of caudal fin filaments and by a completely different color pattern; it differs from C. kotthausi in its shorter main tip of preopercular spine, shorter first spine of first dorsal fin, shorter caudal fin (especially FRICKE: KAIANUS-GROUP OF GENUS CALLIONYMUS 363 Ficure 10. Callionvmus kotthausi, ZIM 5535, holotype, 114.8 mm SL, off Cochin (India): (A) lateral view; (B) left pre- opercular spine; (C) right preopercular spine. ZIM 5536, paratype, specimen 1, male, 81.2 mm SL, Cochin: (D) first and second dorsal fins. in males), and various color markings (e.g., col- or patterns of first and second dorsal fins, and anal fin, head, and sides of body). Callionymus kotthausi new name (Figure 10) Callionvmus kaianus: (non Ginther, 1880) ALcock 1899:74 (Malabar Coast, India, 102 fms [187 m]}). Diplogrammus indicus KorrHaus, 1977:40-41, figs. 423b, 424b, 425 (wsw of Cochin, India). MATERIAL EXAMINED.—Holotype: ZIM 5535, 2, 114.8 mm SL. A. Kotthaus, R/V MeTeEorR, 10 Feb. 1965, about 40 km wsw of Cochin, India (09°40’N, 75°38.8’E to 09°45.3’N, 75°38.5'E), 211-138 m. Paratypes: ZIM 5536, 2 36,9 &, 71.3- 87.0 mm SL; same data as holotype. DiaGNosis.—A Callionymus of the kaianus- group with a small branchial opening; short head (3.84.1 in SL); preopercular spine with a small antrorse and one or two large curved points, and a long main tip; second dorsal fin with a nearly straight distal margin; caudal fin without fila- ments: anal fin with a broad dark margin; sides of body with a row of small distinct black blotch- es: and colorless lower part of caudal fin. DESCRIPTION.—Counts and measurements 364 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 14 (see also Table 1): D, IV; D, viii,1; A viti,1; P, Ilo By 5 Cou, 331, 351i. Body elongate, depressed. Head depressed, 3.8-4.1 in SL. Eye large, about 2.4 in head. Pu- pil about 2.8 in eye. Branchial opening of normal size, about 2-3 times in pupil. Occipital region with a low bony protuberance. Preopercular spine about 1.4 in eye diameter, with a long up- curved main tip, a small antrorse and one or two large curved points on its dorsal side, and a large a antrorse spine at its base (formula | la SEG Fig. 10B,C). Lateral line reaching from area be- hind eye to end of median unbranched caudal fin ray, with a long branch at its ventral side in postorbital region; the line of the opposite side is interconnected by a transverse branch across the occipital region and another across the cau- dal peduncle. Caudal peduncle length about 6.5 in SL; minimal caudal peduncle depth about 24.0 nm SL. First spine of first dorsal fin long (longer than first ray of second dorsal fin), but not filamen- tous. Distal margin of second dorsal fin nearly straight. Anal fin beginning on the vertical through third ray of second dorsal fin. Distal margin of caudal fin convex or slightly pointed, without filaments. Distal margin of pelvic fin convex; longest pelvic fin ray reaching only to anal papilla when laid back. Pectoral fin reaching to middle of third membrane of second dorsal fin when laid back. Color in alcohol. Head and dorsal side of body yellowish brown; back with some dark-edged whitish blotches. Sides of body with a row of dark spots. Thorax, belly, and lower parts of body yellowish white. Eye dark gray. Opercu- lum with dark spots. First dorsal fin brownish: males with one or two black blotches surround- ed by white on second and third membranes; females with an elongate ocellated basal black blotch on first to third membranes. Second dor- sal fin transparent, distally darkish, with two rows of elongate dark spots. Distal margin of anal fin black, tips of rays white. Caudal fin mostly colorless, with 3—4 vertical rows of light brown spots in its upper part. Distal half of fourth and fifth pelvic fin rays dark; pelvic fin basally with irregular darkish spots. DISTRIBUTION.—Southwest coast of India (see Fig. 2); 138-211 m. Discussion.—Kotthaus (1977) assigned Dip- logrammus indicus to the genus Diplogrammus using Smith’s (1963:549) key: “‘A skinny keel along lower flank from tip of pelvic to caudal base,’ in combination with ‘‘an antrorse spine at base of preopercular spine.’’ The latter feature is also valid for species of the genus Calliony- mus, but the specimens of ‘“‘Diplogrammus in- dicus’’ have neither a skinny keel along lower flank of body, as in species of Diplogrammus, nor any other diagnostic feature of Diplogram- mus (e.g., a free flap of skin at the operculum, a lateral line with many branches). Kotthaus’s species belongs in the genus Callionymus, and it posseses all features of the kaianus-group of the subgenus Callionymus. The binomen Cal- lionymus indicus (Kotthaus, 1977), however, becomes a secondary homonym of Callionymus indicus Linnaeus, 1758 (a Platycephalidae now well known as Platycephalus indicus) and must be replaced. ETYMOLOGY.—The species is named after Dr. A. Kotthaus, who first described the species. In accordance with Dr. Kotthaus, who is presently unable to create a new name for the species be- cause of his health, I propose the new name Callionymus kotthausi to replace Callionymus indicus (Kotthaus, 1977). Callionymus africanus (Kotthaus, 1977) (Figure 11) Callionvmus kaianus: (non Giinther 1880) NORMAN 1939:73 (Zanzibar area); SMITH 1963 (part):553 (after Norman). Diplogrammus africanus KOTTHAUS, 1977(part):38—40, figs. 421, 422, 423a, 424a (NE of Mombasa). MATERIAL EXAMINED.—Holotype: ZIM 5533, 2, 102.3 mm SL, A. Kotthaus, R/V METeor, 14 Jan. 1965, about 180 naut. miles NE of Mombasa (01°18’S, 41°56’E to 01°19.8'S, 41°53'E). Paratypes: ZIM 5534, 11 d, 19 2, same data as holotype. Other specimens: BMNH_ 1939.5.24.1422, 1 d, 75.5 mm SL, John Murray Exped., 12 Jan. 1934, near Zanzibar (5°38'54"S, 39°15'42"E to 5°40'18”"S, 39°17'36"E); green mud bottom; bottom temperature 15.52°C; bottom salinity 35.21%c. DiAGNosis.—A Callionymus of the kaianus- group with a small branchial opening; short head (about 4.0 in SL); preopercular spine with a small antrorse and two larger curved points at dorsal side, and a relatively short main tip; near- ly straight distal margin of second dorsal fin; two short median caudal fin filaments; first dorsal fin without a filament and with a normal (not in- cised) second membrane; anal fin with a narrow black distal margin; no vertically elongated dark blotches in median part of caudal fin; back with FRICKE: KAIANUS-GROUP OF GENUS CALLIONYMUS 365 B _— 2mm Ficure 11. opercular spine. dark-bordered light spots; and pectoral fin base with two transverse dark streaks. DESCRIPTION.—Counts and proportions (see also Table 1): D, IV; D, viii,1; A viii,1; P, i- ii, 16—20,i—ii; P, 1,5; C i1,3,0,2,i0. Body elongate and depressed. Head de- pressed, 3.9-4.1 in SL. Eye large, 2.0—2.2 in head. Pupil relatively small, about 3.7 in eye diameter. Branchial opening of normal size, about 3 in pupil. Occipital region with two low bony protuberances. Preopercular spine 1.8 in eye diameter, with a short, slightly upcurved main tip, a small antrorse and two large curved points on its dorsal side, and a large antrorse spine at its base (formula: 1 1; see Fig. 11B). Lateral line reaching from area behind eye to end of third branched caudal fin ray (counted from above); the line of the opposite side is in- terconnected by a transverse branch across oc- cipital region and another across dorsal side of caudal peduncle. Lateral line with a long branch at its ventral side behind eye. Caudal peduncle length 6.0-6.1 in SL, minimal caudal peduncle depth 19.9-21.3 in SL. First spine of first dorsal fin somewhat longer than first ray of second dorsal fin, but not fila- mentous. Distal margin of second dorsal fin nearly straight. Anal fin beginning on the vertical through second ray of second dorsal fin. Distal Callionvmus africanus, BMNH 1939.5.24.1422, male, 75.5 mm SL, Zanzibar: (A) lateral view; (B) left pre- margin of caudal fin convex; males with two rel- atively short median filaments, females without filaments. Distal margin of pelvic fin convex; longest pelvic fin ray reaching to base of first anal fin ray when laid back. Pectoral fin reaching to fourth ray of second dorsal fin when laid back. Color in alcohol. Head and body brown; ven- tral side of body lighter, belly and thorax white A row of paired dark brown blotches on side of body. Back with light spots bordered with dark brown. Pectoral fin base with two transverse dark lines. Eye dark blue, dorsally with dark brown blotches. First dorsal fin brown; a large white-edged black blotch basally on third mem- brane, a basal and a distal branch of which reaches to second spine; occasionally, also with a small distal black blotch on third membrane. Second dorsal fin with three rows of white and two rows of darkish spots; these rows alternate- ly arranged. Anal fin pale, with a narrow black distal margin. Lower part of caudal fin with a broad curved dark bar, upper part scattered with dusky spots. Distal parts of fourth and fifth pel- vic fin rays dark; upper part of pectoral fin with three vertical rows of dark spots. SEXUAL DIMORPHISM.—As described by Kotthaus (1977:40). DISTRIBUTION.—Known from Zanzibar and southern Somalia (see Fig. 1), possibly distrib- 366 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 14 uted along entire east African coast; 212 m, mud bottom. Discussion.—The differences from Callio- nymus regani are discussed in the description of that species. The two paratypes described by Kotthaus (1977:39) from the southern Red Sea are small specimens of Callionymus bentuviai. Callionymus bentuviai new species (Figure 12) Diplogrammus africanus KOTTHAUS, 1977 (part):39 (s Red Sea). MATERIAL EXAMINED.—Holotype: HUJF 9935, 87.8 mm SL, A. Ben-Tuvia, 1957, Eritrea (Ethiopia; s Red Sea). Para- types: HUJF 8068, 2 spec., 79.7-85.2 mm SL, same data as holotype; ZIM 5532, 2 spec., 34.2-46.9 mm SL, A. Kotthaus, R/V METEOR, 6 Dec. 1964, s Red Sea, 70-75 m. DiaGnosis.—A Callionymus of the kaianus- group with a small branchial opening; short head (3.6—4.0 in SL); preopercular spine with a small antrorse and two larger curved points at its dor- sal side, and a relatively long main tip; second dorsal fin with a nearly straight distal margin; two very long median caudal fin filaments; first dorsal fin without a filament and with a deeply incised second membrane; and second dorsal fin with vertical dark streaks. DESCRIPTION.—Counts and proportions (see also Table 1): D, IV; Ds vin,1; A viti,1; P, i- ii, 1S—17,i-11; P, 1,5; C 11,2—4,1-11,2—3 ,1—-i11. Body elongate and depressed. Head de- pressed, 3.6—4.0 in SL. Eye large, 2.15—2.85 in head. Pupil relatively small, about 2.7 in eye. Branchial opening of normal size, about 1.5 in pupil. Occipital region with two low bony pro- tuberances. Preopercular spine 1.2—1.6 in eye diameter, with a long slightly upcurved main tip, a small antrorse and two larger curved points on its dorsal side, and _a large antrorse spine at its base (formula: 1————1; see Fig. 12C,D). Lat- eral line reaching from area behind eye to end of third branched caudal fin ray (counted from above); the line of the opposite side is intercon- nected by a transverse branch across occipital region and another across caudal peduncle. Cau- dal peduncle length 4.9-5.8 in SL, minimal cau- dal peduncle depth 23.1—24.9 in SL. First spine of first dorsal fin longer than first ray of second dorsal fin, but not filamentous. Second membrane of first dorsal fin deeply in- cised (not in young specimens). Distal margin of second dorsal fin nearly straight. Anal fin begin- ning on the vertical through midbase of second membrane of second dorsal fin. Distal margin of caudal fin convex, with two very long median filaments. Distal margin of pelvic fin mostly con- vex; longest ray reaching to middle of second membrane of anal fin when laid back. Pectoral fin reaching to fourth ray of second dorsal fin when laid back. Color in alcohol. Head and body dark brown; ventral side of body dark brown, belly whitish. Back nearly monochromatic, with very few small dark spots. A row of small pale blotches along side of body, occasionally also a row of two groups of three dark spots each. Many black spots on operculum and at pectoral fin base. First dorsal fin light brown, with two darkish transverse lines distally between first and third spines. A large, elongate, curved black blotch basally from first to fourth spine, mainly on third membrane. Another black blotch dis- tally on third membrane. Second dorsal fin pale, with nine vertical dark streaks. Anal fin dark brown, with a black distal margin. Caudal fin pale; dorsal part with a narrow curved dark line, ventral part with a broad curved dark bar. Pelvic fin pale; upper one-third with three vertical rows of dark spots. DisTRIBUTION.—Known only from the south- ern Red Sea (see Fig. 2); 70-75 m. Discussion.—Callionymus bentuviai differs from all other species of the kaianus-group in its deeply incised second membrane of the first dorsal fin, the absence of a dorsal fin filament in combination with two very long median caudal fin filaments, and an unusual color pattern (e.g., in the dorsal fins). ETYMOLOGY.—This species is named for Pro- fessor Adam Ben-Tuvia (Hebrew University of Jerusalem), who collected the holotype and sent it to me for examination. Callionymus ochiaii new species (Figure 13) Callionvmus kaianus: (non Giinther, 1880) OcHIAI, ARAGA, AND NAKAJIMA 1955:111-113, figs. 8-10, table 6 (various s Japan localities); MATSUBARA 1955:713 (after Ochiai, Ara- ga, and Nakajima); KAMOHARA 1964:90 (Kochi Pref. to Kagoshima; deep-sea bottom, very rare); MASUDA, ARAGA, AND YOSHINO 1975:261, pl. 84D (Kumano Bay south). MATERIAL EXAMINED.—Holotype: FAKU 23261, ¢, 95.0 mm SL, I-3 Sep. 1954, Shibushi, Kagoshima Pref., Japan. Paratypes: FAKU 23257-23260 and FAKU 23275, 2 6, 3 2, 77.8-116.8 mm SL, same data as holotype. DIAGNOsIS.—A Callionymus of the kaianus- group with a small branchial opening; short head FRICKE: KAIANUS-GROUP OF GENUS CALLIONYMUS ane C Oomm FiGure 12. preopercular spine; (D) right preopercular spine. (3.54.4 in SL); preopercular spine with a small antrorse and two larger curved points at its dor- sal side, and a relatively long main tip; nearly straight distal margin of second dorsal fin; one or two short median caudal fin filaments; first dorsal fin with a long filament (males) or a fila- ment of median size (females); anal fin without a distal dark margin; and long, slender main tip of preopercular spine, which is longer than the longest point on its dorsal edge. 367 D Callionyvmus bentuviai, HUJF 9935, holotype, 87.8 mm SL, Eritrea: (A) lateral view; (B) dorsal view; (C) left DESCRIPTION.—Counts and proportions (see also Table 2): D, IV; D, viii,1; A viii,1; P, 11,16— 18,i; P, 1,5; C ii,3(—4),i,(2-)3, 11. Body elongate and depressed. Head de- pressed, 3.5-4.4 in SL. Eye large, 2.3-3.0 in head. Pupil relatively small, about 3.65 in eye diameter. Branchial opening of normal sIZe, about 2.5 in pupil. Occipital region with two low bony ridges. Preopercular spine 1.3—1.8 in eye diameter, with a long, slightly upcurved main 368 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 14 FiGure 13. Callionyvmus ochiaii, holotype, FAKU 23261, male, 95.0 mm SL, Shibushi (Japan): (A) lateral view; (B) left preopercular spine. Paratype, FAKU 23258, female, 116.8 mm SL, Shibushi (Japan): (C) first dorsal fin. tip, a small antrorse and two larger curved points at its dorsal side, and a large antrorse spine at its base (formula: 1 levsee Fig: 13B). Lateral line reaching from area behind eye (from where a branch runs down to base of pre- opercular spine) to end of third branched caudal fin ray (counted from above); the line of the op- posite side is interconnected by a transverse branch across occipital region and another across dorsal part of caudal peduncle. Caudal peduncle length 5.3-6.1 in SL; minimal caudal peduncle depth 24.3-27.8 in SL. First spine of first dorsal fin filamentous (fil- ament in females relatively short). Second mem- brane of first dorsal fin not incised. Distal margin of second dorsal fin nearly straight. Anal fin be- ginning on a vertical through midbase of second membrane of second dorsal fin. Distal margin of caudal fin irregular, with two short median fila- ments. Distal margin of pelvic fin convex; long- est pelvic fin ray reaching to midbase of first anal fin membrane. Pectoral fin reaching to fourth ray of second dorsal fin when laid back. Color in alcohol. Body brownish yellow above, whitish below. Back with minute black- ish spots, forming rings and blotches. A row of dark brown blotches along sides of body. Pec- toral fin base with a dark area. Head with some irregular dark spots and blotches. First dorsal fin gray, with a black blotch basally or centrally on third membrane, a basal branch of which oc- casionally reaches to second spine. Second dor- sal fin gray, with large irregular dark blotches and submarginal dark bands. Caudal fin pale ex- NS ee FRICKE: KAIANUS-GROUP OF GENUS CALLIONYMUS cept for the darker lower part where the fin is scattered with several pale pearl-white round spots. Anal fin dusky, sometimes indefinitely but broadly edged with brown (but not with black). Pectoral fin uniformly pale; pelvic fin pale, outer posterior margin more or less dark. DIsTRIBUTION.—Southern Japan (see Fig. 1); about 100 m, sand and mud bottoms. DiscUSSION.—From the most similar species, C. moretonensis, C. ochiaii differs in having a somewhat shorter main tip of preopercular spine, an irregular distal margin of caudal fin, and several different color markings (especially the absence of a distal black band in the anal fin; the color patterns of the caudal and second dor- sal fins and of the pectoral fin base; the body color pattern, etc.). Callionymus ochiaii differs from C. kaianus in the shape of the largest point on the dorsal side of the preopercular spine, in the length of the filament in the first dorsal fin, and in various color markings. The first dorsal fin filament of the male spec- imen figured in Ochiai, Araga, and Nakajima (1955:fig. 8) is very short. Because the specimen (91.0 mm SL) is nearly as long as the holotype (95.0 mm SL), the filament might have been bro- ken in that specimen. ETYMOLOGY.—This new species is named for Dr. Akira Ochiai, who first described the species under the name Callionymus kaianus. Callionymus formosanus new species (Figure 14) ?Callionymus kaianus: (non Giinther, 1880) CHu 1957:22 (Pes- cadores Islands). MATERIAL EXAMINED.—Holotype: CAS 46972, 3, 104.0 mm SL, F. B. Steiner, Apr. 1971, Formosa Str. (25°N, 120°E), approximately 90 m. DiaGnosis.—A Callionymus of the kaianus- group with a small branchial opening; short head (about 4.5 in SL); preopercular spine with a small antrorse and two larger curved points at dorsal side; relatively high second dorsal and anal fins with convex distal margins; filamentous first spine of first dorsal fin; large black blotch nearly on entire third membrane of first dorsal fin; anal fin with a distal black streak on each membrane; distal two-thirds of anal fin brown: and white anal fin ray tips. DESCRIPTION.—Counts and proportions of the holotype (see also Table 2): D, IV; D, viii,l; A wii,b: P, a,16—17,1; P, 1,52 /€ 1i,3,11,2,i11. Body elongate and depressed. Head de- 369 pressed, about 4.5 in SL. Eye large, about 2.3 in head. Pupil relatively small, about 3.4 in eye diameter. Branchial opening of normal size. about 2.5 in pupil. Occipital region with two low bony ridges. Preopercular spine about 1.7 in eye diameter, with a slightly upcurved main tip of medium size, a small antrorse and two larger curved points at its dorsal side, and a large an- trorse spine at its base (formula: | lessee Fig. 14B). Lateral line reaching from area behind eye (from where a long branch runs downwards) to middle of upper median unbranched caudal fin ray; the line of the opposite side is intercon- nected by a transverse branch across occipital region and another across dorsal side of caudal peduncle. Caudal peduncle length 4.9 in SL, minimal caudal peduncle depth 13.0 in SL. First spine of first dorsal fin filamentous. Sec- ond dorsal fin relatively high, distal margin con- vex. Anal fin beginning on a vertical through midbase of first membrane of second dorsal fin. Distal margin of caudal fin convex, with one rel- atively long filament. Distal margin of pelvic fin convex; longest pelvic fin ray reaching to mid- base of first membrane of anal fin. Pectoral fin reaching to third ray of second dorsal fin when laid back. Color in alcohol. Body dark brown, belly whitish. Side of body with a row of irregular blackish blotches. Head dark brown, with some lighter spots. A vertical dark streak under the eye. A dark blotch at pectoral fin base. Back and sides of body with whitish dark-edged blotches. First dorsal fin pale, with a large dark blotch on third membrane and three white blotches on first and second membranes. Second membrane with a distal black margin. Second dorsal fin pale, with a vertical white streak on each fin ray. Distal margin of first three rays of second dorsal fin black. Anal fin pale, distal two- thirds brownish, with a distal black streak on each membrane; tips of fin rays white. Caudal fin pale, with a vertical row of dark spots; lower margin and lower distal margin dark. Pelvic fin with irregular darkish spots and a dark distal margin of third, fourth, and fifth rays. Upper part of pectoral fin with three vertical rows of dark spots. DISTRIBUTION.—Known only from Formosa Strait (see Fig. 2): about 90 m. DISCUSSION.—This interesting new species belongs to the subgroup of species with a high second dorsal fin in the kaianus-group. It seems 370 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 14 = = as Se ESI S s “NY SRS 2mm FiGure 14. Callionvmus formosanus, holotype, CAS 46972, preopercular spine. to be a primitive member of that subgroup be- cause the second dorsal fin is still relatively low. The specimen recorded as Callionymus kaianus by Chu (1957) from Pescadores Islands (southern part of Formosa Strait) is probably this species. EtyMOLOGY.—This new species is named af- ter its type-locality (Formosa Strait). Callionymus guentheri new species (Figures 15-17) Callionyvmus curvicornis: (non Valenciennes, 1837) GUNTHER 1880: 53 (Philippines, 82 fms [150 m]); HERRE 1953:777 (af- ter Gunther). MATERIAL EXAMINED.—Holotype: BMNH _ 1879.5.14.567, 2, 87.3 mm SL, CHALLENGER Exped., 26 Oct. 1874, w of Zamboanga, Philippine Is. (7°03'N, 121°48’E; entrance from Sulu Sea into Basilan Str., about 7 miles w of Mindanao I., and to the Nw of Caldera Pt.), 82 fms (150 m). Paratypes (all from the Philippines collected by J. E. Norton, 1966): CAS 46966, 4 spec., 3 Nov., Sandoval Pt., Catanauan, Quezon, 70— 78 fms (128-143 m); CAS 32897, 11 spec. (7 36, 4 2), 51.2- 128.0 mm SL, 27 June, Lemery Town, Balayan Bay, Batan- gas, Luzon, 90-85 fms (165-155 m); CAS 33879, 1 2, 112.8 mm SL, 15 Dec., N of San Andres I., Marinduque, 151-158 fms (276-289 m); CAS 34286, 3 d, 100.0-108.3 mm SL, 14 male, 104.0 mm SL, Formosa Strait: (A) lateral view: (B) left Dec., Nw of San Andres I., Marinduque, 137-139 fms (250- 254 m); CAS 34197, 1 6, 8 2, 90.2-129.3 mm SL, 24 Nov., Siburio Pt., Ragay Gulf, Camarines Sur Prov., 319-324 fms (583-593 m); CAS 32905, 4 6, 3 9, 69.3—117.2 mm SE, 15 Nov., Buri Pt., Ragay Gulf, Camarines Sur Prov., 304-309 fms (556-566 m); CAS 34426, 1 2, 84.1 mm SL, 4 July, s of Bauan Town, Batangas Bay, Batangas Prov., 90-88 fms (165— 161 m); CAS 34278, 4 2, 89.9-124.5 mm SL, 11 Dec., N of Melchor I., Marinduque, 120-126 fms (219-231 m); CAS 33864, 3 2, 90.0-127.3 mm SL, 23 Nov., Caurusan Pt., Ragay Gulf, Camarines Sur Prov., 302-308 fms (552-564 m); CAS 32916, 5 2, 91.0-118.2 mm SL, 11 Nov., Pusgo Pt., Ragay Gulf, Quezon, Luzon I., 60-67 fms (110-123 m); CAS 33703, 1 2, 122.0 mm SL, 25 June, s of Barrio Nomong Casto, Ba- layan Bay, 105-100 fms (183-192 m); CAS 34401, 1 2, 117.7 mm SL, 24 June, se of Calaca Town, Balayan Bay, 65—55 fms (119-100 m); CAS 32997, 1 2, 107.5 mm SL, 15 June, Batan- gas, Balayan Bay, s of Barrio Sinisian, 95-99 fms (174-181 m); CAS 34190, 1 3d (113.2 mm SL), 1 2 (99.7 mm SL), 6 Nov., se of Alibijaban I., Ragay Gulf, 81-88 fms (148-161 m); CAS 34272, 1 d, 108.0 mm SL, 14 Nov., Nagas Pt., Ragay Gulf, 297-299 fms (543-547 m); CAS 34154, 7 2, 91.1-129.2 mm SL, 3 Sep., N of San Andres Pt., Marinduque, 108-112 fms (197-205 m); CAS 32668, 2 2, 94.5—102.0 mm SL, 26 July, seE of Talaga, Batangas Bay, 138-131 fms (253-240 m); CAS 32801, 1 d, 91.9 mm SL, 25 Aug., NE of Salomague I., Mar- induque, 142-150 fms (260-275 m); CAS 34205, 1 2, 127.0 mm SL, 10 Dec., Nw of Baltazar I., Marinduque, 142-150 fms (260-275 m); CAS 34468, 1 2, 102.8 mm SL, 27 July, Lemery | FRICKE: KAIANUS-GROUP OF GENUS CALLIONYMUS 371 B aa 2mm FiGure 15. lateral view; (B) left preopercular spine. Town, Balayan Bay, 85-90 fms (155-164 m); CAS 34074, 1 2, 123.2 mm SL, N of Sayao Bay, Marinduque, 61-70 fms (111-128 m); CAS 33067, 1 2, 125.2 mm SL, 18 July, s of Barrio Salong, Luzon I., 120-114 fms (220-208 m). D1aGnosis.—A Callionymus of the kaianus- group with a small branchial opening; short head (3.9-4.2 in SL); preopercular spine with a smail antrorse and two larger curved points on dorsal side; very high second dorsal fin with a convex distal margin (males) or a relatively low second dorsal fin with a nearly straight distal margin (females); first dorsal fin without a filament (or with a very short one in smaller specimens); long and slender main tip of preopercular spine; anal fin with a black distal half and white fin ray tips; regular distal margin of caudal fin, with filaments only in small specimens (caudal fin elongate in adults); and relatively large central black blotch on third membrane of first dorsal fin. DESCRIPTION.—Counts and proportions of the holotype (female) (see also Table 2): D, IV; D, viii,1; A viii,1; P, i—ii,17—19,i—ni; P, 1,5; C ii,2- 3,i—11,2—4 111. Body elongate and depressed. Head de- pressed, 3.9-4.2 in SL. Eye large, 2.3-2.6 in Callionvmus guentheri, holotype, BMNH 1879.5.14.567, female, 87.3 mm SL, off Zamboanga, Philippines: (A) head. Pupil of normal size, 2.7-3.5 in eye di- ameter. Branchial opening of normal size, about 2.0 in pupil. Occipital region with two low bony ridges. Preopercular spine 1.5—1.7 in eye diam- eter, with a slightly upcurved main tip which is relatively long or of medium size, a small an- trorse and two larger curved points on its dorsal side, and a large antrorse spine at its base (for- mula: 1 1; see Fig. 15B, Fig. 16B). The two large spines on the dorsal side of the pre- opercular spine of the holotype possess small basal hooks (see Fig. 15B). Lateral line usually reaching from area behind eye to end of third branched caudal fin ray (counted from above); because of the different caudal fin formula of the holotype (C ii,2,i,4,iii instead of C 11,3,1i,2,111), the lateral line reaches to middle of median un- branched caudal fin ray in that specimen. The lateral line of the opposite side is interconnected by a transverse branch across occipital region and another across dorsal side of caudal pedun- cle. Caudal peduncle length 5.1-6.0 in SL, min- imal caudal peduncle depth 23.0—26.0 in SL. First spine of first dorsal fin somewhat longer than first ray of second dorsal fin but not fila- 372 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 14 B — 2mm FIGURE 16. lateral view; (B) left preopercular spine. mentous (only occasionally in small specimens a very short filament). Second dorsal fin in males relatively high, with a convex distal margin, in females lower, with a nearly straight distal mar- gin. Anal fin beginning on the vertical through base of second ray of second dorsal fin. Distal margin of caudal fin in small specimens convex, with one or two long filaments; in larger speci- mens all median caudal fin rays elongate (the two median rays longest), but not filamentous; even the longest rays are connected by mem- branes. Distal margin of pelvic fin convex; long- est pelvic fin ray reaching to midbase of first or second membrane of anal fin. Pectoral fin reach- ing to fourth or fifth ray of second dorsal fin when laid back. Color in alcohol. Head and body brown. One or two rows of large, irregular dark blotches along side of body. A large dark area on pectoral fin base. Back with dark brown semicircular lines in groups which are bordered by white on one side. Eye dark gray. Belly and thorax white; lower side of body brown. Head with few irreg- ular darkish spots. First dorsal fin light brown, Callionymus guentheri, paratype, CAS 32897 (specimen 1), male, 128.0 mm SL, Balayan Bay, Philippines: (A) with five brownish cross-lines in the anterior part; a dark blotch on third membrane distally or centrally. Second dorsal fin pale, females with a basal and a median row of dark spots and some curved white lines (see Fig. 15A); males with vertical darkish streaks (see Fig. 16A). Distal half of anal fin black; tips of fin rays white. Cau- dal fin in females with a broad dark band in the lower and lower distal parts and two median short transverse dark bands; in the upper part some irregular dark spots. Males have the same caudal fin color pattern, but less intense. Pec- toral fin dorsally with two rows of dark spots. Distal three-fourths of pelvic fin darkish, with irregular dark brown spots and blotches. DiIsTRIBUTION.—Known from various locali- ties in the Philippine Islands (southern coast of Luzon to northern coast of Mindanao; see Fig. 2); 100-593 m, on mud bottoms. Discussion.—The differences between C. guentheri, C. whiteheadi, and C. kaianus have been discussed in the descriptions of the last two species. Callionymus guentheri differs from C. regani in the caudal fin shape, the high second FRICKE: KAIANUS-GROUP OF GENUS CALLIONYMUS SS SS 373 FiGuRE 17. Callionvmus guentheri, paratypes, CAS 32897, Balayan Bay, Philippines. Female (specimen 2), 108.8 mm SL: (A) first and second dorsal fins; (B) caudal fin. Male (specimen 4), 77.6 mm SL: (C) first and second dorsal fins; (D) caudal fin. Male (specimen 10), 85.3 mm SL: (£) first and second dorsal fins; (F) caudal fin. dorsal fin (in males), and various color markings; from C. ochiaii in the caudal fin shape, the lack of a dorsal fin filament (in males), the high sec- ond dorsal fin (in males), and several color markings. In the extensive collections of J. E. Norton (paratypes of C. guentheri), a great depth range can be found (100-593 m); 593 m is the greatest depth recorded for any species of the kaianus- group. Because few specimens are known of most species of that group, it is possible that in the future, specimens of other species will be recorded from comparable depths. EtyMOoLoGy.—This new species is named af- ter Dr. Albert Giinther, who, in 1880, reported the first specimen under the name Callionymus curvicornis. Callionymus altipinnis new species (Figure 18) Callionvmus kaianus: (non Gunther, 1880) CHU ET AL. 1962:723-724, fig. 585 (South China Sea); BESEDNOV 1968:63 (Gulf of Tonkin). Callionymus huguenini: (non Bleeker, 1858) SHEN 1964:202— 203, fig. 2 (Hong Kong). MATERIAL EXAMINED.—Holotype: MSL 0001, d, 123.8 mm SL, S.-C. Shen, Hong Kong Fish Market. Paratypes: CAS 46967, 10 spec., 104.1-144.5 mm SL, R. L. Bolin, 23 July 1958, South China Sea, E of Hainan (20°32'N, 112°45'30"E); CAS 46968, 1 spec., 135.9 mm SL, R. L. Bolin, 21 July 1958, off Tungku Pt., Hainan (19°31’N, 111°24'30"E); CAS 46969, 1 spec., 124.8 mm SL, R. L. Bolin, 23 July 1958, South China Sea, E of Hainan (20°32'N, 112°51’E). DiaGnosis.—A Callionymus of the kaianus- group with a small branchial opening; short head (about 4.2 in SL); preopercular spine with a rel- 374 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 14 B SS 2mm FiGureE 18. opercular spine. atively short main tip; small antrorse and one (rarely two) larger curved points at dorsal side; very high second dorsal and anal fins with con- vex distal margins; filamentous first spine of first dorsal fin; small distal black blotch on third membrane of first dorsal fin; anal fin with a black distal margin (including tips of fin rays), but without a distal black streak on each membrane surrounded by dark brown; and one or two cau- dal fin filaments of medium size. DESCRIPTION.—Counts and proportions of the holotype (see also Table 2): D, IV; D, vili,1; A Viltd Pt t4—L5,0: P>.1,5;.C 1,3,11,3,i Body elongate and depressed. Head de- pressed, about 4.2 in SL. Eye large, about 2.5 in head. Pupil of normal size, about 3.8 in eye. Branchial opening about 3 in pupil. Occipital re- gion with two low bony ridges. Preopercular spine about 1.7 in eye diameter, with a relatively short, slightly upcurved main tip, a small an- trorse and one (rarely two) larger points on its dorsal side, and a large antrorse spine at its base Callionyvmus altipinnis, holotype, MSL 0001, male, 123.8 mm SL, Hong Kong: (A) lateral view; (B) left pre- Ae (formula: (peed see Fig. 188). Lateral line reaching from area behind eye to middle of up- per median unbranched caudal fin ray; the line of the opposite side is interconnected by a trans- verse branch across occipital region and another across dorsal side of caudal peduncle. Caudal peduncle length about 5.0 in SL; minimal caudal peduncle depth about 25.0 in SL. First spine of first dorsal fin filamentous. Sec- ond dorsal fin very high, distal margin convex. Anal fin begins on vertical through middle of second membrane of second dorsal fin. Distal margin of pelvic fin convex; longest pelvic fin ray reaching nearly to base of first anal fin ray. Pectoral fin reaching to middle of third mem- brane of second dorsal fin when laid back. Distal margin of caudal fin somewhat pointed, with one or two filaments. Color in alcohol. Head, body, and fins pale except for a distal black blotch on third mem- brane of first dorsal fin, a row of dark blotches along side of body, eyes grayish, a black distal FRICKE: KAJANUS-GROUP OF GENUS CALLIONYMUS margin of anal fin, and a darkish area on pectoral fin base. DISTRIBUTION.—Known only from the South China Sea off China, from Hong Kong (type- locality) to the Gulf of Tonkin; island of Hainan (see Fig. 2). The exact collecting depth of spec- imens of C. altipinnis is not known, but the species should occur at about the same depth as other species of the kaianus-group. Discussion.—Callionymus altipinnis differs from C. sokonumeri in the filamentous first spine of the first dorsal fin, the shorter caudal fin filaments, and various color markings. The record of C. huguenini Bleeker (by Shen 1964) from Hong Kong is based on a misidenti- fied specimen of C. altipinnis (the specimen is now the holotype of C. altipinnis). EtyMoLoGy.—From the Latin altus, high, and pinna, fin, in reference to the unusually high second dorsal, anal, and first dorsal fins. Callionymus sokonumeri Kamohara, 1936 Callionvmus sokonumeri KAMOHARA, 1936:448, fig. 2 (Mi- mase Market); 1952:90, fig. 87 (Prov. Tosa); 1955:63, fig.; MATSUBARA 1955:713 (after Kamohara); OCHIAI, ARAGA, AND NAKAJIMA 1955:109-110, figs. 6-7, table 5 (Mimase and Owase); KAMOHARA 1964:90 (Kochi Pref.). D1AGNosis.—A Callionymus of the kaianus- group with a small branchial opening; short head (3.6—4.4 in SL); preopercular spine with a small antrorse and two larger curved points at dorsal side, and a short main tip; high second dorsal fin with a convex distal margin; no filament in first dorsal fin; anal fin with a distal black streak on each membrane; distal two-thirds of mem- branes of anal fin brown, including tips of fin rays; an irregular distal margin of caudal fin; and a very small black blotch on third membrane of first dorsal fin, extremely distal in position. DEscRIPTION.—Counts and proportions (see also Table 2): D, TV; D, viti,1; A viii,1; P, 18— 20 Pes. (C10. Body elongate and depressed. Head de- pressed, 3.6—4.4 in SL. Eye large, 2.1-2.9 in head. Pupil about 3.1 in eye diameter. Branchial opening about 3 in pupil. Preopercular spine with a short main tip, a small antrorse and two larger curved points at its dorsal side, and a large antrorse spine at its base (formula: | 1). Lateral line reaching from area behind eye to end of one of the median caudal fin rays (?); the line of the opposite side interconnected by a transverse branch across occipital region and 375 another across dorsal side of cauda! peduncle. Caudal peduncle length about 4.8 in SL, caudal peduncle depth 23.0 in SL. First spine of first dorsal fin lower than first ray of second dorsal fin, not filamentous. Second dorsal fin very high, with a convex distal margin. Anal fin begins on vertical through second ray of second dorsal fin. Distal shape of caudal fin irregular; two median caudal fin rays elongate, occasionally filamentous. Distal margin of pelvic fin convex; longest pelvic fin ray reaching to midbase of first membrane of anal fin when laid back. Pectoral fin reaching to fourth ray of sec- ond dorsal fin when laid back. Color in alcohol. Body olive-yellow above, whitish below. Back with both blackish spots and blotches, the former forming darkish rings. A row of several dark brown blotches along side of body. Pectoral fin base with a brown area. First dorsal fin gray, with three oblique dark bars; a black blotch distally on third membrane. Second dorsal fin gray, with several vertical dark streaks on membranes. Distal part of anal fin blackish, occasionally with a transverse streak distally on each membrane except the first. Caudal fin gray, mottled with blackish spots and small whitish blotches. Pectoral fin pale, the dorsal part mottled with dark spots. Pelvic fin gray, distal parts of fourth and fifth rays dark. DISTRIBUTION.—Known only from central Honshu, Japan (Pacific coast; see Fig. 2); col- lection depths not known. DiIscUSSION.—I unfortunately had no material of this species. The present description is com- piled from Kamohara (1936) and Ochiai, Araga, and Nakajima (1955). The differences between C. sokonumeri and C. altipinnis are discussed in the description of the latter species. Callionymus sokonumeri dif- fers from C. ochiaii in its higher second dorsal and anal fins, the shape of the caudal fin, a short- er main tip of the preopercular spine, and var- ious color markings. minimal Callionymus sp. Callionymus kaianus: (non Gunther, 1880) WONGRATANA 1968:58 (Gulf of Thailand). DiscussIOoN.—I did not examine material of the kaianus-group from the Gulf of Thailand. Wongratana’s (1968) paper records Callionymus kaianus, but gives no descriptions or figures. 376 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 14 Callionymus kaianus should not occur in the Gulf of Thailand. I have been unable to deter- mine if the specimens from that area belong to any of the known species of the kaianus-group, or if they represent yet another species. ACKNOWLEDGMENTS For discussion of problems, loan of speci- mens, information, or permission to examine specimens in their collections, I thank the fol- lowing persons: H.-J. Paepke (ZMB, East Ber- lin); P. J. P. Whitehead, A. Wheeler, and O. Crimmen (BMNH, London): A. Ben-Tuvia and D. Golani (HUJF, Jerusalem); K.-C. Au (MSL, Hong Kong); H. K. Larson (AMS, Sydney); J. P. Gosse (IRSN, Brussels); W. N. Eschmeyer, L. J. Dempster, T. Iwamoto, T. R. Roberts, P. M. Sonoda, M. Hearne, and G. Raabe (CAS, San Francisco), T. Iwai and T. Nakabo (FAKU, Kyoto), A. Kotthaus (Eppstein, W. Germany), H. Wilkens and R. Dohse (ZIM, Hamburg). W. N. Eschmeyer and T. Iwamoto (CAS, San Fran- cisco) also reviewed the manuscript. I am greatly obliged to P. Fourmanoir (OR- STOM, Nouméa) for the gift of a specimen of Callionymus moretonensis and a further callion- ymid fish specimen. I am also grateful to D. J. Stewart (FMNH, Chicago) who examined a specimen of Callionymus carebares from his collection for me. I thank O. von Frisch (NMB, Braunschweig) for accepting specimens loaned to me. A financial contribution for expenses dur- ing my visit to the California Academy of Sci- ences was provided by that organization. LITERATURE CITED Atcock, A. W. 1890. On the bathybial fishes collected in the Bay of Bengal during the season 1889-1890. Ann. Mag. Nat. Hist. (6)6:197—222. . 1898. Illustrations of the zoology of H.M. Indian sur- veying steamer “‘Investigator.”’ Pt. 4, Fishes. Calcutta: In- dian Museum. . 1899. A descriptive catalogue of the Indian deep-sea fishes in the Indian Museum, collected by the Royal Indian Marine Survey ship Investigator. Calcutta: Indian Museum. 211 pp. BEAUFORT, L. F. DE. 1951. The fishes of the Indo-Australian Archipelago, 9. Percomorphi (concluded), Blennioidea. Lei- den: E. J. Brill. 484 pp. BESEDNOV, L. N. 1968. Ichthyofauna of the Gulf of Tonkin (in Russian). Uchenye Zapiski Dal’nevostocnyi Universitet, Vladyvostok 15(2):47-85. BLEEKER, P. 1858. Vijfde bijdrage tot de kennis der ichthy- ologische fauna van Japan. Act. Soc. Sci. Indo-Neerl. 5:1— 12. Cuu, K. Y. 1957. A list of fishes from Pescadore Islands. Rep. Inst. Fish. Biol. Minist. Econ. Aff. Natl. Taiwan Univ. 1(2):14—22. Cuu, Y. T., AND OTHERS. 1962. Fishes of the South China Sea. [In Chinese] Peking. Pp. i-xxxvii, 1-1184, figs. 1-860. Fow er, H. W. 1928. The fishes of Oceania. Mem. B. P. Bishop Mus. 10:1—540, figs. 1-82, pls. 1-49. Fricke, R. 1980. Neue Fundorte und noch nicht beschrie- bene Geschlechtsunterschiede einiger Arten der Gattung Callionvmus (Pisces, Perciformes, Callionymidae), mit Be- merkungen zur Systematik innerhalb dieser Gattung und Beschreibung einer neuen Untergattung und einer neuen Art. Ann. Mus. Civ. Stor. Nat. “Giacomo Doria’ 83:57-105, figs. 1-14, pls. 1-9. . 198la. On a new species of the family Calliony- midae (Pisces, Perciformes, Callionymoidei), Callionymus stigmatopareius spec. nov. from Mozambique. J. Nat. Hist. 15(1): 161-167, figs. 1-2, tables 1-2. . 1981b. Revision of the genus Synchiropus (Tele- ostei: Callionymidae). Braunschweig: J. Cramer. 196 pp. GUNTHER, A. (C. L. G.). 1880. Report on the shore-fishes procured during the voyage of H.M.S. Challenger in the years 1873-1876. Rep. Sci. Results H.M.S. Challenger, Zool. 1(6): 1-82, pls. 1-32. Herre, A. W. C. T. 1953. A check-list of the fishes of the Philippines. U.S. Dept. Int. Fish Wildl. Serv., Res. Rep. 20: 1-977. JOHNSON, C. R. 1971. Revision of the callionymid fishes re- ferable to the genus Callionymus from Australian waters. Mem. Queens]. Mus. 16(1):103—-140. 1973. Biology of the dragonet Callionvmus kaianus moretonensis Johnson (Pisces, Callionymidae). Zool. J. Linn. Soc. 52:217—230. JORDAN, D. S., AND A. SEALE. 1905. The fishes of Samoa. Description of the species found in the archipelago, with a provisional check-list of the fishes of Oceania. Bull. U.S. Bur. Fish. 25:175—455, figs. 1-111, pls. 33-53. KAMOHARA, T. 1936. Two new deep-sea fishes from Japan. Annot. Zool. Japon., Tokyo 15(4):446—448, figs. 1-2. 1952. Revised descriptions of the offshore bottom fishes of province Tosa, Shikoku, Japan. Rep. Kochi Univ. Nat. Sci. (3):1—122, figs. 1-100. 1955. Coloured illustrations of the fishes of Japan. Osaka. 1964. Revised catalogue of fishes of Kochi Prefec- ture, Japan. Rep. Usa Mar. Biol. Stn. 11(1):1-99. KoTTHAus, A. 1977. Fische des Indischen Ozeans. A. Sys- tematischer Teil XIX: Percomorphi (9). ‘‘Meteor’’ For- schungsergebn., Reihe D(25):24—44, figs. 406-428. LINNAEuS, C. 1758. Systema naturae .... 1. Regnum ani- male. 824 pp. Masuba, H., C. ARAGA, AND T. YOSHINO. 1975. Coastal fishes of southern Japan. Tokyo: Tokai Univ. Press. 379 pp., 143 pls. MATSUBARA, K. 1955. Fish morphology, and hierarchy. Pt. 1. Tokyo: Ishizaki-Shoten. 789 pp. [2nd. ed., Tokyo 1971.] Munro, I. S. R. 1958. The fishes of the New Guinea region. Territ. Papua and New Guinea, Fish. Bull. 1:97—369. [Re- printed from Papua and New Guinea Agric. J. 10(4):97- 369.] NAKABO, T. 1979. A new and two rare species of the genus Callionvmus (Callionymidae) from the western Indian Ocean. Japan. J. Ichthyol. 26(3):231—237, figs. 1-4, table 1. FRICKE: KAIANUS-GROUP OF GENUS CALLIONYMUS NorMaAn, J. R. 1939. Fishes. Sci. Rep. John Murray Exped. 7(1):1-116, figs. 1-41. Ocuial, A., C. ARAGA, AND M. NAKAJIMA. 1955. A revision of the dragonets referable to the genus Callionvmus found in the waters of Japan. Publ. Seto Mar. Biol. Lab. 5(1):95- 132. Peters, W. C. H. 1876. Uebersicht der wahrend der von 1874-1876 unter dem Commando des Hrn. Capitan z.S. Freiherrn von Schleinitz ausgefiihrten Reise der S.M.S. ‘“‘Gazelle’’> gesammelten und von der Kaiserlichen Admi- ralitat der KGniglichen Akademie der Wissenschaften tiber- sandten Fische. Monatsber. Dtsch. Akad. Wiss. Berl. 1876:831-854. REGAN, C. T. 1906. On fishes from the Persian Gulf, the Sea of Oman, and Karachi, collected by Mr. F. W. Townsend. J. Bombay Nat. Hist. Soc. 16:318—333, pls. A-C. . 1908. The Percy Sladen Trust Expedition to the In- dian Ocean in 1905 under the leadership of Mr. J. Stanley Gardiner. Report on the marine fishes collected by Mr. J. Stanley Gardiner in the Indian Ocean. Trans. Linn. Soc. London, Zool. (2)12(3):217—255, pls. 23-32. RICHARDSON, J. 1844-1848. The zoology of the voyage of H.M.S. Erebus & Terror, under the command of Captain 377 Sir James Clark Ross, R.N., F.R.S., during the vears 1839 to 1843. Fishes. London. 139 pp., 60 pls. SHEN, S.-C. 1964. A list of the fishes from Hong Kong. Pt. 1. Quart. J. Taiwan Mus. (Taipei) 17: 193-208 SmitH, J. L. B. 1963. Fishes of the families Draconettidae and Callionymidae from the Red Sea and western Indian Ocean. Rhodes Univ., Dep. Ichthyol., Ichthyol. Bull. (28):547-S64, figs. 1-8, pls. 83-86. Suwarps1. 1965. Notes on the genus Callionymus (Pisces, Callionymidae), mostly from Indonesian waters, with the description of three new species and a new subspecies. Vi- densk. Medd. Dansk Naturhist. Foren. 128:303-323, pls. 46-48. VALENCIENNES, A. in Cuvier, G., and A. Valenciennes. 1837. Histoire naturelle des poissons 12. Paris. i-xxiv + 507 pp. Weser, M. 1913. Die Fische der Siboga-Expedition. Siboga Exped. 57:1-719. Leiden. WONGRATANA, T. 1968. Check list of fishes caught during the trawl surveys in the Gulf of Thailand and off the east coast of the Malay peninsula. Contrib. Mar. Fish. Lab. Bangkok 13:1-96. CALIFORNIA ACADEMY OF SCIENCES Golden Gate Park San Francisco, California 94118 wile? 0 ; i _ We. He em ay sens! Talia willed) Cay, ee ee Ss Rie ALA aa er" Ree Soe Pree ys prd)) 7; aia an) Th pet sae Lhe LF te. Bite «jee —t) wedi > & ~ 1 atts . pists “gti ie ' , Ww ure ie Matas Sutin iT tos, ft f= he ®> ' AG & - ¥ ' Je 5 ihtecl * twisieon “oy Twit ie iA: ee ve Boe ay Pes s «0s y Perurat tt (eal oo Ta Use, aD). |: pol ;) ul? oO grease 1 & OnE eee ty. r ——o waht 2 tw io her Ae idea fee ie © eg ial im, 3 eres ‘> jes Wi Ne —_- [ND SAUITH GAS ——- , — ‘s 7 4 é ' ee | ro | 6 | " 7 ry ' < oe ©) ( Paty Pepi ve hi sh re ate eat Rect, 1 iSties! ove _ ons Dare gre: Te) as == ss Le + PASTORIS, REPRINT MtA TION GF MOMATENT rir, PROPS / PROCEEDINGS : OF THE CALIFORNIA ACADEMY O Vol. 42, No. 15, pp. 379-407, 18 figs., 4 tables. =, ac Re | wi Marine Biological Laboratory | LIBRAR | NOV 19 1981 SWE OSS ole. Mass October 26, 1981 a Ree 3 ae DISTRIBUTION, REPRODUCTIVE ANATOMY, AND VARIATION OF MONADENIA TROGLODYTES HANNA AND SMITH (GASTROPODA: PULMONATA) WITH THE PROPOSAL OF A NEW SUBGENUS By Barry Roth Department of Invertebrate Zoology, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118 ABSTRACT: Shastelix, new subgenus of Monadenia (type-species, M. troglodytes troglodytes) is proposed, based principally on genital characters. The new subgenus is mostly confined to the Klamath Mountains of California and is parapatric with Monadenia, sensu stricto. Monadenia troglodytes is common in a limited area of Shasta County, associated with limestone terrane. Three subspecies are recognized based on morphometry and geography—M©. t. troglodytes along the McCloud River arm of Shasta Lake; M. t. wintu, new subspecies, between the Pit River and Squaw Creek, with one disjunct, outlying population south of Shasta Lake; and M. t. chaceana (formerly ranked as a distinct species) near the confluence of the Shasta and Klamath rivers in Siskiyou County. Sympatric Monadenia churchi and M. troglodytes differ little in reproductive anatomy; elab- orations of the female genitalia are probably not important in species recognition or reproductive isolation. A phylogenetic hypothesis and evolutionary scenario for the three subgenera of Monadenia are presented. INTRODUCTION The western North American helicacean snail genus Monadenia Pilsbry, 1895, includes con- spicuous species that have long attracted the attention of malacologists. Its type-species, He- lix fidelis Gray, 1834, was the first land mollusk described from the Pacific coast. It includes the most northern helicacean species—and some of the most northern large land snails—on the North American continent. Monadenia is one of the few temperate genera of snails with an elaborate color pattern (Comfort 1951), and M. fidelis is the only snail in the far west with a dramatic polymorphism of shell color and band- ing (Roth in press). _ Pilsbry (1939) and Berry (1940a) divided Mon- adenia into two groups of species, based on re- productive anatomy and shell characters—Mon- adenia, sensu stricto, and Corynadenia Berry, 1940. The range of the genus extends from southern Alaska to central California, principal- ly west of the Cascade Range but penetrating inland along major river valleys. One branch reaches south along the east side of the Sacra- mento Valley; the other follows the southern Klamath Mountains and Coast Ranges to the San Francisco Bay region (Figure 1). The sub- genus Corynadenia exists as an apparently dis- junct group of species on the west side of the Sierra Nevada. Roth (1975) showed that Mon- adenia churchi Hanna and Smith, 1933, which occupies a fairly wide range around the north end of the Sacramento Valley, and which Pilsbry (1939) and Berry (1940a) had grouped with the [379] 380 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 15 = A Gs Ba y ‘ L 204, as rynadenia Wi Monadenia, ages: FiGure |. Distribution of the subgenera of Monadenia in California. Diagonal hachure, generalized range of Monaden- ia, sensu stricto; stippling, in Klamath Mountains region, Shastelix, in Sierra Nevada, Corvnadenia. Dashed line rep- resents 36 C mean maximum July isotherm; dot-dashed line, —4 C mean minimum January isotherm (thermal data after Elford 1970). Sierran subgenus, combines anatomic and shell characters of Monadenia, s.s., and Corynaden- ia. A number of other species from northern California described by Berry (1940b) have not yet been dissected or assigned unequivocally to subgenus. Parapatry and possible sympatry of species have been demonstrated (Roth and Eng 1980; and herein), and it is clear that the zoo- geography of Monadenia is more complex than earlier authors imagined. Monadenia is ecologically diverse, compris- ing species of rock crevices and rockslides, oth- ers that inhabit deep leafmold, semi-arboreal forms, and snails that crawl out exposed on low plant cover. The relations between habit and habitat, on the one hand, and morphology and variation, on the other, are important for under- standing the evolutionary history of the group. This paper is one of a projected series of studies aimed at elucidating those relationships. Monadenia troglodytes Hanna and Smith, 1933, was described from fossil shells of pre- sumed Pleistocene age collected in Samwel Cave, Shasta County, California (sec. 5, 1535 N, R. 3 W, Mount Diablo Base and Meridian, USGS Bollibokka Mountain Quadrangle). Other shells of the species, likewise interpreted as Pleistocene, were reported from Potter Creek Cave (sec. 23, T. 34 N, R. 4 W, MDB&M, USGS Lamoine Quadrangle), Shasta County (Hanna and Smith 1933). Empty shells were lat- er found by Stanford University speleologists a short distance outside Samwel Cave (Smith 1957), and in 1963 the species was discovered alive in the same general area (Walton 1970). Smith (1970) and Roth (1972a, 1972b) cited M. troglodytes as rare and of limited distribution. In May 1973 the late Allyn G. Smith of the California Academy of Sciences found the species at a new locality, in limestone rockslides near Ellery Creek (SE!4 sec. 6, T. 35 N, R. 3 W, MDB&M, Bollibokka Mountain Quadran- gle), west of the McCloud River arm of Shasta Lake (Fig. 2). He collected many empty shells in various states of preservation and one living specimen. Since then, additional collections, in- cluding a substantial amount of material secured independently by S. E. Hirschfeld, D. C. Ru- dolph, and R. L. Seib, indicate that the species is fairly common in a limited area in Shasta County and is strongly associated with lime- stone terrane. Dissections of the reproductive system show that M. troglodytes, along with M. churchi, be- longs to a new subgenus, which is named herein. Selected shell characters were measured to ana- lyze shell variability. A new subspecies is de- scribed, distinguished from typical M. troglo- dytes by details of color, shell microsculpture, _and morphometry. Monadenia chaceana Berry (1940b) is similar in general shell character but differs consistently in certain shell measure- ments and is regarded as a third subspecies. One other species of Monadenia occurs within the range of M. troglodytes, permitting a consider- ation of species criteria within the genus. An hypothesis of phylogenetic relationships within Monadenia is presented. The use of a trinomial to designate the nomi- nate subspecies—a convention sparingly ob- served in American land malacology, but one necessary to distinguish the subspecies from the species sensu lato—is here introduced for Mon- adenia troglodytes troglodytes. ROTH: DISTRIBUTION, ANATOMY, VARIATION OF MONADENIA TROGLODYTES 381 FIGURE 2. The following institutional abbreviations are employed: AMNH—Ame rican Museum of Natural History ANSP—Academy of Natural Sciences, Phila- delphia CAS—California Academy of Sciences, Depart- ment of Invertebrate Zoology CASGTC—California Academy of Sciences, Geology Type Collection, Department of Ge- ology FMNH—Field Museum of Natural History LACM—Los Angeles County Museum of Nat- ural History SSB—Private collection of S. Stillman Berry, Redlands, California SUPTC—Stanford University Paleontological Type Collection, now in Department of Ge- ology, California Academy of Sciences UCMP—Museum of Paleontology, University of California, Berkeley USNM—United States National Museum of Natural History Gilman Road crossing of Ellery Creek (Locality 26); typical Monadenia troglodytes habitat in brush-covered talus at foot of prominent limestone outcrop. Monadenia Pilsbry Monadenia PitsBry, 1895:198.—PiLsBRry 1939:31-35. Type-sPecies: Helix fidelis Gray, 1834, by original designa- tion. Shastelix, new subgenus TyPe-sPECcIES: Monadenia troglodytes troglodytes Hanna and Smith, 1933. DraGnosis.—Monadenia with large, globose atrium; mucus gland much longer than dart sac, its lower part adnate to atrium; penial retractor inserted near middle of epiphallus; flagellum (epiphallic caecum) substantially longer than pe- nis plus epiphallus and borne in a series of he- lical coils. Penis sessile on atrium, not invagi- nated into it. Spermatophore helically coiled (?). Shell of moderate size for the genus, smooth or granulose, protoconch sculpture of minute, somewhat confluent granules, tending to align in diagonal series. The genitalia of Monadenia troglodytes (Figs. 4-6) differ most obviously from those of species 382 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 15 TABLE 1. CHARACTER COMPLEXES IN THE SUBGENERA OF Monadenia. Characters Monadenia, Ss.s. Shastelix Corvnadenia Atrium large and globose (+), or small and narrow +: - () Mucus gland many times longer than (+), or about +f of = equal to (—), dart sac Lower part of mucus gland adnate to (+), or free from = + = (—), atrium Dart sac 5 mm or less in length (+), or longer than = +r = 5 mm (—) Flagellum =1.5 times length of (+), or about as long — + + as (—), penis plus epiphallus Flagellum (and spermatophore) helically coiled (+), = ate $F or straight (—) Basal chamber of penis invaginated into (+), or 3 = + sessile on (—), atrium Penial retractor inserted near middle (+), or on =535 + + distal third (—), of epiphallus of the nominate subgenus (M. fidelis, M. infu- mata (Gould, 1855), M. setosa Talmadge, 1952) in the long, helically coiled flagellum. The fla- gellum in Monadenia, sensu stricto, is thick and straight or simply curved and about as long as the penis plus epiphallus (Pilsbry 1939:figs. ISA, B, M. fidelis; Roth and Eng 1980:fig. 3, M. se- tosa; the genitalia of M. infumata are similar). The flagellum is the organ which secretes the spermatophore; the spermatophore of M. fidelis (Webb 1952:fig. 8A) is straight. In the Sierran Monadenia (Corynadenia) hirsuta Pilsbry, 1927, a species with helically coiled flagellum, the spermatophore is coiled like a corkscrew. It seems probable, therefore, that species of Shas- telix also secrete coiled spermatophores. Both Shastelix and Monadenia, s.s., have a large, globose atrium. (I follow Pilsbry [1939], Berry [1940a] and other authors in using the term ‘‘atrium’’ for the large, saccular elabora- tion of the lower genitalia upon which the penis, dart sac, and vagina insert and which when everted forms the copulatory pad or disk. This organ is mainly developed above the insertion of the penis and is homologous to the lower part of the vagina in other helicoid genera. In these other genera the term ‘‘atrium’”’ is convention- ally restricted to the common passage to the ex- ternal genital pore below the insertion of the pe- nis.) In Shastelix the lower ductlike portion of the mucus gland is adnate to the atrium, whereas in Monadenia, s.s., it runs along the surface of the atrium but is not fused to it. The dart sac in Shastelix is smaller than that in the nominate subgenus. In Monadenia, s.s., the basal part of the penis is invaginated into the wall of the atrium, which clasps it like a collar; in Shastelix the basal part of the penis is sessile on the atrium. In the subgenus Corynadenia Berry, 1940a (type-species, Helix hillebrandi Newcomb, 1864; see Pilsbry 1939:fig. 15C), the atrium is smaller, narrow and elongate, and the mucus gland is shorter or very slightly longer than the dart sac. The flagellum is longer than the penis plus epiphallus and, at least in some species, helically coiled. The basal part of the penis is invaginated into the wall of the atrium, as in Monadenia, 8.8. Character complexes differentiating the three subgenera are summarized in Table 1. Monadenia churchi also belongs to Shastelix and resembles M. troglodytes in genitalia (Fig. 8) and protoconch sculpture. It is probable that some undissected species from the Klamath Mountains (particularly M. cristulata Berry, 1940, and M. marmarotis Berry, 1940) will also prove to belong to Shastelix. The known range of the subgenus (Fig. 1) extends from Butte County on the south and east, around the north ROTH: DISTRIBUTION, ANATOMY, VARIATION OF MONADENIA TROGLODYTES 383 FIGURE 3. @ Monadenia churchi e M. troglodytes troglodytes (el M. troglodytes wintu A M. churchi and M. troglodytes Distribution of Monadenia species in vicinity of Shasta Lake, Shasta County, Calif. Stippling indicates surface outcrops of limestone. Open triangles denote peaks of more than 1000 m elevation. Geology after: Diller (1906); Kinkel et al. (1956); Coogan (1960); Albers and Robertson (1961); Demirmen and Harbaugh (1965); Irwin and Galanis (1976). end of the Sacramento Valley, to near Burnt Ranch, Trinity County, on the west, and the Shasta River—Klamath River confluence on the north. Monadenia churchi and M. troglodytes are sympatric in the vicinity of Shasta Lake. In the drainage of the Trinity River, Shastelix is parapatric with Monadenia, s.s. There M. fi- delis and M. setosa tend to occur in riparian woodland close to streambeds, while M. churchi occurs on drier, more exposed slopes and in rockslides. Only the subgenus Corynadenia is known to occur in the Sierra Nevada. The subgenus name is derived from the place- name Shasta—lake, mountain, river, and county—plus the Greek helix, a spiral, hence a snail. Its gender is feminine. DISTRIBUTION, ANATOMY, AND GEOGRAPHIC VARIATION OF Monadenia troglodytes The material now available, summarized in the Appendix, makes it clear that Monadenia troglodytes is not only extant (Walton 1970) but occupies a considerable range in the vicinity of Shasta Lake (Fig. 3). At several of the localities, living individuals were found to be common. The ranges of M. t. troglodytes and a second subspecies described herein lie within the ‘‘Eastern Klamath Belt,’ the easternmost of several concentric lithic belts that constitute the Klamath Mountains physiographic province of northwest California and southwest Oregon (Ir- win 1972). The Klamath Mountains are a rugged 384 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 15 mountain region including clusters of high peaks 1800-2700 m in altitude. Peaks and ridgecrests of 1200-1350 m occur in the area mapped in Fig- ure 3. The canyons of the Sacramento, Mc- Cloud, and Pit rivers and Squaw Creek dominate the area’s topography. Shasta Lake is an artifi- cial impoundment of these rivers; their courses prior to damming may be seen on pre-1940 to- pographic maps. The spillway of Shasta Dam is at an elevation of 325 m, so that local relief of 1000 m and more is present in the area under study. Southward from Shasta Lake, the Sac- ramento River descends through foothills to the broad, alluviated floor of the Sacramento Val- ley. The Sacramento River—Trinity River wa- tershed—closely coinciding with the Shasta County-Trinity County boundary and dividing the north coast drainage from the interior, Great Valley, drainage—is located a few kilometers off the west margin of the map. North, east, and southeast of the map area lie portions of the Cascade Range physiographic province (Wahr- haftig and Birman 1965); due north is the Pleis- tocene stratovolcano Mount Shasta, 4300 m in altitude. The Klamath Mountains are noted for the di- versity of their flora and the number of local endemic plant species (Whittaker 1961; Axelrod 1976). In the region under study, the vegetation is largely northern yellow pine forest with some Sierran montane forest (Kiichler 1977), partic- ularly between the Pit and McCloud rivers. Lower elevations and exposed slopes are cov- ered by a growth of scrub oak and chaparral. The land at higher elevations was originally densely timbered with coniferous forest, but in many areas logging, fires, and the effects of smelter smoke have removed the original tim- ber, so that second-growth forest and brush are widespread. Manzanita (Arctostaphylos) is the dominant brush plant. The normal annual precipitation is between 120 and 180 cm, more than 90 percent of it falling in the months October through April (Elford 1970; Major 1977). Mean monthly temperatures at Shasta Dam range from around 5 C in January to about 25 C in July (Major 1977). Daytime tem- peratures in excess of 40 C are not uncommon from June through September. Local microcli- matic variation is considerable. Geology of the region is complex, including intrusive and extrusive, sedimentary, and meta- morphic rock suites, of ages from Paleozoic to Quaternary. Of particular interest are bodies of limestone that crop out discontinuously in three principal areas. These are finely crystalline, re- sistant limestones that often form prominent outcrops (Kinkel et al. 1956:fig. 20). Local to- pographic names such as ‘‘Gray Rocks’ (La- moine and Bollibokka Mountain quadrangles), ‘*Limerock Gulch,’ ‘‘Marble Creek,’ and ‘*Up- per and Lower Limestone Valley Creeks’’ (La- moine Quadrangle) acknowledge their presence. All samples of Monadenia troglodytes for which adequate data are available were taken on or adjacent to these limestone areas (Fig. 3). The localities fall into two groups: those associated with the Triassic Hosselkus limestone in the vi- cinity of Brock Mountain, between the Pit River and Squaw Creek arms of Shasta Lake; and those on or near the Permian McCloud lime- stone, along the McCloud River arm of the Lake. The type-locality, Samwel Cave, belongs to the latter group; so does Locality 45, south of the Pit River arm, the southernmost definite site for M. troglodytes. (A bleached, broken shell from somewhat farther south—Loc. 46, in the Anderson Quadrangle—is similar but cannot be assigned unequivocally to species.) Locality 35 is from a limestone quarry, evidently in one of the limestone lenses in the predominantly clastic Pit Formation (Albers and Robertson 1961). Limestones in the Shasta Lake region are dis- continuous because of faulting and erosion. The McCloud and Brock Mountain locality groups are separated by approximately 10 km of non- — limestone terrane—chiefly volcanics, pyroclas- tics, and mudstones. Squaw Creek, its canyon, and a watershed of 830 m minimum elevation also stand between the two groups. Morphologic distinctions between the Brock Mountain and McCloud River herds, discussed below, indicate that a certain amount of genetic isolation exists as well. Localities yielding M. troglodytes range in elevation from about 330 to 760 m, so it does not seem likely that the altitude of the watershed alone constitutes an isolating factor. Exposure and seasonal lack of moisture on ridgetops may restrict the snails’ mobility, but since some col- lections of M. troglodytes were made in zones of high insolation, with snails active in shaded spots even during July, limestone substrate seems a more likely limiting factor. ROTH: DISTRIBUTION, ANATOMY, VARIATION OF MONADENIA TROGLODYTES 385 Cooper (1869) remarked on the strong asso- ciation of Monadenia with carbonate terrane in the Sierra Nevada; and Pilsbry (1939) surmised that discontinuity of limestone substrata was in- volved in the formation of local races. Many other records exist of land mollusks whose dis- tribution is more or less tied to areas of high calcium availability (e.g., Boycott 1934; Twee- die 1961; Heller 1975). To the west, limestone outcrops of the De- vonian Kennett Formation in the Mammoth Buttes—Backbone Ridge area west of the Sac- ramento River arm of Shasta Lake have yielded Monadenia churchi but not M. troglodytes. Monadenia churchi also occurs in the McCloud River and Brock Mountain areas. At a minimum of three stations it is sympatric or parapatric with M. troglodytes and was received in the same samples. M. churchi is not restricted to limestone substrata; it has been found elsewhere in lava rockslides (type lot; Hanna and Smith 1933) and coniferous forest debris (Roth and Eng 1980). At each locality on limestone where both M. troglodytes and M. churchi were taken to- gether, M. troglodytes is the more common species. The implications of the sympatry of M. churchi and M. troglodytes for classification of Monadenia are discussed below. No specimens from Shasta County referable to the Sierran Monadenia (Corynadenia) mor- monum (Pfeiffer, 1857) have been found in the course of this study, and it appears that all such records in the literature are based on misiden- tifications of either M. troglodytes or M. chur- chi. (For example: *‘The most northern locality for mormonum now known is at Shasta, Cal., lat. 41° (nearly), alt. 1160 feet, where in the vol- canic region Dr. Yates found a very few stunted specimens with but five and a half whorls and the bristle-granulations of the young very strongly developed’’ [Cooper 1879:285]. ‘‘In Shasta County, far north of the localities men- tioned, a race of mormonum has been found in the Upper Sonoran Zone at and near the junc- tion of the Pitt [sic] with the Sacramento river (Brewer, Gabb). They agree with the typical form in the absence or extreme faintness of spi- ral striae and in coloration; the shell is smaller and the spire generally higher. They are within the area of M. churchi’’ [Pilsbry 1939:56].) A lot collected in the nineteenth century, UCMP 2491, ex D. O. Mills collection no. 290, labeled ‘‘Shasta County, Calif.”’ and formerly identified as M. mormonum, is probably the same as the Shasta County shells referred to by Pilsbry (1939:56). It is M. troglodytes, similar to those from the Brock Mountain area. The original de- scription of M. troglodytes was based on sam- ples with very low spire index (H/D = 0.411- 0.500); the figured holotype is nearly planispiral. In the samples now at hand, intermediate ex- amples connect these very flat shells with the higher-spired, mormonum-like specimens. ANATOMY.—Specimens were prepared for anatomical study by drowning followed by transfer in stages to 70% ethanol. Specimens of M. troglodytes from the following localities were dissected: 10, 26, 28, 34, 39, 43. Except as noted, the data given apply to all individuals ex- amined. Body grayish tan, shaded darker on dorsum; light buff mid-dorsal stripe; sole light buff. Man- tle over lung translucent buff, with gray dendri- tic pencilling occupying 0—35% (usually about 20%) of surface. (All degrees of mantle pigmen- tation present in sample from Loc. 10; other samples less variable.) Mantle collar colored like body. External genital pore a vertical 1-2 mm slit often showing white rim. Right ocular retractor running between male and female systems, passing over crook at penis- epiphallus junction. Genitalia as in Figures 4—6. Penis stout, basal chamber separated by crook from upper chamber, adnate to atrium but not invaginated therein; upper chamber thin-walled, finely ridged internally, containing cylindrical to ovate-conic, slightly rugulose verge 3—3.5 mm long. Tip of verge with slitlike lateral meatus on anterior edge, dorsal facet concavely beveled (Fig. 4b). (End of verge of specimens from Loc. 43 blunt, as broad or broader than stalk, with flaplike expansion of tip opposite meatus.) Wall of upper penial chamber bearing a single large pilaster that fits against beveled facet of verge and extends into basal chamber. Epiphallus from % to % as thick as penis at their junction. Penial retractor (originating on floor of lung) long, narrow, slightly expanded just before in- sertion on medial part of epiphallus. Flagellum (epiphallic caecum) longer than penis plus epi- phallus, as thick as epiphallus, borne in 4 to 7 helical coils; distal end tapering to fine point. Vas deferens with 2—3 convolutions where it passes under atrium. Spermatheca (bursa cop- 386 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 15 FIGURES 4-5. Genitalia of Monadenia troglodytes. Figs. 4a—b. M. t. troglodytes; (a.) dorsal aspect, Loc. 28, atrium rotated 90° downward with respect to oviduct; (b.) detail of verge, Loc. 10. Figs. Sa-b. M. t. wintu, n. subsp., holotype CAS 018431, Loc. 34; (a.) dorsal aspect; (b.) ventral aspect. Abbreviations: a—atrium; ag—albumen gland; ds—dart sac; ep—epiphallus; fi—flagellum; fo—free oviduct; gp—external genital pore; hd—hermaphroditic duct; mg—mucus gland; pb—basal chamber of penis; pr—penial retractor muscle; pu—upper chamber of penis; sp—spermatheca; spd—duct of spermatheca; s—talon (fer- tilization pouch); vd—vas deferens; ve—verge. ulatrix) globose; spermathecal duct long, straight, unbranched, capacious for lower three-fourths, narrower below spermatheca. Dart sac small to moderate-sized, sessile at convergence of two atrial crura which, when everted, form a copu- latory pad or disk (Fig. 7); sac containing a 2-mm, tubular, calcareous dart. Mucus gland large, bent near middle, lower portion adnate to atrium and running along lower edge of superior atrial crus to insertion at base of dart sac. Measurements of selected organs are given in Table 2. All show considerable variation. There appears to be no systematic difference between samples except that those from the Brock Mountain area tend to have more coils to the epiphallus. ROTH: DISTRIBUTION, ANATOMY, VARIATION OF MONADENIA TROGLODYTES 387 TABLE 2. in parentheses). LENGTHS (in mm) OF SELECTED ORGANS OF Monadenia troglodytes AND M. churchi (range, with M. t. troglodytes Loc. (n) 10 (6) 43 (5) Free portion of mucus gland 16.0-21.2 10.9-21.0 (18.82) (17.12) Dart sac 2.6-4.5 2.1—2.4 (3.18) (2.33) No. of coils of flagellum 3.5-5 3.5=5 (4.13) (4) Free (upper) chamber of penis 3.3-4.8 3.44.7 (3.55) (3.84) Epiphallus 5.6-8.7 5.7-8.6 (7.93) (6.72) Base of epiphallus to insertion 2.6-4.9 2.9-4.1 of penial retractor (4.07) (3.5) In their exserted state the atrial crura form a copulatory pad (Fig. 7). The female genital pore is located on the bottom of the pocket between the two crura. The male opening, through which the verge is exserted, is located near the middle of the forward edge of the anterior crus. The pore of the dart sac is at the convergence of the crura; further eversion of the dart apparatus was not observed. The lower genitalia are enveloped in thin tis- sue that binds the male system to the atrium and in some specimens forms a collar around the penis just below its junction with the epiphallus. The jaw is crescentic, golden brown, with four to six major ribs that denticulate both margins. From one to five smaller intercalary ribs are present on some specimens. The radula of a specimen from Locality 28 has 41 teeth in a half row. The central tooth is some- what wider than in M. fidelis (see Pilsbry 1939:fig. 1ISE), the mesocone as long as the basal plate. On the fourteenth tooth an endocone ap- pears, and on the fifteenth an ectocone. The marginal teeth are somewhat spatulate, the cleft between mesocone and endocone deep, and both mesocone and endocone are sometimes bifid or trifid. The outer marginals are very short. SHELL CHARACTERS AND VARIATION.—Han- na and Smith (1933) described the protoconch of Monadenia troglodytes as consisting of 112 whorls marked by wavy radiating riblets. In a sample mean M. t. wintu M. churchi Mammoth 34 (3) 45 (1) Butte (3) 29 (1) 16.1-30.0 5).9/ IS 78a 173 (21.0) (16.77) 72S '=3)./! 725\ N27) 1.9 (2.87) (2.23) 37) 5 35—4 6 (6) (3.67) 3.7-4.6 4.0 3.5—35-0 3.5 (4.23) (3.6) 8.4-8.9 8.1 2—025 6.6 (8.63) (5.63) 3.5—4.7 4.1 Dae—Bee, 2.6 (4.03) (27) group of 19 M. t. troglodytes hatched in captiv- ity from eggs laid by an individual from Locality 10 (slope below Samwel Cave), the protoconch consists of 1.5-1.8 whorls (counting by the method of Diver 1932) and measures 2.8—3.5 mm in diameter. The extreme nuclear tip is smooth. Thereafter, sculpture of the protoconch consists of minute granules, round and somewhat con- fluent, irregularly disposed at first but after about one half whorl tending to form wavy, ra- dial riblets. Those just outboard of the suture retain this character nearly to the end of the em- bryonic shell. Elsewhere on the top surface the granules become discrete and spirally elongated and tend to align in oblique, protractive and re- tractive series. Below the periphery the granules are discrete, spirally elongated, sometimes ar- ranged in diagonal series but more often scat- tered. All other M. troglodytes in which the pro- toconchs are not abraded show essentially the same sculpture. Slight erosion sometimes em- phasizes the effect of wavy radiating riblets, as described by Hanna and Smith for the type lot. The embryonic sculpture is very similar to that of M. fidelis (see Pilsbry 1939). The onset of neanic growth is marked by an abrupt enlargement of whorl diameter. The in- tensity of surface granulation on early neanic whorls is less than that on the protoconch; the granules are sometimes obscure or localized in a few patches. Irregular, axial growth ridges are the dominant sculptural feature. Between these 388 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 15 9a 9b 9c FiGures 6-9. Figs. 6a—b. Genitalia of Monadenia troglodytes wintu, n. subsp., Loc. 45; (a.) dorsal aspect; (b.) ventral aspect, atrium partially everted. Fig. 7. Copulatory pad of M. 1. wintu. Figs. 8a—b. Genitalia of Monadenia churchi, Loc. 29. (a.) dorsal aspect; (b.) detail of verge. Figs. 9a—c. Ontogenetic shape change in Monadenia troglodytes troglodytes, Loc. 10. Scale line = 5 mm for Fig. 9a, 10 mm for Figs. 9b, 9c. Abbreviations as in Figs. 4—S. ROTH: DISTRIBUTION, ANATOMY, VARIATION OF MONADENIA TROGLODYTES 389 ridges there is an extremely fine, wrinklelike, parallel lineolation, particularly evident on the base. Closely set, wavy, spiral striation appears on the base, usually by the end of the first post- nuclear whorl. These striae are irregular in both strength and spacing. Variably expressed, these elements—granu- lation, growth rugae, wrinkle-lineolation, and spiral striae—constitute the microsculpture of adult shells. When not pitted or breached by erosion, the periostracum is smooth and glossy; in adult shells it is often scratched and abraded on base and spire. Fine, wavy, spiral striation on the base is apparently standard but frequently removed by abrasion. On some shells, stronger, incised striae occur on the shoulder of the last whorl and may be prominent behind the lip. After about the three-whorl stage, granulation is rare below the shoulder. It persists on the spire for a varying number of whorls, usually becoming sparser with ontogeny, the granules becoming lower, more elongate, and less clearly defined until, in some cases, they finally merge into the general undulation of the periostracal surface. In most instances, however, granula- tion stops at or near a growth rest, and when growth is resumed the granules are few, irreg- ular, or absent. The last granulose whorl varies from locality to locality (Fig. 10), and there is a half-whorl to two-whorl range in all samples large enough for consideration. The means of the last granulose whorl increase in populations from north to south along the McCloud River. (Far to the north, M. t. chaceana has a mean of 1.83 gran- ulose whorls, seemingly continuing the cline.) Variation in the Brock Mountain herd is less obviously clinal: the westernmost samples, Lo- calities 34 and 35, have few granulose whorls (x = 2.38 and 2.10, respectively); the few mea- surable shells from the more eastern localities 33, 38, 40, and 41 are in the 2.5—3.5 whorl range; and Locality 39, situated geographically amid the other eastern localities, has a predominance of shells pustulose at 4.5 whorls and beyond (x = 4.17). Locality 45, south of Shasta Lake, also has populations whose granulation persists onto the later whorls (* = 3.73). The difference in number of granulose whorls between the McCloud River and Brock Mountain locality groups is highly significant (P < 0.001). In hatchlings, the protoconch is golden tan; MSCLOUD R. LOC. 17,19 n=2I | X= 250 % OF SAMPLE 1 2 3 4 WHORLS LOC 33,38,40,41 n=4 Kies CHACEANA Ficure 10. Histograms of last granulose whorl (to the nearest half whorl) in Monadenia troglodytes samples from along the McCloud River (M. 1. troglodytes), Brock Mountain region and Loc. 45 south of Shasta Lake (M. f. wintu, n. subsp.), and Siskiyou County (M. t. chaceana). some show a faint reddish-brown spiral band at the periphery of the last half whorl. From the first, the teleoconch shell material is lighter tan than the protoconch. Beginning with the first neanic whorl, narrow whitish zones border the reddish-brown band above and below, the lower zone seeming to appear earlier than the upper. Gradually the narrow band becomes darker and more prominent, partly by contrast with the in- creasing whitish opacity of the rest of the shell. The advancing suture partly or wholly obscures the band. olodytes 18 vari- within a sample. Color of adult Monadeni able, although fairly unifor 390 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 15 The basic shell material is white, covered in fresh, uneroded specimens with a transparent, yellowish periostracum. All fresh specimens have a dark reddish-brown band (chestnut of Rayner [1970]; Munsell midpoint 9.5R/2.3/3.8) just above the periphery, ranging from 0.5 to about 1.5 mm wide on the body whorl. Bands wider than | mm occur mainly in the eastern, Brock Mountain, samples and at Locality 45 south of Shasta Lake. Unpigmented zones, either white or showing only the yellowish color of the periostracum, border the dark suprape- ripheral band above and below. These zones are usually narrower than the dark band, but in some instances they are equally wide or wider. The lower zone is commonly wider than the up- per; on shells with a light-colored base, its an- terior edge may be indistinct. Color of the base is generally uniform, some- times slightly darker toward the periphery. The color ranges from nearly white (even with peri- ostracum intact) to a medium reddish brown (rust of Rayner [1970]; Munsell midpoint 9.0R/ 4.3/8.0), with little variation within a sample. The lightest bases occur in samples from Lo- calities 35, 38, 40, and 41 (all, Brock Mountain area); and the darkest at Localities 39 (Brock Mountain) and 45 (McCloud limestone south of Shasta Lake). Samples from elsewhere on the McCloud limestone are intermediate in shade. The shoulder and spire are medium reddish brown, as dark as or darker than the base. They are never as dark as the supraperipheral band. The darkest spires occur in the Brock Mountain area and the lightest along the McCloud River. In both groups there is a tendency for the center of the shoulder to be lighter, producing a shad- owy secondary banding that is most evident on the lighter shells of the McCloud River herd. The secondary banding is never as distinct as that in Monadenia fidelis (see Talmadge 1960: his ‘‘multibanded phase’’). One other color ef- fect is observable: some individuals have radial streaks of darker pigment on the shoulder. These streaks usually precede a growth rest and evidently represent concentrated pigment de- position at times when shell growth is slowing down. They are often followed by a whitish ra- dial streak (as growth starts up again rapidly?). When combined with secondary banding, as in some McCloud River shells, the streaks produce a mottled effect. It seems likely that the same shell pigment, in various dilutions, produces the supraperipheral band, base, and shoulder coloration. The expanded lip of adult specimens is white. Color variation in M. troglodytes segregates geographically: populations in the Brock Moun- tain area have the darkest spires, both the dark- est and the lightest bases, and the broadest bands near the periphery. Populations along the McCloud River have the lightest-colored shells and the greatest incidence of secondary banding and mottling. In the course of ontogeny, the shell of Mon- adenia troglodytes undergoes several significant changes in shape. The protoconch—that portion of the shell which forms within the egg—is al- most hemispherical, with nearly flat spire and deeply convex base. Gould (1969) suggested that mechanical limitations of space within the egg largely determine protoconch shape in the zoni- tid snail Poecilozonites, and in fact, the proto- conch of M. troglodytes is shaped very much like that of Poecilozonites bermudensis bermu- densis (Gould 1969:pl. 6, figs. 6, 7). Post-embryonic juvenile shells are wide, flat- to low-spired, angulate at the shoulder, and tu- mid at the base. Again as in Poecilozonites, postembryonic growth begins with a marked in- crease in relative width. As growth proceeds, the whorl changes in cross section (Fig. 9) from crescentic and taller than broad, in the embry- onic shell, to auriculate and broader than tall in the adult. Also with growth, the periphery, which is above the middle in juvenile shells, migrates downward until it is medial in adults and changes gradually from angulate to rounded. Spire height increases allometrically with re- spect to shell diameter, producing a domed spire. Since from about the third whorl on, the height-diameter ratio of the shell remains prac- tically constant, the allometry is due chiefly to the downward migration of the periphery. In Monadenia, as in most other helicacean snails, growth is determinate. The shell enlarges up to a point, generally coincident with sexual maturity, when the lip is thickened and turned outward. Thereafter, no additional spiral growth takes place. In M. troglodytes, the greatest ab- solute height of aperture and often the greatest total shell height are achieved one-half to one- quarter whorl before the cessation of growth. For the last quarter turn, translation along the ROTH: DISTRIBUTION, ANATOMY, VARIATION OF MONADENIA TROGLODYTES 391 12 Ficures 11-13. n. subsp., holotype CAS 018431, Loc. 34. Figs. 13a-c. M. t. chaceana, paratype CASGTC 10125. vertical axis is halted or reversed, and at the same time, the whorl is compressed apico-ba- sally. The final 3-4 mm increment of growth strikes downward at about a 30° angle to the _ suture, bringing the peristome nearly into tan- gency with the face of the body whorl. Similar terminal growth occurs in many genera of heli- cacean snails, particularly those from xeric en- vironments. It is presumably an adaptation to enhance the snails’ mucous seal to the substra- tum, in order to retard water loss or exclude predators. The final half-whorl “‘leap’’ of growth is most strongly expressed in populations along the McCloud River north of Shasta Lake and contributes to their generally low height-diam- eter ratios compared to Brock Mountain popu- lations. As already noted above, Monadenia troglo- C 1S Cc Shells of Monadenia troglodytes, x1.5. Figs. 1la-—c. M. t. troglodytes, Loc. 26. Figs. 12a—c. M. t. wintu, dytes undergoes a much greater range of variation in shape than was evident from the type lot alone. Data on basic shape measurements and indices—height (H), diameter (D), H/D ratio, number of whorls, umbilical width (U), and U/D ratio—are summarized in Table 3. Data from adult shells (those with reflected lip) only are included. Samples were collected without spe- cial procedures to insure randomness, but there is no reason to suspect bias with regard to any of the dimensions or ratios used here. All shells were measured with a hand-held cal- iper with vernier scale. Height (H) was mea- sured parallel to, and maximum diameter (D) perpendicular to, the axis of coiling of the shell. The expanded lip of adult shells was excluded from these dimensions. Umbilical width (U) is an inside caliper measurement taken parallel to ‘Op = U ‘s]LOYymM Jo JaquNu J0y Lp = U ‘G/M ‘A 4105 ‘8p = UG + H ‘C/H ‘C 10} ‘6p = UH 104 xx ‘67 = U ‘sOYM Jo Jaquinu Joy :7~E = U ‘G/N ‘A ‘Cd 40J OE = UG + H ‘G/H‘H 404 « PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 15 (L67 + 19°9€) (810'0 + 6110) (6700 + PLS'0) 610 + PS'S) (Ig'0 + LL) (66:1 + Leet) COLI + be er) ly 10) TOPS 8C 6t1 0-880 °0 8£9°0-908 0 SS ISSLG AS Olvaeal 0:9¢-0'81 9 rI-S O01 0 Sajsuripend) BYa1A pur YOO1qUIOH DUDAIDYD *) “PW (LOT + 1S°8€) (S100 + 8IT'0) (FI0'0 + 9¢S'0) = (91 0 + 69°) (9¢°0 + $6 °C) (91:0 + 16'¥e) (1¥'0 + 09'ET) L'6t-8 9t Iy1'0-060'0 OLS 0-6cS 0 6 SF's EAS QST9'ET GAIA sl! 8 (6) Y9A4D A901 PAOGY (€7'7 + LSOF) (Z10'O0 + OZI'0) + (870.0 + TES'0) (910 + 19'S) (Ir'0 + 9I'¢) (Qb'1 = Lv'9Z) (OO'L + LO'PI) Loy vy 9t brl 0-160°0 809'0-£8PF 0 GS-6 'S Ovmric £ 6C—-9 CC bp 91-0 Cl # (Ip ‘Or “8e “PE-Te) WW 401g JO AN (tcc + LELE) (S10'0 + OZI'0) +=(970'0 + 97S'0) += (17 0 * L's) (ch'0 + £60) (Lp l + Spc) (88°0 + P8 CI) Scr 6 tt 8h1 0-F60°0 OLS '0-08P 0 BS-CS EK C Da} VC 6-I-c II Sc (Sg) Y2e1Q Menbg aoqy (LIZ + O@'8E) (HIO'O + L710) (870'0 + cHS'0) §~=— (ST 0 * CL’) (pe 0 + PI’) (9E1 + 8L hc) OOT + Cr ETD) CCY9 VE SS1°0~960°0 18S °0- 18h 0 ASS [ENE CE GECEOKCC SOREL 81 (Cp) Attenb JUaWad suaAjed NIUIM * (LUI + $8°Se) (p10'0 + StI'0) §=(P10°0 + C6h'0) = (ST 0 + OP'S) (OF'0 + 99°¢) (08'0 + P02) (Lh0 + I8 TD) ie ULSAAS VAs 181 0-0F1 0 OS 0-697 0 ISOS CVE Ce 6 bC-8 CC WAS ||! 8 (€h) Y9ID 19110 (L61 + LS‘re) (7100 + prl'0) (1700 + LOS) =I 0 + IHS) Whe 3 GE) (861 + S6c¢) (0L0 + C9'TD) OLED It y9l'O-LIT 0 BPS 0-9SPh 0 Sissi Gas WAS © O'SC~9'0C 9 CI-L Ol Ic (87) YOON seyxxed (pL 1 + S88) (E100 + I91'0) (€70'0 + 98b°0) (hI 0 + LS'S) (Ob'0 + 07h) (QAl se fell) WE) Fe ©) 6 IPC Se p81 0-9t1 0 ccS 0-0 0 SASS SAS T8c-L tC OPIS TI cc (LZ “9Z) ¥9AD AYPIIF (QL 1 + 9€°9€) (910'0 + 6S1'0) (E700 + 8Lh'0) = (17 0 + CHS) (6£'0 + 16't) (pI: + 697) (620 + SLIT) O OPT ct 961 O-SEl 0 Tes 0-1 Ph 0 09-0'S SPO CLEBOMNC Tel-c ol * (P7-L1) PAB] JOMUES JO AWUISTA (STI + pc ee) (yIOO + LPl‘0) (7700 + 9660) (SIO + £9) (L¢°0 + L7't) (PSION eGrCO) eS Si OneeaLOMIL) SENEAS WS cLI0-8cl 0 6tS O-SSP'0 gs~0'S OF-8C S'PC~6 0C €cI-0 01 €C (QL) Aav_ JamURg Mojaq ado[s SAIAPO]BOA] “1 “WW aq+H q/n d/H S[TOYM Jo (N) WPM (q) 19} 2uIvIG (H) 14319H u AqB907] JoquinN feotiquiy) “papnyour sjjays 1jNnpe 392 AJuQ ‘sasayjuaied ul uonelAap pivpurjs duo + uvaW ajdwies YUM aBuRI aie SONSHeIG “SasApojdo4 VIuapyUuop NI SOLLVY ANY (WW UI) SNOISNAWIG TIAHS “¢ FTV L ROTH: DISTRIBUTION, ANATOMY, VARIATION OF MONADENIA TROGLODYTES H/D Samwel types Potter Cr types 393 chaceana 20° ae 10 23° i 24 30° 26, 27 22— — ee 28 21 ee 43 8’ a 1 45 18°- a 35 a a 31-34,38,40 41 a et 39 . U/D .08 10 12 14 16 18 20 chaceana 20° 10 23° 11-24 22 26, 27 22 28 Ne Te 43 ee 45 8° —— 35 2 a 31-34, 38, 40,41 47° — == 39 8° a ‘| FiGures 14a (upper), 146 (lower). Relative height of spire (H/D) and relative width of umbilicus (U/D) in Monadenia troglodytes. Line designates sample range; vertical strike, mean; open bar, mean + one standard deviation; solid bar, 95% confidence interval of population mean. the maximum diameter and occasionally re- quired breaking away a portion of the inner lip. Number of whorls was determined, with the use of a binocular microscope, by the method of Diver (1932) except that the final fraction of a whorl was estimated without pencil lines being drawn on the shell. The measurement is rounded to the nearest 0.1 whorl. Pilsbry (1939:xi, fig. B) evidently counted whorls by approximately the same method, although he usually expressed re- sults to the nearest one-quarter or one-third of a whorl and sometimes quoted uncritically the figures of other authors. The simple measurements of size (H, D, H + D) show that as a group the McCloud River herd averages smaller than the Brock Mountain herd, although there is extensive overlap of ranges in all categories. The smallest averages belong to the samples from Localities 10 (below Samwel Cave) and 28 (Dekkas Rock). Shells of large di- ameter (and large H + D) occur near Ellery Creek (Locs. 26, 27). Relative height of spire (H/D) segregates geo- graphically (Fig. 14a), with McCloud River shells significantly lower-spired than those from Brock Mountain. As in most other parameters, shells from Locality 45 segregate with the Brock Mountain herd. The U/D ratio, the relative width of the umbilicus, segregates along the same lines (Fig. 144)—not surprisingly, since the two ratios, H/D and U/D, may be aspects of the same variable. A tapering tube of a given size and rate of expansion may be coiled steeply (high rate of translation along coiling axis and small distance of generating curve from axis: Raup 1966) or flatly (low rate of translation, large distance from axis). The first mode gen- erates a high-spired shell with small umbilicus, the second a low-spired shell with wide umbili- cus. A correlation coefficient of —0.951 for sam- ple mean H/D and mean U/D further attests to the association between the two parameters in M. troglodytes in the Shasta Lake region. The high spire and small umbilicus of M. t. chaceana 394 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 15 % — M.t. chaceana 31-34, M.t. wintu 38, 40, 41% % 35 510 F a S .490 M.t. troglodytes x= M2627 470 4 2: 10 120 130 140 150 160 U/D Ficure 15. Relation between mean relative height of spire (H/D) and mean relative width of umbilicus (U/D) in Mona- denia troglodytes. are in harmony with this association, although the umbilicus is broader and/or the spire higher than regression of the Shasta Lake statistics pre- dicts (Fig. 15). Figure 16 plots the relationship between mean size (H + D) and whorl number. Although the scatter of individual (specimen) points overlaps extensively, again there is geographic segrega- tion of means. Locality 45 sorts with the Brock Mountain herd, nearest to the pustulose shells from Locality 39. Mean size and mean whorl number correlate strongly in the McCloud River herd (r = 0.954); the more heterogeneous Brock Mountain herd displays no such tight correla- tion. Based on the dimensions given by Hanna and Smith (1933), the type lot of M. troglodytes from Samwel Cave and their material from Potter Creek Cave are flatter (lower H/D) than any of the samples here studied. Both the mean diam- eter and mean H/D ratio of the type lot differ significantly from those of the nearest living sample, from Locality 10. It would be prema- ture, however, to conclude that the difference represents evolutionary change. Hanna and Smith accepted a Pleistocene age for the cave specimens, first because they believed the species extinct and, second, because the shells had been associated with bone deposits. (The Samwel Cave and Potter Creek Cave vertebrate mm: 41| 31-34, 38,40,41 39) 26,27 a 39m | M45 35 37 ad a 11-24 @ @1 4-7 + 43 Tt 35) 28 33 | 10 ih ee h nm i 1 52 53 54 55 56 57 Whorls FiGurE 16. Relation between mean size (H + D) and mean number of whorls in Monadenia troglodytes. Diamonds, M. t. troglodytes; squares, M. t. wintu, n. subsp.; circle, M. t. chaceana. faunas have been assigned a Rancholabrean [late Pleistocene] age; Hibbard et al. 1965.) But the burrowing or crevice-seeking habit of land snails makes them unreliable stratigraphically. State of preservation is not very informative: in the sev- eral cave samples now at hand, there are shells in every gradation from slightly dusty to heavily encrusted with calcium carbonate, and there may be a more or less continual influx of shells drifting or washing into the caves. Independent evidence of age, such as radiometric dating, and collections with good stratigraphic control are needed before one can place the cave samples in a time framework. SYSTEMATICS TAXONOMIC CRITERIA IN Monadenia.—In- dividuals of Monadenia troglodytes in the Shas- ta Lake region group strongly into two geo- graphic units based on shell color and texture, microsculpture, and morphometry. Monadenia chaceana constitutes a third such unit. Although its internal anatomy is not yet known, its shell characters are highly similar to M. troglodytes, and it is thus given coordinate rank with the lat- ter two groups. The subspecies category has been employed somewhat unevenly in terrestrial malacology. Early workers responded to the manifold vari- ability of some tropical snail groups, for in- ROTH: DISTRIBUTION, ANATOMY, VARIATION OF MONADENIA TROGLODYTES 395 stance, by naming dozens of subspecies, and the morass of names led in turn to a reaction against taxonomic recognition of infraspecific units (Gould 1969). I agree with Kavanaugh (1979:93) that “‘recognition of the usefulness of a subspe- cies concept . . . [depends] on one’s particular perspective; more specifically, on the distribu- tion of habitats or areas occupied by the organ- isms one studies. Where gaps between areas of suitable habitat are broad and clear, . . . corre- lated discontinuity in variation is more confi- dently recognized and suggestive of active, ef- fective barriers to gene flow.’’ The three groups of Monadenia troglodytes meet the criteria for subspecies. Shell character differences are mainly quantitative and correlated with geog- raphy. Except for the different verge configu- ration at Locality 43 (the significance of which remains unresolved), the identity of the genitalia between the two Shasta Lake region herds sug- gests that interbreeding would be possible if the two were brought together sympatrically in na- ture. Limestone substratum is apparently the required habitat, its absence a probable barrier to gene flow, at least under present environmen- tal conditions. Rivers are at least short-term bar- riers, but over the long term may be important agents of dispersal. The population at Locality 45 may well have been established by river- borne waifs rafted down the Pit River—perhaps from the population of Locality 39, with which it groups morphologically (Figs. 10, 14-16). Cli- nal variation in granulation along the McCloud River arm suggests incomplete genetic isolation between those localities. Populations on the McCloud limestone, exclusive of Locality 45, constitute one unit; since it includes the type- locality, this unit is the nominate subspecies, Monadenia (Shastelix) troglodytes troglodytes. Populations between the Pit River and Squaw Creek arms of Shasta Lake, and the southern population at Locality 45, are named as a new subspecies below. The sympatric or parapatric occurrence of Monadenia churchi and M. troglodytes at sev- eral localities in the Shasta Lake region should, theoretically, permit an estimate of the degree of morphologic difference that can be expected between species of the genus, and by analogy aid in ranking allopatric taxa. Reproductive structures are particularly suitable for such an analysis, because a major difference in genitalia (taking into account the possible effects of sea- sonal variation and genital polymorphism) im- plies functional incompatibility. Solem (1975) used data on reproductive anatomy of sympatric species to define taxonomic criteria in the snail genus Oreohelix Pilsbry, 1904. Like many other helicacean snails, monadenias have elaborate terminal genitalia, replete with accessory organs which are employed in a courtship of some com- plexity (Webb 1952, 1966). Specialized repro- ductive organs and complex precopulatory be- havior are sometimes regarded as isolating mechanisms in other animal groups; theoreti- cally, they prevent unproductive mating be- tween sympatric species. Solem (1978:67) pre- dicted that, among pulmonates, elaboration of stimulatory and glandular or dart structures would occur where the need for species recog- nition signs was the greatest, for example, under conditions of sympatry in areas of historically fluctuating climate and vegetational cover or in island situations involving explosive speciation. The differences in the genitalia of M. churchi (Fig. 8) and M. troglodytes (Figs. 4-6), how- ever, are relatively limited and far less marked than those between members of different sub- genera. The values for organ lengths in M. churchi (Table 2) fall mostly within the range of variation of M. troglodytes. The dart sac averages smaller in M. churchi, but the size range overlaps that of M. troglodytes. Length of the mucus gland is highly variable in both species and may vary according to recency of copulation. M. churchi from Mammoth Butte, west of Shasta Lake, and others from the Trinity River drainage have a thin, cylindric verge with the end squared and compressed, the meatus transverse. The verge of a specimen from near Campbell Creek on the east side of the McCloud River arm (Loc. 29) is also squared at the tip but thicker, with lateral meatus and concave facet as in M. troglodytes (Fig. 8b). All specimens examined have a single large pilaster in the penial chamber. It appears, therefore, that reproductive anatomy in Shas- telix is too variable and too weakly differentiated to offer reliable characters for diagnosis. The two species are much more readily dis- tinguished on shell characters: M. churchi has strong, pustulose microsculpture over the entire shell, even when adult; a dull periostracal sur- face; and uniform brown color on base and 396 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 15 spire. Quantitative differences, not always pres- ent but usually helpful in distinguishing the two species, include higher spire, stronger growth rugae, and more tightly coiled last whorl. I have detected no systematic conchological differences between M. troglodytes found with M. churchi and those occurring alone. In each case the M. troglodytes shells retain the stan- dard characters of their herd. On the other hand, M. churchi from within the range of M. trog- lodytes tend to be larger, shinier, more inflated, and less densely pustulose than specimens from elsewhere. Whether this apparent convergence with M. troglodytes goes beyond independent geographic variation will be examined in a later paper. The finding that in reproductive anatomy M. churchi and M. troglodytes differ from each oth- er chiefly in verge shape also implies that the elaborations of the female side of the genital sys- tem—dart sac, copulatory pad, etc.—may not be involved in species recognition during court- ship or maintenance of reproductive isolation. According to the sexual selection model of Charnov (1979), hermaphroditic animals may copulate not so much to gain sperm to fertilize eggs as to give sperm away (to gain access to another’s eggs). ‘““There must often exist a con- flict of interest between mating partners—as a recipient each should be inclined to accept sperm (not necessarily for fertilization of its own eggs) in order to give its sperm away’ (Charnov 1979:2482). In order to pair with and fertilize a partner, a Monadenia may have to display its OWn apparent receptivity to the partner’s sperm. Like any character that individuals use to choose sperm donors, the organs involved in such a display would be subject to exaggeration through time. Selective pressure toward species- specific differentiation of the organs would de- pend on (1) frequency of the opportunity for in- terspecific mating, based both on degree of sym- patry and the effectiveness of other organ or behavior systems in maintaining isolation; and (2) relative fitness of resulting hybrids. Monadenia (Shastelix) troglodytes troglodytes Hanna and Smith (Figures 4a-h, 9a-c, lla-c) Monadenia troglodytes HANNA AND SMITH, 1933:84-85, pl. 5, figs. 6-8.—Pitspry 1939:54, fig. 22(6-8).—SmItTH 1957:26; 1970:40.—WaLTon 1970:111.—RoTH 1972a:7; 1972b:7; 1979:13. ORIGINAL DESCRIPTION.—‘‘Shell light buff, medium size, widely umbilicate; spire greatly depressed; whorls 5'2 with moderately deep su- ture; the last whorl slightly depressed near the aperture; outer margin expanded very little, the basal margin somewhat more so; one pale brown spiral band appears just above the periphery, which is bounded above and below by white bands that are slightly wider; surface without markings except growth lines; nucleus consist- ing of 142 whorls marked by radiating wavy rib- lets. Diameter 24.2: altitude 10.8 mm” (Hanna and Smith 1933). This description was repeated by Pilsbry (1939). Type MATERIAL.—Holotype: UCMP 32394 (shell): CALI- FORNIA: Shasta County: Samwel Cave (UCMP loc. 1008). Paratype: CASGTC 5842 (shell); same locality as holotype. REFERRED MATERIAL.—CALIFORNIA: Shasta County: Bol- libokka Mountain Quadrangle: Loc. 8 [2 specimens], 9 [5], 10 [32], 11 [6 in addition to holotype and paratype], 12 [2], 13 [2], 14 [2], 15 [2], 16 [1], 17 [4], 18 [2], 19 [315 20 [2]; 20 [els 22 [1], 23 [2], 24 [9], 25 [2], 26 [30], 27 [5], 28 [38], 29 [3]. Lamoine Quadrangle: Loc. 42 [1], 43 [22], 44 [12], 47 [1], 48 [1]. Figure 3 maps these localities. Discussion.—As described above in the sec- tion on geographic variation and graphically shown in Figures 14-16, M. t. troglodytes has the lowest-spired and most broadly umbilicate shells of any subspecies. The shells average smaller than those of the Brock Mountain herd (M. t. wintu, next described), but large shells occur near Ellery Creek. The lightest-colored shells and the greatest incidence of secondary banding and mottling occur in M. t. troglodytes. The nominate subspecies tends to have fewer coils to the epiphallus than M. ft. wintu. Monadenia (Shastelix) troglodytes wintu, new subspecies (Figures Sa—b, 6a-b, 7, 12a-c) DIAGNOsIS.—M. troglodytes with moderately high spire (sample mean H/D = 0.526—0.546), shell solid, lustrous, shoulder reddish brown, strong white zones above and below brown su- praperipheral band, base either nearly white, or reddish brown, as dark as shoulder. Granulation often persisting past third whorl. DESCRIPTION.—Shell of moderate size for the genus, solid, moderately to distinctly thick, lus- trous; spire convexly conic, moderately elevat- ed: whorls 5.2—6.1 in adult, tightly coiled; body whorl slightly expanded over last % turn. Pro- toconch of 1.5—1.9 whorls, nuclear tip smooth, ROTH: DISTRIBUTION, ANATOMY, VARIATION OF MONADENIA TROGLODYTES 397 followed by fine granulation tending to form wavy, radial riblets below suture; granules else- where spirally elongated, in diagonal series or irregularly scattered. Early neanic whorls less densely granulated, granulation becoming sparser with growth, usually persisting past the 2.5-whorl stage and often past the 3-whorl stage in at least some members of each sample. Body whorl sculptured with fine growth rugae and microscopic, wavy, parallel, spiral lineolation, most evident behind lip. Juvenile shells with base tumid, spire low, shoulder angulate; pe- riphery becoming first obtusely angular and fi- nally rounded with maturity. Base of adult shells rounded, umbilicus open, steep-walled, diame- ter 0.090-0.155 times major diameter of shell. Last 3-4 mm of body whorl striking downward at about 30° angle to suture. Aperture broadly auriculate, apico-basally compressed, oblique. Peristome slightly thickened, everted; inner lip covering 10-25% of umbilicus. Color of spire and shoulder medium reddish brown (rust of Rayner [1970]; Munsell midpoint 9.0R/4.3/8.0); dark reddish brown (chestnut of Rayner [1970], Munsell midpoint 9.5R/2.3/3.8) band just above periphery, 1-1.5 mm wide in adult, bordered above and below with white or light tan zones, upper zone either narrower or about as wide as brown band, lower zone usually as wide or somewhat wider; base whitish with light tan suf- fusion or (at some localities) reddish brown as dark as shoulder. Periostracum smooth, color- less or very light yellowish tan. Dimensions: ho- lotype, height 12.8 mm, diameter 25.1 mm, di- ameter of umbilicus 3.0 mm, 5.6 whorls: largest paratype (Loc. 35), height 14.9 mm, diameter 27.6 mm, diameter of umbilicus 2.9 mm, 5.6 whorls. Dimensions of other referred material -summarized in Table 3. Body of animal dove gray to sooty black with a reddish or purplish cast, darker on dorsum: light mid-dorsal stripe; sole light buff with gray margin. Mantle over lung translucent buff with gray pencilling covering less than 10% of sur- face. Genitalia (Figs. 5, 6) as in typical subspe- cies. Jaw as in M. t¢. troglodytes. Radula substan- tially as in M. ¢. troglodytes, with 42 teeth ina half row (Locs. 34, 45), an endocone developed on tooth 21 (Loc. 34) or 14 (Loc. 45), an ecto- cone on tooth 22 (Loc. 34) or 17 (Loc. 45). Bifid and trifid endocones occur sporadically. Type MATERIAL.—Holotype: CAS 018431 (shell, radula. and soft parts); CALIFORNIA: Shasta County: NW!4 NE!4 sec 8, T. 34 N, R. 2 W, USGS Bollibokka Mountain Quadrangle (15-minute Series [Topographic]; ed. 1957), cave between two limestone buttes at south end of Gray Rocks, above Pit River arm of Shasta Lake. S. E. Hirschfeld coll., Oct. 1975. {Loc 34.) Paratypes: CAS 018432, 018433 (shells and soft parts), same locality as holotype. CAS 018434, 16 shells, same locality as holotype. CAS 018435, 5 shells, Loc. 35. USNM, ANSP, AMNH, FMNH, LACM, one shell each, Loc. 35. REFERRED MATERIAL.—CALIFORNIA: Shasta County: Bol- libokka Mountain Quadrangle: Loc. 30 [2 specimens], 31 [19], 32 [8], 33 [1], 36 [3], 37 [5], 38 [7], 39 [13], 40 [14], 41 [20]. Project City Quadrangle: Loc. 45 [40]. Figure 3 maps these localities. ETyMOLOGY.—The subspecies is named for the people native to the region where it is found, the Wintu tribe. Discussion.—The major features of variation within the subspecies and its morphological re- lationship to other subspecies are discussed above and graphically shown in Figures 14-16. In coloration and spire height, M. t. wintu and M. t. chaceana are more similar than either is to M. t. troglodytes, whereas in microsculpture and relation of whorl number to overall size, M. t. chaceana and M. t. troglodytes group more closely. Monadenia (Shastelix) troglodytes chaceana Berry (Figures 13a—c) Monadenia chaceana BERRY, 1940b:9-11, figs 9, 10.—PILsBRy 1948:1092.—SmitH 1960:97.—RorTH 1972a:5; 1972b:6. ORIGINAL DESCRIPTION.—* ‘Shell of but mod- erate size, weight, and thickness; spire low-con- ic to moderately elevated; whorls 5% to 6, subangulate and carinate above the middle during juvenility, subcarinate at adolescence, but becoming obtusely angular and finally quite well rounded at maturity; base tumid, the um- bilicus open, steep-walled, permeable to apex, and contained on the average about 8.4 times (7.45 to 9.88 in those measured) in the major shell-diameter. Aperture somewhat descending above, oblique, rounded to round-ovate, slightly or not at all flattened below; peristome nearly simple above, elsewhere usually little thickened and but moderately everted, terminating below in a very moderate columellar flare which covers only the edge of the umbilicus. ‘‘Embryonic shell swollen, of 134 to 2 whorls; the surface initially smooth, but almost at once breaking into a few, irregular axial waves suc- 398 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 15 ceeded by a close, fine, crowded granulation, which abruptly ceases with the first post-embry- onic whorl, the often heavy but extremely vari- able growth-striae thenceforth becoming the dominant feature; a few indistinct, elongate, and commonly confluent papillae arranged in rather distant forward-descending series appear on the early turns, but gradually give way on the later whorls to a weak and not very regular spiral striation which may become quite indistinct on the base: general surface between the striae and growth-lines very finely microscopically wrin- kled in a cloth-like pattern. ‘*Periostracum smooth and lustrous; deep brown. encircled by a conspicuous dark-brown band about 2 mm. wide just above the periphery, bordered by a much narrower yellowish band and yet narrower band of the same pale tone just above, while on the shoulder some shells show varying traces of yet another band of interme- diate brownish tone, best seen in juvenals” (Berry 1940b). TypE MATERIAL.—Holotype: SSB 8678 (shell); CALIFOR- N1A: Siskiyou County: among rocks about halfway up a spur of Badger Mountain on west side of Shasta River Canyon not far above its mouth (Berry 1940b). Paratypes: CASGTC 10125 (formerly A. G. Smith no. 7102; shell); SUPTC 6555 (shell); according to Berry (1940b), ad- ditional paratypes are in the Berry Collection, with others originally to be deposited in the USNM and the Emery P. Chace Collection. DISTRIBUTION.—Monadenia t. chaceana is known only from the general vicinity of the type- locality, in the Hornbrook Quadrangle, Siskiyou County (Locs. 1—5), and from one site (Loc. 6) in the Yreka Quadrangle, Siskiyou County (see also.Fic- Ii); Locality 7, “near Yreka’ is too generalized to assign to a quadrangle with con- fidence. Limestone areas west of Gazelle and southeast of Scott Valley need additional pros- pecting for Monadenia. Discussion.—In coloration, M. t. chaceana most closely resembles the darker samples of M. t. wintu. The base is as dark as the spire, although neither is as dark as the supraperiph- eral band. At Localities 5 and 7, along with the usual dark-spired individuals, a form is pres- ent with light shoulder and one narrow, medium- brown, secondary band about two-thirds the dis- tance from the suture to the periphery. The substance of the shell is thinner and more translucent than in the Shasta Lake region groups. Monadenia troglodytes chaceana has the highest mean H/D ratio of any subspecies (Table 3, Fig. 14a), differing highly significantly from any sample of M. ¢. troglodytes or M. t. wintu. At equivalent diameters, adult M. t. chaceana average 0.49 whorl more than M. t¢. wintu from the Brock Mountain area. Berry (1940b) compared his M. chaceana to Monadenia mormonum cala (Pilsbry, 1900) and M. churchi. He also intimated a possible rela- tionship to Monadenia fidelis minor (Binney, 1885) and surmised that the type-locality of that subspecies might be the Mount Shasta region. The latter supposition was evidently based on a statement by Binney (1885; Binney and Bland 1869) that small Monadenia fidelis occur at Mount Shasta. Binney (1885:121, fig. 91) proposed ‘‘Aglaia fidelis var. minor’ without a diagnosis and his original figure is unlocalized, but elsewhere (1885:141, footnote; 493) he recorded it from The Dalles, Oregon, collected by Henry Hemp- hill. Henderson (1936) and Pilsbry (1939) ac- cepted The Dalles as the type-locality. The Cal- ifornia Academy of Sciences collection contains two probable syntypes of A. f. var. minor, CASGTC 6001-6002, formerly H. Hemphill Col- lection no. 8598 and labeled **Helix fidelis Gray ,/ var. minor W. G. Binn/depressed near mormon- um/Near The Dalles, Oregon/Types/HH”’ in Hemphill’s handwriting. They are small M. fi- delis, agreeing well with Pilsbry’s (1939) diag- nosis of M. fidelis minor and moderately well with Binney’s original engraving. That figure shows a pale shoulder with a light-centered spi- ral band and an indication of strong radial growth striae on the early whorls—all charac- teristic of the Dalles form. The Binney and Bland Collection of terrestrial mollusks, now at the American Museum of Nat- ural History (Gratacap 1901), contains one spec- imen of M. fidelis labeled *‘H. fidelis/Mt. Shasta/ Cooper.’ An oval, gold-edged, adhesive label on the specimen states ‘‘Mt. Shasta/WHB JGC’’; the shell was evidently collected by Wil- liam H. Brewer and given to James G. Cooper, who passed it on to Binney. This specimen, AMNH 57788, is 25.8 mm in diameter, 16.1 mm high, with 6.0 whorls. The shoulder is unpig- mented except for faint, discontinuous traces of a light-centered spiral band. It is not the speci- men illustrated by Binney (1885). ROTH: DISTRIBUTION, ANATOMY, VARIATION OF MONADENIA TROGLODYTES 399 To my knowledge, M. fidelis does not live at Mt. Shasta proper, the nearest localities being Beaver Creek near the Klamath River, Siskiyou County (subspecies M. f. leonina Berry, 1937), and near the shores of Klamath Lake, Oregon (unnamed subspecies cited by Pilsbry [1939:42, fig. 18e] as M. f. minor). The Binney specimen is similar to the latter but more narrowly umbil- icate. Brewer probably collected this shell on his 1863 trip through the Klamath Mountains, rather than on his 1862 climb of Mount Shasta (Brewer 1930). Cooper (1869) mentioned receiv- ing from Brewer M. fidelis collected at Crescent City, the western terminus of Brewer’s Klamath route. I conclude (1) that the name minor applies to a race of small Monadenia fidelis (Gray) from around The Dalles, Oregon, and (2) that Berry was mistaken about the similarity of his M. cha- ceana to Binney’s *‘Mount Shasta’ material. PHYLOGENETIC HYPOTHESES CONCERNING Monadenia Early statements about evolution within Mon- adenia tended to accord the widespread and conspicuous M. fidelis a central—and in some unspecified way archetypal—position, while the smaller, more remote and cryptic M. hille- brandi, M. mormonum, and similar forms were viewed as derivative (and possibly degenerate) offshoots (for example, see Cooper 1887: Stearns 1900). A questionable exception is that of Cooper (1873), who evidently attempted to adduce general principles for the direction of character-state transformation in west coast snail genera. One can readily speculate as to what extent an ethnocentric outlook colored such views. A second generation of malacolo- gists studying Monadenia (chief among them Pilsbry, S. S. Berry, G D. Hanna, and Junius Henderson) concerned themselves more with description and less with interpretation of evo- lutionary relationships. But in their work too, a typological bias, with M. fidelis the ‘‘type”’ in more than a nomenclatural sense, may be de- tected. In a recently proposed alternative hy- pothesis (Roth 1979), the ancestral Monadenia was seen as a ground-dwelling, low-spired, and somewhat variable form, from which the ex- posed-crawling, partly diurnal, and semi-arbo- real M. fidelis evolved: ‘‘Once emancipated from life in holes in the ground, the fidelis group achieved large size, relatively high spire, and an TABLE 4. CHARACTER COMPLEXES IN Monadenia morphous states listed first). (apo- 1. Atrium small and narrow (large and globose). . Mucus gland about equal to (many times longer than) dart sac. 3. Lower part of mucus gland adnate to (free from) atrium. 4. Dart sac 5 mm or less in length (longer than 5 mm). 5. Flagellum about as long as (=1.5 times length of) penis plus epiphallus. 6. Flagellum and spermatophore straight (helically coiled). 7. Basal chamber of penis sessile on (invaginated into) atrium. 8. Penial retractor inserted near middle (on distal third) of epiphallus. 9. Habit at least partly arboreal (ground-dwelling). 10. Granulose microsculpture present on teleoconch (limited to protoconch). 11. Shoulder band pigmentation monomorphic (polymor- phic). to extensive range which now reaches farther north than any other American helicacean”’ (Roth 1979:13). Roth and Eng (1980) offered a conjec- tural, ‘“‘narrative’’ (sensu Ball 1976) hypothesis for the origin of Monadenia setosa Talmadge. Such narrative explanations are not analytical and have little predictive power. To date, none of these competing models has specified the cri- teria for judging relationships, and none has been couched in testable propositions. Phylogenetic systematics, the methodology of Willi Hennig (1966), seeks to analyze the prob- able direction of evolutionary character trans- formation in a particular group of organisms and, from this analysis, to reconstruct the evo- lutionary history of the group. An important vir- tue of this method is that it generates testable hypotheses about phylogenetic relations. (See Kavanaugh [1972] for an exposition of Hennig’s approach to systematics and Gaffney [1979] for a useful bibliography of papers dealing with cla- distic methodology.) In land malacology, Van Goethem (1977), Breure (1979), and Bishop (1979) have applied these principles to the land snail groups Urocyclinae, Bulimulidae, and American Camaenidae, respectively, and the same approach is implicit in the writings of a number of other malacologists. Of 55 morphological and natural history char- acters studied, 11 (Table 4) were incorporated in a cladistic analysis. Characters excluded were those in which polarity of the transformations could not be interpreted, those that occur only 400 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 15 Monadenia,s.s NX Corynadenia Shastelix 51679 FiGure 17. Suggested cladogram for the subgenera of Monadenia. Numbers refer to apomorphous states of char- acter complexes listed in Table 4. in states judged to be symplesiotypic for the ge- nus, and those that are unique (as apomorphies) to one species and hence provide no information for grouping. Figure 17 illustrates the relationship of char- acter states in the three subgenera of Monaden- ia. It is well to emphasize that the hypothesis presented here is preliminary and undoubtedly will be modified as study of the genus pro- gresses. This cladogram is based mainly on re- productive anatomy. In general, shell characters are much more labile, phenotypic responses to environment are common, and the number of conchologically polytypic species in Monadenia hampers the use of shell features in this kind of analysis. As Bishop (1979:275) noted, snail shells offer relatively few characters for analysis and are difficult to characterize objectively, “though both shape and texture may be most informative to the eye of an experienced worker in a manner which is hard to express in words.” Moreover, shell characters are subject to nu- merous parallelisms and convergences, usually associated with habit and habitat (for example, the often-remarked similarity between the rock crevice-dwelling species, Monadenia circum- carinata (Stearns, 1879) and Oreohelix elrodi (Pilsbry, 1900)). A closer study is needed to dis- criminate analogous from homologous shell character states and extend the cladistic analysis to the species level. Three character complexes, especially, illus- trate the problems left unsolved by this analysis. The arboreal habit in Monadenia, s.s., is re- garded as an apomorphy, but many of the as- sociated apomorphic characteristics (potential for large size, high spire) are differentially ex- pressed in the various subspecies of the poly- typic Monadenia fidelis. Are these characters secondarily lost in such subspecies as the low- spired M. f. scottiana Berry and the small- shelled, unnamed race near Klamath Lake, or do these races preserve the plesiomorphous condition? Perhaps the arboreal habit and its associated character states are better viewed as expressions of a physiologically based eurytopy which permits life in the trees but also allows M. fidelis to inhabit low herbage near the coast (M. f. pronotis Berry) or rockpiles (Klamath Lake race). A specialized arboreal habit, such as the dependence of M. setosa on standing broadleaf deadwood for juvenile habitat (Roth and Eng 1980), is probably a second level of apomorphy, derived from a more generalized arboreal potentiality. A transparent periostracum, through which shell banding is visible, is probably plesiomor- phous for the genus. Banding, which is wide- spread in Helminthoglyptidae and other helica- cean families, presumably evolved in the face of visual predation pressure (Roth, in preparation). An opaque periostracum that masks the banding (still present in the shell underneath) seems to be associated with cryptic habits and indepen- dently derived in Monadenia, s.s., and Cory- nadenia. Carination of the margin of the adult shell is a paedomorphy (compare Gould 1969, Poecilo- zonites), independently derived in Monadenia, s.s., Corynadenia, and probably Shastelix. ‘‘Hypercarination” (carina set off by pinched grooves) is a further derived state among several deep rock-crevice-dwelling forms of the Sierra Nevada. While a cladogram, such as that in Figure 17, is nothing more than a branching diagram de- picting the nested pattern of synapomorphies among the taxa under study, a phylogeny is an explicit statement concerning the exact nature of the evolutionary relationship among the taxa (Eldredge 1979). A phylogenetic tree is ‘‘a dia- gram (not necessarily branching!) depicting the actual pattern of ancestry and descent among a series of taxa’’ (Eldredge 1979:168) and embod- ies assumptions and information not required in (and theoretically excluded from) the construc- tion of a cladogram. With the further addition of nn man ROTH: DISTRIBUTION, ANATOMY, VARIATION OF MONADENIA TROGLODYTES 401 explanatory narrative, one can often devise a scenario to suggest how the phylogenetic rela- tionship, and the pattern of synapomorphies in the underlying cladogram, came to be. Strictly speaking, the construction of phylo- genetic trees is limited to species; species, not genera or other higher taxa, give rise to other species. Therefore, when we speak of a dichot- omy arising between subgenera (as herein), we are really maintaining that the dichotomy arose between two species which would be classed in the separate subgenera. The suggested cladogram for the subgenera of Monadenia specifies three detected autapomor- phies in the subgenus Monadenia, s.s., two synapomorphies shared by Shastelix and Cory- nadenia, and three autapomorphies each in Corynadenia and Shastelix. The effect of this distribution of apomorphies is to falsify all pos- sible phylogenetic trees for the group except that which is isomorphous with the cladogram (see Platnick 1977:440-441, fig. 2). Consequently, the proposed phylogenetic history of Monadenia consists of (1) a dichotomy between Monaden- ia, S.S., and the common ancestor of Coryna- denia and Shastelix, followed in time by (2) the dichotomy between Corynadenia and Shastelix. THE SCENARIO.—In the John Day Formation (late Oligocene to early Miocene) of central Or- egon, three fossil forms referred to Monadenia are present: M. antecedens (Stearns, 1900), large-shelled and so similar in general appear- ance to M. fidelis that it was formerly ranked as a subspecies of the latter; M. dubiosa (Stearns, 1902), said to be another fidelis-like form (Pils- bry 1939); and M. marginicola (Conrad, 1871), which Hanna (1920) believed to be related to “the mormonum group” (i.e., Corynadenia). No anatomical details are available, of course, and the shell microsculpture, if preserved, is not specified in the literature, but the distinction be- tween large fidelis-type shells and smaller shells with the character of present-day ground-dwell- ing forms is consistent with the hypothesized early split between Monadenia, s.s., and the Shastelix-Corynadenia stock. At the time of deposition of the John Day For- mation (36.4—22 million years before present; Hammond 1979), the Cascade Arc had rotated from its earlier northwest-southeast orientation almost to its present north-south position, east of the presumably coastal Klamath Mountains— Coast Range block (Hammond 1979:figs. 8, 9). The John Day accumulated as clayey and tuff- aceous sediments in an inland basin contempo- raneously with formation of the volcanic and pyroclastic rocks of the middle Western Cas- cades Group. During this episode, Cascade vol- canism was centered in randomly spaced strato- volcanoes and calderas; the string of high volcanic edifices that characterizes the modern Cascade Range had not yet developed. Contemporaneous floras from the Coast Range block to the west are similar in floristic composition and leaf phys- iognomy to the Bridge Creek Flora from the lower member of the John Day (Brown 1959; Wolfe and Hopkins 1967), indicating that the Cascade Range was not a significant climatic or vegetational barrier at this time. The Bridge Creek Flora, associated with a ra- diometric date of 31.5 million years (Evernden and James 1964), represents a mixed mesophytic forest dominated by broad-leaved deciduous trees, in a temperate climate with ample summer rainfall (Chaney 1948; Brown 1959; Wolfe and Hopkins 1967). Its greatest similarities are with modern hardwood forests of eastern North America and eastern Asia (Chaney 1948; Whit- taker 1961). Temperature parameters suitable for forests of this composiion are now lacking in the Pacific coast states except in isolated, small, interior valleys (Wolfe 1979). The source of the John Day land mollusks is evidently the vertebrate-rich middle member, of early Arikareean (late Oligocene) age (Wood et al. 1941) and about 25 million years old (Berggren and Van Couvering 1974). Contem- poraneous floras are also mixed mesophytic, possibly somewhat warmer than the Bridge Creek Flora. The diverse vertebrate remains likewise indicate a temperate and wet climate, quite unlike the semiarid interior of Oregon of the present. It seems plausible that such an en- vironment could have supported greater intra- generic snail diversity than now seen in any for- ests of the west, much as the hardwood forests of the eastern United States now support a sub- stantial diversity of snails in genera of the Polygyridae. Part of the early diversification of Monadenia forms probably involved habitat partitioning between ground-dwelling and arbo- real species. The subsequent vegetational and climatic his- 402 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 15 tory of western North America (greatly simpli- fied) includes warming from late Oligocene to middle Miocene, followed by gradual, fluctuat- ing cooling through the Pleistocene, and a shift from summer-wet to summer-dry climate. By the Pliocene the woody flora was depauperate, with extinctions most marked in broad-leaved lineages with paratropical and subtropical affin- ities (Wolfe 1979). Latitudinal and altitudinal zo- nation became pronounced in the later Tertiary, accompanied by a differentiation of forest types into several adaptive zones (Axelrod 1976:fig. 7). As the rising Cascade Range cast a more pro- found rain shadow on the interior, humid forests became confined to the windward, west slope of the Cascades and lands to the west—the modern range of Monadenia, s.s. At this time (late Miocene—Pliocene), a sub- stantial allopatry between the two existing stocks of Monadenia may have arisen, with the Shastelix-Corynadenia stock inhabiting the drier, interior regions, and Monadenia, s.s., perhaps exploiting a tendency toward eurytopy in humid environments. Equability of climate may also have been a factor. Parapatry like that which now occurs in the Klamath Mountains, where M. (Monadenia) fidelis and M. (M.) se- tosa inhabit riparian woodland while M. (Shas- telix) churchi occupies drier, more exposed slopes and rockslides, may have been common where the environments interfingered. (In this connection, according to Whittaker [1961] and Axelrod [1976], the modern Klamath Mountains preserve a climate and vegetation more like that of the later Cenozoic than any other region in the west.) Some time in this interval Monadenia became extinct in central Oregon. On a graph of mean annual temperature ver- sus mean annual range of temperature (compare Wolfe 1979), only the thermal range of Shastelix overlaps that of present-day mixed mesophytic forest (Fig. 18). Because the John Day fossil oc- currence represents only one datum, we do not know what the total Oligocene thermal range of Monadenia might have been; but certainly the large, fidelis-type snails of the typical subgenus have shifted away (toward the right—equable— side of the graph) from the thermal zone of mixed mesophytic forest. Shastelix has shifted less, if at all. The narrow overlap of thermal ranges of Monadenia, s.s., and Shastelix par- allels their narrow geographic zone of parapatry and suggests progressive range/habitat differ- entiation along a climatic gradient. In contrast, the thermal range of Corynadenia overlaps those of both other subgenera extensively. It occupies a considerable range of mean annual temperatures, but a limited zone of mean annual temperature ranges. If Corynadenia differen- tiated from Shastelix along a climatic gradient, it was presumably not chiefly a gradient of mean temperature and equability. The autapomorphies of Corynadenia can be interpreted as adaptations to the rockslide/rock- crevice habitat and prolonged summer drought. The range of Corynadenia receives the least precipitation of any zone inhabited by Mona- denia (Elford 1970). Reduction or partial loss of the genital apparatus is associated with adapta- tion to xeric environments in other groups of Helminthoglyptidae (Gregg 1960; Bequaert and Miller 1973). Retention of granulose microsculp- ture on the teleoconch is evidently a paedomor- phy, and is most strongly developed in species (M. hillebrandi, M. circumcarinata) with pae- domorphic carination of the adult shell. Small size and low-spired, lenticular shape are other crevice-related characteristics that occur in some species and races of Corynadenia. Additional evidence for the relationship of temperature and distribution in Monadenia is seen in the close correspondence between max- imum summer and minimum winter isotherms and the range limits of Shastelix and Coryna- denia (Fig. 1). The 36 C mean maximum July isotherm practically coincides with the *‘down- hill’ limit of the genus in the Klamath Moun- tains and Sierra Nevada, and the —4 C mean minimum January isotherm, more loosely, with the “‘uphill’’ limit. (At this scale, the isotherms are highly generalized and each stands for a range of related microhabitat temperatures.) The southern limit of Corynadenia occurs just north of where these two isotherms pinch together. Similarly, between the southern limit of Shas- telix and the northern limit of Corynadenia, the July isotherm bulges eastward, almost but not quite intersecting the January isotherm. At this point, the 33 C mean maximum July isotherm, which most closely coincides with the very southern limit of Shastelix, is actually east (‘uphill’) of the —4 C January isotherm. If it persisted long enough, a thermal configuration like this could have produced vicariance within ROTH 20 5 Monadenia, s. ON gr sc | eu 10 ca Mean Annual Temp. (°C) cb si : DISTRIBUTION, ANATOMY, VARIATION OF MONADENIA TROGLODYTES er Sa ee 5 fess I EN Te SAT oe mn ee WE Ee n eS ee 1 ee eye ee SEE EE eet ey 1 5 10 15 20 25 30 Mean Annual Range of Temp. (°C) Ficure 18. Graph of thermal ranges of the three subgenera of Monadenia in relation to temperature parameters. Stippling, Monadenia, sensu stricto; diagonal hachure, Corvnadenia; horizontal hachure, Shastelix. Dot-dashed line encloses thermal range of present-day mixed mesophytic forest of eastern Asia (after Wolfe 1979). Abbreviations for stations defining the subgeneric polygons (in California unless otherwise indicated): bb, Big Bar; ca, Cape Blanco, Oreg.; cb, Clallam Bay | NNE, Wash.; dn, Dunsmuir; du, Dudley: er, Elwha Ranger Station, Wash.; eu, Eureka; gf, Giant Forest; gg, Grant Grove; gr, Graton 1 w; ms, Mount Shasta: or, Orleans; pr, Prince Rupert, Brit. Col.; sc, Scotia; si, Sitka, Alaska; sr, Sonora Ranger Station; ti, Tatoosh Island, Wash.; vr, Yreka. Thermal data from Elford (1970), Wolfe (1979). the Shastelix-Corynadenia stock. The present configuration of the isotherms must be short- lived in geologic terms, particularly in the face of the great climatic fluctuations of the Pleisto- cene; while it may now enforce the geographic separation of Shastelix and Corynadenia, their initial cleavage undoubtedly took place earlier. Alone, or in combination with climatic factors, Miocene or later topographic developments could have fostered the vicariance. (Pliocene through Holocene volcanic rocks of the High Cascades Group overlap the edges of the north- ern Sierra Nevada and eastern Klamath Moun- tains, separating the metamorphic and plutonic rocks of these two provinces for a distance of about 80 km [Hammond 1979]. They were ex- truded initially as basaltic flows and later as an- desitic, cone-building eruptions, filling a struc- tural trough and lowland between the Klamath and Sierran blocks. For a calcicolous ancestral Monadenia, these volcanic rocks could have been a barrier leading to allopatric differentia- tion or interrupting a cline along which some differentiation had already proceeded.) Given the complexity of relations between land snails and their environment, it is unlikely that a single event can be pinpointed as the cause, but no details of phylogeny, distribution, or thermal re- lations contraindicate vicariance in the general geographic setting of the present. ACKNOWLEDGMENTS I gratefully acknowledge the friendship and guidance of Allyn G. Smith (1895-1976), who nourished my interest in terrestrial malacology and helped to frame an early draft of this study. In addition I thank S. E. Hirschfeld, R. L. Seib, and D. C. Rudolph, all of whom, knowing of my interest in Shasta County snails, contributed material for study. Special thanks are owed to P. U. Rodda, D. H. Kavanaugh, H. W. Schorn, P. H. Pressley, and D. W. Taylor for helpful discussion; to A. J. Cain, G. M. Davis, and W. L. Lee for stimulus in several areas; to A. J. 404 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 15 Ferreira for radular preparation; to W. K. Emer- son and H. Feinberg for lending historically sig- nificant material; and to L. L. Eng, E. V. Coan, and L. H. Roth for aiding the investigation in various ways. Susan Middleton contributed photographic assistance. Steven Sechovec ren- dered most of the anatomical drawings. LITERATURE CITED ALBERS, J. P., AND J. F. ROBERTSON. 1961. Geology and ore deposits of East Shasta copper-zince district, Shasta County, California. U.S. Geol. Surv. Prof. Pap. 338. 107 p. AXELROD, D. I. 1976. History of the coniferous forests, Cal- ifornia and Nevada. Univ. Calif. Publ. Bot. 70:1—62. Batt, I. R. 1976. Nature and formulation of biogeographical hypotheses. Syst. Zool. 24:407—430. Bequaert, J. C., AND W. B. Mitter. 1973. The mollusks of the arid southwest, with an Arizona check list. Univ. Arizona Press, Tucson. BERGGREN, W. A., AND J. A. VAN COUVERING. 1974. The late Neogene. Elsevier Sci. Publ. Co., Amsterdam. Berry, S.S. 1940a. A proposed dichotomy of the snail-genus Monadenia. Bull. South. Calif. Acad. Sci. 38:203-205. . 1940b. Nine new snails of the genus Monadenia from California. J. Entomol. Zool., Pomona Coll. 32: 1-17. BINNEY, W. G. 1885. A manual of American land shells. U.S. Natl. Mus. Bull. 28. 528 p. , AND T. BLAND. 1869. Land and fresh water shells of North America. Part I. Pulmonata Geophila. Smithson. Misc. Collect. 194. 316 p. BisHop, M. J. 1979. A new species of Caracolus (Pulmonata: Camaenidae) from the Oligocene of Nebraska and the biotic history of the American camaenid land snails. Zool. J. Linn. Soc. London 67:269-284. Boycott, A. E. 1934. The habitats of land Mollusca in Brit- ain. J. Ecol. 22:1-38. Breure, A. S. H. 1979. Systematics, phylogeny and zoo- geography of Bulimulinae (Mollusca). Zool. Verh. Rijks- mus. Nat. Hist. Leiden 168. 215 p. BREWER, W.H. 1930. Up and down California in 1860-1864. The journal of William H. Brewer, Professor of Agriculture in the Sheffield Scientific School from 1864 to 1903. Edited by F. P. Farquhar. Yale Univ. Press, New Haven. Brown, R. W. 1959. A bat and some plants from the upper Oligocene of Oregon. J. Paleontol. 33:125-129. CHANEY, R. W. 1948. The ancient forests of Oregon. Condon Lecture Publ., Oregon State Syst. High Educ. 56 p. CHARNOV, E. L. 1979. Simultaneous hermaphroditism and sexual selection. Proc. Natl. Acad. Sci. U.S.A. 76:2480- 2484. Comrort, A. 1951. Observations on shell pigments of land pulmonates. Proc. Malacol. Soc. London 29:35-43. CooGANn, A. H. 1960. Stratigraphy and paleontology of the Permian Nosoni and Dekkas Formations (Bollibokka Group). Univ. Calif. Publ. Geol. Sci. 36:243-316. Cooper, J. G. 1869. On the distribution and localities of west coast helicoid land shells, &c. Am. J. Conchol. 4:211-240. . 1873. On the law of variation in the banded California land shells. Proc. Calif. Acad. Sci. 5:121-125. . 1879. Notes on some land-shells of the Pacific slope. Proc. Am. Philos. Soc. 18:282—288. . 1887. West coast Pulmonata; fossil and living. Bull. Calif. Acad. Sci. 2:355-376. DEMIRMEN, F., AND J. W. HARBAUGH. 1965. Petrography and origin of Permian McCloud limestone of northern Cal- ifornia. J. Sediment. Petrol. 35:136—154. Dicer, J. S. 1906. Description of the Redding Quadrangle. U.S. Geol. Surv. Geol. Atlas of United States, folio 138. 14 p. Diver, C. 1932. A method of determining the number of whorls of a shell and its application to Cepaea hortensis Mull. and C. nemoralis L. Proc. Malacol. Soc. London 19:234—239. ELDREDGE, N. 1979. Cladism and common sense. Pp. 165— 198 in J. Cracraft and N. Eldredge, eds., Phylogenetic anal- ysis and paleontology. Columbia Univ. Press, New York. Etrorb, C. R. 1970. Climate of California. Climatography of the United States, no. 60-4, U.S. Dept. Commerce. 57 p. EVERNDEN, J. F., AND G. T. JAMES. 1964. Potassium-argon dates of the Tertiary floras of North America. Am. J. Sci. 262:945-974. GAFFNEY, E. S. 1979. An introduction to the logic of phy- logenetic reconstruction. Pp. 79-111 in J. Cracraft and N. Eldredge, eds., Phylogenetic analysis and paleontology. Columbia Univ. Press, New York. GouLbD, S. J. 1969. An evolutionary microcosm: Pleistocene and Recent history of the land snail P. (Poecilozonites) in Bermuda. Bull. Mus. Comp. Zool. 138:407-532. GRATACAP, L. P. 1901. Catalogue of the Binney and Bland Collection of terrestrial air-breathing mollusks of the United States and Territories. ... Bull. Am. Mus. Nat. Hist. 14:335-403. GrecGc, W. O. 1960. Derivation of the Helminthoglyptinae with particular reference to the desert forms. Am. Malacol. Union Inc. Annu. Rep. 25(for 1959):45—46. HAMMOND, P. E. 1979. A tectonic model for evolution of the Cascade Range. Soc. Econ. Paleontol. Mineral. Pacific Coast Paleogeog. Sympos. 3:219-237. HANNA, GD. 1920. Fossil mollusks from the John Day basin in Oregon contained in the Condon Museum of the Univer- sity of Oregon. Univ. Oregon Publ. 1(6):1-8. , AND A. G. SmiTH. 1933. Two new species of Mon- adenia from northern California. Nautilus 4:79-86. HELLerR, J. 1975. The taxonomy, distribution and faunal succession of Buliminus (Pulmonata: Enidae) in Israel. Zool. J. Linn. Soc. 57: 1-57. HENDERSON, J. 1936. The non-marine Mollusca of Oregon and Washington—Supplement. Univ. Colo. Stud. 23:251- 280. HENNIG, W. 1966. Phylogenetic systematics. Univ. Illinois Press, Urbana. HIBBARD, C. W., C. E. Ray, D. E. SAvAGE, D. W. TAYLOR, AND J. E. GuILDING. 1965. Quaternary mammals of North America. Pp. 509-525 in H. E. Wright, Jr., and D. G. Frey, eds., The Quaternary of the United States. Princeton Univ. Press, Princeton, New Jersey. IRWIN, W. P. 1972. Terranes of the western Paleozoic and Triassic belt in the southern Klamath Mountains, Califor- nia. U.S. Geol. Surv. Prof. Pap. 800-C:103-111. , AND S. P. GALANIS, JR. 1976. Map showing limestone and selected fossil localities in the Klamath Mountains Province, California and Oregon. U.S. Geol. Surv. Misc. Field Studies Map MF-749. ROTH: DISTRIBUTION, ANATOMY, VARIATION OF MONADENIA TROGLODYTES 405 KAVANAUGH, D. H. 1972. Hennig’s principles and methods of phylogenetic systematics. Biologist 54:115—127. . 1979. Studies on the Nebriini (Coleoptera:Carabidae), III. New Nearctic Nebria species and subspecies, nomen- clatural notes, and lectotype designations. Proc. Calif. Acad. Sci. 42:87-133. KINKEL, A. R., JR., W. E. HALL, AND J. P. ALBERS. 1956. Geology and base-metal deposits of West Shasta copper- zine district, Shasta County, California. U.S. Geol. Surv. Prof. Pap. 285. 156 p. KUCHLER, A. W. 1977. The map of the natural vegetation of California. Pp. 909-939, map, in M. G. Barbour and J. Ma- jor, eds., Terrestrial vegetation of California. John Wiley and Sons, New York. Majsor, J. 1977. California climate in relation to vegetation. Pp. 11-74 in M. G. Barbour and J. Major, eds., Terrestrial vegetation of California. John Wiley and Sons, New York. PitsBry, H. A. 1895. Guide to the study of helices [continuation]. Manual of Conchology, ser. 2, 9:161—336. 1900. Note on the varieties of Epiphragmophora mormonum. Nautilus 13:128. 1939. Land Mollusca of North America (north of Mexico). Acad. Nat. Sci. Phila., Monogr. 3, 1(1): 1-573. 1948. Land Mollusca of North America (north of Mexico). Acad. Nat. Sci. Phila., Monogr. 3, 2(2):521-1113. PLATNICK, N. 1977. Cladograms, phylogenetic trees, and hy- pothesis testing. Syst. Zool. 26:438-442. Raup, D. M. 1966. Geometric analysis of shell coiling: gen- eral problems. J. Paleontol. 40:1178-1190. RAYNER, R. W. 1970. A mycological colour chart. Common- wealth Mycological Institute, Kew, Surrey. RoTuH, B. 1972a. Rare and endangered land mollusks in Cal- ifornia. The Resources Agency of California, Dept. Fish and Game, Inland Fisheries Admin. Rep., no. 72-10. 21 p. 1972b. Rare and endangered land mollusks in Cali- fornia. Sterkiana 48:4—16. . 1975. On the affinities of Monadenia churchi Hanna and Smith (Gastropoda: Stylommatophora). Bull. South. Calif. Acad. Sci. 74:93-94. 1979. Thoughts on Monadenia and other snails of northern California (abstract). West. Soc. Malacol. Annu. Rep, 11:13: , AND L. L. ENG. 1980. Distribution, ecology, and reproductive anatomy of a rare land snail, Monadenia se- tosa Talmadge. Calif. Fish Game 66:4—16. SmitH, A. G. 1957. Snails from California caves. Proc. Calif. Acad. Sci., ser. 4, 29:21—-46. . 1960. Note on Trilobopsis tehamana (Pilsbry), a rare northern California land snail. Veliger 2:97. 1970. American Malacological Union Symposium, Rare and Endangered Mollusks. 6. Western land snails Malacologia 10:39—46. SoLeEM, A. 1975. Notes on Salmon River Valley oreohelicid land snails, with description of Oreohelix waltoni. Velize 18: 16-30. 1978. Classification of the land Mollusca. Pp. 49-97 in V. Fretter and J. Peake, eds., Pulmonates, vol. 2A, Sys- tematics, evolution and ecology. Academic Press, London. STEARNS, R. E. C. 1900. Fossil land shells of the John Day region with notes on related living species. Proc. Wash. Acad. Sci. 2:651-660. TALMADGE, R. R. 1960. Color phases in Monadenia fidelis (Gray). Veliger 2:83-85. TweepieE, M. W. F. 1961. On certain Mollusca of the Ma- layan limestone hills. Bull. Raffles Mus. 26:49-65. VAN GOETHEM, J. L. 1977. La systématique des Urocyclinae (Mollusca, Pulmonata, Urocyclidae). Malacologia 16:133- 138. WaAHRHAPFTIG, C., AND J. H. BirMAN. 1965. The Quaternary of the Pacific mountain system in California. Pp. 299-340 in H. E. Wright, Jr., and D. G. Frey, eds., The Quaternary of the United States. Princeton Univ. Press, Princeton, New Jersey. WALTON, M. L. 1970. Longevity in Ashmunella, Monaden- ia, and Sonorella. Nautilus 83:109-112. Wess, G.R. 1952. Pulmonata, Xanthonycidae: Comparative sexological studies of the North American land-snail, Mon- adenia fidelis (Gray)—a seeming ally of Mexican helicoids. Gastropodia 1:[1—3]. 1966. Courtship between Monadenia fidelis and M. infumata. Nautilus 79: 103-105. WHITTAKER, R. H. 1961. Vegetation history of the Pacific Coast states and the ‘‘central”’ significance of the Klamath region. Madrono 16:5-—22. Wo FE, J. A. 1979. Temperature parameters of humid to mesic forests of eastern Asia and relation to forests of other regions of the northern hemisphere and Australasia. U.S. Geol. Surv. Prof. Pap. 1106. 37 p. , AND D. M. Hopkins. 1967. Climatic changes record- ed by Tertiary land floras in northwestern North America. Pp. 67-76 in K. Hatai, ed., Tertiary correlations and cli- matic changes in the Pacific. 11th Pac. Sci. Congress, To- kyo, 1966, Sympos. 25. Woop, H. E., II, R. W. CHANEY, J. CLARK, E. H. COLBerrT, G. L. JEpsEN, J. B. REESIDE, JrR., AND C. Stock. 1941. Nomenclature and correlation of the North American con- tinental Tertiary. Bull. Geol. Soc. Am. 52:1-48. 406 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 15 APPENDIX: LOCALITIES Localities for material studied are listed below, grouped according to geographic area. The num- bers assigned are for purposes of this study; samples are on deposit in the mollusk collection of the California Academy of Sciences. Map quadrangle names refer to the most recent edition of the U.S. Geological Survey 7.5- and 15-minute Series (Topographic). The number in brackets following the collection data is the total number of specimens examined from that locality; it does not always agree with sample size numbers cited in the text because not every character could be measured for every specimen and because many of the computations utilized adult shells or anatomies only. Distances are cited as received in the collectors’ notes and have not been converted to the metric system. CALIFORNIA: Siskivou County: Hornbrook Quadrangle I Along creek 1.5 mi sw of Hornbrook. G D. Hanna, Apr. 1928 [5]. SW'4 sec. 18, T. 46 N, R. 6 W, 1000 ft up Klamath River from mouth of Shasta River. D. E. Marsh, 1931 [2]. Among rocks about half-way up a spur of Badger Mountain on west side of Shasta River canyon not far above its mouth. E. P. Chace, 29 Sep. 1937 [3]. (Type-locality, !. t. chaceana.) Shasta River near junction with Klamath River. E. P. Chace [3]. Sec. 25(?), T. 46 N, R. 7 W, banks of Shasta River, 2-3 mi from mouth, in shaded rockslides. A. G. Smith, 11 Sep. 1934 [5]. CALIFORNIA: Siskivou County: Yreka Quadrangle 6 7 CALIFORNIA: 8 9 Chastain’s Quarry, west of Gazelle. E. P. and E. M. Chace, 28 Sep. 1937 [3]. [?] ‘‘Near Yreka.” E. J. Elliott, autumn 1933 [5]. Shasta County: Bollibokka Mountain Quadrangle Near McCloud River bridge, 30 mi NE of Redding. D. C. Rudolph, B. Martin, S. Winterath, 9 Apr. 1979 [2]. Sec. 32, T. 36 N, R. 3 W, west slope of Bollibokka Moun- tain, 0.75 mi E£ of bridge across McCloud River. J. W. Durham, E. C. Allison, 18 Apr. 1964 [5]. Sec. 5, T. 35 N, R. 3 W, slope below Samwel Cave, above McCloud River arm of Shasta Lake. R. L. Seib, 11 Mar. 1978 [32]. SeCa ly SSuNeRe SEW. Samwell Caves chal) Secu2—5 (=UCMP loc. 1008). E. L. Furlong [8]. (Type-locality, M. t. troglodytes.) Sec. 5, T. 35 N, R. 3 W, Samwel Cave, in gravel slope filling grotto at south end, ch. 2, sec. 4(=UCMP loc. 1009). E. L. Furlong [2]. Sec. 5, T. 35 N, R. 3 W, Samwel Cave, with bone matter, 50 ft in from twilight zone. R. E. Graham, S—6 Jan. 1957 [2]: Sec. 5, T. 35 N, R. 3 W, Samwel Cave, twilight zone floor. R. de Saussure, 10 May 1957 [2]. Sec. 5, T. 35 N, R. 3 W, Samwel Cave. R. de Saussure, 5 June 1957 [2]. Sec. 5, T. 35 N, R. 3 W, Samwel Cave, entrance to main cave. R. de Saussure, 5 June 1957 [1]. Sec. 5, T. 35 N, R. 3 W, Samwel Cave, surface, second exit. R. de Saussure, A. Dacey, 5 June 1957 [4]. Sec. 5, T. 35 N, R. 3 W, Samwel Cave. N. Slusser, 4—7 June 1957 [2]. Sec. 5, T. 35 N, R. 3 W, within 25 ft of entrance to Samwel Cave. R. E. Graham, 8 June 1958 [31]. 20 33 34 35 36 337 38 Sec. 5, T. 35 N, R. 3 W, Samwel Cave, bone chamber pit containing bone and rubble matrix, 7 to 12 inches depth. R. E. Graham, 10-14 June 1958 [2]. Sec. 5, T. 35 N, R. 3 W, Samwel Cave. R. E. Graham, Dec. 1958 [6]. Sec. 5, T. 35 N, R. 3 W, Samwel Cave. R. E. Graham, 18 June 1959 [1]. Sec. 5, T. 35 N, R. 3 W, Samwel Cave. R. E. Graham, 19 Dec. 1959 [2]. Sec. 5, T. 35 N, R. 3 W, Samwel Cave and vicinity. D. C. Rudolph, B. Martin, S. Winterath, 9 Apr. 1979 [9]. SE'4 sec. 7, T. 35 N, R. 3 W, limestone outcrop, summit of Hirz Mountain, 2 mi sw of Samwel Cave. R. E. Graham, June 1960 [2]. SE'4 sec. 6, T. 35 N, R. 3 W, Ellery Creek on Gilman Road, 15 mi NE of junction with Interstate Hwy. 5, lime- stone rock slides. A. G. Smith, 17 May 1973; B. Roth, Mar. 1980 [30]. SE'4 sec. 6, T. 35 N, R. 3 W, Ellery Creek, w side of McCloud River arm of Shasta Lake. R. L. Seib, 24 Mar. 1978 [5]. SW'4 sec. 21, T. 35 N, R. 3 W, Dekkas Rock at junction of Dekkas Creek with E£ side of McCloud River arm of Shasta Lake. R. L. Seib, 24 Mar. 1978 [38]. NW'4 NE'4 sec. 32, T. 35 N, R. 3 W, limestone outcrop 0.3 mi ssw of junction of Campbell Creek with E side of McCloud River arm of Shasta Lake. R. L. Seib, 24 Mar. 1978 [3]. Sec. 28, 29(?), T. 25 N, R. 2 W, Low Pass Creek. J. Gor- man, 29 Jan. 1953 [2]. SE'4 NW'4 sec. 4, T. 34 N, R. 2 W, caves at headwaters of Brock Creek drainage. S. E. Hirschfeld, July 1975 [19]. SE'4 NW'4 sec. 4, T. 34 N, R. 2 W, shelter below and to left of Goblin Shelter, headwaters of Brock Creek drain- age. S. E. Hirschfeld, July 1975 [8]. NE'4 SW'4 sec. 3, T. 34 N, R. 2 W, small cave in Hos- selkus limestone in gray rocks on E side of eastern NE- sw-trending ridge. S. E. Hirschfeld, 3 Apr. 1970 [1]. NW'4 NE4 sec. 8, T. 34 N, R. 2 W, cave between two limestone buttes at south end of Gray Rocks, above Pit River arm of Shasta Lake. S. E. Hirschfeld, Oct. 1975 [49]. (Type-locality, M. ¢. wintu.) SW'4 NW14 sec. 7, T. 34 N, R. 2 W, limestone quarry | mi N, % mi E of Brock Mountain Lookout, above Squaw Creek arm of Shasta Lake. R. L. Seib, 25 Mar. 1978 [39]. Squaw Creek. A. M. Strong, 1898 [3]. Squaw Creek. J. Gorman, 14-15 Mar. 1953 [5]. N ctr., SE'4 sec. 9, T. 34 N, R. 2 W, caves in Brock Creek limestone, above Pit River arm of Shasta Lake. S. E. Hirschfeld, July 1975 [7]. ROTH: DISTRIBUTION, ANATOMY, VARIATION OF MONADENIA TROGLODYTES 407 39 NW'4 SE sec. 9, T. 34. N, R. 2 W, lower limestone across from Brock Spring, above Brock Creek, above Pit River arm of Shasta Lake. S. E. Hirschfeld, July 1975 [13]. 40 NE'4 SW sec. 9, T. 34 N, R. 2 W, **Monadenia Cave” and vicinity, above Pit River arm of Shasta Lake. S. E. Hirschfeld, July 1975 [14]. 41 NE“4 SW sec. 9, T. 34. N, R. 2 W, ‘‘Elk Antler Cave,” above Pit River arm of Shasta Lake. S. E. Hirschfeld, July 1975 [20]. CALIFORNIA: Shasta County: Lamoine Quadrangle 42 SW'4 sec. 13, T. 34 N, R. 4 W, Shasta Lake Caverns. D. C. Rudolph, B. Martin, S. Winterath, 10 Apr. 1979 [1]. 43 SE'%4 SE% sec. 23, T. 34 N, R. 4 W, Potter Creek, along McCloud River arm of Shasta Lake. R. L. Seib, 11 Mar. 1978 [22]. 44 Sec. 23, T. 34 N, R. 4 W, Potter Creek Cave, ‘‘past kitch- en’? (=UCMP loc. 1055) [12]. CALIFORNIA: Shasta County: Project City Quadrangle 45 SE'4 sec. 4, T. 33 N, R. 4 W, Calveris Cement Co. quarry above Interstate Hwy. 5 just south of Shasta Lake bridge, 2.5 mi NNE of Mountain Gate. R. L. Seib, 11 Mar. 1978 [40]. CALIFORNIA: Shasta County: Anderson Quadrangle 46 Clear Creek, on road between Redding and Beegum. G D. Hanna, Sep. 1952 [1]. CALIFORNIA: Shasta County: quadrangle uncertain 47 Crystal Shasta Cave. K. Howard, 1958(?) [1]. 48 Chute Cave. R. de Saussure, Nov. 1957 [1]. 49 ‘Shasta County, Calif.,"” ex D. O. Mills collection, UCMP [3]. CALIFORNIA ACADEMY OF SCIENCES Golden Gate Park San Francisco, California 94118 2) 3.2% Gat > : a t 1 bah six ry i i padi 3aares ae watald ce | 2 = a z ~< a PROCEEDINGS OF THE Weeds Hola Macc poereersivemntonicionslil Vol. 42, No. 16, pp. 409-433, 2 figs., 9 tables CALIFORNIA ACADEMY OF SCIENCES FOOD HABITS OF SMALLER MARINE MAMMALS FROM NORTHERN CALIFORNIA By Robert E. Jones Museum of Vertebrate Zoology, University of California, Berkeley, California 94720 ABSTRACT: General food habits are discussed for 11 species of small marine mammals beach-cast in northern California. The collection method allowed tabulation only of the kinds, numbers, and percentages of fish, molluscan, and crustacean prey. Dominant food species for all male Zalophus californianus were the following: Merluccius productus (62.8 percent of occurrence), Engraulis mordax (23.8 percent), Scorpaenidae (6.3 percent), Porichthys notatus (3.2 percent). Octopus and Loligo also occurred in 7 (23.0 percent) of 30 California sea lions examined. Nine of 19 individuals of Eumetopias jubatus contained food remnants of fish; 10 new dietary items are reported. Seven of the nine also fed on invertebrates, including four kinds of cephalopods. Scorpaenidae 31.2 percent of occurrence), Merluccius productus (21.7 percent), Pleuronectidae (17.3 percent), and Chilara taylori (11.8 percent) were the principal fishes identified. Eight of 12 Phoca vitulina stomachs contained food (6 with fish, 2 with invertebrates). Embiotocids (41.9 percent of occurrence), Lycodopsis pacifica (27.9 percent), Pleuronectidae (9.3 percent), and Hexagrammos decagrammus (9.3 percent) were the dominant fishes. The only cephalopod identified from Phoca was Octopus. The stomachs of all seven individuals of Lagenorhynchus obliquidens examined contained food. Osmerid fishes, Porichthys, and juvenile rockfishes were the most frequent items identified. The presence of five genera of cephalopods suggests that Lagenorhynchus can take a wide variety of prey from several habitats. Juvenile Scorpaenidae, Engraulis, Merluccius, and Microgadus made up 97 percent of the diet of 20 individuals of Phocoena phocoena. The abundant cephalopod Loligo was a primary invertebrate food item and was ingested October 26, 1981 whole. INTRODUCTION The objective of this study is to summarize the literature on the diets of smaller marine mammals in the eastern Pacific and to report on the contents of 102 stomachs from five pinniped and six small cetacean species. The purpose has been to make a thorough scrutiny of published dietary knowledge and analyze stomachs of beach-cast specimens. Particular attention has been directed to reviewing all records on feeding controversy. It has been argued for many de- cades that marine mammals compete with com- mercial fishing industries (Anonymous 1901; Townsend 1918). Marine biologists in California have ques- tioned the impact of marine mammals on the state’s fisheries for at least 100 years (Redding et al. 1875: Rutter et al. 1904; California Division of Fish and Game 1927; Hedgpeth 1944; Ander- son 1960; Frey 1971; National Oceanic and At- mospheric Administration 1974). The report of the California Commissioners of Fisheries for the years 1874-1875 (Redding et al. 1875) stated [409] 410 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 16 clearly that sea lions and seals were protected by special enactment, with penalties imposed on any person who should kill or disturb the beasts. On the basis of the observed ‘‘hundred-fold”’ increase in the populations of seals and sea lions at the entrance to San Francisco Bay over the previous ten-year period, the Commissioners recommended the killing of nine-tenths of the existing population. It is unclear as to what ac- tion, if any, resulted from this recommendation. Scammon (1874) stated, ‘‘A few years ago great numbers of sea lions were taken along the coast of Upper and Lower California, and thousands of barrels of their oil were obtained.’’ The state- ment describes the period before the enactment of regulations protecting California pinnipeds. On the west coast of North America, most of the commercial seal and sea-lion killing stopped about 1860 (Bonnot 1928). The problem of a declining fishery was aired before another commission at San Francisco in 1899, and on 27 April 1899, permission was granted to kill sea lions along the coast, includ- ing lighthouse reservations (Bonnot 1928). The data from stomach inspections, gathered by Pro- fessor L. L. Dyche in 1899, were published by C. H. Merriam (1901a, 1901b). These publica- tions were an attempt toward a critical exami- nation of stomachs from slaughtered sea lions. A view held by many was that stomach analysis was utterly useless, for the observed fact was that sea lions pursued salmon through the Gold- en Gate as far as Sacramento (Anonymous 1901). Thousands were killed before the Treas- ury Department prohibited wholesale slaughter on government lands on 31 May 1899 (Merriam 190 1a). The destruction of sea lions was justified by the belief that the declining shad (Alosa sap- idissima), salmon (Oncorhynchus spp.), and striped bass (Morone saxatilis) fisheries would thereby increase, but little evidence was gath- ered to verify or deny such claims. The U.S. Fish Commission, the Secretary of the U.S. De- partment of Agriculture, and the New York Zoological Society opposed the decisions of the California Fish Commission (Anonymous [‘‘Steelhead”’ pseudonym] 1901). The controversy over control of sea lion pop- ulations pitted biologists C. Hart Merriam, Bar- ton W. Evermann, and William T. Hornaday against David Starr Jordan, Charles H. Gilbert, Harvey W. Harkness, and N. Baird Scofield (Starks 1918; Townsend 1918, 1919). The 1904 report of a Federal commission appointed to re- view the situation contains the first published list of stomach contents of a series of California sea lions (Zalophus) and northern sea lions (Eu- metopias) (Rutter et al. 1904). In 1914-1916, the Newcombe Commission gathered basic biological field data in British Columbia (Newcombe and Newcombe 1914; Newcombe et al. 1918). Twelve of the 14 north- ern sea lions examined contained intact herring; however, food items consumed by these sea lions throughout the year were not documented. Early in the 20th century, suggestions were made to harvest young sea lions, following the example of the Newfoundland harp-seal indus- try. Harp seals (Phoca) had been cropped an- nually for 100 years with an annual take of pups exceeding 200,000. Recommendations for con- trol of pinnipeds in California gained the support of many prominent biologists (Townsend 1919). Following an investigation at the lighthouse reservation on Ano Nuevo Island (Evermann 1921), it was stated that surplus bulls on this rookery could be killed (Evermann and Hanna 1925). This is the earliest example of a recom- mendation to harvest marine mammals along the California coast based on biological information. Because sea lions and seals are polygamous, it was felt that 10 percent of the excess males could be killed, but for economic reasons the cropping was not carried out (Rowley 1929). Target clubs obtained military surplus guns and ammunition and ‘‘practiced’’ on the Santa Cruz and San Mateo coast pinniped populations. Ap- parently this activity frightened the pinnipeds enough to cause their departure (Herb Steindorf, local rancher, pers. comm., 1969). Fishermen still ask permission to kill sea lions, and numer- ous reports of damage to the fishery continue to this day. Between the late 1920’s and 1972, when the Marine Mammal Protection Act was passed, certain marine mammals were fully protected, but commercial and sport fishermen (actively fishing from boats) could kill sea lions and har- bor seals interfering with their operations (Cal- ifornia Fish and Game Code, Sections 3002, 4500). Thus fishermen could protect their nets, tackle, and fish from damage by marine mam- mals. The California Fish and Game Commis- sion reserved the right to reduce the population ee JONES: FOOD HABITS OF SMALLER MARINE MAMMALS 411 and to require permits for educational display or scientific taking of sea lions or seals. The sea otter (Enhydra lutris), Guadalupe fur seal (Arc- tocephalus townsendi), and elephant seal (Mi- rounga angustirostris) were fully protected. State laws applied only to the seals, sea lions, and sea otter; no regulations concerning ceta- ceans were in effect until the enactment of the Marine Mammal Protection Act of 1972. This protection placed an immediate moratorium upon the taking and importation of all marine- mammal products into the United States (Na- tional Oceanic and Atmospheric Administration 1974). The oft-repeated complaint about the lack of life history data on most of our common species of marine mammals is still valid (Starks 1918; Anderson 1960; Peterson and Bartholo- mew 1967; Briggs and Davis 1972). Even the age at puberty and physical maturity of the Califor- nia sea lion (Zalophus californianus) is un- known (Harrison 1972). There are few published accounts of marine mammal life histories, and these studies seldom contain precise information on food habits (Bartholomew 1967; Orr and Poulter 1965, 1967; Peterson and Bartholomew 1967; Peterson and LeBoeuf 1969; Odell 1971; Seed 1972). General publications on marine mammals seldom contain significant or specific information on food habits (Sergeant and Fisher 1957; Slijper 1962; King 1964; Evans and Bastian 1969; Daugherty 1972; Orr 1972; Ridgway 1972). Notes on food habits of pinnipeds have been reported in the literature since the early work of L. L. Dyche in 1903 (Bonnot 1928, 1932a, 1932b, 1951; Scheffer and Neff 1948; Scheffer 1950a; Mathisen 1959; Mathisen et al. 1962; Thorstein- son and Lensink 1962; Fiscus and Baines 1966; Morejohn and Baltz 1970; Briggs and Davis 1972). The literature on distribution, food habits, and life histories of small cetaceans is less extensive, and wholly inadequate. While the federal status report to the Secretary of Commerce (National Oceanic and Atmospheric Administration 1974) is the most complete summary of studies to date, natural history data are lacking even for the most common species. Cetaceans have not had a reputation for eating commercially important fishes nor for harassing fishermen. Until 1972 there were no federal laws protecting or regu- lating the capture of small whales or porpoises. Recently large numbers of porpoises have been killed during tuna-fishing operations (Perrin 1970). Papers dealing with food habits of ceta- ceans are widely scattered (Scheffer 1950b, 1953; Brown and Norris 1956; Tomilin 1957; Wilke and Kenyon 1952, 1957; Wilke and Ni- cholson 1958; Norris and Prescott 1961; Fitch and Brownell 1968; Loeb 1972: Perrin et al. 1973). More study is needed on feeding rates and feeding phenomena (Sergeant 1968, 1969). Fast- ing periods and basic metabolic rates obviously affect calculations of the impact of marine mam- mals on the food resources of the sea. The role of marine mammals in overall ocean ecology needs further study. Current knowledge of food chains and trophic relationships of marine mam- mals has only recently been given attention by marine ecologists (Steele 1970). MATERIALS AND METHODS During this study, I examined 102 specimens found dead on California beaches (Table 1). The 11 species of marine mammals studied (Fig- ures | and 2) include two phocids, Mirounga angustirostris and Phoca vitulina, and three otariids, Eumetopias jubatus, Zalophus califor- nianus, and Callorhinus ursinus. The six ceta- cean species represent three families: Phocoe- nidae, Phocoena phocoena and Phocoenoides dalli; Delphinidae, Delphinus delphis, Grampus griseus, and Lagenorhynchus obliquidens; and Physeteridae, Kogia simus. Where feasible, specimens up to about 100 pounds (45 kg) were removed to the laboratory and data were recorded on standardized sheets (Norris 1961; Scheffer 1967). Where terrain or tidal conditions did not allow removal, carcasses were measured in place. All possible standard measurements were taken. The skull, baculum (when present), and stom- ach were collected from specimens too large to move. All material was labeled and taken to the laboratory for careful examination. Osteological material from each specimen was tagged and cleaned by standard museum techniques. Thir- teen stomachs were donated by personnel of the California Academy of Sciences After initial external cleaning, 25 stomachs were filled with cool tap water and measured to obtain an average stomach volume (Table 2). Each water-filled stomach was drained of its 412 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 16 TABLE I. NUMBER OF MARINE MAMMAL STOMACHS EXAMINED IN NORTH-CENTRAL CALIFORNIA, 1968-1973; (n = 102). A total of 68 stomachs contained food items; of these, 61 had fish remains, 33 invertebrates. Phocoe- Lagenorhyn- CETACEANS Phocoena noides chus Delphinus Grampus Kogia Total Examined 20 5 7 l | | 35 With contents 16 2 7 l 1 1 28 Empty 4 3 0 0 0 0 7 With fish 14 2 5 I 0 I 23 With invertebrates 8 0 3 l l l 14 PINNIPEDS Zalophus Eumetopias Phoca Callorhinus | Mirounga Total Examined 30 19 12 4 2 67 With contents 20 9 8 | 2 40 Empty 10 10 4 3 0 27 With fish 20 9 6 I 2 38 With invertebrates 7 7 2 2 0 19 With rock or plant material 1 3 0 0 0 4 contents into a graduated cylinder for accurate volumetric reading. Stomach contents were segregated, using a parasite-recovery technique of flotation and de- canting, and then washed repeatedly with clean water. This method of sedimentation-decanta- tion allows speedy recovery of all items. Small otoliths (lapilli) were occasionally detected, but all otoliths reported here were sagittae. Other workers (Fitch and Brownell 1968; Morejohn and Baltz 1970; Smith and Gaskin 1974) have screened gastro-intestinal materials through cheesecloth gauze or graded wire screens. Disarticulated bones were separated from in- ternal parasites with forceps. Fish bones and otoliths were dried overnight. Soft tissues were preserved in alcohol. The heavy fish otoliths (sagittae) were easily seen by moving the clear glass container over backgrounds of several different colors. Even the tiny otoliths (asterisci or lapilli) could be de- tected when they were gently moved by the cov- ering water. I removed the otoliths with a camel hair brush or fine forceps. They were carefully cleaned of mucus (film), dried, and stored in gel- atin pill containers. John E. Fitch, California Department of Fish and Game, provided iden- tifications. Most cephalopod beaks, which were stored in alcohol, were identified using the pictorial guide developed by the California Department of Fish and Game (Iverson and Pinkas 1971). Several types were identified by Clifford Fiscus, National Marine Mammal Laboratories, U.S. Department of Commerce. I later checked these samples using the keys developed by Clarke (1962) and by comparing them with squid beaks furnished by Jerome L. Spratt of the California Department of Fish and Game. Four hundred thirty-five cephalopod beaks and 2828 fish otoliths were identified. Thirty-one otoliths and 217 cephalopod beaks were not identifiable. I was able to identify most cephalopod beaks only to genus. Few stomachs contained intact fish. Fish skel- etal material was air-dried, and its volume was measured in a graduated cylinder (Table 9). Fish remains were identified with the aid of Clothier 1950; Roedel 1953; Clemens and Wilby 1961; Fitch and Lavenberg 1971; and Miller and Lea 1972. Common names used follow those of the American Fisheries Society (Robins et al. 1980) except for Atka-mackerel and jackmackerel, which follow Hubbs, Follett, and Dempster (1979). All recovered materials (specimens and contents) are deposited in the Museum of Ver- tebrate Zoology, Berkeley, or the California Academy of Sciences, San Francisco. A total of 35 cetacean and 67 pinniped car- casses was beach-cast along north-central Cal- ifornia. Voucher material has been collected and deposited at these institutions: CAS, California Academy of Sciences, San Francisco; MVZ, Museum of Vertebrate Zoology, University of California, Berkeley; HSC, Humboldt State University, Vertebrate Zoology Museum, Ar- cata; PORE, Point Reyes National Seashore, Point Reyes. JONES: FOOD HABITS OF SMALLER MARINE MAMMALS 413 Oo Farallon Isls. Phocoena Phocoenoides Lagenorhyncus Delphinus Kogia Grampus oe o x B O FIGURE |. CETACEAN Humboldt Co.—Phocoena: HSC 68-7, F, 2 Sep. 1968, 122 cm, 12 otoliths; HSC 73-4, M, 21 Aug. 1973, 150 cm est., 14 otoliths —Lagenorhvnchus: HSC 68-9, M, 26 Dec. 1968, 179 cm, 36 otoliths, | cephalopod beak. Sonoma Co.—Phocoenoides: MVZ 153258 (REJ 670), F, 28 June 1973, 102 cm, empty. Marin Co.—Phocoena: CAS 16602 (2385), M, 1 June 1973, 138 cm, 1083 otoliths; CAS 16603 (2385), 1 June 1973, 146 cm, empty; CAS 21380 (REJ 448), F, 3 July 1970, 158 cm, 15 otoliths —Phocoenoides: MVZ 153259 (REJ 678), F, 7 Aug. 1973, 186 cm, empty; CAS 16604 (2385), M, 1 June 1973, 198 cm, empty; CAS 16297 (2335), M, 21 Sep. 1972, 188 cm, 6 otoliths —Lagenorhyvnchus: MVZ 140845 (REJ 218), F, 9 Feb. 1970, 181 cm, no ID.—Delphinus: CAS 16242 (2340), F, 6 Nov. 1972, 168 cm, 11 otoliths, 4 cephalopod beaks. San Francisco Co.—Phocoena: CAS 16629 (2384), M, 26 May 1973, 140 cm, 2 otoliths; CAS 16572 (2398), F, 20 Aug. 1973, 137 cm, 17 otoliths —Grampus: MVZ 153257 (REJ 659), F, 20 May 1973, 275 cm, barnacle and hydroid. San Mateo Co.—Phocoena: CAS 15992 (2237), F, 19 July 1971, 143 cm, 33 otoliths; CAS 16609 (2390), F, 13 July 1973, 159 cm, 26 otoliths and 1 hake; CAS 16633 (2392), F, 13 July 1973, 126 cm, 4 otoliths, 35 pair cephalopod beaks and 13 whole Loligo.—Phocoenoides: CAS (REJ 674), F, 1 Aug. ——» eer ele 4k Monterey Co. Localities of cetacean beach-cast specimens in north-coastal California, 1968-1973. 1973, (79 in.), 223 cm,* 61 otoliths —Lagenorhvnchus: CAS 16593 (2336), M, 28 Sep. 1972, 190 cm, 47 otoliths; CAS 16342 (2380), F, 20 Apr. 1973, 193 cm,* 4 otoliths —Kogia: CAS 16635 (2382), M, 25 May 1973, 204 cm, 2 otoliths and 231 cephalopod beaks. Santa Cruz Co.—Phocoena: CAS 21381 (KB 19-73), M, 13 Apr. 1973, 134 cm, 17 otoliths; (KB 17-73), F, 1973, 104 cm, empty. Monterey Co.—Phocoena: CAS 21387 (REJ 687), F, 22 Sep. 1973, 107 cm, empty; CAS 21383 (REJ 673), M, 24 July 1973, 126 cm, 48 otoliths, 1 pair cephalopod beaks; CAS 21385 (REJ 661), F, 17 June 1973, 104 cm, 16 otoliths, 3 cephalopod beaks; CAS 21386 (REJ 654), F, 24 Apr. 1973, 137 cm, 15 otoliths; CAS 21389 (REJ 653), M, 25 Apr. 1973, 145 cm, 13 cephalopod beaks; CAS 21388 (REJ 450), ? sex, 24 July 1971, no tl, 127 otoliths, 52 cephalopod beaks; CAS 21384 (REJ 449), ? sex, 24 July 1971, 108 cm, empty; CAS 21382 (REJ 241), F, 6 June 1970, 172 cm, 16 cephalopod beaks.—Lage- norhynchus: CAS 21370 (REJ 652), F, 21 Apr. 1973, 187 cm, 4 otoliths; CAS 21378 (REJ 625), M, 14 May 1973, 177 cm, 147 cephalopod beaks; MVZ (REJ 237), F, 29 May 1970, 180 cm, 7 cephalopod beaks. * Indicates original data taken in inches. 414 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 16 Farallon Isls. Zalophus Eumetopias Phoca Callorhinus e pepo Mirounga FIGURE 2. PINNIPEDS Humboldt Co.—Zalophus: CAS 21401 (REJ 248), M, 16 June 1970, 256 cm, 17 otoliths; HSC 3206 (69-34), M, 1 June 1969, 233 cm,* 52 otoliths; HSC 3205 (REJ 69-33) M, 27 May 1969, 228 cm,* 122 otoliths; HSC 3204 (REJ 69-32), M, 23 May 1969, 249 cm,* 93 otoliths; HSC 3125 (REJ 69-31), M, 14 May 1969, 239 cm,* 152 otoliths —Eumetopias: CAS 21391 (REJ 683), F, 16 Sep. 1973, 111 cm, empty; CAS 21390 (REJ 672), M, 20 July 1973, 309 cm, empty; HSC, (REJ 655), F, 1 May 1973, 200 cm, stones only; HSC, (REJ 247), M, 14 June 1970, 257 cm, 76 otoliths, 2 cephalopod beaks; MVZ 140847 (REJ 249), F, 16 June 1970, 259 cm, 66 otoliths, 13 cephalopod beaks; HSC, (REJ 69-8), F, 16 Feb. 1969, 251 cm,* 2 otoliths, 1 cephalopod beak.—Phoca: CAS 21421 (REJ 684), F, 17 Sep. 1973, 147 cm, empty; HSC 1188 (REJ 69-24), F, 25 Apr. 1969, 149 cm, 13 otoliths; HSC, (REJ 68-38), M, 28 July 1968, 162 cm, 3 otoliths; HSC, (REJ 68-14), F, 6 Apr. 1968, 102 cm, 17 otoliths.—Mirounga: HSC 2165 (REJ 69-6), F, 19 Feb. 1969, 152 cm, | Apristurus brunneus egg; HSC 1356 (?), F, 5 May 1970, no tl, 26 otoliths. Sonoma Co.—Zalophus: CAS 21402 (REJ 680), M, 17 Aug. 1973, 238 cm, empty; CAS 21403 (REJ 660), M, 5 June 1973, 215 cm, 98 otoliths, 1 cephalopod beak.—Eumetopias: CAS 21392 (REJ 688), M, 23 Sep. 1973, 152 cm, 4 otoliths; CAS 21393 (REJ 679), F, 17 Aug. 1973, 229 cm, empty. Ize See \ en Ne Ni ORE \5 ye = ‘> Monterey Co. Localities of pinniped beach-cast specimens in north-coastal California, 1968-1973. San Francisco Co..—Callorhinus: MVZ 140846 (REJ 212), F, 5 Feb. 1970, 131 cm, 7 isopods. Marin Co.—Zalophus: CAS 21404 (REJ 700), M, 29 Sep. 1973, 210 cm, empty; PORE 138 (REJ 657), M, 9 May 1973, 230 cm, | otolith; CAS 21405 (REJ 641), M, I Mar. 1973, 220 cm, empty; CAS 16184 (2316), M, 19 July 1973, 158 cm, 1 otolith, 1 cephalopod.—Eumetopias: CAS 21399 (REJ 682), M, 6 Sep. 1973, 325 cm, empty; CAS 21395 (REJ 677), F, 7 Aug. 1973, 235 cm, broken otoliths and rocks; PORE 136 (REJ 676), M, 6 Aug. 1973, 280 cm, empty; CAS 21395 (REJ 668), F, 22 June 1973, 234 cm, empty; ?(REJ 637), 21 Oct. 1972, no tl, 22 otoliths, 4 cephalopod beaks; CAS 21397 (REJ 635), F, 16 Sep. 1972, 232 cm, empty; CAS 21396 (REJ 629), M, 22 July 1972, 228 cm, 59 otoliths, 5 cephalopod beaks; PORE 137 (REJ 453), F, 14 Aug. 1971, 226 cm, empty.—Phoca: PORE 214 (REJ 681), F, 29 Aug. 1973, 140 cm, empty; CAS 21423 (REJ 669), M, 22 June 1973, 97 cm, 3 Crago sp.; CAS 21422 (REJ 642), M, | Mar. 1973, 158 cm, empty. San Mateo Co.—Zalophus: CAS 21412 (REJ 698), M, 28 Sep. 1973, 160 cm, empty; CAS 21411 (REJ 697), M, 28 Sep. 1973, no tl, empty; CAS 21410 (REJ 695), M, 28 Sep. 1973, 218 cm, empty; CAS 21409 (REJ 693), M, 28 Sep. 1973, 213 cm,* empty; ?(REJ 692), M, 28 Sep. 1973, 152 cm (est.), 2 otoliths, 2 cephalopod beaks; CAS 21408 (REJ 691), F, 28 Sep. 1973, no tl, empty; CAS 21407 (REJ 690), M, 28 Sep. JONES: FOOD HABITS OF SMALLER MARINE MAMMALS 415 TABLE2. AVERAGE STOMACH VOLUME OF THREE SPECIES OF MARINE MAMMALS AS MEASURED BY WATER DISPLACEMENT Total length of animal (cm) Volume of stomachs (1) Species n Sex Range Average Range Average Zalophus Adults 6 M 198-256 223 9.42-19.84 12.63 Subadults 4 M 126-160 145 e292 S75 Eumetopias 3 M 280-325 305 22 .20—46.72 33.67 4 F 200-235 . 230 9.80-—23.74 17.78 Phocoena 3 M 126-145 136 1.10-1.43 1.28 5 F 95-159 120 7.00—2.25 2.44 1973, 196 cm, empty; CAS 16302 (2383), M, 27 May 1973, 122 cm, empty; MVZ 139211 (LGB 317), M, 24 Mar. 1970, no tl, 18 otoliths —Eumetopias: ?(REJ 699), F?, 28 Sep. 1973, 192 cm, 24 otoliths, | cephalopod beak; CAS 21400 (REJ 694), M, 28 Sep. 1973, no tl, empty; CAS 21398 (REJ 675), F, 2 Aug. 1973, 220 cm, | otolith—Phoca: ?(REJ 696), F, 28 Sep. 1973, 133 cm, 9 otoliths, 13 cephalopod beaks; CAS 21424 (REJ 689), F, 28 Sep. 1973, 139 cm, 15 Eptatretus stoutii eggs. Monterey Co.—Zalophus: CAS 21415 (REJ 686), M, 22 Sep. 1973, no tl, 7 otoliths; ?(REJ 685), M, 22 Sep. 1973, 142 cm, 18 otoliths; CAS 21420 (REJ 667), M, 19 June 1973, 195 cm, 7 otoliths; CAS 21417 (REJ 666), M, 19 June 1973, 256 cm, 3 otoliths; CAS 21416 (REJ 665), M, 19 June 1973, 198 cm, 48 otoliths, 40 cephalopod beaks: CAS 21419 (REJ 664), M, 19 June 1973, 151 cm, 221 otoliths; CAS 21418 (REJ 662), M, 19 June 1973, 126 cm, | otolith, 3 cephalopod beaks; CAS 21414 (REJ 647), M, 1 Apr. 1973, 115 cm, | otolith, 3 cepha- lopod beaks; UC tag 6588 (REJ 244), M, 6 June 1970, 125 cm, 2 otoliths, 3 cephalapod beaks; CAS 21413 (REJ 68-40), M, 12 June 1968, 233 cm,* 39 otoliths —Phoca: CAS 21427 (REJ 671), M, 13 June 1973, 98 cm, | otolith; CAS 21426 (REJ 663), F?, 19 June 1973, 87 cm, empty; CAS 21425 (REJ 646), F, | Apr. 1973, 142 cm, 10 cc fish bones, 3 cephalopod beaks.— Callorhinus: MVZ 153256 (REJ 645), F, 1 Apr. 1973, 121 cm, empty; ?(REJ 243), F, 6 June 1970, 101 cm, 12 otoliths, 14 cephalopod beaks; MVZ 138677 (REJ 69-36), F, 12 July 1969, 137 cm, empty. RESULTS ° Sixty-eight percent of the 102 stomachs ex- amined in this study contained material (Table 1), approximately the same percentage as re- corded for stomachs from collected living ma- rine mammals. For example, 331 of 437 stom- achs (76 percent) of the fur seals taken off California in 1966 contained food (Marine Mam- mal Biological Laboratory 1969), and 18 of 44 (41 percent) California sea lions taken recently in Oregon contained food (Mate 1973). Normally about 60 percent of the northern sea lions col- lected during daylight hours have food in their stomachs (Spalding 1964), but Mathisen et al. (1962) found food in 82 percent (114 stomachs). Forty percent of 1300 fur seal stomachs exam- ined in Alaska contained food (Scheffer 1950a). Fish otoliths or other dietary remains were recovered from 61 stomachs, and 33 stomachs had remnants of identifiable invertebrates (Ta- bles 3, 4, 5, and 9). The items in the stomachs included fish bones and otoliths, parasites, sea- weeds, fish egg cases, cephalopod tissue and beaks, eye lenses of fish and cephalopods, rocks, wood, and parts of other invertebrates. Fitch and Brownell (1968) have presented a valid case for the use of otoliths in determination of dietary habits. Other investigators also have found undigested parts such as otoliths and cephalopod beaks in the forestomachs of ceta- ceans (Rae 1965; Harrison et al. 1970; Iverson and Pinkas 1971; Loeb 1972; Smith and Gaskin 1974). Phoca vitulina, Harbor seal Harbor seals in the Aleutians contained fewer prey species than in other areas studied. Wilke (1957) examined seven harbor seals collected in March at Amchitka Island and found that octo- pus was the most frequent item, but that gadid and hexagrammid fishes made up the greatest volume of food present. A later study by Ken- yon (1965) found only octopus (Octopus sp.) and Atka-mackerel (Pleurogrammus monoptery- gius) in 11 seals at Amchitka Island. He theo- rized that harbor seals feed during the daylight hours as suggested by the freshness and large volumes of food in the stomachs. Kenyon failed to consider Wilke’s earlier study which indicat- 416 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 16 TABLE 3. TOTAL NUMBER OF FISH OTOLITHS FOUND IN 22 SPECIMENS OF CETACEANS, NORTH-CENTRAL CALIFORNIA, 1968-1973. Phocoena Phoecoenoides Lagenorhynchus Delphinus (14) (2) (5S) (1) No. Percent No. Percent No. Percent No. Percent Prey species otoliths total otoliths total otoliths total otoliths total Rockfishes (Scorpaenidae) 1017 (71.8) 0 = 8 (9.2) 0 ~ Northern anchovy 253 (17.8) 0 = 5 (7) 0 = Pacific hake 74 (5.8) 6l (91.1) 3 (3.5) 0 = Smelts (Osmeridae) 9 (0.15) 0 = 34 (39.0) 1 (9.1) Pacific tomcod 34 (2.3) 6 (8.9) 0 = 0 = Plainfin midshipman 2 (0.14) 0 = 34 (39.0) 0 = Flatfishes (Pleuronectidae) 13 (0.92) 0 = 1 (1.2) 0 = Medusafish 0 = 0) = 0 - 10 (90.9) Surfperches (Embiotocidae) 9 (0.63) 0 = 0 = 0 3 Spotted cusk-eel 2 (0.14) 0 = 0 = 0 = Jackmackerel 0 = 0 = 2 (2.3) 0 = Total 1413 (99.68) 67 (100) 87 (99.9) 11 (100.0) ed that the abundant rock greenling (Hexagram- mos lagocephalus) made up 96 percent of food volume from harbor seals. Total sample size during March for both Wilke (1957) and Kenyon (1965) was 10 specimens, which is hardly ade- quate for the construction of generalities. Yet Kenyon’s (1965) study is the basis for Morejohn and Baltz’s (1970) model for selective feeding. They looked at a single elephant seal and com- pared its feeding behavior to that of harbor seals at Amchitka. The sample sizes in these studies are barely adequate for comparison, and are in- adequate for feeding models. The diet of harbor seals varies greatly with season and location of populations. Pelagic, bot- tom-dwelling, and anadromous fishes have all been reported in its diet. Captain Scammon (1874) noted that the “‘Leopard Seal’’ pursued and devoured small fish. During a two-year study in Alaska (Imler and Sarber 1947), 166 (41.5 percent) of 400 harbor seal stomachs contained identifiable food items. In the Copper River flats of Alaska, 67 seals fed almost entirely on eulachon (Thaleichthys pa- cificus). Ninety-nine other specimens from southeastern Alaska had fed on walleye pollock (Theragra chalcogramma) and Pacific tomcod (Microgadus proximus) (22.6 percent), Pacific herring (Clupea harengus pallasii) (16.4 per- cent), and flounders (11.1 percent). Lesser num- bers (29.5 percent) of salmonids, sculpins, rock- fish, blennies, and skates were reported. Imler and Sarber (1947) also found shrimp and octopus (20.6 percent) in harbor seals from Alaska. Spalding (1964) collected harbor seals throughout the year in British Columbia; 57 of the 126 stomachs were empty. He found that minimal food was consumed during the June to September pupping season. In summer, the stomachs contained cephalopods (35.4 percent), rockfish (22.6 percent), and salmon (16.1 per- cent). Stomachs collected from September to December contained invertebrates (34.8 per- cent), herring (10.8 percent), and salmon (30.4 percent). Fish of commercial value composed 54 percent of the harbor seals’ diet on a yearly ba- SiS. In Puget Sound also the harbor seal is a gen- eralized feeder (Scheffer 1928). Scheffer and Sperry (1931) point out that fishes made up 93.6 percent, molluscs 5.8 percent, and crustaceans 0.6 percent of the total volume of harbor seal stomach contents. Only two percent of the har- bor seal stomachs contained salmon (Scheffer and Slipp 1944). Studies in Washington (Scheffer 1928; Scheffer and Sperry 1931; Scheffer and Slipp 1944; Seed 1972) revealed that the major prey species were flatfishes: English sole (Par- ophrys vetulus), flathead sole (Hippoglossoides elassodon), Pacific herring, Pacific tomcod, Pa- cific hake (Merluccius productus), sculpins (Leptocottus armatus, Myoxocephalus sp.), walleye pollock, surfperches (Cymatogaster ag- gregata, Rhacochilus sp.), Pacific cod (Gadus JONES: FOOD HABITS OF SMALLER MARINE MAMMALS 417 TABLE 4. ToTrAt NUMBER OF FISH OTOLITHS FOUND IN 38 SPECIMENS OF PINNIPEDS, NORTH-CENTRAL CALIFORNIA, 1968-1973. Phoca Zalophus Eumetopias Callorhinus Mirounga (6) (20) (9) (1) (2) No. Percent No. Percent No. Percent No. Percent No. Percent Prey species otoliths total otoliths total otoliths total otoliths total otoliths total Pacific hake | (2.3) $74 (62.8) S15) (21.7) 0 = 26 (96.3) Northern anchovy 0 218 (23.8) l (0.4) 12 (100) = a2 Rockfishes (Scorpaenidae) l (2.3) S77 (6.2) 79 (31.2) 0 = 0 = Flatfishes (Pleuronectidae) 4 (9.3) 1 (0.1) 44 (17.3) 0 = 0 = Spotted cusk-eel 0 = 6 (0.6) 30 (11.8) 0 = 0 bse Plainfin midshipman 0 = 30 (3.2) | (0.4) 0 = 0 we Sablefish 0 = 0 = 22 (8.7) 0 = 0 = Surfperches (Embiotocidae) 18 (41.9) 3 (0.3) 0 = 0 = 0 a Pacific herring 0 = 16 (1.7) | (0.4) 0 = 0 = Lingcod 0 = 1 (0.1) 13 (5.1) 0 = 0 = Blackbelly eelpout 12 (27.9) 0 ~ l (0.4) 0 = 0 = Jackmackerel 0 = 3 (0.3) 3 (JIG) 0) = 0 = Smelts (Osmeridae) 0 = 2 (0.2) 2 (0.2) 0 = 0 = Kelp greenling 4 (9.3) 0 = 0 = 0 = 0 = Pacific tomcod 2 (4.7) 1 (0.1) 0 = 0 = 0 = Brown cat shark 0 = 0 = 1 (0.4) 0 = 1 (3.7) Pacific hagfish 1 (2.3) 0 = 0 = 0 = 0 = Chinook salmon 0 = 1 (0.1) 0 = 0 = 0 = Queenfish 0 = 1 (0.1) 0 = 0 _ 0 _ Blacktail snailfish 0 ~ 0 = | (0.4) 0 = 0 = Total 43 (100) 914 (99.6) 254 (99.6) 12 (100) 27 (100) macrocephalus), and lingcod (Ophiodon elon- gatus). These authors list 13 kinds of crustacea and 4 kinds of mollusca from the stomachs of TABLE 5. MINIMUM NUMBER OF INDIVIDUAL FISH AND harbor seals. Harbor seals eat flounder, sole, herring, eel, goby, cod, whiting, squid, whelks, crab, and mussels (King 1964). Fishes, squid, octopus, and shellfish constitute the diet of har- bor seals in California (Daugherty 1972). Bonnot (1951) indicated that the fishes, molluscs, and crustaceans consumed by harbor seals in Cali- fornia are usually slow-moving or sedentary forms. The above authors do not specify scien- tific names nor document sources for their di- etary information. I examined stomachs of 12 harbor seals of which eight with food had eaten eight kinds of fish (Merluccius productus, Microgadus proxi- mus, Lycodopsis pacifica, Sebastes spp., Hex- agrammos decagrammus, Embiotoca jacksoni, Phanerodon furcatus, Glyptocephalus zachi- rus), one kind of octopus (Octopus sp.), and one kind of shrimp (Crago sp.). Embiotocid perch constituted 41.9 percent of my sample, com- pared with 11 percent in Washington (Scheffer and Sperry 1931). One harbor seal (REJ 689) had CEPHALOPODS FOUND IN ELEVEN SPECIES OF MARINE MAm- MALS, NORTH-CENTRAL CALIFORNIA, 1968-1973. Cephalopods Fishes Mini- Minimum mum no. no. indi- indi- viduals viduals repre- No. repre- No. sented beaks sented otoliths Zalophus 32 51 476 922 (8*) Eumetopias 11 27 132 258 (5*) Phoca 12 16 23 43 (1*) Callorhinus 7 14 6 12 Mirounga 0 0 13 26 Phocoena 92 168 TID 1429 (16*) Phocoenoides 0 0 34 67 Lagenorhvnchus 86 155 45 88 (1*) Delphinus 3 4 6 11 Kogia 112 Pal 1 2 Grampus 0 0 0 0 Totals 355 652 1448 2858 (31) * Unidentifiable. 418 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 16 TABLE 6. NUMBER OF INDIVIDUAL FISH, GROUPED BY NICHE, FOUND IN THE STOMACHS OF MARINE MAMMALS FROM NorTH-CENTRAL CALIFORNIA, 1968-1973. Schooling Bottom- Inshore Sample (open dwelling (school- size water) (rocky) ing) Zalophus 30 296 52 121 Eumetopias 19 28 97 3 Phoca 12 2 24 0 Mirounga 2 13 0 0 Callorhinus 4 0 0 6 Phocoena 20 52 13 646 Lagenorhynchus if 3 22 20 Phocoenoides 5 33 0 0 Delphinus I 5 0 l Grampus | 0 0 0 Kogia | 0 1 0 ingested Pacific hagfish (Eptatretus stoutii) egg cases (15 eggs). I observed that Pacific hagfish or Pacific lamprey (Lampetra tridentata) was a prey item of seals near river mouths in northern California. Scheffer and Sperry (1931) note that two percent of stomachs in which items oc- curred contained Pacific lamprey. At birth, harbor seals average 81.6 cm in length (Bigg 1969). I examined two pups (98 and 97 cm) which contained solid food: an otolith (Sebastes spp.) in one stomach and a number of shrimp (Crago sp.) in the other. This suggests that harbor seals catch their own food at an early age. In summary, knowledge to date indicates that harbor seals feed on shallow-water fishes and bottom-dwelling invertebrates (Tables 1, 6 and 9). Mirounga angustirostris, Elephant seal Pike and MacAskie (1969) reported hagfish eggs (Eptatretus) and probably digested hagfish remains in an elephant seal from Canada. Huey (1930) pointed out that several of the fish species found in elephant seal stomachs inhabit water from 50 to 120 fathoms (ca. 91 to 219 m) deep. Small sharks (Squalus, Cephaloscyllium), skates (Raja), rays (Myliobatidae), and ratfish (Hydro- lagus) have been reported as food items in ele- phant seals (Kenyon and Scheffer 1955; King 1964; Daugherty 1972; Seed 1972). Huey (1925) found that three of four Guadalupe Island ele- phant seals contained squid (Loligo). Contrary to the report of Morejohn and Baltz (1970), teleost fishes had been reported from el- ephant seals. Huey (1930) reported a single bass, and Daughterty (1972) stated that elephant seals eat rockfish. Freiburg and Dumas (1954) found a dead adult elephant seal in Oregon which may have died from bones of Pacific hake (Merluc- cius productus) blocking the internal nasal re- gion. More information is needed on fish ecology and fish population abundance before the nature of feeding behavior of elephant seals can be clar- ified. The elephant seal may be a selective feeder (Morejohn and Baltz 1970), but seasonal, sex, and age factors need to be resolved. Commercial catches are the reference for fish abundance by Morejohn and Baltz. However, commercial fish catches are a poor indication of fish abundance when most reported fish are of commercial value (Bell 1971). Anyone accompanying commercial vessels realizes that many unreported fish are caught besides those marketed. All fishes More- john and Baltz found were bottom or rock- dwelling species (Chilara, Porichthys, Sebastes, and Lyopsetta or Glyptocephalus). Elephant seals may feed during particular pe- riods of the day. If they are nocturnal feeders, they may catch hake high in the water column, but diurnal feeding would suggest that hake are taken in deep water (Nelson and Larkins 1970). Studies are needed to determine when and where elephant seals feed. Large fish otoliths may indicate offshore feeding by the elephant seal. To date, few elephant seals have been exam- ined for stomach contents. One of my seals, an immature female (HSC—1970) found dead in Humboldt County on 5 May 1970, contained 26 otoliths from Pacific hake, representing at least 12 adult fish. Hake migrate north along the coast in spring and summer and might be expected to be present in marine mammal diets at that time of year. Pacific hake form massive schools just above the bottom and show a pronounced daily vertical migration (Nelson and Larkins 1970). Most adult hake are located beyond the conti- nental shelf at depths of 230-410 m. Although Pacific hake are present in commercially abun- dant numbers, they are not being exploited cur- rently by American fisheries. Pacific hake ranked second in abundance in California larval surveys (Ahlstrom 1965). My other specimen, also an immature female, was found in Trinidad, Humboldt County, 19 Feb. 1969 (HSC-69-6). It contained a single egg JONES: FOOD HABITS OF SMALLER MARINE MAMMALS 419 case of the brown cat shark (Apristurus brun- neus), Which was identified using the description and photos in Cox (1963). This species of shark is found in deep water from British Columbia to Baja California. Eumetopias jubatus, Northern sea lion The northern sea lion has been studied more intensively than most other North Pacific pin- nipeds. This sea lion occurs along the eastern Pacific coasts from Alaska to California, where its relationships to commercial fisheries have been studied extensively. In general, these stud- ies reveal that fish and cephalopods are the pre- ferred foods. In 1899 L. L. Dyche inspected 25 sea lion stomachs from near Monterey Bay, California. All specimens referred to by Dyche contained squid or octopus. The original manuscript Dyche sent to C. H. Merriam in 1901 did not specify which sea lion was involved. This partially handwritten docu- ment mentioned only the common identification “California Sea Lion’? (Unpublished manu- script, Dyche 1901. C. H. Merriam file at Mu- seum of Vertebrate Zoology, University of Cal- ifornia, Berkeley). When Merriam (1901a, 1901b) first published Dyche’s data, he did not indicate whether the northern sea lion (Eumetopias) or the California sea lion (Zalophus) was involved. Later, Dyche (1903) used the name **Zalophus californianus Lesson’’ beneath the general title of *‘Food for California Sea Lions,’ suggesting that California sea lions were examined. Thus it was long believed that all sea lions examined by Dyche were Zalophus. Briggs and Davis (1972) have pointed out that at least some of Dyche’s specimens were Eumetopias jubatus. Since only 7 of these 25 sea lions deposited at the University of Kansas are extant today, it is not possible to make positive identifications for all of Dyche’s specimens (R. S. Hoffmann, Cu- rator Univ. of Kansas, pers. comm. [1973]). Al- though the 7 extant specimens are northern sea lions, it is possible that all 25 of the originally collected specimens may not have been this species. Northern sea lions have been reported to feed at night (Rowley 1929; Bonnot 1951; Mathisen 1959; Mathisen et al. 1962; Spalding 1964; Seed 1972; Mate 1973). Fiscus and Baines (1966) sighted feeding groups of up to several thousand animals 8-22 km out in Unimak Pass, Alaska. It would be of interest to know if the individuals observed feeding returned to the hauling-out area each afternoon with engorged stomachs. Daytime feeding behavior has been noted by many fishermen, specifically long-line and drag- boat operators (Kenyon 1952a). An interesting examination at the cannery dock at the mouth of the Klamath River was reported by Bonnot (1951). Two half-grown northern sea lions were killed, and only lam- preys were found in their stomachs. J. C. Snyder also examined sea lions at river mouths and identified the remains of lampreys in their stom- achs (Kenyon 1952b). More recently, Jameson and Kenyon (1978) reported that 82 percent of observed feeding behavior at the Rogue River, Oregon, was on lampreys. Rutter et al. (1904) presented data on 18 north- ern sea lions (6 males and 12 females) from north-central California. The eight female sea lions containing food were taken from Ano Nue- vo Island in July or August. Five male sea lions from Pt. Arena ate at least 147 fish but only 5 squid. The majority of their food was fish (257 fish present in the 13 animals). The only ceph- alopod material was from five sea lions from both areas. This low incidence of cephalopods does not agree with the findings of Dyche (1903), but the identity of Dyche’s specimens is in doubt. In contrast, in southern California Rutter et al. (1904) found only 39 fish but thousands of cephalopods in a sample of 24 Zalophus. They concluded that the northern sea lion feeds chief- ly on fish, and the California sea lion on ceph- alopods, and that both feed opportunistically. Bonnot (1928) examined two northern sea lion stomachs from San Miguel Island, California, on 20 June 1928. The adult male was empty but the female contained three greenish eggs of a skate or shark. Bonnot stated that this was an old, blind sea lion and perhaps she was dying. In northern California, at the Saint George Reef rookery (Del Norte County), Bonnot found thousands of shells of a tiny pelecypod embed- ded in sea lion feces. Other investigators have not reported analysis of scats, perhaps because of the difficulty of locating adequate samples (Bonnot 1928). The literature contains a composite list of 32 fishes from stomachs of northern sea lions (Ta- ble 7). Because of the nonspecific categories of some fish names, I have not attempted to pro- duce an accurate species list. Only the papers No. of stomachs, kinds of food 19—empty, 1—ceph. beak, 1-10 Ibs., sandlance trace-starry flounder, sculpin, 1-20 Ibs. halibut, cod, flounder, pollack, 4—large stones 8-empty, 15—salmon, cod, halibut, pollack, 326—empty, 28—fish invertebrates, 20— squid and octopus, 10—no ID fish, 154— stones and gravel 20—empty, 31—fish invertebrates, 73—rock and sand, 24—no ID 2—empty, I—salmon, ““cod’* and “‘bass”’ 50—empty, 4—fish and squid, 1—herring, 6—empty, 1—salmon, 12—herring, |—clam- shell, 1—crab, octopus 6—empty, 3—herring, 2— rockfish, 2—fish/ octopus, 1—octopus, I—rockfish, I1—squid and rockfish, |—squid 213—empty, 75—fish (17 spp.), 49—no ID, milk 11—lost at sea, 22—fish (15 spp.), 2—no ID fish, I—clamshell and fish, 9—rocks and 5—empty, 13—fish, 6— 420 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 16 TABLE 7. SUMMARY OF FooD CONTENTS OF Eumetopias (modified from Pike (1958) with recent additions). Season No. of stomachs Locality and year Source (size and/or sex) St. Paul I., Alaska July 1949-1951 (6) 22 (2 females) Sitka to Kodiak I., May, July, Aug. (4) 23 Alaska 1945-1946 (adults) 4— octopus Kodiak to Krenitzin May-July 1959 (8) 382 I., Alaska (372 males, (7 spp.), 21— 10 females) Chernaburg I., May, July 1958 (7) 114 Alaska (46 males, (7 spp.), 61— 51 females, 17 yearlings) fish, S—milk Beresford I., B.C. Aug. 1913 (2) 3 (adults) Scott ds. Bae: June, July 1956 (5) 56 (adults) 1—octopus Scottie baGe June 1957 (5) 8 (4 females, I—no ID fish 4 males) Barkley Sound, B.C. Dec. 1915 (2) 14 (11 males, 3 females) Barkley Sound, B.C. Feb., Apr. 1958 (5) 14 (13 females, | male) skate, hake Isnor Rock, B.C. July, Aug. 1957 (5) 3 (young males) British Columbia Feb.—Dec. 1959 (9) 393 (equal numbers, males/females) or kelp Offshore California— Mar., Sep. 1958-1963 (10) 34 N. Pacific (7 males, 15 females) pebbles Pt. Arena and July—Aug. 1901 (1) 18 Ano Nuevo L., Calif. (6 males, 12 females) squid and octopus JONES: FOOD HABITS OF SMALLER MARINE MAMMALS 421 CONTINUED. TABLE 7. Season Locality and year Source Ano Nuevo I., Calif. ‘several years ago” (3) prior to 1918 North-Central 1968-1973 (11) California No. of stomachs, No. of stomachs (size and/or sex) kinds of food 15 7—empty, 8—rock, sardines, salmon, 3—fish and squid (14 females, 1 young male) 19 10—empty, 9—fish, 7— (7 males, . squid and octopus, 2— 12 females) rocks (1) Rutter et al. 1904. (2) Newcombe et al. 1918. (3) Starks 1918. (4) Imler and Sarber 1947. (5) Pike 1958. (6) Wilke and Kenyon 1952. (7) Mathisen et al. 1962. (8) Thorsteinson and Lensink 1962. (9) Spalding 1964. (10) Fiscus and Baines 1966. (11) Current study 1973. of Wilke and Kenyon (1952), Spalding (1964), and Fiscus and Baines (1966) present scientific names and volumetric determinations which en- able me to present a well-documented dietary list. Pike (1958) also presented a table with stom- ach contents. All of these data are updated and presented as Table 7. My study adds 10 genera of fishes to those previously reported from northern sea lions, as follows: Microstomus, Parophrys, Careproctus, Lyopsetta, Eopsetta, Glyptocephalus, Por- ichthys, Engraulis, Spirinchus, and Chilara. These 10 genera constituted 31.1 percent of the otoliths found in Ewmetopias. All northern sea lion stomachs which con- tained cephalopod beaks also had remains of irom, 2 to, 13 species of fish (Table 5). Four species of cephalopods were identified: Loligo opalescens, Octopus sp., Chiroteuthis sp., and Onychoteuthis sp. Most beaks were so thor- oughly digested that specific identifications were impossible. One male sea lion (REJ 629) had eat- en at least 3 octopus plus 13 species of fish. More surprising than the variety of prey eaten was the fact that the sea lion had been eating during the breeding season when most Ewmme- topias males fast (Spalding 1964). In my study demersal fish were found in six of the nine stomachs containing fish. When the 127 identified fishes from northern sea lions are grouped according to schooling (open-water), bottom-dwelling (rocky), and inshore-schooling species (Table 6), it is apparent that the northern sea lion feeds mainly on bottom-dwelling fishes. The rather high incidence of rocks in the stom- achs also suggests a bottom-feeding habit (Ta- bles I and 9). Zalophus californianus, California sea lion California sea lions make annual north-south migrations along the Pacific Coast of North America. Adult and subadult males move north- ward during September and October after the breeding season (Bonnot 1928; Fry 1939; Orr and Poulter 1965, 1967; Bartholomew 1967; Pe- terson and Bartholomew 1967; Peterson and LeBoeuf 1969; Odell 1971) and return south in March to the more southern breeding rookeries. Virtually nothing is known about the feeding behavior of migrating California sea lions. Pe- terson and LeBoeuf (1969) indicated that influx- es of sea lions into northern areas are correlated with periods of abundance of food, but they did not document their statement. No published studies have been reported on California sea lions between Monterey Bay and the Oregon border. B. R. Mate (Oregon State Univ., pers. comm. [1971]) collected 44 male California sea lions in Oregon, but he has not yet identified the fish otoliths. From the data 422 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 16 presented by Mate (1973), one can calculate that 40.9 percent of the combined sample of delib- erately collected sea lions contained food. My method of obtaining stomach data showed that 66.6 percent of the beached California sea lions contained food. Fiscus and Baines (1966) examined six Cali- fornia sea lions taken during a recent study on fur seals. The stomach volume was recorded; thus the estimate for feeding rates and food in- take to body weight can be determined more clearly. Loligo opalescens (common squid), un- identified squid, northern anchovy, and Pacific hake were found in these pelagic California sea lions. Several observations have been made of feed- ing sea lions. Fink (1959) reported a single ob- servation, on 25 Feb. 1959, in Monterey Bay of California sea lions attacking a school of Pacific sardines. Although feeding on sardines is surely not uncommon, his vivid description of harbor porpoises controlling the fish school and the sea lions feeding on the periphery of the school is the only published account of such behavior. Ryder (1957) reported feeding aggregations of pinnipeds and birds. At the Farallon Islands, California, an adult male California sea lion was seen repeatedly eating several jackmackerel during the daylight hours on 17 and 19 Septem- ber 1973 (T. James Lewis and Barbara Lewis, Point Reyes Bird Observatory, pers. comm.). At Cerros (Cedros) Island, Baja California, a fe- male California sea lion was observed feeding beneath the surface by Bonnot (1932b), who watched this cow eat at least six large flying fish (Exocoetidae). Scheffer and Neff (1948) noted that the anal- ysis of only 58 California sea lion stomachs had been reported in the literature. Of these, Dyche’s 25 specimens from the Monterey area are either misidentified or of questionable iden- tity. Scheffer and Neff examined four sea lion stomachs from southern California. Two fe- males were empty, but the other female con- tained evidence of at least 21 small squid. The single male found dead near La Jolla on 26 Nov. 1943 had 36 nearly whole Pacific herring plus fragments representing 30 other herring. All Cal- ifornia sea lions analyzed by Scheffer and Neff were from south of Point Conception. Rutter et al. (1904) examined stomachs of 24 California sea lions, 13 with food, in July and August 1901 at southern California localities. The eight females each had 100 to 300 small squid parts. Squid pens were food remnants in the stomachs of three of the five males. The re- maining stomach contents consisted of hake, rockfish, ratfish, unidentified small fish, and milk. Bonnot (1928) reported on these same specimens and stated that 5 had eaten fish and 11 had eaten squid. Starks (1918) pointed out that stomachs of two of the breeding bulls that Rutter et al. examined did not contain any food. According to a recent survey by scientists at the Scripps Institution of Oceanography and presented to the California Senate Fact Finding Committee on Natural Resources (Anderson 1960), 24 of 30 stomachs from Zalophus con- tained food. The only identified items were fish otoliths, but many unidentified cephalopod beaks also were present. Carl L. Hubbs (Scripps Institution of Oceanography) and John E. Fitch (California Dept. of Fish and Game, Long Beach, pers. comm. [1973]) revealed that this study was done by the late Art Kelly in southern California and northern Mexico. The 424 fish otoliths which Kelly recovered were identified by Fitch as representing 24 kinds of fishes. Pa- cific hake were found in 17 of the 24 stomachs and constituted 48.1 percent of the total otoliths. The other fishes were cusk-eels, midshipmen, and species of rockfish. Sixty-six (15.6 percent) of the otoliths were of Pacific mackerel, ancho- vy, perch, and white croaker. These sea lions also fed extensively on squid and octopus, as represented by beaks in their stomachs. Briggs and Davis (1972) spent 500 hours aboard sport and commercial salmon boats in Monterey Bay from 14 April to 22 September 1969. They observed seven instances of preda- tion on salmon by California sea lions. Of the hooked fish, 4.1 percent were lost to sea lions. I found 10 dead California sea lions along Mon- terey Bay during the commercial salmon season (April to September). I also located an additional 20 California sea lion carcasses farther north (Figure 1). I found a single female California sea lion north of Monterey Bay. Only one salmon otolith was present among the 922 otoliths (461 fish represented) from my sample of California sea lions. Male California sea lions feed on a variety of schooling fishes (Pacific hake, anchovy, rock- fish, flatfish, cusk-eel, midshipmen, herring, JONES: FOOD HABITS OF SMALLER MARINE MAMMALS 423 lingcod, jackmackerel, salmon, and osmerids). Pacific hake and anchovy make up 86.6 percent by frequency of occurrence of otoliths from sea lion stomachs. Schooling fishes, both inshore and open-water types (417 otoliths) were found in California sea lions (Table 6). Callorhinus ursinus, Northern fur seal Wilke and Kenyon (1957) identified five species of fish from 204 seals (114 with food) collected from the Bering Sea and St. Paul Is- land. Seals collected at sea contained large num- bers of capelin (Mallotus villosus) and walleye pollock (Theragra chalcogramma). Only one salmon (Oncorhynchus sp.) was found in the stomachs. Three seals killed on land at St. Paul Island collectively held one salmon (Oncorhyn- chus sp.), one walleye pollock, and two sandfish (Trichodon). Indian hunters took 41 seals (13 empty) 30 miles (about 48 km) off Washington in 1930. Although the stomach contents were digested, squid eyes and beaks were reported from 21, and identifiable herring vertebrae were present in 15 stomachs (Schultz and Rafn 1936). Clemens and Wilby (1933) looked at 25 stomachs from the west coast of Vancouver Island and reported that 8 contained squids, 9 had _ sal- mon, and the rest had small schooling fish. No data on the volume of the stomachs or the number of empty stomachs were reported. Hanna (1951) recorded fur seals in the Gulf of the Farallons during February and April. Seals were described as competitors with W. I. Follett as he dip-netted for myctophid fish (Tarleton- beania, Symbolophorus), sablefish (Anoplo- poma fimbria), and red Irish lord (Hemilepido- tus hemilepidotus). Hanna also theorized that fur seals, porpoises, and sea birds were all feed- ing on pteropods (Mollusca). More recently 437 stomachs from pelagic fur seals taken off California were examined (Ma- rine Mammal Biological Laboratory 1969). An- chovy, saury, hake, and squid constituted 98 percent of the total food volume. Scheffer (1950a) reviewed the dietary litera- ture on fur seals. He presented data on only two northern fur seal stomachs from California, one of which contained an unidentified bird and the other had fed on Pacific saury (Cololabis saira). Fur seals found on California beaches usually have little food in their stomachs. Likewise a fur seal from Southeast Farallon Island contained no food in its stomach (REJ 212), but had seven isopods (Riggia?) which are external parasites on fish. One beach-cast fur seal had 7 beaks of the common squid and 12 northern anchovy otoliths in its stomach (REJ 243). Phocoena phocoena, Harbor porpoise Although the diet of harbor porpoises in Cal- ifornia waters is poorly known, herring, small cods, soles, and squid are food items of harbor porpoises generally (Ridgway 1972). Scammon (1874) wrote, ‘““They feed upon fish, and are oc- casionally taken in seines that are hauled along the shores of San Francisco Bay by the Italian fishermen.” Harbor porpoises seldom are sighted more than 20 miles (about 32 km) offshore and usually are seen near harbor entrances (Fiscus and Nig- gol 1965). Local fish abundance and seasonal fish movements affect the diet of these porpoises (Rae 1965). In Scottish waters, Rae (1965) examined 45 porpoises from November to March and 7 ad- ditional ones in the summer months. A few in- vertebrates were found which Rae thought might have been taken incidentally with other food items. Fish or fish remains were recognized in 41 of the 43 stomachs with recognizable food. One of the two remaining stomachs contained milk and the other the remains of a very small cephalopod. Ten species of fish were present, with herring (Clupea harengus) and whiting (Gadus merlangus) the most common. Most fish were less than 25 cm in length, with the largest individuals 35 cm. These harbor porpoises had been trapped in nets set for cod or salmon. British naturalists have recorded food habits of harbor porpoises in the North Sea for more than 100 years, and small fish, mainly clupeoids (65 percent) and gadoids (30 percent), consti- tute the major foods taken (Rae 1965). Rae con- cluded that harbor porpoises take pelagic forms of fish. Tomilin (1957) found benthic fish pre- dominating in the diet of harbor porpoises from the Black Sea and Sea of Azov. In a study in the Bay of Fundy, small schooling fishes (Clu- pea harengus, Gadus morhua, Scomber scom- brus) were principal food items for harbor por- poises (Smith and Gaskin 1974). These schooling fishes accounted for 78 percent of the total diet. Tomilin (1957) listed dietary items of harbor porpoises in the Black Sea, where 4000 stom- 424 achs were inspected. These porpoises fed upon eight benthic species and six pelagic species of fish. The pelagic fish were consumed when they occurred in large dense schools. One of two har- bor porpoises found at Pt. Barrow, Alaska, con- tained bones of whitefish (Leucichthys) (Hall and Bee 1954). Pike and MacAskie (1969) re- ported a Canadian harbor porpoise caught in a gill net. This animal had one herring in its stom- ach. Scheffer (1953) inspected a female porpoise from Grays Harbor County, Washington, which had fed on 37 capelin. Its death apparently was caused by an American shad blocking its throat passages. Five years earlier Scheffer had found on the same beach another porpoise which also had choked to death on a fish. A female porpoise from Port Townsend, Washington, May 1950, had eaten five Pacific herring (Wilke and Ken- yon 1952). Orr (1937) reported that a porpoise apparently choked to death on a gray smoothhound shark (Mustelus californicus). This single report of sharks as a food item may be abnormal. Fink (1959) observed for 30 minutes several hundred harbor porpoises feeding on a school of Pacific sardines northwest of Pt. Pinos, Monterey Bay. He vividly described the attack and herding of the sardines, and counted from 5 to 12 fish eaten by an individual porpoise in its attack through the sardines. In my study, juvenile rockfish constituted 71.8 percent of the diet of harbor porpoises (Table 3). Five porpoises collected in June and July contained 1017 rockfish otoliths. Northern an- chovy was the second most frequent fish and was found in seven porpoises from April to Au- gust. Over two-thirds of all fish found in stom- achs of harbor porpoises live in open water or are inshore schooling species (Table 6). Juvenile Pacific hake, Pacific tomcod, rockfish, and northern anchovy accounted for 97 percent of all stomach otoliths found during my investiga- tion (Table 3). Invertebrate remains were found in 8 of the 20 porpoises examined (Table 9). A total of 141 Loligo opalescens beaks was identified from 168 cephalopod fragments. These beaks repre- sent at least 92 individual cephalopods com- pared to at least 712 fish represented by 1429 otoliths (Table 5). One harbor porpoise (CAS 2392) had 13 intact Loligo and an additional 35 pairs of beaks in its stomach. The intact bodies PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 16 of the 13 squids indicated harbor porpoises do not chew this food item. Phocoenoides dalli, Dall’s porpoise This porpoise is much more common than ear- ly records indicate. Brownell (1964) reported its occurrence in southern California waters in the winter. Lustig (1948) saw 10 or 12 porpoises feeding on baitfish, anchovy or sauries, on 13 July 1939, in the Anacapa Passage. Deep-water benthic fish and bathypelagic ceph- alopods were reported as major food items from a large sample of Dall’s porpoise stomachs from Japan (Wilke and Nicholson 1958). Eleven per- cent of the food volume was squid: Watasenia, Ommastrephes, and unidentified genera. Myc- tophidae (lanternfishes) composed 70 and 73 percent of the stomach contents in 1949 and 1952, respectively. Cowan (1944) took five (3 males and 2 fe- males) Dall’s porpoise off the coast of British Columbia in the summer of 1939. Four of the five stomachs were full of herring. Pike and MacAskie (1969) examined three males and two females from British Columbia, and they too found mostly herring or squid in three stomachs. Scheffer (1953) recorded the stomach contents of six Dall’s porpoises from Monterey and northward. These contained Pacific hake, squids (Loligo opalescens and unidentified species), jackmackerel, and unidentified fish. Two Dall’s porpoises from Alaska had fed only on capelin. Brown and Norris (1956) mentioned anchovy as a food item of the Dall’s porpoise. An adult por- poise taken in southern California waters had eaten at least 14 Pacific hake, 2 jackmackerel, and 13 cephalopods (Norris and Prescott 1961). These authors also noted the porpoise circling amid schools of sauries, probably feeding. Fis- cus and Niggol (1965) observed Dall’s porpoises off the north coast and collected five specimens off Cape Mendocino, California. Three females and one of the males had only squid beaks in their stomachs; the stomach of the other male was empty. The Dall’s porpoise is present in Monterey Bay all year. Stomach samples, examined each month, indicated that Pacific hake, rockfish, and squid are important food items. Loeb (1972) ex- amined 25 stomachs of Dall’s porpoise from Monterey Bay and found Pacific hake in 23, squid in 16. Most of the cephalopods present JONES: FOOD HABITS OF SMALLER MARINE MAMMALS were Loligo, with lesser numbers of Abraliop- sis, Gonatus, Onychoteuthis, and Octopus. Pa- cific hake, juvenile rockfish, and squid made up 93 percent of the total diet of the Dall’s porpoise from Monterey Bay (Table 5 in Loeb 1972). On 28 June 1973, an immature female Dall’s porpoise was found on the beach north of the University of California Marine Station at Bo- dega Bay. Presumably this animal (REJ 670, 102 cm total length) was dependent on its mother for nourishment, although no milk was noted in its stomach. I examined four adult Dall’s porpoises (2 males, 2 females), and only two had identifiable food remains. One (REJ 674) had 61 Pacific hake otoliths representing 31 fish in its stomach, and the other (CAS 2335) contained 6 juvenile Pa- cific tomcod. The two other porpoises had emp- ty stomachs (CAS 2384, REJ 678). Loeb (1972) did not mention Pacific tomcod as a dietary item from Dall’s porpoises but did note a wide variety of fishes (15 species) eaten by Dall’s porpoises from Monterey Bay. Pacific hake have been reported as important food for Dall’s porpoise (Scheffer 1953; Norris and Pres- cott 1961; Fiscus and Niggol 1965; Loeb 1972). The hakelike fish (Laemonema, family Moridae) occurs in the diet of Dall’s porpoise from Jap- anese waters (Wilke and Nicholson 1958). No cephalopods were found in any of the five Dall’s porpoise stomachs from my northern Cal- ifornia sample. Delphinus delphis, Pacific common dolphin Common dolphins seldom are sighted north of the California-Oregon border and are rare be- yond the 100-fathom (183-m) line (National Oceanic and Atmospheric Administration 1974). ~ Four Delphinus stomachs collected off Califor- nia contained fish and cephalopods (Fiscus and Niggol 1965). One female had unidentified fish otoliths and another stomach contained (by vol- ume) 60 percent squid, 25 percent saury, and 15 percent northern anchovy. One male dolphin taken at sea contained 90 percent Loligo and 10 percent saury. Another stomach contained 60 percent lanternfish (Myctophidae) and 40 per- cent squid (Gonatus sp., 20 percent; Onvcho- teuthis sp., 10 percent; unidentified squid, 10 percent) (Fiscus and Niggol 1965). Observations in California waters indicate that common dolphins are present in inshore 425 waters throughout the year (Norris and Prescott 1961). The major foods seem to be sardines, an- chovies, sauries, small bonito, and squid (Norris and Prescott 1961). Schmidt (1923) removed 15,191 otoliths from the stomach of one Delphinus. These otoliths represented five species of small fish (7596 in- dividuals). Frost (1924) looked at 4338 of these same otoliths and identified six species in three families. Myctophid fish accounted for 4324 of these 4338 otoliths. The fishes represented prob- ably did not constitute a ‘‘full’? meal for this dolphin (Fitch and Brownell 1968). Many common dolphins stranded in southern California had empty stomachs (Robert Brow- nell, Jr., Smithsonian Institution, pers. comm. [1970]). Fitch and Brownell (1968) examined two which had 133 and 119 otoliths. Anchovy re- mains (141 sagittae) were the most abundant. One dolphin had eaten 63 fishes representing six families. Anchovy, myctophids, and saury were represented in the other common dolphin ex- amined. These authors speculated that both Lagenorhynchus and Delphinus feed on meso- pelagic fish at depths exceeding 120 m (Fitch and Brownell 1968). The Pacific common dolphin (CAS 2340) I ex- amined had 11 otoliths assigned to two species; medusafish ([cichthys lockingtoni) and an os- merid. This specimen also had four Loligo beaks. Medusafish are most abundant around jellyfish and in the upper 150 ft (ca. 46 m) of the ocean (John Fitch, California Dept. of Fish and Game, pers. comm. [1973]). Apparently this dol- phin had fed near the surface. Lagenorhynchus obliquidens, Pacific white-sided dolphin This dolphin has received careful attention from west coast biologists during the last 25 years (Scheffer 1950b, 1953; Brown and Norris 1956; Houck 1961; Norris and Prescott 1961). These authors reported sardine, Pacific herring, salmon, northern anchovy, “‘scad’’ (=jack- mackerel), Pacific saury, squid, and jellyfish remnants as food items (Table 8). Large feeding aggregations of California sea lions, elephant seals, common dolphins, and Pa- cific white-sided dolphins have been observed by various authors (Norris and Prescott 1961; Fiscus and Niggol 1965). Mixed schools of com- mon and white-sided dolphins have been noted 426 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 16 TABLE8. STOMACH CONTENTS OF Lagenorhynchus obliquidens FROM THE WEST COAST OF NORTH AMERICA (CALIFORNIA). Locality Sex Date Source Type of contents w Trinidad je 11 Sep 1958 (3) Pacific sauries, ““scad” Humboldt Co. M 26 Dec. 1968 (6) Pacific sanddab, Pacific hake, eulachon, squid Marin Co. le 9 Feb. 1970 (6) no ID (fish) San Mateo Co. M 29 Sepa loT2 (6) shortbelly rockfish, northern anchovy, plainfin midshipman San Mateo Co. F 6 May 1973 (6) Pacific hake, jackmackerel w Santa Cruz M 4 Mar. 1959 (4) trace squid w Santa Cruz F 5 Mar. 1959 (4) northern anchovy, Pacific hake, squid Monterey Co. F 21 Apr. 1973 (6) night smelt Monterey Co. FE 29 May 1970 (6) squid, octopus w Pt. Piedras F 27 Feb. 1959 (4) northern anchovy, squid Nw Morro Bay M 2) iran), ISS (4) northern anchovy, Pacific hake, squid w Morro Bay FE 14 Feb. 1959 (4) trace squid s Anacapa I. M 27 Feb. 1952 (1) jellyfish, squid Santa Monica z 22 Aug. 1963 (5) Pacific hake, northern anchovy, white seaperch, cephalopods s San Pedro F 6 June 1953 (2) anchovies, squid Long beach F 21 Aug. 1967 (5) northern anchovy, Pacific hake, cephalopods, queenfish (1) Scheffer 1953. (2) Brown and Norris 1956. (3) Houck 1961. (4) Fiscus and Niggol 1965. (5) Fitch and Brownell 1968. (6) Current study 1973. off southern California where the white-sided dolphins are common, but north of San Francis- co only small groups of Lagenorhynchus have been seen (Fiscus and Niggol 1965). In southern California waters, white-sided dolphins have a distinct seasonal movement dur- ing the summer and fall which correlates with a shift from anchovies and squids as a principal food to the offshore schools of Pacific saury. This conclusion is from field observations only and from one stomach analysis (Brown and Nor- ris 1956). No seasonal migration of white-sided dolphins has been observed in northern Califor- nia. A migration of white-sided dolphins to northern California may correlate with oceanic current shifts and/or fish migratory patterns. Pacific sanddab, eulachon, night smelt, short- belly rockfish, and plainfin midshipman can now be added to the known species of food fishes reported from white-sided dolphins. I recovered 89 otoliths, only 3 of which represent open- water fishes. The great number of otoliths from inshore schooling fishes (Table 6) tends to indi- cate that this dolphin eats abundant, small fishes (osmerids, midshipman, and juvenile rockfish). The fact that white-sided dolphins feed on five kinds of cephalopods gives the impression that these invertebrates are a major food resource (Tables 5 and 9). My data indicate that the white- sided dolphin apparently is not dependent on cephalopods in northern California; only three of seven stomachs had molluscan remains. One dolphin (REJ 625) contained 147 of the 155 ceph- alopod beaks found. Grampus griseus, Risso’s dolphin Until recent stranding records were published (Orr 1966; Paul 1968; Stroud 1968; Hatler 1971) this dolphin was known only from the type of Grampus stearnsii (Dall) collected at Monterey Bay in 1873. All four previous specimens re- ported were males, and only two contained iden- tifiable food remains. Orr (1966) reported a Ris- so’s dolphin from San Mateo County that contained two pairs of beaks from the squid Dosidicus gigas. Stroud (1968) listed seven cat- egories of cephalopod beaks from a male Risso’s dolphin recovered in Washington: Onychoteu- this (1), Octopodoteuthis (1), Chiroteuthis (16), Gonatus (4), Gonatidae—form A (7), Gonati- JONES: FOOD HABITS OF SMALLER MARINE MAMMALS 427 dae—form B (13), and unidentified beaks (2). No fishes have been found in any Risso’s dolphin. I examined a single specimen at Southeast Farallon Island on 20 May 1973. All standard measurements (275 cm total length) were taken, and the reproductive tract, blood sample, and complete skeleton were saved. No parasites were discovered, but a goose-neck barnacle (Pollicipes polymerus) and a single hydroid (Aglaophenia latirostris) were in the stomach. Orr (1951) theorized that such material is swal- lowed incidentally as the animal thrashes close to shore. The barnacle and hydroid reported here were probably accidentally ingested. Both kinds of invertebrates are plentiful on the inter- tidal shores of the Farallon Islands. Hatler (1971) reported that plant material seems to be an ‘‘herbal remedy”? in Grampus. Risso’s dol- phin is probably an invertebrate feeder like Globicephala, which eats only squid as reported by Sergeant and Fisher (1957). Grampus lack teeth in the upper jaw and may feed solely on soft invertebrates. Kogia simus, Dwarf sperm whale Brownell obtained three dwarf sperm whales from Japanese waters. Although a good com- parative collection of Japanese fishes was lack- ing, Fitch and Brownell (1968) presented a table indicating that 18 different species of fish in 7 families were in the stomachs. Because two families (Macrouridae and Moridae) are inhab- itants of deep water, these authors speculated that Kogia feeds 800 ft (244 m) or more beneath the surface. Other authors have stated that most specimens of Kogia had eaten cephalopods or pelagic crustaceans (National Oceanic and At- mospheric Administration 1974). Scheffer and Slipp (1948) examined a male pygmy sperm whale specimen from Washington. It had 500 cc of nematodes and food fragments in its stomach; |] eye lenses (including 5 from squid), 15 squid beaks, 21 otoliths of an uniden- tified fish, | crab limb, maxillary bones of 2 specimens of Trichodon (?), and fragments of shrimp Pasiphaea, Pandalus, Pandalopsis. Kogia has a discontinuous distribution and is rather poorly known from the west coast. Few have been seen alive. Beach-cast specimens have provided material for anatomical, taxo- nomic, and distributional studies, but essentially nothing is known of its life history (Handley 1966). Because of difficulty of identification prior to Handley’s work (1966), most earlier literature could apply to either the pygmy sperm whale (K. breviceps) or the dwarf sperm whale. Only in recent studies can identification be trusted. One K. simus recovered at Thorton Beach, San Mateo County, by the California Academy of Sciences (CAS 2382, male, 204 cm total length) contained 217 beaks representing 112 individuals of these families: Octopoteuthidae, Onycho- teuthidae, Enoploteuthidae, Histioteuthidae, Gonatidae, Chiroteuthidae. In addition, a single pair of otoliths in its stomach was from a plainfin midshipman (Porichthys). This is the most northern record of Kogia simus along the west coast. DISCUSSION Statements about marine-mammal diets fre- quently have been vague and misleading and give a false impression of the role these animals play in the marine ecosystem. Much more in- formation is needed to understand predator-prey relationships in California waters (Steele 1970). California waters are rich in commercially im- portant fishes, and these fishes are well studied. However, it is also pertinent to consider the stocks of noncommercial fishes and the dynam- ics of such populations. The distribution of both predator and prey, including the availability of the latter throughout the year, is of prime im- portance. Adult marine mammals are usually migratory. The causes of these movements are largely un- known, but to some extent marine mammals re- spond to the seasonal abundance of food. The breeding cycle and its influence on food gath- ering have not been studied. The smaller whales are not associated with breeding rookeries as are pinnipeds. Cetaceans constantly search for prey and apparently lack the feeding-nursing cycle characteristically found in seals and sea lions. The behavior and feeding techniques of all marine mammals are poorly documented. Es- cape strategies of prey species are not well known. The sheer abundance of smaller prey items allows the escape of some individuals from predators. Some fishes and invertebrates reduce predation upon themselves by camouflage or disguise. Armor and spines must also help some prey to reduce losses. Representatives of the following common families of fishes were com- pletely absent from the marine mammal stom- 428 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 16 TABLE 9. STOMACH CONTENTS OF MARINE MAMMALS FOUND IN THIS STUDY (volume of dry fish bones, cephalopod identification, and miscellaneous items). Cephalopod beaks Fish bones Field number (cc) Upper Lower Identification Zalophus 68-40 950 ~ - — 317 LGB 400 - - _ 647 35 2 3 Octopus sp. 660, 4000 - I Octopus sp. 662 trace 3 - L. opalescens 664 250 - - 665 30 23 17 L. opalescens 666 5 - = = 667 trace - = — 685 120 - - — 1—Polinices shell, Ulva, eelgrass 686 820 - - — 692 45 - l Octopus sp. 244 = 1 3 L. opalescens Eumetopias 69-8 600 - | Chiroteuthis 247 - I I L. opalescens 249 ~ 6 7 L. opalescens 629 ~ 2 2 Octopus sp. 637 2850 2 2 L. opalescens 655 I—rock - | Onvychoteuthis 675 trace - - — 677 5—rocks, trace - - — 688 240, - — — 699 3200 = | Octopus sp. Phoca 69-24 trace - - — 646 40 2 I Octopus sp. 671 10 = = pas 696 400 10 3 Octopus sp. Callorhinus 212 7— external fish - = — isopods (Riggia ?) 243 - 1/ 7 L. opalescens Delphinus CAS 2340 3 l l L. opalescens Lagenorhynchus HSC 68-9 - 1 = Abraliopsis 237 - l 1 Octopoteuthis | 3 L. opalescens - | unknown Gonatidae 625 ~ l I Onvchoteuthis 79 66 L. opalescens = I Gonatus sp. 652 trace = = = Phocoena 241 ~ 9 7 L. opalescens 450, 15 35 17 L. opalescens 653 - 8 5 L. opalescens 661 - ~ l L. opalescens 673 75 I l L. opalescens JONES: FOOD HABITS OF SMALLER MARINE MAMMALS 429 TABLE 9. CONTINUED. Cephalopod beaks Fish bones Field number cc Upper Lower Identification HSC 73-4 180 _ = = CAS 2384 = 2 3 L. opalescens CAS 2385 (476) trace - = == CAS 2392 - 35 36 L. opalescens plus 13 whole Loligo CAS 2398 - 2 ~ Morotheuthis sp. achs examined in this study: Cottidae, Agoni- dae, Serranidae, Blenniidae, Clinidae, and Scombridae. These marine fishes are probably detected and perhaps discriminated as nonprey items. Possibly these fishes possess mechanisms to escape. Sick or injured marine mammals will starve rapidly. Animals not showing obvious causes of death presumably were sick or injured and sel- dom had anything in their stomachs. Specimens showing evidence of violent sudden death had intact squid or fish. Others had only digestion- resistant items such as beaks or otoliths. One harbor porpoise (CAS 2390) had swallowed a 46- cm Pacific hake, the anterior end of which was partially digested. No food item in this study showed evidence of having been chewed or cut by the consumer. Field observations indicate that food is torn apart by much head shaking, and the teeth only aid in the capture and holding of prey. The lack of specialized forelimbs with which to manipulate food is evident in the swal- lowing of whole food. Sexual dimorphism in the size of pinnipeds (Scheffer 1958) should be reflected in feeding rates and the species or size of prey selected, but no published data are available to substan- tiate this. However, such resource partitioning is shown between species (Table 6). It appears that prey selection in marine mam- mals is specialized, and this conclusion is sup- ported by the available data. Comparison of the food of Phoca with that of Eumetopias and Zal- ophus (Tables 1, 4, and 7) shows less dependence on pelagic fishes by Phoca. Perch, eelpout, and greenling (80 percent of the fish eaten by Phoca) are typically shallow-bottom species which live near rocky habitats. Zalophus characteristically feeds on Pacific hake, northern anchovy, and rockfish (93 percent of the fish). These open- water fishes are very abundant (Ahlstrom 1965; Bell (1971). Eumetopias, while also using Pa- cific hake and rockfish, relied more heavily on bottom-dwelling flatfish and cusk-eel (29.1 per- cent of the fish). Octopus was the only cepha- lopod found in the stomachs of harbor seals, al- though several other cephalopods are taken by California and northern sea lions (Table 9). Stomachs of Eumetopias frequently had stones in them (Table 9). Apparently there is some se- lection for these nonfood items by these sea lions. The depth of the sea at which these pin- nipeds feed is unknown. All prey items normally eaten by Eumetopias inhabit water less than about 200 m deep. Thus Zalophus feeds on schooling fishes while Eumetopias feeds on bot- tom fishes (Table 6). The mean feeding rate of small cetaceans as described by Sergeant (1969) is 10.8 percent of their body weight per day. No comparative in- formation is available on feeding rates of juve- nile individuals, nor have sexual or seasonal differences in feeding rates been published. Captive Arctic seals (Cystophora, Pagophilus) require food in amounts of 3—5 percent of their body weight per day in order to maintain good health (Blix et al. 1973). Daily food consump- tion of fur seals (10 percent of body weight), northern sea lion (4 percent of body weight), and harbor seal (11 percent of body weight) are recorded on field-collected specimens by Spald- ing (1964). These data do not include informa- tion on body size, reproductive state, or activity factors which influence the food consumption of these pinnipeds. The smaller odontocetes fed on more cepha- lopods (17 percent of the diet) than did the pin- nipeds (5 percent) (Tables 5 and 9). The diet of the sea lions in this study was 95 percent fish, compared with 83 percent in the cetaceans. 430 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 16 Phocoena fed heavily on rockfish, anchovy, and juvenile Pacific hake. Most of these fishes are semi-pelagic, small, and probably occur near the surface. The recoveries of harbor porpoises were limited to the period April to September and therefore do not reflect a year-round dietary sample. These porpoises prey extensively on Loligo, which is present throughout the year. Lagenorhynchus associates with many other species of marine animals and perhaps feeds on a wide variety of prey. Data from this study (Table 8) indicate that white-sided dolphins are generalized feeders. Osmerids and midshipmen, representing two distinct habitats (inshore schooling and bottom-dwelling), were major food items. Although midshipmen live from the intertidal regions to a depth of 170 fathoms (about 311 m), they also leave the bottom in search of food (Fitch and Lavenberg 1971). We do not know where or when this fish is eaten, but it is an important component in the diets of all marine mammals studied. ACKNOWLEDGMENTS So many individuals have contributed to this study that it is impractical to list them all. The telephone calls from federal, state, and county officials and other interested persons reporting carcasses are deeply appreciated. Personnel at the Point Reyes Bird Observatory deserve spe- cial thanks. The facilities of these institutions were used: California Academy of Sciences; Museum of Vertebrate Zoology, University of California, Berkeley; Humboldt State Univer- sity. Personnel of each institution extended aid, especially my dissertation committee at Hum- boldt State University, chaired by Dr. W. J. Houck. The patience showed by the manuscript reviewers is appreciated; thank you, Drs. S. Harris, J. Waters, and W. Z. Lidicker, Jr. The assistance of Mr. John E. Fitch, Research Di- rector, State Fisheries Laboratory, California Department of Fish and Game, was essential for the identification of fish otoliths. Mr. Clifford H. Fiscus, National Marine Fisheries Service, Se- attle, Washington, aided with the identification of the cephalopod beaks. My able field assistants located as many old carcasses as I did. Thanks, Pica, Chepen, Dusty, and Ginger. And lastly, the support of Lise made all the tasks easier. LITERATURE CITED AHLSTROM, E. H. 1965. Kinds and abundance of fishes in the California Current region based on egg and larval sur- vey. Calif. COFI Prog. Rep. 10:31-37. ANDERSON, H. 1960. Sea lions—creatures of controversy. Calif. Senate Fact Finding Committee on Natural Re- sources. Fort Bragg, California. 14 p. ANONYMOUS. 1901. Sea lions as fish destroyers. For. Stream 56(22):428. ANONYMOUS (‘‘Steelhead,’’ pseudonym). 1901. Food of sea lions. For. Stream 56(26):507—S08. BARTHOLOMEW, G. A. 1967. Seal and sea lion populations of the California Islands. Pp. 229-244 in R. N. Philbrick (ed.), Proc. Symp. on the Biology of the California Islands. Santa Barbara Botanic Garden, Santa Barbara. BELL, R. R. 1971. California marine fish landings for 1970. Calif. Dep. Fish Game Fish Bull. 154. 50 p. BiGG, M. A. 1969. The harbor seal in British Columbia. Fish. Res. Bd. Canada Bull. 172. 33 p. Birix, A. S., J. A. IVERSON, AND A. PASCHE. 1973. On the feeding and health of young hooded seals (Cystophora cris- tata) and harp seals (Pagophilus groenlandicus) in captiv- ity. Norw. J. Zool. 21:55-58. BONNoT, P. 1928. Report on the seals and sea lions of Cal- ifornia. Calif. Dep. Fish Game Fish Bull. 14. 61 p. 1932a. Food habits of the Pacific harbor seal. Calif. Fish Game 18:98. 1932b. A note on the fishing of the California sea lion. Calif. Fish Game 18:98—99. 1951. The sea lions, seals, and sea otter of the Cal- ifornia coast. Calif. Fish Game 37:371-389. BricGs, K. T., AND C. W. Davis. 1972. A study of predation by sea lions on salmon in Monterey Bay. Calif. Fish Game 58:37-43. Brown, D. H., AND K. S. Norris. 1956. Observations of captive and wild cetaceans. J. Mammal. 37:311—326. BROWNELL, R. L., JR. 1964. Observations of odontocetes in central California waters. Norsk Hvalfangst-Tidende 53(3):60—66. CALIFORNIA DIVISION OF FISH AND GAME. sea lions eat? Calif. Fish Game 13:119-120. CLARKE, M. R. 1962. The identification of cephalopod ‘beaks’ and the relationship between beak size and total body weight. Bull. Brit. Mus. (Nat. Hist.) Zool. 8(10):419— 480. CLEMENS, W. A., AND G. V. WiiBy. 1933. Food of the fur seal off the coast of British Columbia. J. Mammal. 14:43- 46. 1927. What do , AND . 1961. Fishes of the Pacific coast of Can- ada. Second ed. Fish. Res. Bd. Canada Bull. 68. 443 p. CLOTHIER, C. R. 1950. A key to some southern California fishes based on vertebral characters. Calif. Fish Game Fish Bull. 79. 83 p. Cowan, I. M. 1944. The Dall porpoise, Phocoenoides dalli (True), of the northern Pacific Ocean. J. Mammal. 25:295— 306. Coxe KeaWwe cyclostome from Californian waters. Calif. 49:271-289 DAUGHERTY, A. E. 1972. Marine mammals of California. Second ed. Dept. Fish and Game, Sacramento, California. 87 p. DY GHEE, eae 1963. Egg-cases of some elasmobranchs and a Fish Game 1901. Notes on the food habits of California JONES: FOOD HABITS OF SMALLER MARINE MAMMALS 431 sea lions. [Manuscript in C. H. Merriam files Mus. Vert. Zool. Univ. Calif., Berkeley. 6 p.] . 1903. Notes on the food habits of California sea lions. Zalophus californianus Lesson. Trans. Kansas Acad. Sci. 18:179-182. Evans, W.E., AND J. BASTIAN. 1969. Marine mammal com- munication: social and ecological factors. Pp. 425—475 in H. T. Andersen. The biology of marine mammals. Academ- ic Press, New York. EVERMANN, B. W. 1921. The Ano Nuevo Steller sea lion rookery. J. Mammal. 2:16-19. , AND G D. HANNA. 1925. The Steller sea lion rook- ery on Ano Nuevo Island, California in 1924. J. Mammal. 6:96-99. FINK, B. D. 1959. Observation of porpoise predation on a school of Pacific sardines. Calif. Fish Game 45:216-217. Fiscus, C. H., AND G. A. BAINES. 1966. Food and feeding behavior of Steller and California sea lions. J. Mammal. 47:195-200. , AND K. NiGGOL. 1965. Observations of cetaceans off California, Oregon, and Washington. U.S. Fish Wild. Serv. Spec. Sci. Rep. Fish. 498. 27 p. Fitcn, J. E., AND R. L. BROWNELL, JR. 1968. Fish otoliths in cetacean stomachs and their importance in interpreting feeding habits. J. Fish. Res. Bd. Canada 25:2561—2574. , AND R. J. LAVENBERG. 1971. Marine food and game fishes of California. California Natural History Guides 28. Univ. Calif. Press, Berkeley and Los Angeles. 179 p. FREIBURG, R. E., AND P. C. Dumas. 1954. The elephant seal (Mirounga angustirostris) in Oregon. J. Mammal. 35:129. Frey, H. W. (ed.). 1971. California’s living marine resources and their utilization. Calif. Dept. Fish and Game. Sacra- mento, California. 148 p. Frost, G. A. 1924. Fish otoliths from the stomach of a por- poise. Nature 113:310. Fry, D.H.,Jr. 1939. A winter influx of sea lions from Lower California. Calif. Fish Game 25:245—250. HALt, E.R., AND J. W. BEE. 1954. Occurrence of the harbor porpoise at Point Barrow, Alaska. J. Mammal. 35: 122-123. HANDLEY, C. O., JR. 1966. A synopsis of the genus Kogia (Pygmy Sperm Whales). Pp. 62-69 in K. S. Norris (ed.). Whales, porpoises, and dolphins. Univ. Calif. Press, Berke- ley and Los Angeles. HANNA, G D. 1951. Alaska fur seals, Callorhinus ursinus, observed off San Francisco Bay, California. J. Mammal. 32:364-365. HARRISON, R. J. (ed.). 1972. Functional anatomy of marine mammals. Vol. 1. Academic Press, London and New York. 451 p. , F. R. JOHNSON, AND B. A. YOUNG. 1970. The oe- sophagus and stomach of dolphins (Tursiops, Delphinus, Stenella). J. Zool. 160:377-390. HaTcer, D. F. 1971. A Canadian specimen of Risso’s dol- phin. Canadian Field-Nat. 85: 188-189. HepGPETH, J. W. 1944. Sea lions. Nat. Mag. 37(8):443-445. Houck, W. J. 1961. Notes on the Pacific striped porpoise. J. Mammal. 42:107. Husss, C. L., W. I. FOLLETT, AND L. J. DEMpsTER. 1979. List of the fishes of California. Occas. Pap. Calif. Acad. ScmlS3=5ilip: Huey, L. M. 1925. Late information on the Guadalupe Island elephant seal herd. J. Mammal. 6:126—127. 1930. Capture of an elephant seal off San Diego, California, with notes on stomach contents Mammal. 11:229=231. IMLER, R. H., AND H. R. SARBER. 1947. Harbor seals and sea lions in Alaska. U.S. Fish Wild. Serv. Spec. Sci. Rep 28e2snpe IverSON, I. L. K., AND L. PINKAS. 1971. A pictorial guide to beaks of certain eastern Pacific cephalopods. Calif. Fish Game Fish Bull. 152:83-105. JAMESON, R. J., AND K. W. KENYON. 1978. Prey of sea lions in the Rogue River, Oregon. J. Mammal. 58:672. KENYON, K. W. 1952a. Diving depths of the Steller sea lion and Alaskan fur seal J. Mammal. 33:245—246. 1952b. The Steller sea lion. Pac. Discovery 5(4):4— 13. . 1965. Food of harbor seals at Amchitka Island, Alas- ka. J. Mammal. 46: 103-104. , AND V. B. SCHEFFER. 1955. The seals, sea lions, and sea otter of the Pacific coast. U.S. Fish Wild. Serv. Circ. 32. 34 p. KING, J. E. 1964. Seals of the world. Brit. Mus. (Nat. Hist.) London. 154 p. Logs, V. J. 1972. A study on the distribution and feeding habits of the Dall porpoise in Monterey Bay, California. Unpublished Master’s Thesis, San Jose State College. 62 p. Lustic, B. L. 1948. Sight records of Dall porpoises off the Channel Islands, California. J. Mammal. 29:183. MARINE MAMMAL BIOLOGICAL LABORATORY. 1969. Fur seal investigations, 1966. U.S. Fish Wild. Serv. Spec. Sci. Rep. Fish. 584. 123 p. Mare, B. R. 1973. Population kinetics and related ecology of the northern sea lion, Eumetopias jubatus, and the Cal- ifornia sea lion, Zalophus californianus, along the Oregon coast. Ph.D. Thesis. Univ. of Oregon. 94 p. MATHISEN, O. A. 1959. Studies on Steller sea lion (Euse- topias jubata) in Alaska. Trans. N. Am. Wildl. Res. Conf. 24:346-356. .R.T. BAADE, AND R. J. Lopp. 1962. Breeding habits, growth, and stomach contents of the Steller sea lion in Alas- ka. J. Mammal. 43:469-477. MERRIAM, C. H. 190la. Food of sea lions. Science 8:777— 779. 1901b. Food of sea lions. For. Stream 56(21):428. MILLER, D. J., AND R. N. LEA. 1972. Guide to the coastal marine fishes of California. Calif. Fish Game Fish Bull. [S75 1235: MOREJOHN, G. V., AND D. M. BALtz. 1970. Contents of the stomach of an elephant seal. J. Mammal. 51: 173-174. NATIONAL OCEANIC AND ATMOSPHERIC ADMINISTRATION. 1974. Report of the Secretary of Commerce. Administra- tion of the Marine Mammal Protection Act of 1972. Federal Register 39:23896—23932. NELSON, O. M., AND H. A. LARKINS. 1970. Distribution and biology of Pacific hake: a synopsis. Pp. 23-24 in Pacific hake. U.S. Fish Wildl. Serv. Circ. 332. NEwcomBE, C. F., W. H. GREENWOOD, AND C. M. FRASER. 1918. The sea-lion question in British Columbia. Contrib. Canadian Biol. Sessional Paper 38a. 39 p. , AND W. A. NEwcomBE. 1914. Sea-lions on the coast of British Columbia. Rep. Comm. Fish. B.C. for 1913:131- 145. Norris, K. S. (ed.). 1961. Committee on marine mammals. Standardized methods for measuring the recording data on the smaller cetaceans. J. Mammal. 42:471-476. 432 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 16 , AND J. H. Prescott. 1961. Observations on Pacific cetaceans of Californian and Mexican waters. Univ. Calif. Publ. Zool. 63:291-401. OpeEtt, D. K. 1971. Censuses of pinnipeds breeding on the California Channel Islands. J. Mammal. 52:187—190. Orr, R. T. 1937. A porpoise chokes ona shark. J. Mammal. 18:370. 1951. Cetacean records from the Pacific coast of North America. Wasmann J. Biol. 9(2): 147-148. 1966. Risso’s dolphin on the Pacific coast of North America. J. Mammal. 47:341—343. 1972. Marine mammals of California. Calif. Nat. Hist. Guides 29. Univ. Calif. Press, Berkeley and Los An- geles. 64 p. , AND T. C. PouULTER. 1965. The pinniped population of Ano Nuevo Island, California. Proc. Calif. Acad. Sci., ser. 4, 32:377-404. , AND 1967. Some observations on reproduc- tion, growth, and social behavior in the Steller sea lion. Proc. Calif. Acad. Sci., ser. 4, 35: 193-226. PAUL, J. R. 1968. Risso’s dolphin, Grampus griseus, in the Gulf of Mexico. J. Mammal. 49:746-748. PERRIN, W. F. 1970. The problem of porpoise mortality in the U. S. tropical tuna fishery. Pp. 45—48 in Proceedings 6th Annual Conference on Biological Sonar and Diving Mammals, Stanford Res. Inst., Menlo Park, Calif. , R. R. WARNER, C. H. Fiscus, AND D. B. HO-ts. 1973. Stomach contents of porpoise, Stenella spp., and yel- lowfin tuna, Thunnus albacares, in mixed-species aggre- gations. U.S. Natl. Mar. Fish. Serv. Fish. Bull. 71:1077— 1092. PETERSON, R. S., AND G. A. BARTHOLOMEW. 1967. The nat- ural history and behavior of the California sea lion. Am. Soc. Mammal. Spec. Publ. no. 1. 79 p. . AND B. J. LEBOEuF. 1969. Population study of seals and sea lions. Trans. N. Am. Wildl. Res. Conf. 34:74-79. PikE, G. C. 1958. Food of the Northern sea lion. Pp. 18-20 in Prog. Rep. 112, Fish. Res. Bd. Canada Biol. Sta. Na- naimo, B.C. , ANDI. B. MACASkIeE. 1969. Marine mammals of Brit- ish Columbia. Fish. Res. Bd. Canada Bull. 171. 54 p. RAE, B. B. 1965. The food of the common porpoise (Pho- coena phocoena). J. Zool. (Proc. Zool. Soc. London) 146: 114-122. REDDING, B. R., S. R. THROCKMORTON, AND J. D. FARWELL. 1875. P. 13 in Report of the Commissioners of Fisheries of the State of California, 1874-1875. Sacramento. RipGway, S. H. (ed.). 1972. Mammals of the sea: biology and medicine. Charles C Thomas, Springfield, Illinois, 812 p. Rosins, C. R., R. M. BaiLey, C. E. Bonpb, J. R. BROOKES, E. A. LACHNER, R. N. LEA, AND W. B. Scott. 1980. A list of common and scientific names of fishes from the United States and Canada (Fourth edition). Am. Fish. Soc. Spec. Publ. 12. 174 p. ROEDEL, P. M. 1953. Common ocean fishes of the California coast. Calif. Fish Game Fish Bull. 91. 184 p. Row ey, J. 1929. Life history of the sea lions of the Calli- fornia coast. J. Mammal. 10: 1-36. RuTrer, C., R. E. SNODGRASS, AND E. C. STARKS. 1904. Report on the sea lion investigation. Pp. 116-119 in Hugh M. Smith. Report on inquiry respecting food-fishes and fish- ing-grounds. Rep. U.S. Comm. Fish Fish. Part 18. Ryper, R. A. 1957. Avian-pinniped feeding associations. Condor 59:68—-69. Scammon, C. M. 1874. The marine mammals of the north- western coast of North America, described and illustrated; together with an account of the American whale-fishery. John H. Carmany and Co., San Francisco. 319 p. [Reprinted by Dover Co., New York. 1968.] SCHEFFER, T. H. 1928. Precarious status of seal and sea-lion on our northwest coast. J. Mammal. 9:10-16. , AND C. C. SPERRY. 1931. Food habits of the Pacific harbor seal, Phoca richardii. J. Mammal. 12:214—226. SCHEFFER, V. B. 1950a. The food of the Alaska fur seal. U.S. Fish Wild. Serv. Wild. Leaflet 329. 16 p. 1950b. The striped dolphin, Lagenorhynchus obli- quidens (Gill 1865), on the coast of North America. Am. Midl. Nat. 44:750-758. . 1953. Measurements and stomach contents of eleven delphinids from Northeast Pacific. Murrelet 34(2):27-30. . 1958. Seals, sea lions, and walruses, a review of the pinnipedia. Stanford Univ. Press, Stanford, California. 179 p. (ed.). 1967. Committee on marine mammals. Standard measurements of seals J. Mammal. 48:459—462. , AND J. A. NEFF. 1948. Food of California sea-lions. J. Mammal. 29:67-68. , AND J. W. Stipe. 1944. The harbor seal in Washing- ton State. Am. Midl. Nat. 32:373-416. , AND . 1948. The whales and dolphins of Wash- ington State, with a key to the cetaceans of the west coast of North America. Am. Midl. Nat. 39:257-337. ScHMIDT, J. 1923. Consumption of fish by a porpoise. Nature 112:902. SCHULTZ, L. P., AND A. M. RAFN. 1936. Stomach contents of fur seals taken off the coast of Washington. J. Mammal. ANS =I5)- SEED, A. (ed.). 1972. Seals, sea lions, and walruses in eastern North Pacific and Arctic waters. Pacific Search, Seattle. 40 p. SERGEANT, D. E. 1968. Feeding ecology of marine mammals. Pp. 89-96 in Proc. 2nd Symposium on Disease and Hus- bandry of Aquatic Mammals. Marineland Florida Res. Lab. 1969. Feeding rates of cetacea. Fiskeridir Skr. Ser. Havunders. 15(3):246-258. , AND H. D. FisHer. 1957. The smaller cetacea of eastern Canadian waters. J. Fish. Res. Bd. Canada 14:83— 115. SLUPER, E. J. 1962. Whales, translated by A. J. Pomerans. Basic Books, New York. 475 p. SmitH, G. J. D., AND D. E. GASKIN. 1974. The diet of harbor porpoises [Phoecoena phocoena (L)] in coastal waters of eastern Canada, with special reference to the Bay of Fundy. Canadian J. Zool. 52:777-782. SPALDING, D. J. 1964. Comparative feeding habits of the fur seal, sea lion, and harbour seal on the British Columbia coast. Fish. Res. Bd. Canada Bull. 146. 52 p. Starks, E. C. 1918. The sea lions of California. Am. Mus. J. 18(3):226—237. STEELE, J. H. (ed.). 1970. Marine food chains. Univ. Calif. Press, Berkeley and Los Angeles. 552 p. Stroup, R. K. 1968. Risso dolphin in Washington State. J. Mammal. 49:347-348. THORSTEINSON, F. V., AND C. J. LENSINK. 1962. Biological JONES: FOOD HABITS OF SMALLER MARINE MAMMALS. 433 observations of Steller sea lions taken during an experi- mental harvest. J. Wildl. Manage. 26:353-359. TomILiIn, A. C. 1957. Mammals of U.S.S.R. and adjacent countries. Vol. 9. Cetaceans. Translated from Russian (Israel program for scientific translation, Jerusalem. 1967). TANG Jo: TOWNSEND, C. H. 1918. Sea lions and the fishery industries. N.Y. Zool. Soc. Bull. 21: 1679-1682. . 1919. The utilization of the sea lion. N.Y. Zool. Soc. Bull. 22:32—33. WILKE, F. 1957. Food of sea otters and harbor seals at Am- chitka Island. J. Wildl. Manage. 21:241—242. , AND K. W: KENYON. 1952. Notes on the food of fur seal, sea-lion, and harbor porpoise. J. Wildl. Manage 16:396-397. , AND . 1957. The food of fur seals in the eastern Bering Sea. J. Wildl. Manage. 21:237-238. , AND A. J. NICHOLSON. 1958. Food of porpoises in waters off Japan. J. Mammal. 39:441-443. CALIFORNIA ACADEMY OF SCIENCES Golden Gate Park San Francisco, California 94118 : a = | ; . at oe vee '» ae ¢ as — f * PS THE SERRA (Cf it ot ' t: b¢ | 4 > A on 7 : ~ F 5 Cd 7¥_ POUR werw NER rin »F,. ae 4h | WESTERN NOHTH AMERICS , aVitivGediund 4 ; o : | Het ant ae ait - i bes™~ incite uec Pome. tw ; < - AT Poa A) v G. ctenocephalus oG.ridens 6 : A G. fasciatus ® 4°- * G. lepidogaster ROP ~S a 5 © G. megalepis oN (ee W a i a a7 y f XXSO PAR : iJ ret A o°- , iS ij Lg eae os BS U PP fe ; 4°- FIGURE 3. 8b. Predorsal scales about 32-36; branched dorsal-fin rays 7/2; branched anal-fin rays 414; male with ctenoid tubercles on head G. ctenocephalus Gastromyzon borneensis Gunther (Figures la, b, 4—5) Gastromyzon borneensis GUNTHER, 1874:6 (type-locality “‘sources of Mingalong River’’ = Mengalong River, Beau- fort District, North Borneo). TyPE-MATERIAL.—The type-series of five syntypes (BMNH 1874.11.24:1—5) comprises two species. Four specimens have a prominent secondary rostrum and the abdomen entirely scaleless; they are conspecific. The largest of these (BMNH 1874.11.24:1, 79.1 mm) is hereby designated lectotype of G. borneensis; the other three (BMNH_ 1874.11.24:2-4, 78.3, 72.6, and 65.5 mm) are paralectotypes. The fifth syntype (BMNH 1874.11.24:5, 78.0 mm) lacks a secondary rostrum, has the abdomen entirely covered with fine scales, and is not conspecific with the others; it is now the holotype of G. lep- idogaster new species. Distribution of Gastromyzon (based on material examined). ADDITICNAL MATERIAL EXAMINED.—NORTH BORNEO: MCZ 34793, 34820, 34823, 27:27.9-70.4 mm, Mount Kinabalu; SU 31473, ZMA 114.365, 6:32.1-52.9 mm, Bongan River, Mandu Bay; FMNH 68129, 4:16.5-38.4 mm, Parutan River, Tambunan; FMNH 68128, 5:20.6-46.5 mm, Sungei Kainger- an, Tambunan. SARAWAK: IRSNB 19723, 64:36.6-89.0 mm, Arur Dalan, a torrential tributary of Padapur River, near Bar- io, Baram basin; IRSNB 19724, 30:19.7-68.3 mm, Sungai Ra- mudu, an affluent of Sungai Kalapang, upstream from Para- mudu village, Baram basin; BMNH 1978.3.20:232-235, 4:35.4-S2.9 mm, Medalam River, Gunong Mulu; BMNH 1895.7.2:82, 1:65.1 mm, Akar River; BMNH 1933.8.9:1-3, 3:57.8-66.9 mm, Lejok River, Tinjar River; FMNH 45852, 1:70.4 mm, Truson River, Lawas; FMNH 45853, 2:64.9-70.5 mm, Pa Brayong, Truson River; FMNH 68583, 3:44.1-51.0 mm, Akah River, Meligong. DIAGNOSIS.—Attaining 89 mm, and thus pos- sibly the largest species in the genus, Gastro- myzon borneensis differs from all of its conge- ners in having a secondary rostrum and, in sexually mature males, developing heavy tuber- ROBERTS: GASTROMYZONTINE FISH 503 FIGURE 4. culation on posteroventral portion of body. The secondary rostrum, present in all specimens ex- amined over 30 mm of both sexes, is tuberculate in both sexes, but becomes slightly larger and more tuberculate in males than in females. Post- oral pouch present (absent in all other Gastro- myzon except G. monticola). Snout terminating in secondary rostrum, oth- erwise gently sloping downward in front of eyes and relatively truncate. Gill opening vertical, its length about equal to eye diameter; no suboper- cular groove. Scales in lateral series 51-62; pre- dorsal scales at least 40; scale rows above lateral series about 20; circumpeduncular scales 28-32. Pectoral fin falls far short of pelvic fin, pelvic fin Gastromyzon borneensis, 88.2-mm female (IRSNB 19723). short of anal fin, and depressed dorsal fin far short of level of anal-fin origin. Dorsal and lateral surfaces of body with dull brownish or brownish-gray background broken by pale, interconnected narrow bands forming irregularly hexagonal or pentagonal figures; dor- sal surface of head with hexagonal or pentagonal figures similar to those on body but much small- er and more closely spaced together. Most col- oration on fins narrowly confined along fin rays; dorsal fin with faintly banded pattern; caudal fin with three to six dusky vertical bands. Paired fins without longitudinal bands or pale margins, pale or colorless posteriorly. 504 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 20 FIGURE 5. Gastromyzon contractus new species (Figures 2, 6) TyPpeE-MATERIAL.—The types and only known specimens of this new species were collected in the mainstream and one small tributary of the Sungai Pinoh, a large high-gradient stream in the southern Kapuas basin, by Soetikno Woerjoat- modjo and me in July 1976. Holotype.—MZB 3447, 34.4-mm gravid 2°, Sungai Tebe- lian, small forest stream 3-4 m wide and 50 cm deep, where it flows into Sungai Pinoh, 19 km upstream from Nangapinoh. Paratypes.—CAS 49319, 1:30.2 mm, same data as holotype; MZB 3448, CAS 44186, USNM 230247, RMNH uncat., BMNH uncat., and MNHN uncat., 17:29.1-36.9 mm, main- stream of Sungai Pinoh 20-60 km upstream from Nangapinoh; MZB 3449 and CAS 49320, 2:25.9-29.8 mm, rocky channel in mainstream of Sungai Pinoh 37 km S of Nangapinoh. DIAGNOosIs.—Perhaps the smallest species of Gastromyzon (largest known specimen 36.9 mm), G. contractus differs from all of its con- geners in having depressed dorsal fin as well as pelvic fin reaching to or beyond level of anal-fin origin (depressed dorsal fin not reaching level of anal-fin origin, and pelvic fin reaching at most to anal fin in all other Gastromyzon). Snout sloping downward in front of eyes more strongly than in most other Gastromyzon, but not so strongly as in G. monticola and G. ridens, Gastromyzon borneensis, 71.5-mm male (IRSNB 19723). and broadly rounded. Postoral pouch absent. Gill opening slightly angular, extending ante- riorly ventral to subopercle only a short dis- tance, its length about equal to eye diameter; subopercular groove present, continuous but shallow or interrupted near its middle. Scales in lateral series 52-63; predorsal scales 36-44; scale rows above lateral series 16-20; circum- peduncular scales about 25-30. Pectoral fin overlaps pelvic fin. Male with dense patches of small conical tubercles on dorsal surface of an- teriormost pectoral-fin rays (Fig. 2). Dorsal and lateral surfaces of body with thick, pale, interconnected lines forming irregular hex- agonal or pentagonal figures; dorsal surface of head with numerous small, round, dark spots. Dorsal fin with two or three spotted longitudinal bands; caudal fin with two or three dark vertical bands; coloration in dorsal and caudal fins con- centrated along fin rays. Paired fins with faint markings, pelvic fins sometimes almost entirely colorless or colorless except basal portion dus- ky. Some specimens with dorsal and lateral sur- faces of body and head blotchy or blanched, suggesting physiological or behavioral color change. ROBERTS: GASTROMYZONTINE FISH 505 FIGURE 6. Gastromyzon contractus, 34.4-mm gravid female holotype (MZB 3447). ETYMOLOGY.—From the Latin contractus, drawn together, in reference to the overlapping or nearness of the fins. Gastromyzon ctenocephalus new species (Figure 7) TyPeE-MATERIAL.—This species is known only from the type-series, collected at Senah, Sarawak, by A. Everett. Holotype.—BMNH 1893.3.6:269, 42.3-mm ¢, Senah, Sa- rawak. Paratypes.—BMNH 1893.3.6:261-268, 8:28.9-37.8 mm, same data as holotype. DIAGNOSIS.—Gastromyzon ctenocephalus differs from all other Gastromyzon in having only 8-12 papillae on lower lip, widely spaced on or near its free anterior margin (vs. extremely numerous papillae densely distributed over en- tire lower lip); gill opening strongly angular, continued farther anteriorly ventral to suboper- cle than in any other Gastromyzon; subopercu- lar groove continuous and relatively deep (vs. subopercular groove continuous but relatively shallow, interrupted in its middle, or absent); and sexually mature male with ctenoid tubercles on head and pectoral-fin base (ctenoid tubercles otherwise known only in male Neogastromyzon pauciradiatus). Snout more or less strongly sloping downward in front of eyes, broadly rounded. No postoral pouch. Length of gill opening about 1.5—2 times eye diameter. Scales in lateral series about 50-60; predorsal scales about 32-36; scale rows above lateral series 15-20; circumpeduncular scales 28-30. Pectoral fin reaching or slightly overlapping pelvic fin; pelvic fin reaching anal fin; depressed dorsal fin falling short of level of anal-fin origin. The holotype, a 42.3-mm male, has dense patches of small conical tubercles on dorsal sur- face of anteriormost branched pectoral-fin rays, as well as ctenoid tubercles on dorsolateral por- tion of head and on pectoral-fin base. A 32.6- mm male? has dense patches of small tubercles 506 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 20 FiGuRrE 7. Gastromyzon ctenocephalus, 42.3-mm ripe male holotype (BMNH 1893.3.6.269). on pectoral fin but no ctenoid tubercles. All oth- er specimens, the three largest of which, 36.3-37.8 mm, are gravid females, lack ctenoid tubercles as well as dense patches of small tu- bercles on pectoral fin, but have numerous small conical tubercles widely scattered on dorsal sur- face of head and anteriormost portion of body (one tubercle per scale) as well as in uniserial rows of small conical tubercles on dorsal surface of pectoral-fin rays. Dorsal surface of head with numerous small, round, pale spots; dorsal and lateral surface of body uniformly pale brownish (faded?) or, in one paratype, with small, round, pale spots on dorsal surface of body and pectoral-fin base, similar to those on head but slightly larger and more widely spaced apart. In holotype, dorsal and anal fins with two longitudinal spotted bands, spots centered on rays; caudal fin with two or three faintly spotted vertical bands, and melanophores narrowly distributed along length of all rays; paired fins with faint longitudinal bands. In all paratypes, dorsal-fin rays and in- terradial membranes heavily pigmented except for depigmented areas forming an alternating se- ries of transparent round spots; anal fin with in- ROBERTS: GASTROMYZONTINE FISH 507 FIGURE 8. terradial membranes each bearing a single ver- tical, oblong, darkly pigmented spot and rays without coloration; caudal fin with simple prin- cipal rays and interradial membranes between principal rays unpigmented, but unbranched principal rays and interradial membranes be- tween branches of individual rays darkly pig- mented; paired fins longitudinally banded in some paratypes, without noticeable banding in others; one paratype with pectoral fin spotted. Coloration of dorsal, anal, and caudal fins of paratypes very different from that in other species of Gastromyzon and in holotype. This difference, perhaps due to individual variation or sexual dichromatism, should be investigated when additional material becomes available. ETyMOLOGY.—From the Greek cteno-, comb, and cephalus, head, in reference to the ctenoid tubercles on the head of mature males. Gastromyzon fasciatus Inger and Chin (Figure 8) Gastromyzon fasciatus INGER AND CHIN, 1961:173 (type-lo- cality “‘Sungai Dapu, a tributary of the Baleh River near the mouth of the Sungai Putai, Third Division, Sarawak’’). MATERIAL EXAMINED.—SARAWAK: FMNH 68115, 68120, and BMNH 1960.12.30:1, 6:48.1-65.9 mm, paratypes, Baleh Gastromyzon fasciatus, 59.0-mm mature male (MZB 3450). River, Rajang basin; FMNH 45852, 1:70.4 mm, Truson River, Lawas; FMNH 45853, 2:64.9-70.5 mm, Pa Brayong, Truson River; FMNH 68583, 3:44.1-5S1.0 mm, Akah River, Meligong; SU 32378, 1:45.1 mm, Sadong River; IRSNB 19725, 6:38.6-69.2 mm, Arur Dalan, torrential tributary of Padapur River, Baram basin. NORTH BORNEO: FMNH 68128, 5:20.6-46.5 mm, Sungai Kaingeran, Tambunan; FMNH un- cat., 4:17.2-32.8 mm, Parutan River, Tambunan. KAPUAS BA- sIN: MZB 3450 and CAS 49321, 7:24.5-59.0 mm, Sungai Pinoh, RMNH 7636, 4:22.5-54.1 mm, Bongan. MAHAKAM BA- SIN: RMNH 7638, 2:53.2-56.8 mm, B6; RMNH 7794, 2:39.2-54.3 mm, Bluu or Bloeoe. DIAGNOsISs.—There does not seem to be any single character by which all specimens of G. fasciatus can be distinguished from all other species of Gastromyzon, but it is nevertheless a highly distinctive species. Predorsal scales 40-55 (usually less than 40 in all other Gastro- myzon). Pelvic-fin and anal-fin rays usually more numerous than in other Gastromyzon (Table 1): branched pelvic-fin rays 20-24, modally 22 (vs. 17-22, modally 20 or less); branched anal-fin rays 5-54, modally 514 (vs. 4-514, modally 5 or less). Specimens of G. fasciatus with well-de- veloped coloration differ from all other Gastro- myzon in having dorsal surface of head and body very darkly pigmented, black or bluish black, with narrow vertical white bars or white spots 508 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 20 FiGuRE 9. Gastromyzon lepidogaster on body and disconnected white vermiculations on head. This coloration is well developed in freshly preserved specimens from Sarawak, North Borneo, and the Kapuas basin but is less well developed or indistinct in older material from the Mahakam basin. Snout gently sloping downward in front of eyes, broadly rounded. No postoral pouch. Gill opening angular, extending anteriorly a short distance ventral to subopercle, its length about 1.5 times eye diameter. Subopercular groove always present, continuous or interrupted in the middle of its length. Scales in lateral series 55-68; scales above lateral series 20-23; circum- peduncular scales 30-43. Pectoral fin fails to reach pelvic fin; pelvic fin falls short of anal fin; -, 78.0-mm holotype (BMNH 1874.11.24:5). depressed dorsal fin falls far short of level of anal-fin origin. Male with dense patches of small tubercles on dorsal surface of anteriormost branched pectoral-fin rays. Freshly preserved specimens, including para- types from the Rajang basin and the Kapuas material, have very distinctive coloration differ- ent from all other Gastromyzon: dorsal and lat- eral surfaces of head and body darkly pigment- ed, black or bluish-black, with several pale or white vertical bands (sometimes breaking up into a few large spots) on body; dorsal fin with three or four longitudinal spotted bands, caudal fin with two or three almost solid, thick, vertical or irregularly broken black bands; paired fins irregularly spotted or mottled (not longitudinally ROBERTS: GASTROMYZONTINE FISH 509 FicureE 10. Gastromyzon lepidogaster, 62.9-mm ripe female paratype (FMNH 68113). banded). Specimens from the Mahakam and some other collections, however, have less dis- tinctive coloration, perhaps partly but not en- tirely due to fading after long preservation. Gastromyzon lepidogaster new species (Figures 9-10) Gastromyzon borneensis GUNTHER, 1874:6 (in part); POPTA 1906; WEBER AND DE BEAUFORT 1916 (Fig. 1, text; in part); INGER AND CHIN 1961; 1962 (in part). TyYPE-MATERIAL.—Holotype: BMNH 1874.11.24:5, 78.0 mm, sources of Mengalong River, Beaufort District, North Borneo. Paratypes.—NoORTH BORNEO: FMNH 68111-68114, 7:38.0-64.4 mm, Kinabatangan basin; FMNH 51683, 1:46.4 mm, East Coast Residency. BRUNEI: IRSNB uncat., 1:37.2 mm, upper course of Sungai Temburong. KAJAN BASIN: RMNH 7639, 3:79.0-83.3 mm, upper course of Boeloengan River above Ben Barum. MAHAKAM BASIN: RMNH 7635, 5:39.3-70.1 mm, upper Mahakam; RMNH 7637, 19:22.3-46.2 mm, Howong; RMNH 7794, 7:36.1-83.4 mm, Bluu or Bloeoe; RMNH 7638, 1:71.8 mm, B6. KAPUAS BASIN: RMNH uncat., 1:72.3 mm, Bongan. DIAGNOSIS.—Gastromyzon lepidogaster dif- fers from all other Gastromyzon in having the abdomen more or less extensively covered with fine scales in all specimens examined over 40.2 mm and in many below this size (abdomen en- tirely scaleless in all other Gastromyzon except in some specimens of G. megalepis which have a few relatively large scales between bases of posteriormost pelvic-fin rays). Snout gently sloping downward in front of eyes, gently rounded. No postoral pouch. Gill opening vertical, its length about equal to eye diameter. No subopercular groove. Scales in lat- eral series 54-62; predorsal scales 39-42; scale rows above lateral series 16-20; circumpedun- cular scales 30-32. Pectoral fin reaches or falls short of pelvic fin; pelvic fin reaches anal fin; depressed dorsal fin falls short of level of anal- fin origin. All Gastromyzon lepidogaster from North Borneo and Sarawak have the abdomen com- pletely covered with fine scales up to the level of the anterior margin of the pectoral girdle, ex- cept the smallest specimen (27.7 mm), which has the abdomen entirely scaleless. Specimens of this species from the Mahakam, Kayan, and Ka- puas basins above 40.2 mm, and many below this size, have the abdomen completely covered with fine scales between the bases of the pelvic fins, and sometimes have scattered scales far- ther anteriorly, but never have the abdomen 510 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 20 FIGURE 11. completely covered with scales. They are simi- lar in other respects to G. lepidogaster from North Borneo and Sarawak. Dorsal and lateral surfaces of head and body uniformly brownish, or body with thin pale ver- tical bands (similar but slightly more numerous than those in G. fasciatus); bands on posterior portion of body in a few specimens intercon- nected to form irregularly pentagonal or hexag- onal figures. Distal half of dorsal fin with two longitudinal spotted bands; caudal fin with broad dusky vertical band basally (on portion of fin covered by scales) and three or four lunate spot- ted bands on rest of fin. Anal and paired fins with faint markings, pectoral with spotted lon- gitudinal bands, pelvic and anal with dusky col- oration largely confined along rays. ETYMOLOGY.—From the Greek /epido-, scale, and gaster, stomach, in reference to the exten- sive abdominal squamation. Gastromyzon megalepis new species (Figure 11) Gastromyzon borneensis INGER AND CHIN, 1961:171 (in part). TyPE-MATERIAL.—Holotype: FMNH 68126, 54.6-mm <4, tributary of Baleh River between Sungai Entunau and Sungai Gastromyzon megalepis, 54.6-mm mature male holotype (FMNH 68126). Putai, Rajang basin, Third District, Sarawak, R. F. Inger, 5 Aug. 1956. Paratypes.—FMNH uncat., 5:45.1-60.5 mm, same data as holotype; FMNH 68127, 3:28.6—33.1 mm, Baleh River, Rajang basin, Third District, Sarawak. DIAGNOSIS.—Gastromyzon megalepis has rel- atively larger scales than any other Gastromy- zon; it is also the only Gastromyzon with pelvic fin lacking serrae. Scales in lateral series 44-48 (vs. 50 or more in all other Gastromyzon); pre- dorsal scales 24-28 (vs. 30 or more); scale rows above lateral series 11-12 (vs. 15 or more); and circumpeduncular scales 20-21 (vs. 25 or more). Pelvic-fin rays 4-15 each with a sharp, dorso- posteriorly projecting continuous lamina or flange, without bony projections from individual lepidotrichia. Snout gently sloping downwards in front of eyes, broadly rounded. No postoral pouch. Gill opening vertical, its length about 1.5 times eye diameter. Subopercular groove absent. Pectoral fin extends to pelvic-fin origin; pelvic fin extends almost to anal fin; depressed dorsal fin falls far short of level of anal-fin origin. Male with dense patches of minute tubercles on dorsal surface of anteriormost branched pectoral-fin rays. ROBERTS: GASTROMYZONTINE FISH 511 FIGURE 12. Dorsal and lateral surfaces of head and body uniformly brownish, sometimes mottled on cau- dal portion of body. Dorsal fin with four longi- tudinal spotted bands and caudal fin with about six lunate spotted bands, the spots nearly con- fined to fin rays; anal fin with melanophores uni- formly distributed along rays; paired fins with faint longitudinal bands, pale margins. ETyMOLOoGY.—From the Greek mega-, large, and /epis, scale, in reference to the relatively large scales of this species. Gastromyzon monticola, 70.9-mm ripe female (BMNH 1894.6.30:196). Gastromyzon monticola (Vaillant) (Figure 12) Lepidoglanis monticola VAILLANT, 1889:81 (type-locality ‘*Kina-Balou’’ = Mt. Kinabalu, North Borneo). Gastromyzon monticola VAILLANT, 1891; 1893:94, pl. 1 (de- scription and figures of syntypes). Gastromyzon borneensis WEBER AND DE BEAUFORT, 1916 (in part). This species has not been recognized by ich- thyologists since it was placed in the synonymy of Gastromyzon borneensis by Weber and de 512 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 20 FiGure 13. Beaufort; it was entirely overlooked by Inger and Chin (1961). MATERIAL EXAMINED.—AIll from NORTH BORNEO: MNHN 1889.83-85, 3:63.5-67.1 mm, syntypes, Mt. Kinabalu (Vaillant 1889, indicated 10 syntypes but only these 3 could be located during my visit to the MNHN in November 1979); BMNH 1894.6.30: 196-197, 2:66.9-70.9 mm, Mt. Kinabalu (these two specimens were compared directly with the syntypes during my visit to the MNHN); FMNH 44725, 4:33.2-56.3 mm, Tem- passuk River, Kota Belud District, West Coast Residency. In addition to the specimens just listed, I tentatively identify MCZ 54091, 2:43.147.2 mm, Sarawak, as this species; these specimens were seen briefly during a visit to the MCZ and have not been considered in the account below. DIAGNOSIS.—Gastromyzon monticola differs from all other Gastromyzon except G. borneen- sis in having a postoral pouch; it lacks a sec- ondary rostrum and differs in many respects from G. borneensis. It differs from all other Gastromyzon except G. ridens and G. contrac- tus in having the snout very strongly sloping downward in front of the eyes. Snout broadly rounded or slightly pointed. Gill opening vertical, its length about equal to Gastromyzon punctulatus, 53.0-mm gravid female paratype (FMNH 68117). eye diameter. Subopercular groove absent. Scales in lateral series about 55—58; predorsal scales about 40; scale rows above lateral series 20; circumpeduncular scales 30-32. Pectoral fin overlaps pelvic fin; pelvic fin reaches anal fin; and depressed dorsal fin falls far short of level of anal-fin origin. Dorsal and lateral surfaces of body uniformly brownish; dorsal surface of head with intercon- nected thick pale lines forming irregularly hex-— agonal or pentagonal figures, very numerous and close-set. Dorsal, anal, and caudal fins with nu- merous small round spots centered on rays, forming about four longitudinal bands on dorsal fin, two (faint) bands on anal fin, and up to eight lunate spotted bands on caudal fin. A particu- larly prominent dark spot at dorsal-fin origin (a spot at dorsal-fin origin occurs in other Gastro- myzon but is usually not so noticeable). Paired fins longitudinally banded, with a moderately wide pale distal marginal band, and a dark sub- marginal band of about equal width; pectoral fin spotted internal to these bands. ROBERTS: GASTROMYZONTINE FISH 513 FiGuRE 14. Gastromyzon ridens, 50.0-mm ripe male holotype (MZB 3455). Gastromyzon punctulatus Inger and Chin (Figure 13) Gastromyzon punctulatus INGER AND CHIN, 1961:173 (type- locality “‘Sungai Dapu, a tributary of the Baleh River near the mouth of the Sungai Putai, Third Division, Sarawak’’). This species is known only from the type- specimens collected in tributaries of the Baleh River, Rajang basin, Sarawak. MATERIAL EXAMINED.—FMNH 68117, 3:32.9-53.6 mm, paratypes, same data as holotype. DIAGNOSIS (partly after Inger and Chin 1961).—Larger specimens of Gastromyzon punctulatus differ from all other Gastromyzon except G. ridens in having a relatively truncate snout (vs. relatively more rounded in all other Gastromyzon and in smaller G. punctulatus) and a transverse row of papillae posterior to lower lip (not observed in any other Gastro- myzon). Snout gently sloping downward in front of eyes. No postoral pouch. Gill opening slightly angular, its length about 1.5 times eye diameter. Subopercular groove broadly interrupted or ab- sent in larger specimens, continuous in small specimens. Scales in lateral series about 58-59; predorsal scales about 40-42; scale rows above lateral series 21-22; circumpeduncular scales about 33. Pectoral fin reaches pelvic fin; pelvic fin falls short of anal fin; depressed dorsal fin falls short of level of anal-fin origin. Tubercles on dorsal surface of head, body, and fins minute or absent except for two or three rows of widely spaced small conical tubercles on anterior rim of snout. Dorsal surface of pectoral fin without dense patches of small tubercles. Two largest specimens examined by me (53.0- mm gravid 2 and 53.7-mm ripe ¢) with faint, pale round spots, relatively large, all about the same size, and lying close together, over entire dorsal surface of head and body including caudal peduncle; sides of body otherwise uniformly brownish; paired fins longitudinally banded, with pale (unpigmented) distal margins; dorsal- PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 20 514 I GG VAN ES Clee I Cu SIE I fi fe SudpId “1 Z I Z I € I I I snjpjnjound *5 I I Gr Cc I SG I 9° XG € “a9 DJOINUOU “fy [ees econ al GeO? taal I 8 Lees Oe LT eat sidajpspu “1 I i GG ks I CaerG I I G@ eis iS I VAL ll SIE IE 4aispdopida] *D I I i @L © 8 tl VP I SG fe 87 snipiospf I ¢ € ¢ € I 6 6 snjpydajouaj) “5 v ia I (Ge ete S Gi 6 CG [il SNJIDAJUOI *H) Oy WAS 41A2 AS (AN O83 I (c fi. (AS) £ C s9 ¢ I SISUQIUAO “Y) VC Cam C Cam Cam) Came O)uro Lamia OGG Gms GLC OC mmSG Ga eG “AS S wr YV 6 %*8 8 AL L HI dIAad [e10199g jeuy yesioq “UOZAWOAISDD) NI SLNNOD AVY NIJ GAHONVUg dO SAIONANOAM =“ ATV], ROBERTS: GASTROMYZONTINE FISH fin and anal-fin rays dusky, interradial mem- branes depigmented except for a few oblong dark spots; caudal fin with four or five dark nar- row Vertical bands. A small specimen, 32.9 mm, is similarly colored, but spots are more distinct, and those on head more variable in shape, ver- miculate rather than round. Gastromyzon ridens new species (Figure 14) TypPE-MATERIAL.—Holotype: MZB 3455, 50.0-mm ripe 3d, mainstream of Sungai Pinoh 20-60 km upstream from Nan- gapinoh, Kapuas basin, T. R. Roberts, S. Woerjoatmodjo, 21 July 1976. Paratypes.—CAS 49322, FMNH 94228, USNM 230249, BMNH uncat., MNHN uncat., RMNH uncat., 7:24.3-— 35.7 mm, same data as holotype; BMNH 1893.3.6.270-275, 6:35.4-48.6 mm, Senah, Sarawak; SU 32378, 9:21.2-36.5 mm, Sadong River, Sarawak. DIAGNOSIS.—Gastromyzon ridens differs from all other Gastromyzon in having sublacrimal groove very deep and enlarged, extending pos- teriorly onto cheek (vs. sublacrimal groove re- stricted to ventral surface of head in all other Gastromyzon); length of gill opening less than or barely equal to eye diameter (vs. equal to or greater than eye diameter); and snout relatively truncate, often with a slight ventral concavity at its tip (snout more rounded in all other Gastro- myzon except G. punctulatus, which lacks ven- tral concavity at snout tip). Pectoral-fin rays 25-30, modally 27 (vs. 22-28, modally 26 or less in all other Gastromyzon; Table 1). Snout strongly sloping downwards in front of eyes (more gently sloping in all other Gastro- myzon except G. contractus and G. monticola). No postoral pouch. Gill opening vertical. No subopercular groove. Scales in lateral series 56-78; predorsal scales 32-52; scale rows above lateral series 15-26; circumpeduncular scales about 30-40. Pectoral fin considerably overlaps pelvic fin, perhaps more so than in any other Gastromyzon except G. contractus; pelvic fin falls short of anal fin; depressed dorsal fin falls far short of level of anal-fin origin. Male holo- type, 50.0 mm, has serrae on pelvic fin relatively larger and more heavily developed than in any other specimen of Gastromyzon examined, and tubercles minute or absent on head, body, and fins except for numerous moderately large con- ical tubercles in a narrow band near snout tip. Serration and tuberculation of paratypes similar to that of holotype but less well developed, and some (2 2?) lack tuberculation on snout tip. 515 Dorsal and lateral surfaces of head and body entirely or almost entirely covered with small, pale round spots; some specimens with posterior portion of body, body posterior of head, or en- tire head and body more or less uniformly pale or blanched, suggesting behavioral or physiolog- ical color change (as in G. contractus). Fins dusky or faintly marked, median fins with me- lanophores mainly confined along fin rays. ETYMOLOGyY.—From the Latin ridens, laugh- ing or smiling, in reference to the peculiar coun- tenance of this species caused by the extension of the sublacrimal groove onto the side of its head. Glaniopsis Boulenger Glaniopsis BOULENGER, 1899:228 (type-species Glaniopsis hanitschi Boulenger, 1899, by monotypy). Ficure 15. Ventral surface of head. (a) Glaniopsis han- itschi; (b) Glaniopsis multiradiata (b,, b,, 6; = rostral, max- illary, mental barbels; b;, b, = mandibular barbels; hs = horny sheaths of upper and lower jaws; 11, ul = lower lip, upper lip; mp = mental pad). 516 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 20 ne 12° G. hanitschi So> G. denudata Membikit Baleh Tinjar Mulu 4°- a <4@¢e * © O P We n6e = G. multiradiata, G. gossei —5° 110° 115° FiGure 16. Distribution of Glaniopsis. DIAGNosIs.—Glaniopsis differs from all other Gastromyzontinae in having a subterminal (rath- er than inferior) mouth, and from all other Bor- nean Gastromyzontinae in having two (instead of only one) mandibular barbels at each corner of mouth (Fig. 15). Dorsal fin with two simple and six or seven branched rays; anal fin with two simple and five branched rays; posteriormost dorsal-fin and anal-fin rays not split to base. Pectoral and pel- vic fins widely separated; pectoral fin with one simple and 11-17 branched rays, with an outer ‘“‘adhesive’? and inner ‘‘vibratory’’ portions (Hora and Jayaram 1951), its origin posterior to head. Pelvic fins separate, each with | simple and 7-10 branched rays. Suprapelvic flap ab- sent. Adhesive pads present on first three to five pectoral-fin rays and first three pelvic-fin rays. Sexually mature males with small or minute tu- bercles widely distributed on head, body, and fins; males with most highly developed tuber- culation have median-fin rays and dorsal surface of paired-fin rays with uniserial rows of conical tubercles; tubercles on body minute, pricklelike, usually one per scale (many scales without tu- bercles). Females nontuberculate except for a few small conical tubercles in uniserial rows on dorsal surface of anteriormost four or five pec- toral-fin rays. Key to Glaniopsis la. Dorsal fin with 7 branched rays ________ 2 1b. Dorsal fin with 6 branched rays. 3 2a. Pectoral fin with 15-17 and pelvic fin with 8-10 branched rays _______ G. multiradiata 2b. Pectoral fin with 12-14 and pelvic fin with 6-9 branched rays —_ 33 rae Baleh, Tinjar, and Mulu Glaniopsis (see accounts below for additional information) 3a. Maximum of nine lateral scale rows (mid- lateral scale row plus three scale rows above and five scale rows below it); prin- cipal caudal-fin rays usually less than 9+9 (most often 8-F8) ae G. denudata 3b. At least 20 lateral scale rows; principal caudal-fin rays almost always 9+9 4 4a. Caudal peduncle depth 7.6—9.4; adpressed nasal barbel extends posteriorly at least to posterior margin of eye; eye diameter 30- 42, 3.5-4.6 in interorbital width; horny sheath of upper jaw varying from gently convex to strongly projecting (beaklike) WEE eet G. hanitschi, Membikit Glaniopsis (see accounts below for additional information) 4b. Caudal peduncle depth 9.2—12.2; adpressed nasal barbel extends posteriorly no farther than middle of eye; eye diameter 24-30, 2.2—3.1 in interorbital width; horny sheath of upper jaw varying from slightly concave to strongly notched medially ___ G. gossei ROBERTS: GASTROMYZONTINE FISH 517 FIGURE 17. Glaniopsis denudata new species (Figure 17) TYPE-MATERIAL.—Holotype: BMNH 1957.2.27:1, 34.5 mm. Sungai Kidikarok, North Borneo; altitude 4500 ft [ca 1370 m]; 26 Aug. 1956; Cambridge North Borneo Expedition. Paratypes.—BMNH 1957.2.27:2-15, 14:19.4-34.9 mm. Same collection data as holotype; IRSNB 618, 1:41.8 mm. Sungai Silau Silau, a headwater tributary of Sungai Liwagu, Labuk basin, near Tenompak, Mount Kinabalu, altitude 5000 ft [ca. 1500 m]; 7 Oct. 1971; Léopold III, J. P. Gosse. DIAGNOsISs.—Glaniopsis denudata, perhaps the smallest species of Glaniopsis, differs from all other Glaniopsis in having extremely reduced squamation and usually fewer than 9+9 princi- pal caudal-fin rays. Largest specimen a gravid femaie, 41.8 mm. Squamation consists of lateral scale series plus maximum of about three scale rows above and five below it, or total of about nine scale rows (vs. 20 or more in all other Glan- iopsis). Principal caudal-fin rays variable, most often 8+8, perhaps always less than 9+9. Sex- ually mature males with minute, pricklelike tu- bercles scattered on abdomen in front as well as behind pelvic fins. Dorsal-fin origin distinctly posterior to a vertical through base of last pel- vic-fin ray. Pelvic-fin origin much nearer pec- toral-fin origin than to anal-fin origin. Eye 17-25, 1.9-2.6 in interorbital space. In- terorbital space 9.7-12.7. Branched dorsal-fin rays 6(13). Principal caudal-fin rays variable, most often 8+8: ?9+29(1), 9+8(1), 9+8 or 9+9(1), 8+9(1), 8+8(6), 7+8(1). Pectoral-fin rays 113(6), i14(6), or 115(1). Pelvic-fin rays i7(13). Glaniopsis denudata, 34.5-mm mature male holotype (BMNH 1957.2.27:1). Length of nasal, outer mandibular, and some- times maxillary barbels about equal to or slightly more than eye diameter, otherwise length of bar- bels less than eye diameter. Adpressed nasal barbel failing to reach exposed portion of eye or at most reaching only slightly beyond front mar- gin of eye. Mental barbel flaplike. Lateral scale series incomplete, with 102-111 scales, some scales absent or missing posterior- ly, which, if present, would yield counts of about 115-120 (observations on three specimens lightly stained with alizarin). Dorsum and ab- domen entirely scaleless. Tubed scale series complete or continuous on anterior one-fourth to one-half of body, always arrested before level of dorsal-fin origin and entirely absent poste- riorly. In specimens less than 24 mm, tubed scale series absent or absent except for short anterior segment of up to about six tube-bearing scales. Myotomal muscle masses more sharply defined externally than in any other Glaniopsis, divisions between them clearly visible for entire length of body (vs. scarcely noticeable at all, or clearly visible only on posterior third of body). All specimens in type-series with nearly iden- tical coloration. Head without markings, dusky dorsally and laterally to just below level of eyes, without melanophores ventrally. Ora! barbels and lips without melanophores. Upper fourth to half of body dusky, with fine melanophores uni- formly distributed everywhere except in vicinity 518 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 20 FIGURE 18. of dorsal-fin base: a dark area due to densely clustered fine melanophores circumscribing dor- sal-fin base; a pallid area with few or no mela- nophores extending on dorsum in front of dorsal fin for a distance about equal to dorsal-fin base (some juvenile specimens of G. hanitschi dis- play a tendency toward this coloration, but with much less marked contrast between dark and pallid areas; similar coloration not observed in juveniles or adults of other Glaniopsis). Lower half of body pale, without melanophores except near midline of body, dusky area dorsal to pec- toral-fin base, very faint dusky area lateral to pelvic-fin base, and dark area lateral to anal-fin base due to concentrations of melanophores. Fins unpigmented or dusky, without distinct markings except for concentration of melano- phores on caudal-fin base immediately posterior to hypural fan. ETYMOLOGY.—The feminine adjective denu- data, Latin, “‘nude,’’ refers to the very limited squamation and sharply defined myotomal mus- cles characteristic of the species. Glaniopsis gossei new species (Figure 18) TyPE-MATERIAL.—Holotype: IRSNB 621, 40.5 mm. Arur Dalan, a torrential headwater of Sungai Padapur, Baram ba- sin, near Bario, Sarawak; altitude 3500 ft [ca. 1100 m]; 11 Oct. 1971; Léopold III, J. P. Gosse. Paratypes.—IRSNB 622, 19:28.6-46.5 mm. Same collection data as holotype. Glaniopsis gossei, 40.5-mm mature male holotype (IRSNB 621). DIAGNOsIS.—Glaniopsis gossei differs from all other Glaniopsis in having a series of depig- mented vertical bars on sides of body (these may be lost or obscured in specimens which are pal- lid, as if bleached or faded) and horny sheath of upper jaw strongly notched or concave medially, rather than strongly convex medially (beaklike) or evenly curved. It also differs from G. hanit- schi in having larger eyes, narrower interorbital space, and shallower caudal peduncle; from G. multiradiata in having fewer dorsal, pectoral, and pelvic-fin rays, larger eyes, and longer bar- bels; and from G. denudata in having more ex- tensive squamation. Eye diameter 24-30, 2.2-3.1 in interorbital space. Interorbital space 9.2—11.0. Depth caudal peduncle 9.2—12.2. Branched dorsal-fin rays in- variably 6 (20). Principal caudal-fin rays 9+9 (19; one specimen with broken fin not counted). Pec- toral-fin rays il1 (2), 112 (14), or i13 (3). Pelvic- fin rays 17 (20). Dorsal-fin origin on a vertical through origin of last pelvic-fin ray or slightly posterior to it. Pelvic-fin origin closer to anal-fin origin than to pectoral-fin origin. Length of nasal and outer mandibular barbels about 1.5 times eye diameter. Length of other barbels except mental barbel about equal to eye diameter. Adpressed nasal barbel extends pos- teriorly at least to middle of eye but not beyond exposed portion of eye. Mental barbel barbel- eee ee a ROBERTS: GASTROMYZONTINE FISH 519 us , iY). Ae nt i . DRA RIE Mite VN u : FiGurE 19. Glaniopsis hanitschi. (a) 59.4-mm mature male (IRSNB 17540); (b, c) 59.4-mm mature female (IRSNB 17541). like, about one-half as long as inner mandibular barbel. Sides of body completely covered with scales which tend to be embedded (without free pos- terior margins) and slightly dispersed (not over- lapping), especially on posterior half of body, making them difficult to count. Over 100 scales in lateral series. Dorsum in front of dorsal fin scaleless, behind dorsal fin covered with scales. Entire abdomen scaleless(?) except for a few scales near vent and anal-fin origin. Tubed scale series complete on anterior half of body, varia- bly interrupted on posterior half. ETYMOLOGY.—Named for Jean-Pierre Gosse, who collected much of the material reported upon in this paper, including the type-series of this species. Glaniopsis hanitschi Boulenger (Figures 15a, 19) Glaniopsis hanitschi BOULENGER, 1899:228 (type-locality Ka- damaian River, Mount Kinabalu, altitude 2100 ft [ca. 640 m)). MATERIAL EXAMINED.—IRSNB 17540, 52:31.7-70.8 mm, Sungai Silau Silau, a headwater tributary of Sungai Liwagu, Labuk basin, near Tenompak, Mount Kinabalu, altitude 5000 ft [ca. 1500 mJ]; IRSNB 17541, 28:17.6-69.3 mm, headwater tributary of Sungai Liodan, below Bundu Tahun, Mount Kin- abalu, altitude 4000 ft [ca. 1200 m]; FMNH 68950, 1:53.7 mm, Sungai Kelangaan, near Mesilan base camp, Mount Kinabalu, altitude 5000 ft [ca. 1500 m]; FMNH 47985, 1:58.7 mm, Jes- selton District. DIAGNOosIS.—G. hanitschi is the largest and most robust Glaniopsis, attaining 97 mm, with the deepest caudal peduncle, broadest interor- bital width, smallest eyes, and longest barbels. 520 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 20 Ficure 20. Glaniopsis multiradiata, 44.5-mm female holotype (IRSNB 619). Caudal peduncle depth 7.6-8.9 (vs. 9.2-12.7 in all other species); interorbital width 7.9-9.6 (vs. 9.2-12.7; eye diameter 30-42, 3.74.6 in inter- orbital width (vs. 17-37, 1.9-3.1 in interorbital width); and adpressed nasal barbel invariably extending posteriorly beyond exposed portion of eye. Mental barbel flaplike. Branched rays in dorsal fin 6 (30) (7 in two specimens with posteriormost ray abnormal), in pectoral fin 12 (20) or 13 (12), and in pelvic fin 6 (1), 7 (30), or 8 (1). Principal caudal-fin rays 9+9 (22) or 10+9 (1) (8+9 in one specimen with anomalous vertebral column). Sides of body entirely covered with close-set, regularly overlapping scales with exposed pos- terior margins except for a small scaleless area immediately above base of pectoral fin. Nearly 30 complete scale rows near middle of body and 20 or more on caudal peduncle. Midlateral scale row complete, with about 125-146 scales (19). Dorsal surface of body anterior to dorsal fin varying from scaleless to covered with scales for five-sixths of distance to occiput, but never completely scaled. Dorsal surface of body pos- terior to dorsal fin completely covered with scales, or with only small scaleless areas im- mediately posterior to dorsal fin and on keel-like ridge over upper procurrent caudal-fin rays. Ab- domen scaleless or nearly scaleless except for a few dispersed, embedded scales near anal-fin base and just anterior to vent. Lateral line canal tube-bearing scale series usually complete on anterior half of body, variably interrupted or in- complete on posterior half of body. Head without markings. Body with 7-18 ver- tical or slightly oblique dark bars, mainly on dor- sal half of sides, partially breaking up into upper and lower portions in some specimens, and reg- ularly or irregularly arranged, sometimes drop- ping out or ‘‘fusing’’; some bars may extend be- low midlateral line, and an occasional bar may lie entirely below it, especially in the caudal re- gion, but bars do not continue onto or across dorsum either in front or behind dorsal fin. An- teriormost bar usually immediately behind head, posteriormost just in front of caudal-fin inser- tion; usually a bar near dorsal-fin origin. In some specimens two or more bars “‘fuse’’ to form a single broad mark. Bars absent or faint in spec- imens less than 32 mm. A few larger specimens with a narrow darkly pigmented area paralleling occiput and traversing dorsum immediately pos- terior to head. Two large specimens with sepa- rate markings which resemble bars on side of body and traverse dorsum anterior to dorsal fin. All specimens with a dense cluster of melano- ROBERTS: GASTROMYZONTINE FISH phores, sometimes faint, on lateral surface of pelvic-fin base and sometimes extending onto basal portion of first 1-2 pelvic-fin rays. Dorsum and sides of body with lozenge-shaped clusters of melanophores centered on each scale. Dorsal and lateral surfaces of head, nasal barbel, upper lip, dorsal and caudal fins, and dorsal surface of paired fins dusky. Ventral and ventrolateral por- tions of body, ventral surface of head excluding upper lip, lower lip, all barbels except nasal bar- bel, and ventral surface of paired fins colorless or depigmented (without melanophores). Glaniopsis multiradiata new species (Figures 156, 20) TyYPE-MATERIAL.—Holotype: IRSNB 619, 44.5 mm. Arur Dalan, a torrential headwater of Sungai Padapur, Baram ba- sin, near Bario, Sarawak; altitude 3500 ft [ca. 1100 m]; 11 Oct. 1971; Léopold III, J. P. Gosse. Paratypes.—IRSNB 620, 30:24.3-59.1 mm. Same collection data as holotype. DIAGNOsIs.—Glaniopsis multiradiata has more rays in the paired fins and shorter barbels than any other Glaniopsis: pectoral-fin rays i15—17, usually i116 (vs. ill—14 in other species of Glan- iopsis); pelvic-fin rays usually 19, exceptionally 18 or 110 (vs. usually 17, exceptionally i6 or i8); and length of all barbels equal to or less than eye diameter, adpressed nasal barbel usually failing to reach exposed portion of eye. Eye diameter 20-37, 2.0-3.1 in interorbital space. Interorbital space 9.3-12.6. Depth caudal peduncle 9.7—12.3. Branched dorsal-fin rays in- variably 7 (31). Branched anal-fin rays invariably 5 (31). Principal caudal-fin rays 9+9 (14), 94+8 (1). Pectoral-fin rays i15 (4), i16 (8), or il7 (7). Pelvic-fin rays i8 (2), 19 (28), or i10 (1). Dorsal- fin origin on a vertical through origin of last pel- vic-fin ray or slightly posterior to it. Pelvic-fin origin closer to anal-fin origin than to pectoral- fin origin. Length of all barbels usually less than eye di- ameter. Outer mandibular barbel longest, its length usually less than, but sometimes equal to, eye diameter. Adpressed nasal barbel usually failing to extend posteriorly to exposed portion of eye, at most extending slightly beyond free anterior margin of eye. Mental barbel barbel- like, half as long as inner mandibular barbel. Sides of body completely covered with close- set, regularly overlapping scales with exposed posterior margins, except for a small naked area just above pectoral-fin base; about 30 complete 521 scale rows near middle of body and 20 on caudal peduncle. Lateral scale row complete, with about 106-130 scales (17). Dorsum in front of dorsal fin usually covered with scales almost to occiput, always scaled for at least half its length in front of dorsal fin. Dorsum posterior to dorsal fin completely scaled except for a small scaleless area immediately posterior to dorsal fin in some specimens. Abdomen posterior to pelvic fins covered with embedded scales. Abdomen ante- rior to pelvic fins scaleless except for a few embedded scales median to pelvic-fin bases. Ventral surface of body posterior to anal fin completely scaled. Tubed scale series either complete, complete except for a few scales with- out tubes near end of series, or with short tube- less segments on posterior half of body in a few specimens. Glaniopsis multiradiata has a variably mottled coloration distinct from that of other Glaniopsis. Mottles largest and best defined on dorsum, es- pecially anterior to dorsal fin; fainter and smaller but similar mottles extend onto the head, in- cluding gill covers, cheeks, and snout in a few specimens; almost all specimens have mottles on dorsum, although varying in distinctness, ex- cept for two smallest specimens, 24.3 and 29.2 mm, which are pallid, as if bleached. Majority of specimens partially decolored, either on one or both sides, as if bleached. Path of lateral line marked by a thin dark longitudinal band which is absent or much less distinct in other Glaniop- sis and which persists in specimens otherwise largely decolored. In specimens with mottlings on sides of body, these tend to be less prominent than those on dorsum. Abdomen colorless. Fins darkly pigmented compared to other Glaniopsis. Caudal fin in many specimens with three wavy vertical bands. Dorsal and anal fins with slight concentrations of melanophores suggesting in- cipient barred pattern. Dorsal, anal, and dorsal surfaces of paired fins often with fine melano- phores concentrated along fin rays, giving them a dusky appearance. Dorsal surface of pectoral- fin base dusky or darkly pigmented, more so than in other Glaniopsis. A cluster of melano- phores on lateral surface of pelvic-fin base. Compared to Glaniopsis hanitschi, G. multi- radiata has body slightly more elongate or slen- der, especially posteriorly; snout more de- pressed; ventral surface of head flatter; lips (especially lower) slightly less arcuate; lower lip 522 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 20 interrupted medially by feebly developed mental pad from posterolateral corners of which mental barbels arise; bases of outer and inner mandib- ular barbels more widely separated; dorsal fin slightly smaller; caudal fin (correlated with shal- lower caudal peduncle) much smaller; caudal fin slightly forked, its upper lobe slightly larger (rather than truncate); and paired fins larger. Pectoral fin about 10-15% longer, its surface area when expanded about one-third greater; first pectoral-fin ray noticeably curved or bowed posteriorly instead of nearly straight or only slightly curved; fringes on distal ends of branched rays less well developed; adhesive pads on ventral surface of outermost rays thick- er; and base of pectoral fin fully one-third long- er. ETYMOLOGY.—The feminine adjective multi- radiata comes from multus, Latin, “‘much,”’ plus radiata, Latin, ‘‘rayed,’’ in reference to the more numerous pectoral-fin and pelvic-fin rays characteristic of the species. Membikit Glaniopsis Glaniopsis hanitschi (in part) INGER AND CHIN, 1962: 107-108 (specimens from Sungai Membikit). MATERIAL EXAMINED.—FMNH 68135, 3:46.5-52.4 mm. Sungai Membikit [tributary of Sungai Pegalan, Padas basin, near southern end of Crocker range], Keningau District, Sa- bah, 5°22’N, 116°06’E. These three specimens resemble G. hanitschi more closely than they do any other Glaniopsis but differ from typical G. hanitschi from Mount Kinabalu in having head narrower and more de- pressed, fins smaller, and abdomen more exten- sively covered with scales anterior to pelvic fins. Two of the three specimens also have fewer pec- toral-fin rays and a more incomplete tubed scale series. Eye 34-43, 3.54.1 in interorbital space. In- terorbital space 10.3-10.8 (vs. 7.9-9.6 in G. han- itschi). Depth caudal peduncle 9.0-9.4 (7.6-8.9 in G. hanitschi). Branched dorsal-fin rays 6 (3). Branched anal-fin rays 5 (3). Principal caudal-fin rays 9+9 (2), 9+7 (1, with caudal fin probably injured or abnormally developed). Pectoral-fin rays ill (2) or 112 (1) (12 or more in G. hanit- schi). Pelvic-fin rays i7 (3). Adpressed nasal barbel reaching to or slightly beyond free posterior margin of eye. Upper and lower horny jaw sheaths markedly convex near symphysis. Dorsum and sides of body nearly completely covered with scales, lateral series complete with about 130-142 scales, about 35 complete scale rows near middle of body and 28 on caudal pe- duncle. Entire abdomen posterior to pelvic fins covered with embedded scales; half of abdomen anterior to pelvic fins with embedded scales. Tubed scale series complete or nearly complete on anterior half of body, absent or considerably interrupted on posterior half. Coloration darker (duskier) than in G. hanit- schi, without well-defined vertical bars on a rel- atively pale background; a continuous dark mid- lateral longitudinal band (absent in G. hanitschi). Fine melanophores extending from sides of body further toward and onto abdomen than in G. hanitschi. Nearly all scales with lozenge-shaped cluster of fine melanophores. Largest specimen, 52.4 mm female, uniformly dusky, without ver- tical bars or marks on dorsum or sides of body. Smallest specimen, 46.5 mm female, with about 20 vertical marks in side of body (less well de- fined, and lower lying than vertical bars in G. hanitschi); dorsum with 10 dark transverse marks anterior and 4 posterior to dorsal fin (ab- sent in G. hanitschi). Specimen intermediate in size, a well-tuberculated 47.5 mm male, with markings on sides fewer and less well defined, but similar to those in smallest specimen, and marks on dorsum barely visible or absent. Baleh and Tinjar Glaniopsis MATERIAL EXAMINED.—FMNH 68137, 68151-68153, 69868, 24:20.4-38.0 mm, mainstream and tributaries of Baleh River, Rajang basin, Sarawak; BMNH 1933.8.9.9-10, 15:19.9-39.3 mm, Lejok River, altitude 50-1500 ft [ca. 15-460 m], Tinjar River District, Sarawak. These specimens superficially resemble G. hanitschi but differ from it in their much smaller size, shorter barbels, having seven instead of only six branched dorsal-fin rays, fewer scales, and other minor differences. Eye 24-32, 2.43.0 in interorbital width. In- terorbital width 9.7-11.2. Depth caudal pedun- cle 8.9-10.0. Branched dorsal-fin rays invariably 7. Principal caudal rays usually 9+9. Pectoral- fin rays il12-i14; pelvic-fin rays i7 or i8. Ad- pressed nasal barbel extends posteriorly to middle of eye or almost to posterior border of exposed portion of eye. Upper and lower horny jaw sheaths moderately convex near symphysis. Dorsum and sides of body completely covered with scales, or scales absent from dorsum only a short distance posterior to occiput; lateral ROBERTS: GASTROMYZONTINE FISH scale series with 102-115 scales, about 18-25 scale rows near middle of body, 22-24 on caudal peduncle. Abdomen between pelvic fins covered with scales, otherwise largely scaleless. Dorsal and lateral surfaces of body mottled or with vertical or variably broken vertical bands of equal or variable width. Largest specimen, 39.3 mm, a gravid female. Mulu Glaniopsis MATERIAL EXAMINED.—BMNH uncat., 2:18.4-33.2 mm. Extreme headwater of Sungai Tapin [Baram basin], Gunong Mulu, Sarawak. These two small specimens possibly represent an undescribed species. They differ from all oth- er Glaniopsis examined in having a more slender body. Unlike Glaniopsis from Membikit, Baleh, and Tinjar, which seem closest to G. hanitschi, Mulu Glaniopsis perhaps is closest to G. mul- tiradiata. It agrees with G. multiradiata in gen- eral appearance, color pattern, and in having rel- atively high paired-fin ray counts but differs in having longer barbels, smaller eyes, larger scales, less extensive squamation, and paired fins with slightly fewer rays, as well as a more slender body. Eye 30-31 (20-25 in 5 G. multi- radiata 24.3-36.9 mm), 3.4-3.7 in interorbital space. Interorbital space 8.4-9.0. Body depth 9.5—9.7 (6.4-8.1 in G. multiradiata). Depth cau- dal peduncle 10.2—11.4. Branched dorsal-fin rays 7 (2). Branched anal-fin rays 5 (2). Principal cau- dal-fin rays 9+9 (2). Pectoral-fin rays 113 (1), i14 (1) ((15—17 in G. multiradiata). Pelvic-fin rays i8 (2) (usually 19 in G. multiradiata). Pelvic-fin or- igin closer to pectoral than to anal-fin origin (equidistant between pectoral and anal-fin origins or closer to anal-fin origin in G. hanitschi and G. multiradiata). Length of nasal, rostral, maxillary and outer mandibular barbels equal to or greater than eye diameter. Adpressed nasal barbel reaches at least to middle of eye. Larger specimen, 33.2 mm, with mental barbels barbel-like. Smaller specimen, 18.4 mm, with unusually broad, thin lower lip and large flaps in position normally occupied by mental barbels. Lower lip inter- rupted by a median mental pad as in G. multi- radiata. Larger specimen with dorsum, abdomen, and ventral surface of body posterior to anal fin scaleless; lateral scale series nearly complete with about 100 scales, allowing for 4-5 scales 523 missing or having failed to develop slightly an- terior to hypural fan (125-146 in G. hanitschi, 106-130 in G. multiradiata); at most 6-7 scale rows below and 5 above lateral scale series, or a maximum total of about 13 scale rows on side of body (vs. 25 or more in G. hanitschi and G. multiradiata); tubed scale series complete on anterior half of body, largely discontinuous or incomplete on posterior half. Smaller specimen entirely or almost entirely scaleless. Larger specimen with dorsal and lateral sur- faces of head and body dusky, on body mainly due to large lozenge- or scale-shaped clusters of melanophores overlying nearly every scale; dor- sum anterior to dorsal fin with large mottles sim- ilar to those in G. multiradiata; cluster of me- lanophores lateral to pelvic-fin base; cluster of melanophores toward base of each caudal-fin lobe, fins otherwise without distinct markings. Smaller specimen with fine melanophores uni- formly distributed over dorsal and lateral sur- faces of head and body, and concentrations of melanophores along dorsal-fin base and lateral line canal, otherwise without distinct markings. ACKNOWLEDGMENTS It is a pleasure to thank the following individ- uals who helped in various ways during the course of this study: Oliver Crimmen, Alwyne Wheeler, Keith Banister, and Gordon Howes, BMNH;; Marie-Louise Bauchot and Martine De- soutter, MNHN; Marinus Boeseman and Peter van Helsdingen, RMNH; Han Nijssen, ZMA; Jean-Pierre Gosse and L. Walschaerts, IRSNB; Karsten F. Hartel and William L. Fink, MCZ; Robert K. Johnson, Donald J. Stewart, and Robert F. Inger, FMNH; and Lillian J. Demp- ster, W. I. Follett, and Michael E. Hearne, CAS. Photography is by Alphonse Coleman, MCZ; and Orrin Moon, The Darkroom, San Ra- fael; and scanning electron micrography by Mary-Jacque Mann, SEM lab, National Mu- seum of Natural History. I am especially in- debted to Jean-Pierre Gosse for the opportunity to report on the extensive and beautifully pre- served material of Gastromyzon and Glaniopsis collected by Léopold III and himself, and to Soetikno Woerjoatmodjo and Rajai for helping collect Gastromyzon in the Sungai Pinoh. The ichthyological survey of the Kapuas basin was sponsored by the Museum of Zoology, Bo- gor, Indonesian National Research Council, and 524 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 20 Smithsonian Tropical Research Institute. Most of the research, supported by grant DEB77- 24759 in the Systematic Biology Program, Na- tional Science Foundation, was done at the Cal- ifornia Academy of Sciences and Tiburon Cen- ter for Environmental Studies. LITERATURE CITED BouLENGER, G. A. 1899. Descriptions of two new homal- opteroid fishes from Borneo. Ann. Mag. Nat. Hist. (7)4:228-229. CHEN, Y. 1980. Systematic studies on the fishes of the family Homalopteridae of China. II. Classification of the fishes of the subfamily Gastromyzoninae. [In Chinese, with English summary.] Acta Hydrobiol. Sin. 7:95—120. GUNTHER, A. 1874. Descriptions of new species of fishes in the British Museum. Ann. Mag. Nat. Hist. (4)14:1-8. Hora, S. L. 1932. Classification, bionomics, and evolution of homalopterid fishes. Mem. Indian Mus. (Calcutta) 12:263-330, pls. 10-12. , AND K. C. JAYARAM. 195la. On two new gastromy- zonid fishes from Borneo. Rec. Indian Mus. (Calcutta) 49: 191-196. , AND . 1951b. A note on the systematic position of the genus Glaniopsis Boulenger (Fishes: Cyprinoidea). Rec. Indian Mus. (Calcutta) 48:85-88. INGER, R. F., AND P. K. CHIN. 1961. The Bornean cyprinoid fishes of the genus Gastromyzon Gunther. Copeia 1961(2): 166-176. , AND . 1962. The fresh-water fishes of North Borneo. Fieldiana Zool. 45: 1-268. Popta, C. M. L. 1905. Suite des descriptions préliminaires des nouvelles especes de poissons recueillies au Bornéo Central par M. le Dr. A. W. Nieuwenhuis en 1898 et en 1900. Notes Leyden Mus. 25:171-186. . 1906. Résultats ichthyologiques des voyages scientif- iques de Monsieur le Professeur Dr. A. W. Nieuwenhuis dans le centre de Bornéo (1898 et 1900). Notes Leyden Mus. 27: 1-304, pls. 1-10. RAMASWAMI, L. S. 1948. The homalopterid skull. Proc. Zool. Soc. London 118:515-538. . 1952. Skeleton of cyprinoid fishes in relation to phy- letic studies. IV. The skull and other skeletal structures of gastromyzonid fishes. Proc. Natl. Inst. Sci. India 18(6):519-538. Roserts, T. R. In press. Unculi (horny projections arising from single cells), an adaptive feature of the epidermis of ostariophysan fishes. Zool. Scripta. Sivas, E. G. 1953. Classification, zoogeography and evolu- tion of the fishes of the cyprinoid families Homalopteridae and Gastromyzonidae. Rec. Indian Mus. (Calcutta) $0:173-263, pl. 5. VAILLANT, L. 1889. Sur les poissons d’eaux douces de Bor- néo. C. Rend. Congr. Internat. Zool. Paris, 1889:81-82. . 1891. [Note on Lepidoglanis]. C. Rend. Soc. Philo- math. Paris, 1890-1891, 2:6. . 1893. Contribution a l'étude de la faune ichthyolo- gique de Bornéo. Nouv. Arch. Mus. Paris 5(3):23-114, pls. 1-2. . 1902. Résultats zoologiques de |’expedition scienti- fique néerlandaise au Bornéo Central. Poissons. Notes Ley- den Mus. 24:1-166, pls. 1-2. WEBER, M., AND L. F. DE BEAUFORT. 1916. The fishes of the Indo-Australian Archipelago. III. Ostariophysi: II Cy- prinoidea, Apodes, Synbranchi. E. J. Brill, Leiden. xv + 45S. WICKLER, W. 1971. Verhaltenstudien am einem hochspezial- isierten Grundfisch, Gastromyzon borneensis (Cyprinoidea, Gastromyzonidae). Z. Tierpsychol. 29:467-480. CALIFORNIA ACADEMY OF SCIENCES Golden Gate Park San Francisco, California 94952 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Vol. 42, No. 21, pp. 525-535, 33 figs. May 14, 1982 LATE MIOCENE BALANID CIRRIPEDIA FROM THE BASAL WILSON RANCH BEDS (‘‘MERCED’”’ FORMATION), SONOMA COUNTY, NORTHERN CALIFORNIA By Victor A. Zullo Department of Earth Sciences, University of North Carolina at Wilmington, Wilmington, North Carolina 28403 and Raj B. Guruswami-Naidu Department of Geology, California Academy of Sciences, San Francisco, California 94118 ABSTRACT: The basal conglomerate of the Wilson Ranch beds (Merced Formation of authors) contains abun- dant barnacle remains including three identifiable species of the family Balanidae. Balanus sp. aff. B. nubilus Darwin, 1854, is similar to extant B. nubilus and the California late Miocene-early Pliocene species B. proxi- nubilus Zullo, 1979, but differs in the internal morphology of the scutum. Balanus irradians new species is a member of the Balanus balanus (Linnaeus, 1758) complex distinguished by its lack of radii. Notomegabalanus(?) insperatus new species is a megabalanine with Southern Hemisphere cool-temperate affinities distinguished by its prominently ribbed parietes. A late Miocene age is assigned to the basal conglomerate on the basis of its stratigraphic relationship to a radiometrically dated tuff within the Wilson Ranch beds and on biostratigraphic evaluation of the molluscan fauna. The fauna of the basal conglomerate suggests deposition in an immediately subtidal environment subjected to wave and current action, and a cool temperate marine hydroclimate. INTRODUCTION A fossiliferous marine conglomerate of late Miocene age in southwestern Sonoma County, northern California (Fig. 1) contains an abun- dance of balanid barnacle remains. The speci- mens consist of shells, disarticulated compart- mental plates, and a few dissociated scuta representing four species. One of these species cannot be identified beyond the generic level. A second is similar to, but not conspecific with, the extant Pacific coast species Balanus nubilus Darwin, 1854. The third, based on shell char- acters alone, is sufficiently unique to permit its description as a new species with affinities to the modern Balanus balanus (Linnaeus, 1758) complex. The fourth is a new species of mega- balanine that appears to represent one of the newly delimited extant austral genera. STRATIGRAPHY Outcrops of marine sandstone in southwest- ern Sonoma County were for many years con- sidered correlatives of the type Merced For- mation of the San Francisco peninsula and [525] 526 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 21 122° 30° MENDOCINO =" 10 KM GEOLOGIC SYMBOLS WILSON RANCH BEDS FAULT Si SONOMA @ HEALDSBURG SEBASTOPOL 15° QUAD TWO ROCK 7.5' QUAD @ PETALUMA SAN PABLO BAY 4 General locality map for CASG locality 54135, lower Wilson Ranch beds, southwestern Sonoma County, Cal- ifornia. FIGURE 1. referred to that unit (e.g., Dickerson 1922; Weaver 1949; Travis 1952). Higgins (1960) doc- umented lithologic and faunistic differences be- tween the type Merced Formation and the So- noma deposits, and provided compelling arguments that the two units were deposited in separate basins. Higgins, however, did not pro- pose a separate formational name for the So- noma County deposits. A tuff interbedded with the sandstone was described by Osmont (1905) as the Sonoma Tuff. In ensuing years, following the work of Dick- erson (1922), the name Sonoma has been used to refer to the Neogene volcanic assemblage of eastern Sonoma and Napa counties. The original Sonoma Tuff can be regarded as one of a series of tuffaceous intertongues of the eastern volca- nic assemblage into the western marine se- quence. Osmont (1905) also described a mollus- can fossil locality in the marine sandstone containing the Sonoma Tuff, to which he applied the name Wilson Ranch beds. As this is the ear- liest formal stratigraphic name that has been used for this Neogene marine unit,.we will refer to these deposits as the Wilson Ranch beds. K-Ar dates from the pumiceous, vitric tuff in- terbed described by Osmont have given an age range of 5.7 + 0.6 to 6.1 + 0.1 m.y. (Sarna- Wojcicki 1976). Bartow and others (1973) re- ported molluscan assemblages in the Wilson Ranch beds that ranged in age from the early Pliocene below, to late Pliocene above the dated tuff. The early and late Pliocene molluscan as- semblages were correlated with Hemphillian (late Miocene to early Pliocene) and Blancan (late Pliocene to early Pleistocene) land mammal assemblages, respectively, from the Petaluma Formation immediately to the east. These rela- tive ages are in terms of conventional Pacific coast usage. If, however, 5.0 m.y. is acceptable as the age of the Miocene-Pliocene boundary as proposed by Berggren (1972, 1978) and Van Eysinga (1975), then that part of the Wilson Ranch beds below the dated tuff is of Miocene age. This conclusion is supported by biostrati- graphic evaluation of the molluscan fauna by Barry Roth (California Academy of Sciences, pers. commun., 1981). Roth would correlate the lower Wilson Ranch beds with the Pancho Rico Formation of Monterey County, California, which he and others now regard as late Miocene in age (Addicott 1976; Roth and Guruswami- Naidu 1978). The barnacles described herein are from Cal- ifornia Academy of Sciences Department of Ge- ology (CASG) locality 54135. This locality is in a coquina near the top of the basal conglomerate of the Wilson Ranch beds in the Two Rock quadrangle, U.S. Geological Survey 7.5’ series, 1954 edition, revised 1971, and is in that part of the section referred by Osmont (1905) to the San Pablo Formation of the eastern San Francisco © Bay region. A precise locality description is on file with the Department of Geology. At the ex- posure containing CASG locality 54135, the Wilson Ranch beds unconformably overlie the Franciscan assemblage (Fig. 2). The basal 0.5 m consists of a fining-upwards conglomerate com- posed of coarse sand, rounded pebbles, and an- gular pebble- to cobble-sized fragments of Fran- ciscan-assemblage rocks in a fine-grained sandstone matrix. The upper part of the con- glomerate is the coquina (CASG locality 54135) which is composed of predominantly horizon- tally bedded invertebrate shells. A vertebrate ZULLO & GURUSWAMI-NAIDU: NORTHERN CALIFORNIA MIOCENE BALANIDS 527 fossil lag, corisisting of scattered intact elements and abundant, well-rounded bone fragments, overlies the coquina. The vertebrate lag grades upward into 3.4 m of massive, moderately in- durated, fine-grained sandstone containing scat- tered invertebrates and rare vertebrate remains. This sandstone is overlain by tuffaceous sand- stone to the top of the exposure. The basal con- glomerate at this locality is estimated to be 50 m stratigraphically below the dated tuff in the Wilson Ranch beds (Peter Rodda, California Academy of Sciences, pers. commun., 1981). PALEOECOLOGY The petrology and fauna of the basal conglom- erate suggest that the barnacles inhabited a rocky intertidal or immediately subtidal environ- ment exposed to moderate or heavy surf. The coquina itself, and its relationship to the under- lying fining-upwards conglomerate, suggest a time of stillstand after rapid transgression in a shallow depositional basin affected by wave base and/or current action. According to Barry Roth (pers. commun., 1981) the molluscan fauna suggests a marine hydroclimate similar to that of modern Puget Sound (i.e., cool temperate). These environmental conditions are supported by the composition, morphology, and preser- vation of the barnacle fauna. SYSTEMATIC ACCOUNT Subclass CiRRIPEDIA Burmeister, 1834 Order THoRACICA Darwin, 1854 Superfamily BALANOIDEA (Darwin), Newman and Ross, 1976 Family BALANIDAE Darwin, 1854 Subfamily BALANINAE Darwin, 1854 Balanus sp. aff. B. nubilus Darwin, 1854 (Figures 3-11) Four complete shells, one partial shell, sev- eral disarticulated compartmental plates, one entire scutum, four fragmentary scuta, and a few fragments of basis from CASG locality 54135 are here referred to a species that is similar to, but probably not conspecific with, Balanus nubilus. The shells are conic, the radii are moderately well developed with oblique, jagged summits, and the orifice is toothed. The ribbing of the shell interior is not prominent, and the basis in larger individuals is cup-shaped and profusely porous near the margins. The parietal septa are TUFFACEOUS 9 SANDSTONE MASSIVE, FINE-GRAINED, SANDSTONE WILSON RANCH BEDS BASAL CONGLOMERATE CAS 54135 FRANCISCAN ASSEMBLAGE FIGURE 2. lower Wilson Ranch beds, southwestern Sonoma County, Cal- ifornia. Stratigraphic setting for CASG locality 54135, crowded and thin, and the narrow, elongate pa- rietal tubes rarely possess transverse septa. The scuta that appear to be associated with these shells are higher than wide, slightly bowed outwards near the apex, and with a slightly re- flexed tergal margin. The exterior is ornamented by prominent, closely spaced growth ridges that are faintly crenate, and the basitergal angle is rounded. The articular ridge is short, less than one-half the length of the tergal margin, and slightly reflexed over the narrow, shallow, artic- ular furrow. The adductor ridge is erect, highest along the margin of the large, oval, and deeply impressed adductor muscle pit, but extending both apically and basally beyond the boundaries of the adductor pit. The depressor muscle pit is large, triangular, shallow, and bordered on its occludent margin by a low ridge. The depressor muscle pit has one or more narrow, low longi- tudinal ridges within. The depressor muscle pit located at the basioccludent angle is triangularly 528 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 21 Figures 3-11. Balanus sp. aff. B. nubilus Darwin, 1854. Figs. 3-5. Exterior, interior, and basal views of lateral plate, hypotype CASG 60881; greatest height 47.5 mm. Figs. 6-7. Basal and alar sutural edge views of carinorostral and carinal plates, hypotype CASG 60882; greatest height 61 mm, greatest wall thickness 17 mm (note transverse septa in parietal tubes). Figs. 8-9. Interior and exterior views of scutum, hypotype CASG 60883; height 23 mm. Fig. 10. Side view of shell, hypotype CASG 60884; greatest height 49 mm. Fig. 11. Fragment of basis, hypotype CASG 60885; greatest diameter of fragment 35.5 mm. ZULLO & GURUSWAMI-NAIDU: NORTHERN CALIFORNIA MIOCENE BALANIDS elongate, deeply impressed, and partially over- hung by the occludent margin of the plate. The shell of this species is similar to that of Balanus nubilus and the related fossil species B. proxinubilus Zullo, 1979, from the upper Mio- cene Pancho Rico Formation of central Califor- nia and the upper Pliocene San Diego Formation of southern California. The only qualitative dif- ference is that the radii of the Wilson Ranch species appear to be consistently broader than those of either B. nubilus or B. proxinubilus. The Wilson Ranch scuta differ in having the ad- ductor ridge separate from the articular ridge, and in the much shorter articular ridge. In this regard the internal morphology is somewhat similar to that of the scutum of B. rostratus Hoek, 1883, but the presence of vertical ridges or crests in the depressor muscle pit serves to distinguish the Wilson Ranch scuta and to sug- gest relationship with B. nubilus. Balanus irradians new species (Figures 12-15, 18-21) DIAGNosIs.—Shell of six compartmental plates without radii; alae restricted to sheath area; sutures between compartmental plates ob- scured or represented by linear grooves; pari- etes with large, square to rectangular, trans- versely septate parietal tubes; number of interior ribs greater than number of parietal septa; basis calcareous, solid; distinguished from other members of the Balanus balanus complex by the lack of radii. DESCRIPTION.—Shell thick, low to high conic or dome-shaped, with narrow carinolaterals and a small, untoothed, diamond-shaped orifice; su- tures between the six compartmental plates ob- scured or represented by narrow grooves; radii absent; sutural edges between adjacent com- partmental plates broad, bearing coarse, com- plexly arborescent denticulations; alae narrow, with horizontal summits, and restricted to region of sheath; exterior of parietes usually corroded; uncorroded parietes with external ornamenta- tion of irregular, transverse growth rugae crossed by fine longitudinal striae; some speci- mens bear one to three external ribs on larger compartmental plates; length of sheath at least one-half height of compartmental plate; lower edge of sheath free-standing, acute, with cavity between it and interior of shell wall; interior of shell wall below sheath strongly ribbed, with the 529 largest internal ribs corresponding to the parietal septa, and from one to three smaller ribs be- tween parietal septa; parietal tubes large, nearly square to rectangular, crossed by transverse septa and sometimes secondarily filled in upper third; parietal septa denticulate basally; inner surface of external lamina bears denticulae be- tween parietal septa; basis calcareous, solid. MATERIAL EXAMINED.—Twelve whole shells and 26 disar- ticulated compartmental plates from CASG locality 54135. TyPE MATERIAL.—Holotype, CASG no. 60891; paratypes, CASG nos. 60892 through 60909, and paratype lot CASG no. 60910 in the California Academy of Sciences paleontological type collection. ETYMOoOLoGy.—The specific name is derived from the Latin prefix ir-, without, and radius. DiscUssion.—Balanus irradians and the ex- tant boreo-arctic species B. balanus and B. ros- tratus, constitute a small group in the genus Bal- anus that is characterized by solid, calcareous bases, large, square, parietal tubes, and second- ary ribs between the primary septal ribs of the interior lamella of the parietal wall. The lack of radii and the overall morphology associated with a Shell lacking radii are the major features dis- tinguishing B. irradians from B. balanus and B. rostratus. In other respects, B. irradians shares characteristics with the extant species. The prominent external ribs present on some speci- mens are reminiscent of the ribbed wall of B. balanus, rather than the smooth shell of B. ros- tratus. On the other hand, both B. irradians and B. rostratus have transverse parietal septa that are lacking in B. balanus. The corroded nature of the shells and the type of sediment in which they occur suggest that B. irradians lived in moderate- to high-energy en- vironments. The thick shell wall, the suppres- sion of radii, and the consequent development of a broad, coarsely denticulate sutural area for rigid articulation of the compartmental plates are comparable to shell development in species of Tetraclita Schumacher, 1817, that often inhabit areas of moderate to heavy wave shock and abrasion in the intertidal zone. A consequence of the lack of radii is that the orifice cannot be enlarged by lateral growth along the sutural edges of the plates as the shell increases in basal diameter and height. As Ross (1969) showed for Tetraclita, monometric growth forms enlarge their orifices by abrasion of the older, upper parts of the shell wall. The secondary filling and 530 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 21 FiGurEs 12-21. Balanus irradians new species. Figs. 12-14. Basal, top, and side views of paratype CASG 60892; greatest diameter of base 29 mm, greatest height of shell 15 mm (note fragment of basis in upper part of Fig. 12). Fig. 15. Side view of shell, paratype CASG 60893; greatest height 13 mm. Figs. 16-17. Exterior and interior views of scutum tentatively identified with B. irradians, hypotype CASG 60920; height 10 mm. Figs. 18-20. Side, top, and basal view of shell, holotype CASG 60891; greatest height 24 mm, carinorostral diameter 38 mm. Fig. 21. Top view of eroded shell with prominent ribs, paratype CASG 60894; greatest diameter 38.3 mm. ZULLO & GURUSWAMI-NAIDU: NORTHERN CALIFORNIA MIOCENE BALANIDS 531 the transverse septa of the parietal tubes prevent exposure of the internal tissues of the barnacle when the upper and outer shell wall are re- moved. The affinities of B. irradians with B. balanus and B. rostratus are in keeping with the conclu- sion that the fauna of the basal Wilson Ranch beds lived in hydroclimatic conditions similar to those of modern Puget Sound. Balanus rostra- tus is a subtidal North Pacific boreo-arctic species ranging south to northern Japan in the west and to Puget Sound in the east. Balanus balanus is a lower intertidal and subtidal species of the Arctic, boreal Atlantic, and boreal Pacific, with a distribution similar to that of B. rostratus in the North Pacific. The fossil record of B. ros- tratus is limited to the late Pleistocene on the Pacific coast of North America where it is re- corded as far south as central California (Zullo 1969b), but extends back to the Pliocene in Ja- pan (Yamaguchi 1977). Balanus balanus is found in Pleistocene deposits of both the North Atlantic and North Pacific basins, but has not been recorded from deposits south of southern Oregon (Zullo 1969b). Miocene and Pliocene European records of B. balanus are spurious (Menesini 1968, in part), but the species does occur in glacio-marine sediments of the Miocene part of the Yakataga Formation in southeastern Alaska, and is reported from the Miocene of Ja- pan by Yamaguchi (1971). The morphology of B. irradians, particularly those features peculiar to this species, suggests that B. irradians is a derivative of either B. bal- anus or B. rostratus. The modifications seen in the new species reflect adaptations to life under conditions of pronounced wave shock and abra- sion, perhaps in the intertidal zone in relatively open coastal conditions. Balanus sp. cf. B. irradians (Figures 16-17) A single, nearly intact scutum from CASG lo- cality 54135 is tentatively referred to B. irradi- ans. It is thin, about as broad as high, and slight- ly concave externally between base and apex. The exterior bears closely spaced, semi-erect growth ridges that are finely crenate. The tergal margin is slightly reflexed and the basitergal an- gle is rounded. The articular ridge is convex, reflexed over the articular furrow, and protrudes beyond the tergal margin. The articular furrow is narrow, shallow, and short. The articular ridge is long, fully two-thirds the length of the tergal margin. A short, low, blunt adductor ridge is present along the margin of the large oval ad- ductor muscle pit in the upper half of the scutum and is separated from the articular ridge. The depressor muscle pit is a large triangular area between the adductor muscle pit and the basi- tergal angle. This pit is not bordered by a ridge on its occludent side. The pit for the depressor muscles in the basioccludent angle is large, tri- angular, and deep. This scutum bears some resemblance to those of B. balanus and B. rostratus, but differs from both in its greater breadth, thinness, its short, blunt adductor ridge that is well removed from the articular ridge, and in the greater size and depth of the depressor muscle pit that is not bor- dered by a ridge on its occludent side. This scu- tum differs from others found at CASG locality 54135 and identified as Balanus sp. aff. B. nu- bilus by its greater width, its convex rather than straight articular ridge that is much more re- flexed and much longer, its shorter, blunter ad- ductor ridge, and the lack of striations or ridges in the depressor muscle pit. Balanus sp. (Figures 31-33) Several whole, high conic and cylindric shells cannot be identified beyond the generic level. Their orifices are toothed, the radii are moder- ately sunken with oblique summits, and the pa- rietes are smooth to irregularly plicate, to in- conspicuously ribbed. These shells may represent either B. sp. aff. B. nubilus or Notomegabala- nus(?) insperatus, but their preservation is not conducive to the identification of specific char- acters. Subfamily MEGABALANINAE Newman, 1980 Notomegabalanus(?) insperatus new species (Figures 22-30) D1IaAGNosis.—Shell of six compartmental plates with broad, tubiferous radii; summits of radii oblique; septa of radial sutural edge bearing denticulae on lower sides only; exterior of pa- rietes of larger compartmental plates with three to five prominent ribs, and ornamented by fine, closely spaced radial striae; parietal tubes nu- merous, rectangular, without transverse septa, 532 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 21 7 S Ay 4 y * VAAN Figures 22-30. Notomegabalanus(?) insperatus new species. Figs. 22-24. Interior, exterior, and right radial sutural edge views of rostrum, holotype CASG 60914; greatest height of plate 36.4 mm. Figs. 25-26. Exterior and interior views of lateral plate, paratype CASG 60915; greatest height of plate 20.5 mm. Figs. 27-28. Radial sutural edge and exterior views of lateral plate, paratype CASG 60916; greatest height of plate 28.5 mm. Figs. 29-30. Side view and enlargement of radial sutural edge of lateral plate, paratype CASG 60917; greatest height of plate 18 mm. ZULLO & GURUSWAMI-NAIDU: NORTHERN CALIFORNIA MIOCENE BALANIDS 533 Ficures 31-33. Balanus sp., side views of three shells, hypotypes CASG 60911 through 60913, respectively; greatest height of shells: (Fig. 31) 42 mm, (Fig. 32) 20.5 mm, (Fig. 33) 29 mm. but secondarily filled in upper half; basis un- known, but presumed calcareous; opercular plates unknown; distinguished from other species of Notomegabalanus Newman, 1980, and Austromegabalanus Newman, 1980, by the prominent external ribbing of the shell wall. DESCRIPTION.—Reconstructed shell high con- ic with broad, tubiferous radii and moderately toothed orifice; summits of radii oblique; radial sutural edges septate, with septa bearing dentic- ulae on lower sides only; alae moderately broad with oblique summits, confined to area of sheath; length of sheath less than one-half height of compartmental plates; lower margin of sheath free-standing, with shallow cavity between basal margin of sheath and interior of shell wall; ex- terior of parietes with from three to five promi- nent ribs on the larger compartmental plates, and ornamented by fine, closely spaced radial striae crossed by weak, widely spaced growth lines; interior of shell wall ribbed between base and sheath, internal ribs low, moderately devel- oped, each corresponding to a parietal septum; parietal tubes numerous, without transverse septa, but secondarily filled in upper half; pari- etal septa thin, basally denticulate; outer lamina with one to four thin, half or quarter septa; basis unknown but presumed calcareous from basal denticulation of parietal septa; opercular plates unknown. MATERIAL EXAMINED.—Twenty-four disarticulated com- partmental plates and one possibly complete shell from CASG locality 54135. TyPeE MATERIAL.—Holotype, CASG no. 60914; paratypes, CASG nos. 60915 through 60918, and paratype lot CASG no. 60919 in the California Academy of Sciences paleontological type collection. ETYMOLOGY.—The specific name is Latin for unhoped for or unexpected. Discussion.—Recently, Newman (1980) sub- divided the genus Megabalanus Hoek, 1913, into three genera. Megabalanus s.s. includes most of the Megabalanus tintinnabulum (Lin- naeus, 1758) complex together with related species of modern tropical and warm temperate seas. The two new genera, Austromegabalanus and Notomegabalanus, include fossil and extant species restricted to austral cool temperate and subpolar waters. The only obvious shell char- acter separating Megabalanus s.s. from the two austral genera is that the septa of the radial su- tural edge of Megabalanus are denticulate on both their lower and upper sides, whereas the septa of the austral genera are denticulate only on their lower sides. The two austral genera are separated on characters of their terga; that of Austromegabalanus has a beak and a closed spur furrow, whereas that of Notomegabalanus is not beaked and has an open spur furrow. There are features of the mouth parts as well that can be used to distinguish true Megabala- 534 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42, No. 21 nus from the austral genera, but when dealing with the generic assignment of fossil species, only shell and opercular plate characteristics are available. The revision of Megabalanus proffered by Newman is sound, both from a systematic and biogeographic point of view. However, the Neo- gene fossil megabalanid record from California presents a problem. The species described here and ‘‘Megabalanus’”’ wilsoni (Zullo, 1969a) from the Pliocene San Diego Formation have radial sutural dentitions characteristic of Newman’s austral genera. Furthermore, the tergum of “*M. wilsoni is typical of Notomegabalanus, although the spur furrow is closed in adult specimens, and its resemblance to that of N. algicola (Pilsbry, 1916) from South Africa was noted previously (Zullo 1969a). The Notomegabalanus species closest geo- graphically to the California coast is N. concin- nus (Darwin, 1854) from the Peruvian province. This species was not included by Newman, but it has the sutural dentition, tergum, labrum, and protuberant lower margin of the first maxilla that characterize Notomegabalanus. Are these Cal- ifornia Neogene species true Notomegabalanus derived from some austral, perhaps South American ancestor? Does the Austromegabal- anus-Notomegabalanus complex represent an ancestral, previously worldwide stock from which tropical and warm temperate Megabala- nus s.S. evolved? Or do the California species represent a separate group exhibiting conver- gence of characters with the austral genera? At present it is possible only to consider the Cali- fornia species on the basis of their morphologies and to assign them to Notomegabalanus on the characters delimiting that taxon. The association of N.(?) insperatus with a cool temperate fauna does, however, suggest affinities with the austral megabalanid genera. Notomegabalanus(?) insperatus differs from all other megabalanid species in bearing distinct, prominent, external ribs. Megabalanus validus (Darwin, 1854) and M. zebra (Darwin, 1854) have ribbed shells, but their ribs are low, round- ed, and usually confluent, rather than erect and separate. In addition, both of these species have the sutural dentition typical of Megabalanus s.s. This new species is questionably assigned to Notomegabalanus rather than to Austromega- balanus solely on the basis of the presence of another Notomegabalanus, N. wilsoni, in the California Neogene. ACKNOWLEDGMENTS We thank Peter Rodda, California Academy of Sciences, for information concerning the geo- logic setting and stratigraphy of the Wilson Ranch beds and for his review of the manu- script; and Barry Roth, California Academy of Sciences, for his assessment of the age and de- positional environment of the Wilson Ranch molluscan fauna and for his review of the manu- script. LITERATURE CITED AppicotTtT, W. O. 1976. Neogene molluscan stages of Oregon and Washington. Soc. Econ. Paleont. and Mineral., Pacific Sec., Neogene Sympos., pp. 95-115. Bartow, J. A., A. SARNA-WoOJcICKI, W. O. ADDICOTT, AND K. R. Lasore. 1973. Correlation of marine and continental deposits in northern California tephrochronology. Am. As- soc. Pet. Geol. Bull. 57(4):769. BERGGREN, W. A. 1972. A Cenozoic time-scale—some im- plications for regional geology and paleobiogeography. Le- thaia 5:195—215. . 1978. Marine micropaleontology an introduction. Pp. 1-17 in Haq, B. U., and A. Boersma, editors, Introduction to marine micropaleontology. Elsevier, New York. Darwin, C. 1854. A monograph on the sub-class Cirripedia, The Balanidae, The Verrucidae. Ray Society, London. 684 p. DICKERSON, R. E. 1922. Tertiary and Quaternary history of the Petaluma, Point Reyes and Santa Rosa quadrangles. Proc. Calif. Acad. Sci., ser. 4, 11(19):527-601.. Hicains, C. G. 1960. Ohlson Ranch Formation, Pliocene, northwestern Sonoma County, California. Univ. Calif. Publ. Geol. Sci. 36(3):199-232. Hoek, P. P. C. 1883. Report on the Cirripedia collected by H.M.S. Challenger during the years 1873-1876. Rep. Sci. Results Voy. H.M.S. Challenger, Zool. 8(25):1—169. . 1913. The Cirripedia of the Siboga-Expedition. B. Cirripedia Sessilia. Pp. 129-275 in Siboga-Expeditie 31b. E. J. Brill, Leyden. LINNAEUS, C. 1758. Systema Naturae. 10th Ed. Holmiae. 428 p. MENESINI, E. 1968. Balanus curvirostratus nuova specie del Pliocene della Toscana. Atti Soc. Toscana Sci. Nat. Mem. P-V, ser. A, 75(2):617-632. NEwMaAN, W. A. 1980. On the biogeography of balanomorph barnacles of the southern ocean including new balanid taxa; a subfamily, two genera and three species. Proc. Internatl. Sympos. Mar. Biogeogr. and Evol. Southern Hemisphere, New Zealand DSIR Inform. Ser. 137, 1:279-306. OsmontT, V. C. 1905. A geological section of the Coast Ranges north of the Bay of San Francisco. Univ. Calif. Publ. Bull. Dep. Geol. 4(3):39-87. Pitssry, H. A. 1916. The sessile barnacles (Cirripedia) in the collections of the U.S. National Museum; including a mono- graph of the American species. U.S. Natl. Mus. Bull. 93: 1-366. ZULLO & GURUSWAMI-NAIDU: NORTHERN CALIFORNIA MIOCENE BALANIDS 535 Ross, A. 1969. Studies on the Tetraclitidae (Cirripedia: Tho- racica): revision of Tetraclita. Trans. San Diego Soc. Nat. Hist. 15(15):237-251. Rot, B., AND R. GURUSWAMI-NAIDU. 1978. A new Mio- cene species of Nuttallia (Mollusca: Bivalvia) from the Sa- linas Valley, California. J. Paleont. 52(1):61-66. SARNA-WolcickI, A. M. 1976. Correlation of late Cenozoic tuffs in the central Coast Ranges of California by means of trace- and minor-element chemistry. U.S. Geol. Surv. Prof. Pap. 972:1-30. SCHUMACHER, C. F. 1817. Essai d’un nouveau systeme des havitations des vers testaces. Copenhagen. 287 p. TrAvis, R. B. 1952. Geology of the Sebastopol quadrangle, California. Calif. Div. Mines Bull. 162:1-33. VAN EysINGA, F. W. B. 1975. Geological time table, 3rd Ed. Elsevier, Amsterdam. WEAVER, C. E. 1949. Geology of the Coast Ranges immedi- ately north of the San Francisco Bay region, California. Geol. Soc. Am. Mem. 35:1-242. YAMAGUCHI, T. 1971. Fossil barnacles from the Pleistocene Miyata Formation. Sci. Rep. Yokosuka City Mus. 18:122-130. . 1977. Taxonomic studies on some fossil and recent Japanese Balanoidea. Trans. Proc. Palaeontol. Soc. Jpn, new ser. 107—108:135-201. ZuLLo, V. A. 1969a. Thoracic Cirripedia of the San Diego Formation, San Diego County, California. Nat. Hist. Mus. Los Angeles Cty. Contrib. Sci. 159:1-25. . 1969b. A late Pleistocene marine invertebrate fauna from Bandon, Oregon. Proc. Calif. Acad. Sci., ser. 4, 36(1):347-361. . 1979. Thoracican Cirripedia of the Pancho Rico For- mation, Salinas Valley, Monterey County, California. Nat. Hist. Mus. Los Angeles Cty. Contrib. Sci. 303:1—-13. CALIFORNIA ACADEMY OF SCIENCES Golden Gate Park San Francisco, California 94118 INDEX TO VOLUME 42 (Compiled by Lillian J. Dempster) New names in boldface type. Abraliopsis 425, 428 Apristurus brunneus 414, 419 Acacia 76, 82 Apterichtus flavicaudus 57-58, 63, 66 Acanthurus triostegus 65 Araneae 318 Achillea Arceuthobium Millefolium 20, 24, 31, 33, 45 cyanocarpum 3, 20, 24 sp. 20, 24 sp. 24 Acmaea 324-325, 328 Archaeogastropoda 323 striata 337 Archoleptoneta schusteri 318 Acmaeidae 323-339 Archytas 10 Acmonotus 190 Arctocephalus townsendi 411 chilensis 190 Arctophyto 11 Acyphoderes 70, 72-73, 80 Arctophyto-Milada 11 acutipennis 73, 78-79 Arctostaphylos 384 amoena 69, 73—76 sp. 24 aurulenta 72 Ardistomina 458 cribricollis 73-74 Artemia 214 delicata 73, 78 Aruncus sp. 23 delicatus 78 Ascothoracica 443 fulgida 69, 73, 76-77 Ascothoracida 443-444 parva 69, 73, 77, 79-80 Ascothorax 444 prolixa 69, 73, 78 Aster sp. 24, 45 sexualis 73, 76-77 Asteroidea 443 suavis 73, 78, 80, 82 Ateloglossa 9, 11-12 velutinus 72 Austromegabalanus 533-534 vespiventris 72 Austromegabalanus-Notomegabalanus complex 534 yucateca 73, 79 Aglaia fidelis var. minor 398 Baccharis 3 Aglaophenis latirostris 427 glutinosa 20, 49 Agonidae 429 pilularis 20, 33 Agonina 458 sp. 20, 24, 33, 49 Allium 20 Balanidae 525, 527 Alosa sapidissima 410 Balaninae 527 Amblyarrhena 305-306 Balanoidea 527 Anaphalis sp. 45 Balanus 529 Andira inermis 72 balanus 525, 529, 531 Anillina 455, 458-459, 494 balanus complex 525, 529 Anillus integripennis 485, 487 irradians 525, 529-531 Anoplopoma fimbria 423 nubilus 525, 527, 529 Antimora 341-348 proxinubilus 525, 529 australis 341-343, 348 rostratus 529, 531 meadi 341-342, 348 Sp» 551169533 microlepis 341-344, 347 sp. aff. B. nubilus 525, 527-529, 531 rhina 341-343, 347 sp. cf. B. irradians 531 rostrata 341-344, 346-348 Baltimora 80 spp. 342 Banksula 315-322 Aotea acus 61 californica 315-317, 322 Aphonus 4 californica group 316-319 pyriformis 50 elliotti 315-317, 319-322 Apistus 287 galilei 316-317, 322 Aploactinidae 287 grahami 315-317, 320-322 Aploactis 288 grubbsi 315-317, 319-320, 322 [537] 538 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42 martini 317 martinorum 315-319, 322 melones 315-317, 320-322 melones group 316 rudolphi 315-318, 322 tuolumne 316-317, 332 ?Belone caudimaculata 301 Bembidiina 455, 458 Bembidiini 455—456, 458-459 Bembidion 455-456, 458-460, 462—464 aeger 455, 460, 462-466, 470, 473, 475-476, 478, 491-492 affine group 460, 482 aratum 460-464, 478-480, 491, 493 armuelles 455—460, 462-465, 467, 481-482, 491, 493 barrensis 455, 460-464, 467, 470, 482-483, 491, 493 championi 460-464, 467, 483-484, 491, 493 championi group 460, 483 chiriqui 455, 460, 462-466, 470, 476, 478, 491-492 cognatum group 460, 484 cortes 455, 460, 463-465, 467, 470, 478, 480-481, 491, 493 cyclodes 460-464, 483, 491, 493 (Cyclolopha) 483 definitum 479 diabola 455, 460, 462-466, 470, 473-475, 478, 491- 492 dorsale group 460, 480 edwardsi 455, 460, 462-466, 470, 473, 475-476, 491- 492 (Eupetedromus) 479-480 franiae 455, 460-464, 467, 470, 477, 479, 491, 493 franiae group 460, 477 (Furcacampa) 482 (Hydromicrus) 465 incrematum group 460, 479 ixtatan 455, 460-464, 467, 470, 478, 484-485, 491, 493 lavernae 455, 460, 462-466, 472-473, 475-476, 491- 492 mexicanum 460, 462-466, 469-471, 478, 491-492 nahuala 455, 460-464, 467, 470, 477-478, 491, 493 nahuala group 460, 476-477 (Notaphus) 479-480 (Peryphus) 466, 468, 477 purulha 455, 460, 462-467, 470, 491-492 quetzal 455, 460, 462-466, 470, 473-474, 478, 491- 492 rogersi 460-464, 466-468, 470, 478, 491-492 rogersi group 460, 467 sanctaemarthae 467 satellites 460, 462-466, 470-472, 478, 491-492 semistriatum group 460, 465—466 sparsum 460-464, 467, 478, 481-482, 491, 493 subapterum 467 _ transparens group 484 vernale 457, 460, 462-466, 469-472, 478, 491-492 vernale group 460, 467-469, 477 versicolor group 482, 484 vinnulum 479 vulcanium 460, 462-466, 470, 474-476, 478, 491-— 492 Bembidium 459 affine 482 aratus 479 aztecanum 485 championi 483 cyclodes 483 mexicanum 469 rogersi 467 sallaei 469 satellites 471 scintillans 479 semistriatum 465 sparsum 481 sphaeroderum 483 tetracolum 468 vernale 469 Bigelovia sp. 24 Bixa 70 Blenniidae 429 Brachysomophis 64 henshawi 57, 59, 64, 66 sauropsis 57, 59, 64, 66 Bromiades 70, 72 brachyptera 72 meridionalis 72 Bryx 182, 184, 192-193, 197-199, 220, 222 arctus 181, 184, 193-194, 196, 220-221, 223 balli 193, 220 banneri 193, 220 (Bryx) 192-193, 197, 220, 222 (Bryx) clarionensis 198 (Bryx) veleronis 197-198 clarionensis 181, 184, 193, 197-199 coccineus 181, 184, 193-194, 196, 220-221 darrosanus 193, 220 dunckeri 184, 197 heraldi 181, 192-194, 196, 220 hildebrandi 193, 196 randalli 197 (Simocampus) 192—193, 195-196, 220, 222 (Simocampus) arctus 193, 195-196 (Simocampus) balli 193 (Simocampus) banneri 193 (Simocampus) coccineus 193, 196-197 (Simocampus) darrosanus 193 (Simocampus) heraldi 192—195 (Simocampus) /ildebrandi 193 veleronis 181, 184, 192-193, 197-199 Buddleia 80, 82 wrightii 80 Bulimulidae 399 Byrsonima 70 INDEX Cacalia decomposita 24, 26 Caecula platyrhyncha 63 (Sphagebranchus) flavicauda 63 (Sphagebranchus) platyrhyncha 63 Calathea 483 Callechelys galapagensis 63, 66 luteus 57-58, 63, 66 Callionymidae 349 Callionymus 349-353, 357, 359-361, 363-364, 366, 369, S7ile S735 375 africanus 349-354, 362, 364-366 altipinnis 349-352, 355, 373-375 bentuviai 349-350, 352, 354, 366-367 calauropomus 359 (Callionymus) 349-350, 364 (Calliurichthys) 349 carebares 349-351, 353-354, 356-357, 376 curvicornis 370, 373 formosanus 349-352, 355, 369-370 guentheri 349-353, 355, 361, 370-373 huguenini 373, 375 indicus 349 kaianus 349-353, 355, 357-361, 363-364, 366, 369— 3705 37/2—3715, 375 kKaianus-group 349-377 kaianus moretonensis 349, 359 kotthausi 349-352, 354, 362-364 lyra 349 maculatus 349 moretonensis 349-353, 355, 358-360, 369, 376 ochiaii 349-352, 355, 366-369, 373, 375 regani 349-351, 353-354, 360-363, 366, 372-373 sokonumeri 349-352, 355, 375 sp. 375-376 (Spinicapitichthys) 349 whiteheadi 349-350, 352, 354, 359-362, 372 Callorhinus 412, 414-415, 417-418, 428 ursinus 411, 423 Camaenidae 399 Caprifoliaceae 45 Carabidae 87, 97, 435, 455 Carabus dentellus 479 gyllenhali 87, 111 litorale 468 lividus 113 nivalis 87, 111 rufescens 111 rufescens gyllenhali 111 varius 480 Careproctus 421 Casearia 72, 80 nitida 77 Ceanothus 46 americanus 45 fendleri 24 sp. 24, 25 Cephaloscyllium 418 Cerambycidae 69, 84 Cetonurus 135, 142-143, 147 Chaenopleura 309-312 Characidae 499 Charis 71 euphrosyne 71 Charisia 71 nigerrima 71 Chauliodus eximius 218 Chilara 418, 421 taylori 409 Chiroteuthidae 427 Chiroteuthis 426, 428 sp. 421 Choeroichthys 222 brachysoma 184 Chrysanthemum leucanthemum 45 Chrysothamnus greeni 24 viscidiflorus 20, 33 Cicindela quadrimaculata 459 Cirrhimuraena macegregori 64 playfairii 57, 59, 64, 66 Cirripedia 443, 525, 527 Cirsium sp. 27, 45, 51 Clevelandia ios 219 Clinidae 429 Clupea harengus 423 harengus pallasii 416 Clupeidae 299 Coccoloba 72 Coecilophis 62 Coelocephalus 144 acipenserinus 144 Coelorinchus 135-136, 144, 156 Coleoptera 69, 87, 122, 435 Collembola 321 Collisella 324-326, 328, 336-337 acutapex 328 atrata 326 discors 326 mitella 326, 328, 337 pediculus 326, 335-336 pelta 332 triangularis 328 turveri 328 Cololabis saira 423 Compositae 20, 23-26, 33, 41-42, 45, 49, 51 Convolvulaceae 46 Cordia 70, 72 Coryphaenoides 135 garmani 151-152 misakius 153 539 540 Corythoichthys 222 flavofasciatus 184 intestinalis 223 sp. 184 ‘“‘Corythoichthys”’ 222 Cosmocampus 192, 222 brachycephalus 184, 223 Cottidae 429 Crago sp. 414, 417-418 Cremanium 303, 311 Croton 74-75, 82 sp. 49 Cruciferae 31, 49 Crustacea 443 Cymatogaster aggregata 416 Cypriniformes 497, 500 Cystophora 429 Dalea albiflora 49 sp. 24 Daucus carota 45 Delphinidae 411 Delphinus 412-413, 416-418, 425, 428 delphis 411 ,425 Dendrogaster 443-444, 447-448, 450, 453 arbusculus 443-448, 450, 452-453 cf. arbusculus 448, 450 arctica 450 astropectinis 448 beringensis 448 dogieli 448 fisheri 443-444, 448-450, 452 iwanowi 448 ludwigi 448, 450, 453 murmanensis 447 okadai 448, 453 punctata 443-444, 450-453 repertus 447 rimskykorsakowi 453 tasmaniensis 448 Dendrogastridae 444 Dentirostrum 222 Janssi 184 Dermatostethus 198 punctipinnis 185, 198 Dexia 16 albifrons 1, 42, 44 canescens 38 cerata 1, 42, 44 halone 39 harpasa 24, 37, 41 prexaspes 40 rufipennis 36, 42, 44 Dexiosoma fumipennis 1, 48 Dinera 8, 11-12, 42 robusta 1, 41-42 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42 Diplogrammus 350, 364 africanus 350, 364, 366 indicus 350, 363-364 Diptera 1, 14 Dolichocodia 11 Doryichthys californiensis 188 Doryrhamphus 182, 184, 187-189, 222 californiensis 188 excisus 187, 190 melanopleura 181, 184, 187-190, 221 melanopleura pleurotaenia 189 negrosensis 187 paulus 181, 188-190, 221 Dosidicus gigas 426 Dunckerocampus 222 dactylophorus 184 Echidna uniformis 61 Echinomacrurus 135, 142-143 mollis 142, 144 occidentalis 135, 138, 142-144 Efferia aestuans 41 Elops 295 Embiotoca jacksoni 417 Embiotocidae 416—417 Encelia farinosa 24 sp. 24 Engraulis 409, 421 mordax 409 Enhydra lutris 411 Enneanectes sexmaculatus 213 Enoploteuthidae 427 Entelurus 218 Eogastrophinae 216 Eopsetta 421 Epimelitta 70-72 aureopilis 71-72 meliponica 71 nigerrima 71 Eptatretus 418 stoutii 415, 418 Ericaceae 24 Eriogonum 31 elatum 33 fasciculatum 36 nudum 20 sp. 20, 24, 31 Estheria 16 tibialis 16-17, 36-37 Eumetopias 410, 412, 414-415, 417-421, 428-429 Jubatus 409, 411, 419-421 Eupatorium 41 coelestinum 42 perfoliatum 45 Euphorbiaceae 49 INDEX Fabaceae 24, 42, 45, 49 Farlapiscus 185 Forsteronia 70, 72 Fragaria 107 Gadidae 176 Gadiformes 341 Gadus macrocephalus 417 merlangus 423 morhua 423 Garrinae 499 Gastromyzon 497-515, 523 borneensis 497-504, 509-512, 514 contractus 497-502, 504—S05, 512, 514-515 ctenocephalus 497-498, 501-502, 505-507, 514 fasciatus 498-502, 507-510, 514 lepidogaster 497, 501-502, 508-510, 514 megalepis 497, 499, 501-502, 509-511, 514 monticola 497-499, 501-504, 511-512, 514-515 punctulatus 501-502, 512-515 ridens 497, 501-502, 504, 512-515 Gastromyzontinae 497-501, 516 Gastrophori 184, 218, 222-223 Gastropoda 379 Geocharidius 455-456, 458, 485, 487 gimlii 455, 485, 487-488, 491, 493 integripennis 485, 487-488, 491, 493-494 phineus 455, 485-487, 489-491, 493-494 romeoi 455, 485, 487-488, 491-492 tagliantii 455, 485, 487-488, 491, 493-494 Geraea canescens 24 Glaniopsis 497-500, 515-519, 521-523 denudata 497, 516-518 gossei 497, 516, 518-519 hanitschi 515-516, 518-523 multiradiata 497, 515-516, 518, 520-523 Globicephala 427 Glyptocephalus 418, 421 zachirus 417 Gonatidae 426—428 Gonatus 425-426 sp. 425, 428 Gossypium sp. 49 Grampus 412-413, 417-418, 427 griseus 411, 426-427 stearnsii 426 Gutierrezia sarothrae 25 sp. 24 Gyropleurodus quoyi 282 Halieutichthys 235 Haloporphyrus rostratus 314-343, 347 viola 341-343, 347 Haplopappus sp. 20 Helianthus sp. 24 Heliopsis parvifolia 24-26 sp. 25 Helix fidelis 379, 381 hillebrandi 382 Hellenium hoopesii 27 Helminthoglyptidae 400, 402 Helobia castanipes 87, 111-112, 115 Hemilepidotus hemilepidotus 423 Hemithylacus Petersi 190 Heraldia 222-223 nocturna 184 Hesperodinera 8, 10—12 Hexagrammos decagrammus 409, 417 lagocephalus 416 Hippasteria californica 443-444 Hippocampus 182-187, 22i—223 breviceps 185 ecuadorensis 186 gracilis 186 heptagonus 185 hildebrandi 181, 186—187 ingens 181, 184-187, 197, 221, 224 kuda 184 reidi 187 spinosissimus 185 Hippoglossoides elassodon 416 Hippohystrix 185 Hipposyngnathinae 216-217 Hipposyngnathus 216-218, 221 convexus 217-218 imporcitor 181, 217-218 neriticus 218 Histioteuthidae 427 Histrio histrio 185 Homalopteridae 497 Horologion 455, 459 speokoites 459 Hydrolagus 418 Hymenocephalus 135-137, 140, 149 gracilis 140 (Hymenogadus) 140 (Hymenogadus) gracilis 140 (Hymenogadus) tenuis 140 italicus 140 kuronumai 140 sp. 137, 140-141 (Spicomacrurus) 140 (Spicomacrurus) kuronumai 140 tenuis 140 Hymenogadus 140 Hypsoblennius gentilis 219 Hystrichodexia 16 contristans 16, 25-26 Ichthyapus vulturis 57-58, 63, 66 541 542 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42 Ichthyocampus 222 belcheri 184 Icichthys lockingtoni 425 Ilypnus gilberti 219 Ipomoea 46 Jatropha 74, 80, 82 curcas 75 Jenkinsiella macgregori 64 Kogia 412-413, 417-418, 427 breviceps 427 simus 411, 427 Kumba 135, 147 Labeoinae 499 Labiatae 49 Laemonema 425 Lagenorhynchus 409, 412-413, 416-418, 425-426, 428, 430 obliquidens 409, 411, 425-426 Lampetra tridentata 418 Leiuranus semicinctus 59, 66 Lepidium thurberi 49 Lepidoglanis 500 monticola 500, 511 Lepidospartum squamatum 20 Leptasterias groenlandica 450 Leptenchelys labialis 62 Leptocottus armatus 416 Leptonotus 181-182, 184, 190, 192, 221-222 blaenvillianus 190 blainvilleanus 181, 184, 188, 190-192, 221 Blainvillei 190 blainvillianus 190 elevatus 192 norae 192 semistriatus 192 Leucichthys 424 Liliaceae ‘20 Liocranoides sp. 318 Lionurus 156 cetonuropsis 147 liolepis 157 (Lionurus) barbiger 157 (Lionurus) fragilis 147 (Lionurus) liolepis 157 (Nezumia) convergens 171 (Nezumia) cuspidatus 171 (Nezumia) latirostratus 169 (Nezumia) loricatus 174 (Nezumia) orbitalis 167 (Nezumia) pudens 163 (Nezumia) stelgidolepis 160 (Nezumia) trichiurus 171 nigromaculatus 153 stelgidolepis 160 ventralis 167 Lippia wrightii 24 Loligo 409, 413, 418, 424-425, 430 opalescens 421-422, 424, 428-429 Lonicera sp. 45 Lophiiformes 229 Lophius histrio 243 (Malthe) cubifrons 258-260 nostratus 274 radiatus 258-260 rostratus 251-253, 260, 274 vespertilio 234, 247, 251-252 Loranthaceae 24 Lottia 323-326, 333-334, 337-338 gigantea 325-326, 332, 335-336 mesoleuca 323, 325, 336-338 mimica 323, 325-326, 332-338 mimica-smithi 335, 337 smithi 323, 326, 333, 334-338 stipulata 336 Lycodopsis pacifica 409, 417 Lymnastis 459 Lyopsetta 418, 421 Macrocystis 206 angustifolia 207 pyrifera 207, 220 Macrodactylus 4 subspinosus 1, 4, 42 Macrometopa 49 calogaster 48 Macrouridae 135-136, 177, 427 Macrourinae 135 Macrourus 135 laevis 149 lucifer 152 nigromaculatus 152-153 nigromarginatus 152 serratus 156 Macruroplus ori 152 potronus 153, pudens 163 pulchellus 159 ?Macruroplus 156 nigromaculatus 153 Macrurus 140, 149 barbiger 157-159 convergens 171, 173 cuspidatus 171, 173 fasciatus 152 flagellicauda 147 fragilis 135, 147 gracillicauda 160 (Hymenogadus) 140 latirostratus 168 liolepis 157, 159 (Lionurus) liolepis 157 [55076 INDEX loricatus 174 (Malacocephalus) 149 (Malacocephalus) laevis 149 (Mystaconurus) 140 orbitalis 167 parvipes 147 stelgidolepis 160 tenuicauda 145 trichiurus 171, 173 Malacocephalini 157 Malacocephalus 135, 149, 151, 157 hawaiiensis 149 laevis 137, 149-151 nipponensis 149, 151 sp. 149 Mallotus villosus 423 Malthaea 234 angusta 243, 246-247 cubifrons 258 longirostris 247, 250 nasuta 247, 251, 258, 267 notata 243, 247 truncata 243, 247 Malthe 234 angusta 243 cubifrons 258, 260 nasuta 251, 253, 258 notata 243 radiata 258 truncata 243 vespertilio 234, 243, 247, 258, 274, 276 vespertilio nasuta 258 vespertilio radiata 258 Malthea 234 guacucuja 247 nasuta 258 notata 243 radiata 258 vespertilio 251 Malthopsis 234 Malvaceae 49 Maroubra 218, 222 perserrata 184 Marrubium vulgare 49 Mataeocephalus 135, 144-145 acipenserinus 146-147 adjustus 135, 144-145, 147 microstomus 147 nigrescens 146-147 tenuicauda 138, 145-147, 177 Maxillicosta 221 Medicago sativa 49 Megabalaninae 531 Megabalanus 533-534 tintinnabulum 533 tintinnabulum complex 533 validus 534 wilsoni 534 zebra 534 ““Megabalanus’’ wilsoni 534 Melastomataceae 303, 305 Melilotus alba 42, 45 sp. 24 Melolonthinae 3 Merluccius 409 productus 409, 416-418 Mesobius 135, 141 antipodum 141-143 berryi 137, 141-143 Miconia 303-314 andreana 306 chionophylla 310-311 chiriquiensis 303-305 coloradensis 303, 305—307 concinna 303, 305, 307-309, 314 confertiflora 303, 309-312 gibba 306-307 grandidentata 303, 311-312 longibracteata 303, 308-309, 312-314 micrantha 307 myrtillifolia 308 parvifolia 311 rubens 305, 308 superposita 314 tabayensis 307 wittii 307 Micratopus 459 Microdonophis 64 erabo 65 fowleri 64-65 macgregori 64 polyophthalmus 64 Microgadus 409 proximus 416-417 Micrognathus 222-223 balli 192 brevirostris 184 crinigerus 214 Microphis extensus 189 Microstomus 421 Milada 11 Mioptachys 458 Mirounga 412, 414, 417-418 angustirostris 411, 418-419 Mochlosoma 5-12 Mollusca 323, 423 Monadenia 379-383, 385, 390, 394, 396, 398-403 antecedens 401 chaceana 380, 394, 397, 399 543 churchi 378-380, 382-383, 385, 387-388, 395-396, 398 circumcarinata 400, 402 (Corynadenia) 379-380, 382-383, 400-403 (Corynadenia) hirsuta 382 544 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42 (Corynadenia) mormonum 385, 401 cristulata 382 dubiosa 401 fidelis 378, 382-383, 387, 390, 398-401 fidelis leonina 399 fidelis minor 398-399 fidelis pronotis 400 fidelis scottiana 400 hillebrandi 399, 402 infumata 382 marginicola 401 marmarotis 382 (Monadenia) 379-380, 382-383, 400—403 (Monadenia) fidelis 402 (Monadenis) setosa 402 mormonum 385, 399 mormonum cala 398 mormonum group 401 setosa 382-383, 399-400, 402 (Shastelix) 379, 381-383, 395, 400-403 (Shastelix) churchi 382—402 (Shastelix-Corynadenia) stock 401-403 (Shastelix) troglodytes chaceana 397-399 (Shastelix) troglodytes troglodytes 395-396 (Shastelix) troglodytes wintu 396-397 troglodytes 379-407 troglodytes chaceana 379-380, 389, 391-394, 397— 399, 406 troglodytes wintu 379, 383, 386-389, 391-392, 394, 396-398, 406 Moridae 341, 425, 427 Morone saxatilis 410 Moroteuthis sp. 429 Muraena maculosa 63 Muraenichthys 61, 221 breviceps 61 chilensis 60 cookei 57-58, 63, 66 devisi 61 gymnotus 58, 66 labialis 62 laticaudata 63 macropterus 57, 61 ogilbyi 61 owstoni 61 puhioilo 57—61, 65—66 schultzei 58, 66 (Scolecenchelys) 61 Mustelus californicus 424 Myceropsis 11, 15, 54 Myctophidae 424—425 Myliobatidae 418 Myoxocephalus sp. 416 Myrichthys 63 bleekeri 57, 59, 64, 66 maculosus 57, 59, 63-64, 66 magnificus 63—64 pantostigmius 57, 63 stypurus 64 tigrinus 57, 63 xysturus 57, 63 Myriocladus 444, 448 arbusculus 444 Myrophinae 57-58, 62 Nebria 87-133, 435-442 acuta 98, 131 acuta quileute 87, 98—99, 122, 126, 128, 131 arkansana 100, 112, 131 arkansana edwardsi 87, 100-102, 125-126, 128, 131 arkansana ooowah 87, 102, 126, 128, 131 arkansana uinta 87, 102-103, 125, 127, 131 bifaria 87, 96, 112 carri 87, 105, 121, 123, 132 castanipes 114 catenata 87, 112 columbiana 108 danmanni 435—440, 442 darlingtoni 87, 104, 123, 132 diversa 112-113 elias 90, 112 eschscholtzii 87, 90, 112, 115 expansa 87, 112 fragilis 131 fragilis fragilis 131 fragilis teewinot 87, 103, 127, 131 frigida 116 fusiformis 112 gebleri 87, 105, 113, 132 gebleri cascadensis 87, 105—106, 123, 132 gebleri fragariae 87, 106-107, 123, 127-128, 132 gebleri gebleri 132 gebleri rathvoni 115 gebleri siskiyouensis 87, 107, 123, 132 gouleti 87, 94-95, 121-122, 129 gregaria 87, 113 gyllenhali 90, 97, 111, 130 gyllenhali castanipes 111-112, 114-115, 130 gyllenhali gyllenhali 111 gyllenhali lassenensis 87, 96—97, 121, 135 gyllenhali lindrothi 87, 97-98, 121, 130 hudsonica 87, 113 incerta 87, 113 ingens 87, 113 kincaidi 107, 132 kincaidi balli 87, 107-108, 121, 125, 132 kincaidi kincaidi 132 lacustris 94-95, 112 lacustris bellorum 87, 95—96, 122, 129 lacustris lacustris 129 lituyae 87, 100, 124, 131 livida 87, 112-113 longula 87, 113 mannerheimii 87, 114-115 INDEX 545 meanyi 109, 133 Neogastromyzon 497, 499-501 meanyi giulianii 435-439, 441-442 nieuwenhuisi 499, 501 meanyi lamarckensis 87, 109, 125, 133 pauciradiatus 493, 505 meanyi meanyi 133 Nerophis 182, 218, 222-223 meanyi sylvatica 87, 109, 124, 133 Nezumia 135, 144-145, 151-152, 156-160, 163, 165, melsheimeri 90 167, 169, 171, 173-174, 176 metallica 87, 114 aequalis 168 moesta 87, 114 africana 158, 163 molbis 114 atlantica 158, 163 mollis 87, 114 bubonis 158, 163 muehlenbergii 114 burragei 144, 158, 171 muiihlenbergii 114 condylura 156 navajo 87, 104-105, 127, 132 convergens 139, 156-157, 169-177 nivalis 97, 111, 130 convergens-loricata-orbitalis complex 176 nivalis bifaria 96 cuspidata 171 nivalis gaspesiana 87, 96, 122, 130 cyrano 171 nivalis nivalis 112, 114, 130 dara 158, 171 obliqua 87, 113-114, 116, 132 ectenes 171 obliqua chuskae 87, 105, 123, 125, 132 gracillicauda 160 obliqua obliqua 132 hebetata 158, 171 obtusa 87, 114-115 holocentrus 158, 171 oregona 87, 115 kamoharai 158 ovipennis 87, 115 latirostrata 138, 140, 146, 156-157, 168-177 pallipes 114-115 latirostratus 171 picicornis 113 liolepis 137-139, 154-159, 162-163, 165, 177 rathvoni 87, 115, 132 loricata 135, 139-140, 156-157, 171-172, 174-176 sahlbergi 97, 100 loricata atomos 135, 169, 174-177 sahlbergii 87, 97, 115-116, 131 loricata loricata 135, 174-175 sahlbergii modoc 87, 99, 123, 131 macronema 158 sahlbergii sahlbergii 116, 131 nigromaculata 153 sahlbergii triad 87, 99, 123, 131 obliquata 171 schwarzi 133 orbitalis 139, 156-157, 164, 167-168, 172-173, 176 schwarzi beverlianna 87, 110, 124-125, 133 parini 138, 141, 156-157, 162, 176-177 schwarzi schwarzi 133 propinqua 167, 171 sonorae 435—442 proxima 171 spatulata 112 pudens 139-140, 156-157, 160, 163-165, 177 spatulata sierrae 87, 108-109, 121, 133 pulchella 139, 154-157, 159-160, 163, 165, 177 spatulata spatulata 112, 133 sclerorhynchus 165, 167 suturalis 87, 96, 113, 115 spp. 163 tenuipes 87, 116 stelgidolepis 139, 154-158, 160-163, 165, 173, 176— texana 116 177 trifaria 87, 116, 133 suilla 165, 167, 173 trifaria catenata 112, 133 tomiyamai 171 trifaria trifaria 133 trichiura 171 trifaria utahensis 87, 90, 110-111, 121, 124, 127- ventralis 135, 157, 164-167, 177 128 Nimioglossa 10-12 trifaria vandykei 109 Notoacmea 323-326, 328, 330, 337-338 turmaduodecima 435—440, 442 vandykei 87, 109, 116, 133 vandykei vandykei 133 vandykei wyeast 87, 109-110, 121, 133 violacea 87, 115-116 fascicularis 325 filosa 323, 330, 332, 337-338 immaculata 323, 325, 329-332, 335-336, 338 insessa 337 viridis 87, 116 persona 332 zioni 132 pumila 323, 325, 327-329, 335-336, 338 zioni oasis 87, 103-104, 121, 132 rothi 323, 325, 329-332, 335-336, 338 zioni zioni 132 rothi-immaculata complex 332 Nebriini 87-133, 435-442 ubiquita 323, 325-329, 335-336, 338 546 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42 Notomegabalanus 533-534 algicola 534 concinnus 534 wilsoni 534 Notomegabalanus(?) insperatus 525, 531-534 Novacampus 190 Octopoteuthidae 426-428 Octopus 409, 425, 429 sp. 415, 417, 421, 428 Odontocera 70, 80-91 aurocincta 81 aurocincta arizonensis 82-83 aurocincta aurocincta 81-83 aurocincta var. nigrocapicalis 81 brachyptera 72 carinicollis 82 clara 81-83 fuscicornis 81 vitrea 81 yucateca 79 Ogcocephalidae 229-231, 234, 243 Ogcocephalus 229-285 corniger 229, 235-241, 245, 247-249, 254, 261, 265, 268, 270, 272, 274-276 cubifrons 229, 231-233, 235-240, 242, 245, 247- 249, 252-255, 257-268, 274, 276 darwini 229, 234-240, 243, 245, 248, 279-283 declivirostris 229, 235-241, 245, 248, 261-262, 264— iD 21O= 27, nasutus 229, 235-240, 242-243, 245-261, 264-265, 268, 272, 274, 282 nasutus var. cayennensis 243 notatus 229, 235-249, 254, 265-267, 270, 272—273 pantostictus 229, 235-240, 242, 245, 247-249, 253- 255, 257, 260-262, 264—268, 270 parvus 229, 235-241, 243, 245, 248, 254, 260-261, 265, 268, 270-272, 274, 276-279 porrectus 229, 235-240, 243, 245, 248, 279-283 pumilus 229, 235-243, 245, 247-248, 254, 270-274, 276 radiatus 229, 243, 246, 252, 259, 264 rostellum 229, 235-240, 242, 245, 248-249, 253- 254, 256-257, 260-262, 266-269, 274 sp. 256, 269, 274, 276 truncatus 243 vespertilio 229, 235-240, 242-251, 253, 255, 259, 263-265, 270, 274, 276, 282 Ommastrephes 424 Ommata 70, 80, 83-84 championella 84 cyanea 84 (Eclipta) championella 84 (Eclipta) cyanea 84 elegans 83 (Ommata) rubriventris 84 sallaei 83-84 Onchocephalus nasutus var. cayennensis 243 radiatus 243, 258 Onchococephalus 234 Oncocephalus 234 longirostris 247 notatus 243 parvus 276 porrectus 282 radiatus 243, 259 truncatus 243, 246 vespertilio 252, 274 Oncorhynchus sp. 423 spp. 410 Onychoteuthidae 427 Onychoteuthis 425—426, 428 sp. 421, 425 Oostethus 184-185 brachyurus 184-185 lineatus 185 Ophichthidae 57-67 Ophichthinae 57-58, 63 Ophichthus 62, 64 (Coecilophis) 62 erabo 57, 59, 65-66 garretti 65 kunaloa 57, 59, 61-62, 65-66 pacifici 62 polyophthalmus 57, 59, 64-66 stypurus 63 urolophus 62 Ophichthys playfairii 64 polyophthalmus 64 Ophiodon elongatus 417 Ophisurus fasciatus var. semicinctus 64 semicinctus 64 Optonurus atherodon 152 Oreohelix 395 elrodi 400 Osmeridae 416-417 Ostariophysi 497 Pagophilus 429 Palpigradida 318 Pandalopsis 427 Pandalus 427 Paracetonurus 135, 147, 149 fragilis 138, 147-149 Paralabrax maculatofasciatus 220 Parapercis 221 Parasyngnathus 222-223 elucens 184, 223 Paratachys 455, 459 Parhomaloptera 497, 500 INDEX Parophrys 421 vetulus 416 Pasiphaea 427 Patella scurra 326 striata 337 Patelloida 324 semirubida 328-329 striata 337 Paullinia 70 Pedicellaster magister megalabis 443-444, 449 Penetopteryx 222 taeniocephalus 184 Phaenomonas cooperae 57-58, 63, 66 Phanerodon furcatus 417 Pharomachrus mocinno 473 Phoca 409-410, 412, 414-415, 417-418, 428-429 vitulina 409, 411, 415 Phocoena 412-413, 415-418, 428, 430 Phocoena 409, 411, 423-424 Phocoenidae 411 Phocoenoides 412-413, 416-418 dalli 411, 424-425 Phyllophaga 4, 14-15, 42, 49 farcta 49 rugosa 4, 37 Ryo B/G Se spp. 1 Phyllophichthus 64 xenodontus 47, 59, 64, 66 Phyllopteryx 222-223 foliatus 184, 223 Physeteridae 411 Pisces 57, 135, 181, 287, 349 Pisoodonophis magnifica 63 Platycephalidae 364 Platycephalus indicus 364 Platynus 458 Pleocoma 15 Pleurogrammus monopterygius 415 Pleuronectidae 409, 416-417 Poecilozonites 390, 400 bermudensis bermudensis 390 Polinices 428 Polistes instabilis 80 Pollicipes polymerus 427 Polyderidius 459 Polygonaceae 31, 33, 36 Polygyridae 401 Popillia 4 Japonica 1, 4, 37 Poraniopsis inflata 443-444 Porichthys 409, 418, 421, 427 notatus 409 Pristidoryrhamphus 187 Jacksoni 187 Problacmaea 324 Proctocanthus philadelphicus 45 Progastromyzon 497 Prokoenenia sp. 318 Prosena 9-12 siberita 3 Prosenini 2; 5—7, 9=11> 15-16 Prosenoides 11-12 Prosopodasys 287 asperrimus 287-290, 292 Protomyzon 497, 500 Pseudonezumia 135-136 Pseudophallus 222 elcapitanensis 184 starksi 184 Psocoptera 321 Psydrini 455 Ptilodexia 1-55 547 abdominalis 4, 36-37 agilis 3555105 12-)14=15 17-20) 24.644 agilis group 12-13, 16, 18-23 arida 5, 9-10, 12-13, 15, 19, 45-46 californica |, 6—7, 9, 12, 15, 18-19, 29-31 californica group 12-13, 29-36 canescens 4-8, 12, 14, 16-17, 37-39, 42 carolinensis 1, 4, 9, 11-12, 14-17, 19, 36-37, 42- 43 carolinensis group 12-14, 36-41 cerata 44 confusa 44 conjuncta 1, 3, S—8, 12-14, 23-24, 48-49 conjuncta group 12-13, 23-29 constrictans 26 contristans 1, 5—7, 10912=1385=165) 182527, dubia 44 elevata 50 flavotessellata 12-13, 15-17, 53-54 fumipennis 48 halone 5-6, 8, 10, 12, 14, 17, 39-40 harpasa 1, 4, 6, 9, 12-14, 18, 37-38, 41-44, 48, 50 harpasa group 12-13, 41-47 hucketti 1, 39 incerta 5—7, 12-14, 17-18, 49-51 leucoptera 1, 41-42 maculata 1, 4, 6-7, 10, 12-13, 15, 17, 51-53 major 1, 11-13, 15, 18, 46-50 major group 12-13, 47-54 mathesoni 5-6, 10, 12, 14, 17, 20-22 minor 1, 37 neotibialis 1, 36-37, 43 obscura 9, 12-14, 19, 22-23, 41 omissa 48 pacifica 1, 5, 7, 9-10, 12, 15, 19, 31-34 planifrons 5—6, 8—10, 12-13, 15, 18, 24-25, 37, 42 planifrons-contristans 5 ponderosa 5, 8, 12-13, 17, 46—47 prexaspes 4, 8-10, 12, 14, 17, 40-41 548 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42 proxima 1, 50-51 punctipennis 26 rufipennis 1, 5-10, 12-14, 19, 42-45 sabroskyi 1, 9, 12, 15, 18, 31, 34-36 scutellata 48 simulans 24 spp. 3 tibialis 11, 23, 41-42 tincticornis 48 westi 1, 5, 7, 10, 12-13, 15, 18, 27-29 Ptilodexia-Mochlosoma line 12 Pulmonata 379 Quietula y-cauda 213, 219 Raja 418 [?] Rana piscatrix 251-253 Rhacochilus sp. 416 Rhagium lineatum 4 Rhamnaceae 24, 45—46 Rhamphinina 8, 11-12, 15-16, 49 dubia 16, 48 major 47-48 Rhinotragides 69 Rhinotraginae 69 Rhinotragini 69-85 Rhodopetala 324 Rhynchiodexia 15-16, 44 arida 45—48 dubia 43-44 flavotessellata 15, 53 fumipennis 48 levata 50 mathesoni 20-21 omissa 48 ponderosa 46-47 robusta 41-42 tincticornis 47-48 Rhynchodexia 15-16, 44, 50 confusa 1, 37-38, 42-44 conjuncta 23 dubia | elevata 1, 50 fraterna 1, 48 omissa 1, 48 planifrons 24 punctipennis 1, 25—26 simulans 1, 23 translucipennis 1, 43-44 Riggia? 423, 428 Rosaceae 23 Rubus parviflorus 33 Rudbeckia sp. 24-25 Rutiliini 11 Salangidae 295, 297, 299, 302 Salicaceae 20 Salix 20 Saperda calcarata 4 Sargassum 201, 216 Scarabaeidae 14 Scaritini 458 Schismorhynchus labialis 57-58, 62, 66 Schultzidia johnstonensis 58, 66 Scomber scombrus 423 Scombridae 429 Scorpaenidae 409, 416-417 Scurria 323-326, 337-338 mesoleuca 326 scurra 325-326 stipulata 326 Sebastes 418 (Sebastomus) capensis complex 220 (Sebastomus) exsul-spinorbis complex 220 spp. 417-418 Senecio salignus 24 sp. 24, 27, 49 Senostoma 9, 11, 16 Serranidae 429 Siphonostoma 198 Siphostoma 198 aciculare 207 arctum 193, 195 auliscus 199 bairdianum 211 barbarae 211 blainvilliana 190 californiense 205 californiensis 205 carinatum 202 exile 204 griseolineatum 210 leptorhynchum 210 leptorhynchus 210 sinaloae 201-202 Sitalcina sierra 316 Solidago 3 canadensis 22-23, 42 juncea 37 microcephala 41 rugosa 42 sp: 203 24727, 33, 37, 45,501 trinervata 20, 25, 45 Sphagebranchus flavicaudus 63 vulturis 63 Sphagemacrurus 135, 149 fragilis 147 trichiurus 171 Sphagnum 466 Spiraea latifolia 23 Spirinchus 421 Spondias 72, 76 INDEX Squalus 418 Stanleya pinnata 31 Stenopseutes 70, 80 aeger 80 sericinus 80 Sundasalangidae 295-302 Sundasalanx 297, 299, 301-302 microps 295-302 praecox 295-302 sp. 300-302 Symbolophorus 423 Synchiropus 349-350 altivelis-group 350 calauropomus 360 phaeton-group 350 Syngnathidae 181-227 Syngnathiformes 181 Syngnathoides 222-223 biaculeatus 184, 223 Syngnathus 181-182, 184, 192, 198-202, 204-205, 207- 208, 213, 215, 218-220, 222 abbotti 211-212 acicularis 181, 190-192, 207 acus 184-185, 198, 202, 207, 222 agassizi 220 arcta 195 arctus 192, 195 arundinaceus 211, 214 auliscus 181, 184, 188, 199-202, 218, 220 avus 181, 217-220 bairdianus 185 banneri 192 barbarae 181, 210-212 blainvilleanus 190 blainvillei 190 blainvillianus 190 brevirostris 210-212, 215 (Bryx) 192, 199, 222 (Bryx) veleronis 192 californiensis 181, 184-185, 199-201, 202-207, 211, 218, 220, 223 californiensis avus 219 californiensis californiensis 204, 208 californiensis griseolineatus 210 californiensis leptorhynchus 210 carinatum 202 carinatus 181, 184, 199-204, 207, 219-220 carinatus-macrobrachium-folletti lineage 219-220 coccineus 192, 196-197 darrosanus 192 dimidiatus 210-211 dunckeri 192 emeritus 181, 217-218, 220 euchrous 181, 199-200, 206, 208-209, 211, 219-220 euchrous euchrous 201, 208-211, 220 euchrous intergrades 211 euchrous ollotropis 201, 208-211, 220 549 exilis 181, 199-201, 203-206, 219-220 floridae 214 folletti 203, 207, 219 fuscus 198, 220 griseolineatus 181, 205, 210 griseolineatus leptorhynchus 202 hildebrandi 192 hippocampus 185 independencia 196-197 insulae 181, 199-201, 209, 214-216, 219-220 leptorhynchus 181, 184, 199-202, 205, 208-216, 218-220 leptorhynchus-euchrous-insulae lineage 220 louisianae 214 macrobrachium 181, 199-201, 203, 207-208, 219 melanopleura 188 (Microsyngnathus), 192, 195, 199 norae 190 pelagicus 184-185, 198, 218, 221 phlegon 220 randalli 192 schlegeli 203-204, 214, 219 scovelli 214 spicifer 184-185 spp. 220 tweedlei 201-202 typhle 212 Syrictes 198 exilis 204 Tachinidae 1-2, 5, 9, 11, 36 Tachyina 455, 458-459 Tamaricaceae 49 Tamarix sp. 49 Tarletonbeania 423 Tectura 324-325 rosacea 328 Tetraclita 529 Thaleichthys pacificus 416 Theragra chalcogramma 416, 423 Thoracica 527 Thunnus alalunga 204 albacares 187 thynnus 187 Tomopterus 70 exilis 69-71 staphylinus 70 vespoides 70-71 Topobea 308 micrantha 303, 307-308 Trachonurus 135 Trechini 455 Trichodon 423, 427 Trigonia floribunda 77 Trogloneta paradoxa 318 Ulva 428 550 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 42 Umbelliferae 45 nigromaculata 138, 153-156, 177 Umbellularia californica 33 nigromarginata 152-153 Urocyclinae 399 ori 153 Urophori 184, 222-223 peterson 152 Usofila (Telema) sp. 318 (Sokodara) 135, 151-153 (Ventrifossa) 135, 151-152 Ventrifossa 135, 149, 151-153, 157, 163 Verbenaceae 24 atherodon 151-152 Verbesina encelioides 26 (Atherodus) 152 Verma flavicauda 63 barbiger 157 ctenomelas 152 Watasenia 424 divergens 152 fasciata 153 Xanthoxylum 74 garmani 152 Xenaploactis 287—293 lucifer 152-153 anopta 287, 289-291 (Lucigadella) 152 asperrima 289-290 (Lucigadus) 135, 149, 151-153 cautes 287—288, 291-292 (Lucigadus) nigromaculata 138, 153 (Lucigadus) sp. cf. ori 153 Zalieutes 234 macropogon 152-153 Zalophus 410, 412, 414-415, 417-419, 422, 428-429 misakia 153 californianus 409, 411, 419, 421-422 mucocephalus 152-153 i alah tad Phy » via bf ahh ea)p ee | yal, 1 gE F ’ iw si" ar i 4 A, hi ry a pigs py, PT AE a Dae ni Bcd: i. 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